Essays Upon Heredity and Kindred Biological Problems

Lycaena violacea. According to Mr. Edwards, the first brood of this species lives three to four weeks at the most.

Smerinthus tiliae. A female, which had just emerged from the pupa, was caught on June 24th; on the 29th pairing took place; on the 1st of July she laid about eighty eggs, and died the following day. She lived nine days, taking no food during this period, and she only survived the deposition of eggs by a single day.

Macroglossa stellatarum. A female, captured on the wing and already fertilized, lived in confinement from June 28th to July 4th. During this time she laid about eighty eggs, at intervals and singly; she then disappeared, and must have died, although the body could not be found among the grass at the bottom of the cage in which she was confined.

Saturnia pyri. A pair which quitted the cocoons on the 24th or 25th of April, remained in coitu from the 26th until May 2nd—six or seven days; the female then laid a number of eggs, and died.

Psyche graminella. The fertilized female lives some days, and the unfertilized female over a week (Speyer).

Solenobia triquetrella. ‘The parthenogenetic form (I refer to the one which I have shown to be parthenogenetic in Oken’s ‘Isis,’ 1846, p. 30) lays a mass of eggs in the abandoned case, soon after emergence. The oviposition causes her body to shrivel up, and some hours afterwards she dies. The non-parthenogenetic female of the same species remains for many days, waiting to be fertilized; if this does not occur, she lives over a week.’ ‘The parthenogenetic female lives for hardly a day, and the same is true of the parthenogenetic females of another species of Solenobia’ (S. inconspicuella?). Letter from Dr. Speyer.

Psyche calcella, O. The males live a very short time; ‘those which leave the cocoon in the evening are found dead on the following morning, with their wings fallen off, at the bottom of their cage.’ Dr. Speyer.

Eupithecia, sp. (Geometridae), ‘when well-fed, live for three to four weeks in confinement; the males fertilize the females frequently, and the latter continue to lay eggs when they are very feeble, and are incapable of creeping or flying.’ Dr. Speyer.

The conclusions and speculations in the text seem to be sufficiently supported from this short series of observations. There remains, as we see, much to be done in this field, and it would well repay a lepidopterist to undertake some exact observations upon the length of life in different butterflies and moths, with reference to the conditions of life—the mode of egg-laying, the degeneracy of the wings, and of the external mouth-parts or the closure of the mouth itself. It would be well to ascertain whether such closure does really take place, as it undoubtedly does in certain plant-lice.

VII. Coleoptera

Melolontha vulgaris. Cockchafers, which I kept in an airy cage with fresh food and abundant moisture, did not in any case live longer than thirty-nine days. One female only, out of a total number of forty-nine, lived for this period; a second lived thirty-six days, a third thirty-five, and a fourth and fifth twenty-four days; all the rest died earlier. Of the males, only one lived as long as twenty-nine days. These periods are less by some days than the true maximum duration of life, for the beetles were captured in the field, and had lived for at least a day; but the difference cannot be great, when we remember that out of forty-nine beetles, only three females lived thirty-five to thirty-nine days, and only one male twenty-nine days. Those that died earlier had probably lived for some considerable time before being caught.

Exact experiments with pupae which have survived the winter would show whether the female really lives for ten days more than the male, or whether the results of my experiment were merely accidental. I may add that coitus frequently took place during the period of captivity. One pair, observed in this condition on the 17th, separated in the evening; they paired again on the morning of the 18th, and separated in the middle of the day. Coitus took place between another pair on the 22nd, and again on the 26th.

I watched the gradual approach of death in many individuals: some days before it ensued, the insects became sluggish, ceased to fly and to eat, and only crept a little way off when disturbed: they then fell to the ground and remained motionless, apparently dead, but moved their legs when irritated, and sometimes automatically. Death came on gradually and imperceptibly; from time to time there was a slow movement of the legs, and at last, after some hours, all signs of life ceased.

In one case only I found bacteria present in great numbers in the blood and tissues; in the other individuals which had recently died, the only noticeable change was the unusual dryness of the tissues.

Carabus auratus. An experiment with an individual, caught on May 27th, gave the length of life at fourteen days; this is probably below the average, since the beetles are found, in the wild state, from the end of May until the beginning of July.

Lucanus cervus. Captured individuals, kept in confinement, and fed on a solution of sugar, never lived longer than fourteen days, and as a rule not so long. The beetles appear in June and July, and certainly cannot live much over a month. As is the case with many beetles appearing during certain months, the length of the individual life is shorter than the period over which they are found. Accurate information, especially as to any difference between the lengths of life in the sexes, is not obtainable.

Isolated accounts of remarkably long lives among beetles are to be found scattered throughout the literature of the subject. Dr. Hagen, of Cambridge, Mass., has been kind enough to draw my attention to these, and to send me some observations of his own.

