MEMOIRS
OF THE
AMERICAN ASSOCIATION
FOR THE
ADVANCEMENT OF SCIENCE.
I.
SALEM, MASS.
F. W. PUTNAM, PERMANENT SECRETARY, A.A.A.S.
1875.
PRINTED AT THE SALEM PRESS, SALEM, MASS.
LETTER OF GIFT.
PORTLAND, Aug. 22, 1873.
Mrs. Elizabeth Thompson 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 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 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.
[Signed]
P. H. VAN DER WEYDE.
To the Standing Committee of the American Association for the
Advancement of Science.
REPORT OF THE SPECIAL COMMITTEE ON THE THOMPSON FUND, HARTFORD MEETING,
AUGUST, 1874.
The Standing Committee of the Association at the Portland Meeting
appointed 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 donation of Mrs. Thompson.
In accordance with the duties assigned to them, the Committee have
accepted the Memoir by Mr. Scudder on Fossil Butterflies as the first
paper to be published by the THOMPSON FUND, and while regretting that
the unavoidable delay in engraving the plates prevents their having the
gratification of presenting the work at the present Meeting, they believe
that the Association and its liberal patron will accept the Memoir as one
in every way worthy of the honor thus bestowed.
ASA GRAY, }
JAMES HALL, }
THOMAS HILL, }
P. H. VAN DER WEYDE, } _Committee_.
J. L. LECONTE, }
T. STERRY HUNT, }
F. W. PUTNAM, }
FOSSIL BUTTERFLIES.
BY SAMUEL H. SCUDDER.
TO COUNT GASTON DE SAPORTA, OF AIX IN PROVENCE,
WHOSE EXTENDED MEMOIRS ON THE FLORA OF THE TERTIARIES OF SOUTHERN FRANCE
FORM THE BASIS OF THE BIOLOGICAL CONCLUSIONS OF THIS ESSAY; AND WHOSE
UNWONTED COURTESIES HAVE PERMITTED A CAREFUL EXAMINATION OF THE MOST
IMPORTANT FOSSIL BUTTERFLIES,
THIS MEMOIR IS RESPECTFULLY INSCRIBED BY THE AUTHOR.
CONTENTS.
INTRODUCTION xi
BIBLIOGRAPHY 1
DESCRIPTIONS OF GENERA AND SPECIES OF FOSSIL BUTTERFLIES
NEORINOPIS 9
1. Neorinopis sepulta 14
LETHITES 34
2. Lethites Reynesii 37
EUGONIA 40
3. Eugonia atava 41
MYLOTHRITES 44
4. Mylothrites Pluto 45
COLIATES 51
5. Coliates Proserpina 52
PONTIA 53
6. Pontia Freyeri 54
THAITES 57
7. Thaites Ruminiana 60
THANATITES 62
8. Thanatites vetula 63
PAMPHILITES 66
9. Pamphilites abdita 68
COMPARATIVE AGE OF FOSSIL BUTTERFLIES 70
PROBABLE FOOD PLANTS OF TERTIARY CATERPILLARS 71
PRESENT DISTRIBUTION OF BUTTERFLIES MOST NEARLY ALLIED TO
FOSSIL SPECIES 76
GENERAL RÉSUMÉ, WITH NOTICE OF UNDETERMINED FORMS 83
FOSSIL INSECTS ERRONEOUSLY REFERRED TO BUTTERFLIES 88
EXPLANATION OF THE PLATES AND WOOD CUTS 97
INTRODUCTORY.
The 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. Reynès, 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.
BIBLIOGRAPHY.
1726. HUEBER. Lithographiæ Wirceburgensis specimen primum. Fol.
Wirceburg. This work contains the first reference to fossil Lepidoptera
which I have found. In his Synopsis Tabellarum, he gives on page 94:
“Tabulâ 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. BROMELL. Lithographia Suecana. Acta Litteraria Sueciæ, II. In a
section _de lapidibus insectiferis Seanicis et Gothicis_ (p. 525) he says:
“Praeter umbratiles etenim papilionum vel muscarum quasdam
imagines, lapidi huic leviter sed distincte impressas, multa
scarabæorum figuras, mole totaque facie imitantur;” these were
found in “saxo fœtido” in “Westrogothia.”
