Fossil butterflies

ONTO

TOR

01293524 3

III
61

UNIVERSITY OF

T

3

Tae

$
woe

.
UPD
tair teleet

pnate

Mast 4s
+) Te ta

mg
‘ ut
: 8)
i |
i es
vig ey
; ay
ser
i

ote

4

coe rr

epi ateries
¢
erated
t 3.

sigh

biFpttecers
Se tts

At
ts irpt leet
ietetele!

meTil i rahelet

¥
i

vee
=

we

erpeaers.

—- sta!
Slit ies
= pint

=

ts

papers

rts

3s

r

Fee

‘ = “4 -&)
, = ‘ 2
Stem "coal , Le
ee i -
~*~ =, *
a
- J:

_ OF THE

AMERICAN ASSOCIATION

FOR THE

SALEM, MASS.
F, W. PUTNAM, PERMANENT SECRETARY, A.A.A.S.
1875.

ae ee

LETTER OF GIFT.

-. _ Porrnanp, Aug. 22, 1873.

Mrs. Exizanetu Tompson of New York City, to-day elected a member,
sympathizing with the purposes of our Association in the advancement of science,
and seeing the new crop of young and industrious scientific investigators who
are to form the future basis of this Association following in the footsteps of the
veterans of science who founded it, and being aware of the financial difficulties "a
which often beset the path of those noble men of science who labor more for
truth than for profit’s sake, wishes to place at the disposal of the Permanent :
Secretary the sum of one thousand dollars, to be used according to the directions ce
of the Standing Committee, for the promotion and publication of such original
investigations by members of the Association as may be accepted by the said
Standing Committee, to be published by means of this special donation. B.o6

[Signed | P. H. Van per Weype.

To the Standing Committee of the American

Association for the Advancement of Science.

REPORT Sain
" . . ; # “=e = A
OF THE SPECIAL COMMITTEE ON THE THOMPSON FUND, HARTFORD MEETING, = ae

a

, AUGUST, 1874.

: Tur Standing Committee of the Association at the Portland Meeting ap-
a _ pointed the undersigned a Committee with full power to accept and print such
‘ papers as they might deem of sufficient importance to be published by the rinse * .
CS of Mrs. Thompson. bar “a
ag In accordance with the duties assigned to them, the Committee’ have acc e
a the Memoir by Mr. Scudder on Fossil Butterflies as the first paper to be publ ish
* ~ by the Tompson Funp, and while regretting that the unavoidable delay”
; ' engraving the plates apdivess their having the eign of presoniae

patron will accept the Memoir as one in every way worthy of the ae .
bestowed.

AsA GRAY, \

James HALL, ©
Tuomas Hitt, ee.
P. H. Van per Weypr, ) Committee.
J. L. LeConre, Oe
T. Srerry Hunt,

F. W. Purwam,

a

4 *

a

my.
* ee
age x

a ;

Saad

TO

COUNT GASTON DE SAPORTA,

OF AIX IN PROVENCE,

WHOSE EXTENDED MEMOIRS ON THE FLORA OF THE TERTIARIES OF SOUTHERN FRANCE FORM THE BASIS

3

4

or riba Seeoenieio as. CONCLUSIONS OF THIS ESSAY; AND WHOSE UNWONTED COURTESIES HAVE PERMITTED
” “a? .

4

‘=

eee erce OF THR Riot IMPORTANT FORSL BUTTERFLIES,

#) THIS MEMOIR IS RESPECTFULLY INSCRIBED BY

a
'

at
ro

CONTENTS.

. at .

= &

PAMPHIUITES =... ee OO OR

@aea2 38

r
“

Pet

3

_ PRESENT DISTRIBUTION OF BUTTERFLIES MOST NRARLY ALLIED TO FOSSIL SPECIES

-—Gexemat Résum, WITH NoTICE OF UNDETERMINED FORMS 3) wh cel arcs

Ea | ali, i el es

——
:

<..

.
i

: > - . _ 7 * . . * . . , ’ . . * * eon
* ¢ a oe
2. elem 2 Gs “o> Pr ee eee
aie : Be
. ne « ee ea Maes

- y ee =

. - ‘ . ° — * . s
Rap ae . ' Pee we in

oi . . . . ’ 4 * * ° . , — « * * - *

; 1% . * et a [a -
> r
~ ‘ Oe on Son Nt
4

: eee
4 7 . : xg .-~
: : ; ay Shere f + et ea ee at bi Ls

-
' ‘ = + eta
*
} X
’ ‘ £0) ae + one, Se be
* ard . y : Ar ees ek ave pera
- . . « ~ e & «8 =
> ?
i ea ‘ wee eee ote
a’
b . =
J
; . i. ee 2, |
»
ail - a « ae > oe ee Ter Se

hash d ‘ OF, i ~ 22 e eee Nee an a

~~ % a
a ae J inet ute : F Mose Sarr aA ot otal peel
4 _ % .
a j . i a ee a . ray eee wt 12) oe ee eT Ae Se
« , -
i : . » * = F é 7 ie Beast Te
ve 7
y
" *

INTRODUCTORY.

HE happy discovery in the Museum of Marseilles of a new fossil butterfly

first drew my special attention to this group of extinct insects, and

determined me to make, during my residence in Europe, a careful study of

the original types of all that had been previously described. By the great
courtesy of Count Saporta, Professor Heer, Dr. Reynés, Mr. Oustalet, Mr.
Woodward, the Rev. Mr. Brodie, Mr. Charlesworth, and the authorities of the
Jermyn street Museum, I was able to study not only all the originals of the
Museums of Aix, Marseilles, Zurich, Paris, London, Cambridge and Warwick,
but several new types, described here for the first time. As I was unable
to visit Vienna, Mr. Brunner de Wattenwyl was good enough to procure for
me new drawings, made under his immediate supervision, of the species from
Radoboj, described by Heer and preserved in the museums of that city. So that
I have either personally inspected all the fossils described within recent times as
butterflies, or have procured new and excellent original drawings of them, with
the exception of Heer’s Vanessa attavina (Sphinx atava Charp.), which I was
unable to find, and two fragments of slight value, viz.: the hind wing referred
by Heer to his Vanessa Pluto, and the portion of a hind wing, called Cyllonium
Hewitsonianum by Westwood. In the hope of drawing attention to fossil
butterflies, which have been hitherto so little studied, I have brought together
in this connection all that has been published of this group of fossils, whether
of text or illustration; presenting thus, within a small compass, a complete

account of our knowledge of these insects, as a basis for future investigations.

; : a a: 4 “sy
<eitorriay e Me
Ny! .

: <i cs i
= oie

ae: bead

> ear, =
» ah
-

vi
he i et

sae
. pe 2

>

a

BIBLIOGRAPHY.

1726. Hureser. Lithographize Wirceburgensis specimen primum. Fol. Wirceburg. This work con-
tains the first reference to fossil Lepidoptera which I have found. In his Synopsis
Tabellarum, he gives on page 94:

“Tabula XV. Similium insectorum alatorum Papilionum videlicet diversas species;” but the plates are
too rude to be of the slightest value or even to indicate the suborder to which the insects may
belong.

1729. Bromerz. Lithographia Suecana. Acta Litteraria Suecie, II. In a section de lapidibus insec-
tiferis Seanicis et Gothicis (p. 525) he says:

“«Praeter umbratiles etenim papilionum vel muscarum quasdam imagines, lapidi huic leviter sed distincte
impressas, multa scarabeorum figuras, mole totaque facie imitantur;” these were found in “saxo
fetido” in ‘ Westrogothia.”

In his enumeration of fossils he specifies further:

[528]. “*9. Papilionum majorum ac minorum imagines et impressiones nitide, in lapide calcario com-
muni inodoro, ubi etiam in alio foetido conspicue, ex eisdem Westrogothie locis.”

[529]. 10. Insectorum ovula, an nymphe seu aurelie lapidee? saxo fostido nigricanti immerse. Ex
eadem parecia karabylonga.”

(531). ‘14. Papilionum minorum imagines et impressiones, in ejusdem generis saxo suillo feetido. Ex
eodem loco. He itidem figura sua a papilionibus illis differre haud videntur, quarum superius Num.
9. meminimus.”

I find no later reference to these supposed Lepidoptera.

1742. Senpevivs. Historia succinorum. Fol. Lipsie.

Deyotes a chapter (De EZrucis, pp. 169-171) to supposed remains of caterpillars and chrysalides in amber.
Several forms are figured (pl. 5, figs. 25-28; pl. 6, figs. 1-4), of which it is not impossible that pl. 6, fig.
1, may represent a Papilionid larva; and pl. 6, fig. 4, the chrysalis of a Nymphalid; but the illustra-
tions are wholly insufficient to assert anything of them with confidence.

1828. Marcet pe Serres. Note sur les Arachnides et les Insectes fossiles et spécialement sur ceux
des terrains d’eau douce.’ Ann. Sc. Nat., XV, 98-108.

This is an extract only from the next citation.
@)

oe *

1829. Marcer pe Serres, Géognosie des terrains tertiaires ou Tableau des principaux animaux

rtp = ,
: ~ mg ae _
—— > Slee Mie a L's et) 4 ie! :
ell _ fe z cat <p —_ nen ~~ i
~ : ~ ; ~ We
. ae
r a Sa
' =
2 FOSSIL BUTTERFLIES.

invertébrés des terrains marins tertiaires du midi de la France. 16mo. Montpellier —
et Paris.

Contains a “Tableau des Arachnides et des Insectes fossiles du bassin tertiaire d’Aix (Bouches-du-
Rhéne),” printed in the preceding citation, in which (p. 230; p. 107 of preceding) occurs the genus
“Papilio,” with the remark: ‘Nous citons ici, sous la foi d’autrui, un Lépidoptére diurne de la
division des Satyrus,” doubtless referring to Neorinopis sepulta.

Speaking of the authors who have treated of the fossils of Giningen, he says: (p. 235) ‘*Ces divers natu-
ralistes y ont signalé des Scarabées, des Lucanus (p. 236) fort rapprochés du Lucanus cervus, des
Papillons,” etc.

In a “Tableau général des Arachnides et des Insectes fossiles” he gives on p. 257, the following:

Genres qui se trouvent dans les terrains anormaux.

Pe

Tertiaires. Secondaires.
2 an,
antérieurs a la retraite des mers
5 dans les couches de a
es ee ;
3 = cs]
Le rT peal
é & '
s " z z
Noms des genres. a 5
Papilio. % * * 4

In the “ marnes calcaires” of Aix he has referred already, as we have seen, to one; he previously speaks
of Papilions at CEningen (see above) and may therefore place two in the second column; he quotes
Sendelius as probably figuring caterpillars in amber as follows (p. 242): ‘‘ Des Lépidoptéres (M.
Brongniart). On acra reconnaitre des chenilles parmi Jes insectes du Sucein figurés par Sendelius
Tab, 3, fig. 28-82;”! and this accounts for one in his third column; and the following passage from
the section on “Insectes fossiles des terrains secondaires inférieurs, ou de transitions” (p. 246)
accounts for that in the fifth column: ‘Il se peut que ce soit également dans des formations de la
méme époque qu’existent les vestiges @insectes, Pailes de Papillons et de Scarabées signalés par
Bromel.” ;

1835. Gravennorst. Bericht der entomologischen Section. Uebers. a Arbeit u. Verand. Schlesisch.
Gesellsch. Vaterl. Cultur, 1854, 92-93.

Gives a general enumeration of the collection of fossils from amber in the museum of the K6nigsberg
Society, specifying a few Lepidoptera.

1 Probably an error for Tab. 5, fig. 28a, 28b, Which seems to represent a Tenthredinidous larya.

-

BIBLIOGRAPHY. 3

1836. Hore. Observations on Succinic Insects. Trans. Ent. Soc. Lond., I, iii, 133-147.

In a list of insects observed in amber we find the following on p. 146:

GENUS. AUTHOR. SUBSTANCE. COLLECTION,
“Papilio. | Hope and Berendt, | Animé and amber. | Mr. Strong.”

1838. Bronn. Lethea Geognostica, 2d ed., II. 8vo.

In a tabular list of fossil insects, with localities, he gives (p. 814):
Papilis [Papilio] (Bernstein), Satyrus (Gyps formation yon Aix).

1838. DuponcHet. Ann. Soc. Ent. France, VII, Bull. 51-52.

Re-announces the discovery of Neorinopis sepulta, referring it to Nymphalis.

1839. BorspuvaL. Ann. Soc. Ent. France, VIII, Bull., 11-12. °*

Gives a verbal report on the characteristics of Neorinopis sepulta, drawn from an inspection of a drawing
sent by Fonscolombe to Audouin, refers the insect to the genus Cyllo and says that the species is
allied to Europa and others.

1840. BorspuvaL. Rapport sur une empreinte de Lépidoptére trouvée dans les marnes des environs
d’ Aix, en Provence, et communiquée par M. de Saporta. Ann. Soc. Ent. France, IX,
371-374. Accompanied by a plate (viii) which appeared in the second livrasion. j

Describes Neorinopis sepulta from the specimen, referring it to the genus Cyllo, and the neighborhood
of the species Rohria, Caumas and Europa, and giving it the specific name sepulta.

1843. Marcet pe Serres. Notes géologiques sur la Provence. Actes Linn. Soc. Bord., XIII, 1-82;
Note additionelle, 83-90; Deuxidthe note additionelle, 170-2. 2 planches.

In a list of the plants and animals found at Aix, the author gives on p. 41: **Lépidoptéres Diurnes.
Papilio de la division des Satyrus. Cette espéce conserve encore en partie ses couleurs.” On p. 172
is a Note relative au Lépidoptére figuré (Cyllo sepulta), in which Boisduval’s opinion of its relationship
is given." The author's review of the plants and animals leads him to the generalization that they
are analogous to those which now live in dry and arid spots in the south of France. :

1843. CHarpentier. Ueber einige fossile Insecten aus Radoboj in Croatien. Acta Acad, Leop.
Carol., XX, 401-410.

Describes (p. 408) and figures (Tab. xxii, fig. 4) Eugonia atava under the name of Sphinx atavus.

1845. Cogquaxp. Bull. Soc. Geol. France [2], II, 384-386.
Refers to and quotes a portion of Boisduval’s description of Neorinopis sepulta ; nothing new is added,

1 The plate, however, is wanting, both in the copy belonging Boston Society of Natural History, so that I cannot tell whether
to the Smithsonian Institution and in that in the Library of the __ it is copied from Boisduval’s figure or is an original.

1845.

1847.

1849.

1849.

1850.

1851.

FOSSIL BUTTERFLIES.

Marcet pe Serres. Sur les fossiles du bassin d’Aix (Bouches-du-Rhéne). Ann. Se. Nat.
[3], IV, 249-256.

Uses the discovery of Neorinopis sepulta as an argument in support of his theory that there is an
intimate relation between the tertiary fauna and flora of Aix and the animals and plants now existing
in southern France; and that the climate of the two epochs was the same. Recalling the then recent
discovery of many butterflies new to the fauna of Europe, he suggests that N. sepulta may yet be
found alive.

'

Hore. Observations on the fossil insects of Aix in Provence, with descriptions and figures of
three species. Trans. Ent. Soc. Lond., IV, 250-255.

Gives a list of genera published by Bronn with some additions; on p. 252, under Lepidoptera, we have
«85, Satyrus B[ronn].”

Heer. Die Insektenfauna der Tertiargebilde von Giningen und von Radoboj in Croatien.

2" Theil. 4to. Leipzig. Extracted from the Neue Denkschr. allg. Schweiz. Gesellschaft
fir Naturw., XI (1850).

Contains (pp. 177-188, Taf. xiv, figs. 3-6) descriptions and illustrations of Eugonia atava (Vanessa
attavina), Mylothrites Pluto ( Vanessa Pluto) and Pontia Freyeri ( Pierites Freyeri).

Heer. Zur Geschichte der Insekten. Verhandl. Schweiz. naturf. Gesellsch., XXXIV, 78-97.

Refers to the late epoch at which Lepidoptera appeared, and adds, pp. 87-8: ‘‘ Merkwiirdig ist, dass von
diesen Schmetterlingen 2 Arten grosse Aehnlichkeit [88] mit ostindischen Arten haben, wihrend eine
mit unserm Distelfalter, eine andere mit unserem Grassacktriger zu vergleichen ist.”

Heer. Zur Geschichte der Insekten. Neues Jahrb. fiir Mineral., 17-33. :
On, the History of Insects. Quart. Journ. Geol. Soc. Lond., VI, ii, 68-76. Translated by

T. R[ymer] J[ones].

Essentially the samé as the preceding. The quotation given above is found on p. 24 of the Jahrbuch, p.
72 of the Journal. ‘‘Schmetterlinge” is everywhere translated Butterflies instead of Lepidoptera.
Aix in Provence is nearly always given as Aix-la-Chapelle.

Leresvre. Observations relatives 4 l’empreinte d’un Lépidoptére fossile (Cyllo sepulta) du
docteur Boisduval. Ann. Soc. Ent. France [2], IX, 71-88, pl. 3, No. II.

Criticises at length the opinion of Dr. Boisduval on the systematic position and structure of Neorinopis

sepulta, maintaining that the fore and not the hind wing was furnished with a tail, and while

confessing his inability to decide upon its relationship, inclines to the opinion that the insect was
more nearly allied to Vanessa. His studies were wholly taken from the plate published by Boisduyal.

Borspuvat. Quelques mots de réponse 4 M. Alex. Lefebvre sur ses observations relatives 4 la
Cyllo sepulta. Ann. Soc. Ent. France [2], IX, Bull. 96-98.

Defends his views against the criticisms of Lefebvre.

"a

1852.

1854.

1854,

1856.

1856.

1856.

1858.

1859.

1860.

BIBLIOGRAPHY. ; 5

Greset. Deutschland’s Petrefacten. p. 644. 8vo. Leipzig.

Catalogues the three butterflies described by Heer from Radoboj.

Westwoop. Contributions to Fossil Entomology. Quart. Journ. Geol. Soc. Lond., X, 378-96,
pl. 14-18.

Represents on pl. 17, fig. 17, and pl. 18, fig. 27, two fragments of wings, which he considers as belonging to
butterflies, and to which, on pp. 395-6, in the explanation of the plates, he gives the names of Cyllo-
nium Boisduvalianum and C. Hewitsonianum.

Picret. Traite de Palzontologie, II, pp. 392-393, pl. 40. 8vo. Paris.

Gives a brief account of the fossil butterflies then known, and reproduces excellently the figures of
Neorinopis sepulta, and Mylothrites Pluto given by Boisduval and Heer.

Gieset. Fauna der Vorwelt, II. pp. 185-7. 8vo. Leipzig.

Gives a similar but fuller account of the butterflies described by Heer and a brief notice of otlfers.

Gieset. Geologische Uebersicht der vorweltlichen Insekten. Zeitschr. gesammt. Naturw.,
VIII, pp. 174-188.

Gives lists of Lepidoptera summarized from his previous work.

Heer. Ueber die fossilen Insekten von Aix in der Provence. Vierteljahrsschr. naturf.
Gesellsch. Zurich, I, 1-40.

Simply mentions in his introductory remarks the occurrence of Neorinopis sepulta at Aix, and says that
most of the insects from this locality present a Mediterranean aspect.

Heer. Ueber die Insectfauna yon Radoboj. Bericht 32° Versamml. Deutsch. Naturf., 118-121.

A cursory review of Radoboj insects, mentioning the rarity of Lepidoptera, and specifying EZugonia atava
( Vanessa attavina) and Mylothrites Pluto (Vanessa Pluto). He remarks that the former resembles
V. cardui and probably fed on thistles, although these had not yet been found in a fossil condition
in that locality; and that the latter was nearly allied to Papilio. Hadena.

‘

Heyrpen. Fossile Insecten aus der Rheinischen Braunkohle. Dunk. u. Mey. Paleontogr., VIII,
1-15, Taf. 1-2.

Contains pp. 12-13, Taf. I, fig. 10, description and figure of Thanatites vetula (Vanessa vetula).

Heer. Untersuchungen iiber das Klima und die Vegetations Verhaltnisse des Tertiarlandes.
4to. Winterthur.

Refers to some of the fossil butterflies described from Radoboj and Aix.

MEMOIRS A. A. A. 8. 3

6 FOSSIL BUTTERFLIES.

1861. Heer. Recherches sur le climat et la Végétation du pays tertiaires ; traduction de Gaudin. 4to.
Winterthur.

The same as the previous; and also (on p. 205; not in the original edition) the following refer-
ence: ‘un cinguidme (Thaites Ruminiana) est tres voisin du genre Thais qui appartient A la faune
méditerranéene.”

1868. Burien. Catalogue of Diurnal Lepidoptera of the family Satyride in the collection of the
British Museum. 8vo. London.

Gives an appendix (pp. 189-190) on fossil species, in which he discusses the zoological position of
Neorinopis sepulta (Cyllo sepulta). ;

1869. Butter. Catalogue of Diurnal Lepidoptera described by Fabricius in the collection of the
British Museum. 8vo. London.

Discusses briefly (p. 109) the relationship of “Vanessa Pluto” to Argynnis Diana and Junonia Hedonia.

1872. Scupper. Description d’un nouveau papillon fossile (Satyrites Reynesii) trouvé & Aix en
Provence. Rev. et Mag. de Zool., 62-71, pl. 7. Also separate, pp. 7.

Description of a New Fossil Butterfly (Satyrites Reynesii) found at Aix in Provence. This is
a translation of a portion of my paper. Geol. Mag., IX, 532-533, pl. 13, figs. 2-3.
The same, separate, pp. 2.

Describes and figures Lethites Reynesii.

1872. Sarorta. Etudes sur la végétation du Sud Est de la France a l’epoque tertiaire. Suppl. I.
Révision de la flore des gypses d’Aix. 1 fascicule, Généralites. Ann. Sc. Nat. [5],
Bot. XV, 277-351.

Discusses (p. 342) the probable food of the caterpillars of Neorinopis sepulta and Thaites Ruminiana.

1873. Burter. On Fossil Butterflies. Lepidoptera Exotica, part xv, pp. 126-8, pl. 48.

On a Fossil Butterfly belonging to the family Nymphalide from the Stonesfield slate near
Oxford ; with notices of two other foreign forms from France and Croatia. Geol. Mag.,
X, No. ciii, 2-4, pl. 1.

Describes the genus Paleontina and species oolitica (a supposed fossil butterfly), refers Cyllo sepulta
Boisd. to a new genus, Neorinopis, and Vanessa Pluto Heer, doubtfully, to Junonia, adding remarks
upon the relationships of each.
1873. Axon. The oldest Fossil Butterfly in the World. The [London] Graphic. Feb. 22.

A popular account of the preceding paper, accompanied by a woodcut of Paleontina oolitica.

Gives a brief notice (pp. 8-9) of the various fossils referred to butterflies, especially of Palwontina
oolitica and Lethites Reynesii, and publishes an opinion expressed to him by me that the former was
Homopterous. nee

1874, Scupprr. Proc. Bost. Soc. Nat. Hist., XVI, 112. !

Doubts the lepidopterous character of Butler’s Palwontina, and refers it, probably, to the Cicadine.

1874. Borter. Notes on the impression of Paleontina oolitica in the Jermyn Street Museum. Geol.
Mag. [2], I, 446-449, pl. 19.

Defends the lepidopterous character of Palzontina and gives new illustrations of the same.

1874. Ssrrn. Discovery of Remains of Plants and Insects. Nature, XI, 88.

Enumerates fossils found at Gurnet Bay, and specifies among them “butterflies.”

‘

DESCRIPTIVE.

NYMPHALES— PRETORES—OREADES.
Genus NEORINOPIS Butter.

Neorinopis Butler, Lepid. Exot., i, 127 (1873) ;—Ib., Geol. Mag. x, 3.

In the shape of the wings (PI. I, fig. 8) this genus closely resembles Neorina
(Pl. II, fig. 13). The fore wings are arched and roundly produced at the apex,
though not so strongly as in Neorina, rather as in Antirrhza or Ceelites, the costal
margin is regularly, but not, as in Neorina, very strongly arched, and the apex is
well rounded; the outer border is sinuous and scarcely crenulate, the upper por-
tion, above the middle of the subcosto-median interspace, very strongly convex
and particularly prominent at the tip of the second inferior subcostal nervule;
below, the margin is again convex, starting from the middle of the upper median
interspace; at first (over one interspace) gently, afterward more fully, but still
rather broadly, to the well rounded lower angle; the inner margin is slightly
concave. The hind wings resemble those of Neorina far more than those of
any other genus, but are long and proportionally rather more produced than in
Neorina, with less crenation of the outer border, and a shorter and slenderer
tail; the costal margin is strongly and abruptly convex next the base, but beyond
this passes with a regular and gentle convexity to the outer angle, which is
larger than a right angle and somewhat rounded off; above the tail the general
trend of the, outer border forms scarcely more than a right angle with the gen-
eral course of the costal margin and is gently crenate; the tail, which lengthens
the upper median neryule by about one-fourth, is about the width of an interspace

at the base and tapers to a rounded point, at first rapidly, afterward slightly; the
(9)

10 FOSSIL BUTTERFLIES.

border is slightly angulated at the tip of the middle median nervule, and still
more strongly at the tip of the lowest median nervule, causing in the latter a
very broad angular projection, beyond which the margin slopes off and is rounded
at the angle. The inner margin has a very broad and extensive basal pro-
jection, and the course of the internal nervure renders it probable that it was
even more extensive than represented in the plate; it reaches more than half-way
along the inner border, and at the broadest exceeds the cell in width; be-
yond it the inner margin has a nearly straight course, parallel and adjacent
to the sub-median nervure.

