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Title: Fossil Butterflies - Memoirs of the American Association for the Advancement of Science, I.
Author: Scudder, Samuel H.
Language: English
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                                 OF THE
                          AMERICAN ASSOCIATION
                                 FOR THE
                         ADVANCEMENT OF SCIENCE.


                              SALEM, MASS.



                                                 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


                                                     P. H. VAN DER WEYDE.

    To the Standing Committee of the American Association for the
    Advancement of Science.

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,        }







  INTRODUCTION                                                xi

  BIBLIOGRAPHY                                                 1


    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



     FOSSIL SPECIES                                           76





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.


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,

    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

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

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,

    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

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.

    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.




    _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.


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

    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.

    Par ce fait, à la place qu’occupe cette lunule, la seconde
    aile serait donc encore transparente? Observons en passant que
    dans les espèces où une semblable lunule ou tache oculaire, se
    remarque en dessous, vers l’angle externe des deux ailes (comme
    à _Melanitis Undularis_, Dr.; _Protogenia_, Cr., _par ex._),
    cette tache qui est toujours placée _un peu avant_ la marge,
    qui ne _l’interrompt jamais_ comme ici, est toujours (comme
    ici, du reste) entre les deux dernières supérieures, et non
    entre la dernière supérieure et la costale.

    S’il est des exceptions à cette règle, elles ne sauraient être
    qu’en bien petit nombre, et lorsque les lunules marginales y
    sont présentes en nombre considérable; mais s’il n’y en a plus
    qu’une ou deux, celle de l’angle externe sera placée ainsi que
    je viens de la dire, et non ailleurs.

    Toujours à l’appui de cette transparence, que j’attribue à
    la seconde aile de la _Sepulta_, si j’interroge le peu de la
    charpente alaire qu’on y distingue, et qui est suffisant pour
    la restituer telle qu’elle devait être, ou à bien peu de chose
    près (comme à la fig. C [Pl. I, fig. 15]), on voit que la
    tache semi-orbiculaire et obscure de l’angle interne y semble
    partagée par une nervule de la seconde aile très bien écrite,
    par la deuxième inférieure. Or, l’étude de cette partie de
    la [77] ptérologie, qui a pour objet les lois relatives à la
    position, à la forme, comme à la présence et à l’absence des
    dessins et des taches, nous apprend que jamais une lunule ou
    une tache orbiculaire marginale n’est divisée par une nervule,
    mais plutôt par le pli internervulaire, les nervules séparant
    d’habitude ces sortes de taches, et ne les scindant pas.

    Par induction, je dirai donc que cette tache orbiculaire n’est
    pas encore ici à sa vraie place sous la seconde aile. Mais
    si je l’attribue à la première, ainsi que tout à l’heure je
    l’ai fait pour la petite lunule, à son tour elle s’y adapte
    merveilleusement bien (fig. C [Pl. I, fig. 15]), entre la
    troisième 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 à 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 à
    la première aile (fig. C [Pl. I, fig. 15]), elles viennent y
    compléter cet ensemble, qui paraît alors fort rationnel, des
    plus habituels, et dont au besoin nous trouverions un exemple
    dans la _Van. Alcithoe_, Cr., etc.

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

    [78] Or, si la nervule dont s’agit (la deuxième inférieure) est
    à sa place normale, la tache orbiculaire qu’elle divise n’y est
    pas. Donc, elle doit appartenir forcément à 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 eût précisément
    conservé ses écailles à cette place. C’est chose possible, mais
    chose peu probable.

    D’après ce qui précède, je suis donc porté à croire, comme je
    l’ai déjà avancé, à la dénudation presque complète 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
    à l’autre éclat de cette marne qui nous est inconnue, n’a pu
    atteindre les portions de la première aile qu’elle abrite.

    En plus, par l’analogie et le faciès de la _Sepulta_, ayant
    tout lieu de penser que le dessus de toutes ses ailes devait
    être d’un brun sombre, uni et privé de tout dessin tranché,
    ou varié de vives couleurs, par cela même, j’en induis [79]
    que la surface supérieure de la seconde aile n’a pu empêcher
    les dessins qu’elle recouvrait de paraître, sans confusion
    aucune, à travers la couche unicolore des écailles du dessus,
    généralement très fines dans les Satyrides. Leur adhérence
    intime à la surface inférieure de la première aile aura même dû
    augmenter la transparence de la seconde.

    Mais avec assez de raison, on pourrait me demander à mon
    tour, par quel privilége, ce qui reste de non recouvert de la
    première aile n’a pas été altéré par ce même frottement, ou
    plutôt par son impression sur l’éclat qui a mis à jour cette
    empreinte? De cet argument _ad hominem_, je ne pourrais me
    tirer je l’avoue, qu’en arguant que nous ne voyons 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 _l’intactum_
    des écailles du dessous de cette deuxième aile, je les admets
    ou les repousse pour mieux soutenir l’opinion que j’avance.…

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

    De toute manière, il est impossible de l’admettre comme dessin
    appartenant au dessous de la première aile, ainsi qu’a dû le
    comprendre M. Boisduval, par une erreur d’optique, que déjà
    sans doute il a reconnu lui-même.

    L’absence bien regrettable de l’éclat qui recouvrait cette
    _Sepulta_ est cause de tant d’incertitude, car je ne mets pas
    en doute qu’il devait conserver, à son tour, la majeure partie
    des écailles de toutes ces ailes, avec lesquelles il était en

    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 le dessin original.

    Après cette série de taches marginales, il existe sur la côte
    elle-même, avant l’angle externe de la seconde aile, une large
    éclaircie blanche, quelque peu ovalaire, nettement dessinée
    en dedans, et posée sur la place qu’à la première aile doit
    occuper la disco-cellulaire et le commencement des deux
    premières supérieures.

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

    Ces dessins maculaires peuvent, à la rigueur, y exister, comme
    n’y pas être, de même qu’ils ne sont guère acceptables à leur
    autre surface; car ce que nous voyons est bien un dessous
    d’aile et non un dessus.

    Les dentelures externes de la tache basale, sont en dessous
    des plus naturelles, 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 l’original: qu’y voyons-nous?

    Déjà, sur le bord costal de la première aile, nous apercevons
    en effet, sur notre gauche, un commencement de cette blanche
    éclaircie qui succède aux lunules, et à notre droite, le bord
    intérieur de cette éclaircie y est des plus évident! (Voy. fig.
    B [Pl. I, fig. 16]).

    Comment donc se fait-il que ces ailes ainsi ployées, ces
    vestiges de la première aile viennent s’adapter d’une manière
    si complète avec toute la portion blanche qui se continue sous
    la seconde aile? cela se peut rencontrer, je l’avoue, mais
    c’est peu fréquent.

    Bien mieux, le large sommet de la tache blanche anguleuse
    et obscure qui lui succède, se voit aussi sur le bord un
    peu diffus de la côte, 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 basale, hachée à son dehors, ne se continue-t-il
    pas sur la côte de la première aile?

    Enfin, si cette dernière tache appartenait à la seconde
    viendrait-elle, ainsi qu’elle le fait, s’arrêter précisément
    sur le bord intérieur de la première, que par transparence
    nous pouvons suivre parfaitement à partir du moment où il
    est recouvert par la seconde aile? En l’attribuant à cette
    dernière, ce serait agir contre toute apparence plausible,
    contre toute disposition naturelle de ces sortes de taches, et
    venir l’interrompre bénévolement et sans motifs spécieux, bien
    avant l’angle anal de la seconde aile, sur lequel elle devrait
    venir s’appuyer pour demeurer dans la forme la plus normale!

    Cette interruption nous fixe donc aussi bien que le
    commencement de toutes les taches du haut, sur l’attribution
    que nous devons en faire à la première aile, et non à la
    seconde, et le peu qui reste de ces divers dessins sous cette
    dernière, si toutefois il en reste, doit se confondre avec
    elles, sans contribuer beaucoup à nous égarer.

    D’ailleurs, nombre de Lépidoptères diurnes des groupes, près
    desquels doit venir se ranger la _Sepulta_, présentent sous
    leurs premières ailes de semblables taches costales et basales,
    ainsi placées, ainsi dentelées, ainsi conformées; d’habitude
    même, elles y sont les vestiges plus ou moins complets de
    ces larges bandes transversales qui couvrent ces mêmes ailes
    d’une manière plus ou moins accusée; assez souvent elles vont
    se répétant sous les secondes ailes, et s’y continuent d’une
    manière parfois assez suivie, et selon l’expansion donnée aux
    ailes. Elles y sont même, à mon avis, 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.

    Voici donc les taches et les dessins qui, après nous avoir
    aidés à reconnaître la forme et la nature plus ou moins opaque
    de ces ailes, sont actuellement eux-mêmes contrôlés par la
    constitution physique de ces organes, restitués à leur places
    voulues, et sous l’aile qui les doit comporter.

    Voyons actuellement si l’étude du système 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 reconnaître que ces précieux vestiges sont
    parfaitement indiqués là où ils doivent être, sur [83] cette
    copie de la pièce originale, et que le dessinateur nous les
    laisse suivre assez facilement, tant à une aile qu’à l’autre.

    Que reste-t-il de la charpente alaire de la première aile?
    D’abord, des traces de la costale; puis, au-dessus de la
    lunule blanche de l’apex, les premier et deuxième rameaux des
    trois apicales qui doivent jaillir de la troisième 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 deuxième supérieure, et plus bas, enfin,
    la première.

    Sur le bord extérieur, je devine encore l’extrémité des deux
    premières inférieures; à 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 deuxième
    inférieure; enfin, je suis non moins facilement, entre les deux
    taches noires orbiculaires, la troisième inférieure, un peu
    moins accusée.

    Ces deux nervules se relient visiblement à la portion très
    lisible de la médiane qui, sur le dessin, coupe le bas de la
    première tache blanche costale.

    Toujours à la première aile, la troisième inférieure s’y
    reconnaît parfaitement à la place voulue, au milieu de la
    dent qu’elle soutient. En effet, le plus souvent, quand une
    dentelure, pareillement située, affecte le bord extérieur des
    premières ailes, ainsi qu’on le peut remarquer dans les _Van.
    Progne_, _Archesia_, _L.-album_, _Anglica_ et autres, cette
    troisième inférieure a la prérogative de lui servir de support.

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

    Plus bas, avant l’angle interne, un autre léger faux trait me
    semble encore inexplicable, car la saillie dentée de la marge
    au dehors, précise assez la place, qu’à la première aile, doit
    occuper l’extrémité de la sous-médiane qui d’habitude reste
    volontiers assez distante de la médiane. En plus, il ne saurait
    exister ici d’intermédiane, dont la présence ne se révèle que
    dans les tribus trop éloignées de celle dont la _Sepulta_ fait
    partie, pour nous en préoccuper ici.

    La nervulation de notre première aile se trouve donc ainsi être
    au complet, ou à peu près, et déjà je la peux réhabiliter avec
    le crayon, telle qu’elle doit être.

    Passons à la seconde.

    La côte seulement se soupçonne, on voit parfaitement comme des
    traits blancs, la costale qui est ici très brève en son trajet;
    puis la sous-costale, la médiane, et le dé-[85] part des trois
    supérieures qui se relient très bien en blanc sur la marge
    dentelée de la tache basale (la deuxième moins facilement).

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

    Vient enfin la troisième inférieure, représentée, peut-être,
    par une forte ligne blanche, et qui doit s’attacher à la
    médiane, peu avant l’articulation fémoro-tibiale de la patte.

    Dans les bords postérieur et abdominal, je ne peux distinguer
    ni la sous-médiane, ni l’interne, qui sont disparues dans la
    portion restante, et évidemment diaphane de cette aile qui 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 [Pl. 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 transparence, ou bien sur ceux de
    la fig. C, qui est l’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 à la
    première aile, ainsi qu’aux vestiges de la charpente alaire
    de la seconde aile. Ainsi, on pourra facilement contrôler mes

    Si donc la Solénoptérologie vient à 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

    Comme plus haut je l’ai dit, c’est évidemment une espèce aux
    premières ailes fortement échancrées et dentelées, tandis
    que les secondes y sont arrondies et simples, à méplats bien

    Avons-nous dans nos espèces 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

    Evidemment la _Sepulta_ ne saurait être une _Cyllo_ proprement
    dite. Serait ce donc une Vanesside? Si la forme des ailes
    s’y prête quelque peu, son faciès, l’agencement des ses
    dessins alaires, me porterait à en faire avec M. Boisduval un
    _Satyride_, appartenant à un de ces genres inter-[87] médiaires
    de ces deux familles nombreuses, déjà si peu éloignées à leur
    état parfait.…

    A essayer de caser cet insecte, j’abuserais à n’en pas douter
    de la patience du lecteur; cependant, en peu de mots, je
    pourrais lui faire observer (en ne nous occupant que de la
    première aile, la seule que nous connaissons, à mon avis) que
    la large tache basale qui se voit ici, comme à tant de Diurnes,
    est avec les autres dessins de sa robe, le propre de nombreux
    Satyrides de cette taille et de cette coloration, qu’avec
    justesse M. Boisduval reconnaît devoir être d’une teinte
    terreuse, 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 à l’endroit (entre les
    première et deuxième supérieures) où cette tache existe le plus
    habituellement dans nombre de Satyrides de ce faciès, 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
    qu’elle se voit encore entre la dent et l’angle interne.

    La nervulation si peu différente parfois entre nombre de
    Satyrides et de Vanessides, ne permet pas, sous ce rapport,
    d’assigner un poste bien fixe à 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 renflements vésiculeux si communs à divers groupes de
    Satyrides. [88]

    Nous ne savons 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 côte, s’accorde moins avec la marche plus
    volontiers prolongée de cette même nervure dans les Satyrides,
    de l’apparence de la _Sepulta_, etc., etc.

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

    _Explication des figures de la planche._

    A [reproduced in our Pl. I, fig. 14]. _Cyllos epulta_, telle
    qu’elle a été comprise par M. le docteur Boisduval, en
    attribuant un appendice caudal à la deuxième aile.

    B [reproduced in our Pl. I, fig. 16]. _Sepulta_, telle qu’elle
    devrait être d’abord comprise sous le rapport de la forme des
    deux ailes.

    C [reproduced in our Pl. I, fig. 15]. _Sepulta_, telle qu’elle
    doit être jugée, tant pour la forme des ailes que pour la
    distribution de leurs dessins et leur nervulation, selon M. A.

