GENERAL RESUME, WITH NOTICES OF UNDETERMINED FORMS.
Nine well authenticated fossil butterflies are now known, all from the European Tertiaries; five of these have been found in the gypsum beds of Aix in Provence, southern France, belonging to the Ligurian, a division of the upper eocene; one in the lignites of Rott in the Rhenish Provinces of Prussia, belonging to the Aquitanian, or lower miocene; and three in the marls of Radoboj in Croatia, Austria, appertaining to the Mayencian or middle miocene. Our present knowledge, then, places the apparition of butterflies towards the end of the lower tertiaries.
As a general rule the specimens thus far discovered are in a fair state of preservation, and especially are those parts preserved which enable us, with considerable confidence, to determine their exact affinities. Three of these insects belong to the highest family of butterflies, Nymphales, four to the Papilionidæ, and two only to the Urbicolæ. If it be considered probable that the lowest of these families was the oldest, we can reasonably account for the scarcity of its members in the tertiary strata by the fact that their almost universally robust and muscular frame enables them to maintain flight when they have lost all but the merest stubs of wings. They would thus seldom meet their end by falling into pools of water, or if at last they did, it would be with fragments of wings whose affinities could not be traced. This supposition would be strengthened on noticing that one of the two fossil forms classed here, Thanatites vetula, belongs to a group of genera which comprises the very feeblest flyers in the family; and by the further consideration that two of the three fossil Nymphalids belong to the weak-winged Oreades. Eugonia, as well as Pamphilites, were doubtless strong and bold flyers; while the genera of Papilionidæ were moderately endowed. To proceed further in the analysis of their structural relations, two of the three Nymphales belong, as we have said, to the highest group of butterflies, the Oreades, represented now by the dark brown butterflies of our meadows; the remaining one to the Præfecti, a group of gaily attired butterflies with angulated wings like our common thistle butterfly, the cosmopolite. Of the four Papilionidæ, three belong to the Danai; two of these three to the group Fugacia, represented by our common yellow brimstone butterflies; the third to the Voracia, or white butterflies of the garden, so destructive to cabbages and other cruciferous plants. The fourth Papilionid belongs to the lower subfamily Papilionides; not, however, to that group which contains our swallow-tailed butterflies, but rather to an allied tribe, represented in America only by the Parnasii of the Rocky Mountain region. The two Urbicolæ are divided between the Hesperides and Astyci, the former closely related to the dingy, sylvan hesperians of early spring, seldom seen but by the naturalist; the latter to the tawny, brisk little skippers busy around the flowers in June.
But a single family of butterflies, then, is unknown in a fossil state,—that of Rurales; and since this comprises, in the main, insects of exceedingly delicate structure and of small size, their absence is by no means unaccountable. Yet, as we shall see further on, there are intimations of the presence of some of their caterpillars in amber, and an obscure and doubtful reference to a fossil Polyommatus from the beds of Aix.
If we enquire where the allies of these nine fossil butterflies are now living, we must seek for those of four of them in the East Indies; for those of three of them in America, and especially in that part lying on the confines of the tropical and north temperate zones; for those of one of them in the north temperate zone of both Europe-Asia and America; and for those of one in the Mediterranean district; for those of two only, therefore, out of the nine, or less than one-fourth, in the region where the fossils were discovered. Analyzing this point still further, we notice that three out of the four species whose living allies are to be sought in the East Indies come from the older deposits of Aix, and that only one of the two remaining Aix species shows special affinities to American types; we thus find here, as among other insects and among the plants, a growing likeness to American types as we pass upward through the European tertiaries.
The study of the floras of the European tertiaries has proceeded so far that in most cases we are able to find, in the very beds where the butterflies occur, plants which we may reasonably judge to have formed the food of these insects in their earlier stages. In but a single instance is the family of plants, upon which it was necessary, or almost necessary, to suppose the caterpillar fed, entirely absent from tertiary strata; and since this family is the Cruciferæ, which in its very nature could scarcely have left a recognizable trace of its presence, the exception has no force.
After presenting these facts, for convenience sake, in a tabular form, we will pass on to the enumeration of those fossils which have been referred to butterflies, but whose exact position is still unsettled.