As to the neuration (Pl. I, fig. 9) this genus approaches more closely the
genera Zophoessa (Pl. Il, fig. 1), Neorina (Pl. II, fig. 8), Debis (Pl. I, fig.
10), and Lethe (Pl. U, fig. 6), than any others, although it differs from any of
them more than they do among themselves. The most noticeable marks of dis-
tinction are these: in the fossil genus the first superior subcostal nervule of
the fore wing is thrown off just at the extremity of the cell while the second
and third are far beyond it; in the recent genera the first neryule is always
emitted some distance before the tip of the cell and the second either at or
before the extremity; in agreement with this, the cell is much shorter in Neori-
nopis than in the others, being but two-fifths the length of the wing, while
in the others it is about one-half its length; in Neorinopis the nervule closing
the cell of the fore wing unites with the median nervure at its last divarica-
tion, while in the others it strikes it a long distance beyond. In the hind
wing the vein closing the cell strikes the median at its last divarication, as in
Zophoessa, while in the others it meets the last branch of that vein at a slight
distance from its origin.

In the fore wings the costal nervure terminates at a little distance beyond
the middle of the costal border. The subcostal terminates, as in the recent
genera mentioned, near the tip of the wing, and has four superior and two
inferior branches; the four superior nervules and the costal nervure terminate at
nearly equal distances apart on the costal border; the first superior nervule is
emitted from the very tip of the upper border of the cell, at two-fifths the distance

— or”

NEORINOPIS. 11

from the base to the apex of the wing, the second beyond the cell, but scarcely
beyond the middle of the wing; the third at a less distance from the base of
the second than that is from the first, and directly below a point midway
between the tip of the costal nervure and that of the first superior subcostal
neryule; the fourth near the extremity of the wing and but little before the tip
of the third superior nervule, or at about two-thirds the distance from the base
of the third superior subcostal nervule to the tip of the subcostal nervure; the
first inferior subcostal nervule originates of course at the tip of the cell, and
separates but narrowly from the main stem, from which it diverges very gradu-
ally as far as the base of the outer superior nervule, where the main stem ap-
proaches it again; the lowermost inferior subcostal nervule arises from the first
inferior scarcely beyond its base, curves inward, downward and then outward be-
fore taking a course parallel to the nervule above, from which it is separated at
its base by twice the distance that the former is there distant from the sub-
costal neryure; the vein closing the cell can scarcely be called a vein, but
rather a break in the membrane such as is often seen in recent butterflies, and
is indicated in the fossil by a curving granulated streak; it arises from the
final curve of the lowermost inferior subcostal nervule opposite and directly
below its origin; it passes thence in a slightly curved line, opening outward,
to the very base of the upper branch of the median nervure. The median ner-
vure runs in a straight line as far as its first divarication, which is a little
beyond the middle of the cell; thence it is bent parallel to the subcostal ner-
yure and exactly at the lower tip of the cell forks, the branches parting but
gradually from each other, the upper gently curved, the lower nearly straight.
The submedian nervure is parallel to the lowest median nervule, as in Neorina,
ete. None of the veins are swollen at the base. The cell is three and a half
times longer than broad.

In the hind wing the neuration is almost precisely that of Neorina Lowii (Pl.
Il, fig. 8). The costal and subcostal veins are confluent for a short distance, when
the costal parts from its neighbor at nearly right angles and immediately

thereafter sends up the basal shoot, which, after passing in a straight line half

12 FOSSIL BUTTERFLIES.

way toward the basal angle of the costal margin, curves slightly outward and
fades away; the costal nervure, on approaching the border, curves outward and
meets the border near the middle of its outer two-thirds; the subcostal breaks
into three branches, exactly as in Zophoessa. The median nervure and its middle
branch form a continuous, almost exactly straight line, from which the lowermost
branch parts opposite the union of the vein closing the cell with the lowest
subcostal nervule; and the uppermost at exactly the tip of the cell, or as far
beyond the origin of the lowest nervule ‘as the upper limit of the vein closing
the cell is from the base of the upper subcostal nervule; the vein closing the
cell is a very weak one and originates on the lowest subcostal nervule, as far
from the second divarication of the subcostal nervure as that is from the first,
and passes in a gentle curve, opening outward, to the second divarication of the
median nervure. The submedian and internal nervures are united for a short
distance beyond the base of the cell; the submedian passes with a gentle regular
curve to the outer border, at the lower outer angle; the internal parts from this
with an opposing curve and terminates somewhere below the middle of the inner
flap of the wing, probably approaching again the submedian nervure near its
extremity. None of the veins are swollen at the base. The cell is two and
three-quarters times longer than broad.

In the disposition of its markings (PI. I, fig. 8) this genus does not seem to
show any strong affinity with any living butterflies, although it has some features
in common with the genera already referred to (Pl. II, figs. 3, 9, 11, 13, 14).
The base of the wing is dark, followed by paler spots and bands, differing greatly
in the front and hind wings, followed again by a belt of dusky scales, which
separates from the rest of the wing a paler submarginal band, enclosing roundish,
interspaceal, often pupillated spots of varying size, and whose outer limits are at
least an interspace’s distance from the outer border; the latter is margined, on
the hind wings, with alternating darker and lighter lines. The middle portions
of the two wings differ; the hind wings have simply a broad pale field, gradually
merging on either side into the darker parts and varied by a cloudy, wavy,

narrow, transverse belt near the middle; the fore wing, on the other hand, is

NEORINOPIS. 13

marked by two large diagonal light patches, whose interior edges are well defined,
but whose exterior are powdered at their confluence with the darker parts; one
of these patches crosses the subcostal interspaces at a little distance beyond the
cell, and reaches from the subcostal to the median nervure; the other crosses
the middle of the outer half of the cell and covers a great part of the basal half
of the lower median interspace; while a third roundish patch, united with it,
occurs near the middle of the medio-submedian interspace. The two diagonal
patches have their inner distinct edges nearly parallel and straight, following lines
which run at nearly right angles to the costal margin; in this respect they agree
with the diagonal disposition of markings upon the upper and under surface of
some species of Zophoessa (PI. IT, figs. 3, 11) and Lethe (PI. IT, fig. 9), while the
nature of the broad patches themselves may best be compared to such masses of
color as we see in Neorina Lowii (Pl. I, fig. 13) and some other species; the
marginal markings of the hind wings and the submarginal spots are common to
very many Oreades, but the nature and disposition of those of Neorinopis and the
disparity of their character. on the two wings are best seen on a comparison with
the types we have already alluded to, and which are represented on the plates.
The small round pale spots accompanying larger dark ones on the fore wing may
be seen in Weorina Lowi, though the relation of the two is different from what we
see in Neorinopis, while the greater importance of the ocellus in the lower median
interspace of the hind wings finds an exaggerated counterpart in Neorina Lowit.
In general, the design of the fore wings approaches that of Neorina Lowi more
nearly than that of the upper surface of any other butterfly I have seen, although
there is a distant resemblance to the markings of Antirrhzea and Anchiphlebia, as
Butler has remarked, as well as toward certain species of Zophoessa. ‘The mark-
ings of the upper and under surface of butterflies have nearly always some and
often a close relation to each other, and therefore we may reasonably look at the
under surface of living insects to find the nearest counterpart to our fossil; in this
respect the under surface of Lethe Dyrta (P1.II, fig. 9) may well be studied, where
in a lighter submarginal band we find a series ofspots, in the principal interspaces,
far from the border; these are ocellated instead of double as in Neorinopis; there

MEMOIRS A. A. A. 8. 4

—— eT a eS a ee ee rn)

14 FOSSIL BUTTERFLIES.

are two large patches of pale color in the upper half of the wing as in Neorinopis,
but the inner is much obscured by a dark bar crossing the middle; and the outer in-
stead of the inner patch is connected with the lighter parts of the lower half of the
wing, and is separated from the parts within by a long line whose general course is
at right angles to the costal border; in the markings of the hind wings it is by no
means unlike Zophoessa Sura (Pl. I, fig. 3), and resembles less conspicuously
Debis Sinorizx (Pl. Tl, fig. 14), with which also it agrees admirably in the form
and neuration of the wing; in the shape of the tail particularly, and in the size of —
the insect also, Neorinopis agrees better with Debis Sinorix than with any but-
terfly I have been able to examine. In neuration and in markings, although not
at all in the form of the wings, this fossil shows no distant alliance to our own
Enodia Portlandia.

The other parts of the body are not sufficiently preserved to admit of their
use in generic description, if we except the hind legs; these are slender, the tarsi
(which are barely shorter than the thorax) being of the same length as the tibize
and a very little longer than the femora.

NEORINOPIS SEPULTA (BoispuvaL) But Ler.
Plate I, figs. 8-17.

Nymphalis sp. Dup., Bull. Soc. Ent. France, 1838, 51-52.
Cyllo sp. Borsp., Bull. Soc. Ent. France, 1839, 11-12.

Cyllo sepulta Borsp., Ann. Soc. Ent. France, ix, 871-374, pl. viii (1840); Is., Bull. Soc. Ent. France,
1851, 96-98; Serres, Act. Linn. Soc. Bord., xiii, 172, pl. ii (1843); Wesrw., Gen. Diurn. Lep.,
861 (1851); Ler., Ann. Soc. Ent. France [2], ix, 71-88, pl. iii, If (1851); Pricr., Traite Pal., ii, 393,
pl. xl, fig. 11, 1854; Burs., Cat. Satyr. Brit. Mus., 189-190 (1868 ).

Antirrhea? sepulta Kirs., Syn. Cat. Diurn. Lep., 39 (1871).

Neorinopis sepulia Butt., Lep. Exot., 127, pl. xlviii, fig. 3 (1873); In., Geol. Mag., x, 3, pl. i, fig.
8 (1878).

The earliest notice of this fossil butterfly, the first species ever described
and illustrated, the most perfectly preserved and the best known to the world at
large, was given by Marcel de Serres in 1828, in the Annales des Sciences
Naturelles; and in 1829 in his Géognosie des terrains tertiaires; where he simply
cites on the authority of some one else the occurrence in the beds of Aix of
a butterfly belonging to “la division des Satyrus.”

NEORINOPIS SEPULTA. 15

The earliest definite mention of the insect is given by Duponchel in the
Bulletin of the Entomological Society of France, as follows :1

“M. Duponchel entretient ensuite la Société d’un fait extraordinaire, et peut-
étre enti¢rement nouveau dans les annales de [52] la science: c’est l’existence d’une
impression trés remarquable de Lépidoptére fossile, qui a été trouvée dans une
platriére des environs d’Aix (en Provence), et acquise par M. de Saporta. Ce
Lépidoptére, suivant M. de Saporta, parait appartenir au genre Vymphale, et a une
espéce étrangére a celles qui vivent aujourd’hui en Europe. Le corselet en est
parfaitement conservé; les couleurs des ailes sont trés-bien indiquées; le dessin de
ces ailes est enti¢rement reconnaissable. Les deux ailes d’un des cédtés du corps
sont repliées en grande partie l’une sur l’autre; la place du ventre est trés distincte ;
Tautre cété manque tout-d-fait.”

The subject seems to have been referred to Dr. Boisduval, for we find in
the following year® that

*M. Boisduval rend un compte verbal du rapport que la Société avait chargé
de faire, sur un dessin envoyé a M. Audouin, par M. de Fonscolombe, et qui repré-
sente une empreinte de lépidoptére fossile trouvée dans les environs d’Aix. M. Bois-
duval déclare, qu’ aprés un examen attentif, il a reconnu que ce lépidoptére devait
appartenir 4 son genre Cyllo, et qu’ il se rapprochait beaucoup des espéces décrites
par les auteurs sous les noms de Satyrus Europa, Caumax, Rhosia et plusieurs
autres lépidoptéres indiens. Le méme membre ajoute que ce Rhopalocére ne peut se
rapporter exactement 4 aucune des espéces vivantes déja connues. Toutefois,
avant de décider si cet insecte doit étre regardé comme un véritable fossile, M. Bois-
duval pense qu’il serait indispensable, que la Société pit avoir sous les yeux [12]
la pierre qui a servi de modéle au dessin envoyé par M. de Fonscolombe.”

The next year a very fair illustration of the insect, reproduced in our
Plate I, fig. 17, was given, and shortly afterward a written report upon the
subject by Dr. Boisduval, in which he furnishes, not only his views upon its
affinities, but a brief historical account of the insect, which is giyen below :*

“Tl y a bient6t un an que je fus chargé par la Société d’examiner le dessin d’un
Lépidoptére fossile trouvé dans les platriéres des environs d’Aix en Provence, et
appartenant 4 M. le Comte de Saporta. Au premier coup d’ceil, ce dessin me
parut deyoir étre rapporté 4 une espéce de Saryrmes du genre Cyllo, 4 cété des
Satyrus Rohria, Caumas et Luropa, de YEncyclopédie; mais la découverte @’un

1 Bull. Soc. Ent. France, 1838, 51-52. 8 Annales Soc. Ent. France, ix, 371-374 pl. 8.
2 Bull. Soc. Ent. France, 1839, 11-12.

16 FOSSIL BUTTERFLIES.

Lépidoptére fossile me sembla un fait tellement neuf, et l’espéce si rapprochée de
celles connues, que je n’osai pas faire de rapport avant d’avoir yu la pierre en
nature. La Société partagea cet avis, et engagea M. Duponchel a écrire a M. de
Fonscolombe pour lui faire part du doute de quelques membres sur l’authenticité
de cette empreinte. Ce fut alors que M. le comte de Saporta, naturaliste fort dis-
tingué et propriétaire du fossile en question, m’écrivit la lettre que jai commu-
niquée a la Société; lettre dans laquelle il prétendait qu’il n’avait pu étre victime
de la supercherie de qui que ce soit, et [372] que par conséquent il n’y avait pas
lieu 4 conserver le moindre doute sur l’exactitude scrupuleuse du dessin commu-
niqué par son beau-pére, M. le Baron de Fonscolombe; qu’on pouvait voir @ail-
leurs au Muséum un Polyommate fossile qu’il avait envoyé depuis plusieurs années
avec des empreintes d’insectes de différents ordres.

Cependant la Société emit de nouveau le désir de connaitre en nature ce lépi-
doptére fossile. M. Duponchel écrivit une seconde fois 4 M. de Fonscolombe: ce
fut alors que M. le comte de Saporta consentit 4 se dessaisir pour quelques jours
de ce précieux échantillon en nous ’envoyant en communication.

Le morceau de calcaire qui porte reéllement Pempreinte parfaite Vun lépidop-
tere conforme au dessin de M. Fonscolombe, est un fragment assez volumineux de
marne gypseuse bituminifére, telle qu’on en rencontre dans une grande partie des
environs d’Aix en Provence. . . satiate

Le Lépidoptére, qui fait le ase le ce S vanpoet fait arte cas [373] Ay ces
genres dont les espéces assez peu nombreuses sont confinées aujourd’hui dans
les files de V’archipel indien ou dans les contrées les plus chaudes du continent
asiatique. D’aprés ce que j’ai pu apprendre de M. Blum de Leyde, ils voltigent
ci et li a Ventour des palmiers, dont peut-étre ils se nourrissent 4 Vetat de
chenille. *

L’individu communiqué par M. de Saporta, et que nous avons nommé
SEPULTA, pour rappeler son origine antédiluvienne, appartient au genre Cyllo,
et se rapproche de Rohria, Caumas et autres espéces voisines; mais il ne peut
étre rapporté 4 aucune de celles conhues de nos’ jours, ce qui est d’autant plus
vraisemblable, que les marnes schisteuses sont de beaucoup plus anciennes que
la derniére catastrophe diluvienne admise par tous les géologues.

Le dessin et la forme de cet insecte sont si bien conservés, que l’on croi-
rait qu'il a été lithographie sur un schiste; seulement il n’existe que le cété
droit, lequel est parfaitement intact, une portion du corselet et une légére em-
preinte de l'abdomen. L/aile supérieure est en grande partie cachéé par Vin-
férieure, et il est impossible de dire si elle offre @autre dessin qu’un cil apical
surmonté d’un point blane; autre, dont on voit toute la surface, est d’une couleur
gris brunitre, comme dans les espéces voisines, avec une tache costale blanche, une
bande transverse, médiane, sinuée, de la méme couleur, suivie de deux yeux noirs

NEORINOPIS SEPULTA. 17

encadrés de blanc, s’alignant extérieurement avec deux points blancs. L’extrémité
de cette méme aile est un peu plus pale, presque blanchitre, et divisée, comme chez
la plupart des espéces vivantes, par deux lignes marginales brunes, paralléles.
L’appendice caudal est un peu plus long que dans Rohria, mais situé de la méme
manicre.

M. le comte de Saporta a émis plusieurs opinions géologiques sur la cause qui
a produit les empreintes d’insectes dans les terrains des environs d’Aix. .

[374] Il admet . . que ces marnes ont été formées couches par couches, ou
plutét feuillets par feuillets, par des dépéts fluviatiles. . . . . . . Selon cer-
taines circonstances, les différentes couches ont varié de couleur, comme on peut
s’en conyaincre par l’échantillon que la société a eu sous les yeux. Les plus infé-
rieures sont colorées par du bitume et des oxydes métalliques; celle ot se trouve le
Lépidoptére est blanche et presque pure, ce qui permet de distinguer le dessin et
probablement la véritable couleur du papillon tel qu’il était avec son incrustation.”

The plate accompanying the Report of Dr. Boisduval has been several times
copied,’ and his statements reproduced in part or referred to, as will be seen by
the Bibliography at the commencement of this essay. But the most extraordinary
of all is an acute criticism by Lefebvre, eleven years subsequently, of which I give
the following extracts,’ from a copy of the paper in my possession slightly cor-
rected by the author.

“ [72] Si de l’ceil on suit les bords de la seconde aile [ PI. I, fig. 17] qu’avec
le docteur je reconnais couvrir en grande partie la premiére, je trouve qu’elle est,
cette seconde aile, totalement arrondie dans ses contours, et je ne peux concevoir
par quelle aberration d’optique il lui a vu la moindre analogie avec la seconde aile
dune de nos Cyllo; comment il lui attribue un appendice caudal, propre volontiers
aux [73] espéces de ce groupe, et qui, selon lui, la termine a la maniére de ceux de
la Cyllo Rohria de Fab. (Voy. f. A [PI. I, fig. 14]).

Pour parler ainsi que le fait M. Boisduval de cet appendice, il faut nécessaire-
ment qu’il ait confondu avec cette seconde aile le dernier contour de la. premiére,
qui l’excéde a partir de leur point de jonction sur le bord externe, lui attribuant
comme appendice caudal cette forte dent de la premiére aile, qui succéde 4 une
forte échancrure, ainsi qu’il en existe dans tant de Vanessides, et qui le plus
souvent y est soutenue par la troisiéme inférieure, ainsi que tout a VPheure elle l’y
sera pour nous dans la Sepulta.

J’ayoue donc que je ne puis, avec le meilleur vouloir, envisager cette em-

1 By Marcel de Serres, Actes Linn. Soc. Bord., Vol. xiii, pl. 2; 2 See particularly papers by Coquand, Marcel de Serres, Heer,

Pictet, Traite de Palaeont., LI pl. 40, fig. 11; Butler, Lep. Exot. Saporta, Giebel, Westwood and Butler.
I, pl. 48, fig. 2; Ib., Geol. Mag. x, pl. 1, fig. 3. % Ann. Soc, Ent. France (2) ix, 71-88 pl., 3, ii.

18 FOSSIL BUTTERFLIES.

preinte autrement que je ne le fais, et que pour la considérer sous le méme point de
vue que notre docte confrére, il me faut faire trop violence 4 mes pauvres yeux . . ;
il me faut enfin donner un démenti aux contours si bien écrits de ces deux ailes
superposées. : ;

Et, en effet, ne voit-on pas se dessiner les bords de la premiére aile dans tous
leurs contours; la transparence de la seconde, avant son angle anal, ne permet-elle
pas de suivre encore le bord inférieur de la premiére, qui est un peu falqué et qui,
dans l’empreinte, passe précisément sous l’articulation femoro-tibiale de Punique
patte postérieure qui existe encore?

Toute la seconde aile ne vient-elle pas de ses bords nettement tranchés, et
surtout dans le bord extérieur, couper toute la premiére aile sur laquelle elle est
appliquée? A partir du point le plus proche de ce méme bord [74] avec celui de la
premiére aile, et presque au centre de sa forte échancrure, ne s’en détache-t-elle
pas, comme au-dessus, par une marge obscure et trés nettement tracée? Enfin,
cette méme aile ne se continue-t-elle pas seule et détachée sur le fond de la pierre,
avec ses méplats voulus dans les bords postérieur et abdominal, jusqu’au-dessus
du fémur de la patte déja citée?

Je ne crois pas qu’on me puisse répondre par la négative, tant les faits sont
patents.

Cet examen nous donne donc pour résultat:

1° Une aile de dessus fortement dentée et échancrée en dehors, 4 son bord
extérieur. (Voy. fig. B [ Pl. I, fig. 16]).

2° Une aile de dessous, simple, arrondie, et sans vestige d’appendice caudal.

Si c’est chose convenue, qu’en déduire? Si ce n’est que par cette seule confor-
mation, nous sommes actuellement en droit de décliner déja toute espéce d’analo-
gie entre la Sepulta et le genre Cyllo, proprement dit, et de ’éloigner des Caumus,
Beroe, Rohria, et autres; et cela, d’abord, par la rondeur inerme de la seconde aile,
et ensuite a la premiére, par cette forte échancrure, suivie dune dent non moins
énorme que soutient la troisiéme inférieure, caractéres que n’offrent guéres les
Satyrides de cette section, et oi Ja dent la plus proéminente du bord extérieur,
comme 4 Banksia God., se prononce a l’extrémité de la premiére supérieure, quand
il en existe une.

Je ne connais que des Vanessides qui puissent présenter en méme temps des
premiéres ailes déchirées de cette maniére a leur bord extérieur, et des secondes
ailes arrondies et sans dentelures. La Van. Archesia, Cr. pourrait, entre autres,
nous en offrirun example. Et cependant chez les Vanessides, lorsque les premiéres
ailes y sont ainsi dentelées et découpées, les secondes le sont égale-[75] ment,
plus ou moins, par la régle assez générale qui veut que chez les Lépidopteres les
secondes ailes y soient toujours plutét munies de dentelures que les premicres.

Voici donc, pour la forme des ailes, un argument en fayeur de mon opinion.
Passons aux dessins.

NEORINOPIS SEPULTA. 19

Avant que d’assayer de les réhabiliter dans cette espéce, il me faut décider
une autre question, a savoir si ces dessins appartiennent a la premiére aile ou a
la seconde. Notre confrére les tient pour étre propres 4 cette derniére. Je ne
suis pas de son avis, et voicé pourquoi:

Je pense que la seconde aile est en grande partie dénudée de ces écailles 4
sa face inférieure, celle que nous voyons.

Ce qui me le fait croire, c’est que déja dans sa marge abdominale, ainsi que je
Yai déja dit, on suit 4 travers la membrane le contour intérieur de la premiére aile,
et dune maniére trop distincte pour admettre que l’adhésion des deux ailes le piit
permettre, si les deux faces de la seconde étaient revétues de leurs écailles.

Ce qui me le fait croire encore, c’est qu’da cette seconde aile, la petite lunule
blanche de VYangle externe (fig. B [ PI. I, fig. 16]), qui est située sur le bord lui-
méme, et qui y est extérieurement coupée par lui, ne saurait devoir y exister a
cette place, si on en juge par la loi suivie dans leur position normale parmi la
majeure partie des Diurnes. En effet, 4 aucun, ou 4 bien peu du moins, je ne
connais pas 4 cet angle de lunule extréme, ainsi placée sur le bord lui-méme des
secondes ailes, et dans cette position, rejetée en arricre de celle qui la précéde.

Régle assez générale, la série marginale de taches lunulaires ou autres, pupil-
lées ou non, qui affectent ces ailes, est d’habitude concentrique a leur base, et la
lunule en [76] question serait sur cette seconde aile placée contre cette régle.

A examiner cette aile dans la fig. B [Pl. I, fig. 16], on comprend de suite
que cette lunule n’y est pas 4 sa place normale; elle choque méme 1a ow elle est
située, tandis que si je la reporte (sans la bouger, bien entendu) sous la premiére
aile (ainsi que je le fais a la fig. C [PI. I, fig. 15]), elle s’y adapte tout natu-
rellement dans lordre que lui est le plus rationnel avec les autres.