To this Dr. Boisduval at once responded, in the following language:[I]—

    M. Al. Lefebvre, après avoir étudié avec soin la position des
    nervures, la disposition des écailles et celle des taches, …
    est arrivé à conclure que j’avais pris l’aile inférieure pour
    la supérieure, et que cet appendice caudal, si manifeste dans
    l’espèce en question, était au contraire un angle appartenant à
    l’aile antérieure. Pour donner plus de poids à cette opinion,
    il a refait une planche où il ressuscite à sa manière notre
    _Cyllo sepulta_. Avec la queue que nous avons attribuée avec
    MM. Boyer de Fonscolombe, de Saporta, Duponchel, et avec
    tous les entomologistes qui ont vu l’échantillon à l’aile
    inférieure, il fait un angle très aigu d’une saillie tout à
    fait insolite, qu’il place au milieu de l’aile supérieure,
    tandis qu’il a fait une aile inférieure complètement arrondie.
    A côté de celle figure, il en donne une autre où il développe
    notre _Cyllo_ comme il prétend que nous l’avons compris. J’en
    demande bien pardon à mon estimable ami, mais jamais je ne
    l’ai compris de cette façon. Je conviens du reste que cet
    intéressant Lépidoptère 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’à présent nous n’avons jamais vu une seule espèce
    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
    anguleuses, les ailes inférieures l’étaient aussi d’une manière
    très manifeste; mais ce que personne de vous ignore, Messieurs,
    c’est que très 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 out leur contour simplement
    sinué. A l’appui de son opinion d’ailes inférieures arrondies,
    aves des supérieures anguleuses, notre collègue a cherché à
    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 des ciseaux, car nous
    en possédons un très bel exemplaire, pris par M. Drege au pays
    des Hottentots, que nous mettons sous les yeux de la Société,
    afin qu’elle s’assure bien qu’au contraire cette espèce est une
    des plus fortement appendiculée. Le choix de cet exemple est
    malheureux. Nous persistons donc tout à fait dans l’opinion que
    nous avons émise 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:[J]

    This very interesting species was described and admirably
    figured by Dr. Boisduval in the French “Annales de la Société
    Entomologique” (1840); that gentleman considered it to be a
    Satyride allied to _Satyrus rohria_, _caumas_, etc.,[K] 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 Vanessidæ, as an example of which he instanced _Vanessa_
    (_Junonia_) _Archesia_ of Cramer. This remarkable 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 espèce 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,[L] 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 Pl. I, fig. 10), although his
fault is primarily due to the inaccuracy of the engraving given by
Boisduval, an inaccuracy which is slightly accentuated in our copy of
it (Pl. I, fig. 17). But by far the larger part of his paper is made up
of a detailed argument, drawn from the position and character of the
markings and from the direction of the nervures, in which he endeavors
to prove, and in most cases really does prove (though he errs in some
of his statements concerning the neuration), that these markings belong
to the front and not to the hind wing. He argues, for instance, that
the two oval dark spots are plainly traversed by the nervures of the
hind wing, and therefore cannot belong to that wing; that the minute
white spot apparently on the outer border of the hind wing is only
half a spot and must belong to the fore wing, and that the markings on
and near the costal border traverse both wings and must belong to the
one to which they certainly belong in part, the front wing. To this
Boisduval makes no sort of answer, and Butler, to judge from his silence
in the matter, and the comparative illustrations he gives on a plate
published subsequently,[M] considers it unproven. All of these writers
are, however, entirely wrong in supposing that the under surface of the
wings is exposed to view, and that the hind wing covers the front wing.
Boisduval does not distinctly state this; but the whole tenor of his
remarks shows that this was the view taken by him; and when Lefebvre
says: “Si de l’œil on suit les bords de la seconde aile, qu’avec le
Docteur je reconnais couvrir en grande partie la première,” no objection
is offered in Dr. Boisduval’s response; nor does he demur to Lefebvre’s
statement, when the latter speaks of the “face inférieure, celle que
nous voyons.” As we shall show later, however, the upper surface of the
wings is that exhibited on the stone, and the front wing almost entirely
conceals the hind one; compare Pl. I, fig. 13, drawn anew from the fossil.

In the same place to which we have just referred Mr. Butler adds the
following remarks on the probable affinities of this fossil:[N]

    The true position of _C. sepulta_ is undoubtedly in the family
    _Satyridæ_; and, so far as can be judged from the beautiful
    figure in the “Annales,”[O] it is exactly intermediate in
    character between three nearly allied genera now existing,
    viz.:—_Neorina_, _Antirrhæa_ and _Anchiphlebia_, its more
    immediate allies being the commonest species in each of the
    above genera. Its characters are distributed between these
    three species as follows:—

                                 _Neorina_   _Antirrhæa_   _Anchiphlebia_
                                  _Lowii_,  _Philoctetes_,  _Archæa_,
                                  Boisd.      Linn.           Hübn.
    Form of front wings,              *
    Form of hind wings,              . .         *
    Tails of hind wings
      (intermediate in
      character between),             *          *
    Submarginal spots of
      front wings,                    *          *              *
    Black disco-submarginal
      spots of hind wings,           . .         *
    Pale costal and discal
      banding of wings,               *?        . .             *
    Limitation of dark dentated
      basal area of hind wings,      . .        . .             *
    Submarginal line of hind wings,   *

    The venation appears to be nearly similar to that of
    Anchiphlebia. It is doubtful, however, whether the drawing
    of the veins has been sufficiently attended to, to offer any
    reliable characters.

In this paper he quotes Boisduval’s locality “Aix en Provence,” but when
he next refers to this insect[P] he gives it as from “Aix-la-Chapelle,
White sandstone,” a mistake, however, corrected subsequently. In this
latter paper he remarks:

    I have discussed the position of this species in my catalogue
    of Satyridæ, pp. 189, 190; showing that its nearest ally is
    _Neorina Lowii_, a common Bornean species, but that it also has
    a slightly more distant relationship to _Antirrhæa Philoctetes_
    and _Anchiphlebia Archæa_, two common tropical American
    forms; the amount of affinity, as regards the first two of
    these species, may be seen on my plate, figs. 4 and 5; the
    resemblance to Anchiphlebia is less striking, and the affinity
    more doubtful; it has nothing to do with Cyllo.

That Butler should have so nearly pointed out the exact affinities of
this insect from the simple study of Boisduval’s plate, is unquestionably
due to his extended familiarity with butterflies, and especially with the
forms of this subfamily; but it also shows the essential harmony between
the markings of the under and upper surface of the wings of butterflies,
notwithstanding their frequent great dissimilarity; for Butler compares
this fossil with the recent forms on the assumption that the under
surface of the wings is seen in Boisduval’s plate.

The actual condition of the fossil, for an opportunity of examining which
I am indebted to the courtesy of Count Saporta, is this (see Pl. I, fig.
13): The thorax, hind legs and both pair of wings of the left side are
preserved, almost completely; all the rest is lost. The thorax is viewed
from above and somewhat on the left side; the hind coxæ seem to be almost
torn away from their immediate connection with the trunk. The two hind
legs are stretched out bent at the femoro-tibial articulation; the left
leg lies above both the wings and is apparently attached throughout,
although its base is covered a little by the crushed body; the right
leg lies below both the wings and is apparently partially detached,
though but slightly, from the coxæ; the tibio-tarsal articulation can
be distinguished (Pl. I, fig. 11) but not the tarsal joints. The wings
are bent over downward in a position the reverse of that of repose. The
fore wing covers the hind wing as in nature, but to such an extent as to
conceal the greater part of it; the guttered portion of the inner margin
of the hind wings is almost fully expanded, but apparently has a fold
next the submedian nervure. The fringe of the fore wing seems to be gone,
but that of the hind wing is preserved nearly throughout. Head, fore and
middle legs, wings of the right side and abdomen are wholly wanting.

The upper surface of the wings is, therefore, the part which attracts
most attention. That it is the upper and not the under surface which
is exposed to view is shown by the relation of the wings to each other
(Pl. I, fig. 10), by their unquestionable attachment to the thorax, of
which we certainly see only the upper portion with its smooth arched
dome marked by the sutures which separate the portions which compose
it; and by the design itself of the wings, which is such as pertains
to the upper rather than to the under surface of butterflies of this
group. These markings are most wonderfully preserved; and the careful
and prolonged study I have given every part of the fossil has enabled me
to separate, with a considerable degree of certitude, the markings which
appertain to the fore wing and those which belong to the hind wing. Those
of the latter are generally to be traced through the semi-diaphanous
fore wing and are given in Pl. I, fig. 8. One is aided greatly in this
investigation by following the lines and series of markings which extend
over both the exposed and covered portions of the hind wing; and then
by comparing the fainter and obscurer tints of the covered portion with
equivalent marks on other parts of the stone covered by both the wings;
in this way the markings of the hind wing may be separated from those
of the front wing, but subject, certainly, to some degree of doubt. In
the figure upon the plate (Pl. I, fig. 8) the portions to which the
least degree of doubt attaches are the outer halves of the two wings. I
am inclined to consider these as almost absolutely accurate. The parts
on the other hand which are more likely to be inaccurate are the basal
halves of the median interspaces of the fore wing and the contiguous
portion of the medio-submedian interspace. Assuming, however, that the
drawing faithfully represents the real markings of this extraordinarily
preserved fossil, a detailed description of its features follows.

The basal portion of the fore wing (Pl. I, fig. 8) is very dark, and
increases in intensity toward the border of the innermost light patch;
the latter is bounded by a line running in a nearly straight course
from the costal nervure, opposite the middle of the upper border of the
cell, toward the middle of the apical half of the submedian nervure;
but it extends slightly outward on reaching the lowest median nervule
and just below this turns baseward and makes a large ovoid curve of
an interspace’s diameter, returning to its course when it has nearly
completed the circuit and reached the middle of the medio-submedian
interspace; the outer limit of this large pale patch, which crosses the
cell and extends nearly to the middle of the lower median interspace,
nearly follows a line running from the upper extremity of the inner
border to and along the middle median nervule. Beyond this the upper
half of the wing, half-way to the apex, is nearly as dark as the basal
part, excepting in a large light patch which crosses the lowest two
subcostal and the subcosto-median interspaces, is broadest in the middle,
but twice as broad at the upper as at the lower extremity, and rounded
throughout excepting at the angular upper basal corner; its interior
margin is sharply defined, and is nearly parallel to the interior border
of the inner light patch, extending in a straight line from the subcostal
nervure midway between the origin of the first and second superior
nervules to the upper median nervule, about as far from its origin as
it is from the base of the first median nervule; the exterior border is
powdery, strongly convex and, starting from the subcostal nervure midway
between the bases of the second and third superior nervules, joins the
other border on the last median nervule; this patch is twice as long as
broad. Extending from the next to the lowest subcostal nervule to the
internal nervure, parallel to the outer border, is a broad indistinct
pale band, broadening below, and on either side merging indefinitely
into the darker parts of the wing, separated from the light patches by
only a narrow belt of dark scales, which becomes narrower and fainter
in the lower half of the wing; at its broadest the pale band is a
little broader than an interspace, and it contains in its middle and
at the middle of each interspace, as well as in the next to the lowest
subcostal interspace, a series of large circular dark spots, of nearly or
quite half the width of the interspaces in which they fall, often, and
especially in the upper interspaces, enclosing a small black pupil; these
spots are almost exactly parallel to the outer border, that in the lowest
median interspace with its outer border at an interspace distance from
it; with the exception of that in the lowest subcostal interspace, they
are each surmounted interiorly by a much smaller circular light spot, the
centre of which is near the circumference of the larger spot, so as to
infringe upon it; with the exception of the uppermost, which is nearly as
large as the spot on whose summit it is placed, the light spots are of
nearly equal size and about one-third of an interspace in diameter; or if
anything the two lower, seated on the largest spots, are smaller than the
others; the wing must have been wrinkled between the nervules next the
outer border, as shown by the dark lines running from the border to the
centre of the dark spots. The outer edge and the apex of the inner are
uniformly dusky and rather lighter than the other dark parts of the wing;
the fringe is evidently lost.

The hind wing is very dark at the base, like the fore wing, nearly as
far as the extreme tip of the cell; this dark area merges gradually into
a lighter portion, which crosses the wing as a very broad equal band
having its outer limit at a narrow, dark, regular belt, with ill defined
outline, which crosses the wing subparallel to the general course of the
outer border a little within the middle of the outer half of the wing;
within this broad light band are two narrow transverse powdery streaks
of dark scales, one extending from the extreme tip of the cell, and
broadening a little in its course, running in a curve opening inward to
the inner border; the other starting from the same point in an opposite
direction, and passing in a sinuous course, with varying width, toward
the middle of the basal two-thirds of the upper subcostal nervule,
hardly separate from the outer limits of the dark base of the wing. The
darkest part of the narrow band in the middle of the outer half of the
wing has a regular curve and strikes the borders in the middle of their
outer halves; there is a submarginal slender dark streak, separated by
scarcely more than its own width from the outer border, becoming narrower
toward the costal and inner borders, and especially towards the costal;
it is broken at the upper median nervule, where the upper portion joins
a second broader band, separated by a space nearly equal to itself from
the submarginal band; this leaves a nearly equal light band in the outer
part of the wing, broadest above and reaching from the costal border,
almost to the inner; along the middle of this belt is a series of six
round dark spots and ocelli, one in each of the interspaces excepting
the costo-subcostal; the largest is in the lower median interspace, and
is a spot nearly as broad as the interspace, deepening toward the centre
to a black pupil; the next largest, in the upper median interspace, is
an ocellus with a black pupil, immediately followed by a pale annulus,
again surrounded by a dark ring of equal diameter, the whole a little
more than half the width of the interspace; next larger are two spots of
less intense depth of color, one in the upper subcostal, the other in the
subcosto-median interspace, about one-third the width of the interspace,
the upper deepening, the lower becoming paler at the centre; the spot
in the lower subcostal and the medio-submedian interspace are equal and
smallest, about one-fourth the width of the interspace, and consist only
of rather faint, powdery marks, a little darker towards their centres.
The fringe of this wing seems to be preserved and is short, nearly equal,
dark, resembling a repetition of the submarginal streak.

Length of fore wing, 37mm.; breadth of fore wing, 20·5mm.; length of hind
wing, 31·75mm.; length of tail, 4mm.; distance of the base of the second
superior subcostal nervule of hind wing from the divarication of the
costal and subcostal nervules, 5·55mm.; rows of scales in the subcostal
region of the fore wings, ·075mm. apart; length of thorax, 5mm.; of hind
femora, 4·6mm.; of hind tibiæ 4·8mm.; of hind tarsi, 4·9mm..

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


    _Satyrites_ Scudd. (nee Blanch.-Brullé), Rev. et Mag. de Zool.,
    1871-72, 66.

The costal border of the fore wing (Pl. I, fig. 5) is gently and equably
curved, the apex moderately acute but well rounded, the outer margin,
except at its extremities, nearly straight, and the inner border straight
or almost so; the outer border is a little shorter than the inner and
about three-fifths the length of the costal margin.

