criticisms by Lefebvre, to the second of which Boisduval only alludes
in the most general way, and does not meet, while Butler makes no
reference to it at all. As far as regards the position of the tail,
Lefebvre is unquestionably wrong (see Pl. I, fig. 10), although his
fault is primarily due to the inaccuracy of the engraving given by
Boisduval, an inaccuracy which is slightly accentuated in our copy of
it (Pl. I, fig. 17). But by far the larger part of his paper is made up
of a detailed argument, drawn from the position and character of the
markings and from the direction of the nervures, in which he endeavors
to prove, and in most cases really does prove (though he errs in some
of his statements concerning the neuration), that these markings belong
to the front and not to the hind wing. He argues, for instance, that
the two oval dark spots are plainly traversed by the nervures of the
hind wing, and therefore cannot belong to that wing; that the minute
white spot apparently on the outer border of the hind wing is only
half a spot and must belong to the fore wing, and that the markings on
and near the costal border traverse both wings and must belong to the
one to which they certainly belong in part, the front wing. To this
Boisduval makes no sort of answer, and Butler, to judge from his silence
in the matter, and the comparative illustrations he gives on a plate
published subsequently,[M] considers it unproven. All of these writers
are, however, entirely wrong in supposing that the under surface of the
wings is exposed to view, and that the hind wing covers the front wing.
Boisduval does not distinctly state this; but the whole tenor of his
remarks shows that this was the view taken by him; and when Lefebvre
says: “Si de l’œil on suit les bords de la seconde aile, qu’avec le
Docteur je reconnais couvrir en grande partie la première,” no objection
is offered in Dr. Boisduval’s response; nor does he demur to Lefebvre’s
statement, when the latter speaks of the “face inférieure, celle que
nous voyons.” As we shall show later, however, the upper surface of the
wings is that exhibited on the stone, and the front wing almost entirely
conceals the hind one; compare Pl. I, fig. 13, drawn anew from the fossil.
In the same place to which we have just referred Mr. Butler adds the
following remarks on the probable affinities of this fossil:[N]
The true position of _C. sepulta_ is undoubtedly in the family
_Satyridæ_; and, so far as can be judged from the beautiful
figure in the “Annales,”[O] it is exactly intermediate in
character between three nearly allied genera now existing,
viz.:—_Neorina_, _Antirrhæa_ and _Anchiphlebia_, its more
immediate allies being the commonest species in each of the
above genera. Its characters are distributed between these
three species as follows:—
=====================================================================
_Neorina_ _Antirrhæa_ _Anchiphlebia_
_Lowii_, _Philoctetes_, _Archæa_,
Boisd. Linn. Hübn.
---------------------------------------------------------------------
Form of front wings, *
Form of hind wings, . . *
Tails of hind wings
(intermediate in
character between), * *
Submarginal spots of
front wings, * * *
Black disco-submarginal
spots of hind wings, . . *
Pale costal and discal
banding of wings, *? . . *
Limitation of dark dentated
basal area of hind wings, . . . . *
Submarginal line of hind wings, *
=====================================================================
The venation appears to be nearly similar to that of
Anchiphlebia. It is doubtful, however, whether the drawing
of the veins has been sufficiently attended to, to offer any
reliable characters.
In this paper he quotes Boisduval’s locality “Aix en Provence,” but when
he next refers to this insect[P] he gives it as from “Aix-la-Chapelle,
White sandstone,” a mistake, however, corrected subsequently. In this
latter paper he remarks:
I have discussed the position of this species in my catalogue
of Satyridæ, pp. 189, 190; showing that its nearest ally is
_Neorina Lowii_, a common Bornean species, but that it also has
a slightly more distant relationship to _Antirrhæa Philoctetes_
and _Anchiphlebia Archæa_, two common tropical American
forms; the amount of affinity, as regards the first two of
these species, may be seen on my plate, figs. 4 and 5; the
resemblance to Anchiphlebia is less striking, and the affinity
more doubtful; it has nothing to do with Cyllo.
That Butler should have so nearly pointed out the exact affinities of
this insect from the simple study of Boisduval’s plate, is unquestionably
due to his extended familiarity with butterflies, and especially with the
forms of this subfamily; but it also shows the essential harmony between
the markings of the under and upper surface of the wings of butterflies,
notwithstanding their frequent great dissimilarity; for Butler compares
this fossil with the recent forms on the assumption that the under
surface of the wings is seen in Boisduval’s plate.
