Lepidoptera

Structure.—The head in the Lepidoptera is sub-globular in shape with the compound eyes exceedingly well developed, and with a pair of ocelli or “simple eyes” often present on the vertex. It is connected to the thorax by a relatively broad and membranous “neck.” The feelers are many-jointed, often they are complex, the segments bearing processes arranged in a comb-like manner and furnished with numerous sensory hairs (fig. 2). The complexity of the feelers is carried to its highest development in certain male moths that have a wonderful power of discovering their females by smell or some analogous sense. Often the feelers are excessively complex in male moths whose maxillae are so reduced that they take no food in the imaginal state. The nature of the jaws has already been briefly described. Functional mandibles of peculiar form (fig. 3, A) are present in the remarkable small moths of the genus Micropteryx (or Eriocephala), and there are vestiges of these jaws in other moths of low type, but the minute structures in the higher Lepidoptera that were formerly described as mandibles are now believed to belong to the labrum, the true mandibles being perhaps represented by rounded prominences, not articulated with the head-capsule. Throughout the order, as a whole, the jaws are adapted for sucking liquid food, and the suctorial proboscis (often erroneously called a “tongue”) is formed as was shown by J. C. Savigny in 1816 by two elongated and flexible outgrowths of the first maxillae, usually regarded as representing the outer lobes or galeae (fig. 4, A, B, g). These structures are grooved along their inner faces and by means of a series of interlocking hair-like bristles can be joined together so as to form a tubular sucker (fig. 4, C). At their extremities they are beset with club-like sense-organs, whose apparent function is that of taste. The proboscis when in use is stretched out in front of the head and inserted into the corolla of a flower or elsewhere, for the absorption of liquid nourishment. When at rest, the proboscis is rolled up into a close spiral beneath the head and between the labial palps (fig. 4, A, p). Only in the genus Micropteryx mentioned above is the lacinia of the maxilla (as A. Walter has shown) developed (fig. 3, B, c). The maxillary palp is usually a mere vestige (fig. 4, B, p) though it is conspicuous in a few families of small moths. A considerable number of Lepidoptera take no food in the imaginal state; in these the maxillae are reduced or altogether atrophied. The second maxillae are intimately fused together to form the labium, which consists only of a reduced mentum, bearing sometimes vestigial lobes and always a pair of palps. These have two or three segments and are clothed with scales. The form and direction of the terminal segment of the labial palp afford valuable characters in classification.

Fig. 4.—Arrangement of the jaws in a typical Moth. Somewhat diagrammatic and in part after E. Burgess and V. L. Kellogg (Amer. Nat. xiv. xxix.).
A, Front view of head. c, Clypeus. e, Compound eye. m, Vestigial mandible. l, Labrum. g, Galeae of 1st maxillae. p, Labial palp. Magnified, B. b, Base of first maxilla dissected out of the head. p, Vestigial palp.	g, Galea. Further magnified. C, Part transverse section showing how the channel (A) of the proboscis is formed by the interlocking of the grooved inner faces of the flexible maxillae. t, Air-tube. n, Nerve. m, Muscle-fibres. Highly magnified.

In the thorax of the Lepidoptera the foremost segment or prothorax is very small, and not movable on the mesothorax. In many families it carries a pair of small erectile plates—the patagia—which have been regarded as serially homologous with the wings. The mesothorax is extensive; its scutum forming most of the dorsal thoracic area and small plates—tegulae—are often present at the base of the forewings, as in Hymenoptera. The tegulae which are beset with long hair-like scales are often conspicuous. The metathorax is smaller than the mesothorax. The legs are of the typical hexapodan form with five-segmented feet; the shins often bear terminal and median spurs articulated at their bases and the entire limbs are clothed with scales.

After A. S. Packard, Mem. Nat. Acad. Sci. vol. vii.

Fig. 5.—Wing-neuration of a Notodont Moth. 2, Sub-costal; 3, radial; 4, median; 5, cubital; 7, 8, anal nervures. a, Discoidal areolet or “cell”; f, frenulum. Note that the forewing has five branches (1-5) of the radial nervure, the hindwing one only. The first anal nervure (No. 6) is absent.

The wings of the Lepidoptera may be said to dominate the structure of the insect; only exceptionally, in certain female moths, are they vestigial or absent (fig. 17). The forewing, with its prominent apex, is longer than the hindwing, and the neuration in both (see figs. 5 and 6) is for the most part longitudinal, only a few transverse nervures, which are, in fact, branches of the median trunk, marking off a discoidal areolet or “cell” (fig. 5, a). The five branches of the radial nervure (figs. 5, 6, 3) (see Hexapoda) are usually present in the forewing, but the hindwing, in most families, has only a single radial nervure; its anal area is, however, often more strongly developed than that of the forewing. The two wings of a side are usually kept together during flight by a few stout bristles—the frenulum—(fig. 5, f) projecting from the base of the costa of the hindwing and fitting beneath a membranous fold or a few thickened scales—the retinaculum—on the under surface of the forewing. In butterflies there is no frenulum, but a costal outgrowth of the hindwing subserves the same function. In the most primitive moths a small lobate outgrowth—the jugum (fig. 6, j.)—from the dorsum of the forewing is present, but it can be of little service in keeping the two wings together. A jugum may be also present on the hindwing. The legs, which are generally used for clinging rather than for walking, have five-segmented feet and are covered with scales. In some families the front pair are reduced and without tarsal segments.

After Packard, Mem. Nat. Acad. Sci. vol. vii.

Fig. 6.—Wing neuration of a Swift Moth (Hepialid). j, Jugum. Nervures numbered as in fig. 5. Note that there are five branches to the radial nervure (No. 3) in both fore- and hindwing, and that the median trunk nervures (No. 4) traverse the discoidal areolet.

Fig. 7.—A, Arrangement of scales in rows on wing of Butterfly. n, Nervure; c, collar-like outgrowths of cuticle. Magnified. B, single scale, and C, an androconium more highly magnified.

Ten abdominal segments are recognizable in many Lepidoptera, but the terminal segments are reduced or modified to form external organs of reproduction. In the male, according to the interpretation of C. Peytoureau, the lateral plates belonging to the ninth segment form paired claspers beset with harpes, or series of ridges or teeth, while the tergum of the tenth segment forms a dorsal hook—the uncus—and its sternum a ventral process or scaphium. In the female the terminal segments form, in some cases, a protrusible ovipositor, but the typical hexapodan ovipositor with its three pairs of processes is undeveloped in the Lepidoptera.