In his enumeration of fossils he specifies further:
[528]. “9. Papilionum majorum ac minorum imagines et
impressiones nitidæ, in lapide calcario communi inodoro, ubi
etiam in alio fœtido conspicuæ, ex eisdem Westrogothiæ locis.”
[529]. “10. Insectorum ovula, an nymphæ seu aureliæ lapideæ?
saxo fœtido nigricanti immersæ. Ex eadem paræcia karabylonga.”
[531]. “14. Papilionum minorum imagines et impressiones, in
ejusdem generis saxo suillo fœtido. Ex eodem loco. Hæ itidem
figura sua a papilionibus illis differre haud videntur, quarum
superius Num. 9. meminimus.”
I find no later reference to these supposed Lepidoptera.
1742. SENDELIUS. Historia succinorum. Fol. Lipsiæ.
Devotes a chapter (_De Erucis_, 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 illustrations are wholly insufficient to
assert anything of them with confidence.
1828. MARCEL DE 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.
1829. MARCEL DE SERRES. Géognosie des terrains tertiaires ou Tableau des
principaux animaux invertébrês 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-Rhône),” 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 Lépidoptère diurne de la division des
_Satyrus_,” doubtless referring to _Neorinopis sepulta_.
Speaking of the authors who have treated of the fossils of
Œningen, he says: (p. 235) “Ces divers naturalistes 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.
/-----------------------/\---------------------\
Tertiaires. Secondaires.
/--------------/\-----------------\ /--/\---\
| | antérieurs à la |
| | retraite des mers |supérieurs
| | dans les couches de |jurassiques.
|postérieurs|/---------/\-------\ | |inférieurs.
| à la | | | |
Noms des | retraite | marnes | lignite | | | Nombre
genres. | des mers. |calcaires.|et succin.| | |d’espèces.
---------+-----------+----------+----------+----+----+----------
| | | | | |
Papilio. | | * | * | | * | 4
=========+===========+==========+==========+====+====+==========
In the “marnes calcaires” of Aix he has referred already, as
we have seen, to one; he previously speaks of _Papillons_
at Œningen (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 Lépidoptères
(M. Brongniart). On a cru reconnaître des chenilles parmi
les insectes du Succin figurés par Sendelius Tab. 3, fig.
28-82;”[A] 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 même époque
qu’existent les vestiges d’insectes, d’ailes de Papillons et de
Scarabées signalés par Bromel.”
1835. GRAVENHORST. Bericht der entomologischen Section. Uebers. d. Arbeit
u. Veränd. Schlesisch. Gesellsch. Vaterl. Caltur, 1854, 92-93.
Gives a general enumeration of the collection of fossils from
amber in the museum of the Königsberg Society, specifying a few
Lepidoptera.
1836. HOPE. 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. Lethæa Geognostica, 2d ed., II. 8vo.
In a tabular list of fossil insects, with localities, he gives
(p. 814):
Papilis [Papilio] (Bernstein), Satyrus (Gyps formation von Aix).
1838. DUPONCHEL. Ann. Soc. Ent. France, VII, Bull. 51-52.
Re-announces the discovery of _Neorinopis sepulta_, referring
it to Nymphalis.
1839. BOISDUVAL. 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. BOISDUVAL. Rapport sur une empreinte de Lépidoptère 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.
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. MARCEL DE SERRES. Notes géologiques sur la Provence. Actes
Linn. Soc. Bord., XIII, 1-82; Note additionelle, 83-90; Deuxième note
additionelle, 170-2. 2 planches.
In a list of the plants and animals found at Aix, the author
gives on p. 41: “Lépidoptères Diurnes. Papilio de la division
des Satyrus. Cette espèce conserve encore en partie ses
couleurs.” On p. 172 is a _Note relative au Lépidoptère
figuré_ (_Cyllo sepulta_), in which Boisduval’s opinion of its
relationship is given.[B] 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. COQUAND. 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.
