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Chapter VIII: Hemiptera, or Bugs—anoplura 532 (23)

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FAM. 15. CIMICIDAE.—_Ocelli absent; elytra very short and broad, so that the broad abdomen is left uncovered. Head short and broad. Rostrum received in a groove beneath the head. Tarsi three-jointed._—Although this family consists of only a dozen species, it is the most notorious of all the Order, as it includes the detestable _Cimex lectularius_ or common Bed-bug. This Insect is now peculiar to the habitations of man, and is said not to trouble savage races; or rather it is supposed to be present only when the habitations have a certain degree of comfort and permanence. It has no fixed period of the year for its development, but the generations succeed one another so long as the temperature {560}is sufficiently elevated; during too cold weather the Insects merely become stupefied, their lives being as it were interrupted till warmth returns. It is a favourite food with other Insects, and is destroyed by cockroaches and ants as well as by _Reduvius_; the small black ant _Monomorium_ will, it is said, clear a house of the bed-bug in a few days. Nothing is really known as to the origin of this Insect; it is now very widely distributed. The other species of the family frequent birds and bats, and are very similar to the common bug. The genus to which the bed-bug belongs is in many works called _Acanthia_ instead of _Cimex_. Other authors apply the term _Acanthia_ to _Salda_, but it is better to allow the name _Acanthia_ to fall into disuse.

FAM. 16. ANTHOCORIDAE.—_Minute bugs, usually with ocelli and with elytra; the latter occasionally abbreviated, but usually fully developed, with membranous tip. Head prolonged in the middle in front much beyond the insertion of the antennae; eyes not far from the thorax. Rostrum free._—These small and obscure Insects appear to be rather numerous in species, and to be chiefly connected with woods and forests. Some of the species live in ants' nests. We have 27 British species belonging to 11 genera. About 200 species of the family are known. The members of the sub-family Microphysides are remarkable from the great dissimilarity of the sexes, for which it is not possible to assign any reason.

FAM. 17. POLYCTENIDAE.—_Proboscis-sheath three-jointed, tarsi four-jointed, antennae four-jointed. Tegmina quite short, of one consistence._—The four or five anomalous species forming this family are parasites on bats of the genus _Molossus_, and have been found in both the Eastern and Western hemispheres. Westwood, who first described {561}them,[493] treated them as aberrant Anoplura or Lice, but there do not appear to be any sufficient grounds for removing these parasites from Hemiptera-Heteroptera. The condition of their alar organs reminds one of what exists in _Cimex_ and _Aëpophilus_, and the mouth is not known to possess any very peculiar structure. We have had no opportunity of making a thorough examination of _Polyctenes_, and therefore speak with some diffidence.

FAM. 18. CAPSIDAE.—_Moderate-sized or small bugs, of delicate consistence, without ocelli; the elytra and wings usually large in proportion to the body, the former with two cells (occasionally only one) in the membrane. Antennae four-jointed, the second joint usually very long, the terminal two more slender than the others. The proboscis not received in a groove. Scutellum exposed, moderately large. Tarsi three-jointed. Female with an ovipositor capable of exsertion._—This family is one of the most extensive of the Hemiptera; we have about 170 species in Britain, where they are most abundant in the south. The exotic species have been but little collected. Their colours are usually delicate rather than vivid, and are never metallic. They frequent plants of all kinds, and many of them skip by the aid of their wings with great agility in the sunshine. The majority probably suck the juices of the plants, but some are known to prey on other Insects. The species of the Indian genus _Helopeltis_ (Fig. 274) are remarkable by possessing a knobbed spine projecting straight up from the scutellum, making the individual look as if it were a specimen with a pin through {562}it: they attack the tea-plant and do considerable damage. They are known as Mosquito-blight. The egg is of comparatively large size, and is placed by the bug in the stems of the tea-plant, but attached to one end of the egg are two long slender threads that project externally. A similar egg (Fig. 275) and method of oviposition have been described by Mr. Dudgeon as occurring in another species of Capsidae, called Moesa-blight, in India.[494]

FAM. 19. SALDIDAE.—_Head short and broad, with large, prominent eyes. Ocelli present. Proboscis not applied to under surface of head or breast in repose. Scutellum large, not covered. Elytra covering the upper surface of the abdomen, formed of three distinct parts. Tarsi three-jointed._—These little bugs run with velocity over mud in damp places, or live in wet moss; some of them can jump; they are all of dark or obscure colour. There are only three genera: _Salda_, of which we have numerous British species, being the principal one.

SERIES 2. CRYPTOCERATA.

The remaining families of Heteroptera are of aquatic habits, and form in nearly all works a separate division called Hemiptera Cryptocerata (or Hydrocorisae, or Hydrocores), distinguished by the antennae being apparently absent; they are, however, really present, being situate on the under side of the head, to which they are closely pressed, or in some cases placed in a pocket in front of each eye. There are six of these families. Schiödte is doubtless correct in treating this division as an unnatural one; it is, however, generally adopted, and is convenient for the purposes of nomenclature and arrangement.

FAM. 20. GALGULIDAE or PELOGONIDAE.—_Form short and broad; head very broad, with prominent eyes, ocelli present. Hind legs thin, formed for running._—The Insects of this family are but little known; they are only sub-aquatic in habits, frequenting damp places at the margins of streams and waters. The presence of ocelli distinguishes them from other water-bugs, with which indeed the Galgulidae appear to be but little related. There are only about twenty species of the family known. We possess none in Britain; but one, _Pelogonus marginatus_, occurs {563}in South Europe. The other members of the family are very widely scattered over the surface of the earth.

