(Featured image: Dr. Tom Thurow and Somalia National University students collecting rangeland composition data on the coastal plains of Central Somalia.)
Because Its rangelands are Somalia’s primary natural resource, a considerable amount of research has been carried out over the last several decades to determine their productive potential and how they may best be managed. Unfortunately, much of the resulting information was not widely published and is, therefore, now difficult to obtain. This post is the first of several to at least partially remedy this situation by making available digital copies of relevant publications and reports for free downloads
Featured image: Eleusine jaegeri, a large tussock grass, dominates the highland grasslands of the Ngorongoro Conservation Area. It is unpalatable except when young. Consequently, grazing pressure is confined to an underlying mat of palatable grasses (Andropogon, Cynodon, Digitaria, Sporobolus) which is kept low by constant usage.
Several months ago I made digital copies of the Range Management Handbook of Kenya available online. The response was so positive that I’ve decided to continue with other difficult-to-obtain publications on eastern Africa’s rangelands. Today I’m happy to announce the online availability of several digitized publications on the rangelands of Tanzania!
During the mid 1960’s and 70’s I was involved in, among other things, surveys of vegetation in Tanzania. Whenever possible, I also obtained copies of other surveys and studies, six of which I have digitized. Published between 1967 and 1978, they are now out of print and hard to obtain. Nonetheless, they may still have some value, be that for planning, instructional, research, or historical purposes.
Also, if you have paper copies of additional publications on the rangelands of Tanzania and would like me to digitize them and make them available, please send me an email.
Outside the house a prowling hyena whooped mournfully, waking me from my reverie. The fire which, last time I noticed, had been burning merrily away in the fireplace, was now.a bed of glowing coals. The short wave radio, having wandered off frequency, was whining and crackling with static. The feeble, slowly pulsating light from the Petromax lantern showed that it needed pumping. Again, the hyena whooped, but farther away this time. I looked at my watch and realized the lateness of the hour . . .
Buffaloes by My Bedroom: Tales of Tanganyika
Many years later, when I think about my nights at Ngorongoro, whether in the Crater Highlands, on the floor of Ngorongoro Crater, or somewhere on the Serengeti Plains, the first thing that comes to mind is the querulous oooo-WHUP (I am here) of a prowling spotted hyena.
The spotted hyena only occurs in Africa. (Graphic: Ngorongoro Hyena Project)
At the time I gave this little thought. But now, having investigated the matter, I know it was because the spotted hyena is not only Africa’s most abundant large carnivore, but also mostly hunts at night.
Something I did know, however, even then, was that the spotted hyena had an image problem. Less than handsome (although its small black cubs are cute), it has long been regarded as a skulking, slinking, nocturnal, weird-sounding, odd-looking, unclean scavenger too cowardly to prey on any but the weak and young. Furthermore, its role in disposing of human corpses encouraged the feeling that the animal is something of a living mausoleum. (Many East African tribes placed their dead, and in some cases, near-dead, in the bush for just this purpose.) Thus, it’s not surprising that the spotted hyena, whose behavior sometimes seems to verge on the demonic, also is associated with death and witchcraft. Consequently, it plays a more important role in African witchcraft than any other animal. (https://en.wikipedia.org/wiki/Cultural_depictions_of_spotted_hyenas#:~:text=In%20the%20culture%20of%20the,branded%20with%20an%20invisible%20mark.)
“Where’s my public relations person?“
(Photo: David Bygott)
That said, any spotted hyenas worried about their poor fan base can take heart from the fact that scientific research is casting them in a more positive light. Two attributes stand out: (a) high intelligence, and (b) speed and efficiency in utilizing prey carcasses. Together, these traits have made the spotted hyena a very successful carnivore. Details follow.
But first, two interesting spotted hyena facts.
Spotted hyenas look like dogs but are taxonomically more closely related to cats (most closely to genets and mongooses).
Hyenas are more closely related to mongooses than foxes, which, like dogs, are in the Suborder Caniformia.
(Graphic: Ngorongoro Hyena Project)
Dwarf mongooses.
(Photo: David Bygott)
Also interesting, and rather strange, is that males and females look so alike (females have a pseudopenis and false scrotum), that spotted hyenas were long thought to be hermaphrodites.
Female? Male?
(Photo by David Bygott)
Now, down to business:
Contrary to their reputation, spotted hyenas are highly intelligent, capable of outsmarting chimpanzees in laboratory problem-solving tests. Some everyday examples of this intelligence include (a) exceedingly cunning and suspicious behavior after escaping from traps, (b) use of deceptive behavior, and (c) an ability to plan for hunts of certain prey species in advance.
Regarding (b) and (c) above, Hans Kruuk, who studied spotted hyenas in Ngorongoro Crater and the Serengeti in the 1960’s, once observed a spotted hyena which upon finding a carcass, sounded the alarm call to keep other hyenas away, allowing it to keep the carcass for itself (Spotted hyena mothers sometimes show similar behavior by sounding alarm calls when other hyenas attempt ro kill their cubs). Hans also could often tell, from their behavior, when a group of hyenas had decided to hunt zebras, even when none were in sight and other prey were more easily available.
Other examples of spotted hyena intelligence occur throughout this report.
Complex behaviors reflect high intelligence, and spotted hyena behavior is the most complex of all African carnivores. An aspect to this is a high degree of behavioral flexibility. For instance, hyenas don’t always stick together, They may act communally, as when hunting dangerous prey, and defending clan territories, or individualistically (and highly competitively) as when caring for their young, foraging, and hunting smaller prey. This allows the species to exploit many different resources efficiently.
The spotted hyena social system differs from other social carnivores in that there is no communal sharing of food (or care of the young, each female caring only for her own).
(Photo: Bernard Dupont. CCAS 2.0 A license*)
Females rule. Larger than males (unusual in mammals), they take the lead in territorial marking exercises, group hunts, and battles with other packs. Females also remain in the clan / pack while males emigrate (at about two years of age). Males not only defer to females, but also play no parental role, and are often not even allowed near the otherwise communal dens, Less closely knit than wild dogs, spotted hyenas more often forage and hunt alone.
“Darn females won’t let me in the house! “
(Photo: David Bygott.)
