The African lion appears to be more gregarious than any other of the Felidae, and the male is certainly addicted to polygamy. Often a lion or a lioness may live and hunt for a time by itself, and very old animals are probably always solitary, as an old lion would be driven away from the females by younger males, and an old female would probably be badly treated by younger animals of both sexes. Sometimes two or even three males will hunt together for a time. More often a male lion may be met with accompanied by from one to four females, some of which latter may be followed by cubs of different ages and sizes. A family party consisting of one old male lion, three or four adult females, and several cubs, some of which may stand almost as high at the shoulder as their mothers, would constitute what the old Boer hunters would have called "en trop leeuws" (a troop of lions). In parts of Africa where game is, or was, very abundant, there are many authentic records of over twenty lions having been seen together. In his article on "The Lion," published in the Badminton Library Series, Mr. F. J. Jackson, C.B., has noted the fact that on August 7, 1890, he and Dr. Mackinnon came across a troop of twenty-three lions near Machakos in East Africa. This troop consisted of three male lions with splendid dark manes, five or six lionesses, and the rest cubs. I have come to the conclusion that such large assemblages of lions as this, in which there are several full-grown males, are, in all probability, only of a very temporary nature, the chance meeting and fraternisation of several families which, as a rule, live and hunt apart; since I believe that the passions of love and jealousy would not allow two or more males to live permanently in the company of lionesses without fighting. When a troop of lions is met with, in which, besides a full-grown male and some females and small cubs, there are also one or two good-sized young males with small manes, I believe that they are the offspring of the old male and one or other of the adult females, and that they have lived and hunted with the troop since cubhood. Such young males are probably not driven away to hunt by themselves until they commence to aspire to the affections of one of the females of the party. In 1879 I encountered two pairs of male lions hunting in company in the Mababi country to the north of Lake N'gami. I shot the first pair, and should certainly have killed both the others had I only had a rifle and a few cartridges with me when I first saw them, as they were right out on an open plain from which the grass had been burnt, far away from the nearest bush, and I was riding the best hunting horse I ever possessed. The two lions which I shot were large and heavy, apparently just in their prime, and the other pair also appeared to be full-grown animals. Now the Masarwa Bushmen living near the Mababi plain—and these wild people are extraordinarily acute observers—declared that they knew both these pairs of lions well, and said that each pair were the cubs of one mother, and had been hunting together since cubhood. Curiously enough, in the case of both these pairs of lions the two animals living and hunting together differed from one another very much. In each case one was of a very dark colour all over, with a dark mane, whilst the body of the other was of a pale yellow, and it had scarcely any mane at all. A few days after encountering the second pair of lions, a friend and myself came upon two lionesses on the same open plain, both of which we shot. One of these lionesses was on the point of giving birth to three cubs, which we cut out of her womb. Two of these cubs were males, and they differed very much one from another in colour even before birth. One was very dark indeed, owing to the blackish tint of the tips of the hairs of its little fluffy coat. The other was of a reddish yellow. The fur of the female cub was also of a much lighter colour than in the dark male. Now I cannot but adhere to the opinion which I wrote down in my diary at the time, that these two male lion cubs would, had they lived, have grown up into animals differing very much in appearance one from the other. The dark cub would have become a dark-skinned, dark-maned lion, the lighter coloured one a yellow lion with probably very little mane.
Commenting upon such a case as the above, Mr. R. Lydekker, in one of his recently published zoological essays, says that when light- and dark-maned cubs are met with in the same litter it is due to crossing between lions of different races. Mr. Lydekker has also stated that "with regard to the lion, it has now been ascertained that the black-maned and tawny-maned specimens belong, in most cases at any rate, to distinct local races."
