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*IS',7&OS

Volume 13 Number 1 March, 1974

published by

The Lepidoptera Research Foundation, Inc.
at

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Edited by WILLIAM HOVANITZ

Volume 12

1973

published by

The Lepidoptera Research Foundation, Inc.
at

1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006

Journal of Research on the Lepidoptera

13(l):l-20, 1974

1160 W. Orange Grove Avc., Arcadia, California 91006, U.S.A.
© Copyright 1975

STUDIES ON NE ARCTIC EUCHLOE

PART 7.

COMPARATIVE LIFE HISTORIES, HOSTS AND THE
MORPHOLOGY OF IMMATURE STAGES

PAUL A. OPLER ^

Division of Entomology,

University of California,

Berkeley, California

This part of the series deals with certain aspects of the life
history of the Nearctic Eiichloe. Although the 'life histories”
of all except Eiichloe creiisa have appeared in the literature, no
comparisons have been made. This study emphasized the im-
mature stages, important differences in oviposition sites, larval
behavior, larval color patterns, larval setation, and pupal con-
figuration.

LIFE HISTORY

Host. — All Euchloe oviposit and feed upon maturing plants
of the family Cruciferae. The food plants utilized are the result
of selection by ovipositing females.

Among the species of Cruciferae which occur at any locality,
female Euchloe may select more than one species as suitable
hosts, but never appear to utilize all of them. Those species
selected must have had several prior requirements. These are:
( 1 ) the plant should grow in full sunlight, ( 2 ) the plant should
be more than five inches in height with an erect configuration,
and ( 3 ) the clusters of unopened flower buds should be terminal
and tightly clustered (except some Euchloe hyantis hosts).

The role of allyl isothiocyanate levels in determining the
suitability of various crucifers as hosts for Euchloe is unknown.

The groups of hosts used by most, well-observed species are
broad. Qualitative differences in host utilization are pronounced

1

2

PAUL OPLER

/. Res. Lepid.

only between the two species groups. The three members of the
“Ausonides group,” i.e. Euchloe ausonides, E. creusa, and E.
olympia, tend to utilize species of Arabis and “mustard-like”
crucifers, i.e. Arabis glabra, Barbarea, Brassica, and Sisymbrium,
while the “Hyantis group,” i.e. Euchloe hyantis, tends to utilize
Caulanthus, Desciirainia, and Streptanthus with greater fre-
quency.

That host specificity levels within the Cruciferae are broad
is shown by the frequent utilization of introduced species of
the family. Swales (1966) briefly discusses the range extension
of some pierids due to the introduction of crucifers into dis-
turbed areas around villages in Arctic Canada. Opler (1969)
explained the role of introduced species of Brassica as a pos-
sible factor permitting the facultative bivoltinism of Euchloe
ausonides in coastal central California.

All pertinent locality and host data for all Nearctic Euchloe
are summarized in Table 1.

Oviposition site. — Members of the “Ausonides group” almost
always place their eggs on unopened flower buds of the host.
This behavioral feature has been reported for E. ausonides by
Coolidge and Newcomer (1908) and Remington (1952), while
Shull (1907) and Meiners (1938) reported the same pattern for
E. olympia. 1 have observed many hundreds of eggs of E.
ausonides and only rarely were eggs found anywhere but on
unopened buds. Eggs of Euchloe creusa near Banff were all
found on unopened flower buds of Draba lanceolata.

Although Euchloe hyantis females may select flower buds as
oviposition sites, other portions of the plant are utilized with
greater frequency. Coolidge ( 1925 ) reported that eggs of Euchloe
hyantis lotta were found on flower buds, leaves, or the stems of
host plants. Near El Portal, Mariposa County, California, I ob-
served that the eggs of Euchloe hyantis were most frequently
placed on the lower surface of the saggitate clasping leaves of
Streptanthus polygaloides.

Larval behavior. — Upon emerging from the egg, larvae of
Euchloe seek unopened flower buds upon which to feed. J. A.
Scott, Division of Entomology, University of California, Berke-
ley, (Personal communication) states that first instar larvae of
Euchloe ausonides die if fresh unopened buds are not available.
Upon hatching, larvae of E. hyantis, emerging from eggs laid on
leaves or stems of the host, must find the reproductive portions
of the plant soon after hatching. To do this the larvae make

13(1):1~20, 1974

NEARCTIC EUCHLOE

3

TABLE I.

HOST AND LOCALITY DATA FOR VARIOUS SPECIES OF EUCHLOE

Euchloe ausonides

Arabia dri-umnondi Gray

Canada, Ontario, Geraldton Forest (Thunder Bay District), 28
June 1966. J.C.E. Riotte. Jour. Lepid. Soc., 22(1): a (1968).
Colorado, Boulder County, Nederland 6 July 1949* C.L. Remington.
Psyche, 59(2): 63 (1952).

Colorado, Gimnison County, Gothic, 9600', 10 July 1967, J. Emmel
and 0. Shields. J. Res. Lepid., 8(1) :31 (1970).

Arabia fendleri (Wats.) Greene var. soatifolia (Rvdb.) Rollins

Colorado, Boulder County, Nederland, 6 July 1949, GL. Remington.

Psyche, 59(2): 63(1952).

A rabi s glabra (L.) Berh.

Colorado, Boulder Coimty, Spring Gulch, 7 July 1949, C.L. Remington.
Psyche, 59(2) :62 (1952).

California, Contra Costa County, Berkeley Hills, NE Oakland,

3 June 1963 J.A. Powell. Preserved larvae in California Insect Survey.

Barbarea vulgaris (L.)

California, Contra Costa County, Russelmann Park, Mt. Diablo,

24 April 1966, P.A. Opler. Personal observation.

Brassica campestris L.

California, Contra Costa County, Briones Regional Park, 28 March
1970, P.A. Opler. Personal observation.

Brassica kaber (D.C.)

California, Alameda County, Strawberry Canyon, 18 April I964,

P.A. Opler, Personal observation.

California, Contra Costa County, Briones Hills Regional Park,

28 March 1970, P.A. Opler. Personal observation.

California, Contra Costa County, Tilden Regional Park, 18 April
1968, P.A. Opler. Personal observation.

California, Santa Clara County, Alum Rock Park, 28 February 1964, P.A.
Opler. Personal observation.

Brassica nigra (L.) Koch

California, Alameda Coimty, Strawberry Canyon, 10, 15 April 1970,

P.A. Opler. Personal observation.

California, Contra Costa Coiinty, Pt. Richmond, 18 April 1969,

J.A. Scott, (preserved larvae).

California, Santa Clara County, New Almaden, 4 May I964, P.A. Opler.
Personal observation.

Descurainia Californica (Gray) Shultz

Colorado, Gunnison County, Schofield Pass, 10,400', I4, 18 July
1967, J. Emmel and 0. Shields. J. Res. Lepid., 8(1) :31 (1970).

4

PAUL OPLER

/. Res. Lepid.

Erysimum capitatum (Dougl.) Greene

Colorado, Boulder Coimty, Nederland, 6 July 1949, C.L. Remington.
Psyche, 59(2): 63 (1952).

I satis tinctoris. L.

California, Modoc County, Buck Creek Ranger Station, 11 June 1970, P.A.
Opler. Personal observation.

Raphanus sativa L.

California, Contra Costa County, Briones Regional Park, 28 March 1970,
P.A, Opler. Personal observation.

Si symbrium altissimum L.

Colorado, Boulder County, Boulder Canyon, 6500', Mt. Flagstaff,
Nederland, 8-9 July 1949, C.L. Remington. Psyche, 59(2) :62 (1952).

Euchloe creusa

Draba lanceolata Royle

Canada, Alberta, Moraine Lake, 6800', Banff National Park, 27
June 1965, P.A. Opler. Personal observation.

Euchloe olympia

Arabis dirmimondii Gray

Michigan, Montcalm County, T12N, RlOW, Sec. 19, 20 June 1966,

M.C. Nielsen. Preserved larvae in Michigan State University Collection.
Michigan. Roscommon County, T24N, RIW, Sec. 3. 5 July 1967,

MC, Nielsen. Preserved larvae in Michigan State University collection.

Arabis lyrata L.

Indiana, Lake Michigan dunes between Clarke Junction and Pine,

C.A. Shull. Ent. News, 18(3): 73 (1907).

Michigan, Berrien County, T53, R19W, Sec. 29, 29 May, 3-5 June
1967, M.C. Nielsen. Preserved larvae in Mich 1 gan State University
Collection.

Arabis missouriensis Greene

Missouri, St. Louis County, Ranken, 28 April 1935, E.P. Meiners,

Proc. Missouri Acad. Sci., 4:154-156 (1938).

Si symbrium sp.

West Virginia, 1891, W.H. Edwards, Butterflies of North America.

Euchloe hy antis

Arabis glabra (■L.)Bemh.

California, Sierra County, Shenanigan Flat, 14 miles W Downieville,
15 May 1970, P.A. Opler. Personal observation.

Argibis holboelii var. pinetorum (Tides.) Roll.

California, San Bernardino County, Sugarloaf Peak, 8000', July
1970, W. Hovanitz. J. Res. Lepid., 8(1): 17 (1970).

13(l):l-20, 1974

NEARCTIC EUCHLOE

5

Caulanthus amplexicaulis Wats.

California, Kern County, between Mojave and Randsburg, Coolidge.

Ent. News, 19:204-210 (1925).

Caulanthus crassicaulis (Torr.) S. Wats.

California, Modoc County, 5 miles S Ft. Bldwell, 11 June 1970, P.A. Opler,
Personal observation.

Descurainia pinnata (Walt.) Britton

California, San Bernardino County, Newberry Mountains, 3/4 mile
NW Kane Spring Road, 27 March 1964, J.F. and T.C. Emmel. Letter
of 24 November 1964.

California, San Bernardino County, Phelan, J.A. Comstock and C.M.

Dammers, Biill. So. Calif. Acad. Sci., 31(2): 35-37 (1937).

Descurainia pinnata var. nelsonii (Rydb.) Detl.

Washington, Benton County, Vemita, 6 April 1966, E.J. New comer.

Letter of 20 May 1967.

I satis tinctoria L.

California, Modoc County, Buck Creek Ranger Station, 11 June 1970, P.A.
Opler. Personal observation.

Sisymbrium altissimum L.

Washington, Benton County, Vemita, 22 April 1967, E.J. Newcomer.

Letter of 20 May 1967.

Stanleya pinnata (Pursh) Britton

California, Inyo County, Wildrose Station, Panamint Mountains,

15 May 1969, P.A. Opler. Personal observation.

Streptanthella longirostris (Wats.) I^db.

California, San Bernardino County, Phelan, J.A. Comstock and
C.M. Dammers. Bull. So. Calif. Acad. Sci., 31(2): 35-37 (1935).

St rep tan thus bemardinus (Greene) Parish

California, San Bernardino County, Lake Arrowhead, 5000', 29 June
1966, C. Henne. Letter of 7 July 1966.

Streptanthus polygaloides Gray

California, Mariposa Coimty, 2 miles W El Portal, 12 April I964,

P.A. Opler. Personal observation.

Streptanthus tortuosus Kell.

California, Sierra County, Shenanigan Flat, I4 miles W Downieville,

15 May 1970, P.A. Opler. Personal observation.

Streptanthus sp.

California, Alpine Coimty, Hope Valley, 9 July 1949, J.W. MacSwain,
Preserved larvae in California Insect Survey.

California, Siskiyou County, Little Castle Lake, 20 July I969,

P.A. Opler. Personal observation.

California, Tuolumne County, Lodgepole Campground, Sonora Pass,

2 July 1966, P.A. Opler. Personal observation.

6

PAUL OPLER

/. Res. Lepid.

Fig. 1.

Left, egg of Euchloe ausonides, 100 X. Strawberry Canyon, Alameda Co.,
Calif.; right, same, 500 X. Stereoscan electron photomicrographs by Wayne
Steele.

13(l)a~20, 1974

NEARCTIC EUCHLOE

7

geo-negative or photo-positive movements. On Streptanthm
polygaloides yoong larvae of Euchloe hyantis bore through
the clasping leaves lying between the site of oviposition and
flowering portions of the plant. During this movement the
larvae do not pause to feed on leaf material.

Young larvae of Euchloe feed upon unopened flower buds
and flowers. During this period the larvae are comparatively
hidden. The young larvae of E. ausonides station themselves
vertically amongst the flower cluster and always cover the por-
tion of the plant upon which they rest with loosely spun silk.

The first instar larvae of Euchloe hyantis were found to
have another anomolous habit which seems to be an adaptation
to feeding upon Streptanthus, The calyx of Streptanthus flowers
is almost closed distally. In order to reach the inside of the
calyx the young larva bores a small hole through the side of the
calyx, enters, and feeds upon the flower from within.

Older larvae feed in more exposed positions and include the
seed pods in their diet. While feeding upon seed pods the larvae
orient with their anterior end directed toward the apical end
of the pods.

Pupation. — The final stages of ecdysis which lead directly
to pupation in Euchloe seem to occur in a characteristic man-
ner that holds true for all species of Euchloe, and probably take
place in much the same fashion for all members of the tribe
Eiichloini. When the larva has finished feeding it commences
a search for a suitable pupation site which is usually located on
the food plant. Upon finding a pupation site the larva attaches
itself by means of a caudal button of silk and silk girdle with
the anterior portion a,t a higher elevation than the posterior.
Pupation then takes place between 24 and 72 hours later. At
a time varying from just prior to the wandering phase to a time
subsequent to being stationed in the pupation position, the
larva becomes purplish in color. A quantitative lack of obser-
vations prevents the determination of the specifioity of any
of the above phenomenon. The appropriate passages from the
literature upon which the above narration was based are cited
below. Shull (1907), in regard to the mature larva of Euchloe
olympia, said “Shortly a purplish tinge makes its appearance at
the posterior end and about the thorax. The color gradually
extends anteriorally until the whole body shows it. This change
immediately precedes and accompanies the wandering of the
larva seeking a place to pupate.”

First instar larvae, segments T 1-3, A 9-10. Fig. 2, Euchloe ausonides.
Alum Rock Park, Santa Clara Co., Calif.; Fig. 3, Euchloe cretisa, Moraine
Lake, Banff National Park, Alberta, Canada; Fig. 4, Euchloe olympia,
Berrien Co., Mich.

13(1); 1-20, 1974

NEARCTIC EUCHLOE

9

Second instar larvae, segments T 1-3, A 9-10. Fig. 5, Euchloe ausonides,
Buck Creek Ranger Station, Modoc Co., Calif.; Fig. 6, Euchloe olympia,
Montcalm Co., Mich.

10

PAUL OPLER

/. Res. Lepid,

Third instar larvae, segments T 1-3, A 9-10. Fig. 7, Euchloe ausonides,
Buck Creek Ranger Station, Modoc Co,, Calif.; Fig. 8, Euchloe oltjmpia,
Montcalm Co., Mich.; Fig. 9, Euchloe hyantis, Buck Creek Ranger Station,
Modoc Co., Calif.

13(l)a-20, 1974

NEARCTIC EUCHLOE

11

Mead (1877) described the pupation of Euchloe hyantis
based upon observation of larvae collected in Yosemite Valley,
Mariposa County, California, as follows, “Just before the change
to chrysalis the caterpillar turns dull purple. The chrysalis re-
tains this color for a day or two and then gradually assumes a
waxy grayish white color.” Coolidge (1925) noted that the
larva of Euchloe hyantis lotta “becomes solid purplish red”
just prior to pupation, Comstock and Dammers (1932), while
describing the larvae of Euchloe hyantis lotta, stated that “a
short time prior to pupation they turn a mottled dark maroon
over the dorsal and lateral surface above the stigmatal line.
Pupation occurs on the food plant as in Anthocharis cethura!’
(i.e. with a girdle, caudal button, and the head pointing up-
ward, fide Comstock and Dammers).

MORPHOLOGY OF IMMATURE STAGES

Egg. The eggs are typical of Euchloini in being columnar
with the micropylar area broadly rounded. The eggs have from
15-20 prominent vertical ridges which are interconnected by less
prominent horizontal ridges (Fig. 1.).

The eggs undergo a series of progressive color changes as
the embryo develops. Coolidge and Newcomer (1908), while
describing the life history of Euchloe ausonides, stated that the
“color when first laid (was) light bluish green changing in 24-
30 hours to light orange. By the third day the color is almost
vermilion, and about the sixth day the egg turns dirty yellow
brown, especially so apically.”

Larva. — The larvae of Euchloe differ interspecifically and
interstadially in features of color and setation. These differences
will be systematically discussed below.

The sequence of color changes undergone by the larvae of
Euchloe ausonides (Coolidge and Newcomer, 1908) and Euchloe
olympia (Shull, 1907) is parallel.

The first instar larva is orange-yellow with a black head,
while the second instar larva is greenish with a black head.
Markings typical of the final three instars are discernible on the
third instar larva: a gray-green dorsal stripe, yellow-green sub-
dorsal stripes, gray-green supraspiracular stripes, yellow spiracu-
lar stripes, and subspiracular and ventral areas green. The head
of the third instar larva is green-black. The fourth instar larva
of the two species has the same markings, but the head capsule

12

PAUL OPLER

J. Res. Lepid.

Fourth instar larvae, segments T 1-3, A 9-10. Fig. 10, Euchloe ausonides,
Buck Creek Ranger Station, Modoc Co,, Calif.; Fig. 11, Euchloe olympta,
Montcalm Co., Mich.

13(1): 1-20, 1974

NEARCTIC EUCHLOE

13

is greenish gray. The spiracular stripe of the fifth instar larva
is white and is subtended with pale yellow. Plate 1, figs. 1, 6.

Larvae of Euchloe hyantis undergo a similar series of
changes, but have a different color pattern. The color pattern
of the last three instar larvae is distinctive. The dorsal area
is green (occasionally with a narrow purplish line). The sub-
dorsal area is also green, while the supraspiracular area is
purplish. The spiracular area is white or yellow. The subspirac-
ular and ventral areas are green. The head capsule is green.
Interpopulation variation of larval color pattern in Euchloe
hyantis was noticed, while larval coloration of Euchloe ausonides
and E. olympia varies within very narrow limits. Plate 1, figs. 3, 5.

The spiracular stripe is white in most populations of Euchloe
hyantis, but may be yellow in some populations in the western
foothills of the Sierra Nevada. The color of the dorsal and sub-
dorsal areas is also variable. In most populations (Vernita,
Wash.; cismontane Calif.) these areas are green, while these
areas are grayish in some Great Basin populations (John Emmel,
1117 9th St., Apt. 207, Santa Monica, Ca. 90403, per. com.) and
yellowish-green in the San Bernardino Mountains population
(Chris Henne, P. O. Box 1, Pearblossom, Calif., personal com-
munication ) .

Chaetotaxic line drawings of Euchloe larvae are presented
as figures 2-15. These drawings portray the three thoracic seg-
ments for all larvae as well as the eighth and ninth abdominal
segments for first instar larvae. The microscopic primary setae
(Hinton, 1946) are not necessarily portrayed.

Certain primary setae on the dorsal, subdorsal, and lateral
areas are thickened and may be cleft at their apices. Coolidge
(1925) reported that the setae of Euchloe hyantis supported
“hyaline drops of fluid,” while Shull (1907) reported that the
setae of Euchloe olympia were glandular. If the forked setae
do exude secretions, it would be of extreme interest to know
what is their funtion and composition.

Prothoracic segment setae XDl, D2, XD2, SDl, SD2, and LI,
mesothoracic setae Dl, D2, SDl, SD2, and LI are thickened and
cleft in the Euchloe larvae.

The subdorsal group of primary setae on the prothoracic seg-
ment occurs on the same sclerotized chalaza for almost all larvae
of the “Ausonides group,” but occur on separate chalazae on

Euchloe hyantis larvae.

14

PAUL OPLER

]. Res. Lepid.

Fourth instar larvae, segments T 1-3, A 9-10. Fig. 12, Euchloe hyantis, Buck
Creek Ranger Station, Modoc Co., Calif. Fifth instar larvae, segments T 1-3,
A 9-10, Fig. 13, Euchloe ausonides, Buck Creek Ranger Station, Modoc
Co., Calif.

13(l):l-20, 1974

NEARCTIC EUCHLOE

15

Color Plate 1.

1. Euchloe ausonides, 5th instar, Briones Regional Park, Contra Costa Co.,
Calif.; 2. Euchloe ausonides, pupa, Briones Regional Park, Contra Costa
Co., Calif,; 3. Euchloe hyantis, 5th instar. Shenanigan Flat, Sierra Co.,
Calif.; 4. Euchloe hyantis, pupa, Shenanigan Flat, Sierra Co., Calif.; 5.
Euchloe hyantis, 5th instar, near Fallon, Nevada, collected by J. F. Emmel;
6. Euchloe hyantis, prepupa. Shenanigan Flat, Sierra Co., Calif, (upside
down ) .

16

PAUL OPLER

/. Res. Lepid.

Fifth instar larvae, segments T 1-3, A 9-10. Fig. 14. Euchloe olympia,
Roscommon Co., Mich.; Fig. 15, Euchloe hyantis, Buck Creek Ranger Sta-
tion, Modoc Co., Calif.

13(1):1~20, 1974

NEARCTIC EUCHLOE

17

Pupae, lateral aspect. Fig. 16, Euchloe ausonides. Strawberry Canyon,
Alameda Co., Calif.; Fig. 17, Euchloe olympia, Montcalm Co., Mich.;
Fig. 18, Euchloe hyantis, El Portal, Mariposa Co., Calif.

PAUL OPLER

/. Res. Lepid.

18

B

■ I fc LLh

c

(MM)

Fig. 19 —Bar graph of larval head capsule widths. A. Euchloe ausonides,
San Francisco Bay area (Alameda, Contra Costa, and Santa Clara Counties);

B, Euchloe ausonides. Buck Creek Ranger Station, Modoc Co., Calif.;

C. Euchloe oltjmpia, Michigan; D. Euchloe hyantis, California (Mariposa,
Modoc, and Sierra Counties).

13(l);l-20, 1974

NEARCTIC EUCHLOE

19

The successive instars of each Euchloe species display dis-
tinct trends in setal coloration and size. With each successive
instar, size of primary setae becomes greater on Euchloe auson-
ides larvae, remains about the same on Euchloe olympia larvae,
and becomes smaller on Euchloe hyantis larvae.

On first instar Euchloe larvae all primary setae are melanized.
Ultimate instar larvae have dorsal and subdorsal primary setae
darkened and the primary setae of lateral, subventral, and ven-
tral areas demelanized.

The secondary setae which appear on Euchloe larvae in in-
creasing numbers on successive instars also vary in size and
prominence. The length of secondary setae becomes longer
on larvae of Euchloe olympia, remains about the same on larvae
of Euchloe ausonides, and become shorter on larvae of Euchloe
hyantis.

The apparent color of secondary setae on ultimate instar
larvae is uniformly pale on Euchloe ausonides larvae, and dark
on dorsal and subdorsal areas of Euchloe olympia and Euchloe
hyantis larvae.

The extent and melanization of the sclerotized setiferous
chalazae and pinaculi also differs between species and succes-
sive instars of the same species. The sclerotized chalazae of
Euchloe ausonides and Euchloe olympia larvae become greater
in extent with each succeeding instar, remaining about the same
proportion to larval size. The chalazae of Euchloe hyantis
larvae become progressively smaller with each succeeding instar,
paralleling the size decrease of the primary setae.

The sclerotized chalazae or pinaculi bearing primary setae
are darkened on all areas of all instars of Euchloe larvae, with
the exception of lateral areas on ultimate instar larvae of Euchloe
hyantis.

The bases of secondary setae on all ultimate instar larvae are
darkened on dorsal and subdorsal areas and pale on the lateral
areas. On the subventral and ventral areas they are darkened
on Euchloe ausonides and Euchloe hyantis, while most are pale
on Euchloe olympia larvae.

Pupa. — Pupal marking and coloration is extremely variable.
Some idea of the pupal coloration may be seen by referring to
Plate 1, figs. 2 and 4.

Although the recurvature of Euchloe pupae is somewhat
variable, the relative proportions of the pupae are slightly dif-
ferentiated. Line drawings of the pupae (Figs. 16-18) show
that the ratio of the cephalic portion to pupal length is greater

20

PAUL OPLER

/. Res. Lepid.

for Euchloe ausonides (.28) than for Euchloe olympia (.25) and
Euchloe hyantis (.24). The dorsal surface of the cephalic portion
of Euchloe hyantis pupae is slightly emarginate when viewed
laterally.

SUMMARY

Certain features of the life history and morphology of the
immature stages of Euchloe emphasize the division of the Nearc-
tic members into two species groups. In particular, location of
oviposition site, larval behavior, larval color pattern and chaeto-
taxy are characters whose condition in the two species groups
indicates genetic isolation of long standing.

ACKNOWLEDGMENTS

The following individuals provided information or material
representing the immature stages of Euchloe which proved in-
valuable in the construction of this paper: J. F. Emmel, Idyll-
wild, California; T. C. Emmel, Gainesville, Florida; Chris Henne,
Pearblossom, California; E. J. Newcomer, Yakima, Washington;
M. C. Nielsen, Lansing, Michigan; and Fred Thorne, El Cajon,
California. Roland Fisher, Michigan State University, East
Lansing, Michigan, kindly loaned larval material representing
Euchloe olympia.

I am indebted to Wayne Steele, Livermore, California, for
his preparation and provision of scanning electron photomicro-
graphs of the egg of Euchloe ausonides.

Evert Schlinger, Division of Entomology, University of Cali-
fornia, Rerkeley, reviewed the manuscript of this paper and
provided suggestions and corrections which were incorporated
into the final draft.

LITERATURE CITED

( In addition to those cited in Part 2 )

HINTON, H. E. 1946. On the homology and nomenclature of the setae
of lepidopterous larvae, with some notes on the phylogeny of the Lepi-
doptera. Trans. Roy. Ent. Soc. London, 97(1): 1-37.

HOVANITZ, W. 1970. Habitat — Euchloe hyantis andrewsi. Jour. Res.
Lepid., 8(1): 16-17.

MEINERS, E. P. 1938. The life history of Euchloe olympia Edwards,
with some notes on its habits. Proc. Missouri Acad. ScL, 4(6): 154-
156.

OPLER, P. A. 1966. Studies on Nearctic Euchloe. Parts 1 and 2. Jour. Res.
Lepid., 5(1): 39-50.

, 1969. Studies on Nearctic Euchloe. Part 5. Distribution. Jour.

Res. Lepid., 7 (2): 65-86.

REMINGTON, C. L. 1952. The biology of Nearctic Lepidoptera. I. Food-
plants and life-histories of Colorado Papilionoidea. Psyche, 59(2): 61-

70.

SHIELDS, O. A., J. F. EMMEL, and D. E. BREEDLOVE. 1970. Butterfly
larval foodplant records and a procedure for reporting foodplants.

Jour Res. Lepid., 8(1): 21-36.

SWALES, D. E. 1966. Species of insects and mites collected at Frobisher
Bay, Baffin Island, 1964 and Inuvik, N.W.T., 1965, with brief eco-
logical and geographical notes. Ann. Soc. Entornol. Quebec, 11(3):
189-199.

Journal of Research on the Lepidoptera

13(l):21-22, 1974

1160 W. Orange Grove Ave., Arcadia, California. U.S.A.
© Copyright 1975

3 STACKS OF THE EGGS OF
HEMISTOLA
HATCHING

NOEL McFarland

129 Gloucester Ave., Belair, South Australia 5052

Eggs of Hemistola chrysoprasaria (Esper) {GEOMETRh
DAE-^GEOMETRINAE) (det. F. Benz), SHOWING LARVAE
IN THE PROCESS OF EMERGING: These eggs were kindly
given to me by Dr. F. Benz of Binningen, Switzerland, who ob-
tained them from a female taken at mv. light, 26 JUNE 1968, at
Ober-Zeihen AG, Switzerland. (The eggs were deposited 28
JUNE 68, and hatching took place 10 days later ) . The mode of
oviposition, in precise stacks, is characteristic. Perhaps the
procryptic value of these stacks lies in the way they simulate
the appearance of tightly-coiled immature tendrils on the stems
of the foodplant (Clematis vitalba L. - RANUNCULACEAE)?
The photograph was made when the larvae were just in the
process of chewing their way to freedom. The third one down
from the top (in the left stack) had already gone, leaving the
(characteristic) clean-cut exit-hole in the end of its empty egg
shell. The second one down (same stack) had nearly finished
its exit-hole; the dark-colored object filling the hole is the head
of the young larva. The top egg (same stack) shows a very
small hole just being started. In the other 2 stacks can be seen
the (dark) larval heads, aligned in rows, still inside the eggs.
The egg dimensions for this series were: 0.90-0.85 x 0.65 x 0.40
mm. (N. McFarland photo; species code-number Gm. 195).

21

22

NOEL McFarland

/. Res. Lepid

Journal of Research on the Lepidoptera

13(l):23-42, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

AN ANNOTATED CHECKLIST OF

THE MACROHETEROCERA OF
SOUTH-EASTERN ONTARIO

P. S. WARD', R. HARMSEN“, and P. D. N. HEBERT'

Biology Department, Queens University, Kingston, Ontario

INTRODUCTION

Progress in theoretical ecology, community evolution and
basic pollution research is seriously retarded by the absence of
systematically collected, carefully documented population sam-
ples of sufficient size in both number of individuals and in species
number. This is especially true for the animal kingdom.

Lepidoptera, especially night flying ones, are an admirably
suitable taxocene for such research. Yet, even here, few collec-
tions are adequate in sampling technique, quantification and
consistency, making it very difficult to use these collections for
the formulation of generalizations concerning ecosystem or com-
munity structure.

In the past several years we have been conducting an inten-
sive complete-season collecting program in an attempt to survey
the macroheterocera fauna of the Kingston region. This forms part
of a broader study on the effects of environmental disturbance
on species diversity, and the evolution of community structure,
the results of which will be published elsewhere. In this publica-
tion we wish to record the moth species collected, their flight
periods and relative abundance, and notes on their life histories.

The list covers the south-eastern portion of southern Ontario,
an approximately triangular area bordered by 75° 45' W longi-
tude, 45° Ol/N latitude, and the St. Lawrence River System
( figure 1 ) . This is a physiographically diverse area, with variably
overlain Paleozoic sediments interrupted in the center of the

^School of Biological Sciences, University of Sydney, Sydney, Australia.
^Biology Department, Queen’s University, Kingston, Ontario.

23

24

WARD, ET AL

J. Res. Lepid.

Fig. 1. — South-eastern Ontario, with the principal collecting sites G, P, and
C. The fourteen other localities are as follows: 1) Portland, Leeds Co.,
2) Westport, Leeds Co., 3) Bedford Mills, Frontenac Co., 4) Lake Opini-
con, Frontenac Co., 5) Cranberry Lake, Frontenac Co., 6) Washburn,
Frontenac Co., 7) Kingston, Frontenac Co., 8) Amherstview, Lennox &
Addington Co., 9) Westbrook, Frontenac Co., 10) Harrowsmith, Frontenac
Co., 11) Marlbank, Hastings Co., 12) Belleville, Hastings Co., 13) Frank-
ford, Hastings Co., and 14) Wooler, Northumberland Co.

13(1):23=42, 1974 ONTARIO MACROHETEROCERA

25

region by a protrusion of the Precambrian Shield. The predomi-
nant forest of white pine and northern hardwoods contains a
considerable infusion of austral (Carolinian) elements (Snyder
et al., 1941; Soper, 1962; Harmsen, Hebert and Ward, 1974).
Much of the original vegetation has been replaced by second
growth forest and agricultural land.

The macroheterocera fauna of the area exhibits a remarkable
species richness which coincides with a great regional diversity
of plants (Beschel et ah, 1970) and birds (Quilliam, 1973). The
south-eastern Ontario total of 820 + macroheteroceran species
may be compared to Ferguson’s (1955) listing of 764 species
from the entire province of Nova Scotia and Moore’s (1955)
tabulation of 1006 species from the state of Michigan. Single-
locality species totals for south-eastern Ontario (see below) are
far in excess of those normally reported in the literature for
temperate regions (e.g. Preston, 1948; Williams, 1964; McFar-
land, 1965; LaFrance, 1968).

METHODS

The bulk of the records is based on daily collections of noc-
turnal macrolepidoptera made by the authors over at least one
complete collecting season (spring to autumn) at each of three
collecting sites: Glenburnie, Perth Road and Chaffey’s Locks,
hereafter designated G, P, and C respectively (Table 1). The
former is located in marginal agricultural land on Ordovician
limestone, while P and C are situated in areas of mixed forest on
Precambrian metamorphic rock. All specimens were collected
with a 20- watt blacklight ( GE FT20T12-BL ) set against a white
cloth background. The light was run all night and moths were
collected off the screen at dawn.

We consider one complete sample of the fauna to consist of
all macroheterocera collected over an entire season at one light.
Thus site G was sampled once, C and P each four times. Alto-
gether 101,862 individuals comprising 778 species were identified
and recorded. The total numbers of species recorded in this
manner from each collecting site are 427 (G), 571 (G), and
701 (P).

Additional material from G, C, and P was obtained by rearing
immature stages, collecting diurnally, and collecting with light
and bait outside the prescribed collecting schedule. Sporadic
collecting has also been carried out at 14 other localities in

26

WARD, ET AL

/. Res. Lepid.

south-eastern Ontario (Figure 1). Most of these collections were
made by the authors but some records from the Royal Ontario
Museum, Toronto (ROM) (mostly collected by J.C.E. Riotte)
and the Canadian National Collection, Ottawa (CNC) have also
been included. These additional data bring the total number of
recorded species to 822.

The majority of the material was identified by P.S.W. and
P.D.N.H., utilizing particularly Forbes (1948, 1954, 1960), Fer-
guson (1955), McGufBn (1967, 1972), and access to the Cana-
dian National Collection, Ottawa. Some difficult material was
determined by specialists mentioned in the acknowledgements.

The families, genera, and species are arranged alphabetically,
and subspecies names are not used. Nomenclature essentially
follows McDunnough (1939), in conjunction with recent revi-
sionary changes. Authors names have been omitted since all
species names may be found in McDunnough (1938) or Forbes
(1948, 1954) with the exception of the following: Anisota
finlaysoni Riotte (1969); Eupithecia rindgei McDunnough
(1949), Semiothisa marmorata Ferguson (1972), Semiothisa
pinostrohata Ferguson (1972), Semiothisa signaria (Hiibner)
(=S. dispuncta (Walker)). Thera juniperata (Linnaeus) {=T.
procteri Brower), Helicoverpa zea (Roddie) (=H. obsoleta of
authors, not Fabricius), and Hypenodes palustris Ferguson
(1954).

THE CHECKLIST

In the following list, the numbers in parentheses after G, C,
or P refer to the total numbers of individuals collected at each
site under the prescribed collecting schedule. This gives a rough
idea of relative abundances, but it is worth remembering that G
was only sampled one year. If a species was only recorded from
G, C, or P outside the collecting program this is indicated by a
dash in parentheses. Records from the fourteen other collecting
localities are usually only included if less than half a dozen
individuals have been collected at G, C, and P.

13(1):23»42, 1974 ONTARIO MACROHETEROCERA

27

AGARISTIDAE
Alypia octomaculata

ARCTIIDAE
Apantesis anna

Apantesis arge

Apantesis celia
Apantesis nais
Aj^antesis parthenice
Apantesis phalerata
Apantesis phyllira
Apantesis virgo
Apantesis virguncula
Arctia caja

Cisthene plumbea

Clemensia albata
Crambidia casta
Crambidia pallida
Crambidia pura
Cycnia tenera
Estigmene acraea

Euchaetias egle
Euchaetias oregonensis
Halysidota caryae
Halysidota maculata

Halysidota tesselaris
Haploa confusa
Haploa contigua
Haploa lecontei
Holomelina aurantiaca
Holomelina ferruginosa
Holomelina opella
Hyphantria cunea
Hypoprepia fucosa
Hypoprepia miniata
Lycomorpha pholus

Phragmatobia assimilans
Phragmatobia fuliginosa

Pyrrharctia isabella

Spilosoma congruum
Spilosoma primum
Spilosoma virginicura

Trigrioides bicolor

CITHERONIIDAE
Anisota finlaysoni

Anisota rubicunda
Anisota virginiensis

Eacles imperialis

DREPANIDAE
Drepana arcuata

Drepana bilineata
Eudeilinea herminiata

Oreta rosea

EPIPLEMIDAE
Callizia amorata

C(-), P(~). Adults diurnal, May 10-Jun 26;
reared from larvae on Vitis riparia

C(18), P(130). Jun 8-Jul 9.

G(20), C(5), P(4). May 8-14; Jul 23-Sep 6.
Predominantly in late summer.

G(l), C(-), P(28). May 31-Jun 24.

C(58), P(228). May 21- Jul 14; Aug 19.

G(3). C(2), P(19). Aug 10-Sep 8.

G{65), C(-). Jun 2-17; Aug 6-Sep 7; Oct 13.

G(2). Aug 5-Sep 5.

G(7), C(2), P{6). Jul 19-Aug 3.

G(29), C(-), P(17). Jun 13-Jul 27.

C(-), P(l). Aug 7. Two specimens at light at
C: Aug 18, 1964; Aug 10, 1966.

C(-). Two specimens at light; Jul 14, 1963;

Jul 17, 1967.

G(2), C(5), P(4). Jun 26-Jul 28.

C(-). One specimen at light; Aug 23, 1969.
C(10), P(168). Jul 27-Sei. 4.

G(22), P(1). Jun 12-26; Aug 6-31.

G(10), C(24), P(71). Jun 2-Jul 30.

G(103), C(9), P(51). May 24~Sep 4. Probably
bivoltine .

G(32), C(22), P{29). Jun 4-Aug 2.

G(4), C(12), P(21). May 31-Jul 13; Jul 16-Aug 9.
G(20), C(19), P(300). May 24-Jul 11.

Belleville. Aug 10, 19c>2, (Carleton University
Collection) ,

G(75), C(374), P(2734). Jun 3-Aug 27.

G(47), c(20), P{57). Jul 19-Aug 18.

G(l), C(ll), P(3). Jul 15-28.

C(l), P(14). Jun 17- Jul 23.

C(9), P(58). Jun 18; Jul 6-Aug 23.

G(154), C(50), P(50). Jun 2-Jul 10; Jul 7-Sep 6.
C(18), P(158). Jun 23-Aug 29.

G(118), C(19), P(192). Jun l~Jul 21.

G(20), C(137), P(832). Jul 9-Sep 13.

G(3), C(17), P(505). Jul 13-Sep 15.

C(-)j P(3). Jul 14-Aug 10. Common diurnal
species, frequently visiting Solidago flowers,

Jul 29-Aug 22.

G(3), C(22), P(168). May 2-Jun 3.

G(68), C(12), P(55). May 17-Jun 7; Jul 20-
Sep 15. Second brood larger.

G(73), C(83), P(165). May 31-Aug 6; A.ug 26-
Sep 12.

G(l), C(181), P(186). May 14-Jul 4; Aug 22.

G(3), C(45), P(115). May 7-Jun 29.

G(252), C(230), P(463). May 1-Sep 16; Oct 22.
Several overlapping generations.

P(10). Jul 24-Aug 6,

Kingston; Jun 27, 1969 (12) (ROM); Otter Lake,
Frontenac Co.: Jul 6, 1974 (Id). Eggs and
larvae collected on Quercus in vicinity of
Belleville .

C(297), P(1463). May 19- Jul 29; Aug 17-Sep 10.
C(4), P(29). May 30- Jul 17. Only females at
light; males diurnal.

C(43), P(76). Jun 12-Aug 8.

G(2), C(309), P{636). May 4-Jul 10; Jul 3-
Aug 30.

G(1), C(91), P(353). May 2-Jun 28; Jul 5-Sep 5.
C{1), P(3). May 31-Jun 12; Aug 12. Locally
common in hardwood swamps at P. Jun 7-16.

G(l), C(13), P(65). Jun 8-Jul 21; Jul 30-
Sep 15.

C(3) , P(l). May 31-Jun 29.

/. Res. Lepid.

28

WARD, ET AL

EUCHKOMIIDAE
Cisseps fulvicollis

Ctenucha virginica

GEOMETRIDAE

Abbottana clemataria
Aethalura anticaria
Alsophila pometaria

Anacamptodes ephyraria
Anacamptodes vellivolata

Anagoga occiduaria
Anavitrinella pampinaria

Antepione thisoaria
Apicia confusaria
Bapta glomeraria
Bapta semiclarata

Bapta vestaliata
Besma endropiaria
Besma quercivoraria
Biston cognataria

Brephos infans

Campaea perlata

Caripeta angustiorata
Caripeta divisata
Caripeta piniata
Catopyrrha coloraria

Cepphis armataria
Cepphis decoloraria
Chlorochlamys chloroleucaria

Cingilia catenaria

Cleora projecta

Cleora sublunaria
Coryphista meadi

Cosymbia pendulinaria

Deilinea erythremaria

Deilinea variolaria

Diactinia silaceata
Dichorda irridaria
Drepanulatrix liberaria
Dyspteris abortivoraria
Dysstroma hersiliata
Dysstroma sp.

Earophila vasaliata
Ectropis crepuscularia

Ematurga atomaria

Ennomos magnarius
Ennomos subsignarius
Epirrhanthis substriataria
Epirrhoe alternata
Epirrita autumnata
Erannis tiliaria
Euchlaena effecta
Euchlaena irraria
Euchlaena johnsonaria

G(96), C(5), P(a). Jun 9-Jul 11; Jul 17-
Sep 8: Oct 11.

G(243), C(25), P{45). Jan l-aug 5; Dep 15.

G(2), C(154), P(92). May 1-Jun 24.

C(64) , P(33). May 7-Jun 2b.

0(83), C{-), P(4). npr 8-Hay 7; Nov 4.

Common, diurnal spring species in Kingston,

Mar 21-Hay 1. Fewer adults emerge in the
autunm .

C(42), P(108). Jun 25-Aug 10.

G(75), C(18), P(4b). May 2-Jul 4; Jul 14-
Aug 10; Sep 4-23.

0(1), P(25). May 4-Jun 14.

G(186), 0(28), P(80). May 15-Jul 8; Jul 15-
Sep 8.

P(32) . May 23-Jun 25.

0(15), 0(6), P(57). Jul 6-Aug 6.

P(2). May 8.

C(-), ?(1). May 14. Collected during the day
at C, P, and Harrowsmith, May 17-29.

G(lb), C(16), P(89). May 17-Jun 28; Jul 17.
C(2), P(33). May 30- Jul 15.

C(9), P(85). May 11- Jul 10; Jul 27-aug 21.
0(34), 0(39), P(115). May 2D-HUg 26. Fre-
quency of melanic specimens in 1971 at P was
12/73 or 16.4f=.

O(-), P(-). Diurnal, early spring species,

Apr 11 -May 9.

0(12), 0(136), P(303). Jun 2-Jul 29; i^ug 15-
Sep 15.

G(l), 0(22), P(43). Jul 15-Aug 15.

C(2) , P(2) . Jul 19-Aug 1.

0(2), 0(42), P(179). May 30-Jul 31.

Cooler. May 9, 1972 {L^dd, 229), flushed from
Ceonothus .

G ( 1 ) , /'{T ) . Jun 10; aug 14.

C(2). Jun 6-18,

0(34), 0(24), P(35). May 25- Jul 17; Jul 25-
Aug 31.

0(5), P(859). Sep lO-Oct 2. Larvae collec-
ted on Myrica gale and on ericaceous bog
plants; in outbreak proportions on sphagnum
bogs at Harrowsmith and westport in 1973 .

P(3). May 8-Jun 17. Also reared from larvae
on Myrica gale in small sphagnum bog at P.
PdTT^y 19.

G(o), C(-), P(1). AUg 9-Sep 13. Larvae on
Berberis vulgaris at C. also attacks orna-
mental Berberis in urban areas, including
Kingston.

G(l), 0(64), P(227). May 14-Sep 14. Two,
partially overlapping broods with peaks in
mid- June and early nugust.

G(l), C(2), P(21). Hay 29-Jul 2; Jul 26-

AUg 30.

G(5), 0(22), P(57). Jun 8- Jul 15; Jul 13-
Sep 3.

C(b). May 25- Jun 2; Jul 29-Aug 12.

C(13), P(27). May 19- Jun 22; Jul 31-Aug 17.

C (4) . Sep b-Oct 12 .

C(l) , P(l). Jun 2-15.

G(l), C(7), P(19). Jun 17-Jul 20.

Bedford Mills, ‘J\festport. Two specimens at
light in sphagnum bogs, Jun 23-29, 1973.

G(l), C(4), P(22). Apr 20-Kay 30.

G(9), 0(60), P(92). May 1-Jun 16; Jun 16-
AUg 16.

P(-). Harrowsmith, vv'estport . Diurnal bog
species, May 24-Jun 26.

G(53), 0(63), P(3b7). Jul 28-Oct 22.

G(10), C(134), P(246). Jul 6-Aug 20.

Wooler. One specimen at light; May 9, 1972.
C(l), P(18). Jun 2-29; Aug 6-Sep 9.

C(45), P(18). Oct 5-24.

G(47), C(3b), P(47). Oct 8-Nov 4.

G(2), C(4), P(1U). Jun 27-Jul 2b.

C(l) , P(24) . Jun 2-Jul b.

0(3), 0(2), P(39). Jul b-Aug 10.

13(1):23~42, 1974 ONTARIO MACROHETEROCERA

29

Euchlaena marginaria
Euchlaena obtusaria
Euchlaena serrata
Eudule mendica
Eufidonia discospilata

Eufidonia notatoria
Eugonobapta nivosaria
Eumacaria latiferrugata
Euphyia centrostrigaria

Euphyia multiferata
Euphyia unarigulata

Eupithecia affinata
Eupithecia castigiata
Eupithecia coloradensis
Eupithecia coagulata
Eupithecia columbiata
Eupithecia filmata
Eupithecia fletcherata
Eupithecia furnosa
Eupithecia gibsonata
Eupithecia luteata
Eupithecia miserulata

Eupithecia misturata
Eupithecia mutata
Eupithecia palpata
Eupithecia perfusca
Eupithecia ravocostaliata
Eupithecia rindgei
Eupithecia russeliata
Eupithecia satyrata
Eupithecia sheppardata
Eupithecia sobrinata

Eupithecia strattonata
Eupithecia swetti
Eustroma nubilata
Guenaria basiaria
Haematopis grataria

Heliomata cycladata

Hesperumia sulphuraria
Hethemia pistaciaria
Heterophleps triguttaria

Homochlodes fritillaria
Horisme intestinata
Hydrelia albifera
Hydria undulata
Hydriomena divisaria
Hydriomena renunciata
Hypagyrtis piniata
Hypagyrtis subatomaria
Hyperetis alienaria

Isturgia truncataria

Itame andersoni complex

Itame brunneata

Itame exhauspicata
Itame pustularia
Itame ribearia

Itame subcessaria
Itame sulphurea

Lambdina fiscellaria
Lobophora nivigerata

Lozogramma subaequaria

C(6), P(25). May 14- Jun 20.

G(l). C(5), F(2}. Jun 2»30.

G{3iK C(1o), P(7). Jul 9-MUg 5.

C(l), P(8). Jul 6-nug 5.

C(-). Harrowsmith, Westport. Diurnal species,
common in bogs, May 24- Jul 8.

G(5), P(47). May 19-Jul 2.

0(2), C(33), P(189). Jul 2-Aug 8.

0(5), P(4). Jun 6-18; Jul 4-KUg 5.

0(9), 0(14), P(34). Jun 24-Oct 22. Appar-
ently two or more generations, commonest In
August - October. Possibly a migratory
species .

P(3). Jun 7-20.

0(46^, C(19), P(25). May 16- Jul 13; Aug 2-
Sep i9.

C(2), P(9). Aug 9-Sep 3.

C(l). Jun 13.

P(l) . Jun 16.

P(6). nug 15-27.

C(31), P(244). May 1-Jun 11.

P(4). May 7-19.

P(146). Aug 8-Sep 2.

0(5), 0(4), P(27). Kay 9-Jun 22; nug 3-Ser 4.
0(19), P(23). May 7-Jun 28.

P(5). Jun 10-26.

G(13), 0(12), P(45). Hay 1-Jul 1; Aug 3-Nov 3.
Much commoner in autumn; possibly overwinters
as adult.

0(1), P(l). May 8; Jul 17.

P(3). Jun 30- Jul 5.

0(1), 0(24), P(89). May 14- Jun 27.

P(6). Jun 17-27; Jul 27.

G(2), C(d), P(14). May 1-Jun 29.

P(l). Jun 2.

P(3). Jun 26-29.

0(1). Aug 25.

C(l) 1(2). May 29-Jun 19.

G(l4), C(27), P(77). AUg 14-Oct 13. L<-rvae
collected on Juniperus comnur.is .

G(l), C(3), P(13). May 19-Jun 19.

P(2). May 16-18.

G(2). Jul 10-Aug 1.

G(l), C(3), P(6). Jun 22~Aug 5.

G(51), 0(1), P(2). Jun 11-15; Jul 23-Sep 16.
Second generation much more prevalent.

C(l). Jun 14. One other record from C:

Jun 14, 1967.

C(-). Two specimens at light: Jul 9-12, 1967.
0(3), 0(69), P(79). May 10-Jun 26.

C(-), P(l). Jul 27. Collected diurnally at
C; Jul 30, 1969.

0(1), P(15). Jun 3-Jul 8.

0(1), c(39), P(10). May 21-Jul 1; Jul 14-Aug 24.
P(4). May jO-Jun 17; A.ug 13.

0(1), C(-), P(28). Jun 1-Aug 21.

0(1). May 3.

C(13), P(58). May 7-Jun 15; Jul 12-26.

C(14) , P(46) . Jun 9-Jul 30.

0(1) P(108). Jun 9- Jul 28; Aug 27.

C(36), P(247). May 8-Jun 26; Jul 30-Aug 6.

Only three second generation individuals.

P(-). Harrowsmith, V/estport. Common diurnal
bog species, May 6-Jun 26.

0(5), P(83). Jun 9- Jul 26. Reared from larvae
collected on Ribes and Vaccinium .

P(l). Jun 26. Common in sphagnum- bog at West-
port, Jun 26-27, 1973.

C(6), P(30). Jun 7-Jul 14.

0(5), C(59), P(536). Jun 30-Sep 4.

0(3), C(-), P(8). Jul 18-30. Larvae collected
on a cultivated Ribes sp.

C(5), P(44). Jul 6-Aug 1.

P(-). Bedford Mills, Portland. Adults diurnal
in sphagnum bogs, the males occasionally at
light, Jun 11-Jul 29. Also rearea from larvae
collected on Myrica gale.

C(6) , P(3). Sep 3-Oct 5.

G(2), C(42), P(108). May 5-Jun 28; Jul 10;

Jul 23.

C(l) , P(29) . May 8-Jun 11.

/. Res. Lepid.

30

WARD, ET AL

Lycia ursaria
Lygris diversilineata

Lygris explanata
Lygris serrataria
Lygris testata
Lytrosis unitaria
Melanolophia canadaria
Melanolophia signataria
Mesoleuca ruficillata
Mesothea incertata

Metanema determinata
Metanema inatomaria
Metarranthis duaria
Ketarranthis hypochraria
Metarranthis indeclinata
Metarranthis obfirmaria

Metarranthis refractaria
Metarranthis warneri
Nematocampa f ilamentaria

Nemoria bistriaria

Nemoria mimosaria
Nepytia canosaria
Nyctobia atroliturata
Nyctobia limitaria
Operophtera bruceata

Orthofidonia flavivenata
Orthofidonia tinctaria
Paleacrita vernata
Percnoptilota obstipata

Perizoma basaliata
Pero honestarius

Pero marmorata
Pero morrisonarius
Phaeoura que maria
Phigalia olivacearia

Phigalia titea

Plagodis alcoolaria
Plagodis nigrescaria

Plagodis phlogosoria

Plagodis serinaria
Pleuroi-rucha insulsaria

Prochoerodes transversata
Protitame virginalis
Protoboarniia porcelaria
Rheumapter'a hastata
Scopula cacuminaria
Scopula enucleata
Scopula inductata

Scopula junctaria
Scopula persimilis

Selenia alciphearia
Semiothisa aeniulataria
Semiothisa bicolorata
Semiothisa bisignata
Semiothisa gnophosaria
Semiothisa marmorata
Semiothisa mellistrigata
Semiothisa minorata
Semiothisa neptaria
Semiothisa ocellinata
Semiothisa orillata
Semiothisa oweni
Semiothisa pinostrobata
Semiothisa sexmaculata

G(8), C(5), P(157). upr 12-Jun 5-
G(32), c(40), P(3B). Jul 9-Sep 15. In-
cludes form gracilineata . Reared from larvae
on Vitis.

C(20) , P(77) . Jun 30-KUg 14; Sep 23.

G(l) , P(o) . Jul 9-27.

C(l), P(2). Sep 14-30.

G{2), C(19), P(30). Jun 24-Jul 19.

0(4) , P(18) . May 29-Jun 23 .

G(2), C(21), P(22). May 4- Jun 24.

C(2), P(2). Hay 21- Jun 7; aug 9-13.

C(-), P(-). Harrowsmith , Washburn, vvestport,
Portland, adults diurnal, on bogs and dry
heaths, apr 2b- Jun 2.

C(-). One specimen at light: May 23, 1973.
0(43), P(117). May 14- Jul 6; Jul 14-aug 25.
G(5), C(3), P(22). May 4-Jun 18.

C(2) , P(7) . May 21-Jun 10.

0(1), 0(12), P(3b). May 24-Jur 23.

C(-), P(l). Jun 7. adults normally diurnal,
on dry heaths or bogs at C, P, and Westport,

May 10- Jun 5.

0(1), C(l^), P(19). May 31-Jun 20.

0(3 ) , CU) , P(ll) . Jun 10-29.

0(4), 0(20), P(45). Jul 3-aug 3. Includes
three of form chagnoni from P.

0(10), 0(170), P(730). May 1-Jun 17; Jul 9-
aug 17; Sep 15.

0(12), P(13). May 28-Jun 19; Oct 27.

0(4), 0(20), P(4). Aug 8-Sep 2o.

O(-), P(30). apr. 19-May 19.

0(1), 0(12), P(15). May 3-28.

0(4), G(-), P(-). Nov 2-3. Oonimon late fall
species at 0 (Oct 31-Nov 8).

0(7), P(188). May 1-29.

P(l) . Jun 9.

0(66), 0(20), P(28). apr 8-May 29.

0(47), 0(9), P(3b). May 18-Nov 3. Oommonest
in Sejitember and uctobei’; possibly migratory.
0(1) . Jul 20.

0(820), 0(41), P(33). May 10-Jul 4; Jul 26-
Oct 11.

P(2). Aug 11-12.'

0(7) , P(2) . May 31-Jun 25.

0(1), 0(1), P(ll). May 23-Jun 30.

0(4), 0(7), P(70). Apr. 12-May 19. all
males, no melanics.

0(3B), 0(31), P(118). apr 12-May 27. all
males; frequency of melanic specimens in 1971
at P was 5/100 or 5.0%.

0(38), P(152). May 8-Jun 18.

O(-), P(l). Jun 10. One specimen at light at
0: Jun 20, 1973.

0(2), 0(174), P(1470). May 1-Jun 25; Jul 5-
aug 14.

0(26), P(IOO). May 10- Jun 17.

0(21), 0(55), P(36). May 31-Oct 25. Appar-
ently several overlapping generations.

0(1), 0(53), P(136). Jul 29-Sep 15.

0(1), 0(8), P(3b8). May 8-Jul 16; Jul 23-Sep 4.
0(6) , P(9) . Jun 27-Jul 23.

C(-). One specimen at light: Jul 1, 1973.

0(7), 0(1), P(-). Jul 4-«ug 14.

0(11), 0(55), P(260). Jun 29-aug 19.

0(170), 0(27), P (8). Hay 31- Jul 17; aug 4-Sep
14.

P(l). Jun 18.

O(-). One male reared from mature wandering
larva collected apr 6, 1970.

P(l). May 8

O(-), P(57). May 29-Jun ?9; Jul 24-Sep 2.
0(124). May 31-aug 28.

0(7), 0(394), P(2741). May 25-Ser. 16.

0(1). Jun 16.

0(2) . Jul 18-20.

0(3), 0(3), P(l). May 31-Aug 14.

0(9), 0(100), P(1051). May 14-Sep 2.

0(1) , P(l) . Jun 2-7.

0(1) , P(l) . aug 16-20.

0(3), P(20). May 25- Jul 3; Jul 26-nUg 22.

0(1), P(15). May 31-Jul 8; Jul 29-aug 13.

C(SOl, P(119l. I>tiy 29-Sep 14.

0(3), 0(4), F(36). May 19-Jun 30; Jul 23-
Sep 5.

13(1):23^42, 1974 ONTARIO MACROHETEROCERA

31

Semiothisa signaria

Semiothisa submarmorata

Semiothisa transitaria
Semiothisa ulsterata
Sicya raacularia
Spargania magnoliata
Steuaspilatodes antidiscaria

Stenoporpia polygrammaria
Sterrha demissaria
Synchlora aerata

Tacparia detersata
Tacparia zalissaria
Tetrads cachexiata
Tetrads crocallata
Thera contracta

Thera juniperata

Trichodezia albovittata

Triphosa haesitata

¥enusia comptaria
Xanthorhoe emendata
Xanthorhoe ferrugata

Xanthorhoe iduata
Xanthorhoe lacustrata
Xanthotype sospeta
Xanthotype urticaria
Xystrota ferruminaria

LASIOCiJ-iPILUE
Epicnaptera americana

Kalacosoma americanum
Malacosoma disstria
Tolype laricis
Tolype velleda.

LMAWTRIIDaE

Orgyia definita
Orgyia leucostigna

Parorgyia

Paror-gyia

Parorgyia

Parorgyia

Parorgyia

NOCTUIDAE

Abagrotis

Achatodes

Acronicta

Acronicta

Acronicta

Acronicta

Acronicta

Acronicta

Acronicta

basifla¥a

dorsipennata

obliquata

plagiata

vagans

alternata

zeae

af flicta

americana

dactylina

fragilis

grisea

haesitata

hasta

Acronicta

Acronicta

Acronicta

Acronicta

Acronicta

Acronicta

impleta

irapressa

sp. near impressa
inclara
innotata
interrupta

acronicta laetifica

Acronicta lanceolaria
Acronicta lepusculina

Bedford Mills, V/estport . i%t light
in sphagnum bogs, Jun 23-26, 1973.

Westport , Conunon at light in sphag-
num bog, Jun 27-29, 1973.

C(l). Jul 20.

C(2), P(9). Jun 13-Jul 9; Aug 9.

G(l) , C(25) , F{16) . Jul 2-23.

P(l). Jun 7.

C(-). One specimen at light: Jun 5,

1967.

P(13). Jun 5- Jul 3.

P(37). Jul 25-Aug 20.

G(BK C(3), P(3). Aug 11-31. Once

in the spring at C; Jun 20, 1973.

C{5) , P(33) . May 18- Jul 2.

C(l) P(8). Jun 1-29,

G(24), C(47), P{9H). May Ib-Jul 1.

G{4), C(-), P(4). Jun 2-29.

G(3), C(2), Pf9). Sep 11-Oct 1;

May 2,

0(17} , C(113), P{41). Oct 3-«ov 3.

Lar¥ae common on Juniperus communis.

C(“), P(“). Common diurnal species
in wooded locations, May 18-Sep 14.

C(-.), P(l). KUg 16. Three specimens at light
at C; May 1-15, 1971. adults found hibernating
in ca¥e near F.

C(339), P{435). Apr 29-Jun 1.

C{7). P{17). aug 13~Sep 9.

G(20), C(30), P(72). Hay 14-Jun 24; Jul 15-
Sep 20.

P{1). KUg 30.

C(2), P{7). May 15- Jun 2; Jul 4-AUg 5.

G{9), C(8), F{17). Jun 23-Jul 2?.

G{8}. C(7), P(24), Jun 17-nug 24,

C{16), P{21). Hay 10- Jul 7; Jul 20-Aug 7.

G(8), C(60), P(154). Hay 1-Jun 30; Jul 7-
aug 12.

G(169), C{201), P{1190). Jun 15-Aug 1.
G(2) C(36), P{132). Jun 26-Jul 31,

G{23), C(299), P{526). Jul 23-Sep 30.
G(59), C{348), P{167). nag 15-Uct 15.

C(20), P(25). Aug 15“0ct 9.

Pf3). Aug 5-Oct 22. Cominon in Kingston in
the fall of 1970, egg masses conspicuous on
shade trees, especially Acer saccharinum,
G(l), C(3), P(32). Jul l-Aug“Tf7~“‘~
C(l) , P{48) . Jul 5~Aug 27.

G(9), 0(40), P(460). Jul 5-Sep 1,

G(3), C(48), P(lOl). Jun 26-Aug 15.

G(7), C{44), P(200), Jun 11-aug 2?.

C(4), P{33). Jul 23-Sep 23.

G{2), P(l). Jul 23-«ug 19,

C(2), P(16). Jun 4-Aug 19.

G(9). G(86)j P(1215). Jun 1-Aus 25.

0(14), C(27), P(168), May lO-Aug 20.

C(l). Aug 9.

C(l). Jul 17.

C(13), P(277). Jun 1-Aug 28.

G(7), C(27), F(105). Hay 21-.Jul 16; Jul 13-
Aug 29.

C(15), P{5?). May 8-Jul 5.

G(l), P{17). May 8- Jun 29; Jul 24-aug 26.

G (1 ) . Jun 13 .

G(2), C(24), F(106), May 31~Jul 20; Aug 10.
G(3), C(36), P{109), Jun 1-Aug 8.

G(ll), C(21), P{55). May 10~Jun 29; Jul 29-
Aug 24, Frequency of melanic or partly melanic
individuals in 19?1 at P was 28/39 or 71,8%.
G(ll), C(55), P(145). May 14-aug 10. Frequen-
cies of melanic or partly melanic individuals;
14/32 or 43.85; (C, 1970); 20/5b or 35*7% (P,
1970); and 34/89 or 34.3% (P, 1971).

P(l). May 31.

C{18), P{39). May 18- Jul 9; Jul 26-Aug 16.

/. Res. Lepid.

32

Acronicta lithospila
Acronicta lobeliae
Acronicta morula
Acronicta noctivaga
Acronicta oblinita
Acronicta ovata
Acronicta pruni
Acronicta radcliffei
Acronicta retardata
Acronicta sperata
Acronicta superans
Acronicta tristis
Acronicta tritona
Acronicta vinnula

Adita chionanthi
Agriopodes fallax

Agroperina dubitans
Agroperina helva
Agrotis gladiaria
Agrotis venerabilis
Agrotis vetusta
Agrotis volubilis
Agrotis ypsilon

Alabama argillacea
Aletia oxygala

Amathes badinodis
Amathes bicarnea
Amathes c-nigrum

Amathes collaris

Amathes normaniana
Amathes opacifrons

Amathes smith!

Amathes tenuicula
Amolita fessa

Amphipoea americana
Amphipoea velata
Amphipyra glabella
Amphipyra pyramidoides
Amphipyra tragopoginis
Amyna octo

Anagrapha falcifera

Anaplectoides prasina
Anaplectoides pressus
Anathix puta
Anathix ralla
Anepia capsularis

Anomis erosa

Anomogyna dilvcida
Anomogyna elimata
Anomogyna youngi
Anticarsia gemmatilis
Anytus privatus
Apamea alia
Apamea amputatrix
Apamea cariosa

Apamea finitima
Apamea impulsa
Apamea inficita
Apamea inordinata
Apamea lignicolora
Apamea verbascoides

Apamea vultuosa
Apharetra purpurea
Aplectoides condita

WARD, ET AL

P(10). Jun 14-Jul 27.

G(l), C(6), P(9). May 16-Jun 30.

0(3), C(24), P(104). May 20-HUg 9.

G(16), C(19), P(82). May 14-Jul 6; Jul 23.
G(l), C(13), P(55). May 29-Jul 16; Aug 7-29.
G{1). C(52), P(875). Jun 9-Sep 7.

G(22), C(-), P(3). Jun 1-Jul 10; Aug 13-Sep 6.
P(4). May 21-Jul 3.

G(2), C(3), P(19). Jun 10-Jul 30; Aug 20.
0(49), 0(1), P(20). May 24-Jul 20.

C{4), P(16). May 25-Jul 15; Jul 27-Aug 6.
0(51), P(253). May 31-Aug 22; Sep 20.

C(l), P(29). May 31-Jun 22; Jul 23-aue 21.
0(25), C(7), P(16). May 21-Jun 26; Jul 23-
Aug 18.

G(3), 0(14), P(35). Aug 18-Oct 13.

C(-), P(2). Jul 3-9. One specimen at light
at C: Jul 12, 1967.

G(3), C(2), P(6), Jul 3-Sep 12.

P(8). Aug 8-19.

G(32), C(2). Jul 30; Sep 5-23.

G(15), C(ll), P(8). Sep 5-Oct 2.

G(l), C(l), P(l). Aug 27-Sep 13.

G(131), C(31), P(62). May 24-Jul 5; Jul 24.
G(226), C(48), P{112). May 8-Nov 3. Commonest
in late summer and fall; apparently several
overlapping generations.

0(3). Oct 2. Migrant from the south.

G(238), C(3), P(19). Jun 9-Jul 14; Jul 25-
Oct 7.

G(21), C(-). Sep 5-Oct 2.

G(4), 0(2), P(ll). Jul 31-Sep 3.

G(5866), 0(291), P(447). Jun 1-Nov 3. Several
overlapping generations; probably more than one
species involved.

C(-)'. Two specimens at light: nug 28, Sep 1,
1967.

0(5), P(42). Jul 13-Sep 27.

P(-). One female at light in sphagnum bog:

Aug 16, 1973.

G(5), 0(4), P(3). Aug 12-Sep 12.

G(29), P(l). Aug 8-Sep 7.

0(3), P(-). Jul 14-25. Two specimens at light
in sphagnum bog at P: Jul 8, 1970.

0(91), 0(8), P(9). Jul 24-Sep 12.

0(3), 0(2), P(9). Jul 13-Aug 17.

0(1), 0(20). Jul 23-Sep 25.

0(26), 0(31), P(121). Aug 5-Oct. 16.

0(228), 0(2), P(ll). Jul lO-Oct lb.

C(-). Two specimens at light: Sep 17, 1972,
Stray from the south.

0(68), 0(3), P(ll). May 29-Jul 15; Jul 23-
Oct 25.

0(2), 0(3), P(19). Jun 23-Aug 29.

0(1), P(5). Jun 18-Aug 8.

0(1), 0(1), P(l). Aug 14-Sep 2.

0(18), P(8). Aug 14-Sep 10.

o(-), P(l). Jul 26. One specimen at light

at 0: Jun 10, 1967.

O(-). One female at light: Oct 11, 1969.

Stray from the south
P(10). Aug 24-Sep 14.

0(1), 0(32), P(47). Jul 26-Sep 18.

0(1), P(l). Jul 20; Aug 25.

0(1 ), 0(3). Oct 2-12. Migrant from the south.
0(1), G(-), P(14). Sep 9-30.

P(l), Jul 7.

0(30), 0(13), P(34). Jun 29-Aug 14.

C(-), P(5). Jun 29-Jul 14. Once at 0: Jun
17, 1967.

0(5), 0(2), P(3). Jun 10-29.

0(6), 0(2), P(l). Jun 26-Jul 25.

0(1), P(38). Aug 8-Sep 8.

0(6) . Jun 14-26.

0(55), 0(10), P(53). Jun 16-Aug 17.

P(2). Jul 3. One specimen at bait at P:

Aug 10, 1971.

0(1), p(3). Jun 15-24: Jul 31.

P(l). Jul 13.

P(3). May 31-Jun 9.

13(1):23^42, 1974 ONTARIO MACROHETEROCERA

33

Archanara oblonga
Archanara subflava
Argyrostrotis anilis
Arzama diffusa
Arzama obliqua
Autographa ampla
Autographa biloba
Autographa mappa
Autographa precationis

Baileya dormitans
Baileya doubledayi
Baileya ophthalmica
Balsa labecula
Balsa malana

Balsa tristigella
Bleptina caradrinalis
Bombycia algens
Bomolocha abalienalis

Bomolocha baltimoralis
Bomolocha bijugalis
Bomolocha deceptalis
Bomolocha edictalis
Bomolocha madefactilis
Bomolocha manalis
Bomolocha palporia
Bomolocha sordidula
Caenurgina crassiuscula

Caenurgina erechtea

Caloe canadensis
Capis curvata
Catabena lineolata
Catocala amatrix
Catocala arnica
Catocala antinympha
Catocala blandula
Catocala briseis

Catocala cerogama
Catocala clintonii

Catocala coccinata
Catocala coelebs

Catocala concumbens
Catocala crataegi
Catocala epione
Catocala gracilis
Catocala habilis
Catocala ilia
Catocala meskei

Catocala mira
Catocala neogama
Catocala obscura
Catocala paleogama
Catocala parta
Catocala piatrix
Catocala praeclara
Catocala relicta
Catocala retecta
Catocala similis
Catocala sordida
Catocala subnata
Catocala ultronia
Catocala unijuga
Ceramica picta
Cerastis tenebrifera
Cerma cora
Chaetaglaea cerata
Chaetaglaea sericea
Chamyris cerintha
Charadra deridens
Chrysanympha formosa
Chrysaspidia contexta
Chrysaspidia putnami
Chrysaspidia venusta

G(5) , P(3) . Aug Ib-bep 14.

P(l). Jul 31.

G(4), C(2}, P(70). May 31-Jul 25.

C(5). Aug 2-12.

C(12) , P(2) . May 29-Jun 29.

C(-) , P{6) . Jul 8-23.

G(-). One specimen at light: apr 2b, 1970.

P(l). Jul 23.

G(32), C(15), P(17). May 24-Jul 7; Jul 26-
Oct 22.

C(23), P(123). May 19-Jul 23.

C(3), P(21). May 31-Jun 12; Jul 13-29.

G(l), C(29), P(220). May 4-Jun 30.

G(30), C(7), P(185). May 20-Aug 8.

G(30), C(9), P(16). May 31-Jun 17; Jul 22-
Aug 24.

G(l), C(5), P(14). Jun 1-Jul 9; Jul 31-Aug 13.
G(9), C(18), P(137). Jun 11-Aug 13.

P(l). Aug 21.

G(3), C(10), P(54). May 21-Jul lb; Jul 25-
Sep 12.

C(18), P(331). May 19-Jul 23; Jul 27-Sep 19.
0(2) . Jul 14; Aug 22.

C(73) , P(289) . May 15-Aug 25.

C(l) , P(l) . Jun 2; Jul 15.

C(9), P(ll). May 29-Jul 19; Aug 9.

C(4), P(7). May 21-Jun 26; Jul 27-Aug 20.

C(35), P(703). May 21-aug 25.

C(5). Jun 15-Jul 9; Jul 31.

G(598) C(llO) , P(43). May 10-Jun 26; Jul 13-
Sep 19.

G(460), C(27), P(25). May 2-Jun 24; Jul 16-
Oct 20.

C{1). Jul 19.

C(3), P(ll). Jun 20-Jul 25; Sep 4.

G(5), C(l), P(ll). May 24-Jul 8; Jul 21-Sep 6.
G(11), C(5), P(7). Aug 28-Oct 9.

C(-). One specimen at light: Aug 10, 1967.

C(-), P(2). Jul 22-30. Once at C: Aug 9, 1967.
C(l), P(7). Jul 13-Aug 24.

C(-), P(2). Sep 15-24. One specimen at light
at C: Aug 14, 1967. Four specimens collected at
bait at P; Aug 12-31, 1971.

C(16), P(46). Aug 5-Sep 23.

G(l), C(-). Aug 5. One specimen at light at C:
Jul 24, 1967. Also collected at Westbrook, Aug
11, 1963.

G(2) , C(5) , P(15) . Jul 20-Aug 20.

C(-), P(7). Jul 23-Aug 17. Larvae collected on
Myrica gale in sphagnum bog at P.

GTgT7~GTbTT P(21). Aug 10-Sep 28.

G(6), C(5), P(6). Jul 14-Aug 14.

C(4), P(142). Jul 16-Aug 25.

C(l) . Jul 20.

G(lb), C(88), P(378). Aug 15-Oct 27.

C(15) , P(70). Jul 13-Sep 23.

C{-). One specimen at light: Sep 6, 1967;
also one male at light at iunherstview: Aug 10,
1972.

P(-). One female at bait: Aug 9, 1971.

G(2), C(30), P(102). Aug 8-Oct 21.

G(l), C(l), P(65). Aug lO-Oct 1.

C(10), P(47). Jul 30-Sep 15-
G(4), C(l), P{2). Aug 7-Oct 2.

G(2), C(l), P(9). AUg 8-Sep 30.

P ( 1 ) . Aug 7 .

G(l), C(4), P(30). Jul 31-Oct 22.

G(4), C(2b), P(141). Aug 14-Oct 25.

C(4), P(35). Jul 15-Aug 25.

C(l), P(10). Jul 13-Aug 14.

C(19), P(58). Aug 7-Oct 9.

G(3), C(9), P(7). Jul 10-Aug 26.

G(l), C{3), P(18). Jul 23-Oct 9.

G(2). Jun 14; Aug 17.

G(-), C(10), P(219). Apr 19-May 28.

C(-), P(12). May 30- Jun 18.

G(3). Oct 2-13.

G(l) P(6) . Sep 20-0ct 22.

G(10), C{3), P(10). Jun 15-Jul 28.

G(4), C(74), P(71). May 21-Aug 12.

C(l)- P(9). Jul 2-28.

G(51), C(2), P(4}. Jun 3-Jul 4; Aug 9-Oct 9.
C(2), P(13). Jun 10-Jul 28; Aug 16-Sep 11.

G(l), C{1). Aug 5; Sep 15.

34

WARD, ET AL

/. Res. Lepid.

Chytolita morbidalis
Chytonix palliatricula

Chytonix sensilis
Colocasia flavicornis
Colocasia propinquilinea
Conservula anodonta
Copivaleria grotei
Cosmia calami
Crocigrapha normani
Crymodes devastator
Cryphia villificans
Cryptocala acadiensis

Cucullia asteroides
Cucullia convexipennis
Cucullia intermedia
Cucullia postera
Diarsia jucunda
Diarsia rubifera
Dipterygia scabriuscula

Dyspyralis illocata

Dyspyralis nigellus

Elaphria festivoides
Elaphria versicolor
Enargia decolor
Enargia infumata
Enargia mephisto
Eosphoropteryx thyatyroides

Epiglaea apiata
Epiglaea decliva

Epizeuxis aemula
Epizeuxis americalis
Epizeuxis concisa
Epizeuxis diminuendis
Epizeuxis forbesi
Epizeuxis Julia
Epizeuxis lubricalis
Epizeuxis rotundalis
Epizeuxis scobialis
Erastria albidula
Erastria bellicula

Erastria carneola

Erastria concinnimacula
Erastria muscosula
Erastria synochitis
Erebus odora

Euagrotis forbesi
Euagrotis illapsa
Eucirrhoedia pampina
Euclidia cuspidea
Eueretagrotis perattenta
Eueretagrotis sigmoides
Euherrichia monetifera
Euparthenos nubilis
Euplexia benesimilis
Eupsilia morrisoni

Eupsilia sidus
Eupsilia tristigmata
Eupsilia vinulenta
Eurois occulta
Euthisanotia grata
Euthisanotia unio
Eutolype electilis

Eutolype rolandi

C(27), P(87). May 31-Jul 23.

G(l), C(5b), H(302). May 19“Aug I4. Frequency
of f. iaspis at P was 1/101 or 1.0^ (1970) and
4/201 or 2.QP/0 (1971).

C(2), P(28). Jul 25-Aug 22,

0(2), C(107), P(341). May 1-Jun 18-; Jul 29.

G(b), C(45), P(37). May 4-Jun 23.

P(6). Jul 13-2b.

G(l), C(2), P(35). Apr 12-May 29.

G(l), C(25), P(7). Jul 14-Aug 4.

G(13), C(80), P(812). Apr 29-Jun 13.

G(423), C(7), P(7). Jul 3-Sep 20.

P(17). Jun 22-Jul 28.

C(-), P(l). Jul 29. One specimen at light at
C: Aug 2, 19b8.

G(b). Jul 10-27.

C(l), P(5). Jun 2b; Jul 23-Aug 19.

G(87) , C(24) , P(27) . May o-Jun 29; Jul 4-Sep 11.
P(2) , Jun 2b- Jul 7.

P(l). Jul 18.

P(4). Aug 10-2b.

P(l). Jul 20. One specimen at bait; Aug 13,

1971.

P(4). Jul 31-Aug 2b. Collected more commonly
at bait at P, especially at dusk, Aug 10-29,

1971.

P(4). Jul 11-Aug 1. Also collected at bait at
P. Lake Opinicon, one specimen resting among
leaf litter: Jul 15, 1973.

C(b8), P(991). May 14-Jul 22.

C(l) , P(4) . May 25-Jun 15.

C(13), P(8). Aug 5-Sep 23.

C(l). Jul 7.

C(l). Jul 19.

G(l), C(-), P(2). Aug 2-Sep 5, One specimen at
light at C: Sep 17, 1972.

P(l). Oct 2.

0(3), C(-). Nov 2-3. One specimen at light at
C: Nov 8, 1969.

G(30), C(521), P(2087). Jun 8-Oct 14 .

G(10), C(lb8), P(229). Jun 9-Oct 14.

C ( 1 ) . Aug 3 .

C(3), P(60). Jul b-Aug 12.

C(7), P(24). Jul 11-Aug 9.

C(l), P(5b). Jul 19-Aug 29.

G(4), C(b), P(lOb). Jul 10-Sep 3.

G(l), C(162), P(1900). Jul 3-Sep I4.

C(5), P(15). Jul 13-Aug 20.

G(81), C(51), P(120). Jun 18-Aug I4.

G(2), C(-), P(3). Jun 27-Jul 2. "One specimen
flushed in a willow field at C: Jul 10, 19b9. A
common diurnal species in sphagnum bogs at P,
Bedford Mills, Harrowsmith, and Westport. May
25- Jul 3.

G(b2), C(6b), P(107). May 24-Jul 15; Jul 16-
Sep 19.

G(l), P(l). Jun 13-19.

G(4), 0(99), P(170). Jun 13-Aug 29.

G(43), C(55), P(35). May 31-Aug 4.

Kingston, two specimens: Sep 23, 1958; Sep, 1961.
Stray from south.

C(5), P(89). Jun 28-Aug 10.

G(l8), C(3), P(20). Jun 9-Jul 4; Aug 8-Sep 7.
C(l) , P(18) . Sep 2-Oct 16.

G(7), C(9), P(ll). May 28-Jul 17; Aug 18.

P(2). Jul 9-15.

P(7). Jul 6-27.

C(9) , P(89) . May 30-Aug 12.

C(-). One specimen at light: Jul 9» 1973.

C(l), P(14). Jun 3-Jul 4; Jul 28-Aug 22.

G(4), C(l), P(46). Oct 6-30; Apr 8-May 18.
Overwinters as an adult, and like the other
congeners, appears to be commoner in the spring,
C(2) P(79). Oct 14-30; Apr 12-May 18.

P(17). Oct 27; Apr 12-May I5.

C(l), P(23). Oct 8; Apr 12-May 9.

G(2), C(2), P(16). Jun 10-13; Aug 10-Sep 15.
G(l), C(2), P(3). Jul 8-30; Sep 9.

G(3), C(2), P(6). Jun 10; Jul 5-Aug 26.

G(-), C(5), P(ll). Apr 19-May 19, All form
depilis . Typical form at Frankford: May I5,

1972.

Wooler, four specimens at light: May 8, 1972.

13(1):23^42, 1974 ONTARIO MACROHETEROCERA

35

Euxoa

albipennis

G{1).

Sep 6.

P(6). Sep 20~0ct 2.

Euxoa

bostoniensis

G(l),

Euxoa

detersa

G(l),

C{1). Sep 4-7.

Euxoa

divergens

P(l}.

Jul 27.

Euxoa

messoria

G(2),

P{3). Aug 19-Sep 15.

Euxoa

obeliscoides

G(5),

C{3) , P(18) . Jul 28-Sep

2.

Euxoa

perpolita

G(l).

Sep 5.

Euxoa

redimicula

G(6),

C(l) , P(32) . Jul 22-Sep

4.

Euxoa

scandens

G{2),

P(l). Jun 29“ Jul 10.

Euxoa

scholatica

C(4),

P(10). Jul 8-Aug 15.

Euxoa

servita

C(-).

One specimen at light: Jul

Euxoa

tesselata

G{3),

C(10), P(8). Jul 10-Sep

7.

Euxoa

velleripennis

G(3),

P(l). Aug,29-Sep 2.

Euxoa

sp. prob Ontario

P(2).

Aug 30“Sep 3.

Euxoa

sp.

P(3).

Aug 9-13.

Euxoa

sp.

G(l).

Jul 16.

Exyra

rolandiana

P(-).

Amherstview, Harrowsmith, ]

Faronta diffusa
Feltia ducens
Feltia geniculata
Feltia herilis
Feltia subgothica
Feralia comstocki
Feralia jocosa

Feralia major
Fishea enthea
Galgula partita

Westport and Portland. Larvae collected in
Sarracenia purpurea leaves, Feb 15“Jun 30, and
Jul 24-Oct 20. Overwinters in third instar.
G(33), C(2), P(25). May 31-Jul 10; Jul IJ-Sep 5.
G(196), C(20), P{18). Jul 25-Sep 21.

G(l), C(3), P(93). Aug 7-Sep 7.

G(2), C{1), P(8). Jul 20-Aug 26.

G(61), C(14), P(20). Jul lO-^Sep 12.

C(2), P(l). May 3-28.

C(~)s P{2), May 11-16, Two specimens at light
at C: May 1, 1970.

G(l), C{-)j P(9). May 2-15.

C(-), P(23). Sep 17-Oct 15.

G(15), C(9), P(15)« Jun 2-Oct 26. Commonest in

Graphiphora haruspica
Haploolophus mollissima
Harrisimerana trisignata
Kelicoverpa zea
Heliothis phloxiphaga
Helotropha reniformis
Hemipachnobia monochromatea

Heptagrotis phyllophora
Homoglaea hircina
Homohadena badistriga
Homohadena infixa
Homorthodes furfurata
Hormisa absorptalis
Hormisa bivittata
Hormisa litophora
Hormisa orciferalis
Hydroecia micacea
Hypena humuli

Hypenodes fractilinea
Hypenodes palustris

Hypocoena inquinata
Hyppa xylinoides

Lacinipolia anguina
Lacinipolia implicata
Lacinipolia lorea

Lacinipolia raeditata
Lacinipolia renigera

Lacinipolia vicina
Lascoria ambigualis
Lemmeria digitalis

Leucania commoides,
Leucania inermis
Leucania insueta
Leucania phragraatidicola

Leucania pseudargyria
Leuconycta diphteroides

late summer and fall.

G(3), P(8). Jul 23-iiUg 12.

G(l), C(4), P(33). Jun 10-Aug 1.

C(-) , P(13) . Jun 11-Aug 2.

G(4), C(6), P(20). Sep 4-Oct 16.

G(4), P(l). Jun 17: Jul 31~Aug 16.

G(2) , C(l) , P{2). Aug 17-Sep 5-
P(l). Jun 25. Common at light in sphagnum bogs
at P, Bedford Mills, and Westport, Jun 10-Jul 7.
C(l), P(ll), Jul 6-31.

G(~), C(l), P(17). Apr 12-May 11.

G{3), C(8), P{44). Jul 3-Aug 9.

C(2), P(l). Jul 6-18.

C(7), P{15). Jul 4-Aug 7.

G(7), C(-), P(2). Jul 5-Aug 15,

G(7), C(l), P(l). Jul 14-Aug 14.

C(l). Jul 29.

G{17), C{5), P(4). Jul 6-Sep 18,

G (1 ) . Aug 18.

G(15), C(3), P(12). May 4-Jul 2; Aug 2-Sep 7.
Overwinters as an adult; appears commonest in
the spring. Two adults found hibernating in
limestone cave near Belleville, Sep 11, 1970,
C(-), P(48). Jun 7~Jul 8; Jul 28-Sep 4,

A small dark form, probably this species, is
common in sphagnum bogs at P, Harrowsmith, Bed-
ford Mills, and Westport, Jun 21- Jul 8, and
Aug 15-Sep 9, at light and easily disturbed in
the daytime,

P(l), Aug 3,

G(42), C(60), P(99). May 14-Jun 29; Jul 14-
Sep 16.

G(l). C(30), P(166). May 10-Jun 24.

C(23), P(457). Aug 9-Sep I5,

G{3), C(~). Jul One specimen at light at

C: Jun 15, 1967.

G{42), C(8), P(27). Aug 7-Sep 5.

G(155l, C{23), P(32). Jun l6-0ct 13. Several
overlapping generations.

G{27), C(13), P(27). Jul 8-iiug 9.

P{5). Jun 6-18; Aug 18-Sep 2.

C(l), P(5). Sep 20-0ct 2. One specimen at light
at Cranberry Lake: Oct 12, I969,

G(143), C(68), P(138). Jun 18-Aug 25.

C(22), P(41). Jun 1-Aug 7.

G(24). C(7}, P{92). Jun 16-Aug I4,

G(324), C(35), P{34). Jun 1-Jul I5: Aug 12-
Oct 13 .

G(13), C{28), P{3). Jul 7-Aug b.

G(6), C(ll), P(ll), Jun 2-Jul 28.

/. Res. Lepid.

36

Leuconycta lepidula
Lithomoia solidaginis
Lithophane amanda
Lithophane antennata

Lithophane baileya
Lithophane bethunei
Lithophane disposita
Lithophane fagina
Lithophane grotei
Lithophane hemina

Lithophane innominata
Lithophane laticinerea
Lithophane oriunda
Lithophane patefacta
Lithophane pexata
Lithophane querquera
Lithophane semiusta
Lithophane tepida
Lithophane unimoda
Lithophane sp.
Lomonaltes eductalis
Luperina passer
Macronoctua onusta
Magusa orbifera
Marathyssa basalis
Marathyssa inficita
Metalectra discalis

Metalectra quadrisignata
Metalepsis fishii
Metalepsis salicarum
Metaxaglaea inulta
Mocis texana
Morrisonia confusa
Morrisonia distincta
Morrisonia evicta
Nedra ramosula

Neoerastria apicosa
Neoerastria caduca
Neperigea costa

Nephelodes emmedonia
Ochropleura plecta

Ogdoconta cinereola
Oligia chlorostigma
Oligia exhausta

Oligia fractilinea
Oligia illocata
Oligia mactata

Oligia minuscula

Oligia modica
Oligia semicana
Oncocnemis saundersiana
Oncocnemis viriditincta
Orthodes crenulata
Orthodes cynica
Orthosia alurina
Orthosia hibisci
Orthosia revicta
Orthosia rubescens
Palthis angulalis

Pangrapta decoralis
Panopoda carneicosta
Panopoda rufimargo
Panthea pallescens

WARD, ET AL

0(2), P(7). Jun 7-Jul 10; Aug 13.

0(1), 0(1), P(3). Sep 8-23.

P(l). Apr 20.

0(1), 0(3), P(10). Sep 21-Oct 13; Apr 30-May
22. Adults of this species overwinter and, like
other members of the genus, are more prevalent
in the spring months.

C(-), P(33). Sep 13-30; Apr 12-May 18.

0(1), P(3). Apr 8-May 19.

P(2). Apr 27-May 27.

0(3), P(33). Sep 22-Oct 13; Apr 12-May 17.

0(7), 0(2), P(24). Oct 1-11; Apr 12-May 19.

0(5), P(31). Oct 21; Apr 19-Jun 5. Frequency
of f. lignicosta at P was 20/31 or 83.9%.

P(l) . May 2.

0(5), 0(1), P(16). Oct 21; Apr 8-May 21.

0(1). Apr 8.

P(l). May 11.

P(14). Apr 23-May 25.

P(l). May 11.

0(2), P(8). Apr 27-May 23.

P(l). Apr 23.

0(0), O(-), P(A). Oct 10; May 3-15.

0(1) . Apr 8.

0(2) , P(2) . Jun 27; Aug 2-13.

0(o). Jun 14-Jul 1/+.

0(3) , P(4) . Sep 7-Oct 7.

P(137). Sep O-Oct 10. Migrant from the south.
0(1), 0(37), P(55). May 8-Jun 18.

0(5), 0(00), P(170). May 30-Aug 29.

O(-), P(3). Jul 29-Aug 10. One specimen at
light at 0: Jul 20, 1968; another at bait: Jul
15, 1969.

0(1), 0(1), P(ll). Jun 10-Aug 7.

P(0). May 11-21.

O(-), 0(1), P(13). Apr 30-May 18.

0(1) , P(2) . Sep I7-2I0

P(l). Sep 13. Stray from the south.

0(55) , P(133) . May 1-Jun 12.

O(-), 0(033), P(363), May 1-Jun 7.

0(3), 0(8), P(80). May 1-29.

0(22), 0(120), P(200). May 0-Jun 27; Jul 3-
Sep 22.

0(1). Jun 10.

0(3), P(0). Jun 1-18; Jul 17-31.

O(-), P(13). Jun 27-Aug 2. One specimen at
light at 0: Jun 28 1972.

0(83), 0(05), P(91). Aug 19-Sep 30.

0(80), 0(02), P(136). May 21-Jul 15; Jul 25-
Sep 18.

0(00), 0(0), P(22). Jun 10-Jul 31; Aug 3-Sep 16.
P(l) . Jul 23 .

0(2), P(l). Jul 20-Aug 9. One specimen at light
at Amherstview: Aug 2, 1972.

0(lb), P(l). Jul 20-Aug 22.

P(10). Sep 15-29.

0(1), P(7). Sep 15-Oct 1. Larva collected on
flower of Oypripedium calceolus (Jun 2, 1973 at
P), and reared on C. acaule ; pupated Jun 23, one
male emerged Sep 12.

P(25). Aug 15-Sep 15. Also collected in sphag-
num bogs at P, Bedford Mills, and Westport, Jul
29-Sep 6, mostly at light. Form grahami rare at
P, common at Westport.

0(21), 0(1), P(6). Jul 23-Sep 26.

P(6). Jun 28-Jul 26.

0(1), 0(1). Aug 30-Sep 0.

0(1). Sep 5.

0(3), 0(1), P(72). Jun 25-Sep 5.

0(1), 0(5), P(8o). May 18-Jul 11.

0(1). May 1.

0(38), 0(41), P(107). Apr 12-Jun 10.

0(5), 0(16), P(124). Apr 20-May 31.

0(1) 0(21) P(173). Apr 19-May 24.

0(12), 0(64), P(171). May 21-Jul 6; Jul 20-
Sep 15.

P(47). Jun 10-Aug 28.

0(2) 0(31), P(87). Jun 20-Aug 19.

0(46) P(89). Jun 9-Aug 13.

0(126), P(422). May 18-Jul 1; Jul 5-Aug 28.
Apparently two generations, the second much more
prevalent .

13(1):23^42, 1974 ONTARIO MACROHETEROCERA

37

Papaipema appassionata P(3). Aug 2b-Sep 8, Also collected at light,

or as larvae in the roots of Sarracenia pur-
purea . in sphagnum bogs at P, Marlbank, Amherst-
view, Bedford Mills, Westport and Portland.
Larvae: Jun 30-Jul 18; adults: Aug 25-Sep 6.

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

Papaipema

cataphracta

eupatorii

furcata

impecuniosa

inquaesita

lysimachiae

marginidens

nebris

nepheleptena

pterisii

purpurifascia

speciosissima

Papaipema unimoda
Paradiarsia littoralis
Parallelia bistriaris
Parastichtis discivaria
Peridroma saucia
Phalaenostola larentioides
Philometra eumelusalis
Philometra hanharai
Philometra metonalis
Phlogophora iris
Phlogophora periculosa

Phoberia atomaris
Plathypena scabra

Platyperigea meralis

P(21)

. Aug :

Platyperigea multifera

0(4),

P(56).

Platypolia anceps

P(3).

Oct 1'

Platysenta vecors

0(1),

P(19).

Platysenta videns

0(19)

, 0(7),

Plusia aerea

0(3),

0(2), .

Plusia aeroides

0(1) ,

0(1), :

Plusia balluca

C(-),

P(i).

Plusiodonta compressipalpus
Polia adjuncta

G(3). Oct 2-7.

P(2). Sep 27-30.

C(l) , P(A). Sep 6-27.

0(4). Oct 2-6.

P(5). Sep 15-Oct 9.

P(l). Sep 11.

0(1) . Oct 11.

0(1). Sep 5.

0(1), P(l). Sep 14-23.

0(5), P(4). Aug 22-Sep 28.

0(1), 0(7), P(12). Aug 26-Oct 25.

P(25). Sep 9-Oct 9. Frequency of f. regalis
was 6/25 or 24.0^.

C(-). One specimen at light: Sep 10, 1968.

0(7) . Jun 17-Jul 3.

0(1), P(33). Jun 1-Sep 7.

0(1), P(2). Aug 7-12.

0(11) , 0(5) , P(65) . Jun 23; Jul 28-Oct 27.
0(6), 0(7), P(3). Jul 10-31; Sep 3-13.

P(l). Jul 28.

0(1), Jul 17.

P(l) . Aug 9.

0(13), 0(16), P(30). May 25-Jul 25.

0(4), P(23). Aug 15-Sep 13. Frequency of f.
v-brunneum at P was 8/23 or 34.8%.

OTI), 0(34) , F(284). Apr 29-May 22.

0(98), 0(15), P(12). Jun 5; Jul 10-Nov 3.
Commonest in late summer and fall.

17-Sep 4.

Aug 15-Sep 15.

-14.

May 25-Jun 24; Aug 1-19.

P(22). Jun 2-30; Jul 28-Sep 4.
P(10). Jun 9-Jul 15: Aug 27-Oct 2.
P(3). Jun 26- Jul 28,

Jul 23. One specimen at light at
C”. Jul 23, 1967. Also collected once at King-
ston: Jul 24, 1963.

0(1). Jul 16.

0(l6l), 0(63), P(83). May 12-Jul 13; Jul 24-

Polia assimilis
Polia atlantica
Polia detracta
Polia grandis
Polia imbrifera
Polia latex
Polia legitima
Polia lilacina
Polia lutra
Polia nimbosa
Polia obscura
Polia purpissata
Polia rugosa

Polia segregata
Polia subjuncta
Prodenia ornithogalli

Protagrotis niveivenosa
Protocryphia secta
Protolampra brunneicollis
Protorthodes curtica
Protorthodes oviduca
Proxenus miranda

Psaphida resumens
Psectraglaea carnosa
Pseudaletia unipuncta

Pseudeva purpurigera
Pseudoplusia includens
Pyreferra citromba
Pyreferra pettiti
Pyrrhia umbra
Raphia frater

Sep 7.

0(2), P(8). Jun 28-Aug 1.

0(47), P(3). Jun 2-Jul 5; Aug 8-Sep 4.

0(11), P(63). Jun 15-Jul 29.

0(1), 0(1), P(16). May 30-Jul 10.

P(l). Jul 30.

0(9), P(77). May 15-Jul 8.

0(25), 0(6), P(ll). Jun ll~Aug 5.

0(9), C(-), P(l). Jun 22-Jul 24.

0(1), 0(2), P(46). Jun 1-Jul 15.

0(2), P(5). Jul 15-28.

0(4), P(36). May 19-Jun 28.

0(1), 0(2), P(14). Aug 7-27.

P(“). One male at light in sphagnum bog at
P: Jun 21, 1973.

0(2), P(26). Apr 12-May 19.

0(37), 0(2), P(16). Jun 7-Sep I4.

0(2), C(-). Sep 7-Oct 11. One specimen at
light at 0: Nov 1, 1969; another at bait:

Oct 12, 1970.

0(10), P(l). Aug 2-19.

C(-), P(78). Jun 9-Aug 5.

0(4), 0(1). Jul 3-13; Aug 18-26.

0(1). Jul 15.

0(15), 0(26), P(30). May 23-Jul 12.

0(67), 0(7), P(28). May 28-Jul 1; Jul 29-
Sep 4.

0(3), P(39). Apr 12-May 18.

P(3). Sep 17-27.

0(649), 0(137), P(67). May 8-Nov 4. Several
overlapping generations.

0(10), 0(6), P(ll). Jul 7-Sep 5.

0(8), 0(4), P(19). Sep 6-Oct 9.

C(l), P(6). Apr 12-May 18.

0(7), P(43). Apr 12-May 29.

0(1), P(l). Jun 20-Jul 17.

0(7), 0(224), P(1135). May 14-Aug 23.

38

WARD, ET AL

J. Res. Lepid.

Renia factiosalis
Renia f lavipunctalis
Renia sobrialis
Rhynchagrotis anchocelioides
Rhynchagrotis brunneipennis
Rhynchagrotis cupida
Rivula propinqualis

Schinia florida

Schinia marginata
Schinia trifascia
Scoliopteryx libatrix

Scotogramma trifolii
Senta defecta
Sideridis maryx
Sideridis rosea
Simyra henrici
Spaelotis clandestina
Spargaloma perditalis
Spargaloma sexpunctata
Spodoptera exigua
Spodoptera frugiperda
Sunira bicolorago
Syngrapha alias
Syngrapha altera
Syngrapha epigaea

Syngrapha rectangula
Syngrapha selecta
Tarache terminimaculata
Tarachidia candefacta
Tarachidia erastrioides

Trachea delicata
Tricholita signata

Trichoplusia ni
Trichoplusia oxygramma
Ufeus satyricus
Ulolonche culea
Ulolonche raodesta
Xanthia flavago
Xylena cineritia
Xylena curvimacula

Xylena nupera

Xylomiges alternans
Xylomiges dolosa
Xylomoia chagnoni
Zale aeruginosa
Zale cingulifera
Zale duplicate
Zale galbanata
Zale helata
Zale horrida
Zale lunata

Zale minerea
Zale obliqua
Zale phaeocapna
Zale unilineata
Zanclognatha cruralis
Zanclognatha jacchusalis
Zanclognatha laevigata
Zanclognatha lituralis
Zanclognatha ochreipennis
Zanclognatha protumnusalis
Zenobia pleonectusa

NOLIDAE

Celama cilicoides

Nola ovilla
Sarbena minuscula

C(10), P(255). Jul 23-Sep 4.

G(7), 0(144), P(823). Jul 10-Sep 15.

P(l). Aug 2.

P(l) . Aug 14.

P(6). Jul 24-Sep 12.

0(2) , P(6) . Jul 30-Sep 28.

0(9b), 0(17), P(2y). Jun 11-Jul 25; Aug 6-
Sep 15.

0(2), P(l). Jul 21-29. Adults common in flowers
of Oenothera, the larval foodplant.

0(1) , 0(2) , P(3) . Jul 27-Aug 14.

0(1), 0(2), P(l). Jul 17-Aug 13.

0(3), 0(1), P(2). Sep 1-7; May 1-16. Adults
found hibernating in caves at P, Lake Opinicon,
and Westport, Aug 30-Oct 25.

0(10), 0(2). Jun 10-Sep 6.

P(l). Jul 24.

P(l) . Jun 20.

0(21), 0(3), P(86). May 23-Jul 5.

G(53), 0(4), P(9). May 16-Jun 21; Jul 24-Sep 5.
0(42), C(-), P(9). Jun 20-Jul 24; Aug 6-Oct 3.
0(6), P(14). May 30-Jul 13; Aug 3.

0(1), 0(34), P(96). May 30-Sep 2.

0(1). Oct 2.

0(131), 0(5), P(62). Aug 17-Oct 24.

0(50), 0(30), P(113). Sep 14-Nov 2.

0(1) . Jul 20.

P(12) . Aug 10-Sep 3 .

0(1), P(-), 0(1). Aug 14-Sep 5. Reared from
larvae on Myrica gale in bogs at P and Portland.
0(1), P(l). Jul
P(3). Aug 13-31.

0(2), P (2). Jun 29- Jul 10; Aug 13.

0(83), 0(2), P(17). Jun 1-Jul 3; Jul 15-Aug 31.
0(88), 0(4), P(24). Jun 1-Sep 16. Probably two
overlapping generations.

P(l). Jul 22o

0(2), P(l). Jul 28-AUg 14. One specimen at
bait at 0: Jul 19, 1969.

0(24), C(-). Aug 16-Oct 11.

P(l). Sep 11.

0(1), 0(3), P(5). Sep 9-Nov 4.

0(5), 0(30), P(180). May 18-Jul 2.

0(1), 0(6), P(60). May 14- Jun 17.

0(1), C(-), P(7). Sep 17-Oct 11.

0(5) , P(2) . Apr 8-May 15.

0(6), 0(4), P(49). Oct 11; Apr 12-May 26.

Adult overwinters.

0(2), C(-), P(l). Apr 30-May 4. One specimen
at bait at 0: Sep 21, 1969.

Pa5) . May 8-30.

O(-) , 0(16) , P(26) . May 1-30.

0(1) , P(5) . Jun 29-Jul 28.

P(8). May 30-Jul 7.

0(7), P(19). May 3-30.

0(11), P(41). May 6- Jun 6.

0(6), 0(4), P(l). May 16-Jun 22; Jul 25.

0(1), 0(38), P(16). May 19-Jun 29.

0(1), 0(2), P(5K Jun 7-Jul 15.

0(1), O(-), P(l). Aug 4; Oct 11. One specimen
at light at 0; Nov 2, 1969; another at bait:

Oct 24, 1970.

0(1), 0(168), P(152). May 1-Jun 30.

0(2). May 8- Jun 1 .

0(31) , P(42) . May 1-Jun 7.

0(4), P(3). May 11-Jun 11.

0(11), P(10). Jun 13-Jul 9.

0(4), P(4). Jul 12-Sep 9.

0(16), P(llO). Jul 11-Aug 25.

0(2) , P(21) . Jun 23-Aug 2.

0(5), 0(125), P(887). Jul 10-Sep 14.

0(4), 0(18), P(22). Jul 2-Sep 2.

0(1), 0(5), P(3). Jul 16-Sep 11.

0(1), O(-). Aug 5. Two specimens at light at
0: Jul 6-10, 1970 (ROM).

0(3). May 12-25.

0(3), 0(9), P(67). Jun 12-Aug 3. Possibly
more than one species involved.

l3(l):23-42, 1974 ONTARIO MACROHETEROCERA

39

NOTODONTIUAE

Cerura borealis
Cerura cinerea

Cerura modesta
Cerura multiscripta

Cerura occidentalis
Clostera albosigma

Clostera apicalis

Clostera strigosa
Dasylophia thyatiroides
Datana angusi
Datana contracta
Datana integerrima

Datana ministra
Datana perspicua
Ellida caniplaga
Gluphisia avimacula
Gluphisia lintneri
Gluphisia septentrionis

Heterocampa bilineata

Heterocampa biundata
Heterocampa guttivitta

Heterocampa manteo
Heterocampa obliqua
Heterocampa umbrata
Hyperaeschra georgica
Macrurocampa marthesia
Nadata gibbosa
Nerice bidentata
Notodonta simplaria

Notodonta stragula

Odontosia elegans

Oligocentria lignicolor
Peridea angulosa

Peridea basitriens
Peridea ferruginosa

Phaeosia rimosa

Schizura apicalis
Schizura badia
Schizura concinna
Schizura ipomeae
Schizura leptinoides

Schizura semirufescens
Schizura unicornis
Symmerista albifrons
Symmerista canicosta
Symmerista leucitys

SATURNIIDAE

Actias luna
Antheraea polyphemus
Automeris io
Callosamia promethea

Hyalophora cecropia

SPHINGIDAE
Amphion nessus

C(3), P(33). May 31-Jun 15} Jul 1-Aug 13.

G(3), C(49), P(128). May 25-Jul 25; Jul 23-

Aug 25.

G(5), c(13), p(43). May 6-Jun 7; Jul 15-Aug 15.
C(2), P(2). May 30-Jun 19. All females, males
apparently diurnal. Reared from larvae collected
on Populus grandidentata ,

G(5TTcTr7)7 P (30). May 8-Jul 1; Jul 11-Aug 20.
G(3), C(191J, P(254). May 3~Jun 25; Jun 22-
Aug 23.

G(3), C(20), P(19). May 16-Jun 19; Jul 27-
Aug 20.

C(2), P(4). May 30-Jun 22.

G(4), P(5). May 23-Jun 18; mg 5-16.

C(23), P(39). Jun 14-Aug 1.

G(9), C(5), P(38). Jun 7-Jul 31.

C(l). Jul 18. More common in previous years
at C; Jun 13-Aug 10.

G(7), C(26), P(lOl). Jun 2-Aug 2.

G(ll), C(l), P(13). Jul 7-Aug 13.

G(7), C(218), P(120). May 1-Jul 25; Oct 2.

C(35j, P(106). May 2- Jun 11.

G(3), C(-), P(7). Apr 23-May 11.

G(21), C(209), P(535). May 8- Jul 5; Jul 3-
Aug 26.

G(21), 0(12), P(23). Jun 2-Jul lo; Jul 23-
Aug 26; Sep 27.

C(20), P(28). Jun 8-Jul 31.

G(2), C(522), P(484). May 5- Jul 21; Aug 20;

Sep 10.

C(-). One specimen at light: Jul 23, 1968.

C(17), P(65). Jun 2-Aug 8.

C(49), P(89). May 31-Jul 28.

G(l), C(18), P(51). May 14-Jul 14; Aug 10.

G(8), C(106), P(386). Jun 19-Aug 25.

G(2), C(215), P(586). May 21-Aug 22.

G(26), C(62), P(104). May 14-Aug 22.

G(2) , C(5) , P(5) . May 16-Jun 13; Jul 16-
Aug 12.

G(9), C(20), P(60). May 31-Jul 10; Jul 18-
Aug 17.

G{2), C(38), P(46). May 21-Jul 11; Jul 15-
Aug 25.

C{24), P(244). Jun 13-Aug 16; Sep 3.

G(12), C(169), P(543). May 31-Jul 5; Jul 3-
Aug 31.

C(8), P(38). Jun 13-18; Jul 10-Aug 2.

G(2), C(165), P(202). May 29- Jun 28; Jun 29-
Aug 24.

G(4), C(113), P(195). May 8-Jul 18; Jul 16-
Sep 15.

C(-), P(9). Jun 1-10; Jul 13-26.

C(5), P(38). Jun 11-Aug 9.

G(l), C(2), P(5). Jul 23-Aug 12.

G(4), C(29), P(51). Jun 19-Aug 7.

G(4), C(512), P(1610). May 21-Sep 14. Probab-
ly two or more overlapping generations.

G(5), C(41), P(117). Jun 12-Aug 16.

G(33), c(83), P(138). May 19-Aug 22.

C(4) . Jun 21-22.

C(26), P(66). Jun 8-Jul 23.

C(30), P(39). Jun 7-Jul 17.

G(l), C(142), P(750). May 14-Jul 30.

G(ll), C(19), P(89). Jun 2-Jul 28.

G(6), C(lOl), P(l68). May 30-Jul 4.

C(-), P(-). Several females at light in 1972
at C. Diurnal males attracted to caged, labor-
atory-bred females at C .and P, Jun 10- Jul 8.
Larvae reared on Prunus . Previously common
in the city of Kingston, feeding on Syringa .
G(2), C(19), P(36). May 29-Jun 20.

G(-), C(-), P(-). Diurnal, May 24-Jul 10,
visiting flower blossoms (especially Syringa )
and carrion. Larvae collected on Vitis riparia .

/. Res. Lepid.

40

WARD, ET AL

Ceratomia amyntor
Ceratomia undulosa

Cressonia juglandis

Darapsa myron

Darapsa pholus
Darapsa versicolor
Deidamia inscriptum
Dolba hylaeus
Eumorpha achemon
Eumorpha pandorus

Hemaris diffinis

Hemaris thysbe

Hyles lineata
Lapara bombycoides
Manduca quinquemaculata
Pachysphinx modesta
Paonias excaecatus
Paonias myops
Smerinthus cerisyi
Smerinthus jamaicensis

Sphecodina abbottii

Sphinx canadensis
Sphinx chersis
Sphinx drupiferarum
Sphinx eremitus
Sphinx gordius
Sphinx kalraiae

THYATIRIDAE

G{11), C(27), P(b7). May 31-Aug 16,

G(6), C{176), P(1026). May 8-Sep 7. Bimodal
flight period, with peaks in mid- June and mid-
to late July.

G(l), C(41), P(144). May 14-Jun 19; Jun 29-
Aug 1; Aug 26,

G(19), C(39), P(34). May 31-Aug 10. Larvae
common on Vitis riparia ,

G(2), C(19T7^92K~tey 21-Aug 12.

C(l) , P(14) , Jun 28-Jul 29.

G(2), C(4), P(6). May 6-30.

P(5). Jun 17-Jul 14.

G(7), P(l), Jul 3-Aug 14,

C(l), P(7). Jul 10-Aug 14. One larva col-
lected on Vitis riparia at Lake Opinicon;
formerly common on Parthenocissus in Kingston.
C(-), P(-). Common diurnal species, May 12-
Jul 31, frequently visiting Syringa blossoms;
observed ovipositing on Lonicera .

C(-), P(-). Diurnal, May 29-Jun 9, visiting
mostly Syringa blossoms.

C{1), Aug 15.

G(3), C(265), P(664). May 31-Aug 10.

G(12), C(-). Jul 16-Aug 12,

C(27), P(48). May 31-Aug 15.

G(13), C(108), P(238). May 29-Aug 18.

G(31), C(00), P(140). May 24-Aug 17.

C(58), P(38), May 4-Jun 26; Jul 11; Jul 30.
G(10), C(23), P{75). May 23-Aug 22. All f.
norm, geminatus except for two typical .jamai-
censis from P.

C(4), P (13 ) . May 6-Jun 19. Larvae collected
on Vitis riparia .

G(2) , CTl) , P(^. Jun 12-Jul 29.

G(14), C(7), P(102). Jun 10-Aug 22.

G(37), C(4), P(3). May 31-Aug 5.

C(-). One male at light: Jul 8, 1974»

G(5), C(26), P(126). May 19-Jul 28; Aug 15.
G(6), C(9), P(32). Jun 1-Aug 13.

Euthyatira pudens

Habrosyne scripta
Pseudothyatira cymatophoroides
Pseudothyatira expultrix

C(2), P(26), May 1-30. Includes tv/o indivi-
duals of form pennsylvanica from P.

G(l), C(l), P(2T: Jul 13-28.

C(2), P(2), Jul 14-Aug 1.

C(2), P(l), Jun 22-Jul 28. Other records:

P, Aug 10, 1971 (at bait), Jun 21, 1973 (at
light in bog); V/estport, Jun 29, 1973 (at
light in bog).

ZANOLIDAE

Apatelodes angelica

C(26), P(272). Jun 8-Aug 11.

ONTARIO MACROHETEROCERA

41

ACKNOWLEDGEMENTS

The authors wish to express their gratitude to the following
persons for their assistance in the determination of some species:
Dr. J. G. Franclemont, Cornell University (Noctuidae); Dr. D.
F. Hardwick, Biosystematics Research Institute, Ottawa (Noc-
tuidae); Dr. W. C. McGuflBn, Biosystematics Research Institute,
Ottawa ( Geometridae ) , and J. C. E. Riotte, Royal Ontario
Museum, Toronto (Arctiidae. Lymantriidae ) .

REFERENCES

BESCHEL, R. E. et al. 1970. List of the vascular plants of the Kingston
region. Queen’s University: Fowler Herbarium.

FERGUSON, D. C. 1954. A revision of the genus Hypenodes Doubleday
with descriptions of new species ( Lepidoptera, Phalaenidae). Can. Ent.
86: 289-298.

FERGUSON, D. C. 1955. The Lepidoptera of Nova Scotia. I. Macrolepi-
doptera. Bull. Nova Scotia Mus. of Sci. No. 2.

FERGUSON, D. C. 1972. Two new conifer-feeding species of the genus
Semiothisa (Lepidoptera: Geometridae). Can. Ent. 104: 563-565.

FORBES, W T. M. 1948. Lepidoptera of New York and neighbouring states.
Part II. Cornell Univ. Agric. Exp. Sta. Memoir 274.

FORBES, W. T. M. 1954. Lepidoptera of New York and neighbouring states.
Part III. Cornell Univ. Agric. Exp. Sta. Memoir 329.

FORBES, W. T. M. 1960. Lepidoptera of New York and neighbouring
states. Part IV. Cornell Univ. Agric. Exp. Sta. Memoir 371.

HARMSEN, R., P. D. N. HEBERT & P. S. WARD. 1974. On the origin of
austral elements in the moth fauna of south-eastern Ontario, including
a number of species new for Canada. J. Res. Lepid. 12:127-134.

LA FRANCE, J. 1968. The nocturnal insect catches at predetermined time
intervals in the organic soil district of Ste. Clotilde, southwestern
Quebec. Ann. Ent. Soc. Quebec, 13: 32-54.

McDUNNOUGH, J. 1938. Checklist of the Lepidoptera of Canada and the
United States of America, Part 1. Macrolepidoptera. Mem. S. Calif.
Acad. Sci. Vol. I.

McDUNNOUGH, J. 1949. Revision of the North American species of the
genus Eupithecia (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat.
Hist. 93(8): 533-728.

McFarland, N. 1965. The moths (Macroheterocera) of a chapparal
plant association in the Santa Monica mountains of southern California.
J. Res. Lepid., 4: 43-74.

42

WARD, ET AL

/. Res. Lepid.

McGUFFIN, W. C. 1967. Guide to the Geometridae of Canada (Lepi-
doptera). I. Subfamily Sterrhinae. Mem. Ent. Soc. Canada, No. 50.

McGUFFIN, W. C. 1972. Guide to the Geometridae of Canada (Lepi-
doptera). II. Subfamily Ennominae. 1. Mem. Ent. Soc. Canada, No. 86.

MOORE, S. 1955. An annotated list of the moths of Michigan, exclusive of
Tineoidea ( Lepidoptera ) . Misc. Publ. Mus. Zool., Univ. Mich, No. 88.

PRESTON, F. W. 1948. The commonness and rarity of species. Ecology
29: 254-283.

QUILLIAM, H. R. 1973. History of the birds of Kingston, Ontario.
Kingston; Kingston Field Naturalists.

RIOTTE, J, C. E. 1969. Fine neue Art der Gattung Anisota (Lep. Satur-
niidae) in Nordamerika. Entomol. Zeitschr. 79(13): 141-146.

SNYDER, L. L., E. B. S. LOGIER, T. B. KURATA, F. A. URQUHART, &
J. F. BRIMLEY. 1941. A faunal investigation of Prince Edward
County, Ontario. Univ. of Toronto Studies, Biol. Series No. 48.

SOPER, J. H. 1962. Some genera of restricted range in the Carolinian flora
of Ontario. Trans. Roy. Can. Inst. 34(1): 2-56.

WILLIAMS, C. B. 1964. Patterns in the balance of nature. London:
Academic Press.

Journal of Research on the Lepidoptera

13(l):43-48, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A NEW SPECIES OF HYPAGYRTIS
(GEOMETRIDAE)

ROGER L. HEITZMAN

3112 Harris Avenue, Independence, Missouri 64052^

ABSTRACT

A new species, Hypagyrtis brendae, is described from Arkansas, Ken-
tucky and Missouri. This new species varies from all other known Hypa-
gyrtis species in three characteristics: the coloration, the angular relation
of the postmedial and medial lines to the inner margin and the postmedial
line formation. The male and female types and the male genitalia are
figured. The male and female types have been deposited in the United
States National Museum.

INTRODUCTION

After several years of extensive investigation of the Hypa-
gyrtis, it was revealed that there was an undescribed species
occuring in scattered colonies throughout the Midwestern United
States. This is supported by three outstanding characteristics:
the coloration, the angular relation of the postmedial and medial
lines to the inner margin and the postmedial line formation. The
genitalia of this new species shows no satisfactory differences
from the other members of the genus. The new species exists
sympatrically with H. subatomaria (Wood) 1839. It is a multi-
voltine species with two broods, the second brood being the
least common. Its habitat is an acid soil, forest environment in
Missouri and Arkansas. It has only been captured in North-
western Arkansas, North Central Kentucky and Eastern Missouri.

Hypagyrtis brendae R. L. Heitzman, new species
MALE: Head: Vertex and front heavily scaled with gray; eyes
black; palpi short and gray, extending beyond front about one
third diameter of eyes; antennal stalk gray with sprinkling of
black scales, pectinations brown.

Thorax: Above, gray, patagia gray extending beyond base
of hindwings; below, gray with fine hairs, legs, gray with scat-
tered black scales.

Abdomen: Above, gray with scattered black scales; below,
uniformly gray.

^Research Associate, Florida State Collection of Arthropods, Division of
Plant Industry, Florida Department of Agriculture and Consumer Services,
Gainesville, Florida.

43

44

ROGER L. HEITZMAN

/. Res. Lepid.

3

Fig. 1. — Hypagyrtis hrendae, new species, holotype male, Washington State
Park, Washington Co., Missouri, 5 June 1973 (J. R. Heitzman) X 1.75

Fig. 2.~~Hypagyrtis hrendae, new species, allotype female, Washington
State Park, Washington Co., Missouri, 7 June 1973 (R. L. Heitzman) X 1.75

Fig. 3.- — Diagrammatic drawing of right forewing of typical Hypagyrtis spp.

Fig. 4.— Diagrammatic drawing of right forewing of Hypagyrtis hrendae.

13(l):43-48, 1974

NEW HYPAGYRTIS

45

Wings: Dorsal surface: Forewings, ground color gray with
scattered black scales except where invaded by small and re-
stricted areas of brown, brown in varying amounts occurs be-
tween the subterminal and postmedial lines, along costal margin
and basal area; fringe, alternating patches of dark and light
scales on outer margin, long hairs on inner margin; subterminal
line, white, rarely complete and with white subterminal spot in
cell Rg; postmedial line, black, complete, convex to subterminal
spot and extending nearly straight beyond to inner margin where
it enters at a basally inclined angle; medial line, black, diffuse,
usually incomplete and entering inner margin obliquely; distal
spot, black; antemedial line, black, usually complete. Hindwings,
concolorous with forewings but paler at costal and basal areas
with brown present only between postmedial and subterminal
lines; fringe, as fore wings; subterminal line white, rarely com-
plete; postmedial line, black, fading at costal margin; distal spot,
black; medial line, black, broad and diffuse, fading at costal
margin; gray hairs present in basal area. Ventral surface: Fore-
wings, pale gray scattered with black scales; postmedial line,
black, always present; distal spot, black; weak diffuse line some-
times present inside distal spot, outwardly inclined in contrast
to postmedial line. Hindwings, concolorous with forewings;
postmedial line, black, complete, no fading at costal margin;
distal spot, black; medial line, black, sometimes incomplete,
again no fading at costal margin.

FEMALE: Same as male; except larger, often lighter and outer
margins more scalloped.

ETYMOLOGY: I take pleasure in naming this species for my
sister Brenda Heitzman.

FORE WING LENGTH OF TYPE SERIES: Spring brood: Aver-
age for 275 males 16.40 mm, range 14-17 mm; for 30 females
19.86 mm, range 18-21 mm. Summer broods: Average for 25
males 12.75 mm, range 12-14 mm; for 3 females 16.50 mm, range
15-17 mm.

TYPES: Holotype, male, Washington State Park, Washington
Co., Missouri, 5 June 1973 (J. R. Heitzman); allotype, female,
Washington State Park, Washington Co., Missouri, 7 June 1973
(R. L. Heitzman); 299 male and 32 female paratypes: from the
type locality: five males and two females, 6 June 1972 (J. R.
Heitzman); 70 males and five females, 5 June 1973 (J. R. Heitz-

\\

46

ROGER L. HEITZMAN

/. Res. Lepid.

Fig. 5. — Male genitalia, Hypagyrtis hrendae, new species, holotype, ventral
view. X 31

13(l);43-48, 1974

NEW HYPAGYRTIS

47

man); 40 males and seven females, 5 June 1973 (R. S. Funk);
13 males, 7 June 1973 (J. R. Heitzman); 123 males and two fe-
males, 7 June 1973 (R. L. Heitzman); nine males and one female,
18 August 1973 (R. L. Heitzman); five males, 18 August 1973
(J. R. Heitzman); five males, 22 August 1973 (J. R. Heitzman);
five males and one female, 22 August 1973 (R. S. Funk); from
Dr. E. A. Rabler State Park, St. Louis Co., Missouri: seven males,
6 June 1973 (J. R. Heitzman); from Blue Springs State Park,
Washington Co., Arkansas: one male and one female, 29 May
1966 (R. L. Heitzman); two females, 29 May 1966 (J. R. Heitz-
man); one female, 27 May 1967 (R. L. Heitzman); one female,
27 May 1967 (J. R. Heitzman); two females, 31 May 1971 (R. L.
Heitzman); one male and two females, 4 June 1971 (J. R. Heitz-
man); one female, 4 June 1971 (R. L. Heitzman); two females, 6
June 1971 (J. R. Heitzman); one male and one female, 21 August

1971 (J. R. Heitzman); three males and one female, 27 May

1972 (R. L. Heitzman); seven males, 27 May 1972 (J. R. Heitz-
man); from Leslie Farm, Nelson Co., Kentucky: two males, 23
June 1971 (G. Florence); from Bardstown, Nelson Co., Ken-
tucky: one male, 29 April 1970 (G. Florence); from Horner Bird
Sanctuary, Oldham Go., Kentucky: one male, 22 June 1966 (C. V.
Govell, Jr.).