Complex behavior requires good communication. Spotted hyenas are excellent communicators. This is because every individual is, to another hyena, a potential competitor (even dangerous enemy) or collaborator, which makes the signaling of moods and intentions very important. Thus, the spotted hyena has an enormous array of calls (whoops, moans, grunts, giggles, whines, yells, growls), expressions, postures, and attitudes. This can lead to a massive amount of noise when they compete with one another over a carcass. Consequently,the spotted hyena is one of Africa’s noisiest animals.
Highly gregarious, the spotted hyena is the most social of all carnivore species, with the largest social groups. For instance, a spotted hyena clan may comprise 35 to 80 adults. (In contrast, the largest recorded pride of lions, the other major social large carnivore, is 30 animals, including cubs.)
Social group size, however, varies with the availability of food, as does the size of spotted hyena territories, and the degree to which the group (clan / pack) defends them. For instance, in the Kalahari Desert spotted hyena densities are so low that they forage and hunt within territories too large (500-2,000 sq. km / 193-772 sq. miles) to defend against other hyenas. In contrast, where wild ungulate prey is most abundant, as in Tanzania’s Ngorongoro Crater, hyenas are numerous, forming large social groups and aggressively defending territories less than 40 sq. km (15 sq. miles) in area. (The much larger Serengeti-Mara ecosystem, which includes the crater, is home to 7,200-7,000 animals, the largest spotted hyena population in Africa.)
Ngorongoro Crater’s large resident population of 25,000 ungulates supports high densities of from 380-470 spotted hyenas.
(Photo: David Bygott.)
One night in the Serengeti, two friends of mine, staying in a guest house, experienced at close hand the aggressive defense of a spotted hyena clan territory. Hearing hyenas, they went outside and played a recording of spotted hyena calls from another area. Minutes later they had to scramble atop a nearby Land Rover to escape a crowd of angry spotted hyenas. Hearing hyenas from another clan in their territory, they had rushed over to expel them.
Another example of the spotted hyena’s complex behavior is a greater plasticity in foraging and hunting behavior than exhibited by other African carnivores. for instance, spotted hyenas both scavenge and hunt, the former usually during the day because they use vultures as indicators of kills, and the vultures only fly during the day.
A spotted hyena waits for lions to finish eating before scavenging the remains.
(Photo: David Bygott)
However, when carcasses are scarce, spotted hyenas also hunt, usually at night, and, depending on the circumstances, either alone, in small parties, or in large groups. A common technique is to lope toward a herd or flock, forcing its members to flee, revealing easy to catch stragglers (weak, young, sick).
Wildebeest calves in Ngorongoro Crater and on the Serengeti Plains are a favorite prey of spotted hyenas. As they are easy to catch, they are hunted by single hyenas.
(Photo: David Bygott)
Like wild dogs, spotted hyenas simply run their prey to exhaustion, usually within 1.5-5 km (1-3 miles). A single hyena can catch and kill healthy prey the size of a bull gnu, but only as a last resort. When hyenas are numerous, other pack members may join in near the end of a chase to help pull down larger animals like wildebeests. However, usually led by a female, they also stage deliberate pack hunts of dangerous prey, such as zebra families guarded by sharp-hoofed stallions.
Zebras fight back, so spotted hyenas must hunt them in groups.
(Photo: David Bygott)
Often eating their prey alive, spotted hyenas, unlike wild dogs (https://storiesofeastafrica.com/2023/09/13/wild-dogs-wildlife-of-ngorongoro/), compete with other pack members by eating as much and as fast as they can, with individuals swallowing up to a third of their weight (A lion can only swallow a fourth of its weight). Twenty hyenas, and another group of 35, were recorded finishing off carcasses weighing 100 kg (220 lbs.) and 220 kg (485 lbs.) respectively, in 13 minutes. And they do this with remarkably little fighting. Instead, there’s lots of noise, which attracts other clan members (up to 65 seen on a kill in Ngorongoro Crater). Only then do all clan members ever come together. (These competitive scrambles are less common in the Serengeti where there is a better ratio of prey to hyena.)
When many spotted hyenas are on a kill, some quickly eat what they can before taking a chunk of meat and bone elsewhere for a quiet meal.
(Photo: David Bygott)
Spotted hyenas also excel in that they eat almost the entire carcass of their prey. They can, for instance, crack quite large bones, such as those of buffaloes and giraffes, noisily splintering them before they are swallowed. Furthermore, their digestive system can dissolve bones, and even teeth, within hours. (Dry hyena scats, composed of ground up bones, are a chalky white.). Thus, virtually everything is eaten except the rumen contents (Grass! Yuck!), and horn bosses of larger antelopes. The hair, and hooves, which cannot be digested, are disgorged. Probably no other carnivore utilizes vertebrate prey so efficiently. Other species waste up to 40% of their kills.
“The good stuff’s finished; time to eat the hide.”
(Photo: David Bygott)
Hans Kruuk found the spotted hyena to be a formidable predator. This to the extent that lions often scavenge hyena kills. Direct competitors with lions for food, spotted hyenas, unless present in large groups, generally give way to the larger carnivores, allowing them to appropriate their kills. In turn, spotted hyenas frequently steal kills made by cheetahs and wild dogs (although the latter, being more socially cooperative, are sometimes able to successfully defend themselves) and, given the opportunity, also kill their cubs, as do lions. In the Serengeti, lions and hyenas have exerted such pressure on wild dogs that they have pushed them into outlying parts of the greater Serengeti ecosystem.
A single hyena being harassed by a pack of wild dogs.
(Photo: Kruger sightings HD. CCA 3.0 U**)
Furthermore, spotted hyenas can be dangerous to livestock and people, especially when other food is scarce, but also when an opportunity arises for an easy meal, such as encountering an unattended child, or someone sleeping in the open. A relevant example (see below) recently appeared in my local newspaper.
Furthermore, spotted hyenas are easily kept and trained. Witch doctors sometimes add to their persona by keep them as pets. Hans Kruuk and his wife, Jane, successfully raised a young hyena as a family pet in the Serengeti National Park, although they eventually put it in a zoo when it learned how to open doors and steal bacon from the chief park warden’s breakfast table.
Spotted hyenas: Complicated creatures. Just like us.
REFERENCES
Estes, R.D. 1992. The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates. The University of California Press.
Fosbrooke, H. 1972. Ngorongoro–The Eighth Wonder. Andre Deutsch.