The objection to this theory is that you cannot classify all African lions under two heads, the black-maned and the tawny-maned. Dealing with this subject in 1881, and referring only to the skins of lions I had seen which had been killed in the country between the Limpopo and the Zambesi, I wrote as follows: "I cannot see that there is any reason for supposing that more than one species (of lion) exists, and as out of fifty lion skins scarcely two will be found exactly alike in the colour and length of the mane, I think it would be as reasonable to suppose that there are twenty species as two. The fact is, that between the animal with hardly a vestige of mane and the far handsomer but much less common beast with a long flowing black mane every possible intermediate variety may be found." Since that time I have seen a great many more skins of lions shot in the country to the south of the Zambesi, as well as a number from limited areas of country in East Africa and in Somaliland, and it appears to me that the lions of these two latter very limited areas show exactly the same variations as regards colour and profuseness of mane as their congeners in the more southerly parts of the continent.
I have seen the skins of many lions and lionesses in South Africa, which seemed to be those of full-sized animals though they may have been young in years, showing very well-defined red-brown spots on the legs, flanks, and belly. The old Boer hunters, indeed, had a name for such lions, "bont pod leeuws" (spotted-footed lions), which some of them maintained belonged to a distinct species. I once, however, showed the skins of five lions, which I had recently shot in Mashunaland, to a well-known Boer hunter. One was that of a large male with a fine dark mane. This he declared to be the skin of a "swart voer-leif leeuw" (lion with the front part of the body black); whilst the skin of a lioness which showed a good many spots on the legs and belly, he declared to be that of a "bont pod leeuw, de kwai sort" (spotted-footed lion, the vicious kind). As, however, these two animals were consorting together when I shot them, I do not believe that they belonged to different species or even races. I am inclined to think that lions showing spots on the legs and belly, when adult but still not old, might very likely lose them in later life.
In regard to wild lions, it may be said, as a general proposition, that the mane usually grows round the neck and on the chest only, with a prolongation from the back of the neck to behind the shoulder-blades. Sometimes large full-grown male lions will be practically maneless. Occasionally specimens will be met with in which the entire shoulders as well as the neck will be covered with mane. When writing of lions in 1881, I stated that I had never seen the skin of a wild lion in which the whole belly was covered with long hair, as is so often the case with lions in captivity in this country, though I had seen full-maned wild lions with large tufts of long dark hair on the elbows and in the flanks. A few years later, however, Lo Bengula, the last chief of the Matabele, gave me the skin of a lion which had been killed near the upper course of the Umzingwani river, not far from Bulawayo, with a very fine mane. In this specimen the tufts of hair in the flanks were very profuse, almost meeting across the belly, and there were a few long hairs all over the under parts of the skin. There is also, I think, good evidence to show that in the more southerly portions of South Africa lions not infrequently developed a growth of long hair all over their bellies; for not only are all the lions figured by Captain (afterwards Sir Cornwallis) Harris so adorned, but there is now in the Junior United Service Club in London a mounted specimen of a South African lion with not only an extraordinary wealth of mane covering the whole of the fore-part of the body, but also with a thick growth of long hair all over its belly. This lion is said to have been killed near the Orange river about 1830, probably, I should think, on the bontebok flats, near Colesberg, in the Cape Colony, though possibly on the plains to the north of the river. Now, personally I believe that cold has more to do with the development of a lion's mane than anything else. The winter cold of the high plateaus of the Cape Colony, the Orange Colony, and the Southern Transvaal is much more severe than in any part of Africa where lions exist to-day, and Harris's drawings and the mounted specimen of the lion I have above referred to, which was killed near the Orange river long ago, show that wild lions sometimes attained very profuse manes and had their bellies covered with long hair in that part of Africa. To-day, lions with really fine manes are never found except in countries where the nights are cold during the winter months, such as the Athi plains, the Uas N'gishu plateau, the high downs of Matabeleland and Mashunaland, and the Haud of Somaliland, as well as other elevated regions. In the Pungwe river district some few lions attain fairly good, but never, I believe, extraordinarily profuse manes. Only a certain proportion of the lions found on high and cold plateaus have, however, fine long dark manes. Many have very poor manes, but it seems to me impossible that there can be more than one species of lion in so confined an area. In the hotter parts of Africa, lying below the level of the more elevated plateaus, I think I am correct in saying that lions never get fine manes, and the hotter the climate, the poorer on the average the manes will be. The fact that the high, cold plateaus are always open grasslands free from thorn-bush, whilst the lower parts of the country are usually covered with scrubby bush and thorny thickets, has led many people to think that lions have poor manes in bush-covered countries because the thorns tear out the hair; but I think that this is quite a mistaken idea, for in the western part of Matabeleland, in the neighbourhood of the Ramokwebani and Tati rivers, where the winter nights are very cold, although the whole country is covered with forest, much of it dense thorn-bush, the lions used sometimes to grow very fine long manes. Personally, therefore, I am convinced that climate is the main factor in the production of a lion's mane, and possibly very high feeding may help to produce certain exceptionally fine animals. As the high plateaus of Southern and Eastern Africa have, before the advent of Europeans, always teemed with great multitudes of zebras and antelopes, and in some cases buffaloes as well, the lions of the high and cold plateaus have most certainly always been well fed. The lions living in the Pungwe river district too must, before the advent of Europeans, have been exceptionally well fed.