The actual condition of the fossil, for an opportunity of examining which
I am indebted to the courtesy of Count Saporta, is this (see Pl. I, fig.
13): The thorax, hind legs and both pair of wings of the left side are
preserved, almost completely; all the rest is lost. The thorax is viewed
from above and somewhat on the left side; the hind coxæ seem to be almost
torn away from their immediate connection with the trunk. The two hind
legs are stretched out bent at the femoro-tibial articulation; the left
leg lies above both the wings and is apparently attached throughout,
although its base is covered a little by the crushed body; the right
leg lies below both the wings and is apparently partially detached,
though but slightly, from the coxæ; the tibio-tarsal articulation can
be distinguished (Pl. I, fig. 11) but not the tarsal joints. The wings
are bent over downward in a position the reverse of that of repose. The
fore wing covers the hind wing as in nature, but to such an extent as to
conceal the greater part of it; the guttered portion of the inner margin
of the hind wings is almost fully expanded, but apparently has a fold
next the submedian nervure. The fringe of the fore wing seems to be gone,
but that of the hind wing is preserved nearly throughout. Head, fore and
middle legs, wings of the right side and abdomen are wholly wanting.
The upper surface of the wings is, therefore, the part which attracts
most attention. That it is the upper and not the under surface which
is exposed to view is shown by the relation of the wings to each other
(Pl. I, fig. 10), by their unquestionable attachment to the thorax, of
which we certainly see only the upper portion with its smooth arched
dome marked by the sutures which separate the portions which compose
it; and by the design itself of the wings, which is such as pertains
to the upper rather than to the under surface of butterflies of this
group. These markings are most wonderfully preserved; and the careful
and prolonged study I have given every part of the fossil has enabled me
to separate, with a considerable degree of certitude, the markings which
appertain to the fore wing and those which belong to the hind wing. Those
of the latter are generally to be traced through the semi-diaphanous
fore wing and are given in Pl. I, fig. 8. One is aided greatly in this
investigation by following the lines and series of markings which extend
over both the exposed and covered portions of the hind wing; and then
by comparing the fainter and obscurer tints of the covered portion with
equivalent marks on other parts of the stone covered by both the wings;
in this way the markings of the hind wing may be separated from those
of the front wing, but subject, certainly, to some degree of doubt. In
the figure upon the plate (Pl. I, fig. 8) the portions to which the
least degree of doubt attaches are the outer halves of the two wings. I
am inclined to consider these as almost absolutely accurate. The parts
on the other hand which are more likely to be inaccurate are the basal
halves of the median interspaces of the fore wing and the contiguous
portion of the medio-submedian interspace. Assuming, however, that the
drawing faithfully represents the real markings of this extraordinarily
preserved fossil, a detailed description of its features follows.
The basal portion of the fore wing (Pl. I, fig. 8) is very dark, and
increases in intensity toward the border of the innermost light patch;
the latter is bounded by a line running in a nearly straight course
from the costal nervure, opposite the middle of the upper border of the
cell, toward the middle of the apical half of the submedian nervure;
but it extends slightly outward on reaching the lowest median nervule
and just below this turns baseward and makes a large ovoid curve of
an interspace’s diameter, returning to its course when it has nearly
completed the circuit and reached the middle of the medio-submedian
interspace; the outer limit of this large pale patch, which crosses the
cell and extends nearly to the middle of the lower median interspace,
nearly follows a line running from the upper extremity of the inner
border to and along the middle median nervule. Beyond this the upper
half of the wing, half-way to the apex, is nearly as dark as the basal
part, excepting in a large light patch which crosses the lowest two
subcostal and the subcosto-median interspaces, is broadest in the middle,
but twice as broad at the upper as at the lower extremity, and rounded
throughout excepting at the angular upper basal corner; its interior
margin is sharply defined, and is nearly parallel to the interior border
of the inner light patch, extending in a straight line from the subcostal
nervure midway between the origin of the first and second superior
nervules to the upper median nervule, about as far from its origin as
it is from the base of the first median nervule; the exterior border is
powdery, strongly convex and, starting from the subcostal nervure midway
between the bases of the second and third superior nervules, joins the
