GENERAL RESUME, WITH NOTICES OF UNDETERMINED FORMS.

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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.

TABULAR VIEW OF FOSSIL BUTTERFLIES.

Names of Species (and families). First referred to genus. First described by When described. Found in Geological Horizon. Preserved in Museum of Nearest living allies found in Probable food of caterpillars. Parts of wings preserved.
(Nymphales).
Neorinopis sepulta Butl.
Cyllo. Boisduval. 1840 Aix. Ligurian, (upp. Eocene). Count Saporta, Aix. E. Indies. GramineÆ. Perfect wings of one side.
Lethites Reynesii Scudd. Satyrites. Scudder. 1872 Aix. Ligurian,(upp. Eocene). Marseilles. E. Indies. GramineÆ. Both fore-wings nearly perfect, superimposed.
Eugonia atava Scudd. Sphinx. Charpentier. 1843 Radoboj. Mayencian, (mid. Miocene). ? North temperate Zone. Salix, Populus or Betula. Upper half of one fore-wing.
(PapilionidÆ).
Mylothrites Pluto Scudd.
Vanessa. Heer. 1849 Radoboj. Mayencian, (mid. Miocene). Hof mineralien-Kabinet, Vienna. E. Indies. LeguminosÆ (Capparis?). Both fore wings nearly perfect.
Coliates Proserpina Scudd. —— Scudder. 1875 Aix. Ligurian, (upp. Eocene). Count Saporta, Aix. E. Indies. Smilax. Two fore-wings superimposed.
Pontia Freyeri Scudd. Pierites. Heer. 1849 Radoboj. Mayencian, (mid. Miocene). Hof mineralien-Kabinet, Vienna. Temperate America. CruciferÆ? Terminalia?? One fore-wing nearly perfect, but neuration obscure.
Thaites Ruminiana Heer. Thaites. Scudder. 1875 Aix. Ligurian, (upp. Eocene). Professor Heer, Zurich. Mediterranean district. Aristolochia. All the wings; those of one side nearly perfect.
(UrbicolÆ).
Thanatites vetula Scudd.
Vanessa. Heyden. 1859 Rott. Aquitanian, (low. Miocene). British Museum. Subtropical N. America. LeguminosÆ (HÆmatoxylon Gleditschia). All the wings, but superimposed and very obscure.
Pamphilites abdita Scudd. —— Scudder. 1875 Aix. Ligurian, (upp. Eocene). Marseilles. Tropical America. GramineÆ. One fore-wing perfect.

In the earliest accounts that we have found, including all those in the last century, the generic term Papilio was used for all Lepidoptera, and therefore we cannot be certain whether butterflies or moths are meant. Hueber’s plates, even, are so inferior that they afford no additional aid; but those of Sendel possibly represent, as we have noticed in the Bibliography at the commencement of this memoir, the early stages of butterflies preserved in amber. The only other direct references to butterflies preserved in amber are the following: Gravenhorst,[AV] in his enumeration of amber insects, gives under the Lepidoptera forty specimens referable to TineÆ and Tortrices, and besides these “mehre Raupen, sÄmmtlich, wie es scheint, Schildraupen, denen des Papilio W. album Ähnlich.” The probable nature of the ancient forest yielding amber renders it unlikely that any butterflies in their perfect state would be found in it. As a rule, butterflies are eminently fond of the light. This has already been remarked by Menge:[AW]—“Das fehlen grÖszerer Schmetterlinge im bernstein deutet auf einen finstern undurchdringlichen urwald, den die kinder des lichts gemieden haben.” Yet as some Theclas do feed upon coniferous trees, it is not impossible that the onisciform larvÆ, referred to by Gravenhorst, may belong to this group. As far as we can discover, no further reference is made to them, excepting by Giebel and Bronn in some of their lists and enumerations of fossil insects. The writings of Berendt, Menge and others, all bear testimony to the great rarity of Lepidoptera in amber, and most of those which have been discovered belong to the lowest two families, above referred to.

Dr. Hagen informs me that he has himself seen specimens of large butterflies in amber, but that these proved to be falsifications, recent European insects like Pieris rapÆ, etc., having been enclosed between slabs of amber, which were then fastened together and the edges roughened, all in so clever a manner that one would not suspect them to be spurious. These specimens were manufactured many years ago, and it is not impossible that it is to one of them that Hope refers in 1836, as found in the collection of Mr. Strong, though why he should quote Berendt as authority I cannot discover.

Heer, in the introduction to the lepidopterous portion of his “Insektenfauna der TertiÄrgebilde von Œningen,” says (p. 175): “Karg erwÄhnt zwar eines sehr schÖnen Œninger-Schmetterlings, der nach ZÜrich gekommen sein soll. Allein hier findet sich dieser nicht und die Angabe verliert noch mehr an Werth, wenn wir berÜcksichtigen, dass Karg das Thier nicht selbst gesehen hat.” Karg’s memoir in the “Denkschriften der SchwÄbischen Gesellschaft der Aerzte und Naturforscher,” T. I., I have been unable to examine.

Boisduval, in his final report upon Neorinopis sepulta, remarks that Count Saporta had written him that many years previously he had sent to the Paris Museum a “Polyommate fossile” from Aix. Count G. de Saporta, in reply to my inquiries concerning this specimen, says that his father can give me no further information concerning this specimen; nor could M. Oustalet and myself, in our search through the fossil insects of the Jardin des Plantes, discover any such relic.

In a recent number of “Nature” (No. 266), Mr. E. J. A’Court Smith writes of the discovery at Gurnet Bay in the Isle of Wight, of an insect bed in which were found, among other things, “a variety of flies, butterflies, and one or two grasshoppers;” no further information has yet been published concerning these relics, and my inquiries upon the subject have not, as yet, elicited any definite response.


                                                                                                                                                                                                                                                                                                           

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