Herlocker, D.J. 2009. Buffaloes by My Bedroom: Tales of Tanganyika. iUniverse.
Featured image: Production risk due to lack of rainfall in Marsabit District
Now available for free downloads: digitized copies of the 22 reports (3 volumes) comprising the Range Management Handbook of Kenya.
For information on rangeland resources, climatology, livestock diseases, constraints facing range development, and much more, go to the sidebar and click on Range Management Handbook of Kenya. Alternatively, click on storiesofeastafrica.com/range-management-handbook-of-kenya/
Some examples of newly available reports / maps appear below.
District-wide maps provide inventories of natural resources and their ecological status
Environmental degradation is ongoing in the rangelands of East Africa. This guide shows how to identify and monitor soil erosion in arid and semi-arid lands.
Camels, which typify arid rangelands, have, in recent years, also become a presence in semi-arid rangelands. See this report for tips on how to manage them.
Featured image: African wild dog pack, Kruger National Park, South Africa. Bart Swanson. Creative Commons Attribution-Share Alike 3.0 unsorted license
“Wild dogs!” John yelped. Pointing out the right window of his Land Rover, he exclaimed “Over there–ten of them.” Then he really got excited: “And they’re chasing something! See how they’re trotting, strung out in a long line? Fantastic!” Abandoning our search for rhinos, we promptly sped off to follow the hunt.
It was 1965 and I, the assistant conservator (forests) for the Ngorongoro Conservation Area (NCA), was temporarily without official transport because the NCA had prematurely exhausted its government-imposed monthly fuel allowance. For the time-being I would be unable to supervise the crew constricting a road around the western and northern rim of Ngorongoro crater or visit forest guards patrolling the Northern Highlands Forest Reserve; they were all too far away. What I could do, however, was accompany my neighbor, John Goddard, a Canadian biologist studying black rhinos, into the Crater. John had his own source of funding, which meant he was unhampered by government fuel allowances.
Rattling across the crater floor in John’s Land Rover, we followed the pack for about a mile before it brought down its prey, an adult Grant’s gazelle. Then, instead of resting from their exertions, the wild dogs immediately started ripping it apart; John and I arrived to find one dog pulling on a foreleg of the gazelle, another yanking the other direction on a hind leg, and two others tugging at its stomach while the rest of the pack danced about uttering excited twitters and whines. John quickly took several photographs, but then surprised me by leaving the vehicle to approach the frenzied melee taking place only thirty feet away. What was he thinking? They’ll eat him for dessert! But then another surprise: instead of aggressively defending their kill, the wild dogs warily backed away, allowing John to walk right up to it.
Wild dog pack tearing into a Grant’s gazelle in Ngorongoro Crater, Tanzania.
In his 1972 book, Ngorongoro: The Eighth Wonder, Henry Fosbrooke remarks on how wild dogs in the crater always gave way when John approached. He did this, Henry said, to collect the prey’s mandibles–aging its teeth showed an animal’s age at death. John wanted undamaged specimens so he collected them as soon as possible after the wild dogs made their kill. However, in this case at least, there also was a fringe benefit: Returning with both the gazelle’s head and part of one of its hindquarters, John announced, “This takes care of dinner, tonight!”
My three years (1964-1967) at Ngorongoro exactly coincided with the presence of a wild dog pack in Ngorongoro Crater. Few were seen prior to this time and the pack left the crater in 1967. But then African wild dogs have a reputation for being rare and elusive. For instance, Henry Fosbrooke saw a wild dog only once during his 30 years of on and off acquaintance with Ngorongoro. George and Lory Frame, who studied cheetahs and wild dogs on the Serengeti Plains in the 1970’s, often spent days or weeks searching for dogs to study, then, having found a pack and studied it for a few days, woke up the next morning to find the dogs had vanished.
“Where’d they go?”
Being rare and elusive makes the African wild dog difficult to study. Nonetheless, it’s worth the effort, and not just because the wild dog occupies its own taxonomic genus (Lyacon) differing from the genus Canis (jackals, wolves, coyotes, domestic dogs) by dentition highly specialized for a hyper-carnivorous diet, and in having four toes on each foot. Both attributes support survival, the first by enhancing the shearing of meat, which increases the speed at which prey is consumed (thereby lessening the chance that lions and hyenas can steal the kill), and the second increases an animal’s stride and speed, allowing long distance pursuit of prey.
To the average viewer, however, the African wild dog’s most distinctive features are its large, round ears, and a splotchy black, white, and brown (sometimes verging on yellow) body–hence its other name, the painted dog. Happily for those who study this species, each animal has its own unique, readily distinguishable coat color pattern. African wild dogs also apparently really stink, although, unlike John Goddard, I never got close enough to tell.
Painted dogs, Kruger National Park, South Africa. Unlike those pictured here, the wild dog in East Africa generally has a white-tipped tail. (The Maasai call it Oloibor kidongoi, the white-tipped one.)
Bernard Dupont. Creative Commons Attribution-Share Alike 2.0 Generic License.
The African wild dog is a habitat generalist able to survive in a wide range of environments (An extreme, although undoubtedly short-lived, example is the sighting of a pack near the summit of Tanzania’s 19,341 ft Mt. Kilimanjaro). However, wild dogs are most commonly found in relatively open habitats which provide good views and running conditions.
Almost exclusively carnivorous, killing most of what it eats, the species is specialized as a pack hunter, concentrating on whatever medium-sized antelopes are most abundant. More enduring than its prey, it pursues the latter at up to 35 m/h (56 km/h), one dog leading , and the rest strung out behind, until the prey is exhausted, usually within 3 miles (5 km). Their ability to run their prey down without having to conceal their approach allows African wild dogs to be conspicuously colored, and like cheetahs, hunt only during the day.
African wild dogs chasing prey in Botswana’s Okavango Swamp.
Probably one of the most successful African carnivores, African wild dogs are such effective hunters that when prey is abundant, a pack can regularly have both breakfast and dinner. For example, wild dogs in Ngorongoro Crater killed twice a day, catching 85% of the animals they chased. Even the lowest recorded success rate for this species (39%) exceeded those of all other large predators except cheetahs. Furthermore, the entire pack benefits from a kill because it is shared among them.