The costal nervule terminates slightly beyond the middle of the costal
margin, its basal two-fifths presenting a considerable and almost uniform
expansion, which tapers rather rapidly at the tip, and reaches nearly
to the middle of the upper border of the cell. The subcostal nervule is
very slight on the basal half of the wing, closely connected with the
posterior surface of the swollen portion of the costal nervure and only
divaricating from that vein after the latter has lost its tumidity; it
emits its first superior nervule at slightly more than three-fifths the
distance from the tip of the bulbous portion of the costal nervure to
the upper apex of the cell; its second at midway between the origin of
the first and the tip of the cell; its third at midway between the upper
apex of the cell and the origin of the fourth, which arises at about
two-fifths the distance from the base of the third to the outer border of
the wing. The first superior nervule terminates near the middle of the
outer two-thirds of the costal border, the second midway between the apex
of the first and third; the third terminates just above, and the fourth
at or scarcely below, the tip of the wing. The first inferior subcostal
nervule arises at a very short distance beyond the base of the second
superior nervule, and curving rather strongly, terminates in the middle
of the upper half of the outer border; the second inferior nervule is
emitted from the first inferior as far beyond the base of the latter as
that is beyond the base of the second superior nervule; at its origin it
is directed inward as well as backward (forming the upper termination
of the cell) and passes backward in a small, narrow and rather strongly
curved bow, bent below more than above, beyond which it assumes a course
nearly parallel to the first inferior nervule; just beyond the arcuate
portion it is connected by a rather long, straight, oblique nervule,
directed considerably outward as well as downward, to the origin of
the upper median nervule. The median nervule is slightly enlarged at
the base, and diminishes gradually and regularly in size to its first
divarication, which is scarcely beyond the middle of the cell; the origin
of its middle branch is slightly nearer the origin of the basal than of
the terminal nervule; the latter strikes the middle of the outer border.
The submedian nervure is straight and not swollen at the base. The cell
is three times as long as broad, and scarcely more than half as long as
the wing.

The article from which the above is quoted, as originally written, closes

“The neuration of the fore wing does not seem to me to accord
sufficiently with that of any known genus of Oreades to admit of its
being classed with them. It undoubtedly has close affinities with the
characters of the genus Debis (= Lethe Hübn.) as laid down by Westwood
and Hewitson, if we exclude therefrom, as we should, the _Papilio
Portlandia_ of Fabricius. It is not a little interesting to notice
that these authors have arranged this group in immediate proximity to
the genus Cyllo (= Melanitis Fabr.), in which Dr. Boisduval placed
the fossil species from Aix, named by him sepulta. Nor is it less
interesting to find that in both genera all the living representatives
(even including those discovered since the publication of the ‘Genera of
Diurnal Lepidoptera’) are natives of the East Indies; so that the fossil
butterflies of Provence have their nearest living allies in the far East.”

Although differing from Neorina (Pl. II, fig. 8) very strikingly in the
form of the wing and the swollen base of the costal nervure, this genus
has some striking points of agreement with that in the neuration of the
fore wing. The nervure closing the cell indeed is straight in Lethites
and strongly curved in Neorina, but, as there, two of the superior
subcostal nervules arise before the tip of the cell, and the other two
are thrown off at about equal distances between the apex of the cell and
of the wing; the vein closing the cell meets the median nervure in both
cases as far beyond its second divarication as that is beyond the first;
the shape and proportionate length of the cell is nearly the same in the
two, but the costal nervure appears to be much shorter in Lethites.

With Lethe (Pl. II, fig. 6) and Debis (Pl. II. fig. 10) the fossil genus
can better be compared, as far as the form of the wing, the dilated
costal vein, and the position and direction of the straight vein closing
the cell are concerned; but in both these genera only a single superior
subcostal nervule is emitted before the apex of the cell; the form of
the cell again shows rather closer affinity between Lethites and these
genera, although the difference in these respects is but slight. It
is by no means distantly related to Enodia, in which two subcostal
nervules are emitted before the tip of the cell, but differs from it in
the much greater and more abrupt swelling of the costal vein, and in
the much greater distance beyond the second divarication of the median
nervure at which the vein closing the cell meets this nervure. It even
exhibits no small affinity to Cercyonis, and especially to those species
in which there is little dilation of the median nervure; the costal
nervure is swollen in precisely the same way, and the superior nervules
of the subcostal nervure are much the same; but the form of the wing is
strikingly different, and the lowest subcostal interspace much wider at
the base, in comparison with the width of the base of the subcosto-median
interspace, in Cercyonis than in Lethites; and this seems to be a
character of considerable importance. It may be noted in this connection
that the markings of the fossil must have closely resembled _Cercyonis

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.


Plate I, figs. 2, 5.

    _Satyrites Reynesii_ SCUDD., Rev. et. Mag. de Zool., 1871-72,
    66-72, pl. vii (1872); IB., Descr. Pap. Foss. 1-7, pl. (1872);
    IB., Geol. Mag. ix, 532-33, pl. xiii, figs. 2, 3 (1872); IB.,
    Descr. Foss. Butt. 1-2, pl., figs. 2, 3 (1872); BROD., Distr.
    Corr. Foss. Ins. [_Satyrites Reynesii_], 8-9 (1873).

I give below the original of the first paper cited above, excepting the
portion which was quoted under the genus.

In a recent examination of the rich collection of fossil insects from
Aix, preserved in the Museum of the city of Marseilles, my attention
was attracted by two little slabs containing the traces of a fossil
butterfly. Although by no means so well preserved, nor so perfect as the
remains of a butterfly from the same beds, described by Dr. Boisduval
more than thirty years ago, a glance showed that it could not be referred
to that species, since the costal nervure of the fore wings was greatly
swollen. No such form having to my knowledge been described from these
beds, Dr. Reynès, the accomplished director of the establishment,
courteously placed the best specimen in my hands for closer study; and
from it the following account and illustrations have been drawn. The
second specimen is very imperfectly preserved, but since it exhibits in
all its features an exact resemblance to similar parts in the better
specimen it undoubtedly belongs to the same species.

The fossil (Pl. I, fig. 2) is a natural cast of a butterfly lying upon
its side, the wings folded back to back, the legs extended as if hanging,
the tongue uncurled and, with the antennæ, drooping in a direction
similar to that of the legs. The right fore wing, which lies beneath, is
pushed a little outward and also forward, even at its base, showing that
the specimen must have been greatly macerated in very quiet water, before
being covered by the deposits which have preserved its more essential
features. The condition and position of all the parts also lead us to
conjecture that it was swept into its final resting place by a gentle
current, which left the slighter appendages lying in the direction of its
final action.

It is evident that the object is a cast, for the veins of the wing which
lie uppermost on the stone are impressed as we see them on the upper
surface of the wings of living Oreades, while those of the wing lying
beneath (veins which are plainly covered by the impressed nervures where
the two come in contact) are in relief, as seen on the under surface of
the same butterflies; that is, we have here the reverse of what would be
the case, were we examining a living butterfly in this position.

The parts before us are: a poorly preserved body, vague indications of
the terminal palpal joint, an antenna (probably a portion only), an
unrolled tongue, the hinder pair of legs and portions of the other pairs,
the greater part of the two front wings and fragments of the base of
the hind wings. Of the latter, no border remains and only the base of a
few of the nervules, which give scarcely any additional information as
to the pterology of the insect. The only portion of the margin of the
front wings which can be determined with certainty is the most essential
part, the apex and the upper half of the outer border of the left wing,
enough to show that its general contour was similar to that of the
European Satyrids of the present epoch; but throughout the remainder all
the nervules can be exactly traced. This being then the best preserved
portion of the insect, we will consider its structure in detail,
subsequently adding whatever can be gleaned from the examination of the
other parts.[Q] [The account of the structural framework of the wing is
given under the genus].

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 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 insertion for
the scales, outlines of the latter of which I have failed to discover.
The wing is 28·5mm. long, the tip of the cell being distant 15mm. from
the base of the wing; the costal nervure is inflated for a distance
of 6·5mm., and the extreme width of this portion is 1mm.; the rows of
punctures indicating the former insertion of the scales are ·12mm. 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 specimen 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 ·75mm. beyond the
head and is of a nearly uniform width (·2mm.), scarcely tapering, with a
rounded tip. The basal portion of an antenna, 5mm. long, is slender and
apparently begins to increase slightly and very gradually in size, as in
the genus Œneis Hübn. A finely impressed line, 7·25mm. long, appears to
be the unrolled, though slightly curved tongue.

One of the hind femora projects 2·5mm. 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 apparently 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 5·6mm.; but if both hind pairs are present,
their probable length is 4·5mm.. 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 our Pl. I, fig. 5, which, as well as fig. 2, is taken from the
originals of my former plate.

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



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 receding at a little more than a right
angle to the middle of the subcosto-median interspace, 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.


Plate I, fig. 1, 3, 7.

    _Sphinx atava_ CHARP., Acta Acad. Leop.-Carol., xx, 408-9, Tab.
    22, fig. 4 (1843).

    _Vanessa attavina_ HEER, Insekt. Tert. Œning., ii, 177-79, Taf.
    14, fig. 3 (1849); IB., Nouv. Mem. Soc. Helv., xi, 177-79, Tab.
    14, fig. 3 (1850); GIEB., Deutschl. Petref., 644 (1852); IB.,
    Faun. der Vorw., ii, 186 (1856).

    _Vanessa? atovina_ KIRB., Syn. Cat. Diurn. Lep., 185 (1872).

    _Nymphalis? atovina_ KIRB., 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:[R]

    Ungemein interessant, und ich möchte sagen, ein Unicum ist der
    in oben bemerkter Figur abgebildete Schmetterlingsflügel. Dass
    es ein solcher sei, zeigt sogleich der erste Anblick, so wie
    sich bei näherer Ansicht herausstellt, dass es unbezweifelt der
    Oberflügel einer Sphinx Art sei. Er ist in seiner Form nicht
    gut gehalten, sondern vorn etwas eingerissen, seine Zeichnung
    ist aber bewundernswerth erhalten, und erinnert sehr an den
    fast im ganzen mittleren und nördlichen Europa vorkommenden
    _Sphinx Tiliæ_, doch ist er wohl specifisch von demselben
    verschieden. Die drei grossen dunklen Flecke, die sich von
    äussersten Vorderrande, fast bindenartig, über einen grossen
    Theil des Flügels ziehen, sind unstreitig die Reste ehemaliger
    Zeichnung und Färbung 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
Pl. 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 redescribed it in
the following manner, referring it to the genus Vanessa, and changing
slightly the specific name:[S]

    Alis anterioribus lividis, basi, fasciis maculisque nigris.
    Long 16⅞ Lin.

    Radoboj. Ein Oberflügel, dessen Innenrand aber nicht erhalten

    Charpentier hat diesen Flügel einem Sphinx zugesprochen und
    ihn mit dem Sphinx Tiliæ L. verglichen; allein schon die
    ziemlich stark gebogene Randlinie (vena marginalis) spricht
    gegen Sphinx, bei welcher Gattung sie bis über zwei Drittel
    Flügellänge fast gerade verläuft und dann erst gegen die Spitze
    sich zubiegt; ebenso aber auch das Geäder. Bei Sphinx haben
    wir nämlich ein geschlossenes Mittelfeld und der Ast der
    vena externo-media, welcher neben der Flügelspitze ausläuft,
    verästelt sich nicht. In der Form des Flügels, im Geäder und
    Färbung stimmt unser Schmetterlingsflügel, wie mir scheint,
    am besten mit der Gattung Vanessa F. überein. Wir bemerken
    nemlich, gerade wie bei den Vanessen, zunächst eine starke vena
    scapularis, welche weit vorn in die v. marginalis ausläuft;
    eine schwächere vena externo-media, welche noch näher der
    Flügelspitze mit dem Rande sich verbindet; diese bildet nach
    Innen zunächst einen Ast[T], der flügelspitzwärts in zwei
    weitere Aeste sich spaltet; der äussere von diesen läuft zur
    Flügelspitze, der innere aber trennt sich nochmals in zwei
    Gabeläste, welche zum Hinterrande verlaufen und von denen jeder
    in einen schwachen, stumpfen Zahn des Flügelrandes ausgeht.
    Auf diesen Gabelast folgen weiter nach Innen zwei Längsadern,
    welche am Grunde sich wahrscheinlich verbinden, und in die
    vena externo-media eingefügt sind. Diese beiden Adern (es
    sind diess die fünfte und sechste Ader von Herrich Schaeffer)
    gehen bei [178] den Vanessen getrennt bis zur v. 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 Einmündung des äusseren Astes in die
    vena externo-media, indem der innere am Grunde ganz verwischt
    ist, wie denn überhaupt die Adern in Folge des starken Druckes,
    dem der Flügel unterworfen war, äusserst schwach hervortreten
    und nur mit Mühe zu erkennen sind. Die vena interno-media
    verläuft wie bei den Vanessen, sie sendet nämlich nach dem
    Hinterrande zwei Aeste aus, so dass im Ganzen drei Längsadern
    zuletzt in parallelen Linien nach dem Rande verlaufen. Die
    vena analis ist nur am Grunde angedeutet, indem der Innenrand
    grossentheils zerstört ist. Das Mittelfeld ist offen,
    wenigstens ist keine Spur eines Verbindungsastes zwischen v.
    externo-und interno-media zu finden. In allen diesen Punkten
    stimmt also das fossile Thier mit den Vanessen überein.
    Ebenso stimmt ferner der zackige Hinterrand, indem wir, wie
    schon bemerkt, an der Ausmündung des äusseren Gabelastes der
    v. externo-media kleine Zacken bemerken, wobei freilich zu
    bedauern, dass von dort an der Flügel zerrissen ist, so dass
    die Randbildung nur an jene kleinen Stelle bestimmt werden
    kann. In der Färbung zeigt der Flügel viel Uebereinstimmendes
    mit demjenigen der Vanessa Cardui L. Wir bemerken nemlich
    zunächst dem Grunde eine dunklere Stelle, welche fast bis
    zu ⅓ Flügellänge hinausreicht; dieser dunklere Flügelgrund
    ist indessen wieder in der Mitte durch einige unregelmässige
    hellere Stellen unterbrochen. Auf diese dunkle Stelle folgt ein
    helles Querband von 1¾ Linien Breite, welches aber nicht bis
    zum Innenrande reicht, wenigstens 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 folgt wieder ein 3 Linien breites dunkles Querband,
    welches mit mittleren schwarzen Querband der V. cardui
    entspricht; bemerkenswerth ist, dass dieses bei der V. attavina
    von der Nahtseite her ebenfalls durch einen helleren Flecken
    getheilt wird, welcher helle Flecken nicht his zum Aussenrand
    hinausreicht. Auf dieses dunkle Querband folgt wieder ein
    helles Band von 1¼ Lin. Breite, und darauf wieder ein dunkler,
    3¼ Lin. breiter Flecken, der aber sehr kurz ist, indem weiter
    nach Innen an jener Stelle der Flügel wieder hellgelb braun
    gefärbt ist; auf diesen dunklen Flecken folgt wieder ein
    kleiner heller Flecken; weiter flügelspitzwärts ist der Flügel
    dunkelbraun gefärbt, welche Farbe allmählig heller wird, so
    dass der Flügelrand wieder hellbraun wird; die Zackenspitzen
    dagegen sind schwarz.