Par ce fait, i la place qu’occupe cette lunule, la seconde aile serait done en-
core transparente? Observons en passant que dans les espéces oti une semblable
lunule ou tache oculaire, se remarque en degssous, vers langle externe des deux
ailes (comme 4 Melanitis Undularis, Dr.; Protogenia, Cr., par ex.), cette tache
qui est toujours placée un peu avant la marge, qui ne Pinterrompt jamais comme
ici, est toujours (comme ici, du reste) entre les deux derniéres supérieures, et non
entre la derniére supérieure et la costale.

Sil est des exceptions 4 cette régle, elles ne sauraient étre qu’en bien petit
nombre, et lorsque les lunules marginales y sont présentes en nombre considéra-
ble; mais sil n’y en a plus qu’une ou deux, celle de Vangle externe sera placée
ainsi que je viens de la dire, et non ailleurs.

Toujours 4 l’appui de cette transparence, que j’attribue 4 la seconde aile de la
Sepulta, si jinterroge le peu de la charpente alaire qu’on y distingue, et qui est
suffisant pour la restituer telle qu’elle devait étre, ou 4 bien peu de chose prés
(comme 4 la fig. C [PI. I, fig. 15]), on voit que la tache semiorbiculaire et obscure

20 FOSSIL BUTTERFLIES.

de langle interne y semble partagée par une nervule de la seconde aile trés bien
écrite, par la deuxiéme inférieure. Or, l'étude de cette partie de la [77] ptéro-
logie, qui 4 pour objet les lois relatives 4 la position, 4 la forme, comme 4 la
présence et a l’absence des dessins et des taches, nous apprend que jamais une
lunule ou une tache orbiculaire marginale n’est divisée par une neryule, mais
plutét par le pli internervulaire, les nervules séparant d’habitude ces sortes de
taches, et ne les scindant pas.

Par induction, je dirai done que cette tache orbiculaire n’est pas encore ici d sa
vraie place sous la seconde aile. Mais si je l’attribue a la premiére, ainsi que tout
i Pheure je lai fait pour la petite lunule, 4 son tour elle s’y adapte marveilleusement
bien (fig. C [ Pl. I, fig. 15]), entre la troisiéme inférieure et la sous-médiane, et
en plus, son rejet en dehors, qui nous choquait il y a un instant, n’a actuellement
rien que d’assez normal.

Allant plus loin, si la grande tache orbiculaire, fort noire, qui la surmonte, et
qui a la place qu’elle occupe sous la seconde aile peut y exister sans discussion,
ainsi que l’autre petite lunule blanche qui se voit au-dessus, sont reportées a la
premiére aile (fig. C [ Pl. I, fig. 15]), elles viennent y compléter cet ensemble, qui
parait alors fort rationnel, des plus habituels, et dont au besoin nous trouverions un
exemple dans la Van, Alcithoe, Cr., ete.

Et ici, il n’y a pas a s’y tromper les nervures encore existantes a cette
seconde aile, sont bien représentées a leur place voulue, selon les lois de la So-
lénoptérologie.

[78] Or, si la nervule dont s’agit (la deuxiéme inférieure) est 4 sa place nor-
male, la tache orbiculaire qu’elle divise n’y est pas. Done, elle doit appartenir
forcément a l’autre aile.

Puisque nous voici fixés sur la position plus que probable de ces deux autres
taches de la seconde aile, convenons que pour les y maintenir il faudrait que cette
aile efit précisément conservé ses écailles a cette place. C’est chose possible, mais
chose peu probable.

D’aprés ce qui précéde, je suis done porté a croire, comme je Vai déja
avancé, 4 la dénudation presque compléte du dessous de cette seconde aile,
et que l’action des eaux sédimenteuses qui a agi sur cette face, vu l’adhérence
de toutes les écailles 4 autre éclat de cette marne qui nous est inconnue, n’a
pu atteindre les portions de la premicre aile qu’elle abrite.

En plus, par lanalogie et le faciés de la Sepulta, ayant tout a de pen-
ser que le dessus de toutes ses ailes deyait étre d’un brun sombre, uni et privé
de tout dessin tranché, ou varié de vives couleurs, par cela méme, j’en induis
[79] que la surface supérieure de la seconde aile n’a pu empécher les dessins
qu’elle recouvrait de paraitre, sans confusion aucune, a travers la couche uni-
colore des écailles du dessus, généralement trés fines dans les Satyrides. Leur

NEORINOPIS SEPULTA. 21

adhérence intime 4 la surface inférieure de la premiére aile aura méme di
augmenter la transparence de la seconde.

Mais avec assez de raison, on pourrait me demander 4 mon tour, par quel
privilége, ce qui reste de non recouvert de la premiére aile n’a pas été altéré
par ce méme frottement, ou plutét par son impression sur Véclat qui a mis a
jour cette empreinte? De cet argument ad hominem, je ne pourrais me tirer
je Vavoue, qu’en arguant que nous ne yoyons que par transparence les taches
et dessins, fort admissibles, de la face supérieure.

Par ce que je vais ajouter encore, on pourrait en déduire que selon le
besoin que j'ai de la dénudation, ou de Vintactum des écailles du dessous de
cette deuxiéme aile, je les admets ou les repousse pour mieux soutenir lopinion
SS 0 a a ce

Il est de fait que par la marge obscure de la seconde aile qui se découpe si
nettement sur la premiére, je suis forcé de reconnaitre que les écailles de ses
bords ont di y étre plus respectées, peut-étre, qu’ailleurs, pour nous apparaitre
encore avec une pareille vigueur; mais peut-étre aussi la concordance d’une
semblable marge en dessus, et qui n’aurait rien que de normal, concourt ainsi
i [80] la rendre aussi visiblement nette que nous la voyons aujourd’hui?

De toute maniére, il est impossible de l’admettre comme dessin apparte-
nant au dessous de la premiére aile, ainsi qu’a di le comprendre M. Bois-
duval, par une erreur d’optique, que déja sans doute il a reconnu lui-méme.

L’absence bien regrettable de Véclat qui recouvrait cette Sepulta est cause
de tant d’incertitude, car je ne mets pas en doute qu'il devait conserver, 4
son tour, la majeure partie des écailles de toutes ces ailes, avec lesquelles il
était en contact.

Tant bien que mal, nous voici donc édifiés sur la portion extérieure de
ces ailes. Continuons cet examen en marchant vers leur origine.

Je reprends Je dessin original.

Aprés cette série de taches marginales, il existe sur la céte elle-méme,
avant langle externe de la seconde aile, une large éclaircie blanche, quelque
peu oyalaire, nettement dessinée en dedans, et posée sur la place qu’d la pre-
miére aile doit occuper la disco-cellulaire et le commencement des deux premiéres
supérieures.

La position de cette tache blanche a la seconde aile n’a rien de réfutable,
non plus que celle trés obscure qui lui succéde, puis autre tache blanche, et
enfin la masse obscure qui couvre toute la base.

Ces dessins maculaires peuvent, 4 la rigueur, y exister, comme n’y pas étre,
de méme qu’ils ne sont guére acceptables 4 leur autre surface; car ce que nous
yoyons est bien un dessous d’aile et non un dessus.

Les dentelures externes de la tache basale, sont en dessous des plus natu-

MEMOIRS A. A. A. 8. 5

22 FOSSIL BUTTERFLIES.

relles, et dans nos Satyrides, dans [81] nos Vanessides, nous en retrouvons
de nombreux exemples.

Mais un instant, ne nous pressons pas de juger: examinons attentivement
Yoriginal: qu’y voyons-nous?

Déja, sur le bord costal de la premiére aile, nous aperceyons en effet, sur
notre gauche, un commencement de cette blanche éclaircie qui suecéde aux lu-
nules, et 4 notre droite, le bord intérieur de cette éclaircie y est des plus évi-
dent! (Voy. fig. B [ PI. I, fig. 16] ).

Comment done se fait-il que ces ailes ainsi ployées, ces vestiges de la pre-
miére aile viennent s’adapter d’une maniére si compléte avec toute la portion
blanche qui se continue sous la seconde aile? cela se peut rencontrer, je
Pavoue, mais c’est peu fréquent. ,

Bien mieux, le large sommet de la tache blanche anguleuse et obscure qui
lui succéde, se voit aussi sur le bord un peu diffus de la céte, dans la partie
externe, et se relie également bien avec celle que la seconde aile nous laisse,
selon moi, apercevoir. Plus loin encore, le commencement de la grande tache ba-
sale, hachée a son dehors, ne se continue-t-il pas sur la céte de la premiére aile?

Enfin, si cette derni¢re tache appartenait 4 la seconde viendrait-elle, ainsi
quelle le fait, s’arréter précis¢ément sur le bord intérieur de la premiére, que
par transparence nous pouvons suivre parfaitement a partir du moment ow il
est recouvert par la seconde aile? En Ilattribuant a cette derniére, ce serait
agir contre toute apparence plausible, contre toute disposition naturelle de ces
sortes de taches, et venir ’interrompre bénévolement et sans motifs spécieux, bien
avant langle anal de la seconde aile, sur lequel elle devrait venir s’appuyer
pour demeurer dans la forme la plus normale! [82]

Cette interruption nous fixe done aussi bien que le commencement de
toutes les taches du haut, sur l’attribution que nous devons en faire 4 la pre-
miére aile, et non a la seconde, et le peu qui reste de ces divers dessins sous
cette derniére, si toutefois il en reste, doit se confondre avec elles, sans con-
tribuer beaucoup 4 nous egarer.

Dvailleurs, nombre de Lépidopterés diurnes des groupes, prés desquels doit
venir se ranger la Sepulta, présentent sous leurs premiéres ailes de semblables
taches costales et basales, ainsi placées, ainsi dentelées, ainsi conformées; d’hab-
itude méme, elles y sont les vestiges plus ou moins complets de ces larges
bandes transversales qui couvrent ces mémes ailes d’une maniére plus ou moins
accusce; assez souvent elles vont se répétant sous les secondes ailes, et s’y
continuent d’une maniére parfois assez suivie, et selon expansion donnée aux
ailes. Elles y sont méme, a mon ayis, un indice de celle que la nature a
entendu leur accorder dans le vol, quand les bandes du dessous des deux ailes
s’y rajustent bien exactement.

NEORINOPIS SEPULTA. 23

Voici donc les taches et les dessins qui, aprés nous avoir aidés 4 reconnaitre la
forme et la nature plus ou moins opaque de ces ailes, sont actuellement eux-mémes
contrélés par la constitution physique de ces organes, restitués 4 leur places °
voulues, et sous Vaile qui les doit comporter.

Voyons actuellement si étude du systéme nervulaire viendra confirmer ou
détruire ces suppositions. Cet examen anatomique a bien son prix actuellement
qu’on en comprend mieux l’importance.

Avant tout, je dois reconnaitre que ces précieux vestiges sont parfaitement
indiqués la ot ils doivent étre, sur [83] cette copie de la piéce originale, et que le
dessinateur nous les laisse suivre assez facilement, tant 4 une aile qu’d l’autre.

Que reste-t-il de la charpente alaire de la premiére aile? D’abord, des traces
de la costale; puis, au-dessus de la lunule blanche de l’apex, les premier et
deuxiéme rameaux des trois apicales qui doivent jaillir de la troisiéme supérieure.
Diverses stries s’échappant du premier, accusent sans doute ici les restes d’un
dessin perdu ou quelques plis anormaux; ¢c’est sans importance. Puis, au-dessus
de la lunule noire, on distingue fort bien la deuxiéme supérieure, et plus bas, enfin,
la premiére.

Sur le bord extérieur, je devine encore l’extrémité des deux premiéres in-
férieures; a travers la seconde aile, un trait noir qui passe entre la lunule blanche
et la large tache noire orbiculaire, m’indique bien la position de la deuxi¢me infé-
rieure; enfin, je suis non moins facilement, entre les deux taches noires orbicu-
laires, la troisiéme inférieure, un peu moins accusée.

Ces deux nervules se relient visiblement a la portion trés lisible de la médiane
qui, sur le dessin, coupe le bas de la premiére tache blanche costale.

Toujours a la premieére aile, la troisiéme inférieure s’y reconnait parfaitement
a la place youlue, au milieu de la dent qu’elle soutient. En effet, le plus souvent,
quand une dentelure, parcillement située, affecte le bord extérieur des premicres
ailes, ainsi qu’on le peut remarquer dans les Van. Progne, Archesia, L.-album,
‘Anglica et autres, cette troisiéme inférieure a la prérogative de lui servir de
support.

Au-dessus de son extrémité nous voyons un faux trait, sans doute, car la pré-
sence d’une nervule me parait impossible 4 cet endroit. Plus bas, au dessous d’elle,
le pli [84] qui, selon moi, doit traverser la tache orbiculaire la plus inférieure,
précéde encore un trait, sans valeur 4 mes yeux, puisquwil m’est inanalysable; et,
en définitive, on voit la sous-médiane qui se projette a travers la seconde aile,
se confondant avec les traces de la seconde inférieure de cette derniére.

Plus bas, avant langle interne, un autre léger faux trait me semble encore
inexplicable, car la saillie dentée de la marge au dehors, précise assez la place, qu’a
la premiére aile, doit occuper l’extrémité de la sous-médiane qui Whabitude reste
volontiers assez distante de la médiane. En _ plus, il ne saurait exister ici d’inter-

24 FOSSIL BUTTERFLIES.
médiane, dont la présence ne se réyéle que dans les tribus trop éloigneés de celle
dont la Sepulta fait partie, pour nous en préoccuper ici. ‘

La nervulation de notre premiére aile se trouve donc ainsi étre au complet, ou
ai peu prés, et déja je la peux réhabiliter avec le crayon, telle qu’elle doit étre.

Passons a la seconde.

La céte seulement se soupconne, on voit parfaitement comme des traits
blancs, la costale qui est ici trés bréve en son trajet; puis la sous-costale, la mé-
diane, et le dé-[85] part des trois supérieures qui se relient trés bien en blanc sur
la marge dentelée de la tache basale (la deuxiéme moins facilement).

Ensuite vient la médiane, dont on suit le parcours, ainsi que sa ramification
qui forme la premiére inférieure et qui passe sous la plus grande des deux taches
orbiculaires; puis la deuxiéme (celle qui, contre toutes les lois de la Spiloptérol-
ogie, couperait la deuxiéme tache orbiculaire, si on l’attribuait 4 la seconde aile).

Vient enfin la troisiéme inférieure, représentée, peut-étre, par une forte ligne
blanche, et qui doit s’attacher a la médiane, pea avant Varticulation fémoro-tibiale
de la patte. :

Dans les bords postérieur et abdominal, je ne peux distinguer ni la sous-
médiane, ni interne, qui sont disparues dans la portion restante, et évidemment
diaphane de cette aile qni se détache ici sur le fond de la pierre.

Maintenant, si sur le tracé de la charpente alaire, scrupuleusement calqué sur
celui de la Sepulta, et que je donne ici (fig. C [ PI. I, fig. 15]) rétablie dans son
entier; si, dis-je, on calque cette nervulation sur un papier végétal et qu’on
reporte cette copie sur le dessin de la pierre originale, qui s’y verra par trans-
parence, ou bien sur ceux de la fig. C, qui est V’insecte tel que je le comprends, on
sera frappé de la précision avec laquelle ces diverses nervures s’adapteront au dessin
et aux taches que j’attribue a la premiére aile, ainsi qu’aux vestiges de la charpente
alaire de la seconde aile. Ainsi, on pourra facilement contrédler mes assertions.

Si done la Solénoptérologie vient a son tour confirmer mes rectifications, je
dois croire que si je me trompe, je ne m’abuse que de bien peu. . . . . .
[86] Maintenant . . . . . . que faire de ce Diurne?

Comme plus haut je l’ai dit, c’est évidemment une espéce aux premicres
ailes fortement échancrées et dentelées, tandis que les secondes y sont arrondies
et simples, 4 méplats bien accusés.

Avons-nous dans nos espéces vivantes quelques-unes qui nous offrent cette
coupe peu commune, et dont les ailes des Van. Archesia et Iphita de Cramer
peuvent nous donner un exemple?

Cette Sepulta me semble tenir beaucoup, tout bien consulté, et des Vanessides
et des Satyrides, telles que nous les comprenons.

Evidemment la Sepulta ne saurait étre une Cyllo phonretian: dite. Serait ce
done une Vanesside? Si la forme des ailes s’y préte quelque peu, son faciés,

NEORINOPIS SEPULTA. 95

Vagencement des ses dessins alaires, me porterait 4 en faire avec M. Boisduval un
Satyride, appartenant a un de ces genres inter- [87| médiaires de ces deux familles
nombreuses, déja si peu Gloignées 4 leur état parfait.

A essayer de caser cet insecte, j’abuserais a n’en pas dourer ds li patience du
lecteur; cependant, en peu de mots, je pourrais lui faire observer (en ne nous
occupant que de la premiére aile, la seule que nous connaissons, 4 mon ayis) que
la large tache basale qui se voit ici, comme a tant de Diurnes, est avec les autres
dessins de sa robe, le propre de nombreux Satyrides de cette taile et de cette
coloration, qu’avee justesse M. Boisduval reconnait devoir étre d'une teinte ter-
reuse, seulement variée de blanc et de noir.

La petite lunule noire me ferait penser qu’en dessus il devait exister une tache
oculaire, dont elle est la simple répétition en dessous, et précisément a lendroit
(entre les premiére et deuxiéme supérieures) ot cette tache existe le plus habitu-
ellement dans nombre de Satyrides de ce faciés, quand elle y est unique.

Certes, il devait y avoir en dessous, le long du bord extérieur. et jusque
dans l’apex, une série disparue d’arceaux internervulaires, formant une double
ligne marginale, ainsi quelle se voit encore entre la dent et langle interne.

La neryulation si peu différente parfois entre nombre de Satyrides et de
Vanessides, ne permet pas, sous ce rapport, d’assigner un poste bien fixe a la
Sepulta; en plus, l'état de son empreinte ne nous permet pas de savoir si la
base de ses nervures est affectée, en tout ou partie, d’entre elles, de ces renfle-
ments vésiculeux si communs a divers groupes de Satyrides. [88]

Nous ne sayons rien non plus de l’absence ou de la présence des disco-
cellulaires, et la perte assez prompte de la costale aux deux ailes, dans la céte,
s’accorde moins avec la marche plus volontiers prolongée de cette méme_ ner-
vure dans les Satyrides, de ’apparence de la Sepulta, etc., etc.

Bref, m’abstenir pour décider rigoureusement de quel genre elle peut étre,
ou méme approximativement, est ce que j’ai de plus prudent a faire; mieux que
moi, d’autres lépidoptéristes pouvant s’acquitter de ce soin. Et a ceux qui,
fatigués de tant de lignes sans ce résultat désiré, me diraient: “Concluez done,” je
répondrais—je ne sais pas!

Explication des figures de la planche.

A [reproduced in our PI. I, fig. 14]. Cyllos epulta, telle qu’elle a été comprise
par M. le docteur Boisduval, en attribuant un appendice caudal @ la deuxiéme aile.

B [reproduced in our PI. I, fig. 16]. Sepulta, telle qu’elle devrait étre
dabord comprise sous le rapport de la forme des deux ailes.

© [reproduced in our PI. I, fig. 15]. Sepulta, telle qu’elle doit étre jugée,
tant pour la forme des ailes que pour la distribution de leurs dessins et leur nervu-
lation, selon M. A. Lefebvre.

26 FOSSIL BUTTERFLIES.

To this Dr. Boisduval at once responded, in the following language :!—

M. Al. Lefebvre, aprés avoir étudié avee soin la position des nervures, la dis-
position des écailles et celle des taches, . . . . . . est arrivé 4 conclure que
javais pris Vaile inférieure pour la supérieure, et que cet appendice caudal, si man-
ifeste dans lespéce en question, était au contraire un angle appartenant a Vaile an-
térieure. Pour donner plus de poids 4 cette opinion, i] a refait une planche ow il
ressuscite d sa mani¢re notre Oyllo sepulta. Avec la queue que nous avons attri-
buée avec MM. Boyer de Fonscolombe, de Saporta, Duponchel, et avec tous les
entomologistes qui ont vu l’échantillon 4 Vaile inférieure, il fait un angle trés aigu
une saillie tout a fait insolite, qu’il place au milieu de Vaile supérieure, tandis qu’il
a fait une aile infériewre complétement arrondie. A cété de celle figure, il en
donne une autre ot il développe notre Cyllo comme il prétend que nous lavons
compris. J’en demande bien pardon a i non estimable ami, mais jamais je ne l’ai
compris de cette fagon. Je conviens du reste que cet intéressant Lépidoptére
fossile serait bien plus antédiluvien tel que M. Alexandre Lefebvre le représente,
que comme nous le supposons, car nous ne lui trouverions aucun analogue,
attendu que jusqu’d présent nous n’ayons’ jamais vu une seule espéce avec des
ailes supérieures anguleuses et appendiculées, et des ailes inférieures arrondies
comme avec un compas, il faut croire que la nature n’en produit plus. Nous
avons toujours observé au contraire que lorsque les ailes supérieures étaient angu-
leuses, les ailes inférieures V’etaient aussi d’une maniére trés manifeste; mais ce
que personne de vous ignore, Messieurs, c’est que trés souvent au contraire les
ailes inférieures, surtout dans le genre dont il est ici question, présentent des
appendices caudiformes plus ou moins saillants, et que parfois les ailes supérieures
ont leur contour simplement sinué. A l’appui de son opinion d’ailes inférieures
arrondies, ayes des supérieures anguleuses, notre collégue a cherché 4 trouver un
exemple dans les figures de Cramer, et il cite en consequence la Vanessa [98]
Archesia qui effectivement présente cette forme; mais Cramer a figuré un individu
mutilé, que probablement on avait arrondi avec hes ciseaux, car nous en possédons
un trés bel exemplaire, pris par M. Drege au pays des Hottentots, que nous met-
tons sous les yeux de la Société, afin qu’elle s’assure bien qu’au contraire cette
espéce est une des plus fortement appendiculée. Le choix de cet exemple est
malheureux. Nous persistons donc tout 4 fait dans Vopinion que nous avons
emise lors de la publication du rapport qui nous a été demandé.

Sometime subsequently Mr. A. G. Butler refers to this dispute between the
two French writers in the following manner:

This very interesting species was described and admirably Pac by Dr.
Boisduval in the French “Annales de la Société Entomologique” (1840); that

1 Bull. Ent. Soc. France, 1851, 97-8. * Cat. Satyr., 189-199.

NEORINOPIS SEPULTA. oT

gentleman considered it to be a Satyride allied to Satyrus rohria, caumas, ete.,'
which it somewhat resembles in the form of the wings.

In the French “Annales” (for 1851) M. Lefebvre published a note upon the
species, in which he criticised Dr.-Boisduval’s paper, and stated that the fossil
species, instead of being allied to rohria, was evidently a Vanessa—that the
strong, tail-like projection belonged to the front, and not to the hind wings, and
represented the angular projection which occurs in all true Vanesside, as an ex-
ample of which he instanced Vanessa (Junonia) Archesia.of Cramer. This re-
markable note was, moreover, accompanied by figures of the species, representing
the tail both upon the front and hind wings.

In the same volume of the “Annales” Dr. Boisduval gives an excellent answer
to M. Lefebvre’s observations, in which he well remarks, “Nous n’avons jamais
vu une seule espéce avec les ailes [190] supérieures anguleuses et appendiculées,
et les ailes inférieures arrondies comme avec un compas;” and certainly, did such
an insect ever exist, its wings would be utterly useless as organs of flight, for
they would invariably carry it downwards. In all insects which have small and
rounded hind wings, the costa of the front wings always far exceeds the inner
margin in length and strength,’ whereas in M. Lefebvre’s insect the reverse would
be the case.

It should be borne in mind, however, that there are two distinct 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 PI. I, fig,
10), although his fault is primarily due to the inaccuracy of the engraving given
by Boisduyal, an inaccuracy which is slightly accentuated in our copy of it (PI. I,
fig. 17). But by far the larger part of his paper is made up of a detailed argu-
ment, 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 neura-
tion), 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
neryures 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

1 Species of Lethe. * As, for instance, in the Sphingide, Heliconide, etc.

e ¥ ;
= ——= — = = ~ 77 > » Pine ‘ i a
a “ae Se ; ne Head
+ =. a =e -_
’ 7 ~

28 FOSSIL BUTTERFLIES.

costal border traverse both wings and must belong to the one to which they cer-
tainly 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,’ 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. Bois-

duyal 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’eeil on suit les
bords de la seconde aile, qu’avec le Docteur je reconnais couvrir en grande partie
la premiére,” no objection is offered in Dr. Boisduyal’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 PI. I, fig. 13, drawn anew from the fossil.
In the same place to which we have just referred Mr. Butler adds the fol-
lowing remarks on the probable affinities of this fossil :?