1845. MARCEL DE SERRES. Sur les fossiles du bassin d’Aix
(Bouches-du-Rhône). Ann. Sc. 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.
1847. HOPE. 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].”
1849. HEER. Die Insektenfauna der Tertiärgebilde von Œningen und von
Radoboj in Croatien. 2er Theil. 4to. Leipzig. Extracted from the Neue
Denkschr. allg. Schweiz. Gesellschaft für Naturw., XI (1850).
Contains (pp. 177-183, Taf. xiv, figs. 3-6) descriptions
and illustrations of _Eugonia atava_ (_Vanessa attavina_),
_Mylothrites Pluto_ (_Vanessa Pluto_) and _Pontia Freyeri_
(_Pierites Freyeri_).
1849. 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: “Merkwürdig ist, dass von diesen
Schmetterlingen 2 Arten grosse Aehnlichkeit [88] mit
ostindischen Arten haben, während eine mit unserm Distelfalter,
eine andere mit unserem Grassackträger zu vergleichen ist.”
1850. HEER. Zur Geschichte der Insekten. Neues Jahrb. für 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 same 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.
1851. LEFEBVRE. Observations relatives à l’empreinte d’un Lépidoptère
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 Boisduval.
1851. BOISDUVAL. Quelques mots de réponse à M. Alex. Lefebvre sur ses
observations relatives à la _Cyllo sepulta_. Ann. Soc. Ent. France [2],
IX, Bull. 96-98.
Defends his views against the criticisms of Lefebvre.
1852. GIEBEL. Deutschland’s Petrefacten. p. 644. 8vo. Leipzig.
Catalogues the three butterflies described by Heer from Radoboj.
1854. WESTWOOD. 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 _Cyllonium Boisduvalianum_
and _C. Hewitsonianum_.
1854. PICTET. Traite de Palæontologie, 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.
1856. GIEBEL. 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 others.
1856. GIEBEL. Geologische Uebersicht der vorweltlichen Insekten.
Zeitschr. gesammt. Naturw., VIII, pp. 174-188.
Gives lists of Lepidoptera summarized from his previous work.
1856. 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.
1858. HEER. Ueber die Insectfauna von Radoboj. Bericht 32e Versamml.
Deutsch. Naturf., 118-121.
A cursory review of Radoboj insects, mentioning the rarity
of Lepidoptera, and specifying _Eugonia atava_ (_Vanessa
attarina_) 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_.
1859. HEYDEN. Fossile Insecten aus der Rheinischen Braunkohle. Dunk. u.
Mey. Palæontogr., VIII, 1-15, Taf. 1-2.
Contains pp. 12-13, Taf. I, fig. 10, description and figure of
_Thanatites vetula_ (_Vanessa vetula_).
1860. HEER. Untersuchungen über das Klima und die Vegetations
Verhältnisse des Tertiärlandes. 4to. Winterthur.
Refers to some of the fossil butterflies described from Radoboj
and Aix.
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 reference: “un cinquième
(_Thaites Ruminiana_) est très voisin du genre Thais qui
appartient à la faune méditerranéene.”
1868. BUTLER. Catalogue of Diurnal Lepidoptera of the family Satyridæ 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. BUTLER. 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. SCUDDER. Description d’un nouveau papillon fossile (_Satyrites
Reynesii_) trouvé à 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. SAPORTA. Études sur la végétation du Sud Est de la France à
l’époque tertiaire. Suppl. I. Révision de la flore des gypses d’Aix. 1er
fascicule, Généralités. Ann. Sc. Nat. [5], Bot. XV, 277-351.
Discusses (p. 342) the probable food of the caterpillars of
_Neorinopis sepulta_ and _Thaites Ruminiana_.
1873. BUTLER. On Fossil Butterflies. Lepidoptera Exotica, part xv, pp.
126-8, pl. 48.
On a Fossil Butterfly belonging to the family Nymphalidæ 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 Palæontina 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. ANON. The oldest Fossil Butterfly in the World. The [London]
Graphic. Feb. 22.