Cerambyx heros. One individual lived in confinement from August until the following year14.

Saperda carcharias. An individual lived from the 5th of July until the 24th of July of the next year15.

Buprestis splendens. A living individual was removed from a desk which had stood in a London counting-house for thirty years; from the condition of the wood it was evident that the larva had been in it before the desk was made16.

Blaps mortisaga. One individual lived three months, and two others three years.

Blaps fatidica. One individual which was left in a box and forgotten, was found alive when the box was opened six years afterwards.

Blaps obtusa. One lived a year and a half in confinement.

Eleodes grandis and E. dentipes. Eight of these beetles from California were kept in confinement and without food for two years by Dr. Gissler, of Brooklyn; they were then sent to Dr. Hagen who kept them another year.

Goliathus cacicus. One individual lived in a hot-house for five months.

In addition to these cases, Dr. Hagen writes to me: ‘Among the beetles which live for more than a year,—Blaps, Pasimachus, (Carabidae)—and among ants, almost thirty per cent. are found with the cuticle worn out and cracked, and the powerful mandibles so greatly worn down that species were formerly founded upon this point. The mandibles are sometimes worn down to the hypodermis.’

From the data before me I am inclined to believe that in certain beetles the normal length of life extends over some years, and this is especially the case with the Blapidae. It seems probable that in these cases another factor is present,—a vita minima, or apparent death, a sinking of the vital processes to a minimum in consequence of starvation, which we might call the hunger sleep, after the analogy of winter sleep. The winter sleep is usually ascribed to cold alone, and some insects certainly become so torpid that they appear to be dead when the temperature is low. But cold does not affect all insects in this way. Among bees, for example, the activity of the insects diminishes to a marked extent at the beginning of winter, but if the temperature continues to fall, they become active again, run about, and as the bee-keepers say, ‘try to warm themselves by exercise’; by this means they keep some life in them. If the frost is very severe, they die. In the tropics the period of hibernation for many animals coincides with the time of maximum heat and drought. This shows that the organism can be brought into the condition of a vita minima in various ways, and it would not be at all remarkable if such a state were induced in certain insects by hunger. Exact experiments however are the only means by which such a suggestion can be tested, and I have already commenced a series of experiments. The fact that certain beetles live without food for many years (even six) can hardly be explained on any other supposition, for these insects consume a fair amount of food under normal conditions, and it is inconceivable that they could live for years without food, if the metabolism were carried on with its usual energy.

A very striking example, showing that longevity may be induced by the lengthening of the period of reproductive activity, is communicated to me by Dr. Adler in the following note: ‘Three years ago I accidentally noticed that ovoviviparous development takes place in Chrysomela varians,—a fact which I afterwards discovered had been already described by another entomologist.

‘The egg passes through all the developmental stages in the ovary; when these are completed the egg is laid, and a minute or two afterwards the larva breaks through the egg-shell. In each division of the ovary the eggs undergo development one at a time; it therefore follows that they are laid at considerable intervals, so that a long life becomes necessary in order to ensure the development of a sufficiently long series of eggs. Hence it comes about that the females live a full year. Among other species of Chrysomela two generations succeed each other in a year, and the duration of life in the individual varies from a few months to half a year.’

VIII. Hymenoptera

Cynipidae. I have been unable to find any accurate accounts of the duration of life in the imagos of saw-flies or ichneumons; but on the other hand I owe to the kindness of Dr. Adler, an excellent observer of the Cynipidae, the precise accounts of that family which are in my possession. I asked Dr. Adler the general question as to whether there was any variation in the duration of life among the Cynipidae corresponding to the conditions under which the deposition of eggs took place; whether those species which lay many eggs, or of which the oviposition is laborious and protracted, lived longer than those species which lay relatively few eggs, or easily and quickly find the suitable places in which to deposit them.

Dr. Adler fully confirmed my suppositions and supported them by the following statements:—

‘The summer generation of Neuroterus (Spathegaster) has the shortest life of all Cynipidae. Whether captured or reared from the galls I have only kept them alive on an average for three to four days. In this generation the work of oviposition requires the shortest time and the least expenditure of energy, for the eggs are simply laid on the surface of a leaf. The number of eggs in the ovary is also smaller than that of other species, averaging about 200. This form of Cynips can easily lay 100 eggs a day.

‘The summer generation of Dryophanta (Spathegaster Taschenbergi, verrucosus, etc.) lives somewhat longer; I have kept them in confinement for six to eight days. The oviposition requires a considerable expenditure of time and strength, for the ovipositor has to pierce the rather tough mid-rib or vein of a leaf. The number of eggs in the ovary averages 300 to 400.