As already mentioned, the characteristic scales on the wings, legs and body of the Lepidoptera are cuticular structures. A complete series of transitional forms can be traced between the most elaborate flattened scales (fig. 7, B) with numerous longitudinal striae and a simple arthropod “hair.” Either a “hair” or a scale owes its origin to a special cell of the ectoderm (hypodermis), a process from which grows through the general cuticle and forms around itself the substance of the cuticular appendage. The scales on the wings are arranged in regular rows (fig. 7, A), and the general cuticle is drawn out into a narrow neck or collar around the base of each scale. The scales can be easily rubbed from the surface of the wing, and the series of collars in which the scales rest are then evident (fig. 7, A, c) on the wing-membrane. On the wings of many male butterflies there are specially modified scales—the androconia (fig. 7, C)—which are formed by glandular cells and diffuse a scented secretion. In some cases, the androconia are mixed among the ordinary scales; in others they are associated into conspicuous “brands” (see fig. 66). The admirable colours of the wings of the Lepidoptera are due partly to pigment in the scales—as in the case of yellows, browns, reds and blacks—partly to “interference” effects from the fine striae on the scales—as with the blues, purples and greens.

A few points of interest in the internal structure of the Lepidoptera deserve mention. The mouth opens into a sub-globular, muscular pharynx which is believed to suck the liquid food through the proboscis, and force it along the slender gullet into a crop-like enlargement or diverticulum of the fore-gut known as a “food-reservoir” or “sucking-stomach.” The true stomach is tubular, and beyond it lies the intestine into which open the three pairs of excretory (Malpighian) tubes. The terminal part of the intestine is of wide diameter, and in some cases gives off a short caecum. The brain and the sub-oesophageal ganglia are closely approximated; there are two or three thoracic and four (rarely five) abdominal ganglia. In the female each ovary has four ovarian tubes, in which the large egg-cells are enclosed in follicles and associated with nutritive cells. There is a special bursa which in the Hepialidae opens with the vagina on the eighth abdominal sternum. In the Micropterygidae, Enocraniidae and the lower Tineides, the duct of the bursa leads into the vagina, which still opens on the eighth sternum. But in most Lepidoptera, the bursa opens by a vestibule on the eighth sternum, distinct from the vagina, whose opening shifts back to the ninth, the duct of the bursa being connected with the vagina by a canal which opens opposite to the spermatheca. In the male, the two testes are usually fused into a single mass, and a pair of tubular accessory glands open into the vasa deferentia or into the ejaculatory duct. In a few families—the Hepialidae and Saturniidae for example—the testes retain the primitive paired arrangement. These details have been worked out by various students, among whom W. H. Jackson and W. Petersen deserve special mention. Summing up the developmental history of the genital ducts, Jackson remarks that there is “an Ephemeridal stage, which ends towards the close of larval life, an Orthopteran stage, indicated during the quiescent period preceding pupation, and a Lepidopteran stage which begins with the commencement of pupal life.”

There are ten abdominal segments—the ninth often small and concealed; prolegs are usually present on the third, fourth, fifth, sixth and tenth of these segments. The head of the caterpillar (fig. 9) is large with firmly chitinized cuticle; it carries usually twelve simple eyes or ocelli, a pair of short feelers (fig. 9 At) and a pair of strong mandibles (fig. 9, Mn), for the caterpillar feeds by biting leaves or other plant-tissues. The first maxillae, so highly developed in the imago, are in the larva small and inconspicuous appendages, each bearing two short jointed processes,—the galea and the palp (fig. 9, Mx). The second maxillae form a plate-like labium on whose surface projects the spinneret which is usually regarded as a modified hypopharynx (fig. 9, Lm). The silk-glands whose ducts open on this spinneret are paired convoluted tubes lying alongside the elongate cylindrical stomach. In the common “silkworm” these glands are five times as long as the body of the caterpillar. They are regarded as modified salivary glands, though the correspondence has been doubted by some students. The body of the caterpillar is usually cylindrical and wormlike, with the segmentation well marked and the cuticle feebly chitinized and flexible. Firm chitinous plates are, however, not seldom present on the prothorax and on the hindmost abdominal segment. The segments are mostly provided with bristle or spine-bearing tubercles, whose arrangement has lately been shown by H. G. Dyar to give partially trustworthy indications of relationship. On either side of the median line we find two dorsal or trapezoidal tubercles (Nos. 1 and 2), while around the spiracle are grouped (Nos. 3, 4 and 5) supra-, post-, and pre-spiracular tubercles; below are the sub-spiraculars, of which there may be two (Nos. 6, 7). The last-named is situated on the base of the abdominal proleg, and yet another tubercle (No. 8) may be present on the inner aspect of the proleg. The spiracles are very conspicuous on the body of a caterpillar, occurring on the prothorax and on the first eight abdominal segments. Various tubercles may become coalesced or aborted (fig. 10, B); often, in conjunction with the spines that they bear, the tubercles serve as a valuable protective armature for the caterpillar. Much discussion has taken place as to whether the abdominal prolegs are or are not developed directly from the embryonic abdominal appendages. In the more lowly families of Lepidoptera, these organs are provided at the extremity with a complete circle of hooklets, but in the more highly organized families, only the inner half of this circle is retained.

B, after Grote, Mitt. aus dem Roemer Museum, No. 6.	Fig. 11.—Pupa of a Butterfly (Amathusia phidippus).
Fig. 10.—Abdominal segments of Caterpillars, to show arrangement of tubercles; the arrows point anteriorly. A, Generalized condition; B, specialized condition in the Saturniidae. s, Spiracle; the numbering of the tubercles is explained in the text. Note that in B No. 2 is much reduced and disappears after the first moult. 4 and 5 are coalesced, and 6 is absent.

