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.”
[Illustration: Fig. 4.
Palæontina oolitica Butl. The neuration, after Butler’s first sketch.]
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.
[Illustration: Fig. 5.
Palæontina oolitica Butl. Facsimile of Butler’s first sketch.]
“_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.
[Illustration: Fig. 6.
Palæontina oolitica Butl. Facsimile of Butler’s second sketch.]
“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.
[Illustration: Fig. 7.
Palæontina oolitica Butl. The neuration, after Butler’s second sketch.]
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.
In his description of the insect Mr. Butler compares the neuration to
that of Caligo, and says its nearest allies are Caligo, Dasyophthalma and
Brassolis. In his latter paper he figures the wing of a Dasyophthalma by
way of comparison. In the genera named all the branches of the subcostal
nervure are simple, and are thrown off from the superior surface,
excepting the single set which is emitted from beneath, and which marks
(as in all butterflies) the limit of the discoidal cell; this corresponds
fairly with the first set of inferior veins emitted by the subcostal vein
in the fossil; for the other sets, however, no counterpart will be found
in the living types.
[Illustration: Fig. 8.
Palæontina oolitica Butl. Corrected sketch of the neuration.]
It was probably Mr. Butler’s want of familiarity with fossils that led
him to overlook several features which can be seen in these originals.
Having first traced the outline of the wing and the general course of
the veins directly from the specimens, I subsequently filled in by
measurement all the other parts which I could follow, studying each vein,
or supposed vein, with the utmost care, from one end to the other of its
course. The result of that study is presented in fig. 8, which differs
essentially in its details from the illustrations given by Butler, and
looks, as he himself confesses, “exceedingly anti-lepidopterous.” In the
first place, the wing is much narrower than depicted by him; and at the
extremity of a vein (the submedian vein of Butler’s sketch) there is a
slight but decided bending inward of the membrane, as very frequently
occurs at the line of demarcation between the middle and inner area
of the wing in all or nearly all the lower suborders of insects, but
never, so far as I am aware, in Lepidoptera. What he has given as a
simple costal vein is neither swollen at the base nor simple, but has
two inferior branches near the middle of the wing, united near their
origin by an oblique cross vein. Branching of the costal vein is unknown
in Lepidoptera; but if it should be claimed that this might be the
subcostal, just as much difficulty will be encountered with the structure
and relationship of the veinlets below, which must then be considered
as belonging to the median vein; in no Lepidoptera can any such
irregularity be shown, nor so disproportionate a magnitude of the area
covered by the median nervure and its branches; a branched internal vein
and cross-veins, which probably united all the longitudinal nervures at
no great distance from the outer border (but which can only be certainly
predicated for the lower three median interspaces), place this insect
wholly beyond the pale of the Lepidoptera. It is but fair to say that
Mr. Butler, having examined the original after he had in his possession
a tracing of fig. 8, denies the existence of the cross-veins; there is
one point, however, which an unprejudiced examination of the fossil
cannot fail to show; that Butler’s “fourth branch” of the subcostal[BE]
arises not from his third branch, but from his upper discoidal vein;
if he can reconcile either this or the points already referred to (on
the supposition that his sketch is otherwise an accurate one) with the
neuration of any group of butterflies, the writer will be the first to
acknowledge it.
As our only purpose in this place is to deny the lepidopterous character
of Palæontina, it is unnecessary to say anything in defence of the view
we have expressed of its homopterous affinities; the superior position
of the cell, the position and character of the lower cross veins (which
we believe really traversed the entire wing), with their origin at the
indentation of the lower border, suggest such a relationship, although
there are not a few points in which it differs somewhat strikingly from
living types.
The discovery of a fossil in the cabinet of the Rev. Mr. Brodie, which
was found in England at the same or nearly the same horizon, as _P.
oolitica_, and which seems to be a pupa case of one of the Cicadida of
rather unusual size, renders my suggestion more worthy of credence.
At the conclusion of his latter paper Mr. Butler draws attention to
the fact that Messrs. Westwood and Bates had expressed their agreement
with his views. It should, however, be borne in mind, that, so far as
appears from any facts which have been published, these gentlemen, whose
well considered views upon the subject would unquestionably be of great
weight, expressed this assent only upon a brief evening examination of
a very obscure fossil in a poorly lighted hall, and before any one had
questioned its lepidopterous character.
