WILLIAM D. FIE Butterflies of the

Genus Vanessa and of

• the Resurrected Genera

. Bassaris and Cynthia

, (Lepidoptera:

Nymphalidae)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 84

William D. Field Butterflies of the

Genus Vanessa and of

the Resurrected Genera

Bassaris and Cynthia

(Lepidoptera:

Nymphalidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Field, William D. Butterflies of the Genus Vanessa and of the Resurrected Genera Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smithsonian Contributions to Zoology, number 84, 105 pages, 1971.—The genus Cynthia with nine

species (including C. annabella, new species) and the genus Bassaris with two

species are recognized as separate from the genus Vanessa, with five species remaining. All taxa are keyed and redefined with characters, including the male

and female genitalia, tarsi, and wing habitus. Knowledge of the distribution and

biology of most species is extensive and is given herein (for distribution) or cited

(known food plants are listed and references to the literature of the biology are

given). Unusual modified setae on the ovipositors are described for five Cynthia

species.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

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For sale by the Superintendent of Documents, U.S. Government Printing Office

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William D. Field Butterflies of the

Genus Vanessa and of

the Resurrected Genera

Bassaris and Cynthia

(Lepidoptera:

Nymphalidae)

Introduction

In this paper the genera Cynthia and Bassaris are

resurrected and removed from the genus Vanessa

and all three genera are redefined on the basis of

characters in male and female genitalia, tarsi, and

habitus.

These butterflies collectively are found nearly all

over the world. The genus Vanessa contains five

species and is mainly, and was perhaps originally,

an Old World genus. One Vanessa species is, however, Holarctic; another is distributed over a great

part of Asia and even occurs in southeast Europe,

Madeira, and the Canary Islands; and the three

remaining species are found in limited island areas

of the Old World. The genus Cynthia contains nine

species and is mainly New World except for one

nearly cosmopolitan species and for one which

occurs in Australia. The genus Bassaris, on the

other hand, is confined to Australia and New

Zealand and a few islands in the southern Pacific

region.

There is a considerable amount of available information concerning the immature stages and the

life histories of the genera treated in this paper.

William D. Field, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

The eggs of all species have not been described,

but from those that have been we can say that they

are barrel shaped, broad at the base, short, and

ovate with prominent laminate ribs that increase

in height to the summit, where they end abruptly.

In Cynthia eggs, there are from fourteen to nineteen

of these ribs, in Bassaris there are eight (B. itea)

or nine (B. gonerilla), and in Vanessa there are

nine.

The eggs are laid singly on the underside of the

leaves of the food plant in Bassaris and usually on

the upper side of the leaves in Vanessa and Cynthia.

In four species of Cynthia (C. virginiensis, C.

altissima, C. braziliensis, and C. myrinna) the ovipositors contain a number of greatly modified,

cephalically directed, teethlike setae (Figures 26-

30). These modified setae (shown enlarged in Figure 26a) may be an aid in stabilizing or anchoring

the ovipositor as it is worked carefully through the

thickly matted hairs on the surface of the leaf of

the host plant prior to the laying of the egg under

this carpet of hairs. Scudder (1889, p. 464) describes

this peculiarity of egg laying for C. virginiensis, but

apparently was not aware of structures that could

aid in this careful procedure.

No such attempt at egg concealment has been

described for species of Vanessa or Bassaris, nor for

other Cynthia species; I would, however, expect C.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

altissima, C. braziliensis, and C. myrinna to act

similarly to C. virginiensis.

The ovipositor setae of Cynthia kershawi (Figure

25) are slightly modified in having their apical

portions bent at nearly right angles and with these

bent portions cephalically directed. The purpose of

this modification is not known, and nothing peculiar about its egg laying has been reported.

The caterpillars of Bassaris and Vanessa species

feed almost exclusively upon the Urticaceae, while

those of Cynthia prefer the Compositae and Malvaceae.

As far as is known, the caterpillars of all three

genera are solitary, living usually in vertical nests

made of leaves tied together with silken strands. In

Vanessa and one species of Bassaris (B. itea) the

nests are made of a single leaf, and feeding takes

place in this nest until the leaf is completely devoured. In Cynthia and the other species of Bassaris

(B. gonerilla), the caterpillars construct nests composed of several leaves and in Cynthia at least they

feed only on the upper surface of these leaves.

The most troublesome taxonomic problem that

a worker faces in revisionary work on many popular

groups of butterflies is the question of how to deal

with the vast horde of infrasubspecific names.

Because this was a major problem in the present

study, I wish to discuss it further in order to explain

the various decisions which were adopted.

Over the past fifty years there has been a considerable amount of controversy among taxonomists,

particularly lepidopterists, as to the value of giving

names to various types of taxons below the rank of

subspecies. Thousands of such names have been

given to "aberrations," "seasonal forms," "color

forms," "sexual forms," and the like. For example,

forty-seven names have been proposed for forms and

aberrations of Vanessa atalanta, and Cynthia cardui

has received nearly the same number of such names.

The record for this kind of name proposing is

probably held by Lysandra coridon (Poda) (Lycaenidae), which has received some seven hundred

and thirty-seven aberrational names.

Brown (1956, pp. 140-142), following Bang-Haas

(1915, pp. 181-185) and Eisner (1955, pp. 177-

179), believes that form and aberrational names

"serve a purpose for the serious student of variation

fully as great as species and subspecies names" and

proposes that their system embodying some sixty-six

"names" crossing subspecific and specific lines be

used in the genus Parnassius. This would mean

that each species and subspecies of Parnassius could

receive sixty-six of these "names." Brown goes on

to say that these "names are purely descriptive and

when used do not take an author's name. They

have no nomenclatorial status." If students of variation wish to adopt such a system, I certainly have

no objection as long as its users realize that they

are really using names as terms to help write about

variation within the various species and subspecies.

In the genera Vanessa, Bassaris, and Cynthia, as

well as in almost all other Rhopalocera, no such

system has ever been proposed or used, and so we

are confronted with the problem of what to do

about the numerous names given to individual variations. Fortunately aberrational and other infrasubspecific names were removed from our formal

nomenclature by the International Code of Zoological Nomenclature Adopted by the XV International Congress of Zoology. Article 1 of the Code

states that "This Code is concerned with such

names" (i.e., scientific names) "in the family-,

genus-, and species-groups" and that "names given

to .. . infrasubspecific forms as such . . . are excluded" from zoological nomenclature. Article 5

states that "the name of a species consists of two

words (binomen) and that of a subspecies of three

words (trinomen); in each case the first word is

the generic name, the second word is the specific

name, and the third word, when applicable, is the

subspecific name." Article 45 (a) states that "the

species-group for the purposes of this Code, includes

the categories species and subspecies."

It could hardly be stated more clearly than it is

stated in the Code, but to emphasize this point,

the Code recognizes names for categories only down

to and including the subspecies and since subspecies

names are necessarily in the form of trinomials, all

quadrinomials are excluded as well as all infrasubspecific names. Names proposed for "sub-subspecies"

or "forms" of subspecies are not recognized.

We need to understand clearly what is meant by

subspecific names and infrasubspecific names and

we need to know how to recognize such names.

Article 45 (d) of the Code dictates three situations

under which we are to accept a proposed name as

a subspecific name:

1. The original status of any name of a taxon of
NUMBER 84

lower rank than species is determined as subspecific,

if the author, when originally establishing the name,

clearly stated it to apply to a subspecies. Obviously

the best way to propose a subspecific name is to

state that it is a subspecies.

2. The original status of any name of a taxon of

lower rank than a species is determined as subspecific, if the author, when originally establishing

the name, did not state its rank. This clearly means

that if an author proposed a trinomial name without explaining the trinomen in any way, we are

to accept it as a proposal of a subspecies.

3. The original status of any name of a taxon of

lower rank than a species is determined as subspecific, if the author, when originally establishing

the name, stated the taxon to be characteristic of a

particular geographical area (or geological horizon)

and did not expressly refer it to any infrasubspecific

category. This clearly covers all names proposed in

the past as races, local forms, altitude forms, and

the like, provided they were proposed as trinomial

names.

Article 45 (d) (iii) gives two ways of recognizing

when a taxon is of infrasubspecific status:

1. The original status of any name of a taxon of

lower rank than species is determined as infrasubspecific, if the author, when originally establishing

the name, expressly referred the taxon to an infrasubspecific rank. This necessarily includes names

given to all categories lower in rank than the subspecies and includes all names given to individual

specimens and segments of populations such as

aberrations, transition forms, seasonal forms, wet

and dry forms, cold forms, color forms, sexual

forms, and the names given to the separate generations of the same population.

2. The original status of any name of a taxon of

lower rank than species is determined as infrasubspecific, if the author, when originally establishing

the name, after 1960, did not clearly state that it

was a subspecies.

What to do about the terms "variety" and "form"

are taken up in paragraph (e) of article 45. In this

paragraph on the "interpretation of the terms

'variety' and 'form' " it is stated that " (i) Before

1961, the use of either of the terms 'variety' or

'form' is not to be interpreted as an express statement of either subspecific or infrasubspecific rank."

(ii) "After 1960, a new name published as that of

a 'variety' or 'form' is to be regarded as of infrasubspecific rank." (This is also stated in article 15.)

For publications dated before 1961, we have to

study each author's text to determine what he

meant by his use of the terms "variety" and "form."

If in any way this text conveys the modern idea of

the term subspecies, then we accept the proposed

name. If an author clearly indicates in his text that

he is writing about an individual variant (such as

an aberration, color form, or the like) that is a part

of a population, then we have to reject the name.

On the other hand, if the original author in no way

shows what he meant by the term "variety" or

"form," or if it is not clear that he is naming either

subspecies or individual varients of such populations, then we have to accept any such proposed

trinomial name as an acceptable trinominal under

article 45 (d) (i).

In summary then there are five kinds or types of

names that I regard as acceptable subspecific names

under the Code:

1. Names given as trinomials (article 5), as subspecies (article 45 (d) (i)).

2. Names given as trinomials (article 5), before

1961, where no expressed category or rank is indicated (article 45 (d) (i)).

3. Names given as trinomials (article 5), before

1961, as races, local forms, altitude forms, or given

as trinomials, to any other geographically based

population (article 45 (d) (ii)) .

4. Names given as trinomials (article 5), before

1961, as "varieties" and "forms" where the author

indicates or even hints that they represent geographically based populations (article 45 (d) (ii)

and (e) (i)). Many authors, perhaps most authors

that used the terms "variety" and "form," indeed

used those terms for subspecies. Article 45 (e) (i)

allows us to accept these terms as subspecific or as

infrasubspecific. When it is obvious that subspecies

is meant, as in the case where a geographical area

is involved, we accept the terms as of subspecific

rank.

5. Names given as trinomials (article 5), before

1961, as "varieties" and "forms" where the author

in no way indicates what he meant by the use of

these terms (article 45 (d) (i) and (e) (i)). Article

45 (e) (i) allows us to accept these terms as subspecific or as infrasubspecific. Where we cannot

determine an author's meaning, we may accept
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the terms "variety" and "form" as of subspecific

rank.

There are seven types of names that are excluded

by the Code or that may be excluded under the

Code, and these seven types of names I treat as

excluded names in this paper:

1. All names proposed as quadrinomials (article

5 by recognizing only the generic name, the specific

name, and, when applicable, the subspecific name).

2. All names given to aberrations as such, transition forms as such, seasonal forms, wet and dry

forms, color forms, sexual forms, generation forms

as such, and similar forms (article 1; article 45 (d)

(iii) and glossary of the code: definition of the term

infrasubspecific).

3. All names given to "varieties" and "forms,"

before 1961, where the author clearly indicates that

he is dealing with an individual variant such as

those mentioned above under number 2. (Some

authors used these terms in a very general way to

apply to what we now call subspecies and for

individual variants, using these terms now for one

and again for the other type of variation. Article

45 (e) (i) allows us to interpret these terms either

as infrasubspecific or as subspecific. When it is

obvious that an author meant an aberration or one

of the other types of individual variants in his use

of these terms, then we may exclude the name he

proposed for such a variation.)

4. All names proposed as trinomials, after 1960,

where it is not clearly stated that such names are

subspecific names (article 45 (d) (iii)).

5. All names proposed for "varieties" or "forms"

after 1960 (article 15, article 45 (e) (ii) ) .

6. All names proposed for races, local forms,

altitudinal forms, or to any geographically based

populations, after 1960, where they are not expressly called subspecies (article 45 (a) and article

45 (d) (iii)).

7. All names given to hybrids (article 1).

Names rejected or excluded under the Code may

later become available, for article 10(b) states that

"a name first established with infrasubspecific rank

becomes available if the taxon in question is elevated to a rank of the species-group, and takes the

date and authorship of its elevation." For this

reason all of the literature dealing with the infrasubspecific names used in the species treated in the

present paper had to be examined carefully to

determine if a name which had been excluded because of its treatment in an earlier work might be

available because of a different treatment in a later

work.

The terminal bibliography is selective. A complete bibliography to the literature of the genera

Vanessa and Cynthia would total many times the

number of titles given at the end of this paper.

Hopefully all papers that were taxonomically significant were included and an attempt was made

to include most of the literature giving life-history

information (except for the large part of this literature that is repetitive). The literature was searched

particularly for distribution data for those species

of which only a relatively few specimens were available for study. Thus the greater amount of the

literature for V. atalanta and C. cardui, for example,

was ignored (distribution records within their normal range, state and local lists, and the like), while

an effort was made to be as complete as possible

for the less abundant and more restricted species.

Acknowledgments

I wish to thank Mr. Jose" Herrera of the University

of Chile for the loan of material and for his valuable aid given a number of years ago while he and

I were studying the Chilean species of Cynthia. I

greatly appreciate the many courtesies shown to me

by Drs. Frederick H. Rindge and Alexander B.

Klots during my visits to the American Museum

of Natural History, and my especial thanks go to

the former for allowing me to study specimens in

his care. For the same reasons thanks are due to

Drs. P. J. Darlington and John M. Burns of the

Museum of Comparative Zoology, Harvard University. I am indebted to Dr. Paul H. Arnaud, Jr.,

of the California Academy of Sciences, for the loan

of a large series of the new species of Cynthia described in this paper. I am especially grateful to Dr.

J. W. Tilden of San Jose State College, San Jose,

California, for advice and to his student, Mr.

Thomas Dimock, for a fresh series of the new species

of Cynthia, from which the primary type, the allotype, and the paratypes were chosen (the latter very

kindly offered by Mr. Dimock for distribution to

other museums). Mr. Allan Watson and Mr. T. G.

Howarth of the British Museum (Natural History)

have my thanks for the loan of material formerly
NUMBER 84

unobtainable to me and Mr. Howarth especially

receives my thanks for his critical remarks on part

of my manuscript. Dr. Pierre Viette of the Museum

National d'Histoire Naturelle, Paris, supplied me

with information on certain types in his care for

which I am especially grateful. Dr. F. Kasy of the

Naturhistorisches Museum of Vienna very kindly

supplied me with the news of the destruction by

fire of much of the Hubner collection over a century

ago, a fact not generally known.

Explanation of Figures

In the figures of the male genitalia (Figures 1-16)

the lateral aspect of the whole genital capsule

(minus the aedeagus) is shown on the left-hand side

of the illustrations. The aedeagus in lateral view is

shown in the lower right-hand corner of each figure,

and the uncus and gnathos are shown in ventral

view in the upper right-hand corner. The figure in

the middle is of the ventral element (all that remains) of the anellus (sometimes called juxta or

furca) shown in a ventral view.

In the figures of the female genitalia (Figures

17-32) the bursa copulatrix, seventh and eighth abdominal segments, and ovipositors are shown in ventral view.

These figures were drawn under the author's supervision by Mr. Andre D. Pizzini, staff artist.

Figures 33 through 160 are photographs of the

adult butterflies. These were made by Mr. Andrew

Wynn and Mr. Victor E. Krantz of the Smithsonian

Institution photo laboratory.

Key to the Genera Treated in This Paper

1. Mid and hind tarsi with paronychia simple, consisting of a single slender lobe, sometimes with

a slight basal projection; both surfaces of forewing with middle of cell Cu2 occupied

by a large light-colored spot, greatly contrasting with the fuscous color on either side

of this spot; upper surface of hindwing with a contrasting light-colored spot adjacent

to end of cell, which may be part of a light-colored band crossing wing near middle; forewing underneath with area opposite end of cell and between there and subapical lightcolored bar dark brown or black and divided by white or yellowish-white lines; under

surface of hindwing with very distinct ocular spots contrastingly greatly with surrounding

area Cynthia Fabricius

Mid and hind tarsi with paronychia bifid, lower element slightly shorter than upper element;

both surfaces of forewing not as described above, or if a light-colored spot is present it is

small, not extending more than halfway to anal vein, or if larger it is part of a single

central band running across wing from costa to anal vein; upper surface of hindwing not

as described above, with end of cell concolorous with surrounding area; forewing underneath with dark area opposite end of cell and between there and subapical light-colored

bar solidly dark brown or black, not divided and with one or two blue bands or a complete

circle of blue; under surface of hindwing with submarginal ocular markings obscured, not

constrasting greatly with surrounding area 2

2. Upper surface of hindwing without submarginal blue-centered ocular spots; under surface

of forewing with dark area opposite end of cell and between there and subapical lightcolored bar with blue marking not forming a circle; male genitalia with uncus bifurcate

distally; female genitalia with signum bursae entirely divided or very weakly sclerotized in

middle and relatively short, not more than three times as long as broad .. Vanessa Fabricius

Upper surface of hindwing with a series of submarginal blue-centered ocular spots; under

surface of forewing with dark area opposite end of cell and between there and subapical

light-colored bar with a nearly perfect circle of blue; male genitalia with uncus not birfucate distally; female genitalia with signum bursae not divided in middle, and more than

three times as long and broad Bassaris Hubner

Genus Vanessa Fabricius

FIGURES 1-5, 17-21, 33-80

Vanessa Fabricius, 1807, p. 281,.—Latreille, 1810, pp. 354, 400.—

Boisduval, 1840, p. 21.—Humphreys and Westwood, 1841,

pp. 47-55.—Herrich-Schaffer, 1843, pp. 38-41.—Duponchel,

1844, pp. 6-7.—Lucas, 1845, p. 57—Poey, 1847, pp. 121-124.

—Duponchel, 1849,pp. 92-109—Blanchard, 1852, pp.25-26.

—Stainton, 1857, pp. 36-38.—Rambur, 1858, pp. 13-14.—

Humphreys, [1859], p. 22.—Scudder, 1875, pp. 287-288.—

De Waldheim and Eversmann, 1881, pp. 104-110.—Scudder,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1889, pp. 430-441.—Dale, 1890, pp. 135-154.—Meyrick, 1899,

pp. 193-194.—Moore, 1899-1900, pp. 102-107.—Dyar, 1903,

p. 23.—Bingham, 1905, pp. 363-367.—Lhomme, 1923, p.

51.—Antram, 1924, pp. 175-176.—Barnes and Benjamin,

1926, p. 11.—Hudson, 1928, pp. 34-36.—Holland, 1930, p.

46; 1931, pp. 153-154.—Evans, 1932, p. 177.—Frohawk, 1934,

pp. 16, 134-147.—Hemming, 1934 pp. 60-68.—Bates, 1935,

pp. 164-165.—Peile, 1937, pp. 139-140.—Field, 1940b, pp.

73, 80-88, 274.—Hoffmann, 1940, p. 640.—Pierce and Beirne,

1941, p. 20.—Bryk, 1944, pp. 10-11.—Comstock, 1944, pp.

449-451.—Aubert, 1949, pp. 129-135.—Pinhey, 1949, pp.

82-83.—Verity, 1950, pp. 327-340.—Clark and Clark, 1951,

pp. 13-14, 43-45.—Hayward, 1951, p. 195.—Klots, 1951, pp.

107-108.—Forster and Wohlfahrt, 1955, pp. 55-56.—Corbet

and Pendlebury, 1956, pp. 203, 207.—Lempke, 1956, pp.

185-194.—Wynter-Blyth, 1957, pp. 209-212.—Zimmerman,

1958, pp. 452-472.—Forbes, 1960, pp. 158-159.— dos Passos,

1964, p. 77.—Gifford, 1965, p. 115.—Niculescu, 1965, pp.

185-196.—Hemming, 1967, p. 455.

Nymphalis Latreille [not Fabricius 1804], pp. 184, 199.—

Hemming, 1934, pp. 68-69.—Scudder, 1875, pp. 229-230.—

dos Passos, 1964, p. 77.—Hemming, 1967, p. 316.

Pyrameis Hubner, [1819], p. 33.—Doubleday, 1849, pp. 202-

205.—Horsfield and Moore, 1857, pp. 138-139.—Lucas, 1857,

pp. 542-544.—Morris, 1860, p. 8; 1862. pp. 58-60.—Trimen,

1862-1866, pp. 117-121.—Butler, [1870], pp. 77-78.—Kirby,

1871, p. 185.—Scudder, 1875, p. 260.—Reed, 1877, pp. 678-

680.—Strecker, 1878, pp. 135-138, 190.—Moore, 1880-1881,

pp. 49-51.—Butler, 1881, pp. 466-167.—Godman and Salvin,

1882, pp. 217-219.—Saalmuller, 1884, p. 77.—Marshall and

de Niceville, 1886, pp. 225-229.—Trimen, 1887, pp. 198-

203.—Scudder, 1889, pp. 434, 435, 440, 441.—Dale, 1890, pp.

137-152.—Maynard, 1891, pp. 92-93.—Miskin, 1891, p. 39.—

Leech, 1892-1894, pp. xv, 349-350.—Aurivillius, 1898, p.

130.—Denton, 1898, pp. 264-268.—Holland, 1898, pp. 169-

171.—Miyajima, 1899, pp. 3, [117]—[119].—Cannaviello,

1900, p. 17.—Staudinger and Rebel, 1901, p. 24.—Wright,

1905, pp. 177-178.—Stichel, 1908, pp. 198-200; 1909, pp.

198-200.—Rebel, 1910, p. 20.—Reuss, 1910a, pi, 1; 1910b,

pp. 62-^7.—Fruhstorfer, 1912a, pp. 524-526; 1912b, pp. 524-

526.—Aurivillius, 1913a, p. 227; 1913b, p. 227.—Seitz, 1914a,

pp. 458--159; 1914b, pp. 458-459.—Waterhouse and Lyell,

1914, pp. 55-56.—Holland, 1915, pp. 99-100; 1920, pp. 143-

144.—Bang-Haas, 1926, pp. 55-56; 1927, pp. 55-56; 1928,

p. 26; 1929, p. 11; 1930, p. 162.—Gaede, 1931a, p. 343 —

Hayward, 1931, pp. 68-72.—Clark, 1932, pp. 72-73, 84-91.—

Hemming, 1934, p. 69.—Barrett and Burnes, 1951, pp. 132-

135.—Paulian, 1956, pp. 67-69.—Hemming, 1967, p. 398.

Pyrameides Hiibner, [1826], p. 7.—Hemming, 1967, p. 389.

Amiralis Rennie, 1832, p. 10.—Scudder, 1875, p. 109.—Hemming, 1934, p. 69; 1967, p. 40.

Phanessa Sodovskii, [1837], p. 80.—Humphreys, [1859], p. 22.

—Scudder, 1875, p. 247.—Hemming, 1934, p. 70; 1967, p.

356.

Vanesja Philippi [misspelling of Vanessa], 1859, pp. 1089-

1090.

Payrameis Reed [misspelling of Pyrameis], 1877, p. 735.

Zanessa Hemming, [misspelling of Vanessa], 1967, p. 249.—

Cowan, 1970, p. 70.

Type species: Papilio atalanta Linnaeus. Type by

reason of subsequent selection by Latreille (1810,

pp. 440, 354).

The generic names Vanessa and Cynthia with

closely related type-species (atalanta L. and cardui

L. respectively) were both described by Fabricius

in the same paper on the same page (1807, p. 281).

Because it was feared that Cynthia might take precedence over Vanessa, the latter name was accorded

precedence in Opinion 156 (1944, Opinions Rendered by the International Commission on Zoological

Nomenclature 2: 239-250). At the same time Vanessa Fabricius was placed on the "Official List of

Generic Names in Zoology" as Name No. 601. This

action was deemed necessary because Cynthia and

Vanessa were usually thought to be congeneric. In

the present paper they are regarded as two separate

genera.

Vanessa differs from Bassaris and Cynthia in the

habitus and in the structures of the male and female

genitalia. From Cynthia it differs also in the paronychia.

GENITALIA.—Male (Figures 1-5) with uncus

(in both dorsal and ventral view) bifurcate distally;

lateral arms of gnathos sharply bent, forming a distinct "heel" before or near middle and with distal

part after bend, usually longer than or nearly as

long as the base; aedeagus sharply bent downward

near middle (V. indica, V. dejeani, V. samani),

slightly bent (V. tameamea), or nearly straight with

distal end pointed (not semi-spade shaped); valva

with clasper near middle of inner face, downward

directed; clasper usually small, sometimes large and

overlying cuiller.

Female (Figures 17-21) with genitalia consisting

of a central pod-shaped structure on eighth sternite,

surrounding the ostium bursae, and with two or

more lateral plates on either side of this central

structure; each signum bursae entirely divided or

much more weakly sclerotized in middle and relatively short, not more than three times as long as

broad.

TARSI.—Paronychia of mid and hind tarsi bifid,

lower element slightly shorter than upper element.

HABITUS.—In the genus Vanessa the upper surface of the forewing is of a uniform dark fuscous

ground color with a single red, orange, or yellowish

band crossing this wing from just below middle of

costal margin almost to the anal angle (except in
NUMBER 84

V. dejeanii mounseyi, where this band is lacking

and is replaced by the fuscous ground color); with

middle of interspace Cu^ adjacent to anal vein of

the same uniform color as the base of the wing,

with a white, yellow, or tawny subapical bar opposite end of cell, and with a series of from three to

seven submarginal and subapical white spots in the

interspaces between costa and vein Cuj. Basal two

thirds to three fourths of the upper surface of the

hindwing is uniformly dark fuscous (or as in V.

samani a uniform yellowish color) without the submarginal ocular markings found in Cynthia, these

being replaced by spots that are somewhat darker

than the ground color, that are triangular or crescent shaped, or round or lunular shaped, and that

occur at the outer edge of the fuscous or yellowish

basal and discal area; the light-colored area of hindwing is submarginal in position and lies between

veins Rg or M, and 1st A; through the middle of

this light-colored area there is a series of from four

to six fuscous and lunular, triangular, or round

spots. On the under surface of the forewing, Vanessa differs from Cynthia in having a great amount

of blue in the fuscous area opposite end of cell and

between there and the white subapical bar, and in

having the base of wing below cell and out to at

least the origin of vein Cu2 black or fuscous, much

darker than in Cynthia. On the under surface of

hindwing the overall color is much darker, and the

ocular spots are greatly obscured by the reticulated

dark ground color and are not round in shape but

lunular, or round and flattened along their outer

sides.

LIFE HISTORY.—Some life-history information has

been recorded for all of the species except V.

samani and the subspecies V. indica pholoe, V.

indica buana, and V. dejeanii mounseyi. The food

plants are various genera and species of the Urticaceae (order Urticales). The commonest and most

widely distributed species V. atalanta also feeds

upon Humulus (Moraceae, order Urticales) and

upon occasion has been reported as reared upon

Helichrysum (Compositae, order Asterales) and

Salix (Salicaceae, order Salicales), but these last

two plants are not its usual food and the question

as to how readily it takes to them should be investigated if indeed these records are authentic.

DISTRIBUTION.—The five Vanessa species are

inhabitants of the Old World except for V. atalanta,

which is represented by a distinct subspecies found

over most of the northern half of the New World.

List of Included Species and Subspecies

1. Vanessa atalanta (Linnaeus), 1758

la. Vanessa atalanta atalanta (Linnaeus)

lb. Vanessa atalanta rubria (Fruhstorfer), 1909

2. Vanessa tameamea Eschscholtz, 1821

3. Vanessa indica (Herbst), 1794

3a. Vanessa indica indica (Herbst)

3b. Vanessa indica pholoe (Fruhstorfer), 1912

3c. Vanessa indica nubicola (Fruhstorfer), 1898

3d. Vanessa indica buana (Fruhstorfer), 1898

3e. Vanessa india vulcania Godart, 1819

4. Vanessa dejeanii Godart, 1824

4a. Vanessa dejeanii dejeanii Godart

4b. Vanessa dejeanii mounseyi (Talbot), 1936

5. Vanessa samani (Hagen), 1895

The numbers preceding the above names are also

used before these species and subspecies in the keys

and in the headings.

Key to the Species of Vanessa

(based upon male genitalia)

1. Cuiller smooth; dasper directed toward distal margin of valva (Figure 1)

1. V. atalanta (Linnaeus)

Cuiller with its distal one fourth or more toothed; clasper directed toward ventral margin

of valva or toward ventral arm of valva 2

2. (1) Valva without ventral arm (Figure 2) 2. V. tameamea Eschscholtz

Valva with a very distinct ventral arm 3

3. (2) Acute lobe on costal margin of valva very large, much larger than harpe; ventral arm of

valva without this tooth *

Acute lobe on costal margin of valva small, about size of harpe; ventral arm of valva with

a subapical upward directed blunt tooth (Figure 3) 3. V. indica (Herbst)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Vanessa—Continued

(based upon male genitalia)

4. (3) Upper distal angle of valva bluntly pointed; acute lobe on costal margin of valva not

heavily sclerotized (Figure 4) 4. V. dejeanii Godart

Upper distal angle of valva a rounded lobe; acute lobe on costal margin of valva heavily

sclerotized (Figure 5) 5. V. samani (Hagen)

Key to the Species of Vanessa

(based upon female genitalia)

1. Ostium bursae lobe large, as large as lateral plates, and with opening near posterior

margin of eighth sternite (Figure 17) I. V. atalanta (Linnaeus)

Ostium bursae lobe not so large and with opening placed near anterior margin of eighth

sternite 2

2. (1) Plates on either side of ostium bursae lobe very large and winglike in shape 3

These plates greatly reduced with no heavy sclerotization beyond middle one fourth of

eighth sternite (Figure 18) 2. V. tameamea Eschscholtz

3. (2) Posterior margin of seventh sternite deeply incised in middle, allowing for intrusion of

ostium bursae lobe from anterior margin or eighth sternite 4

Posterior margin of seventh sternite not incised in middle (Figure 21)

5. V. samani (Hagen)

4. (3) Plates on either side of ostium bursae lobe tapering laterally to a fine line (Figure 19)

3. V. indica (Herbst)

Plates on either side of ostium bursae lobe not as described above (Figure 20)

4. V. dejeanii (Godart)

Key to the Species and Subspecies of Vanessa

(based upon habitus)

1. Hindwing above yellow in color except for costal margin, abdominal margin, and marginal and two submarginal rows of fuscous spots (Figures 76-80) 5. V. samani (Hagen)

Hindwing above fuscous, dark brown or black, or sometimes an orange brown, with submarginal band of red, orange, or yellow, enclosing a series of submarginal fuscous spots. 2

2. (1) Forewing above and below with a red, orange, or yellowish band extending from just

below middle of costal margin to near anal angle 3

Forewing above and below without this band, the area which it would occupy being replaced by three separate white spots (Figures 70, 71)

4b. V. dejeanii mounseyi (Talbot)

3. (2) Forewing above with this bright-colored (red, orange, or yellow) band surrounding or

partially surrounding three large fuscous, dark brown, or black spots (one a bar in

middle of cell, one at and above and below base of vein Cu2, and one above and below

this vein near middle) 5

Forewing above with this bright-colored band not surrounding or partially surrounding

such spots (bar in cell only faintly indicated, the other two absent, being completely

absorbed in the dark ground color of the base of wing) 4

4. (3) Forewing above and below with subapical white bar extending from costa to vein M2,

narrow; black spots in marginal orange or orange-red band on hindwing reduced in

size (Figures 36-40; 44-46) lb. V. atalanta ntbria (Fruhstorfer)

Forewing above and below with subapical white bar broader; black spots in marginal

orange or orange-red band on hindwing larger (Figures 33-35; 41-43)

la. V. atalanta atalanta (Linnaeus)

5. (3) Forewing above with band extending across wing from just below costal margin to near

anal angle, yellowish in color and narrow; hindwing above with a submarginal yellowish band (Figures 66-69) 4a. V. dejeanii dejeanii (Godart)

Forewing above with this band orange or orange red in color and broad; hindwing above

with this submarginal band orange or orange red in color 6
NUMBER 84

Key to the Species and Subspecies of Vanessa—Continued

(based upon habitus)

6. (5) Hindwing above with ground color through base of wing dark orange brown or reddish

brown; with outer half of costal margin and area below this dark brown or black, this

color extending along both outer and inner sides of the submarginal orange band (Figures 72-75) 2. V. tameamea Eschscholtz

Hindwing above with ground color not as described above, being entirely fuscous or dark

brown in color except for submarginal orange band and a row of black somewhat

cheveron-shaped spots along inner side of this band 7

7. (6) Forewing above with an orange or orange-red line or band along inner side of black or

fuscous bar in middle of cell 8

Forewing above without this orange or orange-red line or band (Figures 64, 65)

3d. V. indica buana (Fruhstorfer)

8. (7) Forewing above with orange or orange-red color forming a thin line on inner side of

black or fuscous bar in middle of cell 9

Forewing above with orange or orange-red color forming a thicker band on inner side

of black or fuscous bar in middle of cell 10

9. (8) Extreme margin of hindwing above entirely fuscous or black (Figures 50-53)

3c. V. indica nubicola (Fruhstorfer)

This margin of hindwing with some orange between the veins (Figures 54-55)

3b. V. indica pholoe (Fruhstorfer)

10. (8) Forewing above and below with subapical white bar extending from costa to vein M2

narrow; with round white spot below vein M2 greatly reduced and with black bar in

middle of cell usually smaller (Figures 56-63) 3e. V. indica vulcania Godart

Forewing above and below with this subapical white bar broader; with white spot below

vein M2 and black bar in middle of cell larger (Figures 47-49)

3a. V. indica indica (Hcrbst)

1. Vanessa atalanta (Linnaeus)

FICURES 1, 17, 33-46

Papilio atalanta Linnaeus, 1758, p. 478; 1767, p. 776.— Fabricius, 1775, p. 504; 1787, p. 49; 1793, pp. 118-119.—Esper,

1777, pp. 182-197.—Herbst, 1794, pp. 171-175.—Godart,

1821, p. 99.

Vanessa atalanta Godart, 1819, pp. 319-320.—Boisduval,

Rambur and Gaslin, 1832, p. 50.—Boisduval and LeConte,

1834, pp. 175-177.—Humphreys and Westwood, 1841, p.

55.—Doubleday, 1844, p. 78.-Lucas, 1845, pp. 57-58.—

Wagner, 1870, p. 170.—Scudder, 1889, pp. 443, 447.—Dale,

1890, p. 149.—Barrett, 1893, p. 149.—Merrifield, 1893, pp.

58-62.—Favre, 1899, p. 99.—Dyar, 1903, p. 23.—Comstock

and Comstock, 1904, p. 156.—Kirkaldy, 1906, p. 139-

Comstock, 1927, p. 131.—Williams, 1930, pp. 223-225 —

Holland, 1931, p. 153.—Frohawk, 1934, p. 136.—Bates,

1935, p. 165.—Field, 1940b, p. 81.—Macy and Shepard, 1941,

p. 123.—Pierce and Beirne, 1941, p. 20, pi. 3; fig. 1.-

Richards, 1946, pp. 21-22.—Aubert, 1949, pp. 130-132.—

Williams, 1949b, p. 39.—Klots, 1951, p. 107.—Forbes, 1960,

p. 159.

Pyrameis atalanta Doubleday, 1849, p. 204.—Lucas, 1857, p.

542.—Morris, 1862, pp. 58-59.—Trimen, 1862-1866, p. 118.

—Butler, 1870, p. 78.—Kirby, 1871, pp. 185, 648.—Strecker,

1878, p. 135—Staudinger, 1885, p. 97.—Shatz and Rober,

1887, p. 125—Dixey, 1890, pp. 92, 93, 94, 96, 98, 99, 100,

102, 106, 107, 109, 111, 115, 117, 118, 119, 122, 123, 126, 127,

plate 1: figs. 3, 6, 7, pi. 3: fig. 42; 1893, pp. 69-73.—Tutt,

18%, p. 361.—Holland, 1898, p. 170.—Skinner, 1898, pp.

23-24.—Stichel, 1900, pp. 124-125; 1908, pp. 124-125; 1909,

pp. 124-125.—Reuss, 1910b, pp. 62, 63, 64, 65.—1910c, pp.

85, 86, 87, 88; 1910d, p. 303.—Verity, 1913, p. 181.—Seitz,

1914a, p. 458; 1914b, p. 458.—Holland, 1915, p. 100.—

Schultz, 1916, p. 27.—Verity and Querci, 1924, p. 42.

This species is one of the commonest and best

known butterflies in the Northern Hemisphere.

The upper surface of the forewing is mainly dark

fuscous, very dark brown or black, with the Vanessa

orange or red band crossing this wing from just

below middle of costal margin almost to anal angle,

more narrow than in the other Vanessa species except in V. dejeanii, where it is yellow and more

irregular in its traversal of the wing or is even absent as in V. dejeanii mounseyi. It has the white

subapical bar opposite end of cell and normally has

seven submarginal white spots in the interspaces

between costa and vein Cux on this surface of forewing. On the upper surface of the hindwing the

basal three fourths of the area is uniformly dark

fuscous, very dark brown or black, with the dark

submarginal round to triangular shaped spots in

the orange to red marginal band much smaller than
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in the other Vanessa species. V. atalanta differs from

the other species (except V. dejeanii) on the under

surfaces in again having a more narrow reddish or

orange band across the forewing and in having the

hindwing much darker in overall color, usually with

a very contrasting yellowish or whitish triangular

or subtriangular spot near middle of costal margin.

MALE (Figures 33-39).—Forewing on upper surfaces as mentioned before, mainly dark fuscous to

black, with a narrow orange to red band; with a

large pure white subapical bar opposite end of cell;

with a series of seven submarginal white spots between costa and vein Cux and with the first three

of these down to vein R5 very close together, often

completely fused and appearing as one white spot;

between the margin of the wing and these submarginal white spots there are four pale bluish to purplish spots in interspaces R4, R5, M2, and M3; the

white spots in interspaces R5 and Mt are small but

distinct, not indistinct to absent as they are in the

other species of Vanessa.