The typical Lepidopteran pupa, or “chrysalis,” as shown in the higher families, is an obtect pupa (fig. 11) with no trace of mandibles, the appendages being glued to the body by an exudation, and motion being possible only at three of the abdominal intersegmental regions, the fifth and sixth abdominal segments at most being “free.” A flattened or pointed process—the cremaster—often prominent at the tail-end, may carry one or several hooks (fig. 1, d) which serve to anchor the pupa to its cocoon or to suspend butterfly-pupae from their pad of silk (fig. 11). In the lower families the pupa (fig. 1, c) is only incompletely obtect, and a greater number of abdominal segments can move on one another. The seventh abdominal segment is, in all female lepidopterous pupae, fused with those behind it; in the male “incomplete” pupa this becomes “free” and so may the segments anterior to it, in both sexes, forward to and including the third. The presence of circles of spines on the abdominal segments enables the “incomplete” pupa as a whole to work its way partly out of the cocoon when the time for the emergence of the imago draws near. In the family of the Eriocraniidae (often called the Micropterygidae) the pupa resembles that of a caddis-fly (Trichopteron) being active before the emergence of the imago and provided with strong mandibles by means of which it bites its way out of the cocoon. The importance of the pupa in the phylogeny and classification of the Lepidoptera has lately been demonstrated by T. A. Chapman in a valuable series of papers. Sometimes organs are present in the pupa which are undeveloped in the imago, such as the maxillary palps of the Sesiidae (clearwing moths) and the pectination on the feelers of female Saturniids. E. B. Poulton has drawn attention to the ancestral value of such characters.

“The habit of the species is always to rest on a twig and among dead or dried leaves, and in this position, with the wings closely pressed together, their outline is exactly that of a moderately sized leaf slightly curved or shrivelled. The tail of the hindwings forms a perfect stalk and touches the stick, while the insect is supported by the middle pair of legs, which are not noticed among the twigs and fibres that surround it. The head and antennae are drawn back between the wings so as to be quite concealed, and there is a little notch hollowed out at the very base of the wings, which allows the head to be retracted sufficiently.”

Three families are included in this group having in common certain primitive characters of the wings and neuration (see fig. 6), as well as of the larva and pupa. There is a membranous lobe or jugum near the base of the wing, and the neuration of the hindwing is closely like that of the forewing, the radial nervure being five-branched in both. The pupa has four or five movable segments, and the larval prolegs have complete circles of hooklets.

The three families of the Jugatae are not very closely related to each other. The Micropterygidae (often known as Eriocephalidae), comprising a few small moths with metallic wings, are the most primitive of all Lepidoptera. They are provided with functional mandibles, while the maxillae have distinct laciniae, well-developed palps and galeae not modified for suction (see fig. 3). The larva is remarkable on account of its long feelers, the presence of pairs of jointed prolegs on the first eight abdominal segments, an anal sucker beneath the last segment and bladder-like outgrowths on the cuticle. These curious larvae feed on wet moss. The family has only a few genera scattered widely over the earth’s surface (Europe, America, Australia, New Zealand).

The Eriocraniidae resemble the Micropterygidae in appearance, but the imago has no mandibles, and the maxillae, though short and provided with conspicuous palps, have no laciniae and form a proboscis as in Lepidoptera generally. The abdomen of the female carries a serrate piercing process, and the eggs are laid in the leaves of deciduous trees, the white larvae, with aborted legs, mining in the leaf tissue. The fully-fed larva winters in an underground cocoon and then changes into the most remarkable of all known lepidopterous pupae, with relatively enormous toothed mandibles which bite a way out of the cocoon in preparation for the final change. These pupal mandibles of the Eriocraniidae, together with the nature of the imaginal maxillae in the Micropterygidae (Eriocephalidae) and the wing-neuration in both families, point strongly to a relationship between the Lepidoptera and the Trichoptera.

The Hepialidae or swift moths—the third family of the Jugatae—are in some respects specialized. The moths are of large or moderate size with the maxillae in a vestigial condition, no food being taken after the attainment of the perfect state. The larvae (fig. 12) feed either on roots or in the wood of trees and shrubs, not attaining their growth in less than a year and some large exotic species living for two or three. The family is world-wide in range, and Australia possesses some almost gigantic and strangely coloured genera.

Tineides.

A large assemblage of moths, mostly of small size, are included in this group. The wings have no jugum, but there is a frenulum on the hindwing, which has, as in all the groups above the Jugatae, only a single radial nervure. Three anal nervures are present in the hindwing in those families whose wings are well developed, but in several families of small moths the wings of both pairs are very narrow and pointed, and the neuration is consequently reduced. The sub-costal nervure of the hindwing is usually present and distinct from the radial nervure. The egg is flat except in the Cossidae and Castniidae in which it is upright. The larval prolegs, with few exceptions, have a complete circle of hooklets, and the larvae usually feed in some concealed situation. The pupa is incompletely obtect, with three (in some females only two) to five free abdominal segments, and emerges partly from the cocoon before the moth appears. The cremaster serves to anchor the pupa to its cocoon at the correct degree of emergence, and thus facilitates the eclosion of the imago.

Fig. 18.—Stygia australis. S. Europe.	Fig. 19.—Zeuzera scalaris. India.

The Cossidae are a small family of large moths (figs. 8, 18, 19) belonging to this section, characterized by their heads with erect rough scales or hairs, the pectinate feelers of the males, their reduced maxillae so that no food is taken in the perfect state, and their wings with the fifth radial nervure arising from the third, and the main median nervure forking in the discoidal areolet. The larvae feed in plant stems, often in the wood of trees, forming tunnels and galleries, and usually taking a year or more to reach maturity. The pupa which has three or four free segments in the male and four or five in the female, rests in a cocoon within the food plant, often strengthened by chips of wood, or in a subterranean cocoon. The family is fairly well represented in the tropics; the British fauna possesses only three species, of which the “goat” (Cossus cossus) and the “leopard” (Zeuzera pyrina) are well known, the caterpillars of both being often injurious to timber and fruit trees.

The Tortricidae are a large family of small moths (see fig. 1), nearly allied to the Cossidae. The fifth radial nervure does not arise from the third, the maxillae are well developed, but their palps are obsolete; the head is densely clothed with erect scales; the terminal segment of the labial palp is short and obtuse. The female pupa has three, the male four, free segments. All the larvae of these moths have some method of concealing themselves while feeding. A frequent plan is to roll up a leaf of the food-plant, fastening the twisted portion with silken threads so as to make a tubular retreat; this is the habit of the caterpillar of the green bell moth (Tortrix viridana) which often ravages the foliage of oak plantations. The larvae of the pine-shoot moths (Retinia) shelter in solidified resinous exudations from their coniferous food-plants, while the codlin-moth caterpillar (Carpocapsa pomonella) feeds in apples and pears, growing with the growth of the fruit which affords them both provender and home. The antics of “jumping-beans” are due to the movements of tortricid caterpillars within the substance of the seed.

