II Longevity in the Animal Kingdom
Longevity in the lower animals—Instances of long life in sea-anemones and other invertebrates—Duration of life of insects—Duration of life of “cold-blooded” vertebrates—Duration of life of birds—Duration of life of mammals—Inequality of the duration of life in males and females—Relations between longevity and fertility of the organism
It is wonderful to what an extent the duration of life varies amongst animals, the slightest examination of the facts showing that very many factors must be involved.
As the higher animals are nearly always larger than invertebrates, if there be a definite relation between longevity and size, one would expect to find that vertebrates live longer than invertebrates. However, this is not the case. Amongst animals of extremely simple organisation, there are some which reach a great age. A striking example of this is found in sea-anemones. These animals have a very simple structure, without a separate digestive canal, and with a badly developed, diffused nervous system, and yet have lived very long in captivity. More than forty years ago, I remember having seen in the possession of M. Lloyd, the Director of the Aquarium at Hamburg, an anemone that he had kept alive for several dozen years in a glass bowl. Another sea-anemone, belonging to the species Actinia mesembryanthemum, is known to have lived 66 years. It was captured in 1828 by Dalyell, a Scottish zoologist, and was then quite adult, and probably about 7 years old. It survived its owner for 36 years, and died in Edinburgh in 1887, the cause of death being unknown. Although they are thus capable of living so long, the rate of growth of members of this species is rapid, and their fertility is very high. According to Dalyell, these anemones reach the adult condition in 15 months. The specimen in his possession, in the 20 years from 1828 to 1848 produced 334 larvæ, then after a period of sterility it gave birth, in one night (1857) to 230 young anemones. This extraordinary prolificness decreased with age, but even when it was 58 years old it used to produce from 5 to 20 at a time. In the seven years from 1872 onwards, it gave birth to 150 young anemones.35 This animal, which certainly was not more than the fortieth or the fiftieth of the weight of an adult rabbit, lived six or seven times as long.
Ashworth and Nelson Annandale have published their observations on another sea-anemone, of the species Sagartia troglodytes, which was 50 years old. It differed from younger examples only in being less prolific.
There are other polyps, such as Flabellum, which do not live more than 24 years, although we have no knowledge as to the cause of the different duration of life.
The variation in the length of the life of molluscs and insects is extremely great. Some species of gasteropods (Vitrina, Succinea) live only a very few years, whilst others (Natica heros) can reach thirty years. Some of the marine bivalves, as for instance, Tridacna gigas, can live to sixty or a hundred years.36
Insects are animals as variable in their duration of life as they are in other respects. Some live only a few weeks; some of the plant-lice, for instance, die in a month. In the same order of Insects, however, (Hemiptera) there are species of cicada which live thirteen to seventeen years, that is to say, much longer than such little Rodents as rats, mice, and guinea-pigs. The larva of an American species spends seventeen years buried in the ground in orchards, where it feeds on the roots of apple trees, and the species is known as Cicada septemdecim, because of this duration of life. In the adult stage the insect lives little more than a month, just time enough to lay the eggs, and bring into the world the new generation, which in its turn will not appear above ground until after another period of seventeen years.
Between these extremes of long and short life, there is to be found amongst insects almost every gradation of longevity. Science, in its present state, has failed to find any law governing these facts. Rules which hold good up to a certain point in the case of the higher animals break down in their application to insects. The large grasshoppers and locusts, for instance, live a much shorter time than many minute beetles. Queen bees, the fertility of which is very great, live two or three years and may reach a fifth year, whilst worker bees, which are infertile, die in the first year of their existence. Female ants, although these are small and extremely prolific, reach the age of seven years.37
We know so little about the physiological processes of insects, that we cannot as yet make even a guess at the cause of this great variation in their longevity. It is more probable that we shall find some explanation in the case of vertebrates concerning which we know much more.
Analysis of the facts shows that whilst in the evolution from fish to mammal there has been a great increase in complexity of organisation, there has at the same time been a reduction in the duration of life. As a general rule, it may be laid down that the lower vertebrates live longer than mammals.