Hindwing on the upper surface also mainly dark

fuscous to black, with a marginal orange to red

band about the same width or somewhat wider

than the orange or red band of the forewing and

lying between veins R8 and the anal angle; with a

series of four small, round to triangular-shaped

black spots crossing the center of this band between

veins M, and Cu2 and with a relatively large submarginal blue spot below vein Cu2 and between

there and the anal angle.

Under surface of forewing with most of the pattern of the upper surface repeated except that the

subapical area between apex and subapical white

bar is mottled, with several shades of light and dark

brown, the inner margin of the red band is much

more irregular and is invaded by the dark fuscous

or black color of the base of wing, and there are

two large bluish lines or bars or sometimes a large

bluish spot in the black ground color between cell

and subapical white bar.

The hindwing on the under surface is greatly

mottled and intricately lined with numerous shades

of brown, reddish brown, and black and with a

considerable amount of lilac-colored scaling. The

large subtriangular to triangular white or yellowish

spot near middle of costal margin is usually very

distinct, contrasting greatly with the overall dark

color of most of the rest of the wing. There is a

series of five submarginal, somewhat obscured, dark

ocular spots between veins R8 and Cu2 and pale

streaks between these spots and the margin of the

wing, these streaks being of a lilac color and paler

between veins M2 and Cu2 than elsewhere.

Length of forewing, 25-34 mm (average 28.7 mm).

Male genitalia as illustrated in Figure 1 (of V.

atalanta atalanta, my preparation no. 6179), uncus

terminating in a small hook; gnathos in lateral view

with lateral arms forming a boot-shaped structure;

valva with a small, sharp anteriorly directed lobe

at apex and with an obtuse lower angle; clasper

with outwardly directed acute spur; cuiller a spikelike process ending in a single spur or sometimes in

two spurs; aedeagus nearly straight, bent slightly

downward at base, and with distal end pointed;

anellus consisting of a ventral plate, forked both

dorsally and ventrally where it is attached to the

valves. The two subspecies V. atalanta atalanta and

V. atalanta rubria show no differences in the male

genitalia.

FEMALE (Figures 40-46).—Not distinguishable in

habitus from the male sex except that the outer

margin of forewing near vein Cu2 is slightly more

produced and the marginal orange to red band on

upper surface of hindwing is often slightly wider.

Length of forewing, 25-35 mm (average 30.5 mm).

Female genitalia as illustrated in Figure 17 of

V. atalanta atalanta drawn from my preparation no.

3667), with lateral plates located along anterior

margin of eighth sternite deeply concave; with

sclerotization around ostium bursae large and podlike in appearance, as large as each lateral plate and

projecting posteriorly, with ostium bursae near

posterior margin of eighth sternite; signa deeply

indented both anteriorly and posteriorly near middle. There are no differences displayed between the

female genitalia of the two subspecies V. atalanta

atalanta and V. atalanta rubria.

GEOGRAPHICAL VARIATION.—As just indicated, I

recognize two subspecies of V. atalanta, the nominate subspecies V. atalanta atalanta found in the

Old World and V. atalanta rubria of the New

World.

la. Vanessa atalanta atalanta (Linnaeus)

FIGURES 1, 17, 33-35, 41-43

Papllio atalanta [including the synonym ammiralis] Linnaeus,

1758, p. 478 [Article 11 (d), International Code of Zoologi-
NUMBER 84 11

cal Nomenclature states that "a name first published as a

synonym is not thereby made available." Hence ammiralis

is not an available name.]; 1767, p. 779.—Fabricius, 1775,

p. 504; 1787, p. 49; 1793, pp. 118-119.—Esper, 1777, pp.

182-187, pi. 14: fig. 1; pi. 86: fig. 4.—Herbst, 1794, 171-

175, figs. 3-6.

Vanessa atalanta. Godart, 1919, pp. 319-320.—Boisduval,

Rambur and Graslin, 1832, pp. [49]-[50], pi. [13].—Boisduval and LeConte, 1834, p. 177.—Boisduval, 1840, p. 21.—

Humphreys and Westwood, 1841, pp. 54, 55, pi. 15: figs.

1-4.—Herrich-Schaffer, 1843, pp. 38, 41.—Doubleday, 1884,

p. 78.—Duponchel, 1844, p. 7.—Lucas, 1845, pp. 57-58.—

Noel, 1845, pi. 1 .—Duponchel, 1849, pp. 105-106, pi. 12:

figs. 41 a-c.—De Waldheim and Eversmann, 1851, pp. 109-

110, pi. 12: figs. 3, 4.—Stainton, 1857, pp. 37-38.—Rambur,

1858, p. 15—Humphreys, [1859], pp. 21-22, pi. 11: figs.

5-8.—Berge, 1863, p. 53, pi. 3: fig. 3.—Berce, 1867, pp. 164-

165, pi. 11: fig. 1.—Milliere, 1868, pp. 27-28, pi. 88: fig. 3.—

Morris, 1868, pp. 64-66, pi. 27.—Wagner, 1870, p. 170.—

Staudinger 1871, p. 17.—Frey, 1880, p. 26.—Kirby, 1882, p.

11, pi. 6: fig. 1.—Kane, 1885, p. 62.—Buckler, 1886, pp.

176-179, pi. 8: figs. 2, 2a, 2b.—Jordan, 1886, pp. 84-85 —

Failla-Tedaldi, 1887, p. 69.—Scudder, 1889, p. 447.—Dale,

1890, pp. 148-152.—Barrett, 1893, pp. 145-149, pi. 20: figs.

2, 2a-2d.—Fischer. 1895, pp. 17, 18, 22, 24, 33.—Schille,

1895, p. 217.—Horsbrugh, 1896, p. 360.—Standfuss, 1896,

pp. 252-256, pi. 7: fig. 7.—Favre, 1899, p. 99.—Dyar, 1903,

p. 23.—Comstock and Comstock, 1904, p. 156.—Kirkaldy,

1906, p. 139.—Lhomne, 1923, p. 51.—Comstock, 1927, p.

131.—Holland, 1931, pp. 153-154.—Evans, 1932, p. 177.—

Frohawk, 1934, pp. 16, 24, 25, 33, 134-140, 144, pi. 11: figs.

7-12.—Bates, 1935, p. 165.—Peile, 1937, p. 139.—Field,

1940b, pp. 81-82.—Macy and Shepard, 1941, p. 123.—Pierce

and Beirne, 1941, p. 20, pi. 3: fig. 1.—Stokoe, 1944, pp. 92-

93, pi. 11: fig. 7, pi. 12: fig. 3.—Ford, 1945, pp. 45, 60, 70,

90, 103, 154, 247, pi. 2: fig. 4, pi. 6: figs. 1-6.—Richards,

1946, pp. 21-22.—Aubert, 1949, pp. 130-132.—Paul and

Robert, 1949, pi. 16.—Verity, 1950, pp. 334-340, pi. 52: figs.

7-13, pi. 18: fig. 3.—Klots, 1951, p. 107.—Agenjo, 1952, p.

282.—Forster and Wholfahrt, 1955, p. 56, fig. 21 (p. 55),

pi. 15: fig. 12.—Lempke, 1956, pp. 189-191.—Wiltshire,

1957, p. 31.—Wynter-Blyth, 1957, p. 212.—Forbes, 1960, p.

159.—Chalmers-Hunt, 1960-1961, pp. 59-60, 140.—Niculescu, 1965, pp. 191-196, pi. 6: figs. 3, 4.

Pyrameis atalanta.—Doubleday, 1849, p. 204.—Lucas, 1857,

p. 542.—Morris, 1862, p. 59.—Trimen, 1862-1866, p. 118.—

Behr, 1864, p. 125.—Butler, 1870, p. 78.—Kirby 1871, pp.

185, 648.—Newman, 1874, pp. 62-64, fig. 16.—Strecker,

1878, p. 135.—Alpheraky, 1885, p. 598.—Marshall and de

Niceville, 1886, p. 226.—Leech, 1892-1894, p. 352.—White,

1894, pp. 52-53, 63, pi. 3: fig. 7.—Tutt, 1896, pp. 355-361,

fig. 38 (p. 356) .—Holland, 1898, p. 170.—Skinner, 1898,

pp. 23-24.—Cannaviello, 1900, pp. 17-18—Stichel, 1900,

pp. 124-125, pi. 2: fig. 3.—Staudinger and Rebel, 1901, p.

24.—Alpteraky, 1908, pp. 572-573.—Stichel, 1908, pp. 198-

199, pi. 62: fig. c 2; 1909, pp. 198-199, pi. 62: fig. c 2.—

Rebel, 1910, p. 20.—Reuss, 1910a, fig. 18; 1910b, pp. 62, 63,

64, 65; 1910c, pp. 85, 86, 87, 88, 90; 1910d, p. 303.—Fritsch,

1913, p. 25.—Verity, 1913, p. 181.—Seitz, 1914a, p. 458;

1914b, p. 458.—Holland, 1915, p. 100.—SchulU, 1916, p.

27.—Reuss, 1918, p. 41.—Ragusa, 1919, p. 145.—Sevastopulo,

1920, p. 283.—Stephan, 1923, pp. 35-36.—Frohawk, [1924],

pp. 158-165, pi. 27.—Stephan, 1924, p. 25.—Verity and

Querci, 1924, p. 42.—Bang-Haas, 1926, p. 55; 1927, p. 55;

1928, p. 26; 1930, p. 162.—Gaede, 1930a, pp. 199-200; 1930b,

pp. 199-200.—Bryk, 1940, p. 16.—South, 1941, pp. 81-«2,

pis. 46-48.—Zerkowitz, 1946, p. 246.

Pyrameis atalanta atalanta.—Verity and Querri, 1924, p. 42.

Vanessa atalanta atalanta.—Verity, 1950, p. 339.

Papilio ammiralis Godart, 1821, pp. 99-101.—Lucas, 1845,

p. 57.

Papilio amiralis Retzius [misspelling of ammiralis'}, 1783, p.

31, index, p. 23.—Kirby, 1871, p. 648.—Strecker, 1878, p.

136.

Pyrameis atalanta aberration parisiensis Girard [excluded

name, type 2], 1866, pp. 567^568.

Pyrameis atalanta form parisiensis.—Bang-Hass [excluded

name, type 3], 1927, p. 55.

Vanessa atalanta aberration parisiensis.—Verity, 1950, p. 377.

Pyrameis atalanta aberration fracta Tutt [excluded name,

type 2], 1896, p. 355.—Lambillion, 1903, p. 23.—Stichel,

1908, p. 198.—Rebel, 1910, p. 20.—Fritsch, 1913, p. 25.—

Lambillion, 1913, p. 127.—Ragusa, 1919, p. 145.—Wize,

1922, p. 261.—Stephan, 1925, p. 36.—Pionneau, 1924, p. 59.

Vanessa atalanta aberration fracta.—Lhomne, 1923, p. 51.—

Fletcher, 1936, p. 273.—Peile, 1936, p. 181.—Verity, 1950,

p. 336.—Chalmers-Hunt, 1960-1961, p. 59—Niculescu, 1965,

p. 196.

Vanessa atalanta form fracta.—Agenjo [excluded name, type

3], 1952, p. 282.—Lempke, 1956, p. 192.

Vanessa atalanta aberration klemensiewiczi Schille.—Fischer

[excluded name, type 2], 1895, pp. 217-218.—Fischer, 1903.

p. 224.—Lhomme, 1923, p. 51.-Chalmers-Hunt, 1960-1961,

p. 59.

Pyrameis atalanta aberration klemensiewiczi.—Tutt, 1896, p.

p. 357—Stichel, 1900, pp. 126-127.—Staudinger and Rebel,

1901, p. 24.—Stichel, 1908, p. 198; 1909, p. 198.—Rebel,

1910, p. 20.—Reuss, 1910a, figs. 15, 16; 1910b, p. 66; 1910c,

pp. 89, 90.—Fritsch, 1913, p. 25.

Cynthia atalanta transition form klemensiewiczi.—Gunder

[excluded name, type 2], 1927c, p. 270, pi. 8: fig. 1.

Pyrameis atalanta form klemensiewiczi.—South [excluded

name, type 3], 1941, p. 82.

Vanessa atalanta form klemensiewiczi.—Verity, 1950, p. 337.

Vanessa atalanta aberration merrificldi Standfuss [excluded

name, type 2], 1895, p. 90; 1896, p. 387, pi. 7: fig. 8.

Pyrameis atalanta cold variety merrifieldi.—Fischer [excluded

name, type 3], 1900, p. 5.

Pyrameis atalanta aberration merrifieldi.—Stichel, 1900, pp.

124, 126.—Fischer, 1901, p. 326.—Stichel, 1908, pp. 198-

199; 1909, pp. 198-199.—Reuss, 1910b, p. 64; 1910c, pp.

85-86; 1910d, p. 304.

Vanessa atalanta warm variety merrifieldi.—Fischer [excluded

name, type 3], 1903, p. 224.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vanessa atalanta cold variety merrifieldi.—Fischer, 1907, p.

170.

Pyrameis atalanta form merrifieldi.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta form merrifieldi.—Verity, 1950, p. 337.

Vanessa atalanta aberration merifieldi Standfuss [misspelling

of merrifieldi'], 1896, p. 387.

Vanessa atalanta aberration klymene Fischer [excluded name,

type 2], 1896, pp. 18, 57, pi. 5: fig. 6b; 1903, p. 224; 1907,

pp. 170. 201, figs. 9, 11.

Pyrameis atalanta aberration klymene.—Fischer, 1900, p. 5.

—Stichel, 1900, pp. 127-128; 1908, p. 198, pi. 62: fig. c 3;

1909, p. 198, pi. 62: fig. c 3.

Vanessa atalanta form klymene.—Verity [excluded name, type

3], 1950, p. 337.

Pyrameis atalanta italica Stichel [new synonymy], 1900, pp.

125-126; 1908, p. 199; 1909, p. 199.—Lamibillion, 1913, p.

127.—Verity, 1916a, p. 102.—Ragusa, 1919, p. 145.—Verity,

1920, pp. 61-62.—Verity and Querci, 1924, p. 42.

Pyrameis atalanta aberration italica.—Rebel, 1910, p. 20.—

Fritsch, 1913, p. 25.

Vanessa atalanta italica.—Verity, 1950, pp. 339-340.—Niculescu, 1965, p. 196.

Pyrameis atalanta variety cabeauensis Lambillion [excluded

name, type 3], 1903, pp. 22-23.

Pyrameis atalanta aberration cabeauensis.—Bang-Haas [excluded name, type 2], 1926, p. 55.

Pyrameis atalanta form cabeauensis.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa atalanta aberration cabeauensis.—Verity, 1950, p.

336.—Chalmers-Hunt, 1960-1961, p. 59.—Niculescu, 1965,

p. 196.

Vanessa atalanta form cabeauensis.—Lempke, 1956, p. 193.

Pyrameis atalanta aberration and variety nana Schultz [excluded name, type 2 and type 3], 1905, p. 67.

Pyrameis atalanta aberration nana.—Stichel, 1908, pp. 198,

248; 1909, pp. 198, 248.—Rebel, 1910, p. 20.—Fritsch, 1913,

p. 25.—Ragusa, 1919, p. 145.—Stephan, 1923, p. 36.

Pyrameis atalanta form nana.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration nana.—Lhomne, 1923, p. 51.—

Verity, 1950, p. 336.—Chalmers-Hunt, 1960-1961, p. 59.

Vanessa atalanta form nama.—Lempke, 1956, p. 191.

Pyrameis atalanta aberration cyclops Stichel [excluded name,

type 2], 1908, p. 198; 1909, p. 198.

Vanessa atalanta form cyclops.—Verity [excluded name, type

3], 1950, p. 337.

Vanessa atalanta aberration cyclops.—Niculescu, 1965, p. 196.

Pyrameis atalanta aberration umbrosa Fischer [excluded

name, type 2], 1908, p. 130.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form umbrosa.—Gaede [excluded name,

type 3], 1930a, p. 199.—Gaede, 1930b, p. 199.

Vanessa atalanta form umbrosa.—Verity, 1950, p. 337.

Pyrameis atalanta aberration aestiva Reuss [excluded name,

type 2], 1910, p. 303.

Pyrameis atalanta aberration cabeanensis Rebel [misspelling

of cabeaunensis], 1910, p. 20.—Fritsch, 1913, p. 25.

Pyrameis atalanta aberration merrifieldoides Reuss [excluded

name, type 2], 1910d, pp. 303-304.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form merrifieldoides.—Gaede [excluded

name, type 3], 1930a, p. 199; 1930b, p. 199.

Pyrameis atalanta aberration merrifieldoides.—Verity, 1950, p.

336.

Pyrameis atalanta aberration rosea Reuss [excluded name,

type 2], 1910, p. 88.

Pyrameis atalanta form rosea.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta form rosea.—Lempke, 1956, p. 191.

Pyrameis atalanta aberration atalantoides Lambillion [excluded name, type 2], 1911, p. 91.—Bang-Haas, 1926, p. 199.

Pyrameis atalanta form atalantoides.—Gaede [excluded name,

type 2], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration atalantoides.—Verity, 1950, p.

336.

Pyrameis atalanta aberration bialbata Cabeau [excluded

name, type 2], 1911, p. 22.—Bang-Haas, 1926, p. 55.

Vanessa atalanta variety bialbata.—Lhomne [excluded name,

type 3], 1923. p. 51.

Pyrameis atalanta form bialbata.—Gaede [excluded name,

type 3], 1930a, pp. 199-200; 1930b. pp. 199-200.

Vanessa atalanta form bialbata.—Lempke, 1956, p. 192.

Vanessa atalanta aberration bialbata.—Verity, 1950, p. 336.—

Chalmers-Hunt, 1960-1961, p. 59.

Pyrameis atalanta aberration octocyanea Cabeau [excluded

name, type 2], 1913, p. 25.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form octocyanea.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration octocyanea.—Verity, 1950, p. 336.

Pyrameis atalanta aberration cyanosticta Fritsch [excluded

name, type 2], 1913, p. 26.—Bang-Haas, 1926, p. 55.

Pyrameis atalanta form cyanosticta.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration cyanosticta.—Verity, 1950, p. S36.

Pyrameis atalanta aberration eos Fritsch [excluded name, type

2], 1913, p. 25.—Bang-Haas, 1926, p. 55.

Pyrameis atalanta form eos.—Gaede [excluded name, type 3],

1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration eos.—Verity, 1950, p. 336.—Chalmers-Hunt, 1960-1961, p. 59.

Vanessa atalanta form eos.—Lempke, 1956, p. 191.

Pyrameis atalanta aberration flavescens Fritsch [excluded

name, type 2], 1913, p. 25.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form flavescens.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration flavescens.—Verity, 1950, p. 337.

Chalmers-Hunt, 1960-1961, p. 59.

Vanessa atalanta form flavescens.—Lempke, 1956, p. 192.

Pyrameis atalanta aberration hyensis Lambillion [excluded

name, type 2], 1913, pp. 126-127.—Bang-Haas, 1926, p. 56.
NUMBER 84 13

Vanessa atalanta aberration hyensis.—Lhomne, 1923, p. 51.—

Verity, 1950, p. 336.—Chalmers-Hunt, 1960-1961, p. 59.

Pyrameis atalanta form hyensis.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa atalanta form hyensis.—Lempke, 1956, p. 192.

Pyrameis atalanta aberration ochrobrunnea Fritsch [excluded

name, type 2], 1913, p. 25.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form ochrobrunnea.—Gaede, [excluded

name, type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration ochrobrunnea.—Verity, 1950, p.

337.—Chalmers-Hunt, 1960-1961, p. 59.

Pyrameis atalanta aberration pallida Fritsch [excluded name,

type 2], 1913, p. 25.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form pallida.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration pallida.—Verity, 1950, p 337.—

Chalmers-Hunt, 1960-1961, p. 59.

Pyrameis atalanta aberration rubra Fritsch [excluded name,

type 2], 1913, p. 25.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form rubra.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration rubra.—Verity, 1950, p. 336.—

Chalmers-Hunt, 1960-1961, p. 59.

Vanessa atalanta form rubra.—Lempke, 1956, p. 191.—Niculescu, 1965, p. 196.

Pyrameis atalanta aberration sordida Fritsch [excluded name,

type 2], 1913, p. 26.—Bang-Haas, 1926, p. 56.

Pyrameis atalanta form sordida.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration sordida.—Verity, 1950, p. 337.

Vanessa atalanta form sordida.—Lempke, 1956, p. 191.

Pyrameis atalanta aberration bipunctata Gussich [excluded

name, type 2], 1917, p. 214.—Bang-Haas, 1927, p. 55; 1928,

p. 26.

Pyrameis atalanta form bipunctata.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration bipunctata.—Verity, 1950, p. 336.

Pyrameis atalanta variety turcica Wize [excluded name, type

3], 1917, p. 5.—Wize, 1922, p. 261.

Vanessa atalanta form turcica.—Lempke [excluded name,

type 3], 1956, p. 193.

Pyrameis atalanta aberration albipunctata Ragusa [excluded

name, type 2], 1919, p. 145.

Vanessa atalanta aberration albipunctata.—Verity, 1950, p.

336.

Pyrameis atalanta form septiespupillata.—Verity [excluded

name, type 3], 1919 pp. 198-199.—Gaede, 1930a, p. 199;

1930b, p. 199.

Pyrameis atalanta aberration septiespupillata.—Bang-Haas

[excluded name, type 2], 1927, p. 55.

Vanessa atalanta aberration septiespupillata.—Field, 1940b,

p. 83.—Verity, 1950, p. 336.

Vanessa atalanta form septiespupillata.—Lempke, 1956, p.

192.

Pyrameis atalanta form sexiespupillata Verity [excluded

name, type 3], 1919, pp. 198-199.—Gaede, 1930a, p. 199;

1930b, p. 199.

Pyrameis atalanta aberration sexiespupillata.—Ban-Haas [excluded name, type 2], 1927. p. 55.

Vanessa atalanta aberration sexiespupillata.—Field, 1940b,

p. 83.—Verity, 1950. p. 336.

Vanessa atalanta form sexiespupillata.—Lempke, 1956, p. 192.

Pyrameis atalanta variety leliva Wize [excluded name, type

3], 1922, p. 261.

Vanessa atalanta form leliva.—Lempke [excluded name, type

3], 1956, p. 193.

Pyrameis atalanta aberration albimaculata Pruffer [excluded

name, type 2], 1923, p. 281.—Bang-Haas, 1927, p. 55; 1928,

p. 26.

Pyrameis atalanta form albimaculata.—Gaede [excluded

name, type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration albipunctata.—Verity, 1950, p.

336.

Pyrmeis atalanta aberration martha Stephen [excluded name,

type 2], 1923, p. 36.—Bang-Haas, 1927, p. 55.

Pyrameis atalanta form martha.—Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration martha.—Verity, 1950, p. 336.

Pyrameis atalanta aberration millierei Cabeau [excluded name,

type 2], 1923, p. 14.—Bang-Haas, 1927. p. 55.

Vanessa atalanta aberration millierei.—Lhomne, 1923, p. 51.

—Verity, 1950, p. 337.

Pyrameis atalanta form millierei.—Gaede [excluded name,

type 3], 1931a, p. 343; 1931b, p. 343.

Pyrameis atalanta aberration balbiata Pionneau [misspelling

of bialbiata], 1924, p. 59.

Pyrameis atalanta aberration flava Eitel [excluded name,

type 2], 1924, p. 141.—Bang-Haas, 1927, p. 55.

Pyrameis atalanta form flava.—Gaede [excluded name, type

3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration flava.—Verity, 1950, p. 337.

Pyrameis atalanta minutior Verity and Querci [new synonymy], 1924, p. 42.—Bang-Haas, 1927, p. 55.

Pyrameis atalanta form minutior.—Gaede, 1930a, p. 199;

1930b, p. 199.

Vanessa atalanta italica subrace minutior.—Verity, 1950, p.

340.

Pyrameis atalanta aberration testacea Pionneau [excluded

name, type 2], 1924, p. 59^-Bang-Haas, 1927, p. 55; 1930,

p. 162.

Pyrameis atalanta form testacea.—Gaede [excluded name,

type 3], 1931a, p. 343; 1931b, p. 343.

Vanessa atalanta aberration testacea.—Verity, 1950, p. 336.—

Niculescu, 1965, p. 195.

Pyrameis atalanta aberration dewalschei Derenne [excluded

name, type 2], 1926, pp. 90-91.—Bang-Haas, 1927, p. 55.

Pyrameis atalanta form dewalschei.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vanessa atalanta aberration dewalschei.—Verity, 1950, p. 337.

Cynthia atlanta.—Gunder [misspelling of atalanta], 1927c, p.

270, pi. 8: fig.l.

Pyrameis atalanta form angustata Gaede [excluded name,

type 3], 1930a, p. 199; 1930b, p. 199.

Vanessa atalanta aberration angustata.—Verity [excluded

name, type 2], 1950, p. 336.—Niculescu, 1965, p. 196.

Vanessa atalanta aberration albopunctura Frohawk [excluded

name, type 2], 1938, p. 87, pi. 20: figs. 1, 2.

Pyrameis atalanta aberration flavomaculata Lempke [excluded

name, type 2], 1939, pp. 120-121.

Vanessa atalanta form flavomaculata.—Lempke [excluded

name, type 3], 1956, p. 192.

Vanessa atalanta form ocellata Stammeshaus [excluded name,

type 3], 1954, p. 271.—Lempke, 1956, p. 193.

Vaness atalanta form angustifasciata Lempke [excluded

name, type 3], 1956, p. 192.

Vanessa atalanta aberration angustifasciata.—Chalmers-Hunt

excluded name, type 2], 1960-61, p. 59.

Vanessa atalanta form caerulocellata Lempke [excluded name,

type 3], 1956, p. 193.

Vanessa atalanta aberration caerulocellata.—Chalmers-Hunt

[excluded name, type 2], 1960-1961, p. 59.

Vanessa atalanta form fuscescens Lempke [excluded name,

type 3], 1956, p. 191.

Vanessa atalanta aberration fuscescens.—Chalmers-Hunt [excluded name, type 2], 1960-1961, p. 59.

Vanessa atalanta form infranigrans Lempke [excluded name,

type 3], 1956, p. 192.

Vanessa atalanta form reducta Lempke [excluded name, type

3], 1956, p. 192.

Vanessa atalanta form largomarginata Groenendijk [excluded

name, type 5], 1966, p. 22.

Vanessa atalanta form virescens Groenendijk [excluded name,

type 5], 1966, p. 22.

This is the subspecies found in northern Africa

along the Mediterranean coast and in Asia Minor,

Europe, and central Asia.

MALE (Figure 33-35).—V. atalanta atalanta differs from V. atalanta rubria solely in having the

subapical white bar opposite end of cell on both

surfaces of forewing considerably larger, sometimes

as much as twice as large as it is in the latter. Usually

the submarginal white spot in interspace M2 on

the forewing is also slightly larger. Frequently the

orange or red bands on both fore- and hindwings

on the upper surfaces are slightly broader, and the

central black spots in the marginal orange band of

hindwing above are larger; these last two differences, however, are not constant enough to be

considered diagnostic characters.

Length of forewing, 27-31.5 mm (average 28.8

mm).

FEMALE (Figures 41-43).—This sex is very much

like the male and it differs from the females of V.

atalanta rubria in the same way that the males do.

Some females have a larger subapical white bar

and a larger submarginal white spot in interspace

M2 on the forewing than do the males, and some

have a slightly larger wing expanse.

Length of forewing, 27-34 mm (average 30.5

mm).

INDIVIDUAL AND ABERRATIONAL VARIATION.—The

ordinary individual variation in this subspecies

often consists of differences in color of the lightcolored bands on the upper surfaces of the wings,

this color may vary from yellow to orange to red

to crimson. Other individual differences are caused

by black scaling, which sometimes breaks this same

band on the forewing into two or three distinct

parts. Also specimens are common in both sexes

with an extra white dot in this band in interspace

Cu,, and Richards (1946, pp. 21-22) reports that

out of five hundred specimens, one hundred had

this extra white spot. Aberrational and individual

variations have been studied extensively in this

subspecies, unfortunately chiefly by persons who

gave formal names to the individuals they described. A perusal of the literature and synonymy

citations given previously shows that some twentysix writers have given forty-seven formal names to

what are sometimes remarkable aberrations and to

what are most often slight individual variants.

Specimens lacking a particular submarginal white

spot, having an extra white spot, having a misplaced

marking, or showing any slight difference from the

"normal" have received names ad infinitum. These

names serve no useful purpose and all are excluded

from our formal nomenclature or synonymized in

the preceding text. Workers who are interested in

these named aberrations should consult Verity

(1950, pp. 334-340) and Lempke (1956, pp. 189-

191).

SEASONAL VARIATION.—Seasonal variation or variation in habitus between the two broods (where

two broods occur) does not seem to exist in this

subspecies.
NUMBER 84 15

METHOD OF IDENTIFICATION.—Since Linnaeus in

his original description of atalanta (1758, p. 478)

refers to his 1746 work "Fauna Svecica," I consider

Sweden to be the type locality of V. atalanta atalanta. This consideration and his references toother

previous descriptions and illustrations, all of which

refer to the European population, leave no doubt

about the identification of this name. Verity (1913,

p. 181) in his study on the Linnean types states of

atalanta: "The example labelled by Linnaeus is of

the commonest form, with moderately wide crimson

bands." This specimen is in the Linnean collection

of the Linnean Society, London.

SYNONYMY.—See literature citations and previous

synonymy. Papilio ammiralis Linnaeus (1758, p.

478), a name used earlier by Linnaeus (and therefore not available), is listed by him as a synonym.

This name was used by Godart (1821, p. 99) and

in a variant spelling, P. amiralis, by Retzius (1783,

p. 31). Stichel (1900, pp. 125-126) described

Pyrameis atalanta italica as a subspecies from south

and central Italy, stating that it is characterized by

the reduced width of the red band of the forewing.

This is an individual variant. Verity and Querci

(1924, p. 42) describe Pyrameis atalanta minutior

as a race from "Quercianella, near Leghorn" in

Tuscany, Italy. It is doubtful if they had any

reasonable idea of what the term "race" should

mean. At any rate their name is a synonym of V.

atalanta atalanta.

LIFE HISTORY.—The larval food plants are: Urtica

dioica, Urtica urens, and Parietaria officinalis (all

Urticaceae, order Urticales). Larvae feed also upon

Humulus (Moraceae, order Urticales) and have

been reported upon Helichrysum (Compositae,

order Asterales) and Salix (Salicaceae, order Salicales) I These last two plants are not its usual food

and perhaps represent forced rear ings.

Brief descriptions of the larva and pupa were

given as long ago as 1775 (Fabricius, p. 504). Lifehistory notes and descriptions of the immature

stages were given by Esper (1777, pp. 185-186, pi.

14: fig. 1), Herbst (1794, pp. 174-175), Godart

(1819, p. 320), Boisduval, Rambur and Graslin

(1832, pp. [49]-[50], pi. [13]), Humphreys and

Westwood (1841, p. 54, pi. 15: figs. 3, 4), Duponchel (1849, pp. 105-106, pi. 12: fig. 41), Stainton

(1857, pp. 37-38), Morris (1868, p. 66, pi. 27),

Newman (1874, pp. 62-64), Buckley (1888, pp.

176-179, pi. 8: figs. 2, 2a, 2b), Dale (1890, pp. 149-

152), Barrett (1893, pp. 147-148, pi. 20: figs. 2c,

2d), Tutt (1896, pp. 358-360), Frohawk ([1924],

pp. 158-165, pi. 27); (1934, pp. 137-139, pi. 11:

figs. 7, 10), South (1941, p. 82, pi. 46), Stokoe

(1944, pp. 92-93, pi. 11: fig. 7; pi. 12: fig. 3),

Niculescu (1965, pp. 193-195, figs. 100, 101), and

numerous other writers.

DISTRIBUTION.—This species is found in the

Canary Islands, along the Mediterranean coast of

Africa and from Asia Minor east to Pakistan and

Kashmir (northern India) and north throughout

all Europe (except the northernmost portions)

through central Asia north of the Himalayan

Mountains to the Amur River. Very few individuals

survive the winter in the northern part of its range

(even in England) and it reintroduces itself each

year by migrations from the south.

MATERIAL STUDIED.—Twenty males and thirty-five

females were studied from the following localities:

CANARY ISLANDS: Port Orotava, Teneriffe (July).

SPAIN: Sodelleta (September); Segovia (July);

Sierra de Alfacar. FRANCE: Alsace; Royan (July,

October). ITALY: Isola d'Elba (Poggio), Toscana

(May, 500 m). AUSTRIA. CZECHOSLOVAKIA: Prague

(September); Friedland (September). HUNGARY:

Falgamdcsu (August); Baja (September). TURKEY:

Anlakya (December). SYRIA: Beirut.

lb. Vanessa atalanta rubria (Fruhstorfer)

FIGURES 36-10, 44-46

Papilio atalanta.—Fabricius, 1775, p. 504; 1787, p. 49; 179S.

pp. 118-119.

Vanessa atalanta.—Godart, 1819, p. 319.—Boisduval, Rambur and Graslin, 1832, p. 50.—Boisduval and LeConte,

1834, pp. 175-177.—Humphreys and Westwood, 1841, p.

55.—Doubleday, 1844, p. 78.—Lucas, 1845, p. 58.—Poey.

1847, p. 122.—Wagner, 1870, p. 170.—Scudder, 1888, pi. 2:

fig. 6, pi. 78: figs. 58, 59, pi. 83: figs. 52, 53, 55; 1899, pp.

441-456, pi. 12: fig. 5; pi. 20: fig. 7; pi. 33: figs. 29, 29a;

pi. 61: figs. 29, 37; pi. 64: fig. 24; pi. 70: fig. 10; pi. 74:

fig. 35; pi. 81: figs. 2, 6; pi. 86: figs. 61-65.—Dale, 1890, p.

150.—Barrett, 1893, p. 149.—Favre, 1889, p. 99.—Meyrick,

1899, p. 193.—Dyar, 1903, p. 23 Comstock and Comstock,

1904, pp. 154-156, pi. 26: figs. 1, 2.—Barnes and McDunnough, 1917, p. 11.—Comstock, 1927, p. 131, pi. 43: figs.

1, 2.—Holland, 1931, pp. 153-154, pi. 3: fig. 35; pi. 4: figs.

52, 53, 55; pi. 43: fig. 4.—Frohawk, 1934, p. 136.—Bates,

1935, p. 165.—Davenport and Dethier, 1938, p. 163.—McDunnough, 1938, p. 20.—Wild, 1939, p. 26, pi. 7: fig. 1.—

Field, 1940b, pp. 81-84, 274.—Hoffmann, 1940, p. 680.—
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Macy and Shepard, 1941, pp. 122-123.—Pierce and Beirne,

1941, p. 20, pi. 3: fig. 1 Comstock, 1944, p. 451.—

Leighton, 1946, p. 59.—Aubert, 1949, p. 132.—Clark and

Clark, 1951, pp. 13, 43-44, 205, pi. 8: fig. /.—Klots, 1951,

pp. 49, 107, 113, 114, pi. 6: fig. 7, pi. 14: fig. 9, pi. 17:

fig. 2.—Ferguson, 1954, p. 194.—Brown, 1955, p. 100 2 figs.

—Zimmermann, 1958, pp. 453, 454, 455, 456-457, figs. 388,

389, 390, 395, 399— Forbes, 1960, p. 159.—Dos Passos, 1964,

p. 77.

Pyrameis atalanta.—Doubleday, 1849, p. 204.—Lucas, 1857,

p. 542.—Morris, 1860, p. 8; 1862, pp. 58-59.—Girard, 1868,

p. 290.—Kirby, 1871, pp. 185, 648.—Strecker, 1878, pp.

135-136, 190 Godman and Salvin, 1882, p. 219.—Edwards,

1889, p. 25.—Maynard, 1891, pp. 92, fig. 32c (p. 95).—Tutt,

1896, p. 361.—Denton, 1898, pp. 264-266, pi. [54].—Holland, 1898, p. 170, pi. 3: fig. 35, pi. 4: figs. 52, 53, 55,

pi. 43: fig. 4.—Skinner, 1898, pp. 23-24.—Stichel, 1900, p.

125.—Wright, 1905, p. 177, pi. 22: fig. 227.—Kirkaldy, 1906,

p. 139.—Stichel, 1908, p. 199; 1909, p. 199.—Reuss, 1910b,

pp. 62, 63, 64, 65; 1910c, pp. 85, 86, 87, 88, 90.—Verity,

1913 p. 181.—Seitz 1914a, p. 458, pi. 94: fig. A 1; 1914b, p.

458, pi. 94: fig. A 1.—Holland, 1915, p. 100, pi. 33 —

Schultz, 1916, p. 27.

Cynthia atalanta.—Barnes and Benjamin, 1926, p. 14.

Pyrameis atalanta rubria Fruhstorfer, 1909, p. 94.—Martin,

1922, p. 44.—Bang-Haas, 1926, p. 56.

Vanessa atalanta rubria.—Verity, 1950, p. 339.

Vanessa atalanta variety edwardsi Grinnell [excluded name,

type 3], 1918, pp. 113, 114, pi. 4: fig. 3.

Cynthia atalanta aberration edwardsi.—Barnes and Benjamin [excluded name, type 2], 1926, p. 14.

Vanessa atalanta aberration edwardsi.—Comstock, 1927, p.

131, pi. 43: figs. 4, 7.—McDunnough, 1938, p. 20.—Dos

Passos, 1964, p. 77.

Cynthia atalanta hybrid edwardsi.—Gunder [excluded name,

type 7], 1927c, p. 270, pi. 8: figs. 2, 3.

Vanessa atalanta italica aberration edwardsi.—Field [excluded

name, type 1 and type 2], 1940b, pp. 82, 84.

Vanessa atalanta italica Stichel, Field not Stichd [a misidentification], 1940b, pp. 82-84, 264.—Klots, 1951, p. 107.

Forbes, 1960, p. 159.

Vanessa atalanta italica form minutior Verity and Querci,

Field not Verity and Querci [a misidentification], 1940b,

pp. 82, 84, 274.

This is the subspecies found in North America.

Several authors who have mentioned that the North

American population differed from that of European are: Humphreys and Westwood (1841, p. 55),

Girard (1868, p. 290), Fruhstorfer (1909, p. 94),

Seitz (1914a, 1914b, p. 458), Bang-Haas (1926, p.

56), and others. Fruhstorfer is the first and only

author to give a name to this population. All of

these authors, however, mentioned characters for

the two populations that do not hold up and all

missed the real difference between them.