    In der Färbung des Oberflügels stimmt also der fossile
    Schmetterling am meisten mit Vanessa Cardui L. überein, dennoch
    kann er nicht als analoge Art betrachtet werden, denn fürs
    erste war er beträchtlich grösser [179], fürs zweite ist die
    Randader stärker gebogen, zeigt eine regelmässige Bogenlinie,
    während sie bei Vanessa Cardui in mehr gerader Linie verläuft.

The only subsequent notice of this insect, not directly copied or
abbreviated from the above is by Butler, who remarks[U]; “I think it just
possible, from the great resemblance which _V. Attavina_ of Heer bears to
the under surface of _J.[unonia] Hedonia_, that it is the reverse of _J.

I have been unable to see this fossil, or even to find out where it is
preserved. Charpentier states that he received it for description from
Dr. Unger through Professor Göppert 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
remarks 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 European vau-album or our own j-album. A comparison of the neuration
of _Eugonia j-album_ (Pl. I, fig. 4) with that of Heer’s figures of the
fossil (reproduced on Pl. 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 same in
position, direction and intensity, as in _E. j-album_ (Pl. 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 connection 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.



Of the form of the fore wing (Pl. 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 (Pl. II, fig. 7) is very similar to that
of Mylothris.[V] 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 nervule is emitted
from the subcostal nervure at less than half the distance 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;
the median nervure is of course three-branched and scarcely curves upward
at all to meet the subcostal.

The design of the upper surface of the fore wing (Pl. 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 preservation 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 original, 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.


Plate II, figs. 2, 7, 17 (15?).

    _Vanessa Pluto_ HEER, Insekt. Tert. Œning., ii, 179-82, Taf.
    14, fig. 4, 5 (?) (1849); IB., Nouv. Mem. Soc. Helv., xi,
    179-82, Tab. 14, figs. 4, 5 (?) (1850); GIEB., Deutschl.
    Petref. 644 (1852); IB., Faun. der Vorw., 186-7 (1856); Pict.,
    Traite de Palæont., ii, 393, pl. 40, fig. 21 (1854); LYELL,
    Elem. Geol., 6th Ed., 243, fig. 179 (1865).

    _Argynnis Pluto_ EDW., Butt. N. Amer., I, Argynnis I, fig.
    (1868); Kirb., Syn. Cat. Diurn. Lep., 155 (1871).

    _Junonia? Pluto_ BUTL., Lep. Exot., 127-28, pl. 48, fig. 7
    (1873); IB., Geol. Mag., x, 3-4, pl. 1, fig. 7 (1873).

Heer’s description of this insect is as follows:[W]—

    Alis griseo-nigris, anterioribus margine posteriore ocellis sex

    Länge des Vorderflügels wahrscheinlich 15 Lin; er ist erhalten
    bis zu 14¼ Lin; grösste Breite 8¾ Lin.

    Radoboj. Ein ausgezeichnet schönes Exemplar in dem k. k.
    Hofkabinet zu Wien; leider fehlt aber der Kopf, der Hinterleib,
    der grösste Theil der Hinterflügel und die Spitze der
    Vorderflügel [Pl. II, fig. 2].

    Der Brustkasten ist länglich oval, in der Mitte zwei Linien
    dick, an der Oberseite von ein paar Streifen durchzogen. Der
    Oberflügel ist am Grunde schmal, nach dem Hinterrande hin
    aber stark verbreitert und erreicht daselbst seine grösste
    Breite. Die Aussenrandlinie (v. marginalis) ist sehr stark
    gebogen, und zwar bildet sie vom Grunde zur Spitze eine
    regelmässige, starke Bogenlinie. Die Schulterader ist am
    Grunde stark ¾ Lin. vom Rande abstehend und läuft ausserhalb
    der Flügelmitte in denselben; die vena externo-media ist ihr
    sehr genähert und nur mit Mühe zu unterscheiden, sie mündet
    noch näher flügelspitzwärts in die Randader. Sie sendet
    zunächst einen einfachen Ast ab, der mit dem Hauptstamm
    parallel läuft, ihm sehr genähert ist und noch näher der
    Flügelspitze in die Randader mündet; der zweite Ast spaltet
    sich bald wieder in zwei Aeste, von denen der äussere vor der
    Flügelspitze in die Randader auslauft, der innere theilt sich
    nochmals in zwei Gabeläste, welche ohne Zweifel innerhalb der
    Flügelspitze ausmünden; ganz nahe, wo der zweite Hauptast
    der vena externo-media entspringt, läuft 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 Flügelfläche 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
    einmündet. Das Mittelfeld ist verhältnissmässig ziemlich
    klein und nicht geschlossen, indem kein Querast die beiden
    Mitteladern verbindet. Die vena analis ist einfach und läuft
    nahe dem Nahtrande herunter. In den Feldern zwischen je zwei
    Längsrippen sieht man eine schwache Längslinie, welche vom
    Flügelrande bis zum Augenpunkt läuft; sie stellt eine schwache
    Furche oder Falte dar, die dort im Flügel sich befunden hat.
    Der Hinterrand ist leider nicht ganz erhalten, namentlich fehlt
    die Flügelspitze, de-[180] ren Form zur Bestimmung der Gattung
    so wichtig wäre; es ist daher nicht zu ermitteln, ob diese
    ganzrandig oder gezackt war. Der Hinterrand verläuft in einer
    schwachen Wellenlinie, in der ganz schwache, stumpfe Kerbzähne
    an der Ausmündung der Längsadern liegen.

    Die Farbe des Flügels ist ein dunkles Graubraun; am Grunde
    und im Randfelde ist er dunkler, welche dunklere Parthie aber
    allmählig in die hellere verläuft; gegen die Augenflecken
    zu wird die Farbe wieder dunkler; längs des Randes bemerken
    wir eine Reihe (nemlich 6) von runden, hellen Flecken und
    zwar liegt je zwischen zwei Längsadern ein solcher Fleck,
    welcher das ganze Feld zwischen den Adern ausfüllt. Es reicht
    dieser helle Fleck nicht bis zum Flügelrande, welcher wieder
    dunkler graubraun gefärbt ist. In der Mitte jedes Fleckens
    legt ein schwarzer, runder Punkt; ob dieser noch einen 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
    wahrzunehmen ist. Diese hellen Augenflecken scheinen von keinem
    schwarzen Ring eingefasst zu sein.

    Von den Unterflügeln ist nur der Grund erhalten. Wir sehen da
    die, bald in zwei Gabeläste sich spaltende, vena analis und
    die beiden am Grunde ganz genäherten Mitteladern. Die Färbung
    dieses Flügeltheils ist gleich wie am Oberflügel, und zwar nach
    dem Grunde zu auch dunkler werdend.…

    Die Bestimmung der Gattung, zu welchem unser Thier gehört,
    wird sehr dadurch erschwert, dass der Hinterrand nicht ganz
    erhalten ist. Nach [181] der allgemeinen Form und dem Geäder
    der Flügel muss er wohl zu den Nymphaliden gehören. Bei
    den Papilionen, Pieriden, Danaiden und Satyriden ist die
    Mittelzelle der Flügel durch einen starken Verbindungsast
    der vena externo-und interno-media geschlossen, wogegen beim
    fossilen Thiere die Mittelzelle des Oberflügels, und vielleicht
    auch die des Unterflügels, geöffnet ist, wie diess bei vielen
    Nymphaliden vorkommt. Von den Pieriden unterscheidet er sich
    überdiess durch die Art der Verästelung der v. externo-media,
    indem (um mich der Terminologie von Herrich Schäffer zu
    bedienen) die 7te und 9te Rippe, vom Nahtrande an gerechnet,
    aus der sechsten entspringen, und die achte aus der siebenten,
    während beim fossilen Thiere die 8te und 9te Rippe, wie bei den
    Nymphaliden, aus der 7ten entspringen. So weist also das Geäder
    auf einen Nymphaliden. Unter diesen kommen ein paar Gattungen
    vor (nemlich Apatura und Melitæa) mit offener Mittelzelle der
    Hinterflügel; allein bei diesen finden sich keine Arten mit
    Augenflecken, wogegen unter den Vanessen eine Art vorkommt,
    welche in der Fleckenbildung eine auffallende Aehnlichkeit mit
    dem fossilen Thiere hat. Zwar ist bei Vanessa die Mittelzelle
    der Hinterflügel 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 Uetlandsche Kapellen
    T. II, Taf. 69, C. D. und T. VIII, Taf. 374, E. F. Es hat
    diese genau die Grösse des fossilen Thieres, der Aussenrand
    bildet ebenfalls eine starke Bogenlinie; die Oberflügel
    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 überein, 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 vor, hat also im tropischen Asien eine weite

    Von Schmetterlingen mit ähnlicher Färbung können noch in
    Betracht kommen: die Argynnis Diana Cramer II, p. 4, t. 98, D.
    E. Say. Americ. En-[182] tom. 17, welche im südlichen Theile
    der vereinigten Staaten (Neu-Georgien, Westflorida, Arkansas
    and Missouri) lebt. Es hat dieser Schmetterling eine ähnliche
    Tracht, ist schwarz und am Hinterrande mit einer Reihe gelber
    Flecken versehen, welche je zwischen die Längsadern vertheilt
    sind. Diese gelben Flecken reichen aber bis zum Rande, und
    ferner hat jeder zwei schwarze Punkte. Auch ist die A. Diana
    bedeutend grösser. In Grösse und Färbung stimmt daher das
    fossile Thier mehr mit der Hedonia überein, als mit der Diana,
    doch kann mit voller Sicherheit erst darüber entschieden
    werden, wenn einmal ein Exemplar mit vollständig erhaltenem
    Hinterrand gefunden wird; was von diesem erhalten ist, spricht
    aber auch mehr für die Hedonia als die Diana.

Edwards, in his beautiful work on American Butterflies, refers to this
insect in his description of _Argynnis Diana_[X] 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:[Y]—

    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:[Z]—

    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 Butterflies 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 description (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 woodcuts
    by Lyell and Edwards, which have 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 (Pl. II, 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 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 subcostal 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

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 remarks are as follows:[AA]—

    Hierher rechne ich auch ein Stück eines Unterflügels aus der
    Grätzer Sammlung, das bei Taf. XIV, Fig. 5 [Pl. II, fig. 15],
    dargestellt ist. Die Hauptadern treten an diesem Flügelstücke
    alle hervor. Die beiden Mitteladern schliessen ein nicht sehr
    grosses Mittelfeld ein; ob dieses durch einen Verbindungsast
    zwischen den beiden Mitteladern geschlossen ist oder nicht,
    war mir nicht möglich zu ermitteln: bei guter Beleuchtung
    glaubte ich dort einen schwachen Quereindruck zu sehen, der
    als Verbindungsast zu deuten wäre; jedenfalls wäre derselbe
    aber äusserst zart, viel zarter als die übrigen deutlichen
    Adern. Die äussere Mittelader sendet 4 Aeste aus, der erste
    entspringt nahe der Flügelbasis und läuft nach dem Aussenrande,
    die drei folgenden entspringen näher flügelspitzwärts. Die
    v. interno-media zerspaltet sich in 3 Aeste, ganz so wie die
    des Oberflügels, 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 Gabeläste, welche nach
    aussen laufen. Die Farbe des Flügels ist ein helles Graubraun.

[Illustration: Fig. 1.

Fig. 1. The dotted outer border of the hind wing represents the probable
limit of the Gratz fossil. The broken outer border indicates the probable
size of the hind wing of _Mylothrites Pluto_.]

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[AB] 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 remembered
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; so great a difference is at least unusual among individuals of the
same species in this group; moreover, the 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

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 the fossil was drawn up from the illustration and not from
the fossil itself. Both agree in the points in which my drawing (Pl. II,
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[AC] 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 reëxamination 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.


The fore wing (Pl. 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 gradually 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 nervure 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 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 convexity, 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. II, fig. 4) than to any other genus I have
been able to examine. 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 the wing,
the remarkably straight costa[AD] 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 nervules.


Plate II, 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. 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 longitudinal 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 irregularities
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,
·075mm. 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, 21mm.; greatest breadth, 9mm..

Tertiaries of Aix, Provence, France. Collection of Count de Saporta.



    _Pierites_ HEER, Insekt. Tert. Œning., ii, 182; IB., Nouv. Mem.
    Soc. Helv., xi, 182; GIEB., 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 terminating
a little beyond the middle of the margin. Subcostal nervure with three
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 second 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
interspaces; these occur more often in the female than in the male.

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


Plate II, figs. 16, 18.

    _Pierites Freyeri_ HEER, Insekt. Tert. Œning., ii, 182-83, Taf.
    14, fig. 6 (1849); IB., Nouv. Mem. Soc. Helv., xi, 182-83, Taf.
    14, fig. 6 (1850); GIEB., Deutschl. Petref., 644 (1852); IB.,
    Faun. der Vorw., ii, 187 (1856); KIRB., Syn. Cat. Diurn. Lep.,
    509 (1871).

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

    Alis anterioribus lividis, margine maculisque duabus nigris.

    Länge des Vorderflügels 9¾ Lin., Breite 5½ Lin.

    Radoboi. Ein einzelner Vorderflügel, dessen Spitze und
    theilweise auch Hinterrand aber zerstört ist, in der k. k.
    montanistischen Sammlung zu Wien. [Pl. II, fig. 16.]

    Das Geäder ist nicht in seinem Verlauf zu ermitteln und da auch
    der Umriss des Flügels nicht vollständig 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, wofür auch das dünne Schuppenkleid, das
    er gehabt zu haben scheint, angeführt werden kann, wogegen
    die allerdings stumpfen Zacken am Hinterrand eine abweichende
    Bildung zeigen.