TYPE LOGALITY: Washington State Park, Washington Co.,
Missouri, in forested area.

LOCATION OF TYPES: The holotype and allotype will be
deposited in the type collection of the United States National
Museum, Washington, D.C. Paratypes will go to the following
institutions and individuals : The Florida State Collection of
Arthropods, Gainesville, Florida; American Museum of Natural
History, New York, New York; The Entomology Museum of the
University of Missouri, Columbia, Missouri; Central Missouri
State Univesity, Warrensburg, Missouri; Dr. A. E. Brower,
Augusta, Maine; Dr. C. V. Coveil, Jr., University of Louisville,
Louisville, Kentucky; Richard S. Funk, Illinois State University,
Normal, Illinois; Dr. W. C. McGuffin, Biosystematics Research
Institute, Ottawa, Ontario; Laurence R. Rupert, Sardinia, New
York; and the Heitzman collections.

48

ROGER L. HEITZMAN

J. Res. Lepid.

DISCUSSION

The coloration of H. brendae is unique among the Hypagyrtis,
as no other species possesses such a color scheme and pattern.
The angular relation of the postmedial and medial lines of the
forewing to the inner margin of H. brendae is strongly oblique
(fig. 4); whereas, with all the other Hypagyrtis these lines are
almost perpendicular to the inner margin ( fig. 3 ) . The forewing
postmedial line formation of H. brendae is nearly straight beyond
the st. spot (fig. 4), while in the other species it is deeply con-
cave and jutting to the inner margin (fig. 3). Finally, the sub-
terminal line of H. brendae occurs rarely among the rest of the
genus.

ACKNOWLEDGMENTS

I am grateful for the assistance of Dr. A. E. Brower, Augusta,
Maine; Dr. C. V. Covell, Jr., University of Louisville, Louisville,
Kentucky; Dr. Douglas C. Ferguson, United States National
Museum, and Dr. Frederick H. Rindge, American Museum of
Natural History for determinations and comments on the Hypa-
gyrtis. Also, I wish to thank Richard S. Funk, Illinois State Uni-
versity, Normal, Illinois, for his field assistance.

LITERATURE CITED

WOOD, 1839. Index Entomologicus, p. 241, fig. 1673.

Journal of Research on the Lepidoptera

13(l):49-56, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

GENETIC CONTROL OF

MACULATION AND HINDWING COLOR
IN APANTESIS PHALERATA (ARCTIIDAE)'

JACK S. BACHELER and THOMAS C. EMMEL

Departments of Entomology and Zoology, University of Florida,
Gainesville 32601

The tiger moth Apantesis phalerata (Harris) is a common
representative of the family Arctiidae around Gainesville,
Florida. During the course of a recent investigation (Bachelor,
1972) of its biological and systematic relationship with a
sibling species, A. radians Walker, rearing studies revealed
information on the inheritance of certain pattern elements in
the adult stage. The normal adult male of this species has yellow
hindwings and lateral abdominal stripes, and a criss-cross
pattern of cream bands across black fore wings. The normal
female has red hindwings and abdominal stripes, and only one
cream band (with a parallel, short cream bar) on black fore-
wings (Fig. 1). This paper reports data on the genetic control
of the forewing and body maculation and on the sex-limited
expression of the gene controlling hindwing coloration in
Apantesis phalerata.

METHODS

As part of life history studies on A. phalerata, field-collected
females were brought into the laboratory for oviposition. Re-
sulting larval broods were reared through to adulthood on an
artificial diet, modified slightly (Table I) from that used by
Shorey and Hale (1965).

Among the typical offspring of a phenotypically normal
female (red hindwings, full maculation) collected in March 1969
were three unusually light males ( almost devoid of maculation )
and several females with yellow hindwings. The exact pheno-

Tla. Agricultural Experiment Station Journal Series No. 4559.

49

50

BACHELER AND EMMEL

/. Res. Lepid.

typic ratio in these Fi adults was not noted since these moths
were among the first reared in connection with other studies.
However, each of the light males was mated to separate sibling
females having the yellow hindwings. The larvae resulting
from the one successful mating were reared through to adult-
hood. After several additional unsuccessful F2 crosses, the
F2 progeny were then spread, labeled, and stored for later
analysis.

RESULTS

Thirty-nine males and 35 females resulted from the successful
mating of one of the light Fi males to a yellow-hindwinged
female sibling. Female F2 adults were of two patterns. The
first pattern was that of a typical phalerata female, except the
normally red secondaries and abdominal area were light yellow.
All 29 females displaying this pattern were uniform in expres-
sion. In the second pattern the red areas were again replaced
by yellow, but there was also a loss of black maculation, re-
sulting in females that were nearly all yellow. The typical black
marginal and submarginal spots of the secondaries were absent
and the black costal margin faded slightly proximally. One
female of the six showing this loss of maculation had traces of
submarginal spots. Only traces of postcostal maculation re-
mained on the primaries. The patagium was entirely yellow,
one female having a trace of the typical black spot. The
tegulae were likewise yellow, devoid of the usual black bands.
The antennae and underside of the abdomen had a few yellow
scales. Both are normally black. The black dorsal abdominal
stripe was considerably reduced. The two F2 female patterns
are shown in Fig. la-b.

About half of the 39 males showed a pattern gradation from
almost typical phalerata males to those almost completely de-
void of maculation. The other half were normal. In the transition
through the series, the yellow costal stripe and postmedian
transverse, subterminal W-shaped, and submedian longitudinal
bands widened and fused. The black spots of the patagium,
tegulae, and dorsal abdominal stripe also were reduced through
the series. Four males in this transition are shown in Fig. Id-g.

A discontinuity was noted among males which exhibited a
gradation between absent maculation and normal marking. The
lighter forms of one class retained most of the dark, posterior

13(l):49-56, 1974

COLOR IN APANTESIS

51

forewing band, some of the marginal spot enclosed by the W
band and the wing margin, and a distinct dorsal abdominal
stripe as in normal males. The second class of nine males, con-
taining all light forms, lacked the forewing band and marginal
spot, and the dorsal abdominal stripe was reduced. The abdom-
inal stripe, then, easily separated all the males into two distinct
classes: normal (with variable expression of wing maculation
in some of those males) and light. The ventral wing surfaces
also easily separated the two classes. The light class lacks the
one black spot near the forewing margin, and the proximal black
bar along the leading edge of the forewing is broken, not solid
as in the normal class (see Fig. 2). The light Fi male success-
fully mated in the laboratory belonged to the first class, with a
distinct dorsal abdominal stripe and solid forewing bar.

Kimball (1965) reported an aberrant male almost devoid of
maculation from Gainesville, Florida, in 1959. This specimen
appears identical to several in the Fg experimental series of males.
Another light male was found in July 1968 in Gainesville. It
also fits well into the series, but at a slightly different point.
These two wild specimens are shown in Fig. Ih-i. A third light
male was found in Bradenton, Florida, in 1970. These aber-
rations were the only noticeably light phalerata found among
more than 2,000 males collected during this study.

When this rare aberration of phalerata is collected in the
future, it is hoped that an awareness of the simple mutant
character of this strain will avoid taxonomic confusion and the
unnecessary naming of “forms” or “species.”

GENETIG ANALYSIS

The rearing of this aberrant series shows that this light form
is probably inherited in simple Mendelian fashion, though the
74 progeny of the successful cross showed a considerable range
of variation in maculation expressed by the controlling geno-
types. One autosomal gene appears to control maculation. A
simple dominant allele at this locus causes maculation and its
recessive allele in the homozygous state accounts for the rare,
light forms. In males, expression of the heterozygous gentotype
is variable.

52

BACHELER AND EMMEL

/. Res. Lepid.

The genotype of the original field-collected (dark, red hind-
wings) parental female was, according to this genetic hypo-
thesis, heterozygous: DdRr. If it were homozygous for dark
maculation, no light offspring would have resulted, and if it
were homozygous recessive, its own phenotype would have
been light. Likewise, if it were homozygous for red hindwings,
none of the offspring would have had yellow hindwings, and
if it were homozygous recessive it would have had yellow hind-
wings.

The light aberrant male reported by Kimball and examined
by us was probably a recessive homozygote for maculation while
the light males found in Gainesville and Bradenton were hetero-
zygotes.

This genetic explanation on the inheritance of hindwing
coloration and maculation does have the drawback of necessi-
tating the capture of an apparently very rare female in nature,
and further, having this female mate with a rare male. However,
our two-gene explanation was the simplest and the only one
which fit the classes of moths so precisely. A test cross would
have made the hypothesis more convincing, but was not possible
at the time. Since we are not planning future work on this
problem, these partial results are published in the hope of stimu-
lating further genetic investigations on these interesting arctiids.

Hindwing color seems to be controlled by a second gene
locus. The dominant allele, responsible for red hindwings, is
expressed only in the female. Male hindwings are yellow,
regardless of the presence of this dominant gene. The homozy-
gous recessive state of this gene in the female results in yellow
hindwings.

In the following hypothetical analysis, D represents the
dominant gene for expression of maculation and cl its recessive
allele. R designates the gene for red hindwings and r its re-
cessive allele for yellow.

The male and female successfully crossed in the laboratory
can each have only one genotype if one follows this hypothesis.
The parental genotypes, their observed and expected offspring,
and X“ values are shown below. The observed and expected
numbers for both male and female moths of the laboratory
cross do not differ significantly from a 3:1 ratio.

13(l):49-^56, 1974

COLOR IN APANTESIS

53

Parents : d* X

Ddrr

( Dark partly
expressed)

Offspring :

Fi <?

Ddrr

(Dark; yellow
hindwings)

Genotype

Class

Expected ratio

No. Observed

No. Expected

DDrr or Ddrr

Dark

3

29

2 6.25

ddrr

Light

1

6

8.75

Totals

3^ 35

= 0.770

F2 -fcf

Genotype

Class

Expected ratio

No. Observed

No. Expected

DDrr or Ddrr

Dark

3

30

29.25

ddrr

Light

1

9

9.75

Totals

39

39

x2- 0.009

ACKNOWLEDGMENTS

We thank Dr. Dale H. Habeck, Department of Entomology
and Nematology, University of Florida, who provided facilities,
advice, and commented on the manuscript. This research was
supported in part by NSF Grants GB8442 and GB32151 (to
Thomas C. Emmel) and USDA Cooperative Agreement 12-14-
100-9397 (33) (D. H. Habeck, Principal Investigator).

54

BACHELER AND EMMEL

/. Res. Lepid.

4)^ O O

p

o-^
bO o

§ .^s
S §5 -b

s

— .4-r 43 ^

9=«bE'S

slsgjia

i)? S-2^ S

__33 S-3

cd

'5 “'■'

2 ^ '-w p P

^ 5 o .=5 o
S ^ 0 w
P'T5>-

^•^§.25 o

^ 43 S P -■ P

s J s

o

S P a o ^
55 ^1-b X 43

0 jb'
^ 0^
i g 3"2 "g -2^

a2 -

0^ ^ ^
'^'3
S ^

p -p.

CS
ot" 2

I

w w o
^ ^ P p^

■ ■ o o

0 e^

S

0 bO bO SS

§’ g §)2

13(l):49-56, 1974

COLOR IN APANTESIS

55

Fig. 2.— Dorsal and ventral surfaces of (a) normal homozygous males; (b)
heterozygous males, and (c) light homozygous males of Apantesis phaler-
ata. See text for discussion of maculation differences between the classes.

56

BACHELER AND EMMEL

J. Res. Lepid.

LITERATURE CITED

BACHELER, J. S. 1972. Biology and hybridization of Apantesis phalerata
(Harris) and A. radians Walker in Florida ( Lepidoptera: Arctiidae).
Department of Entomology and Nematology, University of Florida.
Ph.D. Thesis. 89 p.

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Division of Plant In-
dustry, Florida Department of Agriculture, Gainesville. 363 p.

ROBINSON, R. 1971. Lepidoptera Genetics. Pergamon Press, Oxford. 687 p.

SHOREY, H. H. and R. L. HALE. 1965. Mass-rearing of the larvae of
nine noctuid species on a single medium. J. Econ. Entomol. 58:522-
524.

Table 1. Modified Shorey and Hale ( 1965) artificial diet for
rearing Apantes is phalerata .

Pinto beans ( soaked overnight)

640

gm

Brewer's yeast=i=

100

gm

Ascorbic acid''=

10

gm

Sorbic acid=''=

3

gm

Methyl p-hydroxybenzoate-’'-‘=

6

gm

Formaldehyde ( 37%)

6

ml

Agar-''

40

gm

Water; with agar

800

ml

with dry ingredients

1000

ml

='=Nutr itional Biochemicals Corp., Cleveland, Oh.
=:=-Fischer Scientific Co., Atlanta, Ga .

Journal of Research on the Lepidoptera

13(l):57-62, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

NATURAL AND LABORATORY OCCURRENCE
OF “ELYMI” PHENOTYPES IN
CYNTHIA CARDUI ( NYMPHALIDAE )

ARTHUR M. SHAPIRO

Department of Zoology,

University of California,

Davis, California

The ' elymi” series of aberrant phenotypes, characterized
by suppression of the discal wing pattern, fusion of the dark
subapical pattern elements, and development of a series of white
submarginal spots, occurs in at least three species of the Holarctic
genus Cynthia: C. annabella Field (= C. carye auct. ), C. virgin-
iensis Drury, and C. cardui Linnaeus. The seasonal distribution
and frequency of these phenotypes in wild populations of C.
annabella in central California were recently reviewed by
Shapiro (1973).

Cynthia cardui in California is a migratory species which rare-
ly, if ever overwinters north of the Transverse Ranges. Transient
populations occur in most of the state most years, generally
shifting northward and upslope in mid-summer and sometimes
showing a definite return flight southward in autumn. Although
specimens representing various stages in the ‘'elymi” series have
been taken in many localities in California, no systematic search
for them has been linked to seasonal movements and abundance
of the species. The exceptionally large 1973 flight of C. cardui
proved an opportunity for such an investigation.

In 1972, an “average” year, C. cardui was present in the
Sacramento Valley from the third week of March to the first week
in October, with the largest numbers flying in April, May, and
the first half of June. No “elymi” phenotypes were reported any-
where in central California in 1972. In 1973 C. cardui first ap-
peared the fourth week of February and reached “outbreak” pro-
portions by mid-April. Very large populations persisted in the
Valley until the third week of June, when most of the insects
emigrated northward and upslope into the Sierra Nevada. During
this period three generations appeared, feeding primarily on

57

58

ARTHUR M. SHAPIRO

]. Res. Lepid.

Fig. 1. • — ■ Aberrant Cynthia cardui collected in central California in 1973,
upper and lower surfaces. For localities see text. A, v.25; B, vi.8; C, vi.2;
D, ix.7.

13(l);57-62, 1974

‘ELYMF’ PHENOTYPES

59

Silijbum marianum (L.) Gaertn. (Compositae), Amsinckia spp.
(Boraginaceae), and Malva, spp. (Malvaceae). C. cardui re-
mained in the Valley at low density through the hottest part of
summer and occasional larvae could be found on Centaurea
solstitialis L. (Compositae). From late August through September
a significant southward movement through the Valley was ob-
served. In the Sierra Nevada C. cardui was abundant in July and
August and a major southward migration occurred in September
at the same time as that in the Valley. Larvae were taken from
Cirsium spp. and Wyethia mollis Grey (both Compositae). Dur-
ing the 1973 season five “elymi” phenotypes were reported in the
Valley and two in the Sierra Nevada. An estimated 30,000 C.
cardui were examined in the field during the season, giving a
frequency of .00023, much below the .001 frequency observed for
the corresponding phenotype of C. annahella (Shapiro, 1973).

Records of ''elymi” specimens follow:

Sacramento Valley: American River, Sacramento (City) Co.,
California, v. 25:73 ( $ ), vi. 2.73 ( $ and second seen, sex
undetermined); vi. 8.73 ( 5 ). Broderick, Yolo Co., California,
vi. 24.73 (seen, sex undetermined). (All A. M. Shapiro)
Sierra Nevada: East of Jerseydale, Mariposa Co., California, vi.
21.73 { $ ) (O. Shields); Donner Pass, el. 7000’: Placer Co.,
California, ix. 7.73 ( ^ ) (A. M. Shapiro)

All of the captures except the Jerseydale specimen are shown
in figure 1.

The most interesting aspect of this series is the cluster of
records from the American River. All four specimens (the one
missed on vi. 2 is definitely not the same individual taken vi. 8)
were taken within an area one-half mile square and were in
similar condition. Since C. cardui had been flying at the Ameri-
can River since March 2 they were probably locally bred, rather
than immigrant individuals. Moreover, no aberrant specimens
were seen at other Valley localities with equally dense cardui
populations, despite comparable search. It is therefore likely that
these four were siblings or at least that the "elymi” phenotype
had a common ( genetic or environmental ) origin in all of them.

Dimock ( 1968 ) reported the induction of "elymi” phenotypes
by holding fresh pupae of C. cardui at 36° F for 14 days. During
the 1973 season numbers of large larvae of C. cardui were col-
lected in the Valley and Sierra and the resulting pupae subjected
to this treatment. Considerable variation occurred among the 37
adults obtained, but only seven were wild-type. In the remaining

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. 2. — Aberrant C. cardui produced by chilling pupae at 36 °F for 14
days. Upper and lower surfaces.

13(l);57-62, 1974

'ELYMI” PHENOTYPES

61

Fig. 3. — Normal C. cardui from pupae held at 90°F for 14 days. Upper
and lower surfaces.

62

ARTHUR M. SHAPIRO

J. Res. Lepid.

30 the pattern was more or less modified in the direction of
“elymi” ( occasionally on only the fore- or hindwings ) ( figure 2 ) .
Four specimens were indistinguishable from the wild “elymi
phenotypes. An additional 28 pupae died. Many of these develop-
ed fully but failed to eclose; all which had developed a pattern
were ‘ elymi,” and a few were as extreme as DimocFs figure. In
the control group of 30 unchilled pupae, 24 adults were obtained,
all wild-type.

The dates and localities of the 1973 “elyrnF (except the ix. 7
specimen ) virtually exclude sustained ( or severe but intermittent )
chilling of the pupa as a causative agent. In an effort to deter-
mine if heating, a more likely factor afield, could induce “elymi”
21 pupae were subjected to 90 °F until they hatched or for 14
days, whichever happened first. Three pupae so treated produced
wild-type adults (Fig. 3) within 7 days; the remainder all died
without depositing adult pigment.

“Elymi” phenotypes are produced with considerable regular-
ity when fresh pupae of C. cardtii are chilled, whether they come
from low or high elevations. The individual batches tested are too
small for any statements to be made as to the variance within
local populations or among sibs with respect to this character, but
it appears that the potential to produce “elymi” under tempera-
ture shock is inherent in normal cardui. The most promising
hypothesis is that wild specimens of “elymi” are produced by
genes or gene combinations which alter the threshold for ex-
pression of “elymi” so that it is produced under ordinary devel-
opmental conditions. This would make the experimental animals
phenocopies, a familiar situation in genetics and one which
predisposes to the evolution of phenotypic switch mechanisms
through the mechanism called “genetic assimilation” (cf. Wad-
dington, 1957 ) . The occurrence of the phenotype in three species
of Cynthia suggests that it may indeed have been a seasonal
phenotype at one time in the evolution of the genus and that its
expression was subsequently suppressed in a manner akin to the
summer phenotype of Pieris virginiensis (Shapiro, 1971).

LITERATURE CITED

DIMOCK, T. 1968. An extreme experimental aberration of Vanessa cardui
(Nymphalidae). J. Lepid. Soc. 22:146.

SHAPIRO, A. M. 1971. Occurrence of a latent polyphenism in Pieris
virginiensis ( Lepidoptera: Pieridae). Ent. News 82: 13-16.

SHAPIRO, A. M. 1973. Recurrent aberration in Cynthia anahella: a review
with four new records. Pan-Pac. Ent., in press.

WADDINGTON, C. H. 1957. The Strategy of the Genes. Allen and Unwin,
London.

Journal of Research on the Lepidoptera

13(l):63-65, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

DAS NATURHISTORISCHE MUSEUM IN WIEN

UND SEINE LEPIDOPTERENSAMMLUNG

Von Dr. FRITZ KASY

Wien, Austria

Das Wiener Naturhistorische Museum gehort zu den al-
testen naturgeschichtlichen Sammlungen der Welt und zu den
grossten dieser Art. Seine Anfange reichen bis in das Jahr 1748
zuriick, in dem vom damaligen Herrscher von Osterreich, Kaiser
Franz Stephan, dem Gatten der bekannten Kaiserin Maria
Theresia, die grosse Sammlung des Florentiner Universalgelehr-
ten Johann von Baillou angekauft wurde. Durch Aufsammlungen
in den Landern des damals sehr ausgedehnten osterreichischen
Kaiserreiches und durch vom Herrscherhaus grossziigig finan-
zierte Reisen nach Ubersee wurden die in verschiedenen Teilen
der Hofburg (der Residenz der osterreichischen Kaiser) unter-
gebrachten Sammlungen in den folgenden Jahrzehnten stark
vergrossert und schliesslich auch in einer wissenschaftlichen
Ordnung aufgestellt. Den drei Reichen der Natur entsprechend
wurde eine Gliederung in drei “Kaiser-konigliche Hof-Naturalien-
kabinett” vorgenommen. Diese entwickelten sich bald zu
Zentren der naturwissenschaftlichen Forschung in Osterreich und
wurden trotz der immer druckender werdenden Raumnot durch
weitere Aufsammlungen bereichert, bespielsweise durch einen
langen Sammelaufenthalt (1817-1835) in Brasilien. Das Revolu-
tionsjahr 1848 brachte einen schweren Verlust: im Verlauf der
Kampfe wurden grosse Teile der Zoologischen Sammlung durch
einen Brand vernichtet, darunter auch die Lepidopteren-
sammlung von Schiffermiller und Denis. Im Jahre 1889 konnte
endlich der Raumnot durch die Eroffnung des neuen Natur-
historischen Museums auf der Ringstrasse gegeniiber der Hof-
burg ein Ende bereitet werden, allerdings auch nur fur einige
Jahrzehnte, denn heute ist auch dieses grosse prachtige Gebaude
bereits wieder zu klein geworden.

63

64

FRITZ KASY

). Res. Lepid.

Die Lepidopterensammlung des Wiener Museums ist auch
heute noch die grosste auf dem europaischen Festland. Sie
umfasst ca. 2 Millionen Exemplare und zahlreiche, insbesondere
auch alte, Typen.

Das Studium der Lepidopterologie hat in Osterreich seit dem
Ende des 18. Jahrhunderts eine rege Pflege erfahren. Im Jahre
1776 erschien das “Systematische Verzeichnis der Schmetterlinge
der Wiener Gegend”, herausgegeben von den Lehrern Ignaz
Schiffermiller und Michael Denis. In diesem Buche wird zum
erstenmal auf Grund der Kenntnis der Jugendstadien die von
Linne geschaffene Anordnung der Lepidopteren abgeandert und
so ein mehr naturliches System geschaffen. Das umfangreichste
Werk liber europaische Schmetterlinge in deutscher Sprache
(17 Bande, 1807-1835)5 namlich das von Ferdinand Ochsen-
heimer und Friedrich Treitschke, entstand vom 3. Band an eben-
falls in Wien, da beide Autoren am Wiener Burgtheater be-
schMtigt waren, ersterer als Schauspieler. Als Nachfolger
Treitschkes, besonders auf dem Gebiet der Mikrolepidopteren,
ist Josef Fischer von Roslerstamm (1787-1866) hervorzuheben,
dessen Sammlung spater an Dr. Herrich-Sehdffer nach Begens-
burg verkauft wurde, der dadurch erst die Moglichkeit erhielt,
sein Werk iiber europaische Schmetterlinge zum Abschluss zu
bringen.

Der erste Lepidopterologe, der in den kaiserlichen NatuTalien-
sammlungen (wie eingangs ervvahnt, den Vorlaufern des Natur-
historischen Museums ) tatig war, namlich als Kustos und spater
auch als Direktor des '‘Zoologischen Hofkabinettes’', war Vincenz
Kollar (1797-1860). Er veroffentlichte unter anderen auch Ar-
beiten, die fremdlandische Faunengebiete betrafen, beispielsweise
solche iiber Brasilien und Venezuela. Von besonderer Bedeutung
fiir die Lepidopterensystematik waren die Arbeiten von Julius
Lederer (1821-1870), dessen an Typen reiche Sammlung an
Dr. Staudinger in Dresden (Deutschland) verkauft wurde. In
Lepidopterenkreisen weit iiber die Grenzen Osterreichs hinaus
bekannt wurde auch der Name des Biirgermeisters von Wien
Cajetan Freiherr von Felder (1814-1894). Neben Veroffent-
lichungen iiber verschiedene Tagfaltergruppen ist besonders die
Bearbeitung der Lepidopteren aus den Ausbeuten von der
osterreichischen Weltumseglung mit der Fregatte Novara (1857-
1859) hervorzuheben. Seine Sammlungen kamen an Sir Walther
Rothschild in dessen Privatmuseum nach Tring (England).

13(1):63~65, 1974

MUSEUM IN WIEN

65

Der erste Kustos der Lepidopterensammlung am neu erbauten
Naturhistorischen Museum war Alois Rogenhofer (1831-1897),
neben dem durch 45 Jahre am Museum Josef Mann (1804-1889)
als Praparator und insbesondere unermiidlicher Sammler wirkte;
er beschrieb auch zahlreiche neue Arten, vor allem Mikrolepidop-
teren. Der Nachfolger Rogenhofers am Wiener Museum, Prof.
Dr. Hans Rebel (1861-1940), war der bedeutendste Lepidop-
terologe Osterreichs iiberhaupt. Von ihn stammen neben zahl-
reichen Neubeschreibungen aus den verschiedensten Familien
der Gross- und Kleinschmetterlinge und anderen Arbeiten liber
Lepidopteren, der bekannte Staudinger-Rebel-KsLtalog der pala-
arktischen Lepidopteren und das im deutschen Sprachraum bei
Liebhaberentomologen auch heute noch am weitesten verbreitete
Schmetterlingswerk, namlich die Neubearbeitung von Berges
Schmetterlingsbuch, das zahlreiche Amateure zu einer wissen-
schaftlichen Betatigung mit Schmetterlingen anregte. Von
solchen Sammlern stammt auch der grosste Teil des nach 1920
ans Wiener Museum gekommenen Schmetterlingsmaterials. Ein
weiterer bekannter Lepidopterologe des Naturhistorischen Mu-
seums in Wien war Dr. Hans Zerny (1887-1945). Er untemahm
zahlreiche Sammelreisen, unter anderen auch eine nach Slid-
amerika. Sein Spezialgebiet waren die Syntomiden. Nach seinem
Tod war die Schmetterlingssammlung des Wiener Museums be-
dauerlicherweise durch 15 Jahre nicht mit einem Lepidopterolo-
gen besetzt. 1960, also im Jahre des 11. Internationalen Kon-
gresses fur Entomologie, der damals in Wien stattfand, wurde
sie dem Verfasser dieser Zeilen anvertraut. Leider muss fest-
gestellt werden, dass heute die Ausstattung des Naturhistorischen
Museums in Wien mit Personal und finanziellen Mitteln in
keinem angemerrenen Verhaltnis zu der GrOsse and der wissen-
schaftlichen Bedeutung seiner Sammlungen steht, was nur zum
Teil durch die geringe Grosse des heutigen osterreichischen
Staates zu entschuldigen ist.

Journal of Research on the Lepidoptera

13(1 ):66, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

NOTICE

The PRECEDiiXG PAPER by Dr, Fritz Kasy is the first of what
the editor hopes will be a series devoted to institutions which
have in the past and present spent much effort in furthering
the knowledge of the Lepidoptera, not only in the amassing
of collections but also in the constructive work in the field of
experimental research. It is by means of the latter mentioned
experimental work that the systematic work of the museums
and private collectors can best be understood and appreciated.

It is hoped that present workers in such institutions will
feel free to send historical papers to implement this series.

William Hovanitz

66

Journal of Research on the Lepidoptera

13(1):67~68, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

HABITAT; ADELA BELLA IN FLORIDA
( INCUR VARIIDAE: ADELINAE)'

JOHN B. HEPPNER

Department of Entomology and Nematology,

University of Florida
Gainesville, Florida 32611

Little is known of the biologies of adelid moths and the life
history of Adela bella Chambers, in particular, is unknown.
While beginning studies of the moth in March 1973 at Torreya
State Park in the panhandle of Florida, males were found to
congregate on the leaves of Carpinus caroliniana (Betulaceae)
trees deep in the deciduous forest on the slopes overlooking the
Apalachicola River (females were not found). Male congre-
gations by adelid moths, however, do not necessarily indicate
the oviposition host.

The illustration of the forest at Torreya State Park shows
Carpinus mixed in with other deciduous trees and generally
shows the shaded habitat in which A. bella may be encountered.
The Carpinus leaves from which males were taken is in the
upper foreground left and closest to the viewer near the top of
the picture. The moth is diurnal like other adelids and is widely
distributed in the eastern United States and Canada, occurring
mainly in the Appalachian Mountains in the South. There have
been other records of A. bella in Florida but the species is well
established in this Apalachicola uplift at Torreya State Park
which is ecologically similar to the southern Appalachians of
northern Georgia and Alabama.

Males fly in a bouncing but slow fashion on sunny days ( mid-
March to mid-April in Florida), their long antennae held up-
wards. Moths are encountered flying or resting on Carpinus and
other leaves and also near flowers.

^Florida Agricultural Experiment Station Journal Series No, 5308.

67

68

JOHN B. HEPPNER

J. Res. Lepid.

Fig. 1. — Deciduous forest habitat of Adela hella at Torreya State Park,
Liberty Co., Florida. (19 Mar 1973)

THE JOURNAL ©F RESEA R.CH
©NJ THE LEF1JD©PTERA\

CONTENTS

Volume 12 Number 1

March, 1973

A new species of Papilio

from the eastern United

States ( Papilionidae ) J. Richard Heitzman 1

Specific entities of the subgenus
Icaricia Nabokov (Lycaenidae)

J. W. Tilden 11

The correct name for the

subspecies of Limenitis weidemeyerii
occurring in Arizona ( Nymphalidae)

Cyril F. Dos Passos 21

A review of carrying pair

behavior and mating times in
butterflies

Oakley Shields and John F. Emmel 25

Volume 12 Number 2 June, 1973

The ecological associations of the butterflies

of Staten Island Arthur M. Shapiro and

Adrienne R. Shapiro 65

Volume 12 Number 3 September, 1973(1974)

On the origin of austral elements in

the moth fauna of south-eastern Ontario,
including a number of species new for Canada
R. Harmsen, P. D. N. Hebert and P. S. Ward 127

The study of populations of Lepidoptera
by Capture-Recapture methods

P. M. Sheppard and J. A. Bishop 135

Life history studies of Idaea obfusaria (Walker)

Roger L. Heitzman 145

Survey of ultraviolet reflectance

of Nearctic butterflies James A. Scott 151

Butterflies of St. Croix Charles F. Leek 161

An improved method for rearing the

Monarch butterfly Charles Munger 163

An annotated checklist of the Missouri

Geometridae Roger L. Heitzman 169

Cover Illustration:

Variation in Colias nastes of Lapland

William Hovanitz 180

Volume 12 Number 4 December, 1973 (1974)

Adult behavior and population biology of two
skippers mating in contrasting topographic
sites James A. Scott 181

Acceptance of artificial diet by

Slc^atJjijmiis streckcri (Skinner)

M. A. Pefterson and R. S. \\4elgus 197

Some observations on the eggs of moths and certain
aspects of first instar lar.Val behax ior

Noel McFarland 199

New food plant for Darapsa j)J}olus (Cramer)

]. C. E. Riotte 209

The nomenclature in an important British

check list (1972). Part I. Joraj Paclt 211

A name for Glaucopsi/chc h/a.(Iamus auct., not

Edwards 1862. ' J. Tilden 213

Junonia and Precis. A correction.

J. Tilden 216

The flight periods of se\’eral sibling species

of moths. Charles L. Selman 217

Lifespan of Butterflies James A. Scott 225

Altitudinal migration of butterflies in the central

Sierra Nevada Arthur M. Shapiro 231

Early stages and biology of Phyciocles orseis.

James A. Scott 236

Book Review: Annotated check list of the butterflies
of Illinois by R. R. Irwin and J. C. Downey

R. S. Funk

243

NOTICES

Books on Lepidoptera: more than 300 books for sale. Free
catalogue sent on request. Sciences Nat. , 45 Rue de Alouettes, 7 5019,

Paris, France

Wanted to make exchanges with collectors in California and Florida.
Have Maylaysian butterflies for exchange (Trogonoptera brookiana, etc, )
Masaki NAKAYAMA, Kitakyushu- Wakamatsu- Miyamaru -2-10-14,

Fukuoka pref, , Japan ,

Need aid in preparation of specimens for study of population
structure in butterflies; involves obtaining population samples, mounting,
laboratory breeding, etc. Full or part time. Contact William Hovanitz,
1160 W. Orange Grove Ave. Arcadia, Calif. 91006.

i

I

Volume 13 Number 1 March, 1974

IN THIS ISSUE

Studies on Nearctic Euchloe — Part 7

Paul A. Opler 1

3 Stacks of the eggs of Hemistola hatching

Noel McFarland 21

Checklist of the Macroheterocera of
south-eastern Ontario

P. S. Ward, R. Harmsen, and P. D. N. Hebert 23

A new species of Hypagyrtis

Roger L. Heitzman 43

Genetic control of maculation and

hindwing color in Apantesis phalerata

Jack S. Racheler and Thomas C. Emmel 49

Natural and laboratory occurrence of “Elymi”

phenotypes in Cynthia cardui ( Nymphalidae )

Arthur M. Shapiro 57

Das naturhistorische Museum in Wien
und seine Lepidopterensammlung

Dr. Fritz Easy

63

Notice

William Hovanitz

66

Habitat: Adela bella in Florida

John R. Heppner

67

THE JOURNAL

OF RESEARCH

ONJ THE LEFIOOPTERA

HAilsws

Volume 13

Number 2

published by

The Lepidoptera Research Foundation, Inc.
at

1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz

Associate Editors:

Thomas C. Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601

Maria Etcheverry, Centro de Estudios Entomologicos, Casilla 147, Santiago,
Chile.

T. N. Freeman, Div. of Entomology, Dept, of Agriculture, Ottawa, Ontario,
Canada.

Brian O. C. Gardner, 18 Chesterton Hall Crescent, Cambridge, England.

Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.

Lee D. Miller, The Allyn Museum of Entomology, 3701 Bay Shore Road,
Sarasota, Florida, 33580.

Bjorn Petersen, Ostanvag 52, Malmo, Sweden.

Manuscripts may be sent to the Editor or Associate Editors.

The JOURNAL is sent to all members of the FOUNDATION.

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STATEMENT OF OWNERSHIP AND MANAGEMENT
THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a
year, Spring (March), Summer (June), Autumn (September), and Winter (December)
by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
cation and the general business office are located at 1160 W. Orange Grove Ave.,
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Journal of Research on the Lepidoptera 13:73-82, 115-122, 137-148, 1974

1160 W. Orange Grove Awe., Arcadia, California, U.S.A.
© Copyright 1975

THE BUTTERFLY FAUNA OF THE
SACRAMENTO VALLEY, CALIFORNIA

ARTHUR M. SHAPIRO

Department of Zoology, University of California
Davis, Calif. 95616

INTRODUCTION

The State of California has an extremely rich and varied
butterfly fauna, reflecting its topographic, climatic, and botani-
cal diversity (Comstock, 1927; Munz, 1970; Bakker, 1971). The
montane faunas, which are perhaps best known, are character-
ized by a high proportion of endemic species and subspecies.
In recent years regional faunas have been published for Sierra
Nevada localities: Yosemite National Park (Garth and Tilden,
1963), Mather (Shields, 1966), and Donner Pass (Emmel and
Emmel, 1962 ) . A study by Opler and Langston ( 1968 ) included
both Outer and Inner Coast Ranges in Contra Costa County, as
well as part of the Sacramento-San Joaquin Delta. The least
well-known butterfly fauna in northern and central California
is that of the rather densely populated Sacramento Valley (fig.
1). This is scarcely surprising when the current biotic condi-
tion of the Valley is considered. The following discussion is
drawn primarily from Thompson (1961) and Sculley (1973),
who discussed the climatic, physiographic, and ecological con-
ditions of the pristine and present Valley.

VEGETATION

The vegetation of the Sacramento Valley has been more
thoroughly modified by man than that of scarcely any compar-
ably large area in North America. Prior to European- American
colonization, three natural communities were widespread in
the Valley: bunchgrass-Valley Oak savanna; tule-cattail marsh;
and riparian forest (fig. 2). Perennial bunchgrass, with scattered
groves of Valley Oak {Quercus lobata Nee.), occurred on the
higher sites not subject to regular flooding, and was the com-

73

74

ARTHUR M. SHAPIRO

}. Res. Lepid.

Fig. 1.— Location map of the central Sacramento Valley.

SACRAMENTO VALLEY

75

jSr

13:73-82, 115-122, 137-148, 1974

/HOnry CHAPARKAL.

REO lands ok

©L.D ALt.ev‘UM -

OAK ^ GRASS

alluvial PlAINS-
FfiRNEKLV SAVANNA

r/par»an

FaRE-ST f
‘ j^ATURAL UVCfiS

3^S/M

FLeoo iASiNS-
FORAlERLy -RJlF
MAKshe^

Fig. 2.— Physiographic subdivisions of the central Sacramento Valley ^ with
present biotic communities.

76

ARTHUR M. SHAPIRO

/. Res. Lepid.

monest vegetation type in the Valley, especially on the west
side. The lowlands, which were inundated in normal winters
(and in wet years well into summer), supported vast areas of
cattails (Typha) and common tule (Scirpus acutus MuhL).
Torrential rains on the Sierran west slope fed the periodic over-
flows of the Sacramento River and its tributaries. The major
streams built up “natural levees” of silt deposited during flood
stages, and these supported a lush deciduous forest dominated
by Fremont Cottonwood {Populus fremontii Wats.). Decidu-
ousness in a Mediterranean climate like that of California is a
luxury which can be afforded only when a reliable year-round
supply of ground water is available. The riparian forests of
the Sacramento Valley were unique at low elevation in the state.

Of the first two communities little or nothing remains. Most
of the bunchgrass prairie was put into pasture or under the plow;
either way, the native bunchgrasses were competed out of exist-
ence by introduced annual grasses, mostly from Europe. With
the bunchgrasses most of the native flora, both annual and per-
ennial, also succumbed, to be replaced by weedy Crucifers,
Borages, Mallows, Composites, and other aliens. The marshes
were diked or drained and reclaimed for agriculture, or else
grossly modified as overflow channels for flood control (the
Yolo, Colusa, and Sutter Bypasses). Their original character
has in most places been lost. Relict marshlands still exist in
West Sacramento and south of Sacramento, at Beach Lake and
Stone Lake. Substantial fragments of riparian forest remain in
public and private hands — more or less modified by the deletion
of native species and the addition of weedy ones, and by re-
striction in most places to the immediate riverbank where they
once had reached 1-4 miles inland.

Little of the Sacramento Valley is free of major ecological
disturbance for any significant length of time, and except for
the creek bottoms the successional potential of the area— with its
radically altered flora — is not really predictable. Most of the
former bunchgrass-oak savanna is either intensively farmed or
urbanized. The lowland basins are farmed, or farmed in
summer and flooded in winter. The riparian forests are under
increasing recreational-use pressure, especially from off-road
vehicles.

To a casual visitor, the Valley presents a monotonous vege-
tation due to the ubiquity of early-successional, adventive spe-
cies. These broadly adapted plants tend to obscure soil and
water-table differences which do, however, become apparent in

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

77

more mature stands. The lack of native plants and major topo-
graphic features, and the constant presence of man tend to
discourage butterflies (and Lepidopterists as well). The sandy
American River lowland in Sacramento County is mostly not in
agricultural use and has the richest butterfly fauna in the Valley,
as well as the most mesic vegetation. Thirty-six species—58%
of the total Valley fauna — were recorded flying there on June
2, 1973, and the total number of species ever recorded there is
53, or 85% of the Valley fauna. Nonetheless, most of the species
are highly vagile and occur in foothill canyons on both sides of
the Valley; there is little to necessitate postulating a relict
( pre- American ) origin for any of the present butterfly popu-
lations in the Valley (except perhaps Phyciodes campestris) .

CLIMATE AND BUTTERFLY PHENOLOGY

Sixty-two species of butterflies have been recorded in the
Valley. This is a fairly small fauna by California standards;
134 species are recorded in Yosemite, 84 in Contra Costa County,
74 at Mather (Tuolumne County), about 80 at Donner Pass
(Placer County), 70 at Boreal Ridge and 63 at Marin-Sierra
Camp, both Nevada County (Shapiro, unpubl. ), and about 65
in the east-slope canyons of the Vaca Hills immediately west
of the valley (data in part from Shields, pers. comm.). When
species which fly into or through the Valley but are not known
to breed there are excluded, the fauna drops to 53.

This fauna contrasts strikingly with others in California in
the distribution of voltinism. In climates where rainfall is rela-
tively evenly distributed through the year, the proportion of
univoltine species increases steadily with elevation and with
the shortening of the growing season. In California the summer
drought is reflected in the widespread evolution of vernal uni-
voltinism in butterflies at low elevations. The low proportion
of univoltines in the Sacramento Valley — only 13% of the resident
species ( as compared with 39% of the species in the nearby Vaca
Hills, Table I) — reflects the reliable supply of summer water
associated with the riparian systems and agricultural irrigation.
It also reflects the origins of the fauna, which is largely recruited
from the riparian lands where the summer drought has always
been less severe than elsewhere.

Relatively few California butterflies are facultatively univol-
tine; species with only a single brood anywhere generally have
only one everywhere, despite considerable altitudinal ranges.

78

ARTHUR M. SHAPIRO

/. Res. Lepid.

Species which are flexible in brood sequence are primarily low-
land colonizers which periodically invade higher elevations but
cannot be considered permanent residents there: Vanessa spp.,
Precis coenia, Strymon melinus, Lycaena helloides, Plebeius
acmon^ Pieris rapae, P. protodice, Colias eury theme, Pyrgus
communis, Hylephila phylaeus. These species are adapted to
temporarily unstable habitats, and whatever their geographic
origins their seasonal cycles have probably always included
up- and downslope colonization. They are the most conspicuous
element of the Valley fauna, where they disperse from one
disturbed habitat to another in response to agricultural prac-
tices. In good years Valley populations probably serve as a
source for colonizers which reach the high Sierra.

The climatography of the Sacramento Valley is unusually
well documented for the Far West. Reliable records at Sacra-
mento extend back to 1849-50, and recent compilation of means
and extremes has been prepared (Figgins, 1971). The Valley
has a Mediterranean climate in which rainfall occurs from Sep-
tember to April and is usually concentrated in December, Janu-
ary, and February (see Tables 2 and 3). The Coast Ranges
insulate much of the Valley from direct maritime influence and
the resulting continentality is shown in the high summer maxima
and occasionally low winter minima. The gap in the Coast
Ranges at the Carquinez Straits allows a shallow penetration
of maritime air in summer which terminates periods of extreme
heat after two to four days. Skies are clear in summer, but
widespread and persistent low cloudiness and fog prevail in
winter and are especially heavy in and near the river bottoms.
The highest summer temperatures and the clearest, driest winter
weather are provided by “northers,” strong northerly winds which
develop in response to a north-south pressure gradient and are
warmed and dried by their descent from the Siskiyous.

WINTER AND DIAPAUSE STRATEGIES

Sacramento Valley winters are mild and essentially snowless
except in the extreme north. Freezes occur up to 30 times each
winter, but temperatures below 25° F and continuous freezes of
longer than ten hours are exceedingly rare. Under these con-
ditions non-diapausing butterfly immatures may continue their
development through the winter. Larvae of Pieris rapae from
eggs laid in late November develop slowly to pupation in late
January at Davis and Woodland. The resulting pupae enter

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

79

diapause and eclose in late March or April. Diapausing pupae
formed in December or earlier eclose in February and March.
A similar winter history is reported for P. rapae in England
(Gardiner, 1972).

Diapause is generally considered an adaptation to cold
winters, and indeed is associated with biochemical defenses
against intracellular and extracellular freezing. But in both
cold- and mild-winter areas it serves as a timing mechanism
correlating spring emergence with the onset of weather suitable
for adult activities and— most importantly— likely to be sus-
tained. The limiting factor on butterfly breeding in winter in
the Sacramento Valley is the unsuitability of overcast, humid
weather for flight activity, regardless of temperature. Persistent
fog and low overcast initiate winter in the Valley while tem-
peratures are still relatively high. Winter emergences of Pierid
butterflies during fair, mild periods are less common than in
the northeastern United States. Spring emergences coincide with
decreasing cloudiness and humidity in February and March, and
a temperature regime comparable to that in the emergence
seasons of the same species in cold-winter areas. Availability
of suitable flight weather, rather than nectar sources, seems to
be the predominant factor here; major blooms of nectar-rich
plants, some native, begin in January in dry years and may
continue all winter in wet ones.

The flight season begins several days to two weeks later in
the Valley than in the canyons of the Vacas (up to 500 feet).
These canyons are sheltered from the spring “northers” which
render many otherwise suitable days in the Valley unfit for
flight. They also have less fog and dew (which cools the air
near the ground and may prevent flight activity most or all of
the day if the ambient humidity stays high). The topography
of the Valley floor itself also has a bearing on flight times. The
American River bottomlands become damp and foggy earlier
in fall than the higher ground, and flight activity there ends
earlier as a result. During the winter temperatures are moder-
ated by the rapidity with which the air is saturated, and wind
velocities (and the desiccating effects of “northers”) are braked
by the forest cover. Most species emerge four to twelve days
earlier in spring at the American River than elsewhere. The
local climate there allows Pieris protodice to overwinter re-
liably, something it does only very sporadically in the Valley.

One ecological consequence of the dominance of adventive
weeds is the absence of all but a handful of butterfly species

80

ARTHUR M. SHAPIRO

/. Res. Lepid.

from very extensive tracts of agricultural land. Only one of
these is itself adventive from Europe, Pieris rapae. Under Valley
conditions rapae and its primary hosts, Brassica spp., are not
well-coordinated seasonally. Weedy mustards behave as winter
annuals, seldom sustaining much frost damage, and are past
their prime when the spring brood of P. rapae is peaking. The
phenology of alfalfa (Medicago sativa L. ), the major host of
the Orange Sulphur, Colias eury theme, is similar to that of the
native perennial legumes on which this butterfly originally fed.
C. eury theme overwinters as a third- or fourth-instar dormant
( diapausing? ) larva.

Hibernating Nymphalid adults generally do not fly on the
10-20 potentially suitable days in winter; they appear in Febru-
ary just before the non-hibernating butterflies. The energetics
of Nymphalid hibernation in the mild Sacramento Valley winter
is of considerable interest, especially in the Mourning Cloak
{Nymphalis antiopa) which disappears in early August and
remains dormant for six months. Its failure to rear a second
brood even along the major rivers is rather perplexing. Vanessa
(= Cynthia) annabella departs from the usual pattern of its
relatives by overwintering largely as pupae formed in Decem-
ber and early January. Adults alive in December also hibernate
successfully, and fresh adults may eclose during warm spells
in winter and enter hibernation. This species has been recorded
flying 50 weeks of the year at Davis and is the closest thing to
a year-round breeder in the Valley. In 1973, when it was abun-
dant, V. cardui also bred well into December and continued
to eclose.

RAINFALL AND INTERSEASONAL VARIANCE

One of the most striking aspects of Valley climatology is the
very high interseasonal variance in rainfall. Temperature char-
acteristics are less variable. Some idea of the variance in pre-
cipitation may be obtained from figure 3 and Table 3. The
uncertainty of rainfall would be expected to produce adaptations
for facultative diapause in species whose host plants are rainfall-
dependent. The Papilionid Battus philenor produces some dia-
pause pupae in all broods, regardless of photoperiod; these eclose
the following spring (rarely fall), with no apparent chilling re-
quirement. B. philenor is primarily a foothill species, breeding
in canyons where the host plant grows along intermittent

IMCHCS OF INCHES OF lUlN

Fig. S.—Total seasonal rainfall at Sacramento, 1849/50 through 1968/69.
From Figgins, 1971.

82

ARTHUR M. SHAPIRO

J. Res. Lepid.

streams. A similarly mixed developmental strategy occurs in
Euchloe ausonides and Anthocharis sara in the foothills and may
be an adaptation to interseasonal variance in rainfall itself^
rather than stream flow. On the Sacramento Valley floor, B.
philenor and A. sara are both locally single-brooded, but E.
ausonides is double-brooded. There is preliminary evidence to
suggest that the Davis population of B. philenor is genetically
univoltine, perhaps reflecting founder effect — sampling error
introducing only a portion of the variability of a polymorphic
source population.

Dates of spring flights in both univoltine and multivoltine
butterflies are well known to be related to weather conditions,
both during the emergence season and during the preceding
winter. In the Valley, first-flight dates are less variable than in
either the Sierra Nevada or upstate New York, both cold- winter
areas (cf. Shapiro, 1974a). Even so, dates for the first species,
e.g. Pieris rapae, may vary several weeks among seasons. Pre-
cipitation advances the condition of the vegetation, but over-
wintering larvae and pupae respond to temperature and perhaps
photoperiod. Table 4 presents first-flight dates for 25 common
spring butterflies in the Valley after the very dry but mild
1971-72 and very wet and cold 1972-73 winters. Weather data
for the two seasons appears in Table 5 and may be compared
with the norms in Tables 2 and 3. Although the 1973 flight
season began early, continuing episodes of wet and windy
weather delayed most species relative to 1972. Emergences in
the Vaca canyons were earlier than in the Valley and showed
much less departure from 1972 dates, although population
levels of many species were quite different in the two years.

Later-emerging species are increasingly insensitive to weather
as a determinant of first-flight date. Satyrium sylvinus, for ex-
ample, first appeared v.lO in the Valley in 1973 as against v.l5 in
1972 (v.8.72 in the Vacas), but its period of peak numbers was
the same both years.

Cutoff dates for flight in autumn are also under meteoro-
logical control for the highly multivoltine species. An especially
sensitive species is Papilio zelicaon. In 1972 it was last recorded
on ix.20; in 1973 a full month later, on x.30. That this discrep-
ancy reflects a seasonal difference is well illustrated by data on
this species at Suisun Bay, near Fairfield, where the climate is
more maritime and P. zelicaon normally flies later; in 1972 it
was last seen on xi.5 and in 1973 on xi.l8.

(Continued on page 115)

Journal of Research on the Lepidoptera

13:83-98, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

EXTENDED FLIGHT PERIODS OF COASTAL
AND DUNE BUTTERFLIES IN CALIFORNIA’

ROBERT L. LANGSTON

31 Windsor Ave., Kensington, Calif. 94708

Several of the Rhopalocera that occur close to the Pacific
Ocean in California have very long flight periods, giving the
appearance of more than a single generation per year. Further
inland these same species ( often a differently named subspecies ) ,
have shorter flight periods, leaving no doubt of their being
univoltine.

To illustrate, I have chosen ten examples scattered among the
families Pieridae, Nymphalidae, Riodinidae, Lycaenidae and
Hesperiidae. In each case, one or two examples of the coastal
populations are treated. These are then compared directly with
populations further inland. “Inland” in some cases, may be only
20 miles or so away from the coast — if examples are available
from the middle and inner Coast Ranges. In other cases, the
comparisons are made with the West Slope, High, or East Slope
of the Sierra Nevada.

Figures 1 and 2 name the coastal populations in capital letters,
and those inland in lower case. The flight periods based on
known records are shown within the bars—black for coastal
populations and stippled for inland. The diagonal lines at one
or both ends of most bars show the probable extensions of the
flight periods based on: 1) Abundance at the beginning or end
of the known collections; and 2) Fresh and/or worn specimens
at the known beginning or end.

Although more $ $ are usually present at the beginning, and
more $ $ toward the end of the various flight periods, in general
both sexes were present throughout most of the coastal flights.
Since no Speyeria, Cercyonis or Erebia are involved in this study,
any staggered appearance of the sexes is considered negligible.

Tresented at the Twentieth Annual Meeting of the Lepidopterists’ Society,
Pacific Slope Section, Santa Barbara, Calif., 25 Aug. 1973.

83

84

ROBERT L. LANGSTON

/. Res. Lepid.

Fig. 1.— Flight periods of coastal populations (solid) and inland (stippled).
Diagonal lines indicate possible extensions of seasons.

13:83-98, 1974

FLIGHT PERIODS

85

The most exact measurements of flight periods are the cases
where there are sufficient records for a single season at the same
locality. This data was used whenever possible. However, from
my own records, and the data from several publications, it ap-
pears that most lepidopterists only visit the same location once
or twice in a single season. Hence, the other records are based
on the extremes of several seasons from data already published,
or on the labels of the specimens. Experienced collectors tend
to time their visits to obtain fresh ‘mint” specimens. For a
study of this type, it would be advantageous for the collectors
to also go after worn examples, or at least record them in field
notes. Partly because of lack or worn examples, I have extended
the probable seasons of some populations to later dates.

Pieris napi venosa Scudder

At Partington Canyon south of Big Sur in coastal Monterey
County, fresh specimens of both sexes were taken on 21 Feb.
1965. Other examples were taken various seasons in March, April
and May. My latest date is 15 May 1966-— some worn, but several
also fresh. Therefore, the season for the “true” heavily-veined
venosa would probably extend another three weeks-— into June
(diagonal lines in Fig. 1).

Further inland in the Coast Ranges (Alum Rock Park, Santa
Clara County; Stonybrook Canyon & Redwood Canyon, Alameda
County; and even as far inland as Thompson Canyon, Yolo
County) my personal records extend from mid-Feb. to mid- April.
This is a combination of many seasons, and the flight period is
considerably shorter than on the coast. Inland examples become
smaller and have less dark scaling on the veins. Many grade
into the taxon microstriata Comstock, relegated to a synonym of
venosa in dos Passos (1964).

Even further inland (West Slopes of the Sierra Nevada), I
have recorded it at the lower elevations from mid-March to mid-
April. This is based on fewer records, so I will give it the benefit
of the doubt and extend the season considerably at both ends.

Status of gen. aest. castoria Reakirt: Although stated in the
literature (Comstock, 1927; Tilden, 1965) that this is the second
brood of venosa, there is still some doubt. Many places where I
have found venosa commonly, I never found any castoria. In
fact, in my experience, castoria is a rather rare entity.

On the other hand, castoria has been found flying with the
heavily-veined venosa. Dennis Sorg (in his 1971 Season Sum-
mary contribution) found the two together as early as 20 March
1971 near Jasper Ridge, San Mateo County. I have not seen these

86

ROBERT L. LANGSTON

/. Res. Lepid.

Fig, 2. — Flight periods of coastal populations (solid) and inland (stippled).
Diagonal lines indicate possible extensions of seasons.

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FLIGHT PERIODS

87

specimens. On 15 May 1966, I found castoria flying with fresh
venosa at Partington Canyon. Would these actually be the sec-
ond brood from those flying in February and March? If not a
separate species, why are there still fresh venosa phenotypes
present at the same time?

The season on Fig. 1 for castoria is probably too long, as it
represents all of my records from south to north on the coast.
The mid-May record compares directly as it is also Partington
Canyon, and there are other records on into late June in the
Big Sur area (Bruce Walsh, correspondence). The extension of
the bar into August is based on a single $ from Van Damme
Park on the Mendocino County coast. Therefore, the season is
probably somewhat shorter at any single location.

Boloria epithore epithore (Edwards)

The upper black bar represents the one season of 1970 on the
Mendocino County coast. The earliest date is mine of 28 March
1970 at Russian Gulch State Park. On 30 May 1970 it was also
taken near Russian Gulch by John Emm el and several on the
same date near Caspar Beach by Paul Opler. With some fresh
and others worn, this single season probably extended well into
June. All of this data is cited in Perkins & Meyer (1973). Some
examples from north-coastal California are assignable to B.
epithore chermocki Perkins & Perkins (see Fig. 3).

The lower black bar represents epithore in the Santa Cruz
Mountains. Most dates are in May and June, but some in differ-
ent years include late April to the first of July (Perkins & Meyer,
1973). Since this represents many seasons from the 1890’s to the
present, I cannot extend it much on either end for a single season.

The stippled bars denote Boloria epithore sierra Perkins &
Meyer. The upper one represents four seasons I collected along
the North Fork of the Stanislaus River, Calaveras and Tuolumne
Counties. Worn examples were found in late June, 1973 at
slightly above 5000 ft. I did not take it that low in the other
seasons as it no doubt had already flown. The other seasons
it was considerably higher — almost 7000 ft. at Wet Meadow
(ridge to the south) or at Big Meadows along Highway 4 (to
the north). Therefore, it is in abundance for only about two
weeks at any one spot, and the season is only a little over a
month— even taking into consideration an almost 2000 foot range
in elevation. The lower bar denotes an even higher elevation
(Tioga Pass, Tuolumne Co.) where it extends from mid-July to
early August (Perkins & Meyer, 1973).

88

ROBERT L. LANGSTON

/. Res. Lepid.

Fig. 3.— Examples from coastal populations. Most localities in California,
unless otherwise noted.

Row 1. — Pieris napi venosa: Partington Canyon, 9 mi. S. of Big Sur,
Monterey Co., $ , $ 21 Feb. 1965, RLL. Boloria epithore chermocki: Rus-
sian Gulch, Hwy. 1, Mendocino Co., ^ 28 Mar. 1970, RLL. B, e. epithore:
Santa Cruz Mts., Santa Cruz Co., ^ 15 June 1946, T. W. Davies — CAS.
All following specimens leg. R. L. Langston.

Row 2. — Apodemia mormo mormo: Dunes W. of Seaside, Monterey
Co., ^ 4 Sept. 1969; $ 19 Aug. 1963. Callophrys viridis: San Bruno Mts.,
San Mateo Co., $ 30 Mar. 1968; Point Reyes dunes, Marin Co., $ 25 Apr,
1970; Fort Baker, S. of Sausalito, Marin Co., 9 7 Mar. 1970.

Row 3. — Callophrys viridis: Marina Beach dunes, Monterey Co., ^ 11
May 1969; $ 16 Apr. 1973. Plebefus icarioides moroensis: Dunes nr. Oso
Flaco Lake, 5 mi. S. of Oceano, San Luis Obispo Co., ^ 6 June 1966;
^ 18 Apr. 1973; 9 6 June 1966.

Row 4. — P. i. missionensis: San Bruno Mts., San Mateo Co., 9 2 Apr.
1972; Twin Peaks, San Francisco, ^ TOPOTYPE, 10 Apr, 1954; 9 TOPO-
TYPE, 15 May 1954; Plebejus pheres: Point Reyes dunes, Marin Co., $ 25
Apr. 1970, 9 10 May 1973.

Row 5. — Philotes enoptes smithi: Dunes W. of Seaside, Monterey Co.,
$ 24 Aug. 1962; 9 26 Aug. 1971; 9 4 Sept. 1969. Polites sonora siris:

3 mi. W. of Plantation, Sonoma Co., $ 5 May 1955; 9 23 July 1955.

Row 6. — P. s. siris: Lake Sylvia, Grays Harbor Co., Wash., $ , 9

4 July 1958. Panoquina panoquinoides errans: Mouth of Ventura River,
Ventura Co., ^ , 9 30 Aug. 1971; 1 mi. S. of Solana Beach, San Diego
Co., ^ 1 Sept. 1971.

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FLIGHT PERIODS

90

ROBERT L. LANGSTON

/. Res. Lepid.

Fig. 4.— Examples from inland and high elevation populations. Localities
in California, unless otherwise noted. All specimens leg. R. L. Langston.

Row l.—Pieris napi venosa: 3 mi. NE. of Tuolumne City, Tuol. Co.,
$ 23 Mar. 1966; Rock Creek, 6 mi. NE. of Placerville, El Dorado Co., 9
12 April 1973. Boloria epithore sierra: Wet Meadow, 6900', above N. fork
Stanislaus River, Tuolumne Co., $ 9 July 1969; Big Meadows, 6550', Hwy.
4, Calaveras Co., 9 30 June 1973.

Row 2.—-Apodemia mormo mormo: Del Puerto Canyon, Stanislaus Co.,
$ 21 Aug. 1962; 9 6 Sept. 1962. Callophrys lemherti: Upper Lyons
Creek, 7800', El Dorado Co., $ 11 June 1972; 9 6 June 1970; SE. shore,
Loon Lake, 6352', El Dorado Co., 9 9 June 1973.

Row 3. — Callophrys dumetorum: Dunes 1 mi. E. of Antioch, Contra
Costa Co., 9 11 April 1954. Plebejus icarioides ardea (Edwards):
Angel Lake, 8000', Elko Co., Nevada, $ 26 June 1972; 5 mi. SW. of
Henefer, Summit Co., Utah, 9 27 June 1972. P. i. lycea (Edwards): Pole
Mtn. area, 8640' Albany Co., Wyoming, 9 6 July 1972.

Row 4. — Plebejus pardalis: Hill SW. of Paradise Cay, Marin Co., 9
23 May 1964; Berkeley Hills, Alameda Co., $ 17 April 1954; 9 22 May
1954; Calistoga, Napa Co., $ 24 April 1956; Hill 3 mi. NE. of Vallejo,
Solano Co., 9 11 June 1967.

Row 5. — Philotes enoptes tildeni: Del Puerto Canyon, Stanislaus Co.,
^ PARATYPE, 11 Aug. 1962; $ TOPOTYPE, 9 Sept. 1967; 9 TOPO»
TYPE, 11 Sept. 1963; Polonio Pass, 3 mi. E. of Cholame, San Luis Obispo
Co., ^ , 9 27 Aug. 1973.

Row 6. — Polites sonora sonora: E. shore, Bucks Lake, 5153', Plumas
Co., $ 23 June 1949; King Canyon W. of Carson City, Nevada, 9 22 June
1970; SW. of Genoa, 5700', Douglas Co., Nevada, $ 20 July 1964; Mineral
King, 7831', Tulare Co., 9 3 Sept. 1949; Tioga Pass, 9941', Mono Co., 9
8 Sept. 1955.

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FLIGHT PERIODS

91

92

ROBERT L. LANGSTON

/. Res. Lepid.

Apodemia mormo mormo (Felder & Felder)

Based on numerous records ( Opler & Powell, 1961 ) , A mormo
in central and northern California is almost certainly univoltine,
with adults in greatest abundance in late summer. This is op-
posed to A. mormo virgtdti (Behr) which is multivoltine, with
records from February to November. On this and other bases,
it is felt by myself and others (correspondence, Mike Toliver),
that virgulti may be a separate species.

On the sand dunes west of Seaside, Monterey Co., this “uni-
voltine” mormo has been recorded from 4 July to 14 October in
various seasons ( Opler & Powell, 1961 ) . Both sexes ( with some
worn) were taken on 4 July 1959, so the probable season started
in late June. In several other seasons fresh and worn specimens
were taken in August and September.

Further inland, at Del Puerto Canyon, western Stanislaus
County, mormo were found only from mid-August to mid-
September. This canyon in the Inner Coast Range was also
visited several times in early August and no metal-marks were
found.

Callophrys viridis (Edwards)

The upper ridges of the San Bruno Mountains, San Mateo
County, are four miles inland from the coast. Although not on
the immediate coast like the dunes, they certainly get their share
of wind and cold, chilling fogs. Adults of viridis were recorded
from the San Brunos from 26 Feb. 1963 to 4 June 1963 (Gore-
lick, 1971). This is definite for a single season, with numerous
records for March and April in this and other years. The lower
black bar represents slightly later dates for the populations on
the dunes at Point Reyes, Marin County — 30 March to 18 June
(Gorelick, 1971). Since this is based on fewer records, the
season probably extends from a little earlier to somewhat later.

After the data was compiled for the Gorelick study, viridis
was discovered much further south— at the Marina Beach dunes,
Monterey County by J. F. Emmel and the author on 11 May
1969. Large numbers were subsequently collected here in April
and May, 1970-1973. All of these dates are well within the
seasonal extremes of the northern populations.

Callophrys dumetorum (Boisduval) is found further inland
in the dunes east of Antioch, Contra Costa County. The dates
span from 18 March to 20 April (Gorelick, 1971; Opler & Lang-

13:83-98, 1974

FLIGHT PERIODS

93

ston, 1968). Throughout its extensive range most records are in
March and April. The published records go from mid-February
to early June, but not more than about a month at any single
locality.

Callophrys lemberti Tilden is found much further inland and
at higher elevations. Fewer records are available for this species.
Its peak flight appears to be in early June at the 6 to 8000 foot
levels in El Dorado County, and late June at 9 to 11,000 feet
(Tioga Pass and Hoover Wilderness area. Mono County).

Plehejus icarioides moroensis (Sternitzky)

The dunes near Oso Flaco Lake, San Luis Obispo County
have been visited by various collectors, but usually in different
seasons. The earliest in the season appear to be 1 and 9 March
1972 by Bill Swisher, with moroensis recorded in April and May
in other years. My latest record is 6 June 1966 when long series
of mostly fresh ones were taken by myself and others. Therefore,
the season probably extends at least another two weeks. It was
not found on these dunes in August and September. The types
were taken at Morro Beach, S.L.O. Co. on 27 June 1929. This
is about 40 mi. NNW. of the populations indicated by the long
black bar. J. A. Powell has also taken it sparingly in July on
dunes to the north of Oso Flaco Lake. (These two are shown
by the short black bar.)

The icarioides in the higher Sierra Nevada and the White
Mountains, Mono County are dated from mid-June to mid-July.
According to Downey (1962), . . members of a local popula-

tion can be found in an area for about a 2-month period.” Mid-
June- July records are also prevalent for some of the other named
subspecies in Nevada, Utah and Wyoming (specimens in Fig. 4).

Plebejtis icarioides missionensis Hovanitz

The types were taken 1 April 1934 on Twin Peaks, San Fran-
cisco. Not far to the south, in the San Bruno Mountains, San
Mateo County, I have taken it on 4 April 1971 and 27 June 1971
—these extremes for a single season. There are numerous April,
May and June dates in other years.

For comparison, Plehejus pardalis pardalis (Behr) was chosen
from one colony in the Berkeley Hills— less than 20 miles to the
northeast, but not on the immediate coast. The dates are 17 April
1954 to 22 May 1954— again a single season at the same place.
This correlates with extensive records for pardalis in Contra

94

ROBERT L. LANGSTON

J. Res. Lepid.

Costa County (Opler & Langston, 1968) from six distinct locali-
ties— all in April and May, none in March or June.

Plebejus pheres (Boisduval)

Originally described from San Francisco, it is now presumably
extinct within the City — possibly its demise was similar to that of
Glaucopsyche xerces (Boisduval). A morphologically close pop-
ulation to pheres is still found on the dunes at Point Reyes,
Marin County. It has a flight period extending from mid-April
to early July [2 July 1945, 1 $ ] based on examples at the Cali-
fornia Academy of Sciences.

Further inland, Plebejus pardalis is again compared from the
northeast — Napa and Solano Counties. The flight period appears
to be late April to early June, with almost all of the records in
May.

Philotes enoptes smithi Mattoni

On the dunes west of Seaside, Monterey County, this sub-
species has been recorded from 20 June to 4 Sept, in various
seasons. Most of the records are in August, but long series have
been taken on several dates in July.

Further inland, Philotes enoptes tildeni Langston is found in
the Inner Coast Range adjacent to the San Joaquin Valley, usu-
ally in August and September. The stippled bar represents one
season at Del Puerto Canyon, western Stanislaus County. Only
two fresh S $ were taken 11 Aug. 1962, and the last collections
were made 6 Sept. 1962. This is based on most of the type series
cited in Langston ( 1963 ) . In various other years the known dates
go from 31 July to 11 Sept.

On 23 May 1967, six $ S were taken in Del Puerto Canyon
by Glenn Gorelick, and on 17 May 1973 another four ^ by
Jim Mori, indicating a spring flight (at least in some seasons).
As with almost everything biological, there seem to be excep-
tions or “freak” occurrences.

Both smithi and tildeni could extend into late September or
October, but I have no records of their actually being collected.
This is predicted by correlation with Apodemia mormo, which
is abundant at both localities and has the same Eriogonum hosts.

Polites sonora siris (Edwards)

West of Plantation in coastal Sonoma County, I collected
good series of siris on 5 May 1955 and 23 July 1955™in the same
season. It was present other years in June, early and mid-July.
However, it was not found on visits to the exact same spot in

13:83-98, 1974

FLIGHT PERIODS

95

August and September. July specimens from western Washing-
ton are illustrated for comparison (Fig. 3). These show little
variation from those in coastal northern California.

Polites sonora sonora (Scudder) is found in the Sierra Nevada
at moderate to high elevations. More records are needed, but
based on the few I have available, it flies from mid-June to late
July from 5000 to 7000 feet, and in August to early September
from 7000 to 10,000 feet.

Panoquina panoquinoides errans (Skinner)

This is the only other skipper in the whole Superfamily
Hesperioidea that to my knowledge is restricted to the immediate
coast of Cahfornia. It violates the theme of those previously
mentioned, as it probably has two or more generations per year.
Comstock (1927) indicates it flying in mid- August and Sep-
tember. Most collections ( especially those with large numbers ) ,
have indeed been in late summer. However, myself and others
took it near Solana Beach, San Diego County as early as 19
June 1963, and it was still present on 1 July 1963 — again in the
same season.

Since there are no other species or subspecies of the genus
Panoquina occurring in California, no inland comparison is
made for errans.

DISCUSSION

Climate:

Outer Coast Range and beach dunes of California — mild winters
and cool summers.

Very seldom does the weather go below freezing, or does
frost occur on the immediate coast. On sunny days in January
and February the south-facing slopes and the protected sides
of the dunes can get quite hot. Since it does not get extremely
cold at night, the development of Lepidoptera (and other in-
sects ) can commence quite early in many seasons.

Conversely, in the summer the coastal slopes and dunes are
successively plagued by long periods of cold winds and fogs.
In some places for several days in a row ( or even weeks ) the sun
may not shine. During these periods, new individuals do not
emerge from their pupae, and those already in the adult stage
often remain quiescent. They do not carry on their activities of
taking nectar, mating or ovipositing. When the weather becomes
clear and sunny before the next siege of fog, the adults resume
their activities.

96

ROBERT L. LANGSTON

/. Res. Lepid.

Inner Coast Ranges and Sierra Nevada of California — cold win-
ters and hot summers.

The insect season is shorter inland, particularly at the higher
elevations. Below freezing and winter frosts are common in the
Inner Coast Ranges. Heavy snows and several months of below
freezing are characteristic of the Sierra Nevada.

In the spring, the days may warm up suddenly, even though
the nights may still be below freezing. When the days have
warmed, it generally stays warm and gets progressively hotter
as summer approaches. This will instigate rather quick and near
total emergence of adults in a short period of time. With nothing
to delay them, the adults take nectar, mate and oviposit. With
all activities completed, the flight period is soon finished.

Host plants:

Directly correlated with the climate is the condition of the
larval food-plants. It is not the purpose of this paper to specific-
ally record the hosts of the various Lepidoptera concerned.
Therefore, I will treat only some of the examples, and for brevity
refer to the hosts by genus only.

Pieris napi venosa in the Coast Ranges seems to prefer Den-
taria (milk-maids), although Bruce Walsh (personal correspon-
dence) of Carmel Valley has also found it ovipositing on Arahis
(mustard). This could well be an alternate host for it and
gen. aest. castoria. On the immediate coast the Dentaria stays
in green succulent growth for a long time. Further inland it
grows faster, blooms and dries out rather quickly. Possibly the
populations of venosa inland do not work over to several other
cruciferous plants that would appear 'suitable,” This could
explain the absence of castoria and a shorter adult flight of
venosa in the more inland areas. On the West Slopes of the Sierra
Nevada, one of the hosts appears to be Radicula (water-cress)
found in slow-moving parts of creeks and the seepage from
springs.

Apodemia, coastal Callophrys and Philotes are well docu-
mented in the literature (Opler & Powell, 1961; Gorelick, 1971;
Langston, 1969, respectively) as feeding on various species of
Eriogonum (wild buckwheat). In the coastal areas the Eriogo-
num has considerable leaf growth early in the season. The flowers
develop over an extended period from spring through summer.

13:83-98, 1974

FLIGHT PERIODS

97

By mid-summer the plants in a local area may be in various stages
of bud, blossom or seed. Further inland, each separate species
of Eriogonum tends to bloom “all-at-once,” with the flowers soon
becoming unattractive, even as a nectar source.

The Plebejus species cited are all recorded as having various
species of Lupinus (lupine) as their larval hosts. According to
Downey (1962) the peak adult flight period seems to occur at
about the same time as seed formation in the host lupines. The
eggs are deposited singly on leaves, stems, flowers and seed pods.
On the coastal dunes the lupines (especially the large yellow-
flowered perennial) remain green and succulent all year. The
blooming period is very staggered, with buds, flowers and young
seed pods at the same time on the same and/or adjacent plants.
Further inland and at higher elevations, most lupines are in
bloom for a relatively short time in the “spring,” set pods, and
soon become incompatible as hosts.

SUMMARY and CONCLUSIONS

Several species of univoltine Rhopalocera found along the
coast of California have very long adult flight periods. This is
due to mild winters and cool summers, combined with the re-
sultant staggered development of the various larval food-plants.

Just a short distance inland in the Coast Ranges and further
inland in the Sierra Nevada, these same species ( or closest known
taxa) have much shorter adult flights. This is due to cold win-
ters, warming in spring and hot summers. These short flight
periods are also correlated with increased elevation and a rather
quick and uniform development of the larval food-plants, and
their soon becoming unsuitable.

Examples of ten coastal entities among the families Pieridae,
Nymphalidae, Riodinidae, Lycaenidae and Hesperiidae are com-
pared with their inland counterparts.

ACKNOWLEDGMENTS

Some of the examples in this study were taken from my field
notes while on collecting trips with the California Insect Survey.
I wish to thank Dr. J. A. Powell, University of California, Berke-
ley and others involved with the Survey for the opportunity to
make these collections. Specimens were examined at the Cali-
fornia Academy of Sciences, San Francisco for additional records.
I am indebted to Dr. P. H. Amaud for his cooperation and ac-

98

ROBERT L. LANGSTON

/. Res. Lepid.

cess to the CAS collections. I am grateful to the many who have
contributed to the Zone 1 Season Summary since I became co-
ordinator. Data was gleaned from their contributions, and par-
ticularly the related correspondence of those mentioned in the
text.

LITERATURE CITED

COMSTOCK, J. A. 1927. Butterflies of California. Publ. by author, Los An-
geles. 334 pp.

DOS PASSOS, C. F. 1964. A synonymic list of the Nearctic Rhopalocera.
Lepid. Soc. Mem., No. 1, New Haven, v + 145 pp.

DOWNEY, J. C. 1962. Variation in Plebejus icarioides ( Lepidoptera,
Lycaenidae). II. Parasites of the immature stages. Ann. Ent. Soc. Amer-
ica 55 (4): 367-373.

GORELICK, G. A. 1971. A biosystematic study of two species of Callophrys
(Callophrys) in California (Lycaenidae). J. Lepid. Soc. 25 ( SuppL 2):
1-41.

LANGSTON, R. L. 1963. Philotes of central coastal California (Lycaeni-
dae). 7. Lepid. Soc. 17 (4): 201-223.

1969. Philotes of North America: Synonymic list and distribution

(Lycaenidae). J. Lepid. Soc. 23 (1): 49-62.

OPLER, P. A., and R. L. LANGSTON. 1968. A distributional analysis of
the butterflies of Contra Costa County, California. J. Lepid. Soc.
22 (2): 89-107.

and J. A. POWELL. 1961. Taxonomic and distributional studies on

the western components of the Apodemia mormo complex (Riodini-
dae). 7. Lepid. Soc. 15 (3): 145-171.

PERKINS, E. M., and W. C. MEYER. 1973. Revision of the Boloria epithore
complex, with description of two new subspecies ( Nymphalidae). Bull.
Allyn Museum, No. 11: 1-23.

.TILDEN, j. W. 1965. Butterflies of the San Francisco Bay Region. Calif.
Natural History Guides: 12. Univ, Calif. Press, Berkeley. 88 pp.

APPENDIX

Many of the dates and locations cited in the text are based on
specimens in the two accompanying photographs. These data
are given in the legends to the figures.

Journal of Research on the Lepidoptera

13:99-100, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

HABITAT: BREPHIDIUM PSEUDOFEA
(LYCAENIDAE)*

JOHN B. HEPPNER

Depaiiment of Entomology and Nematology,
University of Florida
Gainesville, Florida 32611

Brephidium pseudofea (Morrison) typically is found in as-
sociation with Salicornia bigelovii ( Chenopodiaceae ) , the only
definitely known host plant, along the coasts of Florida and
somewhat further north. The illustration is of an area along the
west coast of central Florida: Cedar Key, Levy County (19 Dec
1973).

Salicornia virginica is the predominant plant in the fore-
ground, with some Salicornia bigelovii mixed in, left. Perhaps
S. virginica is an alternate host; Rawson ( 1961, J. N, Y. Entomol.
Soc. 69:88-91) has also suggested Batis maritima. B. maritima is
a prostrate plant with succulent leaves growing near the Sali-
cornia. The saltmarsh grasses to the left and in the background
are mostly Spartina alterniflora (Gramineae). Bushes in the
picture include Avicennia germinans ( Verbenaceae) (large bush,
right ) and the smaller Lycium carolinianum ( Solanaceae ) ( cen-
ter, right ) , the latter in bloom and attracting other lepidopterous
species {Danaus, Agraulis, Urbanus, Panoquina) but no Bre-
phidium were observed on the Lycium flowers. B. pseudofea
flies in colonies as many other lycaenids tend to do and has
been reported to be generally uncommon in Florida, encoun-
tered in numbers only occasionally. Flight records are year
round in the southernmost areas except for the months of August
and November (Kimball, 1965, Lep. Fla., p. 49).

^Florida Agricultural Experiment Station Journal Series No. 5309.

99

100

JOHN B. HEPPNER

J. Res. Lepid.

Fig. 1. — Coastal strand with Salicornia spp. in foreground on west end of
Cedar Key, Levy Co., Florida. (19 Dec 1973)

Journal of Research on the Lepidoptera

13:101-»114, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

THE EARLY STAGES OF VARIOUS SPECIES OF
THE GENUS DIRPHIA ( SATURNHDAE )

BRIAN O. C. GARDINER

18, Chesterton Hall Crescent, Cambridge, England.

INTRODUCTION

This paper is one of a series describing the rearing in Eng-
land of various Neotropical Saturniidae and deals with five
species of Dirphia, as defined by Michener (1952). Eggs were
sent to me by airmail; D. avia Stoll from Trinidad by Dr, A. D.
Blest; D. baroma Schaus, D. curitiba Draudt, D. lombardi Bou-
vier, and D. ursina Walker, from St. Catarina, Brazil, by Sr.
Fritz Plaumann. The first three of these were successfully
reared through to adults, but the larvae of the last two died
after several instars. The various stages were kept at 20-25° C.,
with occasional fluctuations of ± 5°C. and under natural day-
light conditions, but with additional light during the evenings
while the stocks were being attended to. The duration of the
various stages is summarized in Table 1.

DIRPHIA AVIA

From 375 eggs received 23.VL61, a total of 205 healthy and
8 deformed pupae were obtained. In addition about 25 larvae
were preserved.

The egg5— Laid in rather irregular batches. Shiny white with
a small black micropyle. Virtually spherical 2.5 mm diameter.
There is no prior indication before the larva hatches.

The larvae— In the 1st instar black with a dark brown head.
Chalazae black, forked like a Y on the first 3 segments, simple,
but a distinct bend two thirds of the way up, on the remainder.
In the 2nd instar they become greyish dorsally, reddish-brown
ventrally. In the 3rd instar the ground color a dirty white with
black markings, these now persisting to the final stage, the
ventral surface becoming less reddish and more like the dorsal
as growth progresses.

101

102

BRIAN O. C. GARDINER

/. Res. Lepid.

The fullgrown larvae 8-10 cms long (the females being the
larger). Ground color, including the head, a light grey with a
tinge of green or violet, the color depending rather on how the
larva is being viewed and the incident light. On each segment
black markings, variable in both number and extent, the most
prominent being a dorsal mark like a broad arrow ( ) facing

backwards and situated on each segment between the chalazae.
The spiracles white, outlined in black. The chalazae grey; the
spires grey with black tips; both have a violet sheen to them.
The armature firm and fairly formidable. The thoracic chalazae
and spines particularly long, overhanging and protecting the
head. The prolegs with black crochets. The head grey with a
broad inverted V above the brown mouthparts.

Larval habits — The larvae are gregarious throughout their
life and wander about in long processionary columns. They
feed by night and rest by day in a dense communal cluster.
This was usually formed at the base of the foodplant which in
this case consisted of a polythene water bottle containing the
branches on which the larvae were being fed. So determined
were the larvae to walk down the stalks that unless the neck
of the bottle was extremely tightly plugged with Kleenex tissue,
several of the larvae would drown themselves. When they are
disturbed the larvae cling tight; they neither drop nor assume
any unusual attitude, except that the head is slightly retracted
and the thoracic spines lowered around it.

Sting — From the second to fourth instar the sting from the
spines is about equivalent to that from a nettle ( Urtica diocia ) ,
or a fullgrown Automeris io Fabricius, That of the fullgrown
larva however is far more severe, as painful as that of wasps
(Vespa vulgaris Linnaeus; V. germanica Fabricius), but shows
only the usual type of Histamine reaction (Jones & Miller, 1954)
bleb which soon wears off although the area is a little tender
for a few days.

Pupation— This species was the first of the Neotropical Hemi-
leucinae to be bred by the author and their unusual habits and
individual idiosyncrasies were still strange to him. The finding
of the correct foodplants and the optimum conditions for pupa-
tion were as yet unknown. The D. avia were therefore given
a standard choice by being given a layer of peat overlain with
moss in their rearing cage. This gives all the conditions required
by species that pupate (a) in rolled leaves; (b) in litter; (c)
underground. This species proved to be one of the few per cent

13:101-114, 1974

DIRPHIA EARLY STAGES

103

that is not satisfied with any of these three usual alternatives and
demand specialized treatment.

When they are ready for pupation the larvae turn a purplish-
red color which gradually darkens as they restlessly hunt for a
suitable site over two or three nights. However, once the cocoon
has been spun, the pre-pupa assumes the original larval color
except for the spines, these remaining purple.

In their cages they wandered endlessly around, but during the
day took their place in the communal resting cluster. Only about
a dozen actually spun cocoons in the cages and over half of
these failed to form normal pupae. Seeing that they were so
obviously unhappy in the cages the larvae were removed and
placed either singly in lb. tins or in pairs in 2 lb. tins. These
tins contained a wad of sphagnum moss on top of a few inches
of moist peat. They were fitted with tight metal lids and were
used on the theory that the larvae were seeking somewhere
totally dark. Be that as it may, all the larvae put into the tins
pupated successfully. They formed a large flimsy papery cocoon
between the peat and the moss, often attached to the side of
the tin.

It is certain that in nature these larvae choose some special-
ized site; it is not unlikely that this is inside some species of
Epiphyte.

Foodplants — The larvae were reasonably polyphytophagous
on various temperate deciduous trees and one evergreen tree.
The following were accepted: Hawthorn (Crataegus oxyacan-
thae); Beech (Fagus sylvatica); Plums, Cherries (Prunus spp.)
Oaks (Quercus spp. — including ilex); Apple (Malus sp.). Black
poplar (Populus nigra) and Privet (Ligustrum ovalifolium)
were refused. Very freshly shooted Willow (Salix sp. ) was
eaten for the first two days by the newly hatched FI. larvae,
but they so readily switched over to Evergreen oak ilex)
that Willow was not considered really suitable. For the sake of
convenience the larvae were mainly reared on Hawthorn and
Beech, the FI on Evergreen oak.

Adults — The majority of the pupae were disposed of and only
a few kept for adult emergence. Two pairs were obtained, both
of females that were three days old and had already laid a
considerable number of eggs. Both these pairs were obtained
when the temperature had been raised from 22° C to 30 °C over
some four hours and then dropped over one hour to 25 °C.
Several other moths placed together failed to pair. It is not

104

BRIAN O. C. GARDINER

J. Res. Lepid.

clear if the temperature fluctuation was of any significance or
not. The two pairs occurred some five hours after sunset, the
moths remained in copula for at least 90 minutes, had separated
by morning.

From these two females a few small egg-batches were laid,
in both cases laying only took place over about the 3 hours
after dusk. About 75 per cent of the eggs hatched.

The adults lived for about 7 days. Females commeneed to
‘call’ the same day as they emerged. Females started to lay
infertile eggs in irregular masses if not paired after two or three
nights, although even after this they would spend a short time
calling each night. Males soon battered their wings to pieees
after whieh they flapped rather helplessly on the cage floor.
DIRPHIA BAROMA

Eggs of this speeies were in a paeket that arrived 29.VIII.63
after being a month in transit from Brazil. There had been
some shaking up and these eggs were mixed with those of D.
Curitiba. They had just hatehed on arrival. All five were reared.

Eggs — The remains of the eggshells were indistinguishable
from those of D. curitiba (see below).

The larvae — In the 1st instar body dark reddish-brown. Head
shiny blaek. Chalazae off-white, blaekish towards tips. No
reeords were kept of intermediate instars, but by the third the
larva had assumed the general pattern of the final. The final
instar larva 8 cms long. The head and anal segments blaek.
Ground color, ventral and dorsal, black, with a brick-red in-
complete band on each segment laterally and dorsally. Legs
black with red spots. Chalazae and spines pale blue. The
chalazae short, the spines arising starlike fairly long. Rather
uniform and regular on each segment. This arrangement in
fact resembles that of Automeris inemusae Walker or A. nycti-
mene Latreille.

Larval habits — Since there were only five specimens the
larvae were kept throughout in a plastic box. Apart from the
fact that they kept bunehed together and fed by night, the
sample was too small to give any clear indication of habits.

Sfing— This was not experienced.

Pupation — When ready to pupate the eolors faded. The
larvae were removed to tins with moss and peat. They pupated
underground in the peat, forming a cocoon similar to that of a
Sphingiid.

13:101-114, 1974

DIRPHIA EARLY STAGES

105

Foodplants — The newly hatched larvae were offered Beech,
Hawthorn and Laburnum {Laburnum anagyroides) on which
they commenced to feed, but after a week they transferred to
Beech. The final instar finished their development on Ever-
green oak. The following were refused: Privet; Plum; Elm
(Ulmus campestris).

Adults — After nearly three months all the pupae produced
adults within a week. Unfortunately all five were females.

DIRPHIA CURITIBA

Eggs of this species were received in 1962 and again in 1963.
Those of the first year were reared with virtually no larval
mortality, but from the second year there was a high larval
mortality from what appeared to be granulosis virus disease.
This disease caused the gradual and total loss of the El. larvae.
The information below is based on the first lot received 29.IX.62,
which were a month in transit.

Eggs — Shaped like a slightly flattened ovoid, color white with
a large black micropyle. Laid in regular groups of several
dozen eggs, fastened strongly to the substratum with a clear
cement. It was noticed that infertile eggs were encircled by a
wide grey band on the upper half. No prior indication to larval
hatch. Eggshells partly consumed by the newly hatched larvae.

The larvae — In the first instar brownish, including spines,
head black. Second instar similar. In subsequent instars green-
ish-grey with black markings. The fullgrown larva 7.5 to 10.0
cms long (the females doubtless being the larger). Dorsallly
the ground color greenish-grey, or brownish, ventrally whitish-
grey. The whole body covered with an intricate series of black
markings which vary very considerably between larvae, in some
coalescing to form large blotches which run continuously from
one segment to the next. The spiracles are white, lightly out-
lined in black. The whole larva densely covered with spined
chalazae. These particularly large and dense on the thoracic
segments, and the anal, being here at least 1.5 cms long, pro-
jecting over the head and to the rear, as in avia.

Larval habits — Similar to avia.

Sting — Similar to that of avia.

Pupation — Also similar to avia and it was necessary to re-
move the larvae individually to tins. They also turn purple when
ready for pupation. The cocoon appeared to be a little flimsier
than that of avia.

106

BRIAN O. C. GARDINER

J. Res. Lepid.

Foodplants — Having been informed by Sr. Plaumann that
they fed on Juglandaceae, especially Walnut {Juglandia regia)
this was offered, together with Plum and Laburnum. They com-
menced to feed on the Walnut two days after hatching but by
the fourth instar the supply failed. Oak and Peach {Prunus
persica) were oflFered and accepted. Holly {Ilex sp.); Privet
and Laurel (P. lusitanica) were refused. Their development
was completed on Evergreen oak.

In the FI generation the newly hatched larvae refused to
start feeding on Walnut. This had been forced and it might
well be the the very fresh soft leaf is unsuitable in some way.
The parents had of course been given late fall leaves. These
FI larvae, with about 20 per cent loss, eventually started feed-
ing on Oak, but after two weeks all the larvae accepted Beech
and completed their development on this. The F3 generation
was fed entirely on Evergreen oak.

The newly arrived larvae in 1963 were given a choice of
Walnut and Beech. All preferred the Beech. The FI gener-
ation of these was offered and fed on Hawthorn (other leaf
being unavailable). As already stated however all these died
of what appeared to be virus disease.

Adults — These behaved in much the same way as avia.
Probably because rather more were available pairing proved
relatively easy, without the temperature fluctuation which oc-
cured before the avia pairing. Pairing took place some 2-3 hours
after dusk and lasted 1-2 hours although two to three pairs
were still coupled the following morning. Not all the pairings
produced fertile eggs. Females either laying fully fertile, totally
infertile, or mixed batches. The females that laid totally infertile
batches behaved as if they were laying fertile eggs. That is,
they laid in regular batches. Virgins, after several nights Tail-
ing’ would lay in a jumbled pile. It seems as if the correct
laying reaction is initiated by the act of coupling, not fertiliza-
tion. In the conditions of captivity 100-150 eggs per female were
laid. This is by no means their full potential, on estimate, less
than half. Only 1 pairing was obtained of the F2 generation.
Two pairings were obtained from the F3 but no viable eggs
were laid. By now of course the stock was brothers and sisters
already one generation inbred, so this is perhaps not surprising.

13:101-114, 1974

DIRPHIA EARLY STAGES

107

DIRPHIA LOMBARDI

Eggs of this species were received 29.IX.62 and were just
hatching on arrival after a month in transit. Sr. Plaumann gave
Lauraceae? as possible food. All the larvae died by the end of
the fourth instar.

Eggs — As received these were laid in a jumbled pile. The
color white with a black micropyle. In shape, a slightly flat-
tened ovoid, 1.75 mm x 1.5 mm x 2.0 mm high.

The larvae — In the first instar the dorsal surface rather black-
ish, a dirty white ventrally and laterally. Head black. After
feeding and growing a little the larvae become overall whitish
with black spines. In the 3rd instar a green lateral stripe has
appeared, ground color now greyish-brown. The ultimate death
of the larvae appeared to be due to granulosis virus disease.

Larval habits— Tiocessionstiy and gregarious in the first three
instars.

Sting — Not experienced.

Pupation— None were pupated.

Foodplants— The newly hatched larvae were offered a choice
of Black locust ( Rohinia pseudo-acacia ) , Plum and Beech. They
commenced to feed on Plum on which they were kept through-
out their life. Evergreen oak, and Laburnum which were offered
in the 3rd instar were refused.

DIRPHIA URSINA

Eggs of this species were received 29.IX.62 and again on
22.X.64. None of the larvae survived beyond the end of the
fourth instar. The first batch apparently died of granulosis
virus disease. The second batch almost certainly died due to
the failure to find a suitable food for them; due to their rather
late fall arrival the plum known to be suitable had already
nearly fallen and the unsuccessful attempts to find a suitable
alterantive led to starvation and resultant debility in the first
instar. None survived beyond the second.

Eggs— As received these were laid in a jumbled pile. The
color off-white with a black micropyle. Almost spherical and
about 2.0 mm in diameter.

The larvae— The newly hatched larvae are purplish in color,
including the spines and chalazae, with a black head. There
was a steady mortality of the 50 or so larvae that commenced
to feed. The three that reached the fourth instar had hardly
changed in appearance from the first.

108

BRIAN O. C. GARDINER

y. Res. Lepid.

Larval habits — Processionary and gregarious in the first few
instars.

Sting — Not experienced.

Pupation — None were pupated.

F oodplants — The newly hatched larvae were offered a choice
of Black locust or Plum. The latter was accepted. Accompany-
ing the second batch was a note from Sr. Plaumann that they
fed on Wistaria (Glyzine). In late October in England the only
Wistaria leaves available were in an advanced state of senes-
cence; once pieked they dried up overnight, even when kept in
a high humidity. There were, not unexpeetedly, refused by the
larvae whieh again accepted Plum. Decent green leaves of this
too, were hard to find. Laburnum, Everlasting pea {Lathyriis
latifoliiis) and Laurel were offered but refused.

DISCUSSION

I have been informed ( Blest, in lit. ) and by others who
have had tropical experience, that many Tropical Lepidoptera,
including specifically Dirphia and Aiitomeris species are ex-
tremely difficult to rear where they occur. Either they refuse to
accept any available foodplant offered, or they prove to have
been attacked by parasites, or die of virus diseases. Larvae col-
lected wild in the forest have been known to refuse to accept
the leaves of an apparently identieal tree to that on which they
were found. Also it is not very easy to collect foodplant from
the canopy of a tropical forest.

When translated to a temperate zone, natural parasites, but
not inheritable virus diseases are eliminated. So too of course,
are the natural f oodplants. Since in the majority of cases these
are unknown, it becomes a question of trying one tree after
another until one that is acceptable to the larvae is offered.
Once they have commenced to feed, other leaf may be offered
later, either because it is more readily available, or simply to
find out what range of plants are acceptable. In the case of
these Dirphia the larvae of three species proved to be reasonably
polyphytophagous on temperate zone deciduous trees. The other
two, ursina and lomhardi only accepted one. Plum, but the
larvae only having been available late in the year, other leaf,
acceptable to the first three, was not available. It is perhaps
significant, however, that Evergreen oak was refused. Without
this the first three species could not have been reared. It would
appear that ursina and lombardi have a different food range to
avia, baroma and curitiba.

13:101-114, 1974

DIRPHIA EARLY STAGES

109

The larvae of ursina and lombardi having died in the early
stages, it is difficult to make a comparison between them and
the other three. Nevertheless their general appearance and the
arrangement of the chaiazae and spines were similar to those
of avia and curitiba in similar instars. The odd one out is
baroma which as already mentioned, has a structure similar to
that of certain Automeris larvae. Its mode of pupation is also
unusual being dissimilar to that of any other Dirpha moth known
to me, but similar to that of Eudyaria venata Butler (Crotch,
1956).

Little appears to be known about the life-cycles, but I have
not carried out an exhaustive search of the literature. This
would be no easy matter, as much of the literature likely to
carry information concerning Dirphia (i.e., local South Ameri-
can journals) is not very readily available. Before and during
this investigation some two dozen enquiries were made of
various institutions and colleagues in the Neotropical area. None
was able to give me any information.

In the conditions of captivity there was a life-cycle, in the
three species reared to adults, of six months and two generations
a year could be obtained. It is difficult to obtain precise infor-
mation on the occurrence and number of broods of many tropi-
cal species, but it does appear that these Dirphia are normally
single-brooded. The related D. jorgenseni Schaus (Schreiter,
1943) is single-brooded in Northern Argentina and my supplier
in Southern Brazil, Sr. Plaumann, tells me that curitiba and
other species are single-brooded. This data quite clearly shows
that these species therefore, have a facultative light-controlled
diapause, but the exact determining conditions were not clearly
defined by the present investigation.

According to Crotch (1956) Dirphia larvae are said to be
long, cylindrical, with long slender single spines emerging di-
rectly from the body; at least one species is thought to have
large lappets; a few species appear to spin cocoons not unlike
Saturnia pyri Schiffermiiller. None of these statements is true
of the five species described in this paper. It is true that Dirphi-
opsis eumedide Stoll has two small paired lappets' (Gardiner,
1966), but these are neither large, nor in the position assumed.
This species was included in Dirphia by Draudt ( 1930 ) .

The following larvae, of which I have traced a published
description, bear a structural similarity to avia and curitiba.
Eudyaria venata Butler ( Burmeister, 1879, as Dirphia consularis

110

BRIAN O. C. GARDINER

/. Res. Lepid.

Fig. 1. — Dirphia imagines. Top, avia; second row, ciiritiba; third row,
ursina; bottom, baroma, female only.

13:101-114, 1974

DIRPHIA EARLY STAGES

111

Fig. 2.- — Larva of Dirphia avia.

Fig. 3. — -Larva of Dirphia curitiba.

Fig. 4.”-Larva of Dirphia lomhardi.

112

BRIAN O. C. GARDINER

/. Res. Lepid.

Burmeister ) ; MoUppa sabina Walker and M. simillima D. -Jones
(D.-Jones, 1907). D. jorgenseni Schaus, however, (Schreiter,
1943) bears a remarkable resemblance to baroma, but pupates
amongst the foodplant, not underground.

The question arises as to whether the present v/ork on the
larvae has any bearing on the confused issue of the classification
of Dirphia and its relatives.

Draudt in Seitz (1930) divided the genus Dirphia into ten
groups. This grouping was largely re-distributed by Michener
(1952), some species being removed by him to separate genera,
other genera of Draudt being brought into the Dirphia orbit.
Although Michener uses a system of subgenera, I prefer, taking
into account the larval characteristics and pupation habits, to
regard his subgenera as separate genera. It is impossible from
larval characters as at present known, to regard Periphoba hircia
(see Gardiner, 1967) as being at all closely allied to for instance
Dirphiopsis eumedide (see Gardiner, 1966) and Dirphia baroma,
or the latter two to each other. Dirphiopsis eumedide shows
similarities to Dirphia jorgenseni and they should perhaps be
placed in the same genus but not Dirphia. On the other hand
E. venata bears such a striking resemblance to avia or curitiba
that it should be transferred back into Dirphia.

The larvae of lombardi and ursina at the time of their death
in the fourth instar clearly resembled contemporary larvae of
avia or curitiba and it is therefore considered that they too
belong in Dirphia.

ACKNOWLEDGMENTS

This work was supported under a grant by the United States
National Institutes of Health (Project GM-07109). Thanks are
due to Dr. A. D. Blest for the photograph illustrating the larva
of baroma.

LITERATURE CITED

BURMEISTER, J. 1879. Atlas de la description physique de la Repiiblique
Argentine contenant des viies pittoresques et des figures d’histoire
naturelle. Lepidopteres 5: Buenos Aires.

CROTCH, W. J. B. 1956. A Silkmoth Rearer’s Handbook. The Amateur
Entomologist 12: 1-165.

DRAUDT, M. in SEITZ, A. 1930. Macrolepidoptera of the World. 6, 713-
827.

DUKINFIELD JONES, E. 1907. On the remarkable resemblance between
two species of Molippa. Trans. Ent. Soc. Lond. 1907: 181-182.
GARDINER, B. O. C. 1967. The life history of Periphoba hircia ( Saturni-
idae) with a note on distribution and larval variation. J. Lep. Soc. 21:
198-204.

13:101-114, 1974

DIRPHIA EARLY STAGES

113

Fig. 5. — Larva of Dirphia haroma.

TABLE 1

The duration in days of the various stages in the life-cycle
of the Dirphia species

Species

Egg

1

2

avia

30

8

8

baroma

34

9

le

Curitiba

32

9

7

lombardi

31

7

7

ursina

31

6

11

3 4 5 6

7 4 4 8

7 IZ 10 13

7 8 12 30-45

10 - -

7

Mean

Total

Pupa

25-37

70

62

0

61

82

0

81

105

11

114

BRIAN O. C. GARDINER

/. Res. Lepid.

- — — 1966. The rearing of Dirphiopsis eumedide ( Saturniidae ) . /. Res.
Lepid. 4: 287-292.

JONES, D. L. & J. H. MILLER. 1959. Pathology of the dermatitis pro-
duced by the urticating caterpillar, Automeris io. Arch. Dermatol. 79,
81-85.

MICHENER, C. D. 1952. The Saturniidae ( Lepidoptera ) of the Western
Hemisphere, Morphology, phylogeny and classification. Bull. Amer.
Mus. Nat. Hist. 98: 335-501.

SCHREITER, R. 1943. Notas Entomo-Biologicas y otros. Acta. Zool. Lillo-
ana 1: 7-44.

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

115

(Continued from page 82)

VOLTINISM AND HOST SPECIALIZATION

Opler and Bucket! (1971), working with the large macro-
moth fauna of Santa Clara County, California, suggest that the
breadth of host specialization is correlated with voltinism; spe-
cifically, that univoltine species tend to be monophagous and
multi voltines polyphagous. This correlation, if valid, suggests
a primary biological basis for the adaptiveness of univoltinism:
synchronization with a host plant which is seasonally limited in
availability. Individual examples of such life cycles are well-
known in North American butterflies. In some cases (e.g. Pieris
virginiensis Edwards on Dentaria ) the host plant is an ephemeral
vernal species, annual or perennial. In others the edible part of
the host is available only at certain seasons, even though the
plant itself is present most or all of the year. This seems to be
true of the insect faunas of tannin-rich plants. Tannins interfere
with protein uptake, and the foliage of such plants is nutrition-
ally useful only in spring before much tannin has accumulated
(first demonstrated for the Geometrid Operophtera on oak by
Feeny, 1970; probably applicable to Satyrium, Erynnis, etc.).
Burns ( 1964 ) notes that the multivoltine Erynnis tristis oviposits
and feeds on the young growth on evergreen oaks and avoids
the older leaves. Such growth is limited in quantity and might
be expected to be the object of intense competition among oak-
feeding insects so long as the disadvantages of competition do
not outweigh the advantages of rearing more than one brood
a year.

Fair to good host plant data are at hand for 37 species of
butterflies in the Sacramento Valley. These may be put into
three categories of host specialization: monophagous, defined
as restricted to a single plant genus in the geographical area in
question; oligophagous, restricted to two or more genera in the
same plant family; and polyphagous, feeding on two or more
plant families. In the Valley all of the univoltine species are
monophagous (Table 6), while just over two-thirds of the multi-
voltines are oligo- or polyphagous. Oligophagy is the commonest
pattern: only three multivoltines {Vanessa cardui, Plebeius
acmon, and Papilio rutulus) are polyphagous. Two of these are
clearly colonizers. Most multivoltines have to change hosts
during the season, but for many (Pieris on Crucifers, Colias
eurytheme and Everes comyntas on Legumes) a seasonal suc-
cession of wild hosts of one family is available even in the
absence of agriculture.

116

ARTHUR M. SHAPIRO

J. Res. Lepid.

These data thus tend to bear out Opler and Buckett’s sug-
gestion. Univoltinism may result from other selection pressures
besides host phenology, of course; perhaps it is better to say
that both host specialization and voltinism are apt to be related
to interspecific competition. Differences in phenology in differ-
ent parts of the ranges of species may offer clues to selective
factors acting on the timing of life history phenomena. Lerodea
etifala, for example, is found only from June to early winter in
both northern and southern California (Emmel and Emmel,
1973). Yet in the southeastern United States it flies all year,
being recorded as early as ii.9 in coastal Georgia (Harris, 1972).
Brephidium exilis flies nearly all year in southern California,
but not before June in the Valley or at Suisun Bay; Opler and
Langston (1968) record it as early as April in Contra Costa
County, as does Tilden (1965a) for the San Francisco Bay area.
In this case, however, it is not certain that B. exilis overwinters
successfully inland at the latitude of Sacramento.

THE VALLEY AS A FAUNAL BARRIER
The Sacramento Valley is about 35 miles wide at the latitude
of Davis. Across this expanse of flat land the habitats and plants
of the foothills are absent, and many characteristic foothill butter-
flies (such as Papilio eurijmedon Lucas, Chlostjne palla Bois-
duval, Satyriiim saepium Boisduval, Lijcaena gorgon Boisduval,
Callophrijs diimetorum Boisduval, Thorijhes pijlades Scudder)
disappear as well. The data on the pristine vegetation of the
Valley (Thompson, 1961) suggest that this barrier may have
been much less dramatic in pre-American times. A few isolated
pockets of foothill vegetation still exist in the Valley (e.g. Buck-
eye, Aescidus calif ornica (Spach) Nutt., and Pipevine, Aristo-
lochia calif ornica Torr. on the north fork of Putah Creek near
BartheFs Ranch, southeast of Davis). Along the intermittent
streams such plants must have come down some distance below
the mouths of the canyons, and they may have formed an ele-
ment of the riparian forest on the permanent streams. Because
of changes in the habitats, it is at present impossible to evaluate
the historical role of the Valley as a barrier to dispersal between
the Sierras and coast ranges. The butterfly faunas at comparable
elevations are strongly similar across it, though at least three
species (Satijriiini saepium, S. auretorum, Chlosyne palla) are
phenotypically distinguishable in the two ranges. At least one
species, Poanes melane, was apparently continuously distributed
across the Valley in riparian habitats but is now restricted to
the two systems of foothills and the major riparian habitats

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

117

around Sacramento; it occurred at Davis ( Willowbank? ) as late
as 1947 (Heppner, 1973). The present distribution of Glauco-
psijche lygdanms hehrii is very similar.

Eight or nine species of foothill butterflies enter the Valley
more or less regularly but do not breed there. Nymphalis cali-
fornica migrates regularly across the Valley floor ( Shapiro,
1973); Limenitis hredowii calif ornica, another strong flyer, is
frequently seen at Davis. It is very unlikely that either of these
species ever breeds in the Valley. In contrast, two relatively
weak-flying Lycaenids, Incisalia iroides and Lycaenopsis argioliis
echo, enter the Valley along the streams and have been recorded
so often that both are suspected of breeding, although no known
hosts are available.

A valid study of the effectiveness of the Valley as a barrier
to faunal exchange must await a survey of the isolated Sutter
Buttes in the mid-north Valley, planned for the next two seasons.

ENDEMISM AND EXTINCTION

The Willow Slough population of Phyciodes campestris is
the most distinctive Valley endemic. Valley populations of
Everes comyntas and Limenitis lorquini are phenotypically
weakly distinguished, and Satyrium californica has an unusual
host {Quercus lobata). On the whole, hovyever, the Valley
fauna is strikingly lacking in endemic populations; it is com-
posed largely of wide-ranging, vagile species with little tendency
toward geographic variation.

It seems inevitable that more distinetive butterflies must have
existed in the pristine Valley, with its well-developed plant
communities of considerable antiquity. The Hesperiidae in par-
ticular are lacking in the modern fauna; there is no Valley
Hesperia of any kind, for example. It is conceivable that relict
populations of undescribed Valley endemics may still exist in
isolated or unexplored areas which have escaped intensive
modification by man. The recent discovery of Ochlodes yuma
in southern Sacramento County indicates that the list of Valley
butterflies may not yet be complete. It is unlikely, however, that
we will ever be able to reconstruct — or even guess intelligently —
the composition of the pre-American Valley fauna.

FAUNAL LIST

Host plants are based on actual oviposition or larval records
in the Sacramento Valley only. Botanical nomenclature is from

118

ARTHUR M. SHAPIRO

]. Res. Lepid.

Munz (1970). Butterfly nomenclature is mostly as in Ehrlich
and Ehrlich (1961) and skippers mostly as in dos Passos (1964).
All records are for Yolo, Solano, and Sacramento Counties only.
Flight periods for the 1972 and 1973 seasons are given in Tables
7 and 8 on a weekly basis.

Family Daiiaidae

1. Danaus plexippus Linnaeus. Monarch.

Common throughout. Four broods in both 1972 and 1973. Open coun-
try, especially along sloughs, ditches, and railroads with the host plants.
No overwintering occurs in the Valley. The spring immigration from the
west is more conspicuous than the autumn emigration. Tagged specimens
recovered in the Valley and the Vacas indicate that a least some of our
population overwinters in Marin County. Host Plants: Asclepiadaceae;
Asclepias fascicularis Dene.

Family Satyridae

2. Coenonympha tullia California Westwood and Hewitson. California
Ringlet.

Frequent but spotty, usually found in the Sacramento and American
River floodplains, but not restricted to them. Throughout the season, as in
the foothills; probably two broods, the second very protracted. Although
breeding has not been confirmed in the Valley, this species is very probably
resident. Generally absent from the heaviest clay soil areas.

Family Helicoiiiidae

3. Agraulis vanillae Linnaeus. Gulf Fritillary.

A rare stray. Since this species bred in the San Francisco Bay area in
the 1960s it is perhaps capable of colonizing the Valley, at least tempor-
arily. The normal hosts, Passifloraceae: Passiflora spp., are occasionally
cultivated but there are no native species here.

Family Nymphalidae

4. Speyeria callippe Boisduval. Callippe Fritillary.

A rare stray; one male, fresh, v.9.72, northwest Davis. It is not known
what subspecies or source area this individual represents. (Note: Speyeria
coronis Behr has been taken at Suisun Bay (ix.23.72) and could stray into
the Valley on the west side. )

5. Euphydryas chalcedona Doubleday and Hewitson. Common Checker-
spot.

Occasional in floodplains, as at the American River and Putah Creek,
probably as a stray from the foothills where it is abundant. No larvae have
been collected in the Valley, where the usual host, Scrophulariaceae: Di-
placus, does not occur.

6. Chlosyne lacinia crocale Edwards. Bordered Patch.

This species was not seen in 1972, but in 1973 about 15 were taken on
the American River floodplain at Camp Pollock. These included all three
color forms illustrated by Emmel and Emmel (1973). There were at least
two broods. The Bordered Patch is indigenous to southeastern California,
in the Imperial Valley and Colorado Desert, and seems to be unreported
even as a stray anywhere north of the Transverse Ranges. Its future history
in the Valley should be very interesting. Host Plant: The usual host, Com-
positae: Helianthus annuus L., is abundant on the floodplain.

7. Phyciodes campestris Behr. Field Crescent.

Very local. The Sacramento Valley populations differ from others seen
in the smaller size and heavier infuscation dorsally (illustrated in Shapiro,
1974b). There is a large, vigorous colony along Willow Slough from north-
west of Davis to the Yolo Bypass, and smaller colonies at north Woodland,
Stone Lake-Beach Lake, and North Sacramento; strays to Davis, Willow-
bank, West Sacramento, mainly in autumn. Three broods. To be sought in

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

119

sandy bottomlands with good stands of the host. Host Plant-. Compositae:
Aster chilensis Nees. I took a female investigating Aster novi-belgii h., an
eastern species, in a garden at Davis.

8. Phyciodes mylitta Edwards. Mylitta Crescent.

Common everywhere; four broods. Larva overwinters, feeding up in
February-early March, host Plants: Compositae: Silybum marianum (L. )
Gaertn.; Cirsium californicum Gray; C. vulgare (Savi) Tenore; Carduus
pycnocephalus L. Centaurea solstitialis L. is suspected.

9. Polygonia satyrus Edwards. Satyr Anglewing.

Uncommon, recorded mostly as singletons. Riparian forest (Southport,
Sacramento River; Northgate, Sacramento State University area, both Amer-
ican River); once in Davis. Not confirmed to breed in the Valley, but the
usual host, Urticaceae; Urtica, is common in the places where it has been
found.

10. Nymphalis californica Boisduval. California Tortoiseshell.

Frequent as a transient in spring and fall, but not known to breed in
the Valley where its usual hosts Rhamnaceae: Ceanothus do not occur.
Completely absent during the hottest weather, and not appearing in spring
until several weeks after it begins flying in numbers in the Vacas. In 1972
a significant west-to-east migration across the Valley occurred, beginning
about v.26; in 1973 the same phenomenon occurred vi.8-11. The return,
east-to-west movement was noticeable in Davis on x.l-x.3.73. (Note: Nym-
phalis milberti Latreille was taken at about 1000 feet in the Spenceville
Recreation Area, Yuba Co., iv.11.73, and may reach the Valley floor on oc-
casion northward. )

11. Nymphalis antiopa Linnaeus. Mourning Cloak.

Frequent to common in bottomlands and riparian forest; occasional
elsewhere. Apparently one brood, becoming dormant by August and not fly-
ing again until February. Larvae colonial and often locally conspicuous.
Host Plants: Salicaceae: Salix lasiandra Benth.; not seen on S. hindsiana
Benth., the commonest willow in the Valley, but reportedly damaging S.
Babylonica L. in gardens in Sacramento. The seeming avoidance of S.
hindsiana may reflect its being widely utilized by larvae of the diurnal Sa-
turniid Pseudohazis eglanterina nuttalli Strecker at the same time.

12. Vanessa atalanta Linnaeus. Red Admiral.

Common in the Sacramento-American River floodplains, and frequent
throughout. Probably three broods. Host Plants: Urticaceae: Urtica holoseri-
cea Nutt., a common understory plant of riparian forest and thickets, absent
in other habitats; also Soleirolia (Helxine) soleirolii (Baby’s Tears), an in-
troduced Urticaceous ground cover, in Davis.

13. Vanessa (Cynthia) virginiensis Drury. Painted Beauty.

Frequent in riparian forest, forest edges and nearby roadsides; rare
elsewhere; the least common Vanessa, seldom seen before midsummer. Ap-
parently four broods. Host Plants: Compositae: Gnaphalium palustre Nutt,
(at Southport), probably other Gnaphalium.

14. Vanessa (Cynthia) cardui Linnaeus. Painted Lady.

Common and general. Immigrant, rearing three broods in 1972. 1973
was an outstanding year for this species, which flew for 44 weeks and bred
in great numbers. Its migrations and variation are described in Shapiro
(1974c). Host Plants: Extremely polyphagous, though in normal years
mostly restricted to the plants marked Compositae: Silybum marianum
(L. ) Gaertn.^; Cynara scolymus L. (artichoke); Carduus pycnocephalus
L*; Cirsium vulgare (Savi) Tenore*; Xanthium strumarium L. var. cana-
dense (Mill.) T. & G. (West Sacramento, v.26. 73, larvae); Centaurea sol-
stitialis L.; Helianthus annuus L. (Davis, vii.6.73, larvae common);,
glabra Gray; Leguminosae: Lupinus bicolor Lindl. (oviposition near Winters,
iii.18.72); Boraginaceae: Amsinckia douglasiana A. DC.*; Malvaceae: Mal-
va parviflora L.*, M. neglecta Wallr.*; M. nicaeensis All.*; Althaea rosea
L.; Sida hederacea (Dough) Torr. (larvae all season, Davis, 1973); Urti-
caceae: Urtica urens L. (Woodland Sinks, larva, v.13.73); Plantaginaceae:

120

ARTHUR M. SHAPIRO

J. Res. Lepid.

Plantago lanceolata L. (Putah Creek, v.16.73, 3 larvae); Hydrophyllaceae:
Eriodictyon californicum (H, & A. ) Torr. (Putah Creek, vi. 15.73; also seen
in Amador Co. ) .

15. Vanessa (Cynthia) annabella Field. West Coast Lady.

Abundant, general, flying all year. Five to six broods. For figures and
discussion of variation see Shapiro, 1974d. Host Plants: Malvaceae: Malva
parviflora L., neglecta Wallr., and nicaeensis All.; Althaea rosea L.; Sida
hederaeea (Dough) Torr. (rarely).

16. Precis coenia Hubner. Buckeye.

Common to abundant in bottomlands and on sandy soils; frequent to
common elsewhere. Restricted to bottomlands early in the season, but gen-
erally distributed by late vi. It is not certain that P. coenia overwinters in
the Valley at all. It is abundant in footliill canyons (where it feeds on Scro-
phulariaceae, especially Diplacus) 3-6 weeks before it appears on the Valley
floor. After late viii specimens, especially females, are very red or purplish
beneath; but some clay-colored butterflies reappear after mid xi and until
hard frost. The February butterflies in the canyons are clay-colored and
undersized, and seem to be a mixture of fresh and worn individuals. Host
Plants: Verbenaceae: Lippia lanceolata Michx. and (in cultivation) L.
nodiflora Michx. vars. canescens (HBK.) Kuntze and rosea (D. Don) Munz;
Plantaginaceae: Plantago lanceolata L. (at Vacaville, in lawns). Tilden’s
speculation (Tilden, 1971) that true P. coenia does not feed on Lippia is
clearly in error; more than 100 wild larvae were collected from this plant at
Davis and reared in 1973, and larvae have also been found on it at Fair-
field.

17. Limenitis lorquini Boisduval. Lorquin’s Admiral.

Common in riparian forest and among young willows along creeks,
sloughs, and levees; very rare elsewhere, once seen on the University of
California campus at Davis. Three broods. Valley specimens are, on the aver-
age, less orange and more bluish beneath than foothill ones. Host Plant:
Salicaceae: Salix lasiandra Benth.; not seen on S. hindsiana Benth.

18. Limenitis hredowii californica Butier. California Sister.

Infrequent and sporadic. Not known to breed in the Valley, although
acceptable oaks are locally available especially near the foothills. Probably
formerly resident when the oak-Buckeye association extended farther down
the streams.

Family Lycaeiiidae

19. Atlides halesus Hubner. Great Blue Hairstreak.

Common in riparian forest; frequent to common in residential neigh-
borhoods with established deciduous trees; rare elsewhere. Three (perhaps
a rudimentary fourth) broods. Host Plant: Loranthaceae: Phoradendron
flavescens (Pursh.) Nutt. var. macrophyllum Engelm., typically on Cotton-
wood along the rivers, on various other deciduous trees elsewhere.

20. Strymon melinus pudica Henry Edwards. Common Hairstreak.

Abundant throughout, breeding continuously; about five broods. A few

specimens lack the postmedian line and shading on the forewing ventrally.
One female with the orange completely replaced by white, Davis, vi.3.73.
Host Plants: Malvaceae: Malva nicaeensis All. and neglecta Wallr.; Sida
hederaeea (Dough) Torr. (preferred). Euphorbiaceae: Eremocarpus
setigerus (Hook.) Benth. suspected.

21. Satyrium californica Edwards. California Hairstreak.

Frequent and apparently breeding along the American and Sacramento
Rivers, in relict stands of Valley Oak. Otherwise unrecorded except as strays
along the edges of the Valley. Sacramento specimens are slightly smaller and
darker than foothill ones. One brood. Host Plant: Fagaceae: Quercus lobata
Nee. strongly suspected; no other likely host occurs in the river bottoms.

22. Satyrium sylvinus Boisduval. Willow Hairstreak.

Extremely abundant among young gray willows on floodplains and along
sloughs and levees; less common in mature riparian forest; rare or absent

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

121

elsewhere. Variable in the spotting of the lower wing surfaces and in the
extent of orange above. Apparently always fully tailed. Host Plant: Salica-
ceae: Salix hindsiana Benth., the common sandbar willow.

23. Satyrium auretorum Boisduval. Gold-Hunter’s Hairstreak.

Six males and one female taken on the Sacramento River at Elkhorn
Ferry, v. 12.73 in mixed riparian forest on horehound flowers. These speci-
mens are not phenotypically distinguishable from Vaca Mountains individ-
uals.

24. Incisalia iroides Boisduval. Western Brown Elfin.

Occasional in riparian forest and as a stray down the creeks from the
foothills. Not known to breed in the Valley, and none of the recorded hosts
is present in spring in the riparian forest, but up to 6 have been taken in one
day. (Note: The Bramble Hairstreak, Callophrys dumetorum Boisduval, is
common in the foothills on both sides of the Valley but is unrecorded on
the Valley floor although both Polygonaceae: Eriogonum and Leguminosae:
Lotus scoparius ( Nutt. ) Ottley occur there. )

25. Lycaena xanthoides Boisduval. Great Copper.

Common to locally abundant along sloughs and ditches, in marshes and
bottomlands, even in alkali marshes. Occasional on roadsides and in agricul-
tural land. One brood in late spring. Females variable, with heavy to light
spotting and much to little yellow flush above. Host Plants: Polygonaceae;
Rumex crispus L., R. conglomeratus Murr., probably other Rumex.

26. Lycaena helloides Boisduval. Purplish Copper.

Abundant throughout; damp places, marshes, sloughs, ditches, alkali
lands, dry waste ground with compacted clay soils, etc. Five broods, all
season. Variable, especially the females; an albino female at Putah Creek,
iv.17.72. Host Plants: Polygonaceae: Polygonum aviculare L. (dry places);
P. lapathifolium L., P. persicaria L., P. punctatum Ell., Rumex crispus L.
( all in wet places ) .