MALE (Figures 36-39).—This subspecies differs

from V. atalanta atalanta mainly in having the

subapical white bar opposite end of cell on both

surfaces and forewing considerably smaller, sometimes half as large as it is in the latter. Usually the

submarginal white spot in interspace M2 on the

forewing is also slightly smaller, frequently the

orange bands on both forewing and hindwing on

the upper surface are slightly more narrow, and the

central black spots in the marginal orange band of

hindwing above are smaller. These last two differences are not constant enough to be considered

diagonostic characters.

Length of forewing, 25-34 mm (average 28.7

mm).

FEMALE (Figures 40, 44-46).—Very much like

the male, differing from the females of V. atalanta

atalanta in the same ways that the males differ from

the males of that subspecies. Some females have a

larger subapical white bar and a larger submarginal

white spot in interspace M2 on the forewing thaji

do the males, and some have a slightly larger wing

expanse. Because this is true, comparisons between

the two subspecies should only be made by sexes.

Length of forewing, 25-35 mm (average 30.5

mm).

INDIVIDUAL AND ABERRATIONAL VARIATION.—The

ordinary individual variation in this subspecies is

quite like that of V. atalanta atalanta (which see).

Fortunately workers have not applied names to

individual variants in this population (probably

because it was not thought to be a separate subspecies) except in one case where a hybrid was

suspected (Grinnell, 1918, pp. 113, 114).

SEASONAL VARIATION.—There is some slight

seasonal variation in the adults of this subspecies

but this is not at all constant. In the summer many

individuals are somewhat darker and larger than

spring individuals, with the band across the forewing more narrow and more reddish in color.

METHOD OF IDENTIFICATION.—After having established that there were two subspecies and that the

name V. atalanta atalanta belonged to the European population, it was only necessary to find the

oldest available name for the North American

population. Fruhstorfer's name V. atalanta rubria

is the oldest name applicable and is the only avail-
NUMBER 84 17

able valid name. His original description (1909, p.

94) establishes that he had specimens of V. atalanta

from Mexico and the United States.

The Fruhstorfer collection in the Museum National d'Histoire Naturelle, Paris, contains four

syntypes of V. atalanta rubric From these, I select

as the lectotype, the lectotype candidate labeled

"lectotype." It is a male specimen from "Mexico."

SYNONYMY.—See previous literature citations.

There is one excluded name and several misidentifications, but there is no synonymy for this subspecies.

LIFE HISTORY.—It is not known whether there

are differences in the immature stages between the

two subspecies or not, nor is it known whether

there are any differences in the life cycles, and it

is not expected that this will be shown to be the

case. The following information does apply to the

North American population.

The larval food plants are reported to be: Urtica,

Boehmeria cylindrica, Paritaria debilis, and in

Hawaii, Pipturus albidus (all Urticaceae, order

Urticales). This subspecies also feeds upon Humulus lupulus (Moraceae, order Urticales).

Edwards (1889, p. 25) and Davenport and

Dethier (1938, p. 163) have listed most of the

references to the life-history notes and to the descriptions of the immature stages. Probably the

single most important reference is Scudder (1889,

pp. 445-453, pi. 64: fig. 24; pi. 70: fig. 10; pi. 74:

fig. 35; pi. 78: figs. 58, 59; pi. 81: figs. 2, 6; pi. 86:

figs. 61-65; pi. 83: figs. 52, 53, 55).

DISTRIBUTION.—This species is found in North

America from Hudson Bay and Great Slave Lake,

south throughout the United States and Mexico

into Guatemala, and in Bermuda, Cuba, and

Hispaniola and has been introduced into the island

of Hawaii.

MATERIAL STUDIED.—One hundred and nine males

and seventy-nine females were studied from the following localities: DISTRICT OF MACKENZIE: Fort Resolution (July). BRITISH COLUMBIA: Victoria (July,

September, October). SASKATCHEWAN: Redvers.

ONTARIO. WASHINGTON: Black Canyon, Brewster

(June) ; Cooney Lake (July) ; Mud Lake, Brewster

(June); Neheotta (August) ; Yakima (October).

OREGON: Crater Lake (July). CALIFORNIA: Camp

Baldy, San Bernardino Mountains (July); Los

Angeles; Mission San Jose, Alameda County (July).

NEVADA: Del Monte Ranch, Reno (August). UTAH:

Provo (July); Stockton (August). COLORADO:

Golden (June). MICHIGAN: Betsy River (August);

Detroit (July, August); Interlochen (August);

Lake Wabekaness (August); Monroe Center (August) . WISCONSIN: Madison (September). ILLINOIS:

Decatur (August). KANSAS: Eureka (April, May,

June, July, October, November); Lawrence (June);

Troy (May). NEW HAMPSHIRE: Mount Washington (July). NEW YORK: Allegany State Park

(July, August); Brooklyn (September); Botanical

Gardens, Brooklyn (June); Franklin County

(July); Jamaica, Long Island; Maspeth, Long

Island (July); Prospect Park (June) ; Staten Island

(September). PENNSYLVANIA: Buena Vista (August) ; New Brighton (September). NEW JERSEY:

Cape May County (July). MARYLAND: Brinklow

City. DISTRICT OF COLUMBIA (June). VIRGINIA:

Alleghany County (May); Apple Orchard Mountain, Bedford County (July, 4,000 ft); Difficult Run

(June); Fairfax County (October); Green Sea

(September); Montgomery County (July); Salem

(June, July, October); Shenandoah National Park

(September); Suffolk (October). NORTH CAROLINA:

Great Smoky Mountain National Park (September) . SOUTH CAROLINA. FLORIDA: Homestead (February) ; Key West (April); Miami; Palm Beach.

LOUISIANA: Harahan (June, July). TEXAS: Brownsville (February); Dallas (September); New Braunfels (March, June); San Antonio (September); San

Benito (May); Tiger Hill. ARIZONA: Hereford;

Huachuca (September); Redington. MEXICO:

Cuernavaca (June); Jalapa; Mexico City (June,

December); Orizaba (July) ; Popocatepetl Park

(June) ; San Luis Obispo. GUATEMALA: Volcan

Santa Maria (June). CUBA: Cerro, Havana (September) ; San Bias, Province Santa Clara (July, 600

ft). HAITI. DOMINICAN REPUBLIC.

2. Vanessa tameamea Eschscholtz

FICURES 2, 18, 72-75

Vanessa tameamea Eschscholtz, 1821, p. 207.—Kirkaldy, 1906,

pp. 138-139.—Perkins, 1913, p. civ.—Williams, 1928, p.

164.—Verity, 1950, p. 329.—Swezey, 1954, p. 157, figs. 26-

28.—Zimmerman, 1958, pp. 452, 453, 454, 456, 463-467,

470-472, figs. 386, 387, 389, 392, 393, 397, 401.—Gorelick

and Wielguns, 1968, pp. 111-114, figs. 1-4.

Pyrameis tameamea.—Tueley, 1878, p. 234.—Alfken, 1903,

p. 569.—Fruhstorfer, 1912a, p. 526, pi. 117: figs, c 3, c 4;

1912b, p. 526, pi. 117: figs, c 3, c 4.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vanessa tammeamea Eschscholtz [misspelling of tameamed],

1821, pi. 5: figs. 8a, 8b.—Blackburn, 1882, pp. 55-56.—Dale,

1890, p. 150.—Meyrick, 1899, pp. 193-194, pi. 5: fig. 9.—

Kirkaldy, 1906, pp. 138-139.—Williams, 1928, pp. 164-169,

figs. 1-3.

Pyrameis tammeamea.—Doubleday, 1849, p. 204.—Trimen,

1862-1866, p. 118.—Kirby, 1871, p. 185.—Standinger, 1885,

p. 98, pi. 37: fig. [d-2].—Fruhstorfer, 1898b, p. 151.—Reuss,

1910c, pp. 85, 86, 87, 88.—Perkins, 1913, pp. xxix, xxxi,

xxxvi, lxix, clv-clvi.

Pyrameis Cordelia Doubleday, 1847, pi. 25: fig. 3.—Doubleday, 1849, p. 204 [synonym of tammeamea].—Meyrick,

1899, p. 193.—Zimmerman, 158, p. 463.

This species is unusual as are two other Vanessa

species (V. samani of Sumatra and V. dejeani of

Java, Bali, Lombok, and Mindanao) in being found

only in a relatively small area of the world, in this

case the Hawaiian Islands. It is the largest species

in the genus and it and V. samani are the two most

lightly colored. The forewing has the centrally

placed orange-yellow band very broad, the broadest

of any species in the genus, and like V. indica and

V. dejeanii has both margins of this wing band very

irregular. Nearly all of the outer and upper part of

this wing entirely to the apex is dark brown or

black. The base of the forewing on this surface is

a light orange brown as is the base of the upper

surface of the hindwing. The ground color of the

outer one third of the hindwing is dark brown, and

within this area there is the usual submarginal band

of orange which, in this species, is much broader

and set in farther from the margin than in the other

species. This band is broken into a series of five

separate spots located between veins Rs and 1st A.

Four of the five fuscous spots near the middle of

these orange spots are triangular in shape, the fifth

spot, located between veins Cu, and 2d A, has blue

along its outer edge and is subdivided into two fused

ocular markings. On the under surface of the forewing and over much of the hindwing the predominant color varies from grayish brown to green,

and these colors alone distinguish V. tameamea

from all other species of Vanessa.

MALE (Figures 72, 73).—Forewing above dark

brown to black, with a broad orange band crossing

wing from below middle of costal margin to near

anal angle, and with both upper and lower margins

of this band very irregular; this band encircling

black spot in middle of cell as it does in V. indica;

subapical light-colored bar opposite end of cell not

pure white, always suffused with orange or entirely

orange or orange brown in color; with series of

submarginal white spots ending just above vein M3,

usually four in number; interspace R5 always lacking such a spot and interspace Mx usually lacking it,

or if present greatly reduced in size and with spot

in interspace between margin M2 sometimes white

and often suffused with orange; between margin of

the wing and these submarginal white spots the

pale bluish spots of interspaces R4, R5, M^ and M3

found in V. atalanta are lacking in this species, although there may be a very faint, pale streak near

this margin.

Hindwing on the upper surface with the basal

one half orange brown and with wing beyond this

dark brown, containing a large submarginal band

of five nearly confluent orange spots, each centered

with triangular-shaped small fuscous spots except

the one located between vein Cu2 and 2d A, which

has blue along its outer edge and is subdivided into

the two fused ocular markings mentioned before.

Under surface of forewing with much of the pattern of the upper surface repeated except ground

color and apex are usually green, there is an additional black spot in the cell, and there are blue

streaks between the end of cell and apical bar. This

apical bar is white, not suffused with orange or

orange brown in color as it is on the upper surface.

Hindwing on under surface with an overall grayish brown to greenish color, greatly mottled, and

lined with shades of white, brown, and usually with

several shades of green.

Length of forewing, 31-37 mm (average 33.4

mm).

Male genitalia as illustrated in Figure 2 (drawn

from my preparation no. 3657), uncus with a broad

blunt distal end; gnathos in lateral view with lateral arms parallel to and as long as uncus; valva

with large rounded lobe at apex and with a short,

sharp projection at lower angle; clasper downward

directed and slightly curved anteriorly; cuiller a

spikelike process with numerous spurlike projections

at distal end; aedeagus slightly curved upward near

base and at distal end, basal one-half slightly undulate; anellus similar to that of V. atalanta, with lateral arms more widely separated and relatively

thinner.

FEMALE (Figures 74, 75).—This sex differs from

the male in having the subapical bar opposite end
NUMBER 84 19

of cell white, not orange or heavily suffused with

orange; the submarginal spot in interspace M2 is

also always white, never sometimes suffused with

orange as it is in the male sex.

Length of forewing, 32—40 mm (average 36.4 mm).

Female genitalia as illustrated in Figure 18

(drawn from my preparation no. 6265), with sclerotization of seventh and eighth sternites greatly reduced, genital plate somewhat trapezoidal in shape;

signa very large with a pale, central area and scabrous elsewhere.

INDIVIDUAL VARIATION.—Zimmerman (1958, p.

465) and Gorelick and Wielgus (1968, p. 114) remark upon the individual variation present on the

under surfaces of the wings, where the predominant

color varies from grayish brown to olive green.

METHOD OF IDENTIFICATION.—The type locality is

Oahu, Hawaiian Islands. The original description

and colored illustrations (Eschscholtz, 1821, p. 207,

pi. 5: figs. 8a, 8b) thoroughly depict the species

known for over one hundred years by this name.

SYNONYMY.—On page 207 of the original description Eschscholtz (1821) spells the name of his new

species as "tameamea" and on the plate in the same

publication (plate 5) as "tammeamea". Kirkaldy

(1906, p. 139) shows that "tameamea" is the more

correct orthography.

Pyrameis cor delta described by Doubleday (1847,

pi. 25: fig. 3) was correctly synonymized by him

(1849, p. 204).

LIFE HISTORY.—The larval food plants are the

following members of the Urticaceae (order Urticales) : Pipturus albidus (the principal food plant),

Boehmeria nivea, Neraudia, Touchardia, and

Urera.

Blackburn (1882, pp. 55-56) described the larva

and Perkins 1913, p. civ) the larva and chrysalis.

Life-history notes and descriptions of the immature

stages were given by Williams 1928, pp. 164-169,

figs. 2, 3), Zimmerman (1958, pp. 463-465, 467, 470,

472), and Gorelick and Wielgus (1968, pp. 111-114,

fig. p. 112).

DISTRIBUTION.—This species is endemic to the

mountainous regions of most the Hawaiian Islands:

Kauai, Oahu, Molokai, Maui, Lanai, and Hawaii.

MATERIAL STUDIED.—Twenty-two males and thirteen females were studied from the following

localities: KAUAI: (January, February, April, June,

July; 3,500 to 4,000 ft); Camp Kokee (500 ft).

OAHU: (March, June); Mount Olympus (May);

Tantalus (May). HAWAII: Kona (September); Olaa

(2,500 ft).

3. Vanessa indica (Herbst)

FIGURES 3, 19, 47-65

Papilio atalanta Linnaeus, Herbst not Linnaeus [a misidentification] 1794, pp. 175-176.

Papilio atalanta indica Herbst, 1794, pi. 180: figs. 1, 2.

Pyrameis indica.—Kirby, 1871, p. 185.—Marshall and de

Niceville, 1886, p. 229.—Shatz and Rober, 1887, p. 125.—

Leech, 1892-1894, pp. xv, 352.—Stichel, 1908, p. 199; 1909,

p. 199.—Fruhstorfer, 1912a, p. 525; 1912b, p. 525.—BangHaas, 1926, p. 56.—Matsumura, 1929, p. 10.

Vanessa indica .—Kirby, 1882, p. 11.—Moore, 1899-1900, p.

104.—Bingham, 1905, pp. 366-367.—Peile, 1937, p. 140.—

Bryk, 1946, p. 39.—Verity, 1950, p. 329.—Niculescu, 1965,

pp. 190, 1%.

This species is the second most common species in

the genus (V. atalanta being the most common) and

is mainly Asiatic with a very broad distribution. The

upper surface of the forewing is somewhat similar

to V. atalanta and V. tameamea, differing from the

latter in having a slightly more narrow orange band

across the wing and like that species differing from

V. atalanta in having both margins of this band

very irregular. Nearly all of the outer and upper

part of this wing is dark brown or black. The base

of forewing on this surface is golden brown or dark

brown suffused with golden brown. The ground

color of the inner three fourths or more of hindwing

on the upper surface is the same color as the base

of the forewing, and as in V. atalanta there is an

orange marginal band, this band usually more narrow and containing a row of dark brown or black

triangular-shaped spots that are much larger than

those found in V. atalanta. V. indica on the under

surface of forewing is more similar to V. atalanta

than to V. tameamea, but the orange band across

this wing is wider than in V. atalanta and intermediate in width to that of V. tameamea. On the

under surface of the hindwing the pattern is very

similar to that of V. atalanta, usually with more

white present in the greatly mottled and lined

pattern.

MALE (Figures 47-48, 50-51, 56, 57, 60, 61, 64,

65).—Forewing on the upper surface dark brown or

black, with the orange band crossing wing much
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

broader than in V. atalanta and somewhat more

narrow than in V. tameamea and, as in the latter,

with both upper and lower margins of this band

very irregular; this band encircling the black spot

in middle of cell as it does in V. tameamea and with

the orange inward from this spot even wider than

in that species; subapical light-colored spots white,

sometimes suffused with a litle orange, especially

near costa; with the series of submarginal white

spots ending just above vein M3, usually six in number; interspace R5 always lacking such a spot and

with those in interspaces R5 and Mx more reduced

in size than in V. atalanta; there are pale streaks

between margin of wing and these submarginal

white spots which, instead of being bluish, are gray

or pale brown and more indistinct.

On the upper surface of hindwing the ground

color is golden brown or dark brown suffused with

golden brown, quite similar to that of the base of

forewing, with a marginal orange band more narrow than the orange band of the forewing and more

narrow than a similar band on hindwing of V.

atalanta; with the series of four black spots crossing

center of this band larger than in V. atalanta and

with two fused blue-edged ocular spots below vein

Cu2; inward the orange marginal band is bordered

by a series of black spots.

Forewing on under surface as in V. atalanta and

V. tameamea, with most of the pattern of the upper

surface repeated except that the subapical area

between apex and subapical white bar is mottled

with several shades of light and dark brown, there

are two black spots in the cell that are surrounded

by orange except along costa, and there is a large

amount of blue in the black ground color between

cell and subapical white bar.

The under surface of hindwing is greatly mottled

and intricately lined with numerous shades of

brown, reddish brown, and black and has a considerable amount of blue and lilac scaling and lines

of white in the basal half of wing. As in V. atalanta

there is a large area of suffused white near middle

of costa, but this pale area is not so clearly triangular or subtriangular in shape; other markings are

very similar to those of V. atalanta.

Length of forewing, 25-34 mm (average 29 mm).

Male genitalia as illustrated in Figure 3 (drawn

from my preparation no. 3649 of V. indica indica),

uncus in lateral view slightly enlarged distally and

distinctly projected toward its lower angle, presenting a somewhat beaklike appearance; gnathos in

lateral view bootlike in shape with a distinct "heel";

valva deeply incised on distal margin, forming an

elongated and usually toothed ventral arm and a

dorsal lobe, with a lower angle that is usually acute,

and with a well-defined acute process near middle

of its outer margin; clasper on valva small and

ventrally directed; cuiller large and armed with two

or more spurlike processes; aedeagus bent upwardly

both anteriorly and posteriorly from just beyond

middle; anellus similar to that of V. atalanta, with

lateral arms more deeply separated from each other.

The five subspecies display no differences in the

male genitalia.

FEMALE (Figures 49, 52-55, 58, 59, 62, 63) .—Females of this species are not separable from the

males except by examination of the genitalia; individuals of this sex, however, often have slightly

wider orange bands on upper surfaces of both foreand hindwing and often have slightly larger black

spots in the middle of the orange band on hindwing.

Length of forewing, 27-37 mm (average 31.3

mm).

Female genitalia as illustrated by Figure 19

(drawn from my preparation no. 6298 of V. indica

indica), with lateral plates of eighth sternite much

larger than in V. atalanta and with podlike sclerotization around ostium bursae small, much smaller

than each lateral plate; posterior margin of seventh

sternite deeply incised, allowing for intrusion of

ostium bursae from anterior margin of eighth sternite; signa divided in middle by pale non-dentate

area.

GEOGRAPHICAL VARIATION.—There are five described subspecies recognized in this paper. The

nominate subspecies V. indica indica has the largest

distribution, being found from northern India to

eastern Siberia and Korea and south through Japan,

Okinawa, and Taiwan to northern Luzon in the

Philippine Islands. The subspecies V. indica vulcania occurs at a great distance from the other

subspecies, being found in the Canary Islands,

Madeira, southern Portugal, and Andalusia in

southern Spain.

From the material I have for study of the three

remaining subspecies, it seems that there is at least

one subspecies, southern in distribution (southern
NUMBER 84 21

India, Ceylon, Celebes), distinct from V. indica

indica and V. indica vulcania. I have been unable

to accumulate enough material of all of these three

to form an opinion as to their subspecific reality,

and so for the present I treat them as separate subspecies. These three are V. indica pholoe, described

from southern India, V. indica nubicola, described

from Ceylon, and V. indica buana, described from

the Celebes.

3a. Vanessa indica indica (Herbst)

FIGURES 3, 19, 47-49

Papilio atalanta indica Herbst, 1794, pi. 180: figs. 1, 2.

Pyrameis indica.—Kirby, 1871, p. 185.—Staudinger, 1885, p.

97.—Marshall and de Nice"ville, 1886, p. 229, pi. 18: fig.

74.—Leech, 1892-1894, pp. xv, 352.—Miyajima, 1899, pp. 3.

[118]-[119], pi. 11: fig. 1.—Staudinger and Rebel, 1901, p.

24.—Stichel, 1908, p. 199, pi. 62: fig. c 4; 1909, p. 199, pi.

62: fig. c 4.—Reuss, 1910a, figs. 13, 17; 1910b, pp. 62, 63, 64,

67; 1910c, pp. 85, 86, 88, 90.—Matsumura, 1929, pp. 9,

10, 19, pi. 5: fig. 2.—Mori, 1934, pp. 72-73.

Vanessa indica.—Kirby, 1882, p. 11.—Moore, 1899-1900, pp.

103-105, pi. 320: figs. 1, la, lb.—Bingham, 1905, pp. 366-

367.—Atram, 1924, p. 176.—Verity, 1950, p. 329.—Yokoyama, 1955, pp. 36, 96-97, pi. 29: fig. 88.—Wynter-Blyth,

1957, p. 212, pi. 37: fig. 7.

Pyrameis indica indica.—Fruhstorfer, 1898a, pp. 60-61; 1912a,

p. 525; 1912b, p. 525.—Cho, 1934, pi. 20: fig. 1.—Doi, 1934,

p. 29.

Vanessa indica indica—Evans, 1932, p. 177, pi. 24: fig. F 36.3.

—Peile, 1937, p. 140.—Okano and Ohkura, 1959, p. 42, pi.

41: fig. 123.

Hamadryas calliroe Hiibner, [1808], pi. 46: figs. 3, 4.

Pyrameis calliroe.—Hubner, [1919], p. 33.

Vanessa calliroe.—Doubleday, 1844, pp. 78-79.—Pryer, 1888,

p. 26; 1889, p. xii, pi. 7: fig. 3.—Gray, 1846, p. 11.—Marshall and de Niceville, 1886, p. 229.—Leech, 1892-1894, p.

352.—Moore, 1899-1900, p. 103.

Pyrameis callirhoe Horsfield and Moore [misspelling of

calliroe], 1857, p. 138.—Trimen, 1862-1866, p. 118.—Bremer,

1864, p. 17.—Leech, 1892-1894, p. 352.—Tutt, 1896, p. 357.

Vanessa collirhoe.—Dale, 1890, p. 150.—Fischer, 1895, pp.

22, 23, 24, 33.

Pyrameis callirrhoe Dixey [misspelling of calliroe], 1890, pp.

94, 107, 112, 117, 118, 123, pi. 1: fig. 10.

Pyrameis indica variety asakurae Matsumura [excluded name,

type 3], 1908, p. 158.—Bang-Haas, 1926, p. 56.

Vanessa indica variety asakurae.—Matsumura, 1908, p. 158.

Pyrameis indica form asakurae.—Fruhstorfer [excluded name,

type 3], 1912a, p. 525—Fruhstorfer, 1912b, p. 525.—Matsumura, 1929, p. 19.

Pyrameis indica ab. horishanus Nire [excluded name, type 2],

1917, p. 145.

Pyrameis indica asakurae ab. horishanus.—Bang-Haas [excluded name, type 1], 1926, p. 56.

Pyrameis indica form horishanus.—Matsumura [excluded

name, type 3], 1929, p. 19.

MALE. (Figures 47, 48).—Upper surface of forewing with ground color on base of wing paler

than in the other four subspecies; with a much

larger subapical white bar opposite end of cell,

a larger submarginal white spot in interspace M2

than in V. indica vulcania, and with these markings slightly larger than in V. indica buana; with

dark brown spots projected into the orange band

from the base along lower vein of cell and into

middle of interspace Cu2 larger than in V. indica

vulcania and, like the latter, differing from V.

indica nubicola and V. indica pholoe in having a

broader band of orange in the cell on the inner

and lower sides of the dark brown or black spot

found in middle of cell.

Upper surface of hindwing with ground color

paler than in the other four subspecies and the

same as in the base of the forewing on this surface; with the triangular to diamond-shaped black

spots through the middle of the marginal orange

band somewhat intermediate in size, larger than

those of V. indica vulcania and smaller than those

of V. indica buana, V. indica nubicola, and V. indica pholoe.

Under surface of forewing with white and dark

markings of upper surface repeated and differing

from the other four subspecies in the same ways

as they differ on the upper surface.

Under surface of hindwing indistinguishable

from V. indica vulcania, not as dark in allover appearance as in V. indica buana.

Length of forewing 25-34 mm (average 30 mm).

FEMALE (Figure 49).—The females of the subspecies are much like the males and differ from the

other four subspecies in the same ways that the

males do. Some females have even larger white

markings on the forewing and have a slightly larger

wing expanse.

Length of forewing, 27-37 mm (average 32.8

mm).

METHOD OF IDENTIFICATION.—The type locality is

India. The original description by Herbst includes

two color figures (1794, pi. 180: figs. 1, 2) which
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

clearly establish this name for the subspecies herein

described.

SYNONYMY.—See literature citations above. Hamadryas calliroe is clearly a synonym of V. indica

indica as is shown by the original colored figures

(Hubner, [1808], pi. 46: figs. 3, 4) and by the type

locality, China (Hemming, 1937, p. 424).

LIFE HISTORY.—The larval food plants are reported to be Boehmeria nivea and Urtica (Urticaceae, order Urticales).

Very little has been written on the immature

stages or the life history. A brief description of the

larva was given by Moore (1899-1900, p. 104) and

a few brief life-history notes were given by Pryer

(1888, p. 26), Matsumura (1929, p. 10), Peile

(1937, p. 140), and Wynter-Blyth (1957, p. 212).

DISTRIBUTION.—This subspecies has a rather

broad distribution from northern India and the

Himalayan Mountains east to Burma, northern

Thailand, China, eastern Siberia, Korea, and Japan

and south from there to Okinawa, Taiwan, and

Luzon of the Philippine Islands. In India it has

been taken in Bombay but is only sporadically found

in the plains, being common in the high hills. There

is one male specimen of this subspecies in the collection of the American Museum of Natural History labeled "Muzo, Columbia." This may represent an introduction but I am inclined to believe

that the label is a mistake.

MATERIAL STUDIED.—Seventy-nine males and

forty-six females were studied from the following

localities: INDIA: Ahwa, The Dangs District, Gujarat States (June, 1,694 ft). Coorg (October);

Darjoeling; Landour; Mussoorie, United Provinces

(May, June, July, 2,000-7,900 feet); Rawalpindi

(March); Simla, Punjab (July); Teesta Valley,

Sikkim. NEPAL: Polhara, Phewa Tal (November).

BHUTAN: Chamsa (September, 13,000 ft); Dotenag

(September, 8,400 ft), CHINA: Central China;

Chengtu; Hsingan Mountain, Tachili (September,

1,500 m); Hunan Province; Kingfoo Shan, Szechuan (August); Kiuhau Shan, Anhwei; Minschan, Tauho, Kansu (July, 2,500 m); Peiping,

Hopei; Suifu, Szechuen; Tsinglingschan Mountains, Kansu (May, 1,500 m), SIBERIA: Vladivostok;

Awatschin Bai, Kamtschatka (July). KOREA:

Whitecapped Mountain, (July); Western Korea

(August). JAPAN: Akashi (October); Kanagawa;

Kyoto (May, June, November); Mount Asama

(September); Mount Hira (July, 1,000 m); Shimojima, Nagano (March). OKINAWA: Kaniku (September) ; Nagatomi (June). FORMOSA; Mount Taihei (November). PHILIPPINES: Baguio, Benguet,

Luzon (February).

3b. Vanessa indica pholoe (Fruhstorfer)

FIGURES 54, 55

Pyrameis indica pholoe Fruhstorfer, 1912a, p. 525; 1912b, p.

525.

Vanessa indica pholoe— Evans, 1932, p. 177.—Peile, 19S7, p.

140.

MALE.—This subspecies was described by Fruhstorfer as somewhat intermediate between V. indica

nubicola and the northern subspecies V. indica

indica. As in the former the orange color outlining

the dark brown bar in the cell on upper surface

of forewing is much more narrow and has much

less orange, sometimes none, in interspace Cu2 near

the cell. The extreme margin of the hindwing on

the upper surface in this subspecies is similar to

V. indica indica in having some orange, although

not quite as much, between the veins.

Length of forewing, 27-29 mm (average 28 mm).

FEMALE.— (Figures 54, 55).—This sex does not

differ in habitus from the male.

Length of forewing, 28—30 mm (average 29 mm).

METHOD OF IDENTIFICATION.—The type locality

is Otocamuna, Nilgiris, India. Two syntypes, a male

and a female, are in the collection of the British

Museum (Natural History). The male specimen

bears the type number Rh. 8711 and the female

the type number Rh. 8712. The original description adequately identifies this subspecies.

I select as the lectotype the male syntype bearing the type number Rh. 8711 and the paralectotype the female syntype bearing the type number

Rh. 8712, both in the collection of the British Museum (Natural History). Labels so indicating these

selections have been placed on these two specimens

by Mr. T. B. Howarth of the staff of that institution.

LIFE HISTORY.—The larval food plants have not

been reported upon and the immature stages have

not been described.

DISTRIBUTION.—This subspecies is found only in

southern India.
NUMBER 84 23

MATERIAL STUDIED.—Three males and two females were available for study and these were from

the following localities: INDIA: Nilgiri Hills; Trichinopoly; Kodaimanal.

3c. Vanessa indica nubicola (Fruhstorfer)

FIGURES 50-53

Pyrameis indica Herbst, Moore not Herbst [a misidentifkation], 1880-1881, p. 50, pl. 27: fig. 2.

Pyrameis indica nubicola Fruhstorfer, 1898a, p. 61; 1898b, p.

151; 1912a, p. 525, pl. 117: fig. c 2; 1912b, p. 525, pl. 117:

fig. c 2.—Martin, 1922, p. 44.—Ormiston, 1924, pp. 32-33.

Vanessa indica nubicola.—Peile, 1937, p. 140.—Evans, 1932,

p. 177.—Woodhouse, 1950, pp. xiii, 50-51, pl. 15: figs. 2, 3.

MALE (Figures 50, 51).—This subspecies was described by Fruhstorfer as having the orange color

outlining the dark brown bar in cell on upper

surface of forewing more narrow than in V. indica

indica, and it is thus quite like V. indica pholoe

in this respect. It is also described as having the

entire margin of the hindwing on the upper surface

dark brown, and this seems to be its only differentiating character. The five males and five females

I have for study also display, as does V. indica

pholoe, a tendency to less orange in interspace Cu2

near cell on upper surface of forewing.

Length of forewing, 26-31 mm (average 28.2

mm).

FEMALE (Figures 52, 53).—The sexes show no

habitus differences.

Length of forewing, 30—33 mm (average 31.2

mm).

METHOD OF IDENTIFICATION.—This subspecies was

originally described from four specimens from "the

mountains of Ceylon" (Fruhstorfer, 1898a, p. 61).

This was later more completely given as Nuwara

Eliya, Ceylon, in the Horton Plains at 3,000-6,000

feet elevation (Fruhstorfer, 1912a, p. 325). Fruhstorfer's original description quite adequately

points out the slight differences displayed by the

Ceylon population.

Three of the original series of four syntypes are

in the Museum National d'Historie Naturelle,

Paris, France. I select as the lectotype, the lectotype

candidate labeled "lectotype" in that collection.

It is a male specimen labeled "Ceylon, 5,000 feet."

LIFE HISTORY.—The larval food plants are reported to be Girardinia heterophylla, G. heterophylla variety palmata, and Urtica neilgherriensis

(all Urticaceae).

The immature stages have not been described and

what little is known about the life history was

given by Ormiston (1924, p. 33) and Woodhouse

(1950, p. 51).

DISTRIBUTION.—This subspecies occurs in the

mountains of Ceylon, seldom below 3,000 feet and

usually from 4,500 feet and above.

MATERIAL STUDIED.—As mentioned above only

five males and five females were available for study

and these are from the following localities: CEYLON:

Haldummulla (2,900-4,000 ft), Nuwara Eliya

(June), Haputale (June).

3d. Vanessa indica buana (Fruhstorfer), new

combination

FIGURES 64, 65

Pyrameis indica buana Fruhstorfer, 1898a, pp. 60-61; 1898b,

p. 151; 1912a, p. 525; 1912b, p. 525-Martin, 1922, p. 44.

MALE (Figures 64, 65).—This subspecies is more

distinctly different from V. indica indica than are

V. indica pholoe and V. indica nubicola and could

be regarded as the extreme end of a step-cline.

Only a single male specimen is available to me for

study and it agrees completely with the original

description. In V. indica buana the dark brown bar

in the cell on the upper surface of the forewing

is fused with the dark brown bar lying along the

lower cellular vein and base of Cu2, and there is

no orange outlining this cellular bar inwardly. The

orange present in interspace Cu2 in middle of wing

in V. indica indica is completely absent in this

subspecies. On the upper surface of the hindwing

there is orange between the veins on the extreme

margin of the wings, as is true of V. indica indica,

and the triangular dark brown spots running

through the middle of the marginal orange band

are larger than in any other subspecies, being

nearly confluent.

Length of forewing, 27.5 mm.

FEMALE.—Unknown. The sex of Fruhstorfer's

specimen was not stated in the original description,

but it is known to be a male (Martin, 1922, p 44).

METHOD OF IDENTIFICATION.—The original description is quite adequate to identify this name as

representing the subspecies treated here.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Martin (1922, p. 44) lists the type (male) in his

list of types in the Fruhstorfer collection before its

sale. This holotype is in the Fruhstorfer collection

in the Museum National d'Histoire Naturelle,

Paris, France. It is labeled: "S. Celebes, Bua-Kraeng,

5,000', Oct. 1895, H. Fruhstorfer."

LIFE HISTORY.—The immature stages, the food

plant, and the life history have not been reported

upon.

DISTRIBUTION AND MATERIAL STUDIED.—Described

from a specimen from the north side of the peak of

Bonthain, South Celebes, at an elevation of about

4,000 feet. The single specimen available to me for

study is from the same locality. This is the way

Fruhstorfer gave the type locality; however, as reported above the holotype is labeled slightly different.

3e. Vanessa indica vulcania Godart

FIGURES 56-63

Vanessa vulcania Godart 1819, pp. 320-321.—Doubleday,

1844, p. 78.-Gray, 1846, p. 11.-Doubleday, 1849, p. 204.-

Horsfield and Moore, 1857, p. 138.

Pyrameis indica vulcania.—Kirby, 1871, p. 185.—Staudinger

and Rebel, 1901, p. 24.-Stichel, 1908, p. 199, pi. 62: fig.

c 5; 1909, p. 199, pi. 62: fig. c 5.

Pyrameis vulcania.—White, 1894, pp. 54, 63, pi. 3: fig. 6.

Pyrameis callirhoe Doubleday [misspelling and misidentification of Hamadryas calliroe Hubner = V. indica indica],

1849, p. 204.-Milliere, 1868, pp. 26-27, pi. 86: figs. 1, 2.-

White, 1894, pp. 54, 63, pi. 3: fig. 6.

Vanessa callirhoe—Kane, 1885, p. 62.

Pyrameis callirhoe variety occidentalis Felder [new synonymy],

1862, p. 473.-Stichel, 1908, p. 199; 1909, p. 199.

Vanessa callirrhoe Staudinger [misspelling and misidentification of Hamadryas calliroe Hubner = V. indica indica],

1871, p. 17.

Vanessa vulcanica Staudinger [misspelling of V. vulcania

Latreille], 1871, p. 17.

Pyrameis indica vulcanica.—Leech, 1892-1894, p. 352.—Reuss,

1910b, p. 63.

MALE (Figures 56, 57, 60, 61) .—Upper surface

of forewing with ground color on base of wing

darker than in V. indica indica; with a much

smaller subapical white bar opposite end of cell

and a smaller submarginal white spot in interspace

M2 than in V. indica indica and the other subspecies; with black spots projected into the orange

band from the base along lower vein of cell and

into middle of interspace Cu^ smaller than in V.

indica indica and, like the latter, differing from V.

indica nubicola and V. indica pholoe in having a

broader band of orange in the cell on the inner

and lower sides of the dark brown or black spot

found in middle of cell.

Upper surface of hindwing with ground color

darker than in V. indica indica and the same as

in the base of the forewing on this surface; with

the triangular to diamond-shaped black spots

through the middle of the marginal orange band

smaller than those of V. indica indica.

Under surface of forewing with white and dark

markings of upper surface repeated.

Under surface of hindwing indistinguishable

from V. indica indica and not as dark in allover

appearance as in the remaining three subspecies.

Length of forewing, 26-32 mm (average 28.8

mm).

FEMALE (Figures 58, 59, 62, 63) .—Much like the

male sex having the same characters. Females have

slightly larger white markings on the forewing and

a slightly larger wing expanse.

Length of forewing, 29-33 mm (average 30.9

mm).

METHOD OF IDENTIFICATION.—The original description by Godart quite adequately gives the

characters that differentiate this subspecies.

SYNONYMY.—See literature citations above. Felder

(1862, p. 473), overlooking Godart's name, described Pyrameis callirhoe occidentalis as a new

subspecies from Madeira.

LIFE HISTORY.—The food plant is reported to be

Urtica dioica (Urticaceae). Descriptions of the immature stages and other life-history information

are not available.

DISTRIBUTION.—This subspecies has a rather

limited distribution, being reported from the Canary

Islands, the Madeira Islands, southern Portugal, the

coast of central Portugal, and from Andalusia in

Spain. It may have evolved from specimens accidentally introduced from India by early Portuguese

traders.

MATERIAL STUDIED.—Nine males and ten females

were studied from the following localities: CANARY

ISLANDS: Canaria; La Palma (November); Monte;

Port Crotava, Tenerife (March); Santa Cruz;

Tenerife. MADEIRA ISLANDS: Porto Santo (February) .
NUMBER 84 25

4. Vanessa dejeanii Godart

FIGURES 4, 20, 66-71

Vanessa dejeanii Godart [1824], pp. 821-822.—Dale, 1890, p.