The true position of C. sepulta is undoubtedly in the family Satyride; and,
so far as can be judged from the beautiful figure in the “Annales,”* it is exactly
intermediate in character between three nearly allied genera now existing, viz.:—
Neorina, Antirrhea and Anchiphlebia, its more immediate allies being the com-
monest species in each of the above genera. Its characters are distributed be-
tween these three species as follows: —

Neorina Lowii, | Antirrhea Phi- Anchiphlebia
Boisd. loctetes, Linn. Archea, Hiibn.
Worm of fomt wings, Sc 6 oo ay 8 0 ee ae ce Se *
Town 6f hind Wiugs,: 2 5 bie Bee ES oe eee *
Tails of hind wings (intermediate in character between), . . . . . . * *
Submarginal spots of front wings, . . . . . . ‘ieee ean ; * . “
Black disco-submarginal spots of hind wings,. . . . . .. 2. 5 *
Pale costal and discal banding of wings, . « - . + + s+ + we ee *? ane *
Limitation of dark dentated basal area of hind wings,. . . . .. . ee o ot
Submarginal line of hind wings, ..... a >. 8p) 0 ee eS, * *
1 Lep. Exot. pl. 48. : * In this figure the neuration has not been very clearly defined,
? Loc. cit., p. 100. : the veining of the hind wings not being continuous,

NEORINOPIS SEPULTA. 29

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’ 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 Satyride,
pp- 189, 190; showing that its nearest ally is Veorina Lowi, a common Bornean
species, but that it also has a slightly more distant relationship to Antirrhea Phi-
loctetes and Anchiphlebia Archea, 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 sub-
family; but it also shows the essential harmony between the markings of the
under and upper surface of the wings of butterflies, notwithstanding their fre-
quent 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 coxe 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 coxe; the tibio-tarsal articulation can be distin-
guished (PI. I, fig. 11) but not the tarsal joints. The wings are bent over

1 Lep. Exot., 127, pl. xlviii. Geol. Mag., x, 3, pl. i.

MEMOIRS A. A. A. S. 6

30 : FOSSIL BUTTERFLIES.

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 PI. I, fig. 8. One is aided greatly in this investigation
by following the lines and series of markings which extend over both the ex-
posed 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

NEORINOPIS SEPULTA. 31

~

represents the real markings of this extraordinarily preserved fossil, a detailed
description of its features follows.

- The basal portion of the fore wing (PI. 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 inter-
space; 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 ex-
tremity, 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 mid-
way 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 neryule; the exterior border is powdery, strongly convex and, starting
from the subcostal nervure midway between the bases of the second and third
superior neryules, 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 ner-
vule 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 seales, 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 con-

32 FOSSIL BUTTERFLIES.

tains 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 in-
terspace with its outer border at an interspace distance from it; with the excep-
tion 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 upper-
most, 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 sub-
parallel 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 trans-
verse 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 direc-
tion, 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

NEORINOPIS SEPULTA. 33

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 sub-
marginal 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 inter-
space, 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, 37"; breadth of fore wing, 20°5™"; length of hind
wing, 31:75""°; length of tail, 4"; distance of the base of the second superior
subcostal nervule of hind wing from the divarication of the costal and sub-
costal nervules, 5°55"™; rows of scales in the subcostal region of the fore
wings, ‘075™™ apart; length of thorax, 5"; of hind femora, 4-6"; of hind tibiz
4-8""; of hind tarsi, 49"™.

Tertiaries of Aix, Provence, France; collection of Count de Saporta.

ia _— _— ee a. Ee ee | rr. =a
> 7 e -- 7

34 FOSSIL BUTTERFLIES.

LETHITES Scupper.

Satyrites Scudd. (nec Blanch.-Brullé), Rey. et Mag. de Zool., 1871-72, 66.

The costal border of the fore wing (PI. 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 ex-
pansion, 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 be-
tween 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 neryule;
at its origin it is directed inward as well as backward (forming the upper ter-
mination of the cell) and passes backward in a small, narrow and rather strongly
curyed bow, bent below more than above, beyond which it assumes a course

LETHITES. 35

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 Hiibn.) as laid down by Westwood and Hewitson, if we exclude there-
from, 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. I, 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 neryure 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

36 FOSSIL BUTTERFLIES.

the same in the two, but the costal neryure appears to be much shorter in
Lethites.

With Lethe (Pl. U, fig. 6) and Debis (PI. 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 res-
pects 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 affin-
ity 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 sub-
costo-median interspace, in Cercyonis than in Lethites; and this seems to be a
character of considerable importance. It may be moted 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 REYNESI. 37

LETHITES REYNESII ScupprEr.
Plate I, figs. 2, 5.
Satyrites Reynesii Scupp., Rev. et. Mag. de Zool., 1871-72, 66-72, pl. vii (1872); Is., Descr. Pap. Foss.

1-7, pl. (1872); Is., Geol. Mag. ix, 532-33, pl. xiii, figs. 2, 3 (1872); Is., Descr. Foss. Butt. 1-2, pl.,
figs. 2, 3 (1872); Brop., Distr. Corr. Foss. Ins. [Satyrites Beynesii], 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 neryure
of the fore wings was greatly swollen. No such form having to my knowl-
edge been described from these beds, Dr. Reynés, 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 un-
doubtedly 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 antennx, 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 haye 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 rest-
ing 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

MEMOIRS A. A. A. 8. 7

- " Pe ai
‘ = - = ba a

38 FOSSIL BUTTERFLIES.

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 neryures 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 exam-
ining 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 pre-
served portion of the insect, we will consider its structure in detail, subsequently
adding whatever can be gleaned from the examination of the other parts.’ [The

account of the structural framework of the wing is given under the genus].

The basal two-thirds of the wing appears to have been more darkly clouded

than the other portions, although in this fuscous area there is apparently a
clearer space towards the upper, outer portion of the cell. There is also a
distinct, darker, uniform and equal rounded spot in the middle of the outer
two-thirds of the lowest subcostal interspace, nearly reaching the nervule on
either side; in the specimen it appears to be broader than long by encroaching
upon the next interspace in front, but this is evidently only apparent, the spots
of the two wings (one of which I have stated to be a little in advance of

1 It should first be premised that throughout this descrip- where the extremities of the costal and the first two upper
tion the fore wing will be spoken of as if it were perfect; for so br hes of the sut tal nervures strike it. For those, there-
completely are the essential parts preserved that one may feel a ‘fore, who would follow the description with a severely critical
strong degree of confidence as to the character of the remainder; eye, the illustrations we have given will correct any apparent
searcely any of the costal margin can be traced on the stone, and overstatement of the text. t
yet one may describe with nearly absolute certainty the point

“

LETHITES REYNESII. 39

the other) being blended. The object is so well preserved that one can see
throughout the parallel series of minute punctures forming the points of inser-
tion for the scales, outlines of the latter of which I have failed to discover.
The wing is 28:5" long, the tip of the cell being distant 15™" from the base
of the wing; the costal nervure is inflated for a distance of 6:5", and the
extreme width of this portion is 1"; the rows of punctures indicating the

former insertion of the scales are *12™" apart.

Of the body itself nothing can be predicated, unless it be that the form of
the abdomen and the appearance of its tip lead us to conjecture that the speci-
men was a female which had deposited most of her eggs, or in which they

were but partially developed.

At the anterior upper extremity of the head is a dark prominence which
seems to be the terminal joint of a palpus; it extends *75™" beyond the head
and is of a nearly uniform width (:2"), scarcely tapering, with a rounded
tip. The basal portion of an antenna, 5": long, is slender and apparently be-
gins to increase slightly and very gradually in size, as in the genus Cneis
Hibn. A finely impressed line, 7-25™" long, appears to be the unrolled, though
slightly curved tongue.

One of the hind femora projects 2-5" beyond the body; its tibia and tarsi
are stretched in a single line, at an angle with it, but as the tip of what is ap-
parently the other hind femur strikes them beyond the tip of their own femur,
it is impossible to say whether they do not overlap, or are not overlaid by, the
tibia and tarsi of the opposite side; their united length on the stone is 5-6™™-;
but if both hind pairs are present, their probable length is 45". There are
also some remnants of the other legs, but in so fragmentary and confused a
state that nothing can be determined from them, nor anything surmised of the
length or structure of the front pair.

In the illustration of the fore wing given in the Revue et Magazin de
Zoologie (fig. B), and copied in the Geological Magazine (fig. 3), the artist

neglected to mark the position of the spot upon the wing. This is given in

40 FOSSIL BUTTERFLIES.

our Pl. I, fig. 5, which, as well as fig. 2, is taken from the ns of my
former plate.

Tertiaries of Aix, Provence, France; Museum of the city of Marseilles.

NYMPHALES —NAJADES — PRA/&FECTI.
EUGONIA Hipner.

Fore wings considerably more than half as long again as broad, the costal
border scarcely bent at a little distance from the base, beyond that nearly

straight to an equal distance from the tip, where it becomes more curved;

outer border with the portion above the middle of the lower subcostal interspace
very slightly concave, having a general direction at a very little less than a
right angle with the central portion of the costal border, beyond suddenly reced-

ing at a little more than a right angle to the middle of the subcosto-median inter- ~

space, and continuing in a deep crenulate curve to just below the lower median
nervure, where a prominent rounded tooth is found, and below which the border
is excised, its angle rounded off; inner border very nearly straight, scarcely

convex on the basal two-thirds. First superior subcostal nervule emitted a little

beyond the middle of the outer two-thirds of the upper margin of the cell; the
second a little more than half way from the origin of the first to the tip of the
cell; the third midway between the tip of the cell and the origin of the fourth;
the latter at three-fifths the distance from the tip of the cell to the apex of the
wing; second inferior subcostal nervule arising scarcely one-third way down the
cell; the cell considerably less than half as long as the wing, and three times
as long as broad; middle of the basal curve of the last median nervule connected
with the vein closing the cell.

The butterflies of this genus, which are generally above the average size,
strongly resemble those of the genus Polygonia, in the form, color and design
of the wings, but on the upper surface of the fore wings the costal markings
are much heavier.

The above characters are wholly drawn from recent species of the genus.

EUGONIA ATAVA. 41

EUGONIA ATAVA (CHARPENTIER) SCUDDER.
Plate I, fig. 1, 3, 7.

Sphinx atava Crarp., Acta Acad. Leop.-Carol., xx, 408-9, Tab. 22, fig. 4 (1843).

Vanessa attavina Herr, Insekt. Tert. Gining., ii, 177-79, Taf. 14, fig. 8 (1849); Is., Nouv. Mem. Soc. Helv.,
xi, 177-79, Tab. 14, fig. 3 (1850); Gres., Deutschl. Petref., 644 (1852); In., Faun. der Vorw., ii, 186
(1856).

Vanessa? atovina Kirs., Syn. Cat. Diurn. Lep., 185 (1872).
Nymphalis? atovina Kirs., Syn. Cat. Diurn. Lep., 648 (1872).

This was the second fossil butterfly known previously to the publication
of Heer’s Tertiary insects. It was first described by Charpentier as a Sphinx,
in the following terms:'

Ungemein interessant, und ich moéchte sagen, ein Unicum ist der in oben
bemerkter Figur abgebildete Schmetterlingsfligel. Dass es ein solcher sei,
zeigt sogleich der erste Anblick, so wie sich bei naiherer Ansicht herausstellt,
dass es unbezweifelt der Oberfligel einer Sphinx Art sei. Er ist in seiner
Form nicht gut gehalten, sondern vorn etwas eingerissen, seine Zeichung ist
aber bewundernswerth erhalten, und erinnert sehr an den fast im ganzen mit-
tleren und noérdlichen Europa vorkommenden Sphinx Tiliw, doch ist er wohl
_specifisch von demselben verschieden. Die drei grossen dunklen Flecke, die
sich yon fussersten Vorderrande, fast bindenartig, tiber einen grossen Theil des
Fligels ziehen, sind unstreitig die Reste ehemaliger Zeichung und Fiarbung des
lebenden Thieres.

The remainder of his remarks apply only to the rarity of fossil remains of
Lepidoptera. The illustration was very poor and is reproduced on PI. I, fig. 3.
The next notice of it is by Heer, who also examined the original type, refigured
[see Pl. I, figs. 1, 7] and xedescribed it in the following manner, referring it
to the genus Vanessa, and changing slightly the specific name:?

Alis anterioribus lividis, basi, fasciis maculisque nigris. Long 163 Lin.

Radoboj. Ein Oberfliigel, dessen Innenrand aber nicht erhalten ist.

Charpentier hat diesen Fligel einem Sphinx zugesprochen und ihn mit dem
Sphinx Tiliz L. verglichen; allein schon die ziemlich stark gebogene Randlinie
(vena marginalis) spricht gegen Sphinx, bei welcher Gattung sie bis tiber zwei

Drittel Fligellange fast gerade verliuft und dann erst gegen die Spitze sich
zubiegt; ebenso aber auch das Geider. Bei Sphinx haben wir namlich ein

1 Acta Acad. Leop.-Carol., xx, 408. 2Insekt. Tert. ning. ii, 177-79.
'

42 FOSSIL BUTTERFLIES.

geschlossenes Mittelfeld und der Ast der vena externo-media, welcher neben der
Flagelspitze ausliuft, veristelt sich nicht. In der Form des Fligels, im Geader
und Farburg stimmt unser Schmetterlingsfliigel, wie mir scheint, am besten mit
der Gattung Vanessa F. tiberein. Wir bemerken nemlich, gerade wie bie den
Vanessen, zunichst eine starke vena scapularis, welche weit vorn in die vy.
marginalis ausliuft; eine schwichere vena externo-media, welche noch niher der
Fligelspitze mit dem Rande sich verbindet; diese bildet nach Innen zuniachst
einen Ast’, der fligelspitzwirts in zwei weitere Aeste sich spaltet; der
aussere von diesen liuft zur Fligelspitze, der innere aber trennt sich nochmals
in zwei Gabeliste, welche zum Hinterrande verlaufen und von denen jeder in
einen schwachen, stumpfen Zahn des Fligelrandes ausgeht. Auf diesen Gabel-
ast folgen weiter nach Innen zwei Liingsadern, welche am Grunde sich wahr-
scheinlich verbinden, und in die vena externo-media eingefiigt sind. Diese bei-
den Adern (es sind diess die fiinfte und sechste Ader von Herrich Schaeffer)
gehen bei [178] den Vanessen getrennt bis zur vy. externo-media hinauf und
divergiren gleich, wie sie aus dieser heraustreten; wahrscheinlich ist diess beim
fossilen Thiere auch der Fall, jedoch sieht man nur die Einmiindung des dus-
seren Astes in die vena externo-media, indem der innere am Grunde ganz
verwischt ist, wie denn tiberhaupt die Adern in Folge des starken Druckes, dem
der Fligel unterworfen war, dusserst schwach hervortreten und nur mit Mthe
zu erkennen sind. Die vena interno-media verliuft wie bei den Vanessen, sie
sendet nimlich nach dem Hinterrande zwei Aeste aus, so dass im Ganzen drei
Langsadern zuletzt in parallelen Linien nach dem Rande verlaufen. Die vena
analis ist nur am Grunde angedeutet, indem der Innenrand grossentheils zer-
stért ist. Das Mittelfeld ist offen, wenigstens ist keine Spur eines Verbin-
dungsastes zwischen y. externo- und interno-media zu finden. In allen diesen
Punkten stimmt also das fossile Thier mit den Vanessen tiberein. Ebenso
stimmt ferner der zackige Hinterrand, indem wir, wie schon bemerkt, an der
Ausmitindung des ausseren Gabelastes der vy. externo-media kleine Zacken be-
merken, wobei freilich zu bedauern, dass von dort. an der Fligel zerrissen ist,
so dass die Randbildung nur an jene kleinen Stelle bestimmt werden kann.
In der Farburg zeigt der Fligel viel Uebereinstimmendes mit demjenigen
der Vanessa Cardui L. Wir bemerken nemlich zunichst dem Grunde eine dunk-
lere Stelle, welche fast bis zu } Fligellinge hinausreicht; dieser dunklere Flt-
gelgrund ist indessen wieder in der Mitte durch einige unregelmissige hellere
Stellen unterbrochen. Auf diese dunkle Stelle folgt ein helles Querband von
1; Linien Breite, welches aber nicht bis zum Innenrande reicht, weingstens ist
an der Stelle, wo die v. interno-media den ersten Ast aussendet, wieder ein,
freilich sehr undeutlich umgrenzter, dunkler Fleck; auf dieses helle Querband

1 Wahrscheinlich ist ausser diesem noch ejn Ast da, der aber verwischt ist.

EUGONIA ATAVA. 43

folgt wieder ein 3 Linien breites dunkles Querband, welches mit mittleren
schwarzen Querband der VY. cardui entspricht; bemerkenswerth ist, dass dieses
bei der V. attavyina von der Nahtseite her ebenfalls durch einen helleren Flecken
getheilt wird, welcher helle Flecken nicht bis zum Aussenrand hinausreicht. Auf
dieses dunkle Querband folgt wieder ein helles Band von 1} Lin. Breite, und
darauf wieder ein dunkler, 3} Lin. breiter Flecken, der aber sehr kurz ist, in-
dem weiter nach Innen an jener Stelle der Fligel wieder hellgelb braun ge-
farbt ist; auf diesen dunklen Flecken folgt wieder ein kleiner heller Flecken;
weiter fligelspitzwarts ist der Fligel dunkelbraun gefarbt, welche Farbe all-
mahlig heller wird, so dass der Fligelrand wieder hellbraun wird 3 die Zacken-
spitzen dagegen sind schwarz.

In der Farbung des Oberfliigels stimmt also der fossile Schmetterling am
meisten mit Vanessa Cardui L. tberein, dennoch kann er nicht als analoge Art
betrachtet werden, denn firs erste war er betrachtlich grésser [179], firs zweite
ist die Randader stirker gebogen, zeigt eine regelmissige Bogenlinie, wah-
rend sie bei Vanessa Cardui in mehr gerader Linie verlauft.”

The only subsequent notice of this insect, not directly copied or abbre-
viated from the above is by Butler, who remarks’; “I think it just possible,
from the great resemblance which V. Attavina of Heer bears to the under
surface of J. [uwnonia] Hedonia, that it is the reverse of J. Pluto.”

I have been unable to see this fossil, or even to find out where it is pre-
served. Charpentier states that he received it for description from Dr. Unger
through Professor Goppert of Breslau. Heer makes no mention of the quarter
whence he received it. Herr Brunner von Wattenwyl searched for it in vain in
the Vienna Museums.

All that can be said, therefore, must be drawn from the illustrations and re-
marks of Professor Heer. These seem to me to leave no doubt that the insect
must be placed in Eugonia, and that it was a little larger than the Huropean
yau-album or our own j-album. A comparison of the neuration of Hugonia
Jj-album (PA. I, fig. 4) with that of Heer’s figures of the fossil (reproduced on PI.
I, figs. 1,7) shows that the last divarication of the subcostal nervure of the fore
wing, and the points of termination of the last two superior nervules and of the
subcostal nervure itself are essentially the same in both; while the position of

all the markings on the fossil, allowing for its natural defects, are quite the

1 Lep. Exot. I, 128. .

44 FOSSIL BUTTERFLIES.

same in position, direction and intensity, as in Z. j-album (PI. I, fig. 6). The
same may be said of the form of the wing, as far as it can be seen, but as this
is true only of the costal margin, and the merest fragment of the outer border,
it cannot be considered to have much weight in itself; still, taken in connec-
tion with all the other features, which agree almost wholly with those of Eugonia,
and but partially with its near ally Vanessa, to which Heer compares it, we must
refer the fossil to Eugonia, at least until a new examination of the fossil shall give
us further facts as a basis for an opinion. This is the position dubiously assigned
to it by Kirby, in his Synonymic Catalogue.
Tertiaries of Radoboj, Croatia.

PAPILIONIDA—DANAI—FUGACIA.

MYLOTHRITES ScuppeEr.

Of the form of the fore wing (PI. II, figs. 7, 17) we can say but little, from
the imperfect nature of the fossil; the costal margin, however, is very regularly
and rather strongly arched, and the direction of the middle portion of the outer
border (probably at a right angle, or at a little less than a right angle, with the
apical portion of the costal margin, and but slightly convex) leads us to presume
that the apex was rather pointed, though not falciform.

The neuration of the same wing (PI. II, fig. 7) is very similar to that of
Mylothris.". The costal nervure terminates at about five-sevenths the distance
from the base of the costal margin to its tip; the subcostal nervure emits two
branches before the cell, the second probably close to the apex of the cell, the
limits of which are not given in the drawing prepared for me, but which could
probably be made out by a sufficiently careful examination of the original; a third
superior neryule is emitted from the subcostal nervure at less than half the dis-
tance from the origin of the second to the outer border, and the emission of the
inferior nervule, if it could be traced, would mark the termination of the cell; thé
median nervure is of course three-branched and scarcely curves upward at all
to meet the subcostal.

} Compare, in this gespect, Butler’s Revision of the Pierine, Cist. Ent., I, ili, pl. i, fig. 8; or Trimen, Rhop. Afr. Austr., PI. ii, fig. 2.

MYLOTHRITES PLUTO. 45

The design of the upper surface of the fore wing (PI. II, fig. 17) is simple,
consisting only of a broad marginal pale band on a dark ground, enclosing small
dark spots in the middle of the interspaces. ;

This fossil was placed by Heer among the Nymphales, and referred, like the
preceding, to Vanessa. Heer lays stress on the non-closure of the cell, but it
appears questionable whether this is not simply the result of the defective preser-
vation of the fossil. Edwards has since referred it to Argynnis, on account of
the general aspect of its markings, and Butler, on the same ground, to Junonia.
But the new drawing of the fossil obtained for me through the kindness of my
friend Herr Brunner von Wattenwyl, and by him carefully compared with the orig-
inal, leave little doubt that it is a Pierid, and belongs in the neighborhood of such
genera as Mylothris and Hebomoia. The latter genus it closely resembles in

the form of the wings. Further comparisons are presented under the species.

MYLOTHRITES PLUTO (Heer) Scupper.
Plate I, figs. 2, 7, 17 (15?).
Vanessa Pluto Heer, Insekt. Tert. Gining., ii, 179-82, Taf. 14, fig. 4, 5 (?) (1849); In., Nouv. Mem. Soc.
Helv., xi, 179-82, Tab. 14, figs. 4, 5 (?) (1850); Gres., Deutschl. Petref. 644 (1852); In., Faun. der

Vorw., 186-7 (1856); Picr., Traite de Palwont., ii, 393, pl. 40, fig. 21 (1854); LyrxL, Elem. Geol., 6th
Ed., 243, fig. 179 (1865).

Argynnis Pluto Evw., Butt. N. Amer., i, Argynnis I, fig. (1868); Kirs., Syn. Cat. Diurn. Lep., 155 (1871).
Junonia? Pluto Butt., Lep. Exot., 127-28, pl. 48, fig. 7 (1873) ; In., Geol. Mag., x, 3-4, pl. 1, fig. 7 (1878).

Heer’s description of this insect is as follows:!—

Alis griseo-nigris, anterioribus margine posteriore ocellis sex pallidis.

Lange des Vorderfligels wahrscheinlich 15 Lin; er ist erhalten bis zu 14}
Lin; grésste Breite 8} Lin.

Radoboj. Ein ausgezeichnet schénes Exemplar in dem k. k. Hofkabinet zu
Wien; leider fehlt aber der Kopf, der Hinterleib, der grésste Theil der Hinter-
fligel und die Spitze der Vorderfligel [ PI. II, fig. 2].

Der Brustkasten ist linglich oval, in der Mitte zwei Linien dick, an der
Oberseite yon ein paar _Streifen durchzogen. Der Oberfliigel ist am Grunde
schmal, nach dem Hinterrande hin aber stark yerbreitert und errcicht daselbst
seine grésste Breite. Die Aussenrandlinie (v. marginalis) ist sehr stark gebogen,
und zwar bildet sie vom Grunde zur Spitze eine regelmissige, starke Bogenlinie.

1 Inseké. Tert. Gining., LI, 179-82.
MEMOIRS A. A. A. 5S. 8

46 FOSSIL BUTTERFLIES.

Die Schulterader ist am Grunde stark } Lin. vom Rande abstehend und liuft aus-
serhalb der Fligelmitte in denselben; die vena externo-media ist ihr sehr genahert
und nur mit Mihe zu unterscheiden, sie mindet noch niher fligelspitzwarts in
die Randader. Sie sendet zunachst einen einfachen Ast ab, der mit dem Haupt-
stamm parallel lauft, ihm sehr genahert ist und noch naher der Fligelspitze in
die Randader mindet; der zweite Ast spaltet sich bald wieder in zwei Aeste, von
denen der iussere vor der Fligelspitze in die Randader auslauft, der innere theilt
sich nochmals in zwei Gabeliste, welche ohne Zweifel innerhalb der Fligelspitze
ausmiinden; ganz nahe, wo der zweite Hauptast der vena externo-media ent-
springt, liuft der dritte aus, der einfach und nach dem Hinterrande geht; auf
diesen folgt ein vierter Ast, dessen Insertion aber nicht zu sehen; es scheint, dass
er auf der Fligelflache entspringe.—Die vena interno-media ist ebenfalls stark
ausgesprochen; sie sendet nach Innen zwei starke, aber einfach bleibende Aeste
aus, so dass sie im Ganzen in drei parallelen Adern in den Hinterrand einmiindet.
Das Mittelfeld ist verhiltnissmassig ziemlich klein und nicht geschlossen, indem
kein Querast die beiden Mitteladern verbindet. Die vena analis ist einfach und
lauft nahe dem Nahtrande herunter. In.den Feldern zwischen je zwei Lingsrip-
pen sieht man eine schwache Langslinie, welche vom Fligelrande dis zum Augen-
punkt lauft; sie stellt eine schwache Furche oder Falte dar, die dort im Fligel
sich befunden hat. Der Hinterrand ist leider nicht ganz erhalten, namentlich fehlt
die Fligelspitze, de- [180] ren Form zur Bestimmung der Gattung so wichtig
ware; es ist daher nicht zu ermitteln, ob diese ganzrandig oder gezacht war.
Der Hinterrand verlauft in einer schwachen Wellenlinie, indem ganz schwache,
stumpfe Kerbzihne an der Ausmitindung der Langsadern liegen.