FAM. 21. NEPIDAE.—_Abdomen furnished behind with a long slender siphon; front legs more or less elongate for capturing prey, placed quite at the front edge of the prothorax._—This family consists of two interesting but very dissimilar genera, _Nepa_ and _Ranatra_. Both are widely distributed over the earth, and are rather numerous in species.[495] We have one species of each genus in Britain. _Nepa cinerea_, the common "water-scorpion," is one of the commonest of Insects in Southern Britain, living concealed in shallow waters when nearly or quite stagnant. _Ranatra linearis_ (Fig. 276) is much less common, and appears to be getting rarer; it is not recorded from farther north than Cambridge.

The nature of the respiratory arrangements in these Insects is of considerable interest; the long tube at the extremity of the body consists of two parts (as shown in Fig. 276) brought together in the middle, one from each side. Lacaze-Duthiers states that the processes are elongated pleurae, but in the young it is far from clear that this is the case. However that may be, they seem to convey air to the true breathing organs, situate inside the cleft on the apical part of the abdomen itself; but details as to the way in which transfer of air is effected along this {564}very protracted passage are not forthcoming. The development in _Nepa_ has been studied to a certain extent. The apical stigmata are the only pair of the abdominal stigmata that exist in the imago of _Nepa_, the other six pairs being obliterated; the third, fourth, and fifth, according to Schiödte, in a very peculiar manner: hence, as Martin says,[496] the respiratory system is metapneustic. In an earlier stage of the life, however, these six pairs of stigmata exist in functional activity placed in a groove on the under surface of the body; so that the condition is that termed peripneustic, and remains so till the final moult, when the long siphon appears. In the early life there is a short prolongation from the end of the body in connection with the pair of grooves alluded to, but it is a single unpaired organ, and does little therefore to explain the appearance of the siphon, which must, at present, be considered as being suddenly developed at the last moult.

The eggs of Nepidae are remarkable objects; that of the common water-scorpion bears seven filaments at one end (Fig. 277); while that of _Ranatra_ is more elongate, and bears only two, very elongate, threads. These eggs are deposited in the stems of water-plants, being introduced therein, so that the body of the egg is concealed while the threads project: those of _Ranatra_ are placed in stems floating on the water, and in consequence of the threads the stems look as if they were infested by some fungus. The structure and formation of the eggs have been investigated with considerable detail by Korschelt.[497] He looks on the filaments as pneumatic, and considers that they supply a coating of air to the body of the egg; they consist of a spongy mass encircled by two layers of egg-shell, both of these latter being peculiar in structure; the spongy mass is continuous with a layer of the same kind of substance placed on the interior of the shell of the body {565}of the egg. It will be recollected that we have described (p. 562) an egg, apparently of the same nature, deposited by Capsids in the stems of land plants, so that it is very doubtful whether the threads are really connected with the aquatic development of the embryo in Nepidae. But the most interesting feature connected with these eggs is, according to Korschelt, the mode of development of the filaments, which is _sui generis_; the shell of the egg is developed in the ordinary manner as an exudation or excretion from epithelial cells; but the shell of the filament is formed as an intracellular product; a mode of chitin-formation that appears to be peculiar to this structure. Korschelt remarks that "it is in the highest degree worthy of attention how by any process of development through a large number of successive generations so complex a condition could be established as the result of adaptation to external conditions; and this becomes even more interesting when we remember that highly peculiar special processes and departures from the usual modes of tissue-formation are necessary to permit the development of this apparatus."[498]

FAM. 22. NAUCORIDAE.—_No ocelli, and no terminal process to the body; front legs inserted on or near the front of the prosternum. Anterior femora usually broad and flat._—The members of this family are truly aquatic, and swim readily in the water. The family is small, including about nine genera and thirty species, but, like many water-Insects, the genera are widely distributed. We have two in Britain—one of them, _Naucoris_, common; the other, _Aphelocheirus_, rare.

FAM. 23. BELOSTOMIDAE.—_No ocelli, and no long terminal tube to the body; front legs inserted near the front of the prosternum. Posterior tibiae not spiny; flattened and provided with swimming hairs._—Although these Insects have been classified with Nepidae they have but little relation therewith; on the other hand, the distinctions from Naucoridae are far less important. The family includes some of the largest Insects. The South American _Belostoma grande_ attains a length of four or four and a half inches. Notwithstanding their considerable size Belostomidae exist in very large numbers in some localities, and frequently destroy young fish by aid of the powerful though short rostrum.