FOOTNOTES
[A] Probably an error for Tab. 5, fig. 28a, 28b, which seems to represent
a Tenthredinidous larva.
[B] The plate, however, is wanting, both in the copy belonging to the
Smithsonian Institution and in that in the Library of the Boston Society
of Natural History, so that I cannot tell whether it is copied from
Boisduval’s figure or is an original.
[C] Bull. Soc. Ent. France, 1838, 51-52.
[D] Bull. Soc. Ent. France, 1839, 11-12.
[E] Annales Soc. Ent. France, ix, 371-374 pl. 8.
[F] By Marcel de Serres, Actes Linn. Soc. Bord., Vol. xiii, pl. 2;
Pictet, Traite de Palæont., II pl. 40, fig. 11; Butler, Lep. Exot. I, pl.
18, fig. 2; Ib., Geol. Mag. x., pl. 1, fig. 3.
[G] See particularly papers by Coquand, Marcel de Serres, Heer, Saporta,
Giebel, Westwood and Butler.
[H] Ann. Soc. Ent. France (2) ix, 71-88, pl. 3, ii.
[I] Bull. Ent. Soc., France, 1851. 97-8.
[J] Cat. Satyr., 189-190.
[K] Species of Lethe.
[L] As, for instance, in the _Sphingidæ_, _Heliconidæ_, etc.
[M] Lep. Exot. pl. 48.
[N] Loc. cit., p. 190.
[O] In this figure the neuration has not been very clearly defined, the
veining of the hind wings not being continuous.
[P] Lep. Exot., 127, pl. xlviii. Geol. Mag., x, 3, pl. i.
[Q] It should first be premised that throughout this description the fore
wing will be spoken of as if it were perfect; for so completely are the
essential parts preserved that one may feel a strong degree of confidence
as to the character of the remainder; scarcely any of the costal margin
can be traced on the stone, and yet one may describe with nearly absolute
certainty the point where the extremities of the costal and the first two
upper branches of the subcostal nervures strike it. For those, therefore,
who would follow the description with a severely critical eye, the
illustrations we have given will correct any apparent overstatement of
the text.
[R] Acta Acad. Leop.-Carol., XX, 108.
[S] Insekt. Tert. Œning. ii, 177-79.
[T] Wahrscheinlich ist ausser diesem noch ein Ast da, der aber verwischt
ist.
[U] Lep. Exot. I. 128.
[V] Compare, in this respect, Butler’s Revision of the Pierinæ, Cist.
Ent., I, iii, pl. i, fig. 8; or Trimen, Rhop. Afr. Austr., Pl. ii, fig. 2.
[W] Insekt. Tert. Œning., II, 179-82.
[X] Butt. N. Amer., i, Argynnis, I.
[Y] Cat. Fabr. Lep., 109.
[Z] Lep. Exot., I, xv, 127-28, Pl. 48, fig. 7; Geol. Mag., x, 3-4, Pl. 1,
fig. 7.
[AA] Insekt. Tert. Œning, II, 180.
[AB] Compare the illustrations referred to in the note on page 44.
[AC] These, it is true, are Danai, but aberrant forms, like Leptidia, etc.
[AD] See, however, the American genus Leodonta.
[AE] Insekt. Tert. Œning., ii, 182-3.
[AF] Palæontogr. viii, 12-13.
[AG] Saporta. Revision de la flore des gypses d’Aix. Ann. Sc. Nat. [5]
Bot., xv, 284.
[AH] “La proportion des Graminées relativement au total des Phanérogames,
qui est de 4·5 sur 100, est en rapport avec les minima relatifs de
cette famille, tels qu’on les observe à la Nouvelle-Guinée et à la
Nouvelle-Grenade.” Saporta, loc. cit., 292.
[AI] “Dans la flore des gypses d’Aix, non-seulement les Légumineuses
occupent le premier rang, comme dans la plupart des flores
intertropicales actuelles de l’ancien et du nouveau continent, mais
elles atteignent une proportion de 13 pour 100, pour l’ensemble des
Phanérogames, proportion parfaitement en rapport avec celle de 12 sur 100
qui est fréquente, selon M. de Candolle, dans certaines régions chaudes,
telles que Timor, le Congo, etc.” Saporta, loc. cit., 292.
