Fossil Butterflies Memoirs of the American Association for the Advancement of Science, I.
Part 10
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 | First | First | | | (and families). |referred to| described | When | | | genus. | by |described. | +--------------------------+-----------+------------+-----------+ | (_Nymphales_). | Cyllo. | Boisduval. | 1840 | | Neorinopis sepulta Butl. | | | | +--------------------------+-----------+------------+-----------+ | | | | | | Lethites Reynesii Scudd. |Satyrites. | Scudder. | 1872 | | | | | | +--------------------------+-----------+------------+-----------+ | | | | | | Eugonia atava Scudd. | Sphinx. |Charpentier.| 1843 | | | | | | +--------------------------+-----------+------------+-----------+ | (_Papilionidæ_). | Vanessa. | Heer. | 1849 | | Mylothrites Pluto Scudd. | | | | +--------------------------+-----------+------------+-----------+ |Coliates Proserpina Scudd.| —— | Scudder. | 1875 | | | | | | +--------------------------+-----------+------------+-----------+ | | | | | | Pontia Freyeri Scudd. | Pierites. | Heer. | 1849 | | | | | | | | | | | +--------------------------+-----------+------------+-----------+ | | | | | | Thaites Ruminiana Heer. | Thaites. | Scudder. | 1875 | | | | | | +--------------------------+-----------+------------+-----------+ | | | | | |(_Urbicolæ_). | Vanessa. | Heyden. | 1859 | | Thanatites vetula Scudd. | | | | | | | | | +--------------------------+-----------+------------+-----------+ |Pamphilites abdita Scudd. | —— | Scudder. | 1875 | | | | | | +--------------------------+-----------+------------+-----------+
+--------------------------+---------+---------------+-----------------+ | Names of Species | | Geological | Preserved in | | (and families). |Found in | Horizon. | Museum of | | | | | | +--------------------------+---------+---------------+-----------------+ | (_Nymphales_). | Aix. | Ligurian, | Count Saporta, | | Neorinopis sepulta Butl. | |(upp. Eocene). | Aix. | +--------------------------+---------+---------------+-----------------+ | | | Ligurian, | | | Lethites Reynesii Scudd. | Aix. |(upp. Eocene). | Marseilles. | | | | | | +--------------------------+---------+---------------+-----------------+ | | | Mayencian, | | | Eugonia atava Scudd. |Radoboj. |(mid. Miocene).| ? | | | | | | +--------------------------+---------+---------------+-----------------+ | (_Papilionidæ_). |Radoboj. | Mayencian, |Hof mineralien- | | Mylothrites Pluto Scudd. | |(mid. Miocene).| Kabinet, Vienna.| +--------------------------+---------+---------------+-----------------+ |Coliates Proserpina Scudd.| Aix. | Ligurian, |Count Saporta, | | | |(upp. Eocene). | Aix. | +--------------------------+---------+---------------+-----------------+ | | | | | | Pontia Freyeri Scudd. |Radoboj. | Mayencian, |Hof mineralien- | | | |(mid. Miocene).| Kabinet, Vienna.| | | | | | +--------------------------+---------+---------------+-----------------+ | | | Ligurian, | Professor Heer, | | Thaites Ruminiana Heer. | Aix. |(upp. Eocene). | Zurich. | | | | | | +--------------------------+---------+---------------+-----------------+ | | | | | |(_Urbicolæ_). | Rott. | Aquitanian, | British | | Thanatites vetula Scudd. | |(low. Miocene).| Museum. | | | | | | +--------------------------+---------+---------------+-----------------+ |Pamphilites abdita Scudd. | Aix. | Ligurian, | Marseilles. | | | |(upp. Eocene). | | +--------------------------+---------+---------------+-----------------+
+--------------------------+-------------+-------------+---------------+ | Names of Species | Nearest | Probable |Parts of wings | | (and families). |living allies| food of | preserved. | | |found in |caterpillars.| | +--------------------------+-------------+-------------+---------------+ | (_Nymphales_). | E. Indies. | Gramineæ. |Perfect wings | | Neorinopis sepulta Butl. | | | of one side. | +--------------------------+-------------+-------------+---------------+ | | | |Both fore-wings| | Lethites Reynesii Scudd. | E. Indies. | Gramineæ. |nearly perfect,| | | | | superimposed. | +--------------------------+-------------+-------------+---------------+ | | North | Salix, |Upper half of | | Eugonia atava Scudd. | temperate | Populus | one | | | Zone. | or Betula. |fore-wing. | +--------------------------+-------------+-------------+---------------+ | (_Papilionidæ_). | E. Indies. | Leguminosæ |Both fore wings| | Mylothrites Pluto Scudd. | |(Capparis?). |nearly perfect.| +--------------------------+-------------+-------------+---------------+ |Coliates Proserpina Scudd.| E. Indies. | Smilax. |Two fore-wings | | | | | superimposed. | +--------------------------+-------------+-------------+---------------+ | | | | One fore-wing | | Pontia Freyeri Scudd. | Temperate | Cruciferæ? |nearly perfect,| | | America. |Terminalia?? | but neuration | | | | | obscure. | +--------------------------+-------------+-------------+---------------+ | |Mediterranean| |All the wings; | | Thaites Ruminiana Heer. | district. |Aristolochia.|those of one | | | | | side nearly | | | | | perfect. | +--------------------------+-------------+-------------+---------------+ | | Subtropical |Leguminosæ | All the | |(_Urbicolæ_). | N. |(Hæmatoxylon | wings, but | | Thanatites vetula Scudd. | America. |Gleditschia).| superimposed | | | | | and very | | | | | obscure. | +--------------------------+-------------+-------------+---------------+ |Pamphilites abdita Scudd. | Tropical | Gramineæ. | One fore-wing | | | America. | | 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.