27. Hemiargus isola Reakirt. Reakirt’s Blue.

Locally frequent to common, multiple brooded, waste ground near the
American River. Host Plants: Leguminosae: Melilotus alba Desr.; Medicago
satin a L.

28. Leptotes marina Reakirt. Marine Blue.

Locally common to abundant, multiple brooded, railroad and highway
embankments and thickets near the American River, and occasional else-
where. Perhaps not a permanent resident; much commoner in 1973 than
1972. Host Plants: Leguminosae: Glycyrrhiza lepidota Pursh.; perhaps also
Melilotus alba Desr.

29. Brephidium exilis Boisduval. Western Pygmy Blue.

Extremely abundant on alkaline soils and in alkaline or saline marshes;
common to abundant on waste ground with the hosts; occasional elsewhere.
Probably does not overwinter in most or all of the Valley, but this needs
to be confirmed. About four broods beginning in early summer, commonest
after mid-August. Host Plants: Chenopodiaceae: Atriplex hastata (L.) Hall.
& Clem.; A. rosea L.: A. serenana A. Nels.; A. coronata Wats.; A. cordulata
Jeps.; A. semibaccata R. Br.; Suaeda fruticosa (L.) Forsk.; Salsola kali L.
var. tenuifolia Tausch.; probably other Chenopods.

30. Everes corny ntas Godart. Eastern Tailed Blue.

Common along ditches and sloughs and in bottomlands; occasional in
moist waste ground and lawns. Five broods; the first with females strongly,
the last with them more or less blue-tinted. The Sacramento Valley popu-
lations seem phenotypically distinct from others, so it is unlikely that
they represent an introduction as has been suggested. Host Plants: Legumi-
nosae: Lotus purshianus (Benth.) Clem. & Clem.; Vida villosa Roth; V.
satina L.; V. cracca L.; V. angustifolia Reich.; Lathyrus jepsonii Greene ssp.
calif ornicus (Wats.) Hitchc.; Trifolium tridentatum LindL; Lupinus nanus
Dough ssp. latifolius (Benth.) D. Dunn.

31. Plebeius acmon Westwood and Hewitson. Acmon Blue.

Common to locally abundant; waste ground, ditches, roadsides, levees.

122

ARTHUR M. SHAPIRO

/. Res. Lepid.

creek bottoms; occasional everywhere. Five broods, the first of form cottlei
Grinnell with blue females, the last with transitional males and largely black
females. Host Plants: Leguminosae: Lotus purshianus (Benth. ) Clem. &
Clem.; L. scoparius (Nutt.) Ottley; Melilotus alba Desr.; Polygonaceae:
Polygonum aviculare L.; Eriogonum gracillimum Wats, (sandbars, Brod-
erick ) .

32. Plebeius icarioides Boisduval. BoisduvaPs Blue.

A single, very worn male. West Sacramento, v.5.73, almost certainly a
stray, but too battered to be determined to subspecies.

33. Glaucopsyche lygdamus behrii Edwards. Behr’s Silvery Blue.

Locally abundant with Lathyrus jepsonii along the rivers; infrequent
to common in creek bottoms and levee thickets throughout the Valley.
Not seen elsewhere. Host Plants: Lathyrus iepsonii Creene ssp. californicus
(Wats.) Hitchc. In the Vacas also recorded on Vida sativa L., V. villosa
Roth, and Lupinus succulentus Dough, all of which also occur in the Valley.

34. Lycaenopsis argiolus echo Edwards. Echo Blue.

Infrequent in the American River bottomlands; entering the Valley
elsewhere as a stray from the foothills. Breeding not established; the re-
corded host plants ( Hippocastanaceae: Aesculus; Ericaceae: Arctostaphy-
los) are generally absent in the Valley.

Family Papilionidae

35. Battus philenor Linnaeus. Pipevine Swallowtail.

Frequent to locally common along the creeks near the host plant;
occasional in mature riparian forest, and straying widely. One brood at
Putah Creek, but perhaps partially four-brooded in Carmichael. Host Plant:
Aristolochiaceae: Aristolochia calif ornica Torr. Sometimes shows a “false
brood” in very late fall or very early spring.

36. Papilio zelicaon Lucas. Anise Swallowtail.

Frequent to locally abundant on sandy floodplains as at Southport,
West Sacramento, and at the American River; occasional throughout. Host
Plant: Umbelliferae: Foeniculum vulgar e Mill. Curiously unrecorded from
other common Umbellifers, such as Daucus pusillus Michx.

37. Papilio rutulus Lucas. Western Tiger Swallowtail.

Common in riparian forest, orchards, and older city neighborhoods;
occasional throughout. Three broods. Host Plants: Oleaceae: Fraxinus spp.
(larvae, Davis); Syringa vulgaris L. (ova, Davis vii.27.73); Rosaceae; Pru-
nus caroliniana Ait. (larva, Sacramento). Probably other trees {Populus?
Salix? Platanus? other Prunus? )

38. Papilio multicaudatus Kirby. Two-tailed Swallowtail.

Frequent in riparian forest; apparently two-brooded. Host unrecorded;
ash (Oleaceae: Fraxinus latifolia Benth.) or Prunus spp. (Rosaceae) sus-
pected.

Family Pieridae

39. Pieris rapae Linnaeus. European Cabbage Butterfly.

Abundant everywhere; six broods, flying 42 weeks in 1972 and 49
weeks in 1973. The spring brood is heavily dusted beneath, with reduced
dark markings above. A mosaic gynandromorph taken at Davis, iii.3.72,
and another bred, iv.22.72. Host Plants: Cruciferae: Brassica nigra (L)
Koch; B. kaber (DC.) Wheeler, B. oleracea L. cultivars., B. geniculata
(Desf. ) J. Ball.; Cardaria draba (L.) Desv.; Sisymbrium officinale (L.)
Scop.; S. irio L.; S. altissimum L.; Raphanus sativus L.; Lepidium latifoli-
um L.

40. Pieris protodice Boisduval and LeConte. Checkered White.

Common to locally abundant on sandy soils, especially on the Ameri-
can River floodplain; generally rare in clay areas, most widespread in viii-x.
Not overwintering in most of the Valley, but colonizing widely in summer.
Five or six broods, the first of form vernalis Edwards, the last transitional
to it. The early spring brood is very local but may Idc common where it
does occur. Host Plants: Cruciferae: Brassica geniculata (Desf.) J. Ball
and Lepidium latifolium L. preferred; rarely on Brassica nigra ( L. ) Koch.,
(Continued on page 137)

Journal of Research on the Lepidoptera

13:123-130, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A NEW SPECIES OF ORMISCODES
(DIRPHIELLA) FROM MEXICO
(SATURNIIDAE: HEMILEUCINAE)

JULIAN P. DONAHUE^ and CLAUDE LEMAIRE^

Until now, the subgenus Dirphiella Michener contained only
a single species. The second species, described here, differs re-
markably in general appearance from O. (D.) alhofasciata, al-
though the two are structurally very similar. The females of both
species are unknown.

Genus ORMISCODES Blanchard, 1852:61

Subgenus DIRPHIELLA Michener, 1949a: 130 (nomen
nudum! ); 1949b: 146. Type: Dirphiopsis albofasciata Johnson
and Michener, 1948:11. Monobasic.

ORMISCODES (DIRPHIELLA) TAYLORI Donahue and
Lemaire, new species
(figures 1 and 2)

DIAGNOSIS

General appearance entirely different from that of O. (D.)
albofasciata (figure 3), the only other member of this subgenus.
O. taylori is easily distinguished from albofasciata by the yellow-
ish brown ground color (dark brownish gray in albofasciata),
completely yellow antennae (orange-yellow shaft and blackish
rami in albofasciata), conspicuous, straight, double black and
white postmedian band on forewing, from inner margin to apex
(all white and sigmoid in albofasciata, from inner margin to
costa), forewing apex acute (rounded in albofasciata), absence
of color bands on dorsum of abdomen (present in albofasciata) ,
presence of epiphysis (absent in albofasciata) , and presence of
apical cornutus on vesica of male genitalia (absent in albofasci-
ata). In general appearance taylori is perhaps more reminiscent
of members of Dirphia ( subgenus Dirphiopsis ) than of any other
species of Ormiscodes.

^Natural History Museum of Los Angeles County, 900 Exposition Boulevard,
Los Angeles, California 90007, U.S.A.

M2 Boulevard Victor Hugo, 92200 Neuilly s/ Seine, France

123

124

DONOHUE AND LEMAIRE

/. Bes. Lepid.

DESCRIPTION: Holotype male (fig. 2)

Head: Antenna pale yellow, scape clothed with small appressed scales,
dark brown dorsally and paler ventrally; flagellum (37 segments) bipectin-
ate to apex, rami with 2 or 3 terminal and subterminal bristles, the longest
one 3 times as long as setae; ventral antennal cones conspicuous only on
terminal 12 segments, increasing in prominence distally; each segment with
2-5 midventral setae, most numerous on distal segments, and 1 or 2 mid-
dorsal setae on proximal 15 segments. Eye large, extending from antennal
socket to ventral edge of frons, eye height three times shortest interocular
distance. Labial palpi with blackish brown, appressed, hair-like scales
which project ventrally. Vestiture of frons hair-like, blackish brown and
erect dorsally, ventrally becoming almost black, projecting ventrad and
curving slightly mesad. Vestiture of vertex consists of loose, spreading,
grayish brown hair-like scales.

Thorax: Vestiture of loose, spreading, hair-like scales as follows:
patagium anteriorly narrowly edged pink, centrally grayish brown, becom-
ing darker laterally to blend with blackish brown of wing base; mesonotum
with appressed brown hair-like scales, overlain by looser, paler scales.
Mesoscutellum and metanotum with long, deep pink scales, extending pos-
teriorly over first two abdominal segments. Tegula pale gray-brown. Venter
of thorax densely clothed with hair-like scales, blackish brown between
forecoxae and in anterior axillary region, deep pink elsewhere including a
narrow longitudinal fringe parallel to and at the base of the posterior
half of the hindwing. Legs: brown dorsally, paler ventrally and posteriorly,
pink ventral fringe on femora and brown fringe on tibiae (posterior on
foreleg, dorsal on mid- and hindlegs); foreleg femoral fringe pale pink, the
hair-like scales paler at base; epiphysis large (nearly 1/2 tibial length),
“J”-shaped, the apex projecting posterad, with pale yellowish white tuft
internally and apically; first two tarsal segments with tapering dorso-
posterior fringe; midleg as in foreleg but femoral fringe darker pink; ex-
ternal (ventral) surface of tibial spurs clothed with appressed pale brown
scales; basitarsus with very weak dorsal fringe; hindleg identical to midleg
except basitarsal fringe more strongly developed.

Abdomen: Dorsum of first segment clothed with long, loose, pink hair-
like scales which overlay and conceal two narrow, transverse, subdorsal
patches of short blackish brown scales on the posterior margin of the seg-
ment; dorsum of segments 2-7 uniformly clothed with small, spatulate,
medium gray, appressed scales, intermixed with and almost totally con-
cealed by much longer brownish black, semi-erect hair-like scales; dorsum
of segment 8 and genitalia with tuft of loose, pink hair-like scales. Dorsal
brownish black ground color exending ventrad to clothe pleural area, inter-
rupted in the posterior fourth of each segment by a tuft of pink scales.
Ventral vestiture of appressed hair-like scales yellowish brown, segments
2-7 with mesal grayish brown line.

Wings: General: ground color of both surfaces of wings yellowish
brown, slightly paler on hindwing upperside; fringes paler; veins brownish
black except across the white postmedial ( PM ) band on forewing upper-
side; discocellular spots absent on upperside, present on underside as nar-
row brownish black lines on R4 5-Mi crossvein of forewing and on anterior
third of M2-M3 crossvein of hindwing. Forewing upperside: antemedial
(AM) line evanescent, a faint blackish shade arising at one fourth on inner
margin, extending in nearly straight line, more or less parallel to outer
margin, to the anterior margin of discal cell at a point approximately 1 mm
proximad of origin of vein Ri, then angling about 90° and continuing to the
costa as a broader, more diffuse but darker shade, its course indicated by
further darkening of veins where it crosses anterior edge of cell, vein Sc,
and the costa; AM line bordered externally by a narrow, diffuse white
shade, which expands abruptly at the anterior edge of the discal cell to

13;123-130, 1974

NEW ORMISCODES

125

form an irregular, squarish, poorly defined whitish spot filling the area
between the discal cell and costa, extending from the AM shade approxi-
mately to the origin of vein Ri, the distal edge more or less perpendicular
to costa. Area basad of tlie AM line is slightly darker between the costa and
the posterior margin of the discal cell. Postmedial ( PM ) band conspicuous,
oblique, blackish, about 1 mm wide, extending from inner margin at about
three-fifths to apex at end of vein Rs; almost straight, but most notably off-
set outward in cell R2-R3; PM band bordered internally by a white band
of similar width along its entire length, and bordered externally by a broad,
obscure, fuscous shade, broadest at the inner margin, extending from tornus
to vein R3, where it fuses with the PM band; outer margin of this shade
dentate: concave on veins, convex in cells, the teeth strongest in cell Cui-
Cu2 and becoming weaker anteriorly. Costal margin dark brown from base
to the white subcostal spot, concolorous with ground color beyond the
spot. Hindwing upperside: postmedial band black, more or less parallel to
outer margin, extending from inner margin at about three-fifths (where it
widens on the margin itself) to the costa at seven-eighths, external edge
diffuse, internally bordered by a narrower, diffuse whitish shade. Ground
color basad of PM band slightly paler than area distad of PM band. Under-
side of both wings similar to upperside, except that the ground color of
both wings is uniformly yellowish brown and the markings are less pro-
nounced. Forewing: AM line and subcostal white spot absent, PM band
weaker and more diffuse than on upperside, with internal white edging
only faintly indicated and the distal shade absent; base of forewing with
diffuse blackish shade corresponding to the weaker shade on upperside.
Hindwing: compared to upperside, PM band slightly wider and more
diffuse, internal white edge broader and more conspicuous; PM band
weakly sigmoid, incurved between veins 2A and Cui, excurved and more
or less parallel to outer margin between Cui and costa. Costa whitish from
base to PM band. Subterminal band extremely faintly indicated, parallel
to outer margin and slightly nearer it than to PM band.

Genitalia (fig. 3): almost identical to that of Ormiscodes (Dirphiella)
albofasciata (as figured by Michener, 1952:488, figs. 328-332), with the
conspicuous exception that the apex of the vesica bears a small, stout
cornutus which that species lacks. In addition, the dorsal lobes of the
valva appear to be slightly longer and basally broader in taylori than in
albofasciata.

Size: forewing length 31mm.

Female: unknown.

Variation: the paratype differs from the holotype in minor details, as
follows: smaller (forewing 28mm long); antennae with more flagellar seg-
ments (38 right, 39 left); proximal 20 (right) or 21 (left) segments with
dorsal setae; markings and ground color of forewing upperside slightly
darker: AM line more pronounced and the dentate shade distad of PM
band much more conspicuous; underside of both wings darker: forewing
with darker basal anterior shading, and the shade distad of PM band evi-
dent (as pronounced as on upperside of holotype); hindwing with sub-
terminal band stronger, extending from vein 2A to vein R .

Additional material: excluded from the type series, but representing
this or a very closely related species, is a slightly rubbed male in the
LACM collection from 24 miles south of Valle Nacional, Oaxaca, Mexico,
elevation 5,600 feet, 24-25 July 1970 (E. Fisher and P. Sullivan). This
locality is approximately 120 air miles south-southeast of the type locality
of taylori. The Oaxacan male bears a very close external resemblance to
the type specimens, and may represent individual or geographic variation-—
a point that can be resolved only after the examinaion of additional ma-
terial. This specimen differs from typical taylori as follows: slightly larger
(forewing length 32.5mm), antenna with 42 segments, only the basal 10 with

126

DONOHUE AND LEMAIRE

J. Res. Lepid.

single dorsal setae. Discocellular scaling conspicuous on upperside of both
wings, and more pronounced on underside than in taylori. Forewing upper-
side with AM line placed farther distad, entering discal cell at vein Cus then
angling across to origin of vein Ri where it angles proximad to costa; white
external edge of AM line broader, more conspicuous and more extensive,
extending to vein 2A, almost completely filling the triangular portion of
discal cell distad of AM line, and expanding to form the whitish costal spot
farther distad than in taylori, the spot extending from origin of vein Ri
halfway to fork of R2 and R3; PM band directed to a point on costa before
apex, then angling on vein Mi and continuing, more or less straight, to a
point just below apex. Forewing underside lacking fuscous suffusion in
discal cell, PM band as above except angled at vein R4 5. Fronto-clypeal
protuberance produced into a truncate, triangular projection. There are
also several slight differences in the shape of the uncus, gnathos, transtilla,
and lobes of the valva of the male genitalia.

TYPES: HOLOTYPE S , 1 mile south of Pueblo Calcahualco,
8 road miles west of Coscomatepec, Veracruz, MEXICO; eleva-
tion 6,200 feet, 26 July 1972 (Terry W. Taylor); collected in the
rain at a 15-watt fluorescent black light, between 10:00 and 10:30
p.m. Central Daylight Time, ambient temperature 45 °F. (C.
Lemaire genitalic preparation no. 2637, in glycerine).

PARATYPE, 1 $ , same locality as holotype, 21 July 1973 (Terry

Taylor and Roy R. Snelling); collected at a mercury vapor
light between 10:00 and 11:00 p.m. Central Daylight Time, am-
bient temperature 59 °F. Holotype and paratype deposited in
the Natural History Museum of Los Angeles County, through the
courtesy of Terry W. Taylor.

DISCUSSION

Michener erected the subgenus Dirphiella for the lone species
Dirphiopsis alhofasciata Johnson & Michener, then known from
a single male from Chiapas, Mexico (“. . . presumed to be from
the vicinity of Comitan.'’). It is unique among the nine sub-
genera of Ormiscodes in that the male antenna is bipectinate in-
stead of quadripectinate. Michener (1952:445-446) later ex-
panded the definition of this subgenus in his revision of the
higher categories of the New World Saturniidae.

One of us (JPD) has examined the type specimen of D. alho-
fasciata, and found that the original description is defective with
respect to several antennal characters, as follows: the antennal
shaft and extreme bases of the rami are orange-yellow, conti'ast-
ing sharply with the brownish black of the rami (not “antennal
shafts brown, rami black”); each antennal segment has two

13:123-130, 1974

NEW ORMISCODES

127

stout, subapical, para-medial setae on the ventral side, plus nu-
merous scattered, small, brown lateral setae which contrast with
the color of the shaft; mid-dorsal setae not evident (not "seg-
ments of antennal shaft of male without setae or with a single
dorsal subapical seta” ) ; terminal bristle of rami about three times
as long as longest setae, subterminal bristle present on some rami
(not "terminal bristles of rami nearly twice as long as setae”).
There are 39 segments in each antennal shaft. Unfortunately, it
was not possible to verify the absence of the epiphysis, as the
forelegs are represented by a single femur, and a patch of scales
on the locality label, where one foreleg had apparently been
glued and subsequently fallen oflF.

However, the junior author has verified the absence of the
epiphysis in several specimens of albofasciata (all males) in his
collection from several localities in Chiapas. These specimens
indicate that albofasciata is very variable in color and macula-
tion. The ground color of the upper surface of the wings varies
from pale to deep black; the postmedian bands ( sometimes part-
ly or even entirely absent) vary in color from pure white to
orange yellow; in specimens with the orange-yellow bands the
median area of the forewing and the postmedian ( and sometimes
the median) area of the hindwing are more or less suffused with
the same orange-yellow color or with yellowish brown; the width
of the fringes varies, as does their white or yellow color, and
there are generally black dots in the fringe at the ends of the
veins.

With the discovery of this second species of Dirphiella it is
necessary to modify the definition of the subgenus, as some of
the original characters, as amended above, now appear to be of
only specific significance. These characters as they occur in
taylori, and their corresponding state in albofasciata, are as fol-
lows: epiphysis present in male (absent in albofasciata); anten-
nal cones conspicuous only on terminal 12 or fewer segments
(recognizable nearly to base in albofasciata) ; eyes large, height
of eye three times shortest distance between eyes (subequal in
albofasciata ) .

These differences, plus the entirely different facies, would
appear to be sufficient to warrant the erection of a new subgenus
for taylori were it not for the extreme similarity in genitalia.

We take pleasure in naming this species in honor of Terry
W. Taylor, who collected the type series and brought them to
our attention. We are further indebted to Charles L. Hogue

128

DONOHUE AND LEMAIRE

J. Res. Lepid.

Fig. 1. — Ormiscodes (Dirphiella) taylori, genitalia of male holotype: a. —
ventral view, aedeagus removed; b. — aedeagus, lateral view.

13:123-130, 1974

NEW ORMISCODES

129

for his valuable comments and suggestions on the manuscript,
and to Frederick H. Rindge of the American Museum of Natural
History, who kindly loaned the type specimen of Dirphiopsis
alhofasciata.

LITERATURE CITED

BLANCHARD, EMILE. 1852. IN Claude Gay, Histona Fisica tj Politica de
Chile. Zoologia, Vol. 7. Lepidoptera; 1-112.

JOHNSON, FRANK and CHARLES D. MICHENER. 1948. New Neo-
tropical saturnioid moths (Lepidoptera). American Mus, Novit. no.
1372: 1-15, 16 figs.

MICHENER, CHARLES D. 1949a. Parallelisms in the evolution of the
saturniid moths. Evolution 3: 129-141, 8 figs.

— . 1949b. New genera and subgenera of Saturniidae (Lepidoptera).

J. Kansas Entomol^^ Soc. 22: 142-147.

. 1952. The Saturniidae (Lepidoptera) of the Western Hemisphere.

Morphology, phylogeny, and classification. Bull. American Mus. Nat.
Hist. 98: 335-502, 420 figs.

130

DONOHUE AND LEMAIRE

J. Rex. Lepicl.

Fig. 2 (top). — Ormiscodes (Dirphiella) taylori, dorsal view of holotype
male. Approximately 1.4 times actual size.

Fig. 3 (bottom). — Ormiscodes (Dirphiella) albofasciata, dorsal view of
holotype male. Approximately 1.6 times actual size.

Journal of Research on the Lepidoptera

13:131-136, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

A NEW SUBSPECIES OF
EUPHYDRYAS EDITH A FROM THE
CHANNEL ISLANDS OF CALIFORNIA

THOMAS C. EMMEL AND JOHN F. EMMEL

Department of Zoology, University of Florida, Gainesville 32611
and 1117 9th Street, Santa Monica, California 90403

In our recent book. The Butterflies of Southern California
(Emmel & Emmel, 1973b: p. 34), we mentioned the occurrence
of an unrecognized dark race of Euphydryas editha ( Boisduval )
on the Channel Islands off the coast of Santa Barbara and Ven-
tura Counties, California. The purpose of this present paper is
to formally describe this new subspecies. Publications already
out (e.g., Emmel, 1967; Singer, 1971, 1972) and in preparation
analyze the ecological, host plant, and evolutionary affinities of
the various geographic sets of populations of Euphydryas editha
in California and the other western states. These data will allow
evolutionarily interesting comparisons to be made with informa-
tion from similar studies in the Euphydryas chalcedona complex
(Hovanitz, 1941, 1942; LeGare & Hovanitz, 1951; Emmel &
Emmel, 1973a; Emmel, T. C., in preparation).

Euphydryas editha insularis, new subspecies
(Figure!: a, b, and c)

Diagnosis: This is a very dusky, blackish subspecies with
melanic scaling greatly reducing the moderate to extensive yel-
low and red spotting present on other coastal and montane
Euphydryas editha subspecies in California. The type series
specimens of five males and one female are slightly larger by one
to three millimeters of fore wing length than the average E. e.
hayensis specimen from coastal central California (Emmel, 1967:
p. 75-76). The two sexes are remarkably similar in wing colora-
tion and pattern.

The forewings above have a faint, nearly marginal row of dull
red spots and a faint submarginal row of small crescent-shaped
cream spots. A wide black submarginal band bears a row of small

131

132

T. C. EMMEL AND J. F. EMMEL

/. Res. Lepid.

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13:131-136, 1974

NEW EUPHYDRYAS

133

cream spots. A nearly continuous row of cream-red spots follows
in a median position, with one or two red spots alternating with
cream spots extending posteriorly from the coastal margin of the
forewing as one progresses basally from the median row. One
cream spot is found posteriorly in the otherwise-black basal half
of the wing.

The hind wings above exhibit five well-isolated rows of red
and cream spots, alternately placed distally to proximally in a
rich black ground color. There is a faint, nearly marginal row of
small, dull red spots, and a submarginal row of slightly larger
cream spots. There is a postmedian row of crisply-bordered, en-
tirely dull red spots, of small size compared to mainland coastal
populations (see Fig. 1). There is a median row of distinct
cream spots (the spots in this row merge and become an essen-
tially continuous band in E. e. bayensis, separated only by black-
pigmented veins). Basal to this is a very faintly marked row of
reddish spots, with the bottom half usually heavily melanic.
Proximal to the body are several irregularly-shaped cream spots.

Ventrally, the fore wing and hindwing patterns are strongly
marked with red and cream with reduction of melanic portions
to along wing veins except in the submarginal black band area
on the forewing and in the postmedian area to the outside of the
median row of cream spots on the hindwing. Only in these two
areas do the underside pattern and coloration diverge notably
from those of mainland populations by being blackish in aspect.

The subspecies is named for the type locality, the Channel
Islands group off the coast of California. The type specimens
were collected by Chris Henne in 1941 on Santa Rosa Island ( one
fresh female on March 31, and five fresh males on April 1).
Henne (personal communication) describes the exact locality
of the colony as being at the top of a rolling hill at perhaps 300
feet elevation, directly behind the ranch house on the Vails’
Ranch. Poor weather prevailed during his visit (made with
Lloyd Martin of the L.A.C.M.). One very worn E. chalcedona
male was taken on Santa Cruz Island, March 27, 1941, by Lloyd
M. Martin; the specimen is labeled with the L.A.C.M. Channel
Islands Biological Survey label (No. 1941-4707), and was placed
in the Museum Collection with the series of six E. editha from
Santa Rosa Island. However, it is clearly chalcedona.

HOLOTYPE Male: Radius of forewing, 22.1 mm.; CALI-
FORNIA, Santa Barbara County, Santa Rosa Island, April 1,
1941. Collected by C. Henne. Los Angeles Museum Channel
Islands Biological Survey. No. 1941-4831.

ALLOTYPE Female: Radius of forewing, 28.5 mm.; CALI-
FORNIA, Santa Barbara County, Santa Rosa Island, March 31,

134

T. C. EMMEL AND J. F. EMMEL

/. Res. Lepid.

1941. Collected by C. Henne. Los Angeles Museum Channel
Islands Biological Survey. No. 1941-4755.

PARATYPES: 4 males, same locality, collector and survey
number data as holotype.

The holotype, allotype, and paratypes are deposited in the
collection of the Natural History Museum of Los Angeles County.

It is worthy of note that despite extensive correspondence
with many California lepidopterists during the preparation of
our book, no additional records or specimens of Euphydryas
from the Channel Islands have come to light. The above seven
butterflies were taken during the Los Angeles County Museum’s
Channel Islands Biological Survey (1941). Robert Langston of
the University of California Medical Center at San Francisco
has done subsequent field work on Santa Cruz Island and did not
find E. editha in April; however, Euphydryas chalcedona
(Doubleday) was found during a visit to this island by Lang-
ston from April 25 to May 1, 1966. None of the many lepidop-
terists who have collected on Santa CataHna Island in all months
throughout the years has found E. editha. For that matter,
E. chalcedona itself surprisingly has not been recorded from
Santa Catalina Island (Meadows, 1936; Emmel & Emmel,
1973b).

A glance at the map of the relative positions of the Channel
Islands in Figure 2 shows that the northern cluster of the four
islands of Anacapa, Santa Cruz, Santa Rosa, and San Miguel is
more susceptible to colonization from the few known coastal
E. editha colonies (Emmel & Emmel, 1973) on the mainland,
than are the relatively well isolated southern islands of Santa
Catalina, San Clemente, San Nicholas, and Santa Barbara, par-
ticularly by a non-migratory and indeed relatively sedentary
species (Ehrlich, 1965; Singer, 1972).

Historically, there is even stronger reason to see why E. editha
would be found in the northern and not the southern Channel
Islands. Thorne (1969) states that the northern group had ter-
restrial conection with the mainland down to perhaps a half
million years ago, and that all these islands were interconnected
with each other (but not with the mainland) during the Iowan
or lower Wisconsin glacial stages from about 11,000 to 20,000
years ago, when sea levels were lower. There seems to be no
evidence for the possibility of recent overland migration to the
southern Channel Islands, due to their prolonged and complete
submergence during the Pliocene and Pleistocene epochs
(Thorne, 1969).

13;131-136, 1974

NEW EUPHYDRYAS

135

Fig. 2. — Map of the Channel Islands and the coastal counties of southern
California.

136

T. C. EMMEL AND J. F. EMMEL

J. Res. Lepid.

One man-influenced factor contributing to the few records of
butterflies from the northern Channel Islands in the past has been
their private or Navy ownership and relatively restrictive policies
on visitation. The establishment of several university laboratories
on the isthmus of Santa Catalina and on Santa Cruz Islands^ as
well as Channel Islands National Monument on Santa Barbara,
San Miguel, and Anacapa Islands, has made these sites easier to
visit. Details of how to secure permission to visit these areas and
the other islands are given by Thome (1969: pp. 406-407). It is
also worth noting that the native vegetation of several islands has
been devastated by feral animals such as goats, and these islands
may presently have no suitable habitats for Euphydryas colonies.
ACKNOWLEDGEMENTS

We are grateful to William Hovanitz for his editorial help in
facilitating the publication of this paper with its color plate. The
photograph was taken by Alice Owens of Instructional Resources,
University of Florida. We thank Charles L. Hogue for the loan
of material in the collection at the Natural History Museum of
Los Angeles County. The map was prepared by Paul Laessle,
Staff Artist, Department of Zoology, University of Florida.
LITERATURE CITED

EHRLICH, PAUL R. 1965. The population biology of the butterfly,
Euphydryas editha. II. The structure of the Jasper Ridge colony.
Evolution 19: 327-336.

EMMEL, THOMAS C. 1967. Genetic and phenetic differentiation in two
complexes of western American butterflies. Stanford University, Ph.D.
dissertation. 101 pp.

EMMEL, THOMAS C., and JOHN F. EMMEL. 1973a. Two new sub-
species of Euphydryas chalcedona from the Mojave Desert of southern
California. J. Res. Lepid. 11: 141-146. (‘4972’’)

1973b. The Butterflies of Southern

California. Natural History Museum of Los Angeles County. 148 pp.
+ xii.

HOVANITZ, WILLIAM. 1941. Parallel ecogenotypical color variation in
butterflies. Ecology 22: 259-284.

1942. Genetic and ecologic analyses of wild populations in

Lepidoptera. I. Pupal size and weight variation in some California
populations of Melitaea chalcedona. Ecology 23: 175-188.

LeCARE, MARY JUDE, and WILLIAM HOVANITZ. 1951. Genetic and
ecologic analyses of wild populations in Lepidoptera. H. Color pattern
variations in Melitaea chalcedona. Wasmann Journ. of Biology 9:
257-310.

MEADOWS, DON. 1936. An annotated list of the Lepidoptera of Santa
Catalina Island, California. Bulletin So. Calif. Acad. Sci. 35: 175-180.
SINGER, MICHAEL C. 1971. Ecological studies on the butterfly Euphy-
dryas editha. Stanford University, Ph.D. dissertation. 80 pp.

1972. Complex components of habitat suitability within a

butterfly colony. Science 176: 75-77.

THORNE, ROBERT F. 1969. The California Islands. Ann. Missouri Bot.
Card. 56: 391-408.

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

137

(Continued from page 122)

B. kaber (DC.) Wheeler, Raphanus sativus L., and probably others.

41. Pieris sisymbrii Boisduval. California White.

One male, American River, hi. 14.73, presumably a stray from the Sierra
foothills.

42. Euchloe ausonides Boisduval. Large Marble.

Common on floodplains, along sloughs and ditches, straying elsewhere.
Sacramento Valley specimens average much larger than those from the foot-
hills, at least in late spring; most of the females are orange-tinted {semi-
flava Comstock, flavidalis Comstock). In 1972 seemingly one-brooded with
a very long emergence, but in 1973 clearly double-brooded with two tem-
poral phenotypes. Males patrol territories along gullies or ditches. Host
Plants: Cruciferae: Raphanus sativus L. (preferred); Brassica nigra (L.)
Koch.; B. napus L. According to R. Langston, this species flies much later
into summer in the San Francisco Bay area, but is still presumably two-
brooded.

43. Anthocaris sara Lucas. Sara Orange-Tip.

Frequent in riparian forest along the Sacramento and American
Rivers; occasional elsewhere along fencerows, ditches, etc. The second brood
appears to be rudimentary. Host Plants: Cruciferae: Brassica napus L.; B.
geniculata ( Desf. ) J. Ball; Raphanus sativus L.

44. Colias eurytheme Boisduval. Orange Sulphur.

Abundant throughout, especially in alfalfa fields, reaching greatest
densities in ix. Six broods, the first and last of form ariadne, heavily shaded
and with reduced borders and orange color above. Extremely variable, pro-
ducing cream-yellow, pale green, buff and other unusual females mainly in
autumn; a cream-white male, Davis, ix. 10.72; chrome yellow females,
Willowbank, viii. 13.72, Willow Slough, ix.27.72 and x.26.72; yellow male.
Willow Slough, ix.27.72. Flew 44 weeks in 1972 and 44 in 1973. Host
Plants: Leguniinosae: Medicago sativa L.; Vida angustifolia Reich.; V.
sativa L.; V. cracca L.; V. villosa Roth; Lotus strigosus (Nutt.) Greene;
Glycyrrhiza lepidota Pursh; Lathyrus jepsonii Green ssp. californicus (Wats.)
Hitchc.

45. Colias (Zerene) eurydice Boisduval. California Dog-face.

Rare stray in the Valley (Davis, iv. 26.72, vi.7.72). Not known to breed,
although the usual host, Leguminosae: Amorpha calif ornica Nutt., grows at
Elkhorn Ferry.

Family Hesperiidae

46. Epargyreus clarus Cramer. Silver-Spotted Skipper.

Infrequent, mostly near the host plant (American River, Southport,
Broderick, Clarksburg, Davis). Perhaps triple-brooded. Host Plant: Legumi-
nosae: Robinia pseudoacacia L., the usual host, is widely naturalized in
sandy bottomlands in the Valley. Breeding has not been confirmed, but is
almost certain.

47. Pyrgus scriptura Boisduval. Least Checkered Skipper.

Common to locally abundant on waste ground, roadsides, and levees,
with the host plant; most often on compacted clay or alkali soils. Four to
five broods, the first most contrastingly marked. Host Plant: Malvaceae:
Sida hederacea (Dough) Torr.

48. Pyrgus communis Grote. Common Checkered Skipper.

Common to occasionally abundant throughout, usually in open country;
especially numerous in agricultural land. Four or five broods, the cold
weather specimens smaller, hairier, and whiter. Host Plants: Malvaceae:
Malva neglecta Wallr.; M. parviflora L.; M. sylvestris L.; M. nicaeensis AIL;
Althaea rosea L.; Sida hederacea (Dough) Torr. (Note: Ptjrgus albescens
Plotz, the Common Checkered Skipper of southern California, has not been
recorded from “northern or central California” (Tilden, 1965b ), but its pres-
ence or absence in the Valley should be confirmed by genitalic examination
of many males from a variety of localities. )

138

ARTHUR M. SHAPIRO

/. Res. Lepid.

49. Heliopetes ericetorum Boisduval. Large Checkered Skipper.

Rare immigrant; both broods represented, but not known to breed in
the Valley although a gravid female was taken in West Sacramento, v.5.73.

50. Pholisora catullus Fabricius. Sooty Wing.

Common, but seldom abundant, on waste ground, levees, and roadsides,
often on compacted or alkali soils. Four or five broods. Host Plants: Amaran-
thaceae: Amaranthus hybridus L. preferred; A. retroflexus L.; Chenopo-
diaceae: Chenopodium album L.; C. murale L.; Atriplex rosea L.

51. Erynnis persius Scudder. Persius Dusky-Wing.

Frequent in sandy floodplains, as at Putah Creek and Willow Slough.
Three (perhaps four) broods. Host Plant: Unknown; Leguminosae: Lotus
purshianus (Benth.) Clem, & Clem, a possibility.

52. Erynnis propertius Scudder and Burgess. Propertius Dusky-Wing.
Infrequent in tlie Valley; mostly strays along the edges, but perhaps

breeding along the American River where a very fresh specimen was taken
hi. 9.73. Flies only in spring.

53. Erynnis tristis Boisduval. Sad Dusky-Wing.

Common throughout, wherever planted or native oaks occur, and stray-
ing widely. Four broods. Host Plants: Fagaceae: Quercus suber L. (pre-
ferred at Davis, although introduced); Q. lobata Nee.; Q. douglasii H. & A.;
Q. wislizenii A. DC. In all cases only tender new growth is eaten.

54. Erynnis zarucco funeralis Scudder and Burgess. Funereal Dusky-Wing.
Infrequent summer stray from the San Joaquin. Valley, where resident.

Not known to breed this far north.

55. Hylephila phylaeus Drury. Fiery Skipper.

Abundant in lawns, waste places, and sandy areas throughout. Five
broods; cold weather specimens darker beneath. Host Plant: Gramineae:
Cynodon dactylon (L. ) Pers.

56. Atalopedes campestris Boisduval. Field Skipper.

Common; most numerous in sandy bottomlands, but straying widely.
Four broods. Host Plant: Gramineae: Cynodon dactylon (L.) Pers.

57. Ochlodes sylvanoides Boisduval. Woodland Skipper.

Abundant in riparian forest; less common to infrequent elsewhere;
absent in agricultural land. Not emerging until midsummer, but at least two
broods. Host Plants: Gramineae: at Broderick, Phalaris calif ornica H. & A.
and P. lemmonii Vasey, perhaps other grasses.

58. Ochlodes yuma Edwards. Yuma Skipper.

Thus far recorded only at Stone Lake and Beach Lake in southern
Sacramento County, in fresh-water marsh. Two broods. These records are
the northernmost in the Valley; the species is widespread at Suisun Bay
and in the Sacramento-San Joaquin Delta. Host Plant: Gramineae: Phrag-
mites communis Trin. var. Berlandieri (Fourn. ) Fern, (at Suisun Bay).

59. Ochlodes agricola Boisduval. The Farmer.

Rare stray along the streams from the foothills; not known to breed in
the Valley.

60. Polites sabuleti Boisduval. Sandhill Skipper.

Abundant on sandy soils, as in river bottoms and on levees; widely
distributed on waste ground and lawns in West Sacramento and Broderick,
etc.; also in saline and alkali marshes; occasional elsewhere. Five broods,
spring and fall specimens more heavily marked and often undersized, re-
sembling the high-elevation subspecies tecumseh Grinnell. Host Plants:
Gramineae: Distichlis spicata (L.) Greene; Cynodon dactylon (L. ) Pers.

61. Poanes melane Edwards. Umber Skipper.

Frequent to common in riparian forest and levee thickets along the
Sacramento and American Rivers; not seen elsewhere. Two broods. Host not
known.

62. Lerodea eufala Edwards. Eufala Skipper.

Abundant throughout after late June, occasionally seen earlier; num-
bers variable from year to year. At least two broods, A buff-colored aberrant

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

139

was taken at Willow Slough, ix.4.72. Host Plants: Gramineae: Cynodon
dactylon (L. ) Pers.; Sorghum halepense (L.) Pers.; S. bicolor (L. )
Moench.; Setaria verticillata (L. ) Beauv.; Echinochloa cnis-galU (L.)
Beauv.; Oryza sativa L. The exact date when this southern species first
entered the Valley is unknown, but it has been at Davis since the mid-
1950s, at least.

ACKNOM/LEDGMENTS

This survey was possible only with the help of students and
staff of the University of California, Davis. Dr. E. W. Jameson,
Jr. and Messrs. Allen Allison, Steve Sims, and Steve Strand of
the Zoology Department were especially helpful. Mr. Oakley
Shields of the Entomology Department provided biological in-
formation on many species as they occur in the Vacas. Mr.
Robert L. Langston of Kensington, Ca. assisted in determinations
and provided much useful information. Plant determinations
were made using the resources of the U.C. Davis herbarium.
Mrs. Adrienne R. Shapiro accompanied me on many field trips
and provided a substantial number of records.

LITERATURE CITED

BAKKER, E. 1971. An Island Called California. University of California
Press, Berkeley, Los Angeles and London. 357 pp.

BURNS, J. M. 1964. Evolution in the skipper butterflies of the genus
Erynnis. Univ. of Calif. Publ. Ent. 37; 1-214.

COMSTOCK, J. A. 1927. Butterflies of California. Published by the author,
Los Angeles. 334 pp.

DALE, R. F. 1966. Climate of California. U.S. Department of Commerce,
E.S.S.A. Climates of the States — -California. Publication 60-4. 46 pp.
DOS PASSOS, C. F. 1964. A Synonymic List of Nearctic Rhopalocera.

Lepidopterists Society, Memoir 1. 145 pp.

EHRLICH, P. R. and A. H. EHRLICH. 1961. How to Know the Butterfies.

W. C. Brown Co., Dubuque, Iowa. 262 pp.

EMMEL, T. C. and J. F. EMMEL. 1962. Ecological studies of Rhopalocera
in a high Sierran community — Donner Pass, California. I. Butterfly
associations and distributional factors. /. Lepid. Soc. 16: 23-44.

— — and 1973. The Butterflies of Southern California. Los Angeles

Co. Mus. Nat. Hist., Science Series 26: 1-148.

FEENY, P. P. 1970. Seasonal changes in oak leaf tannins and nutrients as
a cause of spring feeding by winter moth larvae. Ecology 51: 565-581.
FIGGINS, W. E. 1971. Climate of Sacramento, California. NOAA Techni-
cal Memorandum NWS-WR65. 63 pp.

GARDINER, B. O. C. 1972. Pieris rapae L. larvae throughout the winter.
Ent. Record J. Var. 84: 112.

GARTH, J. S. and J. W. TILDEN. 1963. Yosemite buterflies. J. Res. Lepid.
2: 1-96.

HARRIS, L. 1972. Butterflies of Georgia. University of Oklahoma Press,
Norman. 324 pp.

HEPPNER, J. B. 1973. The distribution of Paratrytone melane and its
spread into San Diego County ( Hesperiidae ) . J. Res. Lepid. 10: 287-

140

ARTHUR M. SHAPIRO

/. Res. Lepid

MUNZ, P. A. 1970. A California Flora. University of California Press,
Berkeley, Los Angeles and London. 1681 pp.

OPLER, P. A. and J. S. BUCKETT. 1971. Seasonal distribution of “Macro-
lepidoptera” in Santa Clara County, California. J. Res. Lepid. 9: 75-88.
OPLER, P. A. and R. L. Langston. 1968. A distributional analysis of tlie
butterflies of Contra Costa County, California. J. Lepid. Soc. 22: 89-
107.

SCULLEY, R. 1973. The natural state. Davis New Review 1 ( 1 ) : 3-13.
SHAPIRO, A. M. 1973. Movement of Nymphalis calif ornica in 1972 (Nym-
phalidae). J. Lepid. Soc., in press.

— . 1974a. The butterflies and skippers of New York ( Lepidoptera:

Papilionoidea, Hesperioidea ) . Searchl Agriculture, in press.

1974b. Ecotypic variation in montane butterflies. Wasmann ]. Biol.,

in press.

. 1974c. Natural and laboratory occurrence of “elymi” phenotypes in

Cynthia cardui ( Nymphalidae). J. Res. Lepid., in press.

. 1974d. Recurrent aberration in Cynthia annabella (Lepidoptera:

Nymphalidae), with four new records. Pan-Pac. Ent., in press.
SHIELDS, O. 1966, The butterfly fauna of a yellow pine forest community
in the Sierra Nevada, California. J. Res. Lepid. 5: 127-128.
THOMPSON, K. 1961. Riparian forests of the Sacramento Valley, Cali-
fornia. Ann. Assoc. Amer. Geogr. 51: 294-314.

TILDEN, J. W. 1965a. Butterflies of the San Francisco Bay Region. Uni-
versity of California Press, Berkeley and Los Angeles. 88 pp.

, 1965b. A note on Pyrgus communis and Pyrgus albescens ( Hesperi-

idae). J. Lepid. Soc. 19: 91-94.

— . 1971. Comments on the Nearctic members of the genus Precis

Hubner. J. Res. Lepid. 9: 101-108 (“1970”).

13:73-82, 115^122, 137-^148, 1974 SACRAMENTO VALLEY 141

Table 1

Distribution of univoltinism in some
California butterfly faunas

Locality and

Elevation :

Vaca Mts.
(100-2500')

Contra
Costa Co.
(<3850')

Sacramento

Valleyl

(100’)

Boreal

Ridge

(7000')

Marin-

Sierra

Camp

(5000')

Number of species

65

84

62 (53)

70

63

Number of univoltines

25

27

14 (8)

47

38

Per cent univoltine

39%

32%

23% (13%)

67%

60%

The first number refers to the total fauna. The number in parentheses
refers to breeding residents only. Immigrants are from the Vacas and
Sierra foothills.

142

ARTHUR M. SHAPIRO

J. Res. Lepid.

Table 2. Climatic Summary for
Data from Dale, 1966.

Sacramento Airport Station, NWS.

(For precipitation data, see Table 3)

3SOjg mm\

O O O O O CM IT)

3og: AABaq

CMOOOO OOCMv^lcy.

“ IIHJUTBJ q5M|

00 m cn

cn \£> as m

fO fO 00

es vD eo

^

M O
cd rH
Pm a

m m 00

in so s£>

^osuns-asrauns ‘sq^uaijl o
UT jaAoo Aqs UBajql

aupqsunsl
aqqTssod jo %j ^

esj

en m sD

m CO

00 0^ ON

XSd Rd <7

% ‘A^TPTnmq aAT^eqa^l

0)

eo <£>

tH •
•H 6
Cd

Q a

X

CM vD CM

«sa)«dQ.a332

4J > O

o o <u

o a o

Yeats of Record

13:73-^82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

143

Table 3.

Normal

total

and maximimi and minimum monthly and annual

precipitation

, Sacramento. Data from Figgins, 1971, which

gives

station

and observer notes; covers

period July 1849-

July 1970.

Month

Mean

Maximum and Year

Minimum and Year

JAN.

3.58

15.04-1862

0.15-1889

FEB.

3.47

9.25-1940

0.04-1899

MAR.

2.46

10.00-1850

0.03-1956

APR.

1.62

14.20-1890

*

Trace-1949

MAY

0,62

3.25-1889

0 -1920*

JUNE

0.12

1.45-1884

0 -1969*

JULY

Trace

0.63-1860

**

0 -1970

AUG.

0.04

0.67-1953

**

0 -1969

SEPT.

0.22

3.62-1904

0 -1968*

OCT.

0.80

6.85-1962

0 -1966*

NOV.

1.54

11.34-1885

0 -1933*

DEC.

3.55

13.40-1852

0 -1876

Annual

18.02

36.35-1852/53

4.71-1850/51

indicates year is most recent of record; minimum has occurred in
prior years.

It

minimum has occurred in 50% or more of years of record.

144

ARTHUR M. SHAPIRO

/. Res. Lepid.

Table 4. First flight dates of 25 common butterflies in the dry 1972
and wet 1973 seasons.

Species

1972

1973

Difference

Vanessa carye

ii.lO

ii.7

(+3)

Nymphalis antlopa

ii.lO

il.7

(+3)

Pieris rapae

il.26

1.20

(+37)

Pyrgus scriptura

iil.l

iii. 24

(”23)

Everes comyntas

ill. 3

ill. 26

(-23

Pholisora catullus

lii.3

ill. 27

(-24)

Strymon melinus

iii.3

iii. 9

(-6)

Vanessa atalanta

lii. 4

li.l8

(+17)

Phyciodes mylitta

iii. 4

iii. 9

(-5)

Plebeius acmon

ill. 4

iii. 14

(-10)

Atlides halesus

iii. 5

iv.3

(-29)

Colias eurytheme

iii. 5

ii.25

(+11)

Papilio zelicaon

iii. 5

iii. 14

(-9)

Papilio rutulus

iii. 6

iii. 18

(-12)

Pyrgus communis

iii. 7

iii. 24

(-17)

Vanessa cardui (immigrant)

lii. 8

ii.22

(+17)

Euchloe ausonides

iii. 8

iii. 2

(+6)

Precis coenia

iii. 11

iv.3

(-23)

Erynnis tristis

iii. 16

iv.8

(-23)

Lycaena helloides

iii. 20

iv.29

(-40)

Phyciodes campestris

iii. 23

V.2

(-39)

Hylephila phylaeus

iv.l

lv.12

(-11)

Polites sabuleti

iv.2

v.2

(-30)

Limenitls lorquini

iv. 16

iv.27

(-11)

Atalopedes campestris

iv.l9

V.5

(-16)

Mean departure from 1972 dates, 1973

-10.3

13:73-82, 115-122, 137-148, 1974

SACRAMENTO VALLEY

145

Table 5. Climatic comparison of 1971-72 and 1972-73 winters at Sacramento
Airport Station, NWS.

1971-72

Sept

Oct

Nov

Dec

Jan

Feb

Mar

Total

Precipitation, inches

T

0.13

0.87

4.05

0.81

1.28

0.29

7.43

Mean high temperature, °F

88.7

76.1

64.7

50.7

48.5

61.0

71.6

65.9

Mean low temperature, ®F

55.6

46.3

40.7

35.2

33.5

41.8

45.5

42.7

Mean monthly temperature,

72.2

61.2

52.7

43.0

41.0

51.4

58.6

54.3

®F and deviation from
normal

+0.6

-2.3

-0.2

-3.4

-4.2

+2.2

+5.2

-0.3

Days below 32®

0

0

1

11

13

3

0

28

Days with precipitation
>.01"

0

2

6

13

6

9

5

41

Days with heavy fog

0

1

5

12

15

8

1

42

% sunshine

95

86

79

54

48

64

89

73

Mean sky cover in tenths ,
sunrise-sunset

1.1

2.9

4.5

7.2

7.2

7.5

4.6

5.0

1972-

■73

Sept

Oct

Nov

Dec

Jan

Feb

Mar

Total

Precipitation, inches

0.90

1.75

5.14

1.88

6.87

5.64

2.76

24.94

Mean high temperature, °F

84.3

72.5

56.5

46.8

51.6

60.3

60.0

61.7

Mean low temperature, °F

54.6

51.5

42.8

34.3

37.0

45.9

42.2

44.0

Mean monthly temperature.

69.5

62.0

49.7

40.6

44.3

53.1

51.1

52.9

®F and deviation from
normal

-2.1

-1.5

-3.2

-5.8

-0.9

+3.9

-2.3

-1.7

Days below 32°

0

0

0

13

18

0

0

31

Days with precipitation
>.01"

3

8

14

11

17

16

11

80

Days with heavy fog

0

5

10

7

7

6

1

36

% sunshine

89

67

42

46

53

51

72

60

Mean sky cover in tenths.

2.6

4.9

7.6

6.2

6.7

8.2

5.9

6.0

sunrise-sunset

Table 6

Host adaptations of Sacramento Valley butterflies
grouped by voltlnism.

Monophagous

Oligophagous

Polyphagous

Univoltine species

5

0

0

Multivoltine species

10

19

3

Totals

- 8.56

15

19

.025 > p >.010

3

146

ARTHUR M. SHAPIRO

J. Res. Lepid.

Itvf

nvx

S-^x

zz-a-

8X
♦■r. ,'<•

Ol’Xi

n}>'^

f.l'fiA

tnvk

Z JfA
9l*A
%-'A
IfA
tA

It A!

6A!

zr!ii

>»t !>

r,!!'.

411

Si"!

■Ubl

SZ»x

intx

lt*n

tl IX
OVK
91 X

r/K
SI w
4-x?

IZ'IMA

OIJiA
9r‘tA
HriA
At A
JJI'A
I A
ilM
f -.M
Ot-ll!
9-m
IZI!
4-}!

■ZLbi

X K

I XX

lx

xl

-I

X X

5l

X X

II

p

p

>%CO (U

ss

111

c<l -^3

§

H .5 CO

13:73-82, 115-122, 137-148, 1974
Table 7 (continued).

SACRAMENTO VALLEY

X ^

1

X

X

X X

>

55

X

X

X

X

n

147

NOTICES

B ooks on Lepidoptera: more than 300 books for sale. Free
catalogue sent on request. Sciences Nat. , 45 Rue de Alouettes, 75019,

Paris, France

Wanted to make exchanges with collectors in California and Florida,
Have Maylaysian butterflies for exchange (Trogonoptera brookiana, etc. )
Masaki NAKAYAMA, Kitakyushu- Wakamatsu- Miyamaru -Z- 10- 14,
Fukuoka pref. , Japan ,

Need aid in preparation of specimens for study of population
structure in butterflies; involves obtaining population samples, mounting,
laboratory breeding, etc. Full or part time. Contact William Hovanitz,
1160 W. Orange Grove Ave. Arcadia, Calif. 91006.

THE JOURNAL ©F RESEAR.CHJ
©NJ THE LEF!JD©PTEFIA\

Volume 13

Number 2

June, 1974

IN THIS ISSUE

The butterfly fauna of the

Sacramento Valley, California

Arthur M. Shapiro 73

Extended flight periods of coastal and
dune butterflies in California

Robert L. Langston 83

Habitat: Brephidium pseudofea (Lycaenidae)

John B. Heppner 90

The early stages of various species of
the genus Dirphia (Saturniidae)

Brian O. C. Gardiner 101

A new species of Ormiscodes (Dirphiella) from
Mexico (Saturniidae: Hemileucinae )

Julian Pe Donahue and Claude Lemaire 123

A new subspecies of Euphydryas editha
from the Channel Islands of California

Thomas C. Emmel and John F. Emmel

131

THE JOURNAL j ^

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ON THE LEPIDO'PTERA,

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Journal of Research on the Lepidoptera

13(3):149-=156, 1974

1160 W. Orange Grove Ave,, Arcadia, California 91006, U.S.A.
© Copyright 1975

STUDIES OF THE OVA AND FIRST INSTAR

LARVAE OF GEOMETRIDAE
(ENNOMINAE). I.

ROGER L. HEITZMAN'

3112 Harris Avenue, Independence, Missouri 64052

ABSTRACT

Four species of the subfamily ENNOMINAE {Pero marmoratus
Grossbeck, Syssaura puber Grote & Robinson, Apicia confusaria (Hubner)
and Tetrads crocallata Guenee) are studied, and illustrations of setal maps,
anal plates and head cases, and photographs of ova, are presented.

INTRODUCTION

This is the first in a series of studies of the ova and first
instar larvae of GEOMETRIDAE. It is hoped that these studies
will provide further understanding of species, genus and sub-
family relationships existing in the GEOMETRIDAE.

MATERIALS AND METHODS

Ova were secured by placing females in small vials contain-
ing paper tissue. I have found that most geometridae lay quite
readily under these conditions, usually the very night they are
captured.

A WILD M-5 microscope with drawing tube attachment
was used throughout the study. Photographs of ova were taken
with phototube attachment and a Polaroid camera in conjunc-
tion with a double iris diaphram for increased clarity.

The descriptions are based on ova and larvae from these
female captures: Pero marmoratus, taken at Coolie Lake, Clay
Co., Missouri, 3 May 1972; Syssaura puber, taken at Blue Springs
State Park, Washington Co., Arkansas, 26 May 1972; Apicia con-
fusaria, taken at Blue Springs State Park, Washington Co.,
Arkansas, 26 May 1972; Tetrads crocallata, taken at Warsaw,
Benton Co., Missouri, 1 June 1972.

^Research Associate, Florida State Collection of Arthropods, Division of
Plant Industry, Florida Department of Agriculture and Consumer Services,
Gainesville.

149

150

ROGER L. HEITZMAN

/. Res. Lepid.

A

Fig. 1. — Pero marmoratus Grossbeck; (A) setal maps; (B) anal plate;
(C,D) head, frontal and left lateral aspect, SOX.

13(3):149-156, 1974

OVA AND LARVAE

151

RESULTS

Pero marmomtus Grossbeck

OVUM: Height: 0.82mm. Width: 0.65mm. Grass green with yel-
low areas. Laid singly and in groups on end or side. Eclosion
in seven days.

FIRST INSTAR LARVAE: Length: approximately 4mm. Head
very light brown, height: 0.38mm, width: 0.42mm. Dorsal sur-
face: green with dark green bands encircling thoracic segments
and fold between abdominal segments one through nine. Anal
area yellowish green. Lateral surface: green. Anal as dorsal.
Ventral surface: yellow green. Anal as dorsal.

Syssaura puber Grote & Robinson
OVUM: Height: 0.74mm. Width: 0.58mm. Yellow changing to
pale orange. Laid singly and in small groups on end or side.
Eclosion in eight days.

FIRST INSTAR LARVAE: Length: approximately 3mm. Head
very light yellow brown, height: 0.33mm, width: 0.36mm.
Dorsal surface: thorax and last five abdominal segments yellow;
first five abdominal segments light yellow. Dark brown square
on first five abdominal segments. Lateral margins of squares
joined by stripe running from first thoracic to eighth abdominal,
paralleled by suprastigmatal brown stripes. Light brown band
emerges from each square to encircle the body. Lateral surface:
colors same as dorsal. First five abdominal stigma centered in
a very dark brown irregular circle. A brown stripe joins each
circle about the stigma. This stripe begins on first abdominal
and ends on sixth abdominal. Ventral surface: colors same as
dorsal. Dark brown square on the first five abdominal segments.
A mid-ventral stripe joins the squares. Lateral margins of
squares joined by stripes and paralleled by substigmatal stripes
extending from first to sixth abdominal segments.

Apicia confusaria (Hubner)

OVUM: Height: 0.83mm. Width: 0.67mm. Green changing to
dark golden brown. Laid loose without adhesive. Eclosion in
ten days.

FIRST INSTAR LARVAE: Length: approximately 3mm. Head
very dark brown almost black, height: 0.42mm, width: 0.38mm.
Dorsal surface: first thoracic brown, second and third lighter
with white blotches and markings. First six abdominal segments
dark brown, last four lighter brown. First five abdominal seg-
ments with white V-shaped marking, base of V posteriorly
oriented. White around base of setae of all abdominal segments.
Lateral surface: thorax almost entirely white. Large white area

152

ROGER L. HEITZMAN

/. Res. Lepid.

Fig. 2.-—Syssaura puber Grote & Robinson; (A) setal maps; (B) anal plate;
(C,D) head, frontal and left lateral aspect, 90X.

13(3):149-156, 1974

OVA AND LARVAE

153

A

Fig. 3.—Apicia confusaria (Hubner); (A) setal maps; (B) anal plate
(C,D) head, frontal and left lateral aspect, 85X.

154

ROGER L. HEITZMAN

/. Res. Lepid.

Fig. 4. — Tetrads crocallata Guenee; (A) setal maps; (B) anal plate;
(G,D) head, frontal and left lateral aspect, 90X.

13(3):149-156, 1974

OVA AND LARAVAE

155

Fig. 5.— Ova; (A) Tetrads crocallata; (B) Apida confusaria; (C) Syssaura
puber; (D) Pero marmoratus, all 50X.

156

ROGER L. HEITZMAN

/. Res. Lepid.

in center of each abdominal segment where setae are located.
Abdominal colors same as dorsal. Ventral surface: thorax white
and light brown. First five abdominal segments with white
blotches and striations. Sixth abdominal segment with two white
dashes. Abdominal colors same as dorsal.

Tetrads crocallata Guenee

OVUM: Height: 0.90mm. Width: 0.82mm. Very deep green.
Laid loosely without adhesive. Eclosion in eleven days.

FIRST INSTAR LARVAE: Length: approximately 3mm. Head
very dark brown almost black, height: 0.36mm, width: 0.42mm.
Dorsal surface: thorax dark brown, second and third segments
with white blotches and markings. First six abdominal seg-
ments very dark brown almost black; last four segments brown.
First five segments with white heart-shaped marking, bottom
of heart posteriorly oriented. Last four segments have some
vague creamy patterns. Lateral surface: thorax mostly white
except for first segment which is mainly dark brown. Large
white area in center of each abdominal segment where setae
are located. Area is creamy yellow on last five segments. Ab-
dominal colors same as dorsal. Ventral surface: thorax white and
brown. First five abdominal segments with white blotches and
striations. Abdominal colors same as dorsal.

Journal of Reasearch on the Lepidoptera

13(3):157»161, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

ALTITUDINAL MIGRATION OF
CENTRAL CALIFORNIA RUTTERFLIES
ARTHUR M. SHAPIRO

Department of Zoology, University of California, Davis 95616

Butterfly species which are not mass migrants in the sense
of the Monarch {Danaus plexippus L., Danaidae) or the Painted
Lady {Cynthia cardui L., Nymphalidae) nonetheless are often
recorded as “strays” far beyond their usual ranges. Such “strays”
may be geographic or altitudinal. The occurrence of individual
“strays” could reflect active or passive dispersal which could be
density-related or not. In a previous paper (Shapiro, 1974b)
data were presented on lowland California butterflies collected
at 5000 and 7000 feet above sea level in the central Sierra Nevada
in 1972. It was noted that several of the species displaying alti-
tudinal dispersal in California are well-known northward dis-
persers on the Atlantic coastal plain in the eastern United States.
Regular collections of butterflies and skippers were made at
Donner Pass, Placer County, California, elevation approximately
7000 feet, from 27 June to 16 October 1973, and from 31 May to
16 October 1974. In the course of this work, several additional
records of altitudinal dispersal were obtained, as reported
below. In all of these cases, the insects appear to have dispersed
upslope. In addition, some evidence suggesting downslope dis-
persal by one species, Nymphalis milberti Latreille (Nymphali-
dae), is presented.

Donner Pass Records

Donner Pass is well documented, thanks to assiduous col-
lecting by Emmel and Emmel (1962a, b; 1963a, b; 1974) and
appears to have the richest butterfly fauna of any investigated
montane area in temperate North America of comparable size—
about 100 species in a four-square-mile area. The geography
and vegetation of Donner Pass are discussed by Emmel and

157

158

ARTHUR M. SHAPIRO

J. Res. Lepid.

Emmel (1962a), who note that part of its richness stems from
the mixture and overlap of east-slope (Great Basin) and west-
slope (Californian) biotic elements. In evaluating the signifi-
cance of altitudinal dispersal at Donner, it is often difficult — or
impossible— to decide whether the immigrants are from the
east or west slope. The descent to the lowlands is much more
precipitous on the east slope, which is also less heavily forested,
so that dispersal ( especially if aided by thermals ) may be much
more rapid from the east.

The records which follow do not exhaust the list of potential
dispersers: only especially striking examples are enumerated.
LYCAENIDAE: Brephidiiim exilis Boisduval. A female Pygmy
Blue in good condition was taken 5 October 1973 feeding on the
ffowers of Chnjsothammis nauseosus (Pall.) Brit. (Compositae)
about 1000 feet east of Norden. B. exilis occurs abundantly on
the ffoor of the Sacramento Valley at the western base of the
Sierras, and has been taken up to about 2800 feet in the foot-
hills in El Dorado County and at 4600 feet in Nevada County.
It is also common on the Great Basin ffoor east of Sierraville
in late summer and in desert conditions near Reno, Nevada.
Colonization could thus occur from either side of the mountains.
Among potential hosts of B. exilis (Shapiro, 1973) only the intro-
duced weed Salsola kali L. var. tenuifolia Tausch. (Cheno-
podiaceae) occurs at Norden, along with two other undeter-
mined Chenopods.

Leptotes marina Reakirt. A male in excellent condition was taken
by S. R. Sims on C. nauseosus ffowers at the Donner Lake over-
look on the Norden-Truckee road, 28 September 1973. This blue
also occurs at low elevations on both sides of the Sierra. It is,
however, very local in the Sacramento Valley as compared with
the Great Basin and seems much more likely to have reached
Donner Pass from the east than the west. A female L. marina
was taken at Tahoe City, 18 miles south of the Pass at the same
elevation, on 24 September 1974. Tahoe City has a very marked
east-slope character.

Lycaena helloides Boisduval. The Purplish Copper is a fairly
frequent capture in the mid-elevation Sierras. In 1973, three
males were taken at Donner, on 17 August, 13 September, and
16 October. In 1974, single males were taken on 26 June and
24 July. The 16 October, 1973 specimen was taken on C. nause-
osus ffowers at the same spot where B. exilis was found the
previous week. L. helloides is common in the lowlands on both

13(3):157^161, 1974 ALTITUDINAL MIGRATION

159

sides of the Sierran rtiassif . Potential host plants ( Polygonaceae )
are numerous at Donner. On the west slope, L. helloides be-
comes sporadic above about 2500 feet, but in 1974 it was found
breeding at Lang Crossing, Nevada County, at about 4600 feet.
Habrodais grunus Boisduval. One female, 18 July 1973, on the
Lake Mary road. This is characteristically a species of the upper
foothills and through the Transition Zone, always associated
with its host, Quercus chrysolepis Liebm. (Fagaceae). This
tree is rarely seen above about 5000 feet at the latitude of
Donner Pass and has never been observed in the Pass area.
The specimen must have reached Donner from the west.

NYMPHALIDAE: Limenitis bredoivii calif ornica Butler. Emmel
and Emmel (1974) report three females, 21 June 1970. Individ-
uals (sex undetermined) were seen at Donner Pass on 11 July
and 28 September 1973, and on 9 June, 24 July, and 16 October
1974. It thus appears that L. b. calif ornica reaches Donner fairly
frequently, but its breeding status is unknown. It is a very
strong-flying species which is common to at least 5000 feet on
the west slope.

Precis coenia Hubner. In 1972, this was seen only once at 7000
feet (Boreal Ridge). In 1973, specimens were seen or taken
on 11 and 25 July, 17 August, 28 September, and 5 and 16
October at Donner. In 1974, it was not seen at Donner at all.
Emmel and Emmel considered it common at Donner in 1960.
Hosts ( Scrophulariaceae, Plantaginaceae ) are numerous and it
seems very likely that it breeds in years when gravid females
arrive before late summer. The Buckeye is abundant on the
floors of the Sacramento Valley and Great Basin, and it breeds
every year on the west slope to about 3000 feet.

HESPERIIDAE: Hylephila phylaeus Drury. Females: Soda
Springs, 28 September 1973, and Norden, 5 October 1973, the
latter on flowers of C. nauseosus. The Fiery Skipper is ex-
tremely abundant in the lowland Sacramento Valley, and locally
so around Reno in the Great Basin. In 1972 it was taken once
at 5000 feet on the west slope (female, 29 September). Immi-
gration from the west seems more probable for this species.

Atalopedes campestris Boisduval. One of each sex taken at Nor-
den, 24 July 1974. It seems very unlikely a priori that this lowland
species breeds or overwinters at Donner Pass, It is common in
the Sacramento Valley and unknown at Truckee.

160

ARTHUR M. SHAPIRO

/. Res. Lepid.

Donner Pass may be an exceptionally good accumulator of
altitudinal dispersers for topographic and meteorological reasons.
The Pass is noted for its high winds throughout the year, which
occur when barometric pressure is significantly higher on one
side of the range than on the other and the denser air rushes
through the Pass to displace the lighter air across it. The direc-
tion of this flow is more often from west to east than the
reverse. In autumn, however, Canadian high pressure may build
into the Great Basin and produce sustained easterly flow for
several days.

Movements of Nymphalis milberti

In general, records of altitudinal dispersers are of lowland
species at high elevations rather than the reverse. Similarly, in
the eastern United States, many more butterflies appear to
disperse northward than the reverse. The Tortoiseshells, genus
Nymphalis, provide exceptions to this generalization. In the
east, N. milbeHi Latreille and N. j~album Bdv. & LeC., while
basically northern, undergo periodic range extensions which
carry them far to the south of their permanent breeding grounds,
and to lower elevations. This occurs on a much larger scale in
the mass-migratory N. California Bdv. (Shapiro, 1974a).

Nymphalis milberti is generally a rare insect in central Cali-
fornia. Garth and Tilden (1963) report it from Transition Zone
up to 13,090 feet in Yosemite. Transition Zone, the zone of yellow
pine, incense cedar, and Douglas fir, has its lower limit around
3000 feet.

On 11 April 1973, a female N. milberti was taken at the
Spenceville Recreation Area, Yuba County, at an elevation of
about 1000 feet. It was found in a colony of the introduced
nettle, Urtica tirens L., at the edge of a picnic area in cotton-
wood-willow forest along Dry Creek.

On 6 April 1974, a female N. milbeHi was taken in North
Sacramento, Sacramento County, at an elevation of about 20
feet. It was on flowers of wild radish, Raphanus satimis L., at
the edge of riparian cottonwood forest with an extensive under-
story of U. urens; the native U. holosericea Nutt, is also present.
With the N. milberti were several Vanessa atalanta L., and
larvae of this species were found on the native nettle. It is
noteworthy that in both lowland records of N. milberti the
colony of U. urens was the only occurrence of this plant for
many miles in all directions. The close association of two females
with such plants suggests at least a significant possibility of
breeding at low elevations in spring.

13(3):157-161, 1974

ALTITUDINAL MIGRATION

161

As a working hypothesis it may be proposed that N. milberti,
like N. californica, may overwinter and rear a spring brood at
low elevations, disperse upslope for summer breeding, and
return to the lowlands in autumn. Because of its scarcity, this
movement could easily escape notice in normal years.

LITERATURE CITED

EMMEL, T. C. and J. F. EMMEL. 1962a. Ecological studies of Rhopalocera
in a high Sierran community — ^Donner Pass, California. I. Butterfly
associations and distributional factors. /. Lepid. Soc. 16: 23-44.

1962b. Note regarding habitats and Rhopalocera of Donner Pass,

California. Ibid. 16: 136.

— — ~ 1963a. Ecological studies of Rhopalocera in a high Sierran com-
munity— ^Donner Pass, California. IT Meteorologic influence on flight
activity. Ibid. 17: 7-20.

1963b. Composition and relative abundance in a temperate zone

butterfly fauna. /. Res. Lepid. 1: 97-108.

— 1974. Ecological studies of Rhopalocera in a Sierra Nevadan com-
munity— ^Donner Pass, California. V. Faunal additions and foodplant
records since 1962. J. Lepid. Soc., in press.

GARTH, J. S. and J. W. TILDEN. 1963. Yosemite butterflies. /. Res.
Lepid. 2: 1-96.

SHAPIRO, A. M. 1973. Host records for Brephidium exilis ( Lycaenidae ) .
J. Lepid. Soc. 27: 157-158.

1974a. Movements of Nymphalis calif ornica ( Nymphalidae) in

1972. /. Lepid. Soc. 28: 75-78.

— 1974b. Altitudinal migration of butterflies in the central Sierra

Nevada. J. Res. Lepid. 12: 231-235.

Journal of Research on the Lepidoptera

13(3): 162-168, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

THE CHROMOSOMES OF

APANTESIS PHALERATA, A. RADIANS,

AND THEIR HYBRID

IN FLORIDA POPULATIONS (ARCTHDAE)*

JACK S. BACHELER and THOMAS C. EMMEL

Departments of Entomolosu and Zoolosu, University of Florida,
Gainesville 32601

OcuRRiNG WIDELY in the southeastern portion of North Amer-
ica are two very similar and commonly confused species of
Apantesis, or “tiger moths.” Apantesis phalerata (Harris), a
more broadly distributed moth, has recently been shown in
northern Florida populations (Bachelor, 1972) to be specifically
distinct from A. radians Walker in several biological and mor-
phological characters. The adults of these two species and their
hybrid are illustrated in Fig. 1. We wish to report here the
surprising differences in chromosome numbers between the two
closely related species, and their experimentally-produced hy-
brids. These chromosome counts represent the first published
data for species in this arctiid genus.

METHODS

Meiotic divisions in the Lepidoptera are most easily observed
in testes of male adults (Emmel, 1969) but spermatogenesis was
essentially completed in almost all Apantesis adults examined.
Further preliminary studies showed that germ cell division in
A. phalerata and A. radians occurred to the greatest degree
during the penultimate instar of male larvae. Thus the standard
procedure for chromosome preparations previously reported
(Emmel, 1969) was modified as follows.

Testicular tissues of late penultimate A. phalerata, A. radians,
and A. phalerata $ X A. radians S hybrid larvae were fixed
by injecting a 3:1 absolute ethyl alcohol : glacial acetic acid
mixture into the sixth abdominal segment with a no. 27 hypo-
dermic syringe. The injected larvae were placed into four ounce
jars containing the same fixative, labeled, and refrigerated at
about — 20° C for later dissection.

Tla. Agricultural Experiment Station Journal Series No. 4560.

162

163

03

.§T.

ft

13(3):162-168, 1974 CHROMOSOMES OF APANTESIS

164

J. S. BACHELER AND T. C. EMMEL

/. Res, Lepid.

For chromosome study the testes, usually found in the dorsal
area of the sixth abdominal segment of male larvae, were re-
moved with No. 5 watchmaker’s forceps and placed onto a
standard microscope slide. The testes were then macerated and
a few drops of Lacto-Aceto-Orcein Stain (Emmel, 1969) added.
This preparation was allowed to stand for about 10-12 min. The
stained testes were next covered with a coverslip and the prepa-
ration squashed with about 400 Ib/in.^ pressure between two
pieces of blotting paper. The perimeter of the coverslip was
then sealed with clear lacquer.

Slides were inspected with a Zeiss Research Microscope
STANDARD WL fitted with 25X and 40X plan-apochromatic
field objectives. An oil-immersion Planapo lOOX objective was
used for critical observations.

Chromosome counts were made during meiotic division I
when chromosomes were paired in synapsis. Photographs were
taken of unusually clear chromosomes sets and of any interesting
anomalies.

All larval material was reared in the laboratory from wild
females of both species collected in populations in the vicinity of
Gainesville, Alachua County, Florida. Hybrid larvae were ob-
tained from laboratory crosses of virgin adults of the two species.

RESULTS AND DISCUSSION

Definitive chromosome counts with photographic confirma-
tion were obtained from microscopic slide examination of phal-
erata, radians, and hybrid larval testes squashes. The apparent
haploid chromosome numbers of phalerata, radians, and the
hybrid cross are n = 29, 30, and 29 respectively (Fig. 2). Since
no other counts have been reported in the genus the interpre-
tation of the chromosomal evidence for the phylogenetic re-
lationship between these two species is necessarily limited and
speculative. However, there are two principal ways in which
a difference in lepidopteran chromosome numbers of this type
may be explained and some evidence exists to indicate which
hypothesis is more tenable.

1. Reduction in Number by Fusion or Loss

The karyotypic origin of phalerata (n = 29) could have
resulted from the fusion of two of radians' (or an ancestor of
radians) 30 chromosomes, or a loss of one ancestral chromo-
some. A reduction in chromosome number from a characteristic

13(3):162-168, 1974 CHROMOSOMES OF APANTESIS

165

lepidopteran haploid set of 31 is not infrequent in the arctiids
and closely related noctuids; in counted species of both families
the mean haploid number is 31 chromosomes (Robinson, 1971).
The reported arctiid counts show one species with a haploid
number of 26, four with 28, one with 29, two with 30, seventeen
with 31, and one species with a number varying between 30
and 33. The noctuids show the same apparent tendency toward
reduction in chromosome number for the probable ancestral
number of 31, with seven species having less than 31 and three
exceeding 31.

Thus it is possible that radians resulted from the fusion of
two chromosomes in an n = 31 ancestor, and that phalerata was
the result of a further fusion of two radians chromosomes. These
fusions would reduce slightly the amount of possible genetic
recombination and thus the general adaptive flexibility of the
species. But fusion would be an evolutionarily favorable event
for a well adapted species where the need for genetic recom-
bination would be minimally valuable, even selectively disad-
vantageous.

If phalerata had indeed evolved with fusion from an ances-
tor with a haploid number of 30, such as radians, microscopic
examination of its 29 chromosomes might show one double-
sized or exceptionally large chromosome. However, photo-
graphic examination did not disclose a significantly larger chro-
mosome. Thus it seems more probable that loss of a chromosome
from the radians complement or loss of two chromosomes from a
common ancestral complement represents a plausible explana-
tion of the origin of phalerata’ s n = 29 set of chromosomes.

2. Hypothesis Involving Increase in Number by Fission

(Fragmentation)

The possibility also exists that phalerata (or an n = 29 an-
cestor of phalerata) gave rise to radians (or an ancestor of
radians) by fragmentation of one of its chromosomes. The
evolution of many lepidopteran species has been accompanied
by an increase in chromosome number (Emmel, 1972; Emmel
and Trew, 1973). Thus, one of phalerata’ s 29 chromosomes
could have split, with both parts being retained in the new
genotype, resulting in a new chromosome complement of 30.
The selective advantage of this fragmentation would be in-
creased potential for genetic recombination, allowing the spe-
cies to be more generally adaptable. (Naturally, the ancestral
number of 29 in phalerata would likely have been derived
originally by reduction through loss or fusion. )

166

J. S. BACHELER AND T. C. EMMEL

/. Res. Lepid.

Fig. 2. — Photomicrographs of testes squashes at lllOX magnification;
chromosomes in metaphase during first meiotic division, (a-c) Apantesis
phalerata, brood L-60 (n = 29); (d) A. radians, F^^ brood D-62 (n =
30); (e,f) A. radians $ x A. phalerata $ , brood DL-6261 (n = 29).

13(3):162~168, 1974 CHROMOSOMES OF APANTESIS

167

There is some support for the above fragmentation hypothesis
in phalemta and radians. If radians' karyotype had resulted
from the fragmentation of one of the ancestral phalerata (or
similar n = 29 species) chromosomes, t’wo smaller chromosomes
might be found among the 30. Tv^o radians chromosomes (Fig.
2d) do appear smaller than the others.

If radians had evolved from phalerata, one probably would
expect radians range to be more restricted, having split oflE
rather recently ( suggested by their almost identical morphology,
life history, and occasional laboratory hybridization). Such is
the case, with radians occupying about a third the area that
phalerata does.

EVIDENCE FROM THE HYBRIDS OF THE TWO SPECIES

If one accepts the ancestral fragmentation hypothesis as
more likely, the hybrid chromosome number of 29 is most easily
explained by the two short chromosomes of the radians parent
pairing with their “former” chromosome from the phalerata
parent in the hybrid larvae testes (at meiosis I). A uniform
count of 29 in meiosis II cells of the hybrid larvae would result
from the loss of one of the two small chromosomes after pairing
in meiosis I (the loss of both small chromosomes would have
meant the probable additional loss of the unpaired phalerata
chromosome, resulting in a hybrid count of 28, or at least a
variable count of 28 and 29 in meiotic (II) cells of the hybrid
adult ) .

If one follows the ancestral fusion hypothesis (Section I),
the picture in the hybrid would be superficially similar (n = 29
at meiosis I ) , only the two complete chromosomes of the radians
parent would pair with the large, “fused” chromosome of the
derivative phalerata parent in the hybrid larval testes cells. The
testes of such a hybrid male larva should have a variable num-
ber of chromosomes (28 or 29) in meiotic cells, or if all cells
were n = 29 (due to the loss of just one radians parental chro-
mosome), about half the cells in meiosis II should have one
chromosome noticeably smaller than the other 28 and half the
cells should have all 29 chromosomes appearing as uniformly-
sized.

Unfortunately, our material did not provide sufficient plates
of material in meiosis II to ascertain whether such a chromo-
somal loss was indeed the case.

168

J. S. BACHELER AND T. C. EMMEL

J. Res. Lepid.

SUMMARY

Apantesis phalerata and A. radians are two closely related,
phenotypically similar arctiid species found sympatrically in
northern Florida. Examination of meiotic divisions in testes of
penultimate instar larvae indicated that the haploid chromo-
some numbers for the two species are different: n = 29 for
phalerata and n = 30 for radians. Their hybrid has a chromo-
some number of n = 29. These are the first cytological obser-
vations reported from the genus Apantesis.

ACKNOWLEDGMENTS

This research was supported in part by a USDA Cooperative
Agreement (12-14-100-9397(33) D. H. Habeck, principal in-
vestigator) and by grants from the National Science Foundation
(GB8442 and GB32151, T. C. Emmel, principal investigator).
We thank Dr. Dale H. Habeck, Department of Entomology
and Nematology, University of Florida, for advice, support, and
facilities.

LITERATURE CITED

BACHELER, JACK S. 1972. Biology and hybridization of Apantesis phal-
erata (Harris) and A. radians Walker in Florida ( Lepidoptera: Arc-
tiidae). Department of Entomology and Nematology, University of
Florida, Ph.D. thesis. 89 pp.

EMMEL, THOMAS C. 1969. Methods for studying the chromosomes of
Lepidoptera. Joiirn. Res. Lepid. 7: 23-28 (“1968”)-

, 1972. Mate selection and balanced polymorphism in the tropical

nymphalid butterfly, Anartia fatima. Evolution 26: 96-107.

EMMEL, THOMAS C., and HOUSTON R. TREW. 1973. The chromo-
somes of skipper butterflies from southwestern North America (Lepi-
doptera, Hesperiidae). Cytologica, 38: 45-53.

ROBINSON, ROY. 1971. Lepidoptera Genetics. Pergamon Press, Oxford.
687 pp.

Journal of Research on the Lepidoptera

13(3):169-178, 207-216, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A PARTIAL BIBLIOGRAPHY

OF THE WORLD DISTRIBUTION AND
ZOOGEOGRAPHY OE BUTTERFLIES

OAKLEY SHIELDS

Department of Entomology,

University of California, Davis, California 95616

INTRODUCTION

This bibliography ( albeit incomplete ) is intended as a guide
to the world’s geographical distribution of butterflies, with
emphasis on broader, more comprehensive works. Additional
sources of major importance are contained in Lepidopterorum
Catalogus, Das Tierreich, Genera Insectorum, Seitz’ Macrolepi-
doptera of the World, Zoological Record (Insecta), and “Recent
Literature on Lepidoptera” series in Lepidopterists News
{—Journal Lepidopterists' Society). Perhaps others will con-
tribute further to the task of completing a thorough listing.

With the past decade of a major revolution in geology,
“continental drift” has reappeared in the form of plate tectonics
and sea-floor spreading. Butterflies are such a well-studied group
zoogeographically that their distributional patterns eventually
may be illuminated by the new ideas from geology (e.g. see
Holloway, 1969). This bibliography was compiled in conjunc-
tion with my own studies (in progress) of the biogeography of
organisms and plate tectonics, and is offered as an aid to others
engaged in zoogeographical work.

I would like to thank the Interlibrary Loan Department at
U.C. Davis for tolerating my many requests. This study was
financed in part by an N. S. E. graduate traineeship.

BIBLIOGRAPHY

AHMAD, T., 1949. Insect fauna of Afghanistan IV. Lepidoptera. Indian
Journal Entomology, 8 (2): 202-223.

ANDREWS, C. W., 1900. A Monograph of Christmas Island (Indian
Ocean): Physical Features and Geology. 337 pages, 21 plates. London:
Stephen Austin & Sons. (A. G. Butler — Lepidoptera, Rhopalocera, pp.
60-63, pi. IX)

169

170

OAKLEY SHIELDS

/. Res. Lepid.

ANNANDALE, N., and C. DOVER, 1921, The butterflies of Barkuda
Island. Indian Museum Records, 22: 349-375.

ASAHINA, E., 1932. Butterflies of the Bonin Islands. Zephyrus, 4; 219.
[In J[3,p3.n0S0]

AVINOFF, A., and N. SHOUMATOFF, 1946. An annotated list of the
butterflies of Jamaica. Annals Carnegie Museum, 30; 263-295, 1 plate.
AVINOFF, A., and W. R. SWEADNER, 1951. The Karanasa butterflies, a
study in evolution. Annals Carnegie Museum, 32: 250 pages, 17 plates.
BANG-HAAS, O., editor, 1929-1930. Novitates Macrolepidopterologicae.
Katalog der N eubeschreibungen von Palaearktischen Macrolepidop-
teren. Band IV (1929), 80 pages; Band V (1930), 207 pages. [Good
Rhopalocera bibliography]

BARGANT, M., 1970. Butterflies of Trinidad and Tobago. 314 pages, 28
plates. London: Gollins.

BARRETT, G., and A. N. BURNS, 1951. Butterflies of Australia and New
Guinea. 187 pages. Melbourne: N. H. Seward.

BARTHOLOMEW, J. G., W. E. GLARKE, and P. H. GRIMSHAW, 1911.
Atlas of Zoogeography. Bartholomew’s Physical Atlas, vol. 5. Edin-
burgh: John Bartholomew & Go.

BATES, H. W., 1867. A catalogue of Erycinidae, a family of diurnal Lepi-
doptera. Journal Linnean Society London, Zoology, 9: 367-372.

BATES, M., 1935. The butterflies of Guba. Bulletin Museum Comparative
Zoology, 78 (2): 63-258.

BEATTY, H. A., 1944. The insects of St. Groix, V. I. University Puerto
Rico, Journal Agriculture, 28: 114-172. [see pp. 156-158]

BERGER, L., 1939. Resultats scientifiques des croisieres du navire-ecole
Beige “Mercator”. Lepidopteres-Rhopaloceres. Mem. Mus. Roy. Hist.
Nat. Belgique, 19 (15): 181-207.

BERNARDI, G., 1954. Revision des Pierinae de la faune malgache (Lepid.
Pieridae). Mem. Inst. Sci. Madagascar, Tananarive (E), 5; 239-375,
4 plates, 141 figures.

BERNARDI, G., 1961. Biogeographie et speciation des Lepidopteres Rho-
paloceres des lies Mediterraneennes. France, Centre Nat. Rech. Sci.
Coll. Intern., 94: 181-215. [Extensive bibliography]

BETHUNE-BAKER, G. T., 1925. A revision of the Liphyrinae together with
a description of the structure of the puparium of Liphyra brassolis and
of the pupae of Aslauga vininga and A. lamborni ( Lepidoptera ) .
Transactions Entomological Society London, 72: 199-238.

BIEZANKO, G. M. de, A. RUFFINELLI, and G. S. GARBONELL, 1957.
Lepidoptera de Uruguay: lista anotada de especies. Revta Fac. Agron.
Montivideo, 46: 1-152.

BODENHEIMER, F. S., 1935. Animal Life in Palestine; an Introduction
to the Problems of Animal Ecology and Zoogeography. 506 pages.
Jerusalem: L. Mayer.

BOISDUVAL, J. A., 1832. Voyage de I’Astrolabe. Faune entomologique de
I’ocean pacifique. Premiere partie. Lepidopteres. Paris,

BOURNE, G. G., 1886. General observations on the fauna of Diego Garcia,
Ghagos Group. Proceedings Zoological Society London, 1886: 331-334.
BRITTON, E. B., and 29 others, 1970. The Insects of Australia. 1029 pages.

Carlton, Victoria: Melbourne University Press.

BROOKS, C. J., 1950. A revision of the genus Tenaris Hiibner (Lepidop-
tera: Amathusiidae ) . Transactions Royal Entomological Society Lon-
don, 101: 179-238, 8 plates.

BROWN, F. M., 1971. Review: A Field Guide to the Butterflies of Britain
and Europe, by L. G. Higgins and N. D. Riley. Journal Lepidopterists’
Society, 25: 152-154.

BROWN, F. M. and B. HEINEMAN, 1972. Jamaica and its Butterflies.
478 pages, 10 plates. London: E. W. Classey Ltd.

13(3):169-178, 207-216, 1974

BIBLIOGRAPHY

171

BROWN, K. S., and O. H. H. MIELKE, 1967. Lepidoptera of the Central
Brazil Plateau, I. Preliminary list of Rhopalocera: introduction, Nym-
phalidae, Libytheidae, Journal Lepidopterists" Society, 21 (2): 77-106.

1967. Lepidoptera of the Central Brazil Plateau. I. Preliminary list

of Rhopalocera ( continued ) : Lycaenidae, Pieridae, Papilionidae, Hes-
periidae. Journal Lepidopterists’ Society, 21: 145-168.

— 1968. Lepidoptera of the Central Brazil Plateau. III. Partial list for

the Belo Horizonte area, showing the character of the southeastern
“blend zone.” Journal Lepidopterists Society, 22: 147-157.

BRYAN, E. H., Jr., and collaborators, 1926. Insects of Hawaii, Johnston
Island and Wake Island. B. P. Bishop Museum Bulletin, 31: 1-94.
BURNS, A., 1969. Australian Butterflies in Colour, 112 pages. Sydney:
A. H. and A. W. Reed.

BUTLER, A. G., 1869. Catalogue of Diurnal Lepidoptera. 303 pages. Lon-
don: Taylor and Francis.

1874. List of the diurnal Lepidoptera of the South-Sea islands. Pro-
ceedings Zoological Society London, 1874: 274-291.

— — 1875. On a collection of butterflies from the New Hebrides and

Loyalty Islands, with descriptions of new species. Proceedings Zoologi-
cal Society London, 1875: 610-619.

— 1875. On a small collection of butterflies from Fiji. Proceedings

Zoological Society, London, 1875: 619-620.

- — — ■ 1876. The butterflies of Malacca. Journal Linnean Society London,
Zoology, 13: 196-197.

— — — 1878. On the butterflies of New Zealand. Transactions Proceedings
New Zealand Institute, 10: 263-276.

— — — 1883. The Lepidoptera collected during the recent expedition of
H. M. S. ‘Challenger.’ Annals Magazine Natural History London, Series
5, 11: 402-428.

— 1884. The Lepidoptera collected by Herr Palisa at Caroline Island

in May, 1883. Washington National Academy Memoirs, 2: 93-96.

- — — - 1884. Lepidoptera from the island of Nias. Annals Magazine Natural
History, 13: 58-60.

1884. The Lepidoptera collected during the recent expedition of

H. M. S. ‘Challenger’ Part II. Annals Magazine Natural History,
Series 5, 13: 183-203.

— • 1885. Lepidoptera collected by Mr. C. M. Woodford in the Ellice

and Gilbert Islands. Annals Magazine Natural History, Series 5, 15:
238-242.

1887. Lepidoptera, In: Report on the Lepidoptera collected by the

officers of H. M. S. ‘Flying-Fish’ on Christmas Island. Proceedings
Zoological Society London, 1887 : 522-523.

— — 1888. On the Lepidoptera of Christmas Island. Proceedings Zoologi-
cal Society London, 1888: 542-546.

1898. On a collection of Lepidoptera made in British East Africa by

Mr. C. S. Betton. Proceedings Zoological Society London, 1898: 395-
444.

— 1900. On two consignments of butterflies collected by Mr. Richard
Crawshay in the Kikuyu Country of British East Africa in 1899 and
1900. Proceedings Zoological Society London, 1900: 911-946.

— 1901. On some butterflies from St. Lucia, W. Indies, collected by

Major A. H. Cowie, R. E., F. Z. S. Proceedings Zoological Society
London, 1901 (II): 711-714.

BUTLER, A. G., and H. DRUCE, 1874. List of the butterflies of Costa
Rica, with descriptions of new species. Proceedings Zoological Society
London, 1874: 330-370.

1879. The butterflies of Malacca. Transactions Linnean Society Lon-
don, Series 2, vol. 1, Zoology, 1879: 533-568.

172

OAKLEY SHIELDS

/. Res. Lepid.

CAMPOS, R. E., 1927. Catalogo preliminar de los lepidopteros del Ecuador,
Primera parte, Rhopalocera. Rev. Colegio Nac. Vicente Rocafuerte, 9:
1-106.

CARCASSON, R. H., 1961. The Acraea butterflies of East Africa (Lepidop-
tera, Acraeidae). Journal East African Natural History Society, Special
Supplement no. 8, 45 pages.

1964. A preliminary survey of the zoogeography of African butter-
flies. East African Wildlife Journal, 2: 122-157.

CARPENTER, G. D. H., 1935. The Rhopalocera of Abyssinia: a faunistic
study. Transactions Royal Entomological Society London, 83 (III):
313-440, plates VIII-XII.

1939. Preliminary account of an investigation of the Euplosine but-
terflies of Melanesia. Proceedings Sixth Pacific Science Congress, 4:
305-309.

1953. The genus Eiiploea ( Lep. Danaidae) in Micronesia, Mela-
nesia, Polynesia and Australia. A zoo-geographical study. Transactions
Zoological Society London, 28: 1-184, 9 plates. [Excellent bibliogra-
phy]

CARPENTER, G. D. H., and C. B. Lewis, 1943. A collection of Lepidop-
tera (Rhopalocera) from the Cayman Islands. Annals Carnegie Mu-
seum, 29: 371-396.

CHERMOCK, R. L., 1950. A generic revision of the Limenitini of the
world. American Midland Naturalist, 43: 513-569.

CHNeOUR, a., 1954. Macrolepidopteres de Tunisie I, II. — Rhopalocera,
Grypocera. Societe Sciences Naturelles Tunisie, Bulletin, 7: 207-239,
plates 29-34.

CLARK, A. H., 1904. Notes on the insects of Barbados, St. Vincent, the
Grenadines, and Grenada. Psyche, 11: 114-117.

1927. Geography and zoology. Annals Association American Geogra-
phers, 17: 101-145. [Section on butterflies]

1946. Two new butterflies from the Admiralty Islands. Proceedings

Biological Society Washington, 59: 119-120.

1951. Butterflies of the Marshall Islands ( Lepidoptera ) . Proceedings

Entomological Society Washington, 53: 43-44.

CLARK, J. F. G., 1971. The Lepidoptera of Rapa Island. Smithsonian Con-
tributions Zoology, no. 56, 282 pages.

CLENCH, H. K., 1942. The Lycaenidae of the Bahama Islands (Lepidop-
tera, Rhopalocera). Psyche, 49: 52-60.

1955. Revised classification of the butterfly family Lycaenidae and

its allies. Annals Carnegie Museum, 33: 261-274.

1963. A synopsis of the West Indian Lycaenidae, with remarks on

their zoogeography. Journal Research Lepidoptera, 2: 247-270.

1964. Remarks on the relationships of the butterflies ( excluding skip-
pers) of the Cayman Islands. Occasional Papers on Mollusks, 2 (31):
381-382.

1965. A collection of butterflies from western Chihuahua, Mexico.

Entomological News, 76: 157-162.

CLENCH, H. K., and N. SHOUMATOFF, 1956. The 3rd Danish Expedi-
tion to Central Asia. Zoological Results 21. Lepidoptera Rhopalocera
(Insecta) from Afghanistan. Vidensk. Medd. fra Dansk naturh. Foren.,
118: 141-191, 1 plate.

COCKERELL, T. D. A., 1939. The insects of the Californian islands. Pro-
ceedings Sixth Pacific Science Congress, 4: 283-295.

COGAN, B. H., A. M. HUTSON, and J. C. SHAFFER, 1971. Preliminary
observations on the affinities and composition of the insect fauna of
Aldabra, Philosophical Transactions Royal Society London (B), 260:
315-325.

COLE, A. C., 1951. Insects from Bikini Atoll, Marshall Islands. Journal
Tennessee Academy Science, 26: 246-248.

13(3):169-178, 207-216, 1974

BIBLIOGRAPHY

173

COLLENETTE, C. L., 1925. The present status of Danaida plexippus L.
in the Pacific Islands. Entomol. Monthly Magazine, 61: 198-202.

COMMON, I. F. B., 1964. Australian Butteijlies. 131 pages. Brisbane: The
Jacaranda Press.

COMSTOCK, J. A., 1966. Lepidoptera of American Samoa with particular
reference to biology and ecology. Pacific Insects Monograph, 11: 1-74.

COMSTOCK, W. P., 1944. Insects of Porto Rico and the Virgin Islands:
Rhopalocera or butterflies. New York Academy Science, Scientific Sur-
vey of Porto Rico and the Virgin Islands, 12 (pt. 4): 421-622.

1961. Butterflies of the American Tropics: the Genus Anaea (Lepi-
doptera: Nymphalidae). 214 pages, 30 plates. New York: The Ameri-
can Museum of Natural History.

COMSTOCK, W. P., and E. I. HUNTINGTON, 1943. Lycaenidae of the
Antilles (Lepidoptera, Rhopalocera). Annals New York Academy Sci-
ences, 45: 49-130, 1 plate.

— — 1949. Origins and relationships of Mexican and Antillean Papiliono-
idea (Lepidoptera). An. Inst. Biol. Mex., 20: 385-391.

CORBET, A. S., 1941. The distribution of butterflies in the Malay Penin-
sula (Lepid.). Proceedings Royal Entomological Society London (A),
16: 101-116.

— 1944. Some notes on the butterflies of the Langkawi Islands. The
Entomologist, 77: 39-42.

— — - 1948. Papers on Malaysian Rhopalocera, V: the conspecificity of the
American Precis lavinia (Cramer) with the Oriental Precis orithya
(Linnaeus). The Entomologist, 81: 54-56,

— — 1948. Observations on the species of Rhopalocera common to Mada-
gascar and the Oriental region. Transactions Royal Entomological
Society London, 99: 589-607.

- — — 1948. Papers on Malaysian Rhopalocera, III.: The butterflies of
Singapore Island. The Entomologist, 81: 9-14.

CORBET, A. S., and H. M, PENDLEBURY, 1956. The Butterflies of the
Malay Peninsula, 2nd edition. 537 pages, 55 plates. London: Oliver
and Boyd.

COSTA LIMA, A. Da, 1949 [1950]. Insectos do Brazil. VoL 6, part 2.
Lepidoptera. Escola Nacional de Agronomia, Serie Diddtica, no. 8, 420
pages.

COUCHMAN, L. E., 1955 [1956]. A catalogue of the Tasmanian Lepidop-
tera-—Rhopalocera. Papers & Proceedings Royal Society Tasmania, 90:
1-33.

D’ABRERA, B., 1971. Butterflies of the Australian Region. 415 pages,
numerous colored plates. Melbourne: Lansdowne Press.

DEWITZ, H., 1877. Tagschmetterlinge von Porto Rico. Stett. Entomo-
logische Zeitschrift, 38 : 233-245.

DR. DIEHL, 1954. Liste des Lepidopteres Rhopaloceres rencontres au
sud de Madagascar et surtout sur le plateau du sud. Naturaliste Mal-
gache, 6: 61-70.

DILLON, L. S., 1947. Some new subspecies of butterflies from Dominica,
B. W. I. Bulletin Brooklyn Entomological Society, 42: 97-102.

DIXEY, F., 1894. On the phylogeny of the Pierini, as illustrated by their
wing markings and geographical distribution. Transactions Entomo-
logical Society London, 1894: 249-334.

DOHERTY, W. A., 1886. A list of butterflies taken in Kumaon. Journal
Asiatic Society Bengal, 55: 103-140.

1891. The butterflies of Sumba and Sambava, with some account of

the Island of Sumba. Journal Asiatic Society Bengal, 1891: 141-197.

dos PASSOS, C. F., 1958. The satyrid butterflies of northwestern North
America (Lepidoptera: Satyridae). Proceedings 10th International
Congress Entomology, 1: 673-681.

174

OAKLEY SHIELDS

/. Res. Lepid.

1964. A synonymic list of the Nearctic Rhopalocera. The Lepidop-

terists Society Memoir no. 1, 145 pages.

dos PASSOS, C. F., and L. P. GREY, 1945. A genitalic survey of Argyn-
ninae ( Lepidoptera, Nymphalidae). American Museum Novitates, no.
1296, 29 pages.

DOWNES, J. A., 1966. The Lepidoptera of Greenland: some geographic
considerations. Canadian Entomologist, 98: 1135-1144.

DRUCE, H. H., 1873. A list of the collections of diurnal Lepidoptera made
by Mr. Lowe in Borneo, with descriptions of new species. Proceedings
Zoological Society London, 1873: 337-360.

1876. List of the butterflies of Peru, with descriptions of new species.

Proceedings Zoological Society London, 1876: 205-250.

1891. On the Lycaenidae of the Solomon Islands. Proceedings Zoo-
logical Society London, 1891: 357-373, plates XXXI-XXXII.

1892. A list of the Lycaenidae of the South Pacific Islands east of

the Solomon Group, with descriptions of several new species. Pro-
ceedings Zoological Society London, 1892: 434-446, plate XXVII.

1895. A monograph of the Bornean Lycaenidae. Proceedings Zoo-
logical Society London, 1895: 556-627.

1896. Further contributions to our knowledge of the Bornean Lyca-
enidae. Proceedings Zoological Society London, 1896: 650-683.

DYAR, H. G., 1915. Lepidoptera of the Yale-Dominican Expedition of 1913.
Proceedings United States National Museum, 47 : 423-426.

EBERT, H., 1969. On the frequency of butterflies in eastern Brazil, with
a list of the butterfly fauna of Pocos de Galdas, Minas Gerais. Journal
Lepidopterists Society, 23, Supplement 3, 48 pages.

EBNER, J. A., 1971. Some notes on the Papilionidae of Manus Island, New
Guinea. Journal Lepidopterists’ Society, 25: 73-80.

EDWARDS, W. H., 1884. The Butterflies of North America. Vol. I, II,
III. Unnumbered pages. Boston: Houghton, Mifflin, and Company.

EHRLICH, P, R., 1958. Problems of arctic alpine insect distribution as
illustrated by the butterfly genus Erebia (Satyridae). Proceedings 10th
International Congress Entomology, 1: 683-686.

EHRLICH, P. R., and A. H. EHRLICH, 1961. How to Know the Butter-
flies. 262 pages. Dubuque, Iowa: Wm. C. Brown Co.

EHRLICH, P. R., and P. H. RAVEN, 1965. Butterflies and plants: a study
in coevolution. Evolution, 18: 586-608.

ELIOT, J. N., 1969. An analysis of the Eurasian and Australian Neptini
(Lepidoptera: Nymphalidae). Bulletin British Museum Natural His-
tory, Entomology Supplement, 15: 1-155.

1973. The higher classification of the Lycaenidae (Lepidoptera): a

tentative arrangement. Bulletin British Museum Natural History, En-
tomology, 28 (6): 371-505, 6 plates.

ELIOT, N., 1946. Continental drift and Precis lavinia. The Entomologist,
79: 225-228.

1947. More on continental drift, Precis lavinia Hb. and P. villida F.

The Entomologist, 80: 230-234.

1947. More on continental drift, Precis lavinia Hb. and P. villida F.:

a postscript. The Entomologist, 80: 283.

— 1949. The significance of the wing pattern of Precis (Nymphalidae)

in America. The Entomologist, 82: 176-184.

ELWES, H. J., 1881. On the butterflies of Amurland, North China, and
Japan. Proceedings Zoological Society London, 1881: 856-916.

1891. On butterflies collected by Mr. W. Doherty in the Naga and

Karen Hills and in Perak.— Part 1. Proceedings Zoological Society
London, 1891: 249-289.

1894. [Geographical distribution of butterflies.] Transactions Ento-
mological Society London, 1894: lii-lxxxiv.

13(3);169-178, 207-216, 1974

BIBLIOGRAPHY

175

1903. The butterflies of Chile. Transactions Entomological Society

London, 1903: 263-301.

EMSLEY, M. G,, 1965. Speciation in Heliconius (Lep., Nymphalidae) :
morphology and geographic distribution. Zoologica, 50; 191-254.

ESAKI, T., 1933-1934. Butterflies of the northern Kurile Islands. Bulletin
Bio geo graphical Society Japan, 4: 83-85.

EVANS, W. H., 1912. A list of Indian butterflies. Journal Bombay Natural
History Society, 21: 553-584, 969-1008.

— — — [1926]. Fauna Buruana. Lepidoptera, fam. Grypocera (Hesperi-
idae). Treuhia, 7(1); 371-375.

— 1932. The butterflies of Baluchistan. Journal Bombay Natural His-

tory Society, 36: 196-209.

1932. The Identification of Indian Butterflies, 2nd edition. 454

pages, 32 plates. Madras: The Diocesan Press.

1947. Hesperiana. Proceedings Entomological Society Washington,

49: 162-163.

— 1949. A Catalogue of the Hesperiidae from Europe, Asia and Aus-

tralia in the British Museum (Natural History). 502 pages, 53 plates.
Norwich: Jarrold and Sons, Ltd.

— 1951. A Catalogue of the American Hesperiidae in the British Mu-
seum (Natural History). Part I. Pyrrhopyginae. 92 pages, 9 plates.

— 1952. A Catalogue of the American Hesperiidae in the British Mu-

seum (Natural History). Part 11. Pyrginae. Section 1. 178 pages, plates
10-25. Norwich: Jarrold and Sons, Ltd.

1953. A Catalogue of the American Hesperiidae in the British Mu-
seum (Natural History). Part HI. Pyrginae. Section 2. 246 pages, plates
26-53. Norwich: Jarrold and Sons, Ltd.

1955. A Catalogue of the American Hesperiidae in the British Mu-
seum (Natural History). Part IV. Hesperiinae and Megathyminae. 499
pages, plates 54-84. Norwich: Jarrold and Sons, Ltd.

FERRAR, M. L., 1948. The butterflies of the Andamans and Nicobars.
Journal Bombay Natural History Society, 47: 470-491.

FERREIRA D’ ALMEIDA, R., 1966. Catdlogo dos Papilionidae Ameri-
canos. 366 pages. Sao Paulo: Sociedade Brasileira de Entomologia.

FIELD, W. D., 1971. Butterflies of the genus Vanessa and of the resur-
rected genera Bassaris and Cynthia (Lepidoptera: Nymphalidae),
Smithsonian Contributions Zoology, no. 84, 103 pages.

FIELD, W .D., C. F. dos PASSOS, and J. H. MASTERS, 1974. A bibliog-
raphy of the catalogs, lists, faunal and other papers on the butterflies
of North America north of Mexico arranged by state and province
(Lepidoptera: Rhopalocera ) . Smithsonian Contributions Zoology, no.
157, 104 pages. [Comprehensive bibliography]

FLETCHER, T. B., 1909. Lepidoptera, exclusive of the Tortricidae and
Tineidae, with some remarks on their distribution and means of dis-
persal amongst the islands of the Indian Ocean. Transactions Linnean
Society London, Series 2, Zoology, 13: 265-323, plate 17.

FORBES, W. T. M., 1941. The Lepidoptera of the Dry Tortugas. Psyche,
48: 147-148.

1947. Buckeyes and Wegener. The Entomologist, 80: 56-58.

FORD, E. B., 1957. Butterflies. 368 pages. London; Collins.

FORSTER, W., 1965. Rhopalocera et Hesperiidae Ergebnisse der zoologi-
schen Forschungen von Dr. Z. Kaszab in der Mongolei (Lepidoptera).
Reichenbachia, 7: 165-178.

FOX, R. M., 1940. A generic review of the Ithomiidae. Transactions Ameri-
can Entomological Society, 66: 161-207.

1956. A monograph of the Ithomiidae (Lepidoptera), part 1. Bulle-
tin American Museum Natural History, 111; 1-76, 9 plates.

FOX, R. M., A. W. LINDSEY, Jr., H. K. CLENCH, and L. D. MILLER,
1965. The butterflies of Liberia. Memoirs American Entomological
Society, no. 19, 438 pages.

176

OAKLEY SHIELDS

/. Res. Lepid.

FRANZ, E. and H. SCHROEDER, 1954, Tagfalter (Lep. Rhopalocera) aus
El Salvador, Senckenbergiana Biol., 35: 75-87, 2 figures.

FREEMAN, H. A., 1969. Systematic review of the Megathymidae. Journal
Lepidopterists Society, Supplement I, 23: 58 pages.

FREEMAN, T. N., 1956. The distribution of arctic and subarctic butterflies.
Proceedings 10th International Congress Entomology, 1: 659-672.

FROGGATT, W. W., 1907. Australian Insects. 449 pages. Sydney: William
Books and Gompany.

FRUHSTORFER, H., 1902. Beitrag zur Kenntniss der Lepidopteren der
Viti-Inseln. Stett. Ent. Zeit., 63: 350-359.

— 1908. Lepidopterologisches Pele-Mele. Ent. Zeit., 22: 48-49.

— — — 1912. Rhopaloceren der Isel Matty. Entomologische Rundschau, 29:
33-35

FRYER, J. G. F., 1912. The Lepidoptera of Seychelles and Aldabra, ex-
clusive of the Orneodidae and Pterophoridae and of the Tortricina and
Tineina. Transactions Linnean Society, Series 2, Zoology, 15: 1-28,
plate I,

FULDA, O., 1931. Sammeln in Haiti. Entomologische Rundschau, 48: 169-
170, 176-179, 194-196.

GALVANGI, E., 1912. Bericht fiber Marianem-und Samoa-Schmetterlinge.
Zool.~Bot. Ges. Wien, Verb., 62 (3, 4): 119-121.

de la GARDE, P., 1895. African Rhopalocera. The Entomologist, 28: 153-
155.

GERHARD, E., 1883. tiber die geographische Verbreitung der Macro-
Lepidopteren auf der Erde. Berlin Entomologische Zeitschrift, 27 : 173-
185. [Butterfly genera peculiar to various parts of the* world]

GIBBS, G. W., 1961. New Zealand butterflies. Tuatara, 9: 65-76.

GIFFORD, D., 1965. A List of the Butterflies of Malawi. 151 pages, 9
colored plates. Malawi: Published by the Sociey of Malawi.

GILLHAM, N. W., 1956. Nymphalis vau-album ( Schiffermuller & Denis),
a Holarctic species (Lepidoptera: Nymphalidae). Psyche, 63: 27-29.

GODMAN, F. D., and O. S ALVIN, 1878. Descriptions of some apparently
new species of butterflies from New Ireland and New Britain, sent by
the Rev. G. Brown. Proceedings Zoological Society London, 1878: 733-
735.

1879-1901. Biologia Centrali- Americana. Insecta. Lepidoptera — Rho-
palocera. Vol. I, 487 pages; vol. II, 782 pages. London: Published for
the editors by R. H. Porter.

— 1879. On some hitherto unrecorded diurnal Lepidoptera, from Duke-

of-York Island and New Ireland, with descriptions of some apparently
new species. Proceedings Zoological Society London, 1879: 652-655.

1880. Descriptions of some supposed new species of butterflies from

New Guinea. Proceedings Zoological Society London, 1880: 609-614.

1882. On some Rhopalocera from New Ireland. Proceedings Zoologi-
cal Society London, 1882: 754-755.

1884. A list of the Rhopalocera collected by Mr. G. French Angas

in the Island of Dominica. Proceedings Zoological Society London,
1884: 314-320, plate XXV.

— 1888. New species of butterflies collected by Mr. C. M. Woodford

in the Solomon Islands. Annals Magazine Natural History, 1888: 209-
214.

- — — 1896. On tlie butterflies of St. Vincent, Grenada, and the adjoining
islands of tlie West Indies. Proceedings Zoological Society London,
1896 (I): 513-520.

GRAPES, G. J., 1888. Insects caught at sea. The Entomologist, 21: 161-162.

GRAVES, P. P., 1925. The Rhopalocera and Grypocera of Palestine and
Transjordania. Transactions Entomological Society London, 73: 17-125.

GROTE, A. R., 1886. On the geographical distribution of North American
Lepidoptera. Canadian Entomologist, 18: 162-174.

13(3);169-178, 207-216, 1974

BIBLIOGRAPHY

177

1889. The lepidopterous faunae of Europe and North America.

Canadian Entomologist, 21; 21-24.

GUICHARD, K. M., 1967. Butterflies of the Ganary Islands. The Entomolo-
gist, 100: 293-299.

HALL, A., 1925. List of the butterflies of Hispaniola. The Entomologist,
58: 161-165, 186-190.

— 1936. The butterflies of St. Kitts. The Entomologist, 69: 274-278.

— — — 1939. Gatalogue of the Lepidoptera Rhopalocera (butterflies) of

British Guiana. Journal British Guiana, 1; 25-41, 96-104, 146-169,
215-254.

1940. Gatalogue of the Lepidoptera Rhopalocera (butterflies) of

British Guiana. Entomology Bulletin British Guiana, Department Agri-
culture, 3; 88 pages.

HAWKINS, G. N., 1942. The insects of Norfolk Island, including a pre-
liminary report on a recent collection. Annals Magazine Natural His-
tory, 11th Series, 9: 865-902.

HAYWARD, K. J., 1950. Gatalogo sinonimico de los Rhopaloceros argen-
tinos. Excluyendo Hesperiidae. Acta Zoologica Lilloana, 9: 281 pages.

HERBERT, W. H., 1867. Butterflies at sea. The Entomologist, 3: 226.

HERING, M., 1921. Die geographische Verbreitung der Libytheiden.
Archie fiir naturgeschichte. Berlin, Abt. A: Original - arbeiten. Vol. 87,
heft 4: 248-296, 2 plates.

HERMS, W. B., 1925. Entomological observations on Fanning and Wash-
ington Islands, together with general biological notes. Pan-Pacific
Entomologist, 2: 49-54.

HIGGINS, L. G., 1950. A descriptive catalogue of the Palearctic Euphy-
dryas (Lepidoptera: Rhopalocera). Transactions Royal Entomological
Society London, 101: 435-499.

— 1966. [Checklist of Turkish butterflies.] The Entomologist, 99: 209-

222.

HIGGINS, L. G., and N. D. RILEY, 1970. A Field Guide to the Butterflies
of Britain and Europe. 380 pages, 60 colored plates, 371 maps. Lon-
don: Collins.

HIROSE, T., 1934. [Butterflies from Marshall and Caroline Islands.] Trans-
actions Kansai Entomological Society, 5: 29-31. [In Japanese]

Hobby, B. M., 1940. Results of the Oxford University Expedition to Sara-
wak (Borneo), 1932. — Rhopalocera (Lepidoptera). Entomol. Monthly
Magazine, 76: 217-224.

HOFFMANN, C. C., 1936. Zoogeographische Beziehungen der Lepidop-
teren in Mexiko. Ann. Inst. Biol., 7 : ?

— 1940. Gatalogo sistematico y zoogeografico de los Lepidopteros

Mexicanos. Primera Parte. Anales Inst. Biol. Univ. Nac. Mex., 11:
639-739.

— 1941. Gatalogo sistematico y zoogeografico de los Lepidopteros Mexi-

canos. Segunda Parte. Hesperioidea. Anales Inst. Biol. Univ. Nac. Mex.,
12: 237-294.

HOLLAND, R., 1972, Butterflies of middle and southern Baja California.
Journal Research Lepidoptera, 11: 147-160.

HOLLAND, W. J., 1895. List of the Lepidoptera from Aldabra, Seychelles,
and other East African islands, collected by Dr. W. L. Abbott. Pro-
ceedings United States N ational Museum, 18: 265-273.

1900. The Lepidoptera of Burn. Part I. — Rhopalocera. Novitates

Zoologicae, 7: 54-86.

1916. The Lepidoptera of the Isle of Pines, being a list of the

species collected on the island by Mr. J. L. Graf and Mr. G. A. Link,
Sr., in 1910 and 1912-1913. Annals Carnegie Museum, 10: 487-518,
plate 31.

1920. Lepidoptera of the Congo. Bulletin American Museum Natural

History, 43: 109-369.

178

OAKLEY SHIELDS

J. Res. Lepid.

1930. The Butterfly Book, revised edition. 424 pages, 77 plates.

New York: Doubleday, Page and Company.

HOLLOWAY, J. D., 1969. A numerical investigation of the biogeography
of the butterfly fauna of India, and its relation to continental
drift. Biological Journal Linnean Society, 1: 373-385.

1970. The Application of Numerical Methods of Classification in

Biogeography: Exemplified by Studies on the Lepidoptera of the Indo-
Australian Area. Ph.D. thesis, University of Cambridge.

1973. The affinities within four butterfly groups (Lepidoptera: Rho-

palocera) in relation to general patterns of butterfly distribution in
the Indo- Australian area. Transactions Royal Entomological Society
London, 125 (2): 125-176.

HOLLOWAY, J. D., and N. JARDINE, 1968. Two approaches to zoo-
geography: a study based on the distributions of butterflies, birds and
bats in the Indo- Australian area. Proceedings Linnean Society London,
179 (2): 153-188.

HOPKINS, G. H. E., 1927. Butterflies of Samoa and some neighboring
island-groups. British Museum (Natural History), Insects of Samoa,
Part III, fasc. 1. Lepidoptera, 1-64 pages,

HOVANITZ, W., 1945. The distribution of Colias in the equatorial Andes.
Caldasia, 3: 283-300.

1945. Comparisons of some Andean butterfly faunas. Caldasia, 3:

301-306.

1950. The biology of Colias butterflies I. The distribution of the

North American species. Wasmann Journal Biology, 8: 49-75.

1952. Amphi-Atlantic study of Colias hecla, Colias nastes and Colias

palaeno. Yearbook American Philosophical Society, 1951: 142-144.

1958. Distribution of butterflies in the New World. Zoogeography,

AAAS, pp. 321-368.

HOWARTH, T. G., 1962. The Rhopalocera of Rennell and Bellona Islands.
Natural History Rennell Island, British Solomon Islands, 4: 63-83, 10
plates.

1963. The butterflies of the Windward Islands. The Windward

Islands Annual, 1963: 50-53.

HUDSON, G. V., 1898. New Zealand Moths and Butterflies (Macro-
Lepidoptera). 144 pages. London: West, Newman and Company.

de JOANNIS, J., 1894. Mission scientifique de M. Ch. Alluaud aux lies
Sechelles ( Mars-Avril-Mai 1892). Lepidopteres. Ann. Soc. Ent. France,
63: 425-438.

JOHNSON, F., and W. P. COMSTOCK, 1941. Anaea of the Antilles and
their continental relationships with descriptions of new species, sub-
species and forms (Lepidoptera, Rhopalocera, Nymphalidae). Journal
New York Entomological Society, 49: 301-343.

JOHNSTON, E. C., 1950. Lepidoptera of the Pribilof Islands, Alaska.
Lepidopterists News, 4: 27-30.

JONES, F. W., 1909. The fauna of the Cocos-Keeling Atoll, collected by
F. Wood Jones. Proceedings Zoological Society London, 1909 (I):
132-160.

KALIS, J. P. A., 1933. Lepidoptera Rhopalocera van Nederlandsch-Indie.
Tijdschr. Ent., 76: 47-86.

KAYE, W. J., 1925. The butterflies of Jamaica. Transactions Entomological
Society London, 73: 455-504.

KENRICK, G. H., 1911. Some undescribed butterflies of Dutch New
Guinea. Transactions Entomological Society London, 59: 16-20, plates
III-VI.

KHEIL, N. M., 1884. Die Lepidoptera der Isel Nias. Berlin.

KIMOTO, S., 1971. [Regional differences of the butterfly-fauna of Japan
and Ihe Ryukyu Archipelago.] Kontyu, 39 (3): 310-319. [In Japanese,
English summary]

( Continued on Page 207 )

Journal of Research on the Lepidoptera 13(3) :179-180, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

THE NOMENCLATURE IN AN
IMPORTANT BRITISH CHECK LIST (1972)
PART 2; CORRECTIONS OF FAMILY-GROUP
NAMES FOR GEOMETRIC AE (LEPIDOPTERA)
JURAJ PACLT

Inst, of Exptl Phytopathol. and Entomology,

Slovak Academy of Sciences,

Ivanka pri Dunafi, Czechoslovakia

A COMPARISON OF THE SECOND EDITION with the Original Check
List by Kloet and Hincks (1945) reveals among other things
great differences in names used for subfamilies of Geometridae:
First Edition Second Edition

(1945) (1972)

No. 53 Geometridae

No. 13 Brephidae : Archiearinae

(= Monocteniidae) : Oenochrominae

No. 11 Geometridae : Geometrinae

No. 10 Sterrhidae : Sterrhinae

No. 12 Hydriomenidae : Larentiinae

No. 14 Selidosemidae : Ennominae

As long ago as 1844, Duponchel divided his tribe Phalenides,
i.e. Geometridae sensu lato, into 18 subtribes (I-XVIII). The
Phalenides of Duponchel, however, did not include Brephos
Ochsenheimer, 1816, a genus dealt with by him, Duponchel, as
pertaining to another tribe, namely Phalenoides.

Duponchel’s subtribes Chlorochromites (II) and Acidalites
(XVI) correspond to the modern subfamilies Geometrinae and
Sterrhinae (= Scopulinae) respectively, while Eubolites (X),
Cidarites (XI), Larentites (XII) and Melanthites (XIII) do all
to the modem Larentiinae.

The modern Selidosematinae = Ennominae = Boarmiinae
were disposed by Duponchel in not less than 12 subtribes:
Ennomites (I), Gnophites (III), Boarmites (IV), Cleorites (V),
Amphidasites (VI), Hihernites (VII), Fidonites (VIII), Aspi-
latites (IX), Zerenites (XIV), Caberites (XV), Sionites (XVII),
and Dasydites (XVIII). Of these Ennomites and Boarmites sur-
vived alone and changed later to Ennomidae j Ennominae and
Boarmiidae I Boarmiinae, these being recognized as two distinct
family-groups up to the time of Packard (1876). Since both
divisions have been united thereafter, Ennomites and Boarmites
became equally old nomenclatorial rivals (Article 23d).