Wild dogs in the Serengeti primarily prey upon Thompson’s gazelles and, in season, wildebeest calves (pictured here).
One reason for their rare and elusive nature is that wild dog packs are frequently on the move, averaging 10 km (6 miles) / day, or when game is scarce, up to 40 km (25 miles) / day. In the latter case, hunting ranges can be huge, up to 1500-2000 sq. km / 580 – 770 sq. miles (The largest recorded range is greater than the total area of London), exceeding in area even those of cheetahs. However, a range can be smaller when prey is resident and numerous.
The African wild dog has specialized on an abundant food resource which it can only exploit efficiently by hunting in packs. In East Africa these typically consist of about 10 animals but can go as high as 20 or even 60. Social bonds are strong; when separated from its pack, an African wild dog becomes so depressed that it may die. Each pack has only a single breeding pair, composed of the dominant male and female, which needs assistance from the other adults to provision large litters of up to 10 pups during an extended (12-14 month) period of dependence. Food sharing, by regurgitation of meat obtained from a kill, is supported by an emphasis on submissive, begging behavior within a pack. Aggressive behavior is rare. In-breeding is prevented by the emigration of females to other packs whereas the males, related to one another but not the breeding female, remain.
Wild dog pups. Litter sizes , larger than any other canid, are enough to form a new pack every year.
David Bygott
Despite being such effective hunters, and having an exceptionally efficient reproductive system, African wild dogs are the least common large predator in Africa. Furthermore, their population is declining. With under 7,000 animals remaining in the wild (there are fewer wild dogs than cheetahs) and having disappeared from much of their former range, African wild dogs are the continent’s 2nd most endangered large carnivore, after the Ethiopian wolf. Reasons given include:
(a) infectious diseases: Wild dogs are highly susceptible to canine diseases spread by domestic dogs;
(b) competition from lions and spotted hyenas, which appropriate wild dog kills, and in the case of lions, also kill their pups and adults;
(c) habitat fragmentation and loss.
The fragmented continental habitat of African wild dogs. The full extent of the original habitat can be roughly approximated by the distribution of smaller relict ranges. (Approximately 700 wild dogs live in northern Botswana.)
IUCN. CCA-SA 4.0 International license.
The results are smaller, less efficient and viable wild dog packs. Once African wild dog packs are reduced to small sizes, and suitable habitats are fragmented and altered by humans, wild dog populations seldom recover.
Lions are bad news for wild dogs.
David Bygott
So are domestic dogs, which carry infectious diseases.
Bothar at English Wikipedia CCASA 3.0 Unported.
Thus, the most effective way to ensure the conservation of African wild dog populations is thought to be by creating and protecting areas connecting isolated habitats. A good example of the importance of extensive, connected, ecologically diverse wild dog habitats is provided by the Serengeti-Mara Ecosystem in Tanzania and Kenya. Probably due to a combination of disease and competition from lions, (the major source of wild dog mortality in the Serengeti), and from spotted hyenas, African wild dogs disappeared from the 5,700 sq. mile (14,763 km) Serengeti National Park in the early 1990’s. However, the wild dog populations survived by moving into other parts of the greater (15,444 sq. mile / 40,000 sq. km) Serengeti-Mara Ecosystem. These include Ngorongoro Crater where wild dogs have returned after a 30 year absence, but primarily the Loliondo Game Controlled Area where a more hilly habitat provides greater security from larger predators while the wild dogs are denning and raising their young. Currently, the Serengeti-Mara Ecosystem contains about 120 African wild dogs.
The Loliondo Game Controlled Area (dark green) lies east of the Serengeti National Park.
Abrah Dust. CCA-SA 4.0 International license.
REFERENCES
Estes, R.D. 1991. The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates. The University of California Press. Fosbrooke,H. 1972.
Fosbrooke, H. 1972. Ngorongoro: The Eighth Wonder. Andre Deutsch.
Frame, G. & L. Frame. 1981. Swift & Enduring: Cheetahs and Wild Dogs of the Serengeti. E.P. Dutton.
A cheetah cub purred as it chewed on a windshield wiper; another admired itself in the Land Rover’s fender mirror; a third, sitting on the roof, dangled its tail beside an open window . . . George and Lory Frame, who studied cheetahs and wild dogs in the Serengeti in the late 1970’s, recount how, one day, five large, playful cheetah cubs, members of a family they were studying, turned the tables on the two researchers and investigated them, instead. For the Frames it must have been a welcome break from their usual routine of hours and hours of careful observation and note-taking.
A nice patch of shade.
George and Lory certainly had an interesting animal to study: An inhabitant of wooded savanna, open plains, and desert, the cheetah is a striking cat: lightly built, with long, thin legs, small feet with blunt, unsheathed claws, and small rounded head, it’s built for speed–the field version of a greyhound, as one researcher put it. Capable of top speeds of 60-70 mph (90-112 kph), it’s the fastest animal in the world, a specialist in hunting small, but fast antelopes, such as Thomson’s gazelles (and hares), seldom killing animals larger than itself.
Built for speed. Photo by David Bygott.
However, becoming so speedy has involved tradeoffs. A sprinter with little stamina for chases beyond 300 m / 985 ft., a cheetah must use whatever cover (tall grass, shrubs, trees, ravines) is available to get as close as possible to its prey before attacking it. Furthermore, the cheetah has sacrificed not just stamina, but also the strength (it’s less than a fourth the weight of a full-grown lion) and weapons (note those dull claws) needed to protect itself from other predators. Consequently, cheetahs must avoid large predators, especially lions, which often kill cheetah cubs as well as steal kills. Hyenas also frequently appropriate prey killed by cheetahs. Therefore, cheetahs do not hunt at night, which is when lions and hyenas are most active.
Cheetahs also are unique in that males are social while females are solitary and shy (but still highly promiscuous). Furthermore, they have large home ranges, the largest, up to 400 sq. miles (1,036 sq. km), being those of individual females. Groups of males, called coalitions, defend smaller territories, 14-62 sq. miles (36-160 sq. km) in area, within female home ranges.
Cheetah stalking a Thomson’s gazelle, its favorite prey on the Serengeti Plains. The adult zebra and wildebeests in the background are too large. Photo by David Bygott.