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They appear to be unable to resist the attraction of artificial light, and are consequently sometimes destroyed in large numbers. It has long been known that species of the genera _Diplonychus_ and _Zaitha_ carry their eggs on their backs. There is no special receptacle for the purpose, but the eggs are kept in their peculiar position by means of a cement insoluble in water. It has been stated by Dimmock that they are placed in position by means of a long, flexible ovipositor. Schmidt, however, found that a specimen of _Diplonychus_, bearing eggs and examined by him, was a male, and he subsequently found that this was the case with other egg-bearing individuals of other species, so that the mode in which the eggs are placed in this position and the object of so curious a habit, remain uncertain. The species of _Belostoma_ are highly remarkable on account of the curious and complex structure of their antennae, in respect of which the nearest analogy is to be found in the large Coleoptera of the genus _Hydrophilus_. A very deep, ear-like pocket, exactly suited to the form of the antennae, exists on the under side of the head; hence in repose no sign of the peculiar shape of the antennae exists. When the antennae are placed in this ear-like pocket only the one side of the basal joints is exposed, the long processes being received into the deep pocket. In _Hydrophilus_ the antenna is used as an accessory organ of respiration, and it will be interesting to learn whether this is also the case in _Belostoma_. Belostomidae have patches of air-carrying pubescence, analogous with those of _Hydrophilus_, on the under sides of the body, elytra and wings, but we do not {567}know how they are charged. Another extremely interesting analogy is found in the manner in which the elytra are locked to the body; a projection from the thoracic side-pieces, forming a long pouch, into which a fold on the inner side of the elytra fits, the two being subsequently locked by the action of some special projections. This arrangement is similar to that which exists in the anomalous family of water-beetles Pelobiidae. In order to make this mechanism more perfect the side-pieces in _Belostoma_ form free processes. Martin has informed us that the young have the metasternal episternum prolonged to form a lamella that he thinks may be for respiratory purposes.[499] About twelve genera and upwards of fifty species of Belostomidae are known. None exist in our isles, but several species extend their range to Southern Europe. In the waters of the warm regions of the continents of both the Old and New Worlds they are common Insects, but as yet they have not been found in Australia.

FAM. 24. NOTONECTIDAE.—_Prosternum short, so that the legs are placed near the back part of it as well as near the front; back of the head overlapped by the front of the pronotum._—The water-boatmen are extremely common in our ponds, where they may be seen rising to the surface and raising the posterior extremity of the body for breathing. They swim on their backs instead of in the usual position, and have an elaborate arrangement of long hairs on the body to assist them to carry about an air-supply. They are said to be lighter than the water, and to have some difficulty in keeping away from the surface. _Notonecta glauca_ is the only British species, but we have a second minute Insect, _Plea minutissima_, belonging to the family. It lies in the mud at the bottom of shallow waters, and may sometimes be fished up in great numbers. It is considered by some authors to form a distinct family. The oviposition of _Notonecta_ has been observed by Regimbart; the eggs are inserted into the stems of aquatic plants.

FAM. 25. CORIXIDAE.—_Prosternum short, as in Notonectidae; summit of the head free from the thorax._—We have numerous species of the genus Corixa in Britain; and others extremely similar in appearance occur in various parts of the world. The head is remarkably free, and capable of great rotation. On dissection it is found to be attached to the thorax only by a {568}narrow area; in this respect it differs widely from _Notonecta_, which possesses an extremely large occipital foramen, and the head of which possesses but little freedom of movement. The extremely short proboscis is more or less retractile, and therefore frequently appears absent. A second British genus consists of a single species, _Sigara minutissima_. These Insects, unlike _Notonecta_, are quite at home beneath the water, where they scurry about with extreme rapidity, and occur sometimes in enormous numbers. In Mexico the eggs of _Corixa americana_ and of _C. femorata_ are used as food, and are said to be very nice. The Insects themselves are used as food in both Mexico and Egypt. The species of this family can make a noise beneath the water by rubbing the front feet against the proboscis.[500] The males have a very complex asymmetry of the terminal segments, and in some species possess on one side of the dorsal surface a curious asymmetrical organ consisting of rows of very closely-packed, intensely black, comb-like plates, called by Buchanan White a strigil. This organ seems to be similar to the peculiar structures found on the terminal segments of certain species of Scutellerides.

Sub-Order II. HOMOPTERA.[501]

FAM. 1. CICADIDAE.—_Head with three ocelli, placed triangularly on the summit between the compound eyes; antennae consisting of a short basal joint, surmounted by a hair-like process divided into about five segments. Front femora more or less thick, armed with teeth. Peduncle (or basal joints) of antennae without sensitive organs._—This important family consists chiefly of large Insects, few being as small as one inch across the expanded wings, while in some the expanse is as much as seven inches. As a rule the four wings are transparent and shining, with the nervures remarkably distinct and dark coloured; but there are numerous forms where the whole creature, including the wings, is highly pigmented in a showy manner; frequently in black and yellow. Cicadas are said to be without any special protection, and to be destroyed in considerable numbers by birds and other animals. The body is broad and robust, and is never shaped into the extravagant forms we meet with in some of the other families of Homoptera.