The facts about the longevity of fish are not very numerous, but it seems clear that these animals reach a great age. The ancient Romans, who used to keep eels in aquaria, have noted that these fish would live for more than sixty years. There is reason to believe that salmon can live for a century, whilst pike live much longer. There is, for instance, the much quoted instance of the pike stated by Gessner to have been captured in 1230 and to have lived for 267 years afterwards. Carps are regarded as equally long lived, Buffon setting down their period of life as 150 years. There is a popular idea that the carp in the lakes at Fontainebleau and Chantilly are several centuries old, but E. Blanchard throws doubt on the accuracy of this estimate, inasmuch as during revolutionary times most of the carp were eaten when the palaces were overrun by the populace. There is no doubt, however, that the life of carp may be very long indeed. Not very much is known about the duration of life in batrachians, but it is certain at least that some small frogs may live twelve or sixteen years, and toads as many as thirty-six years.
More is known about the life of reptiles. Crocodiles and caymans, which are large and which grow very slowly, attain great ages. In the Paris Museum of Natural History there are crocodiles which have been kept for more than forty years without showing signs of senescence. Turtles, although they are smaller than crocodiles, live still longer. A tortoise has lived for eighty years in the garden of the Governor of Cape Town, and is believed to have reached the age of two hundred years. Another tortoise, a native of the Galapagos Islands, is known to be 175 years old, whilst a specimen in the London Zoological Gardens is 150 years old. A land tortoise (Testudo marginata) has been kept in Norfolk, England, for a century. I am informed that in the Archbishop’s palace at Canterbury, there is to be seen the carapace of a tortoise which was brought to the Palace in 1623 and which lived there for 107 years.38 Another tortoise, brought to Fulham by Archbishop Laud, lived in the Palace for 128 years. I have already referred to a specimen of Testudo mauritanica, the history of which is known for 86 years, but which is probably much older.
Very little is known as to the longevity of lizards and serpents, but it may be inferred from what I have said about other reptiles that reptiles as a class are able to reach great ages.
It is an easy inference that the great duration of life in cold-blooded animals is associated with the slowness of the physiological processes in these creatures. The circulation, for instance, is so slow, that the heart of a tortoise beats only 20 to 25 times in a minute. Weismann has suggested that one of the factors influencing the duration of life is the rapidity or slowness of the vital activities, the times taken by the processes of absorption and nutrition.
On the other hand, the blood is hot and the vital activities are rapid in birds, and yet birds may attain great ages. Although in the last chapter I gave a number of examples, the subject is so important that I propose to go further into details. The possibility of this is due to an admirable set of details brought together by Mr. J. H. Gurney.39 In his list, in which are included more than fifty species of birds, the lowest figures are from eight and a half to nine years (Podargus cuvieri, Chelidon urbica), and a duration of life so short is an exception, a period of from fifteen to twenty years being more common. Canaries have lived in captivity from 17 to 20 years, and goldfinches up to 23 years. Field larks have lived for 24 years, the Lesser Black-backed Gull 31 years and the Herring Gull 44 years. Birds of medium size may live for several dozens of years, whether they live on animal or on vegetable food, whether they are prolific or lay very few eggs. I will quote only a few instances. Of forty parrots the minimum and maximum ages were respectively 15 and 81 years, and the average 43 years. Without accepting the truth of the story mentioned by Humboldt according to which certain parrots survived an extinct race of Indians, at least we may be certain that great ages have sometimes been reached by these birds. Levaillant mentions a parrot (Psittacus erithaceus) which lost its memory at the age of 60 years, its sight at 90 years, and which died aged 93 years. Another individual, probably of the same species, is reported by J. Jennings to have reached the age of 77. Jones, Layard, and Butler are the authorities for instances of Sulphur-crested Cockatoos having reached respectively 30, 72 and 81 years. M. Abrahams states that an Amazon (Chrysotis amasonica) lived 102 years. I myself have observed two cases of great longevity in the same species of parrot. One of these birds died at the age of 82 years, apparently simply from old age, whilst the other, which was in my possession for several years before it died at the age of 70 to 75 years, was vigorous, showing no signs of senility, but died of pneumonia.