Large home ranges, and mortality from larger predators so intense it limits cheetah numbers even when prey is abundant (less than half of all cheetahs live beyond three months), means that cheetahs never achieve high densities–they are always less abundant than other African carnivores. For instance, the 15,444 sq. mile (40,000 sq. km) Serengeti-Mara Ecosystem (see map at end of post), an area larger than the state of Maryland, and almost twice the size of Wales, has 275 cheetahs (the world’s highest density of these cats), but also 3,000 lions and at least 7,200 hyenas. (The Ngorongoro Conservation Area is part of the Serengeti-Mara Ecosystem.)
Two of the Serengeti-Mara Ecosystem’s 275 cheetahs. Photo by David Bygott.
Most other protected areas are too small to protect viable cheetah populations. Of the roughly 10,000 cheetahs in Africa, about 2/3 live in unprotected areas where. due to persecution and loss of habitat, their numbers are declining. Low populations, the need for young males to sometimes travel large distances to establish new territories, and an extraordinarily low degree of genetic diversity means that cheetahs, of all carnivores, are most vulnerable to habitat loss and fragmentation. (Low genetic diversity also raises the possibility that a disease could devastate wild populations.). Thus, steps are needed to conserve them outside protected areas.
Some conservationists see the cheetah’s future linked to the survival of traditional pastoral livestock management. This is because, historically, there has been little conflict between cheetahs and traditional pastoralists, who minimize stock losses to carnivores by seldom leaving their animals unattended during the day and corralling them at night in protected enclosures.
Traditional pastoralists seldom leave their livestock unattended. Photo by David Bygott.
MAPS
Vegetation of the Ngorongoro Conservation Area, Tanzania. Forest (darkest green–far right) is the poorest of these habitats for cheetahs. Graphic by David Bygott and Jeannette Hanby.
The seasonal movements of the Serengeti-Mara wildebeest populations occur within all or part of each protected area. Graphic by Abrah Eust. Creative commons 4.0 international license.
MAJOR REFERENCES
Estes, R.D. 1991. The behavior guide to African mammals: Including hoofed mammals, carnivores, primates. The University of California Press.
Fosbrooke, H. 1972. Ngorongoro: The eighth wonder. Andre Deutsch.
Frame, G. & L. Frame. 1981. Swift and enduring: Cheetahs and wild dogs of the Serengeti. E.P. Dutton.
Durant. S. 2004. Survival of the fastest: The cheetahs of the Serengeti. Africa Geographic. Pages30-32.
Zoological Society of London. Cheetah conservation in Africa. ZSL.org
“Tembo!” exclaimed one of the men. A large elephant had unexpectedly materialized from the bamboo a few hundred feet away. Like a spirit, it moved silently across an arm of the glade and vaporized into the thicket on the other side. If the laborer hadn’t happened to look up when he did, we would have missed it completely.
Buffaloes by My Bedroom: Tales of Tanganyika
Elephants in a glade of manyatta grass.
It was 1964. My African crew and I were scouting a route for a track that would allow tourists to view wildlife on the rim of Ngorongoro Crater. The proposed route ran through light green bamboo thicket, spotted by dark-leaved pillar wood trees, and punctuated by sun-filled glades of tall manyatta grass. That this was buffalo and elephant habitat was evident from the abundance of their dung, tree trunks worn smooth by the rubbing of large bodies, trails forced through the dense bamboo, and mud wallows. (Refer to last page of blog for maps of the Ngorongoro Conservation Area, and of the vegetation of Ngorongoro Crater.)
Smoothing a stump while easing an itch.Photo by David Bygott.
Sometimes we heard them, especially elephants: snapping branches as they browsed, the low rumbles of their stomachs, an occasional sharp, thrilling trumpet. . . But we also saw them, and not just the elephant in the first paragraph, but also buffaloes.
Three times, while sticking tall lengths of bamboo into the ground to mark where the new track would go, we jumped buffalo herds of forty to fifty animals. Luckily however, we scared them more than they scared us. What a noise they made as they blundered away, grunting and snorting, heavy hooves rumbling across the ground–they were big animals. They always paused at the edge of the bamboo to look back at us. We could see the sun glinting off their horns. They were mean-looking critters!
Buffaloes by My Bedroom: Tales of Tanganyika
Mean-looking critters.
Elephants and buffaloes also used open, grassy habitats. For instance it was not unusual to see elephants grazing the grasses, sedges, and herbs of Munge and Gorgor swamps on the floor of Ngorongoro Crater. However, most of my encounters with the two species were either in or near forest.
Gorgor Swamp. The large plants are Cyperus immensus, a species of sedge.
Elephant in Ngorongoro Crater’s Lerai Forest. Fed by a stream flowing down the crater wall, the forest is dominated by yellow barked fever trees (Acacia xanthophloea).
For instance, there was that cool sunny afternoon, following a rainstorm, when several elephants on the crater rim, seemingly stimulated by the rain, became amusingly animated: trumpeting, and pushing one another about. I could hear the click of their tusks–ivory on ivory–as the large animals playfully sparred. The dry season was coming to an end and they were happy to see the rain.
Elephant near my house on the rim of Ngorongoro Crater.
“In or near forest” included my small house, around which buffaloes often grazed at night.
The rooms of the house (a main room, two bedrooms, and a bathroom) didn’t interconnect. Instead, each opened directly onto the yard. On foggy mornings, after carefully checking for the presence of buffaloes, I groped my way to the bathroom through heavy mist. At night, I checked again with a flashlight before going to bed.
Buffaloes by My Bedroom: Tales of Tanganyika
My “yard” was mown at night by grazing buffaloes.
Not what one wants to meet while groping through fog to the privy. Photo by David Bygott.
At night, buffaloes also grazed the grounds of the Ngorongoro Crater Lodge. To prevent mishaps, the lodge hired Maasai warriors, armed with spears, to escort guests to their scattered cabins, following the evening meal in the main building.
Tourist cabins on the grounds of the Ngorongoro Crater Lodge. The main building, which included the dining room and bar, is at the far right.