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Cicadidae are almost confined to the warmer regions of the earth, but we have one species, a great rarity, in the extreme south of England; altogether there are about 800 species known. These Insects are seen above ground—so far as the life-histories are at present known—only in the perfect condition, the creatures in their earlier stages being subterranean and living on roots. As soon as the individual comes out of the ground it splits open the nymph-skin, and the perfect Cicada emerges. One species—the North American _Cicada septendecim_—is a most notorious Insect owing to its life-cycle of seventeen years. It is considered that the individual, after nearly seventeen years of underground existence, comes to the surface and lives for a brief period the life of a noisy Insect. This is the only Insect at present known having so considerable a longevity. This fact, and several other peculiarities, have attracted much attention, so that there is an extensive literature connected with the seventeen-year Cicada. It has a wide distribution over the United States, but does not confine its appearance to every seventeenth year, being found somewhere or other—frequently in numerous localities—almost every year. The evidence as to its periodicity has been obtained by taking the locality and other points into consideration as well as the year of appearance. {570}By so doing it has been found possible to establish the existence of twenty-two broods which are distinguished by consecutive numeration. This being done, the evidence as to the years during which Cicadas have appeared in any given locality is examined, and the result is believed to bear out the view that the life-cycle of the individual Insect is really one of seventeen years. According to this view there are, underground, in certain localities individuals of different ages that will appear on the surface as mature individuals in different years. Thus in 1885 it was understood that there were underground in Alabama two broods, viz. brood xviii. that would appear on the surface in 1894, and brood iv. that would appear on the surface in 1896. The predictions made as to the years in which Cicadas would appear in some given locality are considered to have proved correct. Moreover, particular entomologists have in certain localities verified by personal examination the appearance of the Insects for several consecutive periods of seventeen years. These facts appear fairly conclusive, but they are much complicated by another point, viz. that in certain localities the period is one of thirteen, not of seventeen, years. This is to some extent a question of climate, the thirteen-year interval being chiefly characteristic of the Southern States. It is not, however, entirely so, for there are localities in which the broods have an interval of either thirteen years or seventeen years. Another fact should be remembered, viz. that it is admitted that not quite all the individuals of a particular brood are true to their proper time of appearance; in other words, a few specimens may appear precociously a year or two before their comrades, while some may lag behind to a considerable extent. It is therefore a matter for great surprise that, under these circumstances, the broods should keep distinct at all, for one would suppose that time-variation of this kind would lead to completely obscuring the distinctness of the broods. We must also call attention to the fact that both the seventeen-year and the thirteen-year broods have a dimorphic form, or sub-species, called _C. cassinii_ which accompanies the ordinary form, with which it is apparently as a rule not connected by intermediates.[502]

{571}Cicadidae are provided with powerful ovipositors. The eggs of _C. septendecim_ are deposited in the woody stems of bushes; after remaining there a few weeks the young hatch out, drop to the ground, and, as previously stated, disappear for nearly seventeen years, nearly the whole of which time is passed in the larval state, the nymph-condition existing for only a few days. They feed on the roots of various trees; it has been said that they are injurious in this way, but other authorities maintain that they suck only a moist exudation from the roots. It is very difficult to obtain information as to their strange, prolonged, subterranean life; it said that the Insects sometimes penetrate to a great depth—ten feet, even twenty feet are mentioned;—and as great changes may take place on the surface during their long lives, these Insect Rip Van Winkles sometimes emerge in very strange conditions, and may appear even in deep cellars. When the pupa comes to the surface it hooks itself on to the stem of some plant or other object, the skin of the back splits, and the Cicada emerges. Among the inexplicable peculiarities of this Insect must be mentioned the fact that when emerging it sometimes constructs chimneys, or flues, extending several inches above the surface of the ground. The reason for this is much disputed; it was said that they are for refuge against inundations, but this appears to be very doubtful. Certain of the broods consist of an almost incalculable number of individuals, and it is very strange to hear woods, or other localities, that have been for many years free from these Insects, all at once resounding with their noisy song. The seventeen-year Cicada is considered to be doomed to a speedy extinction; the extension of cultivation and building, and the introduction to America of the English sparrow, are likely to prove too much for the Insect.

Although Hemiptera are classified by many among the Ametabola or Insects without metamorphosis, it is impossible to deny that the Cicadidae exhibit a considerable amount of metamorphosis, and they are usually mentioned as exceptional. The young (Fig. 280, A) is totally unlike the adult in form and colour, and maintains, to a certain extent, its existence by the aid of a different set of implements. The larva of the Cicada is colourless, with an integument of very feeble consistence, rather large antennae, and a remarkable pair of fossorial legs; {572}the wings are totally wanting. The mode of passage from the larval to the pupal state has not been recorded. The pupa, or nymph, differs from the larva by its much shorter, compressed form; by being encased in a remarkably hard shell; and by the antennae approximating in form to those of the adult. It has short wing-pads at the sides of the body; the front legs are remarkably powerful, and the creature is capable of moving about; the imago escapes from the pupa by the splitting dorsally of the middle of the thoracic segments. The empty pupa-skin does not shrivel, but retains its form, and in countries where Cicadas occur, frequently attracts attention by the strange form it presents, being often placed in a conspicuous position.

SONG.—Cicadas are the most noisy of the Insect world; the shrilling of grasshoppers and even of crickets being insignificant in comparison with the voice of Cicada. Darwin heard them in South America when the _Beagle_ was anchored a quarter of a mile from the shore; and _Tympanoterpes gigas_, from the same region, is said to make a noise equal to the whistle of a locomotive.[503] A curious difference of opinion prevails as to whether their song is agreeable or not; in some countries they are kept in cages, while in others they are considered a nuisance. The Greeks are said to have decided in favour of their performances, the Latins against them. Only the males sing, the females being completely dumb; this has given rise to a saying by a Greek poet (so often repeated that it bids fair to become immortal) "Happy the Cicadas' lives, for they all have voiceless wives."[504] The writer considers the songs of the European species he has heard far from unpleasant, but he is an entomologist, and therefore favourably prepossessed; and he admits that Riley's description of the performances of the seventeen-year Cicada is far from a satisfactory testimonial to the good taste of that Insect; Riley says, "The general noise, on approaching the infested woods, is a combination of that of a distant threshing-machine and a distant frog-pond. That which they make when disturbed, mimics a nest of young snakes or young birds under similar circumstances—a sort of scream. They can also produce {573}a chirp somewhat like that of a cricket and a very loud, shrill screech prolonged for fifteen or twenty seconds, and gradually increasing in force and then decreasing." The object, or use of the noise is very doubtful; it is said that it attracts the females to the males. "De gustibus non est disputandum!" perhaps, however, there may be some tender notes that we fail to perceive; and it may be that the absence of any definite organs of hearing reduces the result of a steam-engine whistle to the equivalent of an agreeable whisper. No special auditory organs have been detected[505] as we have already intimated; and certain naturalists, amongst whom we may mention Giard, think that the Insects do not hear in our sense of the word, but feel rhythmical vibrations; it is also recorded that though very shy the Insects may be induced to approach any one who will stand still and clap his hands—in good measure—within the range of their sensibilities. There is a good deal of support to the idea that the males sing in rivalry.