The Psychidae are a small but widely-distributed family of moths whose males have the head, densely clothed with rough hairs, bearing complex, bipectinated feelers, but with the maxillae reduced and useless. The larvae live in portable cases made of grass, pieces of leaf or stick, with a silken lining, and these cases serve as cocoons for the pupae which agree in structure with those of the Tortricidae. But the most remarkable feature of the family is the extreme degradation of the female, which, wingless, legless and without jaws or feelers, never emerges from the cocoon.

Fig. 20.—Castnia acraeoides. Brazil.	Fig. 21.—Neurosymploca concinna. S. Africa.

The Castniidae are a small family of large, conspicuous, day-flying exotic moths (fig. 20) whose clubbed feelers and bright colours give them a resemblance to butterflies, although their wing-neuration is of the primitive tineoid type; the smooth larvae feed on the stems or roots of plants and the pupal structure agrees with that of the Tortricidae and Psychidae. The distribution of the family is confined to Tropical America and the Indo-Malayan and Australian regions.

Fig. 22.—A, Sesia asiliformis (Gad-fly Hawk Moth). Europe. B, Larva.

The Zygaenidae (burnet moths) are a large family of day-flying moths (fig. 21) adorned with brilliant metallic colours. The feelers are long, stout in the middle and tapering, bearing numerous long or short pectinations. The well-developed maxillae have vestigial palps. The larvae—often very conspicuously coloured—are remarkable among the Tineides in having incomplete circles of hooks on the prolegs, and they feed exposed on the leaves of various plants. The pupa, enclosed in a silken cocoon, has four or five free segments. The Limacodidae are a small family of brownish nocturnal moths, allied to the Zygaenidae and agreeing with them in the structure of the pupa. The larva in this family also is an exposed feeder, but it is remarkable in form, being flattened and slug-like, without prolegs and adorned with curious spinous processes.

The Sesiidae are a large family of small, narrow-winged moths, the sub-costal nervure of the hindwing being absent and the wings being for the most part destitute of scales (fig. 22). The maxillae are developed but their palps are vestigial, while the terminal segment of the labial palp is short and pointed. Many of these insects have their bodies banded with black and yellow; this in conjunction with the transparent wings makes some of them like wasps or hornets in appearance. The larvae feed in the woody stems of various plants. The pupa, with three or four free abdominal segments, remains within its cocoon, formed with chips of wood, until the time for its final change draws near; then it works itself partly out of the tree by means of the spines on its abdominal segments.

The Nepticulidae are the smallest of all the Lepidoptera, measuring only 3-8 mm. across the outspread wings, which are all lanceolate and pointed at the tip. The sucking portions of the maxillae are vestigial, but the palps are long and jointed. The larvae, without thoracic limbs or prolegs, but sometimes with paired rudimentary processes on some of the segments, mine in the leaves of plants. The pupa, with four free abdominal segments in the female and five in the male, rests in a cocoon usually outside the mine.

Fig. 23.—Adela degeerella. Europe.	Fig. 24.—Euplocampus anthracinus. Europe.	Fig. 25.—Tinea tapetzella (Clothes Moth). Europe.

The Adelidae are a family of delicate, but larger, moths with very long feelers (fig. 23) especially in the males. The larvae feed, when young, in flowers; later, protected by a flat case, they devour leaves; the pupa resembles that of the Nepticulidae in structure. The female has an ovipositor adapted for piercing plant tissues.

The Tineidae are a large and important family of small moths (figs. 14, 24, 25) with rough-haired heads, and with the maxillae and their palps usually well developed. Many of the genera have narrow pointed wings with degraded neuration. The larvae differ in their habits, some—Gracilaria for example—mine in leaves, while others, like the well-known caterpillars of the clothes moth (Tinea) surround themselves with portable cases (fig. 14) formed by spinning together their own excrement. The female pupa has three, the male four free abdominal segments.

Plutellides.

This group includes a few large families of small moths that are linked by their imaginal and larval structure to the Tineidae (in which they have often been included) and by their pupal structure to the higher groups that have yet to be considered. The moths have labial palps with slender pointed terminal segments, and narrow pointed wings, but the neuration (except in the Elachistidae) is less degenerate than in most Tineidae. The hairy covering of the head is smooth, and the maxillary palps are usually vestigial. The egg is flat, and the larval prolegs have complete circles of hooklets. The pupa is obtect with only two free abdominal segments (fifth and sixth) in both sexes and does not move out of the cocoon.

Fig. 26.—Cerostoma asperella. Europe.	Fig. 27.—Psecadia pusiella.

Four families are included in this group. The Plutellidae (fig. 26) have the maxillary palps developed, in some genera, as slender threadlike appendages directed straight forward. The larvae do not usually mine in leaves, but feed openly, keeping to the underside for protection (Plutella), or spinning by their united labour a mass of web over the food-plant (Hyponomeuta). In the other three families the maxillary palps are vestigial or obsolete. The Elachistidae have remarkably narrow, pointed wings and their larvae mine in leaves or form portable cases and feed among seeds. In the Oecophoridae (fig. 27) the sub-costal nervure of the hindwing is free and distinct throughout its length, and the larvae usually feed among spun leaves or seeds, or in decayed wood. The Gelechiidae are a large family with similar larval habits; the moths are distinguished by the sinuate termen of the hindwing and the connexion of its sub-costal nervure with the discoidal areolet.

Pyralides.

Fig. 28.—Pterophorus spilodactylus. Europe.	Fig. 29.—Orneodes hexadactylus (24-plumed Moth). Europe.

This group includes a number of moths of delicate build with elongate legs, the maxillae and their palps being usually well developed. The forewings have two anal nervures, the hindwings three (fig. 30, h, i); in the hindwing the sub-costal nervure bends towards and often connects with the radial, and the frenulum is usually present. The egg is flat. The larva has complete circles of hooklets on its five pairs of prolegs, and the pupa (usually completely obtect) does not move at all from its cocoon. This group includes the only Lepidoptera that have aquatic larvae.