Mr. Gurney found that parrots were not the only birds capable of reaching a great age. One raven reached 69 years and another 50, an Eagle-owl (Bubo maximus) 68 years, another 53, a condor 52, an imperial eagle 56, a common heron 60, a wild goose 80, and a common swan 70 years. None of these examples approaches the legendary three centuries attributed to the swan, but it is evident that many different kinds of birds may attain great age. I can add some cases to those of Mr. Gurney. In the Royal Park at Schönbrunn, near Vienna, a white-headed vulture (Neophron percnopterus) died aged 118 years, a golden eagle (Aquila chrysaëtus) aged 104, and another aged 80 (according to Oustalet). Mr. Pycraft (Country Life, June 25th, 1904) reported that a female eagle, captured in Norway in 1829, had been brought to England and had lived for 75 years. In the last thirty years of its life, it had produced ninety eggs. The same writer mentions the case of a falcon having lived to 162 years.
The collection of facts that I have passed in review make it manifest that birds may have a great duration of life, but that reptiles surpass them in this respect. Birds certainly do not reach the very great ages of crocodiles and tortoises.
Longevity, therefore, is reduced as we ascend in the scale of vertebrate life. We find a still greater reduction when we turn from birds to mammals. Some mammals, it is true, may live as long as birds. Elephants are a good instance. It used to be thought that these giant mammals could live three or four centuries, but I can find no confirmation of the legend, which seems as mythical as that relating to the life of swans. There are no exact data as to the ages reached by wild elephants, but it has been stated that in captivity an elephant rarely but occasionally has completed its century. In zoological gardens and in good menageries, where elephants are well cared for, they seldom live more than 20 to 25 years. Chevrette, an African elephant presented to the Jardin des Plantes by Mehemet Ali, in 1825, lived for only 30 years. In the official list of the Indian Government, which gives the deaths of elephants, it appears that of 138 examples, only one lived more than 20 years after it had been purchased (Brehm’s Mammals).
Flourens, using his own formula, assigned the age of 150 years to elephants as their epiphyses do not fuse with the long bones until the age of 30. So far, I know of no fact to support the conclusion, although it seems fairly well established that occasionally an elephant may reach a century. It is stated that one elephant was in service throughout the whole period of more than 140 years in which Ceylon was occupied by the Dutch. This elephant was found in the stables in 1656. Natives with special knowledge of elephants set down their duration of life as from 80 to 150 years, but say that they begin to grow old at from 50 to 60 years of age. My general conclusion from the facts is that the life of these very large mammals is about the same as that of man who is very much smaller.
Centenarians, extremely rare amongst elephants, do not appear to exist in any other kind of mammals except man. The rhinoceros, another large mammal which is a native of the same countries as the elephant, does not reach a great age. According to Oustalet an Indian rhinoceros died in the menagerie of the Paris Museum at about the age of 25 years, and showed all the signs of senility. Another Indian rhinoceros lived for 37 years in the London Zoological Gardens. Grindon has stated his opinion that the rhinoceros may live for 70 or 80 years, but this seems rather an inference from the slowness of growth than a statement of observed fact.
Horses and cattle are large animals, but do not enjoy very long lives. The usual duration of life in horses is from 15 to 30 years. They begin to grow old about 10 years, and in very rare cases may reach 40 or more. A Welsh pony is said to have reached the age of sixty, but such a case is excessively rare. Two other extreme cases are that of a horse belonging to the Bishop of Metz which died at the age of 50 years, and the charger of Field-Marshal Lacy which died at 46.
The duration of life of cattle is still shorter. Domestic cattle show the first sign of age, a yellow discoloration of the teeth, when five years old. In the sixteenth to eighteenth year the teeth fall out, or break, and the cow ceases to give milk, whilst the bull has lost reproductive power. According to Brehm, cattle live for 25 to 30 years or more. Although the duration of life is short, cattle are not prolific. The gestation period of a cow approaches that of the human race (242-287 days), and there is only one birth a year. The total period of reproductivity lasts only a few years.
The sheep, another domesticated Ruminant, has a life even shorter. According to Grindon, sheep do not live longer than 12 years as a rule, but may reach 14 years, which in their case would be extreme age, as they generally lose their teeth at from 8 to 10 years.
Some Ruminants, such as camels and deer, apparently live longer than sheep or cattle, but I do not know exact facts about them.
