LOCUST. 1 In its general acceptation this term is applied only to certain insects of the order Orthoptera, family Acridiidae. The family Locustidae is now viewed zoologically in a sense that does not admit of the species best known as " locusts " being included therein. The idea of a very destructive insect is universally associated with the term; therefore many orthopterous species that cannot be considered true locusts have been socalled; in North America it has even embraced certain Hemiptera-Homoptera, belonging to the Cicadidae, and in some parts of England cockchafers are so designated. In a more narrow definition the attribute of migration is associated with the destructive propensities, and it therefore becomes necessary that a true locust should be a migratory species of the family Acridiidae. Moreover, the term has yet a slightly different signification as viewed from the Old or New World. In Europe by a locust is meant an insect of large size, the smaller allied species being ordinarily known as " grasshoppers," hence the " Rocky Mountain locust " of North America is to Eastern ideas rather a grasshopper than a locust.
In Europe, and a greater part of the Old World, the best known migratory locust is that which is scientifically termed Pachytylus cinerascens with which an allied species P. migratorius has been often confounded. Another locust found in Europe and neighbouring districts is Caloptenus italicus, and still another, Acridium peregrinum, has once or twice occurred in Europe, though its home (even in a migratory sense) is more properly Africa and Asia. These practically include all the locusts of the Old World, though a migratory species of South Africa known as Pachytylus pardalinus (presumed to be distinct from P. migratorius) should be mentioned. The Rocky Mountain locust of North America is Caloptenus spretus, and in that continent there occurs an Acridium (A. americanum) so closely allied to A . peregrinum as to be scarcely distinct therefrom, though there it does not manifest migratory tendencies. In the West Indies and Central America A. peregrinum is also reported to occur.
The females excavate holes in the earth in which the eggs are deposited in a long cylindrical mass enveloped in a glutinous secretion. The young larvae hatch and immediately commence their destructive career. As these insects are " hemimetabolic " there is no quiescent stage; they go on increasing rapidly in size, and as they approach the perfect state the rudiments of the wings begin to appear. Even in this stage their locomotive powers are extensive and their voracity great. Once winged and perfect these powers become infinitely more disastrous, redoubled by the development of the migratory instinct. The laws regulating this instinct are not perfectly understood. Food and temperature have a great deal to do with it, and there is a tendency for the flights to take a particular direction, varied by the physical circumstances of the breeding districts. So likewise each species has its area of constant location, and its area of extraordinary migration. Perhaps the most feasible of the suggestions as to the causes of the migratory impulse is that locusts naturally breed in dry sandy districts in which food is scarce, and are impelled to wander to procure the necessaries of life; but against this it has been argued that swarms bred in a highly productive district in which they have temporarily settled will seek the barren home of their ancestors. Another ingenious suggestion is that migration is intimately connected with a dry condition of the atmosphere, urging them to move on until compelled to stop for food or procreative purposes. Swarms travel considerable distances, though probably generally fewer than 1000 m., though sometimes very much more. As a rule the progress is only gradual, and this adds vastly to the devastating effects. When an extensive swarm temporarily settles in a district, all vegetation rapidly disappears, and then hunger urges it on another stage. The large Old World species, although undoubtedly phytophagous, when compelled by hunger sometimes attack at least dry animal substances, and even cannibalism has been asserted as an outcome of the failure of all other kinds of food. The length of a single flight must depend upon 1 The Lat. locusta was first applied to a lobster or other marine shell-fish and then, from its resemblance, to the insect.
circumstances. From peculiarities in the examples of Acridium peregrinum taken in England in 1869, it has been asserted that they must have come direct by sea from the west coast of Africa; and what is probably the same species has been seen in the Atlantic at least 1 200 m. from land, in swarms completely covering the ship; thus, in certain cases flight must be sustained for several days and nights together. The height at which swarms fly, when their horizontal course is not liable to be altered by mountains, has been very variously estimated at from 40 to 200 ft., or even in a particular case to 500 ft. The extent of swarms and the number of individuals' in a swarm cannot be accurately ascertained. They come sometimes in such numbers as to completely obscure the Sun, when the noise made by the rustling of the wings is deafening. Nevertheless some idea on this point may be formed from the ascertained fact that in Cyprus in 1881, at the close of the season, 1,600,000,000 egg-cases, each containing a considerable number of eggs, had been destroyed; the estimated weight exceeding 1300 tons. Yet two years later, it is believed that not fewer than 5,076,000,000 egg-cases were again deposited in the island.
In Europe the best known and ordinarily most destructive species is Pachytylus cinerascens, and it is to it that most of the numerous records of devastations in Europe mainly refer, but it is probably not less destructive in many parts of Africa and Asia. That the arid steppes of central Asia are the home of this insect appears probable ; still much on this point is enveloped in uncertainty. In any case the area of permanent distribution is enormous, and that of occasional distribution is still greater. The former area extends from the parallel of 40 N. in Portugal, rising to 48 in France and Switzerland, and passing into Russia at 55, thence continuing across the middle of Siberia, north of China to Japan; thence south to the Fiji Islands, to New Zealand and North Australia; thence again to Mauritius FIG. i. Pachytylus migratorius. This and the other figures are all natural size.
and over all Africa to Madeira. The southern distribution is uncertain and obscure. Taking exceptional distribution, it is well known that it occasionally appears in the British Isles, and has in them apparently been noticed as far north as Edinburgh ; so also does it occasionally appear in Scandinavia, and it has probably been seen up to 63 N. in Finland. Looking at this vast area, it is easy to conceive that an element of uncertainty must always exist with regard to the exact determination of the species, and in Europe especially is this the case, because there exists a distinct species, known as P. migratorius, the migratory area of which appears to be confined to Turkestan and eastern Europe.