VOCAL STRUCTURES.—Although we may not be able to pronounce a final opinion as to the value to the Insect of the sounds, yet we cannot withhold our admiration from the structures from which they proceed. These are indeed so complex that they must be ranked as amongst the most remarkable voice-organs in the animal kingdom. They are totally different from the stridulating organs that are found in many other Insects, and are indeed quite peculiar to the Cicadidae. Some difference of opinion has existed as to the manner in which the structures act, but the account given by Carlet, some of whose figures we reproduce, will, we believe, be found to be essentially correct. The structures are partly thoracic and partly abdominal. On examining a male _Cicada_ there will be seen on the under surface two plates—the opercula—usually meeting in the middle line of the body and overlapping the base of the abdomen to a greater or less extent according to the species, sometimes nearly covering this part of the body; these are enlargements of the metathoracic epimera; they can be slightly moved away from the abdomen, and, as the latter part is capable of a still greater extent of movement, a wide fissure may be produced, displaying the complex {574}structures. In order to see the parts it is better to cut away an operculum; underneath it three membranes can be seen, an external, the timbal; an anterior, the folded or soft membrane; and a posterior, the mirror. This last is a most beautiful object, tensely stretched and pellucid, yet reflecting light so as to be of varied colours; there are also three stigmata, and some chambers connected with the apparatus. The sound is primarily produced by the vibrations of the timbal, to which a muscle is attached; the other membranes are probably also thrown into a condition of vibration, and the whole skeleton of the Insect helps to increase or modify the sound, which is probably also influenced by the position of the opercula. The stigmata probably play an important part by regulating the tension of the air in the chambers. In the female some of the structures are present in a rudimentary form, but there are no muscles, and this sex appears to be really quite voiceless.

FAM. 2. FULGORIDAE.—_Ocelli two (rarely three, or entirely obsolete), placed beneath the eyes or near the eyes, usually in cavities of the cheeks, antennae placed beneath the eyes, very variable in form; usually of two joints terminated by a very fine hair, the second joint with a peculiar texture of the surface, owing to the existence of sensitive structures_ (Hansen). _Form of head very diverse; vertex and face forming either a continuous curve, or the planes of the vertex and face forming an acute angle, or both {575}prolonged so as to form a projection or growth that may be monstrous. Prothorax neither armed nor unusually developed._

This family is of large extent, and includes at present so great a variety of forms that it is really almost impossible to frame a definition that will apply to all. The unusual situation of the ocelli and the peculiar second joint of the antennae must at present be taken as the best diagnostic characters: occasionally a third ocellus is present. Some of the Fulgoridae are amongst the largest Insects, others are quite small. The family includes the so-called Lantern-flies, in which the front of the head forms a huge misshapen proboscis that was formerly believed to be luminous. Many of the species are of brilliant or beautiful coloration. A great many—and of very different kinds—have the curious power of excreting large quantities of a white, flocculent wax. This is exhibited by our little British Insects of the genus _Cixius_, and in some of the exotic forms is carried to an extent that becomes a biological puzzle. The Tropical American genus _Phenax_ may be cited as an example; being about an inch long it flies about with a large mass of this waxy substance twice as long as itself; indeed, in the Mexican _P. auricoma_, the waxy processes are four or five inches long. This wax forms a favourite food of certain kinds of Lepidoptera, and two or three larvae of a maggot-like nature may frequently be found concealed in the wax of the live Fulgorids; this has been recorded by Westwood as occurring in India; and Champion has observed it in the New World.[506] {576}The wax of Fulgorids is used by the Chinese for candles and other purposes; and this white Insect-wax is said to be much esteemed in India. Very curious chemical substances have been obtained from it, but its importance in the economy of the Insects that produce it is quite obscure. We have about seventy species of Fulgoridae in Britain. They belong to the subfamilies Tettigometrides, Issides, Cixiides, and Delphacides, which by many authors are treated as separate families. The exotic subfamily Flatides is highly peculiar. In some of its members the head is very different from that of the ordinary forms, being narrow, and the vertex and front forming a continuous curve. Some of these Insects are remarkably like butterflies or moths (_e.g._ the African _Ityraea nigrocincta_ and the species of the genus _Pochazia_), but the young are totally unlike the old, the posterior part of the body bearing a large bush of curled, waxy projections, several times the size of the rest of the body.