Die Farbe des Fhigels ist ein dunkles Graubraun; am Grunde und im Rand-
felde ist er dunkler, welche dunklere Parthie aber allmahlig in die hellere verlauft;
gegen die Augenflecken zu wird die Farbe wieder dunkler; langs des Randes
bemerken wir eine Reihe (nemlich 6) von runden, hellen Flecken und zwar liegt je
zwischen zwei Lingsadern ein solcher Fleck, welcher das ganze Feld zwischen den
Adern ausfillt. Es reicht dieser helle Fleck nicht bis zum Fligelrande, welcher
wieder dunkler graubraun gefairbt ist. In der Mitte jedes Fleckens legt ein
schwarzer, runder Punkt; ob dieser noch eitien weissen Augenpunkt besessen habe
oder nicht, ist nicht mit Sicherheit zu ermitteln, doch ist es wahrscheinlich, indem
wenigstens bei zwei dieser Punkte in der Mitte eine kleine, hellere Stelle wahr-
zunehmen ist. Diese hellen Augenflecken scheinen von keinem schwarzen Ring
eingefasst zu sein.

Von den Unterfliigeln ist nur der Grund erhalten. Wir sehen da die, bald in
zwei Gabeliste sich spaltende, vena analis und die beiden am Grunde ganz genih-
erten Mitteladern. Die Farburg dieses Fligeltheils ist gleich wie am Oberfltigel,
und zwar nach dem Grunde zu auch dunkler werdend.

——— a

MYLOTHRITES’ PLUTO. 47

Die Bestimmung der Gattung, zu welchem unser Thier gehort, wird sehr
dadurch erschwert, dass der Hinterrand nicht ganz erhalten ist. Nach [181] der
allgemeinen Form und dem Geader der Fligel muss er wohl zu den Nymphaliden
gehéren. Bei den Papilionen, Pieriden, Danaiden und Satyriden ist die Mittel-
zelle der Fligel durch einen starken Verbindungsast der vena externo- und interno-
media geschlossen, wogegen beim fossilen Thiere die Mittelzelle des Oberfltigels,
und yielleicht auch die des Unterfliigels, gedffnet ist, wie diess bei vielen Nympha-
liden yorkommt. Von den Pieriden unterscheidet er sich tiberdiess durch die Art
der Varistelung der y. externo-media, indem (um mich der Terminologie yon
Herrich Schiffer zu bedienen) die Tte und 9te Rippe, vom Nahtrande an gerechnet,
aus der sechsten entspringen, und die achte aus der siebenten, wihrend beim
fossilen Thiere die 8te und 9te Rippe, wie bei den Nymphaliden, aus der 7ten
entspringen. So weist also das Geaider auf einen Nymphaliden. Unter diesen
kommen ein paar Gattungen vor (nemlich Apatura und Melita) mit offener
Mittelzelle der Hinterfliigel; allein bei diesen finden sich keine Arten mit Augen-
flecken, wogegen unter den Vanessen eine Art vorkommt, welche in der Flecken-
bildung eine auffallende Aehnlichkeit mit dem fossilen Thiere hat. Zwar ist bei
Vanessa die Mittelzelle der Hinterfliigel geschlossen, aber durch einen so zarten,
feinen Querast, dass dieser sich leicht verwischen konnte. Jene dem fossilen Thiere
nahe verwandte Art der Lebenwelt ist die Vanessa Hedonia L. F. Cramer de Uet-
landsche Kapellen T. II, Taf. 69, C. D. und T. VIII, Taf. 374, E. F. Es hat diese
genau die Grésse des fossilen Thieres, der Aussenrand bildet ebenfalls eine starke
Bogenlinie; die Oberfltigel sind grauschwarz und haben am Hinterrande eine
Reihe von 6 Augenflecken; es sind diese roth und mit einem schwarzen Punkt
in der Mitte versehen; dieser schwarze Punkt umfasst einen kleinen. weissen
Punkt. In der Vertheilung und Stellung dieser Flecken stimmt Pluto ganz
mit Hedonia tiberein, nur sind bei letzterer die Flecken kleiner und von einem
schwarzen Ring umfasst; ferner sind sie etwas weiter vom Rande abstehend.
Die Vanessa Hedonia kommt auf Ceylon, Amboina, Java und den Phillippinen
yor, hat also im tropischen Asien eine weite Verbreitung.

Von Schmetterlingen mit ahnlicher Farbung kénnen noch in Betracht kom-
men: die Argynnis Diana Cramer II, p. 4, t. 98, D. EH. Say. Americ. En- [182]
tom. 17, welche im siidlichen Theile der vereinigten Staaten (Neu-Georgien, West-
florida, Arkansas and Missouri) lebt. Es hat dieser Schmetterling eine ahnliche
Tracht, ist schwarz und am Hinterrande mit einer Reihe gelber Flecken versehen,
welche je zwischen die Liingsadern vertheilt sind. Diese gelben Flecken reichen
aber bis zum Rande, und ferner hat jeder zwei schwarze Punkte. Auch ist die A.
Diana bedeutend grésser. In Grosse und Farbung stimmt daher das fossile Thier
mehr mit der Hedonia tiberein, als mit der Diana, doch kann mit voller Sicherheit
erst dartiber entschieden werden, wenn enimal ein Exemplar mit vollstandig erhalt-

48 FOSSIL BUTTERFLIES.

enem Hinterrand gefunden wird; was yon diesem erhalten ist, spricht aber auch
mehr fiir die Hedonia als die Diana.

Edwards, in his beautiful work on American Butterflies, refers to this insect
in his description of Argynnis Diana‘ and reproduces, from Lyell’s Elements of
Geology, Heer’s figure of the insect. He remarks: “It is called Vanessa Pluto
in the text, but is plainly an Argynnis.”

Butler, when cataloguing the same insect, remarks :*—

It is quite possible, as Mr. Edwards suggests, that the so-called “ Vanessa
Pluto” may be the ancestor of P. Diana, though in the narrower banding of its
wings, with but one row of submarginal spots, it more nearly resembles some of
the East Indian forms of Junonia Hedonia: the two genera to which these species
belong agree in many respects, and are perhaps nearly allied.

Later, he figures the fossil and refers it doubtfully to Junonia, appending the
following remarks :°—

I have noticed this species at p. 109 of my catalogue of Fabrician Diurnal
Lepidoptera; Mr. W. H. Edwards of W. Virginia having decided in his Butter-
flies of N. America that it is unquestionably an Argynnis allied to A. Diana,
notwithstanding the important discrepancies which Heer points out [128]. That
it may bear some distant relationship to A. Diana is quite possible, but that it is
“plainly an Argynnis” is quite another thing; to my mind it is plainly a Vanessid,
probably a Junonia near to J. Hedonia, and I think some points in Heer’s descrip-
tion (of which Mr. Edwards takes no notice) are very important, as evidencing
its near relationship to J. Hedonia rather than to A. Diana [here he quotes Heer’s
description of the submarginal spots].

The ocelli are well shown in Heer’s figure, but in the woodeuts by Lyell and
Edwards, which haye in other respects been made much darker than the original,
the indication of the lower edge of the ocelli has been omitted altogether, and,
consequently, the resemblance to the species of Junonia is rendered less evident.
I think it just possible, from the great resemblance which V. Attavina of Heer
bears to the under surface of J. Hedonia, that it is the reverse of J. Pluto.

This species is very simple in its markings (PI. IT, fig. 17), the whole upper
surface, excepting a broad space next the outer border of the fore wings (the
equivalent part of the hind wings is not preserved) being of an uniform dusky
tint; a broad belt of a lighter shade margins the (fore) wings, growing less

1 Butt. N. Amer., i, Argynnis, I, 5 Lep. Exot., i, xv, 127-28, Pl. 48, fig. 7; Geol. Mag., x, 3-4,
* Cat. Fabr. Lep., 109. Pl.1, fig. 7.

j
#
-

— !. eo

MYLOTHRITES PLUTO. 49

distinct from the darker base above the next to the lowest subcostal nervule; this -
belt darkens toward the outer border, especially in slight dusky fleckings along
the nervures and down the middle of the interspaces; the latter streaks reach
small, round, blackish spots about one-quarter the width of the interspaces, in
the middle of the basal two-thirds of their lighter parts. Heer represents them
too far from the outer margin of the wing, and as often crowned above with a
dark semicircular line, which is not at all indicated in the drawing made for me;
these spots are found in all the interspaces below the outermost superior sub-
costal nervule, but they are very indistinct and minute above, faint below and only
distinct and as large as stated in the three interspaces next above the lowest
median nervule. The light belt is two interspaces wide in the upper median
interspace, but widens a little above this and is separated from the darker base
by a vague and very slightly crenate line (less crenate than in the representation
by Heer), which approaches the outer margin at the nervures and to a slightly
greater extent in the lower part of each interspace than in the upper.

Pierids with so dark a coloring as appears in this fossil are not unknown,
particularly in the genera Archonias and Pereute; compare for example the figure
given in Doubleday and Hewitson’s Genera of Diurnal Lepidoptera, Pl. V, fig.
2. And that markings of this character are not unknown, compare some species of
Ixias, Hebomoia and allied genera; if the colors of Hebomoia Leucippe, as given
by Doubleday and Hewitson, were reversed, the resemblance to Pluto would be
rather close; and while light spots in a dark border are the rule in this subfamily,
dark spots on a light ground are not unknown, and the reversal of tints is a
not uncommon occurrence in nearly related Lepidoptera.

A second fossil, which I have been unable to see or to have redrawn, is given
by Heer as probably representing the under surface of the same insect. His re-
marks are as follows :'—

Hierher rechne ich auch ein Stick eines Unterfitigels aus der Gratzer Samm-
lung, das bei Taf. XIV, Fig. 5 [ Pl. II, fig. 15], dargestellt ist. Die Hauptadern
treten an diesem Fliigelstiicke alle hervor. Die beiden Mitteladern schliessen ein
nicht sehr grosses Mittelfeld ein; ob dieses durch einen Verbindungsast zwischen

1Insekt. Tert. ning, II, 180.

50 FOSSIL BUTTERFLIES,

den beiden Mitteladern geschlossen ist oder nicht, war mir nicht méglich zu ermit-
teln; bei guter Beleuchtung glaubte ich dort einen schwachen Quereindruck zu se-
hen, der als Verbindungsast zu deuten wire; jedenfalls wire derselbe aber ausserst
zart, viel zarter als die ibrigen deutlichen Adern. Die fiussere Mittelader sendet 4
Aeste aus, der erste entspringt nahe der Fligelbasis und liuft nach dem Aussen-
rande, die drei folgenden entspringen niher fligelspitzwarts. Die v. interno-media
zerspaltet sich in 3 Aeste, ganz so wie die des Oberfliigels, welche auch in gleicher
Weise verlaufen. Alle 3 Aeste sind fast gleich weit von einander entfernt und
entspringen nicht von einem Punkt. Die vena analis zerspaltet sich bald nach
ihrem Ursprung in zwei Gabeliste, welche nach aussen laufen. Die Farbe des
Fligels ist ein helles Graubraun.

As far as the neuration is concerned (excepting that of the costal nervure,
which is certainly incorrectly rendered, and does not accord with the description)
it agrees sufficiently with the general neuration of Mylothris' to suppose it may
belong to the allied genus Mylothrites, but that it can belong to M. Pluto is
exceedingly improbable, as one may judge by tracing the
probable extent of the broken hind wing, and placing the
tracing in juxtaposition with the fore wing of Pluto, as in
the accompanying woodcut (fig. 1); for it must be remem-
bered that in all the genera of this subfamily, the cell
extends at least to the middle of the wing; the hind wing

of M. Pluto must, therefore, have certainly been fully one-

sixth longer than the wing conjectured to belong to it;
Fig. 1. The dotted outer border of

the hind wing represents the proba- §0 great a difference is at least unusual among indi-
ble limit of the Gratz fossil. The

broken outer border indicates the 1 } j } *
ch in pacer serag viduals of the same species in this group; moreover, the

I OTe neuration is not quite what we should expect, although

the appearance of veins on the drawing we have reproduced must be in part
due to extraneous causes; we will, therefore, make no attempt to decipher the
present condition of the fossil, trusting that some of the Austrian lepidopterists
will give the subject early attention.

A study of the original description and illustration of the front wing of this
butterfly leads me to the conclusion that the description of the neuration of

1 Compare the illustrations referred to in the note on page 44.

,
a
*

COLIATES. 51

the fossil was drawn up from the illustration and not from the fossil itself. Both
agree in the points in which my drawing (PI. I, fig. 7) differs from them; and
since in these very points they will not harmonize with the neuration of any living
Lepidoptera, while the drawing I present agrees as well as could be desired with
certain of them, I am forced to believe the original drawing published by Heer,
and the accompanying description, presumably founded upon it, to be incorrect.
I am acquainted with but very few living butterflies' in which a nervule is emitted
from the inferior side of the subcostal nervure nearer the base of the wing than
any of the superior nervules of the same vein; this is the manner in which the
neuration of this butterfly is represented in Heer’s plate and in his description, if
read carefully in connection with the plate; although he does not tell us on which
side of his zweite Hauptast his dritte Hauptast originates.

The description given by Heer of the markings of the fore wing is more
complete than I have been able to offer from an inspection of drawings alone; it
differs, too, in one somewhat important point, in that what I have called a broad
lighter belt with blackish dots in each interspace, he has described as a series
of pale circular spots as broad as the interspaces, each containing a blackish pupil.
A reéxamination of the fossil upon this, point is desirable; the only indication of
such circular pale spots in my drawing is the curved boundary in each interspace
between the darker and lighter portions.

Tertiaries of Radoboj, Croatia. Fore wing, Hof-Mineralien Kabinet, ‘Vienna.
Hind wing, Museum of Gratz, Austria.

COLIATES Scupper.

The fore wing (PI. II, fig. 5) is slightly more than twice as long as broad;
the costal border is straight for fully two-thirds its length, and then curves grad-
ually and slightly downward, the apex rounded off; the outer margin has a nearly
regular and slight convexity, but is nearly straight in the middle half; the lower
outer angle is rounded and the inner margin slightly convex. The costal neryure

1 These, it is true, are Dareai, but aberrant forms, like Leptidia, etc.

52 FOSSIL BUTTERFLIES.

scarcely reaches the middle of the costal border; the discoidal cell is but little
more than half the length of the wing; the subcostal nervure has but two superior
branches, although the outer is not only itself forked, but its upper fork is
branched at the extreme tip of the wing; the first superior nervule is emitted at
some distance previous to the tip of the cell, or opposite the base of the first
median nervure; it terminates in the middle of the outer half. of the costal
margin, and the forked branch of the outer superior nervule supports the extreme
apex of the wing; the inferior subcostal nervule arises midway between the bases d
of the two superior nervules, and terminates about one-third way down the outer
border; the vein closing the cell strikes it near the base and has an inward con-
vexity, meeting the upper median nervule farther from its origin; the first median
nervule originates at some distance beyond the middle of the cell.

In the form of the wing and its neuration this fossil group is more nearly
allied to Delias (Pl. I, fig. 4) than to any other genus I have been able to ex-
amine. It is plain at first glance that it must be placed in the vicinity of
Delias, Thyca, Prioneris and similar East Indian Fugacia, in which there are but
two superior subcostal nervules, and in which the outer of these is forked; but I
have met with no instance among these in which one of these forks is itself
branched; and this insect differs notably from them all in the elongate form of ‘a
the wing, the remarkably straight costa’ and the shorter discoidal cell; and from
all Pierids in the shortness of its costal nervure and the basal extension of the first
superior subcostal nervule; this latter nervure always originates, in every living
type I have examined, at or beyond a point opposite the middle of the space
between the bases of the first and second median neryules.

COLIATES PROSERPINA Scupper.
Plate I, fig. 5.

The fossil to which I have given this name is exceedingly obscure, having no
color whatever distinct from the stone in which it is imbedded; this is of a chalky
gray color. I have seen both impression and reverse, the latter a little in relief.

1 See, however, the American genus Leodonta,

PONTIA. 53

The fossil consists of both anterior wings, one beneath and slightly in advance of
the other, thus complicating very greatly the study of the already indistinct
neuration; in addition to this the wings are crumpled and additional longitu-
dinal lines are present, scarcely distinguishable from the longitudinal nervures.
On this account it should be stated that there may be some doubt about the exact
position of the lowest three branchlets of the subcostal nervure. The stone has
been broken next the edge of the wing, and its form can thus be traced where
the real border is wanting, although again the drawing presented may be slightly
inaccurate next the inner margin; but the probabilities are great that it is correct
throughout. The spots which are represented on our plate in the middle of the
lower median, subcosto-median and lower subcostal interspaces, are only irregu-
larities of surface on the stone, but as they appear in regular position are not
improbably dark spots, upon a light ground. A few points for the insertion
of the scales can be detected near the apex of the wings, ‘075"" apart. The
neuration of the fossil agrees better with that of Delias Pasithoe than with
that of any other butterfly I have examined. Length of wing, 21"; greatest
breadth, 9". :
Tertiaries of Aix, Provence, France. Collection of Count de Saporta.

PAPILIONID2 — DANAI— VORACIA.
PONTIA Fapricivs.

Pierites Herr, Insekt. Tert. Gning., ii, 182; Is.; Nouv. Mem. Soc. Helv., xi, 182; Gres., Faun. der Vorw.,
ii, 187.

Fore wings fully three-quarters as long again as broad, the costal margin
slightly convex at the basal and apical fifth, scarcely bent at an angle with the nearly
straight middle portion, the outer angle abrupt but softened. Outer margin nearly
straight and inclined at an angle of forty-five degrees with the middle portion of
the costal border, above the middle subcostal nervule receding slightly in a gentle .
curve. Inner margin straight, the outer angle well rounded. Costal nervure ter-
minating a little beyond the middle of the margin. Subcostal nervure with three

MEMOIRS A. A. A. 8. 9

54 FOSSIL BUTTERFLIES.

superior branches; the first arising shortly before the middle of the outer half of
the cell, a little nearer the apex of the cell in the female than in the male; the sec-
ond arising scarcely (male), or a very little (female), beyond the tip of the cell;
the third emitted at about two-fifths the distance from the apex of the cell to the
outer margin, forked at the extreme tip in the male. Cell half as long as the

wing and nearly four times as long as broad.

The butterflies are scarcely larger than those of the genus Pieris, but have
more pointed fore wings; like them they are white, but extensively spotted with
deep brown; the fore wings have a broad bar at the tip of the cell, and midway
between this and the outer border a widely interrupted transverse series of similar
but smaller spots; the outer border, especially the upper half, is also more or less
distinctly margined with triangular, frequently confluent spots seated in the inter-
spaces; these occur more often in the female than in the male.

The characters given above are drawn wholly from recent species of the

genus.

PONTIA FREYERI (Heer) ScupprEr.
Plate LI, figs. 16, 18.

Pierites Freyeri Heer, Insekt. Tert. Gining., ii, 182-83, Taf. 14, fig. 6 (1849); Isb., Nouv. Mem. Soc. Hely.,
xi, 182-83, Taf. 14, fig. 6 (1850); GirB., Deutschl. Petref., 644 (1852); Is., Faun. der Vorw., ii, 187
(1856); Kirs., Syn. Cat. Diurn. Lep., 509 (1871).

The original description of this insect we owe to Heer; it is as follows:*

Alis anterioribus lividis, margine maculisque duabus nigris.

Lange des Vorderfligels 93 Lin., Breite 5} Lin.

Radoboi. Ein einzelner Vorderfligel, dessen Spitze und theilweise auch Hin-
terrand aber zerstort ist, in der k. k. montanistischen Sammlung zu Wien. [ PI. II,
fig. 16.]

Das Geider ist nicht in seinem Verlauf zu ermitteln und da auch der Umriss
des Fligels nicht vollstandig vorliegt, ist die Gattung nicht mit Sicherheit zu
ermitteln. In Form und Farbe scheint er am meisten mit manchen Pieriden,
_namentlich Pieris Daplidice, zu stimmen, woftir auch das diinne Schuppenkleid,
“das er gehabt zu haben scheint, angefahrt werden kann, wogegen die allerdings
stumpfen Zacken am Hinterrand eine abweichende Bildung zeigen.

41 Insekt. Tert. Gning., ii, 182-3.

.
‘

PONTIA FREYERI. 55

Der Fhigel ist am Grunde stark verschmilert, nach dem Hinterrande zu stark
verbreitert; der Hinterrand ist stumpf gekerbt; in die Bucht der Kerbe lauft eine
Langsfalte, in die Mitte derselben eine Ader aus, die man aber nicht bis zur Inser-
tionsstelle verfolgen kann; die aussere Fliigelspitze fehlt; ebenso ein Stiick des
Hinterrandes an der [183] Nahtseite. Von den Adern kann man nur die einfache
y. analis in ihrer ganzen Lange verfolgen; sie ist dem Nahtrande sehr genihert.
Die Farbe des Fligels ist hell gelbbraun und war im Leben wohl weiss oder gelb-
lich. Der Hinterrand ist von der Mitte an bis zum Aussenrand schwarz, und zwar
wird diese dunkle Parthie auswirts breiter; ungefaihr in-der Fligelmitte geht vom
Aussenrand ein viereckiger, dunkler Fleck aus, welcher dieselbe Grésse, Form
und Stellung hat, wie der schwarze Fleck bei Pieris Daplidice; ein zweiter klein-
erer, rundlicher Fleck liegt naher dem Hinter- und Nahtrande und entspricht
dem, an derselben Stelle liegenden, Flecken der Unterseite von Pieris Daplidice.—
Am Fligelgrunde bemerkt man den Schenkel und Schiene eines dinnen Beines,
das wohl diesem Thiere angehort hat.

This insect evidently belongs to the genus Pontia, judging from the disposi-
tion of the markings of the upper surface of the fore wings (PI. I, fig. 18); this
is the only thing we have here to guide us, although the drawing made for us in
Vienna seems to show that with great pains the neuration of at least a part of the
apex might be traced and lead to more positive determination. The fossil species
seems best comparable with P. Protodice (P1. I, fig. 12) of N. America, although,
as suggested by Heer, most nearly resembling P. Daplidice, of European species.
The dark spot at the apex of the cell appears to cover a larger area than in P.
Protodice, extending with equal breadth almost to the costal margin, and also
covering a considerable space at the base of the subcosto-median interspace, equal
indeed to the entire width of the portion of the spot within the cell. The region
below this spot, next the base of the lower median interspace, is also rather faintly
suffused with griseous tints. The precise extent of the subcostal spots midway
between the cell and the apex cannot be determined, owing to the imperfect state
of the fossil; but they evidently form a connected series as much larger than the
similar spots in P. Protodice as the cellular spot, and extend from the costal
margin to the lowest subcostal nervures, expanding considerably baseward in the
upper half of their course. There is no spot in the upper median interspace, as
in P. Protodice, but, instead, a precisely similar one in the middle of the apical

56 FOSSIL BUTTERFLIES.

three-fifths of the lower median interspace, where it does not occur in P. Proto-
dice; and this affords the principal ground for supposing the insect to be generi-
cally distinct from Pontia, no distinctive premarginal spot occurring in this inter-
space in any species of Pontia which we have seen. The dusky premarginal
fleckings of the nervures terminating on the outer border, often enlarging into
distinct spots, which are so usual in Pontia, especially in the upper half of the
wing, are also absent from the fossil species; but in their place the whole outer
margin appears to be almost uniformly, though not heavily, griseous, a little more
distinctly so in the upper than in the lower half of the wing. The spot just
beyond the middle of the medio-submedian interspace, distinct in P. Protodice,
but deepest in shade on the lower half of the interspace, and in other species
sometimes wholly confined to it, is seen in the fossil species, but is far less
distinct, confined to the lower half and situated exactly in the middle. There
are indications also of dark markings following the basal third of the sub-
median nervure; and apparently the basal half of the costal edge, as far as the
costal nervure, is darker than any part of the wing, excepting in a sudden and
rather broad, distinct break in its middle. This darker portion is considered by
Heer as the femur of one of the legs, superimposed upon the base of the wing;
perhaps, however, this is due to an accidental folding of the wing at this point,
which seems the more probable, because if we suppose this darker portion to be
turned back, the curve of the costal border would approximate much more closely
to its condition in P. Protodice; while its present form is much straighter, exhib-
iting only a very slight and regular convexity. As far as can be judged from
the fragment, the form of the other parts of the wing coincides with that of P.
Protodice.