I50.-Moore, 1899-1900, p. 107.

Pyrameis dejeanii.—Doubleday, 1849, p. 204.—Trimen, 1862-

1866, p. I18.-Kirby, 1871, p. 185.

Pyrameis dejeani Fruhstorfer [misspelling or unjustified

emendation of dejeanii], 1898a, p. 62; 1898b, p. 151.—Reuss,

1910c, pp. 85, 86, 88.-Fruhstorfer, 1912a, p. 525; 1912b,

p. 525.

This species as mentioned before is one of three

Vanessa species that does not enjoy a wide distribution, being found in two subspecies, one found only

on the islands of Java, Bali, Lombok, and Sumbawa,

and the other found only on Mindanao. These two

subspecies differ greatly in habitus, so I offer here

a description to show their similarities and their

essential differences from the other four Vanessa

species.

MALE (Figures 66, 67 = V. dejeanii dejeanii;

Figures 70, 71 = V. dejeanii mounseyi).—Forewing

above with the usual Vanessa light-colored bar

opposite end of cell, with a series of six submarginal

white spots beginning at costa and ending just

above vein M3; with the orange or red of the lightcolored band found in other species of Vanessa

here replaced by yellow, or this band greatly broken

up and represented by individual white spots.

Hindwing on the upper surface with marginal

light-colored band greatly reduced in width and

outlined on inner side with five isolated small, dark

brown or black, round to triangular spots and with

triangular spots found through middle of this marginal band nearly as large as in V. indica.

Under surface of forewing as elsewhere in Vanessa,

with most of the pattern of the upper surface repeated, with all dark coloration much paler, especially in apex of wing, and with a blue bar and a

narrow blue line between the cell and subapical

white bar.

Hindwing on under surface resembling V. indica

more than the other species, mottled and intricately

lined in a similar fashion, with area of suffused

white near middle of costa even more obscure than

in V. indica.

Length of forewing, 23-26 mm (average 24.9

mm).

Male genitalia as illustrated in Figure 3 (drawn

from my preparation no. 3653 of V. dejeanii dejeanii) , uncus, gnathos, aedeagus, and anellus very

similar to those of V. indica, but differing from

that species in the valva, where the ventral arms

are more slender, distinctly upcurved with rough

surfaces, but unarmed (not armed with teeth), and

in having the cuiller on each valva slightly downwardly bent near distal end and armed with numerous spines. The subspecies V. dejeanii dejeanii

and V. dejeanii mounseyi display no differences in

the male genitalia.

FEMALE (Figures 68, 69).—I have seen only the

females of the nominate subspecies and they differ

very slightly from the male sex in having a paler

ground color on upper surfaces of both forewing

and hindwing.

Length of forewing, 23-27 mm (average 25.7

mm).

Female genitalia as illustrated by Figure 20

(drawn from my preparation 6257 of V. dejeanii

dejeanii), similar to those of V. indica, with seventh

and eighth sternites fused and shallowly concave;

with ostium bursae from podlike swelling near

middle of anterior margin of eighth sternite, the

latter protruding into posterior margin of seventh

sternite; each signum elongate and only slightly

paler through middle. Females of V. dejeanii

mounseyi were not available for study.

GEOGRAPHICAL VARIATION.—As indicated above

there are two very different-appearing subspecies,

the nominate subspecies V. dejeanii dejeanii from

several of the Indonesian Islands and V. dejeanii

mounseyi from Mindanao, Philippine Islands.

4a. Vanessa dejeanii dejeanii Godart

FIGURES 4, 20, 66-69

Vanessa dejeanii Godart [1824], pp. 821-822.—Dale, 1890, p.

I50.-Moore, 1899-1900. p. 107.

Pyrameis dejeanii.—Doubleday, 1849, p. 204.—Trimen, 1862-

1866, p. I18.-Kirby, 1871, p. 185.

Pyrameis dejeani Fruhstorfer [misspelling or unjustified

emendation of dejeanii], 1898a, p. 62; 1898b, p. 151.—Reuss,

1910c, pp. 85, 86, 88.

Pyrameis dejeani dejeani.—Fruhstorfer, 1912a, pp. 525-526,

pi. 117: fig. c 1; 1912b, pp. 525-526, pi. 117: fig. c 1.

Pyrameis dejeani sambaluna Fruhstorfer [new synonymy],

1898b, pp. 150-151; 1912a, p. 526.-Martin, 1922, p. 44.
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pyrameis dejeani satnbulana Fruhstorfer [misspelling of

sambaluna], 1912a, p. 526.-Talbot, 1936, p. 112.

This, the nominate subspecies, is found on the

islands of Java, Bali, Lombok, and Sumbawa.

MALE (Figures 66, 67).—Upper surface of forewing with ground color of base and inner four

fifths of anal interspace light brown with a dull

greenish-gold suffusion; ground color of remainder

of wing dark brown with a subapical white bar

opposite end of cell, with six submarginal white

spots down to vein M3, and with a yellowish to

whitish-yellow band across middle of wing from

costa to near anal angle; this band irregular in

shape and broken by black streaks along veins Cuj

and Cu2.

Upper surfaces of hindwing with ground color

like that of base of forewing, light brown with a

dull greenish-gold suffusion; with a submarginal

band of dirty yellow to dirty orange enclosing a

row of triangular to diamond-shaped dark brown

spots and with a row of similar but smaller spots

along its inner edge.

Under surface of forewing with yellow band

through middle of wing and submarginal white

spots and subapical white bar repeated; ground

color of apex of wing grayish brown with blue

streaks in dark brown area opposite end of cell

and between there and subapical white bar, and

with a U-shaped yellow and orange bar outlining

lower and inner sides of dark brown spot in middle

of cell.

Under surface of hindwing grayish brown, greatly

mottled, and reticulated with lines and spots of

darker brown, gray, and white; with a series of

triangular submarginal blue spots between veins

Sc and anal angle, and inward from this series a

series of eye spots between veins Rs and Cu2, two

of these in interspaces M2 and M3 are blue in color,

the other three are dark brown.

Length of forewing, 23-26 mm (average 24.9

mm).

FEMALE (Figures 68, 69).—The females are quite

similar to the males, with a slightly paler ground

color on the upper surfaces of the wings.

Length of forewing, 23-27 mm (average 25.7

mm).

METHOD OF IDENTIFICATION.—Godart's original

description ([1824], pp. 821-822) is very clear and

applies only to the subspecies treated herein as V.

dejeanii dejeanii. The type locality is Java.

SYNONYMY.—Fruhstorfer (1898b, pp. 150-151)

describes from Lombok what he took to be a different subspecies as Pyrameis dejeani (sic!) sambaluna.

His character, a darker and narrower median band

on the forewing above that, is sometimes heavily

suffused with black and is wholly within the range

of individual variation of Vanessa dejeanii dejeanii.

I have therefore synonymized his name to the latter.

This synonym was described by Fruhstorfer from

twelve specimens "vom Plateau von Sambalun, Insel

Lombok, gefangen in April 1896." Martin (1922,

p. 44) indicates both sexes were represented. A

holotype designation was not published by Fruhstorfer. Grabriel (1927) does not list this name

among the types in the collection of the British

Museum (Natural History), but a male specimen

bearing a type label is in that collection. A second

specimen bearing a type label is in the Museum

National d'Historie Naturelle, Paris. I do not see

a need for a lectotype designation at the present

time.

LIFE HISTORY.—The immature stages and the life

history have not been reported upon except that

the larva were observed by Fruhstorfer (1912a and

1912b, p. 526) on the island of Lombok, feeding

upon a common large member of the Urticaceae,

which obtained a height of from 3 to 6i/2 feet. This

plant has not been further identified.

DISTRIBUTION.—As mentioned before, this subspecies is known only from the Indonesian islands

of Java, Bali, Lombok, and Sumbawa.

MATERIAL STUDIED.—Fourteen males and four females from the following localities were studied:

JAVA: Tosari. LOMBOK: Sambalun (4,000 ft.).

SUMBAWA.

4b. Vanessa dejeanii mounseyi (Talbot),

new combination

FIGURES 70, 71

Pyrameis dejeanii mounseyi Talbot, 1936, pp. 112-113.

MALE (Figures 70, 71).—Upper surface of forewing with ground color of base of wing and of anal

interspace dark brown, not light brown as in Vanessa

dejeanii dejeanii, and without the gold suffusion

also found in the latter; ground color of remainder
NUMBER 84 27

of wing even darker brown, almost black; with a

subapical white bar opposite end of cell and with

six submarginal white spots down to vein M3. The

yellowish to whitish-yellow band across middle of

wing in V. dejeanii dejeanii is here replaced by

three greatly separated white spots, a large one at

end of cell, another large one in middle of interspace Cuj, and a much smaller one in interspace

Oil,, near anal angle.

Upper surface of hindwing with ground color

dark brown like that of base of forewing, with submarginal band dark red orange in color, and with

other submarginal markings approximately as in V.

dejeanii dejeanii.

Under surface of forewing with separate white

spots in cell and in interspaces Cuj and Cu2 repeated on this surface as are also the submarginal

white spots and the white subapical bar; ground

color of apex of wing darker than in V. dejeanii

dejeanii, with blue streak in dark brown area

opposite end of cell, between there and subapical

white bar obscure, and with U-shaped bar outlining

lower and inner sides of dark brown spot in middle

of cell entirely white, not orange or yellow as it is

in V. dejeanii dejeanii.

Under surface of hindwing similar to that of the

nominate subspecies but much darker.

Length of forewing, 26 mm.

FEMALE.—As indicated before, the females are

unknown to me. Talbot (1936, p. 113) describes

this sex as having a slightly darker ground color,

with the yellow-brown area on upper surface of

hindwing, which forms a border to the submarginal

spots, not reduced as it is in the male, and with the

under surface as in the male.

METHOD OF IDENTIFICATION.—The type locality is

Mindanao, Philippine Islands. The holotype is a

female specimen in the collection of the British

Museum (Natural History). Talbot's original description (1936, pp. 112-113) clearly describes this

very striking subspecies.

LIFE HISTORY.—Nothing is known concerning the

food plant or the immature stages of this subspecies.

DISTRIBUTION.—This subspecies is known only

from the island of Mindanao, Philippine Islands.

MATERIAL STUDIED.—A single male specimen was

available for study from Mount Apo, Mindanao

(July, 6,000 ft) .

5. Vanessa samani (Hagen)

FIGURES 5, 21, 76-80

Pyrameis samani Hagen, 1895, pp. 359-360; 1896, p. 165, pi.

1: fig. 7.-Fruhstorfer, 1898a, p. 62; 1898b, p. 151; 1912a,

p. 525, pi. 117: fig. B 5; 1912b, p. 525, pi. 117: fig. B 5.

Vanessa samani.—Moore, 1899-1900, p. 107.

V. samani is the rarest and most isolated species in

the genus, being found only in the mountains of

Sumatra and apparently known only from two

specimens up to the year 1912. It is a relatively

small species and the most lightly colored species

of Vanessa, having nearly all of the hindwing and

lower half of forewing, execpt extreme base, ocherous yellow in color.

MALE (Figures 77, 78).—On the forewing above,

this species has, as just mentioned, the light-colored

band, ocherous yellow in color, and greatly widened

toward base of wing; the black spot in middle of

cell is more narrow than in the other species; the

light-colored subapical bar opposite end of cell is

ocherous yellow or heavily suffused with that color;

the usual black spot covering middle of interspace

Cux is almost gone, represented only by a small,

very faint spot, only slightly darker than the ocherous yellow ground color; there are only three very

small submarginal white spots in the black apical

area, one each in interspaces R4, Mx and M2, those

in interspaces R4 and Mx being mere points.

Hindwing on upper surface entirely ocherous

yellow except for the fuscous, dark brown or black

along outer two thirds of costal margin, except for

two rows of parallel, round to triangular-shaped

submarginal fuscous spots, and except for this same

fuscous color along the outer margin, especially at

the ends of the veins.

Forewing underneath similar to upper surface,

with most markings repeated, with apical area

much paler than it is above, and with narrow blue

semicircle or two nearly parallel lines of blue in

dark area found inward from subapical bar of

ocherous yellow opposite end of cell.

Hindwing on the under surface very similar to

that of V. dejeanii dejeanii, with overall color even

paler.

Length of forewing, 25 mm.

Male genitalia as illustrated in Figure 5 (drawn

from my preparation no. 6281), quite similar to

those of V. indica and V. dejeanii, differing from
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

V. indica, as does V. dejeanii, in having ventral

arms of valva more slender, distinctly upcurved

and with rough surfaces, but unarmed; it differs

from V. dejeanii mainly in having cuiller distinctly

constricted before distal end.

FEMALE (Figures 76, 79, 80).—Entirely like the

male in habitus.

Length of forewing, 23 mm.

Female genitalia as illustrated in Figure 21

(drawn from my preparation no. 6282), with

seventh and eighth sternites not as extensively

sclerotized as those of V. indica and V. dejeanii,

with these sternites fused and shallowly concave on

either side of ostium bursae, which is swollen and

podlike and does not intrude into posterior margin

of seventh sternite; with each signum completely

divided by a broad pale band.

METHOD OF IDENTIFICATION.—Hagen's original description (1895, pp. 359-360) and his subsequent

illustration in color (1896, pi. 1: fig. 7) clearly

depict this species. The type locality is given as,

"Karo-Lanndern (1000-1200 m), Sumatra."

According to Hagen's subsequent description only

two specimens were known to him and he did not

indicate the sex. These syntypes may be found in

the collection of the Zoologisches Museum of the

Humboldt Universitat, Berlin.

LIFE HISTORY.—Nothing is known about the food

plants, immature stages, or life history of this

species.

DISTRIBUTION.—This species is found only in the

mountains of Sumatra.

MATERIAL STUDIED.—One male and one female

from the collection of the British Museum (Natural History) were available for study. The data

on the male specimen is Sungei, Kumbang, Korinchi,

west Sumatra (April, 4,500 ft). The data on the

female specimen is Danan Bento Morass, foot of

Korintji Peak, southwest Sumatra (August, 5,000

ft).

Vanessa aequatorialis Wagner, nomen nudum

Vanessa aequatorialis Wagner, 1870, p. 171.

Pyrameis aequatorialis.-Reuss, 1910b, p. 66.

Pyrameis myrinna form aequatorialis.—Seitz, 1914a, p. 459;

1914a, p. 459.

This name used by Wagner for a species found on

Mount Chimborazo, Ecuador, without one word

of description is therefore a nomen nudum. It was

next used by Staudinger who stated only that it is

a questionable and unknown species. Reuss used

the name also but did not mention a single character, only stating that it was found in Ecuador.

The most recent use of this name was by Seitz, who

assigns it to Pyrameis myrinna as a form and in the

next sentence states that it may be identical to

Pyrameis huntera braziliensis or a synonym of the

preceding species (P. terpsichore). He then goes on

to say that it would be well to strike out the name

entirely. Because it is a nomen nudum and cannot

be assigned to any recognized species, the name

should be dropped from nomenclature. From its

supposed country of origin it seems likely that it

was intended for a species of Cynthia.

Genus Bassaris Hiibner, resurrected genus

FIGURES, 6, 7, 22. 23, 81-88

Bassaris Hubner, [1821], pi. [24].-Scudder, 1875, p. 126.-

Hemming, 1934, p. 70; 1967, p. 74.

Type species: Papilio itea Fabricius. Type by reason

of being the sole included species.

Scudder (1875, p. 126) synonymizes Bassaris to

Vanessa and Hemming (1934, p. 70) concurs. Later

Hemming (1967) does not indicate that he considered it to be a synonym of Vanessa, nor does he

indicate anything other than that it is a valid and

available generic name. The two species here assigned to Bassaris have heretofore nearly always

resided in the genus Vanessa.

Bassaris differs from Vanessa and Cynthia in the

habitus and in the structures of the male and female

genitalia. From Cynthia it differs also in die paronychia, which in Vanessa and Bassaris are alike.

GENITALIA.—Male (Figures 6, 7) with uncus

blunt, not bifurcate distally; lateral arms of gnathos

rather blunt and sharply bent beyond middle, forming a "heel" on ventral margin; extreme distal end

of each lateral arm bent downward as in Cynthia

(except C. carye and C. annabella) so that the whole

aspect of gnathos is directed downward; aedeagus

with basal one half or more straight, sharply bent

upward beyond middle, and with distal end sharply

pointed; valva with clasper near middle of inner

face, large and downward directed; dorsal margin

nearly straight or only with a shallow lobe and
NUMBER 84 29

terminating at apex, with an upward directed lobe;

distal margin continuous with lower margin, lacking any pronounced lobe; cuiller unarmed and upward directed in one species (B. itea) and caudally

directed in the other species (B. gonerilla).

Female (Figures 22, 23) with genitalia consisting

of a central pod-shaped structure on eighth sternite,

surrounding the ostium bursae, and with lateral

plates either winglike (B. itea) or very large, embracing most of the remainder of eighth sternite,

and with rounded posterior margins (B. gonerilla);

with lateral saucer-like depressions (B. itea) or deep

pockets to each side of ostium bursae (B. gonerilla);

signum bursae very long, similar to those of

Cynthia.

TARSI.—Paronychia of mid and hind tarsi bifid

as in Vanessa, lower element slightly shorter than

upper element.

HABITUS.—The forewing on upper surface is of

a uniform fuscous to black color except for the

usual subapical and submarginal white to yellowish

spots and a band of red or yellow running across

the wings from below middle of costal margin to

near anal angle. The upper surface of the hindwing

is mainly fuscous or brown with four submarginal

ocular spots, thus resembling Cynthia in the latter

character. On the under surface of the forewing

there is a blue circle in the fuscous area opposite

the end of cell and between the cell and the subapical yellow or white bar. The pattern of the hindwing on the under surface resembles Vanessa, with

the submarginal ocular markings a little more

round, not so flattened along their outer markings.

LIFE HISTORY.—Brief notes and descriptions on

the life histories of the two Bassaris species have

been published and their hosts are reported to be

Urtica (Urticaceae).

DISTRIBUTION.—This genus is known only from

Australia, New Zealand, and a few islands in the

south Pacific region.

List of Included Species and Subspecies

1. Bassaris itea (Fabricius), 1775

2. Bassaris gonerilla (Fabricius), 1775

2a. Bassaris gonerilla ida (Alfken), 1889

2b. Bassaris gonerilla gonerilla (Fabricius)

Key to the Species of Bassaris

(based upon male genitalia)

Cuiller greatly swollen at base, this base armed with teeth along its inner margin, and with its

distal end directed toward apex of valva (Figure 6) 1. B. itea (Fabricius)

Cuiller with base not swollen, this base armed with teeth along its inner surface, and with its

distal end directed downward (Figure 7) 2. B. gonerilla (Fabricius)

Key to the Species of Bassaris

(based upon female genitalia)

Ostium bursae opening before middle of eighth sternite; with deep lateral pouches on each side

of eighth sternite (Figure 23) 2. B. gonerilla (Fabricius)

Ostium bursae opening from anterior margin of eighth sternite, which is projected into seventh

sternite; without these deep pouches (Figure 22) I. B. itea (Fabricius)

Key to the Species and Subspecies of Bassaris

(based upon habitus)

Forewing above with a large, broad yellow band near middle of wing between costa and anal

vein; subapical bar extending from costa to vein M2 yellow or yellowish white; hindwing

above with submarginal blue-centered ocular markings very small and with ground color

of base of this wing concolorous with area surrounding these ocular spots (Figures 81-

g4\ 1. B. itea (Fabricius)

Forewing above with this yellow band replaced by an orange or orange-red band; subapical

bar white in color; hindwing with submarginal blue-centered ocular markings large and
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species and Subspecies of Bassaris—Continued

(based upon habitus)

with ground color of base of wing darker, not concolorous with area surrounding these

ocular spots ^

2. Hindwing above with orange-red band surrounding blue-centered ocular spots larger; whole

aspect of under surface of hindwing paler (Figures 87, 88) . .2b. B. gonerilla ida (Alfken)

Hindwing above with this band smaller; whole aspect of under surface of hindwing darker

(Figures 85, 86) 2a. B. gonerilla gonerilla (Fabricius)

1. Bassaris itea (Fabricius), resurrected combination

FIGURES 6, 22, 81-84

Papilio itea Fabricius, 1775, p. 498; 1787, p. 45; 1793, p. 103.

Bassaris itea.—Hiibner, [1821], pi. [24],—Scudder, 1875, p.

126.—Hemming, 1967, p. 74.

Vanessa itea.—Godart, 1819, p. 321.—Boisduval, 1832, pp.

121-122.—Doubleday, 1844, p. 78.—Dale, 1890, p. 150.—

Hudson, 1898, pp. 107-108, pi. 12: figs. 3, 4; 1928, pp. 35-

36, pi. 4: fig. 8—Verity, 1950, p. 329.—Gaskin, 1966, p. 88,

pi. 1: fig. [6].—Burnes, 1969, pp. 80, 82, fig. 59.—Pritchard,

1969, p. 12.

Pyrameis itea.—Doubleday, 1849, p. 204.—Trimen, 1862-1866,

p. 118.—Butler, 1870, p. 78.—Semper, 1878, p. 148.—Kirby,

1871, p. 185.—Staudinger, 1885, p. 98.—Mathew, 1888, pp.

145-146, pi. 6: fig. 10.—Dixey, 1890, p. 113.—Miskin, 1891,

p. 39.—Fruhstorfer, 1898b, p. 151.—Waterhouse, 1903, p.

11.—Reuss, 1910c, pp. 85, 88.—Fruhstorfer, 1912a, p. 526,

pi. 117: figs, E 1, D 5; 1921b, p. 526, pi. 117: figs, E 1, D

5.—Waterhouse and Lyell, 1914, p. 55, pi. 5: fig. 34.—

Barrett and Burnes, 1951, pp. 134-135.

MALE (Figures 81, 82).—Upper surface of forewing with ground color of apical and outer half

dark brown, almost black, and containing a subapical bar of yellow or yellowish white opposite

end of cell; with four subapical white spots between

costa and vein R5 and in interspace M2; base of

wing reddish brown suffused with dull gold; with

a black bar through middle of cell and with a large

yellow band lying over middle of wing from costa

to anal vein.

Hindwing on upper surface with apical area and

outer margin dark brown, almost black; with cell

area and area between vein M2 and first anal vein

colored reddish brown; base of wing suffused with

dull gold and with four small submarginal ocular

spots pupillated with blue, lying between veins

Mx and Cu2. The ocular spot lying in interspace

M, is at least partially obscured by the dark brown

color of the apical part of the wing.

Under surface of forewing with most of the pattern of the upper surface repeated, with much gray

in the apex; with a blue circle between the yellow

subapical bar and the yellow band of middle of the

wing; base of cell reddish brown in color and base

on interspace Cu2 and anal margin dark brown.

Under surface of hindwing intricately lined and

mottled with shades of black, dark brown, gray,

and lilac and with a series of five obscure submarginal dark ocular spots between veins Rs

 and Cu2.

Length of forewing, 23-31 mm (average 26.3 mm).

Male genitalia as illustrated by Figure 6 (drawn

from my preparation no. 3655), with distal end of

uncus rounded in lateral view, much as in B. gonerilla; gnathos downward directed and quite similar

to B. gonerilla; aedeagus differing from that species

in having a spadelike distal end; valva with dorsal

margin nearly straight, with a slight lobe beyond

middle near apex, and with a large upward directed

lobe at apex; clasper a large, long, and finger-like

lobe, downward directed so that it can and sometimes does cover the cuiller; cuiller with a large

swollen base, which is anteriorly toothed, greatly

reduced and constricted beyond middle, upward

bent, smooth, and spikelike.

FEMALE (Figures 83, 84).—Hardly differing from

the male sex except in having the outer margin of

hindwing slightly more rounded and in having

the area on either side of vein Cu2 on the forewing

slightly more produced outwardly.

Length of forewing, 23-30 mm (average 27.3 mm).

Female genitalia as illustrated by Figure 22

(drawn from my preparation no. 6273), with seventh

sternite only slightly sclerotized through the central

area; eighth sternite with pod-shaped structure surrounding the ostium bursae, with wing-shaped lateral plates, and without the deep lateral pouches

found in B. gonerilla; signa very long and narrow.

METHOD OF IDENTIFICATION.—The type locality

is New Zealand. A single original male specimen,

the holotype, is in the collection of the British Museum (Natural History) and bears the type number
NUMBER 84 31

Rh. 8719. The original description of Fabricius

(1775, p. 498) is adequate to identify the name

Bassaris itea for the species treated here.

LIFE HISTORY.—The larval food plant is Urtica

incisa (Urticaceae).

Brief life-history notes were given by Hudson

(1928, p. 35), Barrett and Burnes (1951, p. 135 and

figures of larva and pupa, p. 134), and Gaskin

(1966, p. 88).

DISTRIBUTION.—This species is found in Australia,

Tasmania, Norfolk Island, New Zealand, Loyalty

Islands, and on Rapa in the Tabuai Islands. In

Australia it occurs in northeast Queensland near

Herberton and from MacKay, Queensland, south

to Victoria and west to Adelaide in South Australia

and also in the southwest part of Western Australia.

In New Zealand it is more widely distributed in

North Island than in South Island.

MATERIAL STUDIED.—Eighteen males and twentythree females were studied from the following localities: QUEENSLAND: Atherton (January) ; Brisbane.

NEW SOUTH WALES: Colo Vale (January). VICTORIA:

Newmarket (September, November); Sandringham.

NORFOLK ISLAND (October). NORTH ISLAND, NEW

ZEALAND: Haumoana, Hawkes Bay (February).

RAPA: Haurei (November).

2. Bassaris gonerilla (Fabricius), new combination

FIGURES 7, 23, 85-88

Papilio gonerilla Fabricius, 1775, p. 498; 1782, p. 82; 1793, p.

103.—Gabriel, 1927, p. 56.

Vanessa gonerilla.—Godart, 1819, p. 321.—Boisduval, 1832,

p. 121.—Hudson, 1883, pp. 217-219.—Dale, 1890, p. 150.—

Hudson, 1898, pp. 105-107, pi. 3: figs. 1, 2, 31, 32, pi. 12:

figs. 5, 6.—Quail, 1900, pp. 153-156.—Hudson, 1928, pp.

34-35, pi. 4: figs. 2, 9.—Williams, 1930, p. 231.—Hudson,

1939, p. 388.—Verity, 1950, p. 329.—Gaskin, 1966, pp. 87-

88, pi. 1: figs. [8], [9].—Pritchard, 1969, p. 12.

Pyrameis gonerilla.—Doubleday, 1849, p. 204.—Trimen, 1862-

1866, p. 118.—Butler, 1870, p. 78.—Kirby, 1871, p. 185.—

Staudinger, 1895, p. 98, pi. 37: fig. [d-1].—Distant, 1882-

1886, p. 173.—Dixey, 1890, pp. 94, 98, 100, 107, 112, 114,

118, 122, 123, 127, pi. 1: fig. 11, pi. 2: fig. 25.-Fruhstorfer,

1898b, p. 151.—Alfken, 1899, pp. 5-6; 1903, p. 602, pi. 32:

fig. 11.—Reuss, 1910c, pp. 85, 86, 88; 19l0d, p. 303.—Fruhstorfer, 1912a, p. 526, pi. 117: figs, D 3, D 4; 1912b, p. 526,

pi. 117: figs, D 3, D4 .

Papilio generilla Fabricius [misspelling of gonerilla], 1787,

p. 24.

MALE (Figure 85).—Upper surface of forewing

with apical half black and with an orange or orangered band crossing middle of wing from near costa

to a point at about the outer one third of anal vein;

this band outlined by black along its inner surface;

base of wing dark brown, heavily suffused with

bronze; with a subapical white bar opposite end of

cell and with a series of six white and blue submarginal spots between veins R3 and Cux; the latter

in interspaces R5, Mlf and M3 always blue, those in

interspace R3 and R4 sometimes blue, and that in

interspace M2 much larger than the rest and with

blue around its outer edge.

Upper surface of hindwing dark brown with a

large orange or orange-red submarginal bar enclosing four ocular black spots pupillated with blue;

these ocular markings located between veins Mt and

Cuj,, with the uppermost two of these spots (those

in interspaces Mx and M2) closer together and placed

more inward than those in interspaces M3 and Cux;

with a submarginal blue line or very thin bar

running between veins Cu2 and anal angle.

Under surface of forewing with orange or red bar

of upper surface repeated, replaced with white near

costa, and broader in interspaces Cux and Cu-j than

it is above; with a crescent-shaped yellow or white

subapical bar opposite end of cell and with a blue

ocular spot, pupillated with a large black spot between this bar and end of cell; with a blue marginal

stripe along outer margin of wing and a second

broad submarginal blue band or stripe between

vein M2 and the anal angle; with four rather indistinct ocular markings between veins R3 and M2,

the two above vein R5 obscure, very small, and

with a submarginal white spot faintly outlined by

blue along its outer margin in interspace M2.

Under surface of hindwing fuscous, sometimes

nearly black, with an overall purplish-blue cast,

heavily reticulated, and marked with lines and spots

of several shades of brown, black, and yellowish

white; with a series of five distinct and rather dark

submarginal ocular spots between veins Rg and Cu,.

A statement on the length of forewing cannot be

made for the species, since only the nominal subspecies is available for study (see paragraph on wing

length under Bassaris gonerilla gonerilla).

Male genitalia as illustrated by Figure 7 (= B.

gonerilla gonerilla drawn from my preparation no.

6192), with uncus and gnathos very much as in B.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

itea; aedeagus differing in having a simple pointed

tip; valva with dorsal margin having a distinct but

shallow lobe near middle; apex of valva with upward directed lobe that is not swollen along its

caudal margin, this lobe much smaller than in B.

itea; clasper a blunt subtriangular lobe and downward directed; cuiller not swollen at base, caudally

directed, evenly thick throughout, and lacking

teeth.

FEMALE (Figures 86, 87 ( $ ?), 88 ( $ ?)) .—Distinguishable from the male sex only in having the

apex of forewing and the hindwing on under surfaces pale brown to tan in color.

As in the male, a statement on the length of

forewing cannot be made (see above).

Female genitalia as illustrated by Figure 23 ( = B.

gonerilla gonerilla, drawn from my preparation no.

6194), with seventh sternite heavily sclerotized;

eighth sternite forming a large genital plate covering entire sternite, divided in middle of posterior

margin; ostium bursae large and from before middle

of eighth sternite; with deep lateral pouches on

either side of ostium bursae; signa ribbon-like, very

large and long, more than half the length of bursa

copulatrix.

GEOGRAPHICAL VARIATION.—Two subspecies are

recognized, but since specimens of B. gonerilla ida

Alfkin were not available for study, the characters

for separating the nominate subspecies from Alfkin's

subspecies are based upon the original description of

the latter.

2a. Bassaris gonerilla ida (Alfken), new

combination

FIGURES 87, 88

Pyrameis ida Alfken, 1899, pp. 5-9; 1903, p. 602, pi. 32: fig.

12.

Pyrameis gonerilla ida.—Fruhstorfer, 1912a, p. 526, pi. 117:

figs, D 1, D 2; 1912b, p. 526, pi. 117: figs, D 1, D 2.

Vanessa gonerilla form ida.—Hudson, 1928, p. 35, pi. 5: fig.

31.

Pyrameis ida variety argentata Alfken [excluded name, type

3], 1903, p. 602, pi. 32: fig. 13.

As mentioned above, since no specimens of B. gonerilla ida were available for study, I can only offer

statements comparing it with the subspecies B.

gonerilla gonerilla that are based upon the original

description of the former and other statements

from the literature.

Alfkin did not specify the sex of his ten specimens but I suspect that they were females because

of his subsequent colored illustration of the under

surface of ida (1903, pi. 32: fig. 12 and Figures 87,

88 in present work). In Alfkin's illustration the

under surface of forewing at apex and the under

surface of the hindwing are very pale in color and

thus quite similar to females of typical B. gonerilla.

If his material should prove to be entirely of the

female sex, I suspect that his name will probably

fall as a synonym of B. gonerilla.

Until Alfkin's material is found and studied or

until additional material from the type locality (Te

One and Maunganui, Island of Warekauri, Chatham Islands) is studied, I provisionally recognize

B. gonerilla ida.

In this subspecies the upper surfaces are said to

be more brownish black in color and the red more

yellowish, with the black border of the ocular spots

more diffused. The fringes are said to be shorter

and more indistinctly checkered than in B. gonerilla gonerilla.

On the under surface of the forewing the pattern

is said to be blurred, not distinct as in B. gonerilla

gonerilla. The two ocular spots standing directly

over the round white spots on the apex of forewing

are hardly distinguishable, whereas in B. gonerilla

gonerilla they are said to stand out clearly.

The under surfaces of the hindwing are said to

be almost uniformly red brown, sometimes indistinctly marbled, sometimes without pattern, and

the ocular spots are absent or if present are hardly

distinguishable. In B. gonerilla gonerilla, on the

other hand, the under surface of the hindwing is

said to be marbled with black and brown and in

a few places dusted with whitish, and four distinct

ocular spots are present.

Fruhstorfer (1912a, 1912b, p. 526) offers another

character for B. gonerilla ida by stating that it is

easily distinguished from B. gonerilla gonerilla by

the enlarged red area on the upper surface of the

hindwing (compare Figures 86, 87). If this statement holds true, this character may be the best way

to distinguish the two subspecies.

Hudson (1928, p. 35) states: "On the Chatham

Islands a form of Vanessa gonerilla occurs (Plate

V. fig. 31), in which the ground colour of the upper

side may be slightly bluer than usual, and the
NUMBER 84 33

sprinkling of golden scales on base of wings slightly

less pronounced. On the underside, the apex of the

fore-wings and the whole of the hind-wings are

strongly suffused with bright reddish-brown."

METHOD OF IDENTIFICATION.—As indicated above,

my statements on this subspecies are based entirely

upon the original description and subsequent descriptions and illustrations.

The location of the original series is unknown

to me.

LIFE HISTORY.—Nothing has been reported on the

immature stages or food plant on this subspecies.

An interesting observation reported by Hudson

(1928, p. 35) is: "Mr. C. Lindsay, who recently

collected quite a number of these butterflies at

the Chathams, informs me that they are much more

timid and harder to catch than the ordinary Vanessa

gonerilla, and that their manner of flight is somewhat different." It is hoped that observations of this

nature will be repeated and more thoroughly described and documented.

DISTRIBUTION.—This subspecies is known only

from the Chatham Islands, more than six hundred

miles east of New Zealand.

(Natural History). This specimen bears the type

number Rh. 8918.

The recognition of the subspecies B. gonerilla

gonerilla is dependent entirely upon the continued

recognition of B. gonerilla ida, and as explained

under the last named subspecies it is characterized

provisionally by characters mentioned in the original and subsequent descriptions of B. gonerilla ida.

LIFE HISTORY.—The larva feeds upon Urtica

incisa and Urtica ferox (Urticaceae).

Descriptions of the immature stages of life-history

notes are given by Hudson (1883, pp. 217-219; 1898,

pp. 106-107, pi. 3: figs. 1, 2, 31, 32 and 1929, pp.

34-35), Quail (1900, pp. 153-156), and Gaskin

(1966, p. 87).

DISTRIBUTION.—This subspecies is distributed

throughout most of New Zealand.

MATERIAL STUDIED.—Ten males and ten females

were available for study from the following New

Zealand localities: NORTH ISLAND: Haumoana,

Hawkes Bay (November, January); Little Bush,

Hawkes Bay (October). SOUTH ISLAND: Canterbury;

Christchurch; Lyttleton (March); Nelson (February).

2b. Bassaris gonerilla gonerilla (Fabricius)

FICURES 7, 23, 85, 86

Pyrameis gonerilla gonerilla.—Alfken, 1899, pp. 5-8; 1903, p.

602, pi. 32: fig. 11—Fruhstorfer, 1912a, p. 526, pi. 117: figs.

D 3, D 4; 1912b, p. 526, pi. 117: figs, D 3, D 4.

Vanessa gonerilla gonerilla.—Hudson, 1928, p. 35, pi. 4: figs.

2,9.

MALE (Figure 85) and FEMALE (Figure 86).—

For the supposed characters of this subspecies see

description above under B. gonerilla ida. This, the

nominate subspecies, is the basis for the above

description of the species.

Length of forewing in male sex, 28—31 mm (average 29.3 mm).

Length of forewing in female sex, 27-33 mm

(average 29.2 mm).

METHOD OF IDENTIFICATION.—The species B.

gonerilla received a rather more extensive original

description than most butterflies described by Fabricius (1775, p. 498) and was easily identified as the

New Zealand (= type locality) butterfly treated

herein. There is a single original male specimen, the

holotype, in the collection of the British Museum

Genus Cynthia Fabricius, resurrected genus

FIGURES 8-16, 24-32

Cynthia Fabricius, 1807, p. 281.—Westwood, 1840, p. 87 —

Humphreys and Westwood, 1841, pp. 56-57.—Stainton.

1857, pp. 36-37.—Humphreys, [1859], p. 21.—Scudder, 1875,

p. 152.—Holland, 1930, pp. 46-47.—Hemming, 1934, p. 70;

1967, p. 136.

Neopyrameis Scudder, 1889, pp. 434, 435, 440, 441, 456 —

Dixey, 1890, p. 94.—Hemming, 1967, p. 310.

Cyntia Costa Lima [misspelling of Cynthia], 1923, p. 150;

1928, p. 135.

Zynthia Hemming [misspelling of Cynthia], 1967, p. 93.—

Cowan, 1970, p. 70.

Type species: Papilio cardui Linnaeus. Type by reason of subsequent selection by Westwood (1840,

p. 87).

The generic name Vanessa (with type species P.

atalanta L.) was accorded precedence over Cynthia

(with type species P. cardui L.) in Opinion 156

(1944) of the International Commission on Zoological Nomenclature (see notes above under Vanessa).

In 1954 in its Direction 4 (same publication, no. 2,

pt. 53, pp. 629-652) the Commission endorsed the

name Cynthia Fabricius as a name "to be used by
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

any specialist who may consider that the type-species

of this genus is generically distinct from Papilio

atalanta Linnaeus, 1758, but not to be used in preference to the name Vanessa Fabricius, 1807." The

name Cynthia Fabricius was thereupon placed on

the Official List of Generic Names in Zoology as

Name No. 805.

Cynthia differs from Vanessa and Bassaris in the

habitus, in the structures of the male and female

genitalia, and in the paronychia and is here recognized as generically distinct.