ARCHIEARINAE

Type-genus ( subfamiliotype ) : Brephos Ochsenheimer, 1816
(nec Huebner, 1813, Samml. exot. Schmetterl. 1, pi. 90!). Re-
placed as junior homonym by Archiearis Huebner, 1823. Hence
the change of Brephinae to Archiearinae.

179

180 JURAJ PACLT /• Res. Lepid.

OENOCHROMATINAE

Type-genus ( subfamiliotype ) : Oenochroma Guenee, 1857.
This subfamily is often wrongly named ‘Oenochrominae (oeno-
chroma, -tos), instead of Oenochromatinae. Monocteniidae,
based on Monoctenia Guenee, 1857 would be a junior synonym
of Oenochromatinae.

GEOMETRINAE

Type-genus (subfamiliotype): Geometra Linnaeus, 1758 (vali-
dated as of this date under suspension of the Rules— see Opinion
450).

SCOPULINAE

Type-genus (subfamiliotype): Acidalia Treitschke, 1825 (nee
Huebner, 1819). The oldest valid name of this genus is Scopula
Schrank, 1802. Accordingly, the family-group name Acidaliinae
has been replaced by Scopulinae, as proposed by Hampson
(Hampson and Durrant, 1918).

LARENTIINAE

Type-genus ( subfamiliotype ) : Larentia Treitschke, 1825. The
family-group name Hydriomeninae must be sunk as a junior
synonym of Larentiinae.

BOARMIINAE

Type-genus (subfamiliotype): Boarmia Treitschke, 1825.
Since Ennomites and Boarmites have been published simultane-
ously (Duponchel, 1844), their relative priority is determined
by the action of the first reviser, viz. Hampson (1898?) to whom
the modern concept of Boarmiinae has been attributed (Hering,
1932), Ennominae as conceived by Dyar (1903), as well as
Selidosematinae, based on Selidosema Huebner, 1823 and pro-
posed first as a family-group name by Meyrick {‘Selidosemidae )
seem not to be more than junior synonyms of Boarmiinae.

LITERATURE CITED

DUPONCHEL, P. A. J. 1844, Catalogue methodique des Lepidopteres
d’Europe. Mequignon-Marvis Fils, Paris. XXXII + 524 p. [Published
first in parts: pp. 1-64 in 1844, the rest probably not earlier than in
1845.]

DYAR, H. G. 1903. A list of North American Lepidoptera and key to the
literature of this order of insects. Bull., US. not. Mus. 52. XIX + 723 p.
[For the year of publication see Bull., U.S. not. Mus. 123:2 (1923).]
HAMPSON, G. F. and J. H. DURRANT. 1918. List of the families and sub-
families of Lepidoptera. Novit. zool. 25:366-394.

HERING, M. 1932. Die Schmetterlinge nach ihren Arten dargestellt. In:
P. Brohmer, P. Ehrmann and G. Ulmer (eds.) Die Tierwelt Mittel-
europas, Suppl. 1. Quelle und Meyer, Leipzig. X + 545 p.

KLOET, G. S. and W. D. HINCKS. 1945. A check list of British insects.

Stockport (privately printed). LX + 484 p.

MEYRICK, E. 1892a. On the classification of the Geometrina of the Euro-
pean fauna. Trans, entomol. Soc. London (1892): 53-140. [Not seen;
reference received with best thanks from D. S. Fletcher, London.]

1892b. Revision of the Australian Lepidoptera. Part V (Geometri-

dae). Proc. Linn. Soc. New South Wales 6:581-678. [Not seen; refer-
ence received with best thanks from D. S. Fletcher, London.]
PACKARD, A. S. 1876. A monograph of the Geometrid moths or Phalaeni-
dae of the United States. Rep. U.S. geol. Surv. 10:1-607.

Journal of Research on the Lepidoptera

13(3):181-190, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

ILLUSTRATIONS AND DESCRIPTIONS OF

SPECIES OF SOME PYRRHOPYGINAE
FROM PANAMA (HESPERIIDAE)

S. S. NICOLAY

1500 Wakefield Drive, Virginia Beach, Va.

The past ten years’ collecting in the Republic of Panama
and in the Canal Zone have produced new information on pre-
viously described species of the HESPERIIDAE and a number
of new species (Nicolay & Small, 1969 and Nicolay, 1973), The
purposes of this paper are to describe new sub-species in the
genera Zonia and Myscelus, provide additional notes on species
already named, and to illustrate species not figured previously.

Zonia zonia panamensis Nicolay, new subspecies
Figures If, 2f, 5, 6, 7

Male: Length of forewing, 29 mm. Upperside: forewing black, a
discal row of conjoined white hyaline spots from vein 1 through cell to
coastal margin in spaces 11 and 12, the latter as narrow streaks, but tri-
angular in interspace lb, rectangular in space 2, almost square across cell;
a narrow, small triangular white hyaline spot in interspace 3, the apex not
reaching vein 4; white hyaline apical spots in spaces 4 and 5, three smaller
spots across the upper edge of spot in space 5 in spaces 6, 7 and 8. A nar-
row bluish-white basal band from costa to inner margin, another pale blue
parallel sub-basal band from costa to inner margin continued as a broken
blue bar in space la to termen, then as a sub-marginal blue line to spot in
space 3 with a few blue scales toward apical spot in space 5. Hindwing
black, upper and lower bluish-white basal streaks, separate and not reach-
ing the central white hyaline band which is 1 mm wide, extending from
vein 2 into interspace 7, but not reaching vein 8; a submarginal bright blue
macular band from interspace lb to vein 7. Fringes dark brown. Underside:
forewing dull black, hyaline markings of the upperside repeated in like
manner; sparse pale blue scaling at wing base; a scattering of blue scales
submarginally from end of spot in space 3 to inner margin. Hindwing black,
inner margin blue-scaled; a sub-basal blue-white band from mid vein 1 to
costa; central hyaline band as on upperside with an added double blue spot
in space Ic; submarginal macular blue band narrow, broken outward at
vein 4, widening toward costal margin. Fringes white, checked brown at each
vein end. Palpi and forecoxae white stripped; head and patagia white
spotted; thorax -with 4 bluish-white stripes; abdomen with alternate
bluish-white and black bands; anal tuft dark brown.

Female: Unknown.

181

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S. S. NICOLAY

13(3);181-190, 1974

PYRRHOPYGINAE

183

Fig. 2. — ■ Upper row, left to right; underside of the same species as illus-
trated in figure 1.

Lower row, left to right; underside of the same species as illustrated in

184

S. S. NICOLAY

J, Res, Lepid.

Holotype male, Madden Forest, Panama Canal Zone, 6 Feb.

1968, collector, S. S. Nicolay. Paratypes, same locality; 1 ^ 15
Aug. 1968, 1 ^ 15 Dec. 1968, 1 ^ 23 July 1969, 1 ^ 24 July

1969, G. B. Small, collector. The holotype will be deposited in
the American Museum of Natural History, New York, New York,
the paratypes, in the collections of the author and Mr. G. B.
Small.

Panamensis has been one of our most interesting catches in
the Canal Zone in recent years. Evans (1951) described the
nominate species zonia for which he erected the genus. Known
only from the single male type taken at Teffe (Ega) Amazonas
in 1879, it has not been figured previously. Photographs of the
type, (Figures 3 & 4) were provided by Mr. R. E. Vane-Wright
of the British Museum (Natural History). He also sent photo-
graphs of an additional male specimen now in the BM which
was taken recently by C. B. Roberts/ BM on the Potaro River,
British Guiana in 1969; it agrees completely with the type.

1 took panamensis for the first time in February, 1968 at a
hilltop in the Madden Forest Preserve. Here, with other “hill-
topping” companion species of the genera Elbella and Phocides,
it swept back and forth across a broad expanse of open grassy
hilltop meadow with an incredibly swift and booming flight.
The method for collecting was relatively simple-stand at an
appropriate place in their flight pattern and, with a long-handled
(about 12 feet) net, attempt to intercept them in full flight! The
percentage of successful intercepts is very low, but none-the-less,
rewarding.

Gordon Small’s experience on many subsequent efforts in this
same local reveals that panamensis was present very infrequently,
if at all, around 11:30 A.M., and only singly. It also occasionally
landed on a small bushy tree where he took all of his specimens.
The Elbella species were also taken after alighting; Phocides
species flew unceasing, never seeming to land.

Panamensis differs from the nominate species zonia in that
there is no hyaline spot in space 9 on the forewing, and the width
of most hyaline spots is about one-third less; the spot in inter-
space 3 on the forewing is, in panamensis, only a fraction of the
width and size of that in zonia. Other less obvious differences
may be noted by comparing the appropriate figures.

The genitalia are identical, with the exception of the valvae;
those in the nominate form are tapered rather gradually and with-
out interruption to the terminal teeth; those of panamensis are

13(3):181-190, 1974

PYRRHOPYGINAE

185

wider thru“0ut more of their length, finally tapering rather
abruptly to the terminal teeth, particularly from the dorsal sur-
face. The terminal teeth are somewhat blunt and there is a slight
asymmetry in the valvae of both species; this latter varies slightly
with each specimen thus far examined.

Zonia and its subspecies panamensis are both apparently very
uncommon insects; I have been unable to find additional speci-
mens in the many collections I have searched; nor have I noted
its presence in the various lists that are published relevant to
specific regions where it may occur. It bears a striking resembl-
ance to Jemadia hewitsonii albescens Rober in size, wing shape
and pattern, but a close inspection will quickly reveal the
notable differences.

Myscelus assaricus michaeli Nicolay, new subspecies
Figures: la, lb, 2a, 2b, 8

Male; Length of forewing, 22 mm. Upperside: forewing with a
yellow-orange basal area, remaining 3/4 of the wing, blue-black with an
indigo sheen; a discal row of hyaline white spots in spaces lb, 2 and across
cell with the widest spot at 2!4 mm in space 2; all spots clearly separated
by black veins; smaller white hyaline spots in spaces 3, 4, 5 and three small
rectangular apical spots in spaces 7-9, all forming a vague, elongate “S'L
shape; a line of vaguely defined, pale blue sub-marginal spots from first
apical spot in space 7 through lb. Hindwing basal 3/4 yellow-orange, outer
quarter shining blue-black with an indigo sheen; black post-discal spots from
space lb through 7, conjoined in spaces lb to 3 and in spaces 4 and 5; an
inner black spot in space 7 and inner cell, almost invisible beneath the
yellow-orange overscaling; termen deeply excavate between veins 1 through
6; fringes white below vein 6. Underside: forewing dull black, white
hyaline spots of the upperside repeated; two pale bluish-white dashes above
cell spot in spaces 11 and 12; a row of bluish- white spots at base of costa
and sub-basal in inner cell, interspace 2 and all of la; a row of wide, lightly
scaled blue submarginal spots from apex through space lb. Hindwing pale
blue with a macular black band near base of wing; a wider sub-median and
post-median band of macular fused black spots, the two joined with a black
line in space 8; a vaguely defined black outer margin; abdominal fold pale
blue; fringes white from vein 7 to tornus, except black at vein ends.

Female; Length of forewing, 26 mm. Upperside: forewing maculation
the same as in male, wings broader, outer margin straight. Hindwing
maculation the same as in the male except the dark outer margin wider, the
postdiscal spots closer to its inner edge, the wing broader and more
rounded.

Above, head and collar black and white striped, tegumen, tegula and
thorax densely covered with long yellow-orange hairs overlapping anterior
segments of abdomen; abdominal segments ringed alternately black and
pale yellow; anal tuft of long, brown pale-tipped hairs: Palpi black above,
pure white below, pectus pale yellow, legs black with pale scales and long
hairs; abdominal segments ringed black and white. Antennae black, pale
yellow inside bend of apiculus and club.

186

S. S. NICOLAY

J. Res. Lepid.

Holotype male, Madden Forest Preserve, Panama Canal Zone,
15 July 1969, G. B. Small, collector. Allotype female, same local-
ity, 16 July 1968, G. B. Small, collector. Paratypes: 1 $ same
locality as holotype, 20 July 1968; 1 $ Gatun, C. Z., 30 Jan. 1970,
1 $ Gatun, C. Z., 25 April 1971, G. B. Small, collector. Two
specimens in the Smithsonian’s National Museum collection, a
male, Carillo, Costa Rica (no date) and a female, Cayuga,
Guatemala, September are also included in the type series. The
holotype will be deposited in the American Museum of Natural
History, New York, New York. The allotype female and one para-
type male remain in the author’s collection, the remainder of the
paratypes, with the exception of the two National Museum speci-
mens, in the collection of Mr. G. B. Small.

The subspecies michaeli extends the known range of the
species complex of Myscelus assaricus Cramer into Central
America and Mexico. Michaeli is most similar to the nominate
form in that the hyaline spots of the forewing are approximately
the same size although smaller than in assaricus and less than
half the width of those in the subspecies mapirica Strand. The
spots of the “S”-shaped apical row are more rectangular, almost
square in michaeli, but very narrow and linear in the nominate
form and in mapirica. The dark spots in the disc of the hindwing
are much smaller in michaeli than in either of the other two sub-
species, being less than half the size of those in mapirica, and
placed closer to the inner edge of the dark margin.

The pale areas of the underside of the hindwing are bright
pale blue or bluish-white in michaeli rather than the greenish-
white of the nominate form assaricus and are wider and more
extensive in michaeli with the black spots and macular bands
wider and heavier in both mapirica and assaricus. The male
genitalia of those few specimens available for examination ap-
pear identical. This, together with the geographical separation,
consistent and obvious differences in wing maculation, estab-
lishes the basis for this subspecies identity with M. assaricus
Cramer.

This beautiful and interesting insect is named for my oldest
son Michael Darrell Nicolay who dearly loved all of nature’s
creatures and particularly the much maligned and misunder-
stood reptiles.

13(3);181~190, 1974

PYRRHOPYGINAE

187

Aspitha leander Boullet

Figures Id, le, 2d, 2e, 9

Yanguna leander Boullet, 1912, Bull Soc. Ent. France, p. 92.
Yanguna parima Plotz, Mabille and Boullet, 1908, Ann. des

Sciences Nat., Paris, 9th Series, pp 187-88, pi 13, fig 3.
Yanguna parima Plots, Draudt in Seitz, 1921, Mac. Lep. of the

World, vol 5, p 842, pi 164d.

Yanguna leander Boullet, Bell, E. L., 1933, Hesperiidae, Jour.

N. Y. Ent. Soc., vol XLl, Sept, pp 288-89.

Aspitha leander Boullet, Evans, W. H., 1951, A Cat. of the Am.

Hesp. in the B. M. (Nat. Hist.) Part 1, p 84.

There has been considerable past confusion regarding this
species. Described by Boullet in 1912, it none-the-less was figured
in color in a paper co-authored by Boullet (1908) four years
earlier as Yanguna parima Plotz. Seitz’ (1921) figure of parima
is in reality that of Aspitha leander Boullet as noted by Evans
(1951). Bell (1933), although he had no specimens before him,
authored an excellent, brief translation of Boullet’s original
description of the male leander that fits very well the species
found in Panama. Aspitha parima Plotz, although obviously very
closely related, is a separate species found in Surinam.

Apparently no females of leander had been taken or asso-
ciated with the male. Evans ( 1951 ) does not mention the female,
nor are characters of the female included as part of his keys.
Early in 1973 Mr. G. B. Small took a pair of A. leander at Gatun
in the Ganal Zone. A description of the female follows:

Female: Length of forewing, 26 mm. Upperside: fore and hindwing
color blue-black; forewing with a broad (3-4 mm) central white hyaline
band from vein 1 in interspace lb (triangular) through upper cell with a
thin, broad spot in interspace 11; two tiny white hyaline spots midway
between the central hyaline band and outer margin in interspace 4, the
lower adjacent to vein 4, the upper, next to vein 5, a third spot above this
in interspace 5; fringes whitish from vein 1 to vein 3. Hindwing with a red
tornal spot 2 mm wide from inner angle through interspace Ic, with a thin,
black lined outer margin, wider at end of vein lb; fringes white from vein
1 through vein 7. Underside: all wings blue-black; markings on both wings
the same as above with an additional thin line of white scales in interspace
12 above spot in interspace 11 on the forewing; wing basal area faintly
paler.

Head black with small white spots at the base of and between the
antennae; tegulae black, patagia at end of tegulae red; thorax and abdomen
black above and below, a few whitish hairs in anal tuft. Mid and hind legs
black, forelegs black with a few white hairs on femur; palpi black, a thin
line of white scales in the center; a narrow line of white scales across fore-
head between tips of the palpi. Antennae black.

188

S. S. NICOLAY

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13(3):181-190, 1974

PYRRHOPYGINAE

189

Fig. 7-9. — Male genitalia, reading from top to bottom (7) Zonia sonia
panamemis Nicolay; (8) Myscelus assaricus michaeli Nicolay; (9) Aspitha
leander Boullet. All drawings with left valva removed, aedeagus in position,
and inner surface of right valva detailed.

190

S. S. NICOLAY

/. Res. Lepid.

It is indeed fortunate that the two specimens figured were
caught en copula. The sexual dimorphism apparent in this very
uncommon species is great enough to make matching the sexes
very difficult. In addition to the obvious differences in wing
fascia, as illustrated in figures 1 and 2, the hindwing in the fe-
male is not as irregular nor as sharply produced at the tornus and
at the ends of veins lb and 2; the forewing is less sharply pro-
duced at the apex and overall, more rounded and full. Another
female collected near Colon in the Republic of Panama in 1969 is
without the sub-apical dots of the specimen figured and des-
cribed herein, but is otherwise identical.

Aspitha leander has not been previously recorded from the
Republic of Panama. The type locality is Muzo, Colombia, 800
meters.

ACKNOWLEDGMENTS

I wish to express my appreciation to Gordon B. Small, Balboa,
Canal Zone for his generous logistic support of my collecting
efforts and for his invaluable assistance in the field work that
made this study possible. I am especially grateful to Mr. R. I.
Vane-Wright on the staff of the British Museum (Natural His-
tory) for his helpful suggestions, and excellent photographs and
sketches of the Evans’ zonia type material. My thanks also to
Wm. D. Field, U. S. National Museum and Dr. F. H. Rindge,
American Museum of Natural History for providing the assistance
and cooperation that allowed me to examine and study the
collections in their care.

Photographs of the panamensis type were made with the
help of WO G. G. Thomas, and SSgt. Richard E. Banzai of the
U. S. Marine Gorps. The color transparencies and line drawings
were made by the author.

LITERATURE GITED

BELL, E. L. 1933. Studies in the Pyrrhopyginae, with Descriptions of New
Species ( Lepidoptera, Rhopalocera, Hesperiidae ) . Jour. N. Y. Ent. Soc.
vol. XLI, Sept-Dec. 1933. 152 p.

1934. Studies in the Pyrrhopyginae (Lepidoptera, Rhopalocera).

Jour. N. Y. Ent. Soc. vol XLIl, Dec. 1934. 48 p.

BOULLET, E. 1912. Bull. Soc. Ent. France. Paris, p 92.

CRAMER, P. 1782. Papillons Exotiques. vol. 3. Amsterdam, pp 120-121,
pi 261, figs F, G.

DRAUDT, M. 1921. American Rhopalocera. In Seitz, A. Macrolepidoptera
of the World, vol 5. Stuttgart.

EVANS, W. H. 1951. A catalogue of the American Hesperiidae. Part I,
Pyrrhopyginae. British Museum of Natural History. London. 92 p.
MABILLE, P. & E. BOULLET. 1908. Ann. des Sciences Nat. Zool. Paris.
9th Series, pp 187, 188; pi 13, fig 3.

NICOLAY, S. S. & G. B. Small. 1969. A New Subspecies of Pyrrhopyge
creon (Hesperiidae) from Panama. J. Lepid. Soc. 23: 127-130.
NICOLAY, S. S. 1973. Descriptions of New Neotropical Hesperiidae. J.
Lepid. Soc. 27: 234-257.

Journal of Research on the Lepidoptera

13(3): 191-206, 1974

1160 W. Orange Grove Ave,, Arcadia, California 91006, U.S.A.
© Copyright 1975

BUTTERFLIES OF THE
SUISUN MARSH, CALIFORNIA
ARTHUR M. SHAPIRO

Department of Zoology,

University of California, Davis 95616

At the time of European colonization, California had an
estimated five million acres of wetlands. By 1973 only a tenth
of that acreage remained, with a current rate of loss of one
percent per year. California, unlike the eastern United States,
is considered to have a depauperate marsh fauna of butterflies,
and its wetlands have received very little attention from col-
lectors.

The Suisun Marsh is a complex of tidally influenced wetland
habitats covering some 44,000 acres in Solano County, west-
central California, and is one of the largest marsh tracts remain-
ing in a reasonably natural state on the entire Pacific coast.
The Marsh occupies a structural basin hemmed in by hills on
three sides. These hills, part of the Coast Ranges, are arid in
summer and are covered with annual grassland or open oak
woodland. They apparently confer sufficient isolation on the
Marsh to produce several endemic taxa of plants and animals;
among these the most prominent are the Suisun Shrew {Sorex
sinuosus) and the showy Composites Grindelia paludosa, Cirsium
hydrophilum, and Aster chilensis ssp. lentus. As may be seen
from Figure 1, the Marsh communicates with San Francisco-
San Pablo Bays to the west and the Sacramento-San Joaquin
Delta to the east, and through the latter with the Central Valley
of California. A biotic connection with the San Francisco Bay
salt marshes is shown by the ranges of two endemic species
shared by them, the Salt-Marsh Harvest Mouse (Reithrodon-
tomys raviventris) and the Composite Grindelia humilis.

191

VACAVlLLiT

192

ARTHUR M. SHAPIRO

/. Res. Lepid.

-Location map of the Suisun Marsh.

13(3):191-206, 1974

BUTTERFLIES OF SUISUN MARSH

193

CLIMATE OF THE MARSH

The Suisun Marsh differs from the Central Valley in being
exposed to a stronger maritime climatic influence, which enters
the basin by the Carquinez Straits. At the same time the climate
of the Marsh is more continental than those of the valleys which
open directly to the coast. Aside from moderating both summer
and winter temperature extremes, the maritime effect increases
the frequency of morning fog and low cloudiness as compared
to the Central Valley, e.g. Vacaville ten miles to the east; but
unlike immediate coastal sections, the Marsh very rarely has
persistent fog or cloud in summer. More dramatically, the
maritime air produces frequent sustained high winds as it rushes
through the Straits to fill up the thermal low-pressure trough
in the interior. Winds in excess of 20 knots probably occur on
100 days or more each year in the Marsh, blowing usually from
the west (“Suisun” means “west wind” in the local Indian dia-
lect, and is still used locally as a name for this wind).

The U.S. Weather Bureau maintains a reporting station at
Fairfield. This location receives from 30-50 days with maximum
temperatures greater than or equal to 90°F (32.2°C) and 15-40
days with minimum less than or equal to 32°F (0°C) per season.
Temperatures in excess of 100°F (37.7°C) occur 3-5 times/sea-
son and below 30 °F (—l.l^C) a similar number of times. Data
for 1971 and 1972 monthly temperatures are given in Table 1.
Precipitation of about 24 inches falls from October to May;
there is little or no precipitation in summer. Relative humidity
in summer is, however, consistently higher than in the Sacra-
mento Valley and the Marsh is sheltered from the full force
of the hot, dry “northers” which produce the most extreme
summer weather in the Valley.

HABITATS OF THE MARSH

Most of the Suisun Marsh is under the control of hunt clubs
whose management practices vary widely. Portions of the Marsh
are cut over or burned from time to time; through an extensive
system of levees, ditches, and gates the water level may be
manipulated on individual parcels of land. Soil salinity is thus
extremely variable from parcel to parcel. The overall result is
a complex vegetational mosaic which for convenience may be
divided into “high marsh,” “low marsh,” and “landfill.”

Table 1. Temperature data In °F for Fairfield Fire Station, California, 1971-72.

194

ARTHUR M. SHAPIRO

/. Res. Lepid.

'T3| v|

ds

a I

enoooooooo

oooO'd-cocyiCN

-al v|

I ^

cop

'dl A I

G
•H

e

H

d

r>*csloOOOOOOOOO

oooooLnr-jr^iocNioo

*Although 1972 was a *'cool'* year, a brief period of record-breaking heat occurred in July.

13(3): 19 1-206, 1974

BUTTERFLIES OF SUISUN MARSH

195

“High Marsh” — This group of associations occurs in areas
of low salinity: high points (islands) within the Marsh, levees
and the edges of sloughs, ditched marshland and fresh-water
pools. It has the highest plant species diversity in the Marsh,
and is characterized by the aspect dominance of showy Com-
posites in late summer. Some characteristic plant species are:
Grindelia humilis, G. paludosa. Aster chilensis ssp. lentus, Bac-
charis douglasii, Cirsium hydrophilum, Solidago occidentalism
Achillea borealis ssp. arenicola, Pluchea purpurascens, Cicuta
bolanderim Apium graveolenSm Potentilla egedeb Atriplex patula
var. hastatam Phragmites communis var. berlandierim Scirpus
olneyb S. koilolepiSm S. acutus, J uncus effusus var. brunneuSm and
Typha latifolia.

“Low Marsh” — This association, or group of associations,
covers the most extensive part of the Marsh. It occupies areas of
high salinity which are saturated or shallowly inundated at high
tide. The dominant plants throughout are Distichlis spicata and
Salicornia virginica. Characteristic associates are: Atriplex
patula var. hastata, Lilaeopsis occidentalism Polygonum aviculare
var. littoralem P. fowler^ Cotula coronopifoliUm Frankenia grandi-
folia, Eryngium articulatumm Rumex crispus, Juncus acutus
var. sphaerocarpuSm J. bufoniuSm and Scirpus olneyi.

“Landfill” — In many places marshland has been filled in
with construction debris or other refuse, or with clay soils from
the uplands. Plants found in such sites are typical weeds of
highly disturbed lowland sites in California: Centaur ea sohtiti-
alis, Cichorium intybus^ Lactuca serriolUm Cirsium lanceolatum,
Silybum marianum, Malva rotundifoliUm M. nicaeensiSm Sida
hederaceUm Convolvulus arvensis, Epilobium paniculatumm Atri-
plex rosea, A. semibaccata. Polygonum aviculare, Foeniculum
vulgare, Bromus rubens, B. rigidus, Avena barbata, etc. The
transition from marsh to annual weeds occurs at about four feet
above sea level, the mean highwater mark at Suisun. Baccharis
piluhris ssp. consanguinea and Tamarix pentandra form thickets
just above this level.

THE BUTTERFLY FAUNA

So far as I have been able to determine, the Suisun Marsh
has never been collected systematically for butterflies before
1972. In 1972 and 1973 I visited it at frequent (usually 2-4
weeks) intervals and recorded all butterfly species present. The
seasonal distributions of the 40 species recorded are given in
Figures 2 and 3. It is likely that one more resident and five to

196

ARTHUR M. SHAPIRO

/. Res. Lepid.

ten immigrant or stray species (from the hills) may be found
in the Marsh. All of the resident species recorded in the Marsh
occur in both the San Francisco Bay area and the Central Valley
(except Ochlodes ijuma, unrecorded from the Bay but to be
expected). Two endemic populations of widespread species
occur in the Suisun Marsh. They are the P otentilla-ieedimg strain
of Lycaena helloides and the large, richly colored Phyciodes
campestris associated with Aster chilensis ssp. lentiis. Both have
been sought, but not yet found, in salt marshes around the Bay
area and in Marin County. They are discussed individually,
below.

SYSTEMATIC LIST

Dancim plexippiis ( L. ) — Occasional to frequent, especially in
autumn; throughout. Larvae on Asclepias fascicularis on land-
fill. Does not overwinter at Suisun.

Coenonympha tidlia California West. — Rare stray from the hills;
not known to breed in the Marsh. Is is striking that the C. tullia
complex has not evolved salt-marsh populations on the Pacific
Coast as it has elsewhere in its vast range.

Speyeria coronis (Behr). — One female, certainly a stray from
the hills. No violets are recorded in the Marsh, but Viola pedun-
culata occurs in the Potrero Hills and on Mt. Diablo.
Euphydryas chalcedona (Dbldy). — One male, a stray from the
hills.

Phyciodes mylitta (Edw. ). — Frequent on levees and high marsh;
multiple-brooded. Larvae common on Silybum on landfill, but
not yet found on Cirsium hydrophilum.

Phyciodes campestris (Behr). — Unaccountably rare. The 7
Marsh specimens are phenotypically very distinct from the usual
lowland P. campestris (Fig. 4), particularly in the lack of con-
trast between the median spot-band and the ground color; the
very well-developed submarginal spot-band; the reduced amount
of black above; and the larger size. They were taken on flowers
of Aster chilensis ssp. lentils and may represent an endemic
population developed in association with this plant, but until
more information and specimens become available it seems best
to defer naming them.

13(3):191-206, 1974

BUTTERFLIES OF SUISUN MARSH

197

D. plexippus

C. t. California
S. coronis

E. chalcedona
P. mylitta

P. campestris
N. antiopa

N. californica
V. atalanta

C. virginiensis
C . carye
C. cardui
P. coenia
L. loiquini
L. bredowii
S. melinus
S. sylvinus
S. californica
L. helloides
L. xanthoides

B. exilis
E. comyntas
P. acmon

E. ausonides

C. eury theme
P. rapae

P. protodice
B. philenor
P. zelicaon
P, rutulus
E. tristis
P. communis
P. scriptura
P. Catullus
H. phylaeus
A. campestris
P. sabuleti

O. sylvanoides
0. yuma

L, eufala
Total species ;

vi.25 vii.l6 vii.23 viii.25 ix.23 x.21

X X XX

XXX

X

X X

XXX
X X

XXX
X X

XXX

X

X

X

XXX

XXX

X

X XXX

X XX

X

X

XXX

XXX
X X

XXX
XXX
X X

XXX
X

X X

X

16 18 19

X

X XXX

X X

X XXX

X X

X XXX

X X

X XXX

18 19 lA 11

Fig. 2. — Seasonal distribution of butterflies at Suisun, 1972.

■Seasonal distribution of butterflies at Suisun, 1973.

J. Res. Lepid.

198

ARTHUR M. SHAPIRO

XX X X X X X

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X

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californica

comyntas

13(3):191-206, 1974

BUTTERFLIES OF SUISUN MARSH

199

XXX XXX X XXXX X x^

CM

X X XXX X XXXXXXX Xvo

CM

X X XXX X XXXXXXXXXXr^

X X XXX

X X XX

XX XXX

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sylvanoides

200

ARTHUR M. SHAPIRO

J. Res. Lepid.

Fig, 4, — Phyciodes campestris from lowland central California, a-c, g-i,
Suisun Marsh, X. 12.73; d-f, j-1, Willow Slough, Yolo Co., X. 19.73.

13(3): 191-206, 1974

BUTTERFLIES OF SUISUN MARSH

201

Nymphalis antiopa (L.) — Occasional throughout. Larvae oc-
casionally abundant on Salix lasiandra along ditches.

Nymphalis calif ornica ( Bdv. ) . — Occasional as a migrant through
the Marsh, but not breeding; no Ceanothus occur in the basin.
Vanessa atalanta ( L. ) . — Occasional throughout, mostly in high
marsh; multiple-brooded. Larvae on Urtica holosericea (Joice
Island ) .

Cynthia virginiensis (Drury). — Occasional throughout; the
least common Cynthia, but still commoner than in most of the
Sacramento Valley. Larvae on Gnaphalium bicolor (Joice Island;
levees near Suisun City).

Cynthia carye Hbn. (= annahella Field).— Common everywhere;
multiple-brooded. Larvae common on Malva rotundifolia, M.
nicaeensis. Althaea rosea, and Sida hederacea (all Malvaceae)
on landfill. A striking aberration was taken in the Marsh, v. 19.73,
figured in Shapiro, 1973.

Cynthia cardui (L.). — Common throughout. Does not appear
to overwinter. Larvae on Silyhum marianum, Cirsium lanceo-
latum, Centaurea solstitialis, Malva spp., and Amsinckia doug-
lasiana on landfill; on Cirsium hydrophilum on high marsh.
Precis coenia (Hbn.). — Common to abundant everywhere,
especially in late summer and autumn; September-October speci-
mens are often very red beneath. Larvae on Plantago coronopus
in low marsh, P. lanceolata and Lippia nodiflora on landfill.
Limenitis lorquini (Bdv.). — Occasional along sloughs near
willows, probably breeding.

Limenitis bredowii calif ornica (But!.). — Recorded once; stray
from foothills.

Strymon melinus Hbn. — Frequent to common throughout, all
season.

Satyrium sylvinus ( Bdv. ) . — Locally frequent in clumps of Salix
hindsiana along ditches, otherwise not seen.

Satyrium calif ornica (Edw. ). — One male, in high marsh on
celery blossoms, certainly a stray from the hills.

Lycaena helloides (Bdv.). — Abundant. On landfill associated
with Polygonum aviculare and Rumex crispus; in high marsh,
especially deep in the Marsh, with Potentilla egedei. The two
strains are slightly differentiated phenotypically in a statistical
sense, and are not altogether synchronized in brood sequence.
See Shapiro, 1974 for a fuller discussion. A bilateral gynandro-

202

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. 5.^ — Bilateral gynandromorph of Lycaena helloides, Suisun, X. 12.73.

13(3):191-206, 1974

BUTTERFLIES OF SUISUN MARSH

203

morph of the Potentilla strain taken in high marsh, X. 12.73 is
shown in Figure 5.

Lycaena xanthoides (Bdv. ). — Abundant on landfill and low
marsh, associated with Riimex crispus; one brood. Suisun speci-
mens average smaller and duller in color than Central Valley
ones, and show some local differentiation.

Brephiditim exilis (Bdv.). — Scarce early in the season, but be-
coming very abundant in September-October. Throughout,
commonest on low marsh and landfill. Larvae abundant on
Atriplex patula var. hastata, A. rosea, and A. semibaccata.
Everes comyntas (God.). — Locally frequent along ditches;
occasional in high marsh; in spring common on landfill. Larvae
on Lotus purshianus, L. strigosiis, and Vida sativa.

Plebeius acmon (West. & Hew.). — Common throughout. Spring
and fall males, and spring females of f. vern. ''cottlef. Larvae
on Polygonum aviculare on landfill.

Euchloe ausonides Lucas. — Frequent to locally common on
landfill; occasional on high marsh. Two broods. Larvae on
Raphanus sativus and Brassica nigra.

Colias eury theme Bdv. — Common everywhere, especially in
September and October. Larva on Vida sativa on landfill-
doubtless on a wide variety of legumes as elsewhere.

Pieris rapae (L. ) — Common throughout. Larvae on Brassica
nigra and B. arvemis and Raphanus sativus on landfill.

Pieris protodice Bdv. & LeC. — Occasional on landfill near Suisun
City, in annual grassland; strays into marsh.

Battus philenor hirsute ( Skinner ) . — Occasional stray from near-
by canyons; once fresh in Suisun City, xii.3.73.

Larvae abundant on Foeniculum vulgar e on landfill and levees.
Papilio zelicaon Lucas. — Abundant on high marsh and landfill.
Not recorded on Cicuta or Apium in high marsh, although both
are acceptable in the laboratory.

Papilio rutulus Lucas. — Occasional in high marsh; not known
to breed, but suspected of doing so on cottonwoods or willows,
as on Joice Island.

Erynnis tristis (Bdv.). — Infrequent on landfill and on high
marsh; presumably a stray from the hills.

Pyrgus communis (Grote). — Abundant on landfill; occasional
on high marsh; all season. Larvae common on Malva rotundi-
folia, M. nicaeensis, and (occasionally) Sida hederacea.

Pyrgus scripture (Bdv.). — Abundant on landfill near the host
plant, Sida hederacea.

204

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. 6.”Variation in male Polites sabuleti. a-c, e-g, Suisun Marsh, X. 12.73.
d, h, P. s. tecumseh, Dormer Pass, Placer Co., el. 6975’, viii. 17.73.

13(3):191-206, 1974

BUTTERFLIES OF SUISUN MARSH

205

Pholisora catullus (Fabr. ). — Common on landfill; occasional on
low marsh. Larvae on Amaranthus hybridus on landfill.
Hylephilu phylaeus (Drury). — Common throughout, especially
in September and October.

Atalopedes campestris (Bdv. ). — Frequent on landfill; occasional
elsewhere.

Polites sabuleti (Bdv.). — Very abundant on low marsh and
landfill. Extremely variable; a few specimens (cf. Fig. 6) indis-
tinguishable from F. 5. tecumseh of the high Sierras. Larvae
common on Distichlis spicata.

Ochlodes sylvanoides (Bdv.). — Common throughout; at least
two broods.

Ochlodes yuma (Edw. ). — Locally frequent around stands of
Phragmites, as at the Suisun City marina and along Highway 21
north of the Fairfield City Limit. Two broods.

Lerodea eufala (Edw.). Frequent in all habitats in late sum-
mer and autumn. Larvae on Sorghum halepense, Echinochloa
crus-galli, and Cynodon dactylon on landfill and along ditches.

DISCUSSION

There is surprisingly little phenological difference in the
butterflies of the Suisun Marsh and the Central Valley. The only
conspicuous example is Papilio zelicaon, which flies into mid-
November at Suisun 6-8 weeks after it has disappeared in the
Valley. It is also more abundant at Suisun than in the Valley.
Brephidium exilis, which is largely a species of saline and alka-
line situations, is near its northern limit in both areas and may
not always overwinter. In 1972 it flew earlier at Suisun than in
the Valley, but in 1973 the situation was reversed. Flight ac-
tivity at Suisun is limited by wind probably to about the same
degree as it is prevented by heat and dryness in summer in the
Valley. In most of California the maximum numbers of indi-
viduals and species fly in spring or early summer; at Suisun this
peak occurs in September and October.

The Suisun Marsh receives no summer rain, but its year-
round water supply and the halophytic habit of many of its
plants provide a constant supply of butterfly host plants and
nectar sources. In most of lowland California there is a summer
die-off of annuals; most such places have a fauna including many
vernal univoltine species. In the nearby Vaca hills in Yolo and
Solano Counties 25 of 65 recorded taxa are univoltine (39%), for
example. In the Sacramento Valley the moist riparian habitats

206

ARTHUR M. SHAPIRO

/. Res. Lepid.

and agricultural irrigation allow for a much more multivoltine
fauna (8 of 47 breeding taxa are univoltine, 17%). The Suisun
Marsh has the most strongly multivoltine fauna recorded in
California — only 2 of 33 breeding taxa are univoltine {Satyrium
sylvintis, Lijcaena xanthoides) , or 6%. (Of the 6 immigrant taxa
from the hills, however, 3 are univoltine — S. coronis, E. chalce-
dona, S. calif ornica.) Except for Euchloe ausonides, Lerodea
eufala and the two Ochlodes, the remaining species are appar-
ently continuously brooded all season. The brood sequence of
Phyciodes campestris is unknown.

Only one additional resident species, Polygonia satyrus
(Edw. ) is likely to be found. Its host, UHica holosericea, is
locally common in the Marsh, Almost any common foothill
species, and some of the less common ones, may turn up even-
tually in the Marsh. There is a remote possibility that Cercyonis
pegala ariane (Bdv. ) may turn up somewhere in the Marsh; it
is unrecorded in Solano County but is found only 15 miles away
in Contra Costa County in similar habitats.

The Suisun butterfly fauna eontains a large proportion of
weedy or adventive species. This is the case with most of low-
land California. Although locally differentiated populations of
two species occur at Suisun, the lack of butterflies and skippers
associated with the many distinctive plants — especially grasses
and sedges — is as striking there as elsewhere. The reclamation
of marshland in the Central Valley may have destroyed some
endemic Lepidoptera, but the absence of relict populations
from places as little disturbed as parts of the Suisun Marsh
argues for a genuine lack of wetland species in California.

ACKNOWLEDGMENTS

Several of the records reported here were obtained by Mrs.
Adrienne R, Shapiro. Mr. Paul Flores showed us collecting
areas on Joice Island. Plants were determined in the U.C. Davis
Herbarium.

This research was aided by Grant D-804 from the Committee
on Research of the Davis Division, Academic Senate.

LITERATURE CITED

MUNZ, P. A. and D. D. KECK. 1970. A California Flora. Univ. of Cali^
fornia Press, Berkeley. 1681 pp.

SHAPIRO, A. M. 1973. Recurrent aberration in Cynthia annabella: a review
with four new records. Pan-Pacific Entomol.

SHAPIRO, A. M. 1974. A salt-marsh population of Lycaena helloides (Lepi-
doptera: Lycaenidae) feeding on a Potentilla (Rosaceae). Entomol.
News.

13(3): 169-178, 207-216, 1974

BIBLIOGRAPHY

207

(Continued from Page 178)

KIRBY, W. F., 1872. On the geographical distribution of the diurnal Lepi-
doptera as compared with that of the birds. Journal Linnean Society
London, Zoology, 11: 431-439.

1896. A Hand-Book to the order Lepidoptera. Part I. Butterflies. —

Vol. 11. 332 pages. London: Edward Lloyd Ltd.

1903. The Buttei-flies and Moths of Europe. 432 pages, 54 plates.

London; Cassel and Companv.

KIRIAKOFF, S., and H. STEMPFFER, 1952. Un difficile probleme de
repartition geographique. Revue Francaise de Lepidopterologie Paris,
13; 229-235.

KISHIDA, K., 1933. A summary of the butterflies in the inner South Seas.
Rigakkai [Scientific World], 31 (9): 789-794. [In Japanese]

KLOTS, A. B., 1931. A generic revision of the Pieridae (Lepidoptera), to-
gether witli a study of the male genitalia. Ent. Amer., 12: 139-242,
plates V-XIII.

— 1936. The interrelationships of the species of the genus Lycaena

Fabricius (Lepidoptera, Lycaenidae). Bulletin Brooklyn Entomological
Society, 31: 154-171.

1951. A Field Guide to the Butterflies. 349 pages, 40 plates. Boston:

Houghton Miffiin Company.

KOCH, G., 1870. Die geographische Verbreitung der Schmetterlinge liber
die Erde. Petermann, Mittheil., 16: 20-25, 52-57.

KORSHUNOV, YU. P., 1964. Rhopalocera (Lepidoptera) of the mountain
region and the southern coast of Crimea. Entomological Review, 43;
303-309.

1972. A catalog of the Rhopalocera (Lepidoptera) in the fauna of

the USSR [I and II]. Entomological Review, 51: 83-98, 212-223.

KOSTROWICKI, A. S., 1969. Geography of the Palaearctic Papilionoidea
(Lepidoptera) Zaklad Zoologii Systematyeznej Polskiej Akademii Nauk,
380 p.

KRUG, H. C., 1877. Tagschmetterlinge von Portorico. Stett. Ent. Zeit., 38:
_233-245.

KRuGER, E., 1932. Verbreitung und Ableitung einiger Tagfalterfamilien
des tropischen Amerikas (Rhop., Lep.) Deutsche Entomologische Zeit-
<trhriif 1939 ( 9 • 149- 194

KURENTZbv, A. L, i970. The Butterflies of the Far East USSR. 163
pages, 14 colored plates. Academy of Sciences of USSR, Siberian
Branch.

KUSNEZOV, N. J., 1925. Some new Eastern and American elements in the
fauna of Lepidoptera of Polar Europe. Comp. Rend. Acad. Sci. URSS
(A): 119-122.

1930. On the absence of certain elements from the lepidopterous

fauna of the Crimea. The Entomologist, 63: 103-106, 128-132.

1935. The origin of the lepidopterous fauna of the arctic Eurasia.

Arctica, 3: 115-136.

LAIDLAW, W. B. R, 1970. Butterflies of New Zealand. 48 pages. Auck-
land: Collins.

LATHY, P. L, 1904. A contribution towards the knowledge of the Lepi-
doptera— Rliopalocera of Dominica, B. W. I. Proceedings Zoological
Society I^ondon, 1904 (I): 450-454.

de LATTIN, G., 1950. Turkiye Kelebekleri Hakkinda I. Tiirkische Lepi-
dopteren I. Istanbul, Universite Fen fakultesi Mecmuasi. Series B.
Tabu ilimler, 15; 301-331, plates I-II.

LEECH, J. H., 1887. On the Lepidoptera of Japan and Corea. Proceedings
Zoological Society London, 1887: 398-571; 1893-4: 297-681.

LEGRAND, H., 1965. Lepidopteres des lies Seychelles et D’Aldabra.
Memoirs du Museum National D'Histoire Naturelle, Ser. A, Zoologie,
37: 1-210, 16 plates.

208

OAKLEY SHIELDS

/. Res. Lepid.

LEWIS, H. L., 1973. Butterilies of the World. 312 pages, 208 plates. Chi-
cago: Follett Publishing Company.

LINDROTH, C. H., 1957. The Faunal Connections between Europe and
North America. 344 pages. New York: John Wiley and Sons.

LINDSEY, A. W., 1926. Lepidoptera collected by the Barbados-Antigua
Expedition from tlie University of Iowa. Iowa (State) University
Studies in Natural History, 11 (7) : 33-35.

1942. A preliminary revision of Hesperia. J. Sci. Labs. Denison

University, 37: 1-50, 6 plates.

LONGSTAFF, G. B., 1912. Butterfly-Hunting in Many Lands. 729 pages.
London: Longmans, Green and Company.

LUH, C., 1945. A list of butterflies from southern Yunnan. Journal West
China Border Research Society, 15 (B): 20-27.

McCUBBIN, C., 1971. Australian Butterflies. 206 pages. Melbourne:
Thomas Nelson Ltd. [Extensive bibliography]

MACKWOOD, F. M., 1913. Some notes on butterflies and their distribu-
tion. Spolica Zeylanica, 9 (pt. 33): 36-40.

MacNElLL, C. D., 1962. A preliminary report on the Hesperiidae of Baja
California (Lepidoptera). Proceedings California Academy Sciences,
4tli Series, 30: 91-116.

M ANDERS, N., 1907. The butterflies of Mauritius and Bourbon. Trans-
actions Entomological Society London, 1907: 429-454, plate XXIX.

MANLEY, W. B. L., and H. G. ALLCARD, 1970. A Field Guide to the
Butterflies and Burnets of Spain. 192 pages, 40 colored plates. Hamp-
ton: E. W. Classey Ltd.

MARSH, J. C. S., 1968. Hong Kong Butterflies. 113 pages, 34 plates. Hong
Kong: Published by the Shell Company of Hong Kong Ltd.

MATHEW, G. F., 1885. An afternoon among the butterflies of Thursday
Island. Proceedings Linnean Society New South Wales, 10: 259-266.

1886. An afternoon among the butterflies of Thursday Island. The

Entomologist, 19: 33-36, 84-97.

1886. Descriptions of some new species of Rhopalocera from the

Solomon Islands. Proceedings Zoological Society London, 1886: 343-
350.

1889. Descriptions and life-histories of new species of Rhopalocera

from the western Pacific. Transactions Royal Entomological Society
London, 1889: 311-315.

MATSUMURA, S., 1915. [Micronesian butterflies.] Entomol. Mag. Kyoto
(Ent. Soc. Japan), 1 (2); 63-68, 93-96. [In Japanese]

1934. Geographical distribution of Lepidoptera in Pacific countries.

Procedings Fifth Pacific Science Congress, 5: 3503-3514.

MEADOWS, D., 1937. An annotated list of the Lepidoptera of Santa Cata-
lina Island, California. Part I. — Rhopalocera. Bulletin Southern Cali-
fornia Academy Sciences, 35: 175-180.

MICHENER, C. D., 1942. A generic revision of the Heliconiinae (Lepi-
doptera, Nymphalidae). American Museum Novitates, no. 1197, 8
pages.

1943. Some systematic notes on the Libytheidae (Lepidoptera).

American Museum Novitates, no. 1232, 2 pages.

MILLER, L. D., 1965. A review of he West Indian “Choranthus”. Journal
Research Lepidoptera, 4: 259-274.

1965. Systematics and zoogeography of the genus Phanus (Hesperi-
idae). Journal Research Lepidoptera, 4: 115-130.

1968. The higher classification, phylogeny and zoogeography of the

Satyridae (Lepidoptera). Memoirs American Entomological Society,
24: 174 pages.

1970. Reports on the Margaret M. Carey-Carnegie Musemn Expedi-
tion to Baja California, Mexico, 1961. 7. The family Hesperiidae (Lepi-
doptera). Annals Carnegie Museum, 41: 169-200.

13(3):169-178, 207-216, 1974

BIBLIOGRAPHY

209

MONROE, R. S., G. N. ROSS, and R. N. WILLIAMS, 1967. A report on
two recent collections of butterflies from Honduras. Journal Lepidop-
terists’ Society, 21: 185--197.

MOORE, F., 1874. List of diurnal Lepidoptera collected in Cashmere
Territory by Capt. R. B. Reed, 12th Regt., with descriptions of new
species. Froceedings Zoological Society London, 1874: 264-274.

-- — 1877. The Lepidopterous fauna of the Andaman, and Nicobar
Islands. Proceedings Zoological Society London, 1877 : 580-632, plates
58-60.

— 1878. A list of the lepidopterous insects collected by Mr. Ossian

Limborg in Upper Tenasserim, with descriptions of new species. Pro-
ceedings Zoological Society London, 1878: 821-858.

— 1886. List of Lepidoptera of Mergui and its archipelago. Proceed-

ings Zoological Society London, 1882: 234-263.

— 1889. List of the Lepidoptera of Mergui and its archipelago collected

for the trustees of the Indian Museum, Calcutta, by Mr. John Ander-
son, F. R. S., Superintendent of the museum. Journal Linnean Society,
21: 29-60, plates 3-4.

MOSELEY, H. N., 1892. Notes by a Naturalist: an Account of Observations
made during the Voyage of H. M. S. ^‘Challenger ’ . 540 pages. London:
John Murray, [p. 116 — on Tristan da Cunha — a “Vanessa!’^ sp. and an
“Argynnis” sp.]

MOULTON, J. C., 1915. Butterflies of Borneo. Sarawak Museum Journal,
2: 197-266.

MUNROE, E. G., 1949. Some remarks on the orithya group ofthe genus
Junonia (Lep., Nymphalidae), The Entomologist, 82: 157-158.

1950. The systematics of Calisto (Lepidoptera, Satyrinae), with

remarks on the evolutionary and zoogeographic significance of the
genus. Journal New York Entomological Society, 58: 211-240.

1950. The occurrence of a butterfly in the Pribilof Islands. Lepidop-

terists News, 4: 44.

— 1951. The genus Junonia in the West Indies (Lepidoptera, Nym-
phalidae). American Museum Novitates, no. 1498, 16 pages.

— 1957. Comparison of closely related faunas. Science, 126; 437-439.

1963. Characteristics and history of the North American fauna:

Lepidoptera. Proceedings 16th International Congress Zoology, 4:
21-27.

1971. Distribution and geographical differentiation of Marpesia

eleuchea Hiibner (Nymphalidae), with descriptions of two new sub-
species. Journal Lepidopterists Society, 25 (3): 185-193.

date? The classification of the Papilionidae (Lepidoptera). Canadian

Entomologist, Supplement, 17: 51 pages.

MUSGRAVE, A., 1947. Some butterflies of Australia and the Pacific. The
Swallowtails VI. Australian Museum Magazine, 9: 133-135.
NAKAMURA, Y., 1929. On the butterflies from Pelew Islands. Lansania,
1 (2): 26-29. [In Japanese]

— 1929. On the butterflies from Saipan Island. Lansania, 1 (6): 87-90.

[In Japanese]

1933, Papilionidae from the inner south seas. Lansania, 4 (35):

67-71. [In Japanese]

— 1934. A supplement to the knowledge of the swallow-tailed butter-

flies, papilionids of Pacific islands under Japanese Mandate. Lansania,
6 (58): 113-115. [In Japanese]

NEAVE, S. A., 1904. On a large collection of Rhopalocera from the shores
of the Victoria Nyanza. Novitates Zoologicae, 11: 323-363.
de NICeVILLE, L., 1890. On new and little-known butterflies from the
Indian region, with descriptions of three new genera of Hesperiidae,
Journal Bombay Natural History Society, 5: 199-225, plates D and E.

210

OAKLEY SHIELDS

J. Res. Lepid.

1900. The food-plants of the butterflies of tlie Kanara District of

tlie Bombay Presidency, with a revision of the species of butterflies
there occurring. Journal Asiatic Society Bengal, 69: 187-278, plate 1.

de NICeVILLE, L., and H. J. ELWES, 1898. A list of the butterflies of
Bali, Lombok, Sambawa and Sumba. Journal Asiatic Society Bengal,
66 (pt. 2): 668-724.

de NICeVILLE, L., and H. KiiHN, 1898. An annotated list of tlie butter-
flies of the Ke Isles. Journal Asiatic Society Bengal, 67 (pt. 2): 251-283.

de NICEVILLE, L., and L. MARTIN, 1896. A list of the butterflies of
Sumatra, with special reference to the species occurring in the north-
east of die island. Journal Asiatic Society Bengal, 64: 357-555.

NIKITIN, M., 1935. The geographical distribution of Lepidoptera in Man-
churia. Comptes Rendus 12th International Congress Zoology, 2: 1109-
1126.

NYSTROM, V., 1958. Macrolepidoptera from die Cape Verde Islands.
Societas Sci. Fennica, Commentationes Biologicae, 17 (7): 1-36.

OGILVIE-GRANT, W. R., 1903. Insecta: Rhopalocera. In: H. O. Forbes,
The Natural History of Sokotra and Ahd-el-Kuri. Liverpool.

OLLIFF, A. S., 1887. Insecta. In: Report on a small zoological collection
from Norfolk Island. Proceedings Linnean Society New South Wales,
12: 1001-1014.

1891. Stray notes on Lepidoptera. Proceedings Linnean Society New

South Wales, 16: 27-30.

OWEN, D. F., 1971. Tropical Butterflies. 214 pages. Oxford: Clarendon
Press.

PAGENSTECHER, A., 1899. Die Lepidopterenfauna des Bismarck-Archi-
pels. 268 pages. Stuttgart: Erwin Nagele.

1900. tiber die geographische Verbeitung der Tagf alter im malayi-

schen Archipel. Jahrb. Nassau Ver., 53: 85-200.

1907. Die Lepidopterenfauna der Antillen. Jahrb. ver. Naturk.

Wiesbaden, 60: 91-102.

1909. Die geographische Verbreitung der Schmetterlinge. 451

pages. Jena: Gustav Fischer.

PATTERSON, D., and J. A. POWELL, 1959. Lepidoptera collecting in die
Sierra San Pedro Martir, Baja California. Journal Lepidopterists' So-
ciety, 13: 229-235.

PA LILIAN, R., 1951, Papillons conimuns de Madagascar. 90 pages. Institut
de Recherche Scientifique Tananarive-Tsimbazaza.

1956. Faune de Madagascar. II. Insectes Lepidopteres. Danaidae,

Nymphalidae, Acraeidae. 102 pages.

PEILE, H. D., 1923. The butterflies of Mesopotamia. Journal Bombay
Natural History Society, 28: 50-70, 345-369.

PENDLEBURY, H. M,, 1933. On a small collection of butterflies from
Christmas Island, Indian Ocean. Bulletin Raffles Museum, 8: 94-97.

1947. Lepidoptera (Heterocera) and Lepidoptera (Rhopalocera).

In: Contributions of die natural history of Christmas Island, in the
Indian Ocean. Bulletin Raffles Museum, no. 18: 58-80.

PETERS, J. V., 1969. The butterflies of Lord Howe Island. Proceedings
Royal Zoological Society New South Wales, 1967-68 (1969): 63-64.

1969. Notes on the distribution of Australian Hesperioidea and

Papilionoidea (Lepidoptera). Australian Zoologist, 15: 178-184.

PETERS, W., 1952. A Provisional Check-List of the Butterflies of the
Ethiopian Region. 201 pages. Middlesex: E. W. Classey Publishers.

PETERSEN, B., 1968. New records of Rhopalocera (Lep.) from Rennell.
Natural History Rennell Island, British Solomon Islands, 5: 105-110,
plates 3-4.

PHILLIPS, W. W. A., 1964. Some observations on the fauna of the Mal-
dive Islands. Part VH — butterflies. Journal Bombay Natural History
Society, 61: 396-401.

13(3):169~178, 207-216, 1974

BIBLIOGRAPHY

211

PINHEY, C. G., 1949. Butterflies of Rhodesia. 208 pages, 21 plates. Salis-
bury: Published by the Rhodesia Scientific Association.

1965. Butterflies of South Africa. 240 pages. Johannesburg: Nelson

and Sons Ltd.

POLAGEK, E. R., 1935. Some closely related butterflies of Europe and
America. Bulletin Natural History Society Maryland, 6 (5): 27-29.

POULTON, E. B., 1900. Hijpolimnas misippus, Linn., taken in the Atlantic
Ocean. Entomologists' Record, 12: 80-81, 315-316,

POULTON, E. B., and N. D. RILEY, 1928. The Rhopalocera of the “St.
George” Expedition, from French Oceania. Transactions Entomological
Society London, 76: 453-468.

1935. Butterflies from the Marquesas. B. P. Bishop Museum Bulletin,

114' 299-303

RAINBOW, W, J., 1896-1900. The insect fauna. In: R. Etheridge, The Atoll
of Funafuti, Ellice Group: its zoology, botany, ethnology, and general
structure. Australian Museum, Memoir III, 609 pages (see pp. 90-95).

RAMOS, J. A., 1946. The insects of Mona Island (West Indies). /. Agr.
Univ. Puerto Rico, 30: 1-74 (see pp. 52-54).

REBEL, 1939. Rhopalocera of Madeira. Ark. Zool. 32 A (3): 1-5, 2 plates.

REED, E. C., 1877. Monografia de las Mariposas Ghilenos. Anales Univ.
Santiago, 1766: 647-736.

REMINGTON, G. L., 1954. Order Lepidoptera. In: G. T. Brues, A. L.
Melander, and F. M. Carpenter, Classification of insects. Bulletin
Museum Comparative Zoology, 108: 206-305. [Good bibliography]

RIBBE, C., 1889. Beitrage zur Lepidopteren Fauna von Gross-Ceram. Iris,
2: 187-265.

1898. Beitrage zur Lepidopteren-Fauna des Bismarck-und Salomon

Archipels in der Siid-See. Iris, 11: 35-133.

RIDLEY, H. N., 1888. Notes on the zoology of Fernando Noronha. Journal
Linnean Society London, Zoology, 20: 473-570 (see pp. 473, 543).

RILEY, N. D., 1923. The Rliopalocera of the Mt. Everest 1921 Expedition.
Transactions Entomological Society London, 1922: 461-483.

1956. The genera of Holarctic Theclinae: a tentative revision. 10th

International Congress Entomology, 1: 281-288.

RINDGE, F. H., 1948. Contributions toward a knowledge of the insect
fauna of Lower California. No. 8. Lepidoptera: Rhopalocera. Pro-
ceedings California Academy Sciences, 4th Series, 24: 289-312.

1952. The butterflies of the Bahama Islands, British West Indies

(Lepidoptera). American Museum Novitates, no. 1563, 18 pages.

1955. The butterflies of the Van Voast- American Museum of Natural

History Expedition to the Bahama Islands, British West Indies. Ameri-
can Museum Novitates, no. 1715, 19 pages, 1 plate.

ROBER, J,, 1922. Lepidoptera, Rliopalocera. Nova Guinea, 13 (Zoologie
3): 43-50.

ROEPKE, W., 1955. The butterflies of tlie genus Delias Hiibner (Lepi-
doptera) in Netherlands New Guinea. Nova Guinea, n. s., 6: 185-260,
plates V-VIII.

ROJAS, E. U., 1964. Catalogo de Lepidopteros de Chile. Bol. Museo Nac.
Hist. Nat., 28 (2) : 67-140, + introduction and index.

ROSS, G. N., 1967. A Distributional Study of the Butterflies of the Sierra
de Tuxtla in Veracruz, Mexico. #67-14,010. Unpublished dissertation,
Louisiana State University. Ann Arbor: University Microfilms, Inc.

ROTHSCHILD, L., 1915. On the Lepidoptera in the Tring Museum sent
by Mr. A. S. Meek from the Admiralty Islands, Dampier, and Vulcan
Islands. Novitates Zoologicae, 22: 192-208, 387-402.

1925. Critical list of the collection of Algerian Lepidoptera of the

late Captain N. J. E. Holl. Novitates Zoologicae, 32: 195-229.

ROTHSCHILD, W., 1895. A revision of the Papilios of the Eastern Hemi-
sphere, exclusive of Africa. Novitates Zoologicae, 2: 167-463.

212

OAKLEY SHIELDS

/. Res. Lepid.

SAALMtiLLER, M., 1884. Lepidopterin von Madagascar. 531 pages, 14
plates. Frankfurt: Werner und Winter.

SAFFORD, W. E., 1919. Natural history of Paradise Key and the nearby
Everglades of Florida, Annual Report Smithsonian Institution, 1917:
377-434 (see pp. 399-404).

SALVIN, O., and F. D. GODMAN, 1877. On a collection of Lepidoptera
made by the Rev. G. Brown on Duke-of-York Island and its neighbor-
hood. Proceedings Zoological Society London, 1877: 139-149.

SCHULTZE, W., 1925. New and rare Philippine Lepidoptera. Philippine
Journal Science, 28: 567-576.

SGHWEIZER, F., and R. G. W. KAY, 1941. Lepidopteres del Uruguay.
II. Catalogo sistematico, Part 1: Rhopalocera & Grypocera. Anales
Museo Hist. Nat. Montevideo (2d. Serie), 5 (3): 4-24, 1 map.

SGOTT, J. A., 1970 [1971]. A list of Antillean butterflies. Journal Research
Lepidoptera, 9: 249-256. [Excellent comprehensive bibliography]

1972. April. Biogeography of Antillean butterflies. Biotropica, 4(1):

32-45.

SCRIBER, J. M., 1973. Latitudinal gradients in larval feeding specialization
of the world Papilionidae (Lepidoptera). Psyche, 80 (4): 355-373.

SGUDDER, S. H., 1875. The introduction of Danaida plexippus into the
Pacific islands. Psyche, 1: 81-84.

1876. A cosmopolitan butterfly. American Naturalist, 10: 392-396,

602-611.

1876. A brief comparison of the butterfly faunas of Europe and

eastern North America, with hints concerning the derivation of the
latter. Proceedings American Association Advancement Science, 1876:
268-273.

1889. The Butterflies of the Eastern United States and Canada,

with Special Reference to New England. 1958 pages, 89 plates. Cam-
bridge, Mass.: W. H. Wheeler.

1889. Cosmopolitan butterflies. Psyche, 5: 190-192.

— 1893. The tropical faunal element of our southern Nymphalinae

systematically treated. Proceedings American Academy Arts Sciences,
27: 236-251.

SEMPER, G., 1886-1902. Reise im Archipel der Philippinen, Zweiter Theil.
Bde. V & V/. Rhopalocera und Heterocera. Wiesbaden.

1906. (1905). Beitrag zur Lepidopterenfauna des Karolinen-Archi-

pels. Iris (Deutsche Entomologische Zeitschrift), Dresden, 18: 245-
267.

SEOK, D. M., 1939. A Synonymic List of Butterflies of Korea (Tyosen).
391 pages. Seoul, Korea: Published by the Korea Branch of the Royal
Asiatic Society.

SHARPE, E. M., 1893. Descriptions of new species of butterflies from the
Island of St. Thomas, West Africa. Proceedings Zoological Society
London, 1893: 553-558.

SHELFORD, R., 1904. A list of the butterflies of Borneo with descriptions
of new species. Part I. Journal Straits Branch Royal Asiatic Society,
41: 81-111.

1906. A list of the butterflies of Borneo, and Nymphalinae. Part II.

Journal Straits Branch Royal Asiatic Society, 45: 89-136.

SHIROZU, T., 1947. A critical study on the zoogeography of Japanese but-
terflies with special reference to the importance of the West-Chinese
elements. Matsumushi, 2: 1-8. [In Japanese]

1960. Butterflies of Formosa in Colour. 481 pages, 76 plates. Osaka,

Japan: Hoikusha.

SLANSKY, F., Jr., 1972. Latitudinal gradients in species diversity of the
New World swallowtail butterflies. Journal Research Lepidoptera, 11:
201-217.

13(3):169-178, 207-216, 1974

BIBLIOGRAPHY

213

SMITH, H. G., 1894. An account of a collection of diurnal Lepidoptera
made by Mr. W. Doherty at Humboldt Bay, Dutch New Guinea, and
in neighboring islands, in the Museum of the Honourable Walter
Rothschild at Tring, with descriptions of new species. Novitates
Zoologicae, 1: 332-365.

SMITHERS, C. N., 1970. Observations on Lord Howe Island butterflies.
Australian Zoologist, 15 (3): 377-379.

— — 1971. A note on Lord Howe Island butterflies. Journal Australian
Entomological Society, 10: 299-300.

SMITHERS, C. N., and J. V. PETERS, 1969. The butterflies of Norfolk,
Philip and Nepean islands. Australian Zoologist, 15 (2): 185-187.
SNELLEN, P. C. T., 1878-1879. Lepidoptera van Gelebes. Tijdschr. Ent.,
21: L43; 22: 61-126.

SONAN, J., 1939. On the butterflies from Botel Tobago Island. Transactions
Natural History Society Formosa, 29; 257-263. [In Japanese]

— 1940. M. Yanagihara's collection from Daito-Islands, Okinawa: But-

terflies. Transactions Natural History Society Taiwan, 30: 369-375,
376-380, + illustrations.

SOUTH, R., 1888. Distribution of Lepidoptera in the Outer Hebrides,
Orkney, and Shetland. The Entomologist, 21: 28-30, 98-99.
STAUDINGER, O., 1888. Lepidopteren der Insel Palawan. Berlin Ento-
mologische Zeitschrift, 32: 273-280.

1889. Lepidopteren der Insel Palawan. Deutsche Entomologische

Zeitschrift Lep., 1889: 3-180.

STEMPFFER, H., 1967. The genera of African Lycaenidae (Lepidoptera:
Rhopalocera ) . Bulletin British Museum (Natural History), Entomology
Supplement, 10: 322 pages.

STEMPFFER, H., and T. H. E. Jackson, 1962. A note on the Rhopalocera
of Bugalla Island, Sesse Isles, Uganda. Proceedings Royal Entomo-
logical Society London (B) 31: 33-37.

SWANEPOEL, D. A., 19^. Butterflies of South Africa: Where, When
and How They Fly. 320 pages, 17 plates. Gape Town: Maskew Miller
Ltd.

SWEZEY, O. H., 1920. The butterflies of the Samoan Islands, Proceedings
Hawaiian Entomological Society, 4: 601-605.

— 1926. Lepidoptera, pp. 73-79. In: E. H. Bryan, Jr., Insects of

Hawaii, Johnston Island and Wake Island. B. P. Bishop Museum
Bulletin, 31: 1-94.

■ 1939. Distribution of Lepidoptera in Pacific island groups. Pro-

ceedings Sixth Pacific Science Congress, 4: 317-324.

1942. Butterflies of Guam. B. P. Bishop Museum Bulletin, 172:

31-38.

SZENT-IVANY, 1955. Butterflies of the Port Morsby area in the light of
zoogeography. Papua New Guinea Science Society Annual Report
Proceedings, 1954: 91-94.

TALBOT, G., 1939. The Fauna of British India, including Ceylon and
Burma. — Butterflies. Vol. 1, 2nd edition.

TORRE Y CALLEJAS, S. L. de la, 1954-1955. An annotated list of the
butterflies and skippers of Cuba ( Lepidoptera, Rhopalocera ) . Journal
New York Entomological Society, 62: 1-25, 113-128, 189-192, 207-249.
TOXOPEUS, L. J., 1926. Lycaenidae Australasiae I. Treubia, 8: 365-375.
— — 1927 (?). Lycaenidae Australasiae III. Treubia, 9: 423-435.

— 1929. Beschreibung einiger schmetterlinge (Riodinidae und Lyca-

enidae) von Pulu Weh bei Sumatra. Tijdschr. Ent., 72: 204-214.

- — — - 1929. De Riodinidae en Lycaenidae van het eiland Java. Tijdschr.
Ent., 72: 215-244.

— 1944. Results of the Archbold Expedition to New Guinea. Lepi-
doptera: Riodinidae ( Erycinidae ) . Treubia, “Hors Serie”, 1944: 156-
193.

214

OAKLEY SHIELDS

J. Res. Lepid.

1950. The geological principles of species evolution in New Guinea.

Proceedings Eighth International Congress Entomology: 508-522.

1951. Java in Sumatra ( Lepidoptera ) . Idea, 8: 102-103. [In Dutch;

discusses great convergence of Rhopalocera of Kaliandra region in S.
Sumatra witli fauna of Java.]

TSUTSUMI, M., 1940. Butterflies of the South Sea islands. Saishu to
Shiiku, 2 (2): 54-55. [In Japanese]

TUCKER, R. W. E., 1952. The insects of Barbados. Univ. Puerto Rico,
Jour. Agr., 36: 330-363.

TURNER, A. J., 1925. The present distribution and past history of our
Australian butterflies. Memoirs Queensland Museum, 8: 115-122.
UICHANCO, L. B., 1938. Experience with a raised coral reef as hunting
ground for butterflies in Mindanao, Philippines. Verh. VII. Intern.
Kongr. Ent., 1 : 437-443.

VALLETTA, A., 1971. The Butterflies of the Maltese Islands. 64 pages.
Malta: Giov. Muscat and Company.

VAN DUZEE, E. P., 1933. The Templeton Crocker Expedition of the
California Academy of Sciences, 1932. No. 12. The diurnal Lepidoptera
of the expedition. Proceedings California Academy Sciences, 4th Series,
21 (12): 139-145.

VAN EECKE, R., 1922. Studies on Indo-Australian Lepidoptera II. The
Rliopalocera, collected by tire Third New Guinea Expedition. Nova
Guinea, 13 (Zoologie3): 55-79, plates I-III.

1924-1932. List of the Lepidoptera, collected by Mr. W. C. Van

Heurn during an exploration-expedition in Dutch North New Guinea.
Nova Guinea, 15 (Zoologie 4) : 33-56, plate I.

1926, Fauna Buruana. Lepidoptera Rhopalocera. Treubia, 7 (1):

351-370, plate VIII.

1932. On some Lepidoptera from the Riouw- Archipelago. Treubia,

14: 3-9.

VERITY, R., 1929. The Asiatic origins of the western Palaearctic Rhopa-
locera exemplified by Melitaea didyma, Esp. Entomologists'" Record
& Journal Variation, 41: 31-34.

VIETTE, P. E. L., 1950. Lepidopteres Rliopaloceres de L’Oceanie Francaise.
Faune de L’ Union [Empire] Francais, 13: 1-101.

1952. Lepidoptera. Results of the Norwegian Scientific Expedition

to Tristan da Cunha 1937-1938, no. 23, 19 pages, 3 plates.

1953. Lepidopteres recoltes aux lies Gilbert ( Pacifique Central).

Ann. Ent. Soc. France, 122: 123-130.

1955. Notes on the Lepidopterofauna of Madagascar. Lepidopterists’

News, 9: 61-66.

1956. Faune de Madagascar. II. Insectes Lepidopteres Hesperiidae.

85 pages.

1960. Les Lepidopteres Rliopaloceres de Madagascar: biogeographie

et repartition. Bull. Soc. Zool. France, 85: 353-360.

VINSON, J., 1938. Catalogue of the Lepidoptera of the Mascarene Islands.

Mauritius Institute Bidletin, 1 (4): 1-69.

WAGNER, W. H., Jr., and D. F. GRETHER, 1948. The butterflies of the
Admiralty Islands. Proceedings United States National Museum, 98:
163-186.

WAKABAYASHI, H., 1937. Butterflies in March of the Bonin Islands.
Zephyrus, 7: 76-77. [In Japanese]

WALKER, J. J., 1881. Entomological collecting on a voyage to the Pacific.

Ent. Monthly Magazine, 18: 81-86.

1883. Entomological collecting on a voyage to the Pacific. Ent.

Monthly Magazine, 20: 91-96.

1886. Anosia plexippus, L. ( Danais archippus, F. ) : a study in geo-
graphical distribution. Ent. Monthly Magazine, 22: 217-224.

13(3):169-178, 207-216, 1974

BIBLIOGRAPHY

215

1914. The geographical distribution of Danaida plexippus, L.

( Danais archippus, F. ) witli especial reference to its recent migra-
tions. Ent. Monthly Magazine, 50: 181-193, 224-237,

1919. The fringes of butterfly life. Transactions Entomological So-
ciety London, 1919: Ixxxix-cxvii, 1 map.

1922. On the occasional occurrence of butterflies in Iceland; with

notes on the Lepidopterous fauna of tlie North Atlantic islands. Ent.
Monthly Magazine, 58: 1-7.

WALL, F., 1921. Butterflies at sea. Journal Bombay Natural History So-
ciety, 28: 293.

WALLACE, A. R., 1865. On the phenomena of variation and geographical
distribution as illustrated by the Papilionidae of the Malayan Region.
Transactions Linnean Society London, 25: 1-71.

■ 1867. On the Pieridae of the Indian and Australian Regions. Trans-

actions Entomological Society London, (3) 4: 300-416, plates VI-IX.

WARREN, B. C. S., 1926. Monograph of the tribe Hesperiidi. Transactions
Entomological Society London, 1926: 1-170.

— 1944. A review of the classification of the Argynnidi with a system-
atic revision of the genus Boloria (Lepid.: Nymphalidae ) . Transactions
Entomological Society London, 94: 1-101.

WARREN, W., 1905. Lepidoptera collected by W. R. Ogilvie-Grant on the
Azores and Madeira in 1903. Novitates Zoologicae, 12: 439-447.
[Mentions only Pieris hrassicae and Lycaena phlaeas.]

WATERHOUSE, G. A., 1897. The Rhopalocera of Lord Howe Island.
Proceedings Linnean Society New South Wales, 22: 285-287.

1904. On three collections of Rhopalocera from Fiji, and one from

Samoa. Transactions Entomological Society London, 52: 491-495.

1920. Descriptions of new forms of butterflies from the south Pacific.

Proceedings Linnean Society New South Wales, 45: 468-471.

1932. What Butterfly is that? A Guide to the Butterflies of Australia.

291 pages, 34 plates. Sydney: Angus and Robertson, Ltd.

WEGNER, A. M. R., 1953. On a collection of Rhopalocera from Panaitan
Island, with description of a new species. O. S. R. - Publication, No.
36, 7 pages.

WEST, B. K., 1966. Butterflies of New Providence Island, Bahamas. Ento-
mologists Record, 78: 174-179, 206-210.

WHEELER, L. R., 1942. Butterflies of Penang Island. Entomologists’ Rec-
ord, 54: 39-41, 53-57.

WILLIAMS, C. B., 1930. The Migration of Butterflies. 473 pages. Edin-
burgh: Oliver and Boyd.

WILLIAMS, C. B., G. F. COCKBILL, M. E. GIBBS, and J. A. DOWNES,
1942. Studies in the migration of Lepidoptera. Transactions Royal
Entomological Society London, 92: 101-283.

WILLIAMS, F. X., 1911. The butterflies and hawk moths of the Galapagos
Islands. Proceedings California Academy Sciences, 4th Series, 1: 289-
322.

WILLIAMS, J. G., 1969. A Field Guide to the Butterflies of Africa. 238
pages. London: Collins.

WILTSHIRE, E. P., 1957. The Lepidoptera of Iraq. 162 pages, 17 plates.
London: Nicholas Kaye Ltd.

1962. Studies in the geography of Lepidoptera, VII: Theories of the

origin of the West Palearctic and world faunae. Entomologists’ Record,
74: 29-39.

— 1964. The Lepidoptera of Bahrain. Journal Bombay Natural History

Society, 61: 119-141,

WISE, K. A. J., 1967. Taxonomy of two New Zealand butterfly species
(Nymphalidae: Satyrinae). Transactions Royal Society New Zealand,
9: 39-44, 1 plate.

WOLFF, N. L., 1964 The Lepidoptera of Greenland. Meddelelser om
Gronland, 159 ( 11 ) : 74 pages, 21 plates.

216

OAKLEY SHIELDS

/. Res. Lepid.

WOODFORD, C. M., 1885. Remarks upon Lepidoptera collected in the
Ellice and Gilbert Islands, Annals Magazine Natural History, Ser. 5,
15: 414-416.

1895. The Gilbert Islands. Geogr. J., 6: 325-350 (see p. 348).

WOODHOUSE, L. G. O., 1949. The Butterfly Fauna of Ceylon, 2nd edi-
tion. 231 pages, 55 plates. Golombo: The Golombo Apothecaries’ Go.
Ltd.

WU, G. F., 1938. Catalogus Insectorum Sinensium. The Fan Memorial In-
stitute of Biology, Peiping. Vol. 4, pp. 695-941. [Rhopalocera]

WYATT, G. W., 1956. Lepidoptera collecting in the Atlas Mountains of
Morocco. Lepidopterists News, 10: 214-222.

1961. Additions to the Rhopalocera of Afghanistan with descrip-
tions of new species and subspecies. Journal Lepidopterists’ Society,
15: 1-18.

WYNTER-BLYTH, M. A., 1957. Butterflies of the Indian Region. 523
pages, 72 plates. Bombay: The Bombay Natural History Society.

YASHIRO, H., 1939. Insects of Minami-Daito Island (Loo-choo). Biogeo-
graphica,3i 127-130. [In Japanese]

YOKOYAMA, M., 1955. Coloured Illustrations of the Butter-flies of Japan.
136 pages, 63 colored plates. Osaka: Hoikusha. [In Japanese]

ZEUNER, F. E., 1943. Studies in the systematics of Troides Hiibner (Lepi-
doptera, Papilionidae ) and its allies; distribution and phylogeny in
relation to the geological history of the Australasian Archipelago.
Transactions Zoological Society London, 25: 107-184.

ZWALUWENBURG, R. H. van, 1943. The insects of Ganton Island. Pro-
ceedings Hawaiian Entomological Society, 11: 300-312.

THE JOURNAL OF RESEARCH
ON THE LEPJDOPTERA

CONTENTS

Volume 11 Number 1 March, 1972

THE BUTTERFLIES OF NEBRASKA

KURT JOHNSON

Volume 11 Number 2 June, 1972

The Neotropical Nymphalid Butterfly,

Anartia amalthea Michael K. Fosdick 65

On the distribution of

some Skippers in Ontario J. C. E. Riotte 81

Observations on life history of

Oarisma powesheik (Parker) 1870

Wilbur S. Me Alpine 83

Habitat: Oeneis jutta ascerta
Masters & Sorensen

John H. Masters 94

Dispersal in a cosmopolitan

butterfly species {Pieris rapae)
having open population structure

Thomas C. Emmel 95

Mating of Butterflies James A. Scott 99

Book Review —

Butterflies of Georgia F. A. Urquhart 128

Volume 11 Number 3 September, 1972

IN THIS ISSUE
Gynandromorphism in Pieris brassicae L.

Brian O. C. Gardiner 129

Two new subspecies of Euphydryas

chalcedoria from the Mojave desert of
Southern California

T. C. Emmel and J. F. Emmel 141

Butterflies of middle and southern

Baja California Richard Holland 147

Vital staining as evidence for wing

circulation in the cabbage butterfly

Pieris rapae John M. Kolyer 161

A dwarf form of Euptoieta ckiudia

Russell A. Rahn 174

Butterfly records for three northwest

Wisconsin counties John H. Masters 152

Variations of Parasemia parthenos

A. E. Brower 183

A search for Speyeria nokomis coerulescens

(Holland) (Nymphalidae) in southern Arizona

Ronald S. Wielgus 187

A bilateral gynandroinorph of Limenitis
tveidemeyerii latifascia (Nymphalidae)

E. M. Perkins and T. F. Perkins 195

An interfamilial courtship (Nymphalidae, Pieridae)

Arthur M. Shapiro 197

Addition to : “A checklist of Utah butterflies
and skippers

Curtis J. Callaghan and Norman B. Tidwell 199

Volume 11 Number 4 December, 1972(1973)

IN THIS ISSUE

Latitudinal gradients in species diversity
of the new world swallowtail butterflies

Frank Slansky, Jr. 201

Lack of melanism in Colias John H. Masters 218

A population study of Euptychia
hermes in northern Florida

Thomas S. Kilduff 219

Limenitis weidemeyerii angustifascia

X L. ostyanax arizonensis = (?) ab. doudoroffi
( Gunder ) 1934.

E. M. Perkins, Jr. and John S. Garth 229

A new subspecies of Gonepteryx amintha

(Pieridae) from Yunnan, mainland China.

Yuri P. Nekrutenko 235

Reduction of abdominal scales of the Monarch
butterfly imago as a result of cauterizing
the Abd PPM of the pupa. F. A. Urquhart 241

Some techniques for the rearing of
Megathymus larvae

Ronald S. and Dale Wielgus 245

Concerning Heliconius cydno aberration

“larsenf Niepelt John H. Masters 251

Polymorphism in two species of

Alaskan Boloria C. D. Ferris 255

A new species of Nephelodes Guenee

from the Great Basin John S. Buckett 260

NOTICES

B ooks on Lepidoptera: more than 300 books for sale. Free
catalogue sent on request. Sciences Nat. , 45 Rue de Alouettes, 7 5019,

Paris, France

Wanted to make exchanges with collectors in California and Florida,
Have Maylaysian butterflies for exchange (Trogonoptera brookiana, etc. )
Masaki NAKAYAMA, Kitakyushu- Wakamatsu- Miyamaru -Z-10-14,
Fukuoka pref. , Japan .

Need aid in preparation of specimens for study of population
structure in butterflies; involves obtaining population samples, mounting,
laboratory breeding, etc. Full or part time. Contact William Hovanitz,
1160 W. Orange Grove Ave, Arcadia, Calif. 91006.

Volume 13 Number 3 September, 1974

IN THIS ISSUE

Studies of the ova and first instar

larvae of Geometridae (Ennominae). L

Roger L. Heitzman 149

Altitudinal migration of central California butterflies

Arthur M. Shapiro 157

The chromosomes of Apantesis phalerata,

A. radians, and their hybrid in Florida populations
(Arctiidae) Jack S. Bachelor and

Thomas C. Emmel 162

A partial bibliography of the world distribution
and zoogeography of butterflies

Oakley Shields 169

The nomenclature in an important British checklist (1972)

Part 2: Corrections of family-group names for
Geometridae ( Lepidoptera ) Juraj Paclt 179

Illustrations and descriptions of

species of some Pyrrhopyginae from Panama
( Hesperiidae ) S. S. Nicolay 181

Butterflies of the Suisun Marsh, California

Arthur M. Shapiro 191

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Journal of Research on the Lepidoptera

13(4):217~238, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

TOWARD A THEORY OF
BUTTERFLY MIGRATION

OAKLEY SHIELDS

Department of Entomology
University of California, Davis, California 95616

INTRODUCTION

It is difficult to know what observations are important or
critical in explaining why and how butterflies migrate, but since
so many have been made, some of these are bound to be of
significance. Regrettably, there is a nearly total lack of experi-
mental work. A good working model is sorely needed; the lack
of this has no doubt severely hampered progress in this field.
This report is my attempt to extract a working hypothesis from
the morass of published data.

Butterfly migrations can reach spectacular proportions. In
1926 in East Africa about 36 million Belenois mesentina crossed
each mile of front per day, and in 1924 in southern California
about 3 billion Vanessa cardui crossed a 40-mile front (Williams,
1949)1 On the other hand, flights may be “so thin that each
butterfly may be hundreds of yards from its nearest companion,”
out of sight of each other (Williams, 1949).

The two questions which most commonly occur are: (1)
Why do they migrate?, and (2) What determines the direction
of migration? (Abbott, 1950).

Various authors have offered their definition of butterfly
migration. (Poulton, 1902, says that emigration is a better term
than migration, for masses of insects moving out of an over-
crowded area, but the term does not specify that a undirectional
flight also occurs.) Wiltshire (1946) says as a general rule,
migrant Lepidoptera do not perform a diapause. “A species can
be considered a migrant if it fulfills most or all of the following
provisions: — (1) it has been observed performing a mass move-
ment; (2) it has been taken, or seen on the wing, at sea; (3) it
shows no geographical variation over a wide and diverse area;
( 4 ) both sexes occur occasionally, or sporadically, far from where
it is known to breed and reside.” “A migration is recognized

217

218

OAKLEY SHIELDS

/. Res. Lepid.

when the butterflies, whether few or many, are all at the same
time flying in the same direction, as if guided by a compass”
(Abbott, 1951). Williams (1957) says that insect migration
refers “to movements of animals in a direction and for a distance
over which they have control, and which result in a temporary
or permanent change of habitat.” “Migration is a persistent,
straightened-out movement with some internal inhibition of the
responses that will eventually arrest it” (Kennedy, 1961). John-
son (1960) says that insect migration is so variable that defini-
tions based on what is or is not migration or what is active
migration and passive dispersal become arbitrary and unsatis-
factory when applied to all migratory insects.

EXAMPLES

Details of five species of migrants are presented below which
are well-known in the literature, in hopes of revealing under-
lying causes.

VANESSA CARDUI

Williams (1925) says that V. cardui in Europe in the late
spring fly mostly NW. These flights come from North Africa
and Asia Minor, although not from the countries immediately
surrounding the Mediterranean (probably from the deserts to
the south ) . “The movement appears to continue for many weeks
or even months at irregular intervals.” He says there is no evi-
dence of a return flight in this region in autumn. It cannot
overwinter in the north in any stage. “So far as we can see, all
those butterflies which fly to the north of the area in which they
can breed throughout the year are lost completely to the species,
as either they or their offspring perish during the winter.”
Migrant individuals have large fat body reserves and unde-
veloped internal sex organs.

Abbott (1950) records that 1924 was a great migration year
for V. cardui in southern California, while in 1925 only 2 were
seen the entire spring. 1926 was another migration year, though
not in such large numbers as 1924. (Thus perhaps 1925 was a
population crash. ) “To maintain their straight path of migration,
butterflies regularly rose over obstacles such as hedges and
trees, even tall eucalyptus trees and three story buildings, yet
they consistently kept within a few feet of the ground other-
wise.” The beginning of the daily flight was noted at 6:50 am
on April 25, 1941, when 6 V. cardui which apparently had been

13(4):217~238, 1974

BUTTERFLY MIGRATION

219

“sleeping” on the lawn flew up, circled around more or less,
and flew off to the northwest. (Similarly, migrant V. cardui
I released in a room lighted by a ceiling light flew up to the
light in a spiral flight, at night.) Abbott (1951) notes that in
1941 there were three waves: March 9-March 28, April 4-7, and
April 15-22. In 1945 waves occurred on March 19-21, March
24-25, and March 27-29. “The same territory is covered in every
migration, the extent varying with the total numbers.” Flights
are from February to April, averaging just under 2 months in
extent. Migrations occur during years of vegetation abundance,
though significantly not every “wildflower year” is a migration
year. Southern California migration years were 1924, 1926,
1941, 1945, 1949. Tilden (1962) noted V. cardui moving NNE
on May 18-19, 1958, in Oregon, and on May 20-21, moving SSW,
perhaps due to meeting a cold front. He saw one V. cardui
migration in light to heavy rain, and rather cold, though he
thinks it unlikely the flight was initiated in the rain. Sampling
of 2 flights by Thorne and Sette, and Tilden, yielded a 50-50
sex ratio. Tilden thinks V. cardui must reproduce along the
line of flight.

On April 15-20, 1973, I encountered V. cardui migrating
NNW (even against a wind) in vast numbers on the Eastern
Mojave Desert, E of Ludlow 20-40 mi., by the thousands. The
desert was carpeted with wildflowers. Desert blooms hit by the
larvae included Amsinkia, Lupinus, Salvia, Phacelia campanu-
laria, every Boraginaceae, and even 2 larvae on a crucifer.
V. cardui was noted migrating and feeding in the pm and
avidly feeding in the early am on white flowers in washes.

ASCIA MONUSTE

Hayward (1953) saw a migration of A. monuste in Argen-
tina during the spring and summer of 1951-52. “On fine morn-
ings the overnight ground population began to move with the
first sign of daylight,” seeking flowers and puddles. “Although
the butterflies sometimes commenced their migratory flight di-
rectly from the flowers on which they were feeding, it was more
usual for them to circle and drift upwards till they had attained
certain height and there assemble before moving off.” The mi-
grations flew in various directions: 9 am = SSE then veering E;
11:30 = N (WSW wind). On December 3rd there was a mas-
sive migration that began to lessen at c. 3000', with individuals
seen by airplane pilots as high as 50001 On December 7th at
9:15 am one flight at ground level was going E, while a high-

220

OAKLEY SHIELDS

/. Res. Lepid.

flying swarm that was earlier going E was flying N! On January
7th the flight continued until darkness. On January 14, there
was heavy overcast and no migration was observed. In one
migration on October 28th, 300 specimens were taken at light
on a moonless night. Females were observed laying eggs during
one flight. “Ascia monuste on migration flies close to the ground
when the wind is adverse and high when the wind is favorable.
There is a tendency to fly low on cloudy days and higher in
bright sunshine. The migratory swarms may reach an altitude
of over 3000 ft., and mountain ranges of up to at least 5000 ft.
in height do not cause the migrants to deviate from their course.”

Nielsen & Nielsen (1950) report a migration of A. monuste
at Fort Pierce, Florida, was at a maximum on April 15, and the
maximum of the next outbreak was May 20, some 36 days later.
They observed that a mass outbreak was necessary for a migra-
tion (“a simultaneous hatching of a whole population within a
few days”). A. monuste migrates “en masse while there seems
still to be ample food on which they could continue to breed in
the habitat” (Johnson, 1969, p. 224).

URANIA MOTHS

Smith (1972) notes of Urania moths in the Neotropics, that,
“Aside from the periods of population explosion and migration,
the distribution of Urania moths over the vast area is rather
localized,” in low densities. “The migration is to the south and
east in the autumn and westward and northward in the spring.”
Movement is basically east-west. The eastward movements
( July-October ) involve far more individuals than do the west-
ward ones. “The most frequent interval between large flights
appears to be 8 years, but there are numerous exceptions.” In
general there is no “return spring flight” in years following a
large eastward movement in the autumn. Migrants fly all day
long. Larvae eat the vine Omphalea ( Euphorbiaceae ) . Some
flight had many gravid females, others did not. In 1969, in the
second and third weeks the frequency of gravid females in-
creased. Copulation was rarely observed; no oviposition was
observed during a flight. Release from selection pressure of
predators and parasites combined with food abundance “must
be responsible for the really major population explosions that
have occurred.” Generally when an eruption of Urania fulgens
occurred in Central America, a simultaneous eruption occurred
in U. leilus in South America.

13(4):217-238, 1974

BUTTERFLY MIGRATION

221

DANAUS PLEXIPPUS

Tilden (1962) says that the monarch has a true migration,
involving “a going out and a return,” i.e. a cyclic movement.
The migrants, however, do not fly in mass (Heape, 1931, p. 151).
Migratory flights of monarchs end in aggregations for hiber-
nation and thus differ from other migrant species where the
flights lead to dispersal. In autumn, monarchs migrate south
in the U.S. (Canada and northern U.S.), to southern California,
Florida, Mexico, and hibernate gregariously in trees. They
breed and migrate (disperse) north the following spring. Their
foodplants grow in exposed and often semi-arid areas. Adults
depart from the cold. To survive in situ as adult diapause in
winter would be fatal. There are one to three generations in
the north before next autumn. Flights are up to 1500 miles
(tagged specimens). Urquhart (1960, p. 296) notes that ‘re-
covery lines” for tagged monarchs in eastern U.S. indicate a
general SW flight in the fall. Spring migration records in
California for tagged monarchs show a general NE or NNE
trend (fig. 79, p. 320). Monarchs do seem to be affected by
landmarks, water bodies, etc. in their migration flights.

Funk (1968) found D. plexippus in December at Yuma,
Arizona, as ova, larvae, pupae, and adults. The implication is
that a return flight might be from this new brood rather than
from overwintering adults. (Marked specimens could not be
recaptured in the area.)

Beall (1948) says that D. plexippus “freshly emerged in the
autumn in Ontario start with a considerable fat reserve, some
30 per cent of the lean weight.” Young migrants add even more;
they have fat as high as 125% of lean weight. Migrants taken in
New Orleans, presumably from a long journey, have fat only
2% of lean.

NYMPHALIS CALIFORNICA

Yothers ( 1913 ) reported that in 1912, N. calif ornica larvae
at a number of places in Washington, Idaho, and British Colum-
bia were “by the millions,” defoliating Ceanothus velutinus.
At Clayton, Washington, the larvae had eaten all the leaves,
and the branches were entirely bare except for the millions of
larvae. On July 7, 1912, he found millions of larvae on Moscow
Mt., Idaho. Then on July 13th, all the larvae and pupae were
gone from the place of infestation. “The caterpillars had evi-

222

OAKLEY SHIELDS

/. Res. Lepid.

dently not migrated, for all around as far as I could see the
Ceanothus had not been touched. Even had the caterpillars
migrated that would not explain the absence of the chrysalids.
I think that the total disappearance of these caterpillars and
chrysalids was no doubt due to birds.”

On May 11, 1973, in Del Puerto Canyon, 22-23 rd. mi. W
of Patterson, Stanislaus Co., Calif., I noted N. calif ornica larvae
all over most of the Ceanothus integerrimus bushes, stripping
some bare, although some bushes were untouched (all stages
of instars above ca. second or third). (Adult hibernants were
fairly common there on March 11, 1973.)

William Neeley (pers. comm.) saw “millions” of N. call-
fornica one day during the first week of August, 1962, on the
summit and sides of Triple Divide Peak, Yosemite, California,
flying east from the SW San Joaquin drainage in a band %-l
mile, 114-30' in height, from noon to three pm; some were
landing.

EXTRA SPECIES JOINING MIGRATION

One puzzling aspect of butterfly migrations, especially those
in the tropics, has been the accompaniment of many species in
the migration. Eor example. Welling (1959) notes one migration
consisting “of just about everything imaginable,” flying from
N to S in the Yucatan Peninsula, especially Anteos clorinde and
A. maerula, with Eunica tatila and E. monima, made up the bulk
of the migration. Also Papilio sp., Heliconius charitonius,
Etirema species, many Libytheana carinenta, Danaus sp., a few
blues, Phoebus, He mentions (1964) Eunica monima, Agraulis
vanillae, “and a few other odds and ends” accompanying a large
migration of Kricogonia castalia lijside in Yucatan. Such obser-
vations appear to be explained by the following observation
recorded by Poulton (1929, 1930): Mr. F. Muir observed “the
sweeping up of the non-migrating butterflies on a Papuan island
when a migratory flight from another island passed over it. In
this instance, in which more than a single species was involved,
it is evident that the social stimulus, and this alone, availed to
compel the non-migrating butterflies to become migrants— with
such success indeed that the island was comparatively depleted
of these species after the migratory stream had passed.”

13(4);217-238, 1974

BUTTERFLY MIGRATION

223

MAJOR SPECIES OF MIGRANTS

The following species from Williams (1930) appear to be
frequent, conspicuous migrants: Catopsilia florella (Africa), C.
sennae (Americas), C. statira (South America), C. pyranthe
(India, Ceylon), C. pomona (Ceylon, India, Java), C. crocale
(Ceylon, India), Belenois severina (Africa), B. mesentina (Af-
rica, India), B. java teutonia (Australia), Appias albina (Cey-
lon, India), A. paulina (Ceylon), Ascia monuste (Americas),
Danaus plexippus ( circumcontinental ) , Vanessa cardui (world-
wide), Nijmphalis calif ornica (U.S. and Canada), Libythea
bachmannii (U.S.), L. carinenta (Americas), L. labdaca (Af-
rica), and the moths Urania leilus (Neotropics) and U. fulgens
(Central America). It is perhaps of significance that these
species belong to the Pieridae, Libytheidae, and Nymphalidae,
a set of families that are evolutionarily inter-related (Shields,
unpublished libytheid study.) Also, all, of course, are common
to very common species, and many are known to be strong, fast
fliers with two or more broods and comparatively rapid gener-
ation time. One migrant female Vanessa cardui produced 685
larvae in captivity (Schrader, 1928).

It is known that Ascia monuste has a dark form (phileta)
that is the migratory phase (Klots, 1951). I have noticed that
Vanessa cardui migrants are larger with brighter colors than
the non-migrant phase, generally. A parallel might be drawn
here with Uvarov’s phase theory of locusts, with solitary and
migratory phases of the same species differing in morphology,
coloration, physiology, and behavior (Poulton, 1929; Williams,
1957).

START OF A MIGRATION

Despite innumerable reports of butterfly migrations in prog-
ress, rarely have observations of their actual commencement
been recorded. The circumstances surrounding the start of the
flight are expected to shed the most light on the illusive cause
of butterfly migration and perhaps the origin of the navigation
of the flight as well. Accordingly, the details of published ac-
counts of the start of migration that I have been able to locate
are given in full.

Sir Guy Marshall recorded that the migration of Catopsilia
florella was actually taking place from a Rhodesian valley
(Poulton, 1931).

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Pitman (1928) gives an account of the crowded breeding-
grounds of Belenois mesentina in the West Nile Province of
Uganda, from which start the great southward migrations in
Uganda and Kenya. In a grassy plain some 2 miles broad and
7-10 miles in length their foodplant {Maerua oblongifolia, Cap-
paridaceae) was abundant “in full, tender leaf and flower,” in
March 1928 following a few heavy thunder-showers after which
the weather continued exceptionally hot and dry. The area
had evidently been burned out some weeks previously. Most
of the foodplants were leafless; “larvae and chrysalises were to
be seen everywhere, as also countless thousands in cop” soon
after emergence. “I have never before seen such countless
myriads of butterflies, and as far as the eye could see there was
a shimmer of white just above the surface of the ground.” Pre-
ceding this emergence the West Nile was in drought for nearly
twelve months. The adults were of small size, evidently due to
starvation from the over-population. No actual migration of the
adults, however, was observed.

Heape (1931, pp. 148-150) records a migration of Catopsilia
(?) in the states of Ceara and Piauhy, Brazil, the end of Janu-
ary or early February, 1915, after the rains had started, from
the mangrove swamps on the right bank of the River Camocim;
“the flight was somewhere to the north of west.” 36 hours after
the flight began, it was followed on horseback by two men for
iYz days and 125 miles. “They diminished markedly in numbers
the further westward they travelled.” By the time the town of
Sao Pedro was reached, some 50 miles further, the flight had
ceased (“petered out”). One observer returned to Camocim and
learned “there that the flight continued to emerge from the
mangrove swamp for forty-eight hours after he had left the
place, and then a heavy storm put an end to those which still
remained in the swamps and the flight stopped.”

“It is of particular interest to note that the route followed
by these gentlemen passed over the high ground east of the
Paranahyba River, and from there on to Sao Pedro, through
sandy hills and rocky flats where the scrub was dried up,
although it was the rainy season; and all over this country large
numbers of the butterflies were seen lying dead. They most
certainly had not laid eggs there, and it was judged they fell
from exhaustion while en route, leaving a trail of dead bodies.”

Collenette (1928) witnessed the start of a migration of
Libijthea carinenta. Between Corumba and Urucum, Matto
Grosso, Brazil, November 1927, “extraordinary numbers” covered

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muddy areas in the road all day long from November 16-21.
November 22-23 was rainy and cool, with the mud practically
deserted. Then on November 24, a bright and clear day, the
migration started, many flying S and SSE starting about 7:30
am, while large assemblages were still on the mud. “At one
such mud gathering, two or three in a minute rose and joined
the passing stream, taking the general direction at once, without
hesitation. In the instances which I was able to watch closely,
there seemed to be no preliminary fluttering or walking about
before taking to wing, and those which rose did not chase or
play with those already in flight.” None of the passing migrants
were seen to settle on the mud, though they “frequently paused
and fluttered over the gathering.” He noted only three or four
instances of pairing. At 9:45 am he left for Corumba in the
direction from which the carinenta were coming “in much the
same numbers.” “The insects quickly became fewer, and in
about a mile I reached the drier vegetation, in which practically
none were present, and where no migration was visible.” He
believed the swarm had “bred locally or had been attracted
from the drier country in the neighbourhood.”

Skertchly (1879) discovered the pupae of Vanessa cardui
very numerous on grass blades between the mountains and
desert in March 1869 west of Sowakin in the Sudan of Africa,
soon after daybreak. “Presently the pupae began to burst, and
the red fluid that escaped sprinkled the ground like a rain of
blood. Myriads of butterflies limp and helpless crawled about.
Presently the sun shone forth, and the insects began to dry
their wings; and about half-an-hour after the birth of the first,
the whole swarm rose as a dense cloud and flew away eastwards
towards the sea. I do not know how long the swarm was, but
it was certainly more than a mile, and its breadth exceeded a
quarter of a mile.”

Abbott (1951) records that John Garth found Vanessa cardui
perhaps near the start of a migration at Santa Maria Bay, Baja
California, 600 miles SSE of San Diego. “Thousands” of adults
were seen, many of them emerging. Many flew south a short
distance to the edge of the cliff, then north again to the breed-
ing area, but no migration as such was in progress. It is prob-
ably significant that they were observed in a migration year,
and at about the same dates as in California.

Emmel and Wobus (1966) noted Vanessa cardui near Floris-
sant, Colorado in large densities (but no mass movement) from
mid-July to mid-August, 1965. Then on August 22nd they ‘Tegan

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flying about due south and south-southwest in vast numbers, and
this mass movement, involving every individual seen, continued
until August 25. By this date, almost all V. cardui had left this
area but scattered individuals were observed flying south
through August 28, when a cold front moved in and stopped all
butterfly activity for several days.” The flight started at 7:00
am to nearly dusk. Between September 1-19, V. cardui con-
tinued moving southward in reduced numbers. On September
20th a severe cold front moved in, apparently curtailing the
migration.

END OF A FLIGHT

Williams (1930, p. 353) says migrants “frequently pass over
large areas of land apparently entirely suitable for their inhabi-
tation, but continue on and on as long as the migration hysteria
lasts, and may end up in the open sea or in entirely inhospitable
lands where breeding is impossible.” “There are records of
swarms of butterflies reaching the coast and flying steadily on
out to sea” (p. 389). At Arolla, Switzerland, August 6, 1903,
V. cardui in large numbers of big faded specimens “invaded
the district and settled down, each on its own piece of land,
to alternate duties of fighting with a neighbour and egg-laying”
(Williams, 1930).

LACK OF RETURN FLIGHTS
It seems a paradox that migrations of V. cardui and others
are unidirectional and nearly constant when they occur, yet
return flights to replentish the migrant gene pool appear rare
or non-existent. If accurate, somehow the species is repetitive
in behavior in migration years (a mass exodus serving as a
“safety valve” on the population). Thus perhaps most but not
all of the migrant population departs during years of migration.

EFFECTS OF OVERCROWDING
According to Poulton (1902), “The [migration] instinct . . .
compels the individuals to move together in vast masses in the
same direction, rather than to scatter and fly in all directions.”
“The sugegstion is made that the crowded masses, resulting
from over-production and inability of enemies to cope with the
increase, are injurious to the species, because it is likely that
food-plants would be checked for years or even killed out
altogether in certain localities, while the heaps of dead indi-
viduals would encourage the attack and rapid spread of bac-

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terial foes.” The advantages derived from removal of the surplus
from an overcrowded area would account for the development
by natural selection of the instinct to move. Otherwise, there
would be a destruction of the species in the area of over-
production.

Poulton (1921) quotes Trimen as saying, “The instinct to
emigrate probably exists in a dormant state in all species liable
to outrun the food-supply in any part of their range.” If the
overcrowding is unchecked, it would threaten the existence of
the foodplant over a wide area. Near Mombasa, Africa, Liby-
thea laius and other butterflies “appear only in occasional years
. . . after a period of prolonged and severe drought,” [when
parasites, diseases, predators at a low, vegetation lush, and
numbers would thus build quickly.] In some cases the migrant
has stripped the leaves and shoots of its foodplant, such as
Catopsilia florelh on Cassia arachoides in Griqualand West in
1881.

Heape (1931) says, “All the evidence I have gleaned from
investigation of the voluntary movements of the higher animals
points, to my mind, definitely to the conclusion that emigration
[of butterflies] is induced by overcrowding and the scarcity of
food” (p. 158.) “Whatever may be the cause of these mass
movements the result is that thereby huge surplus populations
are effectually disposed of and the world saved from the incal-
culable devastation they would cause if they lived” (p. 171).
“I have already suggested that the access of virility which
attends the growth of the gonads and the development of repro-
ductive cells in the higher animals, may well be the exciting
cause of that stimulus which urges towards adventure of such
kind [migration]; and that the resulting condition of a horde
of individuals so affected my attain to what we call hysteria.
I further suggest that, in spite of the radical differences both
in the sense organs and in the central nervous system of insects,
a condition comparable to hysteria may similarly be induced
in them” (p. 170). “These conditions [i.e., the periodicity of
migrations] are probably of climatic origin and affect the quality
of the food supply and so govern reproductive activity.”

In observations of migrating Lepidoptera often the “genital
products” are in a backward condition (i.e., immature), and a
migratory flight is necessary for their full development. Thus
the stimulus to migrate may be a physiological one (Poulton,
1929).

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Chapman (1939) mentions that Cause talks of “relaxation
oscillations” arising from a relaxation or a decrease of environ-
mental resistance, followed by a sudden population increase
causing outbreaks. In insects having a high biotic potential,
this relaxation during a single generation may result in a popu-
lation increase of outbreak proportions.

Alexander (1961, fig. 63) suggests that field crickets under
high density, crowded conditions showed no male territoriality
and dispersed widely, compared with individuals in isolation
which developed male territoriality and dispersed little.

It should be mentioned that Colias eury theme often forms
dense swarms numbering as much as some migrations of other
species, but is not known to migrate unidirectionally (F. T.
Thorne, pers. comm,).

INHIBITION OF STIMULI

Kennedy (1961) thinks “Migrants are distinguished by a
transient accentuation of locomotor functions with depression
of vegetative functions, such that the insect now travels.” “The
Nielsens listed the reflex patterns of adult Ascia monuste ‘living
in a territory’ as feeding, mating, egg-laying, basking and
cleaning, and reported failure to respond to the appropriate
stimuli for any of these when the butterflies were engaged in
unidirectional flight elsewhere.” “Ecologically, migration some-
times looks like an alternative to diapause when conditions will
no longer permit growth, development or reproduction.” “The
internal inhibition of certain reflexes is the behavioural com-
ponent of a whole physiological syndrome,”

POST-TENERAL EXODUS FLIGHT

According to Johnson (I960), “Whenever the start of a
‘mass migration’ of insects has been described — and this is very
rare in spite of a voluminous literature— the insects have always
been making either their first flight as new adults or one very
soon afterwards.” He makes reference here to Catopsilia crocale,
C. pyranthe, Ascia monuste, Vanessa cardui, and Nymphalis
californica doing so, from the breeding site. There was only
ca. I hour between emergence and migration in two instances
of Vanessa cardui, “The initial orientation of the insect in flight
is probably ‘imprinted’ during the teneral period, or determined
at take-off.” “Too long an enforced wait before take-off, leading
to sexual maturation or fertilization, apparently inhibits migra-
tion altogether in A. monuste”

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OVIPOSITION

In Catopsilia pyranthe in Ceylon, “every female seemed
possessed with the one insane idea of getting rid of her eggs
with the utmost expedition . . . and then madly continuing her
flight” (Poulton, 1921). Belenois mesentina females in a migra-
tion at Peradeniya, Ceylon, were often seen “to stop and lay
eggs on bushes of Capparis pedtmculosa, rejoining the migra-
tory flight as soon as a few eggs had been laid” (Poulton, 1929).

SEX RATIO

Williams (1930, p. 344) says males mostly predominate in
pierid migrations checked (24 cases), while females predomi-
nated in two instances (of Catopsilia). In five instances of
V. cardui, females predominated (Europe), while in two in-
stances males predominated (p. 345). A migration of Catopsilia
pyranthe and C. pomona in Ceylon in November 1905 had the
main flight 87% males, but later stragglers were almost entirely
females (p. 346). (Males generally emerge first before females
in butterflies.)

PARALLEL BETWEEN
MAMMAL AND BUTTERFLY MIGRATIONS

Deevey (1960) delves into cause of lemming migrations;
they are neurotically sick during migration. The “shock disease”
causes them to die in numbers “from their own excitement.”
Food, predators, disease, and weather periodically relax a hold
on lemming numbers. It is the younger lemmings that migrate.
Also, the end result is a wipe-out (population crash) of migrat-
ing individuals largely, in lemmings (and certain insect migra-
tions). When there are large numbers of rats, these display a
“pathological togetherness” (i.e., prefer to live together, low
fertility, shortened lives). In mammals, Christian (1950) re-
ported that there is a sudden die-off in late winter and early
spring following a population peak, due to the exhaustion of
the adreno-pituitary system (“shock disease”). “The terminating
factor is the attainment of a population above the carrying
capacity of the environment.” The population is under highly
stressed conditions and taxes the adreno-pituitary system to
the maximum.

Direct experimental evidence that butterfly migrations under-
go a similar “shock disease” is lacking. However, there is
observational evidence that this syndrome does in fact take

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place. Bemheim ( 1917 ) records a massive Libythea bachmannii
migration near Eagle Pass, Texas, September 4, 1916. “There
were literally millions of them and many of them had evidently
completed their alotted span of life as they were dropping from
the air in large numbers. One particular specimen flew past,
almost brushing my face, and, as I thought, alighted upon the
ground. As I stooped to examine it the wings suddenly folded
down tightly in front of the body and 1 picked the insect up
quite dead.” Heape (1931, pp. 148-150) reported large numbers
of a Catopsilia (?) sp. lying dead after its migration through
the territory, in Brazil. On April 15-20, 1973, 1 noticed fresh
Vanessa cardui bodies forming windrows along the hiway dur-
ing a migration in the Eastern Mojave. Some appeared to fall
dead to the pavement from the flight merely from the wind of
passing cars. Hayward (1953) says that on December 26, 1951,
there was an Ascia monuste migration in Argentina present in
the millions, with thousands of dead on the ground, many in
fresh condition.

POSSIBLE COBBELATION WITH SUN-SPOTS

Grant (1937) has noted a correspondence in years of out-
break (40 out of 60) in Celerio lineata moth migrations in
Europe and North America. “In seeking for an explanation of
the outbreaks of L Uvornica and 1. lineata, one must look for
some large cause, wide enough in its effects to have influenced
both continents simultaneously.” She found some relationship
between the outbreaks and 11-year sunspot cycles over a 100
year period (fig. 5). “The number of outbreaks would seem
to rise from the minimum towards the maximum, and then
fall away towards the next minimum . . . ,” with some excep-
tions. She also found a strong tendency for outbreaks to occur
when a wet year has followed a dry year. Williams (1965,
p. 158) found a tendency for simultaneous abundance or rarity
in North America and Europe in V. cardui from 1865-1938,
statistically more significant than Grant’s Celerio lineata cor-
relations! Williams, however, said he could find no apparent
correlation in C. lineata and V. cardui outbreaks to sun-spot
cycles. With V. cardui there were above average numbers at
both the maximum and minimum sun-spots, with below-average
numbers on rising and on falling periods in the cycle.

Brown (1974) mentions there are recent suggestions of
interrelationships between solar activity, the Earth’s magnetic
field, and the weather. Periods of maximum sun-spot activity

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show an 11-year cycle. Wood & Lovett (1974) found a highly
positive correlation of drought years in Ethiopia and sunspot
minima ( from 1540 to 1974 ) . Over a 72 year period there, “The
rainfall clearly follows a cyclic pattern in which rainfall peaks
and troughs precede the sunspot peaks and troughs by a few
years” (i.e., to 2 years). “Thus, there is a strong statistical cor-
relation between Addis Ababa rainfall and sunspot number, the
rainfall peaks leading the sunspot peaks by an average of 1.3
yr.” Thus there would indeed appear to be some correlation
between V. cardui outbreaks and the sunspot cycle, i.e., occur-
ring after several years following rainfall peaks and extreme
droughts.

RAINFALL

Williams (1965, p. 23) says, “Painted Ladies breed during
the winter along the edges of the great North African desert
belt, and then move north in the spring across the rest of North
Africa, the Mediterranean and Europe.” In Ceylon, the NE
monsoon starts in about October with very heavy rainfall. “This
is followed immediately by great flights of butterflies,” lasting
for about 3 months, “and then, after a lull, there is a recrudes-
cence of activity in March and April corresponding to the end
of the north-east and the beginning of the south-west monsoons”
(p. 53). At Portachuelo Pass, 3500', Venezuela, Beebe noted
250 species of butterflies “migrating” through; “the flights seem
to be associated with the rainy season, which lasts roughly from
April to November” (p. 44).

WIND

Of 367 recorded observations of directional flight by butter-
flies in which the direction of flight and wind had been recorded,
there were little or no significant differences in proportion among
flights with the wind, diagonally with and against the wind,
across the wind, and against the wind (Williams, 1949).

DIRECTION OF FLIGHT

Williams (1949) notes that in the tropics, flights of the same
species tend to recur in the same direction at the same season
in different years. “On reaching an obstacle most migratory
butterflies pass over it rather than round it. . . .” “Migratory
butterflies have also at times been observed to follow the wind-
ings of deep valleys, of roads through forest, and of shore-
lines. . . .” “There are ... a few observations of simultaneous
flights of two or more species, each flying in a different direc-

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tion through the same locality for many days, and each keeping
to their own direction.” The movements of both such species
had no apparent eflPect on the other. It is often noticed that
butterfly migrations cease or are much reduced when the sun
is clouded. ‘'Orientation is quite definite at midday in the tropics
when the sun is so near the zenith as to be practically useless
for horizontal orientations” (Williams, 1957).

Flights of migrants that follow the shore line and do not
fly out to sea, e.g., Ascia monuste, Erynnis zarucco funeralis,
and Urania fulgem, suggest that topographic features play a
part in determining the success of a migration, rather than it
being a totally suicide flight.

Welling (1964) found Calpodes ethlius in Panama in its
migrations “does not adhere to one single direction, mention
being made of it passing in almost every direction’ in the course
of a single afternoon and evening.” He saw one migration going
N to S, slowly changing from W to E, then finally changing
from SW to NE in one hour. They were therefore flying circles
within that particular grassy swamp.

A Catopsilia sennae migration continued to fly during a
tropical thunderstorm with heavy rain but oriented randomly
(Johnson, 1969, p. 155). In a migration of Catopsilia statira
in Trinidad, Williams noticed that when he attempted to catch
examples, “any butterfly narrowly missed was put off its direction
by the excitement and flew off wildly in any direction. Other
butterflies close at hand meeting this butterfly flying out of the
general order would in turn become confused and sometimes
follow it in its new direction. So that after several misses in
succession I was surrounded by a number of butterflies flying
in all directions. If I stopped attempting to catch specimens
these would gradually pass away, and the regular direction of
flight would be resumed” (Poulton, 1921).

I reported (1967, p. 112) that migrations of Vanessa cardui,
Danaus plexippus, and a Delias sp. are known to pass over
summits of hills and mountains. No “staying” at the hilltop was
shown by these. One migration of V. cardui on Dictionary Hill
summit flew upslope from the south, flew across the summit,
and instead of flying down the other side flew up and out off
the north slope. Others were seen to pass over the hill’s shoul-
ders. No concentration toward the summit by the flight was
noted. This could be a “breakdown in territoriality” (F. T.
Thorne, pers. comm.), as resident cardui continued to “hilltop”
that day. Wright (1906, p. 37) records a migration of V. cardui

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on top of “Grayback,” 11,600', that was going due east. They
flew up the slope of the mountain and upon reaching the top,
where the crest dropped off suddenly, “they kept straight on
as far as the eye could follow them, right up into the sky. . .

A V. cardui migration was seen flying over a 1300m summit in
Austria in 1877 (Williams, 1930).

Tulloch (1912) reports an Atella phalantha migration on
Mauritius, on the Trou-aux-Cerfs, an extinct volcano. Immense
numbers flying up the crater towards the east; at the top they
flew to the other side of the crater, then downhill to the bottom,
where they went “right-handed round the base of the volcano'"
until they reached where he had been before, where they went
uphill. “In fact, the butterflies were all going on an endless
round, up the hill, across the top, down again, and round to
where they started." The atmosphere was very clear, bright,
and hot; a westerly breeze was blowing.

Abbott (1951) says V. cardui in southern California fly to
the NW mostly, but also NNW or N. He says one would
suppose butterflies following a definite compass direction would
fly across a canyon from rim to rim, but instead V. cardui flies
down one side and up the other, “keeping about the same
distance from the ground all the time.” “Butterflies which ap-
proached the lee side of a hill, flying NW, changed their course
when they came to the top of the hill, turning and flying against
the SSW wind. While they were turning, one single butterfly
was seen to keep on toward the northwest.” He notes that some
upon encountering a wall in their path rose in the air in a
“spiral” until clearing the roof, proceeding then NW.

NAVIGATION

Vleugel (1952) believes that the sun is the governing factor
in the orientation of butterflies, based on bird migration studies.
He supposes that somehow the “migrating butterflies allow for
the change of angle necessitated by the apparent movement of
the sun.” However, Williams (1965, p. 126) defeats the argu-
ment for a sun-compass direction that allows for the angle
change necessitated by the apparent movement of the sun:

(1) the migrant would have to allow for the rising of the sun
each morning at a 180° angle from sunset as flights resume
each morning in the same direction as the previous evening,

(2) when delayed for several consecutive days by bad weather
they still resume their flight in the correct direction, and (3)

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they can migrate at mid-day in the tropics when the sun is
overhead and therefore useless for direction finding. No known
organ in insects seems likely to be sensitive to the Earth's mag-
netic field, according to Williams.

Perhaps a means of circumventing these difficulties is to
assume that the freshly emerged migrant adults navigate by
using polarized light at the time of day they would normally
start exhibiting territoriality and “freezing” upon this angle
(D. A. Watts, pers. comm.). This fixed polarization would over-
come the problem of having to adjust to a changing sun angle
throughout the day. Vanessa cardui spring migrations in north-
ern Africa-Europe and in California both trend mainly NW
(compare Williams, 1925, fig. 2 with Abbott, 1951, fig. 1), while
in the southern hemisphere in SE Australia, V. cardui kershaivi
migrations fly predominantly S to SW during spring (August
to November for some 7-8 weeks) (Smithers, 1969). Vanessa
normally exhibits “hilltopping” and territorial behavior in the
afternoon starting from 12:30-3:15 pm PST (Shields, 1967) but
peaking in late afternoon, while feeding at flowers in the morn-
ing. The sun in the afternoon would be in the SW quadrant
of the sky in the northern hemisphere (above the Tropic of
Cancer) and in the NW quadrant in the southern hemisphere
(below the Tropic of Capricorn) in the spring, so that the
V. cardui migration direction would be “fleeing” at nearly right
angles from the sun’s incident light. This is along the direction
of the plane of polarized light, as can be seen with a Polaroid
sheet, l.e., in the morning in the northern hemisphere above
the Tropic of Cancer, the plane of polarization rotates from
N to E and S to W, while in the afternoon it rotates from
W to N and E to S. For Libythea bachmannii the majority of
the migration records are toward the E or SE in July-September
(SE predominant) in Texas., and Libythea labdaca flies in
March-May to the SW, SSW, S, SE, ESE in Nigeria and Gold
Coast, with one record of L. laius in May in Tanganyika flying
N (Williams, 1930). Here the Libythea would be using the
plane of polarization to fly toward the sun. A summary of ca.
25 migrant species from Kodaikanal, S. India by Evershed (Wil-
liams, 1930, p. 297) showed primarily N-NE, S, SE movements
in six species in Febmary-March [sun slightly southward],
northward and NE-E-SE movements in May-June [sun slightly
north], and 100% southward movements of nearly all the species
in August-November (primarily October) [sun southward to far

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southward]. Waterman (1955) says that light from the sun as
transverse electromagnetic waves is partly polarized by the
scattering of light by air molecules. To the human eye, this
polarization is nearly invisible.

Mazokhin-Porshnyakov (1969) says that sunlight is polar-
ized up to 70-80%. It is known that various aquatic crustaceans,
the horseshoe crab, some terrestrial arachnids, and winged in-
sects ( certain Coleoptera, Hemiptera, Diptera, Hymenoptera,
Trichoptera, and Lepidoptera) react to light polarization; both
faceted eyes and simple ocelli are sensitive to polarized light.
Honeybees can quickly learn to distinguish between two light
beams polarized in different planes. Impulses within the optic
lobes arose following a rapid 90° rotation of the polarized plane
in honeybees. “According to Autrum and Frisch, the insect’s
ability to recognize different polarization planes is due to the
radial distribution of the visual cells in the forms of a rosette.
If we suppose that each visual cell is sensitive to polarized
light in a given plane, then the whole rosette would act as a
polarization analyzer.” “Electron microscopy investigations of
the rhabdom in insects and other arthropods . , . revealed that
polarization analysis is a property related to the rhabdomeres’
periodic ultrastructure which repeats itself in different omma-
tidia. . . . Organic molecules absorb light when it is polarized
in the same direction as the longitudinal axis of the chromophore.
Since the visual pigment molecules are regularly oriented within
the rhabdomeres . . . and absorb more strongly when the light
is polarized in preferred planes (dichroism), they could . . . act
as a polarization analyzer” (p. 141).