As in all species of Pontia there is a slight wrinkling of the membrane in the
interspaces, forming slight channels running from the outer border inward, nearly
to the depth of two interspaces, indicated in the fossil by dark lines as heavy as
the nervures, and caused by their filling with sedimentary material. The extreme
length of the part of the wing preserved is 24" and the greatest width 22.5.

The markings lead one to conjecture that the individual was a male.

i

THAITES. 57

PAPILIONID 2 — PAPILIONIDES— PARNASSII.

THAITES Heer, MS,

Body rather robust (Pl. II, figs.9 and 10). Vertex of head large, broad,
convex. Eyes pretty large, short ovate, their longer diameter vertical. Palpi (Pl.
II, fig. 7) slender, resembling those of Thais, but rather longer, extending far
beyond the eye, rather thinly clothed with hairs. Antenne (PI. ITI, fig. 8) resem-
bling those of Sericinus more than those of Thais, being about half as long as the
body, slender and equal on the basal three-fifths, gradually expanding beyond into
a club, which is more than twice as broad as the stem, and stoutest just before the
well rounded, slightly upturned tip; in the middle of the antenne the joints are
half as long again as broad, broader than long at the base of the club, and three
or four times as broad as long in the middle of the club and beyond; on the apical
half of the club, and perhaps a little further, the joints of the club are furnished
with a double row of minute shallow pits, such as are seen in Eurymus. The
tongue was at least as long as the thorax.

The thorax is well arched and pretty stout; the paraptera (PI. ITI, fig. 6) are
a little more than twice as long as broad, their outer edge nearly straight, the pos-
terior extremity broad and well rounded. The legs are not well enough preserved
to state anything concerning them with certainty, but the middle (?) pair are
probably of the length of the antenne.

. The fore wings (PI. II, fig. 3) are only a little more than half as long again
as broad, the greatest breadth beyond the middle; the costal border is pretty regu-
larly and not greatly arched throughout; the outer margin is more strongly arched
but with a similar regularity, and the general direction of its upper half is at right
angles to the outer third of the costal border, the apex scarcely rounded off; the
inner border is nearly straight. The proportions of the hind wing, as to length
and breadth, are nearly the same as those of the fore wings, making it unusually
long and narrow, as in Thais (PI. II, fig. 4), and also, as there, nearly as broad
toward the base as at tip. The costal border is rather strongly convex next the
base of the wing, but beyond is nearly straight, sloping apically so as to make a

58 FOSSIL BUTTERFLIES.

uniform curve with the outer border, which is almost entire as in Parnassius (PI.
I, fig. 5), rather than as in Thais (PI. IT, fig. 4), strongly arched, especially
near the last median nervule, and angulated below where it meets with the regu-
larly and broadly concave inner margin.

In the neuration of the fore wings (PI. III, fig. 1) this genus is peculiar for
the shortness of its cell, which is less than half as long as the wing, and is broad-
est in the middle of its distal half, beyond which it narrows rather rapidly. The
costal nervure terminates a little before the middle of the outer two-thirds of the
costal border. The subcostal nervure emits two superior branches before the cell;
the first is thrown off near the middle of the outer half of the cell and terminates
as far beyond the tip of the costal nervure as it is beyond the middle of the costal
border; beyond the emission of the first superior nervule the subcostal nervure
curves downward away from the costal nervure, with which it had hitherto been
parallel, and throws off the second superior nervule shortly before the apex of the
cell; this nervule terminates exactly at the apex of the wing, but, just before the
tip, divides, sending a short branch to the outer border; about two-fifths of the dis-
tance from the tip of the cell to the outer border, the subcostal nervure divides
into two branches which reach the outer border near the middle of its upper half;
the inferior subcostal nervule leaves the nervure nearly at right angles, but almost
immediately turns and runs subparallel to it and its lower ultimate branch. The
median nervure throws off its first nervule a little beyond the middle of the cell;
its second midway between this and the base of the fourth, and the third midway
between its two neighbors; beyond the emission of the second nervule the nervure
bends upward, and still more on throwing off the subsequent one; the first two
nervules are straight, the upper two arched, and the base of the last is united to
the short basal fragment of the inferior subcostal nervule by a curving vein open-
ing outward, whose general course is nearly at right angles to the costal border.

In the hind wing the relation of the cell to the length of the wing is as in the
front pair; it is broadest at the first divarications of the bordering neryures and
narrows rapidly beyond. The first branches of the subcostal and median nervures
are emitted near the middle of the distal half of the cell, and that of the subcostal

THAITES. 59

is a nearly straight continuation of the basal portion of the nervure; the outer sub-
costal and median nervules are twice as close at base as any of the others, and the
middle nervules divide the space between the first and third; the submedian ner-
vure is parallel to, and scarcely removed from, the inner border.

In the pattern of their markings (Pl. III, fig. 3) the wings of Thaites are
rather simple. The fore wing is provided with four nearly equidistant, nearly
straight, transverse, pale stripes, depending at about right angles from the subcos-
tal nervure, unequal in length and width, the third from the base situated in the
middle of the wing; and also with a submarginal curving row of moderately large,
transversely ovate spots, one in each interspace opening on the outer border, ex-
cepting the subcosto-median and medio-submedian interspaces, all ranged in .a
series curving more strongly than the outer border. The hind wing is nearly uni-
form on the basal half, but beyond is crossed by transverse, curving, dark, cloudy
bands, broadening on the nervures and enclosing between them roundish or trans-
versely ovate pale spots.

The abdomen is stout, half as long as the hind wings, well arched, and the
terminal segment (of the female?) half as long as broad, the segments provided
with a latero-dorsal and pleural row of very small, vertically ovate, pale spots.

This genus differs from Thais (PI. II, figs. 2,4) and the other genera allied
to the swallow-tails in about the same degree as they do among themselves. It
is closely allied to Thais in most particulars; the antennz resemble those of Thais,
more than they do those of other genera, if we except only Sericinus; in the
form of the wings it lies midway between Thais and Archon; as to neuration
the discoidal cell of the fore wings has the form seen in Sericinus, being broadest
apically, while in Parnassius (PI. ITI, fig. 5), Thais and Eurycus it is largest in the
middle; but it is shorter than half the length of the wing, while in Sericinus, as in
all the other genera, it is considerably more than half the length of the wing; the
tip of the cell is limited above, in most of these genera, by the vein closing the cell;
that is, the inferior subcostal nervule originates beyond the tip of the cell; but in
Thais it originates at the tip of the cell, while in Thaites the cell is limited by the
inferior subcostal nervule and the vein closing the cell originates from it; in other
particulars of its neuration it resembles the tailed Sericinus. -

60 FOSSIL BUTTERFLIES,

In design (Pl. II, fig. 3) Thaites recalls none of the recent genera very
closely. In the fore wings it approaches Thais (PI. ITI, fig. 4) rather than the
others, and in the hind wings some species of Parnassius (PI. III, fig. 5). It has
none of the eccentric spots of Parnassius and a darker ground than any of the
modern types. It is wholly unprovided with the strongly marked crescentic spots
of Thais, but in the position, form and arrangement of the principal markings
rather recalls Archon. Excepting Eurycus and some species of Thais, no modern
genera resemble Thaites in the extension of a distinctive pattern upon the hind
wings to or nearly to the extremity of the cell. Whether any of the markings
were accompanied by the brilliant spots often seen in Thais, Archon and Parnas-
sius cannot be determined, but we may presume that they were not, since in these
genera the markings are dark upon a lighter ground, while in Thaites they are
light upon a dark ground,—a combination found among the Papilonid genera,
only in some of the swallow tails.

In the markings of the abdomen, I do not know that we find anything parallel
to Thaites among the Parnassians, but among the neighboring Equites there are
similar examples of rows of small light spots on a dark ground. I haye not been

able, however, to examine this point carefully.

THAITES RUMINIANA Heer MS.
Plate III, figs. 1, 3, 6-10.

Thaites Ruminiana Herr, Climat pays tert., trad. Gaudin, 205 (1861) [absq. descr.]; Sap., Ann. Sc. Nat.
[5], Bot., xv, 348 (1872) [ibid.].

The wings were evidently dark with light markings. On the fore wings the
first transverse stripe (PI. III, fig. 3) extends from the subcostal nervure, midway
between its first divarication and the base of the wing, almost to the middle of the
basal two-thirds of the inner border; it is slender, nearly equal and straight, the
portion within the cell about four times as long as broad; the second transverse
band is the largest, and lies midway between the first and the third, parallel to
them, reaching from the subcostal nervure almost to the inner border; it is straight
and equal, and the portion within the cell (which is half of the whole, although

THAITES RUMINIANA. 61

traversing the cell at its broadest part) is three times as long as broad; the third
transverse bar is in the middle of the wing, smaller than the first and equally slen-
der, extending from the subcostal nervure, just beyond the tip of the cell, almost to
the upper median nervule; it is equal and straight excepting above, where it curves
inward following the border of the cell; the outermost is broader and more irreg-
ular, depending from the first superior subcostal nervule and extending nearly to
the upper median nervule, so that its exterior border just strikes the subcostal ner-
vure at its divarication far beyond the cell; the inner margin is straight and the
spot thus forms a transverse bar, straight and equal above the subcostal nervure,
but with the outer border sloping away so that the lower extremity is twice as
broad as the upper. The submarginal series of spots are of nearly equal size, the
uppermost largest, the next two smallest; each set of three forms a nearly straight
line, but all together they follow a strong curve which approaches close to the
border in the lowest subcostal interspace, being separated from it by but its own
width; above this they recede rapidly from the border, the outer edge of the in-
nermost being next the fork of the second superior subcostal nervule; but below,
the spots are parallel to the outer border and separated by about an interspace’s
width from it; the upper spots are transversely broad ovate; the lower transversely
subquadrate; apparently the fringe is exceedingly short and concolorous as in
Parnassius.

The basal parts of the hind wing are almost uniformly dark, excepting that
there is a paler suffusion in the outer part of the cell; beyond, the wing is clouded
with darker, transverse, strongly curving, powdery stripes; the most conspicuous
of these is one which crosses the wing a little outside the middle of the portion
beyond the cell; it takes its rise in a darker spot, which borders the wing just
above the tip of the upper subcostal nervure, and runs in a nearly straight line,
widening as it goes, to the lowest subcostal nervule, where it reaches its greatest
width, and scarcely narrowing curves around to the inner border a little before its
tip; on the nervules it reaches further baseward and borderward. Between this
belt and another similar but much less conspicuous band, half way between it and
the tip of the cell, are enclosed circular pale spots, one occupying the entire width

of each interspace below the middle subcostal nervule and a portion of the one
MEMOIRS A, A. A. 8. 10

62 FOSSIL BUTTERFLIES.

above it; following the principal dark band are two alternating sets of dark and
light, narrow, inconspicuous, transverse stripes, more or less confused in the middle
of the wing, the dark bands broadening and deepening at the nervures, breaking
the paler bands to a greater or less extent into broad transverse spots; the fringe
appears to be as on the fore wings. Judging from the form of the last abdominal
segment, and the great size of the abdomen, this specimen was probably a female.
Length of fore wing, 25"; breadth of the same, 14:3"; length of antennz, about
6"; breadth of antennz in middle of stem, ‘2; breadth of antenne toward tip
of club, -5™™.

Tertiaries of Aix. Collection of Professor Heer; Zurich, Switzerland.

URBICOLZA = HESPERIDES.
THANATITES Scupper.

Very much of the general appearance of Thanaos Boisd. (PI. II, fig. 2) but
with somewhat differently formed wings and markings which will not accord with
those of the latter genus, although the two genera are certainly nearly allied.

The body (PI. II, fig. 12) is fully as stout as in Thanaos (PI. III, fig. 11),
the tongue at least as long as the thorax, the eyes ovate and larger. than in
Thanaos, and the palpi with the terminal joint proportionally larger, which is an
unusual feature in the Urbicole. The legs are apparently short, the wings ample.
The costal margin of the fore wings is nearly straight, being scarcely arched on
the apical half, the upper half of outer border as in Thanaos, the rest not pre-
served; the costal fold of the male is narrow and extends a very little beyond the
middle of the costal border, while in Thanaos it reaches considerably further; the
hind wings have the general shape of Thanaos, but the upper outer angle is much
more produced, and the base of the costal border is arched only to the degree that
the apex is, and the portion between them is but slightly convex; the outer border
is almost precisely as in Thanaos and the inner border is, doubtless, folded in the
fossil so as to conceal its true character. Very little of the neuration can be
determined, and what can be made out is comparatively unimportant and agrees

‘ with the neuration of Thanaos; the third superior subcostal nervule strikes the

THANATITES VETULA. 63

apex of the fore wing as in that genus. As to the markings, the agreement with
Thanaos is less striking, although the pattern resembles that of Thanaos more
closely than it does that of any other genus. In the fore wings the spot in the
cell of Thanaos is wanting in the fossil, but in its stead there is a costal spot at
the extremity of the costal fold; the subapical spots of Thanaos depending from
the costa are distinctly repeated in Thanatites, and in addition there is a submar--
ginal series of small round spots of which the upper two, in the uppermost in-
terspaces opening on the outer border, are the only ones visible on the fossil
by its mode of preservation. On the under surface of the hind wings of Thana-
tites, there is a regular submarginal series of equal, rather small, round spots, one
in each interspace, placed between the location of the marginal and submarginal
spots which occur in Thanaos, often distinctly, occasionally as faint blurred bands,
as in JZ. Juvenalis (PI. III, fig. 11); the inner of these two series in Thanaos,
which corresponds best to the submarginal series of Thanatites, is irregular instead
of parallel to the border, being always bent inward opposite the cell. Instead of
the spot, placed in the costo-subcostal interspace of Thanaos near the middle of the
wing, and seen distinctly in 7’. Juvenalis, there are two spots, which, with a third
near the base of the wing above the costal nervure, are placed at equal distances
apart and from the costal border; in addition there are two spots, seldom even
indicated in Thanaos, near the centre of the wing, the larger of which is near the
apex of the cell. These differences alone would suffice to show that the fossil can-
not be referred to Thanaos, and, with the other indications we have given, compel
us to place it apart, but in the immediate vicinity of this group of Urbicole.

THANATITES VETULA (Hrypen) Scupper.
Plate IIT, figs. 12, 16.

Vanessa vetula Heyv., Paleontographica, viii, 12-13, Taf. i, fig. 10 (1859).
Araschnia vetula Kirs., Syn. Cat. Diurn. Lep. 179 (1871).

The only notice of this insect that has been published is the original figure

and description of von Heyden. ‘The figure is reproduced in our PI. II, fig. 16.

The description is as follows:'—
1 Paleontogr. viii, 12-13.

64 FOSSIL BUTTERFLIES.

Es scheint diese Art in die Nahe der bei uns lebenden Vanessa Levana zu
gehoren. Sie ist kleiner als diese, indem der Vorderfligel von seiner Basis bis
zur Spitze nur 6}'” misst. Der Schmetterling liegt auf der rechten Seite, wobei
der linke Hinterfligel den linken Vorderfligel véllig bis auf die Spitze und einen
Theil des Aussenrandes deckt. Von diesen Fligeln ist daher nur die Unterseite
sichtbar. Der rechte Vorderfligel ist mehr vorgeschoben und daher ein grosser
Theil seiner Oberseite sichtbar.

Die Flagel sind im Allgemeinen gut erhalten und scheinen am Aussenrande
an einigen [13] Stellen schwach ausgerandet gewesen zu seyn. Sie zeigen auf
der Grundfarbe gréssere, undeutlich schwarze und viele weisse Flecken yon ver-
schiedener Grosse. Auf den Vorderfliigeln zeichnen sich ein grésserer weisser
Flecken, etwa ein Drittel von der Spitze entfernt und nach dem Vorderrande
hinzielend, sowie drei weisse Fleckchen aus, die in einer Reihe in der Nahe des
Aussenrandes stehen. Auf den Hinterfliigeln, etwa ein Drittel vom Aussenrand
entfernt, bilden sechs weisse Fleckchen eine Querreihe. Es ist nicht unwahrschein-
lich, dass die Grundfarbe der Fligel im Leben braun oder rothbraun war, und man
glaubt sogar noch einen schwachen Schimmer von dieser Farbe wahrzunehmen.

Der Kopf ist etwas zerdriickt und zeigt zwei ziemlich lange, zugespitzte, in
die Hohe gerichtete Taster, von denen der eine vom Kopf getrennt liegt. Oben
am Kopf ist noch ein Auge und unten die in einen Bogen aufgerollte Zunge
sichtbar. Die Brust ist undeutlich, der Hinterleib fast ganz durch die Fligel
gedeckt, und von den Beinen sind nur Bruchstiicke vorhanden.

Dark brown or blackish with light markings. On the upper half of the fore
wing (PI. ITI, fig. 12), both above and below, the following markings are found:
a small quadrate spot on the costal border at the extremity of the costal fold;
depending from the costal border between the tips of the second and third superior
subcostal neryures a confluent series of spots extending to the cell at right angles
to the costal margin, narrowing a little in passing downward; and midway between
this and the outer border, in the upper two subcostal interspaces opening on the
outer border, a small round spot; probably similar spots belong in some of the
interspaces below. On the under surface of the hind wings there is a submarginal
series of three small spots along the costa at equal distances apart, the central
one near the middle of the costa, and the basal one nearly midway between it and
the base of the wing; there is also a larger spot near the tip of the cell and a
second smaller one a little below and beyond it; also a submarginal series of spots
as large as that in the cell parallel to the outer border, at about an interspace’s

————L—S ll

THANATITES VETULA. 65

distance from it, one in each interspace. Length of fore wing, 14""; length of
hind wing, 13-65""-; extreme breadth of hind wing, 11:25™™.

The single fossil represented by von Heyden under the name of Vanessa vetula,
is preserved on a greasy, dark brown, thin and exceedingly fragile sheet of “brown
coal,” and is likely to become so affected by weathering as to be almost or quite
indistinguishable in the course of time. Indeed it is excessively obscure at the
present time, and no fossil object I have ever studied has proved so difficult to de-
cipher as this. It represents an insect (PI. III, fig. 12) lying upon its side in a
somewhat natural attitude (compare fig. 11), so that one can see the whole of the
under surface of the left hind wing, the costal quarter of the under surface of the
left fore wing, and a little more than a quarter of the upper surface of the right
fore wing, also of the costal area; the thorax and head with the eyes, the denuded
palpi, the partially unrolled tongue and fragments of the legs in a confused medley
may also be seen, but there is no trace of the antenne, nor of the right hind wing
(nor of the abdomen?). The left hind wing has an immaterial part of its outer
border remoyed, and a small portion of the outer border of the left fore wing is
also wanting, but the corresponding portion of the right fore wing is present.
The markings can only be made out by extreme care, and a very meagre portion
of the neuration, especially toward the borders of the wings, by great patience
and the closest examination; but most of what can be seen of the neuration adds
but very little to our actual knowledge of the animal; it simply adds its testimony
in the same direction as other features of the object.

The illustration of yon Heyden (PI. III, fig. 16) is faulty in several particulars,
but this is not surprising when we consider the excessively obscure nature of the
fossil; it represents the insect as if the under surface of both wings of one side
were seen, the fore wing concealing a portion of the hind; a break in the stone is
taken for the outline of the wing (just above the extremity of the costal border of
the hind wing) and the markings of the two front wings are blended into one; an
abdomen is represented and above it an outline of the inner border of the hind
wing. ‘The fossil has at first sight this appearance, but I think this view is errone-
ous, although on this point one may not speak with confidence, and it is compara-

66 FOSSIL BUTTERFLIES.

tively unimportant. It is remarkable, however, that von Heyden, in his description,
takes the same view of it as I havedone. I have not attempted to give the shading
of the darker parts of the wing, partly from its obscure nature, partly from a
doubt whether they really represent the original markings of the insect; for the
basal half of the under surface of the hind wings, where most of the dark mot-
tling in Von Heyden’s figure occurs, is usually devoid of any such variegation in
the insects of this group; they are almost always of a uniform grayish or brownish
hue. Von Heyden’s figure does not show the division of the palpal joints.
Tertiaries of Rott, Rhenish Provinces of Germany. British Museum.

URBICOLA — ASTYCI.

PAMPHILITES Scupprer.

This genus belongs to the Astyci and falls in the neighborhood of Pansydia
and Carystus, if we take as an illustration of the latter group the Hesperia Lucasii
of Fabricius. The former genus has a male with a discal dash, the latter without
one. As the fossil species is represented by a single fore wing of what is probably
a female, it is impossible to say into which category it would fall. The costal border
(Pl. IH, fig. 18) is almost exactly straight throughout; next the base, however, it is
arched a little and it slopes slightly downward on the apical fifth to a rather sharply
defined apex; the outer margin is gently and almost regularly convex, but with its
greatest convexity a little above the middle, and at its upper end is at right angles
to the tip of the costal margin; the lower angle is rounded off and the inner mar-
gin is slightly sinuous, being hollowed in the middle; the wing is slightly more than
twice as long as broad. In all these respects it agrees far better with Pansydia
(Pl. TH, fig. 15) than with Carystus (PI. IIT, fig. 13). Indeed, excepting in the
greater length of the wing and the lack of any change of direction in the outer
border at the tip of the lowest median nervule, the form of the wing scarcely dif-
fers from that of Pansydia Mesogramma.

In neuration it agrees better with Pansydia than with Carystus. Poey’s fig-
ure, which for want of better material I have been forced to copy in illustration, is

PAMPHILITES. 67

not executed with sufficient care, for of the first and second superior subcostal
nervules he has made but one. The principal difference between Pansydia and the
fossil genus is in the fourth superior subcostal nervule; in Pansydia this terminates
upon the costal border just before the apex of the wing, while in Pamphilites it
terminates on the outer border just below the apex of the wing, bringing the latter
into a different interspace in the two genera. From Carystus it differs, not only
in having a proportionally shorter cell, but in the same point as that in which it
is distinguishable from Pansydia; and further in the uppermost median nervule,
which in Carystus is thrown off abruptly from the nervure just beyond its second
divarication and which, by curving strongly, makes the upper median interspace
of nearly equal width throughout; while in Pamphilites, the nervule parts gently
from the neryure like the others, and at some distance beyond its second divari-
cation, passing in a regular curved line to the outer border, and causing the upper
median interspace to increase in breadth throughout the whole of its basal half.

In the disposition of its spots, Pamphilites (PI. II, figs. 14, 17) agrees per-
haps better with Carystus (PI. III, fig. 19) than with Pansydia (PI. III, fig. 15).
This is especially true of the large spots in the cell and in the lower two median
interspaces; although in Carystus the spots of the median interspaces are further
removed from the base than in Pamphilites, while the opposite is true of the spot
surmounting the submedian nervure; the submarginal spots beyond the cell of
Pamphilites are wanting in Carystus, and the latter genus has but two of the three
subcostal spots of Pamphilites. The spots of Pansydia are smaller and far less
conspicuous than in Pamphilites, that of the cell being reduced almost to a dot;
the median spots are howeyer large, though removed farther from the base, as in
Carystus; there is also a small spot in the upper median interspace, but further
from the margin than in Pamphilites and unaccompanied by any spot in the
interspace beyond the cell; as in Carystus, the spot surmounting the submedian
neryure is further from the outer margin than in Pamphilites, but the subcostal
spots accord very well with those of the fossil.

By these considerations it would appear that Pansydia is to be placed between
Carystus and Pamphilites, the latter being more nearly related to Pansydia than

68 FOSSIL BUTTERFLIES.

to Carystus, leading us to believe it more probable that we are dealing with a
female, whose partner was possessed of the ornament of a discal dash of spe-
cialized scales. The species of Pansydia are smaller than those of most of the
neighboring genera, but Pamphilites abdita is somewhat smaller even than Pan-

sydia mesogramma.

PAMPHILITES ABDITA Scupper.
Pl. III, figs. 14, 17, 18.