‘The summer generation of Andricus, which belongs to the extensive genus Aphilotrix, have also a long life. I have kept the smaller Andricus (such as A. nudus, A. cirratus, A. noduli) alive for a week, and the larger (A. inflator, A. curvator, A. ramuli) for two weeks. The smaller species pierce the young buds when quite soft, but the larger ones bore through the fully grown buds protected by tough scales. The ovary of the former contains 400 to 500 eggs, that of the latter over 600.

‘The agamic winter generations live much longer. The species of Neuroterus have the shortest life; they live for two weeks at the outside; on the other hand, species of Aphilotrix live quite four weeks, and Dryophanta and Biorhiza even longer. I have kept Dryophanta scutellaris alive for three months. The number of eggs in these agamic Cynipidae is much larger: Dryophanta and Aphilotrix contain 1200 and Neuroterus about 1000.’

It is evidently, therefore, a general rule that the duration of life is directly proportional to the number of eggs and to the time and energy expended in oviposition. It must of course be understood that, here as in all other instances, these are not the only factors which determine the duration of life, but many other factors, at present unknown, may be in combination with them and assist in producing the result. For example, it is very probable that the time of year at which the imagos appear exerts some indirect influence. The long-lived Biorhiza emerges from the gall in the middle of winter, and at once begins to deposit eggs in the oak buds. Although the insect is not sensitive to low temperature, for I have myself seen oviposition proceeding when the thermometer stood at 5° R., yet very severe frost would certainly lead to interruption and would cause the insect to shelter itself among dead leaves on the ground. Such interruptions may be of long duration and frequently repeated, so that the remarkably long life of this species may perhaps be looked upon as an adaptation to its winter life.

Ants. Lasius flavus lays its eggs in the autumn, and the young larvae pass the winter in the nest. The males and females leave the cocoons in June, and pair during July and August. The males fly out of the nest with the females, but they do not return to it; ‘they die shortly after pairing.’ It is also believed that the females do not return to the nest, but found new colonies; this point is however one of the most uncertain in the natural history of ants. On the other hand it is quite certain that the female may live for years within the nest, continuing to lay fertilized eggs. Old females are sometimes found in the colony, with their jaws worn down to the hypodermis.

Breeding experiments confirm these statements. P. Huber17 and Christ have already put the life of the female at three to four years, and Sir John Lubbock, who has been lately occupied with the natural history of ants, was able to keep a female worker of Formica sanguinea alive for five years; and he has been kind enough to write and inform me that two females of Formica fusca, which he captured in a wood together with ten workers, in December 1874, are still alive (July 1881), so that these insects live as imagos for six and a half years or more18.

On the other hand, Sir John Lubbock never succeeded in keeping the males ‘alive longer than a few weeks.’ Both the older and more recent observers agree in stating that female ants, like queen bees, are always protected as completely as possible from injury and danger. Dr. A. Forel, whose thorough knowledge of Swiss ants is well known, writes to me,—‘The female ants are only once fertilized, and are then tended by the workers, being cleaned and fed in the middle of the nest: one often finds them with only three legs, and with their chitinous armour greatly worn. They never leave the centre of the nest, and their only duty is to lay eggs.’

With regard to the workers, Forel believes that their constitution would enable them to live as long as the females (as the experiments of Lubbock also indicate), and the fact that in the wild state they generally die sooner than the females is ‘certainly connected with the fact that they are exposed to far greater dangers.’ The same relation seems also to obtain among bees, but with them it has not been shown that in confinement the workers live as long as the queens.

Bees. According to von Berlepsch19 the queen may as an exception live for five years, but as a rule survives only two or three years. The workers always seem to live for a much shorter period, generally less than a year. Direct experiments upon isolated or confined bees, or upon marked individuals in the wild state, do not prove this, but the statistics obtained by bee-keepers confirm the above. Every winter the numbers in a hive diminish from 12,000-20,000 to 2000-3000. The queen lays the largest number of eggs in the spring, and the workers which die before the winter are replaced by those which emerge in the summer, autumn or during a mild winter. The queen lays eggs at such a variable rate throughout the year that the above-mentioned inequality in numbers is explained. The workers do not often live for more than six to seven months, and at the time of their greatest labour, (May to July), only three months. An attempt to calculate the length of life of the workers and drones by taking stock at the end of summer, gives six months for the former and four months for the latter20.

The drones do not as a rule live so long as four months, for they meet with a violent death before the end of this period. The well-known slaughter of the drones is not, according to the latest observations, brought about directly by means of the stings of the workers, but by these latter driving away the useless drones from the food so that they perish of starvation.