    Der Flügel ist am Grunde stark verschmälert, nach dem
    Hinterrande zu stark verbreitert; der Hinterrand ist stumpf
    gekerbt; in die Bucht der Kerbe läuft eine Längsfalte, in die
    Mitte derselben eine Ader aus, die man aber nicht bis zur
    Insertionsstelle verfolgen kann; die äussere Flügelspitze
    fehlt; ebenso ein Stück des Hinterrandes an der [183]
    Nahtseite. Von den Adern kann man nur die einfache v. analis
    in ihrer ganzen Länge verfolgen; sie ist dem Nahtrande sehr
    genähert. Die Farbe des Flügels ist hell gelbbraun und war im
    Leben wohl weiss oder gelblich. Der Hinterrand ist von der
    Mitte an bis zum Aussenrand schwarz, und zwar wird diese dunkle
    Parthie auswärts breiter; ungefähr in der Flügelmitte geht
    vom Aussenrand ein viereckiger, dunkler Fleck aus, welcher
    dieselbe Grösse, Form und Stellung hat, wie der schwarze Fleck
    bei Pieris Daplidice; ein zweiter kleinerer, rundlicher Fleck
    liegt näher dem Hinter-und Nahtrande und entspricht dem, an
    derselben Stelle liegenden, Flecken der Unterseite von Pieris
    Daplidice.—Am Flügelgrunde bemerkt man den Schenkel und Schiene
    eines dünnen Beines, das wohl diesem Thiere angehört hat.

This insect evidently belongs to the genus Pontia, judging from the
disposition of the markings of the upper surface of the fore wings (Pl.
II, 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_ (Pl. II, 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 three-fifths of the lower
median interspace, where it does not occur in _P. Protodice_; and this
affords the principal ground for supposing the insect to be generically
distinct from Pontia, no distinctive premarginal spot occurring in
this interspace 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 submedian 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, exhibiting 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 24mm. and the greatest width 22.5mm.

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



Body rather robust (Pl. III, figs. 9 and 10). Vertex of head large,
broad, convex. Eyes pretty large, short ovate, their longer diameter
vertical. Palpi (Pl. III, fig. 7) slender, resembling those of Thais,
but rather longer, extending far beyond the eye, rather thinly clothed
with hairs. Antennæ (Pl. III, fig. 8) resembling 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 antennæ 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 (Pl. III,
fig. 6) are a little more than twice as long as broad, their outer edge
nearly straight, the posterior 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

The fore wings (Pl. III, 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 regularly 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 (Pl. III, 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 uniform curve with the outer border, which
is almost entire as in Parnassius (Pl. III, fig. 5), rather than as
in Thais (Pl. III, fig. 4), strongly arched, especially near the last
median nervule, and angulated below where it meets with the regularly and
broadly concave inner margin.

In the neuration of the fore wings (Pl. 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 broadest 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 tip of 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 distance 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 opening 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 nervures 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 is a nearly straight
continuation of the basal portion of the nervure; the outer subcostal
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 nervure is parallel to, and scarcely removed from, the inner

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 subcostal 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,
excepting the subcosto-median and medio-submedian interspaces, all ranged
in a series curving more strongly than the outer border. The hind wing is
nearly uniform on the basal half, but beyond is crossed by transverse,
curving, dark, cloudy bands, broadening on the nervures and enclosing
between them roundish or transversely 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 (Pl. III, 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 antennæ 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 (Pl. III, 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.

In design (Pl. III, fig. 3) Thaites recalls none of the recent genera
very closely. In the fore wings it approaches Thais (Pl. III, fig. 4)
rather than the others, and in the hind wings some species of Parnassius
(Pl. 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
Parnassius 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 have not been able, however, to examine this point carefully.


Plate III, figs. 1, 3, 6-10.

    _Thaites Ruminiana_ HEER, Climat pays tert., trad. Gaudin, 205
    (1861) [absq. descr.]; Sap., Ann. Sc. Nat. [5], Bot., xv, 343
    (1872) [ibid.].

The wings were evidently dark with light markings. On the fore wings the
first transverse stripe (Pl. 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 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 slender, 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 irregular, depending from the first superior subcostal nervule and
extending nearly to the upper median nervule, so that its exterior border
just strikes the subcostal nervure 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 innermost 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
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, 25mm.; breadth of the same, 14·3mm.; length of antennæ, about
6mm.; breadth of antennæ in middle of stem, ·2mm.; breadth of antennæ
toward tip of club, ·5mm..

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



Very much of the general appearance of Thanaos Boisd. (Pl. III, fig.
11) 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 (Pl. III, fig. 12) is fully as stout as in Thanaos (Pl. 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 Urbicolæ. 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 preserved; 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 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
submarginal series of small round spots of which the upper two, in the
uppermost interspaces 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 Thanatites, 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 _T. Juvenalis_ (Pl. 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 _T. 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 cannot 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 Urbicolæ.


Plate III, figs. 12, 16.

    _Vanessa vetula_ HEYD., Palæontographica, viii, 12-13, Taf. i,
    fig. 10 (1859).

    _Araschnia vetula_ KIRB., 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 Pl.
III, fig. 16. The description is as follows:[AF]—

    Es scheint diese Art in die Nähe der bei uns lebenden Vanessa
    Levana zu gehören. Sie ist kleiner als diese, indem der
    Vorderflügel von seiner Basis bis zur Spitze nur 6½‴ misst.
    Der Schmetterling liegt auf der rechten Seite, wobei der linke
    Hinterflügel den linken Vorderflügel völlig bis auf die Spitze
    und einen Theil des Aussenrandes deckt. Von diesen Flügeln ist
    daher nur die Unterseite sichtbar. Der rechte Vorderflügel ist
    mehr vorgeschoben und daher ein grosser Theil seiner Überseite

    Die Flügel sind im Allgemeinen gut erhalten und scheinen am
    Aussenrande an einigen [13] Stellen schwach ausgerandet gewesen
    zu seyn. Sie zeigen auf der Grundfarbe grössere, undeutlich
    schwarze und viele weisse Flecken von verschiedener Grösse.
    Auf den Vorderflügeln zeichnen sich ein grösserer 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 Nähe des Aussenrandes stehen. Auf
    den Hinterflügeln, etwa ein Drittel vom Aussenrand entfernt,
    bilden sechs weisse Fleckchen eine Querreihe. Es ist nicht
    unwahrscheinlich, dass die Grundfarbe der Flügel im Leben braun
    oder rothbraun war, und man glaubt sogar noch einen schwachen
    Schimmer von dieser Farbe wahrzunehmen.

    Der Kopf ist etwas zerdrückt und zeigt zwei ziemlich lange,
    zugespitzte, in die Höhe 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 Flügel
    gedeckt, und von den Beinen sind nur Bruchstücke vorhanden.

Dark brown or blackish with light markings. On the upper half of the fore
wing (Pl. III, 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 nervures 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 distance from it, one in each interspace. Length of fore
wing, 14mm.; length of hind wing, 13·65mm.; extreme breadth of hind wing,

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 decipher
as this. It represents an insect (Pl. 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 antennæ, nor of the right hind
wing (nor of the abdomen?). The left hind wing has an immaterial part
of its outer border removed, 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 von Heyden (Pl. 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
erroneous, although on this point one may not speak with confidence, and
it is comparatively unimportant. It is remarkable, however, that von
Heyden, in his _description_, takes the same view of it as I have done. 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 mottling 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.



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. III,
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 margin 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. III, fig. 15) than with Carystus (Pl. III, 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 differs from that of _Pansydia

In neuration it agrees better with Pansydia than with Carystus. Poey’s
figure, which for want of better material I have been forced to copy
in illustration, is 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 nervure like the others, and at some distance beyond its second
divarication, 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 (Pl. III, figs. 14, 17)
agrees perhaps better with Carystus (Pl. III, fig. 19) than with Pansydia
(Pl. 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 however 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 nervure 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 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 specialized scales. The species of Pansydia are
smaller than those of most of the neighboring genera, but _Pamphilites
abdita_ is somewhat smaller even than _Pansydia mesogramma_.


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 (Pl.
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 superior 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, subtriangular, filling the whole breadth of the interspace,
half as long again as broad, its inner tapering extremity situated just
below the final divarication of the median nervure. 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 interspaces, 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 border of the medio-submedian interspace. Length of
wing, 15·75mm., length of inner border, 9·5mm.; breadth of wing across
the middle, 7·25mm., breadth of wing across outer margin, 9·5mm..

Tertiaries of Aix, Provence, France. Museum of the City of Marseilles.


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 therefore, at least until we
have more precise knowledge concerning the remains of butterfly larvæ 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 platrières même, c’est
à dire des galeries qui servent à 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 espèces; dans ce cas,
elles ne proviennent par des ouvriers mais je les ai recueillies moi même
en suivant les lits sur les points où 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 _Eugonia atava_,
_Mylothrites Pluto_, another fragment possibly referable to Mylothrites,
and _Pontia Freyeri_, all described by Heer. Two of the genera of these
more recent beds contain representatives 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 Œningen,
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 remains of Lepidoptera (one
species) and none whatever of butterflies.


Of the five butterflies from Aix, two belong to the Oreades (_Neorinopis
sepulta_ and _Lethites Reynesii_) the food of whose caterpillars at
the present epoch has invariably been found to be either Gramineæ or,
occasionally, Cyperaceæ. Both of these groups are present in the deposits
of Aix, the former being represented by ten species of Poacites, and the
latter by a Cyperites;[AG] 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, Gramineæ, although some species have
been found also upon Althea, Malva and Lavatera (Malvaceæ), Trifolium,
Coronilla and ?Lespedeza (Leguminosæ), Plantago (Plantaginaceæ), and
Maranta (Scitamineæ). Of these families the Leguminosæ 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 Gramineæ, 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 obtains in New Guinea
or New Grenada, countries the least favored in this respect.[AH] The
proportion of the Gramineæ and Cyperaceæ 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 Gramineæ, etc., to the Phanerogamia being
five per cent., while the proportion of its grass-feeding butterflies 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 Indian aspect, feeding upon plants
essentially temperate, but, as in tropical countries, numerically

The Danai, to which the fourth species from Aix (_Coliates Proserpina_)
belongs, feed almost exclusively upon Leguminosæ, and these have recently
been found in great abundance at Aix. Count de Saporta enumerates one
species each of ?Trifolium, Caragana, Ervites, Sophora, Micropodium,
Cercis and Gleditschia, two of Phaseolites and six of Cæsalpinites,
belonging to the Papilionaceæ, besides nine Acacias and a Mimosa of the
Mimoseæ, and four species of uncertain relations; making a series larger
than he has found in any other family.[AI]

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. 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 rotundiloba_ 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_.[AJ]

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 neighboring genera. Thais feeds principally at least
upon Aristolochia[AK] 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 Crassulaceæ; but nothing very closely allied to this is specified
by Saporta from Aix; neither, also is Aristolochia, but it has been
found not only in Radoboj[AL] in the Mayencian, but also, according to
Heer, at Hohe Rhonen in Switzerland, which belongs to the Aquitanian,
and has at least one plant (_Laurus primigenia_ 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,[AM] and may then be considered, as with
little question, the food-plant of _Thaites Ruminiana_. If it be deemed
hazardous to venture 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:[AN] “Dass indessen auch Weiden oder Pappeln
[Populus] sich vorhanden, dürften der Bythoscopus muscarius und die
Aphrophora spumifera [Homoptera] anzeigen, deren analoge lebende Arten
besonders auf den Blättern und Zweigen dieser Bäume sich umhertreiben.”
The second is a note in the errata to the translation of Heer’s work on
the Climate and Vegetation of the Tertiaries[AO] by Gaudin: “Le Poacites
ciliatus Sap. n’est pas une glume de Graminée, mais plutôt une bractée
ciliée de Peuplier.… Elle doit être probablement rapprochée d’une
empreinte … provenant des mêmes couches et qui se rapporte également au
genre Populus. Les organes voisins de ceux du Pop. Euphratica Oliv. dans
la nature actuelle dénotent l’existence, à l’époque des gypses d’Aix,
d’une espèce de Peuplier dont les feuilles sont encore inconnues, comme
celles de l’Alnus cryptophylla 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 prêter
en paléontologie les diverses branches de l’histoire naturelle.”

The single species from Rott, _Thanatites vetula_, is closely allied to
the modern Thanaos, whose species are numerous and feed upon a variety
of plants, belonging to the families Cruciferæ, Leguminosæ, Umbelliferæ,
Cupuliferæ, Betulaceæ and Salicaceæ. Most of the genera belonging to
its tribe feed upon Leguminosæ, 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 Leguminosæ, mostly belonging to the
Papilionaceæ; they are Templetonia (1 species), Robinia (2), Colutea
(1), Phaseolites (2), Sphinctolobium (1), Dalbergia (1), Hæmatoxylon
(2), Gleditschia (2), Cassia (3), Ceratonia (1), and Acacia (2). It is
probably among these, and perhaps with greatest probability among the
species of Hæmatoxylon 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 habitations of _Thanaos Tages_; and should
traces of silken fastenings be found in connection 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
_Mylothrites Pluto_, and this is a member of the same general group as
Coliates, and feeds probably upon Leguminosæ; for it is not so closely
allied to Delias as Coliates is, but is more nearly related to Hebomoia,
one of whose species, found in the East Indias, feeds upon Capparis.[AP]
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 Cassiæ to have afforded nourishment
to _Mylothrites Pluto_. Moreover, no less than thirty-one species of
Leguminosæ 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 abundance of palatable food.

The food of _Pontia Freyeri_ is doubtful. All the living species of the
genus so far as known, feed upon Cruciferæ; 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. Cruciferæ, however, are
excessively rare in the tertiaries of Europe, two species only being
recorded, and this from the comparatively recent beds of Œningen. 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 Cruciferæ found near the horizon of _Pontia Freyeri_ are a species
of Nelumbium from Günzbourg in the Mayencian, and of Terminalia (_T.
radobojensis_ Ung.)—one of the Calycifloræ, 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, _Eugonia 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., _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 hods 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.


To discuss this question properly we must consider the butterflies of
each geological horizon separately.


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. Cœlites 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 Antirrhæa, a Brazilian genus, but this is too
distant to be given much weight. The closest allies of _N. sepulta_ are
to be found in the Indo-Malayan region.

The same is true, but not to so striking a degree, of _Lethites
Reynesii_. 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, 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 appertaining 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
carefully for very closely allied forms among East Indian Urbicolæ;
but, while it doubtless 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


_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.