Upon a dark, uniform, probably blackish brown ground, the fore wing of this
butterfly was provided (in the female?) with three large spots, three small spots,
and two dots of a vitreous appearance, besides other light streaks or powdery
spots. The three large spots are probably peculiar, in their present extent, to the
female; they consist (PI. III, figs. 14, 17,) of one spot in the cell and one in each
of the lower median interspaces; the cellular spot crosses the cell, is sublunato-
quadrate, its exterior edge concave, extending from the origin of the third supe-
rior subcostal nervule to just beyond the second divarication of the median nervure,
being directed in the upper half of its course toward the base of the second
median nervule; the spot is narrower above than below, the upper half having an
outward as well as upward inclination, the lower margin straight, the interior
margin subsinuate, convex, reaching from midway between the base of the first
and second superior subcostal nervules to just beyond the middle of the space
between the base of the first and second median nervules. The spot in the lowest
median interspace is nearly or quite as large as the previous, but longitudinal
instead of transverse, and as broad as the interspace; excepting for a little spur
above on the inner side, which runs a little way toward the base, the centre of the
spot would lie just below the second divarication of the median nervure, but by
means of this slight spur the spot extends baseward half way from the second
to the first divarication of the median nervure; at the outer extremity the spot
terminates squarely and next the lowest median nervule is two-sevenths the length
of that vein. The spot in the middle median interspace is much smaller, subtri-
angular, filling the whole breadth of the interspace, half as long again as broad,

PAMPHILITES ABDITA. 69

its inner tapering extremity situated just below the final divarication of the median
neryure. The three small spots in the lower three subcostal nervules are seated
one above the other, their inner margins on a line and nearly at right angles to the
costal margin; they are quadrate and increase slightly in size below, the upper one
being square, the lower longitudinally oblong; they are situated midway between
the discoidal spot and the apex of the wing. The two dots are situated one just
above the other in the middle of the upper median and subcosto-median inter-
spaces, midway between the spot in the lower subcostal interspace and the outer
border; the lower is slightly the larger, but not more than one-fourth the size of
the uppermost subcostal spot. Seated upon the submedian nervure, its centre
below the outer edge of the lower median spot, is a pale, powdery spot, twice as
long as broad and about one-third the width of the interspace; outwardly it
merges into the ground color; there are other pale spaces in the wing, looking
somewhat as if due to attrition; especially in the cell on either side of the discoidal
spot, at the extreme base of the lower median interspace, and along the lower bor-
der of the medio-submedian interspace. Length of wing, 15°75™", length of inner
border, 9°5™"; breadth of wing across the middle, 7:25", breadth of wing across
outer margin, 9°5™™.
Tertiaries of Aix, Provence, France. Museum of the City of Marseilles.

MEMOIRS A. A. A. 8. 1l

70 FOSSIL BUTTERFLIES.

COMPARATIVE AGE OF FOSSIL BUTTERFLIES.

All the well determined fossil butterflies come from one of three localities,
Aix, Rott and Radoboj, all belonging to the tertiaries of Europe. Others are
reported, as will be seen further on, to have been found in Prussian amber; and it —
is not in the least improbable that they have been or may be. These would be of
about the same age as the oldest of the others, those of Aix. Of the Aix fossils,
which belong to the upper Eocene, or to speak more definitely, the Ligurian,
Neorinopis sepulta, Lethites Reynesii, Thaites Ruminiana and Pamphilites abdita
(the first described by Boisduval, the rest by myself) come from the calcareous
marls of the gypsum quarries, the only bed in which insects had been found when
visited by Messrs. Murchison and Lyell in 1829. Coliates Proserpina, however,
described here for the first time, was taken from strata beneath these, and there-
fore, at least until we have more precise knowledge concerning the remains of
butterfly larvee in amber, may be considered the oldest known butterfly. Count
de Saporta writes me concerning this fossil, the discovery of which is due to him,
as follows: —“ Cette empreinte ne provient pas des platriéres méme, c’est 4 dire des
galeries qui servent 4 l’exploitation du Gypse; mais d’une assise ou groupe de
couches immédiatement inférieure. Vous verrez cette provenance indiquée pour
un grand nombre de mes espéces; dans ce cas, elles ne proviennent par des ouvriers
mais je les ai recueillies moi méme en suivant les lits sur les points ov ils affleurent
au dehors.”

The next in order, approaching recent times, are the lignite beds of Rott in
the basin of the Rhine, which belong to the Aquitanian or the upper part of the
lower Miocene. Thanatites vetula (described by Hayden) is the only butterfly
known from this division of the Tertiaries.

The most recent beds containing fossil butterflies are the lacustrine deposits
of Radoboj in Croatia, Austria. These belong to the Mayencian or lower portion
of the middle Miocene, and have furnished Hugonia atava, Mylothrites Pluto,

PROBABLE FOOD-PLANTS OF TERTIARY CATERPILLARS. 71

another fragment possibly referable to Mylothrites, and Pontia Freyeri, all de-
scribed by Heer. Two of the genera of these more recent beds contain repre-
sentatives now living in the same region; but none of the older beds have yet
furnished butterflies referable to modern genera. .

It is rather extraordinary that the upper Miocene beds of Giningen, Bavaria,
which, if we except the amber, have furnished almost more insects than all the
other beds of fossil insects of the world together, and which are more recent than
any of those. in which butterflies have been found, have yielded scarcely any re-
mains of Lepidoptera (one species) and none whatever of butterflies.

PROBABLE FOOD-PLANTS OF TERTIARY CATERPILLARS.

Of the five butterflies from Aix, two belong to the Oreades (eorinopis
sepulta and Lethites Reynesii) the food of whose caterpillars at the present epoch
has invariably been found to be either Graminex or, occasionally, Cyperacee.
Both of these groups are present in the deposits of Aix, the former being repre-
sented by ten species of Poacites, and the latter by a Cyperites;! and it is
in the highest degree probable that these formed the sustenance of the Oreades
of that epoch. A third species (Pamphilites abdita) belongs to the Astyci, a group
whose principal food is the same family of plants, Gramines, although some
species haye been found also upon Althea, Malva and Lavyatera (Malvacee), Tri-
folium, Coronilla and ? Lespedeza (Leguminose), Plantago (Plantaginacez), and
Maranta (Scitaminer). Of these families the Leguminose only are found at
Aix, and in abundance, even including a plant doubtfully referred to Trifolium.
It is, however, far more probable that Pamphilites lived upon grasses; and it is
not a little strange that the Graminex, the probable food-plants of three of the
five butterflies known from that fauna, were among the rarest of the plants; that
is, their proportion to the whole phanerogamic flora was about the same as now

1Saporta. Revision de la flore des gypses d’Aix. Ann. Sc. Nat. [5] Bot., xv, 284.

72 FOSSIL BUTTERFLIES.

obtains in New Guinea or New Grenada, countries the least favored in this
respect.!. The proportion of the Gramines and Cyperaceew to the whole of the
Phanerogamia in Europe of to-day is, probably, about the same as in the United
States (more than seventeen per cent.) and much greater than in the East Indies.
The limited number of known fossil butterflies does not give great weight to any
general considerations based upon them, but it may at least be worth while to
remark that Aix, in Eocene times, had, in the point referred to, an assemblage of
plants much better comparable with the East Indian flora of the present day than
with the modern European flora, the proportion of known Graminee, etc., to the
Phanerogamia being five per cent., while the proportion of its grass-feeding but-
terflies to the other rhopalocerous Lepidoptera is sixty per cent. To judge simply
by the catalogue of the East India Museum, the only authority upon East Indian
butterflies extant, the present proportion of gramnivorous to non-gramnivorous
butterflies is as 1 : 5-2, while in Europe it is as 1:3. Eocene Aix, then, had a
European proportion of Satyrids, composed, as will be seen, of species of an In-
dian aspect, feeding upon plants essentially temperate, but, as in tropical countries,
numerically unimportant.

The Danai, to which the fourth species from Aix (Coliates Proserpina) belongs,
feed almost exclusively upon Leguminose, and these have recently been found in
great abundance at Aix. Count de Saporta enumerates one species each of ? Tri-
folium, Caragana, Ervites, Sophora, Micropodium, Cercis and Gleditschia, two of
Phaseolites and six of Cesalpinites, belonging to the Papilionacex, besides nine
Acacias and a Mimosa of the Mimosex, and four species of uncertain relations;
making a series larger than he has found in any other family.”

Of these, two species of Phaseolites, one of Sophora, eight of Acacia and two
of Leguminosites are specified as coming from the lower beds, where Coliates
itself is found. But Coliates is most closely allied, as we have said, to a group of
Indian forms, and the food plants of their caterpillars is almost wholly unknown.

1“La proportion des Graminées relativement au total des flores intertropicales actuelles de l’ancien et du nouveau continent,
Phanérogames, qui est de 45 sur 100, est en rapport avec les min- mais elles atteignent une proportion de 13 pour 100, pour l’en-
ima relatifs de cette famille, tels qu’on les observe & la Nouvelle- semble des Phanérogames, proportion parfaitement en rapport
Guinée et & la Nouvelle-Grenade.” Saporta, loc. cit., 292. avec celle de 12 sur 100 qui est fréquente, selon M. de Candolle,

*“Dans Ia flore des gypses d’Aix, non-seulement les Légu- dans certai régions chaudes, telles que Timor, le Congo, etc.”
mineuses ocenpent le premier rang, comme dans la plupart des = Saporta, loc. cit., 292.

PROBABLE FOOD-PLANTS OF TERTIARY CATERPILLARS. 73
A species of Delias, however, to which genus Coliates has been specially compared,
is stated to feed, not upon a leguminous plant, but upon Dioscorea, one of the Yam
family; and the presence in Aix of a species of a closely allied group, Smilax rotun-
diloba Sap., is announced by Count de Saporta. It is not improbable, therefore,
that Smilax rotundiloba was the food-plant of the larva of Coliates Proserpina.!

The fifth Aix species is Thaites Ruminiana. It is most nearly allied to Thais
of the present day, though it bears certain relations, as we have seen, to neighbor-
ing genera. Thais feeds principally at least upon Aristolochia? and so, too, do
Ornithoptera, Archon and some genera of swallow-tails; indeed, this seems to be
a favorite food-plant with insects of this character. Parnassius, however, feeds on
Sedum, Telephium, Sempervivum and Corydalis, especially on the first-named, one
of the Crassulacese; but nothing very closely allied to this is specified by Sa-
porta from Aix; neither, also is Aristolochia, but it has been found not only in.
Radoboj’ in the Mayencian, but also, according to Heer, at Hohe Rhonen in Swit-
zerland, which belongs to the Aquitanian, and has at least one plant (Zaurus pri-
migenia Ung.) in common with Aix. It seems, therefore, highly probable that
either Aristolochia. nervosa Heer, A. Aesculapi Heer, or a distinct species of the
genus will yet be discovered at Aix,‘ and may then be considered, as with little
question, the food-plant of Thaites Ruminiana. If it be deemed hazardous to ven-
ture such an opinion, attention is called to the two following passages; the first is
from the introduction to Heer’s paper on the fossil insects of Aix:° “Dass indessen
auch Weiden oder Pappeln [Populus] sich vorhanden, dirften der Bythoscopus
muscarius und die Aphrophora spumifera [Homoptera] anzeigen, deren analoge
lebende Arten besonders auf den Blaittern und Zweigen dieser Baume sich umher-
treiben.”
the Climate and Vegetation of the Tertiaries’ by Gaudin: “Le Poacites ciliatus

The second is a note in the errata to the translation of Heer’s work on

1Since this was written, Count Saporta writes me: ‘Le genre
Smilax est on des genres tertiaires les plus frequents. J’ai aussi
signalé dans le dépét voisin de St. Zacharie (étage Tongrien infé-
rieure [and therefore but slightly more recent]) une feuille qui m’a
paru devoir se ranger parmi les Dioscorées.”

2 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, bunt the statement in Fuessly’s Mag-
azin has never, to my knowledge, been confirmed.

3* Nouns p’avons pas encore ré d’Aristolochia dans les
gypses d’Aix; mais existence du genre dans le tertiaire moyen

ne saurait @tre mise en question, depuis que nous avons entre les
mains une superbe empreinte de Radoboj (Aristolochia venusta
Sap.), qui dénote une forme voi des Aristoloches & feuilles
persistantes et demi-coriaces, comme IA. reticulata Nutt. de Vir-
ginie.” Saporta, loc. cit., 342-3.

#In a recent letter from Count Saporta he remarks: “ Relative-
ment au Thaites Ruminiana, je n’ai pas encore decouvert a Aix de
vestiges du genre Aristolochia, mais ce genre devait y exister.”

5 Vierteljahreschrift naturf. Gesellsch. Zurich, i, 12, 1856.

Recherches sur Je Climat et la Vegetation du pays tertiaire,
4to, 1861.

74 FOSSIL BUTTERFLIES.

Sap. n’est pas une glume de Graminée, mais plutét une bractée ciliée de Peuplier.

Elle doit @tre probablement rapprochée d’une empreinte . . . prove-
nant des mémes couches et qui se rapporte également au genre Populus. Les
organes voisins de ceux du Pop. Euphratica Oliv. dans la nature actuelle dénotent
existence, 4 ’epoque des gypses d’Aix, d’une espéce de Peuplier dont les feuilles
sont encore inconnues, comme celles de lAlnus eryptophylla Sap., mais que
M. Heer avait indiqué d’avance, en se fondant sur l’observation d’un insecte fossile,
le Bythoscopus muscarius! Nouvelle preuve du secours que peuvent se préter en
paléontologie les diverses branches de histoire naturelle.”

The single species from Rott, Thanatites vetula, is closely allied to the mod-
ern Thanaos, whose species are numerous and feed upon a variety of plants,
belonging to the families Crucifere, Leguminose, Umbellifere, Cupulifere, Be-
tulacew and Salicacesw. Most of the genera belonging to its tribe feed upon
Leguminosx, and these are the usual food plants of the species Thanaos also;
whence it is probable that Thanatites had a similar taste. Now in the very beds
of Rott, in which this butterfly was found, occur species of Betula, Salix and
Populus, with numerous Querci and no. less than eleven genera of Leguminose,
mostly belonging to the Papilionace; they are Templetonia (1 species), Robinia
(2), Colutea (1), Phaseolites (2), Sphinctolobium (1), Dalbergia (1), Haematoxy-
lon (2), Gleditschia (2), Cassia (3), Ceratonia (1), and Acacia (2). It is proba-
bly among these, and perhaps with greatest probability among the species of
Hematoxylon and Gleditschia, that the food plant of Thanatites must be sought.
Should leaves be found, in which a portion is bent over as if to form a nest, they
should be submitted to the scrutiny of some one familiar with the larval habita-
tions of Thanaos Tages; and should traces of silken fastenings be found in con-
nection with them, or the marks of nibbling at the edges, the plant to which they
belong may be considered with strong probability as the food of Thanatites vetula.

The only butterfly found at Radoboj belonging to an extinct genus is Mylo-
thrites Pluto, and this is a member of the same general group as Coliates, and
feeds probably upon Leguminoss; for it is not so closely allied to Delias as Coli-
ates is, but is more nearly related to Hebomoia, one of whose species, found in the

PROBABLE FOOD-PLANTS OF TERTIARY CATERPILLARS. 75

East Indias, feeds upon Capparis.!. One species of Phaseolites, one of Sophora
and four of Cassia, namely: C. hyberborea Ung., C. phaseolites Ung., C. lignitum
Ung., and C. ambigua Ung., are recorded from Radoboj, and as Cassia is a favorite
food plant among the larger species of Danai at the present day, we may fairly
presume one of these Cassis to have afforded nourishment to Mylothrites Pluto.
Moreover, no less than thirty-one species of Leguminosz in general, or between a
ninth and a tenth of the whole known flora, are given by von Ettingshausen as
occurring in Radoboj; so that in any case our Mylothrites must have found abun-
dance of palatable food.

The food of Pontia Freyert is doubtful. All the living species of the genus
so far as known, feed upon Cruciferze; within this family they do not seem to be
at all particular, making use of a large number of genera, but in only a single
instance are they known to attack the leaves of a genus (Reseda) belonging to an
adjoining family. Cruciferee, however, are excessively rare in the tertiaries of
Europe, two species only being recorded, and this from the comparatively recent
beds of G&ningen. This is unquestionably due simply to the nature of the plants
themselves, which scarcely could leave any trace of their existence; the almost
complete absence of the herbaceous families of plants, even in the later tertiaries,
is doubtless due to this fact. The plants nearest related to the Crucifere found
near the horizon of Pontia Freyeri are a species of Nelumbium from Ginzbourg in
the Mayencian, and of Terminalia (7. radobojensis Ung.)—one of the Calyci-
flor, from Radoboj itself. Perhaps in the absence of better evidence we may
provisionally consider the latter to have been the food plant of P. Freyeri.

A single Radoboj species remains, Hugonia atava. The recent species of
Eugonia feed particularly on Salix, Populus and Betula; also upon Ulmus, and
occasionally on Ribes, and even on Hippophae. The first three seem however to
be their proper food; and since the tertiaries of Radoboj contain fossils of all these
genera, we need look no farther. There are specified: Salix apollinis Ung., sp.,

11 venture to give one more extract from a recent letter re- nourissant de Capparis. Les Capparis ont du exister, mais leur
ceived from Count Saporta, although he writes:—“je vous écris _feuilles sont difliciles & distinguer & cause de Vabsence de carac-
n’ayant sous les yeux ni mes livres ni mes collections, ce qui en- _ teres différentiels; leur forme et leur nervation peu visibles doivent
levera nécessairement un peu de précision & quelques-unesde mes _les fuire confondre avec beaucoup d’autres. Il me semble pourtant

Fd que des Capparis ont été signalés soit & Radoboj, soit & Hoering

“Tl est bien plus difficile de justifier par des exemples tirés en Tyrol, dépét un peu plus anciens [Tongrian], mais en lab-

de la nature des plantes la présence & Radoboj d’un insecte se _— sence de mes livres je ne saurais vous l’affirmer.”

76 FOSSIL BUTTERFLIES.

Populus latior Br., P. mutabilis Heer., P. Heliadum Ung., Betula Dryadum
Brongn. and B. prisca Ett. Three species of Ulmus are also recorded from the
same place.

Excepting in a single case, there is then no difficulty in finding, in the very
beds in which the butterflies occur, remains of plants, which in all probability |
served them as food during the larval stage; and even in this single instance, a
plant not far removed from those upon which species of the genus now feed, occurs
in the same strata.

PRESENT DISTRIBUTION OF BUTTERFLIES MOST NEARLY
ALLIED TO FOSSIL SPECIES.

To discuss this question properly we must consider the butterflies of each

geological horizon separately.

BUTTERFLIES OF THE LIGURIAN (Upper Eocene).

The nearest living ally of Neorinopis sepulta is, with little doubt, Neorina
Lowi, which, like the other members of the genus, is found in the Indo-Malayan
region. The same is strictly true of the species of Zophoessa, Debis and Lethe,
with which we have been obliged to compare this fossil. Ccelites has also been
used in comparison, and most of the species of this group belong to the same
region, although one is described by Felder from Celebes on the confines of the
Austro-Malayan region. We have also pointed out (as Butler has done, but in
incorrect points) its relation to Antirrhea, a Brazilian genus, but this is too distant
to be given much weight. The closest allies of WV. sepulta are to be found in the
Indo-Malayan region.

The same is true, but not to so striking a degree, of Lethites Reynesti.
We have compared this also to Debis, Lethe and Neorina, and especially to the
two former; and all three of these genera, which are certainly its nearest allies,

lod

PRESENT DISTRIBUTION OF ALLIED SPECIES. 17

are strictly confined to the Indo-Malayan region. It is, however, also related, but
in a secondary degree, to Enodia, Cercyonis and Maniola, which are genera apper-
taining to the north temperate zone of both hemispheres.

Coliates Proserpina finds its nearest living representatives in the genus
Delias, which also is strictly confined to the Indo-Malayan region. Thyca and
Prioneris are closely related, the latter of which is limited to the same district and
the former to the Indo-Malayan and Austro-Malayan regions.

Thaites Ruminiana is represented in recent times by the genus Thais, which
is confined to the Mediterranean district, within which Aix lies. An allied
genus, Archon, is also restricted to the same region. Sericinus, however, and
Eurycus, with which we have been obliged to compare it in many points, are
found only in the East, the former in China, the latter in Australia; while on the
other hand, Parnassius, a genus it quite as much resembles, is limited to alpine
and subarctic regions of the northern hemisphere.

The relations of Pamphilites abdita are very different. I have searched care-
fully for very closely allied forms among East Indian Urbicolx; but, while it doubt-
less is not far removed from some of them, its more intimate relationships are
certainly with insects from tropical America and especially with Pansydia and
Carystus.

Three out of the five Aix butterflies, therefore, find their nearest living allies
in the Indo-Malayan region, one is most closely related to forms now found ‘in

tropical America and one is at home in its own resting place.

BUTTERFLIES OF THE AQUITANIAN (Lower Miocene).

Thanatites vetula is the only butterfly yet found from this horizon, and this is
closely related to Thanaos, a genus belonging to the north temperate zones of
both hemispheres, but vastly more developed in the new world, which has at least
four times as many species as the old, some of them extending into the subtropical
regions. The genera adjacent to Thanaos are purely American, although tropical
or subtropical, and therefore the Aquitanian butterfly looks toward subtropical
North America for its relatives of the present day.

MEMOIRS A. A. A. 8. 12

7s FOSSIL BUTTERFLIES.

BUTTERFLIES OF THE MAYENCIAN (Middle Miocene).

Only a single one of these butterflies, Mylothrites Pluto, belongs to an extinct
genus. Its nearest living representatives are to be looked for in the genera Mylo-
thris and Hebomoia, the former of which finds its highest development in torrid
Africa, while the latter is confined to the Indo-Malayan and Austro-Malayan
regions.

The other two belong to modern genera, Eugonia (7. atava) and Pontia (P.
Freyeri). These two genera are very similar in their distribution, spreading, like
Thanaos, above referred to, over the north temperate regions of both hemispheres.
Eugonia, however, is represented equally in Europe and America, while Pontia is
considerably richer in species in the Old World than in the New; yet when we look
into the distribution of the neighboring genera we shall find a result somewhat
similar to the case of Thanaos. Taking into consideration, in the one case, the
present distribution of the genera Hypanartia, Polygonia, Papilio and Hamadryas,’
and on the other of Neophasia, Tatocheila and Leptophobia, we shall find that the
largest development of these groups of genera has been in the New World rather
than in the Old, but in those parts of the New World which lie on the tropical
confines of the temperate zone. -

Two of the more recent species of fossil butterflies are therefore at home
where they are found, although the present development of the group of genera
to which they belong finds its fullest expression in America; while the third
species follows most of those from the lower tertiaries in seeking its allies of to-
day in the tropics of the old world.

Undoubtedly the material at our disposal is, as we have already remarked, far
too meagre to present any generalities of importance, so long as they are unsup-
ported by external proof. This aid we can claim in considering the facts we have
presented concerning the present distribution of the genera of butterflies most
nearly allied to those once living in the neighborhood of Aix. The careful re-

*I use these genera in the sense indicated in my Historical Sketch of generic names. Proc, Am. Acad. Arts, Sci., X, pp. 91-293.

PRESENT DISTRIBUTION OF ALLIED SPECIES. 79

searches of Count Saporta upon the rich flora of this region at the same epoch,
points to very nearly the same results as are here indicated. In his Hxamen des
flores tertiaires de Provence,‘ when writing of the characteristics of the Aix flora,
Count Saporta says (page 150) that about one-fifth of the families represented in it
are now strangers to Europe; that fifty-one genera have an exotic and more or less
tropical aspect, and that forty out of seventy-four, or about one-half, if not exclu-
sively tropical, inhabit the warmer parts of southern regions, or, in small numbers,
temperate extra-European countries. The result is still more striking, if species are
considered, of which there are at least eighty whose individual analogy with living
species is sufficiently clear to yield results of great probability. “De ces espéces,”
to use his own words, “12 seulement correspondent 4 des espéces de l'Europe
moyenne, 6 a des espéces de Europe méridionale, 18 en tout. Les espéces cor-
respondant 4 des formes de Amérique septentrionale ou des régions élevées de
Amérique tropicale, sont au nombre de 10; celles qui répondent a des formes
de Amérique tropicale s’éleventa9 . . . 3; 3 correspondent 4 des espéces du
Cap et 2 a des espéces des iles Atlantiques ‘et de la Barbarie; 14 representent des
formes particuli¢res aux Indes ou aux iles de lArchipel indien et 30, enfin, cor-
respondent 4 des formes australiennes. Le groupe australien est done le plus
considérable, si on les prend isolement. Hn les réunnissant, on voit que sur les
80 et quelques espéces, 28 4 30 seulement correspondent a des formes habitant
aujourd’hui Europe et Amérique du Nord, en y comprenant méme les parties
méridionales de ces continents; tandis que 57 au moins, soit 60 en nombre rond,
representent des formes tropicales ou subtropicales, et dans ce nombre 40 au moins,
c’est-a-dire la moitié du nombre total se rapportent au Cap, aux [151] Indes ori-
entales ot 4 l’Australie; de sorte que le caractére dominant de cette flore est encore
Austro-indien, quoique dans une proportion déja décroissante par rapport 4 Age
précédant.”