GENITALIA.—Male (Figures 8-16) with uncus

(in dorsal or ventral view) not bifurcate distally;

lateral arms of gnathos sharply bent with a distinct

"heel" beyond or near middle of ventral margin,

so that the base is directed downward, and distal

one half or more is directed outward; with distal

part after this "heel" shorter than the base; aedeagus nearly straight or slightly undulate with distal

end pointed (not semi-spade shaped); valva with

clasper near middle, upwardly curved and directed

toward outer margin, or curved inward toward opposite valva.

Female (Figures 24-32) with genitalia consisting

of a central pod-shaped structure located on anterior

margin of eighth sternite, surrounding the ostium

bursae (except in C. carye and C. annabella, where

this structure is enlarged and located near posterior

margin); with two large lobelike lateral plates on

either side of ostium bursae; margin between seventh and eighth sternites undulate or sinuate and

greatly extended forward in middle at location of

ostium bursae (except in C. carye and C. annabella,

where this margin is nearly straight); with a pocket

on each side near end of these lateral plates; each

signum bursae long, much more than three times

as long as broad, and simple, not divided as in

Vanessa.

TARSI.—Paronychia of mid and hind tarsi each

consisting of a single slender lobe, not bifid, sometimes with a slight projection on its base.

HABITUS.—In Cynthia the upper surface of the

forewing is predominently tawny, sometimes pinkish

with fuscous restricted to the extreme base of wing,

to the apical area, to a number of fuscous spots in

the cell, and in interspaces Cut and Cu2; with middle of interspace Cut

 adjacent to anal vein tawny,

not fuscous; with a white or tawny subapical bar

opposite end of cell and a series of five or sometimes

six submarginal and subapical white spots in the

interspaces between costa and vein Cux or vein Cu2;

discal area of hindwing always light colored, usually

tawny, not fuscous as in Vanessa, and with a series

of two or more submarginal ocular spots, although

these may sometimes lack their blue centers and

may be nearly absorbed by a fuscous band (C.

myrinna). On the under surface of the forewing

Cynthia differs from Vanessa in lacking the blue

in the fuscous area opposite end of cell and between

there and the white subapical bar and in having

the base of the wing below cell much lighter in

color, with white or tawny near or at origin of vein

Cu2. On the under surfaces of the hindwing the

overall color of Cynthia is much paler, with large

amounts of white, many white lines and spots, and

often with a white to yellowish band across discal

area from costa to just before anal angle; the ocular

spots usually large, very distinct and round.

LIFE HISTORY.—Of the nine species of Cynthia,

some life-history information has been recorded for

all but C. altissima and C. terpsichore. Of the seven

species for which we have host records, all feed upon

various genera and species of either Compositae

(order Asterales) or Malvaceae (order Malvales).

The commonest species, C. cardui, also has been

reared from three genera of Boraginaceae and a

genus of Solanaceae (both families in the order

Polemoniales) and has even been reported feeding

upon Zornia (a genus of Leguminosae, order Rosales) and upon Urtica dioica (order Urticaceae).

I believe these last two records are open to question.

At any rate V. cardui has been recorded as feeding

upon twenty-one genera of plants, many ranging

over wide areas of the world, and this certainly

helps to explain its own nearly cosmopolitan distribution.

The unusual cephalically directed teethlike setae

found on the ovipositors of four of the five species

of the Virginiensis group of Cynthia and not on

species of Bassaris, Vanessa, or in the remaining

Cynthia species may be explained as structures

necessary as an aid in egg laying. These setae are

perhaps useful to those species which carefully tuck

their eggs under the hairs which cover the leaves

of their food plant so that these eggs are nearly

concealed under a thick mat of hairs. This concealment habit in egg laying was described by Scudder

(1889, p. 464) for C. virginiensis and is probably
NUMBER 84 35

true of the three other Cynthia species which have

these modified setae (C. altissima, C. braziliensis,

C. myrinna), although the egg-laying habits of these

species have not been studied.

DISTRIBUTION.—As just mentioned, V. cardui is

nearly cosmopolitan in distribution. The homeland

of C. kershawi is Australia. The remaining seven

species are entirely New World except for V. virginiensis, which has established itself also in the

Azores, Madeira, the Canary Islands, and the Hawaiian Islands.

List of Included Species

1. Cynthia cardui (Linnaeus), 1758

2. Cynthia kershawi (M'Coy), 1868

3. Cynthia virginiensis (Drury), 1773

4. Cynthia altissima (Rosenberg and Talbot),

1914

5. Cynthia braziliensis (Moore), 1883

6. Cynthia terpsichore (Philippi), 1859

7. Cynthia myrinna (Doubleday), 1849

8. Cynthia annabella, new species

9. Cynthia carye (Hubner), 1812

I arrange the nine species of Cynthia into three

species groups as follows: the cardui group, containing, besides that species, C. kershawaii; the virginiensis group, containing also C. altissima, C. braziliensis, C. terpsichore, and C. myrinna; and the carye

group, which also includes C. annabella. These

groups are not included in the following keys but

are defined later in heading paragraphs in the

treatment of the species.

Key to the Species of Cynthia

(based upon male genitalia)

1. Cuiller smooth 2

Cuiller armed with teeth 6

2. (1) Costal margin of valva nearly straight (Figure 8) 1. C. cardui (Linnaeus)

Costal margin of valva with large upward directed lobe 3

3. (2) Valva with large tooth or acute lobe at lower distal angle 4

Valva without large tooth or acute lobe at lower distal angle 5

4. (3) Cuiller upward directed; with large acute lobe at lower distal angle of valva (Figure

9) 2. C. kershawi (M'Coy)

Cuiller distally directed; with large tooth at lower distal angle of valva (Figure 12)

5. C. braziliensis (Moore)

5. (3) Lobe on costal margin of valva armed with large tooth on its inner surface (Figure

10) 3. C. virginiensis (Drury)

Lobe on costal margin of valva armed with a small tooth on its outer margin near

apex (Figure 13) 6. C. terpsichore (Philippi)

6. (1) Cuiller lightly armed with teeth along its upper edge only 7

Cuiller heavily armed with teeth along the outer half of its inner surface 8

7. (6) Valva with a tooth at apex of upward directed lobe on costal margin (Figure 14)

7. C. myrinna (Doubleday)

Valva with tooth missing at apex of this lobe, but with a large tooth near its base

(Figure 11) 4. C. altissima (Rosenberg and Talbot)

8. (6) Valva with a large club-shaped process at junction of dorsal and distal margins (Figure

16) 9. C. carye (Hubner)

Valva lacking this club-shaped process but with an acute lobe in this position (Figure

15) 8. C annabella, new species

Key to the Species of Cynthia

(based upon female genitalia and the ovipositor)

Ovipositor with all setae hairlike (Figures 24, 25, 29, 31, 32) 2

Ovipositor with at least a few setae short, toothlike, and cephalically directed (Figures

26-28, 30) 6
36 - SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Cynthia—Continued

(based upon female genitalia and the ovipositor)

2. (1) Ovipositor with some setae having apical portions bent at nearly right angles and with

bent portions directed cephalically (Figure 25) 2. C kershawi (M'Coy)

Ovipositor with setae normal, not as described above (Figures 24, 29, 31, 32) 3

3. (2) Opening of ductus bursae placed forward within lobes of seventh sternite 4

Opening of ductus bursae at end of podlike swelling in center of eighth sternite 5

4. (3) Heavily sclerotized lobes of eighth sternite greatly expanded caudally, with a pale central

area leading to opening of ductus bursae nearly same width throughout (Figure

29) 6. C. terpsichore (Philippi)

Heavily sclerotized lobes of eighth sternite not so greatly expanded caudally, with central

area next to opening of ductus bursae not as described above (Figure 24)

1. C. cardui (Linnaeus)

5. (3) Posterior margin of seventh sternite and anterior margin of eighth sternite forming an

even line across central one third or more of ventral surface; podlike area around

opening of ductus bursae without swollen base (Figure 33) 8. .C. armabella, new species

These margins more irregular; podlike area around opening of ductus bursae with base

greatly swollen (Figure 32) 9. C. carye (Hiibner)

6. (1) Eighth sternal plate broadly rounded along posterior margin (Figure 30)

7. C. myrinna (Doubleday)

Eighth sternal plate not broadly rounded along this margin 7

7. (6) Eighth sternal plate abruptly angled laterally (Figure 27)

4. C. altissima (Rosenberg and Talbot)

Eighth sternal plate tapering and becoming rather thin laterally 8

8. (7) With opening of ductus bursae placed deeply forward within the lobes of seventh sternite;

with large lateral pockets between seventh and eighth sternites (Figure 26)

3. C. virginiensis (Drury)

With opening of ductus bursae more shallowly placed within the lobes of seventh sternite;

with lateral pockets between seventh and eighth sternites small (Figure 28)

5. C. braiiltensis (Moore)

Key to the Species of Cynthia

(based upon habitus)

1. Hindwing below with two ocular spots, one each in interspaces Mi and Cui 2

Hindwing below with four or more ocular spots 6

2. (1) Hindwing above with fuscous band crossing middle of wing between anal area and

costal margin straight and broad, enclosing and obliterating submarginal ocular spots;

hindwing below with whitish or yellowish band crossing middle of wing from costa

to anal area straight (Figures 135, 136) 7. C. myrinna (Doubleday)

Hindwing above with this fuscous band narrow and wavy, not enclosing submarginal

ocular spots, sometimes entirely absent; hindwing below with this whitish or yellow

band projected outward along vein M3 3

3. (2) Hindwing above with fuscous band crossing middle of wing between anal area and

costal margin broken into small spots or absent (Figures 109-118)

3. C. virginiensis (Drury)

Hindwing above with this fuscous band not broken into small spots 4

4. (3) Forewing above with light-colored subapical bar just beyond end of cell tawny in color

(Figures 131-134) 6. C. terpsichore (Philippi)

Forewing above with this subapical bar white 5

5. (4) Forewing above with small apical and submarginal spot in interspace Mx white in color;

without distinct white band outlining inner surface of black bar at end of cell; with

fuscous band crossing middle of hindwing not projected outward in interspace M3

(Figures 119-122) 4. C. altissima (Rosenberg and Talbot)
NUMBER 84 37

Key to the Species of Cynthia—Continued

(based upon habitus)

Forewing above with this spot blue; with distinct white band outlining inner surface

of black bar at end of cell; fuscous band crossing middle of hindwing projected outward in interspace M3 (Figures 123-130) 5. C. braziliensis (Moore)

6. (1) Forewing above with light-colored subapical bar just beyond end of cell, white in color .7

Forewing above with this subapical bar tawny 8

7. (6) Hindwing above with five round, submarginal fuscous spots, located between veins R«

and Cu2, very rarely centered with blue; forewing with tawny ground color extended

into base of interspace M2 just beyond bar opposite end of cell (Figures 89-104)

1. C. cardui (Linnaeus)

Hindwing above with four round, submarginal fuscous spots, located between veins Mi

and Cu2, the three spots between veins M2 and Cu2 blue centered; forewing with

tawny ground color not extending into base of interspace M2 (Figures 105-108)

2. C. kershawi (M'Coy)

8. (6) Forewing above with tawny subapical bar just beyond cell slightly larger; forewing

below with yellowish-white postmedial and marginal bar slightly larger; bar below

base of Cu2 on the forewing above distinct; white spot in middle of wing opposite end

of cell on underside of hindwing triangular in shape (Figures 143-147; 155-160)

8. C. annabella, new species

Forewing above with tawny subapical bar just beyond cell smaller; forewing below with

yellowish-white postmedial and marginal bar slightly smaller; bar below base of Cu2

on forewing above very small or absent; white spot in middle of wing opposite end of

cell on underside of hindwing hour-glass in shape (Figures 137-142; 149-154)

9. C. carye (Hubner)

The Cardui Group

The cardui group contains two species, C. cardui

and C. kershawi. Here on the upper surface of the

forewing the fuscous color in interspace CIL,, found

just beyond the middle of anal vein, extends upward into interspace Cux but no farther. On the

upper surface of the hindwing the fuscous and

usually sinuous band crossing the wing from costa

to just above anal angle is broken along vein M2

or found only below that vein and represented by

a hooklike spot that extends upward from the basal

fuscous area. On the under surface of the forewing

the cardui group has most of the area behind end

of cell and between there and the fuscous spots

through middle of cell white and, as on the upper

surfaces, the fuscous color from interspace Cu2 just

beyond the middle of the anal vein does not extend

upward to vein M3 as it does in the virginiensis and

carye groups. On the under surface of the hindwing

there are always five small, submarginal ocular spots

and there is an isolated white spot, subtriangular

in shape, opposite end of cell just above vein M3.

The ovipositor setae are hairlike.

1. Cynthia cardui (Linnaeus), resurrected

combination

FIGURES 8, 24, 89-104

Papilio cardui (including the synonym belladonna) Linnaeus, 1758, p. 475 [Article 11 (d), International Code of

Zoological Nomenclature states that "a name first published as a synonym is not thereby made available." Hence

belladonna is not an available name.].—Fabricius, 1775, p.

499.—Esper, 1777, pp. 132-139, pi. 10: fig. 3.—Fabricius,

1787, p. 45; 1793, p. 103.—Godart, 1821, p. 102.

Vanessa cardui.—Hubner, [1819], p. 33.—Godart, 1819, pp.

323-324.—Boisduval, 1832, p. 119.—Boisduval and LeConte,

1834, pp. 178-179.—Boisduval, 1840, p. 21.—Herrich-Schaffer, 1843, pp. 38, 41.—Doubleday, 1844, p. 79.—Duponchel,

1844, p. 7.—Lucas, 1845, p. 57.—Noel, 1845, pi. 3.—Gray,

1846, p. 11.—Poey, 1847, p. 122.—Duponchel, 1849, pp. 107-

109, pi. 12: figs. 42 ac—Rambur, 1858, p. 15.—Berge, 1863,

p. 51. figs. 1 ac—Berce, 1867, p. 165, pi. 10: fig. 2.—

Wagner, 1870, pp. 169-171.—Staudinger, 1871, p. 17.—

Frey, 1880, p. 26.—De Waldeim and Eversmann, 1881, pp.

107-108, pi. 12: figs. 1, 2.—Kirby, 1882, p. 13, pi. 7: figs.

3 ad.—Kane, 1885, p. 62, pi. 9: fig. 1.—Jordan, 1886, pp.

84-85.—Pryer, 1888, p. 26; 1889, p. xii, pi. 7: fig. 2.— Scudder, 1888, pi. 2: fig. 1, pi. 67: fig. 7, pi. 83: figs. 60-62; 1889,

pp. 470-487, pi. 12: fig. 7, pi. 21: fig. 1, pi. 33: fig. 31, pi.

61: figs. 35, 54, pi. 64: fig. 30, pi. 74: fig. 37, pi. 81: fig. 10,

pi. 86: figs. 69-73—Dale, 1890, pp. 137-148.—Barrett, 1893,
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pp. 149-155, pi. 21: figs. 1, a-e.—Fischer, 1895, pp. 18, 22-

23, 24.—Schille, 1895, p. 218.—Standfuss, 18%, pp. 256-258,

pi. 7: fig. 5.—Aurivillius, 1898, p. 130.—Favre, 1899, p. 99.

—Meyrick, 1899, p. 194.—Moore, 1899-1900, pp. 105-107,

pi. 320: figs. 2 ab.—Dyar, 1903, p. 24.—Comstock and

Comstock, 1904, pp. 158-160, pi. 26: figs. 5, 6.—Bingham,

1905, pp. 365-366.—Kirkaldy, 1906, p. 139.—Bames and

McDunnough, 1917, p. 11.—Grinnell, 1918, p. 113 —

Lhomme, 1923, pp. 51-52—Antram, 1924, pp. 175-176.—

Comstock, 1927, pp. 133-134, pi. 43: figs. 8, 9.—Hudson,

1828, p. 36, pi. 4: fig. 1.—Williams, 1930, pp. 168-212 —

Holland, 1931, pp. 154-155, pi. 1: fig. 1, pi. 3: fig. 37, pi.

4: figs. 60-62.—Evans, 1932, p. 177.—Frohawk, 1934, pp. 16,

24, 25, 33, 140-147, pi. 11: figs. l-«.—Bates, 1935, p. 166.—

Peile, 1937, p. 139.—Davenport and Dethier, 1938, p. 164.

—McDunnough, 1938, p. 21.—Hudson, 1939, p. 388 —

Wild, 1939, p. 26, pi. 7: figs. 4, 5.—Field, 1940b, pp. 85-87,

274.—Hoffmann, 1940, p. 681.—Macy and Shepard, 1941,

pp. 125-126, pi. 4: fig. [7].—Pierce and Beirne, 1941, p.

20, pi. 3: fig. 2.—Comstock, 1944, pp. 450-451.—Stokoe,

1944, pp. 94-96, pi. 9: fig. 3, pi. 10: fig. 3, pi. 15: fig. 1.—

Ford, 1945, pp. 60, 76, 90, 154, pi. 2: fig. 2, pi. VIII: fig. 4,

pi. 25.—Leighton, 1946, p. 59.—Richards, 1946, pp. 21-22.

—Aubert, 1949, pp. 129, 133-135.—Paul and Robert, 1949,

pi. 17.—Pinhey, 1949, p. 83, pi. 12: fig. 3.—Williams, 1949a,

pp. 72-73; 1949b, p. 39.—Verity, 1950, pp. 329-334, pi. 51:

figs. 26-30, pi. 52: figs. 1-6, pi. 18: fig. 2, pi. 19: fig. 14.—

Woodhouse, 1950, pp. xiii, 49, pi. 15: fig. 1, pi. 37, fig. 4;

pi. 42: fig. 4.—Clark and Clark, 1951, pp. 14, 44, 201, pi.

4: figs, d, e.—Klots, 1951, pp. 107-108, 113, pi. 6: fig. 14,

pi. 14: fig. 10.—Agenjo, 1952, p. 283.—Peters, 1952, p. 76 —

Ferguson, 1954, p. 195.—Brown, 1955, p. 102, 2 figs —

Forster and Wohlfahrt, 1955, p. 56, pi. 15: fig. 13—Yokoyama, 1955, pp. 37, 96-97, pi. 29: fig. 89.—Lempke, 1956,

pp. 185-186.—Wiltshire, 1957, p. 31.—Wynter-Blyth, 1957,

pp. 210-212, pi. 38: fig. 8.—Zimmermann, 1959, pp. 453,

454, 455, 456, 457, 463, figs. 388, 389, 391, 396, 400.—Chalmers-Hunt, 1960-1961, pp. 60-62, 140.—Forbes, 1960, p. 159.

—Dos Passos, 1964, pp. 77.—Niculescu, 1965, pp. 186-191,

pi. 6: figs. 1, 2.—Gifford, 1965, p. 115.—Gaskin, 1966, pp.

88-89, pi. 1: fig. [5].—van Son, 1966, p. 66, fig. 1.—Dimock,

1968. p. 146.—Williams, 1970, pp. 157-175.

Cynthia cardui.—Humphreys and Westwood, 1841, pp. 54,

56-57, pi. 15: figs. 7-10.—Stainton, 1857, pp. 36-37, fig.—

Humphreys, [1859], pp. 21-22, pi. 11: figs. 1-4.—Morris,

1868, pp. 81-82, pi. 34.—Buckler, 1886, pp. 49-52, 174-176,

pi. 8: figs. 1 a-e.—Barnes and Benjamin, 1926, p. 15 —

Gunder, 1928, p. 202, pi. 8: figs. Sb, 3c; 1929, p. 9.

Pyrameis cardui.—Doubleday, 1849, p. 205.—Horsfield and

Moore, 1857, pp. 138-139, p. [3], pi. 5: figs. 3, 3a.—Lucas,

1857, pp. 543-544.—Morris, 1860, p. 8; 1862, pp. 59-60.—

Behr, 1864, pp. 125-126.—Bremer, 1864, p. 17.—Trimen,

1862-1866, pp. 119-121.—Butler, 1870, pp. 77-78.—Kirby,

1871, pp. 185-186.—Newman, 1874, pp. 64-66, fig. 17.—

Strecker, 1878, pp. 136-138, 190.—Moore, 1880-1881, pp.

50-51, pi. 27: figs. 1, la.—Godman and Salvin, 1882, pp.

217-218.—Saalmiiller, 1884, p. 77.—AlpWraky, 1885, p.

598.—Staudinger, 1885. p. 97.—Distant, 1882-1886, pp. 48,

91.—Mabille, 1886, pp. 126-127.—Marshall and de Niceville,

1886, pp. 227-228.—Shatz and Rober, 1886, pi. 16: fig. [9];

1887, p. 125.—Trimen, 1887, pp. 200-203.—Edwards, 1889,

p. 26.—Dixey, 1890, pp. 92, 93, 94, 96, 97, 98, 99, 101, 106,

109, 112, 115. 116, 117, 118, 122, 123, 126, 127, pi. 1: figs.

5, 8, pi. 2: fig. 24, pi. 3: fig. 38.—Maynard, 1891, p. 93,

pi. 5: fig. 5.—Leech, 1892-1894, pp. xv, 351— White, 1894,

pp. 54-55, 63, pi. 2: fig. 5.—Hagen, 1896, p. 165.—Skinner,

1898, pp. 24-25.—Tutt, 1896, pp. 350-355, fig. 37.—Denton,

1898, pp. 266-267.—Fruhstorfer, 1898b, p. 151.—Holland,

1898, pp. 170-171, pi. 1: fig. 1, pi. 3: fig. 37, pi. 4: figs.

60-62.—Miyajima, 1899, pp. 3, [118], pi. 11: fig. 2.—Cannaviello, 1900, pp. 18-19.—Staudinger and Rebel, 1901, p.

24.—Wright, 1905, pp. 177-178, pi. 22: fig. 229.—Kirkaldy,

1906, p. 139.—Schultz, 1906, p. 108.—Stichel, 1908, pp. 199-

200, pi. 62: fig. D 1.—Stichel, 1909, pp. 199-200, pi. 62: fig.

D 1.—Rebel, 1910, p. 20—Reuss, 1910a, pi. 1: figs. 1. 5.

9; 1910b, pp. 62, 63, 64, 65, 66, 67; 1910c, p. 90; 1910d. p.

303.—Fruhstorfer, 1912a, pp. 524-525; 1912b, p. 525.—

Aurivillius, 1913a, p. 227; 1913b, p. 227.—Verity, 1913, p.

181.—Seitz, 1914a, p. 458; 1914b, p. 458.—Waterhouse and

Lyell, 1914, p. 55.—Holland, 1915, p. 102, pi. 35.—Schultz,

1916, p. 27.—Verity, 1916b, p. 128.—Reuss, 1918, pp. 41-42.

—Ragusa, 1919, p. 146.—Holland, 1920. pp. 143-144.—

Stephan, 1923, p. 36.—Frohawk, [1924], pp. 151-157, pi. 26.

—Ormiston, 1924, p. 32.—Stephan, 1924, p. 25.—Verity and

Querci, 1924, p. 42— Bang-Haas, 1926, p. 56; 1927. p. 55;

1928, p. 26.—Schrader, 1928, pp. 68-70, pi. 6.-Haas, 1929,

p. 11.—Malsumura, 1929, pp. 9, 11, 19, pi. 5: fig. 5 —

Schrader, 1929, p. 8.—Bang-Haas, 1930, p. 162.—Gaede,

1930a, p. 200; 1930b, p. 200; 1931, p. 343.—Clark, 1932, pp.

13, 14, 22, 88-91, pi. 8: figs. 1-3.—Bryk, 1940, p. 17.—

South, 1941, pp. 78-81, fig. 24, pis: 44, 45.—Zerkowitz, 1946,

p. 246.—Barrett and Bames, 1951, pp. 132-133.^Swanepoel, 1953, pp. 213-215.—Paulian, 1956, pp. 67-69.

Pyrameis cardui cardui.—Fruhstorfer, 1912a, p. 524; 1912b,

p. 524.—Verity, 1919, p. 197.

Vanessa cardui cardui.—Field, 1940b, p. 85.—Corbet and

Pendlebury, 1956, p. 207, pi. 6: fig. 69, pi. 40: fig. 100.—

Okano and Ohkura, 1959, p. 42, pi. 41: fig. 124.

Papilio carduelis Seba, 1765, p. 6, pi. 1.—Cramer, 1775. pp.

40-41, pi. 26: figs, E, F; 1776, p. 152.—Doubleday, 1844, p.

79 [synonym of cardui]; 1849, p. 205.—Horsfield and Moore,

1857, p. 138— Lucas, 1857, p. 544.—Saalmuller, 1884, p.

77.—Mabille, 1886, p. 126.—Marshall and de Niceville.

1886, p. 227.—Leech, 1892-1894, p. 351.—Aurivillius. 1898.

p. 130.—Moore, 1899-1900, p. 105.

Vanessa carduelis.—Dyar, 1903, p. 24 [synonym of cardui].—

McDunnough, 1928, p. 21—Comstock, 1944, p. 450.—Dos

Passos, 1964, p. 77.

Cynthia carduelis.—Barnes and Benjamin, 1926, p. 15 [synonym of cardui].

Pyrameis cardui uniitersa form carduelis.—Verity, 1919, p.

198.

Pyrameis cardui variety carduelis.—Verity, 1920, p. 61.

Pyrameis cardui race carduelis.—Verity and Querci, 1924, p.

42.

Vanessa cardui form carduelis: Verity, 1936, p. 87; 1950, p.

334.
NUMBER 84 39

Vanessa cardui carduelis.—Field, 1940a, p. 342; 1940b, pp. 85,

86, 274.—Klots, 1951, p. 108.

Vanessa cardui aberration carduelis.—Verity, 1950, p. 330.

Papilio belladonna Godart, 1821, pp. 102-104, pi. 14: fig. 2.

Vanessa elymi Rambur, 1829, pp. 256-257, pi. 5: figs. 1, 2.—

Dyar, 1903, p. 24 [synonym of cardui].

Vanessa cardui aberration elymi.—Boisduval, 1840, p. 21.—

Duponchel, 1844, p. 7.—Staudinger, 1871, p. 17.—Jordan,

1886, pp. 84-85.—Scudder, 1889, p. 473.—Fischer, 1896, pp.

17-18, 57, 66, pi. 2: fig. 5 b.—Skinner, 1898, p. 25.—Fischer,

1903, p. 224; 1907, pp. 170, 201, figs. 5, 7—Barnes and

McDunnough, 1917, p. 11.—Lhomme, 1923, p. 51.—Comstock, 1927, p. 134.—Gunder, 1925, p. 198.—McDunnough,

1938, p. 21.—Field, 1940a, p. 87.—Leighton, 1946, p. 59 —

Verity, 1950, p. 332.—Chalmers-Hunt, 1960-1961, p. 61.—

Dos Passos, 1964, p. 77.

Pyrameis cardui variety elymi.—Kirby, 1871, p. 186.

Pyrameis cardui aberration elymi.—Strecker, 1878, p. 137.—

Tutt, 1896, p. 352.—Staudinger and Rebel, 1901, p. 24.—

Schultz, 1906, p. 108.—Stichel, 1908, p. 200; 1909, p. 200.

—Rebel, 1910, p. 20.—Reuss, 1910a, pp. 63, 65, 66.—Fischer,

1900, p. 5.

Vanessa cardui variety elymi.—Dale, 1890, p. 138.

Pyrameis cardui elymi.—Fruhstorfer, 1912a, p. 525.

Vanessa cardui elymi.—Fox, 1921, p. 46, pi. 2: fig. 3.

Cynthia cardui aberration elymi.—Barnes and Benjamin,

1926, p. 15.

Cynthia cardui transition form elymi.—Gunder, 1927, p. 133,

pi. 2: fig. 2a; 1927c, p. 270, pi. 9: figs. 2-4; 1929, p. 9.

Vanessa cardui form elymi.—Dimock, 1968, p. 146, fig. 1.

Pyrameis cardui variety leachiana Doubleday [nomen nudum],

1849, p. 205.

Pyrameis cardui aberration ate Strecker [excluded name, type

2], 1878, pp. 137-138.—Skinner, 1898, p. 25.

Vanessa cardui aberration ate.—Scudder, 1889, pp. 474-475.

—Barnes and McDunnough, 1917, p. 11.—McDunnough,

1938, p. 21.—Field, 1940b, p. 86.—Dos Passos, 1954, p. 77.

Cynthia cardui aberration ate.—Barnes and Benjamin, 1926,

p. 14.

Cynthia cardui transition form ate.—Gunder [excluded name,

type 2], 1927c, p. 270, pi. 9: fig. [6].

Vanessa cardui aberration pallens Noel [excluded name, type

2], 1881, p. 102.—Field, 1940b, p. 86.—Verity, 1950, p. 331.

—Niculescu, 1965, p. 190.

Pyrameis cardui aberration pallens.—Stichel, 1908, p. 199;

1909, p. 199.

Pyrameis cardui form pallens.—Seitz [excluded name, type

3], 1914a, p. 458; 1914b, p. 458.

Pyrameis cardui transition form pallens.—Gunder [excluded

name, type 2], 1927, p. 53.

Vanessa cardui variety pallida Sch0yen, 1881, pp. 77-79.

Pyrameis cardui aberration pallida.—Tutt, 1896, p. 352.—

Rebel, 1910, p. 20.—Pionneau, 1926, p. 4.

Pyrameis cardui variety pallida.—Staudinger and Rebel, 1901,

p. 24.—Schultz, 1906, p. 108.

Pyrameis cardui form pallida.—Stichel, 1908, p. 200; 1909,

p. 200.

Pyrameis cardui cardui form pallida.—Verity, 1919, p. 197.

Vanessa cardui aberration pallida.—Lhomme, 1923, p. 51.—

Verity, 1950, p. 330.—Niculescu, 1965, p. 190.

Pyrameis cardui race pallida.—Gaede, 1930a, p. 200; 1930b,

p. 200.

Vanessa cardui form pallida.—Field, 1940b, p. 84.—Lempke,

1956, p. 187.

Pyrameis kershawi M'Coy, Godman and Salvin not M'Coy

[a misidentification], 1882, p. 218.

Vanessa cardui aberration inornata Bramson [excluded name,

type 2], 1886, p. 284.—Scudder, 1889, pp. 473,474.—Frohawk,

1934, p. 147; 1938, pp. 84, 88, pi. 21: figs. 1, 2.—Field,

1940b, p. 86.—Verity, 1950, p. 331.—Chalmers-Hunt, 1960-

1961, p. 62.

Pyrameis cardui aberration inornata.—Tutt, 1896, p. 352.—

Staudinger and Rebel, 1901, p. 24.—Stichel, 1908, pp. 199-

200; 1909, pp. 199-200.—Rebel, 1910, p. 20.—Pionneau,

1926, p. 4.

Pyrameis cardui form inornata.—Seitz [excluded name, type

3], 1914a, p. 458; 1914b, p. 458.—Van Mellaerts, 1928, p.

105.

Pyrameis cardui variety inornata.—Stephan [excluded name,

type 3], 1924, p. 25.

Vanessa cardui form inornata.—Kuijken, 1967, pp. 224, 225.

Vanessa cardui aberration minor Failla-Tedaldi [excluded

name, type 2], 1887, pp. 70-71.—Verity, 1950, p. 330.

Pyrameis cardui aberration minor.—Ragusa, 1919, p. 146.

Pyrameis cardui universa form minor.—Verity [excluded

name, type 1 and type 2], 1919, p. 198.

Vanessa cardui carduelis form minor.—Field, 1940b, pp. 86,

274.

Vanessa cardui form minor.—Lempke, 1956, p. 187.

Vanessa cardui semisuffusa Cockerell [excluded name, type

2, as an individual variation], 1889, p. 54.

Vanessa cardui form semisuffusa.—Lempke [excluded name,

type 3], 1956, p. 187.

Vanessa cardui aberration semisuffusa.—Chalmers-Hunt [excluded name, type 2], 1960-1961, p. 62.

Vanessa cardui aberration wiskotti Standfuss [excluded name,

type 2], 1895, pp. 90-91; 1896, 392, pi. 7: fig. 6.—Verity,

1950, p. 332.

Pyrameis cardui variety wiskotti.—Fischer [excluded name,

type 3], 1900, p. 5.

Pyrameis cardui aberration wiskotti.—Fischer, 1901, p. 326.—

Schultz, 1906, p. 108.—Stichel, 1908, p. 200; 1909, p. 200 —

Reuss, 1910a, fig. 19; 1910b, p. 64; 1910c, p. 90.

Vanessa cardui warm variety wiskotti.—Fischer [excluded

name, type 3], 1903, p. 224.

Vanessa cardui cold variety wiskotti.—Fischer [excluded name,

type 3], p. 170.

Pyrameis cardui form minor Cockerell [excluded name, type

3 and homonym of minor Failla-Tedaldi], 1890, p. 57.

Pyrameis cardui aberration minor.—Skinner [excluded name,

type 2], 1898, p. 25.
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vanessa cardui aberration minor.—Barnes and McDunnough,

1917, p. 11.—McDunnough, 1938, p. 21.

Cynthia cardui aberration minor.—Barnes and Benjamin,

1926, p. 15.—Gunder, 1928, p. 202, pi. 8: fig. 3 a.

Pyrameis cardui aberration minor Cannaviello [excluded

name, type 2 and homonym of minor Failla-Tedaldi], 1900,

pp. 19-20.—Stichel, 1908, p. 199; 1909, p. 199.—Rebel, 1910,

p. 2O.--Stephan, 1923, p. 36; 1924, p. 25.

Pyrameis cardui variety minor.—Schultz [excluded name,

type 3], 1905, p. 67; 1906, p. 108.

Pyrameis cardui form minor.—Seitz [excluded name, type 3],

1914a, p. 458; 1914b, p. 458.—Gaede, 1930a, p. 200; 1930b,

p. 200.

Vanessa cardui aberration minor.—Lhomme, 1923, p. 51.—

Verity, 1950, p. 330.

Pyrameis cardui aberration carduelis Schultz [excluded name,

type 2 and homoym of carduelis Seba], 1906, p. 108.—

Stichel, 1908, p. 199; 1909, p. 199.—Rebel, 1910, p. 20.

Pyrameis cardui variety carduelis.—Stephan [excluded name,

type 3], 1924, p. 25.

Vanessa cardui aberration carduelis.—Verity, 1950, p. 331.

Pyrameis cardui aberration priameis Schultz [excluded name,

type 2], 1906, p. 108.—Stichel, 1908, p. 199; 1909, p. 199.—

Rebel, 1910, p. 20.—Van Mellaerts, 1928, p. 105.—Zerkowitz,

1946, p. 246.

Pyrameis cardui variety priameis.—Stephan [excluded name,

type 3], 1924, p. 25.

Vanessa cardui aberration priameis.—Verity, 1950, p. 332.—

Chalmers-Hunt, 1960-1961, p. 62.

Pyrameis cardui aberration carduelina Alpheraky [excluded

name, type 2], 1908, pp. 573-574.—Bang-Haas, 1926, p. 56.

Pyrameis cardui form carduelina.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration carduelina.—Field, 1940b, p. 86.—

Verity, 1950, p. 330.

Pyrameis cardui japonica Stichel [new synonymy], 1908, p.

200, pi: 62, fig. D 2; 1909, p. 200, pi. 62: fig. D 2.—Fruhstorfer, 1912a, p. 525; 1912b, p. 525.—Matsumura, 1929, p. 19.

—Cho, 1934, pi. 20: fig. 2.—Doi, 1934, p. 29.—Mori, 1934,

pp. 72-73.—Bryk, 1946, p. 39.

Pyrameis cardui universa form japonica.—Verity, 1919, p.

197.

Vanessa cardui japonica.—Field, 1940b, p. 85.

Vanessa cardui aberration japonica.—Verity, 1950, p. 331.

Vanessa cardui form japonica.—Lempke, 1956, p. 188.

Pyrameis cardui aberration ocellata Rebel [excluded name,

type 2], 1910, p. 20.—Reuss, 1910b, pp. 63, 64.—Bang-Haas,

1926, p. 56.

Pyrameis cardui variety ocellata.—Wize [excluded name, type

3], 1917, p. 5.—Stephan, 1924, p. 25.

Pyrameis cardui form ocellata.—Gaede [excluded name, type

3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration ocellata.—Field, 1940b, p. 86.—

Verity, 1950, p. 331.

Vanessa cardui form ocellata.—Lempke, 1956, p. 187.

Pyrameis cardui aberration carnea Fritsch [excluded name,

type 2], 1912, p. 136.—Bang-Haas, 1926, p. 56.

Pyrameis cardui form carnea.—Reuss [excluded name, type

3], 1918, p. 41.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui variety carnea.—Stephan [excluded name,

type 3], 1924, p. 25.

Vanessa cardui aberration carnea.—Field, 1940b, p. 86.—

Verity, 1950, p. 330.

Vanessa cardui form carnea.—Lempke, 1956, p. 187.

Pyrameis cardui aberration melanosa Cabeau [excluded name,

type 2], 1913, p. 43.—Bang-Haas, 1926, p. 56.

Vanessa cardui aberration melanosa Lhomme, 1923, p. 51.—

Verity, 1950, p. 332.

Pyrameis cardui form melanosa.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui form melanosa.—Lempke, 1956, p. 187.

Pyrameis cardui form brunneaalbimaculata Reuss [excluded

name, type 3], 1916, pp. 130-131.—Bang-Haas, 1926, p.

56.—Gaede, 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration brunneaalbimaculata.—Field [excluded name, type 2], 1940b, p. 86.—Verity, 1950, p. 331.

Pyrameis cardui form rosacea Reuss [excluded name, type

3], 1916, p. 131; 1918, p. 41.—Bang-Haas, 1926, p. 56.—

Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui variety roseacea.—Stephan [excluded name,

type 3], 1924, p. 25.

Vanessa cardui aberration rosacea.—Field [excluded name,

type 2], 1940b, p. 86.—Verity, 1950, p. 330.

Pyrameis cardui form hunteri Reuss [excluded name, type

3], 1918, p. 41.

Vanessa cardui form hunteri.—Lempke, 1956, p. 187.

Pyrameis cardui form ochracea Reuss [excluded name, type

3], 1918, p. 41.

Vanessa cardui form ochracea.—Lempke, 1956, p. 187.

Pyrameis cardui form conjuncta Verity [excluded name, type

1 and type 3], 1919, p. 197.

Pyrameis cardui form conjuncta.—Bang-Haas [excluded

name, type 3], 1927, p. 55.—Gaede, 1930a, p. 200; 1930b.

p. 200.

Vanessa cardui cardui aberration conjuncta.—Field [excluded

name, type 1 and type 2], 1940b, p. 85.

Vanessa cardui aberration conjuncta.—Verity [excluded name.

type 2], 1950, pp. 331, 336.—Niculescu, 1965, p. 190.