A popular account of the preceding paper, accompanied by a
woodcut of _Palæontina oolitica_.
1873. BRODIE. The Distribution and Correlation of Fossil Insects, etc.
8vo. pamph. Warwick.
Gives a brief notice (pp. 8-9) of the various fossils referred
to butterflies, especially of _Palæontina oolitica_ and
_Lethites Reynesii_, and publishes an opinion expressed to him
by me that the former was Homopterous.
1874. SCUDDER. Proc. Bost. Soc. Nat. Hist., XVI, 112.
Doubts the lepidopterous character of Butler’s Palæontina, and
refers it, probably, to the Cicadinæ.
1874. BUTLER. Notes on the impression of _Palæontina oolitica_ in the
Jermyn Street Museum. Geol. Mag. [2], I, 446-449, pl. 19.
Defends the lepidopterous character of Palæontina and gives new
illustrations of the same.
1874. SMITH. Discovery of Remains of Plants and Insects. Nature, XI, 88.
Enumerates fossils found at Gurnet Bay, and specifies among
them “butterflies.”
Sehn wir daher durch das Fenster,
In das alte Schattenreich,
Sehen wir da statt Gespenster,
Wesen, die den jetz’gen gleich;
Sehen nicht des Pluto Schrecken,
Sphinxe und Harpyen Brut,
Nicht Chimären Flammen lecken,
In der Hölle Feuer Glut,
Nein! in diesen stillen Räumen
Wo man sich den Orcus denkt,
Sehn wir tausend Wesen träumen,
Tief in ew’gen Schlaf versenkt.
Haben einst die Welt genossen,
Unterm blauen Himmelszelt,
Jetzt sind sie in Fels verschlossen,
In der schwarzen Unterwelt.
OSWALD HEER.
DESCRIPTIVE.
NYMPHALES—PRÆTORES—OREADES.
Genus NEORINOPIS BUTLER.
_Neorinopis_ Butler, Lepid. Exot., i, 127 (1873);—Ib., Geol.
Mag. x, 3.
In the shape of the wings (Pl. 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 Antirrhæa
or Cœlites, 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 portion, 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 general
course of the costal margin and is gently crenate; the tail, which
lengthens the upper median nervule 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 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 projection,
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; beyond it the inner margin has a nearly straight course, parallel
and adjacent to the submedian nervure.
As to the neuration (Pl. I, fig. 9) this genus approaches more closely
the genera Zophoessa (Pl. II, fig. 1), Neorina (Pl. II, fig. 8), Debis
(Pl. II, fig. 10), and Lethe (Pl. II, fig. 6), than any others, although
it differs from any of them more than they do among themselves. The
most noticeable marks of distinction 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 nervule 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 Neorinopis
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 divarication, 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 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 nervule; 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 gradually as
far as the base of the outer superior nervule, where the main stem
approaches it again; the lowermost inferior subcostal nervule arises from
the first inferior scarcely beyond its base, curves inward, downward and
then outward before 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 subcostal nervure; 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
nervure 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 nervure 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, etc. 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. II, 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 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 (Pl. 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 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
(Pl. II, figs. 3, 11) and Lethe (Pl. II, 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. II, 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 _Neorina
Lowii_, 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 Lowii_. In general, the design of the fore wings approaches that
of _Neorina Lowii_ 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 Antirrhæa and Anchiphlebia, as Butler has remarked, as
well as toward certain species of Zophoessa. The markings 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_ (Pl. II, fig.
9) may well be studied, where in a lighter submarginal band we find a
series of spots, in the principal interspaces, far from the border; these
are ocellated instead of double as in Neorinopis; there 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 instead 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. II, fig. 3), and resembles less conspicuously _Debis Sinorix_
(Pl. II, 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 butterfly 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 tibiæ and a very little longer than the femora.
NEORINOPIS SEPULTA (BOISDUVAL) BUTLER.
Plate I, figs. 8-17.
_Nymphalis_ sp. DUP., Bull. Soc. Ent. France, 1838, 51-52.
_Cyllo_ sp. BOISD., Bull. Soc. Ent. France, 1839, 11-12.