[AJ] Since this was written, Count Saporta writes me: “Le genre Smilax
est un des genres tertiatres les plus frequents. J’ai aussi signalé dans
le dépôt voisin de St. Zacharie (étage Tongrien inférieure [and therefore
but slightly more recent]) une feuille qui m’a paru devoir se ranger
parmi les Dioscorées.”
[AK] An old writer in Fuessly’s Magazin, writing from Italy, says that
Thais feeds in that country upon Quercus. Five species of Quercus are
known from Aix, but the statement in Fuessly’s Magazin has never, to my
knowledge, been confirmed.
[AL] “Nous n’avons pas encore rencontré d’Aristolochia dans les gypses
d’Aix; mais l’existence du genre dans le tertiaire moyen ne saurait être
mise en question, depuis que nous avons entre les mains une superbe
empreinte de Radoboj (Aristolochia venusta Sap.), qui dénote une forme
voisine des Aristoloches à feuilles persistantes et demi-coriaces, comme
l’_A. reticulata_ Nutt. de Virginie.” Saporta, loc. cit., 342-3.
[AM] In a recent letter from Count Saporta he remarks: “Relativement au
_Thaites Ruminiana_, je n’ai pas encore découvert à Aix de vestiges du
genre Aristolochia, mais _ce genre devait y exister_.”
[AN] Vierteljahrsschrift naturf. Gesellsch. Zurich, i. 12, 1856.
[AO] Recherches sur le Climat et la Végétation du pays tertiaire, 4to,
1861.
[AP] I venture to give one more extract from a recent letter received
from Count Saporta, although he writes:—“Je vous écris n’ayant sous les
yeux ni mes livres ni mes collections, ce qui enlevera nécessairement un
peu de précision à quelques-unes de mes réponses.”
“Il est bien plus difficile de justifier par des exemples tirés de la
nature des plantes la présence à Radoboj d’un insecte se nourissant de
Capparis. Les Capparis ont du exister, mais leur feuilles sont difficiles
à distinguer à cause de l’absence de caractères différentiels; leur forme
et leur nervation peu visibles doivent les faire confondre avec beaucoup
d’autres. Il me semble pourtant que des Capparis ont été signalés soit à
Radoboj, soit à Hoering en Tyrol, dépôt un peu plus anciens [Tongrian],
mais en l’absence de mes livres je ne saurais vous l’affirmer.”
[AQ] I use these genera in the sense indicated in my Historical Sketch of
generic names. Proc. Am. Acad. Arts, Sci., X, pp. 91-293.
[AR] Heer et Gaudin, Climat du pays tertiaire, pp. 133-171.
[AS] Ann. Sc. Nat., [5] Bot., xv, 322.
[AT] Climat du pays tertiaire, ed. Gaudin, p. 205.
[AU] Loc. cit., p. 205.
[AV] Arbeit Schlesisch. Gesellsch. Vaterl. Kultur, 1834, 92-3.
[AW] Programm Petrischule Danzig, 1855-56, 4to, p. 30.
[AX] Loc. cit., 387.
[AY] Loc. cit., 390.
[AZ] 1872, xxxi.
[BA] Perhaps Mr. Butler is not altogether to blame in confounding Aix
in Provence with Aix-la-Chapelle; at any rate the mistake had been made
previously by the translator of Heer’s paper in the Quarterly Journal of
the Geological Society of London, VI, 72; the error is corrected by Mr.
Butler at the end of his volume.
[BB] He seems not to have seen the earlier publication of Mr. Brodie.
[BC] Geol. Mag. [2] I, 448.
[BD] These veins have been given a distinct name (discoidal) by the
English Entomologists, as if they hail an independent origin, and had
nothing to do with the subcostal nervure; but by the use of this name,
we wholly lose sight of the simple plan of neuration belonging to the
wings of these insects. I have therefore preferred to speak of them as
the inferior subcostal nervules, in contradistinction to the superior
branches of the same vein.
[BE] In this case he counts from the tip of the wing, in reverse order.
EXPLANATION OF THE PLATES.
[My best thanks are due to my courteous friend Mr. Auguste Sallé, for
his kind agency in securing an artist for the engraving of these plates.