Pyrameis cardui aberration emielymi Verity [excluded name.

type 2], 1919, p. 198.—Bang-Haas, 1927, p. 55.

Pyrameis cardui form emielymi.—Gaede [excluded name.

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration emielymi.—Field, 1940b, p. 86.—

Verity, 1950, p. 331.—Chalmers-Hunt, 1960-1961, p. 62.

Vanessa cardui form emielymi.—Lempke, 1956, p. 189.

Pyrameis cardui variety infrabrunnea Verity [excluded name,

type 3], 1919, p. 198.

Pyrameis cardui inops form infrabrunnea.—Bang-Haas [excluded name, type 1 and type 3], 1927, p. 56.
NUMBER 84 41

Pyrameis cardui form infrabrunnea.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration infrabrunnea.—Verity [excluded

name, type 2], 1950, p. 331.

Pyrameis cardui form infraflava Verity [excluded name, type

3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infraflava.—Bang-Haas, 1927, p.

56.

Vanessa cardui aberration infraflava.—Verity [excluded name,

type 2], 1950, p. 331.

Pyrameis cardui form infragrisea Verity [excluded name, type

3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infragrisea.—Bang-Haas, 1927, p.

56.

Vanessa cardui aberration infragrisea.—Verity [excluded

name, type 2], 1950, p. 331.

Pyrameis cardui form infranigrans Verity [excluded name,

type 3], 1919. p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infranigrans.—Bang-Haas, 1927,

p. 55.

Vanessa cardui aberration infranigrans.—Verity [excluded

name, type 2], 1950, p. 331.

Pyrameis cardui form infraochracea Verity [excluded name,

type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infraochracea.—Bang-Haas, 1927,

p. 56.

Vanessa cardui aberration infraochracea.—Verity [excluded

name, type 2], 1950, pp. 331, 334.

Pyrameis cardui universa form inops Verity [excluded name,

type 1], 1919, p. 198.

Pyrameis cardui race inops Verity and Querci, [new synonymy], 1924, p. 42.—Bang-Haas, 1927, p. 55.—Gaede, 1930a,

p. 200; 1930b, p. 200.

Pyrameis cardui form inops.—Verity, 1936, p. 87.

Vanessa cardui inops.—Field, 1940b, p. 85.

Vanessa cardui aberration inops.—Verity, 1950, pp. 331, 334.

Vanessa cardui form inops.—Lempke, 1956, pp. 186-187.

Pyrameis cardui form sexiespupillata Verity [excluded name,

type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui aberration sexiespupillata.—Bang-Haas [excluded name, type 2], 1927, p. 55.—Van Mellaerts, 1928,

pp. 104-105.

Vanessa cardui aberration sexiespupillata.—Field, 1940b, p.

86.—Verity, 1950, p. 330.—Chalmers-Hunt, 1960-1961, p. 62.

Vanessa cardui carduelis aberration sexiespupillata.—Field

[excluded name, type 1], 1940b, p. 274.

Vanessa cardui form sexiespupillata.—Lempke, 1956, p. 188.

Pyrameis cardui form septiespupillata Verity [excluded name,

type, 3], 1919, p. 198.—Gaede, 1950a, p. 200; 1950b, p. 200.

Pyrameis cardui aberration septiespupillata.—Bang-Haas [excluded name, type 2], 1927, p. 55.—Van Mellaerts, 1928,

p. 104.

Vanessa cardui aberration septiespupillata.—Field, 1940b, p.

86.—Verity, 1950, p. 331.

Vanessa cardui form septiespupillata.—Lempke, 1956, p. 188.

Pyrameis cardui race universa Verity, [new synonymy], 1919,

p. 197; 1920, p. 61.—Verity and Querci, 1924, p. 42 —

Bang-Haas, 1926, p. 56.—Gaede, 1930a, p. 200; 1930b, p.

200.

Vanessa cardui form universa.—Verity, 1950, p. 334.—Lempke,

1956, p. 186.

Vanessa cardui race universa.—Verity, 1936, p. 87; 1937, p.

22.—Field, 1940b, p. 85.

Pyrameis cardui aberration maria Stephan [excluded name,

type 2], 1924, p. 25.

Pyrameis cardui aberration marthamaria Stephen [excluded

name, type 2], 1924, p. 25.—Bang-Haas, 1927, p. 55.

Vanessa cardui aberration marthamaria.—Field, 1940b, p.

86.—Verity, 1950, p. 331.

Pyrameis cardui aberration schoenfellneri Hoffman [excluded

name, type 2], 1924, p. 25.—Bang-Haas, 1927, p. 55.

Vanessa cardui aberration marthamaria.—Field, 1940b, p.

86.—Verity, 1950, p. 331.

Pyrameis cardui aberration schoenfellneri Hoffman [excluded

name, type 2], 1925, pp. 29-30.

Pyrameis cardui litoralis aberration schoenfellneri.—BangHaas [excluded name, type 1], 1927, p. 56.

Pyrameis cardui form schoenfellneri.—Gaede [excluded name,

type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration schoenfellneri.—Field, 1940b, p.

86.—Verity, 1950, p. 332.

Pyrameis cardui aberration subfracta Stack [excluded name,

type 2], 1925, p. 114, fig. 3.—Bang-Haas, 1928, p. 26.

Pyrameis cardui litoralis aberration subfracta.—Bang-Haas

[excluded name, types 1 and 2], 1927, p. 56.

Pyrameis cardui form subfracta.—Gaede [excluded name, type

3], 1930a, p. 200; p. 1930b, p. 200.

Vanessa cardui aberration subfracta.—Field, 1940b, p. 86.—

Verity, 1950, p. 330.—Niculescu, 1965, p. 190.

Vanessa cardui form subfracta.—Lempke, 1956, p. 188.

Pyrameis cardui aberration takesakiana Kato [excluded name,

type 2], 1925, pp. 145-146, fig.

Vanessa cardui aberration takesakiana.—Field, 1940b, p. 86.

Pyrameis cardui litoralis de Souza [new synonymy], 1926, p.

3.—Bang-Haas, 1927, p. 56.

Pyrameis cardui aberration rosea Pionneau [excluded name,

type 2], 1926, p. 4.—Bang-Haas, 1927, p. 55.

Pyrameis cardui form rosea.—Gaede [excluded name, type 3],

1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration rosea.—Field, 1940b, p. 86.

Cynthia cardui jacksoni Clark [as an individual variant, excluded name, type 2], 1927, p. 127.

Pyrameis cardui form jacksoni.—Clark [excluded name, type

3], 1932, p. 89, pi. 8: figs. 2, 3.

Vanessa cardui carduelis aberration jacksoni.—Field [excluded

name, type 1], 1940b, pp. 86, 87, 274.

Vanessa cardui form jacksoni.—Lempke, 1956, p. 187.—Dos

Passos, 1964, p. 77.

Pyrameis cardui aberration browni Meilhan [excluded name,

type 2], 1928, pp. 102-103.
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pyrameis cardui form browni.—Gaede [excluded name, type

3], 1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration browni.—Verity, 1950, p. 331.

Pyrameis cardui aberration flava Bandermann [excluded

name, type 2], 1928, pp. 236-237.—Bang-Haas, 1929, p. 11.

Pyrameis cardui form flava.—Gaede [excluded name, type

3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration flava.—Field, 1940b, p. 86.

Pyrameis cardui aberration rogeri Meilhan [excluded name,

type 2], 1928, pp. 100-103.—Bang-Haas, 1930, p. 162.

Pyrameis cardui subspecies rogeri.—Gaede [new synonymy],

1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration rogeri.—Field, 1940b, p. 86.—

Verity, 1950, p. 331.—Chalmers-Hunt, 1960-1961, p. 62.

Pyrameis cardui aberration varini Meilhan [excluded name,

type 2], 1928, p. 103.—Bang-Haas, 1930, p. 162.

Pyrameis cardui form varini.—Gaede [excluded name, type

3], 1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration varini.—Field, 1940b, p. 86.—

Verity, 1950, p. 331.—Chalmers-Hunt, 1960-1961, p. 62.

Vanessa cardui form varini.—Lempke, 1956, p. 189.

Pyrameis cardui form hemielymi Gaede [excluded name, type

3], 1930a, pi. 13: fig. B 1; 1930b, pi. 13: fig. B 1.

Pyrameis cardui aberration johni Fischer [excluded name,

type 2], 1932, p. 158, fig.

Vanessa cardui aberration johni.—Field, 1940b, p. 86.

Vanessa cardui aberration pallida Frohawk [excluded name,

type 2 and homonym of pallida Sch0yen]. 1938, p. 84.

Vanessa cardui ushuwaia Bryk [new synonymy], 1944, pp. 10-

11, pi. 1: fig. 6; 1946, p. 39; 1953, p. 93.

Vanessa cardui japonica form koreana Bryk [excluded name,

types 1, 2, 3], 1946, p. 39.

Vanessa cardui aberration albicans Verity [excluded name,

type 2], 1950, p. 331.

Vanessa cardui aberration infralutea Verity [excluded name,

type 2], 1950, p. 334.

Vanessa cardui form albipuncta Lempke [excluded name,

type 3], 1956, p. 188.

Vanessa cardui form nigripuncta Lempke [excluded name,

type 3], 1956, pp. 188-189.

This species, probably one of the two or three best

known and most described and illustrated species

of butterflies in the world, hardly needs redescription except to compare it to the other species of the

genus that are not nearly as well known. C. cardui,

as mentioned earlier in the group description, differs

from all other species of Cynthia, except C. kershawi, in having the fuscous color in interspace Cu2

on upper surface of forewing not extending to vein

Cuj. On the upper surface of the hindwing it differs

from C. kershawi in having five, not four, submarginal ocular spots, these located between veins

Rs and Cu2, and all lacking blue centers except for

a rare individual variant that sometimes occurs

with a blue center in the spot located in interspace

Cur

MALE (Figures 89-96).—Forewing above with

subapical white bar just beyond cell, and subapical

and submarginal white spots very much as in C.

kershawi and quite similar to those of the other

Cynthia species except that the white spot found in

interspace Cux in the species of the virginiensis

group is lacking in C. cardui and C. kershawi; with

fuscous bar in interspace Cuj connecting apical and

subapical fuscous areas to central fuscous spot in

interspace Cu2 lacking; with the central fuscous

spot in interspace Cu2 just mentioned usually not

so strongly connected to the fuscous spot at base

and above base of Cu2 as it is in C. kershawi and

with the fuscous spots in middle of cell distinctly

separated, not nearly confluent as is true of C.

kershawi.

Hindwing on the upper surface as mentioned

before with five, not four, submarginal ocular spots

between vein Rg and Cu2, none with blue centers

except very rarely the one in interspace Cux; with

an upwardly projected hooklike fuscous marking

below and beyond cell, all that remains of the

fuscous and usually sinuous band that crosses the

wing from costa to just above anal angle in the

species of the virginiensis group, this hooklike

fuscous spot nearly or sometimes entirely joining

a downward projected fuscous streak from the

fuscous area of the costal margin and base of wing.

Under surfaces of forewing with fuscous of central

area of interspace Cu2 extending upward nearly,

or at least halfway, through interspace Cu2 and with

two fuscous spots in middle of cell just inward from

the white area distinctly separated from each other.

Hindwing on the under surface with a lighter

overall appearance than in C. kershawi, with the

submarginal ocular markings distinctly larger than

in that species, and with the subtriangular white

spot at the end of the cell in interspace M2 also

much larger.

Length of forewing, 22-39.5 mm (average 29.3

mm).

Male genitalia as illustrated by Figure 8 (drawn
NUMBER 84

from my preparation no. 6104), with uncus tapered

distally to a point; gnathos longer, with uncus with

distal part after "heel" nearly at right angles to the

base; valva with costal margin broadly projected

upward, forming a broad obtuse angle with a small

upward projecting spine at its summit just below

uncus; distal margin of valva also broadly projected

and terminating in one or two spines; lower margin

of valva broadly convex and continuous with distal

margin, lacking completely the sickle-shaped projection found in C. kershawi; clasper directed

toward outer margin and curved inward at its

extremity; cuiller smooth with free part curved in

a shallow figure S toward distal margin; aedeagus

very similar to that of C. myrinna, with base slightly

bent downward.

FEMALE (Figures 97-104).—This sex hardly

differs from the male except that most females have

a slightly more rounded outer margin on the hindwing and have the outer margin of forewing near

vein Cu2 slightly more produced.

Length of forewing, 24-37 mm (average 30.9

mm).

Female genitalia as illustrated by Figure 24

(drawn from my preparation no. 3675), with anterior margin of eighth and posterior margin of

seventh sternite greatly undulated and with these

sternites forming two large posteriorly directed

lobes; ostium bursae swollen and located in eighth

sternite between lobes of seventh sternite; with a

pair of laterally placed pockets between seventh

and eighth sternites; ribbonlike signa extending

nearly full length of bursa copulatrix.

INDIVIDUAL AND ABERRATIONAL VARIATION.—Individual variation in this species commonly consists

of differences in the ground color of the upper

surfaces, which varies from a pale tan to a salmon

red. The fuscous markings vary in extent and sometimes there are blue centers in the lowermost submarginal round spots of the upper surfaces of the

hindwings. Aberrational variation has been studied

by numerous persons who unfortunately have given

some forty-nine formal names to individuals displaying this "more rare" type of individual variation. Specimens having an extra white spot, or

lacking one, or displaying any slight variation from

the "normal" have received names. Names have

usually always been given also to any remarkably

different looking specimen. All of these names are

synonymized above or are excluded from nomenclature. Gaede (1930a, 1930b, p. 200), Verity (1950,

pp. 330-332), and Lempke (1956, pp. 186-189)

treat many of these names and should be consulted

by those interested.

SEASONAL VARIATION.—Variation in habitus between the three broods (where there are three

broods) does not seem to occur in this species.

There are "wet" and "dry" forms in material collected in microenvironments that would be thought

of (at least temporarily) as wet and dry areas.

Where there is sufficient rain, specimens occur that

are larger, brighter, and more pinkish above and

darker on the under surfaces.

GEOGRAPHICAL VARIATION.—Verity (1919, p. 197;

1920, p. 21; 1950, p. 334), and Verity and Querci

(1924, p. 42) have maintained that there is a great

deal of geographical variation and have recognized

several subspecies as Vanessa cardui universa, V.

cardui inops, V. cardui carduelis, and V. cardui

cardui. Stichel (1908, p. 200) describes what he

considers to be a subspecies as Pyrameis cardui

japonica. None of these hold up as subspecies. Of

the more than two hundred and fifty specimens examined, the characters supposedly differentiating

these subspecies occur without relation to geography, so all of these proposed subspecies are

synonymized above.

METHOD OF IDENTIFICATION.—The original description of Linnaeus (1758, p. 475) and his references to previous descriptions and illustrations

clearly identify his name with the species here

treated as Cynthia cardui.

Since Linnaeus in his original description refers

to his 1746 work "Fauna Svecica," I consider Sweden

to be the type locality of C. cardui. Verity (1913,

p. 181), in his study of the Linnean types, states of

cardui: "There is nothing noteworthy about the

one typical specimen." This specimen is in the

Linnean collection of the Linnean Society, London.

SYNONYMY.—See synonymy and literature citations above. In addition to the names proposed for

so-called subspecies, which are synonymized above,

Papilio belladonna Linnaeus, a name used earlier

by Linnaeus (and therefore not available), is listed

by him as a synonym (1758, p. 485). Vanessa elymi

Rambur (1829, pp. 256-257), described as a species,

was based upon an aberrant specimen of C. cardui

and is a synonym.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

LIFE HISTORY.—The food plants of the larvae

are principally members of the Compositae and

most especially the various thistles. Among its recorded hosts are: Anaphalis margaritaceae, Arctium

leppa, Artemisia vulgaris, Blumea, Carduus nutans,

C. crispus, C. acanthoides, Centaurea benedicta,

Cirsium, Cnicus lanceatatus, C. acaulis, Filago arvenis, Gnaphalium indicum, Lappa officinalis,

Senecio cineraria, Silybum marianum, and Xanthium, all members of the Compositae (order Asterales). Occasional hosts are: Althaea officinalia,

Malva sylvestris, and M. rotundifolia of the Malvaceae (order Malvales); Anchusa officinales, Borago officinalis, and Echium vulgare of the Boraginaceae (order Polemoniales), and Nicotiana glauca

(Solanaceae, order Polemoniales). It has even been

reported upon Zornia (Leguminosae, order Rosales) and upon Urtica dioica (Urticaceae) ! I believe these last two records are open to question.

Fabricius (1775, p. 499) gave brief descriptions

of the larva and pupa. Life-history notes and descriptions of the immature stages are numerous.

Edwards (1889, p. 26) and Davenport and Dethier

(1938, p. 164) listed many of the references. Some

other references are: Esper (1777, p. 136), Latreille (1824, p. 325), Boisduval and LeConte

(1834, p. 179), Duponchel (1849, pp. 107-109, pi.

12: figs. 42 a-c), Morris (1862, p. 60), Trimen

(1862-1866, pp. 120-121), Newman (1874, p. 64),

Buckley (1886, pp. 49-52, 174-176, pi. 8: figs. 1,

lfl-e), Marshall and de' Nic^ville (1886, pp. 227-

228), Tutt (1886, pp. 352-354), Trimen (1887, p.

202), Dale (1890, pp. 138-140), Barrett (1893, pp.

151-154), Moore (1899-1900, p. 106), Stichel

(1909, p. 200), Frohawk (1934, pp. 143-146, pi. 11:

figs. 1^), Stokoe (1944, pp. 94-96, pi. 9: fig. 3, pi.

10: fig. 3, pi. 15: fig. 1), Woodhouse (1950, p. 49),

and Wynter-Blyth (1957, pp. 210-211).

DISTRIBUTION.—This butterfly is the most cosmopolitan of all butterflies being found in most parts

of the world except Australia, New Zealand, and

only rarely in South America south of Venezuela.

It is absent in the Antartic and the Artie regions. Its migrations have attracted attention for

nearly two hundred years and much of this information has been summarized by Williams (1930,

pp. 168-212).

MATERIAL STUDIED.—One hundred and thirtyeight males and one hundred and twenty-one females were studied from the following localities.

NORTH AMERICA: BRITISH COLUMBIA: Duncans;

Vancouver (August); Victoria (August). ALBERTA:

Calgary (June). MANITOBA: Manchester (July).

ONTARIO: Cedarhurst (September); Hymers.

MAINE: Mount Desert Island (July). MASSACHUSETTS: Essex (July). NEW YORK: Allegany State

Park; Staten Island (September). MARYLAND: College Park (August); Prince Frederick County October) . DISTRICT OF COLUMBIA: (June, August).

VIRGINIA: Bald Mountain (August) ; Limeton (October) ; Salem (July, August, September, October).

NORTH CAROLINA: Asheville (July). SOUTH CAROLINA: Sullivans Island. NORTH DAKOTA: Tower City

(August). MINNESOTA: NO specific locality. MICHIGAN: Chelsey (October) ; Detroit (July); Saint Ignace (July). ILLINOIS: Decatur (September). MISSOURI: Webster Groves (September). NEBRASKA:

Lincoln (September). KANSAS: Concordia (June);

Eureka (April, June, November); Lawrence

(June); Scandia (June). KENTUCKY: Frankfort

(September). LOUISIANA: Baton Rouge (December) . TEXAS: Brownsville; San Benito (May).

WASHINGTON: Brewster (June, July); Cooney Lake

(July); Godman Springs, Blue Mountains (July);

Yakima (October). UTAH: Stockton (June).

WYOMING: Fremont County; Yellowstone Park.

COLORADO: Golden (June). NEW MEXICO: Mount

Tasco (August). ARIZONA: Huachuca Mountains

(September); Palmerlee, Cochise County; Readington. CALIFORNIA: LOS Angeles; San Diego (June.

September); Taft (May). HAWAII: Oahu (March).

MEXICO: Aguascalientes (August); Jalapa; Mazatlan, Sinaloa; Mexico City (June) ; Orizaba (Novvember). GUATEMALA: San Sebastian (June, 3,000

ft.). HONDURAS: NO specific locality. COSTA RICA:

San Jose (January, 4,000 ft.). COSTA RICA: San Jose

(January, 4,000 ft). PANAMA: NO specific locality.

CUBA: Loma del Gato, Sierre del Cobre, Oriente

(September, 2,600 ft); Vinales (June). HAITI:

Beaumont (October).

EUROPE: FRANCE: Royan (October). SWITZERLAND: Zurich. GERMANY: Miesbach. CZECHOSLOVAKIA: Carpathian Mountains; Friedland. HUNGARY: Budapest (July); Galgamacsa (August).

ALBANIA: NO specific locality. GREECE: Athens

(May); Spetsai Island (July, August).

ASIA: TURKEY: NO specific locality. SYRIA:
NUMBER 84 45

Beyrout. JORDAN: Jordan Valley (March). INDIA:

Kasmir; Khasia Hills; Madras; Sikkim (May). BHUTAN: Thimbu (September, 7,700 ft). CHINA: FOOchow; Ishang; near Mupin (14,000 ft) ; Mount

Omei, Szechuan (August); Ts'ao P'o, Szechuan

(August, 6,300 ft); Tschili Province (September).

JAPAN: Kotanijochi (July); Musaki (October);

Oeda (September); Oita (September); Sifu; Tokosanchi (June); Tosari (July). JAVA: Buitenzorg.

SUMATRA: Deli.

AFRICA: UGANDA: NO specific locality. KENYA:

Kibwezi (December); Nairobi (May); Thomsons

Falls (January). RHODESIA: Sinoia. NYASALAND: NO

specific locality. MADAGASCAR: NO specific locality.

2. Cynthia kershawi M'Coy, resurrected

combination

FIGURES 9, 25, 105-108

Cynthia kershawi M'Coy, 1868, p. 76.

Pyrameis kershawi.—Godman and Salvin, 1882, p. 218.—

Dixey, 1890, pp. US, 117, 122, 123.—Waterhouse, 1903, p.

12.—Clark, 1932, p. 91.

Pyrameis cardui kershawi.—Olliff, 1888, pp. 1251-1252.—Fruhstorfer, 1912a, p. 525; 1912b, p. 525.—Reuss, 1910a, pp.

63, 66.—Waterhouse and Lyell, 1914, p. 56, pi. 5: fig. 36.—

Barrett and Burnes, 1951, p. 134.

Pyrameis cardui form kershawi.—Seitz, 1914a, p. 458; 1914b,

p. 458.

Vanessa cardui kershawi.—Williams, 1930, pp. 204-205; 1949a,

p. 73; 1949b, p. 40.—Burns, 1969, p. 80, fig. 58.

Vanessa kershawi.—Field, 1940b, p. 85.—Smithers and Peters,

1966, pp. 67-68.-van Son, 1966, p. 66, fig. 2.—Fox, 1969,

pp. 7, 9, 10.—Pritchard, 1969, pp. 11-12.

Pyrameis cardui kershawii Kirby [misspelling or unjustified

emendation of kershawi], 1871, p. 186.—Semper, 1878, p.

148.—Marshall and de Niceville, 1886, p. 226.—Fruhstorfer,

1898a, p. 151.

Pyrameis kershawii.—Miskin, 1891, p. 39.—Verity, 1916b, p.

128.—Bodley, 1920, pp. 74-75.

Vanessa cardui kershawii.—Dale, 1890, p. 138.—Gaskin, 1966,

p. 88.

Vanessa kershawii.—Niculescu, 1965, p. 191.

Pyrameis lucasii Miskin [new synonymy], 1888, pp. 1515-

1516; 1891, p. 39.

Pyrameis itea lucasii.—Waterhouse, 1903, p. 12.

Pyrameis itea lucasi Fruhstorfer [misspelling or unjustified

emendation of lucasii], 1912a, p. 526; 1912b, p. 526.

Pyrameis cardui kershawi aberration lucasi.—Waterhouse and

Lyell, 1914, p. 56.

Pyrameis cardui aberration lucasi.—Fruhstorfer, 1927a, p.

1112; 1927b, p. 1112.

Pyrameis cardui kershawii aberration suffusa Olliff [excluded

name, type 2], 1888, pp. 1250-1252.—Waterhouse and

Lyell, 1914, p. 56.

Pyrameis kershawii suffusa Miskin [new synonymy], 1891, p.

39.—Waterhouse, 1903, p. 12.

Pyrameis cardui aberration suffusa.—Fruhstorfer, 1912a, p.

525; 1912b, p. 525.

Vanessa cardui Linnaeus, Hudson not Linnaeus [a misidentification], 1898, pp. 108-109, pi. 12: figs. 1, 2.

C. kershawi has been treated frequently as an Australian subspecies of C. cardui, almost from the

time that it was first described. Its specific reality

was settled by Van Son (1966).

As mentioned in the group description above it

differs from all other species of Cynthia except C.

cardui in having the fuscous color in interspace

Cu2 on upper surface of forewing not extending to

vein Cux. On the upper surface of the hindwing it

differs from C. cardui in having four, not five, submarginal ocular spots, these being located between

veins Mt and Ciij,, with the lower three of these

spots having blue centers.

MALE (Figures 105, 106).—Forewing above with

subapical white bar just beyond cell and subapical

submarginal white spots very much as in C. cardui

and quite similar to those of the other Cynthia species except that the white spot found in interspace

Cux in the species of the virginiensis group is lacking in C. kershawi and C. cardui; with fuscous bar

in interspace Cua connecting apical and subapical

fuscous areas to central fuscous spot in interspace

Cu2 lacking; with central fuscous spot in interspace

Cu2 just mentioned usually more strongly connected

to the fuscous spot at base and above base of Cu2

than it is in C. cardui, and with the fuscous spots

in middle of cell confluent or nearly confluent, not

distinctly separate as is true of C. cardui.

Hindwing on the upper surface as mentioned

before with four, not five, submarginal ocular spots

between vein Mt and Coj, the lower three with blue

centers; with an upwardly projected hooklike fuscous marking below and beyond cell, all that remains of the fuscous and usually sinuous band

that crosses the wing from costa to just above anal

angle in the species of the virginiensis group, this

hooklike fuscous spot only very infrequently connected to the fuscous area along costa.

Under surfaces of forewing with fuscous of central

area of interspace Cua not extending above that

vein and with the two fuscous spots in middle of
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cell just inward from the white area much closer

to each other than is true in C. cardui.

Hindwing on the under surface with a darker

overall appearance than in C. cardui, with the submarginal ocular markings distinctly smaller than in

that species, and with the subtriangular white spot

at the end of the cell in interspaces M2 also much

smaller.

Length of forewing, 23-24.5 mm (average 23.8

mm).

Male genitalia as illustrated by Figure 9 (drawn

from my preparation no. 6098), with uncus slightly

tapered distally and blunt; gnathos long, slightly

longer than uncus, and more blunt distally than

in C. myrinna and C. cardui; valva with costal

margin broadly projected dorsally and forming a

broad obtuse angle, with a small upward projected

spine below and distad of this margin's greatest

projection; valva with distal margin concave and,

together with the distal end of costal margin, forming a second angular projection; lower margin of

valva slightly undulate and terminating in a sickleshaped process; clasper from near middle of valva

broad and short and slightly curved toward costal

margin; cuiller smooth and projected toward costal

margin; aedeagus very similar to that of C. cardui

and C. myrinna, not distinctly downward bent near

base.

FEMALE (Figures 107, 108).—Females of this

species, as is true of C. cardui, have a slightly more

rounded outer margin on the hindwing and a

slightly more produced outer margin on the forewing near vein Cu2.

Length of forewing, 22-27 mm (average 25.2 mm).

Female genitalia as illustrated by Figure 25 (drawn

from my preparation no. 6091), similar to those of

C. cardui, with posteriorly directed lobes about as

in that species except those of seventh sternite,

which have at their apices dark colored, downwardly directed, thumblike spurs; pockets between

seventh and eighth sternites much smaller than in

C. cardui; ribbonlike signa over half the length of

bursae copulatrix.

METHOD OF IDENTIFICATION.—The type locality is

given as Australia, "in abundance about Melbourne

and in many other parts of Australia." The original

description gives the major habitus characters which

distinguish this species. The location of the type

specimen is unknown to me.

SYNONYMY.—Pyrameis lucasii, described as a new

species from Victoria by Miskin (1888, p. 1515—

1516), was shown to be an aberration of C. kershawi

by Water house and Lyell (1914, p, 56) and is here

placed in synonymy.

LIFE HISTORY.—The larvae feed upon the following Compositae: Helichrysum, Ammobium alatum,

Acroelinium roseum, Gnaphalium japonicum, and

Cryptostemma calendula, a roadside weed introduced into Australia from South Africa. Very brief

descriptions of the immature stages and some lifehistory notes were given by Bodley (1920, pp. 74-

75) and by Burns (1969, p. 80). Notes on its migrations were given by Williams (1930, pp. 204-205)

and Fox (1969, p. 7).

DISTRIBUTION.—This species is found all around

the coastal region of Australia and for a considerable distance inland. Williams (1949b, p. 40) states

that it may upon occasion migrate to New Zealand.

Pritchard (1969, pp. 11-12) confirms one such migration and states that it has recently been observed

ovipositing on Gnaphalium japonicum in Auckland, New Zealand.

MATERIAL STUDIED.—Seven males and eleven

females were available for study from the following

localities: AUSTRALIA: Birchip (February, March);

Brisbane; Colo Vale, New South Wales (January);

Maribyrnong (September); New Market, Victoria

(September); Tasmania; Victoria (October). NEW

ZEALAND: NO specific locality.

The Virginiensis Group

This group contains five species: C. virginiensis,

C. altissima, C. braziliensis, C. terpsichore, and C.

myrinna. Here on the upper surface of the forewing

the fuscous color from interspace Cu2, just beyond

the middle of anal vein, extends upward at least

through interspace Cux and usually through interspace M3 to merge with fuscous ground color of

the subapical area. On the under surface of the

forewing the virginiensis group has no white behind

the end of cell, or only a small amount, usually a

white line along the inner surface of the fuscous

bar at the end of the cell, and as on the upper surface the fuscous color from interspace Cu2, just

beyond the middle of the anal vein, extends upward

to vein M3. On the under surface of the hindwing

there are only two submarginal ocular spots, these
NUMBER 84 47

being large, usually many times larger than those

in the cardui group. There is a white spot, sometimes peppered with fuscous, subtriangular in shape,

opposite the end of cell just above vein M3, but

this spot is not isolated as it is in the cardui and

carye groups, and it forms the central segment of

the light-colored band running across middle of

wing from costa to abdominal margin just before

anal angle.

In this group all species except C. terpsichore

have on the ovipositors a number of cephalically

directed teethlike spines. These spines are lacking

in the cardui and carye groups.

3. Cynthia virginiensis (Drury), resurrected

combination

FIGURES 10, 26, 109-118

Drury, 1770, pp. 10-11, pi. 5: fig. 1 [without name].

Nymphalis cardui virginiensis Drury, 1773, index p. [1] [name

given to 1770 description].

Papilio cardui virginiensis.—Boisduval and LeConte, 1834, p.

180 [synonym of Vanessa huntera (Fabricius) ].—Humphreys and Westwood, 1841, p. 57 [synonym of huntera].—

Doubleday, 1849, p. 205 [synonym of huntera].—Lucas

1857, p. 543 [synonym of huntera].

Papilio virginiensis.—Doubleday, 1844, p. 80 [synonym of

huntera].

Pyrameis virginiensis.—Morris, 1860, p. 8 [synonym of

huntera].—Morris, 1862, p. 60 [synonym of huntera]; 1868,

p. 83 [synonym of huntera].—Kirby, 1871, p. 186.—Staudinger, 1885, p. 98.—Rebel, Weymer and Stichel, 1901, p.

307.—Staudinger and Rebel, 1901, p. 24.—Alfken, 1903, p.

569.—Stichel, 1908, p. 200, pi. 62: fig. D 3; 1909, p. 200.—

Reuss, 1910a, pi. 1: figs. 3, 7, 11; 1910b, pp. 63, 64, 66, 67;

1910c, p. 90.—Stephen, 1924, p. 25.—Clark, 1932, pp. 21, 26,

86-88, pi. 27: figs. 1, 2.—South, 1941, p. 81.

Vanessa virginiensis.—Kirby, 1882, p. 13.-Dale, 1890, p. 147.—

Barrett, 1893, p. 155 [synonym of huntera].—Dyar, 1903, p.

23 [synonym of huntera].—Barnes and McDunnough, 1917,

p. 11.—Gunder, 1925, p. 198, pi. 5: fig. 10a.-Comstock,

1927, p. 132, fig. D (p. 132), pi. 43: figs. 3, 6.-Williams,

1930, pp. 225-226.—Holland, 1931, pi. 154, pi. 1: fig. 2, pi.

3: fig. 34, pi. 4: figs. 54, 63, 64, pi. 33: fig. 6.—McDunnough, 1938, p. 20.—Wild, 1939, p. 26, pi. 7: figs. 2, 3.—

Field, 1940b, pp. 81, 84-85, 274.—Hoffmann, 1940, p. 681 —

Macy and Shepard, 1941, pp. 123-125.—Comstock, 1944, pp.

449-450.—Bryk, 1946, p. 39.—Leighton, 1946, p. 59.-Verily,

1950, p. 329.—Clark and Clark, 1951, pp. 14, 44-45, 201,

227, pi. 4: figs. /, g, pi. 30: figs. /, k—Klots, 1951, pp. 48,

108, 113, pi. 5: fig. 5, pi. 14: fig. 11.—Ferguson, 1954, pp.

194-195.—Brown, 1955, p. 101, 2 figs—Zimmermann, 1958,

pp. 453, 454, 455, 456, 472, figs. 388, 389, 394, 398, 402.—

Forbes 1960, p. 159.—Dos Passos, 1964, p. 77.—Fulton, 1967,

p. 289.

Pyrameis huntera virginiensis.—Seitz, 1914a, p. 459; 1914b, p.

459.

Cynthia virginiensis.—Barnes and Benjamin, 1926, p. 14.

Vanessa virginiensis virginiensis.—Bates, 1935, pp. 165-166.—

d'Almeida, 1941, p. 308.

Papilio huntera Fabricius, 1775, p. 499; 1787, p. 45; 1793,

p. 104.—Herbst, 1794, pp. 165-166.—Comstock, 1944, p.

449 [synonym of virginiensis].—Zimmermann, 1958, p. 472

[synonym of virginiensis].

Vanessa huntera.—Godart, 1819, p. 324.—Boisduval and

LeConte, 1834, pp. 180-181.—Doubleday, 1844, p. 80 —

Capronnier, 1874, p. 25.—Kirby, 1882, p. 13 [synonym of

virginiensis].—Scudder, 1888, pi. 2: fig. 3, pi. 38: fig. 12, pi.

78: figs. 52, 53, 60, pi. 83: fig. 64; 1889, pp. 457-469, pi. 12:

fig. 9, pi. 20: fig. 8, pi. 33: fig. 22, pi. 53: fig. 5, pi. 74: fig.

34, pi. 81: fig. 11.—Dale, 1890, p. 147 [synonym of virginiensis].—Barrett, 1893, pp. 155-156.—Meyrick, 1899, p. 194.

—Dyar, 1903, p. 23.—Comstock and Comstock, 1904, pp.

156-157, pi. 26: figs. 3, 4.—Kirkaldy, 1906, p. 139.—Barnes

and McDunnough, 1917, p. 11.—Holland, 1931, p. 154

[synonym of virginiensis].—Davenport and Dethier, 1938,

p. 163.—McDunnough, 1938, p. 20.—Verity, 1950, p. 329.—

Klots, 1951, p. 108 [synonym of virginiensis].—ChalmersHunt, 1960-1961, pp. 62, 140.—Dos Passos, 1964, p. 77

[synonym of virginiensis].

Cynthia huntera.—Humphreys and Westwood, 1841, pp. 54,

57, pi. 15: figs. 5, 6.—Morris, 1868, pp. 83-B4, pi. 35.—

Barnes and Benjamin, 1926, p. 14 [synonym of virginiensis].

Pyrameis huntera.—Doubleday, 1849, p. 205.—Lucas, 1857,

pp. 542-543.—Morris, 1860, p. 8; 1862, p. 60.—Butler, 1870,

p. 77.—Strecker, 1878, pp. 138, 190.—Godman and Salvin,

1882, pp. 218-219.—Marshall and de Niceville, 1886, p.

226.—Edwards, 1889, pp. 25-26.—Dixey, 1890, pp. 93, 94,

100, 101, 107, 112, 115, 116, 117.—Maynard, 1891, pp. 92-93,

95 (fig. 32b) .—Godman and Salvin, 1891, p. 101.—White,

1894, pp. 56-57, 63, pi. 2: fig. 4.—Denton, 1898, pp. 267-

268, pi. [31].—Holland, 1898, p. 170, pi. 1: fig. 2, pi. 3:

fig. 34, pi. 4: figs. 54, 63, 64, pi. 33: fig. 6.—Skinner, 1898,

p. 24.—Wright, 1905, p. 177, pi. 22: fig. 228.—Kirkaldy,

1906, p. 139.—Seitz, 1914a, p. 459; 1914b, p. 459.—Holland,

1915, pp. 110-101, pi. 34.—Stephan, 1924, p. 25.—Forster,

1949, p. 106.

Papilio iole Cramer, 1776, p. 153 [as jole, validating iole

Cramer without generic assignment, 1775, pp. 17, 18, pi.

12: fig. E, F].—Boisduval and LeConte, 1834, p. 80 [synonym of huntera].—Humphreys and Westwood, 1841, p. 54

[synonym of huntera].—Doubleday, 1844, p. 80 [synonym

of huntera]; 1849, p. 205 [synonym of huntera].—Lucas, 1857,

p. 543 [synonym of huntera].—Morris, 1869, p. 83 [synonym

of huntera].—Berg, 1882, p. 166 [synonym of virginiensis].

—Comstock, 1944, p. 449 [synonym of virginiensis].

Pyrameis iole.—Morris, 1860, p. 8 [synonym of huntera]; 1862,

p. 60 [synonym of huntera].—Seitz, 1914a, pi. 94: figs, A 6,

B 1; 1914b, pi. 94: figs, A 6, B 1.

Vanessa iole.—Dyar, 1903, p. 23 [synonym of huntera].—

Barnes and McDunnough, 1917, p. 11 [synonym of virginiensis].—Holland, 1931, p. 154 [synonym of virginiensis].
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

—McDunnough, 1938, p. 20 [synonym of virginiensis].—

Dos Passos, 1964, p. 77 [synonym of virginiensis].