other border on the last median nervule; this patch is twice as long as
broad. Extending from the next to the lowest subcostal nervule to the
internal nervure, parallel to the outer border, is a broad indistinct
pale band, broadening below, and on either side merging indefinitely
into the darker parts of the wing, separated from the light patches by
only a narrow belt of dark scales, which becomes narrower and fainter
in the lower half of the wing; at its broadest the pale band is a
little broader than an interspace, and it contains in its middle and
at the middle of each interspace, as well as in the next to the lowest
subcostal interspace, a series of large circular dark spots, of nearly or
quite half the width of the interspaces in which they fall, often, and
especially in the upper interspaces, enclosing a small black pupil; these
spots are almost exactly parallel to the outer border, that in the lowest
median interspace with its outer border at an interspace distance from
it; with the exception of that in the lowest subcostal interspace, they
are each surmounted interiorly by a much smaller circular light spot, the
centre of which is near the circumference of the larger spot, so as to
infringe upon it; with the exception of the uppermost, which is nearly as
large as the spot on whose summit it is placed, the light spots are of
nearly equal size and about one-third of an interspace in diameter; or if
anything the two lower, seated on the largest spots, are smaller than the
others; the wing must have been wrinkled between the nervules next the
outer border, as shown by the dark lines running from the border to the
centre of the dark spots. The outer edge and the apex of the inner are
uniformly dusky and rather lighter than the other dark parts of the wing;
the fringe is evidently lost.
The hind wing is very dark at the base, like the fore wing, nearly as
far as the extreme tip of the cell; this dark area merges gradually into
a lighter portion, which crosses the wing as a very broad equal band
having its outer limit at a narrow, dark, regular belt, with ill defined
outline, which crosses the wing subparallel to the general course of the
outer border a little within the middle of the outer half of the wing;
within this broad light band are two narrow transverse powdery streaks
of dark scales, one extending from the extreme tip of the cell, and
broadening a little in its course, running in a curve opening inward to
the inner border; the other starting from the same point in an opposite
direction, and passing in a sinuous course, with varying width, toward
the middle of the basal two-thirds of the upper subcostal nervule,
hardly separate from the outer limits of the dark base of the wing. The
darkest part of the narrow band in the middle of the outer half of the
wing has a regular curve and strikes the borders in the middle of their
outer halves; there is a submarginal slender dark streak, separated by
scarcely more than its own width from the outer border, becoming narrower
toward the costal and inner borders, and especially towards the costal;
it is broken at the upper median nervule, where the upper portion joins
a second broader band, separated by a space nearly equal to itself from
the submarginal band; this leaves a nearly equal light band in the outer
part of the wing, broadest above and reaching from the costal border,
almost to the inner; along the middle of this belt is a series of six
round dark spots and ocelli, one in each of the interspaces excepting
the costo-subcostal; the largest is in the lower median interspace, and
is a spot nearly as broad as the interspace, deepening toward the centre
to a black pupil; the next largest, in the upper median interspace, is
an ocellus with a black pupil, immediately followed by a pale annulus,
again surrounded by a dark ring of equal diameter, the whole a little
more than half the width of the interspace; next larger are two spots of
less intense depth of color, one in the upper subcostal, the other in the
subcosto-median interspace, about one-third the width of the interspace,
the upper deepening, the lower becoming paler at the centre; the spot
in the lower subcostal and the medio-submedian interspace are equal and
smallest, about one-fourth the width of the interspace, and consist only
of rather faint, powdery marks, a little darker towards their centres.
The fringe of this wing seems to be preserved and is short, nearly equal,
dark, resembling a repetition of the submarginal streak.
Length of fore wing, 37mm.; breadth of fore wing, 20·5mm.; length of hind
wing, 31·75mm.; length of tail, 4mm.; distance of the base of the second
superior subcostal nervule of hind wing from the divarication of the
costal and subcostal nervules, 5·55mm.; rows of scales in the subcostal
region of the fore wings, ·075mm. apart; length of thorax, 5mm.; of hind
femora, 4·6mm.; of hind tibiæ 4·8mm.; of hind tarsi, 4·9mm..
Tertiaries of Aix, Provence, France; collection of Count de Saporta.
LETHITES SCUDDER.
_Satyrites_ Scudd. (nee Blanch.-Brullé), Rev. et Mag. de Zool.,
1871-72, 66.