FAM. 3. MEMBRACIDAE.—_Prothorax prolonged backwards into a hood or processes of diverse forms; antennae inserted in front of the eyes; ocelli two, placed between the two eyes._—This family is of large extent but its members are chiefly tropical, and are specially abundant in America. Although not of large size the Membracidae are unsurpassed for the variety and grotesqueness of their shapes, due to the unusual development of the pronotum. We figure two of these forms (Fig. 283).[507] Very little is known about their {577}habits and life-histories. We have only two species of the family in Britain, and these do not afford any ground for supposing that there are any peculiarities in their lives at all commensurate with the oddness of the Insect's structures. Belt has recorded the fact that in Nicaragua the larvae of certain Homoptera were assiduously attended by ants for the sake of a sweet juice excreted by the bugs, but it is by no means clear that these larvae were really those of Membracidae. In North America _Ceresa bubalus_ and _C. taurina_ place their eggs in an extremely neat manner in the woody twigs of trees. The young have but little resemblance to the adults, the great thoracic hood being absent, while on the back there is on each segment a pair of long, sub-erect processes having fringed, or minutely spiny, margins.[508]

FAM. 4. CERCOPIDAE.—_Ocelli two (occasionally absent) placed on the vertex; antennae placed between the eyes. Thorax not peculiarly formed._—In the characteristic forms of this family the front of the vertex bears a suture, touched on each side by one at right angles to it, or converging to it so as to form a triangle or a sort of embrasure; the hind tibiae have only one to three strong spines. The Cercopidae are much less extraordinary than many of the previously considered families. But some of them have the habit of secreting a large quantity of fluid; and when in the immature stages, certain of them have the art of emitting the liquid in the form of bubbles which accumulate round the Insect and conceal it. These accumulations of fluid are called cuckoo-spits or frog-spits; and the perfect Insects are known as frog-hoppers, their power of leaping being very great. The most abundant of the frog-hoppers in our gardens is _Philaenus spumarius_, a little Insect of about a quarter of an inch long, obscurely coloured, with more or less definite pale spots; it is so variable in colour that it has received scores of names. Some of the Insects do not use their fluid in this manner, but eject it in the form of drops, and sometimes cast them to a considerable distance. The phenomena known as weeping-trees are due to Cercopidae; some of the species make such copious exudations of this kind that the drops have been compared to a shower of rain. In Madagascar it is said that _Ptyelus goudoti_ exudes so much fluid that five or six dozen larvae would about fill a {578}quart vessel in an hour and a half. The frog-spit is considered by some naturalists to be a protective device; the larvae are, however, a favourite food with certain Hymenoptera, which pick out the larvae from the spits and carry them off to be used as stores of provision for their larvae. In Ceylon the larva of _Machaerota guttigera_ constructs tubes fixed to the twigs of the tulip-tree, and from the tube water is exuded drop by drop. According to Westwood, this Insect is intermediate between Cercopidae and Membracidae.[509]

FAM. 5. JASSIDAE.—_Ocelli two, placed just on the front margin of the head (almost in a line with the front of the eyes or more to the front) or on the deflexed frons. Hind tibiae usually with many spines._ This vaguely limited family includes a very large number of small or minute Insects, usually of narrow, parallel form, and frequently excessively delicate and fragile. They are often mentioned under the name of Cicadellinae. Ashmead distinguishes two families, Bythoscopidae, in which the ocelli are clearly on the frons or front, and Jassidae, in which they are on the upper edge thereof. _Ulopa_, _Ledra_, and a few other exceptional forms, are also by many distinguished as representatives of distinct families. Very little is actually known as to the life-histories of these small and fragile Insects, but it is believed that the eggs are usually deposited in the leaves or stems of plants, and more particularly of grasses. In North America the development of _Deltocephalus inimicus_, from hatching to assumption of the adult form, has been observed by Webster to occupy about six weeks. As Jassidae are numerous both in species and individuals it is believed that they consume a considerable part of the vegetation of pastures. Osborn has calculated that on an acre of pasture there exist, as a rule, about one million of these hoppers, and he considers they obtain quite as large a share of the food as the Vertebrates feeding with them.

FAM. 6. PSYLLIDAE.—_Minute Insects with wings usually transparent, placed in a roof-like manner over the body; with three ocelli, and rather long, thin antennae of eight to ten joints. Tarsi two-jointed._—These small Insects have been studied chiefly in Europe and North America, very little information having yet been obtained as to the exotic forms. They are about the {579}size of Aphidae, but in form and general appearance remind one rather of Cicadidae. The wings are in many cases even more perfectly transparent than they are in many Cicadidae. They are sometimes called springing plant-lice, as their habit of jumping distinguishes them from the Aphidae. Löw has called attention to the remarkable variation in colour they present in conformity with either the age of the individual, the food-plant, the climate, and, more particularly, the season of the year.[510] Réaumur long since pointed out that at their ecdyses these Insects go through a remarkable series of changes of colour, and Löw found that this did not take place in the normal manner in the winter generation that hibernates. This has been confirmed by Slingerland in North America in the case of _Psylla pyricola_,[511] which has been introduced there. He finds that there are several generations in the year, and that the hibernating adults differ from the summer adults in size, being nearly one-third larger; in their much darker colouring; and especially in the coloration of the front wings.