Vowles (1950) noted that ants {Cataglyphis, Monomorium,
Myrmica) maintained a constant angle of orientation relative to
the plane of polarization of light, in experiments ( field and lab ) .
With Myrmica, by altering the plane of polarized light with a
Polaroid sheet, a new angle was assumed by the ant to the dorsal
beam of plane-polarized light (his graph), but not to one of
ordinary light. A tortricid larva {Archips cerasivorana) that
crawled toward the sun was induced to turn 90° by placing a
polarizing sheet over it as it moved, with continued maintenance
of the new course under the influence of the polarizing sheet
(Wellington, 1955). “It is common to see both larvae and adults
fall victims of heat stroke on a sheet of paper while they are
engaged in ineffectual circling movements in response to rapid
changes in the sky,” due to clouds or smoke. “. . . the sun is
primarily a heat source, and . . . the plane of polarization is

236

OAKLEY SHIELDS

/. Res. Lepid.

the primary means of maintaining orientation to the sun . . .
when an insect crosses an open space in daylight,” according
to his experiments. “The compound eye of an insect is fixed on
the head and seems to be especially suitable for recognizing
the polarization over the whole sky at once, providing that we
assume that it can act as an analyzer of polarized light” (Frisch,
1950, pp. 90-96).

NIGHT MIGRATION?

Reports of butterflies migrating at night may be spurious
unless actually observed to do so. The only migration claimed
to be observed taking place at night that I am aware of is
“Eagle Clark’s observation . . . that ... V. cardui flew toward
England from the Continent against a wind at nighf' ( Shannon,
1915, p. 618). Kendall & Click (1972) record many instances of
diurnal butterflies collected at visible and ultraviolet light at
night, including some migrant species. “Evidence indicates most
diurnals must be disturbed from their resting places before they
appear at induced light” (p. 275). Thus, claims of butterflies
migrating at night because they were taken at lights are prob-
ably unjustified. Their high density in the vicinity would account
for some turning up at light. An exception would seem to be a
report by Kingdon (1932) of a ship 87 mi. at sea with Vanessa
cardui at lights at midnight thought to have come on board
the previous hour. There was a strong off-shore wind, and had
been all day. However, Williams (1930, p. 342) noted indi-
viduals of a swarm of Pieris hrassicae resting on the surface of
the ocean and taking flight when disturbed, so it is possible the
cardui were roosting nearby on the water. Even large-scale
migrations take place only from sunrise or somewhat later
(start) until sunset or dusk (cease) (Williams, 1930; Hayward,
1953; Smith, 1972). Birds can use celestial navigation in their
migrations (Wallraff, 1960) but butterflies do not appear to
do so.

BIBLIOGRAPHY

ABBOTT, C. H., 1950. Twenty-five years of migration of the Painted Lady
Butterfly, Vanessa cardui, in southern California. Pan-Pac. Ent. 26:
161-172.

, 1951. A quantitative study of the migration of the Painted Lady

Butterfly, Vanessa cardui L. Ecology 32: 155-171.

ALEXANDER, R. D., 1961. Aggression, territoriality, and sexual behavior
in field crickets (Orthoptera: Gryllidae). Behavior 17: 130-223.
BEALL, G., 1948. The fat content of a butterfly, Danaus plexippus Linn.,
as affected by migration. Ecology 29: 80-94.

BERNHEIM, J. L., 1917. Swarms of butterflies. Ent. News 28: 339-340.
BROWN, G. M., 1974. A new solar-terrestrial relationship. Nature 251:
592-594.

13(4);217-238, 1974

BUTTERFLY MIGRATION

237

CHAPMAN, R. N., 1939. Insect population problems in relation to insect
outbreak. Ecol. Monogr. 9: 261-269.

CHRISTIAN, J. J., 1950. The adreno -pituitary system and population cycles
in mammals. /. Mammalogy 31: 247-259.

COLLENETTE, C. L., 1928. A migration of Libythea carinenta carinenta
Cr, Ent. Mo, Mag. 64: 124-126.

DEEVEY E. S., Winter 1960. The hare and the haruspex: a cautionary
tale. The Yale Rev. 49: 161-179.

EMMEL, T. C. & R. A. Wobus, 1966. A southward migration of Vanessa
cardui in late summer and fall, 1965. /. Lepid. Soc. 20: 123-124.

FRISCH, K. VON, 1950. Bees: Their Vision, Chemical Senses, and Lan-
guage. Cornell Univ. Press, Ithaca, 119 p.

FUNK, R. S., 1968. Overwintering of monarch butterflies as a breeding
colony in southwestern Arizona. ]. Lepid. Soc. 22: 63-64.

GRANT, K. J., 1937. An historical study of the migrations of Celerio
lineata lineata Fab. and Celerio lineata Uvornica Esp. ( Lepidoptera ) .
Trans. Roy. Ent. Soc. London 86: 345-357.

HAYWARD, K. J., 1953. Migration of butterflies in Argentina during the
spring and summer of 1951-52. Proc. Roy. Ent. Soc. London, Ser. A,
28: 63-73.

HEAPE, W., 1931. Emigration, Migration and Nomadism. Heffer, Cam-
bridge, 369 p.

JOHNSON, C. G., 1960. A basis for a general system of insect migration
and dispersal by flight. Nature 186: 348-350.

, 1969. Migration and Dispersal of Insects by Flight. Methuen, Lon-
don, 763 p.

KENDALL, R. O., & P. A. CLICK, 1972. Rhopalocera collected at light in
Texas. J. Res. Lepid. 10: 273-283.

KENNEDY, J. S., 1961. A turning point in the study of insect migration.
Nature 189: 785-791.

KINGDON, D., 1932. Pyrameis cardui, L., observed flying at midnight,
between Madeira and Bathurst. Proc. Roy. Ent. Soc. London 7: 56-57.

KLOTS, A. B., 1951. A Field Guide to the Butterflies. H. Mifflin Co., Bos-
ton, 349 p.

MAZOKHIN-PORSHNYAKOV, G. A., 1969. Insect Vision. Trans, by R. &
L. Masironi. Plenum Press, New York, 306 p.

NIELSEN, E. T., & A. T. NIELSEN, 1950. Contributions toward tlie
knowledge of the migration of butterflies. Amer. Mus. Novit. no. 1471,
29 p.

PITMAN, C. R. S., 1928. The area, in the West Nile Province of Uganda,
from which start the great southward migrations of Belenois mesen-
tina. Cram., in Uganda and Kenya Colony. Proc. Roy. Ent. Soc. Lon-
don 3: 45-46.

POULTON, E, B., 1902. The gregarious instinct in hybernation and emi-
gration of insects. Trans. Ent. Soc. London 50: 460-465.

- — = — , 1921. Notes on the migration of Lepidoptera, with a suggestion as
to the cause of the backward and foreward flight occasionally observed.
Trans. Ent. Soc. London 69: xii-xxvii.

—, 1929. The Uvarov theory of locust migration and its possible bear-
ing upon butterfly migration. Proc. Ent. Soc. London 4: 16-20.

— , 1931. The problem of butterfly migration. Nature 127: 807-809.

SCHRADER, W., 1928. Experiments on a species of migrating butterfly.
Bull. So. Calif. Acad. Sci. 27: 68-70.

SHANNON, H, J., 1915. Do insects migrate like birds? Harpers Mag. 131:
609-618.

SHIELDS, O., 1967. Hilltopping: an ecological study of summit congre-
gation behavior of butterflies on a southern California hill. /. Res.
Lepid. 6: 69-178.

238

OAKLEY SHIELDS

/. Res. Lepid.

SKERTCHLY, S. B. J., 1879. Butterfly swarms. Nature 20: 266.

SMITH, N. G., 1972. Migrations of Rie day-flying moth Urania in Central
and South America. Caribbean J. Sci. 12: 45-58.

SMITHERS, C. N., 1969. A note on migrations of Vanessa kershawi (Mc-
Coy) (Lepidoptera: Nymphalidae) in Australia, 1963-1968. Australian
Zool 15: 188-194.

TILDEN, J. W., 1962. General characteristics of the movements of Vanessa
cardui ( L. ) . /. Res. Lepid. 1 : 43-49.

TULLOCH, B., 1912. The immigration of Pyrameis atalanta. The Entomol.
45: 17-20.

URQUHART, F. A., 1960. The Monarch Butterfly. Univ. of Toronto Press,
Ontario, 361 p.

VLEUGEL, D. A., 1962. Wind and orientation of migrating butterflies in
comparison with birds. Verh. XI Intern. Kongr. Ent. 1960, v. 3: 15-19.
VOWLES, D. M., 1950. Sensitivity of ants to polarized light. Nature 165:
282-283.

WALLRAFF, H. G., 1960. Does celestial navigation exist in animals? Cold
Spring Harbor Symposia on Quantitative Biology 25: 451-461.
WATERMAN, T. H., July 1955. Polarized light and animal navigation.
Sci. Amer. 193 ( 1 ) : 88-94.

WELLING, E. C., 1959. Notes on butterfly migrations in the Peninsula of
Yucatan. J. Lepid. Soc. 13: 62-64.

, 1964. Butterfly migrations in southeastern Mexico during 1963 and

1964. J. Lepid. Soc. 18: 229-230.

WELLINGTON, W. G., 1955. Solar heat and plane polarized light versus
the light compass reaction in the orientation of insects on the ground,
Ann. Ent. Soc. Amer. 48: 67-76.

WILLIAMS, C. B., 1925. The migrations of the Painted Lady Butterfly.
Nature 115: 535-537.

, 1930. The Migration of Butterflies. Oliver & Boyd, London, 473 p.

, 1949. Migration in Lepidoptera and the problem of orientation.

Proc. Roy. Ent. Soc. London, Ser. C, 13: 70-84.

— , 1957. Insect migration. Ann. Rev. Ent. 2: 163-180.

— , 1965. 2nd ed. Insect Migration. Collins, London, 237 p.

WILTSHIRE, E. P., 1946. Studies in. the geography of Lepidoptera— III.
Some Middle East migrants, their phenology and ecology. Trans Roy.
Ent. Soc. London 96: 163-182.

WOOD, C. A., & R. R. LOVETT, 1974. Rainfall, drought and the solar
cycle. Nature 251 : 594-596.

WRIGHT, W. G., 1906. The Butterflies of the West Coast. Publ. by the
author, San Bernardino, 257 p., 32 pis.

YOTHERS, M. A., 1913. Eugonia californica Bdv. in the Pacific Northwest.
Canad. Ent. 45: 421-422.

Journal of Research on the Lepidoptera

13(4):239-245, 1974

1160 W . Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

ATRYTONOPSIS HIANNA BIOLOGY
AND LIFE HISTORY IN THE OZARKS

J. RICHARD HEITZMAN' and ROGER L. HEITZMAN'

3112 Harris Avenue, Independence, Missouri

ABSTRACT

The ovum, first and final instar larva, and pupa of Atrytonopsis hianna
(Scudder) are described and illustrated. The biology of the Ozarks popu-
lations is given and the association between A. hianna and Hesperia metea
Scudder is discussed.

INTRODUCTION

In the Missouri and Arkansas Ozarks Atrytonopsis hianna
(Scudder) occurs in local colonies in woodland clearings, cedar
glades, or forest edges in close association with beard grass
( Andropogon gerardi Vitm. ) . This is the only known food plant
for A. hianna in this region. This is also the food plant of Hes-
peria metea Scudder and these two species are consistantly
found together in the same localities. We have reared A. hianna
on numerous occasions from both Missouri and Arkansas popu-
lations. The illustrations are from specimens collected near
Warsaw, Missouri.

MATERIALS AND METHODS

Ova were obtained by confining wild females in nylon chif-
fon bags over potted plants of Andropogon gerardi. Larvae
were reared on potted plants of this grass and on others that
had been planted outdoors and enclosed in screen wire cover-
ings. A Wild M5 stereomicroscope with drawing tube and photo-
tube attachments was used in preparing the illustrations of the
ovum and first instar larva.

DESCRIPTION OF EARLY STAGES
OVUM: Width 1.25 mm, height .80 mm. Color bright lemon
yellow when first layed changing to pale brown with pinkish
accent dorsally. Micropyle coral red. Eclosion in seven to eight
days.

^Research associate, Florida state collection of Arthropods, Division of Plant
Industry, Florida Department of Agriculture and Consumer Services,
Gainesville.

239

240

J. R. HEITZMAN and R. L. HEITZMAN

J. Res. Lepid.

FIRST INSTAR LARVA: Head and prothoracic shield deep
reddish purple, shiny. Freshly emerged larvae yellow, after
feeding all but abdominal segments nine and ten turn pale
green. Surinal plate with long white setae, body thinly covered
with long white hairs, prothoracic color paler. The first instar
larvae are active silk spinners, trailing a silken thread as they
move about. Stadium period: nine days.

SECOND INSTAR LARVA: Head dull orange brown, mandibles
darker brown. Head thickly covered with minute white setae.
Prothorax white, prothoracic shield, slender, shiny black. Body
color dark green, posterior segments dull brown, anal segment
with thick black setae, rest of body sparsely covered with
minute black setae. Integument translucent. The second instar
larval tent is constructed by fastening the top two thirds of two
grass blades together. The top of the tent is left open. Stadium
period: nine days.

THIRD INSTAR LARVA: Head pale orange brown, midcranial
inflection edged with paler lines, stemmata black, mandibles
dark brown. Head thickly covered with a mixture of short and
longer brown setae. Body color pale pinkish brown with green
undertones due to translucent integument, abdominal segments
eight and nine paler, segment ten dull brown. Prothorax creamy
white, prothoracic shield, slender, shiny black. Stadium period:
nine days.

FOURTH INSTAR LARVA: Head dull orange brown, stem-
mata and mandibles black. Head thickly covered with short
pale setae and a few longer white hairs. Body color bright pink
dorsally, anal segment darker, almost black, with some long
black hairs. Rest of body covered with minute white setae,
integument translucent. Abdominal areas dull green. Prothorax
creamy white, prothoracic shield black, slender. Stadium period:
nine days.

FIFTH INSTAR LARVA: Head light brown, Irons pale orange,
labrum dark brown, mandibles and stemmata black. Head thick-
ly covered with short white setae and a few longer white hairs.
Body color lavender pink dorsally, bright pink ninth abdominal
segment, anal segment grayish brown with long black hairs, rest
of body thinly covered with short white setae. Prothorax white,
prothoracic shield, slender, shiny black. Abdomen grayish white.
Spiracles tiny pale yellow dots. The fifth instar larvae live in
a three- to four-inch tube composed of several grass blades
fastened together, open at the top and bottom. These tents are

13(4):239-245, 1974

ATRYTONOPSIS BIOLOGY

241

usually constructed within six inches of the tips of the leaves.
Feeding slows up greatly in this instar with the larvae spending
many days aestivating. Before transforming to the next instar
the larvae seal the tent at top and bottom. Stadium period:
variable up to 19 days.

SIXTH INSTAR LARVA: Head deep red, mandibles black,
frons and area below midcranial inflections deep reddish orange.
Head granulose and thickly covered with short white setae. Body
color light pink dorsally, brighter on abdominal segments eight
and nine, tenth segment darker brown dorsally. There is an in-
conspicuous middorsal gray line. Abdomen and prothorax pale
translucent gray. Spiracles pale orange, inconspicuous. Pro-
thoracic shield, slender, shiny black. The body now has a downy
appearance due to a thick covering of soft white hair. The
larvae in this instar pass through alternate periods of eating and
aestivating. The larval tent is the same as in the fifth instar.
Stadium period: variable, an average of six to seven weeks.
SEVENTH (FINAL) INSTAR LARVA: Head width 3.5 mm,
deep reddish purple, granulose, unmarked but with brown shad-
ing below laterofacial suture lines, deeply cleft at midcranial
inflection, mandibles black. Head covered with short reddish
setae and longer white hairs. Position and size of stemmata as
illustrated. Length of mature larvae 29-32 mm. Body color pale
pinkish lavender dorsally, abdominal segments eight and nine
pale pink, anal segment depressed, pale glossy brown. Pro-
thorax pale gray, prothoracic shield dark brown, surface granu-
lated. Abdomen pale grayish white, thoracic legs pale brown.
Entire body covered with long yellowish white hair, shorter on
the abdominal areas. The prolegs have orange hair and the anal
segment has bright orange setae, coarser and shorter than on rest
of body. Only the last abdominal spiracles are noticeable, they
are pale brown. Final instar larvae have two small wax pad
areas beneath the posterior segments of the abdomen. The final
instar larvae feed consistantly once the cooler damp weather of
September begins and by early October the larvae seal them-
selves in a nest among the leaves at the base of the food plant
to pass the winter as mature larvae, pupating early the follow-
ing spring.

PUPA: Length 18.5-20 mm, width at widest point of wing cases
4.5-4.75 mm, width at eyes 3.50-3.65 mm. Wing cases light
brown with olive tint. Head, eye cases and upper thorax darker
brown. Thoracic spiracles ruby red. Tongue case deep brown,
detached below wing cases. Abdominal segments light orange

242

J. R. HEITZMAN and R. L. HEITZMAN

/. Res. Lepid.

Fig. 1. — Atrytonopsis hianna (Scudder), A, male and female, ventral and
dorsal view of specimens from Warsaw, Missouri. B, C, D, different aspects
of the ovum. E, mature larva.

13(4):239-245, 1974

ATRYTONOPSIS BIOLOGY

243

Fig. 2.- — Atrytonopsis hianna (Scudder), A, first instar suranal plate. B,
pupa, ventral and right lateral aspect. C, first instar setal maps of pro-
thorax, mesothorax, eighth abdominal segment, all in left lateral aspect.

244

J. R. HEITZMAN and R. L. HEITZMAN

/. Res. Lepid.

Fig. 3. — Atrytonopsis hianna (Scudder), A, head of final instar larva,
frontal and left lateral aspect. B, head of first instar larva, frontal aspect.

13(4):239-245, 1974

ATRYTONOPSIS BIOLOGY

245

with pink overcast, intersegmental folds pale orange brown.
Cremaster and anal segments darker brown. Cremaster 2.75 mm
in length, strongly curved ventrally, without hooks but densely
covered at the tip with stiff brown setae. Pupation occurs in the
base of the Andropogon clumps, one to three inches above
ground level in a sealed case of silk and grass leaves.

DISCUSSION

In the introduction it was mentioned that A. hianna occurs
consistantly with H. metea. Both species are univoltine flying
for only a few weeks in the spring but metea normally begins
emergence one or two weeks ahead of hianna. The first hianna
normally appear during the first week in May at Warsaw, Mis-
souri (earlier in Arkansas) and the imagines have disappeared
by the end of May. While both species utilize the same food
plant their different feeding and resting positions seem to effec-
tively preclude any severe competition for the host plant. While
metea in its later instars lives in a tent in the base of the grass
clumps hianna is living in a tent from one to several feet above
ground and does not appear to enter the base of the plants until
ready to hibernate after feeding is completed in the autumn.
A. hianna is an avid flower visitor and females are best collected
by searching the flowers of Verbena, Camassia, and Fragaria
growing in the vicinity of Andropogon. Males exhibit strong
‘perching” tendencies and will dart out at any passing butterfly
or insect that remotely resembles a female hianna. Both sexes
are wary and not easily collected. The numbers of imagines
present from year to year varies greatly but to date no parasites
have been bred from any of the wild larvae collected.

ACKNOWLEDGMENTS

We wish to thank the Missouri State Park Board for furnish-
ing permits pursuant to our studies in the State Parks of Missouri.
Dr. Leo J. Paulissen, Fayetteville, Arkansas provided valuable
field assistance in this project.

Journal of Research on the Lepidoptera

13(4) :246, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

LARVAL MIGRATION OF HYLES LINEATA
(FAB.) (SPHINGIDAE)

JAMES F. WELLS and RICHARD M. BROWN

16711 Marsh Creek Rd. #162, Clayton, California 94517
and

1055 Plaza Dr., Martinez, California 94553.

Larvae of Hyles lineata (Fabricius) were observed in mass
migration on April 4, 1974, 17 miles south of Furnace Creek
Ranch, Death Valley National Monument, Inyo County, Cali-
fornia at an elevation of 300 feet. These larvae were traveling in
a westward direction with a maximum concentration of approxi-
mately fifteen individuals per square meter. The width of the
migration was approximately one fourth mile and the length
is unknown. During the march, larvae fed on the prostrate
lobed leaves of Chorizanthe rigida (Torr) T&G ( Polygonaceae ) ,
a little low growing annual, and then moved on to another plant
when these were gone.

Twenty larvae were taken and reared to adults on apple
( Malus sp. ) . All were in the last instar, although they had much
feeding to do before burying themselves to pupate.

246

Journal of Research on the Lepidoptera

13(4):247=248, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

NEW FOOD PLANT FOR
DARAPSA PHOLUS (CRAMER)
(SPHINGIDAE)

J. C. E. RIOTTE, M.A.

Research Associate, Department of Entomology and Invertebrate Zoology,
Royal Ontario Museum,

100 Queens Park, Toronto, Ontario, Canada, MSS 2C6

Tietz (1972) listed the following plants for Darapsa pholus:
Kalmia angustifolia, Nyssa sylvatica. Rhododendron nudiflora,
Rhododendron viscosa, Tradescantia virginiana. Viburnum den-
tatum, Viburnum lentago. Viburnum opulus.

These records were taken from Beutenmiiller ( 1895 ) , Morris
(1862), Harris (1839), J. E. Smith (1779) and others (Eliot and
Soule, 1902; Fernald, 1886; Baynes-Reed, 1881). I myself
checked additionally the following: Forbes (1948), Holland
(1903), Rothschild and Jordan (1903), Clemens (1859), Fer-
nald (1886, re-checked), Draudt in Seitz (1931) and Hodges
(1971). The food plants for D. pholus mentioned in these pub-
lications are the following: Forbes: Azalea, Viburnum and "re-
ported from other food plants, probably in error for D. myron
and cnotus”; Holland: Viburnum, Azalea; Rothschild and Jor-
dan: same as those of Ampeloeca myron (Ampelopsis, Vitis)
which is certainly in error; Clemens: Azalea nudifiora; Fernald:
Ampelopsis quinquefolia, Viburnum lentago. Viburnum denta-
tum, Viburnum opulus, Nyssa multiflora, Azalea viscosa, Azalea
nudifiora (it seems that the error concerning Ampelopsis origi-
nated with Fernald); Draudt in Seitz: Ampelopsis Vitis (prob-
ably on the authority of Rothschild and Jordan); Hodges:
Azalea Viburnum species.

Tietz (1972) also gave Viburnum as a food plant for D.
myron besides Ampelopsis and Vitis. Hodges (1971) too men-
tioned Viburnum for D. myron and it may be that the erroneous
record of Ampelopsis and Vitis for D. pholus was created in a
kind of exchange between food plants of myron and pholus.
I did not check in which of the many papers quoted by Tietz
for D. myron this Viburnum record originated.

One plant group, however, has never been mentioned among

247

248

J. C. E. RIOTTE

/. Res. Lepid.

the food plants of D. pholus: Vaccinium spp., our common blue-
berries. During field work in 1974 at Queen’s University Biologi-
cal Station at Chaffeys Locks, Leeds Co., Ontario, we found all
stages of D. pholus associated with Vaccinium spp. and it is not
unreasonable to suppose that Vaccinium may be a general food
plant for the species, the more so as several Viburnum species
are also common around Chaffeys Locks but D. pholus was not
found associated with them.

Data for the immatures collected in 1971 are: egg on blue-
berry at Washburn, Ontario, 25 July, hatched and reared suc-
cessfully on blueberry; third instar larva on blueberry on Snake
1. in L. Opinicon, Chaffeys Locks, Ontario, 26 July, reared suc-
cessfully on blueberry; a freshly-emerged, unmated female moth
flushed from a blueberry patch at the Station around noon, 8
July, eggs infertile.

REFERENCES

BAYNES-REED, E. 1881. Sphingidae — Hawk Moths. Twelfth Ann. Rep.
Ent. Soc. Ont.: 48-70.

BEUTENMULLER, W. 1895. Descriptive catalogue of the Sphingidae
found witlrin fifty miles of New York City. Bull. Am. Mus. Nat. Hist.
VII: 275-318.

CLEMENS, B. 1859. Synopsis of North American Sphingidae. Jour. Ac.
Nat. Sci. Philad. N.S. IV, II: 25-51.

DRAUDT, N. 1931. Familie Sphingidae. In Seitz, A., Die Gross-Schmetter-
linge der Erde, VI: Die Amerikanischen Spinner und Schwarmer.
Alfred Kernen, Stuttgart (1940). pp. 845-900.

ELIOT, I. M. and C. G. SOULE. 1902. Caterpillars and their motlis. The
Century Co., New York, 302 pp.

FERNALD, C. H. 1886. The Sphingidae of New England. Sprague and
Son, Augusta. 87 irp.

FORBES, W. T. N., 1948. Lepidoptera of New York and neighboring
states. Part II, Geometridae, Sphingidae, Notodontidae, Lymantriidae.
Cornell Univ. Agric. Exp. Stn., Mem. 274: 1-264.

HARRIS, T. W. 1839. Descriptive catalogue of the North American in-
sects belonging to the Linnaean genus Sphinx in the cabinet of Thad-
deus William Harris, M.D., Librarian of Harvard University. Am. Jour.
Sci. & Arts, 36: 282-320.

HODGES, R. W. 1971. The moths of America north of Mexico including
Greenland. Fascicle 21. Sphingoidea. Hawkmoths. E. W. Classetj and
R. B. D. Publications, London. 158 pp.

HOLLAND, W. J. 1903. The moth book. Doubleday, Page & Co., Garden
City, New York, 480 pp.

MORRIS, J. G. 1862. Synopsis of tire described Lepidoptera of North
America. Smithsonian Institution, Washington, D.C. 358 pp.

ROTHSCHILD, W. and K. Jordan. 1903. A revision of the lepidopterous
family Sphingidae. Novit. Zool. 9, suppl. : 972 pp.

SMITH, J. E. 1797. hi Abbot, J. and J. E. Smith, The natural history of the
rarer lepidopterous insects of Georgia, 2 vol. 214 pp., 104 pi.

TIETZ, H. M. 1972. An index to the described life histories, early stages
and hosts of the macrolepidoptera of the continental United
States and Canada. Publ. by A. C. Allyn for The Allyn Mus. of Ento-
mok, Sarasota, Florida, 2 vol. 1041 pp.

Journal of Research on the Lepidoptera

13(4):249~264, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.

© Copyright 1975

NOTES ON

ARCTIC AND SUB- ARCTIC COLLECTING

CLIFFORD D. FERRIS'

University of Wyoming, Laramie, Wyoming 82071

The ensuing remarks are not intended as an all-inclusive
coverage of arctic collecting. They relate to ^certain areas of
which the author has some knowledge and oflFer comments,
hopefully useful, to those collectors who might be anticipating
a trip to the northlands. Travel to the Far North becomes easier
every year, and more and more collectors will be traveling north
in search of the “treasures” to be found there.

There are several items which every collector going to the
Far North should keep in mind. Of primary importance is
seasonal variation. In any given year, the flight period for a
species may be quite different from the year before or the year
following. Some species appear to be biennial.

Weather conditions are extremely variable from year-to-year,
and from day-to-day. Overnight temperature drops of 60 °F are
not uncommon.

During June and July, the prime collecting months, extended
or continuous daylight occurs depending upon latitude. Thus it
is useless to take along a light trap for moths during this period,
although a trap may be of use in some areas in August.

For personal comfort, protection against biting insects is
essential. Some of the best collecting is in muskegs (bogs).
Entrance into such areas by any warm-blooded body is an open
invitation to a host of mosquitoes and flies. My suggestion for
dress is as follows: Mid-calf rubber boots (for both bogs and
tundra areas), heavy weight but loose Levis, light shirt under
a light weight Nylon windshirt, cap or hat with visor, and an
army-style O.D. headnet. The boots will assure dry feet. The
loose heavy jeans should frustrate the bugs in any attempt to
bite legs. Tuck the jeans inside the boots. For some reason, mos-
quitoes don’t seem to be able to penetrate effectively Nylon
cloth, hence the windshirt. Army-style headnets, available from

^Research Associate, Allyn Museum of Entomology, Sarasota, Florida. Mu-
seum Associate, Los Angeles County Museum of Natural History, Los
Angeles, California.

249

250

C. D. FERRIS

/. Res. Lepid.

surplus outfits, cover head and shoulders and tie under the arm-
pits. This avoids the use of insect repellant. Generally speaking,
I have found repellants nearly useless in arctic regions and in
comparable areas in the Rocky Mountains. Although smelly, the
oldtime Oil of Citronella appears to be more effective than
anything else, with Cutters a close second. The latter, however,
attacks some plastics and one must use it with caution. Light
gloves may be necessary when collecting in deep bogs. Oil of
Citronella rubbed on the hands, especially the backs of the
hands, will generally suffice for protection against the angry
stilettoes.

Clothing should be chosen in anticipation of wet weather
and a wide temperature variation. The mean temperature at
Churchill, Manitoba in the summer is about 50° F. In McKinley
Park, Alaska, I have experienced a temperature drop from nearly
90 °F to 30° F over an eight-hour period at the end of June.

Finally, avail yourself of a “native guide” if one exists. Local
collectors can indicate very quickly the good and bad spots.
Many of the arctic species tend to be quite local in habitat, and
frequent areas that one familiar with more southern latitudes
would bypass. Collectors with experience in the Rocky Moun-
tains (high-altitude species) are perhaps the best prepared to
“smell out” the habitats of the arctic species.

Generally speaking, the arctic mountain passes are at low
elevations ( between 3000-4000' ) with altitude-treeline about
3000' or less. Above certain latitudes, no trees occur, only scrub.
This is true at 58°45'N (sealevel) at Churchill.

Arctic and subarctic collecting can be at the same time, both
very rewarding and very frustrating. The reward is in the variety
of species that can be found; the frustration is in trying to col-
lect them because of weather conditions. In this paper, some
of the prime collecting areas are described along with the prob-
lems associated with visiting these spots.

From my experience, albeit somewhat limited, the best sea-
sons for collecting appear to occur in odd-numbered years fol-
lowing a cool and wet year. In central Alaska, many of the
desirable species of Boloria and Oeneis seem to fly only during
odd-numbered years, although occasional examples may be
taken in even-numbered years. Another feature to recognize is
that the daylight hours are different from more southern lati-
tudes. Twenty-four hour daylight or near daylight is the rule in
June and July. For this reason, the collecting season begins in
mid-June and is virtually over by the end of July in most locali-

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251

ties; a far different situation from the Rocky Mountains, for
example, when prime collecting is from mid-July into August.

Weather conditions are of major importance, and at best are
unstable and unpredictable. Anyone planning an arctic trip
should allow enough time to be in any given collecting area
during the better part of the season. It is not unusual for bad
weather to continue for a week or more. Thus one should have
a month's time, at least, available for arctic trips. Coastal
Alaska, especially the southern coast, the Hudson Bay area
(Churchill), and northeastern Canada (Mt. Albert, Gaspe, etc.)
are subject to stationary low pressure areas during the summer,
which may persist for days or weeks. In the Churchill area,
when the ice breaks up on Hudson Bay at the end of June or
in early July, cold weather (in the 30’s), fog, and drizzle are the
rule. These conditions may persist for three days to a week
with no collecting possible. By contrast, interior Alaska, north
of Fairbanks and south of the Brooks Range can be hot and
clear (temperatures in the 90's) at the same time of year. South
of Fairbanks (McKinley Park, etc.), coastal conditions prevail.
Alaska:

Alaska is accessible by car, boat, and air. For one who has
the time, it is well worth the extra days of travel to drive the
Alaska Highway. Weather permitting, there is good collecting
starting north of Edmonton, Alberta (Hwy. 43). The Alaska
Highway proper begins at Dawson Creek, B.C. The first 88
miles are paved. The section through British Columbia and
the Yukon Territory to the Alaska border is very well maintained
gravel. The last leg of the highway from the border to Fair-
banks is paved. Since the major portion of the road is gravel,
one should take some necessary precautions. These include
extra spare tires, beyond the usual spare, plastic headlight pro-
tectors, a 1/4" hardware cloth or other screen across the front
of the car to deflect stones, chassis undercoating, and a pad over
the fuel tank, for rear-mounted tanks, to prevent stone punctures.
A tool kit and spare parts should be carried, as although there
are gas pumps every 50-100 miles, repair service is hard to find
and expensive. Because of the fine dust, a heavy-duty air cleaner
is strongly recommended. Other than dust and possible flying
stones, when passing or being passed, the only other hazards
are occasional washouts after heavy rains. These are usually
repaired within 24 hours and generally present little problem
for passenger cars, but may cause difficulty for recreational
vehicles and cars with trailers. The main washout danger area

252

C. D. FERRIS

/. Res. Lepid.

appears to be the 300-mile stretch between Dawson Creek and
Ft. Nelson, British Columbia. Extra gasoline should be carried,
as many of the service stations open late in the morning and
close early in the evening, especially on weekends. A VW or
similar car is ideal for the trip, although a bit cramped.

Except for Whitehorse, Yukon Territory, accommodations
along the highway are minimal. There are a number of turnout
areas and several campgrounds, however, and tent or recreation-
al vehicle camping is no problem. One should plan on four
nights camping between Dawson Creek and Fairbanks if any
amount of collecting is to be done along the way. Members of
the Ameriean Automobile Assn, should obtain the booklet avail-
able on the Alaska Highway, as it contains an excellently de-
tailed map.

The best collecting spots that I found are muskegs, willow
bogs, and the emergency air strips. The latter are well-marked
clearings just off the road. They are covered by low vegetation
and usually present a good supply of wildflowers. One must
get out into the centers of the bogs and investigate the little
islands of black spruce, for these will yield Erebia disci and
Oeneis jiitta. Various species of Boloria frequent the willow
bogs.

Figure 1 shows the main road and major collecting areas in
Alaska, in terms of accessibility. Nome, Anatuvuk Pass in the
Brooks Range, and the North Slope are accessible by air via
Wien Airlines. One should check at Fairbanks for flight sched-
ules. One of the major areas is Eagle Summit, which is 109
miles north of Fairbanks on the Steese Highway. As of the
summer 1971, the first 44 miles of the road were paved with the
remainder gravel. The road continues past Eagle Summit to the
Yukon River. Some of the species of interest at Eagle Summit
and 12 Mile Summit just to the south are: Oeneis melissa,
polixenes, bore, Erebia youngi, niagdalena, discoidalis, disa,
fasciata, Boloria polaris, titania, chariclea, youngi, freija, frigga,
napaea, distincta (for the lucky), Papilio machaon, Colias nas-
tes and Parnassiiis eversmanni. The summit areas are above-
treeline tundra. Both north and south of them are below-treeline
muskeg areas which should be investigated for Plebejus optilete,
Colias palaeno, Hesperia manitoba, Boloria, Erebia, and Oeneis.

Generally speaking, the major habitats, and this is true of the
aretic regions in general, are black spruce muskegs and willow
bogs in the Taiga Zone, and the open Tundra Zone. The boreal
forests are usually too dense to permit colleeting and appear

13(4):249~264, 1974

ARCTIC AND SUBARCTIC

253

Fig. l.—Map of Alaska and Yukon Territory, Highways are shown as the
thicker solid lines.

254

C. D. FERRIS

/. Res. Lepid.

to harbor very few species.

South of Fairbanks, the Richardson highway affords good
collecting in spots. At Paxson Jet. (mile 185.5), the Denali
Highway leads west to McKinley Park, 161 miles away.
Although it was being “improved” in 1971, the Denali was an
accumulation of chuckholes loosely connected by gravel. I
would not recommend travel on it except in a light, but tightly-
sprung vehicle. It is by far the worst major road that I have
ever driven. Along the highway, one finds Erebia rossii, Boloria
eunomia denali, and a variety of other species. The Denali
Highway in McKinley Park is well-maintained and collecting
is excellent, especially in the vicinity of the several Pass areas.
Collecting, however, is by permit only from the National Park
Service. Generally these permits are available only to university
or museum personnel engaged in specific studies. Thus unless
one wishes to tour the Park, which is quite beautiful and un-
cluttered, it is not a recommended trip because of the highway
and the restricted collecting.

Toward Valdez and Anchorage, some interesting species can
be obtained, but collecting is difficult because of adverse weather
conditions. Snow at higher elevations (above 2500') is not
unusual in June and July.

I have not found collecting at Haines very productive, al-
though the dark coastal race of Pieris napi occurs there as well
as Anthocaris sara alaskensis. I have collected both species there
in early July. The Haines Highway will be covered in the
Yukon Territory discussion.

Other areas of interest include spots along the Taylor High-
way from the junction with the Alaska Highway to Eagle,
Alaska. Charter flights can be arranged into remote locations,
although bad weather can sometimes maroon the luckless
collector in remote spots for extended periods of time.

Yukon Territory, Northern British Columbia,

Alberta, and Northwest Territories:

The most easily accessible collecting in northern Alberta,
British Columbia, and the southern Yukon Territory is via the
Alaska Highway and Highways 43, 34, and 2 from Edmonton
to Dawson Creek, via Whitecourt, Alberta. An alternative route
to Alaska is from Prince George to Prince Rupert, British Colum-
bia. There are a number of spots along Highway 16 that will
yield interesting species. At Prince Rupert, one enters the
“Alaska Marine Highway.’ This is the ferry system which oper-

13(4). ■249-264, 1974

ARCTIC AND SUBARCTIC

255

ates between Prince Rupert, B.C. and Skagway, Alaska, with
stops at Ketchikan, Wrangell, Petersburg, Juneau, and Haines.
The trip is quite scenic and takes about 36 hours. It is fairly
expensive. The cost from Prince Rupert to Haines for two
people and a small car is in excess of $200, excluding meals.
Ferry transport is also available from Seattle and Vancouver
to Anchorage and other ports, at additional expense.

At Haines, Alaska, one enters the Haines Highway, which is
approximately 160 miles long. The initial portion in the Alaska
panhandle is paved. The remainder through a small corner of
B.C. and the Yukon Territory is well-maintained gravel. The
road joins the Alaska Highway at Haines Junction, Y.T. Enter-
ing the main portion of Alaska by this route eliminates over
900 miles of gravel-road driving on the Alaska Highway. The
best collecting appears to be in the area between Kathleen
Lake, Y.T. (mile 142) and Haines Jet., Y.T. (mile 159). Such
species as Oeneis hore, Euphydryas anicia helvia, Erebia epip-
sodea remingtoni, C. tullia ssp., Colias philodice vitabunda, and
Parnassius phoebus ssp. can be taken. Chilkat Pass, B.C. at
mile 65.3, although appearing to be good habitat, I found com-
pletely devoid of butterflies.

Additional areas in the Yukon Territory are served by the
Klondike Highway, which leaves the Alaska Highway just west
of Whitehorse, Y.T. and meanders northwest to Dawson, and
then west to Chicken where it joins the Taylor Highway to
Eagle, Alaska. A spur goes to Mayo Lake and Keno Hill. Other
areas can be reached by Charter flight. Essentially all areas in
the Northwest Territories must be reached by air. The Mac-
kenzie Highway from Peace River, Alberta does serve a few
areas in the southern portion of the N.W.T. including Dawson
Landing and Fort Providence on Great Slave Lake, Fort Simp-
son on the Mackenzie River (by a spur route), and Yellowknife,
also on Great Slave Lake (north shore).

Two primary collecting areas are the Richardson Moun-
tains in the western section of the N.W.T. and Baker Lake in
the eastern section. A highway has been proposed that will
skirt the Richardson Mts. allowing access to that area by car.
Baker Lake, reached by air, is a choice spot in that Colias
boothi is taken there, as well as a number of other species.

The Pas-Chur chill, Manitoba:

In addition to his own experiences, the author has drawn on
the papers by the Chermocks (1968) and Masters (1971) in

256

C. D. FERRIS

/. Res. Lepid.

preparing this section. Acknowledgment is made here to these
sources without subsequent mention.

The region between The Pas and Churchill is of interest for
a variety of reasons. It is easily accessible. There are a number
of subarctic and arctic species. There are good living accom-
modations.

The Pas is served by highway 10 which continues north
another 100 miles to Flin Flon. Where land has not been cleared
for agricultural purposes, it is a black spruce and muskeg area,
much like northern Maine, dotted with lakes. Some of the
indiginous species are Colias gigantea, Boloria eunomia, Papilio
machaon, Erebia species, Oeneis jutta and macouni, Coeno-
nympha tullicg etc., as well as species associated with more
southern latitudes. It is a community of some 6000 with stores,
motels, and a small museum. It is also the point of embarkation
by rail to Churchill on the west coast of Hudson Bay. No roads
go to Churchill, so one has to take the train from The Pas, or
the plane from Winnipeg. In 1973, the round-trip fare, per
person, was about $134.00 by air from Winnipeg and about
$37.00 by rail from The Pas (coach).

If one possesses an adventurous spirit, then by all means
take the train. Those of a more sedate mien are well advised
to fly. The rail distance from The Pas to Churchill is 510 miles
and requires about 22 hours in transit, as the train makes ex-
tended stops to unload freight and change erews at Thicket
Portage, Thompson, and Gillam. At these stops, the train is
literally invaded by the local Indians, to whom a rail coach is
tantamount to a movie matinee. This is not conducive to sleep-
ing, as the Gillam stop occurs just after midnight. The train
schedule is typically: leave The Pas 10:30 AM Mon., Wed.,
Fri.; leave Churchill 5:30 PM, Tries., Thurs., Sat. In either
direction, the Gillam stop is at midnight.

It is also possible to drive from The Pas to Thompson and
fly from there to Churchill, or board the train. This saves about
175 miles of train travel, but increases driving time. During
the peak of the summer months, there are some flights, but
not on a daily basis, from The Pas to Churehill.

Churchill is actually a dual community of Churchill and Ft.
Churchill. The two are about 3 miles apart and the overall
population is just over 3000. There are several “motels” and
hotels as well as stores in Churchill proper. My wife and I
stayed at the Whalers Co-op “motel” which has comfortable
housekeeping units and makes very reasonable weekly rental

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ARCTIC AND SUBARCTIC

257

rates. It is located between the railroad and the Churchill
River about 100 yards from the tundra. Thus right on the edge
of prime collecting territory, which is between the railroad and
the river from town to about 4 miles south at milepost 505 on
the railroad.

Tent camping on the tundra is possible, and the Chermocks
did so. They wrote an amusing account of some of their ex-
periences. Because of the weather, which can turn miserable
for days on end, and the mosquitoes and flies, this is not recom-
mended. After spending a hot day of collecting and fighting
the biting insects, it is a welcome relief to come back to a hot
shower and a comfortable bed. During sieges of bad weather,
temperatures hovering just above freezing are common. Inci-
dentally to preserve one’s sanity, a headnet is essential at
Churchill, particularly after the sandfly hatch. Figure 2 illus-
trates some of the “friendly” creatures that inhabit the area and
lust for the blood of visiting lepidopterists.

A rough map of the area is shown in Figure 3. The Sub-
climax Tundra area affords the best collecting for Erehia rossii
inornata Leussler, Boloria polaris (Bdv. ), and the endemic
Oeneis. O. jutta is in the little stands of black spruce and
tamarack. The remaining species are in open areas. The Taiga
Zone south of milepost 507 on the railroad offers the best col-
lecting for most of the remaining endemic species, although
there is some exchange of species between the Taiga and Tundra
Zones. When they first appear on the wing, Colias nastes and
hecla can be found a mile or more into the Taiga, along the
railroad right-of-way (cf. Masters, 1971, p. 8).

Although the Climax Tundra to the east and south of town,
and north to Cape Merry is profusely covered with wildflowers,
I found it for the most part, devoid of butterflies, at least until
mid-July when we departed Churchill. Scenically it is a strik-
ing area of granite boulders, flowers, mosses, and lichens. An
excellent view of the ice on Hudson Bay as well as Churchill
Bay is afforded from this area. It is well worth a visit just for
the panorama it provides. One small area just at the edge of
town yielded Erehia theano canadensis Warren, Colias nastes
moina Strecker, and Plebejtis aquilo lacustris Freeman.

To cover the various habitats completely, about 8-10 miles
of walking per day is necessary. This may be reduced, however,
by using the local bus or taxis. In the early morning, I frequent-
ly took the bus to Dene Village and began collecting along the
Ft. Churchill railroad spur to where it joins the main line. From

258

C. D. FERRIS

/. Res. Lepid.

Fig. 2.— Photograph of some of the “friendly” creatures which add to the
“joy” of arctic collecting. These are from Churchill, Manitoba. The head-to-
tail length of the central fly is 2 cm.

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259

Fig. 3.— Map of Churchill, Manitoba area. W indicates the Beluga Sport
Whaling Motel. A and D are respecively Akudlik and Dene villages, n in-
dicates the old naval building. The numbers are mileposts along the rail-
road. The fine dotted lines represent the separation between the Tundra
and Taiga Zones.

260

C. D. FERRIS

/. Res. Lepid.

there, one can walk south along the tracks to mile 505 and then
retrace steps and continue north across the tundra into town.
The North Star Bus Lines, Ltd. provides regular service between
Churchill and Ft. Churchill, with stops at Akudlik and Dene
Villages.

Rental cars are available, but this is a waste of money. I
believe in the philosophy ascribed to A. B. Klots, that is, if you
are riding in a car, you can’t be in the field collecting.

On warm mornings when the temperature is about 50 °F,
collecting begins at 9:00 AM; on cooler days, the bugs don’t
get out until about 10:00 AM. By 3:00 PM on cool days and
4:00 PM on warm days, collecting is over. This is quite different
from interior Alaska, where I have collected from 7:00 AM
until 9:00 PM. At Churchill, as long as there is full sun and
the temperature is above freezing, butterflies will fly. I have
collected in 35 °F temperatures in mid-July. A warm day is
when the temperature reaches 55-60° F.

Because of its interest to the collector and the accessibility
of Churchill, a checklist is now provided including overall
dates of flight. Seasonal variation is extreme so no attempt has
been made to list peaking of flights. Masters ( 1971 ) made an
honest attempt to do this, but when I collected in 1973, the
species peaked much earlier than he indicated. Some years,
however, are quite retarded. In 1973, the ice broke up in the
vicinity of Churchill much earlier than normal and the Beluga
whales returned three weeks ahead of schedule. In 1972-73
winter was exceptionally mild, after a cold and wet summer
in 1972.

CHECKLIST — Churchill, Manitoba Butterflies

30-vi — 3-vii L

( Biennial on even years? )

17-vi — ^14~vii E
25-vi — 9-viii M

Hesperia manitoba borealis Lindsey

( Biee

Pyrgus centaureae freija ( Warren )
Colias g. gigantea Strecker
Colias hecla hela Strecker (rare)
Colias nastes moina Strecker
Colias palaeno chippewa Edwards
Pier is napi ssp.

Glaucopsyche lygdamus ssp.

Ltjcaeides argyrognomon scudderii
( Edwards )

Plebejus aquilo lacustris Freeman
Plebejus optilete yukona ( Holland )

ll~15-vii L ( Type Locality )

26-vi — 9-viii L ( Type Locality )

24-vi — 9-viii M

16-vi — 15-vii E

Several reports,
no dates E

no dates L

15-vii-73 L

9-14-vii-73 L

13(4):249-264, 1974 . ARCTIC AND SUBARCTIC

261

Boloria eunomia triclaris ( Hubner )

23-vi— 9-viii

M

Boloria f. freija ( Thunberg )

10-vi— 15-vii

E

Boloria frigga saga ( Staudinger)

17-vi — 1-viii

E

( Biennial on

Boloria polaris ( Boisduval )

20-vi — 28-vii

E

Boloria titania boisduvalii
( Duponchel )

1-vii— 9-viii

L

odd years )

Erebia disa mancinus Doubleday

9-vii--73

M

( 1 pair ~ only

Erebia d. discoidalis (Kirby)

15-vi — 10-vii

E

record )

Erebia rossii ornata Leussler

10-vi — ^16-vii

E

(Type Locality)

Erebia theano canadensis Warren

2-23-vii

L

(Type Locality)

Oeneis borejtaygete ssp. ( ?? )

15-vi — 16-vii

E

( See Text)

Oeneis jutta ssp.

20-vi — 15-vii

E

Oeneis melissa semplei Holland

12-vi — 2-viii

E

Oeneis p. polixenes (Fabricius)

12-vi — 2-viii

E

Casuals and Strays

Papilio glaucus canadensis Roth. & Jordan (1) G. S. Brooks collector^.
1940.

Colias eurytheme Boisduval G. S. Brooks collector, 1942. Ferris saw what
appeared to be a female of this species in the Taiga Zone, July, 1973.
Pieris protodice ssp. Bdv. & LeC. (1) G. S. Brooks collector^, 1942.

Pieris rapae (L. ) (1) J. H. Masters collector, 3-vii“70.

Boloria selene ssp. ( 1 ) A. B. Klots, in litt.

Cynthia cardui (L.) 30-vi — ^15-vii"73 C. D. Ferris collector (common).
( Migrant )

Nymphalis antiopa ( L. ) ( Stray )

Nymphalis m. milberti (Godart) (Stray)

Lycaena species ( probably dorcas ssp. ) ( 1 ) 3"Vii-52 in R. D. Bird field
notes, see Masters (1971).

E rr species which appear early in flight season; M = species which appear
in middle of flight season; L species which appear late in flight season.
Actual appearance dates vary from season-to-season. ( 1 ) = single speci-
men record.

^These .specimens have apparently been lost or destroyed.

Boloria chariclea ssp. has been recorded from Churchill, but
probably has been confused with titania. At one time, the two
taxa were considered conspecific by some authorities, and I
believe that this has caused some of the nomenclatorial con-
fusion in the Churchill records. Out of a lot of well over 100
specimens of titania which I took in 1973, four could be as-
signed to chariclea based upon facies. I choose to consider these
as pale titania. A similar confusion zone between titania and
chariclea exists in several parts of Alaska.

Masters (1971) asserts that two subspecies of Oeneis jutta
occur at Churchill. O. j. alaskensis Holland, he claims flies on
even numbered years, and O. j. ridingiana Chermock & Cher-
mock flies on odd numbered years. This situation needs more
study, as there is an unpublished Chermock name relating to
the Churchill population of jutta.

262

C. D. FERRIS

/. Res. Lepid.

A strange situation exists with Oeneis horeltaygete. This has
been reported by various collectors as occurring at Churchill.
I took a small series of what I thought was this insect. The
veins on the undersides of the hindwings are clearly outlined
in gray as in taygete. The ground color of the wings dorsally is
paler than normal polixenes. Genitalic examination has proved
these specimens to be polixenes. I have received specimens
from other collectors sent as taygete, which on genitalic exami-
nation are polixenes. It appears then that two phenotypes of
polixenes fly at Churchill. I have yet to see taygete from
Churchill, although it may occur there.

My collecting included two species not recorded at Chur-
chill previously: Erebia clisa mancinus and Plebejtis optilete
lyukona. A pair of the Erebia was taken at about 1:15 PM
(CDT) in full sun in a clearing (cf. Masters, 1972, p. 4) at
mile 506.5 on the railroad right of way on 9-vii-73. P. optilete
yukona was found throughout the Taiga Zone in moist areas.
It and Pyrgiis centaureae freija are probably quite common and
widely distributed, but overlooked because they fly close to the
ground and blend in color so well with the mossy ground cover.
Both are erratic fliers.

I suspect that nearly all of the endemic species recorded
from Churchill, except perhaps for C. hecla, are common in
suitable habitat at appropriate times during the season. My
pair of E. disa may have been introduced via the railroad from
one of the colonies at Gillam or elsewhere. One resident species
which now appears uncommon is Erebia theano canadensis. The
Chermocks found it abundant in the early sixties, but J. A.
Ebner s collector in Churchill failed to find it during three suc-
cessive seasons in the mid-sixties, (in litt.). Masters found it
in 1969-70. In 1973, I did not find it in any of the locales from
which it had been reported in the past. I did find several
colonies at other locations, but it is a very local species and is
tied to a “micro-habitat.” One of Chermocks collecting sites
has been destroyed by an Eskimo village housing development.
Other areas have been ditched for drainage purposes and this
has altered habitat. E. theano is probably common in remote
areas that collectors have not yet penetrated.

The Churchill area is undergoing considerable development.
The Subclimax Tundra Zone especially is being reduced by
housing and radio antenna installations. The area between the
railroad and the Churchill River is still virgin except for the

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263

extreme northern end where the Whaling Centre is located.
Most of the town growth appears toward the east and south
toward the high Climax Tundra Zone. Evident in 1973, was the
construction of a new hospital and a government-financed hous-
ing development. In spite of the encroachments of ‘progress,”
Churchill remains a fascinating place to visit.

Mt. Albert, Gaspe Peninsula, Quebec:

Mt. Albert's attraction lies in several species which are re-
corded from the tundra tableland on its top. The tableland is
oblong, quite extensive, and virtually without distinguishing
features except for a boulder pile at the center. It is very easy
to lose one’s sense of orientation. Oeneis taygete gaspeensis
dos Passos, Oeneis poUxenes ssp., Papilio brevicauda gaspeensis
McDunnough, Hesperia manitoba borealis Lindsey, and Pyrgus
centaureae freija (Warren) are recorded as well as a record in
the 1940’s for Oeneis chryxus ssp. A number of other species
are found at the base of the mountain along the Ste. Anne River.

Mt. Albert is reached by a cut-off road from Highway 6 just
northwest of New Richmond, Quebec. A resort hotel is located
along the road across the river from the mountain. In contrast
to Mt. Katahdin in Maine, as yet there is no restriction to
collecting.

Getting to the top of the mountain used to be somewhat of
a problem, as legally, one was supposed to have a Canadian
guide. This situation has changed recently and access to the
mountain is relatively easy. Trail maps can be obtained by
writing: Direction Generale des Parcs, Ministere du Tourisme,
de la Chasse et de la Peche, Edifice G, Cite parlementaire,
Quebec, Que., Canada, GIA IRS and requesting the brochure
“Senders de Randonnee Pedestre, Parc de la Gaspesie.” The
brochure is published in French. There is a campground close
to the trailhead, or one may stay at the resort hotel. There are
several routes to the top now. The “Plaque Malade” trail is an
easy hike through the trees to the top, but is longer than the
Mt. Albert Trail (“Sentier du Mont Albert”).

The optimum collecting time is quite variable. L. Paul Grey
(Enfield, Maine) has made a study and feels that the best time
is probably between July 6 and July 17 (in lift.). Oeneis taygete
appear to fly for a very short period each year. On the other
hand, John Johnstone (Willowdale, Ont. ), (in lift.) has stated
that there are two flights of taygete; one in the latter part of
June, and another in July. Clear weather is essential for col-

264

C. D. FERRIS

J. Res. Lepid.

lecting on the top of Mt. Albert. Making the trip on an over-
cast day will prove fruitless, as the bugs just don’t fly.

Another region of interest is Labrador-Newfoundland. The
author has not collected in this area, and has been informed
that, at present, access into the most desirable areas is restricted
because of mining interests. Richard Holland (1969) has de-
scribed some collecting in this locality and mentioned the
concomittant problems.

ACKNOWLEDGMENTS

The author wishes to express his appreciation to various in-
dividuals who provided him with general information on arctic
collecting, and specific information on collecting areas prior to
trips to northern locales. Included are James A. Ebner (Okau-
chee. Wise.) and Dr. Kenelm W. Philip (Fairbanks, Alaska).
The author, his wife, and Paul Grey spent a delightful and
amusing several days at Mt. Albert in 1968, and mulled over the
quixotic behavior of O. taygete.

Jim Ebner and Ronald L. Huber (St. Paul, Minn.) were
especially helpful in providing data from Frank Ghermock’s
notes on collecting at Churchill, Manitoba. Warren Kiel ( Grove-
ton, N.H.) kindly provided current information on the access
routes to the top of Mt. Albert.

LITERATURE CITED

CHERMOCK, P. M. and F. H. 1967. Churchill. Bull. Assn. Minn. Entomol.
(field collecting section) 2(2):33-39.

HOLLAND, R. 1969. Notes on Newfoundland butterflies. /. Lepid. Soc.
23(l):33-42.

MASTERS, J. H. 1971. The butterflies of Churchill, Manitoba. The Mid-
Cont. Lepid. Ser. 2(25): 1-13.

1972. Ecological and distributional notes on butterflies of the genus

Erebia in Manitoba. The Mid-Cont. Lepid. Ser. 4 (52). -1-10.

Journal of Research on the Lepidoptera

13(4):265-266, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

KLOET & HINCKS’ CHECK LIST OF
BRITISH INSECTS (LEPIDOPTERA) EDN. 2.

A REPLY TO CRITICISMS.

J. D. BRADLEY, D. S. FLETCHER and P. E. S. WHALLEY

British Museum of Natural History,

Dept, of Entomology, London

Our attention has been drawn by Dr. J. G. Franclemont
of Cornell University to a paper by Juraj Paclt ( 1974, Jl Re-
search Lepid. 12(4) :211-212) proposing corrections to some
of the family-group names used in the Lepidoptera part of
Kloet & Hincks' Check List of British Insects (Edn. 2, 1972).
Since we are jointly responsible for the preparation of this
revised, second edition, a reply from us seems appropriate.

The correction of Evergestiinae to Evergestinae is acknowl-
edged, but the other proposals put forward are contrary to the
provisions of the current International Code of Zoological
Nomenclature (Edn. 2, 1964) and its amendments, 1974, Bull,
zool. Nom. 31: 77-89.

Family No. 50, Endromidae, derives from the suprageneric
term Endromidi proposed by Boisduval, 1828, Eur. Lepid. Index
meth. ( 1 ) : 50, which is based on the validly included genus-
group name Endromis Ochsenheimer, 1810. Article 29(d) of
the Code states: “Incorrectly formed stem. — A family-group
name proposed before 1961 based upon an incorrectly formed
stem is not to be amended for that reason if it is in general
current use. Incorrectly formed names published after 1960 are
to be corrected wherever they are found”. The correct spelling
of the name applied to family No. 50 therefore is Endromidae,
as published in the Check List.

For the same reason the spelling of the family-group names
Syntominae proposed by Herrich-Schaffer [1846] 1845, Syst.
Bearbeitung Schmett. Eur. 2: 49, Cerostominae, proposed by
Borner, 1925, in Brohmer, Fauna Dtl. (Edn. 3): 375, Enico-

265

266

J. D. BRADLEY ET AL

/. Res. Lepid.

stominae, proposed by Heslop, 1938, New bilingual Cat. Br.
Lepid.: 82 and Cemiostominae, proposed by Spuler, 1898, Sber.
phys.-med. Soz. Erlangen 30: 33 are correct as published in the
Check List and are not to be amended.

Pack's proposals to alter the synonymy of a number of the
family-group names are equally ill-founded, as reference to
Article 40 of the Code clearly demonstrates.

‘'Synonymy of the type-genus. — When, after 1960, a nominal
type-genus is rejected as a junior synonym (objective or sub-
jective), a family-group name based on it is not to be changed,
but continues to be the valid name of the family-group taxon
that contains both the senior and junior synonyms.

(a) Conservation of certain names. — If a family-group
name, changed before 1961 because of such synonymy, has won
general acceptance, it is to be maintained in the interests of
stability”.

In family No. 11 for example, Limacodidae is the oldest
suprageneric term, based on a validly included genus-group
name; it was proposed by Duponchel [1845] 1844, Cat. meth.
Lepid. Eur.: 84. It will be seen from Article 40 of the Code that
the validity of the family-group Limacodidae is in no way affec-
ted by the synonymy of its type-genus Limacodes Berthold,
1827 with Apoda Haworth, 1809.

Even if the family-group name were still to be based on the
oldest included genus-group name, as Pack appears to suggest,
then his proposed synonymy of Limacodidae with Apodidae
would still be in error. Heterogenea Knoch, 1783, not Apoda
Haworth, 1809 is the oldest included genus-group name and
this was used by Hampson in 1918 as the basis of the family-
group name Heterogeneidae.

Journal of Research on the Lepidoptera

13(4):267-270, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

THE NOMENCLATURE IN AN
IMPORTANT BRITISH CHECK LIST (1972)

PART 3: CORRECT GENDER FOR GENERIC
NAMES DERIVED FROM CLASSICAL WITHOUT
CHANGE OF TERMINATION.

JURAJ PACLT

Inst, of Exptl Phytopathol. and Entomology,

Slovak Academy of Sciences
Ivanka pri Dunaji, Czechoslovakia

The requirements of the International Code of Zoologi-
cal Nomenclature concerning the agreement of species-group
names in gender seem not to be strange to the authors of the
new catalogue: The endings of the valid specific names, where
adjectival, have been changed in accordance with Article 30 of
the Code of Zoological Nomenclature' (J. D. Bradley, D. S.
Fletcher and P. E. S. Whalley in their Introduction, p. vii).

Let us tell that the problem of correct application of end-
ings in species-group epithets affects not only combinations in
which the gender of the generic name is either somewhat doubt-
ful or at least hard to determine. It will affect, as seen immedi-
ately below, even many of those combinations in which the
generic name is derived from Greek (or Latin) without change
of termination.

Nematopogon (masc. ) with N. swammerdamellus (not
‘swammerdamella), N. panzerellus (not -ella), N. pilellus (not
-ella), and N. metaxellus (not -ella) included.

Diplodoma (neuter) herminatum (not lierminata).

Nemapogon (masc.) with N. granellus (not ‘granelld), N.
cloacellus (not -ella), N. albipunctellus (not -ella), N. person-
ellus (not -ella), iV. ruricolellus (not -ella), N. arcellus (not
-ella), and N. picarellus (not -ella) included.

Archinemapogon (masc.) laterellus (not ‘laterella).

Acrocercops, a latinized Greek noun of masculine gender,
should, according to the Art. 30, be treated as such (provided
that the author of that genus did not decide to use his name
as a noun of feminine gender).

267

268

JURAJ PACLT

/. Res. Lepid.

PhyllQnorycter (masc. ) with about 50 species listed the
majority of which (42) are treated as if Phyllonorycter were of
feminine ( !') gender.

Yponomeuta (masc.: cf. toreuta, a graver) with Y. evony-
mellus (not ‘evonymelld) , Y. padellus (not -ella), Y. malinellus,
Y. cagnagellus (not ‘cagnagelld) , Y. rorrellus (not -ella), Y.
irrorellus (not -ella), Y. plumbellus (not -ella), and Y. sedellum
(not ‘vigintipunctatd) included.

Euhyponomeuta (masc.) stannellus (not ^stannelld).

Ocnerostoma (neuter) piniariellum (not 'piniarielld ) .

Prays (masc.) fraxinellus (not Jraxinelld).

Augasma (neuter) aeratellum (not ‘aeratelld) . This genus is
a member of the family Coleophoridae to which Eupistidae and
‘Atigasmidae were referred to as synonyms. The latter must,
however, be corrected to Augasmatidae.

Goniodoma (neuter) Imioniellum (not dimonielld).

Enicostoma (neuter) obellum (not 'obelld).

Levipalpiis (masc.) hepatariellus (not Eepatarielld ) .

Gnorimoschema (neuter) streliciellum (not ‘strelicielld).

Crocidosema (neuter) plebejanum (not ‘plebejand) .

Olethreutes (masc.) with 14 species listed all of which are
treated as if Olethreutes were of feminine ( ! ) gender.

Archips (masc.) with 12 species listed all of which are
treated again as if Archips were of feminine ( ! ) gender. Note
the correct usage of A. podaniis (not 'podand) in recent litera-
ture.

Clepsis (fern.) melaleucana (not 'melaleucanus') . The re-
maining species listed bear names agreeing in gender.

Ptycholoma (neuter) lecheanum (not decheand).

Aleimma (neuter) loeflingianum (not doeflingiand ) .

Chile (masc.) phragmitellus (not "phragmitelld).

Parapoynx (masc.) with P. stratiotatus (not (‘stratiotatd ) ,
P. obscuralis, and P. stagnutus (not ^stagnatd) included.

Pyrausta (masc.) with P. auratiis (not ‘auratd), P. nigratus
(not ‘nigratd), and P. cingulattis (not ‘cingulatd) included.

Herpeto gramma (neuter) centrostrigale (not ^ centrostrigalis) .

Myelois (myeloeis, masc.) cribrellus (not ‘cribrella) and
M. cirrigerellus (not -ella).

Homoeosoma (neuter) with H. nebulellum (not ‘nebulelld),
H. sinuellum (not -ella), and H. nimbellum (not -ella) included.

Apomyelois (apomyeloeis; masc.) bistriatellus (not ‘bistri-
atelld ) .

13(4):267-270, 1974

NOMENCLATURE, PT. 3

269

Ochlodes (gender prescribed: masculine; Art. 30: ‘Names
ending in . . . -odes . . . are . . .’) with O. venatus (not "venatd)
as the only species listed.

Colias (fern.) crocea (not ^croceus). For details on the
gender see Paclt (1951).

Malacosoma (neuter) castrense (not 'castremis).

Phyllodesma (neuter) ilicifolium (not Hlicifolid) .

Cilix (masc. ) glaucatus (not 'glaucatd).

Camptogramma (neuter) with C. bilineatum (not 'bilineatd)
and four subspecies (bilineatum, atlanticum, hibernicum, isola-
turn) included.

Perizoma (neuter) with P. alchemillatum (not 'alchemillatd),
P. bifasciatum (not -ata), P. minoratum (not -ata), P. blandi-
atiim (not -ata), P. albulatum (not -ata), P. flavofasciatum (not
-ata), P. didymatum (not -ata), P. sagittatum (not -ata); and
including also some distinct subspecies. The remaining two
species listed ( P. taeniatum, P. affinitatum ) bear names agreeing
in gender.

Anticollix (masc.) sparsatus (not ‘sparsatd).

Abraxas (masc.) with A. grossulariatus (not ^ grossulariatd ) ,
A. sylvatus (not -ata), and A. pantarius (not -ata) included.

Ennomos, a Greek noun of masculine or feminine gender,
should, according to the Art. 30, be treated as masculine (pro-
vided that the author of that genus did not decide to use his
name as a noun of feminine gender). Note the prevailing usage
of E. subsignarius (not ‘subsignarid) in recent literature.

Apocheima (neuter) hispidarium (not ‘hispidarid) and A.
pilosarium (not -ria).

Biston (masc.) stratarius (not 'stratarid) and B. betularius
(not -ria).

Selidosema (neuter) brunnearium (not 'brunnearid ) , ssp.
scandinaviarium (not -ria) and ssp. tyronense (not ‘tyronensis) .

Aids (masc.) with A. repandatus (not 'repandatd), three
subspecies (repandatus, murarius, sodorensium^) , and A. jubatus
(not 'jubatd) included.

Pterostoma (neuter) palpinum (not "palpind),

Eilema (neuter) with £. sororculum (not ‘sororculd), E.
griseolum (not ‘griseold), E. caniolum (not -ola), E. pygmae-
olum (not -ola), E. complanum (not ^compland), E. sericeum
(not ‘sericed), E. deplanum (not ‘depland), and E. lurideolum
(not lurideold) included.

270

JURAJ PACLT

/. Res. Lepid.

Eugnorisma (neuter) depunctum (not 'depunctd),

Eurois (euroeis; masc. ) occidtiis (not 'occulta).

Eublemma (neuter) with E. ostrinum (not 'ostrind), E. par-
cum (not ‘pared), and E. noctuale (not ‘noctualis) included.

Pseudoips (masc.) prasinanus (not Jagand). The authors
disregard apparently the revision of the Linnean type (Lempke,
1947), according to which Phalaena Tortrix prasinana Linnaeus
is the Scarce Silver Lines (Pseudoips). On the contrary, Pyralis
fagana Fabricius corresponds to the Green Silver Lines (Bena).

Polypogon (masc.) strigilatus (not ‘strigilatd) . The remain-
ing species listed bear names agreeing in gender.

ERRATA

On page 211 of Part 1 of the present paper (/. Res. Lepid.
12: 211-212), the sentence beginning with ‘Although' (lines 17
to 22) is to be corrected to the following effect:

Although both subfamilies are called actually Plutellinae and
Depressariinae respectively, and although the genus Cerostoma
Latreille, 1802 has been replaced as junior synonym by Ypsolo-
pha Latreille, 1796, the above family-group names should be
referred to as Cerostomatinae and Enicostomatinae.

^ Gen. pi. ex vocis (Insulae) Sodor-enses quae ad Insulas Ebudes vel
Ebudas (= the Hebrides) pertinent.

Journal of Research on the Lepidoptera

13(4):271-277, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

ARTIFICIAL DIET:

THE KEY TO THE MASS REARING OF
MEGATHYMUS LARVAE

RONALD S. WIELGUS1.2

3434 West Augusta Ave., Phoenix, Arizona 85021

Techniques for rearing Megathymus larvae utilizing yucca
caudices were discussed by Wielgus and Wielgus (1973). As a
medium for rearing^ yucca caudices proved to be dijBBcult to
preserve in a wholesome state for prolonged periods, were ob-
tainable only with great labor and generally not available to
others, and were cumbersome and space-consuming.

Petterson and Wielgus (1974) discussed a method of intro-
duction and acceptance of an artificial medium by Megathymus
streckeri (Skinner) larvae. The biology of streckeri larvae made
rearing on yucca caudices extremely difficult (and hazardous
to the larvae during the transfer process) so a more suitable
substitute was devised in the form of artificial diet and man-
made containers. Simultaneous with rearing streckeri larvae,
the artificial diet substitute was used successfully with the larvae
of other Megathymus species.

In 1974, 52 streckeri and 30 Megathymus texanus texanus
B. &. McD. ova were obtained and the newly-hatched larvae
successfully started and established on artificial diet. Later that
same year, several Megathymus ursus ursus Poling ova were
collected and the newly-hatched larvae were also started and
established on artificial diet with no difficulty.

Megathymus species split into two natural groups based
mainly upon larval biologies, but also upon distinct pupal dif-
ferences. The first group (“tent-builders”) includes all of those
species whose larvae construct and maintain tents throughout
the feeding period. The other group (“non-tent-builders”) are
all of those species whose larvae construct tents after the feed-

^Research Associate, Allyn Museum of Entomology.

^Museum Associate in Entomology, Natural History Museum of Los Angeles
County.

271

272

RONALD S. WIELGUS

/. Res. Lepid.

■>* i

Fig. l.-~Plastic ice cream container with four Megathymus ursus ursus
larvae reared on artificial diet in PVC pipe sections. Scale in inches.

Fig. 2,— Closeup view of rolled paper tents shown in Fig. 1. from wmcn
ursus larvae have constructed silken tents. Dark particles adhering to tents
are frass pellets. Scale in inches.

13(4):27 1-277, 1974

ARTIFICIAL DIET

273

ing period and just prior to pupation. The latter group includes
Megathymus cofaqui (Strecker), Megathymus harrisi Freeman,
Megathymus texanus texanus B. & McD., Megathymus texanus
leussleri Holland and Megathymus streckeri (Skinner).

The larvae of both groups present unique problems to rear-
ing on artificial diet based mainly upon behavioral patterns.
The basic problems, however, are 1 ) diet acceptance and estab-
lishment, 2) containment of the diet and larvae, 3) confinement
of the larvae, and 4) replenishment of the diet and/or transfer
of the larvae to fresh diet. The techniques developed and dis-
cussed herein have proven quite adequate for rearing scores of
Megathymus larvae under completely artificial conditions.
Larvae just newly-hatched, or field-collected in various instars,
have been established on diet and successfully reared to adults
in the laboratory utilizing these techniques.

Introduction and acceptance of artificial diet by Megathymus
larvae is easily accomplished by the insertion of a small plug of
yucca pulp into the diet. The yucca used is the native foodplant
species of each Megathymus taxon being reared. One small
(40 mm in diameter by 10 cm in length) yucca caudex will
produce hundreds of plugs, and the unused portion of caudex
may be stored in a household refrigerator for several months if
properly sealed in plastic wrapping (freezer wrap).

TENT-BUILDERS

Containment of the diet differs for both larval groups. For
the tent-builders, the author uses sections of Polyvinylchloride
(PVC) irrigation pipe 24 mm in inside diameter by 80 mm in
length which have been filled with diet. One end of the pipe is
capped with a square of sheet aluminum 27 mm on a side by
0.5 mm in thickness which is taped to the pipe. The other end
of the pipe is capped with a square of clear plastic sheet of
approximately the same dimensions. In the center of each plastic
square, a small hole is cut for the insertion of an artificial tent
of rolled paper. The paper tent is pushed into the diet so that
it surrounds the yucca plug inserted earlier. Then an ovum, still
attached to a portion of the yucca leaf, is placed into the paper
tent to await hatching, or, a larva in any instar is allowed to
make its way into the tent and establish itself. Containers of
this type, properly labeled, are placed upright in various small
boxes, plastic bowls, etc., and maintained at room temperature.
Larval transfers are easily accomplished and later, as the larvae
mature, only a wrapping of polyethylene plastic is used to

274

RONALD S. WIELGUS

J. Res. Lepid.

Fig. 3.— Metal tray container holding several Megathymus streckeri and
Megathymus texanus texanus larvae reared on artificial diet in plastic tubes.
Scale in inches.

Fig. 4. — Closeup view of plastic tube ends shown in Fig. 3. with two short
tents made by streckeri larvae. Note that the cotton plugs have been incor-
porated into the tents. Scale in inches.

13(4):271~277, 1974

ARTIFICIAL DIET

275

encase the PVC pipe ends in order to reduce moisture loss of
the diet through evaporation. In some instances, a few larvae
may begin to bore out of the bottoms of the pipes through the
plastic wrapping. If the pipes have been set upon metal or
tough plastic containers, further boring out is prevented. Tent-
building larvae will defecate outside of the tent and the fecal
pellets will accumulate on the bottoms of the boxes, etc. Mature
larvae will powder up and pupate in their surrogate yucca
caudices (pipes) after several months of feeding. Under labora-
tory conditions, the feeding period is markedly shortened and
diapause may be non-existent or brief (lasting a few weeks
instead of months).

NON-TENT-BUILDERS

For the non-tent-builders, the problems of rearing are more
acute. Containers must be impenetrable to larval mandibles.
In the early instars (up through fourth instar), the larvae of
streckeri and texanus are notorious borers and must be confined
at all costs. Otherwise, they may bore out unnoticed and escape,
never to be found again, or discovered many days or weeks later
in a state of severe desiccation.

Virtually all of the non-tent-building larvae reared by the
author have been ex ova. The initial container is an ordinary plas-
tic drinking straw filled with diet to within 20 mm of one end. The
end of the straw destined to be the bottom is folded over and
stapled down. The other end of the straw receives a small yucca
plug which is pushed into the diet a short distance. Then an ovum,
still attached to a portion of a yucca leaf, is introduced into the
straw and the straw end folded over and stapled. The newly-
hatched larvae will bore into the yucca plugs and ultimately
into the diet. The straws, in clusters of twenty or more, are
stood upright in ordinary glass jelly jars which have been
properly labeled. As the larvae feed on the diet and moult, they
are observed through the clear plastic. Frass is packed into the
straws behind them. Before their diet is all consumed, the larvae
may have outgrown the straws and will need to be transferred.

The next container is an intermediate one and consists of a
semi-transparent polyethylene plastic tube 9 mm in inside
diameter by 30.5 cm in length with a wall thickness of approxi-
mately 2 mm. This size will allow streckeri and texanus larvae
to feed up through the fifth instar. The tube is filled with diet
to within 4 cm of one end. At this end the larva is allowed to

276

RONALD S. WIELGUS

J. Res. Lepid.

make its own way in. Both ends of the tube are then capped
with squares of sheet aluminum 10 mm on a side by 0.5 mm in
thickness taped in place. The aluminum prevents boring out
and escape of the larva. Sufficient air is present in the diet and
the tube to allow for larval respiration. The larva is easily ob-
served through the semi-transparent plastic. Ecdysis is accur-
ately determined and exuviae readily recovered utilizing plastic
tube containers.

The final container is similar to the intermediate one, the
only difference being one of greater inside diameter. In this
case, tubes measuring 12 mm in inside diameter are used.
Larvae well into fifth instar and beyond will still require that
aluminum caps be placed over the bottom ends of the tubes.
The upper ends (tubes are placed upright or with one end
higher than the other in various containers) however, from
which the larvae will construct their tents, are plugged only
with wads of cotton. The larvae will defecate against the cotton
plugs. Later, after diapause, they will push out the cotton plugs
or bore through them when tent building is initiated.

At any time during the rearing, it may become necessary to
transfer the larvae to fresh diet. For non- tent-builders, removal
of each larva is a simple matter. First, uncap both ends of the
tube. Second, hold the tube so that the bottom end is above a
level surface a few centimeters. Then, blow into the upper
end of the tube with gentle but sustained pressure. This will
force the contents of the tube (including larva) out of the
bottom end and onto the surface. Thereupon, the larva may be
collected, inspected and introduced to a new container.

The ideal container for a non-tent-building larva appears to
be one in which the inside diameter of the tube corresponds
exactly to the larval one. Non-tent-building larvae are notorious-
ly wasteful if given a column of diet greater than their larval
diameter.

The habit of plugging the burrow behind them with frass
as they feed (in contrast to the tent-builders) exposes the non-
tent-builders to the hazards of fungus and bacteria which may
culture within the tubes. Aside from vigilance and transfer to
fresh diet when this problem arises, the author has not found
a satisfactory solution. Consequently, losses due to disease are
much greater with laboratory reared non-tent-building larvae
than with the tent-builders.

13(4):271-277, 1974

ARTIFICIAL DIET

277

CONCLUSION

The use of artificial diet for the rearing of Megathymus
larvae affords an opportunity to other workers interested in
this genus. Under controlled laboratory conditions, resources
permitting, there appears to be little limitation to the number
of larvae or taxa involved which one can successfully rear to
adults. Larval duration can be significantly reduced (depending
upon the taxon) from a time span of around ten months to as
little as 120 days from egg to adult by employing the techniques
described herein.

It is hoped that the techniques presented in this paper will
encourage others in the rearing of Megathymus larvae. Those
who will try their hand at rearing these fascinating beasts are
urged to try other techniques. By taking advantage of their
innate larval behavior, the worker will find Megathymus larvae
relatively simple to rear.

ACKNOWLEDGEMENTS

The author is deeply indebted to Mr. Merrill Petterson,
USDA Vegetable Insects, Mesa, Arizona, for providing artificial
diet as needed throughout these rearings.

The special efforts of Don B. and Viola Stallings resulted
in additional ova of streckeri and texanus being made available
for this study and to them the author is ever grateful.

Joseph R. Wielgus of Glendale, Arizona, provided pleasant
companionship during fieldwork in 1974 and assisted in the
rearing program.

Thanks are again due Dr. Frank F. Hasbrouck, Associate
Professor of Zoology and Curator of Insects, Arizona State Uni-
versity, Tempe, Arizona, who critically reviewed the manuscript.

LITERATURE CITED

DOS PASSOS, C. F., 1964. A synonymic list of the Nearctic Rhopalocera.

Lepid. Soc. Mem., No. 1, New Haven, Conn., v -j- 145 pp.
PETTERSON, MERRILL A. and RONALD S. WIELGUS, 1974. Accept-
ance of artificial diet by Megathijmus streckeri (Skinner) (Lepidoptera:
Megathymidae). /. Res. Lepid., 12 (4); 197-198.

WIELGUS, RONALD S. and DON B. STALLINGS, 1974. The laboratory
biology of Megathymus streckeri and Megathymus texanus texanus
(Megathymidae) with as.sociated field observations. Bull. Allyn Mus.,
No. 23, 15 pp., 33 figs.

WIELGUS, RONALD S. and DALE WIELGUS, 1972 (1973). Some tech-
niques for the rearing of Megathymus larvae. J. Res. Lepid., 11 (4):
245-250, 2 figs.

Journal of Research on the Lepidoptera

13(4):278-280, 1974

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

BUTTERFLIES OF THE WORLD, by H. L. Lewis, illustrations
by Lionel Leventhal Ltd. 1973. xvi + 312 p., 208 color plates,
2 drawings, 1 map. Follett Publishing Co., Chicago. Price $29.95.

There has long been a need for a general butterfly book which covers
a broad geographic area. Mr. Lewis’s book accomplishes this admirably,
as tlie coverage includes all families of butterflies on a world-wide basis.
More than 5,000 species of butterflies are represented in the color plates.
The author has made the following geographic divisions: North America,
South America including Mexico, Europe, Africa, Asia, Indo-Australia.
For each region, he has figured representatives of each family including
the Hesperioidea, which are all too often excluded from major works. The
more common species from each region are figured along with many of the
uncommon ones.

The book will be valuable to the amateur and serious collector alike.
Its major value will be enabling the easy identification of an unknown
specimen as to family, and in most cases, genus, if the specimen itself is
not figured. At this point, one can refer to the existing regional works
for in-depth study.

Considering the relatively low cost of the book and the number of
color plates, the plates are of excellent quality. Some plates are slightly off
in color, but on the whole, they are very good. Each plate contains a
small map of the geographic area represented. This is a nice touch and
provides a convenient reference.

The text portion of the work is not really a scientific treatise.
It presents information about some but not all of the specimens shown in
each of tire plates. This inclules scientific name, authority, English name
in some instances, general geographic distribution, similar species, brood
information, and larval foodplant in some cases. There is included a com-
plete index. The book is in reality a pictorial encyclopedia of the world’s
butterflies.

Mr. Lewis is to be complimented on his choice of the speeies figured.
The treatment is well-balanced for each geographic region. He has not
restricted the figures to “showy” species and the plates present a good
general coverage of fauna.

As with any work of this scope, there are some negative comments
to be made about the book. The author should have had his manuscript
reviewed by collectors or authorities familiar with each of the geographic
regions. There are some problems with nomenclature and misidentification
of species and genera.

In the treatment of Nortli American families, there is no consistency
concerning the use of generic and subgeneric names, or in the use of
species and subspecies names. Examples are the use of Stnjmon without
recognition of Chlorostnjmon, Euristnjmon etc., while Icaricia, Agriades
etc. are used rather than Plehejus. There are many instances of this nature.
In plate 15, for example, Spetjeria calgariana and S. callippe are figured
without any indication that two subspecies of the same species are involved.
This is a common occurrence throughout tlie plates.

A “corrigenda” sheet accompanied the book, but a number of items
are not included. Some specific examples are:

278

279

Pi. 12. Papilio daunus is used rather than P. multicaudata;
Dione rather than Agraulis is used; tlie butterfly flgured (Fig.
17) as Libytheana carinenta appears to be L. hachmanii lar-
vata (Strecker). Both L. carinenta and Danaus eresimus
(Fig. 18) should probably be considered casual for the geo-
graphic region (North America exclusive of Mexico) pre-
sented. No mention of this is made in the associated text.

PI. 13. Anartia amathea (Fig. 3) is not found in the region
designated as North America by the author according to the
map on page x.

PI. 14. The genus Metamorpha rather than Siproeta is used.
Fig. 25. Polygonia orcas is presumably oreas.

PL 15. Thessalia leanira is shown as Phyciodes leanira.
Speyeria adiante rather than S. adiaste is used. There is some
question about including the Hawaiian Pyrameis tammeamea
(Fig. 3) in the region represented.

PI. 16. As in PI. 15, species and subspecies names of Speyeria
appear randomly. Vanessa carye is used rather than V. or
Cynthia anabella.

PI. 17. The insect figured (Fig. 17) as Colias hageni is
Colias interior. The former is a ssp. of C. philodice. Fig. 18
appears to depict the underside of Colias scudderi and not C.
interior. Fig. 19, is Colias eurytheme and not C. pelidne.
There is confusion between Anthocharis and Euchloe as the
generic name in several cases. Paramidea is introduced as the
generic name for Anthocharis genutia, and Pontieuchloia for
the genus in P. protodice. Neophasia terlootii is misspelled as
N. terlooii.

PL 18. Fig. 15. Gnodia portlandia in place of Lethe port-
landia. Fig. 16. E. ctjmele should be E. cymela.

PL 19. Fig. 6. Satyrodes eurydice is used rather than Lethe
eurydice.

PL 19. Plebejus (Icaricia) shasta (Fig. 37) is assigned to
Lycaeides. Paramecera xicaque, if from Arizona, should be
P. allyni, (Fig. 6). Two extinct examples are shown in G.
xerces and G. x. fm. antiacis.

PL 20. Fig. 12, Glaucopsyche piasus daunia is shown as
Phaedrotus [sic] sagittigera. The associated text restricts
the butterfly to the Rocky Mts. without mention of the
coastal races. Fig. 17, Plebejus aquila [sic] = aquilo. Here
Plebejus is used for aquilo when in PL 19, Agriades was used
with podarce. The generic names Lephelisca and Lymnas are
used rather than Calephelis and Melanis. All of the Strymoniti
are treated as genus Strymon. Strymon saepium is shown as
Mitoura saepium.

PL 22. Hesperia ruricola (Fig. 8) appears which is a Nomen
dubium. Fig. 23, Pholisora aepheus [sic] = P. alpheus. Fig.
44, S. zampa = pulverulenta and the specimen shown is
probably S. evansi. Fig. 25, P. mejicanus. The associated text
states Canada to Texas for the range. Obviously P. catullus
and P. mejicanus have been confused. The former is found
from Canada to Mexico. The latter occurs in Colorado, New
Mexico, Texas, and Mexico.

PL 24. Fig. 7, depicts a Troides as Papilio xanticles. The
butterfly shown appears to be Troides rhadamantus (Lucas)
from the Philippines. A similar confusion occurs on PL 136 as
noted in the ''corrigenda”.

PL 43. Fig. 27. Dryadula phaelusa [sic] = D. phaetusa.

280

PL 48. Figs. 10, 11 Lycorea ceres and L. cleobaea appear to
be misidentified.

PL 74. The genus Lymnas rather than Melanis is used.

This reviewer has restricted the majority of his critical comments to
the North American region, the area with which he is most familiar. In
the text which accompanies Plates 12-22, many of the listed geographic
ranges are either incomplete or incorrect. A few examples have been noted
above. This is truly unfortunate when such works as Klots’s A FIELD
GUIDE TO THE BUTTERFLIES (1951), Ehrlich and Ehrlich’s HOW
TO KNOW THE BUTTERFLIES (1961), and many regional guides pub-
lished in the past decade are available. The dos Passes Checklist has been
available since 1964 for nomenclature, and several revisions have appeared.
These shortcomings of Mr. Lewis’s work detract from its overall quality
and utility.

In some cases, European generic names have been applied to North
American genera. It is not yet clear that these names correctly apply to
our fauna.

Despite tliese failings, the book is still a useful reference, and one
that many collectors will wish to own.

Clifford D. Ferris, Bioengineering Program, University of Wyoming,
Laramie, Wyoming 82071.

NOTICES

NEW PUBLICATIONS:

BUTTERFLIES OF LEBANON

BY TORBEN B. LARSEN U. S. A. ^22.50

GEOGRAPHIC VARIABILITY IN SPEYERIA (Reprint)

BY ARTHUR H. MOECK U. S. A. ^3.50

both the above available at the indicated prices from

Entomological Reprint Specialists
P. O. Box 77224, Dockweiler Station
Los Angeles, California 90007

Volume 13 Number 4 December, 1974

IN THIS ISSUE

Toward a theory of butterfly migration

Oakley Shields 217

Atrytonopsis hianna biology and life
history in the Ozarks

J. Richard and Roger L. Heitzman 239

Larval migration of Hyles lineata (Fab.)

James F. Wells and Richard M. Brown 246

New food plant for Darapsa pholus (Cramer)

J. C. E. Riotte 247

Notes on Arctic and Subarctic collecting

Clifford D. Ferris 249

Kloet and Hincks’ Check list of British Lepidoptera
Insects (Lepidoptera) Edn. 2. A reply to criticisms
J. D. Bradley, D. S. Fletcher and P. E. S. Whalley 265

The Nomenclature in an Important British
check list (1972). Part 3. Correct gender
for generic names derived from classical
without change of termination Juraj Paclt 267

Artificial Diet: The key to the mass
rearing of Megathymus larvae

Ronald S. Wielgus 271

Review: Butterflies of the World

by H. L. Lewis Clifford D. Ferris 278

THE aOURNAL c) ^ ^
OF FsESEARCHJ
©NJ THE LEPIJOOPTERA

Volume 14 Number 1 March, 1975

published by

The Lepidoptera Research Foundation, Inc.
at

1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz

Associate Editors:

Thomas C. Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601

Maria Etcheverry, Centro de Estudios Entomologicos, Casilla 147, Santiago,
Chile.

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Canada.

Brian O. C. Gardner, 18 Chesterton Hall Crescent, Cambridge, England.

Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.

Lee D. Miller, The Allyn Museum of Entomology, 3701 Bay Shore Road,
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ONJ THJE LEPIJDOPTE^A

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Edited by WILLIAM HOVANITZ

Volume 13

1974

published by

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at

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Journal of Research on the Lepidoptera

14(l):l-40, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1973

MATE-LOCATING BEHAVIOR OF
WESTERN NORTH AMERICAN BUTTERFLIES

JAMES A. SCOTT

Department of Entomology, University of California, Davis, 95616

Abstract. Mate-locating behavior, including the methods used to locate
mates, the location of mating in relation to the larval foodplants and to
topography, and the time of day of mating, was studied in 284 species of
western nortli american butterflies and skippers. 18 pairs of congeneric
species of butterflies and skippers were found to mate at contrasting topo-
graphic sites; tile use of separate mating sites by these congeneric species
certainly prevents, and may result from, interference competition during
mate-locating behavior. Some trends in mate-locating behavior were ob-
served: 1) perching species more often tend to mate in restricted sites in
the habitat, and more often mate during a restricted part of the day,
than do patrolling species; 2) perching rather than patrolling behavior is
more frequent in species that have only 1-2 ratlier than 3-4 broods; 3)
species emerging at the end rather than at the start of the season, and
species with large movements, more often have patrolling rather than
perching behavior and more often mate throughout the habitat rather than
in resricted sites; 4 ) species feeding on slirubs or trees rather than on herbs
more often tend to be perching rather than patrolling species and more
often tend to mate during a restricted part of the day; 5) high mountain
species usually patrol and mate at any time of day. Mate-locating behavior
provides many useful taxonomic characters; closely related species usually
have similar mate-locating behavior. Despite the trends noted above, dif-
ferent mate-locating strategies are often used by species feeding on the
same larval foodplants. The strategy used often seems to be that used by
a ^ taxonomic relative feeding on other plants, but convergence of mate-
locating behavior of taxonomically distant species feeding on the same
plants has sometimes occurred.

Mate-locating behavior is defined as behavior which brings
the sexes together for mating. It includes the methods used to
find mates, the location of mating, and time of day of initiation
of mating ( Scott, 1974a ) .

Elsewhere I presented a general discussion of mate-locating
behavior of butterflies (Scott, 1974a). There are three methods
used to locate mates: perching behavior (males rest at charac-
teristic sites and investigate passing objects by flying out at
them to search for females; females generally fly to these sites
to mate, then they depart ) , patrolling behavior ( males fly almost
continuously in search of females ) , and use of pheromones from
more than a few meters away (this method seems quite rare).

1

2

JAMES A. SCOTT

J. Res. Lepid.

Movement, size, wing color, wing pattern, and odor are stimuli
which can be transmitted during sexual communication in the
approach of a male to a female (Scott, 1973a). Perching males
are highly attracted to moving objects, whereas patrolling males
often are attracted to motionless objects resembling in some way
females. A patrolling male may rarely find a female which gives
off a pheromone when he wanders near her position. Perching
species usually mate in limited areas of the habitat, often during
only part of the day, whereas patrolling species usually mate
throughout the habitat at any time of day.

This paper is a detailed survey of mate-locating behavior of
several hundred western North American butterflies (Papiliono-
idea and Hesperioidea ) . In it I summarize mate-locating be-
havior for each species, and attempt in the discussion to de-
termine major trends of behavior among all the species.

METHODS

To determine mate-locating behavior, adult behavior was
observed, especially interactions between males and between
sexes (including courtship and mating). In perching species,
such interactions occurred predominantly when resting males
investigated moving objects. In patrolling species, interactions
occurred predominantly when resting males investigated mov-
ing objects. In patrolling species, interactions occurred pre-
dominantly when flying males investigated either flying or
resting individuals. Perching and patrolling differ from resting
and flying respectively, by the addition of investigative behavior.
In a few species, noted in the results, both behavior patterns
occur.

Location and time of day of mating were determined from
field observations of such interactions. Varied topographic and
vegetational sites were studied when possible to make sure that
a restricted location of mating was characteristic of the popula-
tion rather than due to limited opportunity.

Time of day of mating in this paper refers to initiation of
mating. Therefore, field observations of copulating pairs were
used with caution when initiation of mating was not seen. This
is because copulation occasionally lasts several hours (very
rarely several days) (Scott, 1973a), so that time of observation
of the pair lags behind time of initiation. It is therefore possible
to observe a copulating pair in the afternoon, if in that species
courtship only occurs in the morning. Therefore time of day of
mate-locating behavior, courtship and initiation of mating, were

14(l);l-40, 1975

MATE LOCATING

3

given priority over time of observation of copulating pairs whose
initiation was not seen. Species which are stated herein to court
and mate during only part of the day were carefully studied to
make sure that the restricted mating period was characteristic of
the population and not a result of inclement or unusual weather.

The ideal method of studying the mate-locating behavior is
to observe and record all details of male-male and male-female
interactions, including dozens of completed courtships. For
some species in this paper such as Precis coenia the data are
based on more than 40 completed courtships in addition to other
courtships and male behavior. However, it is not necessary to
observe many completed courtships to determine mate-locating
behavior. In all the species so far studied in detail ( see literature
cited), the following statements are true. In butterflies, the
mating process always involves an investigation by the male of
other individuals. Mating therefore occurs only at the locations
where males investigate other individuals, and only during the
time of day when males investigate other individuals. The
method of mate-location is determined by observing whether
males fly or sit before interacting with other individuals. These
statements suggest that a complete description of mate-locating
behavior can therefore be based only on male behavior, but
observations of completed courtship are desirable supplements
for confirmation. I include some species in this paper for which
no completed courtships were seen.

Study of mate-locating behavior of butterflies can easily be
carried out by those readers willing to spend field time observ-
ing behavior. The basic method is to observe interactions ( male-
male and male-female) and to record when and where these
occur, and whether the male was resting or flying just prior to
the interaction. Observations of resting or flying alone are of
very litle value without observations of interactions. Again I
emphasize that perching and patrolling as I define them differ
from resting and flying respectively in that perching and patrol-
ling males investigate other individuals, whereas resting and
flying males may not. I emphasize this because some people are
treating the words perching and patrolling as the same as resting
and flying respectively; to avoid such confusion it may eventu-
ally be necessary to propose special latin names for perching
and patrolling behavior.

Some authors would use territoriality prominently in a dis-
cussion of mate-locating behavior. Territoriality may be defined

4

JAMES A. SCOTT

/. Res. Lepid.

as males remaining in a small area to search for females, where
males intentionally drive away other males. It may therefore
be the fourth criterion for classification of mate-locating behavior
(the first three being where mating occurs, when it occurs dur-
ing the day, and the method used to locate females (perching
or patrolling usually ) ) . I do not use “territoriality” because no
butterfly has conclusively been shown to be territorial with my
interpretation of the vertebrate definition of territoriality (Scott,
1974a), and because demonstration of territoriality requires de- '
tailed study of movement of marked individuals which are be-
yond the scope of this paper. Some previous authors have used
“territoriality” for what I call perching behavior, other authors
have used it for what I call patrolling behavior where males
sometimes return to the same spot, and still other authors ap-
parently do not use it in any consistent manner. The methods
and terms used in this paper were operationally constructed for
use with butterflies, whereas “territoriality” was originally used
for vertebrates and has since been applied so loosely that its
meaning has become diffuse and the benefits from its use ( espe-
cially with invertebrates) have become few.

RESULTS

These studies were conducted mainly in Colorado, especially
the eastern slope of the continental divide and adjacent plains,
and to a lesser extent in other western North American states
and provinces, especially California. Eleven species were studied
in detail in Colorado and California from 1969 to 1973 (see
literature cited). During 1972 to 1974 over 200 field days were
spent, mainly in Colorado, gathering data for this paper.

Supporting data including localities and times of observations
of interactions, courtship and mating are not included in this
paper due to space limitations; these data can be obtained from
the author should they prove important to the reader. As a
compromise I have indicated the quality of the observations as
follows (a — about 100 or more observations of interactions, con-
clusions unlikely to change with further study; b — about 50-100
observations of interactions, conclusions may change slightly with
detailed study; c — less than 50 observations of interactions, con-
clusions may change somewhat with further study and are pre-
sented mainly to increase the taxonomic variety studied).
Localities studied are the eastern slope of the continental divide
and adjacent plains of Colorado (Jefferson, Boulder, Douglas,
El Paso, Clear Creek, Park, Chaffee, Custer, Pueblo, Fremont,

l4(l):l-40, 1975

MATE LOCATING

5

and Saguache Counties) unless otherwise stated. An asterisk is
used for species occurring in the Pueblo area (Pueblo, Fremont,
Custer, Chaffee, Huerfano, and Saguache Counties, Colorado)
because these are analyzed in the discussion. Shields & Emmel
(1973) have also compiled a list of mating times, and Shields
(1967) provides information on several species not mentioned
here. Several errors in Table 2 of Scott (1974a) are corrected
herein. All times are reported as 24-hour standard time. The
words ‘'all day” in any context mean that interactions occurred
at least from before 0900 to after 1500.

PAPILIONIDAE

Parnassius clodius. Males patrol all day about meadows and
hillsides near the larval host. They patrol most often in swales,
and often dip down between shrubs to search there; males have
a somewhat bobbing flight (Nevada, El Dorado, and Siskiyou
Cos. Calif. — b).

"^Parnassius phoehus. Males patrol all day in open areas
where the larval host is abundant, slightly more frequently in
sheltered sites (Scott, 1973b). Patrolling is most frequent on the
leeward side of ridges both in Colorado and in California (Al-
pine & Siskiyou Cos.). Males patrol about Vim above ground,
with a faster and steadier flight than P. clodius (a).

^Papilio polyxenes, *F. bairdii, *P. zelicaon (and form
nitra). Males patrol and sometimes perch, throughout the day
(about 0900-1530 for Colorado zelicaon). At the usual fairly low
density, males patrol but often perch, mainly on large hilltops.
At high density and in the absence of hilltops males patrol and
mate near the larval hosts (Guppy, 1969). In riparian forest,
males perch in clearings among the trees (Sacramento, Cali-
fornia, Arthur Shapiro, pers. comm.). Some hilltopping males in
some localities may patrol only in a restricted area of a hilltop;
this behavior might possibly be territorial ( a ) .

Papilio indra. Males perch and patrol all day at about equal
frequency, on somewhat sheltered rocky outcrops just below a
hilltop (Jefferson Co. Colo. & Napa Co. Calif.) (b).

"^Papilio rutulus. Males patrol all day up and down streams
with much riparian vegetation (they rarely patrol on hilltops
for a few minutes in Colorado and Napa Co. Calif., then depart).
In Davis, California, a suburb, males patrol about sunlit open-
ings in wooded parts of the city (b).

"^Papilio multicaudata. Males patrol all day up and down
canyon bottoms, which are usually drier than those frequented

6

JAMES A. SCOTT

/. Res. Lepid.

by P. rutulus (a).

Papilio eurymedon. Males patrol (and sometimes perch) all
day on large hilltops; males often patrol a small area there
(Colorado, Napa Co. Calif.) (b).

Battus philenor. Males patrol all day on hillsides and flats
near the larval host in Contra Costa and Yolo Co. California
(they occasionally patrol on hilltops near Austin Texas and in
Napa Co. California) (c).

PIERIDAE

Anthocaris sara. Males patrol all day up and down valley
bottoms, or more haphazardly on flat land, but always near trees
in forested areas (b).

Anthocaris lanceolata. Males patrol all day mainly in valley
bottoms or steep hillside draws (El Dorado & Nevada Cos.
Calif.) (c).

A. cethura. Males patrol on windless sides of ridgetops at
least from 1130-1430 (Churchill Co. Nevada) (c).

"^Euchloe ausonides. Males patrol all day in open areas,
slightly more frequently in valley bottoms than elsewhere, in
Colorado and Contra Costa Co. Calif, (a; Scott, 1975a).

*£. olympia. Males patrol all day on hilltops (b).

£. hyantis. Males patrol all day usually on hilltops, but
sometimes throughout the habitat or restricted to areas of larval
foodplant (Napa & Nevada Cos. Calif., Churchill & Lander Cos.
Nevada, Pima Co. Arizona) (b).

"^Colias meadi, ^alexandra, eurytheme, "^philodice. Males
patrol all day in open areas regardless of topography, (b).

C. occidentalis. Males patrol all day in valley bottoms, open
woods, or on hillsides near the larval host (Colusa & Lake Cos.
Calif.) (c).

C. cesonia. Males patrol all day, especially in valley bottoms
and on flats (Pima & Santa Cruz Cos. Arizona) (c).

Eiirema nicippe, mexicana, proterpia, boisduvaliana. Males
patrol all day, mainly in gullies and flats (Pima & Santa Cruz
Cos. Arizona) (c).

"^Nathalis iole. Males patrol all day several cm. above ground,
mostly in gullies, roadsides, or flat land (c),

^Pieris chlorodice beckeri. Males patrol all day up and
down usually dry arroyo bottoms (Colorado, and Churchill Co.
Nevada; b).

*P. sisymbri. Males patrol all day, usually on hilltops, hap-
hazardly when on flat land (Colorado, and Napa, Colusa, &

14(1); 1-40, 1975

MATE LOCATING

7

Alpine Cos. Calif.) (a).

*P. callidice Occident alis. Males patrol all day, mainly on
hilltops when they are available (b).

*F. protodice. Males patrol all day, often on hilltops, but
often on flat land when hilltops are unavailable (b).

*F. napi. Males patrol all day along partly shaded streams
in forests. Except for A. sara, the other Pieridae prefer open
areas (b).

*F. rapae. Males patrol all day near larval hosts near riparian
vegetation in native habitats and on flat land in cultivated areas.
They share the slow patrolling flight of F. napi (b).

"^Neophasia menapia. Males patrol all day, circling slowly
about the larval foodplant trees (b).

Phoehis sennae. Males patrol all day, often on flat land
(Mexico and Texas).

NYMPHALIDAE

^Euptychia dorothea. Males patrol all day up and down
narrow dry shaded gullies (b).

E. rubricata. Males patrol all day, usually in semi-shaded
sites such as gullies (Texas) (c).

Euptychia cymela. Males patrol all day, mainly in woods
under shading trees (Ohio, and Yuma Co. Colo.) (c).

^Coenonympha tullia. Males patrol all day in grassy areas
regardless of topography (a).

"^Neominois ridingsii. Males perch (and seldom patrol brief-
ly) from about 0750 to about 1100, strongly from 0830 to 1000,
and rarely later in the day, on small (3m) grassy hilltops, mesas,
or saddles (Fremont Co. Colo., Scott, 1973c). Behavior in Mono
Co. Calif, is similar (a).

"^Cercyonis pegala. Males patrol all day in grassy areas, main-
ly around riparian trees ( b ) .

*C. oetus. Males patrol all day in open grassy areas regard-
less of topography ( b ) .

C. sthenele. Males patrol all day in grassy areas, often in
gullies and valley bottoms (California, Colorado) (c).

*C. meadi. Males patrol all day everywhere in the habitat
but most often in valley bottoms and in woods (b).

Oeneis nevadensis. Males perch all day on fallen trees and
other objects in lanes of valley bottom forest (Siskiyou Co.
Calif., c). Males also perch on hilltops, but I have not seen this.

*0. chryxus. Males perch (and occasionally patrol briefly)
all day, usually on hilltops or on sloping ridgetops (Colo.) (b).

8

JAMES A. SCOTT

/. Res. Lepid.

O. c. ivallda males perch and when disturbed patrol before
settling, all day on ridgetops (Inyo Co. Calif.).

alberta. Males perch, and often patrol 10 m or less, all
day among bunch grasses, usually in somewhat sheltered hollows
of hillsides and valley bottoms (c).

*0. uhleri. Males perch, and occasionally patrol short dis-
tances, all day on slopes at the base of bunch-grass community,
or on a trail below the bunch grass community, usually on south-
facing slopes (b).

O. polixenes. Males mainly patrol but sometimes perch
(patrolling flights are often 10 m or more) all day in swales on
grassy north-facing slopes (b).

^O. melissa. Males perch, and often patrol, all day in rocky
areas of hilltops and ridgetops (b).

O. taygete. Males perch and patrol all day on grassy hill-
sides and sometimes on hilltops (b).

"^Erebia magdalena. Males patrol all day over rockslides (b).
epipsodea. Males patrol all day over wet grassy meadows
or swales (b).

E. theano. Males patrol all day about grassy areas on slopes
or in bogs; males appear to be very local and spend most of their
time sitting among vegetation (Hinsdale Co. Colo., c).

E. callias. Males patrol all day usually over alpine cushion
plant community, often over rounded ridges, but sometimes on
slopes (b).

Gyrocheilus patrobas. Males patrol all day, usually in shady
grassy areas in oak woodland (southern Arizona, c).

^Euptoieta claudia. Males patrol all day in every topo-
graphic situation in open areas, especially on flats (b).

Speyeria nokomis. Males patrol all day in spring-fed mead-
ows, especially in valley bottoms if the spring is on a slope.
Males fly about Yz m above ground and dip into hollows to
search (Mono Co. Calif., Kane Co. Utah, Uinta Co. Utah, Mesa
Co. Colo., b).

*S. aphrodite. Males patrol all day, mainly in open areas,
regardless of topography (a).

*S. edwardsii. Males patrol all day in open areas regardless
of topography, but several males patrolled in a small space just
below the summit of a large hilltop, remaining there for several
hours (b).

*S. coronis. Males patrol all day in open areas regardless of
topography (S. aphrodite, edwardsii, and coronis often patrol

14(l);l-40, 1975

MATE LOCATING

9

the bottoms of valleys) (b).

S. zerene. Males patrol all day in open areas regardless of
topography (Routt Co. Colo.; Idaho) (b).

S. callippe. Mate-locating behavior occurs all day. In the
Front Range of Colorado, males patrol and occasionally perch
on shrubs etc., mainly on hilltops until about 1300, when they
patrol (very rarely perch) more generally on hillsides. Court-
ship and mating occur all day despite this behavioral switch.
Males patrol in fairly open areas close to the ground, in contrast
to the preceding four species of Speyeria which patrol about
1 m above ground (b). In Glenn and El Dorado Counties, Cali-
fornia, males patrol all day about Yz m above ground in the flat
forest (c). At Mt. Diablo, Contra Costa Co. Calif., males some-
times perch on shrubs, grass, etc., and also patrol. Perching was
noticed sometimes until about 1200, after when males would
only patrol. Perching and patrolling occurred only on the upper
100 feet of the hilltop, mostly in clearings on the ridge (c).

S. atlantis. Males patrol all day in wet meadows and along
streams in Colorado (b) and Coconino Co. Arizona, and patrol
about open forest all day in Glenn Co. Calif, (c).

S. egleis. Males patrol all day, mostly along shaded forest
lanes such as abandoned roads, about 1/3 m above ground (Routt
Co. Colo., b). In Mendocino and Alpine Cos. California, males
patrol all day near the ground mostly on hilltops (c).

*S. mormonia. Males patrol all day just above the ground in
open vegetated areas regardless of topography, usually in mead-
ows at lower elevations (b).

^Boloria selene. Males patrol all day in wet meadows and
springs, usually on fairly level land but sometimes on hillsides

(b).

B. eunomia. Males patrol all day among the willows of wil-
low bogs, often near the edge of a bog. (b).

B. frigga. Males patrol all day, in willow bogs, usually in
low spots between a slope and a flat, in mixed willow-grass
parts of the willow bog (but some males can be found through-
out the bog) (b).

freija. Males patrol all day in open areas near the larval
hosts, usually at the edge of a bog or stream, often on hillsides

(b).

*jB. titania. Males patrol all day in open areas, usually swales
and valley bottoms, in meadows or bogs. At willow bog locali-
ties, males patrol the borders of the bog but rarely enter the bog

10

JAMES A. SCOTT

/. Res. Lepid.

center (b).

B. toddi. Males patrol all day in moist meadows (Alberta,
Summit Co. Colo.) (c).

B. epithore. Males patrol all day in moist shrubby meadows
and moist open woods, primarily in valley bottoms (Siskiyou &
Glenn Cos. Calif., b).

B. alberta. Males patrol all day over open nearly barren or
vegetated flats and hillside scree slopes, especially where Dryas
octopetala is abundant (Alberta, c).

B. astaHe. Males patrol all day on ridgetops and hilltops,
usually just below the leeward edge of the ridgetop or plateau
(Alberta, c).

B. kriemhild. Males patrol, at least during morning and mid-
day, over moist meadows, usually where aspen is common
(Montana, c).

Etiphydryas chalcedona. Males perch and occasionally
patrol all day. Perching often occurs on shrubs in hollows below
a slope (Napa Co. Calif., b). Near Rosita, Custer Co., Colo.
(E. c. near eurytion), males patrol all day in meadows, espe-
cially in grassy swales (b). At Aspen, Pitkin Co. Colo., males
patrol all day all over a hill ( c ) . In the foothills of the Colorado
Front Range (E. c. capella), males usually perch but sometimes
patrol, all day on low and large hilltops (c). At Diamond Peak,
Moffat Co. Colo. (E. c. near hernadetta), behavior is very simi-
lar to capella (c). In Sevier Co. Utah, males patrol as in eurytion
(c). In El Dorado and Alpine Cos. Calif, (b), E. c. sierra males
patrol and sometimes perch all day on hilltops at low density,
but mainly patrol throughout the habitat especially over flowers
at high density.

Euphydryas editha. In western Colorado (Moffat Co. and
Pitkin Co., b) males patrol and often perch all day on ridgetops
and hilltops. At Connors Pass, White Pine Co. Nevada (c),
males mainly perch on hilltops. At Jasper Ridge, San Mateo Co.
Calif., males only patrol all day (Labine, 1966). At Rutts
Canyon, Napa Co. Calif, (c) males perch and patrol about
equally on ridgetops all day. At Rlue Ravine, Sierra Co. Calif,
(c), males patrol and sometimes perch on shrubs on a serpen-
tine slope. At Goat Mtn., Colusa Co., Calif, (c) behavior is
similar but is on a ridgetop. At Mono Pass, Inyo Co. Calif, (c),
males patrol and sometimes perch all day on a ridgetop.

^Poladryas minuta. Males perch and rarely patrol on ridge-
tops and hilltops from about 0700 to 1230; from about 1230 to

l4(l);l-40, 1975

MATE LOCATING

11

about 1500 males patrol, usually on hillsides and flats etc. near
flowers. Courtship occurs throughout the day (a; Scott, 1974).

Chlosyne palla. In Gilpin and Routt Cos. Colorado (b),
males perch, and sometimes patrol, all day, on slight prom-
inences such as mine dumps in a valley bottom or on the outer
edge of a dirt road rounding a hillside. In El Dorado County
Calif. ( c ) males patrol and perch about equally, usually in valley
bottoms, roadside ditches, or swales, all day. Males patrolled
near flowers on a ridgetop once in Napa Co. Calif, (c).

Chlosyne definita. Males patrol just above the ground all
day, all over the ridgetops, hillsides, and gulches where a popu-
lation occurs (western Texas, b).

Chlosyne chinatiensis. Males patrol all day just above the
ground, mainly on ridgetops where Agave lecheguilla is abun-
dant; colonies may be somewhat local, being scarce on a ridge
and common on an adjacent ridge (western Texas, c).

Chlosyne theona. Males patrol all day on flats mostly (Mexi-
co, c). In southwest Texas and central Arizona males often occur
on hilltops, however (c).

Chlosyne neumoegeni. Males perch, and sometimes patrol,
in gulch bottoms (like Phyciodes mylitta) at least from 1130 to
1440 when observations were made (Pima Co. Ariz., c). In San
Bernardino Co. Calif, males perch in gulch bottoms all day (c).

Chlosyne hoffmanni. Males patrol all day about ni above
ground about valley bottom meadows, swales, and adjacent
shrubland near the larval host (Siskiyou Co. Calif., c), and in
roadside ditches (El Dorado Co. Calif., c).

"^Chlosyne acastus. Males perch, and sometimes patrol, in
dry washes and gullies all day ( Fremont Co. Colorado, southern
Utah, b).

Chlosyne damoetiis. Males patrol all day over rockslides
where the larval host is abundant (Custer Co. Colo., c). In
Alberta and California (Tuolumne, Mono, and Alpine Counties)
behavior is similar but males often patrol up and down chutes
in the rockslide (c).

"^Chlosyne leanira ftdvia. Males court and mate all day. At
fairly low density, males perch and occasionally patrol on low
hilltops near the larval host. Several times high density was
encountered, and males patrolled and mated in a relatively small
area where the population was concentrated (b). C. 1. alma
males perch, and sometimes patrol some after an encounter, on
hilltops (Kern Co. Calif., c). C. 1. leanira males mainly perch

12

JAMES A. SCOTT

/. Res. Lepid.

and sometimes patrol on hilltops all day (Napa Co. Calif., c).

Chlosyne elada. Males patrol all day about 14 m above the
ground throughout the habitat, often on flats (Arizona, Texas,
Mexico, c).

"^Chlosyne gorgone. Courtship and mating occur all day.
On hilltops, males mainly perch and seldom patrol. On hillsides
and valleys, males almost always patrol (b).

"^Chlosyne nycteis. Males patrol all day along streamsides
near Rudheckia laciniata (b).

Chlosyne lacinia. Courtship and mating occur all day. Like
C. gorgone, males patrol and sometimes perch on hilltops in
Texas, but elsewhere they mostly patrol (Mexico) (c).

"^Phyeiodes tharos. Males patrol all day in wet meadows and
streamsides (b).

"^Phyciodes campestris. Males patrol all day just above the
ground, mostly in meadows, grassy swales and valley bottoms
(Nevada & Yolo Co. California, and Colorado, a).

"^Phyeiodes picta. Males patrol all day just above the ground
in colonies mainly on flat land such as near streams, vacant
fields, beside railroad tracks, etc. (b).

Phyciodes pallida. Courtship and mating occur all day.
Males usually perch in gullies on 1 m tall twigs or other objects.
At Red Rocks, Jefferson County, Colorado, most males perched
in gullies but some males patrolled on slopes near a hilltop and
rarely males are found on hilltops (b). At Connors Pass, White
Pine County, Nevada, all three males observed patrolled on
ridgetops.

Phyeiodes mylitta. Mating occurs all day. In California and
Arizona males usually perch in gullies or between riparian shrubs
and a hillside (and occasionally patrol about 1/3 m above
ground, b). However, in agricultural areas of Idaho, Washing-
ton, and central California, males patrolled near Cirsium arvense,
a larval host growing in waste places (b).

Phyciodes orseis. Males perch all day, and occasionally
patrol, in gullies and between riparian shrubs and hillsides
(Siskiyou Co. California, Douglas Co. Nevada; b).

Phyciodes phaon. Males patrol all day, mostly on flats near
the larval hosts (western Texas, c).

Phyciodes vesta. Males patrol all day, often on flats, often
near waterholes in arid areas (Texas, c).

Phyciodes texana. Males usually perch (and occasionally
patrol about 1 m above ground), all day, mainly in gulches and

14(1): 1-40, 1975

MATE LOCATING

13

dry stream beds (southern Arizona, c).

Microtia dymas. Males patrol slowly all day, most often in
gullies and flats in hilly areas (southern Arizona, western Texas;

c).

Microtia elva. Males patrol slowly all day all over the habi-
tat, although colonies are somewhat local (Mexico, c).

^Polygonia satyrus. Males perch on vegetation in gullies or
along tree-shaded banks, from about 1230 to late afternoon (b).

^Polygonia faunus {'%ylas'). Males perch on shrubs or rocks
etc. in gullies, from roughly about 1300 to late afternoon (b).
My observations on Polygonia and Nymphalis, especially P.
faunus, indicate that hibernating individuals usually mate in
spring.

^Polygonia zephyrus. Males perch on bushes or stones, logs,
etc. in valley bottoms, from roughly about 1230 to late after-
noon (perching most actively after about 1300) (southern Colo-
rado, and Douglas Co. Nevada) (a).

P. oreas. In California, males perch on shrubs and small trees
in small clearings in valley bottoms (sometimes 3m above
ground), at least in the afternoon when observations were made
(c).

^Nymphalis antiopa. Males perch, and occasionally patrol,
in gullies or swales, and in valley bottoms, from about 1130 to
at least 1700 (start of perching behavior is gradual) (b).

^N. californica. Males perch, and occasionally patrol, start-
ing about 1400 and perching until 1700 at least, especially on
ridgetops, or on banks of valley bottoms, and on flat land on
the side of trees, etc. (California, Colorado) (b).

*IV. milberti. Males perch usually behind shrubs on hill-
tops, sometimes on logs etc. next to gully banks. Males arrive
between 1050 (rarely) and about 1400, averaging perhaps 1230,
and then perch until late afternoon ( b ) .

^Vanessa atalanta. Males perch, usually on the leeward side
of shrubs on hilltops, or on porch roofs of farm houses, and in
similar situations (b).

*V. virginiensis. Males perch, often on shrubs on hilltops,
or on the leeward side of shrubs on flats (c).

*V. caryae. Males perch, on shrubs on hilltops, or on flat
land on the ground behind cars or people or other large objects
(c) (Colorado and Yolo Co. Calif., c).

cardui. Males perch and sometimes patrol, usually on
shrubs on hilltops at low density. All four Vanessa species first

14

JAMES A. SCOTT

/. Res. Lepid.

arrive on hilltops at about 1330 plus or minus an hour or more
depending on meteorological conditions (Shields, 1967), and
most mating and courtship and perching occurs after this time
until as late as 1930 in June. The situation is not this simple,
however. In 1973 V. carclui was extremely abundant in Colorado.
Numerous interactions between individuals were seen from 0800
onward, although intensity and frequency of interactions was
greatest in afternoon especially late afternoon. Mrs. Betsy Webb
of the Denver Museum of Natural History observed a com-
plete courtship and mating at about 1200. Interactions occurred
wherever individuals were, including hillsides (a).

Precis coenia. Males perch all day on flat bare spots in fields,
trails, etc. (Scott, 1975b; a).

Adelpha hredowii. Males perch on 1-2 m shrubs usually in
gulches, and sometimes patrol, all day (Napa Co. Calif.; c).
Courtship occasionally occurred on Buckeye flowers.

Limenitis tveidemeijeri. Males perch, and rarely patrol, all
day in gullies and valley bottoms on bushes and treelimbs about
2-3 m above ground ( a ) .

L. lorquini. Males perch all day on shrubs about 2-3 m above
ground (Siskiyou Co. Calif.) in valley bottoms (c).

^L. archippiis. Males patrol all day along willow groves
along streams and irrigation ditches (b).

L. arthemis arizonemis. Males patrol all day along willow
groves; rarely in Texas males patrol a small area of a hilltop
(southern Arizona, western Texas; c).

Asterocampa celtis. Males perch on small trees along gul-
lies, on the rocks and logs and other objects at the side of a
gully, and often on my hat or net in a gully, or on the larval
host trees on flat land. Males perch most actively from early
afternoon to dusk, but interactions occur all day, as in V. cardui.

Asterocampa leilia. Males perch all day on rocks etc. in gully
bottoms, although there may be a gradual change in activity as
in A. celtis (s. Arizona, w. Texas; c).

"^Anaea andria. Males perch all day in clearings among cot-
tonwood groves, on stones, logs, and branch tips up to 3 m above
ground in the clearings (c).

"^Danaus plexipptis. Males patrol in flat areas near the larval
host, and interactions occur all day as early as 0800 (b, Colo-
rado). Dr. Lincoln P. Brower (pers. comm.) has observed many
matings all day in California, where mating occurs in early
spring at the roosting sites. The preference of D. gilippus for
mating in afternoon more than in morning may have been due
to the warmer afternoon weather (Brower et al. 1965).

14(l);l-40, 1975

MATE LOCATING

15

LYCAENIDAE

Calephelis arizonensis. Males perch all day in narrow gully
bottoms, on plants about m above the gully bottom such as
grass inflorescences (Santa Cruz Co. Ariz., c).

C. rawsoni freemani. Males perch all day on vegetation in
narrow gullies, or at the side of broader gulches (Chisos Mts.
Texas, c). Two other Calephelis spp., C. nemesis and C. wrighti,
behave differently. They remain on or near the larval hosts
(Clematis in southern Texas and Bebbia juncea near Blythe,
California respectively), and occasionally perch and may also
patrol there (c).

^Apodemia nais. A population was studied in Custer Co.
Colo, (b) in which individuals occurred on and near Ceanothus
bushes. Males perched usually on 1/5 m tall dead stalks in open
areas partly surrounded by shrubs between the mouth of the
gulch and a roadside ditch, occasionally on the tip of a Cerco-
carpus shrub in a clearing at the mouth of the gulch. Perching
behavior started almost precisely at 1130 in three days observa-
tions and lasted until about 1430. Before and after this perching
period, and seldom during, males feed on Ceanothus flowers
where females oviposit on Ceanothus.