While I commonly saw elephants in the swamps on the floor of Ngorongoro Crater, there were few buffaloes there in the 1960’s. In contrast, in terms of biomass, they are now the dominant herbivore, and for a very interesting reason. This significant change probably resulted from the removal of Maasai from the crater in 1974. With the Maasai went their traditional grassland management in which grazing and controlled burning kept grasslands short, palatable, and suited to small and medium sized grazers, such as wildebeests. Subsequently, grasses on the crater floor became longer, less palatable, and more suited to less selective grazers, such as buffaloes. Consequently, buffaloes became more abundant while the numbers of wildebeests, once the crater’s dominant herbivore, significantly declined. The Ngorongoro Conservation Area Authority has since returned grass fires to the crater ecosystem through a program of controlled burning. (Also see https://storiesofeastafrica.com/2022/09/27/maasai-pastoralists-of-ngorongoro-as-they-are-now/)
Buffaloes are more difficult prey than wildebeests. Will this cause lions to prey more heavily on the latter? Photo by David Bygott.
The Ngorongoro Conservation Area is thought to contain over 300 elephants. However, much of the habitat of Ngorongoro’s elephants is dense forest. This makes them hard to count. Thus, their true number is unknown. Although elephants are endangered throughout Africa by loss of habitat, and by poaching for ivory, little elephant poaching has been recorded in the NCA over the past several years. That said, on my last visit to Ngorongoro, in 2004, I was startled to see a tuskless adult elephant, something I had never encountered before.
Notice something missing ?
In well-protected areas, elephants without tusks may comprise as little as 2% of a population. This may be the case in the Ngorongoro Conservation Area. Let us hope so, anyway, because populations with significantly higher amounts of tuskless elephants (especially if the amounts are increasing), reflect heavy, unrelenting poaching. Simply speaking, elephants without tusks are less apt to be killed by poachers. Over time, therefore, their genes will dominate the population. Even thirty years ago, up to 25-38% of some African elephant populations were without tusks. A more recent study in Mozambique’s Gorongosa National Park reported over half of all female elephants in to be tuskless. For more information on this subject, check out the on-line articles by Garrigan, Preston, and Associated Press in the reference section.
MAPS
Ngorongoro Crater is in the Crater Highlands, east of Tanzania’s famous Serengeti Plains. Graphic by David Bygott and Jeannette Hanby.
Ngorongoro Crater vegetation. Light blue (wet meadows), dark blue (reed swamp), yellow (medium grassland), light yellow (short grassland), light brown (bushland), dark brown (high woodland), green (forest). ( Herlocker, D.J. & H.J. Dirschl. 1972.)
Herlocker, D.J. 2009. Buffaloes by My Bedroom:Tales of Tanganyika. iUniverse.
Herlocker. D.J. & H.J. Dirschl. 1972. Vegetation of the Ngorongoro Conservation Area, Tanzania. Canadian Wildlife Service– Report Series Number 19.
Oates, L. & P.A.Rees. 2013. The Historical Ecology of the Large Mammal Populations of Ngorongoro Crater, Tanzania, East Africa. Mammal Review 43(20013) 124-141.
September 1964. One of the things I remember from my first day as a Peace Corps volunteer in Ngorongoro Crater were the reactions of Henry Fosbrooke, conservator of the Ngorongoro Conservation Area, and John Goddard, a Canadian wildlife biologist, to tourist vehicles tightly clustered around animals of interest, such as lions.
“Why don’t they stay on the main tracks?” Henry complained. “There’s quite enough off-track driving going on. It’s wearing down the grass and disfiguring the crater floor. I must talk with our guides about this.” John wasn’t pleased either. “Crowding the animals makes them edgy.”
Buffaloes by My Bedroom: Tales of Tanganyika.
That year, 12,137 tourists visited Ngorongoro. In 2018, over half a century later, the amount was closer to three quarters of a million. If Henry and John were worried about the impact of tourists on the crater’s environment and wildlife then, I wonder how they would feel now.
Not that I blame people for wanting to visit the Ngorongoro Conservation Area, for there is much to see. Here are some examples:
Lions. Tourist vehicles were most apt to crowd around the big cats, of which lions are both the largest, and usually easiest to find. And who wouldn’t want to see one, for the lion is the epitome of ferocity, the King of the Beasts lording it atop the food chain. Even from a distance, the large carnivore’s booming, moaning roars have a way of concentrating one’s attention, especially when sleeping in the fragile protection of a tent.
Meal time in Ngorongoro Crater. Photo by David Bygott.
Ngorongoro Crater has long been considered one of the best places on earth to see lions. In 1975, at 125 animals, it contained one of the densest populations in Africa. As of 2014, however, only 55 lions, comprising four prides, ruled the crater. Unfortunately, the lions of Ngorongoro Crater have a problem. Back in the early 1960’s most of them died, weakened by a plague of Stomoxys biting flies. Although the population subsequently grew, so few new lions entered the crater to replenish the gene pool over the next three decades that the crater’s lions became inbred. Immigration of new lions into the crater apparently has always been rare. However, a growing belt of Maasai communities may have exacerbated the situation by detering movement of lions between plains and crater (Maasai warriors kill lions to protect their livestock; they also once did it to prove their manhood.) This is one of the reasons why the Ngorongoro Conservaton Area Authority (NCAA) worries about the impact of an increasing human population on wildlife conservation (See MAASAI PASTORALISTS OF NGORONGORO: AS THEY ARE NOW). That said, however, KopeLion https://www.KopeLion.org and The Lion Recovery Fund, https://www.lionrecoveryfund have recently had some success in improving the relationship between lions and the Maasai.
Wildebeests. High-shouldered, broad-muzzled, and with cow-like horns and spindly legs, the wildebeest, or gnu, will ever win a beauty contest. (That said, I must admit to the cuteness of gnu calves.)
Wildebeest calves in Ngorongoro Crater.
In compensation, the wildebeest has a comical personality best displayed when, for instance, one of the funny looking animals cavorts about, kicking it’s heels in the air, or again, when sparring males drop to their front knees and butt heads with their hind ends in the air. Then, there’s those bleating grunts–ngggh. . . ngggh. . . ngggh–which sound like the large antelopes have sinus problems. (The name, gnu, might come from an approximation of that sound. However, the Swahili word for wildebeest is Nyumbu.)
The most spectacular wildebeest population in the Ngorongoro Conservation Area (NCA) comprises the over a million animals of the migratory Serengeti herds which visit the eastern Serengeti Plains in the rainy season to graze nutritious forage and give birth.
Wildebeests on the Serengeti Plains. The animals calve on the eastern plains, which lie within the Ngorongoro Conservation Area. Photo taken in 1972 by Dirk Kreulen.