In the earlier stages, Psyllidae differ greatly in appearance from the adult forms; the legs and antennae in the newly hatched larvae are short, and have a less number of joints. In the nymph the shape is very peculiar, the large wing-pads standing out horizontally {580}from the sides of the body, so that the width of the creature is about as great as the length. The period occupied by the development apparently varies according to season. Witlaczil, who has given an account of many details of the anatomy and histology of various Psyllidae,[512] considers that there are four larval stages; Heeger's account of _Psylla succincta_ is not quite clear on this point, and Slingerland indicates a stage more than this, the perfect Insect being disclosed as the result of a fifth moult; it is probable that he is correct. In these earlier stages the body bears long hairs called wax-hairs; according to Witlaczil in the young larvae of certain species—_Trioza rhamni_, e.g.—these are broad and flat, so as to make the body appear studded with oval processes; he states that these hairs change their form during the growth of the individual. Nothing is more remarkable in Psyllidae than the amount of matter they secrete or exude from their bodies; in some species the substance is a "honey-dew," and the nymph may keep itself covered with a drop of it: in other cases it is solid, as shown in Réaumur's figures of _P. buxi_, where this exudation forms a string several times longer than the body, and attached to it. Another form of exudation is a light downy or waxy matter. Slingerland says that honey-dew was exuded by _P. pyricola_ in such quantities that it "literally rained from the trees upon the vegetation beneath; in cultivating the orchard the back of the horse and the harness often became covered with the sticky substance dropping from the trees. It attracts thousands of ants, bees, and wasps, which feed upon it." The writer last year observed in the New Forest a stunted sloe-bush, about which a large number of Bombi were busily occupied; and examination showed that they were thrusting their proboscides into the curled and deformed leaves, in which were secreted nymphs of a _Psylla_ exuding honey-dew. It must not be assumed that this honey-dew is the excrement of the Insect; this also is known, and is a different substance. Those who have tasted it say that the honey-dew has a clean, good flavour. The source of the honey-dew is not quite certain, but it seems probable that it comes, like the solid matter figured by Réaumur, directly from the alimentary canal, and not from hairs or pores on the body. Psyllidae give rise to definite formations or galls on certain plants; sometimes these Psyllid galls are mere changes in form of a limited part, or {581}parts, of a leaf, giving rise either to crumpling or to growth of a portion in one direction only, so that on one surface of the leaf a swelling is formed, and on the opposite side a more or less deep cavity in which the Insect dwells. A formation of this kind on the leaves of _Aegopodium podagraria_ is described by Thomas[513] who states that the growth is due to the deposition of an egg of the _Psylla_, and is independent of the after life of the Insect; a fungus—_Puccinia aegopodii_—forms similar structures on the leaves. Structures much more definite than this may be the result of the attacks of Psyllidae; for an example the reader may refer to Réaumur's account of _Psylla buxi_.[514] In Australia and Tasmania there are Psyllidae known as Laap or Lerp Insects, the products of which are called leaf-manna or Lerp, and are used as food. This manna is a scale produced by the young Insect on the leaves of _Eucalyptus_ as a covering or protection. The scale is fastened to the leaf by a hinge, and is somewhat like the shell of a cockle. Although the scales are said to be in some cases objects of great beauty, very little is known about these Australian Psyllidae, one of which has, however, been referred by Schwarz to the genus _Spondyliaspis_, Signoret.[515] About 160 species of Psyllidae are known to occur in the Palaearctic region, and about fifty of them have been found in Britain.[516]

FAM. 7. APHIDAE (_Plant-lice or Green-fly._)—_Minute Insects; as usually met with destitute of wings, though many individuals have two pairs of transparent wings. Antennae long, or moderately long, three- to seven-jointed; abdomen frequently with a pair of tubes (siphons), or short processes on the upper side of the fifth abdominal segment. Tarsi two-jointed, first joint sometimes excessively short._—These soft-skinned Insects are frequently called blight, and are so abundant in temperate climates that a garden, however small, is sure to afford abundance of specimens during the warm months of the year. This great abundance is due to peculiarities in the physiological processes that render these obscure little animals highly important creatures; the individual life for several generations is restricted to constant, or at any rate copious, imbibition of food, accompanied by an almost uninterrupted {582}production of young by parthenogenetic females, the young so produced becoming rapidly (sometimes in the course of eight or ten days, but more usually in about twenty days) themselves devoted to a similar process; so that in the comparatively short period of a few months the progeny resulting from a single individual is almost innumerable. This remarkable state of affairs is accompanied by other peculiarities of physiology, with the result that the life-histories of successive generations become very diverse, and complex cycles of series of generations differing more or less from one another are passed through, the species finally returning to bi-sexual reproduction, and thus inaugurating another cycle of generations. The surprising nature of these facts has in the last 150 years caused an immense amount of discussion, but no satisfactory light has yet been thrown on the conditions that really give rise to the exceptional phenomena. These phenomena are (1) parthenogenesis; (2) oviparous and viviparous reproduction; (3) the production of generations of individuals in which the sexes are very unequally represented, males being frequently entirely absent; (4) the production of individuals differing as to the acquirement of wings, some remaining entirely apterous, while others go on to the winged form; (5) the production of individuals of the same sex with different sexual organs, and distinctions in the very early (but not the earliest) stages of the formation of the individual; (6) differences in the life-habits of successive generations; (7) differences in the habits of individuals of one generation, giving rise to the phenomenon of parallel series. All these phenomena may occur in the case of a single species, though in a very variable extent.