NOTICE OF INSECTS WHICH HAVE BEEN ERRONEOUSLY REFERRED IN RECENT TIMES TO BUTTERFLIES.
1. _Cyllonium Boisduvalianum_ WESTW., and _C. Hewitsonianum_ WESTW.
These two insects were figured by Westwood in the Quarterly Journal of the Geological Society of London for November, 1854, the former (reproduced in our fig. 2) on Pl. XVII, fig. 17; the latter (reproduced in our fig. 3) on Pl. XVIII, fig. 27. Of the former he makes the following remarks:[AX] “Pl. XVII, fig. 17 represents a number of fragments of delicate tegument, covered with minute punctures and traversed by straight and somewhat radiating veins, which appear like portions of the hind wing of some species of Butterfly, entirely denuded of scales.” The name is given to it in a note to the explanation of the plates, p. 395. Concerning the second he says:[AY] “Pl. XVIII, figs. 27 and 30, appear to be portions of the hind wings of some species of Butterfly; still they have very much of a vegetable aspect. The surface is covered with minute punctures, which may be the cells for the insertion of the quills of the coloured scales, which are all removed, supposing the specimens to be Lepidopterous.” The name we have quoted is given only to fig. 27, in a foot note on p. 396.
I have not been able to find, even with Mr. Brodie’s help, the first specimen referred to; but an examination of the original of the latter (see fig. 3) proved that, while it is unquestionably an insect, it cannot be referred to the Lepidoptera; the punctures referred to are both too large and much too irregularly disposed to have been the points of insertion of the scales; they are probably the marks of the insertion of hairs, such as are not uncommonly seen irregularly scattered over the wings of insects belonging to the other suborders. As the figure of the first species closely resembles in this particular the one I have seen, I am forced to the conclusion that neither of these wings are lepidopterous. Plainly, the only reason why a new generic name was appended to these forms was that their remains were too fragmentary to afford the slightest guess as to what modern genus they might be referred. The fossils came from the English Purbecks.
2. _Palæontina oolitica_ BUTL.
The first notice I find of this remarkable and very interesting fossil is that published in various literary and scientific London journals reporting remarks given at a meeting of the Entomological Society of London, and which afterward appeared as follows in their Proceedings:[AZ]
“Mr. Butler exhibited a remarkably perfect impression of the wing of a fossil butterfly in the Stonesfield slate. It appeared to be most nearly allied to the now existing South American genus Caligo.”
A full description of this insect soon appeared in the author’s “Lepidoptera Exotica,” accompanied by a plate; both were afterward republished in the “Geological Magazine.” In fig. 4 we reproduce fig. 1 of his plates, representing the neuration of Palæontina; and in fig. 5, fig. 2 of his plates, subsequently copied by “The Graphic.” A description of the genus and species is first given, which it is unnecessary to reproduce here; afterward, the following remarks:
“[126] Though a British insect, this species belongs to a group so completely tropical that I do not hesitate to describe and figure it in the present work; its nearest allies are the genera Caligo, Dasyophthalma and Brassolis, all three essentially tropical American genera.
“_P. oolitica_ is especially interesting, as being the oldest fossil butterfly yet discovered; the most ancient previously known to science having been found in the Cretaceous series (white sandstone of Aix-la-Chapelle[BA]), whilst the bulk of the known species are from the Lower Miocene beds of Croatia; it is also interesting as belonging to the highest family of butterflies, and to a subfamily intermediate in [127] character between two others, namely, the Satyrinæ and Nymphalinæ, whilst the more recently discovered fossils are referable, with one exception, to the two latter groups. The nervures appear to have been impregnated with iron, which will partly account for their well-defined condition.”