Apodemia mormo. Males perch near the larval foodplant
(Eriogonum jamesi in Chaffee County, Colo., where most ob-
servations were), mostly in depressions such as gullies cutting
into hillisides, and depressions at bases of slopes. Perching starts
about 1100 and continues til 1430 or later. In the morning males
mainly feed on the larval host and fly away when disturbed.

"^Hypaurotis crysalus. Males patrol over and around tops of
the larval host trees, starting (gradually) about 1400 and end-
ing about 1730 (Scott, 1974d; a).

Atlides halesus. Males perch on treetops on hilltops, at
least in the afternoon when observations were made (Arizona,
New Mexico) (c).

"^Strymon melinus. Males perch on small trees on hilltops
when available (on tall shrubs elsewhere) from about 1300
( roughly ) to nearly dusk ( b ) .

^Harkenclenus titus. Males perch on small shrubs on hill-
tops (on taller shrubs if a colony occurs on flat land), from
about 1000 (not earlier) to late afternoon. Mating sometimes
occurs on the larval host (Prunus virginiana) (b).

^Satyrium acadica. Males perch (and seldom patrol) on

16

JAMES A. SCOTT

/. Res. Lepid.

small willow or other plants about 1 m tall, especially on low
plants growing out from a willow grove (up to 3-4 m away from
the grove), from about 1350 to dusk (b).

Satijrium calif ornica. Males perch (and occasionally patrol)
on top of 2-6 m trees and shrubs on ridges and hilltops usually,
from about 1400 to dusk (Jefferson Co. Colorado, and Napa Co.
Calif., b). One courtship occurred on a Buckeye flower in Cali-
fornia.

"^Satyriiim sylviniis. Males perch (and seldom patrol) on
small willow or other plants, especially those growing a few m
out from a willow grove, from about 950 (not earlier) to late
afternoon (about 1500) (Colorado, b). The same behavior was
observed in Mono Co. California and Lander Co. Nevada in
afternoon (c).

"^Satyrium liparops. Males perch on bushes or tree limbs
about 1-2 m tall or less (sometimes on the side of a small tree)
in gullies in the Front Range of Colorado (b). Perching and
courting occur all day. In Routt Co. Colo, males perched on
such shrubs on hilltops at least in morning (c).

"^Satyriiim calamis. Males perch all day on small shrubs or
other objects in gullies or depressions in oak groves in southern
Colorado (b). In Routt County, Colo., however, males perched
on small shrubs on hilltops at least in morning (c).

S. auretorum. Males perch and sometimes patrol on top of
about 5 m tall oak trees on ridgetops and hilltops, mainly from
about 1445 to dusk. Courtship sometimes occurs on Buckeye
flowers. I noticed that S. auretorum and S. californica usually
perched on separate trees (Napa Co. Calif., b).

S. tetra. Males perch on the side of shrubs and trees (com-
monly 1-2 m above ground on the shrub). They do this on hill-
tops (from about 1345 to dusk in Napa County, from about
900-1000 to dusk but mainly from 1230 onward in Contra Costa
Co. Calif. (Mt. Diablo)), and they perch also on the side of
tall shrubs on or near the larval host {Cercocarpus betuloides)
in a valley bottom (late afternoon to dusk, Napa Co,), More
work is needed on the time of day of mating; there seems to
be a clear preference for late afternoon to dusk (b).

S. saepitim. Males perch all day about 1 m above ground on
small shrubs or halfway up larger ones on ridgetops and hill-
tops (Jefferson Co., Colorado, c; Contra Costa Co., Calif., c).
In Colusa Co. Calif, males perched all day on the side of shrubs
on the side facing the hillside, in addition to hilltops (c).

14(1):1~40, 1975

MATE LOCATING

17

*S. hehrii. Males perch all day on top of shrubs or small
trees on hilltops (b).

S. fulginosum. Males usually sit on shrubs including the
larval hostplant lupines, and every few minutes patrol erratic-
ally about the canopy of the lupines. They do this usually about
shrubs in the lee of a ridgetop from at least 1130 to 1600
(Tuolumne and Alpine Cos. Calif., c).

"^Callophrys spinetorum. Males perch all day on top of
prominent trees, especially on hilltops (Gilpin Co. Colo., Ari-
zona; c).

"^Callophnjs nelsoni siva. Males perch all day on top and
sides of junipers, especially those on a ridge (often sloping)
rather than those in a valley (Colorado, b). Behavior of C. n.
nelsoni in El Dorado Co. Calif, is very similar; courtship some-
times occurs on flowers.

"^Callophrtjs augustiniis. Males perch all day near the larval
host in small open spots on ridges on the ground or larval host
on the sunny side of the clearing, or on the uphill side of
bushes where the ridgetop slopes downward from the crest
(Douglas Co. Colo., b). In Nevada and Siskiyou and Alpine
Cos. Calif, behavior is similar, males perch on top of shrubs on
slopes or at the top of a slope, starting at 845 or earlier and
continuing through the day (b). Powell’s (1968) observations
are similar although he did not observe perching in early morn-
ing probably due to unsuitable weather.

*C. polios. Males perch all day in swales and narrow valley
bottoms, in small clearings and on trails near the ground there,
on or near the larval host (b).

*C. mossii. Males in Jefferson & Boulder Cos. Colorado
perch all day in gullies, in fairly open level sunny spots next to
a sunlit bank of the gully (b). In San Mateo Co. Calif., males
perch in a similar situation, on shrubs where a grassy hillside
meets on a shrub belt extending to the valley bottom (c). In
Nevada and Siskiyou Cos. California, males perch all day on
top of shrubs sometimes at the upper part of a shrubless sloping
swale or on prominent shrubs on a slope below an open space,
but more often perch on shrubs at the top edge of a precipice
(b).

*C. eryphon. Males perch all day on the side of small trees
(usually Finns ponderosa in Colorado), about 2 m above the
ground, almost exclusively in narrow valley bottoms and gulch
bottoms (Jefferson County Colorado, b; El Dorado, and Siski-

18

JAMES A. SCOTT

J. Res. Lepid.

you Cos. Calif., c).

Callophnjs apama. Males perch all day in gullies on grass
stems and other objects there (a).

C. diimetorum (Napa and Marin Cos. Calif., b) and C.
dumetorum viridis (San Mateo Co. Calif., b). Males perch (and
occasionally patrol) all day on small shrubs or other objects
near the larval host, frequently on small or large hilltops.

C. affinis. Males perch all day on small shrubs (mainly
Artemisia tridentata) on ridgetops and hilltops (Pitkin Co. Colo-
rado; Lander Co. Nevada; Sweet Grass Co. Montana; c).

C. sheridani. Males perch all day near the larval host, in
depressions on grassy hillsides, or occasionally in shallow road-
side depressions, in Colorado (b) and southern New Mexico
(c). In El Dorado and Alpine Cos. California (C. 5. lemberti),
males perch all day near the larval host in sheltered hollows,
on shrubs or rocks or the ground (b). In Churchill Co. Nevada,
C. s. lemheHi-comstocki perched on shrubs or the ground in a
gulch bottom next to a SW-facing slope (c).

Erora laeta quaderna. Males perch on top of trees on hill-
tops, at least in afternoon when observations were made (New
Mexico, Arizona; c).

Phaeostrymon alcestis. Alales patrol about the canopy of
Sapindus dnmwiondi trees, the larval host, and often patrol
during cloudy, windy, or rainy weather (as does Hypaurotis
crysaliis which has an almost identical mate-locating system.)
Patrolling occurs from about 1400 to about 1800. In the morning
males are mostly at flowers and mostly only females are found
on the larval host trees; during the mating period both sexes
occur on larval host trees (Baca Co. Colorado, b).

Hahrodais grumis. Behavior is very similar to H. crysalus.
Males start to patrol about Querctis chrysolepsis about 1400 and
then patrol near the leaves until dusk. They patrol most fre-
quently on the shady side of a group of trees. (Mt. Diablo,
Contra Costa Co. Calif., c).

"^Lycaena arota. Males perch on branches of shrubs and
trees 1-2 m above ground in small clearings in many different
topographic situations, usually valley bottoms. A clearing likely
to have a perching male is about 3-5 m in diameter, reasonably
level, and surrounded by tall trees or steep hillsides. When
abundant, males may perch at the side of large gulches where
a few trees approximate a clearing. Males perch from about
0700 to as late as 1230. In El Paso County, Colorado, males

14(l)tl-40, 1975

MATE LOCATING

19

mostly stopped perching by about 1130, but in Chaffee County
males stopped by about 1230 (part of this difference may be
due to the sun rising earlier in El Paso than in Chaffee County )
(a; Scott, 1974b).

*L. cupreus. Males perch and often patrol all day in hollows
of rockslides near the larval host (Clear Creek & Custer Cos.
Colorado, c). In El Dorado Co. Calif., males patrol and some-
times perch all day mostly in rocky depressions (c).

*L. heteronea. Males patrol all day, like the Plebejini they
resemble, near the larval host {Eriogomim iimbellatum in the
Colorado Front Range) on hillsides etc. regardless of topogra-
phy (b).

*L. xanthoides. Males perch and occasionally patrol, all day
on 1 m or less vegetation beside trails through fields or beside
streams, and often court on flowers in afternoon (b, Jefferson
Co. Colorado and Contra Costa Co. Calif.; Scott & Opler, 1975).

L. editha. Males perch all day in shallow narrow gullies and
in shallow depressions in meadows, usually on low vegetation
(Moffat Co. Colorado, and California, c).

*L. rubidus. Males perch all day on vegetation in dry gullies
and along streams, or along dirt trails through fields when high
density fills more suitable sites (Colorado, b). The same be-
havior occurs in afternoon in Mono Co. California (c).

*L. thoe. Males perch all day on vegetation 1 m or less
above ground along streams and on prominent vegetation in or
bordering a meadow where the larval host is abundant (b).

*L. helloides. Males perch and sometimes patrol all day in
depressions of wet meadows and especially along streams,
usually on m tall vegetation. Males sometimes patrol espe-
cially in the montane meadows of Colorado, and mate-locating
behavior occurs in wetter habitats than in L. rubidus (b). In
Lander Co. Nevada males patrol and perch around Polygonum
plants at least in afternoon (c).

L. nivalis. Males perch all day in shallow usually bare de-
pressions, such as a shallow ( Vz m deep, 2 m broad) gully, and
shallow depressions in an abandoned slightly inclined dirt road
usually next to the valley bottom; the perching sites are hard
to describe verbally, but males remain there often for most of
a day (Gilpin Co. Colorado; Mono, El Dorado, and Siskiyou
Cos. California; b).

L. gorgon. Males patrol and perch, all day on low vegeta-
tion near the larval hosts (California) (c).

20

JAMES A. SCOTT

/. Res. Lepid.

L. mariposa. Males perch in hollows and valley bottoms in
clearings often in dense forest, at least during midday (Yakima
Co. Washington, c) and afternoon (Siskiyou Co. California, c).

"^Hemiargiis isola. Males patrol erratically all day regard-
less of topography, often on flat land and in meadows where a
larval host {Trifolium repens) occurs (c).

Hemiargiis ceraunus. Males patrol erratically all day through-
out the habitat, usually in valleys and on flats (Arizona, Mexico,
c).

Leptotes marina. Males patrol very erratically (with a non-
linear flight) all day throughout the habitat, usually in valley
bottoms where most of the larval foodplants occur (c).

"^Plebejus melissa. Males patrol all day throughout the habi-
tat, mainly near the larval hosts ( b ) .

P. argyrognomon. Males patrol all day near the larval hosts
on hillsides, flats, mainly in bunch-grassland near forest in
Hinsdale Co. Colorado (c) or in moist meadows (Siskiyou Co.
Calif., c).

*P. saepiolus. Males patrol all day near Trifolium, the larval
hosts, mainly in meadows, moist swales, and streamsides where
these plants grow (b).

*P. icarioides. Males patrol all day near Lupinus species,
the larval hosts ( b ) .

*P. acmon. Males patrol all day near the larval hosts, wher-
ever they grow on hillsides and flats, etc. (Colorado, California;

b).

P. lupini. Males patrol all day about the larval foodplant
Eriogonum plants (El Dorado Co., Nevada Co., Tulare Co.
California; b).

*P. shasta. Males patrol all day about 10 cm or less above
the ground near the larval host in alpine cushion plant com-
munities which have about 50% bare ground. These communities
often occur on ridges and saddles and steep slopes ( Colorado, b;
Mono & Tuolumne Cos. Calif., c).

"^Agriades glando7i. Males patrol all day near the larval
hosts {Androsace species) often in valley bottoms.

"^Brephidiiim exilis. Males patrol all day around the larval
hosts, which mainly grow on flats. Males fly weakly among and
a few cm from branches of the. larval hosts (Stockton, Califor-
nia; c).

"^Glaucopsyche lygdamus. Males patrol all day near the larval
hosts in valley bottoms, hillsides, etc. (b).

14(l);l-40, 1975

MATE LOCATING

21

*G. piasus. Males patrol all day near the larval host {Lupinus
argenteus) in valley bottoms or hillsides (b).

Philotes enopteSj hattoides, rita. Males patrol all day
on and between the larval host plants regardless of topography.
Males and females often alight and feed on these plants (b for
each species).

P. spaldingi. Males patrol all day near but not on Eriogo-
num mcemosum (the larval host), patrolling more swiftly and
erratically and slightly higher off the ground than the preceding
three Philotes species. Males patrol in open pinyon-juniper wood-
land on fiats and hillsides (b).

P. sonorensis. Males patrol weakly all day along the base
of cliflFs or sometimes on steep rocky slopes, always near the
larval foodplants (California) (c).

P. speciosa. Males patrol all day about 6" above ground in
swales or gully bottoms near the larval foodplant (Kern Co.
Calif., c).

"^Everes comyntas. Males patrol all day near the ground
near the larval host (Trifoliiim repens in Colorado), mainly in
meadows and streamsides where this plant grows (b).

*£. amyntula. Of all the Plebejini, this species most nearly
approaches a perching species. Males often patrol, but often
perch in depressions such as between a hillside and riparian
shrubs or at the mouth of a tiny gully. Perching males investi-
gate passing individuals, then patrol back and forth before
returning to the vicinity of the previous perch. Mate-locating
behavior occurs all day (Jefferson Co. Colorado; Siskiyou Co.
Calif.; b).

"^Celastrina argiolus. In Napa Co. California males patrol
all day over Aesculus trees throughout the habitat (c). Males
patrol all day about shrubs and low trees, especially in valley
bottoms and seldom on ridges etc. in Colorado (b). In southern
Arizona and western Texas males often patrol about shrubs on
hilltops (c).

HESPERIIDAE

^Epargyreus clarus. Males perch in gullies on vegetation
1-2 m above ground. Perching starts about 0730 depending on
temperature, and ends about 1315; after this period males main-
ly hang upside down from vegetation (b).

Zestusa dorus. Males perch on tips of branches of oak trees
at the edge of a hilltop, at least in afternoon. Males also sit on

22

JAMES A. SCOTT

/. Res. Lepid.

tree limbs overhanging streams where they feed on mud, but I
have not yet seen perching behavior there; further observations
are needed (southern Arizona, c).

"^Thorybes pylades. Males perch all day (few observations
were made in afternoon) in gullies in west Texas and Colorado
(c). In Utah and Arizona (Coconino Co.) and California (Co-
lusa Co.) males perched all day on hilltops (c).

T. diversus. Males perch in tiny (often 3 m in diameter)
forest openings where the presumed legume larval host is com-
mon on flat land and gentle slopes, at least 1330 to 1600 when
observations were made (Tuolumne Co. Calif., b).

*r. mexicana. Males perch all day on stones etc. on hilltops
(Colorado, b; Tuolumne Co. California, c). T. confusis in cen-
tral Texas perches on hilltops at least in afternoon (c) so may
have similar behavior.

"^Erynnis icelus. Males perch and occasionally patrol, all
day in slight depressions and shallow gullies near aspen trees
(Gilpin Co. Colorado, b; Siskiyou Co. California, c).

*£. hrizo. Males perch all day on hilltops, often perching on
the ground, sometimes on small shrubs. Males may fly back and
forth before returning to a perch, which is a pecularity occa-
sionally seen in other Erynnis species also (b).

^E. persius. Males perch all day on hilltops, on the ground
and on low twigs often beside larger shrubs on the hilltop (a).

*E. afraniiis. Males perch all day in gullies and swales, such
as sunlit indentations in gully banks in morning and late after-
noon, small gullies and narrow valley bottoms at midday (b).

*E. maHialis. Males perch all day on hilltops, mainly on the
ground or on low twigs ( b ) .

*E. telemachiis. Males perch all day in gullies, mainly on
vegetation about 1 m above ground there ( Colorado, a ) . In Colo-
rado one copulating pair was found near a hilltop, and in Utah
one male was observed perching on a hilltop.

*E. homtius. Males perch all day on hilltops, usually on
1/3 m twigs, seldom on the ground (Colorado, b). In Texas
males flew about small areas of hilltops before resting again,
whereas Colorado males usually settled more rapidly.

*E. pacuvius. Males perch all day on hilltops on low shrubs
and other objects (in Jefferson and Douglas Co. Colorado, b;
Colusa Co., California, c).

E. tristis. Males perch on hilltops all day, on barb-wire
fence or a twig about 1 m above ground there. Males some-

14(l)-.l-40, 1975

MATE LOCATING

23

times patrol about a few times before resting again (Napa Co.
California, c).

E. propertius. Males perch all day, mainly in clearings on
the sunlit side of ridgetops a few m from the ridgetop, but
sometimes also on banks (such as readouts) in valley bottoms
(California, b). E. p. meridianus males perch on hilltops all
day (Coconino Co. Arizona and western Texas; c).

Autochton cellus. Males perch in gullies on about 2 m tall
vegetation, at least in afternoon when observations were made
(Cochise Co. Arizona and western Texas; c).

"^Pyrgus communis. Males perch, and often patrol, all day,
in swales and valley bottoms and similar situations. Perching
males often patrol about a small area before returning to a
perch. Males perch more when suitable depressions are avail-
able (b).

P. philetas. Males patrol and occasionally perch, in gulches
and valley bottoms, waterholes, etc., all day (southern Arizona;
c).

P. scriptura. Males patrol, and occasionally perch, all day in
low areas of prairie, shallow gullies, and other depressions. Flight
is only about 10-20 cm above ground (b).

^P. xanthus. Males patrol, and often perch especially in
gullies, all day near the larval host at high density, but at the
usual low density males occur in small mostly dry gullies several
meters deep and broad, where they often sit on the bottom or
the sunlit side of the gulch (Colorado, New Mexico; b).

P. ruralis. Males patrol (and occasionally perch) about 10
cm above the ground all day, mainly in valley bottoms or grassy
swales (Siskiyou and El Dorado Cos. California, c).

*P. centaurae. Males patrol and occasionally perch, all day,
in swales, valley bottoms, low spots next to bogs. Males perch
more often at low temperatures, and patrol more often at high
temperatures (c).

Staphylus ceos. Males patrol all day in gulches and valley
bottoms, flying faster than P. catidlus (southern Arizona, west-
ern Texas; c).

"^Pholisora catullus. Males patrol all day just above the
ground in roadside ditches, gulches, dry reservoir bottoms, and
other depressions (b).

*P. mejicanus. Males patrol all day in the same manner and
locations as P. catullus; the two species cannot be distinguished
during flight (c).

24

JAMES A. SCOTT

J. Res. Lepid.

*P. alpheus. Males patrol in gullies and around the larval
host shrubs, apparently all day (few observations were made in
afternoon; c).

P. graciellae. Males patrol in gulches but mainly patrol
through and around the larval host shrubs on flats, at least in
morning and early afternoon (California, c).

P. lihya. Males patrol in gullies and in and about the larval
host shrubs at least in morning and early afternoon (Indio,
California; c).

Systasea evansi. Males perch (they usually fly back and
forth before resting again) in gulches apparently all day (few
observations were made in afternoon) (southern Arizona, west-
ern Texas; c). S. pulveriilenta had very similar behavior when
observed at midday (western Texas; c).

Celotes nesstis. Males patrol all day just above the ground
(flight is rather weak) in gulches and along roadsides and
other depressions. Males rest often so they may perch occasion-
ally (southern Arizona, western Texas; c). C. limpia in western
Texas with a longer distal prong on the male valva has nearly
identical behavior (few observations; c).

CaHerocephalus palaemon. Males perch on grass stalks and
other objects about 1/3 m above ground, and often patrol as
well, all day, in sedge swales in wet valley bottoms (Siskiyou
Co. Calif.; c).

Copaeodes aurantiaca. Males perch all day in gullies on
vegetation about 1 m tall and have a very fast flight ( Coconino
Co. Arizona, southern Arizona and western Texas; b).

*Pinma pirm. Males patrol all day usually in valley bottoms
and gulches; males patrol several cm from the canopy of low
plants averaging 1 m or less above ground, and patrol following
the contours of the vegetation ( b ) .

Ancyloxypha arene. Males patrol weakly all day over grass
at the edges of springs and reservoirs (western Texas, southern
Arizona; b).

A. numitor. Males patrol weakly all day over and among
grasses at the edges of streams and reservoirs and at springs
(Ohio, c; Yuma Co. Colorado, c).

Adopaeoides prittwittzi. Males patrol fairly weakly over
watercress flowers and occasionally over grasses at a spring, at
least in morning when observations were made (southern Ari-
zona, c).

"^Oarisma garita. Males patrol weakly to rapidly all day in

14(l):l-40, 1975

MATE LOCATING

25

tall grass of meadows, valley bottoms, and to a lesser extent on
hillsides (b).

*0. edwardsii. Males patrol often rapidly all day in grassy
openings between shrubs, throughout the habitat but more often
in valley bottoms than on hillsides (c).

^Yvretta rhesus. Males perch all day on small (2 m tall)
prairie mesa tops when these are present, on flat land occasion-
ally (c).

Y. earns. Males perched at least from 1200 to 1400 when
observations were made, on flat bare ground next to the probable
larval host, a short grass growing next to a waterhole, and on
sand bars. Females were mostly on the short grass, and some
individuals fed on mud (Pima Co. Arizona; c).

"^Stinga morrisoni. Males perch all day on hilltops and
ridges, often just below a shrub or between trees on the hilltop
whereas Hesperia pahaska males usually perch in more open
sites (b).

"^Hesperia uncas. Males perch all day on small (2 m tall)
prairie mesa tops or small hilltops, but perch on smaller hill-
tops than most other Hesperia. H. uncas is very similar to Y.
rhesus in behavior and appearance (b).

H. lindseyi. Males usually patrol slowly (about m per
sec.) by fluttering about 15 cm above the grass, and males perch
between these flights, all day, in grassland usually near the top
of a ridge (Napa Co. Calif.; b).

*jFf. comma. Males perch all day in southern Colorado and
central Colorado (b) on hilltops. At high density at some locali-
ties (Chaffee and Jefferson Cos. Colorado, c) males investigate
passing individuals while at mud. In Utah and Nevada and
Mono Co. California, males of H. c. harpalus perch on hilltops
or on flat areas when hilltops are not present, and males court
at flowers (b). In El Dorado Co. California (c) males perched
all day on the outer edge of a road on a steep slope. In Marin
Co. California (c), H. c. dodgei males perched in bunch grass
where the local population occurs and not on adjacent hilltops
where H. Columbia perches.

H. juba. Males perch all day in gullies, rocky depressions,
and valley bottoms, both in Jefferson Co. Colorado (c) and in
El Dorado Co. California (c).

nevada. Males perch all day on small to large hilltops
and ridges (Custer and Park Counties Colorado, b; Tuolumne
Co. California, c).

26

JAMES A. SCOTT

J. Res. Lepid.

viridis. Males perch all day on rocks or bare ground of
gullies and valley bottoms (Colorado, b; Texas, c).

H. Columbia. Males perch all day on ridgetops and hilltops
on low shrubs and rocks (Marin and Napa Counties, Califor-
nia, b).

*//. pahaska. Males perch all day on hilltops and ridgetops
on rocks or short plants (a; Scott, 1973d).

leonardus pawnee and H. 1. montana. Males perch all
day on small to large hilltops, or on flats near Liatris punctata
flowers (b).

H. ottoe. Males perch all day on flowers, especially thistle.
Males almost never occur away from these flowers, and by re-
leasing other individuals near males on flowers it was found
that males investigate other individuals all day. These flowers
occurred in a valley bottom (Boulder Co. Colorado, c).

H. miriamae. Males perch all day on rocks or sometimes on
the ground (especially those rocks that are sunlit and out of
the wind) on alpine hilltops and ridgetops. Males have an ex-
tremely fast flight (Inyo and Mono Co. California, c).

H. woodgatei. Males perch all day on hilltops where they
sit on rocks etc. on or near the ground (southern Arizona, c).

Hijlephila phijleus. Males perch all day on prominent plants
or hedges on lawns, grassy swales, and similar sites (Clark Co.
Nevada, c; Yolo Co., California, c).

Atalopedes campestris. Males perch all day in grassy mead-
ows and swales Fremont Co. Colorado, c; central Texas, c).

Nastra julia. Males perch all day in grassy swales such as
along roads and irrigation ditches (SW of Parker, Arizona; c).

Lerodea eufala. Males perch all day in grassy swales and
flats, along irrigation ditches (SW of Parker, Arizona, c; Yolo
Co. California, c).

L. arahus. Males perch all day in gulches and along streams
near large clumps of the presumed larval host grass. Males
perch on vegetation as much as a meter above ground (Pima
Co. Arizona, c).

Paratrytone melane. Males perch all day in grassy swales,
gullies, lawns (California, c).

"^P elites origenes. Males perch all day in grassy swales and
openings in valley bottoms (b).

*F. themistocles. Males perch all day in grassy swales, de-
pressions in meadows and lawns (b).

P. mystic. Males perch all day in low spots of meadows and

14(1):1~40, 1975

MATE LOCATING

27

grassy streamsides (b).

^P. sonora. Males perch all day in low spots o£ moist grassy
meadows (Park, Chaffee, and Saguache Counties Colorado, c).
In El Dorado and Tulare Counties California, behavior was
similar but at high density courtships were frequent at flowers

(c).

*P. sahuleti. Males perch all day in low spots of alkaline
flats where larval host is abundant, in low spots of meadows,
and on flowers on flat land (Colorado, Clark & Nye Cos. Nevada,
Mono & Inyo Cos. California, Emery & Millard Cos. Utah; a).

*P. draco. Males perch all day in low spots of meadows and
in gullies (b).

P. coras. Males perch all day in grassy swales and low spots
of meadows ( c ) .

Ochlodes sylvanoides. Males perch all day in gullies and
valley bottoms on vegetation about I m above ground (Colo-
rado, b). In Yolo Co. California, males perch in nooks among
shrubs, often on ridgetops (c).

O. agricola. Males perch mainly in the shade of shrubs
about 1/3 m above ground at the edges of clearings throughout
the habitat, and males often court at flowers all day.

O. yuma. Males perch all day in low spots near the larval
host, especially in depressions between this host (Phragmites)
growing at a river edge and the bank. Males perch on large
rocks or I m tall vegetation ( California, c; Mesa Co. California,
b).

*0. snowi. Males perch all day in gullies on rocks or I m
vegetation, and some courtships occur at flowers (a; Scott,
1973d).

"^Poanes taxiles. Males perch all day in gullies and valley
bottoms, especially in shaded areas, on vegetation about a
meter or more high. Males perch in more shaded locations and
in cloudier weather than does Poanes hohomok (a).

*P. hobomok. Males perch all day on vegetation about 2 m
above the bottom of gullies and valley bottoms (b).

"^Atrytone delaware. Males perch all day in grassy swales,
mainly in valley bottoms (b).

A. arogos. The main activity of both sexes is feeding on
flowers. However, feeding males do not investigate individuals
released nearby (contrasting with H. ottoe). After many fail-
ures in observing interactions at flowers I finally observed males
perching on low (about 10 cm) vegetation on a gently sloping
base of a hillside covered with Andropogon, the larval host.

28

JAMES A. SCOTT

/. Res. Lepid.

A few males perehed in a grassy swale below a similar slope. In
three days observations perching started about 1320 and con-
tinued until late afternoon until rain or clouds stopped activity.
This mating period is strange because it occurred during cloudy
and sometimes rainy weather; perching males were often seen
to rapidly vibrate their wings during cloudy periods, apparently
as a heat gain device (shown to raise body temperature by
Krogh & Zeuthen, 1941; Kammer, 1970). Additional observations
are desirable to confirm this mating period (c).

"^Euphyes vestris. Males perch in gullies all day on about
1 m vegetation ( Colorado, a; in California males perch in swales
in coastal meadows, c).

E. himaciila. Males perch all day in low spots of moist
sedge marshes, on about 1 m stalks of clumps slightly taller than
surrounding vegetation (Yuma Co. Colorado, c).

Atrytonopsis hianna. Males perch all day on or near the
ground on 2-4 m wide relatively flat clearings of valley bottoms,
and occasionally in similar situations on slopes near the larval
host (c).

*A. vierecki. Males perch all day in gulches, usually on 1 m
tall vegetation (Colorado, c; Cimarron Co. Oklahoma, c).

A. cestiis and A. ovinia. In two days observations in Pima
Co. Arizona, males perched on rocks on south-facing sides of
gullies, especially gullies 3-7 m high next to hillsides and
flowers. Males perched from early morning to about 1200, and
in afternoon fed on flowers and rested (without interacting)
on gully walls (c for each).

A. pittacus. Males perch all day, almost always in gullies in
southern Arizona and western Texas (c), although near Alpine,
Texas males also perched in a depression on the sloping leeward
side of a hilltop about 3 m from the top ( c ) .

AmblysciHes simiiis. Males perch on top of small prairie
plateaus, ridgetops and hilltops, from about 0740 to about 1050,
with maximum perching from 0830 to 0930 (a; Scott, 1973c).

^A. aenus. Males perch all day in gully bottoms, especially
on rocks in rocky areas of the gully (b).

*A. vialis. Males perch all day in narrow valley bottoms
with much vegetation, usually perching on 1 m vegetation (b).

*A. oslari. Males perch all day in gully bottoms and road-
side ditches, especially in sandy spots on the gully bottom (b).

*A. eos. Males perch all day in gullies and valley bottoms, on
flat usually grassy spots on or just beside the gully bottom (c).

14(l);l-40, 1975

MATE LOCATING

29

^A. phylace. Males perch all day in mostly bare depres-
sions, such as a depression at the mouth of a small gully open-
ing onto a flat, a roadside cut below a small gully, etc. The
perching locations change throughout the day as males choose
sunlit locations (b).

A. nysa. Males perch all day near or on the bottom of nar-
row gullies (Texas, c).

A. texanae. Males perch all day in gullies on rocks or other
objects (western Texas, c).

Agathymus mariae. Males perch in morning near the larval
host (western Texas, c).

A. remingtoni estelleae. Males perch in morning near the
larval host, especially at the base of a slope (western Texas, c).

^ Megathymus streckeri. Males perch on hillsides and flats
near the larval host. My observations were only in morning, but
Scott Ellis (written communication) has observed mating at
1300 in M. s. leussleri and much chasing between males in late
afternoon in M. 5. streckeri in southern Colorado (c). Mr. Kilian
Roever, however, states (pers. comm.) that male activity in
M. s. streckeri and M. yuccae ceases after midday.

*M. yuccae. Males perch near the larval host at least in
morning (c). Scott Ellis (pers. comm.) has observed strong
perching behavior from about 0900-1100, “but the males seem
to disappear in the afternoon.’’

DISCUSSION

Before making generalizations, I will mention some of the
most unusual species. Some species both perch and patrol fre-
quently (most perching species patrol infrequently, and many
patrolling species sometimes perch). The best examples are
most Oeneis and Pyrgus species; others are the Papilio machaon
group (P. zelicaon, bairdii, polyxenes) , P. eurymedon, Limenitis
wiedemeyeri, Nymphalis antiopa, N. californica, Phyciodes
mylitta, P. pallida, P. texana, Euphydryas spp., Pohdryas minuta,
Chlosyne palla, C. acastus, C. gorgone, C. lacinia, C. leanira,
C. theona, Calephelis nemesis, Lycaena cupreus, and Everes
amyntula. P. minuta differs from the others in that males mainly
perch in the morning, but mainly patrol in the afternoon.
Speyeria callippe is unusual in that the location of patrolling
changes somewhat during the day. In several patrolling species,
males seem to patrol about the tops of shrubs late in the day,
apparently seeking roosting females {Boloria titania, Plebejus
melissa ) .

30

JAMES A. SCOTT

J. Res. Lepid.

The study of mate-locating behavior provides numerous use-
ful taxonomic characters. For instance, in Amblyscirtes the three
species groups previously recognized on morphological grounds
also differ in mate-locating behavior. A. simius differs from other
Amblyscirtes by mating only part of the day and on ridgetops
but not in gullies; A. phylace differs by mating at the mouth of
small gulches and in roadside ditches (the other Amblyscirtes
mate in deep gullies). Perching or patrolling behavior and the
time of day of mating seem to be rather conservative characters
that are useful in comparing genera or higher taxa; the location
of mating often differs between congeneric species and is very
useful for intrageneric classification.

Location of mating can be placed into two general cate-
gories: 1) mating primarily occurs near the larval foodplants;
2) alternatively, mating occurs in specific topographic sites
regardless of the distribution of the larval foodplants. If the
larval foodplants are generally distributed within the flight area
of the population regardless of topography (such as throughout
a bog in a bog species, or throughout the habitat) patrolling
behavior seems to be the usual mate-locating method, but often
mating occurs in specific topographic sites such as gullies or
hilltops (such as some Hesperia species that feed as larvae on
widespread grasses). If larval foodplants are spotty in distribu-
tion within the normal flight area of a population (such as Apo~
clemia nais which feeds on Ceanothus fendleri) perching be-
havior seems to be the usual strategy. If the foodplants are
linearly distributed ( such as streamside willows ) , in many species
males patrol along the stream {Limenitis achipptis), and in some
species males perch there (Safyrium sylvimis).

The strategy of mating in topographically distinct sites is
often used for bringing the sexes together in species in which the
larval foodplants occur throughout the habitat. Often several
types of sites appear adequate to this function: closely related
species may mate in different types of sites.

I found that eighteen congeneric pairs of butterflies rendez-
vous for mating in contrasting topographic sites. These eighteen
pairs are listed below. The first species of the pair mates on
hilltops, the second species in gulches, valley bottoms, or, for
P. indra, rocky outcrops just below a hilltop. The foodplants of
the species as larvae are listed. Localities studied are listed.
1) Papilio eurymedon (Ceanothus fendleri; Red Rocks, Jeffer-
son Co. Colorado ) and P. multicaudata ( Prunus virginiana;

14(1);1~40, 1975

MATE LOCATING

31

Red Rocks, and Jarre Canyon, Douglas Co. Colorado, Hard-
scrabble Canyon, Custer Co. Colorado). 2) P. zelicaon {Har-
bouria trachypleiira; Red Rocks ) and P. indra ( H. trachypleura;
Red Rocks). 3) Etichloe olympia (various Cruciferae unde-
termined for Colorado; Red Rocks, Jarre Canyon, and the
Arkansas River Canyon, Fremont Co. Colorado and NW of
Pueblo, Colorado) and E. ausonides (various Cruciferae; Red
Rocks, Jarre Canyon, Hardscrabble Canyon). 4) Pieris sisymbri
(various Cruciferae undetermined for Colorado; Jarre Canyon,
Arkansas River Canyon, and NW of Pueblo) and P. chlorodice
{=beckeri) (Arkansas River Canyon, NW Pueblo; various Cru-
ciferae such as Stanleya pinnata). 5) Oeneis melissa (unknown
monocotyledons; Loveland Pass and Mt. Evans, Clear Creek Co.
Colorado) and O. polixenes (same foodplants and localities).
6) O. chryxus (unknown grasses; Jarre Canyon, and Gregory
Canyon, Boulder Co. Colorado, and Greenhorn Peak, Huerfano
Go. Colorado, and Rosita, Custer Co. Colorado) and O. uhleri
(unknown grasses; same localities except for Rosita). 7)
Speyeria callippe (Viola; Red Rocks) and S. atlantis (same
foodplants and locality, Arkansas River Canyon). 8) Chlosyne
gorgone (Helianthiis pumiliis; Red Rocks, Hardscrabble Canyon,
and Green Mountain, Jefferson Co. Colorado) and C. nycteis
(Rudbeckia laciniata; same localities except for Green Moun-
tain). 9) Nymphalis calif ornica (Ceanothiis fendleri; Jarre
Ganyon) and N. antiopa (Salix, Populiis, Ulmiis, Celtis; Red
Rocks, Jarre Canyon, Gregory, Canyon, Hardscrabble Canyon).
10) Callophrys augiistimis (Arctostaphylos tiva-iirsi; Russel
Ridge, Douglas Co. Colorado) and C. polios (A. uva-ursi; Greg-
ory Ganyon, Russel Ridge). 11) Satyriiim calif ornica (Ceano-
thus fendleri; Red Rocks) and S. acadica (Salix exigua; Ganon
City, Fremont Co. Colorado). 12) Thorybes mexicana (various
Leguminosae undetermined for Colorado; Red Rocks, Green-
horn Peak) and T. pylades (same foodplants; Red Rocks).
13) Erynnis brizo (Querctis gambellii; Red Rocks, Jarre Gan-
yon) and E. icelus (Popultis tremuloides; Gentral City, Gilpin
Co. Colorado). 14) Erynnis persius (Astragalus, Thermopsis,
and other legumes; Red Rocks, Jarre Canyon, Rosita, Arkansas
River Canyon) and E. afranius (Astragalus, Lupinus, Lotus,
and other legumes; Red Rocks, Green Mountain, and Wetmore,
Custer Co. Colorado. 15) Erynnis horatius (Quercus gambellii;
Red Rocks, Jarre Canyon, Oak Creek Canyon S. of Canon City,
Colorado) and E. telemachus (same foodplant; same localities

32

JAMES A. SCOTT

/. Res. Lepicl.

and Hardscrabble Canyon). 16) Hesperia pahaska (Bouteloiia
gracilis; Red Roeks, Green Mountain, Arkansas River Canyon,
Platte River Canyon, Douglas Co. Colorado) and H. viridis
{B. gracilis; same localities except Platte River Canyon, and
Pueblo, Colorado, and Black Mesa, Kenton Co. Oklahoma).
17) Hesperia comma (various grasses; Red Rocks, Arkansas
River Canyon, Rosita, Platte River Canyon) and H. juha (vari-
ous grasses undetermined in Colorado; Red Rocks, Green Moun-
tain). 18) Amblyscirtes simitis (B. gracilis; Arkansas River Can-
yon, and near Saguaehe, Saguache Co. Colorado) and A. oslari
(grasses undetermined in Colorado; Red Rocks, Arkansas River
Canyon, Gregory Canyon, Hardscrabble Canyon).

The use of separate mating sites by these congeneric species
certainly prevents, and may result from, interference competi-
tion during mate-locating behavior. Interference between butter-
flies during mate-locating behavior arises from the generalized
nature of the stimuli involved in visual communication between
the sexes (Scott, 1973a, 1974a). Mate-locating behavior is a
process of screening flying or resting objects for receptive fe-
males. Perception of the insect compound eye is poor for shapes,
but good for movement. The size and color pattern of the two
species of a pair is very similar in most cases. Males probably
cannot distinguish color pattern differences as small as those
between most of the congeners. Male butterflies sometimes
investigate resting individuals, and often investigate aerial
objects of a great variety of shapes and colors including other
insects, birds, etc. Pheromones which might enable interspecific
males to avoid interaction are currently unknown. Both perch-
ing and patrolling species appear to delineate a space around
them in which moving and occasionally resting objects are
investigated (‘'moveable individual space”). Two congeneric
males do not occur in the same space for long because investi-
gative interactions occur after which at most one male remains.
Intense con- and interspecific interactions occur where males
are concentrated. These interactions result in interference com-
petition for time, space, and energy; time is lost (in which a
passing female might be missed), the probability of being in
a favorable mating arena is less, energy is wasted, and court-
ships with receptive females are often disrupted (another male
often investigates a courting pair and the two males then often
interact, ending courtship).

In some of these congeneric pairs interference competition

14(l);l-40, 1975

MATE LOCATING

33

itself may have resulted in the evolution of separate mating sites.
If t'wo species have the same mating sites and if interference
competition between them is great, when a mutation for mating
at another site arises in one species (especially the rarer spe-
cies) mating may be more successful at the second site and the
proportion of individuals of that species mating there will
gradually increase, ultimately resulting in genetic fixation. As
selection proceeds, as the proportion of the selected species
mating at the first site drops the selection against those mating
at the first site will intensify because an increasing proportion
of individuals encountered there will be nonconspecific. Rapid
fixation of mating at the second site for the one species will
then occur due to the continual increase of selection intensity
against mating at the first site. This mechanism would be most
rapid in two species which are postzygotically, but not pre-
zygotically, reproductively isolated, which could occur if post-
zygotic but not prezygotic reproductive isolation developed in
two allopatric populations which then became sympatric.

Most butterfly species mate at any time of day. Restricted
time of day of mating occurs often in some taxa (Nymphalini,
Riodininae, Theclini, Megathyminae), rarely in other taxa
(Satyrinae, Apaturini, Lycaenini, Pyrginae, Hesperiinae), and
with my current knowledge not at all in many taxa (Papilioni-
dae, Pieridae, Argynnini, Melitaeini, Limenitini, Charaxinae,
Danainae, Plebejini).

There is an altitudinal trend in the time of day of mating.
Of 284 species for which mate-locating behavior is reported
herein, only 35 mate during a restricted part of the day (in-
cluded are five species, Vanessa spp. and Asterocampa celtis,
which seem to mate more frequently late in the day). All of
these 35 species are primarily desert, foothills, or plains species
rather than high mountain species. The primarily high moun-
tain species all mate at any time of day.

Inclement weather is frequent in spring and fall. It is sig-
nificant that only two species that mate during only part of
the day are univoltine and mate in the spring {Megathymus
yucca, Polygonia faunus). In P. faunus long life of the hiber-
nating adults allows them to wait for good weather. The other
species are univoltine and emerge in summer, or are multivoltine.

Mate-locating is an energy-demanding process, and greatly
exposes males to predation. Perhaps more dependable weather
in the foothills and plains compared with the alpine zone, and

34

JAMES A. SCOTT

J. Res. Lepid.

in summer compared with spring, allows some species to reduce
the hours devoted to mating ( and sometimes reduced adult feed-
ing) without seriously decreasing the proportion of females
mating. At high altitude and in early spring, however, inclement
weather requires that animals make use of all available periods
of suitable weather, no matter what time of day these occur.
Time of day of mating seems to have evolved as a compromise
between energy (and predation?) losses and inseminating the
maximum number of females.

Those species which mate only in early morning or in after-
noon and early evening mate on sunlit areas such as treetops
and hilltops; species mating at other times can mate in gullies
and other locations that do not receive early and late sunlight.

Scott and Scott (1976) present an ecological analysis of the
butterflies of an area of southern Colorado. I determined mate-
locating behavior of the species in that area, and the following
analysis attempts to determine how the behavioral and ecologi-
cal characteristics of a species influence or are influenced by its
mate-locating behavior.

To quantify mate-locating behavior, the following three
qualitative indices were constructed: a) perching-patrolling in-
dex (1-males almost always perch prior to encounters; 4-males
almost always patrol prior to encounters; 2 and 3 intermediate);
b) location of mating index (1-mate in quite restricted sites of
the habitat such as hilltops; 4-mate almost anywhere in the habi-
tat; 2 and 3 intermediate ) ; c ) time of day of mating index ( dur-
ation of the daily mating period of a species in hours, divided
by 2; a mating period of “all day” is assumed to be 8 hours in
duration ) .

Perching species in southern Colorado have a strong tendency
to mate in restricted sites in the habitat, as predicted by Scott
(1974a). The evidence is that the correlation (product-moment)
between the perching-patrolling index and the location of
mating index for the 159 species is .54 (p < .01), which means
that perching species tend to mate in restricted sites and patrol-
ling species tend to mate more generally in the habitat.

Perching species in southern Colorado also tend to mate
only during restricted times of day rather than at any time of
the day. The correlation between the perching-patrolling index
and the time of day of mating index is .34 (p < .01).

Perching behavior in southern Colorado is much more com-
mon in species that have only 1 or 2 broods during the season

14(l):l-40, 1975

MATE LOCATING

35

than in species with 3-4 broods (Table 1).

No consistent changes of mate-locating behavior with alti-
tude were observed for the southern Colorado butterflies, ex-
cept that above timberline all the species mate all day, and
almost all species patrol there.

Southern Colorado butterflies that emerge at the end of the
season tend to have patrolling behavior and tend to mate
throughout the habitat much more frequently than species that
emerge at the start of the season (Table 2).

The 159 species were placed into five categories depending
on the meagre (usually) to extensive data I have concerning
their movements (Scott, 1976). The only trends found (Table
3) are that species with very large movements more often tend
to be patrolling species and to mate throughout the habitat.

No consistant relationship between the frequency with which
a southern Colorado species feeds on flowers of the larval host
and mate-locating behavior was observed.

Larval foodplants of the southern Colorado butterflies were
given by Scott and Scott (1976). Tables 4 and 5 relate mate-
locating behavior to the larval foodplants. Species feeding on
shrubs or trees rather than on herbs more often tend to be
perching species and more often tend to mate during a restricted
period during the day.

I now attempt to determine the effect of the distribution of
larval foodplants on mate-locating behavior in a different way,
by examining the mate-locating behavior of various western
U. S. butterflies that have the same larval foodplants, to dis-
cover whether convergent evolution occurs. The following com-
parisons were made.

1) Ceanothus fendleri (plants spottily but sometimes gen-
erally distributed on south-facing hillsides and dry valley bot-
toms). Mating generally occurs in hilltops or gulches: Erynnis
pacuvius, E. martialis, Papilio eunjmedon, Satyrium saepitim,
S. californica, and often Nymphalis californica mate on hilltops
( N. californica sometimes mates in gulches etc. ) , and Apodemia
nais mates in low spots in gulches near Ceanothus.

2) Eriogonum umhellatum (generally distributed on steep
hillsides, mesas, and other locations). Two strategies occur.
Plebejus acmon, Philotes enoptes, and Lycaena heteronea patrol
regardless of topography ( L. heteronea is the only Lycaena with
mainly patrolling behavior, and has apparently convergently
evolved patrolling behavior). Callophrys sheridani, C. affinis,

36

JAMES A. SCOTT

J. Res. Lepicl.

and C. apama (this species probably feeds on E. iimbellatum,
but no proof exists) mate in hilltops, gullies, or hillside de-
pressions depending on the species.

3) Salix exigua (linearly clumped distribution along streams
and irrigation ditches). Two strategies occur. Satyrium sylvinus
and S. acadica perch on the plant, while Nymphalis antiopa and
Limenitis tveidemeyeri perch in valley bottoms near willows.
Papilio rutulus and L. archippiis patrol near the willows. The
diflFerence in strategy of the two Limenitis may be due to their
different habitats: L. archippiis occurs on the plains where S.
exigua is more continuously distributed, whereas L. iceidemeyeri
occurs mainly in the foothills (and larvae feed on other Sali-
caceae as well) where the foodplants are more spotty and
there are more varied topographic sites.

4) Qiierciis gamhellii (generally distributed, sometimes in
groves). In Hypaiirotis crysaliis, in which all activities are car-
ried out on the plant (Scott, 1974), males patrol. Satyrium
calanus, S. liparops (which may not feed on this oak species
in Colorado), Erynnis telemachus, E. hrizo, and E. horatius mate
in gulches or hilltops.

5) Primus virginiana (usually in small groves, mainly in valley
bottoms). Papilio multicaudata patrols up and down valley bot-
toms, but Harkenclenus titus perches on hilltops.

6) Sedum lanceolatum (generally distributed on hillsides,
etc.). Parnassius phoehus patrols on hillsides and meadows near
the plants, but Callophrys mossii perches in gullies and is a
foothills species whereas P. phoehus usually occurs in grass-
lands at higher altitude.

7) Pinus species (generally distributed). Neophasia menapia
patrols throughout the habitat, but Callophrys eryphon perches
in gulch bottoms.

8) Ribes species (throughout the habitat, but more often in
valley bottoms). Polygonia zephyrus and Lycaena arota both
perch mainly in valley bottoms.

9) Celtis reticulata (in groves on flats and in valley bot-
toms). Asterocampa celtis, Nymphalis antiopa, and Polygonia
interrogationis all perch in valley bottoms on or near the Celtis
plants.

10) Cirsitim spp. (clumped weeds often on disturbed areas).
Vanessa cardui perches often on hilltops (and feeds on many

14(1):1~40, 1975

MATE LOCATING

37

Other plants as larvae), whereas Phyciodes pallida, P. mylitta,
and P. or sets mainly perch in gullies.

11) Sphaeralcea coccinea (widely distributed, mainly on
flats). Vanessa caryae perches often on hilltops, but Pyrgus
scriptura and P. communis patrol and perch in swales.

12) Atriplex spp. (clumped in alkaline areas usually on flats).
Brephidium exilis, and Pholisora alpheus and its congeners,
patrol near Atriplex, or patrol in gullies (Pholisora) .

13) Astragalus spp. (mostly generally distributed in open
areas). Most species patrol (Colias eurytheme, C. philodice, C.
alexandra, Hemiargus isola, Leptotes marina, Everes amyntula,
Plebejus acmon), but Erynnis persius perches on ridgetops.

14) Lupinus argenteus (clumped, often in valley bottoms
on deep soil). Plebejus icarioides, Glaucopsyche lygdamus, and

G. piasus patrol, but Erynnis afranius perches in gullies.

15) Trifolium (usually clumped in moist meadows except
near timberline). Colias cesonia, C. meadii, Plebejus saepiolus,
and Everes comyntas patrol, but Thorybes pylades, which has
other hosts, perches in gulches.

16) Bouteloua gracilis (very widely distributed). Neominois
ridingsii, Amblyscirtes simius, Yvretta rhesus, Hesperia pahaska,

H. viridis, H. tineas, and H. comma all mate on hilltops or
gulches.

It is clear from the above mate-locating behavior of species
with the same larval foodplants that for a given situation there
is usually more than one strategy of locating females. Behavior
is likely to be similar in taxonomically closely related species.
Distantly related species often have different behavior, but
sometimes they have convergently evolved the same behavior.
Perhaps the most spectacular examples of convergence are
Philotes and Lycaena heteronea, and Neominois ridingsii and
Amblyscirtes simius.

ACKNOWLEDGEMENTS

1 thank Glenn R. Scott, Scott L. Ellis, Ray E. Stanford,
Oakley Shields, Mike Fisher, John F. Emmel, Michael Young,
Andre Meulenbrouk, Maurice Howard, Donald Eff, Fred
Urquhart, Betsy Webb, Kilian Roever, and Arthur M. Shapiro
for providing helpful information.

38

JAMES A. SCOTT

J. Res. Lepid.

LITERATURE CITED

GUPPY, R. 1969. Further observations on ‘Tilltopping” in Papilio zeli-
caon. J. Res. Lepid. 8: 105-117.

KAMMER, A. E. 1970. Thoracic temperature, shivering, and flight in the
monarch butterfly, Danaus plexippus (L). Z. Verd. Phusiol. 68: 334-
344.

KROGH, A., and E. ZEUTHERN. 1941. The mechanism of flight prepar-
ation in some insects. J. Exp. Biol. 18: 1-10.

LABINE, P. A. 1966. The reproductive biology of the checkerspot butter-
fly, Euplujdnjas editha. PhD thesis, Stanford University.

POWELL, J. A. 1968. A study of area occupation and mating behavior in
Incisalia irioides ( Lepidoptera, Lycaenidae). J. New York Ent. Soc.
76: 47-57.

SCOTT, J. A. 1973a. Mating of butterflies. /. Res. Lepid. 11: 99-127.

. 1973b. Population biology and adult behavior of the circumpolar

butterfly, Parnassiiis phoehiis F. (Papilionidae). Ent. Scand. 4: 161-
168.

. 1973c. Convergence of population biology and adult behavior in

two sympatric butterflies, Neominois ridingsii ( Papilionoidea: Nym-
phalidae) and Amhlyscirtes simiiis ( Hesperioidea: Hesperiidae). J.
Anim. Ecol. 42: 663-672.

. 1973d. Adult behavior and i^opulation biology of two skippers

(Hesperiidae) mating in contrasting topographic sites. J. Res. Lepid.
12: 181-196.

. 1974a. Mate-locating behavior of butterflies. Anier. Midi. Nat. 91:

103-117.

. 1974b. Population biology and adult behavior of Lycaena arota.

(Lycaenidae). J. Lepid. Soc. 28: 64-72.

. 1974c. Adult behavior and population biology of Poladryas minuta,

and tlie relationship of the Texas and Colorado populations (Lepidop-
tera: Nymphalidae). Pan-Pacific Ent. 50: 9-22.

. 1974d. The interaction of behavior, population biology, and environ-
ment in Hypaurotis crysalus (Lepidoptera). Anier. Midi. Nat. 91: 383-
394.

. 1975a. Movements of Euchloe ausonides (Pieridae). J. Lepid. Soc.

29: 24-31.

. 1975b. Movements of Precis coenia, a “pseudoterritorial” sub-
migrant (Lepidoptera, Nymphalidae). J. Anim. Ecol. (in press).

. 1976. Movement patterns among eleven butterfly species; inverse

correlation between density and movement. Ecology, in press.

SCOTT, J. A., & P. A. OPLER. 1975. Population biology and adult behavior
of Lycaena xanthoides (Lycaenidae). J. Lepid. Soc. 29: 63-66.

SCOTT, J. A., & G. R. SCOTT. 1976. Niche analysis and distribution of
southern Colorado butterflies. ( under review ) .

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6: 69-178.

SHIELDS, O., & J. F. EMMEL. 1973. A review of carrying pair behavior
and mating time in butterflies. J. Res. Lepid. 12: 25-64.

14(1): 1-40, 1975

MATE LOCATING

39

Table 1. Change of behavioral characteristics with number of broods.
Brephidhmi exilis, with 5 broods, is placed with the 4-brood group. Figures
are averages. Means and standard deviations for all the variables for all 159
species are also given.

Standard

No. of broods

1

Number of Broods
2 3

4

Mean

1.59

Deviation

.98

No. of species

108

22

17

12

159

159

Perching-patrolling

2.32

1.93

2.82

3.58

2.41

1.41

Index

Location of Mating

2.28

1.87

2.29

2.83

2.27

.88

Index

Time of Day of

3.76

3.42

3.85

3.83

3.73

.67

Mating Index

Table 2. Change of behavioral characteristics throughout the season for
single brood species only. Figures are averages.

Time of Emergence

No. of species
Perching-patrolling
Index

Location of

Mating Index
Time of Day of
Mating Index

early

mid

late

early

mid

late

August-

May

June

June

June

July

July

July

early

Sept

12

12

13

10

19

14

16

11

1.83

1.92

1.85

1.60

2.74

2.14

2.87

3.18

1.92

1.92

2.23

1.80

2.42

2.14

2.75

2.86

3.55

4.00

4.00

3.63

3.83

3.66

3.46

4.00

Table 3. Means of behavioral characteristics for species with different magnitude
of movements.

No. of species
Perching-patrolling
Index

Location of Mating
Index

Time of Day of
Mating Index

Magnitude of Movements

Neither

Apparently Can Move

Migrate

Very Local-

Migratory

Can Move

Several

Thousands of

ized Species

or very
Local

Several

Kilometers

Kilometers

Kilometers

24

102

16

15

2

2.29

2.20

2.62

3.80

2.50

2.04

2.18

2.12

3.40

2.00

3.35

3.84

3.35

4.00

3.50

40

JAMES A. SCOTT

/. Res. Lepid.

Table 4. Means of behavioral characteristics for species with different lar-
val foodplants.

No. of

Perching-

Location of

Time of

Larval Foodplants

Species

Patrolling

Index

Mating

Index

Day of Mating
Index

Salicaceae

10

2.40

2.10

3.30

Primus

3

2.00

1.67

3.60

Rumex

5

1.40

1.80

4.00

Asclepias-Croton-Celtis

3

2.00

1.67

3.73

Urtica

4

1.00

2.00

2.35

Malvaceae

3

2.00

2.33

3.50

Chenopodiaceae-

4

4.00

2.00

4.00

Amaranthaceae

Ribes

2

1.00

2.00

2.10

Seduni

2

2.50

3.00

4.00

Androsace

1

4.00

4.00

4.00

Viola

7

4.00

3.43

4.00

Ericaceae

3

2.00

2.00

4.00

Potentilla

2

2.50

2.00

4.00

Scrophulariaceae

3

2.50

2.67

4.00

Compositae

11

3.00

1.91

3.77

Cercocarpus

1

1.00

1.00

4.00

Quercus

5

1.60

1.80

3.56

Pinus-Juniperus-

4

1.75

2.75

4.00

Arceuthohium

Ceanothus

4

1.25

1.25

2.87

Eriogonurn

7

3.14

3.43

3.67

Leguminosae

18

3.11

2.83

3.94

Umbelliferae

2

3.00

2.00

4.00

Criiciferae

9

4.00

2.11

4.00

Polyphagous (S. melinus.

2

2.50

2.00

3.00

C. argiolus)

Grasses or sedges

42

1.81

2.06

3.88

Yucca

2

1.00

3.00

2.65

Table 5. Means of behavioral characteristics for species feeding on herbs
and on shrubs.

Herbs Shrubs-Trees

2.54 2.10

2.29 2.20

3.84 3.46

Perching-Patrolling Index
Location of Mating Index
Time of Day of Mating Index

Journal of Research on the Lepidoptera

14(l):41-48, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

HISTORY OF SCIENTIFIC STUDY ON
A LARVAL COLOR POLYMORPHISM
IN THE GENUS CHLOSYNE ( NYMPHALIDAE )

RAYMOND W. NECK

Department of Zoology, University of Texas at Austin, 78712^

Tresent address: Texas Parks and Wildlife Department, John H. Reagan
Bldg., Austin, Texas 78701.

A STRIKING POLYMORPHISM involving larval color is manifested
in two species of melitaeine butterflies, Chlosijne lacinia
(Geyer) and Chlosijne gorgone (Hubner). Each species exhibits
three morphs: 1) rufa, an orange form, 2) nigra, a black form
and 3) bicolor, a black form with a row of orange spots along
the mid-dorsal line giving the appearance of a broad stripe.
The larval morphs of C. lacinia were illustrated in natural colors
by Neck et al. (1971). The resemblance of the polymorphic
larvae of these two species is remarkable. Genetic studies have
revealed that the inheritance mechanisms are identical in both
species (Neck et ah, 1971; Neck, 1973b). The polymorphism is
believed to be homologous in that it is probable that both species
are derived from a common ancestor which was also polymor-
phic for larval color patterns (Neck, 1973b). Remarkably, there
has been no scientific study of these polymorphisms until the
past several years.

The difference between the comparable morphs of these two
species are diagnostic but very minor. The gorgone rufa is a
yellowish orange while the lacinia rufa is orange to orange-red.
This is true as well for the stripe of the comparable bicolor
morphs. The mid-dorsal spots of the gorgone bicolor are strictly
square-like in configuration while the spots of the lacinia bicolor
are more variable and less distinct in shape and tend toward
ovoid in some individuals. The melanized portions of the bicolor
morph and nigra morph of gorgone appear to be somewhat
darker than the comparable portions of lacinia larvae. This
results in a greater contrast between the dark background and
the lighter stripe of the bicolor morph in gorgone than in lacinia.

41

42

RAYMOND W. NECK

/. Res. Lepid.

Larval Descriptions By W. H. Edwards

Edwards (1893) reported the initial description of lacinia
larvae (as Stjnchloe lacinia Geyer) as quoted from a letter by
T. D. A. Cockerell — “1. nigra, a black form. 2. bicolor, a black
with broad dorsal stripe. 3. rufa, a red form.” Edwards (1893)
also described lacinia larvae in his own words as follows: “There
are at least three distinct types of larva: 1st. — All red- or yellow-
fulvous. 2nd. — The dorsum and lower part of side fulvous, the
sub-dorsal areas dark brown. 3rd. — All black, or black with a
greenish band on each side.” Note the reversal of order of nigra
and rufa.

The following year Edwards ( 1894 ) described samples of
gorgone larvae (reported as Phtjciodes carlota Reakirt) which
he had reared simultaneously with the lacinia larvae. His de-
scription (from three larvae) fits the bicolor morph — “deep
black . . .; a red-fulvous mid-dorsal band . . . .” Although he
received eggs from Colorado and larvae from Montana, most of
his larvae entered diapause before maturing. His original
samples may have contained individuals of one or both of the
other morphs. The color polymorphism is quite muted in dia-
pausing individuals.

It is surprising that Edwards did not refer to a resemblance
of the bicolor larvae of these two species. Although he quoted
from a letter by Cockerell in describing lacinia larvae, Edwards
reared larvae of the two species concurrently. He further stated,
“The eggs of the two are in no way distinguishable, nor are the
larvae in the first two stages; as regards shape and armature they
are alike in the succeeding stages, but differ in coloration.”
Edwards apparently did not consider the larvae to be similar
due to the interspecific differences discussed above. Possibly,
the larvae were affected by rearing conditions, long overland
mail travel, or entered diapause, all of which are known to affect
larval coloration.

Subsequent Descriptions of lacinia Larvae

Other less complete descriptions of the larvae of lacinia have
appeared in the American scientific literature. Comstock (1927:
119) reported the larvae as being “so variable in color as to be
described with difficulty. The range is from a solid black to a
reddish-fulvous, with a variety of stripes and blotches.” Later,
Comstock (1946) reported the larvae of this species to be
“extremely variable in pattern and color.” Tinkham (1944)

14(l):41-48, 1975

LARVAL COLOR

43

merely referred to the “spiny caterpillars” of lacinia. Thorne
(1962) mentioned that “the few orange caterpillars of lacinia
were conspicuous among the dark calif ornica larvae.” It is
quite likely that he overlooked the nigra lacinia larvae as they
look much like the black larvae of Chlosijne calif ornica (Wright)
as described by Comstock ( 1929 ) . Thorne makes no mention of
bicolor-type larvae.

Descriptions of lacinia larvae originating from Latin Ameri-
can populations were also published. Koehler ( 1927 ) reported
upon the biology of lacinia (as Chlosijne saundersi Dbl. & Hew.)
from observations in northern Argentina at the southern edge
of the geographical range of lacinia. He reported that the larvae
vary from bright reddish brown to almost black. The dark form
contained many spots, dorsal, lateral or both (an apparent
reference to the bicolor morph). (“Su color varia de marron
rojizo claro al casi negro. En la forma oscura observamos muchas
veces manchas dorsales o laterales o ambas. . . .” Dyar (1911)
gives a perfect description of a rufa larva as follows: “Body red-
brown, marked transversely with black lines, two on each seg-
ment behind the spines. . . .” This description was made from
an inflated specimen originating from an undefined locality in
Mexico. It is pertinent here to mention that lacinia is not listed
in monographs on the nymphalid larvae of South America
(Muller, 1886) or life histories of Mexican lepidoptera (Com-
stock and Vasquez, 1961).

Subsequent Descriptions of gorgone Larvae

Other larval descriptions of gorgone exist. Only shortly be-
fore the appearance of the paper by Edwards (1894), Dyar
(1893) described what was apparently a rufa larva (“body
brownish-red”) although it might have been a variant bicolor
as he further describes it “with a dorsal and subdorsal black
shaded line.” Handford (1933) described the diapausing be-
havior of gorgone larvae but he gave no phenotypic description.
Leussler (1938-39) mentioned that the larvae of gorgone were
easily reared but made no mention of larval phenotypes. Heitz-
man ( 1963 ) referred to the similarity of the “quite black” larvae
of Chlosyne nycteis (Dbl.) to the larvae of gorgone without
referring to the polymorphic nature of gorgone larvae.

44

RAYMOND W. NECK

/. Res. Lepid.

One reference to the similarity of the larvae of lacinia and
gorgone has been found. Cockerell (1914) described the bicolor
and rufa of gorgone (as Phijciodes ismeria Bdv. & Lee.) from
Boulder, Colorado. It is strange that the nigra morph was not
also described as nigra is the most common morph in Kansas
populations (Neck, 1974). Reference is made to the fact that
the two morphs described for gorgone “nearly correspond to two
varieties” of lacinia. The two morphs which Cockerell described
are the same two which had been previously reported (bicolor
by Edwards, 1894, rufa by Dyar, 1893). Apparently the tri-
morphic nature of the larvae of gorgone was not realized until
the recent publication on the genetics of this polymorphism
(Neck, 1973b).

Most general (Morris, 1862; Scudder, 1889:111, 1811; Holland,
1898, 1931; Klots, 1951) and regional (Macy and Shepard, 1941;
Brown et al., 1957; Ebner, 1970; Harris, 1972) butterfly manuals
repeated a description for gorgone which best fits a rufa larva
but differs from the one above. This description originated from
Boisduval and LeConte (1833:168) whose name ismeria is now
relegated to a synonym of gorgone Hubner (see Dos Bassos,
1969). The description refers to a larva which is “yellowish with
three longitudinal black stripes.” Edwards (1894) states that
the description of ismeria larvae “has no application” to the
gorgone larvae he reared.

This description, however, also closely fits the larva of
Chlosyne harrisii (Scudder). The better fit of the larval descrip-
tion of ismeria may be significant in that the dispensation of
this name has not been completely settled. Higgins (1960) felt
that ismeria more closely resembles aberrant forms of harrisii.
F. M. Brown (1974) felt that ismeria is a synthetic drawing,
possibly a modification of harrisii or gorgone. The description
of the larva gives some credence to the view that ismeria is
some form of harrisii although a harrisii- gorgone hybrid is also
a possibility. However, the supposed type locality of ismeria,
Georgia, is more likely to be gorgone as harrisii is not presently
known from localities that far south. A likely solution to this
nomenclatural problem is that ismeria is an extreme form of
gorgone as some contemporary gorgone are known from Georgia
which closely resemble ismeria (Harris, 1972:264, color plate 7).

14(l);41-48, 1975

LARVAL COLOR

45

Reasons for Lack of Study

It can thus be seen that of the three people who described
the larvae of lacinia in detail, only one (Cockerell in Edwards,
1893) divided the larval forms into three distinct categories.
Apparently only one person (again Cockerell) saw more than
one morph of gorgone. Comstock apparently saw all three
morphs of lacinia, but he did not separate them into three dis-
tinct categories. This may have been partially due to the effect
of environmental factors and/or genetic modifiers which tend
to produce a nearly continuous variation from a basically dis-
continuous genetic polymorphism (Neck, 1974). Koehler, like
Comstock, did not take notice of the three morphs of lacinia,
although it appears that this larval polymorphism of color pat-
terns is also present in Argentine populations.

The phenomenon of polymorphism was not as widely dis-
cussed at the times of the above descriptions as it is now. This
may partially explain the lack of distinction between larval
morphs of lacinia. At the time of most early descriptions of
larvae of these species there was little concept of polymorphism
in the United States scientific community. Mendel’s work had
not yet or had just become known; forms other than the sup-
posedly ubiquitous “wild-type” were considered sports. Later,
the rise of the ecological genetics school in England saw work
on Panaxia, Biston and Cepaea. From this beginning polymor-
phism has become one of the most discussed biological phenom-
ena of today.

Interestingly, one publication (Edwards, 1893) includes the
term polymorphism in the title. Here, however, the term is ap-
plied to variant forms of the adult phenotypes of lacinia
(adjutrix Scudder and crocale Edwards and hybrids thereof),
not to the larval stages. At that time attention for studies was
focused (as it still is today, although to a lesser extent) upon
imaginal forms. Larval descriptions were recorded for identi-
fication purposes, but these stages were often considered to
be merely a stage required to produce the adult form and not
a form with its own adaptive strategy. Cockerell (1914) re-
ferred to the larvae of lacinia as “polychroic.”

The descriptions in Edwards (1893) were not picked up by
most later workers. Higgins (1960) does not refer to this paper
in his discussion of the larvae of lacinia although he does refer
to the paper in other contexts. Holland (1898) in his first
edition refers to the larvae as being “fully described by Edwards”

46

RAYMOND W. NECK

]. Res. Lepid.

and follows with the correct reference (Edwards, 1893), but
he does not repeat the larval descriptions. For some unknown
reason he drops the mention of this description in his revised
edition (Holland, 1931) and says nothing of the larvae of
lacinia. As most subsequent natonal and regional butterfly
treatments are based upon this work, subsequent guides say
nothing of the larvae of lacinia. The reference was not com-
pletely lost, however, as Davenport and Dethier ( 1937 ) in their
bibliography of rhopaloceran life histories listed this reference
with respect to hcinia.

The long, unprecise description of lacinia larvae given by
Comstock ( 1927 ) would not likely be republished by workers
who would prefer a short, precise description for publication in
a guide book. As this description was published in a regional
butterfly book, it would not have received the wide distribution
of a book of the scope of Holland (1898, 1931). In his book,
Comstock (1927) does not refer to the Edwards (1893) refer-
ence, although he does make reference to it in a later paper
(Comstock, 1946).

The lack of study of these larvae may also be related to the
non-occurrence of lacinia and apparent uncommon occurrence
of gorgone in those areas extensively studied by the early
American lepidopterists, i.e. the northeastern states. C. lacinia
occurs from Texas westward to southern California and south-
ward to Argentina. C. gorgone ranges from Texas to Manitoba,
occurring into the eastern Rockies particularly along water
courses. It is also sporadically reported throughout the southern
states to Florida and Georgia. An apparently isolated population
of gorgone occurring in northern New York is the only known
eastern record north of Georgia (Shapiro, 1974).

Both butterflies are opportunistic species of varying abun-
dance whose larvae feed upon species of the family Compositae
which are characteristic of highly disturbed habitats (Neck,
1973a and unpublished data). Much of the ranges of these
species includes areas which are characterized by long periods
of dry weather when butterflies are essentially non-existent.
This is particularly true of the desert and semi-desert areas of
southern California, Arizona, New Mexico and western Texas.
This fluctuating pattern of population size may possibly have
contributed to their being overlooked. Edwards (1894) remarks
on the lack of study as follows: “Considering what a common

14(l):41-48, 1975

LARVAL COLOR

47

species carlota is over at least one-third of the territory of the
United States, it is remarkable that so little has been published
respecting it.” He further notes that some of his correspondents
found this species to be common, while others in the same
region found it rare.

As neither of these species is an agricultural pest, there was
no study by agricultural scientists. Although both feed on wild
sunflowers, the commercial monocephalic sunflowers are not
readily acceptable to lacinia (Neck, 1973a).

SUMMARY

A striking polymorphism involving color polymorphism is
exhibited by the larvae of Chlosyne lacinia and Chlosyne gor-
gone. These polymorphisms have remained unstudied until very
recently because of a combination of factors. Their geographical
ranges occur, for the most part, outside the areas most intensive-
ly studied by early American lepidopterists. A reference de-
scribing the polymorphic nature of one species was not utilized
by later workers. Only recently was the complete polymorphic
status of the other species described. Other descriptions referred
to a single morph (in one case two morphs were described) or
described a continuous variation of larval phenotypes. A prob-
lem of taxonomic nomenclature resulted in an erroneous de-
scription for gorgone in most butterfly texts. Characteristics of
their population biology also tend to lessen the chances of study.

ACKNOWLEDGEMENTS

I wish to thank D. D. Balser for assistance in preparation of
the manuscript.

LITERATURE CITED

BOISDUVAL, J. A. and J. E. LeCONTE. 1833. Historie generale et mono-
graphie des lepidopteres et des chenilles de FAmerique septrionale.
Paris.

BROWN, E. M. 1974. The butterfly called ismeria by Boisduval and
LeConte (with a neotype for Eresia carlota Reakirt). Bull. Allyn Mus.
16:1-12.

, D. EFF and B. POTZGER. 1957. Colorado Butterflies. Denver Mus.

Natur. Hist., Denver.

COCKERELL, T. D. A. 1914. The entomology of Helianthus. Ento. 47:
191-196.

COMSTOCK, J. A. 1927. Butterflies of California. Published by the author,
Los Angeles.

. 1929. Studies in Paciflc Coast lepidoptera (continued). Bull. So.

Cal. Acad. Sci. 28:50-58.

. 1946. A few pests of sunflower in California. Bull. So. Cal. Acad.

Sci. 45:141-144.

and L. G. VAZQUEZ. 1961. Estudios de los ciclos biologicos en

lepidopteros Mexicanos. Annales Inst. Biol. Mex. 31:349-448.

48

RAYMOND W. NECK

/. Res. Lepid.

DAVENPORT, D. and V. G. DETHIER. 1937. Bibliography of the described
life-histories of the rhopalocera of America north of Mexico. Entom.
Amer. 17:155-194.

DOS PASSOS, C. F. 1969. A revised synonomic list of the Nearctic Meli-
taeinae with taxonomic notes ( Nymphalidae). J. Lepid. Soc. 23:115-
125.

DYAR, H. G. 1893, On, some butterfly larvae not hitherto described. Can.
Entom. 25:93-94.

— . 1911. Descriptions of the larvae of some Mexican lepidoptera.

Proc. Entom. Soc. Wash. 13:227-232.

EBNER, J. A. 1970. The butterflies of Wisconsin. Milwaukee Public Mu-
seum, Popular Science Handbook No. 12.

EDWARDS, W. H. 1893. Notes on a polymorphic butterfly, Synchloe
lacinia (in Hub. Zutr. ), with descriptions of its preparatory stages.
Can. Entom. 25:286-291.

. 1894. Description of the preparatory stages of Phyciodes carlota,

Reakirt {Charidryas ismeria, Scudder). Can. Ent. 26:3-8.

HANDFORD, R. H. 1933. Note on tire hibernation of Phyciodes gorgone,
Hbn. (Lepidoptera, Nymphalidae). Can. Entom. 65:95.

HARRIS, L. 1972. Butterflies of Georgia. Univ. Okla. Press, Norman.

HEITZMAN, R. 1963. Record charts for the collector of lepidoptera. J. Lep.
Soc. 17:44-46.

HIGGINS, L. G. 1960. A revision of the melitaeine genus Chlosyne and
allied species (Lepidoptera: Nymphalidae). Trans. Entom. Soc. Lon-
don 112:381-467.

HOLLAND, W. J. 1898, 1931. The butterfly book. 1st and revised editions.
Doubleday, Garden Gity, N.Y.

KLOTS, A. B. 1951. A field guide to the butterflies. Riverside Press, Gam-
bridge, Mass.

KOEHLER, P. 1927. Biologia de Chlosyne Saundersi Dbl. & Hew. Revista
Soc. Ent. Argentina 1-2: 3-4,

LEUSSLER, R. A. 1938-1939. An anotated list of the butterflies of Ne-
braska, with the description of a new species ( Lepid. :Rliopalocera).
Ent. News 49:3-9, 76-80, 213-218, 275-280, 50:34-39.

MACY, R. W. and H. R. SHEPARD. 1941. Butterflies. University of Minne-
sota, Minneapolis.

MORRIS, J. G. 1862. A synopsis of the described lepidoptera of North
America. Smith Misc. Coll. 4:1-357.

MULLER, W. 1886. Sudamerikanische Nymphalidenraupen. Gustav Fisher,
Jena.

NEGK, R. W. 1973a. Foodplant ecology of the butterfly Chlosyne lacinia
(Geyer) (Nymphalidae). I. Larval foodplants. J. Lepid. Soc. 27:22-33.

. 1973b. Homologous polymorphism and niche equivalence in the

butterfly genus Chlosyne. Heredity 31:118-123.

. 1974. Ecological genetics of a larval color polymorphism in the

buterfly genus Chlosyne. Ph.D. dissertation, Univ. Texas at Austin.

, G. L. BUSH and B. A. Drummond III. 1971. Epistasis, associated

lethals and brood effect in larval color polymorphism of the patch
butterfly, Chlosyne lacinia. Heredity 26:73-84.

SCUDDER, S. H. 1889. The butterflies of the eastern United States. Pub-
lished by author, Cambridge, Mass.

SHAPIRO, A. M. 1974. Butterflies and skippers of New York State. Search
( Agric.-Ent. ) 4(3): 1-60.

THORNE, F. 1962. Larval notes on Chlosyne lacinia and C. californica. J.
Lepid. Soc. 16:61.

TINKHAM, E. R. 1944. Faunistic notes on the diurnal lepidoptera of the
Big Bend region of Trans-Pecos, Texas, with the description of a new
Melitaea. Can. Entom. 76:11-18.

Journal of Research on the Lepidoptera

14(l):49-56, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
(c) Copyright 1975

THE BUTTERFLY FAUNAS OF SAN ANDRES
AND PROVIDENCIA ISLANDS
IN THE WESTERN CARIBBEAN

THOMAS C. EMMEL‘

Department of Zoology, University of Florida,

Gainesville, Florida 32611

San Andres Island and Providencia Island are the western-
most islands in the West Indies, lying only about 120 miles east
of the Nicaraguan coast. Both islands are located beyond the
100 fathom level (Fig. 1), however, in the western Caribbean
and there is no evidence for submerged banks between the
islands and the mainland that might have supported butterflies
or other organisms during a past period of lowered sea level
(Corn and Dalby, 1973). Thus this isolated pair of islands and
associated tiny cays are of considerable interest in considering
the colonization and evolution of island butterflies in the West
Indies, and despite excellent recent reviews of Antillean butter-
fly zoogeography (Scott, 1971, 1972), there has been no pub-
lished survey of the San Andres and Providencia butterflies.

In company with other faculty and students in the Organi-
zation for Tropical Studies, Inc. course program in Costa Rica,
I was able to make five visits to these islands at various times
in the dry and wet seasons of 1967 and 1968. I collected on San
Andres during March 25-26 and July 22-30, 1967, and February
19-23, June 21-24, and July 15-22, 1968; collecting on Providencia
was done on June 2 and July 20-21, 1968. The records of butter-
fly species in this report come entirely from this collecting.
Specimens are deposited in the Florida State Collection of
Arthropods and the authors personal collection.

^Research Associate, Florida State Collection of Arthropods and Natural
History Museum of Los Angeles County.

49

50

THOMAS C. EMMEL

/. Res. Lepid.

DESCRIPTION OF THE ISLANDS

San Andres and Providencia lie approximately 202 and 240
km, respectively, east of Nicaragua. For historical reasons, both
islands are possessions of the country of Colombia. San Andres
(120° 35' N, 81° 42' W), the more southerly of the two islands,
is about 13 km long and 3 km wide. The relief is comparatively
gentle, with a central hill rising to a maximum height of about
104 m (338 feet) above sea level. The island is completely
covered by a layer of hard limestone, although underneath it is
assumed to be basically volcanic. Several freshwater ponds (the
largest of which is known as “Big Pond,” placed at the island’s
center) are present. Most of the island is presently devoted to
the cultivation of coconut palms and Colombian tourism, and
the human population density is great ( 17,000 residents in 1967 —
distributed over the 16 square miles of island area). During the
17th and 18th centuries, however, there apparently were ex-
tensive natural stands of “cedar,” possibly Cedrela odorata in
the family Meliaceae (Whitmore and Hartshorn, 1969), which
were decimated by the early colonists.

Providencia (13° 21' N, 81° 22' W) is located about 88 km
NNE of San Andres and differs markedly from it in vegetation,
topography and superficial geology. Approximately 8 km long
and 5 km wide, it is a high, rugged island of andesite and basalt,
with steep, rocky, central peaks rising to 363 m (1100 feet)
above sea level. No freshwater ponds are found; however, several
streams descend from the peaks. They may become intermittent
during the dry season (January- April). In the higher regions of
Providencia, bracken and tree ferns are common, and in the
steeper uncleared areas occur dense thickets of figue (Foiircroya
sp.), calabash {Crescentia sp. ), and other trees. Coconut palms
are found only infrequently, and there are few agricultural areas;
introduced guinea grass {Panicwn maximum) supports some
cattle in the lower valleys. Extensive mangrove swamps occupy
a considerable portion of the northeast shore. Few tourists visit
this island, the transportation being infrequent and the facilities
minimal.

The climate on both islands is tropical, with the average
daily temperature being about 27° C throughout the year. The
dry season, as in Central America, lasts from January to April,
March and April being the driest months. October and November
are normally the wettest months. The average annual rainfall on
San Andres between 1934 and 1943 was 1824 mm (71.81 inches).

14(l):49-56, 1975

ISLAND FAUNAS

51

The interglacials of the Pleistocene Epoch would have con-
siderably decreased the surface area of low-lying San Andres.
The glacial periods, however, clearly greatly modified the sur-
face areas ^of both islands, as indicated by the extensive and
massive coral reefs and shoals bordering parts of each island.
Providencia lies 74 km, and San Andres 83 km, from the 100
fathom depth line off mainland Central America. Soundings of
trenches as deep as 1500 m have been taken between the two
islands.

THE BUTTERFLIES OF SAN ANDReS
Fifteen species of butterflies were taken or observed on this
island. Most were distributed generally around the island, as
were the seven species of dragonflies recorded earlier by Paul-
son (1968). No Papilionidae, Satyridae, Libytheidae, Riodinidae,
or Megathymidae were found. It is strange that Heliconius chari-
tonius and Euptoieta hegisia are apparently absent. A list of
recorded species follows.

DANAIDAE

1. Danaus plexippus Linnaeus.

Scattered and relatively worn individuals were seen on
all dates. No milkweeds were seen on this island.

NYMPHALIDAE

1. Dry as julia Fabricius.

A number of subspecies of this long-winged heliconian
have been named from various Caribbean islands (and
mainland areas), for the species seems to vary distinguish-
ably from island to island (see Brown and Heineman,
1972, for a partial review). No special name is presently
assigned to the San Andres-Providencia populations.

Dryas flies in fair abundance on the slopes around
Big Pond throughout the year and is continuously
brooded here and elsewhere on San Andres with fresh
adults appearing every month. Dryas is relatively rare
at Puerto la Ensenada, on the central west coast of San
Andres. On 21 February 1968, two Dryas males were ob-
served courting different Agraulis vanillae females around
0900 and 0915 hours, in low second-growth vegetation.
Female Dryas have a dark blackish band across the fore-
wings, while males rarely have any indication of this band.

52

THOMAS C. EMMEL

J. Res. Lepid.

2. Agraulis vanillae Linnaeus.

Rare in the Big Pond area ( where Dry as is abundant ) ,
the Gulf Fritillary is quite abundant at Puerto la Ensenada
on the west coast (24 adults in an hour in February). It
resembles typical Florida A. v. nigrior, although the black
forewing spots often form a bar-like continuous row.
Young larval instars were found on Passiflora sp. at the
Puerto area.

3. Victorina (Siproeta) stelenes (Linnaeus).

Several fresh specimens were observed (Big Pond area,
Hotel Casa Blanca) each day in February, but it was
rarer in other months. The species is known to feed on
Blechwn hlechum and Rtiellia coccinea ( Acanthaceae )
in Jamaica (Brown & Heineman, 1972); these are weedy
plants which may well occur on San Andres.

4. Hist oris odiiis Fabricius.

A robust, strong-flying nymphalid, Historis is a rare
sight on San Andres (July 1967, 1968) but its black and
orange-brown coloration is unmistakable. The few adults
seen may be strays from Providencia or even the main-
land because its only known foodplants are trees of the
genus Cecropia, a member of the Urticaceae and charac-
teristic inhabitant of lowland rain forest areas in the New
World tropics.

5. Precis evarete zonalis Felder & Felder.

The West Indian buckeye is a medium-sized dark
brown butterfly with small hindwing ocelli. It also occurs
on the mainland from northern Mexico and the southern
tip of Florida south to Venezuela and Trinidad. It is
relatively rare on San Andres and occurs principally in
the Big Pond area (February, July).

6. Anartia jatrophae Johansson.

This nymphalid is quite variable through its Neo-
tropical range, and the species also shows seasonal vari-
ation in color and pattern. It prefers weedy areas and is
fairly common in July, but rare in February.

PIERIDAE

I. Phoebis sennae sennae (Linnaeus).

This large, rich yellow Cloudless Sulphur has scattered
patches or lines of reddish brown scales on the underside
of the hindwing. Fresh specimens were seen daily, but

14(l):49-‘56, 1975

ISLAND FAUNAS

53

the species is not particularly common in February, March
or July. A low-growing weedy Cassia (larval foodplant)
is found on the coastal margins and around Big Pond.

2. Eurema daira Godart.

This variable Eurema, common in the Antilles and on
the mainland in Nicaragua and Costa Rica, is relatively
rare on San Andres; it was recorded daily in July but not
in February. The larvae feed elsewhere on vetches and
other related Fabaceae. The species is known to be migra-
tory in Costa Rica.

3. Eurema lisa Boisduval & Leconte.

The Little Sulphur is also comparatively rare on San
Andres and was noted only during the wet season in July
(T.C.E., G. N. Ross). Rs larval host are Mimosa and
Cassia; both present on San Andres.

LYCAENIDAE

1. Leptotes cassius (Cramer).

A very common blue throughout the West Indies, this
Leptotes is quite colonial on San Andres. A large popu-
lation exists slightly north of Puerto la Ensenada, centered
around low beach legumes, while none were found at Big
Pond or around the settlements with the exception of one
male seen visiting flowers at the Hotel Casa Blanca.

HESPERIDAE

1. Urbanus proteus (Linnaeus).

This dark long-tailed skipper, with iridescent greenish-
blue wing-scaling near and on the thorax, is common on
the island and the adjacent mainland. Gravid females
with mature eggs were found in both February and July
at Big Pond, where the species was using unidentified
low-growing legumes as larval hosts.

2. Pyrgus oileus (Linnaeus).

The former name of this common checkered skipper in
the Caribbean was P. syrichtus (Brown & Heineman,
1972). Pyrgus oileus was seen sporadically around Big
Pond (February 1968).

3. Small brown skipper, possibly female Hyelephila phyleus
(see Providencia list).

One was seen but not captured, along road above the
east side of Big Pond on 22 February 1968.

54

THOMAS C. EMMEL

/. Res. Lepid.

4. Small black skipper (body length 9 mm).

One specimen was taken in a sweeping study of grass-
inhabiting insects under coeonut palms, near the trail
going up from the east side of Big Pond, at 1000 hours
on 22 February 1968. Unfortunately, it was so damaged
by debris in the sweep net that identification was im-
possible.

THE BUTTERFLIES OF PROVIDENCIA

Loeal inhabitants suggested that butterflies on this island
were at their peak later in the wet season than when collecting
was done in the present study (June and July). From personal
observation, I would agree that the flight peak for resident
species is probably more marked than on San Andres and that
good population numbers likely do not appear until August
through October or November. Every species found on San
Andres also must oecur on Provideneia, and with the greatly
inereased diversity of vegetation on Provideneia, one would
presume that several additional speeies of butterflies should
oceur. The diseovery of a seeond Phoebis speeies and another
skipper speeies, even with very little sampling time, supports
this view. Most of the records reported here were obtained on
the southern and western coastal and moderate-elevation areas.

DANAIDAE

1. Danaiis plexippiis.

NYMPHALIDAE

1. Dry as jiilia.

2. Agraulis vanillae.

3. Hist oris odiiis.

PIERIDAE

1. Phoebis sennae.

2. Phoebis statira Cramer.

A large Phoebis with the basal two-thirds of the wing
yellow and the outer third covered with whitish scaling,
P. statira is a pandemic species in the Caribbean area and
its migratory behavior is well known. Though only two
males were seen in July, the presence of its Cassia food-
plants likely means it is a breeding resident on at least
Provideneia if not also on the more arid island of San
Andres.

14(l):49-^56, 1975

ISLAND FAUNAS

55

LYCAENIDAE

1. Leptotes cassius,

HESPERIIDAE

1. Hylephila phijleus phyleus (Drury).

The Fiery Skipper is found throughout the West Indies
and North, Central, and South America. It is common on
Providencia in lush grassy areas from sea level to over
800 feet elevation.

DISCUSSION

A total of 17 species of true butterflies and skippers were
found on San Andres and Providencia Islands, located about 120
miles east of the coast of Central America. No endemic species
occur on these isolated West Indian islands, nor are there any
uniquely Antillean residents. All of the butterflies are also found
on the adjacent Nicaraguan mainland area, and all represent
species that are migratory or are known to be frequent colonizers.
This situation is typical of the Antilles as a whole, where more
than 60 percent of the relatively impoverished butterfly fauna
of 285 species are widespread continental species of great dis-
persal ability (Scott, 1971, 1972). Further collecting in the high
interior of Providencia and during the wet season will undoubt-
edly uncover several additional resident species of butterflies
and skippers.

LITERATURE CITED

BROWN, F. M., and HEINEMAN. 1972. Jamaica and Its Butterflies. E. W.
Classey, Ltd., London. 478 pp.

CORN, M. J., and P. L. DALBY. 1973. Systematics of the anoles of San
Andres and Providencia Islands, Colombia. /. Herpetology, 7(2) :63-
74.

PAULSON, D. R. 1968. Odonata from Isla San Andres, western Caribbean
Sea. Entomol. News, 79:229-231.

SCOTT, J. A. 1971. A list of Antillean butterflies. /. Res. Lepid., 9(4):249-
256. (“1970”)

. 1972. Biogeography of Antillean butterflies. Biotropica, 4(l):32-45.

WHITMORE, J. L., and G. S. HARTSHORN. 1969. Literature review of
common tropical trees. Inst. Forest Prod. Coll. For. Prod. Univ. Wash-
ington Contrih. No. 8. 113 pp.

56

THOMAS C. EMMEL

/. Res. Lepid.

84° 80°

Fig. 1. — The position of San Andres and Providencia Islands, Colombia, in
the western Caribbean in relation to Nicaragua on the mainland of Central
America. The 100-fathom depth level of the ocean is indicated by a dotted
contour line.

Journal of Research on the Lepidoptera

14(l):57-59, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A LIST OF THE BUTTERFLIES OF
THE WILLOW RIVER STATE PARK,
WISCONSIN

JOHN H. MASTERS

8126 Santa Inez Drive, Buenu Park, California

Willow River State Park, in Saint Croix County, northwest
Wisconsin, was officially opened in 1972 and is the newest ad-
dition to the Wisconsin state park system. The Park consists of
three small impounded lakes on the Willow River, recreational
areas that are for the most part adjacent to these lakes and
several hundred undeveloped acres of woodland, meadows and
old pastures. It is a very good area for the collection and study
of Lepidoptera.

I previously listed (Masters, 1973) 42 species of butterflies
as occurring in St. Croix County where I resided for 3 years.
Heavy collecting in the Willow River Park has added 7 species
to the county list and 12 species of Hesperiidae (not included
in the previous list) were also observed, making a total of 61
species taken in the State Park or its immediate vicinity. The
large number of species taken here is a result of the diverse
habitats available in the Park which include deciduous wood-
land, both wet and dry meadows, remnant prairie and oak
openings. A list of the species taken follows:

HESPERIOIDEA
Poanes hobomok ( Harris )

Atrytone delaware ( Edwards )

Wallengrenia otho ( Smith )

Polites coras ( Cramer )

Polites themistocles ( Latreille )

Polites mystic ( Scudder )

Hesperia metea Scudder
Erynnis icelus ( Scudder & Rurgess )

Erynnis juvenalis ( Fabricius )

Pyrgus communis ( Grote )

Ancyloxypha numitor (Fahricius )

Epargyreus clarus ( Cramer )

57

58

JOHN H. MASTERS

/. Res. Lepd.

PAPILIONOIDEA

Papilio polyxenes asterius Stoll

Papilio glaucus glaucus Linnaeus

Pieris protodice protodice Boisduval & LeConte

Pieris rapae rapae ( Linnaeus )

Colias eurytheme eurytheme Boisduval
C olios philodice philodice Godart
Colias interior interior Scudder
Colias cesonia cesonia (Stoll)

Nathalis iole Boisduval

Euchloe olympia olympia (Edwards)

Libythea bachmanii bachmonii ( Kirtland )

Danaus plexippus plexippus ( Linnaeus )

Lethe anthedon anthedon (Clark)

Lethe eurydice eurydice (Johansson)

Euptychia cymela cymela ( Cramer )

Cercyonis pegala nephele (Kirby)

Asterocampa celtis celtis ( Boisduval & Le Conte )

Limenitis arthemis ! astyanax (Fabricius) hybrid population
Limenitis archippus archippus ( Cramer )

Vanessa atalanta rubria (Fruhstorfer)

Cynthia virginiensis (Drury)

Cynthia car dui (Linnaeus)

Nymphalis niilberti milberti ( Godart )

Nymphalis antiopa antiopa (Linnaeus)

Polygonia interrogationis ( F abricius )

Polygonia comma ( Harris )

Polygonia progne ( Cramer )

Phyciodes tharos tharos ( Drury )

Chlosyne nycteis nycteis (Doubleday)

Boloria selene nr. inyrina ( Cramer )

Boloria bellona bellona (Fabricius)

Speyeria idalia ( Drury )

Speyeria cijbele cybele (Fabricius)

Speyeria aphrodite aphrodite (Fabricius)

Speyeria aphrodite alcestis ( Edwards )

Euptoieta clatidia claudia ( Cramer )

Harkenclemis titus titiis ( Fabricius )

Satyrium edivardsii ( Grote & Robinson )

Satyrium calanus falacer ( Godart )

Satyrium caryaevoriis ( McDunnough )

Satyrium acadica acadica ( Edwards )

14(1) -.57 -59, 1975

LIST OF BUTTERFLIES

59

Incisalia henrici henrici ( Grote & Robinson )

Mitoura gryneus gryneus Hubner
Feniseca tarquinius tarquinius (Fabricius)

Lycaena xanthoides dione Scudder
Lycaena helloides (Boisduval)

Lycaena phlaeas americana Harris
Glaucopsyche lygdamus couperi Grote
Everes corny ntas comyntas (Godart)

Celastrina argiolus pseudargiolus (Boisduval & Le Conte)

Satyrium caryaevorus, which is new to the county list,
seemed to be restricted to Oak Opening communities and
seemed to fly much higher up in the oak and hickory trees than
Satyrium calanus which flew with it. Speyeria idalia actually
did not occur in the state park, but was taken on a patch of
remnant prairie along a railway track just south of the park.
C olios interior was recorded from a single male (very definitely
this species) that was taken at the edge of a field of cultivated
strawberries on the south border of the park!

LITERATURE CITED

MASTERS, JOHN H. 1973. Butterfly records for three northwest Wiscon-
sin Counties. Jour. Res. Lepid., 11(3) ( 1972) : 175-182.

Journal of Research on the Lepidoptera

14(1 ):64, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

LIVE GEOMETRID

(Cover illustration)

NOEL McFarland

P. O. Box 475, Geraldton, W. Australia 6530

Botli upper and lower photos show the same moth, at rest, as viewed
from two different angles. No leg or wing movements have taken plaee.

Identification: Oenochroma vinaria Guen.

(N. McF. code No. G. 77)

Family GEOMETRIDAE— Subfamily OENOCHROMINAE

Locality: South Australia, Mt. Lofty Range, nr. Blackwood. July 30
1967, 0200 hrs. at UV light, Noel McFarland

Photographed by Noel McFarland.

Lower figure: Living adult $ in its natural position, as viewed from
the rear. Right antenna visible tucked along side of the tliorax, beneath
wings. Length of forewing, from base to tip is 24 mm. This species nonn-
ally rests on litter or near the ground among dead leaves; thus, the wings
are held up at rest rather tlian appressed to the substrate. The uplifted
wings cast a shadow as do the variously twisted and bent dead leaves.

60

THE JOURNAL OF RESEARCH
ON THE LEPIJDOFTERA

C ONTE NTS

Volume 13 Number 1

March, 1974

Studies on Nearctic Euchloe — Part 7

Paul A. Opler 1

3 Stacks of the eggs of Hemistola hatching

Noel McFarland 21

Checklist of the Macroheterocera of
south-eastern Ontario

P. S. Ward, R. Harmsen, and P. D. N. Hebert 23

A new species of Hypagyrtis

Roger L. Heitzman 43

Genetic control of maculation and

hindwing color in Apantesis phalerata

Jack S. Bacheler and Thomas C. Emmel 49

Natural and laboratory occurrence of “Elymi”

phenotypes in Cynthia cardui (Nymphalidae)

Arthur M. Shapiro 57

Das naturhistorische Museum in Wien
und seine Lepidopterensammlung

Dr. Fritz Kasy

63

Notice

William Hovanitz

66

Habitat: Adela bella in Florida

John B. Heppner

67

Volume 13 Number 2 June, 1974

The butterfly fauna of the

Sacramento Valley, California

Arthur M. Shapiro 73

Extended flight periods of coastal and
dune butterflies in California

Robert L. Langston 83

Habitat: Brephidium pseudofea (Lycaenidae)

John B. Heppner 90

The early stages of various species of
the genus Dirphia ( Saturniidae)

Brian O. C, Gardiner 101

A new species of Ormiscodes ( Dirphiella ) from
Mexico (Saturniidae: Hemileucinae)

Julian P. Donahue and Claude Lemaire

123

A new subspecies of Euphydryas editha
from the Channel Islands of California

Thomas C. Emmel and John F. Emmel 131

Volume 13 'Number 3 September, 1974

Studies of the ova and first instar

lar\'ae of Geometridae (Ennominae). I.

Roger L. Heitzman 149

Altitudinal migration of central California butterflies

Arthur M. Shapiro 157

The chromosomes of Apantesis phalerata,

A. radians, and their hybrid in Florida populations
(Arctiidae) Jack S. Bacheler and

Thomas C. Emmel 162

A partial bibliography of the world distribution
and zoogeography of butterflies

Oakley Shields 169

The nomenclature in an important British checklist ( 1972)

Part 2: Corrections of family-group names for
Geometridae ( Lepidoptera ) Juraj Paclt 179

Illustrations and descriptions of

species of some Pyrrhopyginae from Panama
( Hesperiidae ) S. S. Nicolay 181

Butterflies of the Suisun Marsh, California

Arthur M. Shapiro 191

Volume 13 Number 4 December, 1974

Toward a theory of butterfly migration

Oakley Shields 217

Atrijtunopsis liianna biology and life
history in the Ozarks

J. Richard and Roger L. Heitzman 239

Larval migration of litjles lincata (Fab.)

James F. Wells and Richard M. Brown 246

New food plant for Darapsa pholus (Cramer)

J. C. E. Riotte 247

Notes on Arctic and Subarctic collecting

Clifford D. Ferris 249

Kloet and Hincks’ Check list of British Lepidoptera
Insects (Lepidoptera) Edn. 2. A reply to criticisms
J. D. Bradley, D. S. Fletcher and P. E. S. Whalley 265

The Nomenclature in an Important British
check list (1972). Part 3. Correct gender
for generic names derived from classical
without change of termination Juraj Paclt 267

Artificial Diet: The key to the mass
rearing of Me^atJuimus larvae

Ronald S. Wielgus 271

Review: Butterflies of the World

by H. L. Lewis Clifford D. Ferris 278

NOTICES

DEFECTIVE COPIES: Some missing pages in volume 13^ no, 1 have
been detected. Please check your copy; correct copies will be sent
to those who received faulty issues and indicating such to us.

INDEXES: Index to volume 1 1 is in volume 13, no. 3, to volume 12
is in volume 13, no, 1.

PUBLICATIONS: The Swallowtail Butterflies of North America.
Hamilton A. Tyler, Naturegraph Publishers Inc. Healdsburg, Calif.
95448. Paper ^5. 95, cloth ^9. 95.

Mites of Moths and Butterflies. Asher E. Treat, Cornell Univ.
Press, 124 Roberts Place, Ithaca, N. Y. 14850, 035,00

Geographical Variability in Speyeria. Arthur H, Moeck. reprint.
Entomologfcal Reprint Sperialists, P. O. Box 77224, Dockweiler Sta. ,
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Butterflies of Lebanon, Torben B. Larsen. Nat'l Council for Sci.
Res., Republic of Lebanon. Avail, in U. S. A. from E. R. S. (see above)
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The Swallowtail Butterflies of East Africa, reprint. R. H.
Carcasson, F. R. E. S. E. W. Classay Ltd. in U. S. A. from E. R. S.

(see above). 04, 95.

Macr olepidoptera of Fiji and Rotuma. Gaden S. Robinson. E. W.
Classey, Ltd. in U. S.A. from E. R. S. (see above). 0Z5,95,

Biology, Ecology and Host Specificity of Microlepidoptera
associated with Quercus Agrifolia. Paul A. Opler. Univ. Calif. Press,
2223 Fulton St. , Berkeley, CA 94720. ^4.25.

Colour Identification Guide to British Butterflies. T. G. Howarth .
Frederick Waine and Co., Ltd. 40 Bedford Square, London. WCIB.

New York 101 Fifth Ave. £3. 00 net.

FOR SALE: Mounted, named local moths. Min. order 1000 at ^400.
per thousand in series up to 20 per species. Similar price for 1976
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SPECIAL NOTICE: In order to return to a normal publication
schedule, the J. Lep. Res. needs manuscripts. Color can be used
within a few months if received shortly.

THE J0UIRNJAL ©F RliEA.R.€HJ
OH THE LEFIJOOFTERA

Volume 14 Number 1 March, 1975

IN THIS ISSUE

Mate-locating behavior of the

western North American butterflies

James A. Scott 1

History of scientific study on
a larval color polymorphism in
the genus Chlosyne (Nymphalidae)

Raymond W. Neck 41

The butterfly faunas of San Andres and

Providencia Islands in the western Caribbean

Thomas C. Emmel 49

A list of the butterflies of the Willow River

State Park, Wisconsin

John H. Masters

57

Live Geometrid

Noel McFarland

60

Volume 14 Number 2 May, 1976

published by

The Lepidoptera Research Foundation, Inc.
at

1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz

Associate Editors :

Thomas C. Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601

Maria JEtcheverry, Centro de Estudios Entomologicos, Casilla 147, Santiago,
Chile.

T. N. Freeman, Div. of Entomology, Dept, of Agriculture, Ottawa, Ontario,
Canada.

Brian O. C. Gardner, 18 Chesterton Hall Crescent, Cambridge, England.

Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.

Lee D. Miller, The Allyn Museum of Entomology, 3701 Bay Shore Road,
Sarasota, Florida, 33580.

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STATEMENT OF OWNERSHIP AND MANAGEMENT
THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a
year. Spring (March), Summer (June), Autumn (September), and Winter (December)
by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
cation and the general business office are located at 1160 W. Orange Grove Ave.,
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Journal of Research on the Lepidoptera

14(2):61»83, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

THE GENETICS OF SUBSPECIFIC
PHENOTYPE DIFFERENCES

IN PIERIS OCCIDENT ALIS REAKIRT AND OF VARIATION
IN P. O. NELSONl W. H. EDWARDS (PIERIDAE)

ARTHUR M. SHAPIRO

Department of Zoology, University of California, Davis, California 95616

The Western White, Pieris occidentals Reakirt, has one of
the widest latitudinal ranges of any nearctic butterfly — from near
36° N above 3000 m in the White Mountains of Mono and Inyo
Counties, California to 69 °N at sea level at Umiat, Alaska. The
northernmost population to be studied biologically is located at
Fairbanks, Alaska (64°49' N); its bionomics are described in
Shapiro (1975a) and its phenotypic and developmental responses
to photoperjod in Shapiro ( 1975b ) . This population corresponds
to the taxon Pieris nelsoni W. H. Edwards (type locality St.
Michael, Alaska), characterized phenotypically by a reduced
black pattern dorsally in the male as compared with nominate
P. occidentals from California or Colorado (figs. 1, 2). Pieris
nelsoni was described from a single male in 1883 and then “lost”
for 91 years-, during which time its female remained unknown
along with its degree of relationship to P. occidentals. Following
its rediscovery at Fairbanks, P. nelsoni was reared in quantity
for photoperiodism studies, which revealed a longer-day thresh-
old for diapause induction and a reduced propensity to pro-
duce the estival phenotype (lightly marked ventrally) as com-
pared with P. occidentals (Shapiro, 1975b). However, its early
stages were indistinguishable from occidentals and the pattern
of the female differed from that taxon only very subtly. These
facts suggested strongly that nelsoni
graphic subspecies of a single
the crosses reported below.

and geO"

s^ahl i^Othesis , tested by

62

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. la —Dorsal surfaces of wild Pieris occidentalis nelsoni from Fairbanks,
Alaska, July 1974. Males at left.

14(2):61-83, 1975 GENETICS OF PIERIS

63

Fig. lb. — Ventral surfaces of same.

64

ARTHUR M. SHAPIRO

/. Res. Lepid.

Pig. 2. — ^Dorsal and ventral surfaces of wild Pieris occidentalis from Don-

ner Pass, California. Males at left.

14(2):61~83, 1975

GENETICS OF PIERIS

65

In breeding F. nelsoni from Fairbanks I obtained a number
of aberrant individuals which were mated and studied in con-
tinuous culture. These are described, figured, and discussed
elsewhere in this paper.

MATERIALS AND METHODS
The initial cross of Alaskan and Californian stocks was made
using a fresh wild male F. occidentalis collected at 3000 m at
the base of Sonora Peak, Mono County, California on 8 August
1974 by S. R. Sims. This male was caged with two virgin female
F. nelsoni of the first generation in continuous culture, ex Fair-
banks. This stock was maintained at 25° C under continuous light
and reared on fresh Lepidium virginicum L. ( Cruciferae ) . The
wild male was not very vigorous but did mate with one female
on the second day, and this female laid 8 eggs, of which 5
hatched. All of these eventually pupated, but two subsequently
died, producing an Fi of only three individuals^ — two males and
a female. A sib mating was, however, obtained, producing 49
ova of which 40 hatched, yielding 37 pupae and 25 adults of
the F2.

All rearing was done in colorless plastic Petri dishes (diam-
eter 14 cm) on L. virginicum at densities of 10 larvae or less/dish
at 25° C under a 60-W incandescent lamp. They pupated in the
dishes, and only non-diapause pupae were obtained. Eclosion
took place in nylon-mesh cages 41 cm x 41 cm x 41 cm, and
matings were obtained in these in filtered sunlight. All adults
used for breeding were provided fresh dandelions as a nectar
source. Mated females were confined in colorless plastic cages
19 X 13 x 10 cm with fresh cuttings of Lepidium and fresh
dandelions and allowed to oviposit.

RESULTS OF HYBRIDIZATION
The Fi consisted of two males, both intermediate between
estival and vernal phenotypes, with complete occidentalis pat-
tern, and one female with a dark-intermediate ventral pattern
and a dorsal occidentalis pattern which, however, resembled
nelsoni in having the spot at the end of the cell of the forewing
narrow. All of these individuals became too battered to be
figured.

The Fo (figs. 3, 4) of 13 males and 12 females was exceed-
ingly variable in all phenotypic characters, with seemingly inde-
pendent assortment at several genetic loci. Within each sex, the

66

ARTHUR M. SHAPIRO

J. Res. Lepid.

Fig. 3a.— Dorsal surfaces of F2 males, Alaska X California Pieris occiden-
talis, reared at 25°C on continuous light.

14(2):61-83, 1975

GENETICS OF PIERIS

67

Fig. 3b.-“Same but ventral,

wff.

68

ARTHUR M. SHAPIRO

/. Res. Lepid.

» ^ f#

Fig. 4a. — Dorsal surfaces of Fg females, Alaska X California Pieris occiden-
talis, reared at 25° C on continuous light.

14(2);61-83, 1975

GENETICS OF PIERIS

69

Fig. 4b. — Same but ventral,

70

ARTHUR M. SHAPIRO

J. lies. Lepid.

phenotypic range includes apparently normal occidentalis and
nelsoni, plus recombinants never seen in wild samples or pure
bloods of either. Because of the small numbers of animals and
the impracticability of large-scale genetic experimentation with
these traits, only a very rough idea of their inheritance can be
given. The traits which appear to be segregating are:

(1) Dorsal pattern of male. In nelsoni, the apical and sub-
apical pattern elements are suppressed. This phenotype appeared
only in one male of the Fo, but several others had these black
markings smaller than most male occidentalis. The sample is too-^
small to discriminate between control by one locus, with the
nelsoni allele recessive (expectation 1/4 nelsoni), and two loci
(expectation 1/16 nelsoni).

(2) Size and shape of spot in discal cell of forewing. This
trait is variable in both parental populations. The most frequent
nelsoni state is “narrow” in both sexes; “narrow” is, however,
rare in pure occidentalis. “Narrow” spot appeared in the Fi
female and in four males and four females of the Fg. The males
included the one with nelsoni dorsal pattern.

(3) Dorsal pattern of female. Nelsoni females do not differ
from California occidentalis in any conspicuous character. They
are more likely to have the black chevrons between the veins of
the hindwing darker than the remainder of the pattern; the
forewing more often has a complete (although narrow) black
outer margin, and the white spots enclosed in the forewing
border are usually broader in nelsoni than in occidentalis. The
spot in cell Cuo is almost always smaller in female nelsoni and
is occasionally lacking. In the Fo, all of these characters are
very variable. In three Fg females and in the one Fi female, the
forewing border is broader than in either parental population,
producing very elongate, narrow white spots enclosed within it.
In five of the 12 Fv females, the spot in cell Cu2 is reduced as
in nelsoni; in three it is heavier than in the average occidentalis.
Both parental populations are variable as to basal dark shading,
as is the F2. Control of female pattern is probably multifactorial
and not an expression of the same major gene(s) involved in the
male phenotypic difference.

(4) Seasonal Phenotypes. As noted in Shapiro, 1975b, nelsoni
produces an estival phenotype (light ventrally), less often than
does occidentalis when reared under continuous light. Again,
the F2 is extremely variable, and the variation is poorly cor-
related with characters on the dorsal surface. On the whole,

14(2):61~83, 1975

GENETICS OF PIERIS

71

ventral melanization in both sexes in the F2 is more similar to
nelsoni than to Sierran occidentalis. In nelsoni, there is a ten-
dency- for the heaviest melanization to be basal, and this is
found in four males and one female of the Fs. One of the males
is the single individual with dorsal nelsoni phenotype.

One F2 female is asymmetrical. This female was not recog-
nized as abnormal until after her death, but her phenotype is
essentially identical to the trait ‘Tilaterah’ previously found in
Pennsylvania-New Jersey stock of the closely related species,
Pieris protodice Boisduval and LeConte (Shapiro, 1970). This
trait is inherited as an autosomal recessive. Some of the modi-
fications produced in the homozygous ‘TilateraF protodice are
related to the 'ray” phenotypes discussed below.

ABNORMAL PHENOTYPES

The Fairbanks stock of F. nelsoni was started with five wild
females, whose ova were pooled. In the Fi, a number of aberrant
individuals of both sexes were obtained, and several matings
set up among them. The aberrant phenotype is apparently in-
herited as a simple autosomal dominant. It is expressed in both
long- and short-day (estival and vernal) phenotypes (figs. 5, 6).
Its differences from "wild type” are: Males: The submarginal
black markings on the dorsal forewing are even more reduced
than in “wild type” nelsoni, but the marginal triangles are present
and may be slightly enlarged. The spot in cell Cu2 is always
absent and that at the end of the cell is slightly enlarged.
Ventrally, the basal markings are more heavily melanized on
the hindwing than “wild type” but the submarginal chevrons are
only very weakly indicated. The spot at the end of the hindwing
cell ventrally is enlarged. Females: The submarginal elements
of the dorsal pattern are more or less reduced; as a minimum,
the chevrons on the hindwing are not darker than the rest of
the pattern and are thinner than in “wild type” nelsoni, making
the enclosed white spots appear larger than normal; in estival-
phenotype females the chevrons are usually obsolete. The spot
at the end of the cell of the forewing is conspicuously enlarged
(and ventrally may be prolonged basad along the cubitus, but
not as a ray actually reaching the base). As in males, the basal
melanization is exaggerated on the ventral hindwing.

In each of the six broods involving this phenotype reared
in this study, a few highly abnormal individuals appeared in
which the characters described above were exaggerated, along
with distortion of the wing shape and (often) a reduction in

72

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. 5a. — Aberrant phenotypes of male Pieris occidentalis nelsoni, dorsal
surfaces, illustrating light (estival) and dark (vernal) pattern.

14(2):61~83, 1975

(;enetics of pieris

73

Fig. 5b. — Same but ventral.

74

ARTHUR M. SHAPIRO

]. Res. Lepid.

Fig. 6a. — Aberrant phenotypes of female Fieris occidentalis neloni, dorsal
surfaces, illustrating light (estival) and dark (vernal) pattern.

14(2):61-H3, 1975

CENETICS OE PIERIS

75

Fig. 6b. — Same but ventral

76

ARTHUR M. SHAPIRO

J. Res. Lepid.

the number of antennal segments. Some of these are illustrated
in figures 7 and 8. These individuals were generally quite
vigorous and lived 2-8 days, but most were unable to fly and all
failed to mate, although the females were courted by normal
males. These extreme phenotypes would presumably be lethal
in Nature.

No wild specimens similar to those produced by this Fair-
banks allele have been seen from any Pieris occidentalis popu-
lation. Apparently, the male which mated with one of the
founder females was heterozygous for it, but no examples were
found among the 18 specimens collected at Fairbanks. The
phenotype has some similarity to that of the mutant 'ray”
described from New York City stock of P. protodice by Shapiro
(1973). The most extreme individuals produced in that line
also somewhat resemble those reported here (fig. 9). However,
“ray” is inherited as recessive. The dark “ray” along Cu on the
forewing ventrally is perhaps produced in a variety of ways
in this species group, as it occurs consistently (in females only)
of Colorado P. o. occidentalis from above timberline and is
inherited as a sex-limited dominant in crosses with Sierran
stock (Shapiro, unpublished data).

Homozygous “ray” females often show stripping of scales
along the outer margin below the forewing apices dorsally
(Shapiro, 1973). This trait was observed in many Fairbanks
specimens, both “wild type” and aberrant, and with equal fre-
quency in both sexes. It may reflect adhesion of the wing to
the pupal cuticle, such as occurs in the “albinensis” trait in
Pieris brassicae Linnaeus (Gardiner, 1962). No difficulties with
eclosion were observed in the pure Fairbanks line, but Gardiner
was able to select for a line of P. brassicae in which “albinensis”
eclosed normally, and the same selection may have occurred
naturally at Fairbanks. When this trait was recombined with
a California genome the system could have been disrupted,
thereby contributing to the unusually high mortality of fully
pigmented, ready-to-eclose pupae in the F2. Only one adult
which eclosed, the nehoni-phenotype male, displays it.

The nature of the abnormalities observed in the Fairbanks
material parallels those found in mass-reared stocks of P.
protodice closely enough that one may assume similar develop-
mental pathways are involved, whether or not the genetic
control is homologous. Curiously, these abnormalities have not
yet been seen in California P. occidentalis.

l4(2);61-83, 1975

GENETICS OF PIERIS

77

DISCUSSION

The hybridization experiments leave little doubt that Pieris
nelsoni Edwards 1883 and P. occidentalis Reakirt 1866 are
conspecific. By the International Code o£ Zoological Nomen-
clature, the former should be considered a subspecies of the
latter, and written as Pieris occidentalis nelsoni (this combi-
nation was used by dos Passes, 1964). Left unresolved is the
relation of both to P. callidice Hiibner 1805, of the Palaearctic
region. This relationship can be clarified only by genetic ex-
perimentation.

The small size of the Fi and Fg Fairbanks x California broods
precludes any convincing statistical analysis of the inheritance
of phenotypic differences between the stocks, but it is evident
that they must differ at several loci influencing color and pattern.
In the case of differences in thresholds for diapause and pheno-
typic induction, these differences are clearly adaptive for the
subarctic nelsoni. Even the aberrant phenotypes recovered from
the Fairbanks stock would be potentially adaptive if the con-
centration of melanin at the hind wing base ventrally contributes
to body warming during lateral basking. Strikingly, the reduc-
tion of the dorsal black pattern in the subarctic nelsoni parallels
the appearance of extreme vernal phenotypes of P. protodice
which ffy in early spring in both eastern and western North
America, suggesting an inherent advantage to this pigmentation
under low-temperature conditions.

The hybrid broods are again too small for much to be said
with confidence concerning the strength of postzygotic barriers,
except that the fertility and viability of the F2 did not depart
significantly from many pure broods of both parental stocks
when reared under high temperatures and continuous light.
The sex ratio of the F2 was normal. In butterfly hybrids in-
volving genetic barriers there is usually a deficiency of females
(the heterogametic sex). As noted previously, the high inci-
dence of pupal mortality in the F2 was not duplicated in the
pure Fairbanks line but has been seen in some lots of Sierran
pupae ' held at high temperatures in continuous culture; in
these cases, however, most of the mortality occurs before adult
pigment is laid down, while in the F2 it was after.

78

ARTHUR M. SHAPIRO

J. Res. Lepid.

Fig. 7a. — Dorsal surfaces of extreme aberrant Fieris occidentalis nehoni
males.

14(2);61-83, 1975

c;enetics of pieris

79

Fig. 7b. — Same but ventral.

ARTHUR M. SHAPIRO

]. Res. Lepid.

Fig. 8a. — Dorsal surfaces of extreme aberrant Pieris occidentalis nelsoni
females.

14(2):61-83, 1975

(;E NET ICS OF PIERIS

81

Fig. 8b.— Same but ventral.

82

ARTHUR M. SHAPIRO

J. Res. Lepid.

Fig. 9. — -Selected examples of extremely aberrant Pieris protodice of the
‘‘Ray'* line for comparison with Fairbanks stock of P. o. nelsoni (see
Shapiro, 1973),

14(2):61-83, 1975

CENETiCS OF PIERIS

83

SUMMARY

1. Hybridization experiments demonstrate the apparent con-
specificity of the taxa “Fieris nelsonf from Alaska and P.
occidentalis from the Sierra Nevada of California.

2. Three bred Fi hybrids between these taxa were basically of
the California phenotype, but within an Fg of 25 individuals
phenotypic variation was very great and recombination
among several loci was evident.

3. An aberrant phenotype of P. o. nelsoni was obtained from
the Fairbanks stock; it is inherited as an autosomal dominant.
A few individuals exhibit functionally lethal exaggerations
of the characteristics of this phenotype.

4. Aspects of the genetic variation of P. o. nelsoni are shown
to parallel the related species P. protodice, which was studied
previously.

ACKNOWLEDGMENTS

Collection of stocks was supported by Grant D-804 from the
Committee on Research of the Academic Senate, University of
California, Davis.

Mr. S. R. Sims assisted in field work in California and Mrs.
A. R. Shapiro did so in Alaska.

LITERATURE CITED

DOS PASSOS, C. F. 1964. A synonymic list of the Nearctic Rhopalocera.

Lepidopterists Society Memoir 1. 145 pp.

GARDINER, B. O. C. 1962. An albino form of Pieris hrassicae Linnaeus
( Lepidoptera, Pieridae). Entomologist’s Gazette 13: 97-100.

SHAPIRO, A. M. 1970. Inheritance of three genes affecting the wing pat-
tern of Pieris protodice Boisduval and LeConte (Lepidoptera: Pieri-
dae). Wasmann Journal of Biology 28: 245-257.

— — - — . 1973. Genetics of two aberrant patterns in Pieris protodice Bois-
duval and LeGonte (Lepidoptera: Pieridae). Wasmann Journal of Bi-
ology 31: 301-312.

. 1975a. The biology of a “weedy” butterfly, Pieris occidentalis

(Lepidoptera: Pieridae) at Fairbanks, Alaska. Arctic and Alpine Re-
search 7: 273-276.

—. 1975b. Photoperiodic control of development and phenotype in a

subarctic population of Pieris occidentalis (Lepidoptera: Pieridae).
Canadian Entomologist 107: 775-779.

Journal of Research on the Lepidoptera

14(2) :84, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

EARLY STAGES OF PHYCIODES PALLIDA,

P. ORSLIS, AND P. MYLZTTA (NYMPHALIDAE)

JAiMES SCOTT

Department of Entomology, University of California, Davis, California

Scott (1973) described early stages of P. orseis, compared
them with published descriptions of early stages of P. mylitta,
and speculated that P. orseis might be most closely related to
F. pallida.

This speculation has proven false. P. pallida from Red Rocks,
Jefferson County, Colorado, was raised on Cirsium vulgare.
P. mylitta from Thompson Canyon, Yolo Co. Calif, and near
Copper, Siskiyou Co. California were raised on Silybum marP
anum. P. pallida differs in having later instar larvae ochre in
color rather than maroon black as in P. orseis or light maroon
brown as in P. mylitta. P. pallida mature larvae have a dorsal
brown line, a brown band through the supraspiracular setae,
and a few other brown spots. Mature larvae of all three species
have the dorsal white stripe on the head. Mature larvae of P.
pallida lack the subdorsal light spot on the head present in P.
orseis, but have a light area just above the ocelli ( absent in
P. orseis). P. mijlitta rarely have the subdorsal light spot and
rarely have the supraocellar light patch.

I conclude that phenetically P. orseis and P. mylitta are most
similar to each other, and P. pallida is more distantly related
but is more similar to P. mylitta than to P. orseis.