P. cinerascens is certainly the most common of the " locusts " occasionally found in the British Isles, and E. de Selys-Longchamps is of opinion that it breeds regularly in Belgium, whereas the true P. migratorius is only accidental in that country.
A South African species allied to the preceding and provisionally identified as Pachytylus salcicollis is noteworthy from the manifesta- FIG. 2. Acridium peregrinum.
tion of the migratory instinct in immature wingless individuals. The families of young, after destroying the vegetation of a district, unite in a vast army and move away in search of fresh pastures, devastating the country as they go and proceeding of necessity on foot, hence they are known to the Dutch as " voetgangers." Travelling northwards towards the centre of the continent, the home of their parents before migration, they are diverted from their course by no obstacles. Upon reaching a river or stream they search the bank for a likely spot to cross, then fearlessly cast themselves upon the water where they form floating islands of insects, most of which usually succeed in gaining the opposite bank, though many perish in the attempt.
Acridium peregrinum (fig. 2) can scarcely be considered even an accidental visitor to Europe; yet it has been seen in the south of Spain, and in many examples spread over a large part of England in the year 1869. It is a larger insect than P. migratorius. There is every reason to believe that it is the most destructive locust throughout Africa and in India and other parts of tropical Asia, and its ravages are as great as those of P. migratorius. Presumably it is the species occasionally noticed in a vast swarm in the Atlantic, very far from land, and presumably also it occurs in the West Indies and some parts of Central America. In the Argentine Republic a (possibly) distinct species (A. paranense) is the migratory locust.
Caloptenus italicus (fig. 3) is a smaller insect, with a less extended area of migration; the destruction occasioned in the districts to which it is limited is often scarce less than that of its more terrible allies. It is essentially a species of the Mediterranean district, and especially of the European side of that sea, yet it is also found in North Africa, and appears to extend far into southern Russia.
Caloptenus spretus (fig. 4) is the " Rocky Mountain locust " or " hateful grasshopper " of the North American continent. Though a comparatively small insect, not so large as some of the grasshoppers of English fields, its destructiveness has procured for it great notoriety. By early travellers and settlers the species was not recognized as distinct from some of its non-migratory congeners. But in 1877, Congress appointed a United States Entomological Commission to investigate the subject. The report of the commissioners (C. V Riley , A. S. Packard and C. Thomas) deals with the whole subject of locusts both in America and the Old World. C. spretus has its home or permanent area in the arid plains of the central region east of the Rocky Mountains, extending slightly into the southern portion of Canada; outside this is a wide fringe to which the term sub- permanent is applied, and this is again bounded by the limits of only occasional distribution, the whole occupying a large portion of the North American continent ; but it is not known to have crossed the LOCUST-TREELODGE, E.
Rocky Mountains westward, or to have extended into the eastern states.
As to remedial or preventive measures tending to check the ravages of locusts, little unfortunately can be said; but anything that will apply to one species may be used with practically all. Something can be done (as is now done in Cyprus) by offering a price for all the egg-tubes collected, which is the most direct manner of attacking them. Some little can be done by destroying the larvae while in an FIG. 3. Caloptenus italicus.
unwinged condition, and by digging trenches in the line of march into which they can fall and be drowned or otherwise put an end to. Little can be done with the winged hordes; starvation, the outcome of their own work, probably here does much. In South Africa some success has attended the spraying of the swarms with arsenic. It has been shown that with all migratory locusts the breeding-places, or true homes, are comparatively barren districts (mostly elevated plateaus) ; hence the progress of colonization, and the conversion of those heretofore barren plains into areas of fertility, may (and probably will) gradually lessen the evil._ Locusts have many enemies besides man. Many birds greedily devour them, and it has many times been remarked that migratory swarms of the insects were closely followed by myriads of birds.
FIG. 4. Rocky Mountain Locust (Caloptenus spretus). (After Riley.)
a, a, a, Female in different posi- d, e show the earth partially re- tions, ovipositing. moved, to illustrate an egg- b, Egg-pod extracted from mass already in place, and ground, with the end broken one being placed, open. [ground. /, shows where such a mass has r, A few eggs lying loose on the been covered up.
Predatory insects of other orders also attack them, especially when they are in the unwinged condition. Moreover, they have still more deadly insect foes as parasites. Some attack the fully developed winged insect. But the greater part attack the eggs. To such belong certain beetles, chiefly of the family Cantharidae, and especially certain two-winged flies of the family Bombyliidae. These latter, both in the Old and New World, must prevent vast quantities of eggs from producing larvae.
The larger Old World species form articles of food with certain semi-civilized and savage races, by whom they are considered as delicacies, or as part of ordinary diet, according to the race and the method of preparation. . (R. M'L.; R. I. P.)