Of the families comprised in this division three deserve special mention. The Pterophoridae (plume moths, fig. 28) usually have the wings deeply cleft—a single cleft in the forewing and two in the hindwing. The hairy larvae feed openly on leaves, while the soft and hairy pupa remains attached to its cocoon by the cremaster, although it is incompletely obtect and has three or four free abdominal segments. The Orneodidae (multiplume moths) have all the wings six-cleft. Our British species, Orneodes hexadactyla (fig. 29), is an exquisite little insect, whose larva feeds on the blossoms of honeysuckle. The pupa is completely obtect, with only two free abdominal segments. The Pyralidae (figs. 13, 30), a large family with numerous divisions, have entire wings, and their pupae are obtect. The caterpillars feed in some kind of shelter, some spinning a loose case among the leaves of their food-plant, others burrowing into dry vegetable substances or eating the waxen cells of bees. Several species of this group, such as the Mediterranean flour moth, Ephestia kühniella (fig. 30), become serious pests in storehouses and granaries, their larvae devouring flour and similar food-stuffs.

After Riley and Howard, Insect Life, vol. 2 (U.S. Dept. Agr.).
Fig. 30.—Flour Moth (Ephestia kühniella).
c, With wings spread. f, At rest. g, h, i, Marking and neuration of wings. a, Larva.	b, Pupa. d, Head and front body-segments of larva. e, 2nd and 3rd abdominal segments.

Noctuides.

In this group may be included a number of families of moths with the second median nervure of the forewing arising close to the third. This feature of neuration characterizes also the Jugatae (see fig. 6), Tineides, Plutellides and Pyralides. But the Noctuides differ from these groups in having only two anal nervures in the hindwing. The maxillary palps are absent or vestigial, and a frenulum is usually present on the hindwing. The larva has usually ten prolegs, whose hooklets are arranged only along the inner edge, while the immobile pupa is always obtect with only two free abdominal segments (the fifth and sixth). The Lasiocampidae and their allies have flat eggs, but in the Noctuidae, Arctiidae and their allies the egg is upright.

Fig. 31.—Claterna cydonia. India.

The Lasiocampidae, together with a few small families, differ from the majority of this group in wanting a frenulum. The maxillae of the Lasiocampidae are so reduced that no food is taken in the imaginal state, and in correlation with this condition the feelers of the male are strongly (those of the female more feebly) bipectinated. The moths are stout, hairy insects, usually brown or yellow in the pattern of their wings. The caterpillars are densely hairy and many species hibernate in the larval stage. The pupa is enclosed in a hard, dense cocoon, whence the name “eggars” is often applied to the family, which has a wide distribution, but is absent from New Zealand. The Drepanulidae are an allied family, in which the frenulum is usually present, while the hindmost pair of larval prolegs are absent, their segment being prolonged into a pointed process which is raised up when the caterpillar is at rest. The hook-tip moths represent this family in the British fauna.

The Lymantriidae resemble the Lasiocampidae in their hairy bodies ana vestigial maxillae, but the frenulum is usually present on the hindwing and the feelers are bipectinate only in the males. Some females of this family—the vapourer moths (Orgyia and allies, fig. 17), for example—are degenerate creatures with vestigial wings. The larvae (fig. 15) are very hairy, and often carry dense tufts on some of their segments; hence the name of “tussocks” frequently applied to them. The pupae are also often hairy (fig. 16)—an exceptional condition—and are protected by a cocoon of silk mixed with some of the larval hairs, while the female sheds some hairs from her own abdomen to cover the eggs. The family is widely distributed, its headquarters being the eastern tropics. To that part of the world is restricted the allied family of the Hypsidae, distinguished from the “tussocks” by the slender upturned terminal segment of the labial palps and by the development of the maxillae.

Fig. 32.—Ophideres imperator. Madagascar.

Fig. 33.—Cyligramma fluctuosa. W. Africa.

From Mally, Bull. 24, Div. Ent. U.S. Dept. Agr.

Fig. 34.—e, f, Heliothis armigera. Europe, c, Larva; d, pupa in cell. Natural size. a, b, Egg, highly magnified.

The Noctuidae are the largest and most dominant family of the Lepidoptera, comprising some 10,000 known species. They are mostly moths of dull coloration, flying at dusk or by night. The maxillae are well developed, the hindwing has a frenulum, and its sub-costal nervure touches the radial near the base. The larvae of the Noctuidae (fig. 34, c) are rarely hairy and the pupa (fig. 34, d) usually rests in an earthen cell, being often the wintering stage for the species; sometimes the pupa is enclosed in a loose cocoon of silk and leaves. In some Noctuidae (fig. 32) the hindwings are brightly coloured, but these are concealed beneath the dull, inconspicuous forewings when the insect rests (fig. 34, f). Nearly allied to the Noctuidae, but very different in appearance, are the gaily-coloured Agaristidae, a family of day-flying moths (figs. 35, 36), confined to the warmer regions of the globe and distinguished by their thickened feelers, those of the Noctuids being thread-like or slightly pectinate.

The Arctiidae (tiger moths, footmen, &c.) are allied to the Noctuidae, but their wing-neuration is more specialized, the sub-costal nervure of the hindwing being confluent with the radial for the basal part of its course. These moths (fig. 37) have gaily coloured wings, and the caterpillars are often densely covered with long smooth hairs. The pupae are enclosed in silken cocoons (fig. 38). The highest specialization of structure in this group of the Lepidoptera is reached by the Syntomidae, a family nearly allied to the Arctiidae, but with the sub-costal nervure in the hindwing absent. The Syntomidae have elongate narrow forewings and short hindwings, usually dark in colour with clear spots and dashes destitute of scales (fig. 40). The body, on the other hand, is often brilliantly adorned. The family, abundant in the tropics of the Old World, has only two European species.

Fig. 35.—Rothia pales. Madagascar.

Sphingides.

Fig. 36.—Aegocera rectilinea. Tropical Africa.	Fig. 37.—Haploa Lecontei. N. America.

After Lugger, Riley and Howard, Insect Life, vol. 2 (U.S. Dept. Agr.).

Fig. 38.—c, Tiger Moth (Phragmatobia fuliginosa, Linn.). Europe. a, Caterpillar; b, cocoon with pupa. Slightly enlarged.

This group includes a series of families which agree with the Noctuides in most points, but are distinguished by the origin of the second median nervure of the forewing close to the first, or from the discocellular nervure midway between the first and third medians (see fig. 5). These neurational characters may appear somewhat insignificant, but such slight though constant distinctions in structures of no adaptational value may be safely regarded as truly significant of relationship. Several of the families in this group have lost the frenulum. In larval and pupal characters the Sphingides generally resemble the Noctuides, but in some families there is a reduction in the number of the larval prolegs. The egg is spherical or flat, upright only in the Notodontidae.