The costal border of the fore wing (Pl. I, fig. 5) is gently and equably
curved, the apex moderately acute but well rounded, the outer margin,
except at its extremities, nearly straight, and the inner border straight
or almost so; the outer border is a little shorter than the inner and
about three-fifths the length of the costal margin.
The costal nervule terminates slightly beyond the middle of the costal
margin, its basal two-fifths presenting a considerable and almost uniform
expansion, which tapers rather rapidly at the tip, and reaches nearly
to the middle of the upper border of the cell. The subcostal nervule is
very slight on the basal half of the wing, closely connected with the
posterior surface of the swollen portion of the costal nervure and only
divaricating from that vein after the latter has lost its tumidity; it
emits its first superior nervule at slightly more than three-fifths the
distance from the tip of the bulbous portion of the costal nervure to
the upper apex of the cell; its second at midway between the origin of
the first and the tip of the cell; its third at midway between the upper
apex of the cell and the origin of the fourth, which arises at about
two-fifths the distance from the base of the third to the outer border of
the wing. The first superior nervule terminates near the middle of the
outer two-thirds of the costal border, the second midway between the apex
of the first and third; the third terminates just above, and the fourth
at or scarcely below, the tip of the wing. The first inferior subcostal
nervule arises at a very short distance beyond the base of the second
superior nervule, and curving rather strongly, terminates in the middle
of the upper half of the outer border; the second inferior nervule is
emitted from the first inferior as far beyond the base of the latter as
that is beyond the base of the second superior nervule; at its origin it
is directed inward as well as backward (forming the upper termination
of the cell) and passes backward in a small, narrow and rather strongly
curved bow, bent below more than above, beyond which it assumes a course
nearly parallel to the first inferior nervule; just beyond the arcuate
portion it is connected by a rather long, straight, oblique nervule,
directed considerably outward as well as downward, to the origin of
the upper median nervule. The median nervule is slightly enlarged at
the base, and diminishes gradually and regularly in size to its first
divarication, which is scarcely beyond the middle of the cell; the origin
of its middle branch is slightly nearer the origin of the basal than of
the terminal nervule; the latter strikes the middle of the outer border.
The submedian nervure is straight and not swollen at the base. The cell
is three times as long as broad, and scarcely more than half as long as
the wing.
The article from which the above is quoted, as originally written, closes
thus:
“The neuration of the fore wing does not seem to me to accord
sufficiently with that of any known genus of Oreades to admit of its
being classed with them. It undoubtedly has close affinities with the
characters of the genus Debis (= Lethe Hübn.) as laid down by Westwood
and Hewitson, if we exclude therefrom, as we should, the _Papilio
Portlandia_ of Fabricius. It is not a little interesting to notice
that these authors have arranged this group in immediate proximity to
the genus Cyllo (= Melanitis Fabr.), in which Dr. Boisduval placed
the fossil species from Aix, named by him sepulta. Nor is it less
interesting to find that in both genera all the living representatives
(even including those discovered since the publication of the ‘Genera of
Diurnal Lepidoptera’) are natives of the East Indies; so that the fossil
butterflies of Provence have their nearest living allies in the far East.”
Although differing from Neorina (Pl. II, fig. 8) very strikingly in the
form of the wing and the swollen base of the costal nervure, this genus
has some striking points of agreement with that in the neuration of the
fore wing. The nervure closing the cell indeed is straight in Lethites
and strongly curved in Neorina, but, as there, two of the superior
subcostal nervules arise before the tip of the cell, and the other two
are thrown off at about equal distances between the apex of the cell and
of the wing; the vein closing the cell meets the median nervure in both
cases as far beyond its second divarication as that is beyond the first;
the shape and proportionate length of the cell is nearly the same in the
two, but the costal nervure appears to be much shorter in Lethites.
With Lethe (Pl. II, fig. 6) and Debis (Pl. II. fig. 10) the fossil genus
can better be compared, as far as the form of the wing, the dilated
costal vein, and the position and direction of the straight vein closing
the cell are concerned; but in both these genera only a single superior
subcostal nervule is emitted before the apex of the cell; the form of
the cell again shows rather closer affinity between Lethites and these
genera, although the difference in these respects is but slight. It
is by no means distantly related to Enodia, in which two subcostal
nervules are emitted before the tip of the cell, but differs from it in
the much greater and more abrupt swelling of the costal vein, and in
the much greater distance beyond the second divarication of the median
nervure at which the vein closing the cell meets this nervure. It even
exhibits no small affinity to Cercyonis, and especially to those species
in which there is little dilation of the median nervure; the costal
nervure is swollen in precisely the same way, and the superior nervules
of the subcostal nervure are much the same; but the form of the wing is
strikingly different, and the lowest subcostal interspace much wider at
the base, in comparison with the width of the base of the subcosto-median
interspace, in Cercyonis than in Lethites; and this seems to be a
character of considerable importance. It may be noted in this connection
that the markings of the fossil must have closely resembled _Cercyonis
Pegala_.