LITERATURE CITED

SCOTT, J. A. 1973. Early stages and biology of Phyciodes orseis (Nym-
phalidae). /. Res. Lepid. 12: 236-242.

84

Journal of Research on the Lepidoptera

14(2):85-89, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

RECENT CAPTURES OF
ANTHOCHARIS CETHURA CATALINA
MEADOWS

LARRY J. ORSAK

Center for Pathobiology, University of California, Irvine, California 92664

The Catalina Orange-tip, Anthocharis cethura catalina
(Pieridae), was described by Meadows (1936) from a series of
eight males and four females, collected from 1928-1933 at various
locations on Santa Catalina Island, California (Grand Canyon,
Whites Landing, Renton Mine area, Salta Verde, and Little
Harbor). The holotype (from Grand Canyon) and allotype are
currently in the Natural History Museum of Los Angeles County
collection, and a series of paratypes (one male, two females)
are in the University of California, Irvine Collection (Charles
Rudkin collection). Other paratypes are housed in a portion of
the Don Meadows collection, currently at the Bowers Museum
(Santa Ana, California). Strangely enough, over forty years
later, these twelve specimens were still the only known (or
publicized) representatives of this subspecies. Some Lepidop-
terists had postulated that the subspecies may have become
extinct in the intervening years since its description (Emmel
and Emmel, 1973). In fact, at the time of this writing, the Office
of Endangered Species (Washington, D. C.) was investigating
the possibility of placing the Catalina Orange-tip on the En-
dangered Species List (Federal Register, 40(55) : 12691).

On a recent trip to the island, the author was able to verify
the continued existence of Anthocharis cethura catalina and
dispel any rumors to the contrary. Since almost nothing is known
about the habits of this small desert butterfly which managed
to attain an island foothold, it seems worthwhile to add my
observations in the hopes that other Lepidopterists will be
moved to initiate other studies on this local subspecies.

The author, accompanied by another collector (C. R. Rose-
land), visited the Avalon area on 20 April, 1975, collecting from
1100 to 1530 PST (daylight savings). Calm, warm (74° F. )

85

86

LARRY J. ORSAK

J. Res. Lepid.

weather prevailed, the sun being obscured but briefly during
this time by haze. Meadows (1936) had stated that “the most
accessible locality where catalina may be collected is a small,
open meadow at the top of a ridge between Renton Mine and
Jewfish Point, two miles south of Avalon.” Thus, it was decided
to try to reach this locality.

We started a grueling cross-country hike up the steep hills
south of Avalon, moving up a fire break at the top of the second
ridge from the ocean. Not one butterfly was seen during the
first hour of hiking. However, as the sun warmed the island, a
few Celastrina argiolus echo (Lycaenidae) began to fly in the
chaparral areas. The first Catalina Orange-tip (so we assumed)
was sighted at time 1215, flying along the steep south-west
slope of the ridge, at an elevation of 500 feet. The slope, covered
with a rich carpet of grass and other annuals at this time of
year, also had vigorous growths of California Sage {Artemesia
calif or nica) , with some sumac (Rhus laiirina) and Opuntia also
present. The specimen which was sighted proved impossible
to collect. We continued up the ridge via the firebreak until
we reached an elevation of approximately 650 feet. It was then
that I saw the first Avalon Hairstreak of the day (Strymon
avalona). At the same time, however, we sighted two orange-
tips flying up the northeast slope and over the ridge. Neglecting
avalona, we managed to capture both butterflies in flight, and
a glance at the ventral wing pattern left no doubt that we had
just captured cethiira catalina! The specimens captured turned
out to be two males. We continued up the ridge to an
elevation of about 1100 feet without additional captures of the
Catalina Orange-tip, although we observed or captured a few
avalona flying about the long-dead blossoms of Rhus lamina,
one Orange-tip, Anthocharis sara gunderi (another island en-
demic), two Papilio zelicaon, and 2-3 additional orange-tips that
may have been catalina. When the ridge twisted, facing South,
we followed, sighting perhaps 2-4 Anthocharis, although none
could be captured, due to their swift and erratic flight over the
rough terrain.

As we started hiking in the southern direction, down a ridge
adjacent to the southwest of the one on which we had captured
our first catalina (a course leading back to Avalon, as we no
longer had time to pursue our hike to the Meadows locality near
Renton Mine), we began to see considerably more Anthocharis.
This ridge was covered with dense impenetrable chaparral flora,

14(2):85-89, 1975

ANTHOCHARIS C. CATALINA

87

broken only occasionally by grassy areas. The majority of the
orange-tips captured, however, were A. sara gunderi. However,
at an elevation of 1100 feet (time approximately 1445), a male
and female catalina were seen flying up the slope and were cap-
tured. The female was somewhat aberrant in that the normal
dorsal orange marking was considerably reduced, being retained
only along the veins in the subapical area, giving a somewhat
streaked appearance. Both specimens were in fresh condition.

The remainder of the trip proved uneventful.

Several statements can be given concerning the habits and
status of cethura catalina, even though our excursion to cata-
linds habitat was brief. First and foremost, the Catalina Orange-
tip is still extant on Santa Catalina Island. Additionally, fear
that the subspecies was presently endangered appears unfound-
ed. The butterfly was found on two ridges, half a mile apart,
adding to the already know extensive, although spotty distribu-
tion of the subspecies on the island (see Figure 1). The Cata-
lina Orange-tip may probably be expected to occur on many,
if not most, of the rather inaccessible ridges in the area near
Avalon. We should note that the butterfly has managed for
years to survive the onslaught of the environment by goats
(introduced by the Spanish hundreds of years ago) and wild
boars, and it is doubtful that the butterfly will succumb to any
analogous habitat changes (grazing, recreational use). Indeed,
the butterflies seemingly were not affected at all by the defoli-
ated firebreak up the first ridge, and were sighted more often
on this ridge than on the more natural appearing adjacent ridge.
Further observations on the populations, however, are required
before the true status of the subspecies can be determined.
Commercial development is not being planned for any catalina
habitats, most of the island now being a preserve, and thus the
butterfly is probably in no danger for many years to come.

No strong hilltopping behavior was observed. Perhaps as
many individuals were seen fifty or more feet down the slopes
of the ridges as were seen anywhere near the top, hence why
so few specimens could be procured. The erratic flight typical of
cethura, however, was observed in catalina.

The date of our captures is of interest, particularly in regard
to the freshness of specimens. Meadows (1936) captured cata-
lina from March 23 to April 9, his last capture being almost two
weeks previous to ours. The flight period given by Emmel and
Emmel (1973) and Meadows (1936) should thus probably be

^ CATAI ;MA i^LAhlh

88

LARRY J. ORSAK

]. Res. Lepid.

14(2);85-89, 1975

ANTHOCHARIS C. CATALINA

89

amended to read “late March to late April”. Late Spring rains
in 1975, however, could have caused an unusual delay in the
adult emergence of catalina, as such delays were reported for
many southern California butterflies during this season.

It is rather surprising that captures of the Catalina Orange-
tip have not been reported in so many years, although numerous
Lepidopterists have visited Santa Catalina Island in order to
procure specimens of the other two endemic butterflies found
on the island. This may be attributed to the past and present
inaccessibility of catalina habitats (physical and/or legal), short
flight period of adults, possible confusion with Anthocharis sara
gunderi, and the tendency of local collectors not to exert much
effort searching for a relatively unspectacular-appearing butter-
fly (versus the amount of effort expended to find the legendary
lost Atossa Fritillary, Speyeria adiaste atossa, of southern Cali-
fornia ) .

I would like to acknowledge the help of Mr. Craig Roseland
in securing specimens, Mr. John Haynie, our pilot to the island,
and the Xerces Society, which has provided financial and moral
support for studies in rare and endangered southern California
Rhopalocera. A voucher specimen of ceihura catalina from this
trip will be deposited in the collection of the Natural History
Museum of Los Angeles County.

AUTHOR’S NOTE: Three additional records of the Catalina
Orange-tip came to light as a result of the distribution of pre-
liminary drafts of the above article, but were received after the
article went to press. Dr. Charles Remington informs me of
the existence of two specimens collected by Charles Ingham
and purchased through the Ingham collection, now housed at
Yale University. The only recent record to come to light in
addition to my collecting experiences is that of Dr. Jerry Powell
of the University of California, Berkeley, and Dr. Paul Opler
(presently of the Office of Endangered Species, Washington,
D.C.). On April I, 1968, a specimen of catalina was taken by
them at Little Harbor during a lull in generally bad weather.
This fairly recent capture indicates the continued existence of
catalina from more than one locality on Santa Catalina Island.

LITERATURE CITED

EMMEL, THOMAS C., and JOHN F. 1973. The Butterflies of Southern
California. Natural History Museum of Los Angeles County. 148 pp.
-f- xii.

MEADOWS, D. 1936 (1937). An Annotated list of the Lepidoptera of
Santa Catalina Island, California. Bull. South. Calif. Acad. Set. 35:175-
180.

Journal of Research on the Lepidoptera

14(2):90-92, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

OLD TIMERS

JOHN ADAMS COMSTOCK

ENRICO PIAZZA

One of the first English lepidopterists that I met here in
Southern California was Enrico Piazza. He was collecting for
Rothschild of the British Museum.

His actual name was Henry Place, but he went to Italy early,
was a grand opera singer, and Italianized his name for profes-
sional reasons. When his voice gave out, he commercialized his
hobby, and came to Southern California. I visited him in a
cottage which he had rented in San Diego, and admired the
fine condition of the moths he was preparing for his customers.

Four species of California moths were named for him, namely:
Tornos piazzata Cassino & Swett.

Cosymbia piazzaria W. S. Wright
Raphia piazzi Hill.

Myelois piazzella Dyar.

He became ill, and started for England, but died in New York
of cancer. Nothing was known of his effects, and his key collec-
tion was lost. His types had probably been sent to England.

I have not been able to locate essential facts concerning his
background.

FORDYCE GRINNELL JR.

Fordyce was our most prominent and active lepidopterist when
I arrived in Southern California. He was living in Pasadena
with his parents. He was born June 17, 1882, in Pine Ridge,
South Dakota, son of Dr. Fordyce Grinnell Jr. and Elizabeth
Pratt Grinnell.

His brother was Joseph Grinnell Ph.D., a noted ornithologist
and Faculty member of the University of California.

Fordyce was the founder of the Lorquin Natural History So-
ciety (now the Lorquin Entomological Society.) He was very
active in collecting lepidoptera and had many local neophytes
whose leadership he followed from about 1900 to 1914, when he
entered Stanford University.

Read by John A. Comstock at the Association Banquet, evening of June 19, 1965.

90

91

JOHN ADAMS COMSTOCK

J. Res. Lepid,

He obtained his Bachelor degree in 1918, and went to Hawaii
in the same year. He taught there until 1921 when he returned
to Stanford.

Then, in 1922 he went to the Philippines where he served
under the Bureau of Education of the Philippines, as a teacher.

He resigned in five months because, in his own words “he
was unsuited to the official red tape and the classroom grind.”

Thereafter he collecter butterflies and beetles for the Bureau
of Science in Manila, and the Hill Museum in Surry, England.
This took him through various portions of the Philippine Islands
until March of 1923, when he sailed for Australia.

He collected specimens in widely separated parts of the con-
tinent, interspersed with odd jobs to replenish his funds, until
1930, when he arrived in Cooktown, Australia. There he received
a letter from his mother asking him to return home.

Mrs. Grinnell was then living in Sausalito, a widow, 80 years
of age. Fordyce lived with her until he married Mary D. Walters
on August 10, 1937. The ceremony occurred in Muir Woods.

I have no further record of his activities until he died in San
Francisco, July 20, 1943.

He was author of the following species of butterflies:

Anthocaris lanceolata australis. Cercyonis behrii. Vanessa
atalanta ab. edwardsi. Vanessa carye ab. intermedia. Van-
essa canje ad. letcheri. Strymon sylvinus desertorum. Ple-
bejus emigdionis. Glaucopsyche lygdamus australis. Erynnis
persius pernigra. Erynnis callidus $ (nec. $ ) (syn. lacustra
callidus 2 partim) Polites sabuleti tecumseh.

WILLIAM SHERMAN WRIGHT

William S. Wright was an early staff member of our own San
Diego Museum of Natural History. He served as Curator of
Insects from 1922 to 1933, in addition to holding the important
post of County Supervisor of Nature Study. He was a specialist
in the Geometridae, and his own collection of insects, with his
types, served as the nucleus of our present Entomological Divi-
sion.

He was a profound student, a capable administrator, and a
beloved teacher and friend.

A bronze plaque commemorating his generosity and helpful-
ness to our Museum may now be seen in the Entomological
Laboratory.

14(2);90-92, 1975

OLD TIMERS

92

He was born in Plaino, Illinois, April 23, 1866, and died in
Laguna Beach, California, July 8, 1933.

I knew him well, and frequently collected with him, but,
regretfully failed to make notes of the background of his history
and family. There is little that I can find in the biographies.

He published several new species, and a number were named
for him, but I have not had time to list them.

WILLIAM GREENWOOD WRIGHT

Most of us have heard of the colorful pioneer lepidopterist of
our early Southwest, but very few have seen his monument
in the shape of his volume on “THE BUTTERFLIES OF THE
WEST COAST”.

The tragic reason is that “at the great fire in San Francisco,
April 18, 1906, all the items going to make up this book . . .
consisting of everything in the printers and the binders hands,
and the finished books . . . all were destroyed.”

I have one of these books, thanks to the fact that Wrights
possession were given to the California Academy of Sciences in
San Francisco, including the few books that were in San Ber-
nardino when the fire occurred. I was able to buy one of these
volumes, and you can now take a peek at it.

W. G. Wright was born in 1830, and died in 1912. There may
have been memorial notices of him, but I have failed to find
only the one by Fordyce Grinnell in the Entomological News,
Vol. 24, pp. 91-92, 1913. My notes contain mainly hearsay from
people who knew him. He was a plaining mill owner, whose
hobby was butterfly collecting, and had had very little formal
schooling. His book evidences that fact, but it also shows that
he had enthusiasm and purpose. Knowing his handicaps, I con-
sider that he did a remarkable piece of work.

There are many errors, to be sure, but others have pointed
these out, as, for instance, Henry Skinner and Fordyce Grinnell
in the Entomological News in Vol. 16, pp. 336-340, 1905.

Journal of Research on the Lepidoptera

14(2):93-97, 1975

1160 W. Orange Grove Ave., Arcadia^ California 91006, U.S.A.

© Copyright 1975

WHY DO CALIFORNIA
TORTOISESHELLS MIGRATE? '

ARTHUR M. SHAPIRO

Department of Zoology, University of California, Davis, California 95616
The California Tortoiseshell butterfly, Nymphalis call-
fornica Boisduval ( Nymphalidae ) , is well known to entomok
ogists and laymen alike on the Pacific Coast for its mass move-
ments. The biological basis for these movements has always been
obscure. After discussing notable outbreaks in the Yosemite
region, Garth and Tilden (1963) say: 'The explanation seems
to be that the California Tortoiseshell is a swarming species
which, like the lemming, has cycles of abundance followed by
a drastic reduction in the population . . .” But this is obviously
no explanation at all. Powell (1972) wrote that "the records
suggest that this species periodically develops an imbalance
with factors in its population equilibrium at isolated sites,
followed by mass emigration of adults in various directions. . . .
Nymphalis californica should not be considered a migratory
species except in the broadest sense.” This is a more definitive
statement, but it also falls short of being an explanation. Mass
movements of butterflies, as Klots (1951) observes, have been
attributed to "population pressure” and “parasite pressure.”
Just what do such “explanations” mean?

Let us assume that the word “migration” is applicable to
N. californica as I intend to show it is. If we ask “Why do
California Tortoiseshells migrate?” we are not asking a simple
question. Ernst Mayr ( 1961 ) pointed out in a classic paper
that any “why” question in biology may be answered at several
levels. Mayr actually addressed himself to a question about
migration: “Why did the warbler on my summer place in New
Hampshire start his southward migration on the night of the
25th of August?” Mayr perceived at least four equally legitimate
levels of causality:

1) an ecological cause. “The warbler, being an insect eater,
must migrate, because it would starve to death if it should try
to winter in New Hampshire.”

‘Address prepared as invitational research lecture, 1975 Summer Advising
Conference, UC Davis.

93

94

ARTHUR M. SHAPIRO

/. Res. Lepid.

2) a genetic cause. “The warbler has acquired a genetic con-
stitution in the course of (its) evolutionary history which
induces it to respond appropriately to the proper stimuli from
the environment.”

3) an intrinsic physiological cause. “The warbler . . . responds
to the decrease in day length and is ready to migrate as soon
as the number df iioiirs of daylight has dropped below a certain
level.”

4) an extrinsic physiological cause. . . sudden drop in tem-
perature and associated weather conditions affected the bird,
already in a general physiological readiness for migration, so
that it actually took off on that particular day.”

Mayr groups (3) and (4) as proximate causes of migration —
the immediate triggering mechanisms. Causes (1) and (2) he
calls ultimate causes — “causes that have a history and that have
been incorporated into the system through many thousands of
generations of natural selection.” Clearly a physiologist, asked
our “why” question, would refer to proximate causes; an evo-
lutionist, to ultimate ones. Equally clearly, cause (1) is the
basis for the natural selection which brought (2) into being,
and (3) is the phenotypic manifestation of the genetic infor-
mation in (2), and is brought into action by (4). This method
of causal analysis is theoretically applicable to any adaptation;
and by its use we may perhaps be spared the travail of endless
controversies over the significance of a phenomenon such as
hilltopping behavior in butterflies and other insects, or of
“territoriality” in anything. Let us now try to analyze levels of
causality in the light of what we know of Nymphalis californica
migrations — which is not much, but is considerably more than
many people think we know.

I have been watching Tortoiseshell migrations for the past
four years, and unlike most Tortoiseshell watchers, I have been
chasing them. When you follow their movements from place to
place — not by individual marking, which would truly be a
needle-in-the-haystack operation, but by keeping track of where
the front of the migration is on consecutive occasions — it becomes
clear that, at least at the latitude of Sacramento, Powell is dead
wrong: Tortoiseshells do not go in all directions; they have a
set seasonal directionality, with a spring-fall reversal. The con-
fusion in many published reports is based at least in part on the
“static observer” effect and on local eddies in the migratory
flow produced by topography. But California Tortoiseshells go

14(2);93-97, 1975

MIGRATION

95

north and east in May and June and south and west in Septem-
ber and October. The generalized pattern is for dispersal out
of the central Coast Ranges in spring — northward in the higher
ranges, especially from Colusa County north, eastward from the
lower ranges in Napa, Yolo, and Solano Counties, crossing the
floor of the Sacramento Valley and going up the Sierra foothills
east of Sacramento. Almost simultaneously Tortoiseshells migrate
out of the Sierra foothills, heading upslope in a N to NE direc-
tion. The two currents generally merge. The pattern is exactly
reversed in the fall; again the migrants can be seen crossing the
Valley floor, where they never breed (there being no hosts).
This pattern, first described in my 1974 paper based on 1972
observations, has been repeated unerringly in successive years;
and as I get more sophisticated at Tortoiseshell-watching, I am
getting better too at predicting the dates. They are rather
variable; for example, the eastward-moving spring front passed
through Davis on 26 May 1972, 9 June 1973, 6 June 1974, and
13 June 1975. Based on this small sample, the warmer and drier
the spring, the earlier the flight. These migrants are, of course,
not the adults which crossed the Valley the autumn before and
overwintered (very few Tortoiseshells seem to remain in the
lower Coast Ranges through the summer, and the hibernators
are mostly or all immigrants from the north or east). They are
their offspring. And the return migrants in fall are their offspring,
or even their grandchildren.

The regularity of this pattern suggests that it is an adapta-
tion, an attribute of the animal which promotes its welfare.
Being an evolutionist, I am most interested in the ultimate
levels of causality — the basis of natural selection resulting in
the acquisition of a genetic program which instructs the animal
to migrate in such and such a direction given such and such
(proximate causality) conditions.

Why should Nymphalis californica leave the lowlands in
both the Coast Ranges (which are often only foothills without
any mountains) and the Sierras? Of course, it could be heat-
intolerant. After all, its host plants, wild lilacs ( Ceanothus
species, Rhamnaceae), are green all summer; presumably it
could breed continuously in the lowlands if it “wanted” to.
Instead it leaves the foothills, with their Ceanothus species, to
go breed in the high country and the north, with a different
set of Ceanothus. It could be heat intolerance, acting directly;
but I think not. Being a Pierid specialist, with a dislike for

96

ARTHUR M. SHAPIRO

J. Res. Lepid.

Nymphalids as experimental animals, I do not want to test the
following hypothesis myself. It would make someone a nice
Ph.D. thesis, as well as being a good exercise in Mayr-style
causal analysis.

In 1970, P. P. Feeny, from Cornell, published a landmark
study of the role of host-plant chemistry in insect phenology.
Feeny found that the spring feeding season of Winter Moth
(Operophtera bnimata L., Geometridae) larvae coincided with
the period of minimal tannin concentration in their food — oak
leaves — and that the higher tannin concentrations characteristic
of mature, summer oak foliage interfered with nitrogen avail-
ability and perhaps leaf palatability to the larvae. Feeny was
thus able to develop a causal explanation of spring feeding by
Winter Moth larvae as an adaptive response to the seasonal
pattern of nutritional “availability” of oak foliage. The insect
faunas of high-tannin plants throughout the Northern Hemi-
sphere appear to show this effect. Shapiro (1975) gives a
schematic representation for some oak-feeding Lepidoptera in
New Jersey, for example. Similar schemata could be prepared
easily for Californian faunas on native oaks or on other high-
tannin plants, such as Cercocarpiis — or Ceanothus.

I am suggesting that California Tortoiseshells leave the low-
lands and go upslope in spring because the lowland Ceanothus
put on all their new growth in late winter-early spring and
become nutritionally unsuitable for breeding by June. The
higher one goes, the later the Ceanothus commence active
growth and hence the later they have young, tender, hypo-
thetically low-tannin foliage available for Tortoiseshell larvae.
The seasonality of lowland Ceanothus of course reflects the
arid-summer climate, so indirectly at least climate may be a
“cause” of Tortoiseshell migrations. But, in Mayr’s sense, the
ultimate cause — the ecological cause — would be the correlation
of geography and availability of Ceanothus foliage. In Cali-
fornia’s progressively drier Quaternary summer climate, a Ceano-
thus feeder unable to handle tannins has a “choice” between
being sedentary and univoltine or migratory and multivoltine;
I propose that N. californica has evolved along the latter course.

Sometimes the proximate and ultimate causes coincide, in
whole or in part. The proximate causes of Tortoiseshell migra-
tions could involve a response to the chemical or textural con-
dition of the plants. They could also be tied to photoperiod,
temperature, or some other seasonal indicator — we know not

14(2):93-97, 1975

MIGRATION

97

what At present, however, there is nothing to suggest that
either population density or ‘parasite pressure” has anything
to do with it. Based on my four years of careful observation,
I am willing to assert that migration occurs in the directions
described with population densities fluctuating by at least two
orders of magnitude. If the host-availability hypothesis holds
up, this would scarcely be surprising; inedible plants are inedible
whether there are a few or a lot of hungry caterpillars.

In recent years major strides have been made in the under-
standing of insect migration, e.g. the sophisticated studies by
Dingle (1968, 1972) and his colleagues on milkweed bugs
(Hemiptera, Lygaeidae) in North America. There is no reason
why butterflies should be any more difficult to unravel, especially
once we realize that explanations at different levels are not
mutually exclusive (but, rather, mutually complementary), and
that naming a phenomenon (“swarming species,” “cycles of
abundance”) is not the same as explaining it.

LITERATURE CITED

DINGLE, H. 1968. The influence of environment and heredity on flight
activity in the milkweed bug Oncopeltus. J. Exp. Biol. 48:175-184.

. 1972. Migration strategies of insects. Science 175:1327-1335.

FEENY, P. P. 1970. Seasonal changes in oak leaf tannins and nutrients as
a cause of spring feeding by winter moth caterpillars, Ecologii 51:565-

581.

GARTH, L S. and I. VV. TILDEN. 1963. Yosemite Butterflies. J. Res.
Lepid. 2:1-96.

KLOTS, A. B. A Field Guide to the Butterflies. Houghton Mifflin, Boston.
349 pp.

MAYR, E. 1961. Cause and effect in biology. Science 134:1501-1506,

POWELL, J. A. 1972. Population e.xpansions and mass movements of
Nymphalis californica ( Nymphalidae ) . J. Lepid. Soc. 26:226-228.

SHAPIRO, A. M. 1974. Movements of Nymphalis californica (Nymphali-
dae) in 1972. J. Lepid. Soc. 28:75-78.

. 1975. The temporal component of butterfly species diversity. In

M. L. Cody and J. Diamond, eds., Ecology and Evolution of Com-
munities (Proceedings of the Robert MacArthur Memorial Symposium,
Princeton, N.J., November 1973) (in press).

Journal of Research on the Lepidoptera

14(2):98-99, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

EARLY WORK ON THE MEGATHYMIDAE

JOHN ADAMS COMSTOCK

My interest in Lepidoptera began in 1894, at 11 years of age,
when living in Evanston, Illinois. This was appreciably stimu-
lated by correspondence with a young collector living in Pasa-
dena, California, named Fordyce Grinnell, whose address I
found in the Entomological News. This contact cherished my
hope that I would someday collect butterflies in California.

At that time I knew nothing of the ‘giant skippers’ called
Megathymiclae, since they did not occur in Illinois.

It was not until about 1920 that interest in the Yucca and
Agave borers was aroused by a chance acquaintance with Com-
mander Charles M. Daminers of Riverside.

Dammers brought his son Carlito to the Southwest Museum,
where I was then Director, and asked my assistance in teaching
the boy how to rear and illustrate the life histories of butterflies.
This led to the father’s interest in excess of the boy’s capabilities,
and resulted in Dammers cooperating with us in joint publication
of numerous articles on the biology of California lepidoptera.

In 1932 we were able to successfully rear Agathymus stephensi
(Skinner) on Agave desert i Engelman, and to publish its life
history in Volume 33, (2): pages 81-86, 1934, Bulletin, So. Calif.
Academy of Sciences.

This led us to an intensive effort to locate larvae of a Mega-
thymus, the images of which had been netted occasionally in
association with Joshua trees — Yucca brevifolia Engelman, on
the Mojave Desert. Commander Dammers and Carlito began a
search on the Joshuas one forenoon, but Carlito soon tired, and
stretched full length on the desert floor while Dammers climbed
assiduously. Suddenly Carlito noticed a female Megathymus
alighting on a baby Yucca plant of about 8 inches in height,
which proceeded to lay an egg. A shout brought the Command-
er to earth. Sure enough, an egg, and by further searching, more
on other young shoots. The problem was solved. The Mega-
thymus larvae were root-feeders underground, never on or in the
leaves.

98

99

JOHN ADAMS COMSTOCK

/. Res. Lepid.

It then became easy to locate the 'cones’ on top of infested
young plants, and dig down to the roots where the larvae were
feeding in late winter, or the pupae resting below the cones in
early spring.

The life history was published and illustrated in the same
issue of the Bulletin as was that of stephensi. It was listed as
Megathymus yuccae navajo, the name that had been given to me
by Dr. Henry Skinner on a specimen sent to him for determin-
ation. Later (1956) the subspecies was named by Stallings and
Turner, in the Academy “Bulletin” as martini, in recognition of
Lloyd’s fine work on the Lepidoptera.

Those of you who have worked on the Megathymidae recall
that our San Diego Museum Curator of Insects, Charles F.
Harbison, has described two species of Agathymus from Baja
California, namely comstocki and dawsoni, and that I have de-
scribed and illustrated the early stages of these. In addition, I
have similarly treated Megathymus evansi. The main reason that
I mention it is that, by comparison with the great amount of
research made by such experts as Don B. Stallings, J. R. Turner,
H. Avery Freeman, Dr. C. J. Remington and Dr. Ernest R. Tink-
ham, I fully realize the shortcomings of my own efforts.

With a complex group such as this, studies require methods
of oviposition, careful association of larvae with specific food
plants, exact location in the leaves or roots of the plants, sap or
plant fiber feeding, setu and method of frass disposal, nature
and placement of the frass, method of silk-lining of the tunnels,
chromosome counts, and many other details.

This means that one must live in close contact with a given
species if they hope to obtain an intimate picture of life activi-
ties. This is an exacting task.

Journal of Research on the Lepidoptera

14(2): 100^102, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

SUPPLEMENTARY RECORDS OF

BUTTERFLIES IN THE

SACRAMENTO VALLEY AND SUISUN MARSH,

LOWLAND CENTRAL CALIFORNIA

ARTHUR M. SHAPIRO

Department of Zoology, University of California, Davis, Ca. 95616

SACRAMENTO VALLEY

The following records are additions to Shapiro, 1975a.
New species are starred.

Chlosyne laciniu crocale Edwards. — This apparent introduc-
tion, which was fairly common in 1973 at one locality (Camp
Pollock, Sacramento County), was not seen in 1974. The habitat
was flooded during much of the winter of 1973-74; it lies in the
American River By-Pass.

^Nijmphalis milberti Latreille. — A single worn female taken
iv.6.74 in North Sacramento, elevation circa 15 feet, in a colony
of the introduced European nettle Uii:ica iirens L. Presumably
immigrant from higher elevations (Shapiro, 1975b).

Polygonia satyrus Edwards. — A second Davis record, female,
vi. 14.74. Since there are no nettles anywhere near Davis, the
occurrence of this species there must be based on rare strays
unless it breeds on the introduced Urticaceous ground cover
Soleiroleia as does Vanessa atalanta L., which however is
common.

Limenitis bredoicii californica Butler. — There is still no
evidence of breeding on the Valley floor, but an unusual number
of observations of adults were made in 1974. At Davis seen v.25,
ix.l, X.27.

Callophrys dumetonim Boisduval. — The first Valley record,
fresh male. Beach Lake, Sacramento County, iii.21.74. Presum-
ably breeding, but host not determined.

Pieris sisymbrii Boisduval. — Second Valley record: female,
Southport, Yolo Co., iv. 14.74. Presumably a stray from the foot-
hills.

100

101

ARTHUR M. SHAPIRO

/. Res. Lepid.

"^Euchloe hijantis Edwards. — A single male taken among
the very common E. ausonides. Beach Lake, Sacramento Co.,
hi. 21. 74, It is possible that this species occurs more widely in
the Valley, being mistaken for the commoner species, but it
could also be a stray; it is usually considered a foothill and
montane species. It flies with atisonides at 1000 feet in both the
Vaca Hills and Sierra foothills.

Colias (Zerene) eunjdice Boisduval. — California Dog-Faces
were unusually common in the foothills in 1974. Two males were
taken at Beach Lake, Sacramento Co., hi. 13 and hi. 21. The usual
host, Amorpha, has not been noted there.

Lerodea eufala Edwards. — One of the rare spring emergents
was taken at the American River, Sacramento Co., iv.28.74. The
main summer emergence was very late, first observed vhi.l9,
not seen at Davis until viii.Sl.

The addition of N. milberti, C. dumetonim, and E. hy antis to
the Valley list raises the total fauna to 65 species.

SUISUN MARSH

The following records are additions to Shapiro, 1975c.

Euphydryas chalcedona Doubleday. — Four seen on vi.7.74,
including a female. Despite this, there is no reason to believe
this species breeds in the Suisun Marsh,

Phyciodes campestris Behr. — Much commoner in 1974 than
in 1973, but seen only from ix.5-x.l9. Clearly associated with
Aster chilensis ssp. lentus, as earlier reported, and successfully
reared in the laboratory on A. chilensis ssp. chilensis Nees. The
distinctive Suisun facies was consistently maintained in both
wild and reared individuals.

Nymphalis calif ornica Boisduval. — Frequent in 1974. Seen
on ii.23, iii.9, and iv.2. On vi.7 about 1000 seemingly fresh indi-
viduals flew across the marsh headed ENE between 1200-1500
hours. Their flight was very steady and strongly directional.
Migrations oriented toward the N were observed the same week
in the North Coast Ranges and at Donner Pass.

Limenitis bredowii californica Butler. — A second stray, fe-
male, ix.28.74.

^Glaucopsyche lygdamus behrii Edwards. — Apparently resi-
dent on landfill near the Marsh, associated with a fencerow
population of Lathyrus jepsoni jepsoni, the endemic Suisun sub-
species of this plant. Collected on iii.9 and iii.20.74.

14(2):100-102, 1975

SUPPLEMENTARY RECORDS

102

Brephidium exilis Boisduval. — In 1974 was still emerging on
xii.24.

Batttis philenor hirsuta Skinner. — Two additional strays,

viii. 6 and ix.5.74.

"^Enjnnis propertim Scudder and Burgess. — One male and
one female both in good condition, taken on celery blossoms in
the Marsh on iv.11.74. Presumably strays from oak woodland in
the nearby hills.

"^Poanes melane Edwards. — Fresh male, in a shrub thicket,

ix. 5.74. The origin of this individual is unknown.

Ochlodes tjuma Edwards. — John H. Lane took a fresh male
at the Suisun City marina x,8.74, the latest this species has been
recorded in the Marsh.

The additions of G. lygdamus, E. propertim, and P. melane
raise the Suisun species total to 43.

LITERATURE CITED

SHAPIRO, A. M. 1975a. The butterfly fauna of tlie Sacramento Valley,
California. /. Res. Lepid., 13:73-82, 115-122, 137-148.

1975b. Altitudinal migration of central California butterflies. Ibid.

13:157-161.

1975c. Butterflies of the Suisun Marsh, California. J. Res. Lepid.

13(3): 191-206.

Journal of Research on the Lepidoptera

14(2): 103-124, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

AN ECOLOGICAL STUDY OF

THE BUTTERFLIES OF
THE SIERRA DE TUXTLA

IN VERACRUZ, MEXICO'

GARY N. ROSS^

Department of Entomology, Louisiana State University
Baton Rouge, Louisiana 70803

^From a dissertation submitted to the Graduate Faculty of the Louisiana
State University in partial fulfillment of the requirements for the degree of
Doctor of Philosophy in the Department of Entomology.

^Present Address: Department of Biology, Southern University, Baton Rouge,
Louisiana 70813.

103

1

14(2):103^124, 1975 BUTTERFLIES OF TUXTLA

105

FRONTISPIECE

Author in Montane Rain Forest on Volcan San Martin Tuxtla. August 1962,
3,500 feet. Photograph by R. F. Andrle.

3

106

GARY N. ROSS

]. Res. Lepid.

ACKNOWLEDGMENTS

Many persons have contributed to and assisted me in the
preparation of this work and I wish to express my sincerest
appreciation to them all. However, I feel that certain individuals
deserve a special acknowledgment and so I wish to enumerate
them. They are: Mr. and Mrs. J. M. Lind and family and Dr.
and Mrs. R. F. Andrle and family for living accommodations,
the wonderful hospitality that they extended to me during my
residence in the Sierra, and for numerous other services subse-
quent to my visits; the Department of Entomology at Louisiana
State University for financial assistance during my 1963 and
1965 expeditions to Mexico; F. M. Brown, H. K. Clench, R. M.
Fox, W. S. McAlpine, W. J. Reinthal, E. C. Welling, and K. H.
Wilson for butterfly determinations and assistance with numer-
ous problems in butterfly systematics; V. E. Rudd and her
associates at the United States National Museum for plant de-
terminations; T. Escalante (Mexico City) for making his exten-
sive collection of Mexican butterflies available to me; M. S. Blum
for his supervision and guidance during the course of this study;
H. B. Boudreaux, W. J. Harman, G. H. Lowery, Jr., L. D. New-
som, and J. S. Roberts for criticism and suggestions regarding
this manuscript; and, lastly, Mrs. G. G. Wynn for her assistance
in proofreading the manuscript and numerous other tasks.

4

14(2):103-124, 1975 BUTTERFLIES OF TUXTLA 107

TABLE OF CONTENTS

Page

TITLE PAGE 1

ACKNOWLEDGMENTS 4

TABLE OF CONTENTS 5

LIST OF TABLES 6

LIST OF FIGURES 7

LIST OF PLATES 8

ABSTRACT 9

CHAPTERS

1. INTRODUCTION 11

IT TOPOGRAPHY AND GEOLOGY 15

III. CLIMATE 19

IV. VEGETATION 21

V. ACCOUNTS OF BUTTERFLY SPECIES

IN THE SIERRA DE TUXTLA 51

Plan of the Species Accounts 51

Accounts 55

Family Papilionidae 55

Family Pieridae 62

Family Ithomiidae 70

Family Danaidae 74

Family Satyridae 75

Family Nymphalidae 81

Family Lycaenidae 105

Family Riodinidae ..120

VI. CORRELATION AND SYNTHESIS .129

Biotic Relationships 129

Faunal-Floral Relationships 141

Climatic Relationships 145

Faunal Relationships 147

REFERENCES CITED 156

APPENDICES ........159

APPENDIX A .159

APPENDIX B ..171

5

108

GARY N. ROSS

]. Res. Lepid.

LIST OF TABLES

Page

1. Plant Formations in the Sierra de Tuxtla 23

II. Plant FoiTnations and Associated Butterflies in the

Sierra de Tuxtla 131

III. Life Zones and Corresponding Plant Fonnations in

the Sierra de Tuxtla 140

IV. Genera of Butterflies Collected in the Sierra de

Tuxtla within Forests with Closed Canopies 142

V. Species of Butterflies Collected in the Sierra de

Tuxtla Known to Occur within the Borders of the

United States 148

VI. Comparison of Butterfly Genera Common to the Sierra
de Tuxtla, the United States, and the Neotropical
Regions 150

VII. New Butterfly Records for the Sierra de Tuxtla

(excluding Endemics) 152

VIII. Butterflies Endemic to the Sierra de Tuxtla 155

6

14(2): 103-1 24, 1975

BUTTERFLIES OF TUXTLA

109

LIST OF FIGURES

Page

1. Location map of the Sierra de Tuxtia 12

2. Vegetation of the Sierra de Tuxtia 24

7

no GARY N. ROSS '

LIST OF PLATES

Page

Frontispiece

1. Lago Catemaco 14

2. Volcan San Martin Tuxtla 16

3. Volcan Santa Marta 18

4. Volcan San Martin Pajapan 20

5. Lower Montane Rain Forest 26

6. Montane Rain Forest or Cloud Forest 28

7. Gum-oak forest 30

8. Montane Thicket 32

9. Elfin Woodland 34

10. Semi-Evergreen Seasonal Forest 36

11. Bursera-Sahal-Orhignya Forest 38

12. Savanna 40

13. Deciduous Woodland 42

14. Pine-oak Forest 44

15. Littoral Woodland 46

16. Swamp Forest 48

17. Mangrove Woodland 50

18. Recently Abandoned Milpa 52

19. Pasture 54

20. Hedgerow 56

8

14(2):103-124, 1975

BUTTERFLIES OF TUXTLA

111

ABSTRACT

The Sierra de Tuxtla is a small and isolated volcanic mountain range
along the Gulf coast of southern Veracruz, Mexico. Because of the geo-
graphic isolation, the range affords excellent conditions for distributional
and ecological investigations. The present study is the first comprehensive
report of the butterfly fauna of the range or for any geographic unit within
the Neotropics.

Fifteen months (representing all seasons of the year) were spent in
the field during 1962, 1963, and 1965. The various relatively widespread
plant communities were classified into 16 distinct types or fonnations. All
of these formations were sampled for butterflies and a total of 3,893 speci-
mens representing 359 species, 133 genera, and eight families were col-
lected. Of these species, 40 are recorded from the Sierra for the first time;
these include ten range extensions within the state of Veracruz, 18 new
state records, nine new national records, three new species, and one new
subspecies. All 359 species are listed in the species accounts along with the
field data — complete ( number of specimens, locales and altitudes, and col-
lection date) for those species that represent new records for the Sierra
but condensed (number of specimens and only ranges in altitude and col-
lection dates) for those species recorded previously from the Sierra.

Various relationships between the butterfly fauna and the environment
are discussed. First, an analysis of the plant formations with their indicator
and characteristic butterfly species indicates that life zone boundaries with-
in the Sierra are vague but still definable. The Sierra can be divided into
two major zones— an Upper Tropical Zone and a Lower Tropical Zone.
Furthermore, the data indicate that the Lower Tropical Zone can be
subdivided into a humid and an arid component.

Second, the majority of the butterfly species were found in the Lower
Tropical Zone in the open and relatively open plant formations whereas
very few species (principally members of the Ithomiidae and Satyridae)
were found in the dark interiors of the forests. Because of the Sierra’s
relatively low altitude and relatively uniform rainfall, it is suggested
that the principal governing factor detennining butterfly areal and alti-
tudinal distributions is the plant formation.

Third, although butterfly zoogeography is not sufficiently advanced to
enable one to determine the origins of most genera and species groups, the
majority of the genera ( 97 % ) and species ( 97 % ) found within the
Sierra’s boundaries appear to have their affinities with forms further south;
consequently, the butterfly fauna is essentially Neotropical.

Fourth, although the climate in the Sierra is relatively mild and uni-
form, enough diversity exists to produce significant variations in the butter-
fly populations. In general, populations of most species reached maximum
densities in late summer and early fall and their minimum densities in
winter and spring. In addition, daily population densities were greatest
between the hours of 10:00 A.M. and noon.

Fifth, butterfly endemism proved to be comparable to endemism in
other groups; three species, one subspecies, and one form (probably a good
subspecies) are endemic to the Sierra de Tuxtla.

9

112

GARY N. ROSS

J. Res. Lepid.

10

14(2);103-124, 1975

BUTTERFLIES OF TUXTLA

113

L INTRODUCTION

The Sierra de Tuxtla or Tuxtla Mountains (Tuxtla being the
Spanish corruption of the Axtec “Toxtli” meaning rabbit) is a
rather restricted highland of volcanic origin situated between
18°10^ and I8°45' N latitude and94°42' and 95°27' W longitude
on the Gulf Coastal Plain of the state of Veracruz in the Repub-
lic of Mexico (Fig. 1). The range trends northwest-southeast
with areal dimensions of approximately 55 by 30 miles and is
isolated from any other highland (the nearest being the Sierra
Juarez in the state of Oaxaca approximately 90 miles away) by
the Veracruz lowlands, principally the drainage basins of the
Papaloapan and Coatzacoalcos rivers. The Sierra is composed of
numerous ridges and volcanic cones and peaks of which four
attain elevations in excess of 3,000 feet, the maximum elevation
being 5,450 feet. These volcanic extrusions encircle a central
basin containing the picturesque Lago Catemaco, the third
largest lake in Mexico (Plate I).

When man first entered and began to settle the Sierra is still
unknown. Sears (1952) states that artifacts dating from approxi-
mately 1500 to 500 R.C. and probably Olmec in origin were
found in archeological sites in and around the range. The Span-
ish reached the Sierra a few years subsequent to their arrival in
Mexico — ca. 1522 (Melgarejo Vivanco, 1960). Today the area of
approximately 2,700 square miles is moderately populated with
both Mexicans and Indians, (Popolucas and Aztecs), the total
population in 1960 being approximately 145,000 (Andrle, 1964).
The people are engaged mainly in subsistence agriculture, which
includes the cultivation of corn, coffee, tobacco, and citrus fruits.
Because of the rather long history of settlement and cultivation,
relatively few undisturbed areas still ^xist. These are found prin-
cipally on the windward (Gulf facing) slopes of the major vol-
canoes and on the leeward slopes above elevations of 2,500 to
3,000 feet.

11

114

GARY N. ROSS

/. Res. Lepid.

12

FIGURE 1

Location map of the Sierra de Tuxtla. Map modified from that of Andrie
( 1964 ) . Basic map used with the permission of the author.

14(2):103-124, 1975

BUTTERFLIES OF TUXTLA

115

Scientific studies in the Sierra have been very limited and
brief; biological investigations have been confined almost ex-
clusively to the vertebrate fauna of the Lago Catemaco basin and
the slopes of Volcan San Martin Tuxtla. Avian and mammalian
studies include those of Sclater (1897), Wetmore (1943), Gold-
man (1951), Davis (1952), Amadon and Eckelberry (1955),
Edwards and Tashian (1959), and Andrle (1964). Herpetologi-
cal studies include those of Firschen (1950), Firschen and
Smith (1956), and Pyburn (1963, 1964, 1966). Previous studies
on arthropods are limited to a single paper on opilionids
(Goodnight and Goodnight, 1959).

My interest in the Sierra de Tuxtla began in 1961 because of
R. F. Andrle, a fellow graduate student who previously had
visited the range and who at the time was preparing for a 12
month return visit in early 1962 to study in detail the mammalian
and avian faunas. Andrlels enthusiasm about the Sierra coupled
with the fact that comprehensive studies on the bionomics of
Neotropical butterflies are practically nonexistent, convinced me
that the Sierra de Tuxtla would be an ideal study area for me.
So in June 1962 and with Andrle as a field partner, 1 began my
investigation of the Sierra’s butterfly fauna. During this first
study period (June through mid-December) my main base of
operations was located at Playa Azul on Lago Catemaco. From
that locale I directed my studies to the Volcan San Martin
Tuxtla massif and the Lago Catemaco-Bahia Sontecomapan
areas. Two incidental papers (Ross, 1963, 1964b) resulted from
this endeavor. The following year, June through August 1963,
I made a return visit to the Sierra and established a base at
Ocotal Chico. During this period my studies were directed to
the Volcan Santa Marta massif. Three incidental papers (Ross,
1964c, 1964d; Hepburn and Ross, 1964) resulted from this study.
In 1965 (February through July) I revisited the Ocotal Chico
site and expanded my investigation of the region to include the
Volcan San Martin Pajapan massif. One incidental paper resulted
from this 1965 trip (Ross, 1966). Transportation during all of
the study periods was provided by four-wheel drive vehicles,
trucks, mules, power boats, canoes, and my own two feet.

This dissertation is based on an assemblage of 3893 butterfly
specimens collected during a total of 15 months of personal field

13

116

GARY N. ROSS

/. Res. Lepid.

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14(2):103~124, 1975

BUTTERFLIES OF TUXTLA

117

work in the Sierra. In sections II (TOPOGRAPHY AND GE-
OLOGY) and III (CLIMATE) I have relied heavily on infor-
mation presented by Andrle ( 1964 ) and the reader is referred
to that work for additional and more detailed discussions of
those topics.

IT TOPOGRAPHY AND GEOLOGY

The Sierra de Tuxtla is an isolated mountain mass dominated
by four relatively large volcanoes. To the north these slope
rather steeply down to the Gulf of Mexico. To the South, West,
and East they slope more gently down to the Gulf Coastal Plain.
All slopes are deeply dissected and a radial erosion pattern is
very evident. The Sierra can be divided by the basin of Lago
Catemaco into a northwest and a southeast massif. The north-
west massif is dominated by Volcan San Martin Tuxtla (5,450
feet; Plate 2) and to a lesser extent by Cerro Tuxtla (2,725 feet),
Cerro Rlanco (2,375 feet), and the elongate Cerro Cintepec
(2,950 feet). Numerous small cones, hills, and crater lakes are
common south, east, and west of Volcan San Martin Tuxtla. To
the north, long, steep-sided ridges radiate down to the Gulf of
Mexico. The southeast massif is dominated by Volcan Santa
Marta (5,250 feet; Plate 3), Volcan San Martin Pajapan (3,750
feet; Plate 4 ) , and Cerro Campanario ( 3,900 feet ) . The southeast
section exhibits more uniformity than does the northwest sec-
tion in the sense that there are very few secondary cones and
crater lakes.

The numerous ravines on the slopes of the volcanoes usually
contain swift-flowing, clear streams, the flow from many of which
is either significantly reduced or stopped during the spring dry
season. The streams usually are fed by cool, clear springs that
issue from rock crevices on the upper slopes of the volcanoes.
At lower elevations the streams join larger streams and rivers
which in turn eventually flow into the Gulf of Mexico.

Murray ( 1961 ) states that the Tuxtla Uplift probably existed
as early as Mesozoic times. He considers the uplift to be high
areas of basement rock in the Mesozoic-Cenozoic geosyncline
that probably was part of the arc-shaped “Tamoulipas-Yucatan
archipelago.” Later, this syncline sank and was subjected to Gre-
taceous and Tertiary deposits of blue clays and shales, tuffs,
sandstones, limestones, and conglomerates, which upon later
emergence of the Sierra, were partially eroded away.

Schieferdecker and Tschopp (1922) suggest that the Sierra

15

118

GARY N. ROSS

/. Res. Lepid.

16

PLATE 2

San Martin Tiixtla. August 1962, 2,000 feet. Photograph by R.

14(2):103-124, 1975

BUTTERFLIES OF TUXTLA

119

rests on a diaritic laccolith of early Miocene or Oligocene age
that lifted and in places folded the Tertiary beds and from which
the volcanic extrusions have emerged. These extrusions comprise
most of the present-day Sierra and consist of older Pliocene
deposits of an acid andesitic character upon which were super-
imposed basalt flows and volcanic plugs of late Pliocene age
following the last marine inundation.

Andrle (1964) recognized seven principal eruption centers
or zones within the Sierra. These are: Cerro Tuxtla, Cerro
Blanco, Volcan San Martin Tuxtla, the Lago Catemaco Basin
(including Cerros Mono Blanco, Las Animas, and Cintepec),
Cerro Campanario, Volcan Santa Marta, and Volcan San Martin
Pajapan. Besides these principal cerros, there are numerous sub-
sidiary lava, ash, and cinder cones in the area, principally in the
vicinity of Volcan San Martin Tuxtla. The true nature of Lago
Catemaco is still debatable. Friedlaender (1923) considered it to
be a caldera but Andrle (1964) suggests that it is simply a
spring and stream-filled low section of the range whose southern
and western borders are effectively blocked by volcanic cones
and debris. Layers of ash, lapilli, and cinders are evident par-
ticularly in the Catemaco basin. Basalt bombs, pumice, and
asphalt cakes are fairly common, particularly along the coast.
Fine-grained olivine basalt rocks are the dominant rock type
throughout the Sierra. These are evident as blocks, both large
and small, and as extensive flows, which in places exhibit colum-
nar faulting.

Only Volcan San Martin Tuxtla has a historical record of
eruptions. This fact tends to support the suggestions of Fried-
laender ( 1923 ) that the northwest massif is of younger geologic
age than the southeast counterpart. Medel & Alvarado (1963)
briefly described an eruption on October 15, 1664, which Fried-
laender ( 1923 ) defined as an ash eruption with a possible re-
stricted lava flow to the north. Mocino ( 1870 ) reported a second
eruption that began in March 2, 1793 and that consisted of vio-
lent explosions, lava flows to the northeast and northwest, and
ash falls that continued intermittently through September.
Garcia (1835) observed fumarolic activity in the crater in 1829
but nothing more.

17

120

GARY N. ROSS

}. ties. Lepid.

PLATE 3

Volcan Santa Marta. Body of water in the foreground is an artifieially

created reservoir. June 1965, 500 feet.

14(2):103-124,1975 BUTTERFLIES OF TUXTLA

121

The four major volcanoes each show well developed oval
and steep" walled craters, which for the most part, are open to
the north indicating the direction of major lava flows. The
crater of Volcan San Martin Tuxtla has a maximum length of
approximately one mile and a maximum depth of approximately
600 feet. The crater of Volcan Santa Marta has a maximum
length of approximately one and a half miles and a maximum
depth of approximately 500 feet. Cerro Campanario and Volcan
San Martin Pa japan have craters smaller and shallower than
those of Volcans San Martin Tuxtla and Santa Marta.

III. CLIMATE

The Sierra de Tuxtla is characterized by rather uniform year-
round temperatures and seasonal rainfall. This rather mild cli-
mate is a result of the moderating effect of the Gulf of Mexico.
Andrle (1964) lists temperature and precipitation data for six
stations in the Sierra. Unfortunately, all of these stations are in
a relatively narrow zone on the southern slopes of the Sierra and
hence, the data can be used only to illustrate general trends.
April and May usually are the warmest months and January and
February the coolest. The average annual temperature is ap-
proximately 75.5° F (average elevation of 955 feet). The aver-
age mean for the coldest month is 68 °F. The lowest temperature
recorded at any station ( San Andres Tuxtla, 1,188 feet, 32 years
of data) was 4L2°F. Medel & Alvarado (1963) reported that on
February 9 and 10, 1899, the peak of Volcan San Martin Tuxtla
was covered with ice; thus freezing conditions are not unknown
on the peaks of the highest volcanoes. Low temperatures usually
occur between October and April after the passage of a mass
of cold air that moves across the Gulf from the north or north-
east. These fronts, which are called “nortes” by the local inhabi-
tants, vary in intensity and duration, some being weak and
lasting for only two or three days, others somewhat stronger and
lasting for as long as seven days.

Precipitation is variable with two pronounced seasons — a wet
season from June through January (maximum rainfall in July
and October) with usually a slight decrease in August, and a
dry season from February through May (minimum rainfall in
March and April). Most precipitation during the winter months
is associated with the passage of fronts. Most summer and fall
precipitation occurs during thundershowers in the night and
early morning, Andrle’s climatic data indicate that rainfall is

19

122

GARY N. ROSS

/ Res. Lepid

PLATE 4

Volcan San Martin Pajapan. May 1965, 2,600 feet.

20

BUTTERFLIES OF TUXTLA

123

14(2):103-I24. 1 :73

principally an orographic type and varies considerably between
each station and even from year to year at any single station.
The minimal early average for any station is 69 inches at Gua-
suntlan (elevation 595 feet) and the maximal yearly average is
163 inches at Coyame (elevation 1,122 feet). Rainfall above
elevations of 1,500 feet on the Gulf slopes is probably in excess
of 170 inches per year, and the peaks of the principal volcanoes
conceivably receive upwards of 200 inches per year because of
the orographic effect.

The cloudiest months are December, January, and July; the
least cloudy are March, April, and May.

Since the mean temperature of the coldest month is greater
than 64.4 °F and the mean precipitation of the driest month is
greater than 2.4 inches, the Sierra falls within the “Tropical Rain
Forest Climate” (Af) of Koppen (1936).

IV. VEGETATION

The Sierra de Tuxtla lies within (but near the northern
border of) the Neotropical Realm of Wallace (1876) and rep-
resents the most northern extension of the relatively uninter-
rupted belt of tropical rain forest that extends (in climatically
favorable areas ) from southern Mexico through Central America
and far into South America (Leopold, 1950; 1959). Hence the
floristic composition of the region is basically tropical with most
components being related to plant groups further south. This
tropical composition coupled with the fact that the Sierra repre-
sents a relatively small geographic area (approximately 2,700
square miles ) would lead one to the conclusion that the flora
of the region is rather homogeneous. Such a conclusion, however,
would be completely erroneous because the Sierra exhibits a
considerable diversification in vegetation. This diversity is a re-
sult of many factors of which some of the most important in-
elude: altitudinal zonation from sea level to a maximum ele-
vation of 5,450 feet resulting in temperature and rainfall gradi-
ents; differences in the composition and ages of soils due to
differential weathering and variances in age of parent material;
the long axis of the range in respect to the prevailing winds
resulting in relatively heav\^ precipitation on the Gulf facing
slopes and a slight rain shadow effect on the leeward slopes; and

124

GARY N. ROSS

J. Res. Lepid.

the agricultural practices of man resulting in the presence of
all stages of plant succession.

Plant Formations

In attempting to define the habitats of butterfly species in
the Sierra, a more subtle ecological division that either “life zone’'
or “biotic province” had to be chosen. Because of the Sierra’s
numerous and frequently widely distributed plant communities,
I decided to empoly a habitat classification based on plant for-
mations.

Andrle ( 1964 ) in his investigation of the Sierra differentiated
between ten distinct types of vegetation. However, because of
the broadness of many of his terms I have found it necessary to
modify this classification. This was accomplished by correlating
(where possible) the apparent climax types with those as out-
lined by Beard ( 1944; 1955 ) . When no correlations were ap-
parent, e.g., the various serai communities and the oak and pine
communities, I have errected new categories, being careful not
to employ any of the Beard terminology. The result of this
effort is that I recognize 16 distinct formations in the Sierra
(Table 1). The geographic locations of the major types are illus-
trated in Figure 2. In actuality the boundaries between each
type and the next are rather arbitrary for oftentimes relatively
wide transitional zones or ecotones exist between formations.

( to be continued )

22

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SPECIAL NOTICE: In order to return to a normal publication
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THE JOURNAL OF RESEARCH
ON THE LEFiJDORTERA

Volume 14 Number 2 May, 197^

IN THIS ISSUE

The genetics of subspecific phenotype differences
in Pieris occidentalis Reakirt and of variation
in P. o. nelsoni W. H. Edwards ( Pieridae )

Arthur M. Shapiro 61

Early stages of Phyciodes pallida, P. orseis,

and P. mylitta (Nymphalidae) James Scott 84

Recent captures of Anthocharis cethura catalina

Meadows Larry J. Orsak 85

Old Timers John Adams Comstock 90

Why do California Tortoiseshells migrate?

Arthur M. Shapiro 93

Early work on the Megathymidae

John Adams Comstock 98

Supplementary records of butterflies in the
Sacramento Valley and Suisun Marsh,
lowland central California

Arthur M. Shapiro 100

An ecological study of the butterflies of

the Sierra de Tuxtla in Veraeruz, Mexico

Gary N. Ross

103

t

THE JOURNAL
OF RESEA ROHJ
ON THE LEFIJDOFTERA

Volume 14 Number 3 September, 1975

I! THE LEPlJDOPTERA

published by

The Lepidoptera Research Foundation, Inc.
at

1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006

EDITOR: William Hovanitz

Associate Editors :

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Canada.

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Journal of Research on the Lepidoptera

14(3):125»-141, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

COURTSHIP AND MATING BEHAVIOR

OF THE FIERY SKIPPER, HYLEPHILA PHYLAEUS
(HESPERIIDAE)

IRENE SHAPIRO

Graduate Group in Ecology,

University of California,

Davis, California 95616^

ABSTRACT

The courtship and mating behavior of Hylephila phylaeus ( Hesperi-
idae) was studied by introducing dead and live tethered wild and labora-
tory reared females to wild males in the field. The hypothesis that wing
movement elicits the male’s initial investigative response in this species
was tested. Entire courtship sequences resulting in successful matings were
obtained with 18 live virgin females, indicating that males are less respon-
sive to previously mated females.

INTRODUCTION

The problem of locating and successfully mating with a con-
specific is of primary importance in the perpetuation of a
species. Not only must potential mates recognize one another as
opposed to closely related secies, but a synchronization of repro-
ductive cycles must occur so as to bring members of the opposite
sex together at the proper time of year. Only by studying the
courtship and mating systems of individual species can theories
pertaining to the significance of various aspects of such systems
be validated.

The present study deals wi h the courtship and mating system
of a Lepidopteran, the fiery skipper, Hylephila phylaeus (Hes-
periidae). This species is abundant in the Sacramento Valley of
California, where it is multiply brooded, having up to five gen-
erations per year. The larval host is Bermuda grass, Cynodon
dactylon (Gramineae). //. phylaeus can be classified as a perch-
ing species as defined by Scott (1974c), i.e. males perch on pro-
jecting objects such as leaves, pieces of paper, and tall blades of

Tresent address: 1005 S. Park Victoria Dr., Milpitas, California 95035.

125

126

IRENE SHAPIRO

J. Res. Lepid.

Map of Study Area (34,000 square meters)

Fig. 1. — Map of Study Area (34,000 square meters)

14(3):125~141, 1975

COURTSHIP AND MATING

127

grass on lawns and vacant lo.s in residential areas. Perching
males fly towards and “investigate” moving objects such as other
butterflies of the same or different species, other insects, and bits
of leaves blown by the wind. The hypothesis that movement
(specifically wing fluttering movement) elicits the male investi-
gative response in this species was tested in the field.

MATERIALS AND METHODS

The study was conducted on the University of California,
Davis campus throughout the periods of September to November,
1973 and June to December, 1974. The 34,000 square meter study
area consisted of a 12,300 square meter lawn area of Cynodon
dactylon, Poa pratensis, Agrostis alba, and Festuca rubra, which
was separated by a road (10 meters in diameter) from a 21,700
square meter flower and vegetable garden that served as a food
source for adult phylaeus (see Map Eig. 1). Male perching and
investigative behavior were observed on the lawn area, while
courtship and mat mg behavior experiments were conducted in
the vegetable gardens because of the greater density of H.
phylaeus there.

In order to determine the behavioral components involved
in a complete courtship sequence, dead and live wild male and
female H. phylaeus were presented to wild males in the field.
A modification of the technique devised by Tinbergen et al.
(1942) was used: butterflies were cooled for about ten minutes
at 12° C so as to facilitate handling, and a quick-drying epoxy
cement was applied to the dorsal surface of the thorax while a
fine piece of fishing line was held in position until the glue dried
(about three minutes). The fishing line was then attached to a
rod which could be dangled in front, above or to the side of
males at a distance of up to 1.8 meters from the observer. When
fishing line was properly mounted, no difference from normal
rapid flight movement was observed, except that individuals
could not fly farther than a 1.8 meter diameter circle around the
observer.

In a second group of experiments, laboratory reared females
and males were presented in the same manner to wild males in
the field in order to determine whether different results could
be obtained with females known to be virgin. Results were
recorded by notes and through use of a portable cassette tape
recorder.

Table 1. 1973 Experiments with Tether^^_ Wild Males and Females Presented to Wild Males in the Field

(n = # males investigating or touching tethered skipper ; N = total # males to which tethered skippers
were presented | * = tethered skipper not flying, data discarded).

128

IRENE SHAPIRO

/. Res. Lepid.

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(No significant difference at ,05 level between female, male, and eufala tethered skippers, using the
t-test of difference between two means).

14(3):125-141, 1975

COURTSHIP AND MATING

129

RESULTS

1973 Experiments with Wild Females
and Males Presented to Wild Males

All experiments with dead male and female phylaeus failed
to elicit courship behavior from wild males in the field. 13 dead
females and 15 dead males presented to 10 wild males apiece
elicited no responses, even when tethered skippers were dangled
from fishing line directly in front of males. 2 dead females
elicited an approach response, investigative behavior, from one
of ten males apiece. In both instances, males immediately de-
parted without touching or following the females. As a control,
all tethered animals were also presented to 10 wild females
apiece, and negative results were obtained in all cases.

Experiments with live phylaeus yielded more positive results
on the preliminary phases of courtship. Data from 8 female and
6 male H. phtjlaeus were obtained. Both males and females
elicited investigative behavior from wild males. When a tethered
skipper was moved past a male at a distance of 0.3 to 0.6 meters,
the male usually flew toward and often touched it with his
head and antennae. A few males followed the tethered skipper
for several feet, but most immediately departed, returning to
an area near their original position. All tethered animals were
presented to 10 wild females apiece, and no responses were
elicited.

Five Eufala skippers, Lerodea eiifala, were used in another
group of experiments. In this species, the sexes are phenotypically
alike, both being an overall brown-gray color with a peppering
of small white spots on the upperside of the forewing. Investi-
gative behavior and touching behavior were again elicited when
live tethered eufala were presented to wild male phijlaeus, and
no responses were elicited when presented to wild females.

Results suggest that the fluttering of the wings during flight
is a stimulus necessary to induce investigative behavior in males,
since in all but two instances where a tethered skipper did not
flutter its wings but merely hung at the end of the line, males
did not respond. Differences in responses to female and male
phylaeus and eufala were not significant at the 95% level using
the t-test of difference between two means. This further sub-
stantiates the hypothesis that the fluttering of the wings may be
more important than color or scent in initiating the first stages
of courtship in H. phylaeus (Table 1).

130

IRENE SHAPIRO

]. Res. Lepid.

1974 Experiments with Virgin Females and
Males Presented to Wild Males

Results obtained in the 1974 experiments support the hypoth-
esis that the fluttering of the wings is an essential stimulus to
elicit investigative behavior in males. 16 females did not fly
when presented to 10 males apiece, and subsequently failed to
elicit a response in all 10 males. One female that failed to
flutter its wings did elicit investigative behavior in three of 10
males, while two more non-flying females stimulated approaches
in one of 10 males apiece. It was also found that feeding males
usually failed to respond to tethered skippers. Out of a total
of 71 trials with 16 female phylaeiis, only 3 feeding males were
stimulated to investigate; the other 68 males continued to feed.

Because no complete courtship sequences could be induced
in the 1973 experiments with wild females, it was hypothesized
that perhaps only virgin females could provide the correct
stimuli necessary to elicit a sequence resulting in a successful
mating. Therefore, in 1974, laboratory reared virgin females
were used in the experiments. As a control experiment, 10
laboratory reared virgin males were also presented to wild
males in the field; results were identical to those of 1973, where
only the first phase of courtship, investigative behavior, was
elicited. Experiments with virgin females, however, produced
18 successful matings. A description of the various phases in the
courtship sequence of H. phylaeiis is as follows:

Phase 1 — Investigative Behavior: the male flies toward and may touch
a passing female.

Phase 2 — Settling Behavior: the male lands behind the female as she
settles on a leaf, flower, or the ground.

Phase 3 — Head-Wing Behavior: the male, while situated at the rear
of the female, thrusts his head between the female’s hindwings, touch-
ing the upper surface of her wings and the rear of her abdomen with
his head and antennae; males often thrust themselves so far forward
that they are able to touch the upperside of the female’s forewings.

Phase 4 — Fluttering Behavior: the male rapidly flutters his wings
while moving laterally to the female and/or in front of her.

Phase 5 — Curving Behavior: the male, in a position posterior to the
female, curves his abdomen in a U-shaped arc, so that his genitalia are
in close proximity to the female’s; the male genitalia are often extruded.

Phase 6 — Coupling Behavior: the male clasps the female with his
harpes and apparently inserts his aedeagus into the female; after sev-
eral seconds to several minutes, the male moves his body around so
that he is facing in the opposite direction from the female ( their bodies
form a 180° angle).

14(3);125-141, 1975

COURTSHIP AND MATING

131

Phase 7 — Uncoupling: copulation was terminated from 40 to 72 min-
utes after coupling except in one instance, when the mating pair sep-
arated after 27 minutes; it is not known whether a spermatophore was
transferred or not, since the female flew away immediately after un-
coupling. After separating, the male and female of all pairs observed
either began feeding or flew away.

In the 18 successful courtships observed, overt female sexual
behavior was minimal. The female usually remained stationary
where she had settled, occasionally moving her antennae or
walking forward several centimeters. In a few instances, females
were observed to extrude genitalia, although this behavior also
occurred during unsuccessful courtships. Shapiro (1970) pro-
posed that such behavior in Pierid butterflies is part of a rejec-
tion display by unreceptive females. An identical display in
Heliconiid butterflies, however, appears to have a pre-copulatory
function, since in this group, the extrusion of the genitalia is
related to pheromone release (Crane, 1957).

The duration of successful courtships prior to coupling
ranged from ten seconds to three minutes and 30 seconds. Most
were of short duration, and only five of 18 were longer than
40 seconds. Similar results were obtained by Brower et al. (1965),
who found that the mean duration for 266 courtships in Danaus
gilippus berenice was 40.4 seconds.

There was considerable variation in the sequence of phases
as well as the repetition of phases among individual courtships.
Nine courtships were single-phased sequences, while the other
nine had sequences with repetitious phases. All but three
courtships began with investigative behavior followed by the
settling phase (Table 2).

Only one successful wild courtship as opppsed to 208 un-
successful wild courtships was ever observed. The duration of
the courtship was 15 seconds, and the duration of time in copula
was 40 minutes. The courtship was single-phased and was
sequentially analogous to successful courtships in the experi-
ments with laboratory reared virgin females.

Seven cases of carrying pair behavior were observed. In all
cases, the female flew while carrying the quiescent male. Such
behavior occurred only when the pair was disturbed in some
way, such as by other males attempting to court the female or
when touched by the investigator.

132

IRENE SHAPIRO

J. Res. Lepid.

Table 2» 197^ Experiments vrith Virgin Females Presented to Vi/lld Males in the

Field - Sequence of Phases in Successful Courtships.

Female #

Sequence of Phases

1

1-2-4- 5-6-M

2

1_2-3-4-5-6-M

3

1-2-5-6-M

4

1-2-5-4-6-M

Courtship Sequences

5

1_2-5-6-4-M

with Single Phases

6

1-2-5-^6-M

7

3-ZP-5-6-M

8

1_2-3-4-5-6-M

9

1-2->5-6-M

10

1- 2-5-5^3l*pl5^-1P-6-M '

11

1-2-5-4-3-5-6-M

12

1- 2-4- > 5-6- P- 5-6-4-FI- 2- 5-6-M

13

2— 5— 3’"4— 6— 4— P— 5'“^^6— S— P— 3~4— 5—4— 6— M

Courtship Sequences with

14

1-2-5-6-P-5-6-M

Repetitious Phases

15

1_2-^5-6-P-5-6-P-5-6-4-H

16

1_ 2- 5- 6- ^ p- 6- 3-4-m

17

1- 2- 5- S-6- p- 5- 3-Z4~ s- 5- 3-^ S-6- 5- 3-4-6-M

18

2-4-3-^5-6-FD-2-6-P-5-6-P-4-6-M

1 = Investigative Behavior

2 = Settling Behavior

3 = Head- Wing Behavior
^ = Fluttering Behavior

5 = Curving Behavior

6 = Coupling Behavior
M = Successful Mating
P = Female Pulled Away
S = Female Shuddered

FI = Courtship Disrupted by Another Male
FD = Female Deserted

14(3):125~141, 1975

COURTSHIP AND MATING

133

Unsuccessful Courtships with Virgin Females

The courtship sequence can be terminated by either the
male or the female after any of the first six phases in any of
several ways. The most common cause of unsuccessful courtship
was disruption by other males, which resulted in the courting
male leaving the female and flying after the other male. Such
investigative behavior often resulted in “chases” between two
or more males to areas far removed from the origin of the
encounter. 41 out of 88 unsuccessful courtships ended in this
type of disruption behavior.

36 out of 88 courtships ended in male desertion behavior,
where the male departed to a nearby flower or leaf. Such be-
havior may be elicited by a female rejection display called
shuddering behavior, in which the female rapidly flutters the
forewings and hindwings, which are opened at about a 45° angle
from one another. Shuddering could be elicited either by head-
wing behavior, fluttering behavior, or curving behavior and was
often accompanied by movement a short distance away from
the male. 28 of 36 courtships terminating by male desertion
occurred after the female had shuddered once or several times.

These observations are in sharp contrast to the successful
matings observed, where there were 16 out of 18 courtships in
which the female did not shudder. It may be significant to note
that the two females that did not shudder had repetitious court-
ship sequences, and that neither shuddered during the final
sequence culminating in a successful mating.

Another female display which may be a rejection is the
raising of the abdomen up and away from the male’s genitalia
during coupling. In all cases such pulling away behavior resulted
in an uncoupling of the pair.

Courtships were also terminated by female desertion, where
the female flew away from the male to another area. Only 11 out
of 88 courtships ended in this manned (Table 3).

Unsuccessful courtships with virgin females were longer
than unsuccessful courtships with mated females. Only the first
phase of courship, investigative behavior, was elicited by mated
females, while virgin females elicited at least one of the later
stages of courtship (settling, head-wing, fluttering, curving).

Table 3» 197^ Experiments with ’Virgin Females Presented to Wild Males in the

Field - Causes of Termination of Unsuccessful Courtships (in # of Observations ) ,

IRENE SHAPIRO

J. Res. Lepid.

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14(3):125-141, 1975

COURTSHIP AND MATING

135

DISCUSSION

Courtship and mating systems of insects have been shown
to consist of progressive exchanges of stimuli between males
and females; one partner’s response to a stimulus elicited by
the other releases the next response, and so on (Frings and
Frings, 1958; Perdeck, 195-7; Crane, 1957; Baerends, 1959).
In Hylephila phylaeus, the initial investigative response of males
appears to be elicited by movement, specifically the fluttering
of the wings. Color of the wing does not appear to play a sig-
nificant role, since males were equally attracted to both male
and female phylaeus, despite the pronounced sexual dimorphism
in this species, as well as to the darker eufala skippers. Size
may possibly be a factor, although male phylaeus were occa-
sionally seen to investigate larger butterfly species, such as
Vanessa cardui, V. carye, Colias eurytheme, and Pieris rapae.

Magnus ( 1958 ) obtained similar results with Argynnis paphia
( Nymphalidae ) . He proposed that the fluttering of the wings
itself was not the decisive factor, but rather the rapid alteration
of color versus absence of color when the wings are fluttered.
He found that the intensity of the male response rose in direct
proportion to the speed of color alteration. It is possible that a
similar response may occur in H. phylaeus, since experiments
thus far indicate that the fluttering motion of the wings is
essential in eliciting a male response.

The investigative reaction appears to be related to the mate-
locating system of various butterfly species: in perching species,
the female must fly past the male to attract his attention; in
patrolling species, it is the male that must fly past and locate
non-flying females (Scott, 1974c). It would thus appear that
color might be an important stimulus to the patrolling male,
while the perching male would be more attracted to movement
or quick flashes of color from fluttering wings. Thus far, the
available data substantiates this hypothesis. In the perchers
HypoUmnas misippus (Stride, 1957, 1958a and b), Neominois
ridingsii, Hesperia pahaska, Ochlodes snowi, Lycaena arota,
and Amblyscirtes simius (Scott, 1973a, b; 1974a), the first phase
of courtship is elicited by movement of the female past the
stationary male. In the patrollers Argynnis paphia (Magnus,
1958), Erebia epipsodea (Brussard and Ehrlich, 1970a,b),
Hypaurotis crysalus (Scott, 1974b), and Pieris rapae (Obara,
1970), color of the female appears to be of primary importance
in attracting males. In the patroller Danaus gelippus berenice,

136

IRENE SHAPIRO

J. Res. Lepid.

Fig. 2. — Settling Behavior of Hylephila phylaeus During Courtship Se-
quence (Male Lands Behind Female).

14(3) -.125-141, 1975

COURTSHIP AND MATING

137

however^ Brower et al. (1965) stated that males are initially
attracted to the female by seeing her in flight or while she is
fluttering on herbage. Such behavior may be an artifact due to
the procedure used by the investigators — females were released
by hand into the air and all females that did not fly past the
male were discarded from the results.

In H. phtjlaeus, the investigative reaction may also be re-
lated to carrying pair behavior. In order to carry the male during
copulation, the female’s wings must dry before mating occurs.
Thus, the evolution of the wing fluttering movement as a stimu-
lus to the male in courtship may also have evolved in response
to the hardening of the female’s wings signaling her readiness
to mate.

Results of the field experiments on H. phylaeus suggest that
in the next phases of the courtship sequence, where the male
follows the female and settles near her, virgin females are con-
siderably more attractive to males than are already mated
females or other males. Firstly, there was no observable differ-
ence in length of the following reaction between tethered male
or mated female phylaeus; secondly, all of the successful mat-
ings observed were with virgin females; thirdly, the unsuccess-
ful courtships with virgin females were longer and more involved
than those with mated females. Color would thus not appear to
play a major role in eliciting the following response of males.
It may instead be that a pheromone emitted by virgin females
attracts males after the investigative approach. Scott (I973a,b;
1974a) found evidence that virgin females of Neominois
ridingsii, Hesperia pahaska, Ochlodes snowi, and Lycaena arota
may possess a pheromone that attracts males at close range
(within about a meter). H. pahaska and O. snotoi are fairly
closely related to H. phylaeus, being in the same tribe.

In later stages of courtship in H. phylaeus, odor again may
be the primary stimulus eliciting the male’s responses, especially
in head-wing behavior, in which the male’s head is thrust
between the female’s hindwings, touching her genital region.
Tactile stimuli may also be functioning. In these later stages,
species-specific pheromones would be adaptive to insure against
hybridization with other species.

The stigma or pouch on the forewing of males of various
butterfly species has been proposed to carry a pheromone which
stimulates the female during courtship (Scott, 1973b; MacNeill,
1964). In H. phylaeus, the fluttering behavior of the male may

138

IRENE SHAPIRO

/. Res. Lepid.

Fig. 3. — -Curving Behavior of Hylephila phylaeus During Courtship Se-
quence (Male Genitalia are Extruded).

14(3):125-141, 1975

COURTSHIP AND MATING

139

serve to disperse the pheromone from the stigma. Males often
moved laterally to and in front of the female while fluttering
their wings, thus possibly surrounding the female with scent.
Similar results have been observed by Scott ( 1973a, b; 1974a)
in Amblyscirtes simius, Hesperia pahaska, Ochlodes snowi, and
Lycaena arota.

The rejection display by females may function in minimizing
energy expenditure. When a rejection display abruptly halts a
male’s advances, wasteful energy expenditure is reduced— the
female’s energy can then be directed towards oviposition. A
rejection display is thus advantageous only if it stimulates the
male to abruptly terminate a futile courtship, as was the case
in H. phylaeus. As a consequence, wasteful energy expenditure
on the part of the male is reduced. Also, in attempting to court
a previously mated female, a male’s chances of successfully con-
tributing to the next generation are decreased (assuming that
sperm precedence occurs, Tayor (1967)), simply because such
a female is older and has a shorter survival time during which
to oviposit than does a newly emerged virgin. However, if virgin
females are in short supply, any male’s fitness is likely to increase
if he attempts to mate any female he encounters. This is likely
to be the case in species like H, phylaeus in which the average
female mates only once but males may mate multiply, so the
frequency of unsuccessful courtships involving non-virgin fe-
males observed in the wild is not surprising.

ACKNOWLEDGEMENTS

1 would like to thank Arthur M. Shapiro for critically reading
the manuscript, James A. Scott for his invaluable explanations
of data analysis, and Frederick J. Shapiro for his excellent
photography.

LITERATURE CITED

BAERENDS, G. P. 1959. Ethological studies of insect behavior. Ann. Rev.
Entomol. 4:207-234.

BROWER, L. P., JANE VAN ZANDT BROWER, and FLORENGE PIT-
KIN CRANSTON. 1965. Courtship behavior of the queen butterfly,
Danaus gilippus berenice (Cramer). Zoologica 50(l):l-40.
BRUSSARD, P. F. and P. R. EHRLICH. 1970a. The population structure
of Erebia epipsodea ( Lepidoptera : Satyrinae ) . Ecology 51 ( 1 ) : 1 19-129.
. 1970b. Adult behavior and population structure in Erebia epipso-
dea (Lepidoptera; Satyrinae). Ecology 51(5) :880-885.

CRANE, J. 1957. Imaginal behavior in butterflies of the family Heliconiidae:

changing social patterns and irrelevant actions. Zoologica 42:135-146.
FRINGS, H. and M. FRINGS. 1958. Ann. Rev. Entomol 3:87-106 (in
Baerends, G. P. 1959. Ethological studies of insect behavior. Ann. Rev.
Entomol. 4:207-234.

140

IRENE SHAPIRO

/. Res. Lepid.

■In Copula Pair of Hylephila phylaeus.

14(3):12S~141, 1975

COURTSHIP AND MATING

141

MacNEILL, C. D. 1964. The skippers of the genus Hesperia in western
North America, with special reference to California ( Lepidoptera:

Hesperiidae ) . Univ. Calif. Pub. Entom. 35:1-230.

MAGNUS, D. B. E. 1958. Experimental analysis of some “overoptimal”
sign-stimuli in the mating behaviour of the fritillary butterfly Argynnis
paphia L. (Lepidoptera: Nymphalidae). Proc. Tenth Internat. Cong.
Entomol. 2:405-418.

OBARA, Y. 1970. Studies on the mating behavior of the white cabbage
butterfly, Pieris rapae crucivora Boisduval (Lepidoptera, Pieridae),
III. Near ultraviolet reflection as the signal for intraspecific communi-
cation. Z. Vergl. Physiol. 69:99-116.

PERDECK, A. C. 1957. The isolating value of specific song in two sibling
species of grasshoppers {Chorthippus hrunneus Thumb, and C. bigut-
tulus L. ). Behaviour 12:1-75.

SCOTT, J. A. 1973a. Convergence of population biology and adult behavior
in two sympatric butterflies, Neominois ridingsii ( Papilionoidea: Nym-
phalidae) and Amblyscirtes simius ( Hesperioidea: Herperiidae ) . /.
Anim. Ecol. 42:663-672.

-. 1973b. Adult behavior and population biology of two skippers (Hes-
periidae) mating in contrasting topographic sites. /. Res. Lepidopt.
12(4):181-196.

—. 1974a. Population biology and adult behavior of Lycaena arota

( Lycaenidae). /. Lepidopt. Soc. 28(l):64-72.

— . 1974b. The interaction of behavior, population biology and environ-
ment in Hypaurotis cnjsalus (Lepidoptera). Amer. Midi. Nat. 91(2):
383-394.

-. 1974c. Mate-locating behavior of butterflies. Amer. Midi. Nat.

91(1):103-117.

SHAPIRO, A. M. 1970. The role of sexual behavior in density-related dis-
persal of Pierid butterflies. Amer. Nat. 104:367-372.

STRIDE, G. O. 1957. Investigations into the courtship behavior of the male
of Hypolimnas misippus L. (Lepidoptera, Nymphalidae), with special
reference to the role of visual stimuli. Brit. J. Anim. Behav. 5:153-167.

— - — -. 1958a. On the courtship behaviour of a tropical mimetic butterfly,
Hypolimnas misippus L. (Nymphalidae). Proc. Tenth Internat. Cong.
Entomol. 2:419-424.

“. 1958b. Further studies on the courtship behaviour of African mi-
metic butterflies. Brit. J. Anim. Behav. 6(3-4) : 224-230.

TAYLOR, O. R. 1967. Relationship of multiple mating to fertility in Atteva
punctella (Lepidoptera: Yponomeutidae ) . Ann. Entomol. Soc. Amer.
60(3):583-590.

TINBERGEN, N., B. J. D, MEEUSE, L. K. BOEREMA, W. W. VAROS-

SIFAU, 1942. Die Balz des Samtfalters, Eumenis (Satyrus) semele
(L.). Z. Tierpsychol. 5:182-226.

Journal of Research on the Lepidoptera

14(3): 142-147, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

VARIABILITY OF COURTSHIP OF
THE BUCKEYE BUTTERFLY, PRECIS COENIA
(NYMPHALIDAE)

JAMES A. SCOTT

Department of Entomology, University of California,

Davis, California 95616

Courtship behavior of butterflies ranges from extremely
stereotyped response in some species, to variability in other
species (Scott, 1973). I have found that P. coenia has perhaps
the most variable courtship of any butterfly studied to date.
Mating of P. coenia was studied at Point Richmond, Contra
Costa County, California, in connection with mark-recapture
experiments (Scott, 1975), mainly by observing natural court-
ships and by releasing females in front of resting males.

Mate-locating behavior. P. coenia is a perching species as
defined by Scott (1974a). Males perch primarily on bare spots
of ground on fairly flat areas such as roadcuts and trails on
hillsides, valley bottoms, and vacant fields. The larval foodplant
{Plantago lanceolata) is most common in such sites, and females
are most common near the foodplant. Males perch all day,
especially from 0800 to 1400 ( 24-hour standard time). After
1400 fewer males show perching behavior, but some individuals
still perch until 1600. Courtship and mating occur at the same
time as male perching. 112 courtships and 27 copulating pairs
occurred from 0815 to 1540, especially during the warmest part
of the day. Virgin females were found at all times of day, and
adults emerge from pupae in morning, indicating that some
females wait until the day after emergence to mate. Females
mate perdominently at the larval feeding sites, where males
usually perch.

On cloudy or cold days few males perch and few courtships
occur, but at temperatures of about 14-18 °C individuals are able
to perform most activities by basking (spreading the wings,
moving the fore wings forward, and orienting with the wings
nearly perpendicular to the sun). Heinrich (1973) shows that

142

14(3):142~147, 1975 COURTSHIP OF THE BUCKEYE

143

basking raises body temperature, and Blest (1975) showed that
it may also reduce predation by displaying the eyespots which
can startle predators.

Perching males sit on the ground and wait for a passing
object. When an object appears, the male approaches near.
Males investigate objects of a very wide size range. Three indi-
vidual males followed for as long as possible investigated the
following objects over a combined time span of two hours: a
car (1 investigation); motorcycle (1); birds (11); a black and
yellow butterfly (Papilio zelicaon Lucas) with 5 cm wingspan
(2); an orange butterfly {CoUas eurytheme Boisduval) with 4
cm wingspan (1); white butterfly {Pieris rapae L. ) with 3 cm
wingspan (2); other Precis coenia (32); a brown 2 cm wingspan
butterfly {Phyciodes campestris Behr) (2); a white 2 cm wing-
span butterfly (Coenonympha tullia Muller) (29); small grass-
hopper (1); small moths (2); small dragonflies (3); small bees
and wasps (7); flies (5). These males sometimes investigated
movement of vegetation due to wind, and made other investi-
gative flights toward objects which 1 did not observe.

If the passing object is not another male or female P. coenia
(of different appearance), the male usually returns to or near
the previous spot. If a male chases another male which is flying
past, the other male usually flies away when the perching male
gets within 10 cm, and the perching male returns to the vicinity
of his starting point. If two males which perch near each other
investigate each other, they may separate or both may rise high
in the air near each other then separate and descend to near
the starting points, unless the encounter carries them too far
away. Objects flying slowly (such as teneral males) seem to be
pursued farther than erratically flying or swift objects, appar-
ently because of greater resemblance to females. If a female
passes by, the male pursues her for several meters until she
lands, and courtship begins. Occasionally movement alone may
be sufficient to attract the male for mating; one deformed fe-
male which could only hop erratically on the ground was found
to be mated. Some females, especially older ones, are not
pursued at all. If two males chase laterally more than ten
meters from their previous sites, they seldom return. The males
that were watched for up to several hours gradually moved
over roughly a 30x30 m area, until they disappeared, usually
because they pursued a female or another object for 20 m or
more and did not return.

144

JAMES A. SCOTT

J. Res. Lepid.

Courtship. Three stereotyped male and one female behavior
patterns can occur in successful courtship: 1) male hovering
consists of the male hovering in air by beating his wings at
small amplitude, usually downwind just above the female;
2) male fluttering appears similar to male hovering but the male
is on the ground behind the female, and the wings are usually
moved with greater amplitude and lower frequency; 3) male
nudging consists of the male, with his wings slightly raised and
antennae pointing backward, pushing his head underneath the
hind wings of the female (which are usually spread also) until
his head is near her abdomen; 4) female flapping consists of
the female on the ground flapping her wings at wide amplitude,
continuously or in bursts (females flick their wings only occa-
sionally when crawling during other activities).

Table 1 shows the association of these behavioral elements.
In the simplest form of courtship, the female lands, raises her
wings, the male lands behind, he moves alongside and bends
his abdomen laterally (either right or left) to copulate. In a
complicated courtship, after landing the female kept her wings
spread, the male hovered then landed, the female flapped her
wings and the male fluttered his, the male moved forward and
nudged her, she raised her wings, he moved beside her and
bent his abdomen to copulate.

Temporal patterns during couttship. Male hovering is most
frequently observed just after the female lands either for the
first time or after her short flights during courtship. The male
then lands, whereupon male fluttering may occur. Male nudging
generally occurs just prior to joining. Female flapping may occur
whenever the male is on the ground just behind her. Virgins
which are not very receptive often crawl or fly a short distance
after being nudged. During courtship the female and male are
sometimes quiescent, and movement by the female often leads
to male nudging and abdominal bending.

Function of behavioral elements. Three behavioral responses
during courtship seem to function similarly to the same responses
in courtship of another Nymphaline butterfly, Poladryas 7ninuta
(Scott, 1974b). Male hovering and male fluttering are some-
what ritualized, but apparently help induce moderately unre-
ceptive females to mate. Male nudging seems to function often
merely as a technique for creeping under her wings if they are
spread to enable the male to join; it may occasionally stimulate
her to raise her wings to facilitate joining, and may also stimu-

14(3):142~147,1975 COURTSHIP OF THE BUCKEYE

145

late her to lower her abdomen so that joining may occur.
Receptive females often raise their wings vertically and un-
receptive females usually keep their wings spread. In one
case the male nudged the female while her wings were raised.
If the female does not lower her abdomen to slightly below
horizontal or if she raises it slightly above horizontal, the male
cannot grasp her. Female flapping is the “rejection dance” used
to deter males. The females in Table I which copulated usually
did not flap their wings or fly, but if they did, they flapped or
flew weakly for a short time. Unreceptive virgin females flapped
or flew less than did unreceptive mated females. When already
mated females are pursued by males, the female generally flaps
her wings vigorously upon landing, which seems to inhibit male
responses somewhat (Table 1); then he usually flies away.
The male may hover over the female for a moment. Female flap-
ping resembles male fluttering closely, so may possibly cause
the male to leave by convincing him that he is courting another
male. Further evidence of this includes several observations in
which a male chased another male, both landed on the ground
and fluttered, then both flew away, and observations of teneral
males flapping their wings when courted by another male.
Females have other rejection behavior: 1) rarely an unreceptive
female eluded a male by flying erratically. 2) an unreceptive
virgin or newly mated female crawls or flies when the male
tries to nudge or join; 3) she may raise her abdomen slightly
so that he cannot copulate; 4) she keeps her wings spread, so
that the male cannot crawl alongside to join and he must resort
to “nudging.” Courtship usually terminates unsuccessfully when
the male departs. Sometimes, when hovering or crawling be-
hind the female, he gets too far away and cannot relocate the
female.

Role of vision and odor in couHship. Movement is necessary
to elicit an investigative response by the male. By coloring all
or part of a female’s wings, I found that the general but not
the detailed color pattern of the female is important in court-
ship. Virgin females with the' upperside made completely black
or red were usually ignored by males after a brief investigation,
even though these females were courted for several minutes by
other males before the females were colored. Females with
several ocelli, white areas, or wings altered mated readily,
however: one with the white areas on the fore wings made
black; one with the hindwing ocelli made green; one with the

146

JAMES A. SCOTT

/. Res. Lepid.

ocelli of the left wings torn off. One female with crippled wings,
and another less than half the normal size, had both mated.
Males which had the dorsal wing surfaces blackened moved
and apparently courted females normally (no copulations re-
sulted because the females had already mated). One would
expect the color and pattern of the male to be relatively un-
important because the male is behind the female during court-
ship.

The female might possess a pheromone. The shortest success-
ful courtships involve nearly motionless females, which may
indicate use of a pheromone for recognition. The most interest-
ing observations concern interactions in which two males
courted the same female; in three instances after one male
joined, the other male continued attempts to join with the
female, but did not attempt to join with the male of identical
appearance. These males might have been able to recognize
the female by following her movements during the interaction,
or they might have used olfactory cues given off by one or
both sexes.

Behavior during and after copulation. Immediately after
joining, the male turns and faces opposite the female. Copu-
lation lasts an average of 27 minutes (14-59, S = 11.5, N = 14).
The female is more active than the male during copulation; she
usually spreads her wings to bask, but the males does so only
in about 30% of the copulating pairs. When disturbed, it is the
female which carries the dangling male in flight; of 12 copu-
lating pairs that were stimulated to fly, females flew 22 times,
males never flew (some copulating pairs flew more than once).
Copulation ends by the female crawling away dragging the
male, and the female kicks with the hind legs and may turn
until the male is broken off. The male then flies and the female
remains for a short time.

Duration of courtship. Successful courtship lasted from only
ten seconds when all wing movements and male nudging were
eliminated, to 10 minutes 20 seconds. Unsuccessful courtship
with virgin females lasted from 30 seconds to 20 minutes ( often ) ,
rarely up to 9 minutes. Courtships with unreceptive mated
females almost always lasted less than a minute.

Number of matings. Males probably can mate many times
since they perch and mate at all ages, but only one male is
known to have mated twice. Number of female matings was
determined by counting spermatophores. Females predominant-

14(3):142^147, 1975 COURTSHIP OF THE BUCKEYE

147

ly mate once, but occasionally mate twice and rarely three
times (Table 2).

LITERATURE CITED

BLEST, A. D. 1957. The function of eyespot patterns in the iepidoptera.
Behaviour 11:209-256.

HEINRICH, B. 1972. Thoracic temperatures of butterflies near the equator.

Comp. Biochem. Physiol. 43A: 459-467.

SCOTT, J. A. 1973. Mating of butterflies. J. Res. Lepid. 11: 99-127.

— , 1974a. Mate-locating behavior of butterflies. Amer. Midi. Nat. 91:

103-117.

— , 1974b. Adult behavior and population biology of Poladryas minuta

( Lepidoptera, Nymphalidae) and the relationship of the Texas and
Colorado populations. Pan-Pacific Ent. 50: 9-22.

— , 1975. Movements of Precis coenia, a ‘'pseudoterritorial” submi-
grant. /. Anim. Ecol. (in press).

Table 1.— -Association of some of the behavioral elements of courtship: suc-
cessful courtship of virgin females (left of the three positions separated by
two dashes); unsuccessful courtship of virgin (middle) and mated (right)
females. Terms are defined in text. Flying refers to a female flying during

courtship,

Male

hover, flutter, nudge

Female
flapping, flying

-2-

flapping

-2-

flying

neither

Totals

hover, flutter

- -1

-2-

-1-

- -1

-3-2

hover, nudge

-3-1

1-2-1

-1-

1-1-

2-7-2

flutter, nudge

2-1-1

-1-

-1-

2-3-1

hover

-3-3

1-5-3

-2-1

-5-

1-15-7

flutter

_ _

-1-2

2-1-

2-1-

4-3-2

nudge

- -1

-4-1

_ .

1- -

1-4-2

neither

-1-

1-3-15

1- -1

2- -

4-4-16

Totals

-9-6

5-20-23

3-6-2

6»8-l 14-43-32

Table 2.— -Relationship between number of matings and apparent age
(wing condition) of wild caught females. Totals are larger than sum of
females rated once or twice because some females were not graded for
wing condition.

Wing Condition

0

Young ■
38

0

0

0

Old

0

Total

38

Number of

1

23

9

12

11

7

95

Spermatophores

2

1

2

5

5

3

17

3

0

0

0

0

1

1

Journal of Research on the Lepidoptera

14(3): 148-157, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

VARIATION IN

COLIAS ALEXANDRA CHRISTINA EDWARDS

(PIERIDAE) IN SOUTHWEST MANITOBA
JOHN H. MASTERS'

8126 Santa Inez Dr., Buena Park, California

Manitoba lepidopterists have traditionally referred Mani-
toba populations of Colias alexandra Edwards to the subspecies
mayi Chermock & Chermock; incorrectly however as mayi actu-
ally refers to Colias gigantea Strecker (e.g. Masters, 1971a),
although they were assigned to Colias Christina (now considered
to be a subspecies of alexandra) by Chermock & Chermock
(1940). The assignment of mayi to gigantea, ostensibly left the
Manitoba population of Colias alexandra without a subspecific
name which, in turn, led me to a series of studies which have
culminated in this paper. In the course of this study I deter-
mined, to my complete satisfaction, that the Manitoba population
of Colias alexandra is best referred to Colias alexandra christina
Edwards of which Colias eurytheme alberta Bowman is a syno-
nym (Masters, 1971b). This paper reports on the variability of
Colias alexandra christina that is displayed in Manitoba popu-
lations.

SUBSPECIATION IN COLIAS ALEXANDRA
Colias christina (fig. 1) is the northern, predominately
orange colored population of Colias alexandra. The orange
christina phenotype occurs in the Great Slave Lake Region of the
District of Mackenzie; south to Lake Athabasca and the Peace
River Region of Alberta; thence eastward through northern Sas-
katchewan to the vicinity of The Pas, Manitoba; then south-
ward along the Manitoba Escarpment to Riding Mountain and
the vicinity of Brandon; and then appears again as an isolated
population in the Black Hills of South Dakota {krauthi Klots).
In southern Alberta and British Columbia, a broad band of
gradual intergradation to the yellow phenotype (alexandra) be-
gins. This intergradation zone continues through Montana,

^Research Associate, Carnegie Museum, Pittsburgh, Pennsylvania.

148

14(3):148-^157, 1975 COLIAS ALEXANDRA

bO

CD

hJ

C3

O

?

s

oT

m

i-O N

Is

M S

c I

e <v

Q4::

i"

O d

• ^

m §

eq

t

tL

150

JOHN H. MASTERS

/. Res. Lepid.

Idaho and northern Wyoming until typical alexandra is encoun-
tered in southern Wyoming and Colorado. The typical alexan-
dra phenotype is found from Colorado; west into Utah; south
into the White Mountains of Arizona and Sango de Cristo Range
of New Mexico; and with an isolated offshoot in the Pine Ridge
of Nebraska. The West Coast populations, are of a distinct albeit
yellow phenotype. They range from the foothills in Southern
California northward, west of the Sierra Nevada Divide and
principally in foothills, to southern British Columbia. They are
known as harfordi and barbara in the south and occidentalis
and chrysomelas in the north. Hovanitz (1950) first allied all
of these populations into one species, although this is still not
universally accepted.

The name astraea Edwards is available to apply to the alex-
andra! Christina blend zone population as a whole. Even though
central Alberta specimens are quite distinct from northwest
Wyoming ones, they are parts of the same dine.

MANITOBA POPULATIONS

In 1967, John Sorensen and I were of the opinion that Mani-
toba populations of Colias alexandra represented an undescribed
subspecies. We were working on the mistaken premise that
typical Christina was represented by Southern Alberta popula-
tions as exemplified by specimens taken near Calgary. This is
the same mistaken premise that many others (including Bow-
man, 1942) have made in dealing with Colias christina. Later
comparisons of Manitoba christina with Edwards types of chris-
tina (at Carnegie Museum), other topotypical christina, Bow-
mans (1942) description and paratypes of alberta have con-
vinced me that there are only statistical distinctions between
them-— and certainly not enough to warrant more than one sub-
specific name. Southern Alberta ‘"christina” as well as populations
from the Alberta Rockies are distinct, however, and they are,
statistically at least, distinct from typical astraea from Wyoming;
they are, as I have previously stated, part of the same dine
(between alexandra and christina) as astraea and the names
christina or alberta in the strict sense cannot apply to them at all.

I recently had the opportunity to examine and classify the
collection of Lepidoptera, now in the Manitoba Museum of
Man and Nature, assembled by Jack Dennis at Birtle, Manitoba
between 1898 and 1944. Of considerable interest in this collec-

14(3):148-157, 1975

COLIAS ALEXANDRA

151

Fig. 2. — Distribution in size of 400 specimens of C alias alexandra Christina
from Birtle, Manitoba.

152

JOHN H. MASTERS

/. Res. Lepid.

tion is a long series of C olios alexandra christina, containing over
400 specimens taken at Birtle over a 40 year span. Dennis was
evidently very fond of C olios christina and dutifully filled at
least one Hiker Mount with 12 to 18 specimens of this species
for each year he collected. This large collection, including speci-
mens collected all season long for a lengthy period of time, pro-
vided an excellent opportunity to examine variation in this
species at one given locality.

SIZE OF BUTTERFLIES. The first variable considered was
the size of specimens, this was determined by measuring the
length of the forewing to the nearest millimeter. The distribution
in size for both sexes fall into typical bell-shaped curves (figure
2.). The median forewing length for males is 25 mm. and for
females 26 mm., the average size for males slightly less than
25 mm. and for females slightly more than 26 mm. A series of
36 males and 36 females of topotypical northern Alberta C olios
christino (at Carnegie Museum) displayed slightly smaller size.
Alberta males had a median forewing length of 24 mm., with
the average slightly more than 24 mm.; the range was between
22 and 27 mm. Alberta females had a median forewing length
of 26 mm., with the average being slightly less than 26 mm.;
the range was between 19 and 29 mm.

FLIGHT PERIOD. Dennis’ capture dates were interpolated
to determine the flight period at Birtle (figure 3.). The male
flight period is between June 12th and July 5th with a peak
between June 16th and 23rd when the females are beginning
to emerge. The female flight period is somewhat later, occurring
between June 16th and July 14th and peaking between June
26th and July 2nd. The lack of Dennis specimens between July
4th and 6th is probably the result of some other activity that
Dennis was engaged in that prevented his from field collecting
at this time (perhaps he always used the week following Do-
minion Day to cut hay) rather than by an absence of individual
butterflies at this time. There is a complete lack of individuals
with collection dates during this time period throughout the
Dennis collection.

154

JOHN H. MASTERS

/. Res. Lepid.

PHENOTYPE VARIATION. C olios alexandra Christina is
remarkable in that the males are extremely regular in appear-
ance while the females are extremely variable. Males are always
bright orange, with a basal yellow area, and have uniform black
borders. Females may have an orange, yellow or white ground
color and the black bands may be absent entirely or complete.

GROUND COLOR OF WINGS. 100% of the males examined
had a uniform orange ground color to their wings. Of the fe-
males 68.1% were orange, 24.6% were yellow and 6.4% were white.
Of the Alberta sample of nominate Christina, only 27.8% of 36
females were orange, while 30.6% were yellow and 41.8% were
white. The variation in the percentage of white females in Colias
populations has been covered in detail by Hovanitz (1950b);
he recorded 9.95% white females in Manitoba Colias alexandra
and from 33.33 to 81.25% white females in various parts of
Alberta. The relative lack of white females in the Manitoba
population is unimportant from a subspeciation standpoint. The
percentage of white females shows a great deal of geographical
variation in all species of North American Colias.

BLACK BANDS OF WINGS. Males of Colias alexandra
from localities throughout their range, display consistant black
bands as in figure 1. Females, however, are quite variable show-
ing a range from immaculate, to apex band on forewing, to
outer band only, to complete band (see figure 4.). By judge-
ment, although the distinctions are not discreet, each of 220
Birtle females was placed into one of the four categories with
the following results: 1.8% showed immaculate wings, 14.5%
had an apex band only, 22.8% had an outer band only and 60.9%
had full bands. By comparison, the Alberta sample of 36 fe-
males contained 19.4% with immaculate wings, 44.4% with apex
bands only, 22.2% with outer bands only and only 13.9% with
complete bands. The bands of Alberta males are identical to
the bands of Manitoba males.

14(3):148-157, 1975

COLIAS ALEXANDRA

155

Fig. 4.— Variation in the dorsal black band patterns of female Colias olex-
andra Christina. A. -immaculate wings. B. -band at forewing apex only.
C. -outer band only. D. -complete band.

156

JOHN H. MASTERS

/. Res. Lepid.

CO

txOl
d
•H

4-1
O

U
O
1-1

o
o

d
d
o

j-i

o

SuxAaj:oj -[esjoa

uo pueg :^oexg
JO uoT:iTpuoo

Fig 5 — Table showing the co-ordinated abundance of the variation in the
background color of the wings and the extent of black maculation of 220
females of Colias dexandra christina collected at Birtle, Manitoba.

14(3):148-1S7, 1975

COLIAS ALEXANDRA

157

CO-ORDINATED DISTRIBUTION OF BACKGROUND
COLOR AND BLACK BANDS, The segregates of wing ground
color and black band condition were platted against each other
(figure 5.) to determine if there was any correlation between
the two. Apparently there isn't and the genetic conditions that
influence the black bands are independent of those that influence
ground color. Nearly half, 44,5% of the Manitoba population
displayed an orange ground color and complete wing bands.
The Alberta sample of 36 females is too small to plot when 12
variables are involved. The largest individual segregate included
7 specimens (19.4%) with a yellow ground color and with apex
bands only.

LITERATURE CITED

BOWMAN, K., 1942. A note on Colias eurytheme Bdv., with description

of a New Race ( Lepidoptera, Pieridae). Canadian Ent., 74: 25.

CHERMOCK, F. H. & R. L. CHERMOCK, 1940. Some new diurnal Lepi-
doptera from the Riding Mountains and the Sand Ridge, Manitoba.
Canadian Entomologist, 72: 81-83.

HOVANITZ, W., 1950a. The biology of Colias butterflies. I. The distribu-
tion of the North American species. Wasmann Jour. Biol., 8: 49-75.

— 1950b. The biology of Colias butterflies. II. Parallel geographic vari-

ation of dimorphic color phases in North American species. Wasmann
Jour. Biol., 8: 197-219.

MASTERS, J. H., 1971a. Concerning Colias christina mayi Chermock &
Chermock. Jour. Res. Lepid., 9(1970): 227-232.

— — — 1971b. Concerning Colias eurytheme alberta Bowman (Pieridae).
Jour. Res. Lepid., 9(1970): 97-99.

Journal of Research on the Lepidoptera

14(3): 158-168, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

THE ROLE OF WATERCRESS,
NASTURTIUM OFFICINALE

AS A HOST OF NATIVE AND INTRODUCED
PIERID BUTTERFLIES IN CALIFORNIA

ARTHUR M. SHAPIRO

Department of Zoology, University of California,
Davis, California 95616

The mustard family, Cruciferae, has a very distinctive in-
sect fauna. The chemical basis for host selection by Crucifer
insects was demonstrated as early as 1911 by Verschaeffelt, and
has more recently stimulated both physiological and ecological
studies (e.g. Thorsteinson, 1953; Schoonhoven, 1967; Feeny,
Paauwe, and Demong, 1970). Root and Tahvanainen (1969)
pointed out that, because of their dependence on mustard oils
as phagostimulants, multivoltine Crucifer insects may require
a seasonal succession of host species in a given locality. Many
Crucifers are short-lived annuals, and even many perennial
species (such as members of the genus Dentaria) are seasonally
ephemeral. In the northeastern United States wintercress,
Barbarea vulgaris R. Br., forms a critical element in the seasonal
succession because it is the only very common Crucifer which
overwinters as a well-developed rosette; it is thus available to
many insects when alternative hosts are not. In lowland Cali-
fornia most Crucifers are vernal species which disappear during
the long, hot, dry summer. On the floor of the Sacramento Valley
the introduced multivoltine cabbage butterfly, Pieris rapae Lin-
naeus (Lepidoptera, Pieridae) and the harlequin stinkbug, Mur-
gantia histrionica Hahn. (Hemiptera, Pentatomidae), are able
to continue breeding through the summer on two introduced
weedy, perennial Crucifers, Lepidium latifoliiim L. and Brassica
geniculata (Desf. ) J. Ball. (Shapiro, unpublished data). Over
a wide elevational range in the Sierra Nevada introduced water-
cress, Nasturtium officinale R. Br. ( = Rorippa nasturtium-

158

14(3):158-168, 1975

WATERCRESS AS HOST

159

aquaticum Schinz. and Thell. ) appears to have a distinctive
ecological role as a host to both native and introduced Crucifer
insects. This paper reports its impact on the seasonality and
spatial distribution of two Pierid butterflies, Piem napi micro-
striata J. A. Comstock and P. rapae.

BIOLOGY OF THE PLANT

Nasturtium officinale is a succulent, perennial aquatic weed,
native to Europe. Unfortunately there appears to be no archeo-
logical evidence to date its introduction into California ( Robbins,
1940). Major waves of weed introduction occurred in the Mission
period (1769-1800) and the Gold Rush ( 1849-1860s ) . Watercress
is widely naturalized in California, forming well-defined colonies
in shaded, quiet, shallow, flowing water from near sea level
(about 40 feet. Fair Oaks, Sacramento County) to at least 7000
feet (Nevada, Placer Counties) (fig. 1). The plant cannot tol-
erate prolonged desiccation and is thus mostly confined to
permanent streams. It colonizes intermittent springs, seeps, or
runoff channels, but behaves as an annual there and is generally
stunted. In the lower part of its elevational range it may have
two seasonal maxima of flowering, one in spring and another in
late summer. At higher elevations the flowering period is in
early- to midsummer. As the seeds ripen the tops die back and
new growth begins from the base and sometimes from nodes on
the creeping stems. The plant overwinters as rosettes, which
commence rapid growth in late winter or very early spring.
Unlike the native perennial Crucifer Dentaria californica Nutt,
which often grows near it on the sides of canyons, N. officinale
is green and available to insects for most of the warm season.
In its lower elevational range in the foothills it is normally the
only Crucifer which is green in summer.

HOST RELATIONS OF PIERIS NAPI

Pieris napi is a Holarctic species or species complex occurring
naturally (not by human intervention) in most of the cooler and
montane parts of Eurasia and North America. The subspecies
P. n. microstriata is confined to riparian habitats, generally in
canyons, on the east slope of the Coast Ranges and the west slope
of the Sierra Nevada in California. It is characteristically spring-
univoltine and monophenic, unlike the coastal fog-belt sub-

160

ARTHUR M. SHAPIRO

/. Res. Lepid.

Fig. 1.— -Habitat of watercress, Nasturtium officinale, in stream bed, Wash-
ington Creek, Nevada Co., California; male Pieris napi fly up and down the
stream bed “patrolling” for newly-emerged females.

14(3):1S8-168, 1975

WATERCRESS AS HOST

161

162

ARTHUR M. SHAPIRO

J. Res. Lepid.

species venosa Scudder (Shapiro, 1975a). In the literature its
usual host association is Dentaria californica, whose vernal-
ephemeral phenology generally fits that of the insect. Shapiro
(1975b) reported P. n. microstriata breeding on Arabis glabra
(L. ) Bernh., a native annual, but not ovipositing on Barbarea
orthoceras Ledeb., a native biennial, at Washington (2600 feet)
and Lang Crossing (4500 feet) (both Nevada County); these
are both spring plants. The development of local oviposition
preferences seems to be routine for this animal. Thus at Baxter,
Placer County (3900 feet) on 31 May 1975 three female P. n.
microstriata were observed for about 20 minutes as they ovi-
posited in a mixed stand of Barbarea verna (Mill.) Asch.,
Lepidium campestre (L. ) B. Br., and Arabis glabra: eggs were
laid only on the last. All three plant species were flowering and
beginning to set fruit, and the females visited the blossoms of
B. verna repeatedly. About 500 feet away many plants of Lepid-
ium virginicum L. var. pubescens (Greene) Thell. (mostly not
yet in flower) were growing in the road in much stronger
sunshine, and they were not visited at all. ( However, on 23 May
1975 several P. napi ova were found on this plant at Washing-
ton; see below. )

P. NAPI ON WATERCRESS

Given this selectivity, it is extremely striking that P. n.
microstriata will apparently breed on N. officinale wherever the
ranges of plant and insect coincide. The use of this plant was
first called to my attention by R. L. Langston, who subsequently
( 1975 ) published it. Langston observed ovipositions on this plant
near Placerville, El Dorado County (1850 feet). This locality
was confirmed in 1975 and, more strikingly, P. napi was found
breeding on watercress in close proximity to the localities where
the host-selection observations reported in Shapiro, 1975b had
been made. The spatial relationship between watercress and
Arabis sites at Washington and Lang Crossing is shown in
figures 2 and 3. Within the immediate vicinity of these sites the
butterfly fails to utilize Barbarea orthoceras and Lepidium cam-
pestre although these plants are growing along napi ‘Tyways”
and may even be visited for nectar. Lepidium virginicum nor-
mally grows in quite different sites in full sun, but at Washing-
ton several plants were found in shade by the roadside a few
feet from the stream bed where P. napi was breeding on water-
cress, and larvae were reared successfully to the adult from wild
ova found on these plants.

At the altitude of Washington or Lang Crossing the phenol-
ogy of N. officinale rather closely matches that of Arabis glabra,

14(3):158-168, 1975

WATERCRESS AS HOST

163

Fig. 3. — Spatial distribution of host records at Lang Crossing, Nevada
County. Symbols as in figure 2 ( see text ) .

164

ARTHUR M. SHAPIRO

}. Res. Lepid.

and the seasonality of P. napi is unlikely to be altered by its
presence. The butterfly is, however, undoubtedly able to main-
tain larger populations with than without it. Moreover, as it is
perennial and occupies the same location from year to year, it
provides a more reliable food source than does A. glabra, a
successional species whose numbers may fluctuate widely from
year to year. In this regard N. officinale resembles the native
host Dentaria.

The situation is rather different in the foothills. Here the
introduction of N. officinale may have allowed P. napi to expand
both its range and its seasonality. The butterfly is common near
Auburn in the American River gorge ( Placer - El Dorado Coun-
ties, circa 650 feet) and is generally univoltine there as else-
where in the Sierra Nevada, flying in early spring (March). In
most of the gorge N. officinale is the only Crucifer. In two sites
(fig. 4) P. napi is partially bivoltine in heavily shaded, cool,
moist ravines where it breeds on lush growths of watercress.
Although this bivoltinism appears to be due to microclimate
rather than any genetic difference from adjacent univoltine
populations ( Shapiro, in preparation ) , neither uni- nor bivoltine
P. napi could exist at all here in the absence of this introduced
plant. Since California P. napi selects shaded ravine-riparian
habitats, the preference of the plant for exactly these situations
preadapted it as a host.

HOST RELATIONS OF PIERIS RAPAE

The European cabbage butterfly is generally considered to
have entered North America in Quebec about 1860, but ReakirPs
description of Pieris yreka, which appears to be this species,
from Cahfornia as 1867 suggests an earlier introduction there,
possibly by the Spanish during the Mission period. P. rapae is
now very widely distributed, from sea level to at least 8000 feet
in disturbed habitats. It has a very long list of host records from
lowland California (Shapiro, 1975c), but individual females
tend to oviposit successively on the same Crucifer species, ignor-
ing other potential hosts (unpublished notes). On the middle
west slope of the Sierra its host preferences broadly overlap
P. napi, including watercress and Arabis glabra but extending
also to Barbarea and the two Lepidiums. Eggs and larvae of
P. rapae have been collected from both watercress and Arabis
glabra at both Washington and Lang Crossing, often from the

14(3):158-168, 1975

WATERCRESS AS HOST

165

166

ARTHUR M. SHAPIRO

}. Res. Lepid.

I

^-i

ii

14(3):158-168, 1975

WATERCRESS AS HOST

167

same individual plants as eggs and larvae of P. napi. Pieris rapae
is at least bivoltine at both stations.

In the American River gorge P. rapae (like P. napi) would
be unable to exist in most sites in the absence of watercress.
(It does breed along the arid roadside on scattered plants of
Brassica geniculata growing in disturbed soil.) In one of the
ravines where P. napi is bivoltine, an attempt was made on
1 July 1975 to collect second-generation napi larvae on water-
cress. Careful search of the entire stand produced only ten
larvae of P. rapae, despite the fact that female napi had been
flying there ten days earlier. Conditions at this site are much
more mesic than on the roadside mustard in full sun, and wild
P. rapae consistently prefers high water content Crucifers (such
as N. officinale) over low water content species (B. geniculata)
where a choice is available.

DISCUSSION

Lees and Archer (1974) group the hosts of P. napi in Britain
according to field preference. They list watercress in group 1,
i.e. plants on which females “were frequently and regularly
observed to oviposit,” and “from which larvae could readily be
obtained by searching.” There is no need to invoke genetic
adaptation to a new host by Californian populations of P. napi;
watercress seems to be intrinsically “desirable” as a napi host,
and its utilization wherever it occurs on the Sierran west slope
argues powerfully for this. It does have the potential disadvan-
tage of liability to flash flooding, but this phenomenon was not
actually observed at any Sierran locality under study in 1974
or 1975. Figure 5 summarizes the localities at which P. napi has
been found breeding on watercress to date.

There is no native, aquatic, perennial Crucifer in the Cali-
fornia flora. The native annual to biennial species Rorippa
curvisiliqua (Hook.) Bessey has a wide elevational range, but
grows in streams and seeps in meadow habitats, where Califor-
nian P. napi do not occur. (It is used by P. rapae at Donner
Pass (7000 feet) (Shapiro, 1975d) but this butterfly has not
been found on it in the Sacramento Valley. There, R. curvisiliqua
is a spring species, very infrequent and inconspicuous compared
to the rank, weedy “mustards.”) The introduction of watercress,
with its unique characteristics, permitted the expansion of the
niches of the native butterfly Pieris napi microstriata and the
introduced, weedy butterfly P. rapae in California. Its aquatic

168

ARTHUR M. SHAPIRO

J. Res. Lepid.

perennial habit ideally preadapts it as a refuge for inultivoltine
species during the hot, dry lowland California sinniner and it
may be expected to fill the same role for other members of the
Crucifer fauna.

ACKNOWLEDGMENTS

I thank Mr. Robert L. Langston for calling watercress to
my attention as a host of P. napi; Mr, William Ikitterson for
showing me the American River localities for bivoltine T. napi;
Dr. William Hovanitz for the photograph of Washington Creek
(fig. 1); Dr. Frances Chew for stimulating discussion; and the
Committee on Research of the Academic Senate, LCD, for its
continuing support of our investigation of seasonality and colo-
nizing ability in Pieris, through grant D-804.

LITERATURE CITED

FEENY, P. P., K. PAAUWE and N. DEMONG. 1970. Flea beetles and
mustard oils; host specificity of Phyllotreta cruciferae and P. striolata
adults. Ann. Ent. Soc. Arner. 63:832-841.

LANGSTON, R. L. 1975. Extended flight periods of eoastal and dune but-
terflies in Galifornia. J. Res. Lepid. 13:83-98.

LEES, E. and D. M. ARGHER. 1974. Ecology of Pieris napi (L.) (Lep.,
Pieridae) in Britain. Ent. Gazette 25:231-237.

ROBBINS, R’. W. 1940. Alien plants growing without cultivation in Gali-
fornia. Univ. Calif. Ag. Expt. Sta. Bull. 637:1-127.

ROOT, R. B. and J. O. TAHVANAINEN. 1969. Role of wintercress, Bar-
harea vulgaris, as a temporal host in the seasonal development of the
Crucifer fauna. Ann.Ent. Soc. Anier. 62:852-855.

SCHOONHOVEN, L. M. 1967. Chemoreception of mustard oil glycosides
in larvae of Pieris hrassicae. Proc. Royal Acad. Sci. ( Ain.sierdani } 70:
556-568.

SHAPIRO, A. M. 1975a. Developmental and phenotypie responses to photo-
period in uni- and bivoltine Pieris napi ( Lepidoptera: Pieridae) in
California. Trans. Roy. Ent. Soc. London 127:65-71.

1975b. Host-plant utilization by Pieris napi populations in Califor-
nia (Lepidoptera: Pieridae). Psyche 81:361-366.

1975c. The butterfly fauna of the Sacramento Valley, California.

J. Res. Lepid. 13:73-82, 115-122, 137-148.

1975d. Ecological and behavioral aspects of coexistence in six Cru-
cifer-feeding Pierid butterflies in the central Sierra Nevada. Amer.
Midi. Nat. 93:424-433.

THORSTEINSON, A. J. 1953. The chemotactic responses that determine
host specificity in an oligophagous insect ( Plutella rnaculipennis j . Can.
J. Zool. 31:52-72.

VERSCHAEFFELT, E. 1911. The cause determining the selection of food
in some herbivorous insects. Proc. Acad. Sci. ( Amsterdam ) 13:536-542

14(3): 169-188, 1975

Journal of Research on the Lepkloptera

( Continued from Vol. 14(2). -124)

TABLE 1

PLANT FORMATIONS IN THE SIERRA DE TUXTLA

A. MONTANE FORMATIONS

( 1 ) Lower Montane Rain Forest ( Terminalia-Dalber^ia
Association )

(2) Montane Rain Forest or Cloud Forest ( Eufi^leJiardtia-
Quercus Association)

(3) Liqiiidambar-Qiiercus Associes

(4) Montane Thicket (Podacarpiis-Thouinidiiini Associ-
ation )

( 5 ) Elfin Woodland ( Q tier cus-Clusia-Poda carpus
Association )

B. SEASONAL FORMATIONS

(6) Semi-Evergreen Seasonal Forest (Bursera-lnga
Association)

(7) Bursera-Sabal-Orbignya Associes

C. SEASONAL-SWAMP FORMATIONS

(8) Savanna {Curatella-Bijrsonima Association)

( 9 ) Deciduous Woodland ( Qiiercus Consociation )

(10) Pinus-Quercus Associes

D. DRY EVERGREEN FORMATION

(11) Littoral Woodland or Dry Evergreen Woodland
( Ficus-Hibiscus Association )

E. SWAMP FORMATIONS

(12) Swamp Forest (Pflc/u'ra-Fict/5 Association)

( 13 ) Mangrove Woodland ( Rhizophora Consociatioii )

F. MISCELLANEOUS FORMATIONS

(14) Recently Abandoned Milpas

(15) Pastures

( 16 ) Hedgerows

169

23

170

GARY N. ROSS

J. Res. Lepid.

oq

24

Vegetation of the Sierra de Tnxtla,

14(3):169-188, 1975 BUTTERFLIES OF TUXTLA

171

The analyses of the vegetation in the following discussion are
based on samplings by both Andrle and myself. In no instance
was an effort made to completely characterize a vegetative type.
Usually only the apparently more common members that were
either in blossom or fruit were collected. (All plant specimens
were donated to the United States National Museum. ) Through-
out the discussion plant species are listed in descending fre-
quency of abundance. Also, several terms belonging to the
science of plant ecology are used, and in order to avoid con-
fusion, are defined below.

1. Dominant~--‘'those members of the community which exert
a controlling influence over the other components” ( Beard, 1944 ) .
These were chosen empirically according to size and abundance.

2. Association—^the largest possible group which has con-
sistent dominants” ( Beard, 1944 ) . When possible I have tried to
characterize the association with the generic names of the two
most common dominants. However, in certain instances this
limitation could not be imposed so three names had to be
employed.