The simple form of Aphid life may be described as follows:—eggs are laid in the autumn, and hatch in the spring, giving rise to females of an imperfect character having no wings; these produce living young parthenogenetically, and this process may be repeated for a few or for many generations, and there may be in these generations a greater or less number of winged individuals, and perhaps a few males.[517] After a time when temperature falls, {583}or when the supply of food is less in quantity, or after a period of deliberate abstention from food, sexual individuals are produced and fertilised eggs are laid which hatch in the spring, and the phenomena are repeated. In other cases these phenomena are added to or rendered more complicated by the intercalated parthenogenetic generations exhibiting well-marked metamorphosis, of kinds such as occur in apterous or in winged Insects; while again the habits of successive generations may differ greatly, the individuals of some generations dwelling in galls, while those of other generations live underground on roots.

PARTHENOGENESIS.—Returning to the various kinds of peculiarities we have enumerated on the preceding page, we may remark that the phenomena of parthenogenesis have been thoroughly established as occurring in Aphidae since Bonnet discovered the fact 150 years ago; and though they have not been investigated in much detail it is known that the parthenogenesis is usually accompanied by the production of young all of the female sex. In other cases males are parthenogenetically produced; but whether these males come from a female that produces only that sex is not yet, so far as the writer knows, established. A note by Lichtenstein[518] suggests that usually only one sex is produced by a parthenogenetic female, but that both sexes are sometimes so produced. There is not at present any species of Aphid known to be perpetuated by an uninterrupted series of parthenogenetic generations. It was formerly supposed that there are no males at all in _Chermes_, but, as we shall subsequently show, this was erroneous. It has, however, been observed that a series of such generations may be continued without interruption for a period of four years, and we have no reason to suppose that even this could not be much exceeded under favourable conditions. The parthenogenetic young may be produced either viviparously or oviparously, according to species.

OVIPAROUS AND VIVIPAROUS REPRODUCTION.—The distinction between these two processes has been extensively discussed, some naturalists maintaining that they are thoroughly distinct _ab initio_. This view, however, cannot be sustained. The best {584}authorities are agreed that in the earliest processes of individualisation the ovum, and the pseudovum[519] giving rise to a viviparous individual, are indistinguishable. Leydig, Huxley, Balbiani, and Lemoine are agreed as to this. Nevertheless, differences in the development occur extremely early. The nature of these differences may be briefly described by saying that in the viviparous forms the embryonic development sets in before the formation of the egg is properly completed. Balbiani says, "In fact at this moment [when the viviparous development is commencing] the germ [pseudovum] is far from having obtained the development it is capable of, and from having accumulated all the matter necessary for the increase of the embryo, so that the evolution of the former coincides, so to speak, with that of the latter. On the other hand, in the true ovum the two processes are chronologically separate, for the rudiment of the new individual never appears before the egg has completed the growth of its constituent parts."[520] As regards the difference in structure of the organs of viviparously and oviparously producing individuals, it is sufficient to remark that they are not of great importance, being apparently confined to certain parts remaining rudimentary in the former. Leydig, indeed, found an _Aphis_ in which certain of the egg-tubes contained eggs in various stages of development, and others embryos in all stages.[521]

As regards the physiology of production of winged and wingless individuals there has been but little exact inquiry. Vast numbers of individuals may be produced without any winged forms occurring, while on the other hand these latter are occasionally so abundant as to float about in swarms that darken the air; the two forms are probably, however, determined by the supply of food. The winged forms are less prolific than the apterous forms; and Forbes has noticed in _Aphis maidi-radicis_, where the generations consist partly of apterous and partly of winged individuals, that when the corn begins to flag in consequence of the attacks of the _Aphis_, then the proportion of {585}winged individuals becomes large.[522] The appearance of winged individuals is frequently accompanied by a peculiar change of habit; the winged individuals migrating to another plant, which in many cases is of a totally different botanical nature from that on which the apterous broods were reared: for instance _Aphis mali_, after producing several apterous generations on apple, gives rise to winged individuals that migrate to the stems of corn or grass, and feeding thereon commence another cycle of generations. The study of this sort of Aphis-migration is chiefly modern, but many very curious facts have already been brought to light; thus _Drepanosiphum platanoides_, after producing a certain number of viviparous generations on maple (_Acer_), quits this food-plant for another, but after two or three months returns again to the maple, and produces sexual young that lay eggs.[523] Histories such as this are rather common. Even more interesting are the cases of those species that, after some weeks of physiological activity on a plant, pass into a state of repose on the same plant, and then after some weeks produce sexual young. On the whole, it would appear that the appearance of winged forms is a concomitant of decreasing nutrition. It is a very remarkable fact that the sexually perfect females are invariably apterous, and this is frequently also the case with the males. It is also highly remarkable that the sexually perfect individuals are of comparatively small size. There are at least three kinds of males in Aphidae—1, winged males; 2, wingless males with mouth well developed; 3, wingless small males with mouth absent. As regards some of these points the conditions usual in Insect life are reversed.[524] Huxley inclined to treat all these products of a fertilised egg, that are antecedent to another process of gamogenesis (_i.e._ production with fertilisation), as one zoological individual: in that case the Aphis zoological individual is winged before attaining the mature state, and is wingless and smaller when mature. Some species may have as a rule two, others three, winged generations in a year.

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The Cambridge natural history, Vol. 06 (of 10)Chapter VIII: Hemiptera, or Bugs—anoplura 532 (23)

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