This was published in 1861, and would accord entirely with what we know of
the butterflies of Aix and their nearest allies. But eleven years later, after study-
ing the great amount of material which had meanwhile accumulated, Saporta seems

1 Heer et Gaudin, Climat du pays tertiaire, pp. 133-171.

80 FOSSIL BUTTERFLIES.

to have reached different conclusions, for in his Revision de la Flore des Gypses
d@ Aix he states that the affinities of the eocene vegetation of Aix are with south-
eastern Asia and with Africa, and lists of analogous species are given, showing
that twenty-two Aix species are to be compared with similar types in Asia, and
forty with those of Africa. So that African forms much surpass the Asiatic in the
eocene flora of Aix. This is particularly true, he says, with reference to the region
of Africa between Abyssinia and the Cape of Good Hope. “C’est la évidemment
le pays qui nous offre le tableau le plus ressemblant de ce que devait étre le midi
de la France, et c’est aussi vers ce méme pays, ne loublions pas, que nous avons été
ramenés par l’examen des autres élémens de la flore, spécialement par la proportion —
relative des deux grandes classes et des familles prédominantes.”’ The African
element seems to be almost altogether wanting in the eocene butterflies, while the
Asiatic predominates. In a chart accompanying Count Saporta’s paper, however,
he represents the present limits of the principal genera noticed in the flora of the
gypsum of Aix by means of colored lines. These lines cluster remarkably along the
southern borders of Asia and extend over a large part of Africa and across the
ocean to America, and particularly toward the southern United States and the
Antilles. Based on the distribution of these principal genera alone, the flora of
the southern border of Asia would show a closer affinity to that of eocene Aix
than would that of any equivalent belt in Africa; and if we may suppose that our
relics of butterflies represent the principal genera then existing, we should trace a
somewhat similar chart, but for the entire absence of African types; for subtropical
American types mingle with those of the Mediterranean district and especially with
those of the Indo-Malayan region. Count Saporta shows in his memoir just
quoted, as before, that the relations of the eocene flora of Aix to that of the present
Mediterranean basin were more restricted than its relations to exotic types, but in
a letter to me he writes: “Ces affinités [les affinités présumées de la flore d’Aix]
sont d’une part avec la région Méditerranéen, de l'autre avec Afrique et les Indes
orientales. Les affinités miocénes avec Amérique sont postéricures.” These later
American affinities are, however, foreshadowed among the plants and also, as we

1 Ann. Se. Nat., [5] Bot., xv, 922.

PRESENT DISTRIBUTION OF ALLIED SPECIES. 81

haye seen, in the Pamphilites of eocene Aix. They appear again, and very de-
cidedly, when we reach the miocene itself, for the affinities of the butterfly from
Rott, and two of the later butterflies from Radoboj (where first we meet with truly
modern types), are certainly with America in the first instance, and secondarily
with the whole north temperate zone. While the last of the Radoboj butterflies
shows still the remains of the earlier affinities of the Aix flora in finding its nearer
existing types in Africa and southeastern Asia. The results we reach in consider-
ing the Aix butterflies are not, however, in accordance with those drawn from the
insects of the same locality by Professor Heer. He writes:'—

“A Radoboj, . . . on rencontre une plus forte proportion de formes
tropicales [than at Giningen].

Cette faune des insectes s’harmonise parfaitement avec le flore de Radoboj qui,
ainsi que nous l’ayons prouvé précédement, a un caractére plus méridional que celle
d’ningen; ce qui s’expliquerait par sa plus grande ancienneté.

Comme il résulte des recherches de M. G. de Saporta qu’Aix appartient 4
Pétage ligurien, on devrait s’attendre 4 y rencontrer encore plus de formes tropi-
cales q’u 4 Radoboj. C’est tout le contraire, si bien qu’en m’appuyant sur la faune
et en yoyant que Aix avait 10 espéces en commun avec Radoboj et 4 avec Ginin-
gen, javais rapporté précédemment les terrains d’Aix 4 la méme époque que ceux
de Radoboj et je les avais rangés dans le Mayencien. Quatres genres ont disparu.

Tous les autres genres vivent encore dans la Provence, mais ce sont,
comme & Ciningen, presque tous des genres qui occupent une aire géographique
trés vaste. . . . On ne peut pas dire que la faune des insectes d’Aix contredise
positivement Vidée que cette localité avait un climat sous-tropical, cas presque tous
les genres que l’on y a obseryés jusqu’a présent s’étendent jusque dans la zéne
sous-tropicale, néanmoins cette faune ne fournit que bien peu de preuves positives,
tandis que, comme M. de Saporta l’a démontré, la flore est riche en formes méridio-
nales.”

It should be remarked, however, that the insect fauna of Aix is as yet little

known; that these observations of the learned Zurich Professor were founded upon

1Climat du pays tertiare, ed. Gaudin, p. 205.

82 FOSSIL BUTTERFLIES.

a material exceedingly meagre, in comparison with the present vast accumulations
of the museums of Marseilles, Paris and Aix; we may hope soon to become familiar
with them through the careful researches of M. Oustalet; and these will show that
the beds of Aix are, perhaps, even richer in fossil insects than those of Giningen.

The American affinities of the Rott butterfly are in entire harmony with what
is known of the other insects of the lignites of the Rhine, where, says Professor
Heer:!—“On retrouve également des types américains, qui appertiennent. a ’Amé-
rique tropicale et sous-tropicale.”

As to the flora of Radoboj, Professor Heer writes in the work just quoted (p.
96): “Les plantes de la zéne tempérée sont représentées plus fortement qu’d
Sotzka,” and of the latter place he says (p. 95), after speaking of types of the
temperate zone: “Cependant ces espéces se trouvent fort 4 l’arriére-plan en com-
parison des formes tropicales et subtropicales, parmi lesquelles prédominent .
les formes indo-australiens; neanmoins les formes américains, loin d’y faire défaut,
sont représentées par des types assez nombreux et nettement accusés.” As a
whole, therefore, the affinities of the tertiary butterflies seem to be precisely what
we should have anticipated from a study of the vegetation of the period.

- We close this portion of our subject with a tabular view of the results we
have reached in considering the affinities of the tertiary butterflies with living
types, in which the countries, where the living allies of the fossil forms are now
found, are placed in the right-hand columns according to the degree of affinity of
their inhabitants to the tertiary species against which they are placed.

1 Loe. cit., p. 205.

GENERAL RESUME. 83

DEGREE OF AFFINITIES.

NAMES OF BUTTERFLIES.
FIRST DEGREE. SECOND DEGREE. THIRD DEGREE. FOURTH DEGREE.
Neorinopis pote, Indo- Malayan. Austro-Malayan. S. American,
g North temperate
3 Lethites Reynesii. Indo-Malayan. Zone 1
a
i Coliates Proserpina. Indo- Malayan. Austro- Malayan.
!
Chinese and Australian.
3 Thaites Ruminiana. Mediterranean. Subarctic and Alpine.
Pamphilites abdita. Tropical America. Indo- Malayan.
z g Subtropical North North temperate
3 Thanatites vetula. Atnarica, Sone
2
4 Ce Subtropical temperate! North temperate
: Engonia 4
3 3 Subtropical temperate} North temperate
cf Pontia Freyeri ‘America. zone.
pie
2 Indo~ Malayan
3 Mylothrites Pluto. African, Austro- Malayan.

GENERAL RESUME, WITH NOTICES OF UNDETERMINED FORMS.

Nine well authenticated fossil butterflies are now known, all from the Euro-
pean 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 knowl-
edge, then, places the apparition of butterflies towards the end of the lower terti-
aries.

84 FOSSIL BUTTERFLIES.

As a general rule the specimens thus far discovered are in a fair state of pres-
eryation, and especially are those parts preserved which enable us, with consider-
able confidence, to determine their exact affinities. Three of these insects belong
to the highest family of butterflies, Nymphales, four to the Papilionidx, and two
only to the Urbicole. If it be considered probable that the lowest of these fami-
lies 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 Papilionidze were moderately endowed. To proceed further in the analy-
sis 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 Prefecti, a group
of gaily attired butterflies with angulated wings like our common thistle butter-
fly, the cosmopolite. Of the four Papilionidsx, 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 destruc-
tive 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 Urbicolw 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

GENERAL RESUME. 85

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 Polyom-
matus 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 dis-
trict; 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 Crucifer, 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 haye been referred to butterflies,

but whose exact position is still unsettled.
MEMOIRS A. A. A. 8. 13

FOSSIL BUTTERFLIES.

Secon cee, smauyurEty Baste *soTTOsaB IT UvLNS TT “19ppnog “pphog wypqu soutudueg
‘amosqo 419A pus | -( 1peTy “noLouly vaunoatt Mo) a # ;
ar 3 *(ousd0TTy “A\0]) Ppnog BINIeA 99 UL
Moukog ur | see | ae pertir savjuuyuby “wopsoH PpOOg sIm904. 901
*goasted Apiwau *qoUASTp “W091 . ‘
- Young, (auaoog *ddn) *
Hasan ayy tty | PORE | tone | MOH tossayora |” ‘uBNsyT oppuog Joo H CUvTUTUMY sonwU,
uonuinow sq é¢ UVa MIOT, “BOLIOUy “BUUdTA ‘euIqeY | “(euos0rny' prim) ;
__— Ayavou ¢ Blazronig yuiedmey, | -ualyetoutu Jor ‘unjousduyy | | “100 *ppnog Heder BIyWOT
{A\-O403 OUQ '
: eens OMT “xsi ‘SOUL "A | cprodvs yanog Badan Poa “*xoppnog “ppnog vuydresorg se}vI109
‘ Suwon | “(gspuddug) | , oq | ‘BUUATA GouIquy | -(ondd0ry-prm) : “pphog OjnTq sam
sdern-esay toa wsouunsory | “SPOT 'A | -uoyusouju jo | ‘uvjouacesy 79°H oprimdog)
“Buy a-oroy “wrnqag 10 oad, (euasory’prm) | , +101)
auo jo sey toddg | snqndog *xyug be ad é uepUStOne -uedavyg “pphog VALE vluOsN
*pasodurjzadns
ad x “ ‘ . *(euac0gq *ddn) - 2 a
pate Bal eouyUBAr) *sorpuy “ar SOTTOSAVTT “uvOsy] roppuos Ppnog yseudoy sojrq0'T
“apis ouo 7 P : ‘(ana00gq *ddn & “pu: dos (00.
Jo supa 1o9p10q sere sour “a |" seodugyanog | 'umaayy TRANS is eromteyy -
“poasosord ‘sxuqdaaqes jo | . UE pUnoS jo umosnyy *u0zO yy Aq *(sorjuryy pure)
sfajm Jo suva | poor aquqoig | SUM FWA | ay possosora Tw9}H0]00H ge er sojoady Jo sourey

“SGUITAVALLOG TISSOA AO MAIA AVTAAVIL

UNDETERMINED FORMS. 87

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 repre-
sent, 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,'in his enumera-
tion of amber insects, gives under the Lepidoptera forty specimens referable to
Tine and Tortrices, and besides these “mehre Raupen, simmtlich, wie es scheint,
Schildraupen, denen des Papilio W. album ahnlich.” The probable nature of the
ancient forest yielding amber renders it unlikely that any butterflies in their per-
fect state would be found in it. As a rule, butterflies are eminently fond of the
light. This has already been remarked by Menge:*—“ Das fehlen grészerer schmet-
terlinge 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 larvex, 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 enu-
merations 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 rape, etc., having been enclosed between slabs.of amber, which were then
fastened together and the edges roughened, all in so cleyer 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 1886, 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

tArbeit. Schlesisch. Gesellsch. Vaterl. Kultur, 1834, 92-3. 2 Programm Petrischule Danzig, 1855-56, 4°, p. 30,

88 FOSSIL BUTTERFLIES.

der Tertiirgebilde von Ciningen,” says (p. 175): “Karg erwihnt zwar eines sehr
schénen (ninger-Schmetterlings, der nach Zirich gekommen sein soll. Allein —
hier findet sich dieser nicht und die Angabe verliert noch mehr an Werth, wenn
wir bericksichtigen, dass Karg das Thier nicht selbst gesehen hat.” Karg’s
memoir in the “Denkschriften der Schwibischen Gesellschaft der Aertze 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 Wxstw.

These two insects were figured by Westwood in the Quarterly Journal of the
Geological Society of London for Noveniber, 1854, the former (reproduced in our
fig. 2) on Pl. XVII, fig. 17; the latter (reproduced in our fig. 3) on Pl. XVII,
fig. 27. Of the former he makes the following remarks:' “Pl. XVII, fig. 17
represents a number of fragments of delicate tegument, covered with minute punc-

1 Loe. cit., 387.

INSECTS ERRONEOUSLY REFERRED TO BUTTERFLIES. 89

tures 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:?
“Pl. XVII, 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 __ Fig. 2.

for the insertion of the quills of the coloured scales, — Cy!lonium Boisduvalianum Westw.

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 re-
ferred to are both too large and much too irregularly dis-
posed 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

oat. ae anum _aSects belonging to the other suborders. As the figure of
Westw. 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 lepidop-
terous. 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. Paleontind oolitica Butt.

The first notice I find of this remarkable and very interesting fossil is that

published in various literary and scientific London journals reporting remarks

1 Loc. cit. 390:

90 FOSSIL BUTTERFLIES.

given at a meeting of the Entomological Society of London, and which afterward
appeared as follows in their Proceedings:!

“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 “Geologi-
cal Magazine.” In fig. 4 we reproduce fig. 1
of his plates, representing the neuration of
Paleontina; 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:
ieee the ae *[126] Though a British insect, this

The neuration, after Butler’s first sketch. species belongs to a group so completely

Fig. 4.

tropical that I do not hesitate to describe and figure it in the present work;

Fig. 5.
Palwontina oolitica Butl. Facsimile of Butler's first sketch.

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 dis-

11872, xxxi.

INSECTS ERRONEOUSLY REFERRED TO BUTTERFLIES. 91

covered; the most ancient previously known to science having been found in the
Cretaceous series (white sandstone of Aix-la-Chapelle), 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 Satyrine and Nymphaline,
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, draw-
ings which I had made from the originals; my comments at that time were pub-
lished very briefly, as I was reserving the proof of my statements for the present
paper. Mr. Butler, however, was induced by this publication? to examine the re-
verse 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

1 Perhaps Mr. Butler is not altogether to blame in confounding Quarterly Journal of the Geological Society of London, VI, 72;
Aix in Provence with Aix-la-Chapelle; at any rate the mistake the error is corrected by Mr. Butler at the end of his volume.
had been made previously by the translator of Heer’s paper in the 2 He seems not to have seen the earlier publication of Mr. Brodie.

92 FOSSIL BUTTERFLIES.

found it impossible to make a tracing of it, and therefore drew the whole by meas-
urement. This sketch is now produced on Pl. XTX, 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

Fig. 4.
Palwontina oolitien Butl. the wing of Dasyophthalma (fig. 1), and

Facsimile of Butler’s second sketch.

Cicada (fig. 2), one may come to a pretty

accurate conclusion as to the group of insects to which it ought to be referred.”?
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, con-

cealing 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 cer- Fig. 7.
e . i : Palxontina oolitica Butl.
tain extent in pairs; the middle pair usually The neuration, after Butler’s second sketch.

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 back-
ward. 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,

1 Geol. Mag, [2] I, 448.

INSECTS ERRONEOUSLY REFERRED TO BUTTERFLIES. 93

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 sef 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.!. 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 neryures,—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 api-
cal 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 ac-
cordance 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 counter-
part 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.

1These veins have been given a distinct name (discoidal) by belonging to the wings of these insects. I have therefore pre-
the English Entomologists, as if they had an independent origin, ferred to speak of them as the inferior subcostal nervules, in
and had nothing to do with the subcostal nervure; but by the use —contradistinction to the superior branches of the same vein.
of this name, we wholly lose sight of the simple plan of neuration

MEMOIRS A. A. A. 8. 14

94 FOSSIL BUTTERFLIES.

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.

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 es-

sentially in its details from the illustrations

m cama si given by Butler, and looks, as he himself

ee ee ee 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 Lepi-;
doptera. 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 re-
lationship of the veinlets below, which must then be considered as belonging

EE —————————EE—E—

INSECTS ERRONEOUSLY REFERRED TO BUTTERFLIES. 95

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’ arises not from his third branch, but from his upper dis-
coidal 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
Paleontina, 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 posi-
tion 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 sub-
ject 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.

1 In this case he counts from the tip of the wing, in reverse order,

oo Ne AY
} ‘ ‘ t-
c et 1 ie Sette Geena aed Dees
— —— -

: ‘ ‘

d eG See oe) a

S ; . ee a 7 1? - ei’
. . tel be 2 et her 23 i} od

: j i Seog ho

5 se eee Se ee afl) 5

; ; : ee ee a ad | ah ,

4 «
a wa 4 ee i a. Pieris rey we

“

EXPLANATION OF PLATES. 97

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.]

17.

Plate I.

Eugonia atava. Copied from Heer, Insekt. Tert: CEning., ii, pl. xiv, fig. 8 (4):

Lethites Reynesii. Drawn by S. H. Scudder (4).

Eugonia utava. Copied from Charpentier, Nov. Act. Leop.-Carol, xx; pl xxii, fig: 4 (4).

Eugonia j.-album. Neuration of fore wing; drawn by S. H. Scudder (4). The second superior subcostal
nervule is carried too far toward the tip of the wing.

LIethites Reynesii. Fore wing; drawn by S. H. Scudder (}).

Eugonia j.-album. Markings of the upper surface of the fore wing; drawn by S. H. Scudder (4).

Eugonia atava. Neuration of tip of fore wing (}); copied from Heer, Insekt. Tert. Gning., ii, pl. xiv,
fig. 3+.

Neorinopis sepulta. Markings of the upper surface of the two wings, #estored; drawn by S. H. Scudder
(}. The drawing represents the genetal effect of the fore wing as darker than the hind wing, and in
so far is inaccurate.

The same. Neuration of the two wings, sepafated; drawn by S. H. Scudder (}).

The same. Neuration of the two wings, as seen in the fossil; drawn by S. H. Scudder (7). The
engraver has unfortunately made the lines of the hind wing the heavier, as if it lay uppermost; they
should have been the lighter.

The same. Right hind leg; drawn by S. H. Scudder (4).

The same. Left hind leg; drawn by S. H. Scudder ().

The same. Drawn by 8S. H. Scudder (}). The spot of thé medio-submedian interspacé of the fore
wings has not been well rendered by the engraver.

The same. Copied from Lefebvre, Ann. Soc. Ent. France [2], ix, pl. iii, II, fig. A (4).

The same. Copied from the same, fig. C (+).

The same. Copied from the same, fig. B (+).

The same. Copied from Boisduval, Ann. Soc. Ent. Fr., ix, pl. 8 (4).

Plate II.

Zophoessa Sura. Neuration of the wings; drawn by G. Willis (+).

Mylothrites Pluto. Copied from Heer, Insekt. Tert. GEning., ii, pl. xiv, fig. 4 (4).

Zophoessa Sura. Markings of the upper surface of the wings; drawn by G. Willis (+).

Delias Pasithoe. Neuration of the wings; drawn by G. Willis (+).

Coliates Proserpina. Neuration and markings of fore wings; drawn by S. H. Scudder (¢). The tip of
the costal nervure has been extended too far toward the apex of the wing.

Lethe Dyrta. Neuration of the wings; drawn by G. Willis (4).

Mylothrites Pluto. Neuration of the wings; after a drawing obtained through Mr. Brunner de Watten-
wyl (}). The second superior subcostal nervule on the left wing should join the nervuré midway
between the bases of the first and third nervules.

“

16.
17.

15.

17.

19.

FOSSIL BUTTERFLIES.

Neorina Lowi, Neuration of the wings; drawn by G. Willis (+).

Lethe Dyrta. Markings of the lower surface of the fore wing; drawn by G. Willis (+).
Debdis Sinoriz. Neuration of the wings; drawn by G. Willis (4).

Zophoessa Sura. Markings of the lower surface of the fore wing; drawn by G. Willis (4).

Pontia Protodice. Neuration and markings of the upper surface of fore wing; drawn by S. H. Scudder (}.)
Neorina Lowi. Markings of the upper surface of the wings; drawn by G. Willis (}). This was drawn

for the pattern of markings only; the neuration is faulty.
Debis Sinoriz. Markings of the upper surface of the wings; drawn by G. Willis (+).
Mylothrites? sp. Copied from Heer, Insekt. Tert. (Ening., ii, pl. xiv, fig. 5 (4).
Pontia Freyeri. Copied from Heer, Insekt. Tert. Gning., ii, pl. xiv, fig. 6 (4).

Mylothrites Pluto. After a drawing from the original, furnished by Mr. Branner de Wattenwy]l (+).

Pontia Freyeri. Drawn from the original under the direction of Mr. Brunner de Wattenwyl (4).

Plate III.

Thaites Ruminiana. Neuration of the wings, restored; drawn by S. H. Scudder (}).
Thais Rumina. Neuration of the wings ; drawn by S. H. Scudder (}).

Thaites Ruminiana. Markings of the upper surface of the wings, restored; drawn by S. H. Scudder (?).

Thais Rumina. Markings of the upper surface of the wings; drawn by S. H. Scudder (+).

Parnassius Smintheus. Markings of the upper surface and neuration of the wings; drawn by S. H.

Scudder (4).
Thaites Ruminiana. One of the wing-covers (patagia); drawn by S. H. Scudder (4).
The same. Portion of the palpi; from a camera sketch by S. H. Scudder (+).
The same. Antenna; drawn by S. H. Scudder (#).
The same. From a camera sketch by S. H. Scudder (#).
The same. Drawn under the camera from the reverse of fig. 9, by S. H. Scudder (#).

Thanaos Juvenalis. Drawn in the position of fig. 12 by J. H. Emerton (}); fig. lla, the palpus,

denuded (#).

Thanatites vetula. Drawn in outline by an artist in the employ of H. Woodward, Esq., of the British

Museum, and filled in by S. H. Scudder (+2). Incorrectly named Thanatites Juvenalis on the plate.

Carystus Lucasii. Neuration of fore wing. Drawn by G. Willis (}).

Pamphilites abdita. Markings of the upper surface of the fore wing; drawn by S. H. Scudder ().-
Pansydia Mesogramma. Neuration and disposition of spots on the fore wing; copied from Poey, Cent.

p. Cuba, 2° Dec. (}).

Thanatites vetula. Copied from Heyden, Paleontogr., viii, pl. i, fig. 10 (4%). Incorrectly named Thana-

tites Juvenalis on the plate.
Pamphilites abdita. Markings of the upper surface of fore wing; drawn by S. H. Scudder (4).

The same. Neuration and disposition of the spots on the fore wing; drawn by S. H. Scudder @).

Carystus Lucasii. Markings of the upper surface of fore wing; drawn by G. Willis (4).

it

LIST OF WOOD CUTS. , 99

LIST OF WOOD CUTS.

Fig. 1 (p. 50). Mylothrites Pluto. Outlines to show the disparity in size of the two insects referred to this species

by Heer; drawn by S. H. Scudder; engraved by S. S. Kilburn. e

** 2 (p. 89). Cyllonium Boisduvalianum. Copied by photography from Westwood, Quart. Journ. Geol. Soc..
Lond., x, pl. xvii, fig. 17; engraved by H. Marsh.

«© 3 (p. 89). Cyllonium Hewitsonianum. Copied by photography from the same, pl. xviii, p. 27; engraved by
H. Marsh.

“« 4 (p. 90). Palwontina oolitica. Copied by photography from Butler, Lep. Exot., pl. xlviii, fig. 1; engraved
by H. Marsh.

* 5 (p.90). Thesame. Copied by photography from the same, fig. 2; engraved by H. Marsh.

“ 6(p.92). Thesame. Copied by photography from Butler, Geol. Mag., [2] i, pl. xix, fig. 4; engraved by H.
Marsh.

“ 7 (p. 92). The same. Copied by photography from the same, fig. 5; engraved by H. Marsh.

“ 8 (p.94). The same. Drawn by S. H. Scudder; engraved by S. S. Kilburn.

ERRATA.

Page 19, line 4; for voicé, read voici.
* 29. The first three lines form a part of the quotation from Butler, and should haye been but single-leaded.
“ 51, note; for Dareai, read Danai.
“ 58, line 9; for before the cell, read before the tip of the cell.
“ @2, line 13; for fig. 2, read fig. 11.

-

tt ya Sees Vere! ote + BF
Ne Pa F

i sale)

s ' esate
: ¥

~ , a

* Memoirs: Ain So. ig cience Is

Plate T.® |
a

Debray so.
ss . * = = ; > = : : ie
> 10,7 Eugonia atava. ky 8 Eugoma J-album. 2, 5% Lethites ‘Reynesii. 8-17 Neopinopis sepultaas
. Y, :
’ _" + , *.
a . 7 » . ”
in ~ .
~~. i * . a) ~ »*% ‘
». ~ - .
7 Ad <
an : c i »
a
> ° Imp. Oh. Ghar Pa, . _<
. yp. Ch, Chardon aine, Paris. a

:

i . : > e

Memoirs Amer. Assoc: Adv. Science. I.

1 ee ee e = cae

a? Plate Il.

~~ °
B30 Zophoessa Sura. 2,717 Mylothrites Pluto. .,Delias Pasithoe.

a *.
; 8 B\Neorina Lowi, 10,1 Debis Simorix. 12 Pontia Protcdice.

eT” y
: 7g! -

Top. th. Chardon winds, Pariv

. Debray sc. |)

.

5. Coliates Prosenpina. 6,9 Lethe Dyrta. id

15 Mylothrites ? 16,18 Pontia Freyeri. ae

Debray sc.

od

-

ll

5 Parnassius Sminth eus.

Rumina.

Thais

2, k

-

f

10 Thaites Ruminiana.

1,5,6

Thanaos Juvenalis.

=

Meso §ramma. f

casii.

GwThanatites Juvenalis. 13.19 Car

-

.

Paris,

uh, 17,18 Pamphilites abdita. 15 Pansydia

.
<_-

.

Seudder, Samuel Hubbard
gee > oa Fossil butterflies
L583 on .

i

i ‘3
nie
tected

bgt at

ea

of,

PEP Mee