Its nearest ally among living European types would seem to be _Maniola
Hermione_, in which the costal and median veins are about equally
swollen. The neuration of Lethites agrees with this genus in much
the same way as it does with Cercyonis, the comparative width of the
interspaces beyond the cell being very different in the living genera
from what it is in the fossil. In the form of the wing Maniola agrees
much better with Lethites than Cercyonis does, but the costa is much more
arched, and the cell is much the longer in Maniola; were there no obscure
spot in the lower median interspace in the male of _M. Hermione_, the
markings of the fossil would agree with it almost perfectly.
LETHITES REYNESII SCUDDER.
Plate I, figs. 2, 5.
_Satyrites Reynesii_ SCUDD., Rev. et. Mag. de Zool., 1871-72,
66-72, pl. vii (1872); IB., Descr. Pap. Foss. 1-7, pl. (1872);
IB., Geol. Mag. ix, 532-33, pl. xiii, figs. 2, 3 (1872); IB.,
Descr. Foss. Butt. 1-2, pl., figs. 2, 3 (1872); BROD., Distr.
Corr. Foss. Ins. [_Satyrites Reynesii_], 8-9 (1873).
I give below the original of the first paper cited above, excepting the
portion which was quoted under the genus.
In a recent examination of the rich collection of fossil insects from
Aix, preserved in the Museum of the city of Marseilles, my attention
was attracted by two little slabs containing the traces of a fossil
butterfly. Although by no means so well preserved, nor so perfect as the
remains of a butterfly from the same beds, described by Dr. Boisduval
more than thirty years ago, a glance showed that it could not be referred
to that species, since the costal nervure of the fore wings was greatly
swollen. No such form having to my knowledge been described from these
beds, Dr. Reynès, the accomplished director of the establishment,
courteously placed the best specimen in my hands for closer study; and
from it the following account and illustrations have been drawn. The
second specimen is very imperfectly preserved, but since it exhibits in
all its features an exact resemblance to similar parts in the better
specimen it undoubtedly belongs to the same species.
The fossil (Pl. I, fig. 2) is a natural cast of a butterfly lying upon
its side, the wings folded back to back, the legs extended as if hanging,
the tongue uncurled and, with the antennæ, drooping in a direction
similar to that of the legs. The right fore wing, which lies beneath, is
pushed a little outward and also forward, even at its base, showing that
the specimen must have been greatly macerated in very quiet water, before
being covered by the deposits which have preserved its more essential
features. The condition and position of all the parts also lead us to
conjecture that it was swept into its final resting place by a gentle
current, which left the slighter appendages lying in the direction of its
final action.
It is evident that the object is a cast, for the veins of the wing which
lie uppermost on the stone are impressed as we see them on the upper
surface of the wings of living Oreades, while those of the wing lying
beneath (veins which are plainly covered by the impressed nervures where
the two come in contact) are in relief, as seen on the under surface of
the same butterflies; that is, we have here the reverse of what would be
the case, were we examining a living butterfly in this position.
The parts before us are: a poorly preserved body, vague indications of
the terminal palpal joint, an antenna (probably a portion only), an
unrolled tongue, the hinder pair of legs and portions of the other pairs,
the greater part of the two front wings and fragments of the base of
the hind wings. Of the latter, no border remains and only the base of a
few of the nervules, which give scarcely any additional information as
to the pterology of the insect. The only portion of the margin of the
front wings which can be determined with certainty is the most essential
part, the apex and the upper half of the outer border of the left wing,
enough to show that its general contour was similar to that of the
European Satyrids of the present epoch; but throughout the remainder all
the nervules can be exactly traced. This being then the best preserved
portion of the insect, we will consider its structure in detail,
subsequently adding whatever can be gleaned from the examination of the
other parts.[Q] [The account of the structural framework of the wing is
given under the genus].