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 Mylothris 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 (_E. 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,[AQ] 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 unsupported 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 researches 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 _Examen des flores tertiaires
de Provence_,[AR] 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 exclusively 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 espèces,” to use his own words, “12 seulement
correspondent à des espèces de l’Europe moyenne, 6 à des espèces de
l’Europe méridionale, 18 en tout. Les espèces correspondant à des formes
de l’Amérique septentrionale ou des régions élevées de l’Amérique
tropicale, sont au nombre de 10; celles qui répondent à des formes de
l’Amérique tropicale s’élèvent à 9 …; 3 correspondent à des espèces
du Cap et 2 à des espèces des îles Atlantiques et de la Barbarie; 14
représentent des formes particulières aux Indes ou aux îles de l’Archipel
indien et 30, enfin, correspondent à des formes australiennes. Le groupe
australien est donc le plus considérable, si on les prend isolement. En
les réunnissant, on voit que sur les 80 et quelques espèces, 28 à 30
seulement correspondent à des formes habitant aujourd’hui l’Europe et
l’Amérique du Nord, en y comprenant même les parties méridionales de ces
continents; tandis que 57 au moins, soit 60 en nombre rond, représentent
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 orientales où à l’Australie; de sorte que le caractère dominant de
cette flore est encore Austro-indien, quoique dans une proportion déjà
décroissante par rapport à l’âge 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 studying the great amount of material which had meanwhile
accumulated, Saporta seems 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 southeastern 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 là évidemment le pays qui nous offre le tableau le plus
ressemblant de ce que devait être le midi de la France, et c’est aussi
vers ce même pays, ne l’oublions 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.”[AS]
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 l’Afrique et les Indes orientales. Les
affinités miocènes avec l’Amérique sont postérieures.” These later
American affinities are, however, foreshadowed among the plants and
also, as we have seen, in the Pamphilites of eocene Aix. They appear
again, and very decidedly, 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
considering the Aix butterflies are not, however, in accordance with
those drawn from the insects of the same locality by Professor Heer. He

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

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

    Comme il résulte des recherches de M. G. de Saporta qu’Aix
    appartient à l’étage ligurien, on devrait s’attendre à y
    rencontrer encore plus de formes tropicales qu’à Radoboj. C’est
    tout le contraire, si bien qu’en m’appuyant sur la faune et
    en voyant que Aix avait 10 espèces en commun avec Radoboj et
    4 avec Œningen, j’avais rapporté précédemment les terrains
    d’Aix à la même époque que ceux de Radobo; et je les avais
    rangés dans le Mayencien. Quatre genres ont disparu.… Tous
    les autres genres vivent encore dans la Provence, mais ce
    sont, comme à Œningen, presque tous des genres qui occupent
    une aire géographique très vaste.… On ne peut pas dire que la
    faune des insectes d’Aix contredise positivement l’idée que
    cette localité avait un climat sous-tropical, cas presque tous
    les genres que l’on y a observés jusqu’à présent s’étendent
    jusque dans la zône 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

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 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 Œningen.

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:[AU]—“On retrouve également des types américains, qui
appertiennent à l’Amérique tropicale et sous-tropicale.”

As to the flora of Radoboj, Professor Heer writes in the work just
quoted (p. 96): “Les plantes de la zône tempérée sont représentées
plus fortement qu’à Sotzka,” and of the latter place he says (p.
95), after speaking of types of the temperate zone: “Cependant ces
espèces se trouvent fort à l’arrière-plan en comparison des formes
tropicales et subtropicales, parmi lesquelles prédominent … les formes
indo-australiens; néanmoins 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.

  |        |            |              DEGREE OF AFFINITIES.             |
  |        |NAMES OF    +--------------+------------+----------+---------+
  |        |BUTTERFLIES.|   FIRST      | SECOND     | THIRD    | FOURTH  |
  |        |            |  DEGREE.     | DEGREE.    | DEGREE.  | DEGREE. |
  |        +------------+--------------+------------+----------+---------+
  |        |Neorinopis  |Indo-Malayan. | Austro-    |          |S.       |
  |        |sepulta.    |              | Malayan.   |          |American.|
  |        +------------+--------------+------------+----------+---------+
  | Aix—   |Lethites    |Indo-Malayan. |            |          |North    |
  | Upper  |Reynesii.   |              |            |          |temperate|
  |Eocene. |            |              |            |          |Zone.    |
  |        +------------+--------------+------------+----------+---------+
  |        |Coliates    |Indo-Malayan. |            |Austro-   |         |
  |        |Proserpina. |              |            |Malayan.  |         |
  |        +------------+--------------+------------+----------+---------+
  |        |Thaites     |Mediterranean.|Chinese and Australian,|         |
  |        |Ruminiana.  |              | Subarctic and Alpine. |         |
  |        +------------+--------------+------------+----------+---------+
  |        |Pamphilites |  Tropical    |            |          |Indo-    |
  |        |abdita.     |  America.    |            |          |Malayan. |
  |        |            |              |            |          |         |
  | Rott—  |Thanatites  | Subtropical  | North      |          |         |
  | lower  |vetula.     |   North      | temperate  |          |         |
  |miocene.|            |  America.    | Zone.      |          |         |
  |        |            |              |            |          |         |
  |        |            | Subtropical  | North      |          |         |
  |        |Eugonia     |  temperate   | temperate  |          |         |
  |        |atava.      |  America.    | Zone.      |          |         |
  |        +------------+--------------+------------+----------+---------+
  |Radoboj—|            | Subtropical  | North      |          |         |
  | middle |Pontia      |  temperate   | temperate  |          |         |
  |miocene.|Freyeri.    |  America.    | Zone.      |          |         |
  |        +------------+--------------+------------+----------+---------+
  |        |            |              | Indo-      |          |         |
  |        |Mylothrites |  African.    | Malayan.   |          |         |
  |        |Pluto.      |              | Austro-    |          |         |
  |        |            |              | Malayan.   |          |         |


Nine well authenticated fossil butterflies are now known, all from the
European 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 knowledge, then, places the apparition of
butterflies towards the end of the lower tertiaries.

As a general rule the specimens thus far discovered are in a fair state
of preservation, and especially are those parts preserved which enable
us, with considerable confidence, to determine their exact affinities.
Three of these insects belong to the highest family of butterflies,
Nymphales, four to the Papilionidæ, and two only to the Urbicolæ. If it
be considered probable that the lowest of these families 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 Papilionidæ were moderately endowed. To proceed
further in the analysis 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 Præfecti, a group of gaily attired
butterflies with angulated wings like our common thistle butterfly, the
cosmopolite. Of the four Papilionidæ, 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 destructive 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 Urbicolæ 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 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 Polyommatus 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 district; 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 Cruciferæ, which in its very nature
could scarcely have left a recognizable trace of its presence, the
exception has no force.

After presenting these facts, for convenience sake, in a tabular form, we
will pass on to the enumeration of those fossils which have been referred
to butterflies, but whose exact position is still unsettled.


  |     Names of Species     |   First   |   First    |           |
  |     (and families).      |referred to| described  |   When    |
  |                          |  genus.   |  by        |described. |
  |  (_Nymphales_).          |  Cyllo.   | Boisduval. |    1840   |
  | Neorinopis sepulta Butl. |           |            |           |
  |                          |           |            |           |
  | Lethites Reynesii Scudd. |Satyrites. |  Scudder.  |    1872   |
  |                          |           |            |           |
  |                          |           |            |           |
  |   Eugonia atava Scudd.   |  Sphinx.  |Charpentier.|    1843   |
  |                          |           |            |           |
  |  (_Papilionidæ_).        | Vanessa.  |   Heer.    |    1849   |
  | Mylothrites Pluto Scudd. |           |            |           |
  |Coliates Proserpina Scudd.|    ——     |  Scudder.  |    1875   |
  |                          |           |            |           |
  |                          |           |            |           |
  |  Pontia Freyeri Scudd.   | Pierites. |   Heer.    |    1849   |
  |                          |           |            |           |
  |                          |           |            |           |
  |                          |           |            |           |
  | Thaites Ruminiana Heer.  | Thaites.  |  Scudder.  |    1875   |
  |                          |           |            |           |
  |                          |           |            |           |
  |(_Urbicolæ_).             | Vanessa.  |  Heyden.   |    1859   |
  | Thanatites vetula Scudd. |           |            |           |
  |                          |           |            |           |
  |Pamphilites abdita Scudd. |    ——     |  Scudder.  |    1875   |
  |                          |           |            |           |

  |     Names of Species     |         |  Geological   |  Preserved in   |
  |     (and families).      |Found in |   Horizon.    |   Museum of     |
  |                          |         |               |                 |
  |  (_Nymphales_).          |  Aix.   |   Ligurian,   | Count Saporta,  |
  | Neorinopis sepulta Butl. |         |(upp. Eocene). |   Aix.          |
  |                          |         |   Ligurian,   |                 |
  | Lethites Reynesii Scudd. |  Aix.   |(upp. Eocene). | Marseilles.     |
  |                          |         |               |                 |
  |                          |         |  Mayencian,   |                 |
  |   Eugonia atava Scudd.   |Radoboj. |(mid. Miocene).|        ?        |
  |                          |         |               |                 |
  |  (_Papilionidæ_).        |Radoboj. |  Mayencian,   |Hof mineralien-  |
  | Mylothrites Pluto Scudd. |         |(mid. Miocene).| Kabinet, Vienna.|
  |Coliates Proserpina Scudd.|  Aix.   |   Ligurian,   |Count Saporta,   |
  |                          |         |(upp. Eocene). | Aix.            |
  |                          |         |               |                 |
  |  Pontia Freyeri Scudd.   |Radoboj. |  Mayencian,   |Hof mineralien-  |
  |                          |         |(mid. Miocene).| Kabinet, Vienna.|
  |                          |         |               |                 |
  |                          |         |   Ligurian,   | Professor Heer, |
  | Thaites Ruminiana Heer.  |  Aix.   |(upp. Eocene). |  Zurich.        |
  |                          |         |               |                 |
  |                          |         |               |                 |
  |(_Urbicolæ_).             | Rott.   |  Aquitanian,  |   British       |
  | Thanatites vetula Scudd. |         |(low. Miocene).|    Museum.      |
  |                          |         |               |                 |
  |Pamphilites abdita Scudd. |  Aix.   |   Ligurian,   |   Marseilles.   |
  |                          |         |(upp. Eocene). |                 |

  |     Names of Species     |   Nearest   |  Probable   |Parts of wings |
  |     (and families).      |living allies|  food of    |  preserved.   |
  |                          |found in     |caterpillars.|               |
  |  (_Nymphales_).          | E. Indies.  |  Gramineæ.  |Perfect wings  |
  | Neorinopis sepulta Butl. |             |             | of one side.  |
  |                          |             |             |Both fore-wings|
  | Lethites Reynesii Scudd. | E. Indies.  |  Gramineæ.  |nearly perfect,|
  |                          |             |             | superimposed. |
  |                          |    North    |   Salix,    |Upper half of  |
  |   Eugonia atava Scudd.   |  temperate  |  Populus    |   one         |
  |                          |    Zone.    |  or Betula. |fore-wing.     |
  |  (_Papilionidæ_).        | E. Indies.  | Leguminosæ  |Both fore wings|
  | Mylothrites Pluto Scudd. |             |(Capparis?). |nearly perfect.|
  |Coliates Proserpina Scudd.| E. Indies.  |  Smilax.    |Two fore-wings |
  |                          |             |             | superimposed. |
  |                          |             |             | One fore-wing |
  |  Pontia Freyeri Scudd.   |  Temperate  | Cruciferæ?  |nearly perfect,|
  |                          |  America.   |Terminalia?? | but neuration |
  |                          |             |             |   obscure.    |
  |                          |Mediterranean|             |All the wings; |
  | Thaites Ruminiana Heer.  |  district.  |Aristolochia.|those of one   |
  |                          |             |             | side nearly   |
  |                          |             |             |  perfect.     |
  |                          | Subtropical |Leguminosæ   |   All the     |
  |(_Urbicolæ_).             |     N.      |(Hæmatoxylon | wings, but    |
  | Thanatites vetula Scudd. |  America.   |Gleditschia).| superimposed  |
  |                          |             |             |  and very     |
  |                          |             |             |  obscure.     |
  |Pamphilites abdita Scudd. |  Tropical   |  Gramineæ.  | One fore-wing |
  |                          |  America.   |             |   perfect.    |

In the earliest accounts that we have found, including all those in the
last century, the generic term Papilio was used for all Lepidoptera, and
therefore we cannot be certain whether butterflies or moths are meant.
Hueber’s plates, even, are so inferior that they afford no additional
aid; but those of Sendel possibly represent, as we have noticed in the
Bibliography at the commencement of this memoir, the early stages of
butterflies preserved in amber. The only other direct references to
butterflies preserved in amber are the following: Gravenhorst,[AV] in
his enumeration of amber insects, gives under the Lepidoptera forty
specimens referable to Tineæ and Tortrices, and besides these “mehre
Raupen, sämmtlich, wie es scheint, Schildraupen, denen des Papilio W.
album ähnlich.” The probable nature of the ancient forest yielding amber
renders it unlikely that any butterflies in their perfect state would
be found in it. As a rule, butterflies are eminently fond of the light.
This has already been remarked by Menge:[AW]—“Das fehlen gröszerer
Schmetterlinge im bernstein deutet auf einen finstern undurchdringlichen
urwald, den die kinder des lichts gemieden haben.” Yet as some Theclas
do feed upon coniferous trees, it is not impossible that the onisciform
larvæ, referred to by Gravenhorst, may belong to this group. As far as we
can discover, no further reference is made to them, excepting by Giebel
and Bronn in some of their lists and enumerations of fossil insects.
The writings of Berendt, Menge and others, all bear testimony to the
great rarity of Lepidoptera in amber, and most of those which have been
discovered belong to the lowest two families, above referred to.

Dr. Hagen informs me that he has himself seen specimens of large
butterflies in amber, but that these proved to be falsifications,
recent European insects like _Pieris rapæ_, etc., having been enclosed
between slabs of amber, which were then fastened together and the edges
roughened, all in so clever a manner that one would not suspect them to
be spurious. These specimens were manufactured many years ago, and it is
not impossible that it is to one of them that Hope refers in 1836, as
found in the collection of Mr. Strong, though why he should quote Berendt
as authority I cannot discover.

Heer, in the introduction to the lepidopterous portion of his
“Insektenfauna der Tertiärgebilde von Œningen,” says (p. 175): “Karg
erwähnt zwar eines sehr schönen Œninger-Schmetterlings, der nach Zürich
gekommen sein soll. Allein hier findet sich dieser nicht und die Angabe
verliert noch mehr an Werth, wenn wir berücksichtigen, dass Karg das
Thier nicht selbst gesehen hat.” Karg’s memoir in the “Denkschriften der
Schwäbischen Gesellschaft der Aerzte und Naturforscher,” T. I., I have
been unable to examine.

Boisduval, in his final report upon _Neorinopis sepulta_, remarks that
Count Saporta had written him that many years previously he had sent to
the Paris Museum a “Polyommate fossile” from Aix. Count G. de Saporta,
in reply to my inquiries concerning this specimen, says that his father
can give me no further information concerning this specimen; nor could
M. Oustalet and myself, in our search through the fossil insects of the
Jardin des Plantes, discover any such relic.

In a recent number of “Nature” (No. 266), Mr. E. J. A’Court Smith writes
of the discovery at Gurnet Bay in the Isle of Wight, of an insect bed in
which were found, among other things, “a variety of flies, butterflies,
and one or two grasshoppers;” no further information has yet been
published concerning these relics, and my inquiries upon the subject have
not, as yet, elicited any definite response.


1. _Cyllonium Boisduvalianum_ WESTW., and _C. Hewitsonianum_ WESTW.