_Cyllo sepulta_ BOISD., Ann. Soc. Ent. France, ix, 371-374, pl.
viii (1840); Ib., Bull. Soc. Ent. France, 1851, 96-98; SERRES,
Act. Linn. Soc. Bord., xiii, 172, pl. ii (1843); WESTW., Gen.
Diurn. Lep., 361 (1851); LEF., Ann. Soc. Ent. France [2], ix,
71-88, pl. iii, II (1851); PICT., Traite Pal., ii, 393, pl. xl,
fig. 11, 1854; BUTL., Cat. Satyr. Brit. Mus., 189-190 (1868).
_Antirrhæa? sepulta_ KIRB., Syn. Cat. Diurn. Lep., 39 (1871).
_Neorinopis sepulta_ BUTL., Lep. Exot., 127, pl. xlviii, fig. 3
(1873); IB., Geol. Mag., x, 3, pl. l, fig. 3 (1873).
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.”
The earliest definite mention of the insect is given by Duponchel in the
Bulletin of the Entomological Society of France, as follows:[C]
“M. Duponchel entretient ensuite la Société d’un fait
extraordinaire, et peut-être entièrement nouveau dans les
annales de [52] la science: c’est l’existence d’une impression
très remarquable de Lépidoptère fossile, qui a été trouvée dans
une plâtrière des environs d’Aix (en Provence), et acquise par
M. de Saporta. Ce Lépidoptère, suivant M. de Saporta, paraît
appartenir au genre _Nymphale_, et à une espèce étrangère
à celles qui vivent aujourd’hui en Europe. Le corselet en
est parfaitement conservé; les couleurs des ailes sont
très-bien indiquées; le dessin de ces ailes est entièrement
reconnaissable. Les deux ailes d’un des côtés du corps sont
repliées en grande partie l’une sur l’autre; la place du ventre
est très distincte; l’autre côté manque tout-à-fait.”
The subject seems to have been referred to Dr. Boisduval, for we find in
the following year[D] that
“M. Boisduval rend un compte verbal du rapport que la Société
l’avait chargé de faire, sur un dessin envoyé à M. Audouin,
par M. de Fonscolombe, et qui représente une empreinte de
lépidoptère fossile trouvée dans les environs d’Aix. M.
Boisduval déclare, qu’après un examen attentif, il a reconnu
que ce lépidoptère devait appartenir à son genre _Cyllo_, et
qu’il se rapprochait beaucoup des espèces décrites par les
auteurs sous les noms de Satyrus Europa, Caumax, Rhosia et
plusieurs autres lépidoptères indiens. Le même membre ajoute
que ce _Rhopalocère_ ne peut se rapporter exactement à aucune
des espèces vivantes déjà connues. Toutefois, avant de décider
si cet insecte doit être regardé comme un véritable fossile,
M. Boisduval pense qu’il serait indispensable, que la Société
pût avoir sous les yeux [12] la pierre qui a servi de modèle 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 given below:[E]
“Il y a bientôt un an que je fus chargé par la Société
d’examiner le dessin d’un Lépidoptère fossile trouvé dans les
plâtrières des environs d’Aix en Provence, et appartenant à
M. le Comte de Saporta. Au premier coup d’œil, ce dessin me
parut devoir être rapporté à une espèce de Satyrides du genre
_Cyllo_, à côté des _Satyrus Rohria_, _Caumas_ et _Europa_, de
l’Encyclopédie; mais la découverte d’un Lépidoptère fossile
me sembla un fait tellement neuf, et l’espèce si rapprochée
de celles connues, que je n’osai pas faire de rapport avant
d’avoir vu la pierre en nature. La Société partagea cet avis,
et engagea M. Duponchel à écrire à 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 distingué et propriétaire du fossile en
question, m’écrivit la lettre que j’ai communiquée à la
Société; lettre dans laquelle il prétendait qu’il n’avait pu
être victime de la supercherie de qui que ce soit, et [372] que
par conséquent il n’y avait pas lieu à conserver le moindre
doute sur l’exactitude scrupuleuse du dessin communiqué par
son beau-père, M. le Baron de Fonscolombe; qu’on pouvait voir
d’ailleurs 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 connaître
en nature ce lépidoptère fossile. M. Duponchel écrivit une
seconde fois à M. de Fonscolombe: ce fut alors que M. le comte
de Saporta consentit à se dessaisir pour quelques jours de ce
précieux échantillon en nous l’envoyant en communication.