The short life of domesticated carnivorous animals is well known. Dogs seldom live more than 16 or 18 years, and even before that, at an age of from 10 to 12, they usually show plain signs of senility. Jonatt has mentioned as an extreme rarity a dog of 22 years of age, and Sir E. Ray Lankester (Comparative Longevity, p. 60) cites another instance, in this case the age being 34 years. The oldest dog that I have been able to procure died at the age of 22.
It is generally believed that cats do not live so long as dogs. The average age which they may attain is usually thought to be 10 or 12 years, but certainly a cat of that age has not the decrepid appearance of an old dog. Thanks to the kindness of M. Barrier, the Director of the Ecole d’Alfort, I have had in my possession a cat 23 years old. It appeared to be quite vigorous, and died from cancer in the liver.
Most rodents, particularly the domesticated kinds, are extremely prolific and very short lived. It is extremely rare for a rabbit to reach the age of 10 years, whilst 7 years is the utmost limit for a guinea-pig. Mice, so far as I can ascertain, do not live more than 5 or 6 years.
It is plain from the facts that I have brought together, that mammals, whether they are large or small, as a rule, have shorter lives than birds. It is probable, therefore, that there is something in the structure of mammals which has brought about a shortening in the duration of their lives.
Whilst most of the lower vertebrates, and all birds, reproduce by laying eggs, the vast majority of mammals are viviparous. As the tax on the parent organism is greater when the young are produced alive than when eggs are laid, it might be thought that in this difference lay the cause of the shorter life of mammals. It is well known that an animal may be made feeble by too great fecundity, and it is conceivable that the kind of parasitic life of the embryos within the body of the mother may weaken her system.
There are many facts, however, which make it impossible to accept such a view. The longevity of mammals is nearly equal in the two sexes, although the tax on the organism caused by reproduction is much greater in the case of females than in males. Longevity, however, cannot be regarded as a character stable in each species and necessarily identical in the two sexes. The animal kingdom presents many cases of disparity in this respect, the difference in longevity in the two sexes being specially striking in species of insects. Generally, the females live longer than the males, as, for instance, amongst the Strepsiptera, where the females have 64 times the duration of life of the males. On the other hand, amongst butterflies, there are cases (e.g., Aglia tau) where the males live longer than the females. In the human race, there is a difference in the longevity of the sexes, the females having the advantage.
As in most cases of disparity in the duration of life the female lives longer than the male, it is plain that the difference cannot be assigned to the drain on the organism caused by reproduction, which, of course, is much greater in females.
Moreover, a closer scrutiny of the facts shows that although mammals do not live so long as birds, the reproductive drain is greater in the case of birds.
It is well known that the productivity of an animal is not necessarily identical with its fecundity. Fish or frogs which lay thousands of eggs at a time (a pike, for example, produces 130,000) are obviously more prolific than, for instance, a sparrow which lays only 18 eggs in a year, or than a rabbit, which in the same time gives birth to from 25 to 50. However, to produce this much smaller quantity of eggs or of young, the sparrow and the rabbit (I have chosen the most prolific bird and mammal) expend a much larger quantity of material than the frog or the fish. The sparrow and the rabbit employ in producing their progeny a bulk of material greater than the weight of their body, whilst the enormous quantity of eggs laid by the frog does not weigh more than one-seventh part of the body of the frog. It may be laid down, as a general rule, that although fecundity, that is to say the number of eggs or of young which are produced, diminishes as the organism becomes more complex, the productivity on the other hand increases, expressed in percentage of weight. The productivity, which is not more than 18 per cent. in batrachia, reaches 50 per cent. in reptiles, 74 per cent. in mammals, and 82 per cent. in birds.
It is plain that if reproduction shortens the life of mammals by weakening the organism, it must be the productivity, not the fecundity, which is the important factor. I have just shown that productivity is greater in birds than in mammals, and in consequence it cannot be on account of any greater burden of reproduction that mammals have a shorter life than birds. The shortness of mammalian life, again, cannot be attributed to the fact that mammals give birth to young, whilst the long-lived reptiles and birds produce eggs, because the longevity of the males, which produce neither young nor eggs, is none the less practically equal to that of the females of the same species. The reason of the short life of mammals must be sought for elsewhere.