Cynthia iole.—Barnes and Benjamin, 1926, p. 14 [synonym of

virginiensis].

Vanessa virginiensis iole.—d'Almeida, 1941, p. 308.

Papilio jole Cramer, 1776, p. 153 [variant in spelling of iole].

Pyrameis huntera jole.—Berg, 1876, pp. 200-201.

Vanessa jole.—Dos Passos, 1964, p. 77 [as synonym of virginiensis].

Vanessa hunteri Hiibner [misspelling of huntera], [1819], p.

33.—Doubleday, 1849, p. 205.—Berg, 1882, p. 166 [synonym

of virginiensis].—Dyar, 1903, p. 23 [synonym of huntera].

—Barnes and McDunnough, 1917, p. 11 [synonym of virginiensis]; 1938, p. 20 [synonym of virginiensis].—Dos Passos,

1964, p. 77 [synonym of virginiensis].

Pyrameis hunteri.—Behr, 1864, p. 126.

Cynthia hunteri.—Barnes and Benjamin, 1926, p. 14 [synonym of virginiensis].

Pyrameis huntera variety fulvia Dodge [excluded name, type

3], 1900, p. 92.

Vanessa huntera fulvia.—Dyar [new synonymy], 1903, p. 23.

Pyrameis virginiensis aberration fulvia.—Reuss, 1910b, p. 66.

Pyrameis huntera form fulvia.—Seitz, 1916a, p. 598; 1916b, p.

598.

Vanessa virginiensis aberration fulvia.—Barnes and McDunnough, 1917, p. 11.—Gunder, 1925, p. 198.—Comstock, 1927,

p. 132.—McDunnough, 1938, p. 20.—Field, 1940b, pp. 84,

85, 274.—Forbes, 1960, p. 159.—Dos Passos, 1964, p. 77.

Cynthia virginiensis aberration fulvia.—Barnes and Benjamin,

1926, p. 14.

Pyrameis huntera form fulva Seitz [misspelling of fulvia],

1914a, p. 459; 1914b, p. 459.

Vanessa virginiensis fulva.—Fox, 1921, p. 46.

Vanessa virginiensis variety ahwashtee Fox [as an aberrant

form, excluded name, type 3], 1921, pp. 45-46, pi. 2: figs.

1, 2.

Vanessa virginiensis aberration ahwashtee.—Gunder [excluded

name, type 2], 1925, p. 198.—Comstock, 1927, p. 133, pi.

42: fig. 6.—McDunnough, 1938, p. 20—Field, 1940b, pp.

84, 85.—Dos, Passos, 1964, p. 77.

Cynthia virginiensis aberration ahwashtee.—Barnes and Benjamin, 1926, p. 14.

Cynthia virginiensis transition form ahwashtee.—Gunder [excluded name, type 2], 1927c, p. 270, pi. 8: figs. 1-5.

Vanessa virginiensis aberration massachusettensis Gunder [excluded name, type 2], 1925, p. 198, pi. 5: fig. 10.—Comstock,

1927, p. 133, pi. 43: fig. 5.—McDunnough, 1938, p. 20 —

Field, 1940b, pp. 84, 85.—Dos Passos, 1964, p. 77.

Cynthia virginiensis aberration massachusettensis.—Barnes

and Benjamin, 1926, p. 14.

Cynthia virginiensis transition form simmsi Gunder [excluded

name, type 2], 1927a, pp. 133-134, pi. 2: fig. 2; 1927c, p.

270, pi. 8: figs. 1-4.

Vanessa virginiensis aberration simmsi.—McDunnough [excluded name, type 2], 1938, p. 20.—Dos Passos, 1964, p. 77.

This species differs from all other species in the

virginiensis group in having the fuscous band on the

upper surface of the hindwing—that usually crosses

the wing from costa to the hind margin near anal

angle—greatly reduced in width and always broken

up into separate interspacial spots and sometimes

even absent. There are also four or five submarginal

ocular spots between veins R8 and Cu2 on the

upper surface of this wing. It differs from C. braziliensis, C. terpsichore, and C. myrinna on the under

surface of the hindwing in having the light spot

(the central segment of the light-colored band

running across middle of wing from costa to abdominal margin just before anal angle) opposite

the end of cell just above vein M3 white peppered

with fuscous scales, not pure white as it is in those

species; from C. altissima, which it otherwise greatly

resembles on this surface of the hindwing, it differs

in having the two submarginal ocular markings respectively larger and in lacking the great amount

of white edging along inner surface of submarginal

blue band, this white edging being reduced to a

very thin line.

MALE (Figures 109, 110, 113-116).—Forewing

above with subapical bar just beyond cell white,

frequently straight, sometimes curved inwardly;

with a series of subapical and submarginal spots

down to vein M3 white, including the spot in interspace Mt; white spot in interspace CUi smaller than

in C. braziliensis, about as in C. altissima and, as in

that species and C. terpsichore, often reduced to a

mere point; inner surface of fuscous bar at end of

cell not outlined with white; marginal bluish-white

line about as in C. altissima and C. braziliensis,

somewhat thinner; light areas of wing tawny, sometimes rather pale.

Hindwing on upper surface tawny, with the fuscous band crossing the middle of the wing greatly

reduced in width, sometimes absent, when present

always broken up into separate interspacial spots

(as mentioned above); with four or five submarginal

ocular spots between veins Rs and Cu2, the two in

interspaces Mi and Cux fairly large with blue centers, those in interspaces Rs

, M2 and M3 smaller and

without blue centers, and the one in interspaces Rs

sometimes reduced to a mere point or entirely

absent.

Under surface of forewing very similar to that of

C. altissima, with tawny color a little brighter, with
NUMBER 84 49

many of the same markings that occur on the upper

surface, especially the white subapical bar just beyond the cell, the series of white subapical and submarginal spots found below costa and between there

and vein R5, the white spots in interspaces R5 and

Mx and the white spots in interspaces M2 and Cujj

with fine, white lines behind the subapical white

bar and between this bar and the fuscous bar opposite end of cell; white bar crossing cell along inner

edge of this fuscous bar less distinct than in C.

altissima and with less white along costal margin

opposite end of cell than in that species; as in C.

altissima the fuscous markings in base of wing near

lower angle are similar to those markings above.

Hindwing underneath with white and peppered

band crossing middle of wing from costa to anal

angle similar to that of C. altissima, and as in that

species with the element of this band opposite end

of cell in interspace M2 also peppered with fuscous,

not pure white as in C. braziliensis, C. terpsichore,

and C. myrinna; with two ocular markings in interspaces Mt

 and Cux rather large, larger than in any

of the other Cynthia species, and, as is true of the

other species in the virginiensis group, with the

ocular spot in interspace Cu! the larger of the two;

all aspects of pattern much as in C. altissima except

as noted and except for the greatly reduced amount

of white edging along inner surface of the submarginal blue band.

Length of forewing, 19-29 mm (average 25.1 mm).

Male genitalia as illustrated by Figure 10 (drawn

from my preparation no. 6122), with uncus distally

blunt; gnathos longer than uncus, very much like

C. cardui, only slightly thicker; valva with anterior

half of costal margin projected upward, forming a

large, rounded lobe and containing a large spinelike process on posterior part of this lobe; costal

margin of valva posterior to this lobe concave,

ending in an acute process at the distal angle (junction of distal margin and costal margin); distal

margin of valva rounded to slightly undulate; lower

margin broadly rounded and with only a small

slightly rounded lobe at junction of distal margin;

clasper from near middle of valva large, but somewhat smaller than in C. myrinna, and directed upward toward distal angle; cuiller larger than in C.

cardui but otherwise very similar, smooth, with free

part curved in a shallow figure S toward distal margin; aedeagus similar to that of C. cardui except

that the base is nearly straight, not bent downward

as it is in that species.

FEMALE (Figures 111, 112, 117, 118) .—Females

of this species definitely show a more rounded outer

margin on the hindwing and usually display a

slightly more produced outer margin on the forewing near the end of the vein Cu2.

Length of forewing, 21-35 mm (average 28.5 mm).

Female genitalia as illustrated by Figure 26

(drawn from my preparation no. 3679), with posterior margin of seventh sternite and anterior margin of eighth sternite as greatly undulated as in C.

myrinna; with ostium bursae on eighth sternite and

placed deeply within lobes of seventh sternite; with

a pair of large laterally placed pockets between

seventh and eighth sternites; lobes of eighth sternite greatly projected posteriorly; signa ribbonlike

and much less than half the length of bursae copulatrix; ovipositors with number of forward-directed

teethlike spines.

SEASONAL VARIATION.—Specimens captured in the

spring (Figures 9, 12) are smaller, paler colored

with fuscous markings reduced in size, and often

more narrow.

METHOD OF IDENTIFICATION.—This species was

thoroughly described and illustrated in color by

Drury (1770, pp. 10-11, pi. 5: fig. 1; 1773, index p.

[1]), and there is no question of its identification

with the species treated here. This species was described from New York, Maryland, and Virginia.

Because of its name I restrict the type locality to

Virginia. The original specimens have not been

located. Because of the original and very excellent

colored figures of both surfaces of the wings (probably made from a female specimen), a neotype

designation is unnecessary.

SYNONYMY.—See synonymy and literature citations above. Fabricius (1775, p. 499) without giving

a reason proposed Papilio huntera in place of

Drury's name. Cramer (1775, pp. 17, 18, pi. 12: figs.

E, F.; 1776, p. 153) describes the species Papilio iole,

which is clearly a synonym of C. virginiensis as was

shown by Boisduval and LeConte (1834, p. 80).

Several names were given to aberrant specimens as

is shown above.

LIFE HISTORY.—The larval food plants are:

Gnaphalium obtusifolium, G. purpureum, G. polycephalum, Antennaria plantaginijolia, Anaphalis
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

margaritacea, and Artemisia, all members of the

Compositae.

The larva was described as long ago as 1770

(Drury, p. 11). Life-history notes and descriptions

of the immature stages were given by a number of

authors. References to this literature were given

by Edwards (1889, pp. 25-26) and Davenport and

Dethier (1938, p. 163). The most important single

reference is that of Scudder (1889, pp. 461-463,

465-468).

DISTRIBUTION.—From Nova Scotia, Alberta, and

British Columbia, over most of North America

south to Colombia and the Antilles, where it occurs

on Cuba, Haiti, and Puerto Rico. It has emigrated

to and is established in Hawaii, the Azores, Madeira,

and the Canary Islands. I have one specimen labeled

"Rio de Janeiro, Brazil" (Figures 115, 116). Further

confirmation is desirable as to its occurence at this

locality.

MATERIAL STUDIED.—One hundred and eighteen

males and one hundred and twenty-three females

were studied from the following localities: ONTARIO:

Nepigon (July); Niagara Falls (August). MAINE:

Augusta (July); Brunswick (July). NEW YORK:

Allegany State Park (July, August) ; Brooklyn

(September); Maspeth, Long Island (July) ; Monticello (July); Newtown, Long Island (July).

PENNSYLVANIA: New Brighton (September, October) ; Tobyhanna (July). NEW JERSEY: Lakehurst

(July); Pluckemin (July); Swartswood, Lake

County. MARYLAND: Beltsville (July); Frostburg

(July); Prince Frederick (October). DISTRICT OF

COLUMBIA: Rock Creek Park (June); Washington

(June, October). VIRGINIA: Dendron (September);

Dismal Swamp (April, May) ; Lake Tecomseh

(July) ; Millwood (May) ; Morrison City (August);

Old Trap near Petersburg (August); Princess Anne

(May, July); Salem (September, October); Sherando (July); White Top (August). SOUTH CAROLINA: Columbia (January, May, June, October).

GEORGIA: Brookhaven (May, July); Calhoun (October) . FLORIDA: Andras (March); Chokoloskee; Dade

City; Glenwood. MICHIGAN: Interlochen (August);

Monroe Center (August); New Hudson (July) ;

Pinckney (August) ; Washington County (July).

WISCONSIN: Madison. NEBRASKA: Lincoln (September) . IOWA: Keokuk. ILLINOIS: Elkhart (October);

Putnum (June). INDIANA: Bloomington (May).

KANSAS: Eureka (April, May, June); Lawrence

(June); Pittsburg (June). MISSOURI: Columbia

(June); St. Louis (May). TENNESSEE: Bedford

County (May); Cades Cove, Great Smoky Mountains National Park (August); Townsend, Great

Smoky Mountain National Park (August). TEXAS:

Brownsville; Dallas (April) ; Laredo (June); San

Benito (May) ; Trinity (March). LOUISIANA: Fort

McComb (June). MISSISSIPPI: Vicksburg. ALABAMA:

Marion (January). OREGON: Reed (July, September, October). WYOMING: Clark County (August) ;

Fremont County (7,000-11,000 ft). COLORADO: Hall

Valley (June). CALIFORNIA: Camp Baldy, San

Bernadino Mountains (July); Los Angeles; Olancha

(September); San Diego (May); Shasta County

(July). ARIZONA: Huachuca Mountains; Palmer lee,

Cochise County; White Mountains. HAWAII: Olaa.

MEXICO: Cordoba (May) ; Jalapa; Mexico City

(June); Saltillo (July) ; Santa Rosa (July); Urupan. GUATEMALA: Volcan Santa Maria; Zacapa.

HONDURAS: Subirana Yoro (February). COSTA RICA:

Mount Poas; San Pedro de Montes de Oca (February) . VENEZUELA: Chorone (September). COLOMBIA: Bogota. CUBA: Loma del Gato (July). HAITI:

La Visite and La Selle Range (September, 5,000-

7,000 ft); Mount Bovrette (September). PUERTO

RICO: Cayey (February). BRAZIL: Rio de Janeiro!

4. Cynthia altissima (Rosenberg and Talbot),

new combination, new status

FIGURES 11, 17, 19-122

Pyrameis huntera altissima Rosenberg and Talbot, 1914, p.

675.—Seitz, 1914a, p. 459; 1914b, p. 459.

Of all the Cynthia species, C. altissima is the most

similar to C. braziliensis. It has on the forewing

the submarginal spot in interspace Mx white, not

blue, however, and lacks the white band outlining

inner surface of black bar at end of cell found in

that species; it differs from all other species of

Cynthia in having the fuscous band crossing the

middle of hindwing not projected outward in interspaces M2 and M3.

MALE (Figures 119, 120).—Forewing above the

subapical bar just beyond cell white, usually considerably wider in interspace R5 than elsewhere;

with the series of subapical and submarginal spots

down to vein M3 white; with the white spot in

interspace Cux small, often reduced to a mere point;
NUMBER 84 51

inner surface of fuscous bar at end of cell not outlined with white; marginal bluish-white line about

as in C. braziliensis; light areas of wing tawny, only

rarely with a slight pinkish hue.

Hindwing above tawny, with a nearly straight

fuscous band crossing wing from costa to just above

anal angle, this band not more obscure near anal

angle; this band not greatly projected outward in

interspaces M2 and M3 as it is in C. braziliensis and

C. terpsichore; with two submarginal ocular markings, one each in interspace Ma and interspace Cu2,

these markings usually smaller than in C. braziliensis

and with blue centers sometimes lacking; with two

small fuscous dots in interspaces M2 and M3 and

sometimes a third dot in interspace Ra.

Forewing underneath hardly distinguishable from

C. virginiensis except as mentioned below, with

many of the same markings that occur on the upper

surface, especially the white subapical bar just

beyond the cell, the series of white subapical and

submarginal spots found below costa and between

there and vein R5, the white spot in interspace

R5 and Mx and the white spots in interspaces M2

and Cu^ the lines found in C. myrinna behind the

subapical white bar and between this bar and the

fuscous bar opposite end of cell are here white, not

bluish white, and are greatly obscured except along

outer side of this fuscous bar; white bar crossing

cell along inner edge of this fuscous bar in C.

virginiensis, C. terpsichore, and C. myrinna is more

distinct than in C. virginiensis and C. terpsichore,

similar to that of C. myrinna and with more white

along costal margin opposite end of cell than in

C. virginiensis; the fuscous markings in base of

wing and near lower angle are similar to those

markings above.

Hindwing underneath with white and peppered

band crossing middle of wing from costa to anal

angle very much as in C. virginiensis, projected outward in interspace M2 as it is in C. virginiensis and

other Cynthia species except C. myrinna; with two

ocular markings in interspaces Mx and Cux rather

small, larger than in C. terpsichore but smaller than

in C. virginiensis, with the one in interspace Cux

slightly the larger of the two; all aspects of pattern

much as in C. virginiensis except for a great amount

of white edging along inner surface of submarginal

blue band.

Length of forewing, 24-27 mm (average 25.5 mm).

Male genitalia as illustrated by Figure 11 (drawn

from my preparation no. 6141), very similar to those

of C. virginiensis, differing chiefly in the valva,

where the distal angle lacks a definite angled toothlike projection, the lobe at junction of distal margin

and lower margin is slightly more pronounced, and

the cuiller is toothed along the middle of its inner

edge, with clasper larger and directed toward costal

margin of valva.

FEMALE.—This sex as is true of most other species

of Cynthia only differs from the male sex in having

a more rounded outer margin on the hindwing

and a slightly more produced forewing in the area

of the lower angle.

Length of forewing, 28 mm.

Female genitalia as illustrated by Figure 27

(drawn from my preparation no. 6465), most similar to C. virginiensis, differing in having the posterior margin of eighth sternite more abruptly

produced posteriorly; signa as in C. virginiensis;

ovipositors with a number of forward directed

teethlike spines.

METHOD OF IDENTIFICATION.—The original description was not sufficient to identify the species

to which the name Cynthia altissima belonged. This

species was described from "a series from Agualani,

S. E. Peru, 9000 feet, July and August 1905" from

"Limbani, S. E. Peru, 9500 feet, April and May

1904" and from "one specimen from Banos, Rio

Pastaza, E. Ecuador, 6100 feet, January 1911." The

study of a paratype from Agualani in the collection

of the Museum of Comparative Zoology, Harvard

University, identified this name as belonging to the

species here treated.

A male labeled type in the collection of the

British Museum (Natural History) is here designated the lectotype, and a female labeled cotype,

also in the collection, is designated the paralectotype. Appropriate labels have been placed on both

of these specimens by Mr. T. G. Howarth of the

aforementioned institution. The lectotype is also

labeled "Agualani, S. E. Peru, 9,000 ft., July

 f05."

LIFE HISTORY.—Seitz (1914a, 1914b, p. 459) gave

a brief description of the larvae but the food plants

are unknown. They are probably Compositae.

DISTRIBUTION.—This species is found only in high

elevations in the Andes of Ecuador, Peru, and

Bolivia.
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MATERIAL STUDIED.—Seven males and two females

were available to me for study from the following

localities: PERU: Agualani, Carabaya (June, August,

October, 9,000 ft, 10,000 ft); north Peru, west

slopes of Andes (June) ; Oroya to Limbani (January) ; Pisac (May). BOLIVIA: Cochabamba (five days

north, August); Okara (April, 7,500 ft); Cusilluni

(May).

5. Cynthia braziliensis (Moore), new

combination

FICURES 12, 28, 123-130

Pyrameis huntera iole (Cramer), Berg not Cramer, [a misidentification], 1875, pp. 69-70, 102.

Pyrameis braziliensis Moore, 1883, pp. 236.—Dukinfield Jones

1883, p. 237.

Pyrameis huntera braziliensis.—Seitz, 1914a, p. 459; 1914b, p.

459.—Hayward, 1931, pp. 71, 186, pi. 12: fig. 3.

Vanessa virginiensis braziliensis.—d'Almeida, 1941, p. 309.—

Hayward, 1951, p. 196.—Biezanko, Ruffinelli and Carbonell,

1957, p. 127.

Pyrameis virginiensis (Drury), von Bayeran not Drury, [a

misidentification], 1901, p. 256.—Stichel, 1909, pi. 62: fig.

D 3.

Pyrameis virginiensis variety rubia Staudinger, 1894, pp. 70-

71.—Hayward, 1951, p. 196 [synonym of braziliensis].

Pyrameis huntera rubia.—Seitz, 1914a, p. 459; 1914b, p. 459.

Pyrameis huntera form rubia.—Hayward, 1931, pp. 71-72.

Pyrameis brasiliensis.—Reuss [misspelling of braziliensis],

1910a, p. 66.

Pyrameis huntera Fabricius, Burmeister not Fabricius [a misidentification], 1878, pp. 149-151.—Berg, 1882, p. 166.—

Hayward, 1931, pp. 70-71, pi. 12: fig. 2; 1951, p. 196.

Vanessa cardui ushuwaia Bryk, Hayward not Bryk [a misidentification as a synonym of V. braziliensis], 1951, p. 196.

Pyrameis huntera aberration dallasi Kohler [excluded name,

type 2], 1945, p. 256, pi. 20: fig. 2.

Vanessa virginiensis braziliensis aberration dallasi.—Hayward,

1951, p. 196.

This species differs from all other species of Cynthia

in having on the forewing above a white band outlining inner surface of the black bar found at the

end of cell; on this wing it also differs from all

species of Cynthia, except C. myrinna, in having

the submarginal spot in interspace Mx blue, not

white.

MALE (Figures 123-126) .—Forewing above with

subapical bar just beyond cell white, slightly inwardly curved; with a series of subapical and submarginal spots down to vein M3 white except for

the spot in interspace Mx which is blue, not white,

as mentioned above; white spot in interspace Cux

larger than in C. altissima, C. terpsichore, and C.

myrinna; as described above and as mentioned in

the key with a white band outlining inner surface

of fuscous bar at end of cell, this bar being much

thicker along base of cell; with a marginal bluishwhite line, about as in C. myrinna, extending from

apex of wing to vein Mlf broken by the veins and

continued between veins M2 and lower angle and

only faintly indicated below vein Cu,; light areas

of wing tawny, with a slight pinkish hue.

Hindwing above tawny, often with a pinkish cast

and with a sinuous fuscous band crossing wing from

costa to just above anal angle, this band more

obscure near anal angle; with two submarginal

ocular markings, one each in interspaces Mt and

Culf usually about equal in size and often with

two small fuscous spots between these ocular markings in interspaces M2 and M3.

Forewing underneath most similar to C. virginiensis and, as in other species of Cynthia, with many

of the markings of the upper surfaces repeated,

especially the white subapical bar and the series of

white subapical and submarginal spots; margin of

apex down to vein M2 of a much less bluish hue

than in C. virginiensis and C. altissima; as in C

virginiensis with less white outlining fuscous bar

at end of cell along its inner edge, especially opposite costal margin; the lines found behind the subapical white bar and between this bar and the

fuscous bar opposite end of cell in C. myrinna and

other Cynthia species are more of a white color and

more obscure than in C. myrinna and C. virginiensis; fuscous markings in base of wing and near

lower angle similar to those same markings found

on the upper surfaces and of about the same size.

Under surface of hindwing with yellowish band,

peppered with fuscous, crossing middle of wing

from costa to anal angle, projected outward along

vein M3; as in C. virginiensis, C. altissima, and C.

terpsichore, this band pure white in interspace M2;

with two ocular markings in interspaces Mx and Cu^

about the size of those in C. altissima, with the one

in interspace Cux slightly larger.

Length of forewing, 20-28 mm (average 24.36

mm).
NUMBER 84 53

Male genitalia as illustrated by Figure 12 (drawn

from my preparation no. 3663), similar to those of

C. virginiensis and C. altissima, differing chiefly in

the valva, where the spinelike process found on

inner side of large lobe of costal margin in those

two species is absent and is replaced by a spine

located on outer edge of this lobe at the distal end

of its base; the small rounded lobe found at the

junction of distal and lower margins in C. virginiensis and C. altissima is in C. braziliensis greatly

enlarged and upward projected, forming a large

acute lobe; cuiller smooth and usually forked at

distal end, rarely simple, and sometimes with three

tines; clasper similar to that of C. altissima—large

and directed toward costal margin of valva.

FEMALE (Figures 127-130) .—The females of C.

braziliensis have a more rounded hindwing along

the outer margin and are slightly more produced

on the forewing along the edges of interspaces Mt

and Cu2.

Length of forewing, 21-30 mm (average 25.41

mm).

Female genitalia as illustrated by Figure 28

(drawn from my preparation no. 3680), similar to

those of C. virginiensis with eighth sternite much

more narrow and broadly divided near middle,

forming two wing-shaped plates; pockets between

seventh and eighth sternites small; signa about as

in C. virginiensis; ovipositor with forward directed

teethlike spines.

METHOD OF IDENTIFICATION.—The original description outlines the characters of this species quite

adequately to fix the name for the species here

called Cynthia braziliensis. The type locality is

Sao Paulo, Brazil.

The type specimen of this species is apparently

in the collection of the Liverpool Free Public

Museum, Liverpool, England.

SYNONYMY.—A study of the original description

of Pyrameis virginiensis rubia Staudinger (1894,

pp. 70-71) shows that this is a synonym.

LIFE HISTORY.—The larvae feed upon the following Compositae: Achyrocline satureioides, Gnaphalium spicatum, G. stachydifolium, and G. obtusifolium. A description of the larva and brief

life-history notes were given by Dukinfield Jones

(1883, p. 237).

DISTRIBUTION.—This species occurs from Merida,

Venezuela, south through the mountains of Colombia, Ecuador, Peru, and Bolivia and east to Rio de

Janeiro, Brazil, south through Paraguay, Uruguay,

and throughout most of Argentina.

MATERIAL STUDIED.—Sixty males and sixty-five

females were studied from the following localities:

VENEZUELA: Merida. COLOMBIA: NO specific locality;

Rio Quirata (December, 2,700 m). ECUADOR: NO

specific locality (April); Banos; Loja; Puyo, Pastaza,

Oriente (3,500 ft). PERU: CUZCO (July, 11,500 ft);

Oroya (May); Tincochaca (August, 7,000 ft);

Tingo Maria (670 m). BOLIVIA: NO specific locality; Chulumani (December). BRAZIL: Blumenau,

Santa Catherina; Cauna, Santa Catherina (April);

Pelotas, Rio Grande do Sul (March, April, May,

October); Petropolis; Rio de Janeiro (April, November) ; Sao Paulo (February); Tijuca. PARAGUAY:

Nueva Italia (July); Sapucay. ARGENTINA: NO

specific locality.

6. Cynthia terpsichore (Philippi), new

combination

FICURES 13, 29, 131-134

Vanesja [sic] terpsichore Philippi, 1859, pp. 1089-1091.

Vanessa terpsichore.—Philippi, 1860, pp. 266-267.—Ureta,

1938b, p. 298; 1938c, p. 127.—Bryk, 1944, p. 10.—Hayward,

1951, p. 196.—Herrera, 1954, pp. 45, 53, pi. 7: figs. 1, 2.—

Herrera and Etcheverry, 1956, pp. 278, 286.—Herrera, Etcheverry, and Barrientos, 1958, pp. 240 (fig. 2), 243, 244,

247 (figs. 15-17), 248 (figs. 20, 21), 250-251, pi. 1: figs. 3, 4.

Pyrameis terpsichore.—Kirby, 1871, p. 186.—Reed, 1877, pp.

679-680.—Butler, 1881, p. 467.—Staudinger, 1885, p. 98.—

Bartlett-Calvert, 1886, p. 314—Dixey, 1890, p. 122.—

Weymer and Maassen, 1890, pp. 14, 36, 48. 58.—BartlettCalvert, 1898. p. 99.—Porter, 1899a, p. 36.—Elwes, 1903,

pp. 287-288.—Reuss, 1910a, pp. 65, 66, 67.—Seitt, 1914a, p.

459, pi. 94: figs, A 4, A 5; 1914b, p. 459, pi. 94: figs, A 4, A

5.—Stephan, 1924, p. 25.—Bullock, 1934. p. 47.—Ureta, 1934.

p. 79; 1935, p. 94.—Ruiz and Stuardo, 1935, p. 315.

Payrameis [sic] terpsichore.—Reed, 1877, p. 735.

Pyrameis virginiensis (Drury), Berg not Drury [a misidentification], 1882, p. 166.

Pyrames [sic] iterpsichore [sic].—Ureta [a misspelling for

Pyrameis terpsichore], 1935, p. 87.

C. terpsichore, like C. annabella and C. carye, has

the subapical bar on forewing above just beyond

cell tawny, not white, in both sexes; it can easily

be distinguished from the latter two species by the

presence of a sinuous and fuscous band on the

upper surface of the hindwing that runs from the
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

costal margin across wing to the hind margin just

above anal angle. It differs from all other species

in the genus, except some specimens of C. braziliensis, by having two, not four, or more submarginal

ocular spots on upper surfaces of hindwing.

MALE (Figures 131, 132).—Forewing above with

subapical bar just beyond cell tawny, not white,

and rather straight, not inwardly curved as it is in

C. myrinna; with a series of subapical and submarginal spots down to vein M3 white and with a

small white spot in interspace Cux; with a marginal

bluish-white line from apex of wing to vein Mlt

greatly obscured below this vein; light areas of wing

tawny in color; fuscous markings in base of wing

and on lower half almost as greatly reduced as in

C. annabella and C. carye.

Hindwing above tawny, with the sinuous fuscous

band described before, and with two submarginal

ocular markings, one each in interspaces M1 and

Cux.

Forewing underneath, as in the other species of

Cynthia, with many of the same markings that occur

on the upper surfaces, particularly the white subapical bar just beyond cell, the series of white

subapical and submarginal spots found below costa

and between there and vein R5, the white spot in

interspaces R5 and Mx (here reduced to mere

points), and the white spots in interspaces M2 and

Cuj (here more reduced than in C. myrinna, C.

altissima, and C. braziliensis); the bluish lines found

in C. myrinna behind the subapical white bar and

between this bar and the fuscous bar opposite end

of cell are here greatly obscured and are replaced

by tawny along lower half of outer side of this

fuscous bar; white bar crossing cell along inner

edge of this fuscous bar in C. myrinna, C. altissima,

and C. braziliensis absent, replaced by the tawny

ground color; fuscous markings in base of wing and

near lower angle similar to those markings above,

more reduced in size.

Hindwing underneath with yellowish band crossing middle of wing from costa to anal angle not

straight as in C. myrinna, projected outward along

vein M3 as it is in C. altissima, C. braziliensis, and

most other species of Cynthia; this band becoming

white, almost silvery in interspace M2; with two

ocular markings, one each in interspaces Mx and

Cu,, very small in size, the one in interspace Cu!

slightly the larger of the two.

Length of forewing, 23-27 mm (average 25 mm).

Male genitalia as illustrated by Figure 13 (drawn

from my preparation no. 3665), similar to those of

C. braziliensis, C. virginiensis, and C. altissima,

differing chiefly in the valva; costal spine of valva

similar to that of C. braziliensis, located just below

the greatest projection of costal lobe; junction of

lower and distal margins lacking the acute lobe

found in C. braziliensis and thus similar to C.

altissima and C. virginiensis; cuiller smooth and

somewhat shorter than in those species and clasper

directed inward toward opposite valva as in C.

cardui; aedeagus similar to that of C. braziliensis.

FEMALE (Figures 133, 134).—This sex shows

very definitely a more rounded outer margin on the

hindwing, and the outer margins of forewing at

ends of interspaces Cuj and Cu2 are more produced

outwardly near end of vein Cu2.

Length of forewing, 25-28 mm (average 27 mm).

Female genitalia as illustrated by Figure 29

(drawn from my preparation no. 6438), similar to

those of C. cardui, with lobes of seventh and eighth

sternites nearly as large as in C. myrinna; with

lateral pockets between seventh and eighth sternites

very small; signa longer than in C. myrinna; ovipositors without teethlike spines.

METHOD OF IDENTIFICATION.—The original description by Philippi (1859, p. 1089) was sufficient

to identify his name for the species here described.

The type locality is given as "Frequens prope

Santiago, in prov. Valdiva etc."

LIFE HISTORY.—The food plants and immature

stages are unknown.

DISTRIBUTION.—This species is found in Chile

from the Province of Atacama south into the

Province of Aysen. Weymer and Maassen (1890)

report it also from Colombia and Ecuador, but I

believe these records should be verified.

MATERIAL STUDIED.—Eight males and five females

were studied from the following localities: CHILE:

Andes (March, 6,000 ft); Angol; Penco.

7. Cynthia myrinna (Doubleday), new

combination

FIGURES 14, 30, 135-136

Pyrameis myrinna Doubleday, 1849, p. 205.—Kirby, 1871, p.

186.—Staudinger, 1885, p. 98.—Dixey, 1890, pp. 93, 101,116,

117, 118, 122.—Longstaff, 1908, p. 118.—Reuss. 1910a. pi. 1:
NUMBER 84 55

figs. 4, 8, 12; 1910b, pp. 62, 64, 65, 66, 67; 1910c, p. 90.—

Seitz, 1914a, p. 459, pi. 94: figs, B 3, B 4; 1914b, p. 459, pi.

94: figs, B 3, B 4.—Stephan, 1924, p. 25.—Gabriel, 1927, p.

84.—Hayward, 1931, p. 72, pi. 12: fig. 1.

Vanessa myrinna.—Capronnier, 1874, p. 25.—Verity, 1950, p.

329.—Hayward, 1951, p. 196.—Biezanko, Rufinelli and

Carbonell, 1957, p. 124.

Vanessa syngenesiae Capronnier, 1874, p. 25 [a Boisduval

manuscript name and nomen nudem, in synonymy of

myrinna].

Pyrameis myrinna aberration incarnata Seitz [excluded name,

type 2], 1914a, p. 459; 1914b, p. 459.

Pyrameis myrinna aberration eunice Hall [excluded name,

type 2], 1917, p. 171.

Pyrameis myrinna aberration merlinde Ziken [excluded name,

type 2], 1937, pp. 386-387.

C. myrinna is easily distinguished from all other

species in the genus by the broad, straight fuscous

band crossing the upper surface of the hindwing

near the middle and by the straight, white to yellowish band crossing the middle of this same wing

on the under surfaces.

MALE (Figure 135).—Forewing above with a

white subapical bar just beyond cell, lying between

costa and vein M3, and inwardly curved; with a

series of subapical and submarginal spots down to

vein M3 white, except for the blue or bluish-white

spot in interspace M,, and as in C. virginiensis, C.

altissima, C. braziliensis, and C. terpsichore with a

small white spot in interspace Cu^ with a marginal

white to bluish-white line from apex of wing to

vein Mlt broken by the veins and continued between

vein M2 and lower angle and only very faintly

indicated below vein Cux; light areas of wing

tawny in color.

Hindwing above tawny, with a broad and straight

fuscous band as mentioned above and as described

in the key; with a submarginal series of four or

five ocular spots very indistinct, these being completely enclosed in the fuscous band and lacking

light-colored centers.

Forewing underneath with many of the same

markings that occur on the upper surfaces, particularly the white subapical bar just beyond cell, the

series of white subapical and submarginal spots

found below costa and between there and vein R5,

the white spot in interspace R5 (here reduced to a

mere point), and the white spots in interspaces M2

and Cux; with two blue or bluish-white lines behind

the subapical white bar and between this bar and

fuscous bar opposite end of cell; with a narrow

white bar crossing cell along inner edge of aforementioned fuscous bar; fuscous markings in base

of wing and near lower angle similar to those

markings above.

Hindwing underneath with the white to yellowish-white straight band crossing its middle as described before and as mentioned in the key; with

two large submarginal ocular markings in interspaces Mx and Cux, these being about equal in size.

Length of forewing, 25-32 mm (average 29.2

mm).

Male genitalia as illustrated by Figure 14 (drawn

from my preparation no. 6147), with uncus blunt

and slightly swollen; gnathos with lateral arms very

long, longer than uncus; valva with distal margin

projected acutely to a spinelike structure below

uncus and rounded from just opposite clasper to

lower margin; clasper directed toward upper part

of outer margin; cuiller armed with numerous teeth

along its dorsal margin and thus differing from all

Cynthia species except C. annabella and C. carye;

aedeagus slightly undulate and sharply downward

bent near base.

FEMALE (Figure 136).—In the females of C.

myrinna the fuscous areas on the upper surfaces of

both fore- and hindwing are slightly larger, the

outer margin of forewing is slightly more produced

in the area on either side of vein Cu,, and the

outer margin of hindwing is more rounded.

Length of forewing, 27-35 mm (average 32 mm).

Female genitalia as illustrated by Figure 30

(drawn from my preparation no. 6441), similar to

those of C. cardui but with posteriorly directed

lobes much larger and lacking the lateral pockets

found in C. cardui and other species; ovipositors

with a large number of forward-directed teethlike

spines; ribbonlike signa not as long as in C. cardui,

extending not more than one half the length of

bursae copulatrix.

METHOD OF IDENTIFICATION.—Doubleday's original

description (1849, p. 205) gives the necessary characters to identify his name with the species here

treated. The type locality is cited as Brazil.

There are two specimens of C. myrinna labeled

types in the collection of the British Museum (Na-
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tural History). I select as the lectotype the male

labeled type Rh. 8713 and as the paralectotype the

female specimen labeled type Rh. 8714. Appropriate

labels have been placed on both of these specimens

by Mr. T. G. Howarth of the aforementioned institution.

LIFE HISTORY.—The larvae feed upon Achyrocline

flaccida (Compositae). Very brief descriptions of

the larvae and pupae were given by Seitz (1914a,

1914b, p. 459).

DISTRIBUTION.—This species occurs from Caracas,

Venezuela, south through the mountains of Colombia, Ecuador, Peru, and Bolivia, and in southern

Brazil and Paraguay.

MATERIAL STUDIED.—Sixteen males and nineteen

females from the following localities: VENEZUELA:

Caracas. COLOMBIA: Sierra Nevada Mountains (8,000

ft). PERU: Chanchamayo. BOLIVIA: Cochabamba (5

days north, September). BRAZIL: Blumenau, Santa

Catherina; Santa Gallo; Nova Friburgo; Pernambuco; Rio de Janeiro (April); Santa Catherina

(February); Sao Paulo (April); Tijuca. PARAGUAY:

No specific locality; Buena Vista (February).

The Carye Group

This group contains the two species, C. carye and

C. annabella. The habitus of the upper surface of

the forewing is most similar to the virginiensis

group, differing in having a marginal tawny spot

in interspace M3. On the upper surface of the hindwing there are always four blue-centered round

ocular spots between veins Mt and Cu2, and the

fuscous sinuous band crossing this wing from costa

to just above anal angle is greatly reduced in width

and entirely absent, at least above vein Mj. On the

under surface of the forewing this group has the

area at the end of the cell, between there and the

fuscous spots, and through the middle of cell entirely tawny, and as in the virginiensis group, as is

true of the upper surfaces, the fuscous color from

interspace Cu2 just beyond the middle of the anal

vein extends upward to vein M3. On the under

surface of the hindwing there are four, sometimes

five, small submarginal ocular spots greatly obscured

by the fuscous of the surrounding area. There is an

isolated triangular or hourglass-shaped white spot

opposite end of cell and just above vein M3 on the

under surface of this wing.

