haunts of the species, where there is no midsummer night, the male closely resembles the female in wing patterns, the development of the conspicuous white being needless. A very interesting sexual dimorphism is seen in the wingless condition of several female moths—the winter moths (Hybernia and Cheimatobia) among the Geometridae and the vapourers (Orgyia and Ocneria) among the Lymantriidae for example (fig. 17). It might be thought that the loss of power of flight by the female would seriously restrict the range of the species. In such insects, however, the caterpillars are often active and travel far.
Distribution and Migration.—The range of the Lepidoptera is practically world-wide; they are absent from the most remote and inhospitable of the arctic and antarctic lands, but even Kerguelen possesses a few small indigenous moths. Many of the large and dominant families have a range wide as that of the order, and certain species that have attached themselves to man—like the meal moths and the clothes moths—have become almost cosmopolitan. Interesting and suggestive restrictions of range can, however, be often traced. Although butterflies have been found in 82° N. latitude in Greenland, they are unknown in Iceland, and only a few species of the group reach New Zealand. Three large sections—the Ithomiinae, Heliconiinae and Brassolinae—of the great butterfly family Nymphalidae are peculiar to the Neotropical region, while the Morphinae, a characteristically South American group, have a few Oriental genera in India and Indo-Malaya. The Acraeinae, another section of the same family, have the vast majority of their species in Ethiopian Africa, but are represented eastwards in the Oriental and Australian regions and westwards in South America. A comparison of the lepidopterous faunas of Ireland, Great Britain and the European continent is very instructive, and suggests strongly that, despite their power of flight the Lepidoptera are mostly dependent on land-connexions for the extension of their range. For example, Ireland has only forty of the seventy species of British butterflies. The range of many Lepidoptera is of course determined by the distribution of the plants on which their larvae feed.
Nevertheless certain species of powerful flight, and some that might be thought feeble on the wing, often cross sea-channels and establish or reinforce distant colonies. Caterpillars of the great death’s head moth (Acherontia atropos) are found every summer feeding in British and Irish potato fields, but it is doubtful if any of the pupae resulting from them survive the winter in our climate. It is believed by Tutt that the species is only maintained by a fresh immigration of moths from the South each summer. Hosts of white butterflies (Pieris) have been frequently observed crossing the English Channel from France to Kent. Migrating swarms of Lepidoptera have often been met by sailors in mid-ocean; thus, Tutt records the presence around a sailing ship in the Atlantic of such a swarm of the rather feeble moth Deiopeia pulchella, nearly 1000 m. from its nearest known habitat. This migratory instinct is connected with the gregarious habits of many Lepidoptera. For example, H. W. Bates states that at one place in South America he noticed eighty different species flying about in enormous numbers in the sunshine, and these, with few exceptions, were males, the females remaining within the forest shades. Darwin describes a “butterfly shower,” which he observed 10 m. off the South American coast, extending as far as the eye could reach; “even by the aid of the telescope,” he adds, “it was not possible to see a space free from butterflies.” Sir J. Emerson Tennent, witnessed in Ceylon a mighty host of butterflies of white or pale yellow hue, “apparently miles in breadth and of such prodigious extension as to occupy hours and even days uninterruptedly in their passage.” Observations at Heligoland by H. Gätke have shown that migrating moths “travel under the same conditions as migrating birds, and for the most part in their company, in an east to west direction; they fly in swarms, the numbers of which defy all attempts at computation and can only be expressed by millions.” The painted lady butterfly (Pyrameis cardui) comes in repeated swarms from the Mediterranean region into northern and western Europe, while in North America companies of the monarch (Anosia archippus) invade Canada every summer from the United States, and are believed to return southwards in autumn. This latter species has, during the last half-century, extended its range south-westwards across the Pacific and reached the Austro-Malayan islands, while several specimens have occurred in southern and western England, though it has not established itself on this side of the Atlantic. It is noteworthy that the introduction of its food-plant—Asclepias—into the Sandwich Islands in 1850 apparently enabled it to spread across the Pacific.
Fossil History.—Our knowledge of the geological history of the Lepidoptera is but scanty. Certain Oolitic fossil insects from the lithographic stone of Solenhofen, Bavaria, have been described as moths, but it is only in Tertiary deposits that undoubted Lepidoptera occur, and these, all referable to existing families, are very scarce. Most of them come from the Oligocene beds of Florissant, Colorado, and have been described by S. H. Scudder. The paucity of Lepidoptera among the fossils is not surprising when we consider the delicacy of their structure, and though their past history cannot be traced back beyond early Cainozoic times, we can have little doubt from the geographical distribution of some of the families that the order originated with the other higher Endopterygota in the Mesozoic epoch.
Classification.—The order Lepidoptera contains more than fifty families, the discussion of whose mutual relationships has given rise to much difference of opinion. The generally received distinction is between butterflies or Rhopalocera (Lepidoptera with clubbed feelers, whose habit is to fly by day) and moths or Heterocera (Lepidoptera with variously shaped feelers, mostly crepuscular or nocturnal in habit). This distinction is quite untenable as a zoological conception, for the relationship of butterflies to some moths is closer than that of many families of Heterocera to each other. Still more objectionable is the division of the order into Macrolepidoptera (including the butterflies and large moths) and the Microlepidoptera (comprising the smaller moths). Most of the recent suggestions for the division of the Lepidoptera into sub-orders depend upon some single character. Thus J. H. Comstock has proposed to separate the three lowest families, which have—like caddis-flies (Trichoptera)—a jugum on each forewing, as a suborder Jugatae, distinct from all the rest of the Lepidoptera—the Frenatae, mostly possessing a frenulum on the hindwing. A. S. Packard places one family (Micropterygidae) with functional mandibles and a lacinia in the first maxilla alone in a suborder Laciniata, all the rest of the order forming the suborder Haustellata. T. A. Chapman divides the families with free or incompletely obtect and mobile pupae (Incompletae) from those with obtect pupae which never leave the cocoon (Obtectae), and this is probably the most natural primary division of the Lepidoptera that has as yet been suggested. Dyar puts forward a classification founded entirely on the structure of the larva, while Tutt divides the Lepidoptera into three great stirps characterized by the shape of the chorion of the egg. The primitive form of the egg is oval, globular, or flattened with the micropyle at one end; from this has apparently been derived the upright form of egg with the micropyle on top which characterizes the butterflies and the higher moths. These schemes, though helpful in pointing out important differences, are unnatural in that they lay stress on single, often adaptive, characters to the exclusion of others equally important. Although it is perhaps best to establish no division among the Lepidoptera between the order and the family, an attempt has been made in the classification adopted in this article to group the families into tribes or super-families which may indicate their probable affinities. The systematic work of G. F. Hampson, A. R. Grote and E. Meyrick has done much to place the classification of the Lepidoptera on a sound basis, so far as the characters of the imago are concerned, but attention must also be paid to the preparatory stages if a truly natural system is to be reached.
Jugatae.
Three families are included in this group having in common certain primitive characters of the wings and neuration (see fig. 6),