Fig. 39.—Halias prasinana. Europe.	Fig. 40.—Euchromia formosa. S. Africa.

The Notodontidae are stout, hairy moths (figs. 5, 41, 42 a) with maxillae and frenulum developed. In the larva the prolegs on the hindmost segment are sometimes modified into pointed outgrowths which are carried erect when the caterpillar moves about. From these structures whip-like, coloured processes are protruded by the caterpillar (fig. 42 b) of the puss moth (Cerura) when alarmed; these processes are believed to help in “terrifying” the caterpillar’s enemies. Allied to the Notodontidae are the Cymatophoridae—a family of moths agreeing with the Noctuidae in appearance and habits—and the large and important family of the Geometridae. The moths (fig. 43) of this family are distinguished from the Notodontidae by their delicate build and elongate feet, the caterpillars (fig. 43, c) by the absence or vestigial condition of the three anterior pairs of prolegs. The two hinder pairs of prolegs are therefore alone functional and the larva progresses by “looping,” i.e. bending the body so as to bring these prolegs close up to the thoracic legs, and then, taking a fresh grip on the twig whereon it walks, stretching the body straight out again. Many of these larvae have a striking resemblance both in form and colour to the twigs of their food-plant. In some of the species the female has the wings reduced to useless vestiges. The family is world-wide in its range. The tropical Uraniidae are large handsome moths (figs. 44, 45), often with exquisite wing-patterns, allied to the Geometridae, but distinguished by the absence of a frenulum in the moth and the presence of the normal ten prolegs in the larva.

Fig. 41.—Notodonta ziczac (Pebble Prominent Moth). Europe.
Fig. 42a.—Cerura borealis. N. America.	Fig. 42b.—Larva of Cerura (Puss Moth).

After Grote, Natural Science (J. M. Dent & Co.).

Fig. 43.—Geometrid Moth (Amphidasys betularia, Linn.). Europe. a, Large grey type; b, dark variety; c, caterpillar in looping attitude.

Fig. 44.—Urania boisduvalii. Cuba.

Fig. 45.—Urania boisduvalii at rest, showing under surface of wings.

The Sphingidae (hawk moths) are insects often of large size (figs. 46a, 47), with spindle-shaped feelers, elongate and powerful forewings and the maxillae very well developed. The hindwing carries a frenulum and has its sub-costal nervure connected with the radial by a short bar. The caterpillars have the full number of prolegs, and, in many genera, carry a prominent dorsal horn on the eighth abdominal segment (fig. 46b). The pupa lies in an earthen cell. On account of their powerful flight the moths of this family have a wide range; certain species—like Acherontia atropos and Protoparce convolvuli—migrate into the British Islands in numbers almost every summer.

Fig. 46a.—Chlaenogramma jasminearum (Jessamine Sphinx). N. America.

Fig. 46b.—Larva.

Fig. 47.—Smerinthus ocellatus (Eyed Hawk moth). Europe.

A group of families in which the first maxillae are vestigial, the feelers bipectinate and the pupa enclosed in a dense silken cocoon, have been regarded as the most highly specialized of all the moths, though according to other views the whole series of the Lepidoptera culminates in the Syntomidae. Of these cocoon-spinning families may be specially mentioned the Eupterotidae, large brown or yellow moths inhabiting tropical Asia and Africa, and represented in Europe only by the “processionary moth” (Cnethocampa processionea). In this family the frenulum is present, and the larvae are protected with tufts of long hair. The Bombycidae have no frenulum, and the larvae are smooth, with some of the segments humped and the eighth abdominal often carrying a dorsal spine. The family is tropical in its distribution, but the common silkworm (Bombyx mori, fig. 48) has become acclimatized in southern Europe and is the source of most of the silk used in manufacture and art. Of commercial value also is the silk spun by the great moths of the family Saturniidae, well represented in warm countries and contributing a single species (Saturnia pavonia-minor) to the British fauna. These moths (fig. 49) have but a single anal nervure in the hindwing and only three radial nervures in the forewing. The wing-patterns are handsome and striking; usually an unsealed “eyespot” is conspicuous at the end of each discoidal areolet. The caterpillars are protected by remarkable spine-bearing tubercles (fig. 10, B).

After C.V. Riley, Bull. 14, Div. Ent. U.S. Dept. Agr.

Fig. 48.—Bombyx mori. China. a, Caterpillar (the common silkworm); b, cocoon; c, male moth.

Fig. 49.—Epiphora bouhiniae. W. Africa.

Grypocera.

Fig. 50.—Tagiades sabadius. S. Africa.

This group stands at the base of the series of families that are usually distinguished as “butterflies.” The feelers are recurved at the tip, and thickened just before the extremity. The forewing has the full number of radial nervures, distinct and evenly spaced, and two anal nervures; the frenulum is usually absent. The larvae (fig. 51) have prolegs with complete circles of hooklets, and often feed in concealed situations, while the pupa is protected by a light cocoon. The affinities of this group are clearly not with the higher groups of moths just described, but with some of the lower families. According to Meyrick they are most closely related to the Pyralidae, but Hampson and most other students would derive them (through the Castniidae) from a primitive Tineoid stock allied to the Cossidae and Zygaenidae.

Three families are included in the section. The North American Megathymidae and the Australian Euschemonidae have a frenulum and are usually reckoned among the “moths.” The Hesperiidae in which the frenulum is wanting form the large family of the skipper butterflies, represented in our own fauna by several species. They are insects with broad head—the feelers being widely separated—usually brown or grey wings (fig. 50) and a peculiar jerky flight. The family has an extensive range but is unknown in Greenland, New Zealand, and in many oceanic islands.

Rhopalocera.

Fig. 51.—Chrysalis and Larva of Nisoniadestages (dingy skipper). Europe.	Fig. 52.—Chrysophanus thoe. N. America.

This group comprises the typical butterflies which are much more highly specialized than the Grypocera, and may be readily distinguished by the knobbed or clubbed feelers and by the absence of a frenulum. Two or more of the radial nervures in the forewing arise from a common stalk or are suppressed. The egg is “upright.” The larvae have hooklets only on the inner edges of the prolegs. The pupa is very highly modified, only two free abdominal segments are ever recognizable, and in some genera even these have become consolidated. The cocoon is reduced to a pad of silk, to which the pupa is attached, suspended by the cremastral hooks; in some families there is also a silken girdle around the waist-region. In correlation with the exposed condition of the pupa, we find the presence of a specially developed “head-piece” or “nose-horn” to protect the head-region of the contained imago. Their bright colours and conspicuous flight in the sunshine has made the Rhopalocera the most admired of all insects by the casual observer.