Owing, however, to the distance at which the work was done, a few errors
have unavoidably occurred, which, to prevent misapprehension, are
mentioned below.]
Plate I.
Fig. 1. _Eugonia atava._ Copied from Heer, Insekt. Tert. Œning., ii,
pl. xiv, fig. 8 (1/1).
“ 2. _Lethites Reynesii._ Drawn by S. H. Scudder (1/1).
“ 3. _Eugonia atava._ Copied from Charpentier, Nov. Act.
Leop.-Carol., xx, pl. xxii, fig. 4 (1/1).
“ 4. _Eugonia j.-album._ Neuration of fore wing; drawn by S. H.
Scudder (1/1). The second superior subcostal nervule is
carried too far toward the tip of the wing.
“ 5. _Lethites Reynesii._ Fore wing; drawn by S. H. Scudder (2/1).
“ 6. _Eugonia j.-album._ Markings of the upper surface of the fore
wing; drawn by S. H. Scudder (1/1).
“ 7. _Eugonia atava._ Neuration of tip of fore wing (2/1); copied
from Heer; Insekt. Tert. Œning., ii, pl. xiv, fig. 3a.
“ 8. _Neorinopis sepulta._ Markings of the upper surface of the
two wings, restored; drawn by S. H. Scudder (2/1). The drawing
represents the general effect of the fore wing as darker than
the hind wing, and in so far is inaccurate.
“ 9. _The same._ Neuration of the two wings, separated; drawn by
S. H. Scudder (2/1).
“ 10. _The same._ Neuration of the two wings, as seen in the fossil;
drawn by S. H. Scudder (2/1). The engraver has unfortunately
made the lines of the hind wing the heavier, as if it lay
uppermost; they should have been the lighter.
“ 11. _The same._ Right hind leg; drawn by S. H. Scudder (2/1).
“ 12. _The same._ Left hind leg; drawn by S. H. Scudder (2/1).
“ 13. _The same._ Drawn by S. H. Scudder (1/1). The spot of the
medio-submedian interspace of the fore wings has not been well
rendered by the engraver.
“ 14. _The same._ Copied from Lefebvre, Ann. Soc. Ent. France [2], ix,
pl. iii, II, fig. A (1/1).
“ 15. _The same._ Copied from the same, fig. C (1/1).
“ 16. _The same._ Copied from the same, fig. B (1/1).
“ 17. _The same._ Copied from Boisduval, Ann. Soc. Ent. Fr., ix, pl.
8 (1/1).
Plate II.
Fig. 1. _Zophoessa Sura._ Neuration of the wings; drawn by G. Willis
(1/1).
“ 2. _Mylothrites Pluto._ Copied from Heer, Insekt. Tert. Œning.,
ii, pl. xiv, fig. 4 (1/1).
“ 3. _Zophoessa Sura._ Markings of the upper surface of the wings;
drawn by G. Willis (1/1).
“ 4. _Delias Pasithoe._ Neuration of the wings; drawn by G. Willis
(1/1).
“ 5. _Coliates Proserpina._ Neuration and markings of fore wings;
drawn by S. H. Scudder (2/1). The tip of the costal nervure
has been extended too far toward the apex of the wing.
“ 6. _Lethe Dyrta._ Neuration of the wings; drawn by G. Willis
(1/1).
“ 7. _Mylothrites Pluto._ Neuration of the wings; after a drawing
obtained through Mr. Brunner de Wattenwyl (1/1). The second
superior subcostal nervule on the left wing should join the
nervure midway between the bases of the first and third
nervules.
“ 8. _Neorina Lowi._ Neuration of the wings; drawn by G. Willis
(1/1).
“ 9. _Lethe Dyrta._ Markings of the lower surface of the fore
wing; drawn by G. Willis (1/1).
“ 10. _Debis Sinorix._ Neuration of the wings; drawn by G. Willis
(1/1).
“ 11. _Zophoessa Sura._ Markings of the lower surface of the fore
wing; drawn by G. Willis (1/1).
“ 12. _Pontia Protodice._ Neuration and markings of the upper surface
of fore wing; drawn by S. H. Scudder (1/1).
“ 13. _Neorina Lowi._ Markings of the upper surface of the wings;
drawn by G. Willis (1/1). This was drawn for the pattern of
markings only; the neuration is faulty.