Happening to be in London not long after the publication of the description and illustration of this insect, I took pains to make a very careful examination both of the original specimen, which Mr. Charlesworth kindly allowed me to study at my leisure, and of its reverse, which is preserved in the School of Mines, Jermyn street. I mentioned to Mr. Butler and to others, my conviction that the insect was to be considered homopterous rather than lepidopterous, and on my return to America, exhibited before the Natural History Society of Boston, drawings which I had made from the originals; my comments at that time were published very briefly, as I was reserving the proof of my statements for the present paper. Mr. Butler, however, was induced by this publication[BB] to examine the reverse at the Jermyn street Museum, and although he had been supplied by me with a rough tracing of the drawing I had taken of it, he failed to be convinced of any mistake, and published a paper in defence of his own view in the Geological Magazine for October, 1874. In this paper he gives new drawings of the insect, quotes portions of letters in which I had expressed my opinions upon the nature of the fossil, gives the remarks referred to from the “Proceedings of the Boston Society of Natural History,” and makes, among others, the following comments.
“Seeing that Mr. Scudder had made his views public, I felt that it was time for me to take similar steps on my side. I therefore availed myself of an early opportunity of again visiting Jermyn street, where, through the courtesy of the officers, I was enabled to make a sketch of the impression in the Museum. I found it impossible to make a tracing of it, and therefore drew the whole by measurement. This sketch is now produced on Pl. XIX, fig. 4 [see fig. 6]; and any body can judge for himself whether or not it is more perfect than that which I previously figured (see Geol. Mag., 1873, Vol. X, p. 2, Pl. I, fig. 2 [see fig. 5]).”
“In order to show the extent to which the Jermyn street example is deficient, I have restored it (fig. 5 [our fig. 7]), filling in the blanks from Mr. Charlesworth’s specimens. By comparing the latter with the wing of _Dasyophthalma_ (fig. 1), and _Cicada_ (fig. 2), one may come to a pretty accurate conclusion as to the group of insects to which it ought to be referred.”[BC]
The neuration of Lepidoptera as a group is the simplest in the whole order of insects, if we except that of the elytra of Coleoptera; this is due, doubtless, to the fact that their wings are heavily scaled, concealing the nervures; just as in Coleoptera, the thickness and opacity of the fore wings often completely masks the neuration.
The normal number of veins in the wings of insects is six, disposed to a certain extent in pairs; the middle pair usually ramify to a greater extent than the others, and support most of the membrane of the wing. In butterflies the foremost vein is always absent and very commonly the hindmost, so that there are but five (often but four) principal veins, usually designated, though not very appropriately: costal, subcostal, median, submedian and (when present) internal, reciting them in their order from in front backward. The costal, submedian and internal nervures are invariably simple and terminate at the margin, or are occasionally lost in the membrane of the wing. The subcostal and median nervures, on the other hand, are as invariably forked, and with their branches support nearly the entire wing; the subcostal nervure curves downward and the median upward so as to meet, or nearly to meet, not far from the middle of the wing, and to enclose between them a large space called the discoidal cell; the branches of the median nervure are all thrown off from its lower edge before union with the subcostal; the principal branches of the subcostal nervure are, on their side, thrown off from its upper edge; but, as the nervure curves downward at the extremity of the cell, another set is thrown off (at least in the fore wings) from the lower edge; and it is these veins, rather than the subcostal nervure proper, which unite with the median to close the cell.[BD] None of the median, nor any of the inferior subcostal nervules are ever branched; but at the apex of the wing, where the play of neuration is usually the greatest, the last superior subcostal nervule is occasionally forked in the front wing. This is the only forked branchlet in either of the wings.
The last figure of _P. oolitica_ given by Mr. Butler agrees in all its essential features with his first illustration. They both represent a front wing with four principal nervures,—costal, subcostal, median and submedian; the costal nervure is swollen at the base and extends, unbranched, to the tip of the wing; the median nervure is three-branched, the three forks simple, equidistant, emitted from the apical half of the vein, which at its extremity is united by a cross vein to a branch of the subcostal, closing the cell; the submedian nervure is simple and divides the space between the median vein and the margin of the wing. So far all is in accordance with the lepidopterous type; but when we examine the subcostal vein, which occupies nearly half the wing, the resemblance ceases altogether. This vein is represented as bearing no superior branches, but as sending out from its _inferior_ surface three distinct veinlets, the first and second of which again emit a tributary from their inferior surfaces. This is a structural anomaly which finds no counterpart whatsoever in any family of butterflies. So that should we accept Mr. Butler’s own sketch of the fossil as correct, it would be impossible to consider the wing that of a butterfly.