A modification that has taken place in several families of butterflies is the reduction of the first pair of legs. H. W. Bates arranged the families in a series depending on this character, but neurational and pupal features must be taken into account as well, and the sequence followed here is modified from that proposed by A. R. Grote and J. W. Tutt.

Fig. 53.—Rathinda amor. India.	Fig. 54.—Cheritra freja. India.

The Lycaenidae are a large family including the small butterflies (figs. 52, 53, 54) popularly known as blues, coppers and hairstreaks. The forelegs in the female are normal, but in the male the tarsal segments are shortened and the claws sometimes are absent. The forewing has only three or four radial nervures (fig. 55), the last two of which arise from a common stalk; the feelers are inserted close together on the head. The larva is short and hairy, somewhat like a woodlouse in shape, the broad sides concealing the legs and prolegs, while the pupa, which is also hairy or bristly, is attached by the cremaster to a silken pad and cinctured with a silken thread. The upper surfaces of the wings of these insects are usually of a bright metallic hue—blue or coppery—while beneath there are often numerous dark centred “eye-spots.” The family is widely distributed. Nearly related are the Lemoniidae, a family abundantly represented in the Neotropical Region, but scarce in the Old World and having only a single European species (Nemeobius lucinia) which occurs also in England. In the Lemoniidae (figs. 56, 57) the forelegs of the male are reduced and useless for walking. The Libytheidae may be recognized by the elongate snout-like palps, the five-branched radial nervure of the forewing, the cylindrical hairy larva, and the pupa attached only by the cremaster.

After Grote, Natural Science, vol. 12 (J. M. Dent & Co.).

Fig. 55.—Neuration of Wings in Lycaena.

2, Sub-costal. 3, Radial. 4, Median. 5, Cubital. 7, 8, Anal nervures.

Fig. 56.—Eurybia carolina. Brazil.	Fig. 57.—Calephelis caenius. N. America.

Fig. 58.—Papilio machaon (Swallow-tail.). Europe.

Fig. 59.—Parnassius apollo (Apollo). European Alps.

The Papilionidae are large butterflies with ample wings, and all six legs fully developed in both sexes. The forewing has five radial and two anal nervures, the second of the latter being free from the first and running to the dorsum of the wing, while the hindwing has but a single anal, and is frequently prolonged into a “tail” at the third median nervure (fig. 58). The larva is cylindrical, never hairy but often tuberculate and provided with a dorsal retractile tentacle (osmaterium) on the prothorax. The pupa, which has a double “nose-horn,” is attached by the cremaster and a waist-girdle to the food-plant in the Papilioninae (fig. 58), but lies in a web on the ground among the Parnasiinae (figs. 59, 60). The latter sub-family includes the well-known Apollo butterflies of the Alps. The former is represented in the British fauna by the East Anglian swallow-tail (Papilio machaon), and is very abundant in the warmer regions of the world, including some of the most magnificent and brilliant of insects.

Fig. 60.—Thais medesicaste. S. France.	Fig. 61.—Colias hyale (Pale clouded Yellow Butterfly). Europe.

Agreeing with the Papilionidae in the six perfect legs of both sexes and the cincture-support of the pupa we find the Pieridae—the family of the white and yellow butterflies (figs. 61, 62)—represented by ten species in the British fauna and very widely spread over the earth’s surface. In the Pieridae there are two anal nervures in the hindwing, while the second anal nervure in the forewing runs into the first; the larva is cylindrical and hairy without an osmaterium. The pupa has a single “nose-horn,” and in the more highly organized genera there is no mobility whatever between its abdominal segments. The wintering pupae of the common cabbage butterflies (Pieris brassicae and P. rapae) are common objects attached to walls and fences and their colour harmonizes, to a great extent, with that of their surroundings.

Fig. 62.—Appias nero (male). Malaya.

Fig. 63.—Dione moneta. Brazil.	Fig. 64.—Larva of Argynnis paphia (Silver-washed Fritillary). Europe.

Fig. 65.—Vanessa io (Peacock) and its pupa.

Fig. 66.—Euploea leucostictos (male). Malaya.

After A. R. Grote, Natural Science, vol. 12 (J. M. Dent & Co.).

Fig. 67.—Neuration of Wings in a Nymphaline Butterfly.

2, Sub-costal. 3, Radial. 4, Median. 5, Cubital. 6, 7, 8, Anal nervures.

Fig. 68.—Nymphalis jason. W. Africa. Upper and under surface.

Fig. 69.—Larva and Pupa of Apatura ilia.	Fig. 70.—Callithea sapphira. Brazil.