8. Cynthia annabella, new species

FIGURES 15, 31, 143-148, 155-160

Vanessa carye (Hubner) , Doubleday not Hubner [a misidentification], 1844, p. 79.—Dyar, 1903, p. 24.—Barnes and

McDunnough, 1917, p. 11.—Coolidge, 1925, pp. 146-147.—

Comstock, 1927, pp. 134-135, fig. A 42 (p. 135), pi. 44: figs.

1-3.—Holland, 1931, p. 154, pi. 20: fig. 12.—Davenport and

Dethier, 1938, p. 164.—McDunnough, 1938, p. 21.—Field,

1940, pp. 81, 87-88, 274.—Hoffmann, 1940, p. 681.—Leighton, 1946, p. 59.—Dos Passos, 1964, p. 77.—Carrillo, Ortego,

and Gibson, 1966, p. 78.

Pyrameis carye.—Morris, 1860, p. 8.—Behr, 1864, p. 125.—

Strecker, 1878, p. 138.—Godman and Salvin, 1882, p. 219.—

Dyar, 1889, pp. 237-238.—Dixey, 1890, pp. 107, 117, 122 —

Maynard, 1891, p. 93, fig. 32a (p. 95) .—Skinner, 1898, p.

25.—Wright, 1905, p. 178, pi. 22: fig. 231.—Schrader, 1929,

pp. 8-11, pi. 4.

Cynthia carye.—Barnes and Benjamin, 1926, p. 15.—Gunder,

1927c, p. 270, pi. 8: fig. 4, pi. 9: fig. 1; 1929, p. 9, pi. 17:

figs. [1] [2].

Pyrameis caryae Edwards [misspelling of carye'] not Hubner

[a misidentification], 1874, p. 329; 1889, p. 26.—Holland,

1898, pp. 170-171, pi. 20: fig. 12.—Huguenin, 1921, pp.

216-217.

Vanessa caryae.—Grinnell, 1918, p. 111.—Brown, 1955, p. 103,

2 figs.

Pyrameis carye variety muelleri Letcher [as an individual

variant, excluded name, type 3], 1898, p. 38, pi. 3.

Pyrameis carye aberration muelleri.—Skinner [excluded name,

type 2], 1898, p. 25.

Vanessa carye aberration muelleri.—Barnes and McDunnough,

1917, p. 11.—Gunder, 1925, p. 198.—Comstock, 1927, p.

135, pi. 44: figs. 5, 6.—McDunnough, 1938, p. 21.—Field,

1940b, pp. 87, 88.—Dos Passos, 1964, p. 77.

Vanessa caryae variety muelleri.—Grinnell [as an individual

variant, excluded name, type 3], 1918, p. 111.

Vanessa caryae form muelleri.—Fox [excluded name, type 3],

1921, p. 46.

Cynthia carye aberration muelleri.—Barnes and Benjamin,

1926, p. 15.

Cynthia carye transition form muelleri.—Gunder [excluded

name, type 2], 1927a, p. 133, pi. 2: fig. 2 b; 1927c, p. 270,

pi. 9: figs. 2-5; 1929, p. 9, pi. 17: figs. [3], [4].

Vanessa caryae variety intermedia Grinnell [as an individual

variant, excluded name, type 3], 1918, pp. 111-112, pi. 4:

fig. 6.

Cynthia carye aberration intermedia.—Barnes and Benjamin

[excluded name, type 2], 1926, p. 15.

Vanessa carye aberration intermedia.—Comstock, 1927, p.

135, pi. 44: figs. 4, 5.—McDunnough, 1938, p. 21—Field,

1940b, pp. 87, 88.—Dos Passos, 1964, p. 77.

Cynthia carye transition form intermedia.—Gunder [excluded

name, type 2], 1927, p. 270.

Vanessa caryae variety letcheri Grinnell [as an individual
NUMBER 84 57

variant, excluded name, type 3], 1918, pp. 112-113, pi. 4:

fig. 8.

Cynthia carye aberration letcheri.—Barnes and Benjamin

[excluded name, type 2], 1926, p. 15.

Vanessa carye aberration letcheri.—Comstock, 1927, pp. 135—

136, pi. 44: figs. 7, 8.—McDunnough, 1938, p. 21.—Field,

1940b, p. 87.—Leighton, 1946, p. 59.—Dos Passos, 1964, p.

77.

Cynthia carye transition form letcheri.—Gunder [excluded

name, type 2], p. 270.

Cynthia carye transition form nivosa Gunder [excluded name,

type 2], 1927, p. 53.

Vanessa carye aberration nivosa.—McDunnough [excluded

name, type 3], 1938, p. 21.—Field, 1940b, pp. 87, 88.—Dos

Passos, 1964, p. 77.

Cynthia carye transition form schraderi Gunder [excluded

name, type 2], 1929, p. 9, pi. 17: figs. [5]-[10].

Vanessa carye aberration schraderi.—McDunnough [excluded

name, type 2], 1938, p. 21.—Dos Passos, 1964, p. 77.

This species and C. carye are easily distinguished

from all other species of Cynthia, except C. terpsichore and some females of C. virginiensis, in having

the bar just beyond cell on upper side of forewing

tawny instead of white. From C. terpsichore and C.

virginiensis they are easily distinguished by having

four, instead of two, blue-centered ocular markings

on upper side of each hindwing.

MALE (Figures 143-148).—Forewing above with

tawny subapical bar just beyond cell slightly larger

in C. annabella than in C. carye; subapical white

spot at costa and interspaces Rx and R2 slightly

larger than in C. carye; basal transverse bar in cell

more distinct in C. annabella; bar below case of

Cu2 usually very small, sometimes absent in C. carye,

present and distinct in C. annabella; usually with

a small tawny spot in interspace M3 just above the

large tawny quadrate spot in interspace Cux in C.

annabella, which is absent in most specimens of C.

carye; submarginal white line composed of three

slender bars on apex of forewing, usually more distinct in C. annabella than in C. carye.

Hindwing above with white spot in middle of

costal margin larger and more distinct in C. annabella, otherwise as in C. carye.

Forewing underneath with the pale yellowishwhite postmedial and marginal bar slightly larger

in C. annabella; with dark spot at base of interspace

Cux smaller and usually separate from postmedial

black bar in this interspace in C. carye, while in

C. annabella it is larger and always connected to the

postmedial black bar.

Hindwing underneath with whole aspect lighter

in color in C. annabella; with pure white spot in

middle of wing opposite end of cell more triangular

in shape in C. annabella and somewhat hourglass

shaped in C. carye.

Length of forewing, 17-27 mm (average 23 mm).

Male genitalia as illustrated by Figure 15 (drawn

from my preparation no. 3716), with uncus in lateral

view downward curved at distal end, in ventral view

large and triangular, with distal end acute and

greatly extended; gnathos with lateral arms smaller

than those of C. carye, gradually curved upward

along dorsal margin and abruptly bent along ventral margin, forming a distinct "heel" near middle

of this margin; ventral process of lateral arms not

showing in lateral view; valva with a large rounded

lobe on middle of dorsal margin, with a large acute

lobe at junction of dorsal and distal margins, instead of the large clublike process found in C. carye,

margin beyond this acute lobe short and nearly

straight; lower margin of valva gradually and

broadly rounded, forming nearly a right angle at

junction of distal margin; clasper a large flaplike

lobe, directed toward lower angle (junction of

distal and lower margins); cuiller heavily armed

with large spinelike teeth along inner surface;

aedeagus sharply constricted just before middle of

ventral margin so that distal half is very thin and

needle-like, base thick and slightly downward bent;

saccus with a caudal element that projects outward

a great distance past base of lower margin of valva.

FEMALE (Figures 155-160).—Not distinguishable

from the male sex except by examination of the

genitalia where the large podlike ostium bursae

lobe can be seen after the scales are brushed off of

the underside of posterior surface of abdomen.

Length of forewing 17—26 mm (average 22.8 mm).

Female genitalia as illustrated by Figure 31

(drawn from my preparation no. 6445), differing

from all other species of Cynthia, except C. carye,

in lacking the undulated margin between seventh

and eighth sternites and in having the seventh

sternite projected posteriorly near middle; it differs

also from those species in having two heavily

sclerotized laterally placed ridges near spiracles of

seventh sternite and in having the eighth sternite

heavily sclerotized with a very large podlike ostium

bursa lobe, this podlike lobe not swollen at its base

as is C. carye.
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

METHOD OF IDENTIFICATION.—When it was discovered that there were two distinct species going

under the name of C. carye Hiibner, a North American species and a South American species, an unsuccessful effort was made to locate the type material of C. carye. Unfortunately none could be

found. The lack of this material was not of great

importance as Hiibner's original colored figures of

the upper and under surfaces ([1812], pi. [45], figs.

1, 2) were found to quite accurately, and in fact

beautifully, portray his species. This turned out to

be the one found in the Andes of Peru, in Bolivia,

Brazil, Chile, Uruguay, and Argentina. See further

notes under C. carye.

TYPE LOCALITY.—First valley west of Arroyo

Verde Park, Ventura, Ventura County, California.

ADDITIONAL TYPE DATA.—Described from the holotype male (USNM 71044), allotype female, and

five male and fifteen female paratypes with data

as follows: holotype, locality as given above, elevation 500 feet, reared from last instar larva, emerged

17-24 January 1965, Thomas Dimock collector;

allotype, same data as holotype except emerged

21-31 March 1965; one male and six female paratypes, same data as holotype; two male and three

female paratypes, same data as allotype; two female

paratypes, same data as holotype except emerged

February 1965; one female paratype, same data as

holotype except collected in flight, not reared, 22

March 1965; one female paratype, collected in

flight, above Arroyo Verde Park, elevation 780 feet,

other data as in holotype; two male and two female

paratypes, Arroyo Verde Park, Ventura, Ventura

County, California, elevation 750 feet, 27 March

1970, Thomas Dimock.

LOCATION OF TYPES.—Holotype (USNM 71044),

allotype, and one pair of paratypes in the National

Museum of Natural History, one pair of paratypes

in the collection of the California Academy of Sciences (San Francisco). The remainder of the paratypes to be deposited in various museums by

Thomas Dimock of San Jose and Ventura, California.

LIFE HISTORY.—The larvae of this species have

been reported feeding upon the following Malvaceae: Althaea, Lavatera assurgentifiora, Malva

horealis, M. parviflora, Malvastrum exile, M. fasciculatum, and Sphaeralcea ambiqua. Larvae have

also been reported feeding upon Ligustrum (Oleaceae, order Oleales), Lupinus arborens (Leguminosae, order Rosales), and Urtica holosericea (Urticaceae, Urticales).

Notes on the life history and immature stages

have been given by Behr (1863, p. 125), Edwards

(1874, p. 329), Dyar (1889, pp. 237-238), Coolidge

(1925, pp. 146-147), Huguenin (1926, pp. 216-217),

and Comstock (1927, pp. 134-135). References to

the literature on the life history and immature

stages were given by Edwards (1889, p. 26) and

Davenport and Dethier (1938, p. 164).

Mr. Thomas Dimock, the collector of the type

series, has offered (in a letter) the following information about the type locality and the species

association therein:

Most of the specimens were taken in the first valley west (by

about one-eighth of a mile) of a city park named Arroyo

Verde Park (Ventura, Ventura County, California). This

place is located in the foot-hills north of eastern Ventura,

and about a half mile from my residence. Costal sage brush

associations are on most northerly facing slopes, and a "weedy

annual grassland" group occupies the south-facing exposed

areas. In this valley west of the Park, mallow, the imported

annual "weed", usually grows lushly on the flat valley bottom,

provided cattle aren't pastured there, and the larvae of this

butterfly can be found in abundance in very early spring

(January to March), occupying the exact same niche as

cardui, also on mallow. When the adults emerge they hilltop.

Both species, and less frequently atalanta, can be found all

year round on some of the hilltops, which are mostly 800 to

1200 feet in elevation. There are two main broods (both

species) : a late January and early February brood, and a late

February-March brood, after which both are less common.

In fall and early winter fresh adults of "carye" become slightly

more frequent in backyards.

DISTRIBUTION.—Vancouver Island and the mainland of British Columbia east into Alberta and

south through California and Mexico (all parts

according to Hoffmann, 1940, p. 681) into Guatemala, and from California east through Nevada,

Utah, Montana, Wyoming, Colorado, Arizona, New

Mexico, and El Paso, Texas. It has strayed into

Scott County, Kansas, from Colorado.

MATERIAL STUDIED.—In addition to the type series

recorded above, I have studied one hundred eightytwo males and one hundred and seven females from

the following localities: ALBERTA: near Mount Assinboine (August). WASHINGTON: Bellingham;

Brewster (August); Godman Springs, Blue Mountains (July); Seattle (July); Seguin (September) ;

Toroda (August). OREGON: Newport (September).
NUMBER 84 59

CALIFORNIA: Alameda County (January, March,

September, December); Albany, Alameda County

(June); Albany Hill, Contra Costa County

(March); Alum Rock Park, Santa Clara County

(May); Antioch, Contra Costa County (May, October) ; Beverly Hills; Berkeley, Alameda County

(March, May) ; Bethel Island, Contra Costa County

(May); Buttonwillow, Kern County Qune);

Cherry Flat, Santa Clara County (September);

Chula Vista, San Diego County (January, March,

April, May, August, September, October, December) ; Claremont (April); Concord (May, October) ; Davis (October); Deerpark, Placer County

(August, 6,200 ft); Fairfax, Marin County (May);

Fillmore (June) ; Glen Alpine Creek, El Dorado

County (July) ; Glendora (February); Golden Gate

Park, San Francisco (January, February); Griffith

Park, Los Angeles County (February); Humbug

Mountain, Siskiyou County (September); Indian

Flat, Mariposa County (May); Inverness, Marin

County (August); Laguna Beach (July) ; Lake Alpine, Alpine County (July); Lake Chabot Road,

Alameda County (May) ; Las Uvas, Santa Clara

County (June) ; Los Alamitos (July); Los Angeles

(January, February, June, November) ; Mariposa

Grove, Mariposa County; "Middle California"; Mill

Valley (December); Mitchell Canyon, Contra Costa

County (May) ; Mocho Canyon, Livermore (April);

Mokel Hill; Mount Bruno Ridge, San Mateo

(April); Napa County (September); Novato,

Marin County (February); Oakland Hills, Alameda

County (September) ; Ontario (March); Palo Alto

(May, September, November) ; Pasadena (June);

Petaluna (June); Pleasant Hill, Contra Costa

County (March—July); Plumas County; Redwood

Canyon, Contra Costa County (June, July); San

Diego (September) ; San Francisco (January—March,

May, July—October, December) ; San Jose; San Juan

(August); San Leandro, Alameda County (May,

June—August) ; San Mateo County; Santa Barbara

Islands; Santa Cruz (May, June, August, September,

October, December); Santa Rosa, Riverside County

(April); Sherman Island, Sacramento County

(September); Sierra County; "Southern California";

Warner Mountains, Modoc County (July); Yosemite Valley, Yosemite National Park (August).

NEVADA: Del Monte Ranch, Reno (August); Reno

(September). UTAH: Bellevue, Washington County

(June, 4,000 ft) ; Vineyard (July). MONTANA: Deerlodge, Powell County (August). COLORADO. NEW

MEXICO: Bear Trap Camp, 26 miles southwest,

Magdalena, Socorro County (July, 8,500 ft); Cedar

Creek Camp, 2 miles north Ruidosa, Lincoln

County (June, 7,000 ft); McMillan Camp, 13 mites

north Silver City, Grant County (July, 6,800 ft);

Mogollon, Catron County (July, 7,200-7,600 ft);

Pine Camp, 2 miles northeast Cloudcroft, Otero

County (July, 8,600 ft); Signal Peak, 22 miles northeast Silver City, Grant County (July, 8,600-8,900

ft) ; Willow Creek Ranger Station, 25 miles east

Alma, Catron County (July, 8,000 ft). ARIZONA:

Alpine Divide Camp, 4 miles north Alpine, Apache

County (July, 8,500 ft); Chiricahua Mountains

(May, 9,000 ft); Coconimo County, 17 miles north

Williams (August, 6,700-6,800 ft); Flagstaff (July,

August, 7,000 ft); Graham Mountains, Graham

County (September, 9,000 ft); Jacob Lakes, Coconino County (July); Keyanta, Navajo County

(July); North Rim, Grand Canyon (July, 8,000-

9,000 ft); Pinal County, near Superior (August) ;

Slade Ranch, near Greer (July). TEXAS: Alpine,

Brewster County; El Paso. MEXICO: Chiapas; Jalapa

(February); La Boguilla (July) ; Llamas Penon,

Federal District (May); Mexico City (June, November) ; Morelia, Micoacan (March); Presidio,

Vera Cruz (May, June); San Pedro de los Pinos,

Federal District (October); Tuxtepec, Oaxaca

(June); Yerbanis, Cuencame', Durango (August,

6,700 ft). GUATEMALA: Mpio, Acatenango, Quisache,

Chimaltenango (December, 1,750 m).

9. Cynthia carye Hiibner, new combination

FICURES 16, 32, 137-142, 149-154

Hamadryas carye Hubner, [1812], pi. 45: figs. 1, 2.

Vanessa carye.—Hubner, 1819, p. 33.—Williams, 1930, p. 231.

—Ureta, 1938b, p. 298; 1938c, p. 127; 1939, p. 231.—

Gutierrez, 1940, p. 34.—Bridarolli, 1944, p. 32.—Breyer,

1945, p. 312.—Zischka, 1947, p. 32.—Verity, 1950, p. 329.—

Hayward, 1951, p. 195.—Herrera, 1954, pp. 45, 53, pi. 6:

fig. 4.—Herrera and Etcheverry, 1956, pp. 277, 285.—Biezanko, Ruffinelli and Carbonell, 1957, p. 124.—Herrera,

Etcheverry and Barrientos, 1958, pp. 240 (fig. 1) , 244, 246-

250, 247 figs. 11-14), 248 (figs. 18, 19), pi. 1: figs. 1, 2 —

Niculescu, 1965, p. 191.—Hayward, 1966, p. 69.

Pyrameis carye.—Doubleday, 1849, p. 205.—Kirby, 1871, p.

186.—Berg, 1875, p. 70; 1876, p. 201.—Burmeister, 1878,

pp. 151-152.—Butler, 1881, p. 466.—Staudinger, 1885, pp.

97, 98.—Bartlett-Calvert, 1886, p. 314.—Weymer and Maassen, 1890, pp. 14, 41, 97.—Godman and Salvin, 1891, p.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

101.—Staudinger, 1894, p. 70.—Izquierdo, 1895, p. 823, pi.

4: figs. 4, IS.—Bartlett-Calvert, 1898, p. 99.—Porter, 1899a,

p. 36.—von Bayeren, 1901, p. 257.—Rebel, Weymer and

Stichel, 1901, p. 305.—Elwes, 1903, p. 287.—Reuss, 1910a,

pi. 1: figs. 2, 6, 10; 1910b, pp. 64, 66; 1910c, pp. 86, 89, 90.

—Seitz, 1914a, p. 459, pi. 94: figs, A 2, A 3; 1914b, p. 459,

pi. 94: figs, A 2, A 3.—Giacomelli, 1922, pp. 203-204.—

Porter, 1930, p. 279.—Hayward, 1931, pp. 69, 186, pi. 12:

fig. 4.—Ureta, 1934, p. 79; 1935, p. 87; 1938a, p. 181.

Pyrames [sic] carye.—Porter, 1899b, p. 181.

Cyntia [sic] carye.—Costa Lima, 1923, p. 150; 1928, p. 135.

Vanessa charie Blanchard [misspelling of carye], 1852, p. 26,

pi. 2: fig- 5.—Hayward, 1951, p. 195 [synonym of carye].

Pyrameis charie.—Reed, 1877, p. 679.—Gazulla and Ruiz,

1928, p. 290.—Hayward, 1931, p. 69 [synonym of carye].

Vanesja [sic] charie.—Philippi, 1859, p. 1089.

Payrameis [sic] charie.—Reed, 1877, p. 735.

Vanessa charye Porter [misspelling of carye], 1898, p. 32.

Pyrameis caryae Verity not Hubner [misspelling of carye],

1916b, p. 128.—Hayward, 1931, p. 69 [synonym of carye, a

misidentification of a misspelling by Holland]; 1951, p. 195

[as before].

Pyrameis caryoides Giacomelli [new synonymy], 1922, pp. 203-

205, fig. [3].

Pyrameis carye aberration caryoides.—Hayward, 1931, p. 70.

Vanessa carye aberration caryoides.—Hayward, 1931, p. 196.

Pyrameis carye form minscula Hayward [excluded name, type

3], 1931, pp. 69-70, pi. 12: fig. 5.

Pyrameis carye minuscula Hayward, 1931, p. 186 [corrected

spelling of minscula]; 1951, p. 196.

Pyrameis caryae aberration bruchi Kohler [excluded name,

type 2], 1945, p. 256, pi. 20: fig. 1.

Vanessa carye aberration bruchi.—Hayward, 1951, p. 196.

C. carye and C. annabella as mentioned in the description of the latter are easily distinguished from

all other species of Cynthia, except C. terpsichore

and some females of C. virginiensis, in having the

bar just beyond cell on upper side of forewing tawny

instead of white. They are easily distinguished from

C terpsichore and C. virginiensis by having four,

instead of two, blue-centered ocular markings on

the upper side of hindwing.

MALE (Figures 137-142).—Forewing above with

tawny subapical bar just beyond cell slightly smaller

in C. carye; subapical white spot at costa and interspaces R.! and R2 slightly smaller than in C. annabella; basal transverse bar in cell less distinct in

C. carye than in C. annabella; bar below base of

Cu2 usually very small, sometimes absent in C. carye,

present and distinct in C. annabella; with the small

tawny spot in interspace M3 just above the large

tawny quadrate spot in interspace Cux found in

C. annabella, absent in most specimens of C. carye;

submarginal white line composed of three slender

bars on apex of forewing much less distinct in C.

carye than in C. annabella.

Hindwing above with white spot in middle of

costal margin smaller and less distinct in C. carye

than in C. annabella, otherwise as in C. annabella.

Forewing underneath with the pale yellowishwhite postmedial and marginal bar slightly smaller

in C. carye; with dark spot at base of interspace Cut

smaller and usually separate from post-medial black

bar in this interspace in C. carye, while in C. annabella it is larger and always connected to the postmedial black bar.

Hindwing underneath with whole aspect darker

in color in C. carye; with pure white spot in middle

of wing opposite end of cell somewhat hourglass

shaped in C. carye, more triangular in C. annabella.

Length of forewing, 20-27 mm (average 22.7 mm).

Male genitalia as illustrated by Figure 16 (drawn

from my preparation no. 3660), with uncus in lateral view very similar to that of C. annabella, a

little thicker, especially just before distal end, in

ventral view constricted before distal end and thus

not so clearly triangular in shape; gnathos with

lateral arms very large, with distal one third sharply

bent upward toward uncus, with lower margin of

these arms having a "heel" near middle of relatively straight portion of lower margin; ventral

process of lateral arm of gnathos bent inward so

that it shows in lateral view; valva with dorsal margin nearly straight or only slightly curved upward

near middle; valva with a large club-shaped process

at junction of dorsal and distal margins instead of

the acute lobe found in C. annabella, this club with

a blunt toother lobe dorsally near its base; distal

margin of valva greatly concave because of this clublike process; lower margin of valva gradually and

broadly rounded with a large acute lobe at junction

of distal and lower margins; clasper similar to but

not as large as that of C. annabella; cuiller also

similar to that species but much more heavily armed

with large teeth; aedeagus similar to that of C. annabella, with base longer; saccus also similar to the

latter species.

FEMALE (Figures 149-154).—As in C. annabella

this species differs from C. carye in the same charac-
NUMBER 84 61

ters as do the males and is not distinguishable from

the male sex except by examination of the genitalia,

where the large podlike ostium bursa lobe can be

seen after the scales are brushed off the underside

of posterior surface of abdomen.

Length of forewing, 15-29 mm (average 25.4 mm).

Female genitalia as illustrated by Figure 32,

(drawn from my preparation no. 3678), similar to

those of C. annabella, differing in the shape of the

large podlike lobe at ostium bursa, which in this

species is swollen at the base.

METHOD OF IDENTIFICATION.—See notes under this

heading in the description of C. annabella. The

lack of type material of C. carye and the fact that

Hiibner did not cite a type locality were found not

to be a deterrant in the identification of this species,

since Hubner's original colored figures of both

surfaces ([1812], pi. [45], figs. 1, 2) were found to

beautifully and quite accurately portray C. carye

as the species found in the Andes and farther south

in South America. Hubner's figures agree with the

population just mentioned in the following nine

particulars: (1) The subapical white spot on forewing above in interspaces R, and R3 is smaller than

it is in C. annabella. (2) The basal transverse bar

in forewing cell is smaller than in C. annabella.

(3) The tawny spot found in C. annabella just

above the large quadrate spot in interspace Cux is

absent in Hubner's figure as it is in the South

American population. (4) The white spot in

middle of costal margin of upper surface of hindwing is absent or greatly reduced as it is in the

South American species, while in C. annabella it is

never absent and is always larger and more distinct

than in the latter. (5) On the under surface of

forewing the pale yellowish-white postmedial bar

is small in Hubner's figure as it is in the South

American species. In the North American species

this bar is distinctly larger. (6) On this same surface the postmedial fuscous bar ends at 2n A as it

does in all of the South American specimens I have

seen. In the North American species this bar continues into interspace 2nd A. (7) The white spot

in middle of hindwing opposite the end of cell on

the under surface is hourglass shaped in Hubner's

figure. This is true also of the South American

species, while in C. annabella this spot is more

triangular in shape. (8) On the under surface the

apex of hindwing is quite dark in Hubner's figure,

with a cheveron-shaped terminal line in interspaces

Rg. In the North American species this area is

yellowish to white in color and lacks a cheveronshaped terminal line in interspace R8. (9) The

tawny spot found in C. annabella just above the

large quadrate spot in interspace Cux, on the under

surface of the forewing, is absent in Hubner's

figure as it is in the South American species.

According to Horn and Kahle (1935, p. 119) the

Hiibner collection went to the Zoologische Abteilung, Naturhistorisches Museum, Vienna. Dr. F.

Kasy of that institution informs me that there are

only a few Hiibner types remaining in their collection, all of Heterocera. The butterfly types and

other missing Hiibner types were probably destroyed by fire in 1848. Because of the excellent

original colored figures, I see no need to designate

a neotype.

LIFE HISTORY.—The larval food plants are Malvaceae: Althaea rosea, Malva nicacensis, M. parviflora, Modiola caroliniana, Sphaeralcea bonariensis,

and S. obtusiloba; Compositae: Cynara cardunculus,

Gaillardia scabiosoides, and Pascalia glauca; Geraniaceae: Pelargonium pelatatum and P. zonale. It

has been reported upon Urtica urens (Urticaceae)

but this was perhaps a forced rearing.

The egg, larva, and pupa have been described by

Burmeister (1878, p. 152), Izquierdo (1895, p.

832), and Hay ward (1931, p. 70).

DISTRIBUTION.—This species is found from the

Andes in Colombia, Ecuador, Peru, and Bolivia

south through Chile and in southwestern Brazil,

Paraguay, and Uruguay south through much of

Argentina. It also occurs in the Juan Ferndndez

Islands of Chile, on Easter Island, and on Mangareva Island of the Gambier Islands, of the

Tuamotu Archipelago on the Tropic of Capricorn,

over halfway to New Zealand from Chile.

MATERIAL STUDIED.—Forty-two males and fortyseven females were studied from the following

localities: COLOMBIA: Guambia,Cauca (December).

ECUADOR: Oriente; Salcedo (March, 2,700 m).

PERU: Chichicastenango (September); Cuzco (July,

11,500 ft); Ollantaitambo, Cuzco (March, 9,200 ft);

Oroya (May); Pisco (May); Puno (March, April);

Usaquen, Cundiamarca (January, 2,800-2,900 ft).

BOLIVIA: Belen Experiment Station (May); La Paz

(April, May, 12,000 ft); Neguejahuica (May, 8,000

ft); Okala-Ankoma (April, 7,500-13,700 ft) ; Oruro
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(March, April, 12,500 ft); Oruro (March, April,

12,500 ft) ; Pitiguaya (May, 5,800 ft); Playa Verde

(December, 13,000 ft). CHILE: Angol (JanuaryMarch) ; Criados, Province Tarapaca (February,

October, November); Huanta, Conquimbo (December) ; Iquiqui, Province Tarapaca (September,

October, November); La Junta, Coquimbo (December) ; Los Lagos; Olmul (March); Macul (March);

Maipu; Pucon (February); San Carlos, Nuble

(December); Santiago; Talca, 22 miles north

(December); Temuco; Valparaiso (January, August) ; Tofo (January, Apirl, November). BRAZIL:

Castro Parana; Ponta Grossa, Parana (August).

URUGUAY: Carmen, Durazno; Montevideo (November) . ARGENTINA: Cordova; La Rioja; Punta Lara,

La Plata (November); Tucumen. JUAN FERNANDES

ISLANDS (CHILE) : Bahia Cumberland, Masatierra

(January, March); Las Chozas, Masafuera (January) . EASTER ISLAND (CHILE) : Hanga Rosa (January) . TUAMOTU ARCHIPELAGO: Rikitea Village,

Mangarewa Island, Gambier Islands (December).

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PLATES

NUMBER 84 79

1 Vanessa atalanta

2 Vanessa tameamea

FIGURES 1-2.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventral viewa; aedeagus in lateral view. 1, Vanessa atalanta (drawn from

preparation number 6179); 2, Vanessa tamzamea (drawn from preparation number 3657).
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3 Vanessa indica

4 Vanessa dejeanii

\.

FIGURES 3-4.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventral view aedeagus in lateral view. 3, Vanessa indica (drawn from

preparation number 3649); 4, Vanessa dejeanii (drawn from preparation number 3653).
NUMBER 84 81

5 Vanessa samani

Bassaris itea

FIGURES 5-6.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventral view; aedeagus in lateral view. 5, Vanessa samani (drawn from

preparation number 6281); 6, Bassaris itea (drawn from preparation number 3655).
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

7 Bassaris gonerilla

8 Cynthia cardui

FIGURES 7-8.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventra view; aedeagus in lateral view. 7, Bassaris gonerilla (drawn from

preparation number 6192); 8, Cynthia cardui (drawn from preparation number 6104).
NUMBER 84 83

9 Cynthia kershawi

10 Cynthia virginiensis

t K.LRF.S y-10.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventral view aedeagus in latera view. 9, Cynthia kershawi (drawn from

preparation number 6098); 10, Cynthia virginiensis (drawn from preparation number 6122).
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

11 Cynthia altissima

12 Cynthia braziliensis

FIGURES 11-12.—Male genitalia consisting of lateral view (on the left side of each figure) of tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral view;

gnathos and uncus in ventral view; aedeagus in lateral view. 11, Cynthia altissima (drawn from

preparation 6141); 12, Cynthia braziliensis (drawn from preparation 3663).
NUMBER 84 85

13 Cynthia terpsichore

14 Cynthia myrinna

FIGURES 13-14.—Male genitalia consisting of lateral view (on the left side of each figure) of

tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral

view; gnathos and uncus in ventral view; adeagus in lateral view. 13, Cynthia terpsichore (drawn

from preparation number 3665); 14, Cynthia myrinna (drawn from preparation number 6147).
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

V 'V /

15 Cynthia annabella

16 Cynthia carye

FIGURES 15-16.—Male genitalia consisting of lateral view (on the left side of each figure) of

tegumen, vinculum, uncus, gnathos, and the right valva; ventral element of anellus in ventral

view; gnathos and uncus in ventral view; aedeagus in lateral view. 15, Cynthia annabella (drawn

from preparation number 3716); 16, Cynthia carye (drawn from preparation number 3660).
17 Vanessa atalanta

19 Vanessa indica

18 Vanessa tameamea

FIGURES 17-19.—Ventral aspect of seventh and eighth abdominal segments and ovipositors showing female genitalia consisting of genital plates, ostium bursae, ductus bursae, bursa copulatrix,

and signa. 17, Vanessa atalanta (drawn from preparation number 3667); 18, Vanessa

tameamea (drawn from preparation number 6265); 19, Vanessa indica (drawn from preparation

number 6298).
00

00

22 Bassaris itea

20 Vanessa dejeanii

FICURES 20-22.—Ventral aspect of seventh and eighth abdominal segments and ovipositors

showing female genitalia consisting of genital plates, ostium bursae, ductus bursae, bursa copulatrix, and sign a. 20, Vanessa dejeanii (drawn from preparation number 6257); 21, Vanessa samani

(drawn from preparation number 6282) ; 22, Bassaris itea (drawn from preparation number 6273).
23 Bassaris gonerilla

24 Cynthia cardui

25 Cynthia kershawi

FIGURES 23-25.—Vental aspect of seventh and eighth abdominal segments and ovipositors showing

female genitalia consisting of genital plates, ostium bursae, ductus bursae, bursa copulatrix,

and signa. 23, Bassaris gonerilla (drawn from preparation number 6194); 24, Cynthia cardui

(drawn from preparation number 3675); 25, Cynthia kershawi (drawn from preparation number

6071).
26 Cynthia virginiensis

27 Cynthia altissima

28 Cynthia braziliensis

FIGURES 26-28.—Ventral aspect of seventh and eighth abdominal segments and ovipositors showing female genitalia consisting of genital plates, ostium bursae, ductus bursae, bursa copulatrix,

and signa. 26, Cynthia virginiensis (drawn from preparation number 3679); 27, Cynthia altissima

(drawn from preparation number 6465); 28, Cynthia braziliensis (drawn from preparation

number 3680).
30 Cynthia myrinna

29 Cynthia terpsichore

31 Cynthia annabella

32 Cynthia carye

FIGURES 29-32.—Ventral aspect of seventh and eighth abdominal segments and ovipositors showing female genitalia consisting of genital plates, ostium bursae, ductus bursae, bursa copulatrix,

and signa. 29, Cynthia terpsichore (drawn from preparation number 6438); 30, Cynthia myrinna

(drawn from preparation number 6441); 31, Cynthia annabella (drawn from preparation

number 6445); 32, Cynthia carye (with ductus bursae and bursa copulatrix removed, drawn

from preparation number 3678). to
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURES 41-49.—Vanessa atalanta atalanta: 41, 9 (Friedland, Czechoslovakia) ; 42, 9 (Baja,

Hungary); 43, 9 (Segovia, Spain). Vanessa atalanta rubria: 44, 9 (Eureka, Kansas); 45, 9

(Salem, Virginia); 46, 9 (Palm Beach, Florida). Vanessa indica indica: 47, $ (Teesta Valley,

Dikkim, India); 48, $ (Coorg, India); 49, 9 (Darjeeling, India).
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURES 58-65.—Vanessa indica vulcania: 58, 9 (Canary Islands); 59, underside of 58; 60, $

(Madeira Islands); 61, underside of 60; 62, 9 (Madeira Islands); 63, underside of 62. Vanessa

indica buana: 64, $ (Bonthain Mountain, south Celebes, 4,000 ft); 65, underside of 64.
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURES 74-80.—Vanessa tameamea: 74, 9 (Tantalus, Oahu, Hawaii); 75, underside of 74.

Vanessa samani: 76, 9 (Kara Mountains, Sumatra, from lithograph published by Hagen, 1896);

77, $ (Sungei, Sumatra); 78, underside of 77; 79, 9 (foot of Korintji Peak, Sumatra); 80,

underside of 79.
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B I 0 0 5

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FIGURES 89-96.—Cynthia cardui, males: 89 (Friedland, Czechoslovakia) ; 90, underside of 89; 91

(Spetsai Island, Greece); 92, underside of 91; 93 (Japan); 94 (Madagascar); 95 (Washington,

District of Columbia); 96, underside of 95.
FIGURES 97-104.—Cynthia cardui, females: 97 (Friedland, Czechoslovakia); 98, underside of 97;

99 (Spetsai Island, Greece); 100, underside of 99; 101 (Oeda, Japan); 102 (Madagascar); 103

(Washington, District of Columbia); 104, underside of 103.
FIGURES 105-116.—Cynthia kershawi: 105, $ (Colo Vale, New South Wales, Australia); 106,

underside of 105; 107, $ (Birchip, Australia); 108, underside of 107. Cynthia virginiensis: 109,

$ (Princess Anne, Virginia); 110, underside of 109; 111, <j> (South Carolina); 112, underside

of 111; 113, $ (Allegany State Park, New York); 114, $ (Princess Anne, Virginia); 115, $

(Rio de Janeiro, Brazil); 116, underside of 115.
FIGURES 117-126.—Cynthia virginiensis: 117

, $ (Sherando Lake, Virginia); 118, $ (Pluckemin,

New Jersey). Cynthia altissima: 119, $ (Agulani, Peru, 9,000 ft, syntype); 120, underside of

119; 121, $ (Gusilluni, Bolivia); 112, underside of 121. Cynthia braziliensis; 123, $ (Brazil);

124, $ (Oroya, Peru); 125, underside of 124; 126, $ (Boliva).

noH»cHO

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FICURES 127-136.—Cynthia braziliensis: 127, $ (Blumenau, Santa Catarina, Brazil); 128, $

(Pelota, Rio Grande do Sul, Brazil); 129, ° (Montevideo, Uruguay); 130, $ (Cuzco, Peru).

Cynthia terpsichore: 131, $ (Andes of Chile, 6,000 ft); 132, underside of 131; 133, $ (Chile);

134, underside of 133. Cynthia myrinna: 135, $ (Santa Catarina, Brazil); 136, $ (Santa Catarina, Brazil). S
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURES 149-160.—Cynthia carye, females: 149 (Angol, Chile); 150, underside of 149; 151 (Belen

Station, Atchacachi, Bolivia); 152, underside of 151; 153 (Masafucra, Las Chozas, Juan Fernandez

Islands, Chile); 154, underside 153. Cynthia annabella, females: 155 (allotype, Arroyo Verde

Park, Ventura, California); 156, upper side of 155; 157 (San Francisco, California); 158, upper

side of 157; 159 (Godman Springs, Blue Mountains, Washington); 160, upper side of 159.

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(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

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"Figure 96." If illustrations (usually tone photographs) are printed separately from the text as

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