It has always seemed to me that in Africa and India, where, although the nights may be cold, the sun is always hot, a heavy mane must be more or less of a nuisance and encumbrance to a lion; and I believe that such a wonderful growth of hair must be a reversion to an ancestral adornment first evolved in a cold climate.
The fossil remains of the so-called cave lion (Felis spelaea), which have been discovered in great abundance in the cave deposits of Pleistocene times in Western Europe, are said by Professor Boyd Dawkins to present absolutely no osteological or dental character by which they can be distinguished from those of existing lions, and I think that we are therefore justified in believing that the lion was first evolved in a cold climate, and that in the course of ages it gradually spread south and east, following the migrations of the game on which it preyed. It probably entered Africa before that continent was separated from Europe by the Mediterranean Sea, at the same time as the ancestors of the giraffes, antelopes, buffaloes, elephants, and rhinoceroses of to-day, and accompanied them through Eastern Africa right down to Cape Agulhas. Some lions remained in Europe long after the separation of Africa from that continent, and even in the time of Herodotus these animals appear to have been still common throughout South-Eastern Europe.
As the ancient cave lions which roamed the woods and plains of Western Europe co-existed with bears, mammoths, reindeer, elk, wild cattle, and other denizens of a cold country, there can be little doubt that their coats were thick and furry in both sexes, whilst a heavy mane would have been an adornment to the males without being an encumbrance.
That the flowing mane and shaggy hair on the belly of the male lion were first evolved in a cold climate is, I think, proved by the undoubted fact that there is an inherited tendency in all lions to grow a mane, which is crippled and dwarfed by a hot climate but encouraged by exposure to cold. Quite recently there was a fine lion in the Zoological Society's Gardens at Regent's Park which was presented by Messrs. Grogan and Sharpe. This animal was caught near the Pungwe river, in South-East Africa, and brought to England by these gentlemen when quite a small cub. When full-grown it developed a very much finer mane than I believe has ever been seen in a wild lion that has come to maturity in the part of Africa from which it was brought. Similarly, some thirty years ago there was a very fine lion in the Society's Gardens which was brought by Colonel Knox from the Soudan. Colonel Knox took me to the Gardens to see this animal, and pointed out to me the fact that it had developed a far finer mane (extending much farther back over the shoulders and under the belly) than any man had ever seen in a wild lion in the country from which it came. Lion cubs brought to this country from India also grow fine manes, though I do not think that there is any record of a lion ever having been shot in India with anything more than a fairly good mane. The fact that lion cubs captured in any part of Africa or Asia, and brought up in the comparatively cool and damp climate of Western Europe, always—or nearly always—grow fine manes, which usually cover the whole shoulders and often extend all over the under-surface of the body, and the further fact that in the hotter parts of Africa lions always have very scanty manes, but on the high, cold plateaus often develop good, and occasionally very luxuriant manes, appears to me to show that a heavily maned lion is a reversion to an ancient ancestral type, first evolved in Pleistocene times in a cold and inclement climate.