The wildebeest population in Ngorongoro Crater contains both resident and migratory animals. Thus, about 7,000 wildebeests are present throughout the year, blending in with a similar number of other ungulates, larger mammals, such as rhinos, and predators. (The present wildebeest population is about half what it was in the 1960’s. This is because removal of Maasai cattle from the crater, combined with a ban on setting grass fires, reduced wildebeest numbers by encouraging taller grass, a habitat more conducive to buffaloes.)
Wildebeests and zebras in Ngorongoro Crater
The year-round presence of wildebeests in Ngorongoro Crater strongly contrasts with that on the Serengeti Plains where the migratory herds provide an overwhelming visual impact in the wet season but are completely absent during the five-month dry season. In fact, few ungulates of any kind use the plains in the dry season.
Dry season on the Serengeti Plains. Photo by David Bygott.
In the 1960’s, I occasionally encountered oryx on the drier parts of the Serengeti Plains. Adapted to dry environments, they also make dry season use of areas which other herbivores only use in the wet season.
Black rhinos. While only half the size of an 1,750,000-year-old ancestor discovered by archeologists at Oldupai Gorge, the black rhino is still a heavy animal, weighing in at about a ton. However, as it is able to gallop at 30 mph (50 kph), and turn in its own length, it is a surprisingly nimble large animal.
Black rhino in Ngorongoro Crater.
The rhino’s inability to distinguish a motionless object beyond 15 yards (14 meters) may explain why, rather than charge a perceived threat, it’s more apt to snort loudly and, tail looped over its rump, trot away. Even its “charges” tend to be more impulsive and confused than aggressive. I once experienced this when, while rapidly retreating from what I thought was a charging rhino, I tripped and fell flat on my face only to find my presumed pursuer trot past fifty or so feet away.
Rhinos also can become habituated to the presence of humans. I remember one, named Horace, who was fast asleep when Henry, John, and I visited him that first day in the crater, and stayed that way as we ate lunch a few feet away.
The only signs of activity from Horace, other than heavy breathing, came from several yellow-beaked tick birds (ox peckers) that hopped and fluttered about as they poked into nooks and crannies of his body in search of ticks.
Buffaloes by My Bedroom: Tales of Tanganyika.
Non-habituated rhinos, however, can be dangerous. For instance, when Horace first appeared in the crater, he vigorously charged everything in sight. John and I barely escaped the wrath of an angry female, who only managed to bump our Land Rover as we sped away. A colleague encountering a grumpy rhino in the Serengeti National Park, did not escape so easily, the big animal hooking it’s horns under his Land Rover’s fender and banging the vehicle up and down several times before disengaging, leaving the fender looking like it had been holed by artillery shells.
A rhino charging a vehicle that came too close.
Sadly, the black rhino, as it is elsewhere in Africa, is the most threatened large mammal in the NCA. Intensive poaching wiped out the entire Oldupai Gorge population of 70 animals by the 1980’s and reduced their numbers in Ngorongoro Crater from over 100 in the late 1960’s to a mere 12 in 1996. Subsequently, however, a program of intensive monitoring and protection carried out by the NCAA and Frankfurt Zoological Society, increased the population to 30 by 2017. Yet another problem is that, as with the crater’s lions, its rhinos also may be genetically isolated and inbred. This has been partially addressed by the translocation into the crater of two female rhinos.
Mobile ranger post used for monitoring and protecting rhinos in Ngorongoro Crater. Photo taken in 2004.
Nonetheless, the black rhinos of Ngorongoro Crater continue to be a major tourist attraction.
Tourists viewing a distant rhino in Ngorongoro Crater. My old colleagues, Henry Fosbrooke and John Goddard, would be startled by the numbers of vehicles, but highly gratified that none have left the road. Photo taken in 2008 by David Bygott.
(TO BE CONTINUED)
Ashley Maberley, C.T. 1962. Animals of East Africa. Howard Timmons, Cape Town.
Estes, R.D. 1991. The Behavior Guide to African Animals. The University of California Press, Berkeley and Los Angeles.
Fosbrooke, H. 1972. Ngorongoro: the Eighth Wonder. Andre Deutsch, London.
Ngorongoro Conservation Area Authority. 1966. General Management Plan. Tanzania Ministry of Natural Resources and Tourism.
Oates, L. & P.A. Rees. 2013. The Historical Ecology of the Large Mammal Populations of Ngorongoro Crater, Tanzania, East Africa. Mammal Review 43 (2013) 124-141.
In the mid-1960’s, the pastoral Maasai of Ngorongoro, proud of their reputation for being fierce warriors, and possessing an abundance of cattle, were content with their way of life. Thus, they were conservative and resistant to change, an attitude that frustrated government officials, both pre-and post-independence, and gave the Maasai a reputation for being backward. (Adapted from the previous blogpost, The Maasai of Ngorongoro: 1960’s.)
Over half a century later their situation has changed–drastically.
For instance, due to better health care, and the immigration into the Ngorongoro Conservation Area (NCA) of other pastoralists (especially during droughts), the human population is now twelve-fold greater.
In contrast, several factors have constrained growth of the livestock population: (a) Valuable grazing lands have been lost to other uses, primarily wildlife conservation and tourism, (b) Livestock carrying capacity has declined due to overgrazing, a ban on setting grass fires, and recurring droughts, (c) Livestock deaths have increased due to diseases and droughts.
DETAILS
Valuable grazing land was lost when the Ngorongoro Conservation Area Authority (NCAA), concerned that the Maasai and their livestock were hindering wildlife conservation, removed them from Ngorongoro, Olmoti, and Empakaai craters in the 1970’s. (Ngorongoro Crater was an especially hard loss.)
Ngorongoro Crater is the largest in the Crater Highlands. Empakaai Crater contains a small, but lovely lake.Graphic by David Bygott and Jeanette Hanby.
Ngorongoro Crater, 97 sq. miles (252 sq. km) of productive rangeland. (View from my house on the crater rim in 1964.)
Also, expanding numbers of wildebeests calving in the wet season on the eastern Serengeti Plains, and the associated risk of cattle catching malignant catarrh fever (MCF), forced herders to keep their livestock in the highlands for extended periods. The reduced ability of the Maasai to use this wet season grazing area caused an estimated 35% reduction in cattle numbers.