[Illustration: Fig. 2.

Cyllonium Boisduvalianum Westw.]

These two insects were figured by Westwood in the Quarterly Journal
of the Geological Society of London for November, 1854, the former
(reproduced in our fig. 2) on Pl. XVII, fig. 17; the latter (reproduced
in our fig. 3) on Pl. XVIII, fig. 27. Of the former he makes the
following remarks:[AX] “Pl. XVII, fig. 17 represents a number of
fragments of delicate tegument, covered with minute punctures and
traversed by straight and somewhat radiating veins, which appear like
portions of the hind wing of some species of Butterfly, entirely denuded
of scales.” The name is given to it in a note to the explanation of the
plates, p. 395. Concerning the second he says:[AY] “Pl. XVIII, figs.
27 and 30, appear to be portions of the hind wings of some species
of Butterfly; still they have very much of a vegetable aspect. The
surface is covered with minute punctures, which may be the cells for the
insertion of the quills of the coloured scales, which are all removed,
supposing the specimens to be Lepidopterous.” The name we have quoted is
given only to fig. 27, in a foot note on p. 396.

[Illustration: Fig. 3.

Cyllonium Hewitsonianum Westw.]

I have not been able to find, even with Mr. Brodie’s help, the first
specimen referred to; but an examination of the original of the latter
(see fig. 3) proved that, while it is unquestionably an insect, it cannot
be referred to the Lepidoptera; the punctures referred to are both too
large and much too irregularly disposed to have been the points of
insertion of the scales; they are probably the marks of the insertion
of hairs, such as are not uncommonly seen irregularly scattered over
the wings of insects belonging to the other suborders. As the figure
of the first species closely resembles in this particular the one I
have seen, I am forced to the conclusion that neither of these wings
are lepidopterous. Plainly, the only reason why a new generic name was
appended to these forms was that their remains were too fragmentary
to afford the slightest guess as to what modern genus they might be
referred. The fossils came from the English Purbecks.

2. _Palæontina oolitica_ BUTL.

The first notice I find of this remarkable and very interesting fossil
is that published in various literary and scientific London journals
reporting remarks given at a meeting of the Entomological Society of
London, and which afterward appeared as follows in their Proceedings:[AZ]

“Mr. Butler exhibited a remarkably perfect impression of the wing of a
fossil butterfly in the Stonesfield slate. It appeared to be most nearly
allied to the now existing South American genus Caligo.”

[Illustration: Fig. 4.

Palæontina oolitica Butl. The neuration, after Butler’s first sketch.]

A full description of this insect soon appeared in the author’s
“Lepidoptera Exotica,” accompanied by a plate; both were afterward
republished in the “Geological Magazine.” In fig. 4 we reproduce fig. 1
of his plates, representing the neuration of Palæontina; and in fig. 5,
fig. 2 of his plates, subsequently copied by “The Graphic.” A description
of the genus and species is first given, which it is unnecessary to
reproduce here; afterward, the following remarks:

“[126] Though a British insect, this species belongs to a group so
completely tropical that I do not hesitate to describe and figure it in
the present work; its nearest allies are the genera Caligo, Dasyophthalma
and Brassolis, all three essentially tropical American genera.

[Illustration: Fig. 5.

Palæontina oolitica Butl. Facsimile of Butler’s first sketch.]

“_P. oolitica_ is especially interesting, as being the oldest fossil
butterfly yet discovered; the most ancient previously known to
science having been found in the Cretaceous series (white sandstone of
Aix-la-Chapelle[BA]), whilst the bulk of the known species are from the
Lower Miocene beds of Croatia; it is also interesting as belonging to the
highest family of butterflies, and to a subfamily intermediate in [127]
character between two others, namely, the Satyrinæ and Nymphalinæ, whilst
the more recently discovered fossils are referable, with one exception,
to the two latter groups. The nervures appear to have been impregnated
with iron, which will partly account for their well-defined condition.”

Happening to be in London not long after the publication of the
description and illustration of this insect, I took pains to make a very
careful examination both of the original specimen, which Mr. Charlesworth
kindly allowed me to study at my leisure, and of its reverse, which is
preserved in the School of Mines, Jermyn street. I mentioned to Mr.
Butler and to others, my conviction that the insect was to be considered
homopterous rather than lepidopterous, and on my return to America,
exhibited before the Natural History Society of Boston, drawings which I
had made from the originals; my comments at that time were published very
briefly, as I was reserving the proof of my statements for the present
paper. Mr. Butler, however, was induced by this publication[BB] to
examine the reverse at the Jermyn street Museum, and although he had been
supplied by me with a rough tracing of the drawing I had taken of it, he
failed to be convinced of any mistake, and published a paper in defence
of his own view in the Geological Magazine for October, 1874. In this
paper he gives new drawings of the insect, quotes portions of letters in
which I had expressed my opinions upon the nature of the fossil, gives
the remarks referred to from the “Proceedings of the Boston Society of
Natural History,” and makes, among others, the following comments.

[Illustration: Fig. 6.

Palæontina oolitica Butl. Facsimile of Butler’s second sketch.]

“Seeing that Mr. Scudder had made his views public, I felt that it was
time for me to take similar steps on my side. I therefore availed myself
of an early opportunity of again visiting Jermyn street, where, through
the courtesy of the officers, I was enabled to make a sketch of the
impression in the Museum. I found it impossible to make a tracing of it,
and therefore drew the whole by measurement. This sketch is now produced
on Pl. XIX, fig. 4 [see fig. 6]; and any body can judge for himself
whether or not it is more perfect than that which I previously figured
(see Geol. Mag., 1873, Vol. X, p. 2, Pl. I, fig. 2 [see fig. 5]).”

“In order to show the extent to which the Jermyn street example is
deficient, I have restored it (fig. 5 [our fig. 7]), filling in the
blanks from Mr. Charlesworth’s specimens. By comparing the latter with
the wing of _Dasyophthalma_ (fig. 1), and _Cicada_ (fig. 2), one may come
to a pretty accurate conclusion as to the group of insects to which it
ought to be referred.”[BC]

The neuration of Lepidoptera as a group is the simplest in the whole
order of insects, if we except that of the elytra of Coleoptera; this
is due, doubtless, to the fact that their wings are heavily scaled,
concealing the nervures; just as in Coleoptera, the thickness and opacity
of the fore wings often completely masks the neuration.

[Illustration: Fig. 7.

Palæontina oolitica Butl. The neuration, after Butler’s second sketch.]

The normal number of veins in the wings of insects is six, disposed to
a certain extent in pairs; the middle pair usually ramify to a greater
extent than the others, and support most of the membrane of the wing.
In butterflies the foremost vein is always absent and very commonly the
hindmost, so that there are but five (often but four) principal veins,
usually designated, though not very appropriately: costal, subcostal,
median, submedian and (when present) internal, reciting them in their
order from in front backward. The costal, submedian and internal nervures
are invariably simple and terminate at the margin, or are occasionally
lost in the membrane of the wing. The subcostal and median nervures,
on the other hand, are as invariably forked, and with their branches
support nearly the entire wing; the subcostal nervure curves downward
and the median upward so as to meet, or nearly to meet, not far from the
middle of the wing, and to enclose between them a large space called
the discoidal cell; the branches of the median nervure are all thrown
off from its lower edge before union with the subcostal; the principal
branches of the subcostal nervure are, on their side, thrown off from
its upper edge; but, as the nervure curves downward at the extremity of
the cell, another set is thrown off (at least in the fore wings) from
the lower edge; and it is these veins, rather than the subcostal nervure
proper, which unite with the median to close the cell.[BD] None of the
median, nor any of the inferior subcostal nervules are ever branched;
but at the apex of the wing, where the play of neuration is usually the
greatest, the last superior subcostal nervule is occasionally forked in
the front wing. This is the only forked branchlet in either of the wings.

The last figure of _P. oolitica_ given by Mr. Butler agrees in all its
essential features with his first illustration. They both represent
a front wing with four principal nervures,—costal, subcostal, median
and submedian; the costal nervure is swollen at the base and extends,
unbranched, to the tip of the wing; the median nervure is three-branched,
the three forks simple, equidistant, emitted from the apical half of
the vein, which at its extremity is united by a cross vein to a branch
of the subcostal, closing the cell; the submedian nervure is simple and
divides the space between the median vein and the margin of the wing.
So far all is in accordance with the lepidopterous type; but when we
examine the subcostal vein, which occupies nearly half the wing, the
resemblance ceases altogether. This vein is represented as bearing no
superior branches, but as sending out from its _inferior_ surface three
distinct veinlets, the first and second of which again emit a tributary
from their inferior surfaces. This is a structural anomaly which finds
no counterpart whatsoever in any family of butterflies. So that should
we accept Mr. Butler’s own sketch of the fossil as correct, it would be
impossible to consider the wing that of a butterfly.

In his description of the insect Mr. Butler compares the neuration to
that of Caligo, and says its nearest allies are Caligo, Dasyophthalma and
Brassolis. In his latter paper he figures the wing of a Dasyophthalma by
way of comparison. In the genera named all the branches of the subcostal
nervure are simple, and are thrown off from the superior surface,
excepting the single set which is emitted from beneath, and which marks
(as in all butterflies) the limit of the discoidal cell; this corresponds
fairly with the first set of inferior veins emitted by the subcostal vein
in the fossil; for the other sets, however, no counterpart will be found
in the living types.

[Illustration: Fig. 8.

Palæontina oolitica Butl. Corrected sketch of the neuration.]

It was probably Mr. Butler’s want of familiarity with fossils that led
him to overlook several features which can be seen in these originals.
Having first traced the outline of the wing and the general course of
the veins directly from the specimens, I subsequently filled in by
measurement all the other parts which I could follow, studying each vein,
or supposed vein, with the utmost care, from one end to the other of its
course. The result of that study is presented in fig. 8, which differs
essentially in its details from the illustrations given by Butler, and
looks, as he himself confesses, “exceedingly anti-lepidopterous.” In the
first place, the wing is much narrower than depicted by him; and at the
extremity of a vein (the submedian vein of Butler’s sketch) there is a
slight but decided bending inward of the membrane, as very frequently
occurs at the line of demarcation between the middle and inner area
of the wing in all or nearly all the lower suborders of insects, but
never, so far as I am aware, in Lepidoptera. What he has given as a
simple costal vein is neither swollen at the base nor simple, but has
two inferior branches near the middle of the wing, united near their
origin by an oblique cross vein. Branching of the costal vein is unknown
in Lepidoptera; but if it should be claimed that this might be the
subcostal, just as much difficulty will be encountered with the structure
and relationship of the veinlets below, which must then be considered
as belonging to the median vein; in no Lepidoptera can any such
irregularity be shown, nor so disproportionate a magnitude of the area
covered by the median nervure and its branches; a branched internal vein
and cross-veins, which probably united all the longitudinal nervures at
no great distance from the outer border (but which can only be certainly
predicated for the lower three median interspaces), place this insect
wholly beyond the pale of the Lepidoptera. It is but fair to say that
Mr. Butler, having examined the original after he had in his possession
a tracing of fig. 8, denies the existence of the cross-veins; there is
one point, however, which an unprejudiced examination of the fossil
cannot fail to show; that Butler’s “fourth branch” of the subcostal[BE]
arises not from his third branch, but from his upper discoidal vein;
if he can reconcile either this or the points already referred to (on
the supposition that his sketch is otherwise an accurate one) with the
neuration of any group of butterflies, the writer will be the first to
acknowledge it.

As our only purpose in this place is to deny the lepidopterous character
of Palæontina, it is unnecessary to say anything in defence of the view
we have expressed of its homopterous affinities; the superior position
of the cell, the position and character of the lower cross veins (which
we believe really traversed the entire wing), with their origin at the
indentation of the lower border, suggest such a relationship, although
there are not a few points in which it differs somewhat strikingly from
living types.

The discovery of a fossil in the cabinet of the Rev. Mr. Brodie, which
was found in England at the same or nearly the same horizon, as _P.
oolitica_, and which seems to be a pupa case of one of the Cicadida of
rather unusual size, renders my suggestion more worthy of credence.

At the conclusion of his latter paper Mr. Butler draws attention to
the fact that Messrs. Westwood and Bates had expressed their agreement
with his views. It should, however, be borne in mind, that, so far as
appears from any facts which have been published, these gentlemen, whose
well considered views upon the subject would unquestionably be of great
weight, expressed this assent only upon a brief evening examination of
a very obscure fossil in a poorly lighted hall, and before any one had
questioned its lepidopterous character.


[A] Probably an error for Tab. 5, fig. 28a, 28b, which seems to represent
a Tenthredinidous larva.

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

[C] Bull. Soc. Ent. France, 1838, 51-52.

[D] Bull. Soc. Ent. France, 1839, 11-12.

[E] Annales Soc. Ent. France, ix, 371-374 pl. 8.

[F] By Marcel de Serres, Actes Linn. Soc. Bord., Vol. xiii, pl. 2;
Pictet, Traite de Palæont., II pl. 40, fig. 11; Butler, Lep. Exot. I, pl.
18, fig. 2; Ib., Geol. Mag. x., pl. 1, fig. 3.

[G] See particularly papers by Coquand, Marcel de Serres, Heer, Saporta,
Giebel, Westwood and Butler.

[H] Ann. Soc. Ent. France (2) ix, 71-88, pl. 3, ii.

[I] Bull. Ent. Soc., France, 1851. 97-8.

[J] Cat. Satyr., 189-190.

[K] Species of Lethe.

[L] As, for instance, in the _Sphingidæ_, _Heliconidæ_, etc.

[M] Lep. Exot. pl. 48.

[N] Loc. cit., p. 190.

[O] In this figure the neuration has not been very clearly defined, the
veining of the hind wings not being continuous.

[P] Lep. Exot., 127, pl. xlviii. Geol. Mag., x, 3, pl. i.

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

[R] Acta Acad. Leop.-Carol., XX, 108.

[S] Insekt. Tert. Œning. ii, 177-79.

[T] Wahrscheinlich ist ausser diesem noch ein Ast da, der aber verwischt

[U] Lep. Exot. I. 128.

[V] Compare, in this respect, Butler’s Revision of the Pierinæ, Cist.
Ent., I, iii, pl. i, fig. 8; or Trimen, Rhop. Afr. Austr., Pl. ii, fig. 2.

[W] Insekt. Tert. Œning., II, 179-82.

[X] Butt. N. Amer., i, Argynnis, I.

[Y] Cat. Fabr. Lep., 109.

[Z] Lep. Exot., I, xv, 127-28, Pl. 48, fig. 7; Geol. Mag., x, 3-4, Pl. 1,
fig. 7.

[AA] Insekt. Tert. Œning, II, 180.

[AB] Compare the illustrations referred to in the note on page 44.