The Nymphalidae are by far the largest and most dominant family of butterflies. In both sexes the forelegs are useless for walking (fig. 63), the tarsal segments being absent and the short shins clothed with long hairs, whence the name of brush-footed butterflies is often applied to the family. The neuration of the wings resembles that found among the Pieridae, but in the Nymphalidae the pupa, which has a double nose-horn (fig. 65)—as in Papilio—is suspended from the cremaster only, no girdling thread being present, or it lies simply on the ground. The egg is elongate and sub-conical in form and ornamented with numerous ribs, while the larva is usually protected by numerous spines (fig. 64) arising from the segmental tubercles. To this family belong our common gaily-coloured butterflies—the tortoiseshells, peacock (fig. 65), admirals, fritillaries and emperors. In most cases the bright colouring is confined to the upper surface of the wings, the under-side being mottled and often inconspicuous. Most members of the group Vanessidi—the peacock and tortoiseshells (Vanessa) and the red admiral (Pyrameis) for example—hibernate in the imaginal state. This large family is divided into several sub-families whose characters may be briefly given, as they are considered to be distinct families by many entomologists. The Danainae (or Euploeinae, fig. 66) have the anal nervures of the forewing arising from a common stalk, the discoidal areolets in both wings closed, and the front feet of the female thickened; their larvae are smooth with fleshy processes. The danaine butterflies range over all the warmer parts of the world, becoming most numerous in the eastern tropics, where flourish the handsome purple Euploeae whose males often have “brands” on the wings; these insects are conspicuously marked and are believed to be distasteful to birds and lizards. So are the South American Ithomiinae, distinguished from the Danainae by the slender feet of the females; the narrow winged, tawny Acraeinae, with simple anal nervures, thick hairy palps and spiny larvae; and the Heliconiinae whose palps are compressed, scaly at the sides and hairy in front. This last named sub-family is confined to the Neotropical Region, while the Acraeinae are most numerous in the Ethiopian. The Nymphalinae include the British vanessids (fig. 65), and a vast assemblage of exotic genera (figs. 68, 70), characterized by the “open” discoidal areolets (fig. 67) owing to the absence of the transverse “disco-cellular” nervules. In the Morphinae—including some magnificent South American insects with deep or azure blue wings, and a few rather dull-coloured Oriental genera—the areolets are closed in the forewings and often in the hindwings. The larvae of the Morphinae (fig. 71) are smooth or hairy with a curiously forked tail-segment. A similar larva characterizes the South American Brassolinae or owl-butterflies—robust insects (figs. 72, 73) with the areolets closed in both wings, which are adorned with large “eye-spots” beneath. The Satyrinae, including our native browns and the Alpine Erebiae, resemble the foregoing group in many respects of structure, but the sub-costal nervure is greatly thickened at the base (fig. 74). This sub-family is world-wide in its distribution. One genus (Oeneis, fig. 75) is found in high northern latitudes, but reappears in South America. The dark, spotted species of Erebia are familiar insects to travellers among the Alps; yet butterflies nearly related to these Alpine insects occur in Patagonia, in South Africa and in New Zealand. Such facts of distribution clearly show that though the Nymphalidae have attained a high degree of specialization among the Lepidoptera, some of their genera have a history which goes back to a time when the distribution of land and water on the earth’s surface must have been very different from what it is to-day.

Fig. 71.—Larva of Amathusia phidippus.

Fig. 72.—Opsiphanes syme. Brazil.

Fig. 73.—Brassolis astyra. Brazil.

After A. R. Grote, Natural Science, vol. 12 (J. M. Dent & Co.).

Fig. 74.—Neuration of wings in Pararge, a satyrid butterfly.

2, Sub-costal. 3, Radial. 4, Median. 5, Cubital. 7, 8, Anal nervures.

Fig. 75.—Oeneis jutta. Arctic Regions.	Fig. 76.—Bia actorion. Brazil.

Bibliography.—The handsome Lepidoptera, with their interesting and easily observed life-histories, have naturally attracted many students, and the literature of the order is enormous. M. Malpighi’s treatise on the anatomy of the silkworm (De Bombycibus, London, 1669) and P. Lyonnet’s memoir on the Goat-caterpillar, are among the earliest and most famous of entomological writings. W. F. Kirby’s Handbook to the Order Lepidoptera (5 vols., London, 1894-1897) should be consulted for references to the older systematic writers such as Linnaeus, J. C. Fabricius, J. Hübner, P. Cramer, E. Doubleday and W. C. Hewitson. Kirby’s Catalogues are also invaluable for the systematist. For the jaws of the Lepidoptera see F. Darwin, Quart. Journ. Mic. Sci. xv. (1875); E. Burgess, Amer. Nat. xiv. (1880); A. Walter, Jen. Zeits. f. Naturw. xviii. (1885); W. Breitenbach, Ib. xv. (1882); V. L. Kellogg, Amer. Nat. xxix. (1895). The last-named deals also with wing structure, which is further described by A. Spuler, Zeits. wiss. Zool. liii. (1892) and Zool. Jahrb. Anat. viii. (1895); A. R. Grote, Mitt. aus dem Roemer-Museum (Hildesheim, 1896-1897); G. Enderlein, Zool. Jahrb. Anat. xvi. (1903), and many others. For scales see A. G. Mayer, Bull. Mus. Comp. Zool. Harvard, xxix. (1896). For internal anatomy W. H. Jackson, Trans. Linn. Soc. Zool. (2) v. (1891), and W. Petersen, Mem. Acad. Imp. Sci. St Petersburg (8) ix. (1900). The early stages and transformations of Lepidoptera are described by J. Gonin, Bull. Soc. Vaud. Sci. Nat. xxx. (1894); E. B. Poulton, Trans. Linn. Soc. Zool. (2) v. (1891); H. G. Dyar, Ann. New York Acad. Sci. viii. (1894); T. A. Chapman, Trans. Entom. Soc. Lond. (1893), &c. For habits and life-relations see A. Seitz, Zool. Jahrb. Syst. v., vii. (1890, 1894); A. Weismann, Studies in the Theory of Descent (London, 1882) and Entomologist, xxix. (1896); F. Merrifield, Trans. Entom. Soc. Lond. (1890, 1893, 1905); M. Standfuss, Handbuch der paläarktischen Gross-schmetterlinge (Jena, 1896); R. Trimen, Proc. Ent. Soc. Lond. (1898); E. B. Poulton, Colours of Animals (London, 1890); Trans. Entom. Soc. (1892 and 1903), and Journ. Linn. Soc. Zool. xxvi. (1898); F. E. Beddard, Animal Coloration (London, 1892). For distribution see H. J. Elwes, Proc. Entom. Soc. Lond. (1894); J. W. Tutt, Migration and Dispersal of Insects (London, 1902); Fossil Lepidoptera, S. H. Scudder, 8th Rep. U.S. Geol. Survey (1889). Among recent general works on the Lepidoptera, most of which contain numerous references to the older literature, may be mentioned A. S. Packard’s unfinished work on the Bombycine Moths of N. America, Mem. Nat. Acad. Sci. Philadelphia, vii. (1895), and Mem. Acad. Sci. Washington, lx. (1905); D. Sharp’s chapter in Cambridge Nat. Hist. vi. (London, 1898); G. F. Hampson, Moths of India (4 vols., London, 1892-1896), and Catalogue of the Lepidoptera Phalaenae (1895) and onwards; S. H. Scudder, Butterflies of New England (3 vols., Cambridge, Mass., 1888-1889); W. J. Holland, Butterfly Book (New York, 1899). Works on the British Lepidoptera are numerous, for example, those of H. T. Stainton (1851), C. G. Barrett (1893-1907), E. Meyrick (1895), and J. W. Tutt (1899 and onwards). For recent general systematic works, the student should consult the catalogues mentioned above and the Zoological Record. The writings of O. Staudinger, E. Schatz, C. Oberthür, K. Jordan, C. Aurivillius and P. Mabille may be specially mentioned.