Wasps. It is interesting that among these near relations of the bees, the life of the female should be much shorter, corresponding to the much lower degree of specialization found in the colonies. The females of Polistes gallica and of Vespa not only lay eggs but take part in building the cells and in collecting food; they are therefore obliged to use all parts of the body more actively and especially the wings, and are exposed to greater danger from enemies.

It is well known from Leuckart’s observations, that the so-called ‘workers’ of Polistes gallica and Bombus are not arrested females like the workers of a bee-hive, but are females which although certainly smaller, are in every way capable of being fertilized and of reproduction. Von Siebold has nevertheless proved that they are not fertilized, but reproduce parthenogenetically.

The fertilized female which survives the winter, commences to found a colony at the beginning of May: the larvæ, which hatch from the first eggs, which are about fifteen in number, become pupæ at the beginning of June, and the imagos appear towards the end of the same month. These are all small ‘workers,’ and they perform such good service in tending the second brood, that the latter attain the size of the female which founded the colony; only differing from her in the perfect condition of their wings, for by this time her wings are greatly worn away.

The males appear at the beginning of July; their spermatozoa are mature in August, and pairing then takes place with certain ‘special females which require fertilization’ which have in the meantime emerged from their cocoons. These are the females which live through the winter and found new colonies in the following spring. The old females of the previous winter die, and do not live beyond the summer at the beginning of which they founded colonies. At the first appearance of frost, the young fertilized females seek out winter quarters; the males which never survive the winter, do not take this course, but perish in October. The parthenogenetic females, which remain in the nest during the nuptial flight, also perish.

The males of Polistes gallica do not live longer than three months—from July to the beginning of October; the parthenogenetic females live a fortnight longer at the outside—from the middle of June to October, but the later generations have a shorter life. The sexual females alone live for about a year, including the winter sleep.

A similar course of events takes place in the genus Vespa. In both these genera the possibility of reproduction is not restricted to a single female in the nest, but is shared by a number of females. In the genus Apis alone is the division of labour complete, so that only a single female (the queen) is at any one time capable of reproduction, a power which differentiates it from the sterile workers.

Note 4. The Duration of Life of the Lower Marine Animals

I have only met with one definite statement in the literature of this part of the subject. It concerns a sea anemone,—which is a solitary and not a colonial form. The English zoologist Dalyell, in August, 1828, removed an Actinia mesembryanthemum from the sea and placed it in an aquarium21. It was a very fine individual, although it had not quite attained the largest size; and it must have been at least seven years old, as proved by comparison with other individuals reared from the egg. In the year 1848, it was about thirty years old, and in the twenty years during which it had been in captivity it had produced 334 young Actiniae. Prof. Dohrn, of Naples, tells me that this Actinia is still living to-day, and is shown as a curiosity to those who visit the Botanical Gardens in Edinburgh. It is now (1882) at least sixty-one years old22.

Note 5. The Duration of Life in

Indigenous Terrestrial and Fresh-water Mollusca

I am indebted to Herr Clessin—the celebrated student of our mollusca—for some valuable notes upon our indigenous snails and bivalves (Lamellibranchiata). I could not incorporate them in the text, for a number of necessary details as to the conditions of life are at present entirely unknown, or are at least only known in a very fragmentary manner. No statistics as to the amount of destruction suffered by the young are available, and even the number of eggs produced annually is only known for a few species. I nevertheless include Herr Clessin’s very interesting communications, as a commencement to the life statistics of the Mollusca.

(1) ‘Vitrinae are annual; the old animals die in the spring, after having produced the spawn from which the young develope. These continue to grow until the following spring.’

(2) ‘The Succineae are mostly biennial; Succinea putris probably triennial. Fertilization takes place from June till the beginning of August, and the young develope until the autumn. Succinea Pfeifferi and S. elegans live through the winter, and the fact is proved by very distinct annual markings. Reproduction takes place in July and August of the following year, and they die in the autumn. They continue to grow until their death.’

(3) ‘The shells of our native species of Pupa, Clausilia, and Bulimus (with the exception of Bulimus detritus) show but faint annual markings. They can hardly require more than two years for their complete development. The great number of living individuals with full-sized shells belonging to these genera, as compared with the number which possess smaller shells, makes it probable that these animals live in the mature condition longer than our other Helicidae. I have always found full-sized shells present in at least two-thirds of the individuals of these genera characterized by much-coiled shells—a proportion which I have never seen among our larger Helicidae. Nevertheless direct observations as to the length of life in the mature condition are still wanting.’

(4) ‘The Helicidae live from two to four years; Helix sericea, H. hispida, two to three years; H. hortensis, H. nemoralis, H. arbustorum, as a rule three years; H. pomatia four years. Fertilization is not in these species strictly confined to any one time of year, but in the case of old animals takes place in the spring, as soon as the winter sleep is over; while in the two-year-old animals it also happens later in the summer.’