Wildebeest calves on the eastern Serengeti Plains, which lie entirely within the NCA. From 1961 to 1977, the Serengeti wildebeest population grew from 250,000 to its present level of approximately 1,277,000. This caused the area used for calving to significantly expand. Photo by Dirk Kreulen.
Afterbirth of a wildebeest calf. If cattle graze grass that has been touched by it they are apt to contract (and die from) malignant catarrh fever (MCF). Photo by Dirk Kreulen.
Forage quality and production dropped (at least for cattle) in parts of the Serengeti Plains because of declining grass and increasing shrub cover associated with a ban on setting grass fires imposed by the NCAA. (Pastoralists typically burn grasslands to kill ticks, remove dry grass, suppress woody plants, and induce greening of the vegetation.) Increasing abundance of unpalatable grass species in the highlands probably reflects overgrazing.
Grass fire on the floor of Ngorongoro Crater. A decrease in fires following removal of Maasai and their livestock from the crater in the 1970’s resulted in taller grasses and lower grassland species diversity. (Also, as I can personally attest from having to pick them off my pants in 2004, more ticks.)
Livestock deaths, especially of cattle, increased when herders, unable to use wet season pastures on the eastern Serengeti Plains, were forced to keep their animals for extended periods on traditional dry season pastures in the highlands. This increased the exposure of cattle to ticks, vectors for East Coast fever (ECF). Major die-offs occurred. A good example is provided by Andrew Clark, who in 1967, described the results of a virulent outbreak of ECF in Loliondo, north of the NCA: “Hundreds of cattle died in a few weeks. The whole area stunk of rotting carcasses. Hyenas, bellies pendulous from gorging, could barely walk. Vultures were so stuffed they could hardly get off the ground.”
Ticks, carriers of East Coast fever (ECF). They become abundant on rangelands which are seldom burned and/or are used for extended periods by livestock. Photo by Dr. Alexey Yakovlev. CC Attribution-SA 2.0 G.L.
Veterinary staff with bones of Maasai cattle killed by East Coast fever in Loliondo, Photo by Andrew Clark.
As a result, Maasai pastoralists were forced to reduce the proportion of cattle in their herds and increase that of goats. This is because goats are less susceptible to disease than cattle.
A mixed herd of goats and sheep. Goats also reproduce more quickly, produce milk throughout the year, utilize a variety of habitats (Cattle are restricted to grasslands), are drought-resistant, and easy to sell and slaughter. Thus, they are the fallback livestock for impoverished pastoralists.
Finally, droughts are becoming more frequent, and lengthy. Consequently, the grasses that provide forage for cattle are less able to recover their vigor between droughts, making them less productive. Thus, they support fewer animals, which tend to be weaker, in poorer condition, and more apt to die during the next drought. The drought ending in 2009, one of the most serious in recent memory, killed 35-40% of all cattle in Ngorongoro District, which includes the Ngorongoro Conservation Area (NCA) and, to the north, Loliondo.
THE CONSEQUENCES OF CHANGE
The result of all this is that, even though animal numbers have increased, the livestock population, especially of cattle, has not grown in accordance with the human population. Livestock biomass per pastoralist, well above subsistence level in 1966, is now below subsistence level.
This has caused the Ngorongoro Maasai, with too few livestock to support themselves, to become so impoverished that they must find other ways to supplement their livestock-based subsistence economy. Presently, they cultivate. Unfortunately, most are still too poorly educated to be employed in the region’s burgeoning wildlife-viewing tourist industry (six tourist lodges in the NCA alone). Those migrating to cities generally only find work as low-paid security guards.
This 2004 scene of Maasai bomas shows two examples of change since the 1960s: (a) cultivation , and (b) huts unprotected by fences (Predators may no longer be a problem, or the Maasai now know how unhealthy it is to live at close quarters with livestock inside the stockades).
Maasai security guards in Zanzibar. Photo by Jack Meyers.
Nonetheless, despite there being too few livestock to adequately support resident pastoralists, the Ngorongoro Conservation Area Authority (NCAA) is still concerned that there are too many for the land to support. It is especially worried about the ecological impact of overgrazing (as well as that of settlements and cultivation) on wildlife-based tourism, a major source of foreign currency (in 2017, 650,000 tourists visited the Ngorongoro Conservation Area, generating about 65 million $ U.S.) https://africasacountry.com/2022/04/people-live-here).
And the government is worried, too: The National assembly recently debated whether the Maasai even have a right, guaranteed in laws as far back as 1959, to live in the NCA. Also, recent reports in the media (denied by the government), state that it is considering relocating 80,000 Ngorongoro Maasai–much, if not most of the total population–outside the NCA. Whether or not this eventually happens, the NCAA/Tanzania government are “encouraging” impoverished herders to go elsewhere. Furthermore, a few hundred Maasai recently have, moving 210 miles (340 km) to Handeni in eastern Tanzania https://www.kbc.co.ke/hundreds-of-masai-ready-to-leave-conservation-area/.
Is this the future of Ngorongoro’s Maasai?
To best secure their future, the Maasai of Ngorongoro must become better educated. Photo by Christopher Michel. CC Attribution 2.0 GL.
ADDITIONAL REFERENCES
Amiyo, T.A. 2006. Ngorongoro Crater rangelands: condition, management, and monitoring. MS thesis, School of Biological and Conservation Sciences, University of Kwazulu-Natal, Pietermaritzburg.
Borges, J. et al. 2022. Landsat time series reveal forest loss and woody encroachment in the Ngorongoro Conservation Area, Tanzania. Remote Sensing in Ecology and Conservation. Open Access https://doi.org/10.1002/rse2.277.
Galvin.. et al. 2015. Transitions in the Ngorongoro Conservation Area: The story of land use, human well-being, and conservation. Pages 483-512 in Serengeti IV: Sustaining biodiversity in a coupled human-natural ecosystem. The University of Chicago Press.
Homewood, K.M. & W.A. Rodgers. 1991. Maasailand ecology: Pastoralist development and wildlife conservation in Ngorongoro, Tanzania. Cambridge University Press.
Ngorongoro Conservation Area Authority. 1966. General Management Plan. Tanzania Ministry of Natural Resources and Tourism.