Le morceau de calcaire qui porte réellement _l’empreinte
parfaite_ d’un lépidoptère 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.…
Le Lépidoptère qui fait le sujet de ce rapport fait partie
d’un [373] de ces genres dont les espèces assez peu nombreuses
sont confinées aujourd’hui dans les îles de l’archipel indien
ou dans les contrées les plus chaudes du continent asiatique.
D’après ce que j’ai pu apprendre de M. Blum de Leyde, ils
voltigent çà et là à l’entour des palmiers, dont peut-être ils
se nourrissent à l’état 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 espèces voisines; mais il ne peut être rapporté à
aucune de celles connues de nos jours, ce qui est d’autant plus
vraisemblable, que les marnes schisteuses sont de beaucoup plus
anciennes que la dernière catastrophe diluvienne admise par
tous les géologues.
Le dessin et la forme de cet insecte sont si bien conservés,
que l’on croirait qu’il a été lithographie sur un schiste;
seulement il n’existe que le côté droit, lequel est
parfaitement intact, une portion du corselet et une légère
empreinte de l’abdomen. L’aile supérieure est en grande partie
cachée par l’inférieure, et il est impossible de dire si elle
offre d’autre dessin qu’un œil apical surmonté d’un point
blanc; l’autre, dont on voit toute la surface, est d’une
couleur gris brunâtre, comme dans les espèces voisines, avec
une tache costale blanche, une bande transverse, médiane,
sinuée, de la même couleur, suivie de deux yeux noirs encadrés
de blanc, s’alignant extérieurement avec deux points blancs.
L’extrémité de cette même aile est un peu plus pâle, presque
blanchâtre, et divisée, comme chez la plupart des espèces
vivantes, par deux lignes marginales brunes, parallèles.
L’appendice caudal est un peu plus long que dans _Rohria_, mais
situé de la même manière.
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 plutôt feuillets par feuillets, par des dépôts
fluviatiles.… Selon certaines circonstances, les différentes
couches ont varié de couleur, comme on peut s’en convaincre
par l’échantillon que la société a eu sous les yeux. Les
plus inférieures sont colorées par du bitume et des oxydes
métalliques; celle où se trouve le Lépidoptère 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,[F] and his statements reproduced in part or referred to, as will
be seen by the Bibliography at the commencement of this essay.[G] But the
most extraordinary of all is an acute criticism by Lefebvre, eleven years
subsequently, of which I give the following extracts,[H] from a copy of
the paper in my possession slightly corrected by the author.
[72] Si de l’œil on suit les bords de la seconde aile [Pl. I,
fig. 17] qu’avec le docteur je reconnais couvrir en grande
partie la première, 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 d’une de nos _Cyllo_; comment
il lui attribue un appendice caudal, propre volontiers aux
[73] espèces de ce groupe, et qui, selon lui, la termine à la
manière de ceux de la _Cyllo Rohria_ de Fab. (Voy. f. A [Pl. I,
fig. 14]).
Pour parler ainsi que le fait M. Boisduval de cet appendice,
il faut nécessairement qu’il ait confondu avec cette seconde
aile le dernier contour de la première, qui l’excède à partir
de leur point de jonction sur le bord externe, lui attribuant
comme appendice caudal cette forte dent de la première aile,
qui succède à une forte échancrure, ainsi qu’il en existe dans
tant de Vanessides, et qui le plus souvent y est soutenue par
la troisième inférieure, ainsi que tout à l’heure elle l’y sera
pour nous dans la _Sepulta_.