[AC] These, it is true, are Danai, but aberrant forms, like Leptidia, etc.

[AD] See, however, the American genus Leodonta.

[AE] Insekt. Tert. Œning., ii, 182-3.

[AF] Palæontogr. viii, 12-13.

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

[AH] “La proportion des Graminées relativement au total des Phanérogames,
qui est de 4·5 sur 100, est en rapport avec les minima relatifs de
cette famille, tels qu’on les observe à la Nouvelle-Guinée et à la
Nouvelle-Grenade.” Saporta, loc. cit., 292.

[AI] “Dans la flore des gypses d’Aix, non-seulement les Légumineuses
occupent le premier rang, comme dans la plupart des flores
intertropicales actuelles de l’ancien et du nouveau continent, mais
elles atteignent une proportion de 13 pour 100, pour l’ensemble des
Phanérogames, proportion parfaitement en rapport avec celle de 12 sur 100
qui est fréquente, selon M. de Candolle, dans certaines régions chaudes,
telles que Timor, le Congo, etc.” Saporta, loc. cit., 292.

[AJ] Since this was written, Count Saporta writes me: “Le genre Smilax
est un des genres tertiatres les plus frequents. J’ai aussi signalé dans
le dépôt voisin de St. Zacharie (étage Tongrien inférieure [and therefore
but slightly more recent]) une feuille qui m’a paru devoir se ranger
parmi les Dioscorées.”

[AK] An old writer in Fuessly’s Magazin, writing from Italy, says that
Thais feeds in that country upon Quercus. Five species of Quercus are
known from Aix, but the statement in Fuessly’s Magazin has never, to my
knowledge, been confirmed.

[AL] “Nous n’avons pas encore rencontré d’Aristolochia dans les gypses
d’Aix; mais l’existence du genre dans le tertiaire moyen ne saurait être
mise en question, depuis que nous avons entre les mains une superbe
empreinte de Radoboj (Aristolochia venusta Sap.), qui dénote une forme
voisine des Aristoloches à feuilles persistantes et demi-coriaces, comme
l’_A. reticulata_ Nutt. de Virginie.” Saporta, loc. cit., 342-3.

[AM] In a recent letter from Count Saporta he remarks: “Relativement au
_Thaites Ruminiana_, je n’ai pas encore découvert à Aix de vestiges du
genre Aristolochia, mais _ce genre devait y exister_.”

[AN] Vierteljahrsschrift naturf. Gesellsch. Zurich, i. 12, 1856.

[AO] Recherches sur le Climat et la Végétation du pays tertiaire, 4to,

[AP] I venture to give one more extract from a recent letter received
from Count Saporta, although he writes:—“Je vous écris n’ayant sous les
yeux ni mes livres ni mes collections, ce qui enlevera nécessairement un
peu de précision à quelques-unes de mes réponses.”

“Il est bien plus difficile de justifier par des exemples tirés de la
nature des plantes la présence à Radoboj d’un insecte se nourissant de
Capparis. Les Capparis ont du exister, mais leur feuilles sont difficiles
à distinguer à cause de l’absence de caractères différentiels; leur forme
et leur nervation peu visibles doivent les faire confondre avec beaucoup
d’autres. Il me semble pourtant que des Capparis ont été signalés soit à
Radoboj, soit à Hoering en Tyrol, dépôt un peu plus anciens [Tongrian],
mais en l’absence de mes livres je ne saurais vous l’affirmer.”

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

[AR] Heer et Gaudin, Climat du pays tertiaire, pp. 133-171.

[AS] Ann. Sc. Nat., [5] Bot., xv, 322.

[AT] Climat du pays tertiaire, ed. Gaudin, p. 205.

[AU] Loc. cit., p. 205.

[AV] Arbeit Schlesisch. Gesellsch. Vaterl. Kultur, 1834, 92-3.

[AW] Programm Petrischule Danzig, 1855-56, 4to, p. 30.

[AX] Loc. cit., 387.

[AY] Loc. cit., 390.

[AZ] 1872, xxxi.

[BA] Perhaps Mr. Butler is not altogether to blame in confounding Aix
in Provence with Aix-la-Chapelle; at any rate the mistake had been made
previously by the translator of Heer’s paper in the Quarterly Journal of
the Geological Society of London, VI, 72; the error is corrected by Mr.
Butler at the end of his volume.

[BB] He seems not to have seen the earlier publication of Mr. Brodie.

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

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

[BE] In this case he counts from the tip of the wing, in reverse order.


[My best thanks are due to my courteous friend Mr. Auguste Sallé, for
his kind agency in securing an artist for the engraving of these plates.
Owing, however, to the distance at which the work was done, a few errors
have unavoidably occurred, which, to prevent misapprehension, are
mentioned below.]

Plate I.

  Fig.  1. _Eugonia atava._ Copied from Heer, Insekt. Tert. Œning., ii,
            pl. xiv, fig. 8 (1/1).

   “    2. _Lethites Reynesii._ Drawn by S. H. Scudder (1/1).

   “    3. _Eugonia atava._ Copied from Charpentier, Nov. Act.
            Leop.-Carol., xx, pl. xxii, fig. 4 (1/1).

   “    4. _Eugonia j.-album._ Neuration of fore wing; drawn by S. H.
            Scudder (1/1). The second superior subcostal nervule is
            carried too far toward the tip of the wing.

   “    5. _Lethites Reynesii._ Fore wing; drawn by S. H. Scudder (2/1).

   “    6. _Eugonia j.-album._ Markings of the upper surface of the fore
            wing; drawn by S. H. Scudder (1/1).

   “    7. _Eugonia atava._ Neuration of tip of fore wing (2/1); copied
            from Heer; Insekt. Tert. Œning., ii, pl. xiv, fig. 3a.

   “    8. _Neorinopis sepulta._ Markings of the upper surface of the
            two wings, restored; drawn by S. H. Scudder (2/1). The drawing
            represents the general effect of the fore wing as darker than
            the hind wing, and in so far is inaccurate.

   “   9. _The same._ Neuration of the two wings, separated; drawn by
           S. H. Scudder (2/1).

   “  10. _The same._ Neuration of the two wings, as seen in the fossil;
           drawn by S. H. Scudder (2/1). The engraver has unfortunately
           made the lines of the hind wing the heavier, as if it lay
           uppermost; they should have been the lighter.

   “  11. _The same._ Right hind leg; drawn by S. H. Scudder (2/1).

   “  12. _The same._ Left hind leg; drawn by S. H. Scudder (2/1).

   “  13. _The same._ Drawn by S. H. Scudder (1/1). The spot of the
           medio-submedian interspace of the fore wings has not been well
           rendered by the engraver.

   “  14. _The same._ Copied from Lefebvre, Ann. Soc. Ent. France [2], ix,
           pl. iii, II, fig. A (1/1).

   “  15. _The same._ Copied from the same, fig. C (1/1).

   “  16. _The same._ Copied from the same, fig. B (1/1).

   “  17. _The same._ Copied from Boisduval, Ann. Soc. Ent. Fr., ix, pl.
           8 (1/1).

Plate II.

  Fig.  1. _Zophoessa Sura._ Neuration of the wings; drawn by G. Willis

   “    2. _Mylothrites Pluto._ Copied from Heer, Insekt. Tert. Œning.,
            ii, pl. xiv, fig. 4 (1/1).

   “    3. _Zophoessa Sura._ Markings of the upper surface of the wings;
            drawn by G. Willis (1/1).

   “    4. _Delias Pasithoe._ Neuration of the wings; drawn by G. Willis

   “    5. _Coliates Proserpina._ Neuration and markings of fore wings;
            drawn by S. H. Scudder (2/1). The tip of the costal nervure
            has been extended too far toward the apex of the wing.

   “    6. _Lethe Dyrta._ Neuration of the wings; drawn by G. Willis

   “    7. _Mylothrites Pluto._ Neuration of the wings; after a drawing
            obtained through Mr. Brunner de Wattenwyl (1/1). The second
            superior subcostal nervule on the left wing should join the
            nervure midway between the bases of the first and third

   “    8. _Neorina Lowi._ Neuration of the wings; drawn by G. Willis

   “    9. _Lethe Dyrta._ Markings of the lower surface of the fore
            wing; drawn by G. Willis (1/1).

   “   10. _Debis Sinorix._ Neuration of the wings; drawn by G. Willis

   “   11. _Zophoessa Sura._ Markings of the lower surface of the fore
            wing; drawn by G. Willis (1/1).

   “   12. _Pontia Protodice._ Neuration and markings of the upper surface
            of fore wing; drawn by S. H. Scudder (1/1).

   “   13. _Neorina Lowi._ Markings of the upper surface of the wings;
            drawn by G. Willis (1/1). This was drawn for the pattern of
            markings only; the neuration is faulty.

   “   14. _Debis Sinorix._ Markings of the upper surface of the wings;
            drawn by G. Willis (1/1).

   “   15. _Mylothrites?_ sp. Copied from Heer, Insekt. Tert. Œning., ii,
            pl. xiv, fig. 5 (1/1).

   “   16. _Pontia Freyeri._ Copied from Heer, Insekt. Tert. Œning., ii,
            pl. xiv, fig. 6 (1/1).

   “   17. _Mylothrites Pluto._ After a drawing from the original,
            furnished by Mr. Brunner de Wattenwyl (1/1).

   “   18. _Pontia Freyeri._ Drawn from the original under the direction
            of Mr. Brunner de Wattenwyl (1/1).

Plate III.

  Fig. 1. _Thaites Ruminiana._ Neuration of the wings, restored; drawn
           by S. H. Scudder (2/1).

   “   2. _Thais Rumina._ Neuration of the wings; drawn by S. H.
           Scudder (1/1).

   “   3. _Thaites Ruminiana._ Markings of the upper surface of the
           wings, restored; drawn by S. H. Scudder (2/1).

   “   4. _Thais Rumina._ Markings of the upper surface of the wings;
           drawn by S. H. Scudder (1/1).

   “   5. _Parnassius Smintheus._ Markings of the upper surface and
           neuration of the wings; drawn by S. H. Scudder (1/1).

   “   6. _Thaites Ruminiana._ One of the wing-covers (patagia); drawn by
           S. H. Scudder (12/1).

   “   7. _The same._ Portion of the palpi; from a camera sketch by S. H.
           Scudder (12/1).

   “   8. _The same._ Antenna; drawn by S. H. Scudder (3/1).

   “   9. _The same._ From a camera sketch by S. H. Scudder (3/1).

   “  10. _The same._ Drawn under the camera from the reverse of fig. 9,
           by S. H. Scudder (3/1).

   “  11. _Thanaos Juvenalis._ Drawn in the position of fig. 12 by J. H.
           Emerton (1/1); fig. 11 a, the palpus, denuded (2/1).

   “  12. _Thanaos 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 (+2/1). Incorrectly named _Thanatites Juvenalis_
           on the plate.

   “  13. _Carystus Lucasii._ Neuration of fore wing. Drawn by G. Willis

   “  14. _Pamphilites abdita._ Markings of the upper surface of the fore
           wing; drawn by S. H. Scudder (2/1).

   “  15. _Pansydia Mesogramma._ Neuration and disposition of spots on
           the fore wing; copied from Poey, Cent. Lep. Cuba, 2e Dec.

   “  16. _Thanatites vetula._ Copied from Heyden, Palæontogr., viii,
           pl. i, fig. 10 (+2/1). Incorrectly named Thanatites Juvenalis
           on the plate.

   “  17. _Pamphilites abdita._ Markings of the upper surface of fore
           wing; drawn by S. H. Scudder (1/1).

   “  18. _The same._ Neuration and disposition of the spots on the fore
           wing; drawn by S. H. Scudder (2/1).

   “  19. _Carystus Lucasii._ Markings of the upper surface of fore wing;
           drawn by G. Willis (1/1).


  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.

   “   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). _Palæontina oolitica._ Copied by photography from
                   Butler, Lep. Exot., pl. xlviii, fig. 1; engraved
                   by H. Marsh.

   “   5 (p. 90). _The same._ Copied by photography from the same, fig.
                   2; engraved by H. Marsh.

   “   6 (p. 92). _The same._ 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.


  Page 19, line 4; _for_ voicé, _read_ voici.

   “   29. The first three lines form a part of the quotation from Butler,
              and should have been but single-leaded.

   “   51, note; _for_ Dareai, _read_ Danai.

   “   58, line 9; _for_ before the cell, _read_ before the tip of
              the cell.

   “   62, line 13; _for_ fig. 2, _read_ fig. 11.

Transcriber’s Note: These errata have been corrected in this e-text.
“Page 51, note” is footnote AC.

Additionally the following changes were made:

  Page 16, le sujet le ce rapport => le sujet de ce rapport

  Page 20, marveilleusement => merveilleusement

  Page 25, de cette taile => de cette taille

  Page 26, non estimable ami => mon estimable ami

  Page 37, Satyrites Beynesii => Satyrites Reynesii

  Page 41, Zeichung => Zeichnung (twice)

  Page 42, Färburg => Färbung (twice, and another time on page 47)

   “   42, bie den Vanessen => bei den Vanessen

  Page 46, dis zum Augenpunkt => bis zum Augenpunkt

   “   46, gezacht => gezackt

   “   46, indem ganz schwache => in der ganz schwache

  Page 47, Varästelung => Verästelung

   “   47, enimal => einmal

  Page 81, précédement => précédemment

   “   81, q’u à => qu’à

   “   81, Quatres genres => Quatre genres

   “   81, note (footnote AT), du pays tertiare => du pays tertiaire

   “   88, Aertze => Aerzte

Minor errors in punctuation, capitalisation, accents etc have been
corrected without note.

[Illustration: Plate I.

  1, 3, 7 Eugonia, atava.
  4, 6 Eugonia. J-album.
  2, 5 Lethites Reynesii.
  8-17 Neorinopis sepulta.

_Imp. Ch. Chardon ainé, Paris._]

[Illustration: Plate II.

  1, 3, 11 Zophoessa Sura.
  2, 7, 17 Mylothrites Pluto.
  4. Delias Pasithoe.
  5. Coliates Proserpina.
  6, 9 Lethe Dyrta.
  8, 13 Neorina Lowii.
  10, 14 Debis Sinorix.
  12 Pontia Protodice.
  15 Mylothrites ?
  16, 18 Pontia Freyeri.

_Imp. Ch. Chardon ainé, Paris._]

[Illustration: Plate III.

  1, 3, 6-10 Thaites Ruminiana.
  2, 4 Thais Rumina.
  5 Parnassius Smintheus.
  11 Thanaos Juvenalis.
  12, 16 Thanatites Juvenalis.
  13, 19 Carystus, Lucasii.
  14, 17, 18 Pamphilites abdita.
  15 Pansydia Mesogramma.

_Imp. Ch. Chardon ainé, Paris._]

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