J’avoue donc que je ne puis, avec le meilleur vouloir,
envisager cette empreinte autrement que je ne le fais, et que
pour la considérer sous le même point de vue que notre docte
confrère, il me faut faire trop violence à 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
première 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 première, qui est un peu falqué
et qui, dans l’empreinte, passe précisément sous l’articulation
fémoro-tibiale de l’unique 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
première aile sur laquelle elle est appliquée? A partir du
point le plus proche de ce même bord [74] avec celui de la
première aile, et presque au centre de sa forte échancrure,
ne s’en détache-t-elle pas, comme au-dessus, par une marge
obscure et très nettement tracée? Enfin, cette même 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 déjà 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:
1º Une aile de dessus fortement dentée et échancrée en dehors,
à son bord extérieur. (Voy. fig. B [Pl. I, fig. 16]).
2º 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 conformation, nous sommes actuellement en droit
de décliner déjà toute espèce d’analogie entre la _Sepulta_
et le genre _Cyllo_, proprement dit, et de l’éloigner des
_Caumus_, _Beroe_, _Rohria_, et autres; et cela, d’abord, par
la rondeur inerme de la seconde aile, et ensuite à la première,
par cette forte échancrure, suivie d’une dent non moins énorme
que soutient la troisième inférieure, caractères que n’offrent
guères les Satyrides de cette section, et où la dent la plus
proéminente du bord extérieur, comme à _Banksia_ God., se
prononce à l’extrémité de la première supérieure, quand il en
existe une.
Je ne connais que des Vanessides qui puissent présenter en
même temps des premières ailes déchirées de cette manière
à leur bord extérieur, et des secondes ailes arrondies et
sans dentelures. La _Van. Archesia_, Cr. pourrait, entre
autres, nous en offrir un example. Et cependant chez les
Vanessides, lorsque les premières ailes y sont ainsi dentelées
et découpées, les secondes le sont égale-[75] ment, plus ou
moins, par la règle assez générale qui veut que chez les
Lépidoptères les secondes ailes y soient toujours plutôt munies
de dentelures que les premières.
Voici donc, pour la forme des ailes, un argument en faveur de
mon opinion. Passons aux dessins.
Avant que d’assayer de les réhabiliter dans cette espèce, il
me faut décider une autre question, à savoir si ces dessins
appartiennent à la première aile ou à la seconde. Notre
confrère les tient pour être propres à cette dernière. Je ne
suis pas de son avis, et voici pourquoi:
Je pense que la seconde aile est en grande partie dénudée de
ces écailles à sa face inférieure, celle que nous voyons.
Ce qui me le fait croire, c’est que déjà dans sa marge
abdominale, ainsi que je l’ai déjà dit, on suit à travers la
membrane le contour intérieur de la première aile, et d’une
manière trop distincte pour admettre que l’adhésion des deux
ailes le pût permettre, si les deux faces de la seconde étaient
revêtues de leurs écailles.
Ce qui me le fait croire encore, c’est qu’à cette seconde aile,
la petite lunule blanche de l’angle externe (fig. B [Pl. I,
fig. 16]), qui est située sur le bord lui-même, et qui y est
extérieurement coupée par lui, ne saurait devoir y exister à
cette place, si on en juge par la loi suivie dans leur position
normale parmi la majeure partie des Diurnes. En effet, à aucun,
ou à bien peu du moins, je ne connais pas à cet angle de lunule
extrême, ainsi placée sur _le bord lui-même_ des secondes
ailes, et dans cette position, rejetée en arrière de celle qui
la précède.
Règle assez générale, la série marginale de taches lunulaires
ou autres, pupillées ou non, qui affectent ces ailes, est
d’habitude concentrique à leur base, et la lunule en [76]
question serait sur cette seconde aile placée contre cette
règle.
A examiner cette aile dans la fig. B [Pl. I, fig. 16], on
comprend de suite que cette lunule n’y est pas à sa place
normale; elle choque même là où elle est située, tandis que si
je la reporte (sans la bouger, bien entendu) sous la première
aile (ainsi que je le fais à la fig. C [Pl. I, fig. 15]), elle
s’y adapte tout naturellement dans l’ordre que lui est le plus
rationnel avec les autres.