3. Consociation^ — “a group of equivalent rank to the associ-
ation where there is only one clear dominant” (Beard, 1944).

4. Associes— -the major unit of a sere (Clements, 1936), a
sere being defined as '‘any community which is patently in a
state of change, development or transition” ( Beard, 1944 ) .

MONTANE FORMATIONS

Although Andrle (1964) classified all of the relatively un-
disturbed and “tall forests” of the range in two categories —
“Rain Forest” and “Cloud Forest”-— he did recognize that the
formations were not homogeneous throughout their extent. My
investigations indicate that the diversification within each type
is sufficiently pronounced to warrant the formations divisions
into subcategories, which can be correlated with the various
“Montane Formations” of Beard (1955).

1. Lower Montane Rain Forest {Terminalia-Dalbergia Associ-
ation); Plate 5.

This formation encompasses the “Rain Forest” of Andrle
(1964) and exists principally on the slopes of the major vol-
canoes. On the Gulf slopes the formation occurs within an alti-
tudinal range of approximately 50 to 3,000 feet. The most
extensive areas remaining south of the high long axis of the range

25

172

GARY N. ROSS

/. Res. Lepkl.

PLATE 5

Lower Montane Rain Forest near Vigia. May 1965, 1,700 feet.

26

14(3):169-188, 1975 BUTTERFLIES OF TUXTLA

173

are in the vicinity of Cerro Cintepec, the Ciirbres de Bastonal,
and southwest of Volcan San Martin Tuxtla (Andrle, 1964).

Until five to seven years ago the forest on the Gulf slopes
was relatively undisturbed by man. But since then there has
been a rather steadily accelerating movement of people from
the more densely populated leeward slopes around and onto the
Gulf facing slopes. This in turn has resulted in forest destruction
so that presently there exist rather extensive ( but disjunct ) areas
that are already cleared or are in the process of being cleared.
On the leeward slopes the forest is very much reduced because
of several factors. First, because of extensive cultivation invol-
ving both corn and coffee, the former ccurring in no restricted
zone and the latter occurring principally in a zone between
2,300 and 2,900 feet in elevation; second, because of a relatively
low annual precipitation due to a rain shadow effect; and third,
because of unfavorable edaphic conditions, particularly on the
Santa Marta massif.

Andrle (1964) stated that the ‘'Bain Forest” seems to have its
nearest affinities with the “Seasonal Evergreen Forest” of Beard
(1944) because of the presence of an understory composed prin-
cipally of palms and because several of the dominant trees
exhibit buttressing. However, as indicated subsequently by
Beard (1949), these two characteristics are also shared by the
Lower Montane formation but that the critical factors that de-
termine the Lower Montane formation are the presence of only
two distinct tree strata and simple leaved dominants, both of
which are characteristic of the forests at relatively low elevations
in the Sierra. Furthermore, since the three formations that exist
on the slopes at higher elevations correlate very well with other
subdivisions in Beards Montane sequence, I conclude that the
forests between 50 and 3,000 feet should be considered a typical
Lower Montane Rain Forest.

The flora within this formation is very rich. Ground vegeta-
tion is relatively sparse and includes Aphelandra aurantiaca and
Didymochlaena truncatula (Andrle, 1964). Above this is a zone
of saplings and shrubs, which include H amelia longipes, Myrio-
carpa longipes, Cephaelis elata (Andrle, 1964), Psychotria sp.,
and Deherainia smaragdina. Because of the relative sparseness
of ground vegetation, walking upright in the formation is not
difficult.

The lowest tree stratum ranges between 10 and 18 feet and
is composed predominantly of palms — Astrocaryum mexicanum

27

174

GARY N. ROSS

J. Res. Lepid

PLATE 6

Montane Rain Forest or Cloud Forest on Volcan Santa Marta. April 1965
3,200 feet.

28

14(3):169~188, 1975 BUTTERFLIES OF TUXTLA

175

and Chamaedarea tepejilote being the most common. The dicot
Aegiphila costaricensis is also very common.

A distinct middle stratum cannot be. differentiated from an
upper stratum of emergents. In most cases the tallest trees —
Terminalia amazonia, Dalbergia sp. and Bernoullia flammea,
Talauma mexicana, Pithecollobium arboreum, Mirandaceltis
monoica, Phoebe mexicana, Engelhardtia guatemalemis, and
Virola guatemalensis (Andrle, 1964)-— although ranging from 90
to 110 feet in height, do not protrude significantly above the
canopy to warrant the title of emergents. Thus it is best to com-
bine them with the slightly lower species such as Pseudolmedia
oxyphyllaria, Stemmadenia galeottiana, Pleuranthodendron mexi-
cana, Calatola sp., Cletha macrophylla, Saurauia sp., Annona
sp., Coccolaba sp. (Andrle, 1964), and Rinorea guatemalensis
into a single upper stratum or canopy layer that ranges between
70 and 110 feet in height.

Lianas and the climbing fern Dryopteris sp. are common on
the trunks of many trees. Epiphytes are relatively uncommon and
occur principally in tree crowns.

Within this formation there appears to be a continual shed-
ding of leaves by the component species although there is a
heavier leaf fall near the end of the dry season (May). Only
Bernoullia flammea was observed to lack leaves for any extended
period during the dry season. Also, there appears to be no dis-
tinct flowering or fruiting season although flowering, like leaf
fall, is more common towards the end of spring.

Where relatively substantial openings exist in the forest, e.g.,
along logging roads and trails, Boeheria sp. and Urera elata
usually form dense thickets, which attain maximum heights of
8 to 12 feet.

2. Montane Rain Forest or Cloud Forest {Englehardtia-Quercus

Association); Plate 6.

This formation is located above the Lower Montane forma-
tion and ranges between approximately 3,000 and 4,100 feet on
Volcans San Martin Tuxtla and Santa Marta but to only 3,500
feet on Volcan San Martin Pajapan because of the latter’s slight-
ly lower elevation.

Ground vegetation is similar to that in the Lower Montane
formation. Above the ground cover is a zone of saplings and
shrubs of which the most common are Cephaelis elata, Chamae-
dorea ernesti-augustii, Deppea excelsa, Rudgea cornifolia, Engel-
hardtia mexicana, and Ceratozamia mexicana.

29

176

GARY N. ROSS

]. Res. Lepid.

PLATE 7

Gum-oak forest on Volcan Santa Marta.

May 1965, 2,700 feet.

30

14(3);169~188, 1975

BUTTERFLIES OF TUXTLA

177

Only two tree strata are present. The lower stratum ranges
between 15 and 30 feet and includes Eugenia sp., Chanuiedorea
elegans, Chamaedorea sp., Eupatorium tuerckheimii, Solanum
schlechtendalianum, and Carpinus caroliniana. Tree ferns,
Cyathea sp. and Akophila schiedeana (Andrle, 1964), are very
common in the numerous ravines and ridge slopes.

The upper stratum ranges between 50 and 70 feet and in-
cludes Engelhardtia mexicana, Quercus skinneri, and Rheedia
edulus.

Lianas and epiphytes are more common than in the Lower
Montane formation; epiphytes are not restricted to tree boles.
Trunks and limbs usually are festooned with mosses, algae, and
ferns.

3. Liquidambar-Quercus Associes; Plate 7.

The gum-oak forest, which appears to be a subclimax com-
munity, is restricted to a narrow zone on the southern slopes of
Valcan Santa Marta between elevations of 2,500 and 3,000 feet.
However, both dominants occur sporadically in the Montane
Rain Forest on Volcan San Martin Tuxtla and on small isolated
hill slopes northeast and northwest of Lago Catemaco.

Ground vegetation is relatively sparse although there is a
dense understory of shrubs, bushes, and saplings of which the
most common are Cephaelis elata, Siparuna andina, Phoebe
bourgeauviana, Enjthroxylon tabascense, Hirtella racemosa,
Rinorea guatemalensis, Croton glabellus, Rondelitia galeoUii,
Persea longipes, Machaonia sp., and Casearia sylvestris.

The understory forms a gradient up to the canopy layer that
ranges between 30 and 50 feet in height and which consists
primarily of Liquidambar styraciflua, Quercus ghiesbrechtii,
Belotia sp., Casearia nitida, and Alchornea latifolia.

The gum trees usually drop their leaves in early February
and remain leafless for approximately two to three weeks after
which time new growth and blossoms appear. The oaks usually
do not lose their leaves until late March or early April; new
growth and blossoms appear immediately thereafter.

Andrle ( 1964 ) suggests that the gum-oak forest exists be-
cause of a combination of factors— -destruction of the preexist-
ing vegetation by man, lowered soil fertility from extensive
weathering, and rainfall that is slightly lower than that in other
sections of the Sierra because of the “broad, high front pre-
sented by the south crater walls of Cerro Campanario and Vol-
can Santa Marta.”

31

178

GARY N. ROSS

]. Res. Lepid.

PLATE 8

Montane Thicket on Volcan Santa Marta. April 1965, 4,700 feet.

32

14(3):169~188, 1975 BUTTERFLIES OF TUXTLA

179

Although all three faetors probably are operational, it is my
opinion that the first two are of greater significance than the
third. The Popoluca Indians have been intensively utilizing the
area within the present-day gum-oak forest for the cultivation
of their corn and coffee for hundreds of years since corn and
coffee do not grow well in the pine and oak forests that surround
the Indian villages. The corn fields are used only for three to
five consecutive years; they then are abandoned and suceession
is allowed to proceed. Within 20 to 30 years a rather substantial
forest . of gum and oak becomes established. This is then cut
and burned and hence the cyele is begun anew. Very few areas
are allowed to proceed beyond the gum-oak community for
arable land is at a premium. However, those few areas that
remain uncut for longer periods develop a forest that gradually
acquires the characteristics of the Montane Rain Forest that
occurs presently slightly higher in elevation. Thus, I conclude
that intensive agriculture with its inevitable lowering of soil
fertility is the primary factor for the existence and maintenance
of the Liquidambar-Quercus Associes in the Sierra and that this
associes is a subclimax community in the Montane Rain Forest
formation.

4. Montane Thicket (Podacarpus-Thoiiinidium Association);
Plate 8.

This formation, which corresponds in part to the “Cloud
Forest” of Andrle (1964), occurs between approximately 4,100
and 4,800 feet on Volcans San Martin Tuxtla and Santa Marta
but is absent on Volcan San Martin Pajapan. In physiognomy
the forest is slightly modified from that described by Beard
(1949) inasmuch as there is a distinct and dense understory of
shrubs and small trees in addition to the canopy layer. This
understory ranges between approximately 15 and 30 feet in
height and consists principally of Clethra siiaveolens, Oreopanax
xalapense, Oreopanax capitatum, Xylosma sp. (Andrle, 1964),
Engelhardtia mexicana, Ardisia sp. (?), Thouinidiuni decan-
drum, Eugenia sp., Deppea excelsa, and Rudgea cornifolia.

The canopy layer ranges between 45 and 60 feet in height
and is composed principally of Podocarpus oleifolius, Thouinid-
ium decandrum, and Engelhardtia mexicana.

Tree trunks (usually with no extensive buttressing) and
branches support luxuriant growths of mosses, lycopodiums,
ferns, bromeliads, and orchids {Elleanthus capitatus being a
common species). Lianas aren’t as common as in the Montane

33

180

GARY N. ROSS

/. Res. Lepid.

PLATE 9

Elfin Woodland on peak of Volcan Santa Marta. April, 1965, 5,100 feet.

34

14(3):169~188, 1975 BUTTERFLIES OF TUXTLA

181

Rain Forest.

The forest on the crests of several of the steep ridges on
Volcan Santa Marta has been cut by the Popoliica Indians in
order to establish hunting trails up to the crater. In these rela-
tively open areas there is a serai community in which the palms
Chamaedorea elegans, Chamaedorea ernesti-augustii, and Cham-
aedorea sp. predominate and which tends to resemble the “Palm
Break” subclimax community of Beard (1944; 1949). However,
because of the restricted distribution of this community in the
Sierra, I think that the community does not warrant the rank of
formation.

This formation (and the succeeding one) frequently are
enveloped in clouds caused by the condensation of moist air
moving in on the north and northeast winds from the Gulf.
Although mist is more prevalent during the rainy season, there
are enough misty days during the dry season to maintain a
relatively high constant humid condition.

5. Elfin Woodland (Querciis-Clmia-Podacarptis Association);
Plate 9.

This formation is the highest in the Montane sequence and
is limited to the upper ridges and crater rims and walls of the
three principal volcanoes. The forest begins approximately at
4,800 feet on Volcans San Martin Tuxtla and Santa Marta and at
3,400 feet on Volcan San Martin Pajapan. However, on ridges
that are very steep and frequently exposed to strong winds
(particularly on Volcan San Martin Pajapan and Cerro Tuxtla),
elements of this formation occur at much lower elevations (as
low as 2,700 feet). The numerous ravines within this formation
contain elements of the Montane Thicket and/or Montane Rain
Forest.

Ground vegetation is very luxuriant and consists of a thick
mat of mosses and lichens that support profuse numbers of
orchids and bromeliads. Where the canopy is relatively open and
light penetration is good, grasses, principally Aulonemia sp. and
Isachne arundinacea, the sedge Rynchospora tuerckheimii, and
numerous small bushes and shrubs such as Miconia glaberrima,
Centropogon affine, and Sokinum spp. are common. The fern
Gleichenia palmata and the cactus Agave sp. are locally common,
especially on open, exposed ridges.

There is but one tree stratum and this consists of a gnarled,
interlaced, many branched, and almost impenetrable growth of
small trees ranging between 8 and 20 feet in height and con-

35

182

GARY N. ROSS

J. Res. Lepid.

PLATE 10

Semi-Evergreen Seasonal Forest. TOP, forest near Barrosa. Area in fore-
ground was cut and burned for corn cultivation. June 1962, 500 feet.
Photograph from Andrle (1964) and used with the author’s permission.
BOTTOM, forest in ravine on Volcan Santa Marta near Ocotal Chico. July
1965, 1,700 feet.

36

14(3);169-188, 1975 BUTTERFLIES OF TUXTLA

183

sisting of Quercus ghiesbrechtii, Clusia salmnii, Podacorpus olei-
folius, Albizia sp., Phoebe psychotrioides, Ardisia sp. (?), Wein-
mannia pinnata, Gaultheria sp., Myrica cerifem, Solarium sp.,
Ceiba pentandra, and Gymnanthes actinostemoides . Andrle
(1964) recorded the following additional species: Senecio sp.,
Hoffmania lenticillata, Viburnum acutifolium, Ilex nitida, Ore-
opanax xalapense, and Clethra suaveolens.

Practically every branch, limb, and trunk is profusely fes-
tooned with mosses, principally Pterobryum densiim and Pilo-
trichelta flexilis (Andrle, 1964), ferns, lycopodiums, bromeliads,
and orchids, principally Elleanthus capitatus. Many of the top-
most branches of the tallest trees are dead. Not all species are
evergreen; Albizia sp. (?) remains leafless during the dry sea-
son. Flowering and fruiting of all species are most common
during the spring dry season.

Landslides resulting from mild earth tremors occasionally
occur along the steep walls of the three primary craters ( particu-
larly on Volcan Santa Marta). These slides create openings in
the forest and produce optimal conditions for primary succession.
One of the most common angiosperms to appear shortly after a
slide is Schistocarpha sp.

As stated previously, this formation and the Montane Thicket
frequently are enveloped in mist, a fact that tends to make
insect collecting very difficult.

SEASONAL FORMATIONS

6. Semi-Evergreen Seasonal Forest ( Bursera-Inga Association);
Plate 10.

This formation corresponds to the “Semi-Deciduous Forest”
of Andrle (1964). Because of man’s agricultural practices, the
forest exists today only as remnants, principally on ridges and
slopes in the southern part of the range where annual precipi-
tation is usually less than 70 inches (primarily south of Cerro
Gintepec), in the vicinity of Lago Catemaco, and in the numer-
ous ravines within the Deciduous Woodland (including the
Pinus-Quercus Associes) on the Santa Marta massif.

Ground vegetation is scanty although there is a dense under-
story of saplings and herbaceous plants. Common species in-
clude Piper spp., Odontonema callistachyum, Acalypha diversi-
folia, Myriocarpa bifurca, and Heliconia latispatha. The palm
Orbignya sp. occurs sporadically throughout the formation.

Two tree strata are present. The lower stratum ranges be-

37

184

GARY N. ROSS

/. Res. Lepid.

38

Bursera-Sabal-Orhigntja Forest near Tibernal. August 1962, 200 feet. Photo-
graph from Andrle (1964) and used with the author’s permission.

14(3):169-188, 1975

BUTTERFLIES OF TUXTLA

185

tween 15 and 30 feet in height and is composed principally of
Cecropia mexicana, Acalypha diversifolia var. carpinifolia, and
Tabernaemontana citrifolia. Along streams Erythrina mexicana
is common.

The upper stratum ranges between 40 and 60 feet in height
and consists primarily of Bursera simaniba, Inga spuria, Inga
leptoloba, Luehea speciosa, Myrcia splendens, Albizia idiopoda,
Dendropanax arboreus, Ilex belizensis, and Roupala borealis.

Trees usually branch low and the boles frequently are
umbrella-shaped. Buttressing is uncommon. Trunks usually sup-
port numerous lianas and vines (such as Anguria tabascensis) .
Epiphytes are relatively uncommon.

During the dry season several of the dominants drop their
leaves and remain leafless until the onset of the summer rains.

7. Bursera-Sabal-Orbignya Associes; Plate 11.

In the extreme southwest section of the range and at slightly
lower elevations than the Semi-Evergreen Seasonal Forest is
found a community that appears to be of subclimax rank. This
forest is composed principally of Bursera simaniba and the
palms Sabal sp. and Orbignya sp. Other trees include Cecropia
mexicana, Inga spuria, Cassia spectabilis, and Cassia occiden-
talis. There is no definite canopy since the trees usually exist in
dense, disjunct stands separated by extensive tracts of coarse
grasses, sedges, and herbaceous plants of which the most com-
mon are Paspalum spp., Sporobolus spp., Rynchospora spp.,
Dichromena ciliata, Asclepias woodsoniana, Melanthera angusti-
folia, and Stemodia durantifolia. These open areas seem to be
the result of, and, to be perpetuated by repeated burnings by
the local Mexicans and intensive pasturing by livestock.
SEASONAL-SWAMP FORMATIONS

8. Savanna (Curatella-Byrsonima Association); Plate 12.

This formation, which seems to correlate well with the
Orchard Savanna of Beard (1953), occupies a rather restricted
area in the Sierra, principally south and southwest of the Santa
Marta massif and at elevations below 450 feet. The formation
intergrades with both the Semi-Evergreen Seasonal Forest and
the Deciduous Woodland where contact exists.

39

186

GARY N. ROSS

J. Res. Lepid.

40

PLATE 12

Javanna on Volcan Santa Marta near Guasuntlan. May 1962, 200 feet,
‘holograph by R. F. Andrle.

14(3):169-188, 1975

BUTTERFLIES OF TUXTLA

187

Ground cover within the savanna is of variable density and
consists principally of grasses, sedges, and woody plants of which
the most common are Paspalum spp., Panicum sp., Dichromena
ciliata, Rynchospora spp., Asclepias woodsoniana, Stemodia
dumntifolia, and Melanthera angustifolia.

The formation is rather open. Common trees include Ciira-
tella americana, Byrsonima crassifolia, Apeiba tibourbou, Quer-
cus oleoides, and Spondias mombin (Andrle, 1964). These attain
maximum heights of 10 to 20 feet.

Epiphytes are uncommon. Although leaves of most trees are
shed annually (usually at the end of the dry season. May, or
after the passage of fire), the trees never remain leafless for any
extended length of time.

The reasons for the existence of this formation are debatable
as they are for most other tropical savannas. Budowski (1959)
states that experimental evidence indicates that all savannas
would revert eventually to forest if fire is precluded from the
area and if a seed source is near. However, there is not uni-
versal agreement on this matter. Beard (1953) states that
‘savanna may be characterized as the vegetation of the highly
mature soils of senile land formations . . . which are subject to
unfavorable drainage conditions in the form of intermittent
perched water tables, with alternating severe periods of water
logging and dessication.” Furthermore, he continues and states
that although the savanna may be swept by regular fires and
the vegetation be adapted as to be fire resistant, the vegetation
is not dependent upon fire for its maintenance but is an edaphic
climax.

The Beard hypothesis seems to be the more reasonable ex-
planation for the existence of the savanna in the Sierra. First,
the formation occurs in one of the most ancient geological areas
in the range (the Santa Marta massif) and the grey to black
clay soils probably indicate severe leaching has occurred
( Friedlaender, 1923 ) . Second, the land is of low relief and there
are numerous outcroppings of bedrock, two factors that proba-
bly make drainage relatively inefficient. Third, the annual pre-
cipitation, as recorded at Guasuntlan, averages approximately
67 inches and there are at least five months of the year that
receive less than four inches of rainfall (Andrle, 1964). Fourth,
the area has very few human inhabitants (and probably has
had very few in the past) and so man-caused fires are relatively
uncommon.

41

188

GARY N. ROSS

]. Res. Lepid.

42

PLATE 13

Deciduous Woodland on Volcan Santa Marta near Soteapan. June 1965,
1,700 feet.

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THE JOURNAL OF RESEARCH
ON THE LEFIJDOFTERA

Volume 14 Number 3 September, 1975

IN THIS ISSUE

Courtship and mating behavior of the

Fiery Skipper, Hylephila phylaeus ( Hesperiidae )

Irene Shapiro 125

Variability of courtship of the Buckeye Butterfly,

Precis coenia (Nymphalidae)

James A. Scott 142

Variation in C alias alexandra Christina Edwards
(Pieridae) in Southwest Manitoba

John H. Masters 148

The role of watercress. Nasturtium officinale as a
host of native and introduced pierid butterflies
in California Arthur M. Shapiro 158

An ecological study of the butterflies of

the Sierra de Tuxtla in Veracruz, Mexico

(continued) Gary N. Ross 169

Volume 14

Number 4

December, 1975

published by

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Journal of Research on the Lepidoptera

14(4):189-212, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A REVISION OF

THE NORTH AMERICAN COMADIA

(COSSIDAE)

RICHARD M. BROWN

1055 Plaza Drive
Martinez, California 94553

This paper is the result of my attempt to confirm and place
in the genus a species that was described as Comadia suaedivora
Brown and Allen (1973). During this work I discovered the
lack of any recent literature and the existence of many speci-
mens, mostly from California, that did not fit the previously
described species.

The material used in this study was borrowed from the fol-
lowing institutions: California Academy of Sciences, San Fran-
cisco; Canadian National Collection, Ottawa, Ontario; Carnegie
Museum, Pittsburgh, Pennsylvania; The American Museum of
Natural History, New York; California Department of Agricul-
ture, Sacramento; Los Angeles County Museum of Natural His-
tory, Los Angeles, California; University of California, Berkeley;
United States National Museum, Washington, D.C.

Approximately 450 specimens were used in this study, in-
cluding the Allotype of Heterocoma albistriga Barnes & Mc-
Dunnough, the genitalia of the Holotype of Comadia bertholdi
bertholdi (Grote), and a number of paratypes of various other
species. Dr. Ronald W. Hodges kindly compared specimens to
additional material in the U.S. National Museum.

COMADIA BARNES AND McDUNNOUGH
Bombyx Linnaeus, 1785, Syst. Nat., 1:496.

Zeuzera Latreille, 1804, Noiw. Diet. d’Hist. Nat., 24:186.
Hypopta Hiibner, 1816, Verz, beck. Schmett., pg. 195; Neu-
moegen & Dyar, 1893, Joiirn. N. Y. EntomoL Soc., 1:32; ibid,
1894, Joiirn. N. Y. Entomol. Soc., 2:164.

189

190

RICHARD M. BROWN

/. Res. Lepid.

Fig. 1, Coniadia Jietirici, Blyth, Riverside Co., California; Fig. 2, Coin-
aclia suaedwora, paratype, Tipton, Tulare Co., California; Fig. 3, Comadia
dolli, Soldier Meadows, Humbolt Co., Nevada; Fig. 4, Comadia intnisa,
Santa Monica, Los Angeles Co., California; Fig. 5, Comadia arenae, Holo-
type, Wheeler Spring, Ventura Co., California; Fig. 6, Comadia suhtermi-
nata, Tonto Creek Camp, Gila Co., Arizona; Fig.- 7, Comadia speratus,
Holotypc, Madera, Madera Co., California.

14(4) -.189-212, 1975

COMADIA REVISION

191

Comadia Barnes & McDunnough, 1911, Contrib. Natur. Hist.
Lepid. of N. Amer., 1:26-29; Barnes & Benjamin, 1923, Con-
trib. Natur. Hist. Lepid. of N. Amer., 5:88-96.

Heterocoma Barnes & McDunnoiigh, new synonymy, 1918, Con-
trib. Natur. Hist. Lepid. of N. Amer., 4(2):179; McDun-
nough, 1939, Check List of Lepid. of Canada and U. S. Amer.,
Part 2, Microlepid. pg. 62.

Head: proboscis absent; clypeus flat, level with or recessed
below level of eyes. Male antennae bipectinate to end, pecti-
nations arising from middle of segments on ventral half, each
pectination with inner surface pubescent, terminating with one
seta, a dorsal nob with one seta three fourths from base; female
antennae serrate, shallowly bifid, each terminating with one or
two setae. Thorax: epiphysis simple, less than tibial length;
hind tibiae swollen; all spurs present. Wings (fig. 15): forewing
Sc. free, Ri from middle of discal cell, R2 from top of areole,
R3 from end of areole, R.i and R.-, stalked from R3, Mi usually
from top angle of discal cell, M2 and M3 from below center of
discal cell, Ai and A2 free; hindwing Sc. free, R and Mi usually
from a point, but may be separated or stalked, Cui from lower
angle of discal cell, Cru from bottom of discal cell, Ai, A2, and
A3 free; cellula intrusa in both wings. Abdomen clothed with
long hairs laterally lengthened. Male genitalia (fig. 17); harpe
upturned, more or less squared posteriorly, inner surface slight-
ly excavated; valvula with strong hooked process at base just
below the costa; tegumen broad, hood shaped with 2 to 6 setae
at anterior corners of base; uncus small, deflexed as a short,
strong hook; gnathos spoon shaped with long diverging arms;
aedeagus as simple sclerotic tube slightly deflexed from middle,
without internal sculpturing. Female genitalia (fig. 16); ovi-
positor simple, long and tapering; posterior and anterior apophy-
sis long, well developed; ovipositor lobe densely covered with
setae, as is posterior edge of segment 8; ductus bursa and bursa
membranous, without any chitinized structure.

KEY TO THE SPECIES OF COMADIA

( Based on male characters )

1. Both wings light buff or cream colored; without dark brown

discocellular spots 2.

Both wings pale gray to dark gray or brown, with or without
dark brown discocellular spots 3.

192

RICHARD M. BROWN

/. Res. Lepid.

Fig. 8, Comadia J)ertholdi indistincta, Holotype, 8 mi. w. Fort Jones,
Siskiyou Co., California; Fig. 9, Comadia hertholdi polingi, Kenwortliy,
Riverside Co., California; Fig. 10, Comadia hettholdi polingi, paratype.
Bent, Otero Co., New Mexico; Fig. 11, Comadia alleni, Holotype, La
Grange, Stanislaus Co., California; Fig. 12, Comadia manfredi, Oro Blanco
Mts., Santa Criiz Co., Arizona; Fig. 13, Comadia redtenhacheri. Big Bend
National Park, Brewster Co., Texas; Fig. 14, Comadia alhistriga, Alamo-
gordo, Otero Co., New Mexico.

14(4):189-212, 1975

COMADIA REVISION

193

2. Wings not more than 10 inin. long, cream colored, immacu-
late, epiphysis one half tibia length ..manfredi.

Wings at least 12 mm. long, light buff with white markings;
epiphysis greater than one half tibia length . henrici.

3. Upper forewing with at least some dark discocellular scal-
ing present; wing not crossed by lines formed by bicolored

scales - - 4.

Upper forewing without discocellular scaling; wing crossed
with numerous lines formed by an alignment of bicolored
scales; epiphysis quite short .....redtenhacheri,

4. Discocellular spot divided by prominent whitish discal bar
extending to Ciu in the tornal area; a white bar along cubi-
tal vein to discal bar; white subterminal line; epiphysis

minute or missing albistriga.

Upper forewing not so divided by white bar; epiphysis well
developed 5.

5. Upper surface of wings dark gray to brown 6.

Upper surface of wings silver gray to white 8.

6. Upper forewing with subterminal line; discocellular spot
covering entire end of discal cell ( Utah, Colorado, Arizona,

New Mexico) ...suhterminata.

Upper forewing usually without postmedial line; discocellu-
lar spot reduced to lower angle of discal cell (California) 7.

7. Upper fore wing heavily dusted with gray; subcosta very
white, discal cell with some white (San Joaquin

Valley) suaedivora.

Upper forewing lightly dusted with brown scaling; subcosta
concolorous to wing, not white (coastal Southern Califor-
nia ) intrusa.

8. Upper forewing pale tan or buff with scattered brown

scales; veins lightly marked; discocellular spots relatively
undefined; usually with some indications of subterminal

lines dolli.

Upper forewing more gray or white; without general scat-
tering of dark scales; discocellular spots well defined, but
may be reduced in size 9.

9. Upper forewing light gray; discocellular spot greatly re-

duced, restricted to lower distal corner of discal cell .arenae.
Upper forewing light gray or white; discocellular spot prom-
inent . - 10.

194

RICHARD M. BROWN

J. Res. Lepid.

Fig. 15, Wing veination: A, right forewing, B, right hindwing; Fig. 16
Female genitalia; Fig. 17, Male genitalia: A, tegumen with gnathos and
uncus, B, left harpe, C, aedeagus.

Distributional maps for Comadia.

14(4):189-212, 1975

COMADIA REVISION

195

10. upper forewing white; veins unlined with dark brown ..-11.

Upper forewing grayish; veins usually lined with dark
brown, except for herthoJdi indistincta 12.

11. Discocellular spot dark, defined and contrasting with white
wing; overlay of gray restricted to submarginal and apical

region speratus.

Discocellular spot poorly defined; upper forewing with over-
lay of gray heavy except for submarginal and discal area,
which is white alleni.

12. Discocellular spot restricted to lower angle of discal cell;

veins lined dark to margin (Great Basin Region) 13.

Discocellular much reduced; veins unlined (North and

Coastal California) hertholdi indistincta,

13. Upper forewing light gray to cream with light suffusion of

dark brown; subcosta and discal cell very white; median
vein at base of discal cell dark brown; fringe well checker-
ed at end of dark veins hertholdi polingi.

Upper forewing gray with heavy suffusion of dark brown;
subcosta and discal cell with general wing suffusion; median
vein at base of discal cell with dark brown greatly reduced
or missing; fringe lightly checkered at end of dark
veins hertholdi heHholdi.

COMADIA HENRICI (Grote)

(fig. 15)

Hijpopta henrici Grote, 1882, Papilio, 2:31; Dyar, 1902, Bull.
U.S.N.M., 52:363; Neumoegen & Dyar, 1893, Journ. N. Y.
Entomol. Soc., 1:33; ibid., 1894, Journ. N. Y. Entomol. Soc.,
2:165; Holland, 1903, Moth Book -.379, .PI 12, Fig. 3; Dyar,
1937, in Seitz, Macrolepidoptera, 6:1275, PI. 182.

Coinadia henrici; Barnes & McDunnough, 1911, Contrih. Natur.
Hist. Lepid. N. Amer., 1:28, PI. 5, fig. 5; ibid., 1917, Check
List of the Lepid. of Boreal Anier.: 195.

Male: Head, thorax and abdomen light cream colored; long-
est antennal pectination four times shaft width; antennal pectin-
ation/ eye diameter ratio average is .42; epiphysis/ tibia ratio
average is .67. Upper surface of wings, costa with buff scales
scattered or in spots to apex; subcosta white; area between radial
and cubital veins to submarginal area buff; remainder with buff
spots between veins; basal half of cell M3 white, remainder buff;
cell Gui buff to fringe; basal two thirds of cell Guo and anal cell

196

RICHARD M. BROWN

J. Res. Lcpid.

14(4);189-212, 1975

COMADIA REVISION

197

buff. All veins white; fringe white. Hindwings uniformly light
cream, veins may be slightly darkened. Under surface of wings,
with forewing costa somewhat darker than upper surface; dark
scaling more diffuse, following general pattern of upper surface;
hindwing concolorous to upper surface. Female; similar to male;
abdomen longer, heavier. Wing size in male 12U5 mm; in
female 16-20 mm.

Range.-— California, Arizona, Utah, see map # 1. Flight
period, March through May.

This species is recognized by its pale cream and buff color.
The antennal pectination in the California specimens are slight-
ly shorter than in those from Arizona and Utah. This species has
the longest epiphysis in the genus. Of the 116 specimens exam-
ined, the majority are from Southern California.

COMADIA SUAEDIVORA Brown & Allen

(fig. 2$ )

Comadia suaedivora Brown and Allen, 1973, Pan-Pacif. Ento-
mol., 49:240.

Male: Head and thorax with mixture of white and brown
scales; longest antennal pectination three times shaft width;
antennal pectination/ eye diameter ratio average is .42; epiphy-
sis/tibia ratio average is .59. Upper forewing mostly fuscous;
subcostal to subapex with discal cell invaded by white; disco-
cellular spots reduced and restricted to lower angle of discal
cell; usually a faint postmedial line present. Hindwing con-
colorous with abdomen. Under forewing costa with basal three-
fifths dark brown, remainder white; dark siibapical spot; post-
medial line present; distally dark between veins. Hindwing
creamy white with dark brown suffusion. Female: Similar to
male, somewhat darker and larger. Wing size in male, 12-16mm;
in female 12-17mm.

Early stages.— Brown and Allen (1973) reported that the
larvae feed gregariously in the crown and roots of Suaeda
fruticosa (L.) Forsk. The larva is rose-lavender colored with a
heavily sclerotized horn on the dorsal surface of the anal flap.
The pupa is dark brown and heavily spined. Pupation occurs in
a subterranean chamber, but prior to emergence the pupa leaves
the chamber and extends part way above the ground surface.

Range.— San Joaquin Valley in California, see map #1. Flight
period, May and June.

198

RICHARD M. BROWN

/. Res. Lepid.

Comadia suaedivora is nearly the darkest species in the genus,
only subterminata is darker. However, suaedivora is differentia
ated by its white subcosta, contrastingly lighter hindwings and
geographical distribution. Number of specimens examined, 99.

COMADIA DOLLl Barnes & Benjamin
(fig. 35 )

Comadia dolli Barnes & Benjamin, 1923, Contrib. Natur. Hist,
Lepid. N. Amer., 5:90.

Hypopta dolli; Dyar, 1937, in Seitz, 6:1275.

Male: Head with cream and brown scales; antennae with
cream colored scaling to end, longest antennal pectination is
2. 5-3.0 times shaft width; antennal pectination/eye diameter
ratio average is .36; epiphysis /tibia ratio average is .59. Thorax
cream colored; collar blackish; partagia tipped in black. Abdo-
men lighter in color than thorax. Upper wings, ground color
creamish-white; costa checkered, subcosta to subapical and up-
per part of discal cell white; rest of wing with scattered brown
scales; discocellular spots reduced, with only the veins outlined
darker, except for Cii, and Cuj which are whitish; end of veins
brown; fringe checkered at end of veins. Hindwings concolorous
or slightly darker, veins lined darker. Under wings cream to
fuscous; fore and hindwings concolorous. Female: Wing length
same as in the male, abdomen longer; upper surface forewing
with discocellular cell wanting; wing scattered with brown
scales, some forming short reticulations between veins. Hind-
wings as in male. Wing size 12-15mm.

Range. — California, Nevada, see map #1. Flight period, April
through July.

This moth has the patches at the end of the discal cell re-
duced and restricted to the lower angle and cubital veins.
Comadia dolli is very close to bertholdi in size and markings,
but has longer antennal pectinations, longer epiphysis and a
tendency to subterminal lines. Also, dolli tends to have a cream
coloration similar to henrici, as opposed to the gray-white of
bertholdi. Number of specimens examined, 16.

COMADIA INTRUSA Barnes & Benjamin
(fig. 45 )

Hifpopta bertholdi; Rivers, 1897, Psyche, 8(249) :10; Dyar, 1897,
' Psyche 8(249);10; Holland, 1903, Moth Book:379, PI. XII
fig. 2.

Comadia intrusa Barnes and Benjamin, 1923, Contrib. Natur.

Hist. Lepid. N. Amer., 5:92.

Hypopta intrusa; Dyar, 1937, in Seitz, 6:1275.

14(4):189-212, 1975

COMADIA REVISION

199

Male: Head and thoracic vestitiire with white and dark-
brown scales; longest antennal pectinations two times shaft
width; antennal pectination/ eye diameter ratio average is .29;
epiphysis/ tibia ratio average is .59; abdomen not darker than
thorax. Upper forewing white with a light scattering of pale
brown; discocelkilar spots pale and diffuse; veins outlined
darker brown; reticulated siibmargin present. Hindwing light
fusciis. Under forewing with co-sta checkered or solid brown on
basal half, remainder white; remainder ventral surface fiiscus.
Under hindwing concolorous with forewing. Female: Larger,
with markings less defined than in male. Wing size in male
13-17mm; female 20mm.

Early stages. — Rivers (1897) in his brief account of the
larvae of ''Hypopta bertholdi Grote” reports that its food “con-
sists of the fibre of the main stem and larger roots of the lilac
flowered lupin,” and that the presence of larvae of different ages
in the same plant implies that larval growth extends beyond one
year. The larvae wander some distance from the plant and pu-
pate in a subterranean chamber more than one foot deep. Just
prior to eclosion the pupa extends part way above the ground
surface.

The color of the larva when about one half inch long is a
yellow white, but when larger it becomes a bright carneliaii red,
heightened by an enameled surface (Rivers, 1897). Additional
characterization of the larvae by Dyar (1897) tells of the body
being ventrally flattened, with a large black recurved horn above
on the anal flap. Thoracic legs are small and pointed; abdominal
legs are very short with crotchets distinct and arranged in two
long parallel transverse rows.

Range. — California, see map #2. Flight period, June through
August.

Dyar (1897) followed Rivers (1897) in recognizing the
southern California larvae as Htjpopta bertholdi Grote, not
realizing an undescribed species was involved. Holland (1903)
figured a female he called H. bertholdi. I have a female intriisa
from the Holland collection labeled “Moth Book Plate XII,
fig. 2,” that matches the specimen in the plate. Some intnisa
strongly resemble suhterminata, but do not have dark suffusion
or the prominent reticulations found in siibterminata. The an-
tennal pectination in this species is the shortest found in Coma-
dia. Specimens examined, 11.

200

RICHARD M. BROWN

/. Res. Lepid.

r

I

K

i

14(4):189-212, 1975

COMADIA REVISION

201

COMADIA ARENAE Brown, new species
(fig. 5« )

Male: Head and thorax whitish-gray with a few brown
scales; antennal shaft dorsally scaled, longest pectination twice
the shaft width; antennal pectination/ eye diameter ratio average
is .29; epiphysis/tibia ratio average is .55. Abdomen concolorous
with thorax. Upper forewing concolorous with body; disco-
cellular spot represented by a few dark brown scales and ocher
shading; base of cell Cui ocher; middle of cell ocher; veins
unmarked; fringe wide and white. Hindwings mouse-gray, veins
lightly marked; fringe as on forewing. Under forewing costa with
white and dark brown scales mixed to three-fifths from base,
remainder white; remainder of wing light fuscous; fringe white.
Hindwing lighter; fringe white. Wing length in Holotype 15mm;
variation, 13-17mm.

Holotype male, California, Ventura County, Wheeler Springs,
29 July 1943, Don Meadows. Paratypes: 4$ , same locality and
collector as holotype, 1 ^ , 21 July 1943, 1 ^ , 27 July 1943,
2 ^ , 29 July 1943. Type disposition: The Holotype and three
paratypes are at the Los Angeles County Museum of Natural
History, Los Angeles, California; one paratype at the California
Academy of Sciences, San Francisco. See distribution map #2.

This species is somewhat variable, with the discocellular
spot greatly reduced or absent. There is no dark scaling on the
veins and no concentration of dark suffusion on the upper fore-
wing; this gives a uniform, smooth appearance. Number of
specimens examined, 5.

COMADIA SUBTERM IN AT A Barnes & Benjamin

(fig. 6s }

Comadia subterminata Barnes and Benjamin, 1923, Contrib.
Natur. Hist. Lepid. N. Amer., 5:91.

Hypopta subterminata; Dyar, 1937, in Seitz, Macrolepidoptera,
6:1275.

Comadia bertholdi fusca Barnes and Benjamin, 1923, Contrib.
Natur. Hist. Lepid. N. Amer., 5:92.

Male: Head, palpi, front and vertex concolorous with or
darker than thorax; thorax with mixture of black and white
vestiture; longest antennal pectination 2.5-3. 0 times shaft width;
antennal pectination/eye diameter ratio average is .36; epiphy-

202

RICHARD M. BROWN

/. Res. Lepid.

sis/tibia ratio average is .56. Upper forewing with ground color
fuscus gray; costa with dark brown checks, in some specimens
becoming a solid border; subcostal area and discal cell whiter;
submarginal area with reticulations, usually consisting of one
line that fades before reaching inner margin; discocellular spots
quite heavy, extending cephalad to radials and caudad to vein
A2; veins lined with brown. Hindwings light fuscous with veins
lined darker. Under forewing costa dark brown, broken by two
or three white spots two thirds out from base; remainder of
wing fuscus. Under hindwing same as upper surface. Female:
Larger than male, markings similar to male, but less distinct.
Wing length in male, 13-18mm; in female 19-21mm.

Range. — Utah, Colorado, Arizona and New Mexico, see map
#2. Flight period, March through July.

This species is closely related to hertholdi, but is distin-
guished from hertholdi by its larger size (wing averages 2mm
longer) and increased amount of dark scaling. Barnes and
Benjamin ( 1923 ) described fiisca as a subspecies of hertholdi.
In a preliminary examination Rodald W. Hodges ( in corres-
pondence) suggested that fiisca and suhterminata were actually
the same species; in my research I have been unable to find any
reason to keep them separate species. Number of specimens
examined, 24.

COMADIA SPERATUS Brown, new species
(fig. 7 S )

Male: Head, thorax white with pale fuscus shading about
the collar; palpi quite black on the outer surface, white in inner
surface; antennal shaft white scaled to tip; longest antennal
pectination three times shaft width; antennal pectination/eye
diameter ratio is .36; epiphysis /tibia ratio is .56. Abdomen white.
Upper surface of wings with ground color very white; costa with
dark brown scaling; discocellular spot at base of cells Mo and
Cui; a brown spot mid-cell A^; heavy suffusion of dark brown
between discal cell and apex; discal cell and subcosta with less
dark scaling; fringe checkered. Hindwing mouse gray, veins
darker. Under forewing costa black three fifths out from base,
remainder white; remainder of wing fuscus except for discal
cell. Hindwing costa white; remainder of wing concolorous to
forewing. Female unknown. Wing length in holotype, 15mm.

Holottjpe male, California, Madera County, Madera, 14 May
1962, H. E. Gleason. Paratypes, 2 $ , same data as Holotype. See
distribution map #2.

14(4):189-212, 1975

COMADIA REVISION

203

Type disposition. — The Holotype is at the California Acad-
emy of Sciences, San Francisco. Two paratypes are at the Cali-
fornia Department of Agriculture, Sacramento.

This is the whitest of all the Comadia. The black shading
on the forewing when viewed together form a diagonal band
across the wing. The white color of this moth readily distin-
guishes it from the other Comadia. Number of specimens exam-
ined, 4.

COMADIA BERTHOLDI (Grote)

Hypopta hertholdi Grote, 1880, Bull. Brook. Entomol. Soc. 3:45.

Male: Head and thorax with black and white mixed vesti-
ture; abdominal color slightly lighter in color. Upper surface
forewing from very light gray to a dark ash, with some black
suffusion; heaviest suffusion obliquely from end of discal cell
to apex; cubitus and end of discal cell caudad of Mo dark brown;
base cell Cui and Mo with scaling as on cubitus; base of cell Ms
usually without dark scaling; basal one third to one half of
radius with dark scaling; remaining veins may or may not be
lined in dark brown; fringe lightly checkered, beige to gray-
brown. Under surface forewing costa dark brown or black;
remainder of forewing and all of hindwing white to gray-brown.
Female: Larger than male; ground color as in male; the dark
color on the cubitus and at end of discal cell less distinct.

The slightly different maculation in each population of the
variable species has caused the proliferation of names under the
nominate subspecies. At present there are three subspecies; the
nominate subspecies is from the Great Basin, C. hertholdi polingi
is from the southwest and a new subspecies is from California.

COMADIA BERTHOLDI BERTHOLDI (Grote), new status

(no figure)

Hypopta hertholdi Grote, 1880, Bull. Brook. Entomol. Soc., 3:45;
Neumoegen & Dyar, 1893, Journ. N. Y. Entomol. Soc., 1:33;
ibid., 1894, Journ. N. Y. Entomol. Soc., 2:165; Rivers &
Dyar, 1897, Psyche, 8:10; Holland, 1903, Moth Book: 379,
PL XII fig. 2; Dyar, 1937, in Seitz, Macrolepidoptera, 6:1275,
PI. 182; Dyar, 1902, Bull. U. S. N. M., 52:363.

Comadia hertholdi; Barnes & McDunnough, 1911, Contrih. Na~
tur. Hist. Lepid. N. Amer., 1:27; ibid., 1917, Check List of
the Lepid. of Boreal Amer.: 195.

204

RICHARD M. BROWN

/. Res. Lepid.

'A

14(4):189-212, 1975

COMADIA REVISION

205

Hijpopta edwardi Neiimoegeii & Dyar, 1893, Journ. N. Y. Ento-
moL Soc., 1:32; Dvar, 1937 in Seitz, Macrolepidoptera,
6:1275; Dyar, 1902, Bull U. S. N. M., 52:363.

Comadia edwardi; Barnes & McDiinnoiigh, 1917, Check List of
the Lepid. of Boreal Amer.: 195.

Comadia engelhardti Barnes & Benjamin, 1923, Contrib. Natur.
Hist. Lepid. N. Amer., 5:89.

Hypopta englehardti; Dyar, 1937, in Seitz, Macrolepidoptera,
6:1275, PL 182.

Comadia stabilis Barnes & Benjamin, 1923, Contrib. Natur. Hist.
Lepid. N. Amer., 5:90.

Hypopta stabilis; Dyar, 1937, in Seitz, Macrolepidoptera, 6:1275.

Male: Longest antennal pectination three times shaft width;
antennal pectination/eye diameter ratio is .34; epiphysis /tibia
ratio is .56. Upper surface of forewing with prominent dark
suffusion; cubitus and end of discal cell caudad of M2 with
heavy dark brown scaling; remainder of wings as described in
general characterization. Female as described. Length of fore-
wing: male 13-16mm; in female 18-19mm.

Range. — California, Colorado, Wyoming, see map #3. Flight
period, June through August.

This is the darkest of the three subspecies and the most
variable. I have not seen C. edwardi, but have placed it as a
synonym of bertholdi bertholdi. Barnes and Benjamin (1923)
doubted the validity of edwardi as a species, believing it to be
a color form or an abberant individual. Neumoegen and Dyar
(1893) in their original description state that the type is from
Colorado, which lends support to this concept. Number of speci-
mens examined, 45.

COMADIA BERTHOLDI INDISTINCTA Brown,
new subspecies

(fig. 8)

Male: Similar to bertholdi bertholdi except cubitus with dark
markings reduced; markings at end of cell less distinct. Female:
Larger than male; dark scaling so reduced that the wings are
nearly immaculate. Wing length in Holotype, 16mm.; in Allo-
type, 18mm,

Early stages. — The Allotype and one female paratype are
labeled “Lupinus”, presumably the host of this species, as the

206

RICHARD M. BROWN

/. Res. Lepid.

perennial lupine is the host for C. intnisa in Southern Cali-
fornia.

Holotype male, California, Siskiyou County, 8 miles west of
Fort Jones, 21 June 1971, F. D. Horn. Allotype, California, Con-
tra Costa County, Orinda, 18 July 1949, E. G. Linsley. Para-
types: California, Contra Costa County: Orinda, 1 9 , 18 July
1949, E. C. Clark; Kern County: Mt. Pinos, 1 S , 12 July 1953,
Lloyed M. Martin; Napa County: Mt. Saint Helena, 2 <5 , 30 June
1956, W. R. Bauer and J. S. Buckett; Siskiyou County: Etna,
4 ^ , 16 July 1970; 3 miles east of Etna, 3^,11 July 1971, F. D.
Horn. Holotype and allotype are deposited at the California
Academy of Sciences, one paratype is in the collection of the Los
Angeles County Aluseum of Natural History, one paratype is in
the California Insect Survey, University of California, Berkeley,
and the remainder are at the California Department of Agricul-
ture, Sacramento. See distribution map #3.

This subspecies is distinguished from b. hertholdi by the
reduced, indistinct brown discocellular spots. Antennal pecti-
nation/eye diameter ratio is .37; epiphysis/tibia ratio is .50.
Number of specimens examined, 13.

COMADIA BERTHOLDI POLINGI Barnes & Benjamin,

new status
( figs. 9 ^ , 10 $ )

Comadia poJingi Barnes and Benjamin, 1927, Pan-Pacif. Ento-

mol., 4.67.

Male: Longest antennal pectination 2.5 to 3.0 times shaft
width; antennal pectination/eye diameter ratio is .36; epiphysis/
tibia ratio .48. Upper surface of forewing with reduced dark
brown suffusion, white scaling prominent; subcosta to near apex
without dark brown scales; costa dark brown, occasionally
slightly checkered; dark brown of distal end of cell reduced;
base of radial with dark brown streak about one fourth wing
length. Hindwing slightly darker than forewing; veins lined
dark. Under surface forewing with heavy brown suffusion except
discal cell, which is paler. Under surface hindwing concolorous
to upper surface. Female: Larger and with heavier brown suf-
fusion than male. Wing length in male, 15-17mm; in female,
18-19mm.

Range.— Arizona, California, Nevada, New Mexico, see map
#3. Flight period. May through August.

14(4);189-212, 1975

COMADIA REVISION

207

The white in polingi almost becomes a silver in some indi-
viduals. The brown scaling is always quite dark, but varies in
amount present, thus, some specimens have the discocellular
marking only on the veins. The over all darkness found in
hertholdi hertholdi is not present. The range for polingi is
around the southwestern edge of the Great Basin. Number of
specimens examined, 40.

COMADIA ALLENI Brown, new species

(fig. no

Male: Head and thorax suffused with white and black scales;
longest antennal pectination two times shaft width; antennal
pectination two times shaft width; antennal pectination/eye
diameter ratio is .43; epiphysis /tibia ratio is .5; antennal shaft
white dorsally to apex. Forewing ground color white; oblique
band of suffused fuscus from apex to end of discal cell; base
of Badial vein with dark bar; Cubital vein and discocellular
spot dark brown surrounded by ochre; middle of cell Ai with
spot of ochre and dark brown; veins outlined dark, checkered at
end; fringe checkered. Hindwing fuscus, veins outlined dark;
fringe white. Under side: fore wing fuscus except for white over
discal cell and subapically at costa. Hindwing lighter but fuscus
at apex; veins dark. Female paler and slightly larger than male,
almost immaculate. Wing length in Holotype 17mm; in allo-
type 18mm.

Holotype male, California, Stanislaus County, La Grange,
30 May 1960, R. P. Allen. Allotype, same locality and collector,
17 May 1968. Paratypes: same locality and collector as Holo-
type, 1^,11 May 1968; 1 $ , 2 June 1960. Holotype and allo-
type are deposited in the California Academy of Sciences, San
Francisco; paratypes are deposited in the California Department
of Agriculture, Sacramento. See distribution map #4.

This species is nearly as white as sperdtus but alleni has
heavier gray shading on the forewing in an area obliquely from
the apex through the discocellular spot and out to the fringe.
I am naming this species in honor of my good friend R. P.
Allen. Number of specimens examined, 4.

208

RICHARD M. BROWN

/. Res. Lepid.

14(4):189~212, 1975

COMADIA REVISION

209

COMADIA MANFREDI (Neumoegen)

(fig. 12^ )

Hypopta manfredi Neumoegen, 1884, Papilio, 3:139; Neumoegen
and Dyar, 1893, Journ. N. Y. Entomol. Soc., 1:33; ibid,
1894, Journ. N. Y. Entomol. Soc. 2:164; Dyar, 1937, in Seitz,
6:1275; Dyar, 1902, Bull U. S. N. M., 52:363.

Comadia manfredi; Barnes and McDunnough, 1911, Contrib.
Natur. Hist. Lepid. N. Amer., 1:28; ibid., 1917, Check List
of the Lepid. Boreal Amer., pg. 195.

Male: Head, thorax and abdomen covered with long cream
colored hair; longest antennal pectination three times shaft
width; antennal pectination /eye diameter ratio is .5; epiphysis/
tibia ratio is .45. Upper fore and hindwings cream colored and
devoid of all markings. Under forewing with a suffusion of dark
along costa and subcosta; remainder of wing and hindwing
concolorous with upper surface. Female unknown. Wing length
in male, 11mm.

Range. — Arizona, see map #4. Flight period. May.

This species is the smallest and palest of all the Comadia,
but has the longest antennal pectinations in the genus. From
henrici, manfredi can be easily separated by its much smaller
size, longer antennal pectination and shorter epiphysis. Speci-
mens examined, 1.

COMADIA REDTENBACHERI (Hammerschmidt),
new combination

(fig. 13 ^ )

Zeuzera redtenbacheri K. E. Hammerschmidt, 1848, in Natur-
ivissenschaftliche Abhandlungen, Gesammelt und Durch
subscription von Wilhelm Haidinger, 2:151-152.

Bombyx agavis I. Blasquez, 1870, La Naturaleza, 1:285-288.
Hypopta agavis; L, Ancona, 1930, Anales Del Institute De Bio-
logia de la Universidad Nacional Autonoma de Mexico,
1:265-277.

Hypopta chilodora H. Dyar, 1910, Proceed. Natur. Mus., 38:270.
Hypopta redtenbacheri; Dyar, 1937, in Seitz, Macrolepidoptera,
6:1275, PI. 181.

Male: Head, dorsal thorax, dorsal abdomen with tan and
dark brown scaling; ventrally less dark; longest antennal pecti-
nations 2.5-3.0 times shaft width; antennal pectination/eye
diameter ratio is .42; epiphysis /tibia ratio is .26. Upper fore-
wing, subcosta with narrow, very white strip from wing base
to just short of apex; general color brown; wing scales tan at

210

RICHARD M. BROWN

/. Res. Lepid.

base, dark brown at tip; two lines of raised scales cross wing;
lines whitish on inner edge, dark brown on outer edge, lines
formed by an alignment of these bicolored scales; inner line
bent inward below discal cell; outer line out curved, parallel
to fringe; terminal with a few scattered white scales; fringe
scales very long, longest white and spatulate at end; shorter
scales concolorous to wing, less spatulate. Hindwing uniform
gray-brown; fringe as in forewing. Under forewing costa dark
brown near base, mixed with white near apex; discal cell with
some long hair-like white scales; remainder of wing uniform
gray-brown. Hindwing as upper surface. Female: Generally
larger and marked similar to male; lines of raised scales less
contrasting and more numerous as short dashes; general ground
color paler than male, due to the lack of dark tiped scales.
Wing length in male, 12-14mm; in female 13-16mm.

Early stages.— The life history has been studied by Hammer-
schmidt (1848), Blasques (1870), and Ancona (1930). The
host is Agavis salmiana Otto, (Amerijllidaceae) (Ancona 1930).
The eggs are laid near the base of the leaves of the host and are
coffee colored at the time of emergence. Larvae are pale red-
dish in the first instar, becoming carmine in the later instars.
The last tergite has a group of five spines forming a horn. The
pupa has an array of spines that aid in digging to the ground
surface prior to eclosion. In the regions of Mexico where Agavis
salmiana is of economic importance, a 15% infestation of the
host has been found (Ancona, 1930).

Range. — In the Big Bend area of Texas and wide spread
within the boundaries of its host in Mexico. See map #4. Flight
period, April and May.

I have placed this moth in this genus on the grounds the
genitalia and wing veination are well within the generic limits.
Ancona (1930), in his treatise of the larvae, placed this species
in Hypopta and made Dyar’s (1910) chilodora a synonym of
agavis. Dyar (1937) placed both chilodora and agavis as syno-
nyms of redtenhacheri.

Comadia redtenhacheri is distinguished by the color, and the
lines of bicolored raised scales crossing the wings. Comadia
henrici and albistriga tend to have some of the dark scales at
the end of the discal cell bicolored, but since redtenhacheri does
not have this discocellular spot there should be no confusion
between species. Number of specimens examined, 16.

14(4):189-212, 1975

COMADIA REVISION

211

COMADIA ALBISTRIGA (Barnes & McDunnough),
new combination
(fig. Us )

Heterocoma albistriga Barnes and McDunnough, 1918, Contrib.
Natur. Hist. Lepid. N. Amer., 4(2): 179; McDunnough,
1939, Check List of Lepid. of Canada and U. S. Amer. Part
2, Microlepidoptera, pg. 62.

Male: Head and dorsal thorax creamy white, mixed with
black and brown scales; collar lightly marked with dark border;
ventral thorax without dark scales; antennal shaft dorsally
scaled, concoloroiis to thorax; longest antennal pectination three
times shaft width; antennal pectination/eye diameter ratio aver-
age .48; epiphysis /tibia ratio average .21. Abdomen with dorsal
anterior two thirds to three fourths mouse gray, remainder paler;
ventral abdomen concolorous to ventral thorax. Upper surface
of wings with fore wing costa with brown checks; subcosta
creamy-white without checks; discal area mouse brown; white
postmedial line formed by staggered bars between veins; distal
postmedial line paler; vein Ai broadly creamy-white to fringe;
discal spot large, creamy- white, extending to tornus; caudad A^
concolorous to submarginal area; fringe pale checked at vein
end; discocellular scaling dark brown. Hindwings uniformly
mouse brown; fringe creamy-white without checks. Under sur-
face of wings with forewing costa and fringe as above; remain-
der of wing mouse brown. Under hindwing marked as above.
Female: Nearly immaculate, brown shading found in male only
faintly visible. Wing length in male ll-14mm; in female 13mm.

Range. — Arizona, New Mexico, Texas, see map #4. Flight
period, April and May.

Barnes and McDunnough (1918) were in error when they
established the genus Heterocoma for albistriga, based mainly
on the lack of a cellula intrusa and the presence of strongly
stalked veins R and Mi. There is variation among the individuals

examined and albistriga is within the range of variation of
Comadia.

Comadia albistriga can easily be recognized by the white bar
crossing the wing from just below the costa through the brown
discocellular spot to the tornal area. This line, when connected
to the other three white lines, will form two triangles. Addition-
ally, the upper forewing has variable reticulations formed by an
alignment of dark brown tiped scales. These reticulations can
be found any place on the wing except for the pale subterminal
area.

212

RICHARD M. BROWN

/. Res. Lepid.

LITERATURE CITED

BROWN, R. M. and R. P. ALLEN, 1973, A New Comadia from the San
Joanquin (sic) Valley of California {Lepidoptera: Cossidae). Pan-
Pacif. EntomoL, 49:240-245.

DYAR, H., 1937, in Seitz, Macrolepidoptera, 6:1275, PI. 182.

ANCONA, L., 1930, Los Chilocuiles O Gusanitos De La Sal De Oaxaca.
Andes del Institute de Biologia, Universidad Nacional Autonoma de
Mexico, 1:265-277.

BARNES, W. and F. H. BENJAMIN, 1923, Cossidae, Cossinae, Contrih.
Natur. Hist. Lepidoptera, 5(2);88-96.

BARNES, W. and J. McDUNNOUGH, 1918, Cossidae, Contrih. Natur.
Hist. Lepidoptera, 4(2): 179.

DYAR, H., 1910, New Lepidoptera from Mexico, Cossidae, Proceedings of
the U. S. National Museum, 38:269-271.

HOLLAND, W. J., 1903, The Moth Book, Doubleday, Page and Co., pg.
379.

RIVERS, J. J., 1897, Some Facts in the Life History of Hypopta Bertholdi
Grote, Psyche, 8(249): 10.

BLASQUEZ, I., 1870, La N aturaleza, Periodico Cientifico De La Sociedad
Mexicana De Historia Natural, 1:285-288.

HAMMERSCHMIDT, K. E., 1848, Beschreibung Eines Neuen Mexicani-
schen Schmetterlinges. Naturwissenschaftliche Abhandlungen, Ge-
sammelt und Durch Subscription, Wien, 2:151-152.

Journal of Research on the Lepidoptera

14(4):213-215, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

A NOTE ON OENEIS MELISSA (FABRICIUS)

IN THE WESTERN UNITED STATES (SATYRIDAE)^
CLIFFORD D. FERRIS^

College of Engineering, University of Wyoming, Laramie, Wyoming 82071

Oeneis MELISSA (Fabricius) in the western United States has
generally been referred to the subspecies lucilla Barnes and
McDunnough in recent publications (Callaghan and Tidwell,
1971; Ferris 1971 a, b). The population in the Canadian Rocky
Mountains was named beani by Elwes in his 1893 revision with
Edwards of the worldwide Oeneis. The type locality is Laggan
(Lake Louise) in Banff National Park, Alberta. The type local-
ity for lucilla is Hall Valley, Park Co., Colorado.

Barnes and McDunnough, 1918, separated lucilla from the
other subspecies of melissa on the basis of the following key
characters: black palpi fringed with white dorsally, virtually nb
median (mesial) band on the ventral surface of the hind wings,
and the “y^l^^w” aspect of many specimens, especially in the
females. Elwes in describing beani noted the blackish-brown
aspect of the insect and its entirely black palpi.

Recent examination of specimens from the Rocky Mountains
in the United States has demonstrated that both subspecies occur
in this region. Material taken in Colorado, New Mexico and
Utah (Leidy Peak, Uintah Co.) is lucilla. In Wyoming, melissa
is found in the Wind River Range, Sublette Co. and on the
Beartooth Pleateau in Park Co. Specimens from these localities
are the typical black-palpi beani found in Alberta. This sub-
species has been recorded from Carbon Co. (Beartooth Plateau
region). Glacier National Park, Montana and from Okanogan
Co., Washington. To date, I have been unable to find any Idaho
records.

^Published with the approval of the Director, Wyoming Agricultural Ex-
periment Station as Journal Article no. JA 831.

^Research Associate, Allyn Museum of Entomology, Sarasota, Florida; Mu-
seum Associate, Los Angeles County Museum of Natural History, Los An-
geles, California.

213

CLIFFORD D. FERRIS L Res. Lepid.

Fig. 1. — Distribution of Oeneis melissa in the western United States with
partial records for southwestern Canada. Black dots = lucilla; black tri-
angles = beani.

14(4):213-215, 1975

OENEIS MELISSA

215

O. melissa heani frequents wind swept outcroppings of black
or black-lichen-covered rocks above treeline. They are swift
fliers and very wary. In the Beartooth Range, they occur at
11,000' and some 3000' lower on Plateau Mountain in southern
Alberta.

Figure 1 illustrates the distribution of melissa in the United
States. The Colorado county records include: Boulder, Chaffee,
Clear Creek, Costilla, Custer, Fremont, Gilpin, Grand, Gunni-
son, Hinsdale, Huerfano, Lake, La Plata, Larimer, Ouray, Park,
Pitkin, Rio Grande, Saguache, San Juan, San Miguel, Teller.
In New Mexico, it is found in Taos Co.

ACKNOWLEDGEMENTS

I would like to thank A. C. Sheppard of the Lyman Museum
at McGill University for questioning the identity of Wyoming
material, which prompted this study. R, E. Stanford, Denver,
Colorado kindly provided extensive Colorado records. L. D.
Miller, Allyn Museum of Entomology, Sarasota, Florida, E. M.
Perkins, Jr., University of Southern California, Los Angeles and
F. H. Rindge, American Museum of Natural History, New York
kindly provided additional records.

LITERATURE CITED

BARNES, W. H. and J. McDunnough. 1918. Contributions to the natural
history of the Lepidoptera of North America. 4(2):61~208,

CALLAGHAN, C. J. and K. B. TIDWELL. 1971 [1973]. A checklist of
Utah butterflies and skippers. /. Res. Lepid. 10(3) :191-202.

ELWES, H. J. and J. EDWARDS. 1893. A revision of the genus Oeneis.
Trans. Ent. Soc. Lond. (Royal Ent. Soc.) Part IV:457-481.

FERRIS, C. D. 1971a. A Key to the Rhopalocera [Butterflies] of Wyoming.
University of Wyoming, Agric. Exp. Sta. Sci. Mongr. 21. xv + 64 p.

— . 1971b. An Annotated Checklist of the Rhopalocera [Butterflies] of

Wyoming. University of Wyoming, Agric. Exp. Sta. Sci. Mongr. 23.
75 p.

Journal of Research on the Lepidoptera

14(4):216-232, 1975

1160 W. Orange Grove Ave., Arcadia, California 91006, U.S.A.
© Copyright 1975

POST-PLEISTOCENE ENVIRONMENTS AND
MONTANE BUTTERFLY RELICTS

ON THE WESTERN GREAT PLAINS

KURT JOHNSON' "

Unification Theological Center,

2628 Davidson Avenue, New York, NY 10468

For several reasons, a knowledge of the paleobotany of the
western Great Plains has become increasingly important in
understanding its butterfly fauna. Recent studies of many of its
regional faunas (Brown et al., 1957; Conway, 1966; Defoliart,
1956; Ferris, 1970a, 1970b, 1971; Johnson and Nixon, 1967; John-
son, 1972 (1973), 1975a; Nordin, 1967, 1968; Puckering and Post,
1960; and Scott, Ellis, and Eff, 1968) have placed a proper em-
phasis on the montane character of butterflies populating its
non-riparian scarp woodlands and remaining climax coniferous
forests. A rationale is needed to characterize these faunas in
relation to their own elements as well as other faunas in the
Nearctic (A. B. Klots and F. M. Brown, pers. com.). It is im-
portant to know whether they represent dispersal eastward from
the Rocky Mountains, or if they are relicts of a former, more
extensive, montane association in which unique allopatric pop-
ulations of some montane Nearctic butterfly species occur, rep-
resenting extreme divergence as peripheral populations. Simi-
larly, understanding current ecological changes effecting their
decline requires a knowledge of their probable origins (Johnson,
1976a).

Expanding distributional knowledge of taxa described from
these areas, research presented in taxonomic studies (Ferris,
1973; Johnson 1976b, 1976c; Johnson and Balogh, 1976; Perkins
and Perkins, 1967), and the recognition of important biological

^Museum Research Associate, Museum of Natural History, University of
Wisconsin, Stevens Point, 54481 .

'Research Associate, Allyn Museum of Entomology, Sarasota, Florida.

216

14(4):216-232, 1975

ENVIRONMENTS

217

distinctions in some populations (Klots, 1975; and pers. com.)
has indicated that many, if not most, of these scattered montane
faunas are more related to each other than to faunas in the
mountain ranges to their west. Thus, it is important to present
paleobotanical evidence establishing this former montane associ-
ation and to comment on its significance.

DISCUSSION

The major remaining conifer forest and montane areas of
the Great Plains occur from western Nebraska northward. They
include the Black Hills of South Dakota, the Pine Ridge of
Nebraska, the Cheyenne Ridge of Nebraska and Wyoming
(called the “three great pine relicts” by Clements, 1945) and
the more limited scarps along the Niobrara River in Nebraska
and the Little Missouri River in Montana and North Dakota.
Of these, only the Black Hills remains in a climax condition.

The history of these areas, and thus a major part of the
history of the Great Plains, is best understood by presenting a
vegetative chronology since the Wisconsin glaciation. This evi-
dence can then be used to explain the occurrence of the montane
elements in its butterfly fauna:

The late Miocene, Pliocene, and early Pleistocene glaciations:

In the late Miocene and Pliocene the entire area was open
woodland (savannah), with Cedrella (Meliaceae), and early
Celtis, Quercus, Mahonia, Piniis, Artemisia, Ambrosia, and many
Asteraceae. Montane areas, like the uplifted Black Hills, were
coniferous forest and remained so throughout the four glaciations
of the Pleistocene. The ice advances and interglacial episodes
caused floral fluctuations probably not unlike those of the Wis-
consin glaciation described below, which are most important in
understanding present faunas.

Source: Wells, 1970.

The developing Wisconsin (before 20,000 years ago):

Boreal forests proceeded in bands before the glaciers. Peri-
glacial winds formed a sand dune desert in central Nebraska
from sand blown in from the Ogallala formation of the northern
Great Plains. The Pine Ridge, Cheyenne Ridge, and other
present-day uplands were not exposed at this time, but were
in fact buried.

Sources: Schuchert, 1955; Smith, 1965; Wright, 1970.

218

KURT JOHNSON

J. Res. Lepid.

14(4):216-232, 1975

ENVIRONMENTS

219

The full-glacial Wisconsin (14-20,000 years ago)

Boreal forest extended west to northeastern Kansas and north-
ward across central Nebraska, but was prevented from reaching
the Black Hills by the sand dunes. Instead, it continued west-
ward across northwest South Dakota and north-central Wyo-
ming. Montane Pimis climax forest covered the uplands west-
ward. The present-day Pine Ridge, Cheyenne Ridge, and western
North Dakota areas had now been exposed as uplands by the
continual movement of loess eastward. Lowland pine-juniper
open woodland occurred concentrically around the upland
forests.

Sources: Wright, 1970; Wright and Ruhe, 1965.

The early glacial retreat (11-12,600 years ago):

Glacial retreat ended the periglacial winds. The sand dunes
gave way to boreal forest which proceeded west to the present
Black Hills. There, and in the Pine Ridge, Cheyenne Ridge,
western North Dakota association, Niobrara River area, and
associated lowland savannahs, it intermixed with montane
coniferous forests. Contributed were still-surviving Betula lenta
L., B. papijrifera Marsh., Populus tremuloides Michx., Quercus
macrocarpa Michx., Ostrya virginiana Willd., Ulmus americana
L. (pi. Clayt. ), Willd., and Corylus americana Walt. The Big-
horn Mountains were connected to the Black Hills by a moist
(compared to today) Pinus-Juniperus savannah.

Sources: Morgan, 1963; Nixon 1967; Wells, 1970; Wright, 1970;
Wright and Ruhe, 1965.

Mid-glacial retreat ( 10,000 years ago ) :

Further glacial retreat caused destruction of the boreal forest.
Relicts survived in the Black Hills, Pine Ridge, and Niobrara
escarpments. The extensive interplay of coniferous forest, open
woodland, and prairie began. Montane coniferous areas were
now nearly as shown in Figure 1.

Sources: Meintosh, 1961; Morgan, 1963; Nixon, 1967; Wells,
1970.

Post Wisconsin — the toarm postglacial interval (9,500 to 1700
years ago ) :

Vast Pinus climax forests were evident as indicated in Fig-
ure 1.

Pinus dominated pine-juniper open woodland covered the
present Sandhills of Nebraska and perhaps west-central North
and South Dakota. Moist open woodland, predominantly Juni-

220

KURT JOHNSON

/. Res. Lepid.

perus with some Finns, connected the Laramie Mountains and
Cheyenne Ridge, and the Bighorn Mountains and the Black
Hills. A trend toward aridity began at the end of this period.
Sources: Deevy and Flint, 1957; Wells, 1970.

The woodland to semi-desert transition ( 1700-200 years ago ) :

Continuous coniferous forest began to recced. Pine climax
retreated upland leaving subclimax relicts throughout various
escarpments, but a climax forest in the Black Hills. In the face
of increased aridity and prairie fires, open woodland began to
give way to prairie.

Source: Wells, 1970.

Demise of the conifers ( 200 years ago to the present ) ;

Continuous juniper and pine-dominated open woodlands
came to an end. Conifers became nearly extinct in most areas,
except remaining montane uplands. Mixed prairie and short-
grass prairie disclimax prevailed over most areas; pine relicts
remained limited to scarp woodland or the Black Hills.

Sources: Clements, 1945; Weaver and Albertson, 1956; Wells,
1965, 1970.

Figure 1 illustrates the probable maximum extent of former
coniferous forest and pine-juniper open woodland on the western
Great Plains in relation to existing nonriparian scarp woodlands.
Distributions of species and subspecies endemic to these conif-
erous forests biomes are shown in figures 2 and 3. Figures 4
through 6 illustrate ranges of other montane entities still in-
habiting the region, and probable relicts of two earlier fauna-
influencing periods.

Examination of these data in relation to the chronology of
paleobotanical events indicates evidence of five major elements
influencing the existing butterfly fauna and its affinities:

1. Relicts of the boreal invasion and its associated deciduous
flora (L. eurydice ssp., P. batesii, N. vau-album j-album).

2. Relicts of the extensive montane coniferous climax after
the recession of the glaciers (C. siva, C. meadii, C. alex-
andra krauthii, E. lucilius afranius, £. persius fredericki,
E. ausonides palaeoreios, E. a. bernadetta, E. amyntula
valeriae, G. piasus, H. pahaska pahaska, L. weidemeyerii
oberfoelli, L. rubidus longi, N. ridingsii, N. menapia, P.
phoebus dakotaensis, P. indra, P. muticaudatus, P. sisym-
brii nordini, P. icarioides lycea, P. shasta minnehaha, P.
satyrus, S. callippe, S. edwardsii, S. zerene).

14(4):216^232, 1975 ENVIRONMENTS

221

Hesperia pahaska

pahaska Leussler

Erynnis lucilius
afrani us°TSaidd¥r
and Burgess)

Erynnis persi us

fredericki H. A.
Freeman

Euchloe ausonides
pal aeoreios Johnson

ns sisy^ril colias alexandra

nordini Johnson cottpTEx

Fig. 2.- — Distributions of species (includes only Hesperia pahaska) and sub-
species ( others ) whose type localities are in one of the disjunct relict wood-
lands. Those clearly endemic are shown as such and usually represent mon-
tane coniferous .forest entities. Others with more expansive distributions are
usually montane grassland entities. Black lines on map indicate location of
existing scarp woodlands in the region.

222

KURT JOHNSON

/. Res. Lepid.

Limenitis weidemeyeri i
oberfoelli Brown

Lycaena rubidus
lonqi Johnson and
Balogh

Plebejus shasta
minnehaha" (Scudder )

Euphydryas anicia
bernadetta Leussler

Parnassi'us phoebus
dakotaensi s Eisner

Fij^. 3. — Same as Fig. 2.

14(4):216-232, 1975

Speyeria zerene
(Boisduval)

Speyeria callippe
meadii (Edwards)

ENVIRONMENTS

223

satyrus Speyeria edwardsii

(Edwards)

Speyeria atlantis

(Edwards)

Speyeria coronis

(Behr)

Fig. 4.— Distributions of other montane butterflies on the western Great
Plains in relation to existing scarp woodlands.

224

KURT JOHNSof^

/. Res. Lepid.

Papilio mul ti caudatus
Ki rby

Papilio ze1 icaon
Lucas

Papilio rutulus
Boisduval

Papi 1 io indra

Roaki rt

Papilio bairdii

brucoi Edwards

Fig. 5. — Distributions of other montane butterflies on the western Great
Plains in relation to existing scarp woodlands.

14(4):216-232, 1975

ENVIRONMENTS

Oeneis uhleri

f Reakirt]

Cercyonis meadi i

(Edwards )

225

Plebejus icarioiaes
lycea (Edwards)

Cercyonis oetus
oetus (Boisduval)

RELICTS OF FORMER EASTERN
DECIDUOUS FOREST and NORTH-
WARD MONTANE INVASIONS

Phyciodes batesii (Reakirt)
Nymphalis vau-album j- album
(Boisduval & LeConteJ
Lethe eurydice (Johannsen)

Asterocampa celtis an ton i a

(Edwards f

Fig. 6. — Distributions of other montane butterflies on the western Great
Plains in relation to existing scarp woodlands. Bottom right: listing of relict
deciduous forest entities found in the Pine Ridge and representing extreme
allopatry from their normal range.

226

KURT JOHNSON

/. Res. Lepid.

3. Relicts of the northward dispersal of western montane
entities (those above [2] but obviously A. c. antonia).

4. Relicts of montane entities once dispersed throughout
the vast open woodlands (C. siva, C. alexandra ssp., L.
eurydice ssp., L. rudibiis longi, P. multicaudatus, P. shasta
minnehaha, S. edwardsii) .

5. The pattern of retreat of the coniferous forest at the be-
ginning of the current aridity and fire-influenced period
(all of the above, determining existing ranges).

These relationships show patterns important in interpreting
the origin and faunal character of the insects of the region. All
entities with habitats characteristic of montane coniferous for-
est, and with type localities within one of the now disjunct pine
relicts, show a north-south distributional relationship within the
former coniferous forest belt. Some are still found eastward along
the Niobrara River where some limited scarp woodlands still
support montane butterflies. Almost conversely, such entities,
whose present distributions span the east-west arid basins, are
grassland-related species, probably indicating the role of the
open woodland in maintaining their ranges during the demise
of the conifers, when montane forest entities retreated upland.
Vast open woodland formerly in the Nebraska Sandhills, south
of coniferous forest now isolated along the Niobrara River,
figures as the origin of montane-related species which still occur
there, in isolated and often unique environs like the post-climax
prairie (Pool, 1914; Tolstead, 1941). In no case does a taxonomi-
cally distinct entity described from a montane conifer niche in
the region show its primary relationship westward to the Rocky
Mountains.

The most limited relict, small scarps which remain of the
Cheyenne Ridge, contains montane representatives mostly with
a northward relationship. However, some of its plains-adapted
montane species evidence distinct interaction with populations
clearly dispersed eastward from the Rocky Mountains. In strong-
ly polymorphic Colias alexandra Edwards, evidence from Ferris
(1973) and A. R. Klots (pers. com.) indicates the unique bi-
voltine and “orange” population called C. a. krauthii in the
Rlack Hills does not extend southward and include the Cheyenne
Ridge, Instead, a “yellow” population ( of unclear voltinism )
occurs here which seems more related to the “yellow” uni-
voltine populations (called C. a. alexandra) on the plains sur-

14(4):216-232, 1975

ENVIRONMENTS

227

rounding the Front Range in Colorado. Similarly, Lijcaena rii-
bidus Behr occurring on the plains south of the Pine Ridge
shows purely northward related populations (L. r. longi) only in
the pine=forested canyons of the region. Remaining populations
on the plains, or in the limited remnants of the Cheyenne Ridge
in Wyoming, show evidence of secondary intergradation with
L. r. rubidus of the Rocky Mountains which has established
many plains populations in eastern Colorado and Wyoming. The
evidence of secondary intergradation in plains populations south
of the Pine Ridge is shown in the geographic affinities of geni-
talic and wing pattern traits illustrated in Figure 7.

The great influence of the former extent of coniferous forest
in the region can also be illustrated by the ranges and affinities
of other montane species occurring on the Great Plains, but
which have apparently not undergone extreme speciation (or
perhaps simply have not been studied). Their distributions are
shown in Figures 4-6.

Faunal resemblance formulae (Long, 1963) were used to
compare the entire faunas of each relict area in this study and
the mountain ranges in Colorado and Wyoming (Johnson, 1968).
Results of such comparisons agree with the general affinities
indicated by the ranges of individual taxa (The Pine Ridge
relates to the Black Hills first, the Bighorn Mountains second;
the Black Hills relates to the Big Horn Mountains first; the
Cheyenne Ridge relates to the Pine Ridge first) but were not
deemed as useful in illustrating real relationships. Their values
are greatly prejudiced by the unique admixtures of biota in these
areas caused by the incursion of prairie biomes into the scarp
woodlands, the occurrence of alpine environs only in the Black
Hills and Bighorn Mountains, and the infusion of southern-
related species into the river drainages of Colorado and western
Nebraska.

Apparent relict populations of butterflies representing earlier
vegetative events in the region were also noted in the study
areas and further attest to these uniquely vegetated scarp wood-
lands providing refugia for species not characteristic of the sur-
rounding plains. In the Pine Ridge, a curious Pohjgonia “hijlas”
(Edwards) [the name htjias is presently of uncertain relation-
ship with more northern faunus (Edwards) but has been tradi-
tionally used for Rocky Mountain populations of the complex]
or extremely green P, progne ( Cramer ) occurs in the same areas
as birch relicts. Birch ( Betula sp. ) is the northern larval food-

Montana

228

KURT JOHNSON

/. Res. Lepid.

14(4):216-232, 1975

ENVIRONMENTS

229

plant of the P. faunus- P. hylas complex.

Also, three deciduous forest entities {Phyciodes batesii and
Nymphalis vau-album j-album of eastern origin, and Astero-
campa celtis antonia of southern origin) occur in the river bot-
toms of the Pine Ridge canyons. These are relicts of the west-
ward deciduous movement and the northward invasion of
southern montane entities respectively. Most interestingly, plains
river bottom populations of A. celtis represent the nominate
subspecies celtis (Boisduval and LeConte) which is of eastern
affinity. The uniqueness of the canyon populations was con-
firmed by studying and rearing the larval stages [Johnson 1972
(1973)]. Also of importance is the occurrence of an extremely
allopatric and disjunctly distributed subspecies of eastern Lethe
eurydice Johannsen along the streams in the deep escarpment
canyons. This population, isolated from the formerly most wes-
tern population of the species, L. e. fumosa Leussler, is clearly
a relict of the former westward extension of the eastern de-
ciduous forest. Unfortunately, many areas in the region highly
suited for such relicts have been ill-studied by lepidopterists
and our knowledge of their occurrence is limited to only those
areas which have received intensive local sampling. Thus, in-
dividuals of the above species ( and others ) may also occur
northward in the present-day disjunct conifer woodlands, as do
many of the montane species mentioned earlier.

CONCLUSIONS

These data support three conclusions important to the history
of butterfly speciation in the central and western Nearctic
Realm.

1. The montane faunas of the remaining coniferous forests on
the western Great Plains represent relicts of former, more ex-
tensive populations (not merely eastward dispersal from the
mountains) and have a disjunct relationship to each other as
opposed to the mountains to their west. 2. These same areas
support allopatric populations of butterfly species from eastern
and southern origins which are relicts of other vegetative in-
vasions of the region. 3. This ancient montane and coniferous
forest area should be considered as having formed a distinct
area of speciation in the evolution of the present butterfly faunas
of the Rocky Mountains and to a lesser extent those of eastern
deciduous forest areas of North America.

230

KURT JOHNSON

/. Res. Lepid.

ACKNOWLEDGMENTS

I am grateful to Drs. Alexander B. Klots and F. Martin
Brown (Ameriean Museum of Natural History) for originally
diseussing the need for this researeh with me, and to Dr. Klots
for reviewing the manuseript. Particular debt is owed Dr.
Frederick H. Rindge (Ameriean Museum of Natural History)
for use of facilities and advice in investigating taxonomic prob-
lems. Thanks are due Mr. Erie Quinter (American Museum of
Natural History) for critique, Dr. Lee D. Miller (Allyn Museum
of Entomology, Sarasota, Florida) for permission to reproduce
from a paper in press there, and Dr. Charles A. Long ( Museum
of Natural History, University of Wisconsin, Stevens Point) for
the museum’s support of this research.

SPECIAL NOTES CONCERNING FORMAT
IN THIS PAPER

I have included the authors’ names for each taxon on the
maps in the figures.

This seemed more ideal than citing them the first time they
were used in the text I. because of the number of names in
the paper 2. the fact that some are mentioned in both species or
subspecies contexts at times 3. the variance in degree of study
in some of these groups taxonomically.

Thus, the only authors’ names mentioned in the text are in use
for the first time at variance with those in the figures (e.g. when
a species is discussed, but a specific trinomen is cited in the
figures). Therefore, the taxonomic names in the figures form a
consistent usage for the region, dependent on the degree of study
each group has received by entomologists.

LITERATURE CITED

BROWN, F. M., D. EFF and B. ROTGER. 1957. Colorado Butterflies.

Denver Mus. Natiir. Hist., Denver, 368 pp.

CLEMENTS, F. E. 1949. The Dynamics of Vegetation. H. W. Wilson, New
York, xxiii + 296 pp.

CONWAY, P. J. 1966. A quick collecting trip to the Black Hills. Newsl.
Assoc. Minn. Entom. 1(1): 16-19.

DEEVY, E. S. and R. F. FLINT. 1957. Postglacial hypsithernial interval.
Science, 1.25: 182-184.

DEFOLIART, G. R. 1956. An annotated list of southeastern Wyoming
Rhopalocera. Lepid. News 10 (3-4): 165-175.

EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus
Cercijonis (Satyridae). I. Characteristics of the genus. /. Lepid. Soc.
23 (3): 165-175.

14(4):216-232, 1975

ENVIRONMENTS

231

FERRIS, C. D. 1970a. A checklist of the Rhopalocera of eastern Wyoming.
Mid-Contin. Lepid. Ser. I (10): 1-11.

. 1970b, Collecting spring diurnal Lepidoptera in southeastern Wyo-
ming. Mid-Contin. Lepid. Ser. 1 (12): 6-10.

. 1971. An annotated checklist of the Rhopalocera of Wyoming.

Agric. Exper. Stat., Univ. Wtjo. Sci. Monog. 21, 64 pp.

. 1973. A revision of the Colkis alexandra complex (Pieridae) aided

by ultraviolet reflectance photography with designation of a new sub-
species. /. Lepid. Soc. 27 ( 1 ) : 57-73.

JOHNSON, K., and E. S. NIXON. 1967. The Rhopalocera of northwestern
Nebraska. Anier. Mid. Nat., 78 (2): 508-528,

JOHNSON, K. 1972 (1973). The butterflies of Nebraska. J. Res. Lepid.
11 (1): 1-64.

. 1976a. Prairie and plains disclimax and disappearing butterflies.

Atal.a, Memoir No. 1, Xerces Society (in press).

. (ms. 1968) The faunal resemblances of butterflies of the western

Great Plains, southern and central Rocky Mountains. Project in Quan-
titative Biology, Univ. Wise., Stevens Point.

. 1976b. Concerning the name Anthocaris coloradensis Hy. Edwards

with designation of a new subspecies (Pieridae). /. Lepid. Soc. 30 (4):
in press.

. 1976c. A new subspecies of Pieris sisymhrii from western Great

Plains relict forests (Pieridae). ]. Lepid. Soc. 30 (3): in press.
JOHNSON, K., and G. BALOGH. 1976. A revision and taxonomic analysis

of the nearctic Lycaena ruhidus complex. Bull. Allyn Mus. Entomol.
( in press ) .

KLOTS, A. B. 1975. in HOWE, W. (coordinating ed.) The Butterflies of
North America. Doubleday and Company, Inc., Garden City, xii, 633
p., 97 pis.

LONG, C. A. 1963. Mathematical formulas expressing faunal resemblance.

Trans. Kans. Acad. Sci. 66 (1): 138-140.

McIntosh, a. C. 1931. a botanical survev of the Black Hills of South
Dakota. Black Plills Eng. 19:159-276.

MORGAN, G. 1963. Relict populations of Betula lenta L. in northwestern
Nebraska. Unpublished M.A. thesis, Univ. Nebr., Omaha.

NIXON, E. S. 1967. A vegetational study of the Pine Ridge of northwestern
Nebraska. Southwest. Nat. 12 (2): 134-145.

NORDIN, J. S. 1967. Hesperiidae records for South Dakota, Newsl. Assoc.
Minn. Entom. 1 (4): 90-92.

. 1968. 1966 Lycaenidae records for South Dakota. Bull. Assoc.

Minn. Entom. 2 (3): 60-61.

PERKINS, S. F., and E. M. PERKINS. 1967. Revision of the Limenitis
iveidemeijerii complex with description of a new subspecies ( Nym-
phalidae). J. Lepid. Soc. 21 (4): 213-234.

POOL, R. J. 1914. A study of the vegetation of the sandhills of Nebraska.
Minn. Bot. Studies 4:189-312.

PUCKERING, D. L., and R. L. POST. 1960. Butterflies of North Dakota
(North Dakota Insects), 1:32 pp. North Dakota State University,
Fargo.

SCHUCHERT, C. 1955. Atlas of paleographic maps of North America.

John Wiley and Sons, Inc. New York. 177 pp.

SCOTT, J. A., S. L. ELLIS, and J. D. EFF. 1968. New records, range ex-
tensions, and field data for Colorado butterflies and skippers. J. Lepid.
Soc. 22 (3): 159-171.

SMITH, H. T. U. 1965. Dune morphology and chronology in central and
western Nebraska. Ecol. Monog. 12:255-292.

TOLSTEAD, W. L. 1941. Vegetation of the northern part of Cherry Coun-
ty, Nebra.ska. Jour. Geol. 73:557-578.

232

KURT JOHNSON

J. Res. Lepid.

WEAVER, J. E., and F. W. ALBERTSON. 1956. Grasslands of the Great
Plains. Johnsen Piibl. Comp., Lincoln, Nebraska.

WELLS, P. V. 1970a. Vegetational history of the Great Plains: a post-
glacial record of conireroiis woodland in southeastern Wyoming, in:
Pleistoeene and recent environments of the central Great Plains ( Dort,
W. D., and J. K. Jones, Edts.). Dept, of Geol., Univ. Kansas, Spec.
Puhl. 3:185-202.

. 1970b. Historical factors controlling vegetation patterns in the cen-

ral plains region of North America, in: Pleistocene and recent environ-
ments of the central Great Plains (Dort, W. D., and J. K. Jones, Edts.).
Dept, of Geol., Univ. Kansas, Spec. Puhl. 3:211-221.

WRIGHT, H. E., JR. 1970. Vegetational history of the eentral plains, in:
Pleistocene and recent environments of the central Great Plains ( Dort,
W. D., and J. K. Jones, Edts.). Dept, of Geol., Univ. Kansas, Spec.
Puhl. 3:157-172.

WRIGHT, H. E., JR., and R. V. RUHE. 1965. The Quaternary of the
United States: Princeton Univ. Press. 922 pp.

14(4):233-252, 1975

Journal of Research on the Lepidoptera

( Continued from 4( 3 ) : 188 )

9. Deciduous Woodland (Quercus consociation); Plate 13.

This formation attains its greatest development on the ridges

on the southern slopes of Volcan Santa Marta between eleva-
tions of 400 and 1,500 feet, i.e., above the Savanna. However,
small, disjunct stands are found above the P inns -Quercus
Associes and on the southern slopes of Volcan San Martin
Pajapan at elevations between 2,300 and 2,500 feet.

Ground cover is of variable density and consists of short
grasses and sedges of which the most common are Eragrostis
sp., Rynchospora globosa, Paspalum pectinatum, Paspaliim
plicatulum, and Sporoboltis cubensis. Oxalis neaei is also com-
mon. Dense stands of Calliandra grandiflora and Conostegia
xalapensis are frequent.

Quercus peduncidaris, which attains a maximum height of
30 to 40 feet is the most common species of tree although
Quercus oleoides and Quercus ghiesbreghtii are common. Other
trees include Byrsonima crassifolia, Miconia argentea, and
Acalypha unibract eata.

Towards the end of the dry season (mid-May), most of the
oaks drop their leaves and blossom. New growth appears shortly
thereafter so that the trees are not leafless for more than one
or two weeks.

Epiphytes are fairly common in the taller trees, especially
at elevations above 2,000 feet.

This formation appears to be a phase of the Orchard Savanna
of Beard (1953) in which species of Quercus predominate. As
stated previously, the oak forest exists principally on ridges
whereas the Savanna is found on less coarse topography. There-
fore, the soils within the oak forest ( and which are thin, sandy,
and range in color from grey to yellow) probably are more
heavily leached and more efficiently drained than those in the
Savanna. These two edaphic factors possibly are responsible for
the existence of the Deciduous Woodland, which might be
termed a modified Orchard Savanna.

10. Pinus-Quercus Associes; Plate 14.

The pine-oak forest is restricted to a relatively small area
on the southern slopes of Volcan Santa Marta. The forest occurs
on the upper slopes and crests of many of the numerous ridges
between elevations of 1,600 and 3,000 feet.

Ground vegetation is of variable density and consists pri-
marily of numerous grasses and sedges of which the most com-
mon are Paspalum pectinatum, Paspahnn plicatulum, Eragrostis

233

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GARY N. ROSS

J. Res. Lepid.

o

u

o

^ C5

G

44

14(4):233-252, 1975 BUTTERFLIES OF TUXTLA

235

sp., Sporobolus cubensis, Rtjnchospora globosa, Bulbostijlis pa-
pillosa, and Dichromena ciliata. Other plants include Croton
repens, Borreria siiaveolens, Crnsea calcocephala, Calliandra
grandiflora, Eupatorium incomptum, Calea zacatechichi, Rtiellia
fluviatilis, Stevia rhombifolia, Cordia spinescens, Lemourouxia
viscosa, Conyza chilensis, Escobedia laevis, Turnera ulmifolia,
Calea longipedicellata. Cassia hispidida, Vernonia leiocarpa,
Salvia shannonii, Calea cacosmoides (?), and Vernonia argyro-
pappa. The cycad Zamia loddigesii is locally common.

Finns oocarpa is the dominant tree species although Quercus
conspersa and Quercus ghiesbrechtii occur rather commonly
throughout the formation. Trees never form dense thickets or
forests but are relatively widely distributed in open stands.

The pine-oak community represents one of the most interest-
ing curiosities and enigmas of the Sierra de Tuxtla for the for-
mation has such a restricted distribution. The nearest com-
parable formation occurs in the Department of Tuxtepec,
Oaxaca, 90 miles away. The Sierra’s Pinus Quercus community
appears to be a subclimax community, which probably cor-
responds to the “Pine Savanna” of Beard (1953) and which is
considered by that author to be a phase of the “Orchard Savan-
na.” Beard states that pines may and usually do invade savannas
when a seed source is near. However, in the Sierra pines do not
occur in either the typical Savanna or the Deciduous Woodland
(modified savanna), formations that exist in areas more proximal
to the pine seed source in Oaxaca. Hence, additional factors
probably are responsible for the limited extent of the pine-oak
community. Two suggestions are offered. First, the community
exists only on the crests of steep ridges, areas in which soils
are extremely thin and lateritic ( Friedlaender, 1923) and hence
relatively poor in nutrients. Furthermore, most of the Sierra has
been settled by Indians — relatively recent!}^ by Popolucas and
Aztecs and formerly by Olmecs — for over 2,000 years (see Boss,
1966). Today most of the Indian villages on the Santa Marta
massif are restricted to ridge crests and information gathered
from Popolucan legends and stories indicates that this pattern
of settlement is an ancient one. Thus the ridge crests have been
subjected to severe soil disturbances for at least hundreds of
years. Numerous works, e.g., those of Pessin (1937), Stoate
(1950), and Merrifield, Foil, and Hansbrough (1964), have
shown that pines can grow well in soils with relatively low
concentrations of nutrients and which are too poor for many

45

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GARY N. ROSS

/. Res. Lepid.

i

46

PLATE 15

Littoral Woodland near Zapoapan; view is towards the southeast. May 1965,
sea level.

14(4):233-252, 1975

BUTTERFLIES OF TUXTLA

237

plant species. Therefore, I conclude that the ridge crests repre-
sent a favorable habitat for pines and a relatively unfavorable
one for many (or even most) other species of trees. Second,
the Popolucas today engage (and have engaged for hundreds
of years) in annual burnings of the pine lands. These burnings
significantly increase the geographic extent of the pine-oak
community for whenever fire is excluded from an area for several
consecutive years a heavy ground cover consisting of numerous
grasses, shrubs, and oak saplings develops. These saplings
develop later into trees; pine seedlings usually are never present
(Ross, 1966). Therefore, I conclude that the pine-oak community
in the Santa Marta area is a serai stage within the Deciduous
Woodland (which in turn is a modified savanna) and which has
become established because of favorable edaphic conditions and
is being maintained in its present extent by man-caused fires.

The ravines within the Pinus-Quercus Associes contain ele-
ments of the Semi-Evergreen Seasonal Forest.

DRY EVERGREEN FORMATION

11. Littoral Woodland or Dry Evergreen Woodland {Ficus-
Hibiscus Association); Plate 15.

This formation occurs along the seashore between the high
water mark and the volcanic headlands fronting the Gulf. Hence
the formation is variable in width, extending inland for only a
few feet or to 500 to 1,000 feet. Strong winds and salt spray
are common throughout.

Along the sandy beaches the following plant species are
common: Cypenis ligtdaris, Cijperus articulatus, Cenchrus in-
certus, Distichlis spicata or Sporoboltis virginicus, Chloris pe-
traea, Vigna luteola, Ipomoea stolonifera, and Ipomoea pes-
caprae.

Farther inland are small, gnarled, and windswept trees and
shrubs, which are found both in small open patches and in
extensive dense thickets. These woody species include Ficus
spp., Hibiscus titiaceous, Pachira aquatica, and Piper cordovan.

SWAMP FORMATIONS

12. Swamp Forest (Pachira-Ficus Association); Plate 16.

The Swamp Forest is located at low elevations near the
Gulf and bordering several of the large streams, e.g., Rio Zapoa-

47

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GARY N. ROSS

J. Res. Lepid.

48

PLATE 16

Swamp Forest near Zapoapan. May 1965, sea level.

14(4):233~252, 1975

BUTTERFLIES OF TUXTLA

239

pan, Rio Carizal, Rio Salado, Rio Yougualta japan, and
Rio Maquina. The ground in this formation is waterlogged
during most of the year and is frequently inundated for long
periods during the rainy season.

The understory is relatively open and is composed princi-
pally of Chamaedorea tepejilote, Piper cordovan, and numerous
saplings.

The single tree stratum ranges between 40 and 60 feet in
height and is composed principally of Pachira aqiiatica, Ficus
spp. (including Ficus obtiisifolia) , Hibiscus tiliaceous, and
Pleuranthodendron mexicana.

Many trees exhibit buttressing and stilting. Lianas and epi-
phytes are common.

13. Mangrove Woodland (Rhizophora consociation); Plate 17.
This formation is restricted to the margins of Bahia Sonte-

comapan. Rhizophora mangle is the only principal species and
forms thick, closed stands up to 40 or 60 feet in height. Stilt
roots and pneumatophores are common.

UNRESTRICTED MISCELLANEOUS FORMATIONS

14. Recently Abandoned Milpas; Plate 18.

When a milpa or corn field is abandoned the field is invaded
by numerous grasses and annuals. The latter include Melam-
poditim divaricatum and Ageratum comjzoides, both of which
blossom in the spring and which are replaced in the summer by
Baltimora recta, Bidens pilosa var. bimucronata, and Melam-
podiiim kunthianum. In subsequent years these species usually
are replaced by more woody plants such as Polymnia maculata,
Cordia spinescens, H amelia patens. Piper auritum, Conostegia
xalapensis (Andrle, 1964), Heliotropium indicum, Calliandra
grandiflora, and Vernonia leiocarpa.

15. Pastures; Plate 19.

Areas that are used consistenly for the grazing of cattle
and/ or horses usually develop rather distinctive floristic charac-
teristics. There are numerous patches or clumps of relatively
short vegetation composed of Cordia alliodora, Croton soliman,
Picramnia andicola, Heliotropium indicum, Crotalaria vitellina,
Urera elata, Solanum ochraceo-ferrugineum, and Piper auritum.
Trees, which are relatively few in number, include Inga spuria,
Inga leptoloba, Cassia spectabilis. Cassia occidentalis, Annona
reticulata, Annona muricata, Trema micrantha, and Pleurantho-
dendron mexicana.

49

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GARY N. ROSS

/. Res. Lepid.

50

PLATE 17

Mangrove Woodland along Bahia Sontecomapan. May 1962, sea level.
Photograph by R. F. Andrle.

14(4): 233-25 2, 1975

BUTTERFLIES OF TUXTLA

241

16. Hedgerows; Plate 20.

Com fields and pastures in the more heavily cultivated sec-
tions of the Sierra, e.g., in the vicinity of the larger cities and
towns, usually are bordered by fences about which is found a
varied but characteristic assemblage of plants. Where best
developed these rows may be as wide as 15 to 20 feet. The
undergrowth usually is a tangle of vegetation composed of
Cordia spinescens, Ur era elata, Passiflora serratifolia, and PassP
flora coriacea. Tree species attain maximum heights of 20 to 30
feet and include Bursera simaniha, Glirtcidia sepium, Enjthrina
americana (Andrle, 1964), Zanthoxijlum elephantiasis, Ficm
padifolia, Inga spuria, Inga leptoloba. Cassia spectabilis, Annona
muricata, and Annona reticulata.

V. ACCOUNTS OF BUTTERFLY SPECIES IN
THE SIERRA DE TUXTLA
Plan of the Species Accounts

Unfortunately, systematics of the Lepidoptera, especially
tropical forms, is in a relatively unstable state. Indeed, between
one-third and one-half of the species listed in the “Catalogo
Sistematico y Zoogeografico de los Lepidopteros Mexicanos”
(Hoffmann, 1940) do not bear today the same generic and
specific names. For this reason I have had to rely on numerous
and isolated taxonomic works in addition to personal corres-
pondence in order to present here a reasonably accurate and
modern systematic arrangement of taxons. In particular, I have
followed Munroe and Ehrlich ( 1960 ) for the arrangement of
the Papilionidae, Klots (1931) for the Pieridae, Fox ^1956) for
the Ithomiidae, Micheiier (1942) and Emsley (1963) for the
Heliconiinae ( Nymphalidae), Forbes (1944) and Higgins
(1960) for the Argynididi (Nymphalidae), Chermock (1950) for
the Liminitidi ( Nymphalidae ) , and Clench ( 1955 ) for the
Lycaenidae and Riodinidae. Thus, it is hoped that this paper,
although not intended to be taxoiiomically oriented, nevertheless
will represent a significant contribution to the systematic litera-
ture and serve as a modern reference for students of Neotropical
Lepidoptera.

Each species account is introduced by the scientific name of
the species. Following this heading and under the caption

51

242

GARY N. ROSS

J. Res. Lepid.

52

PLATE 18

Recently Abandoned Milpa on Volcan San Martin Tiixtla. Field has re-
mained fallow for approximately one year. June 1962, 2,350 feet. Photo-
graph by R. F. Andrle.

14(4);233-252, 1975

BUTTERFLIES OF TUXTLA

243

SPECIMEN or SPECIMENS are the field data. For those species
that previously have been recorded from the Sierra, the data
have been condensed and include number of specimens of each
sex, maximum-minimum altitudes (elevations were measured
with a Taylor “Forecaster-Altimeter” calibrated to measure in
200 foot intervals), and earliest-latest collection dates. For those
species that represent new records for the Sierra or which are
rare, the complete field data are given (mileages represent
straight-line distances ) . If a particular specimen is in a collec-
tion other than my own, initials of the collection are included.
The following initials are employed :KHW = personal collection
of Kent H. Wilson and LSUMZ = Louisiana State University
Museum of Zoology. Sequence of entries is determined primarily
by altitude (lowest elevation listed first) and secondarily by
collection date (the earliest day and month listed first). Thus,
the first and last entries give the altitudinal range of the species.

Following the data is a discussion paragraph. In the first
sentence I express the relative abundance of the species in the
Sierra by employing four general terms — abundant, common,
uncommon, and rare. I consider a species to be abundant if ten
or more individuals of it were noted every (or almost every)
day, common when less than ten individuals were noted each
day, and uncommon when only three to five individuals were
seen at fairly wide intervals of time. Those species collected
only after very long time lapses or only once or twice during
my residence are designated as rare. Unfortunately, these terms
are extremely difficult to standardize when referring to animals
in the tropics because of the overwhelming array of inconspicu-
ous and virtually uncollectible habitats. Therefore, in many
cases the evaluation of the relative abundance of a species may
be very biased and not reflect the actual status of the species.
In the second sentence (and frequently in the latter part of the
first ) I give the principal plant formation or formations inhabited
by the species. Sequence of formations is based on decreasing
relative abundance of the butterfly species. The remaining sen-
tences are devoted to general comments, principally on the
ecology and ethology of the species, that I consider pertinent.
When a species previously has not been recorded from the
Sierra, the nearest recorded locale is given in the last sentence
of the paragraph.

53

244

GARY N. ROSS

J. Res. Lepid.

PLATE 19

f\'istiirf near Zapoapan. May 1965, sea level.

54

14(4) -.233^252,1975 BUTTERFLIES OF TUXTLA

245

Accounts

FAMILY PAPILIONIDAE
SUBFAMILY Papilioninae

TRIBE Graphiini
SUBTRIBE Graphiiti

1. Graphiiim phaon (Boisduval)

SPECIMENS: ms S, 19; 1,1004,800 feet; 12 May^3 Oct
This species is common along hedgerows, particularly in the
vicinity of Lago Catemaca. Most butterflies were collected as
they imbibed moisture from wet sand and soil. The flight is
relatively rapid, erratic, and usually between two and four feet
of the ground. Two of the 11 specimens collected exhibit a loss
of marginal and submarginal greenish dots on the dorsal fore
wings and a replacement of the postmedian-median row of
greenish scales on the dorsal hind wings with red scales. This
morphotype, which has been named form eridamas (Reakirt),
bears a close resemblance to Parides polyzelus (Papilioninae:
Troidini), a species that is abundant along hedgerows and the
margins of forests.

2. Graphium branchus (Doiibleday)

SPECIMENS: 3 ^ 5 ; 2 mi. NE Catemaco, 1,100 feet, 27
July 1962, 1 ^ ; 3 Aug. 1962, 1 S : Ocotal Grande, 1,800 feet,
15 May 1965, 1 ^ .

This tailless Graphium is uncommon and was collected along
the margins of the Semi-Evergreen Seasonal Forest only in late
spring and summer. The flight is relatively close to the ground
(usually within three or four feet) and similar in velocity to
that of Parides spp. This species was recorded previously from
Veracruz only from the ‘'Sierra Madre Oriental” (Hoffmann,
1940).

3. Graphium- helesis (Bates)

SPECIMENS: 18 ^ , 6$ $; 1.5 mi. SSE Sontecomapan,

900 feet, 14 July 1962, 1 $ : 2 mi. NE Catemaco, 1,100 feet, 1
July 1963, is ; 2 July 1962, 1 $ ; 9 July 1962, 1 $ (KHW); 27
July 1962, 2$ S (1 KHW) ; 9 Aug. 1962, 1 ^ ; 31 Aug. 1962, 1 ^ ;

55

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GARY N. ROSS

J. Res. Lepid.

56

14(4):233-252, 1975

BUTTERFLIES OF TUXTLA

247

8 Sept. 1962, 1 $ : 4.5 mi. NE Catemaco, 1,100 feet, 26 June 1962,
1^:4 mi. S Coyame, 1,200 feet, 25 June 1962, 3 S S : 2.5 mi.
SW Sontecomapan, 1,300 feet, 26 June 1962, 1 ^:1 mi. N So-
teapan, 1,400 feet, 28 March 1965, 1 ^ ; 29 March 1965, 2s S ■
1 mi. SSW Vigia, 1,800 feet, 18 March 1965, 1 9 : 1 mi. NNE
Ocotal Chico, 2,000 feet, 1 $ , 1 $ : 2 mi. NNW Ocotal Chico,
3,800 feet, 1 $ . Reared specimens: 2 mi. NE Catemaco, 1,100
feet, emerged 30 July 1962, 1 $ (KHW); emerged 3 Sept. 1962,
1 $ ; emerged 6 Sept. 1962, 1 $ .

This species is abundant during the spring and summer
months along the margins of the Semi-Evergreen Seasonal Forest
and Hedgerows, particularly in the vicinity of Lago Catemaco.
Adults are attracted to wet sand and soil. The flight is usually
slow and within three feet of the ground, a behavior that is
atypical for most members of the genus. Of the 24 specimens
collected, two have faint white blotches on their dorsal fore
wings. This morphotype, named form hephaestion (Felder),
bears a close resemblance to females of Parides spp. Larvae
(described in Ross, 1964b) were found occasionally on Annona
miiricata, a tree called “Cuanabana” by the local residents and
commonly planted around habitations. G. helesis was recorded
previously from Veracruz only from the “Sierra Madre Oriental”
(Hoffmann, 1940).

4. Graphiiim philolaus (Boisduval)

SPECIMENS: 3 ^ ^ , 1 $ ; 1,100-1,200 feet; 20, 27 June.

Although only four specimens were taken, the species is com-
mon during the spring and early summer months, in pastures and
along sunny road sides in the vicinity of Lago Catemaco. The
butterflies are attracted to moist earth and to the flowers of Inga
leptolohci, a tree common in pastures. G. philolaus exhibits the
high, soaring flight that is characteristic of most species in the
genus.

5. Graphhnn epidaus epidmis (Doubleday, Westwood, & Hewit-

son)

SPECIMENS: 11 ^ ^ , 3 $ $ ; 1,100 feet; 20 June-27 July.

Adults are very abundant during the spring and summer
months in Pastures, Recently Abandoned Milpas, and along
Hedgerows and sunny road sides throughout the Sierra but
particularly in the vicinity of Lago Catemaco. This distribution
correlates with the distribution of the larval food plant Annona
reticulata, a tree that produces sweet fruit and which is culti-
vated frequently by the local inhabitants. Adults exhibit the

57

248

(;ary n. ROSS

/. Res. Lepid.

characteristic Graph him flight and are attracted to mud puddles
and damp soil and sand. Immature stages (described in Ross,
1964b) were found commonly on the leaves of the food plant
during the summer months.

6. Graphium agesilaiis neosilaus (Hoffer)

SPECIMENS: 4 S S ; 2 mi. NE Catemaco, 1,100 feet, 2 July
1962, 2s S : I mi. N Soteapan, 1,400 fee, 28 March 1965, 2$ $ .

This “kite swallowtail” is uncommon and was collected only
along sunny road sides during spring and early summer. All
four specimens were taken as they drank from damp sand. The
previous Veracruz record is the “Sierra Madre Oriental” (Hoff-
mann, 1940).

7. Graphium caUiste calliste (Bates)

SPECIMENS: 16 6 6 ,l9; 4,100-5,100 feet; 3 March-7 April.
G. c. calliste is abundant above the canopies of the Alontane
Thicket and Elfin Woodland during March and April. The
butterflies descend to within a few feet of the ground usually
only to visit flowers, particularly those of Schistocarpha sp. (a
composite that is common on the open, sunny ridges and crater
walls ) .

TRIBE Troidini
SUBTRIBE Battiti

8. Battus polijdamas (Linnaeus)

SPECIMENS: 6 6 6,4 9 9; 0-2,700 feet; 10 March-14 Aug.
B. poly (lamas is abundant in the Littoral M7)odland and
common in all other formations except the Montane series.
Adults are attracted to the flowers of Lantana camara and to
moist soil and sand. The flight is relatively rapid and usually
within eight feet of the ground. When pinched, specimens pro-
truded their yellowish abdominal seent glands, which emitted
an acrid odor. Larvae were found on Aristolochia asclepiadifolia,
a vine that is common in the vicinity of the Popoluca Indian
villages on the Santa Marta massif.

9. Battus beliis varus (Kollar)

SPECIMENS: 4 9 9; 700-1,400 feet; 20 May-14 July.

This papilionid is locally common, being seen most frequent-
ly along the margins of the Semi-Evergreen Seasonal Forest in
the ravines within the oak and pine-oak forests. The flight
usually is above ten feet of the ground. Two of the four speci-
mens were taken as they fed on the blossoms of Inp^a spuria. The
abdominal scent glands produce an acrid odor.

58

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BUTTERFLIES OF TUXTLA

249

10. Battus laodanms copanae (Reakirt)

SPECIMENS: 1 5 , 1 $ ; 2 mi. NE Catemaco, 1,100 feet, 24
June 1962, 1 $ ; 28 June 1962, 1 5 .

This rare species was collected only in Pastures; the female
as it fed on the blossoms of Lantana camara and the male as it
flew about the blossoms of Inga spuria. The flight is relatively
rapid and erratic.

SUBTRIBE Troiditi

11. Parides photinus (Doubleday)

SPECIMENS: 8S 6,5 9 9; 700-5,100 feet; 30 March-30 Aug.
This species is common in all the Montane Formations and
the Semi-Evergreen Seasonal Forest on the Santa Marta massif.
The butterflies are particularly numerous along sunny trails
and in bright glades with the forests. The flight is relatively slow
and usually within four feet of the ground. However, when in-
dividuals are disturbed they fly rapidly with powerful wing
beats. The abdominal scent glands emitted an acrid odor when
the butterflies were pinched. The larval food plant is Arista-
lochia asclepiadifolia, the same as that of Battus polijdamas.
Immature stages are described elsewhere (Ross, 1964d).

12. Parides montezuma (Westwood)

SPECIMENS: 4s 6; 1,050-2,000 feet; 1 Aug.-29 Oct.

This tailed Parides is locally common, being found most fre-
quently in the Recently Abandoned Milpas surrounding the
Indian villages of Meeayapan and San Fernando. The butterflies
fly fairly rapidly between four and ten feet of the ground and
are attracted to flowers. The abdominal scent glands produce
an acrid odor.

13. Parides polyzelus poUjzelus (Felder)

SPECIMENS: 2U 6 ,'8 9 9 ; 0-1,900 feet; 13 March-5 Oct.
This papilionid is abundant in the Semi-Evergreen Seasonal

Forest, Lower Alontane Rain Forest, Littoral Woodland, Swamp
Forest, and along Hedgerows. In fact, this species is the most
common forestdnhabiting swallowtail. The flight is relatively
slow, weak, and usually within three or four feet of the ground.
The abdominal scent glands produce acrid odors.

14. Parides sesostris zestos (Gray)

SPECIMENS: 4 6 6,1 9; 0-1,750 feet; 12 July-29 Oct.

This swallowtail is locally common, being found in the Lower
Montane Rain Forest, Semi-Evergreen Seasonal Forest, Littoral

59

250

(;ary n. ROSS

J. Res. Lepid .

Woodland, and Swamp Forest. It and the following two speeies,
P. iphidamas and P. areas mijlotes, in addition to being very
similar in appearanee, seem to be very elosely related eeologic-
ally and ethologieally. Fo rexample, all three speeies prefer the
relatively bright or sunlit sections of mature forests below an
elevation of approximately 3,000 feet and usually are found in
loealized assemblages. The flight is rapid and within three to
five feet of the ground. As reported in Ross (1964a), the butter-
flies appear to have definite “Hyways” from which they very
rarely deviate. These Hyways usually are sections of relatively
wide trails. The abdominal scent glands produce an acrid odor.

15. Parides iphidamas (Fabrieius)

SPECIMENS: 16^ 6,9 9 9 ; 0-2,450 feet; 13 Mareh-18 Oct.
This speeies is locally common (more comn:ion than P. sesos-
tris zestos) in the Lower Montane Rain Forest, Semi-Evergreen
Seasonal Forest, Littoral Woodland, and Swamp Forest. The
Hight behavior is the same as that of P. sesostris zestos. The ab-
dominal scent glands, as those of other members of the genus,
produce an acrid odor.

16. Parides areas rni/lotes ( Bates )

SPECIMENS: IT'^ 6 , 13 9 9 ; 0-2,700 feet; 13 March-24 Oct.
P. areas mijlotes is locally abundant in the Lower Montane
Rain Forest, Semi-Evergreen Seasonal Forest, Littoral Wood-
land, and Swamp Forest. The Hight behavior is the same as that
described under P. sesostris zestos. The abdominal scent glands
emit an acrid odor.

TRIBE Papilionini

17. Papilio pohjxenes asterius Stoll

SPECIMENS: 3 6 6,2 9 9 ; 1,100-2,700 feet; 5 May-14 Sept.
This “Huted swallowtail” is uncommon and was collected
only in the Pinus-Quereiis Associes of the Deciduous Woodland
and in a small, disjunct patch of oak forest NNE of Catemaco.
Two of the five specimens are typical P. polyzenes a.steriiis
whereas the remaining three are of the morphotype named form
amerieiis Kollar. The Hight is characteristic of most members of
the genus Papilio — rather rapid with strong wing beats and
usually between six and 15 feet of the ground.

18. Papilio thoas autoeles Rothschild & Jordan

SPECIMENS: 12 6,5 9 9 ; 0-2,700 feet; 12 March-30 Sept.

This species is the most common swallowtail in the Sierra and

occurs in Recently Abandoned Milpas, Pastures, Semi-Evergreen
Seasonal Forest, Littoral Woodland, and along Hedgerows and

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BUTTERFLIES OF TUXTLA

251

most sunny road sides. The butterflies visit the blossoms of
numerous species of plants and mud puddles. The flight is
typical of Papilio spp. Larvae, described in Ross (1964d) were
found on Piper marginatum and Piper kerberi, both of which are
common along shaded stream banks in the Semi-Evergreen Sea-
sonal Forest on the Santa Marta massif.

19. Papilio androgens epidaurus Godman & Salvin
SPECIMENS: 1 $ <5,1$; 1,100-5,100 feet; 24 March-13 Oct.
This papilionid is common above the canopy of the Elfin

Woodland. Most butterflies were collected as they flew in
circular patterns between eight and 12 feet above the peaks
of Cerro Tuxtla and Volcan Santa Marta. This behavior is com-
monly known as “hilltopping.” Larvae (described in Ross,
1964b) were found during the summer months on Zanthoxylum
elephantiasis, a small tree that is common in the hedgerows
bordering Lago Catemaco.

20. Papilio anchisiades idaetis Fabricius

SPECIMENS: 6 ^ <5,8$ $ ; 500-1,800 feet; 20 June-9 Oct.
This swallowtail is common in Pastures and along Hedgerows
in the vicinity of Lago Catemaco and most towns and villages
throughout the Sierra where citrus trees are cultivated. Citrus
spp. being the larval food plant. The flight is typical of most
species in the genus. Immature stages are described in Ross
(1964b).

21. Papilio victorinus victorinus Doubleday
SPECIMENS: 1 <5 , 1 $ ; 2 mi. NE Catemaco, 1,100 feet, 28

July 1962, 1 S : 2.5 mi. NNW Ocotal Chico, 3,000 feet, 12 April
1965, 1 $ .

This species is rare; the male was collected in a sunny glade
within the Semi-Evergreen Seasonal Forest and the female along
a wide logging road in the Montane Rain Forest. Both specimens
were flying rapidly approximately ten to 12 feet above the
ground when collected.

252

GARY N. ROSS

J. Res. Lepid.

FAMILY PIERIDAE
SUBFAMILY Dismorphiinae

22. Dismorphia (Dismorphia) praxinoe (Doubleday)
SPECIMENS: 8$ $,12 9 9; 0-4,300 feet; 13 March-17 Nov.
D. praxinoe is locally common, being found principally in the

Lower Montane Rain Forest, Montane Rain Forest, Semi-Ever-
green Seasonal Forest, and Swamp Forest. The flight is very
weak and the butterflies very rarely rise more than four feet
above the forest floor. As reported previously (Ross, 1964a), this
slow, weak flight is very atypical for members of the Pieridae
but very similar to that of many species in the family Ithomiidae.

23. Dismorphia (Dismorphia) fortunata (Lucas)

SPECIMENS: 84$ $,19 9 9; 700^5,000 feet; 9 Feb.-30 Oct
This species locally abundant and found principally in the

Lower Montane Rain Forest, Montane Rain Forest, Semi-Ever-
green Seasonal Forest, and Swamp Forest. The flight is weak
and slow, much more so than that of the related species D.
praxinoe. The butterflies usually fly within one to two feet of
the forest floor. This flight behavior is very similar to that of
several species of ithomiids, principally Oleria paula, Ptero7itjmia
cotytto, and Hypoleria cassotis (see Ross, 1964a also).

24. Dismorphia (Dismorphia) euryope (Lucas)

SPECIMENS: 11 ^ ^ , 1 $ ; 3 mi. NNW Ocotal Chico, 4,100

feet, 23 Feb. 1965, 1 <^ ; 12 March 1965, 1 $ ; 4,300 feet, 7 March
1965, 1 ^ ; 4,400 feet, 17 June 1963, U ; 30 July 1963, 3^5;
4,800 feet, 5 April 1965, 1 ^ ; 16 July 1963, 2 $ $ : Peak Volcan
Santa Marta, 5,000 feet, 11 June 1963, 1 $ ; 5,100 feet, 26 May
1965, 1 ^ .

This pierid is local and uncommon; all specimens were col-
lected in the Montane Thicket and Elfin Woodland on Volcan
Santa Marta. The flight, particularly that of the males, is more
rapid and erratic than that of the preceding two species but
still slower than that of most pierids. The butterflies were col-
lected most frequently as they rested on leaves in relatively
bright or sunny glades within the forests. The body fluids of
pinched specimens smelled sour. The previous Veracruz record
is the “Sierra Madre Oriental” (Hoffmann, 1940).

( to be continued )

62

NOTICES

The Smithsonian F oreign Currency Program^ a national research
grants program^ offers opportunities for support of research in
Burma, Guinea, India, Pakistan, Poland (through 1976), and E gypt
in (among others) Systematic and Environmental Biology. Awards
are determined on the basis of competitive scientific review. The
deadline for submission is November 1 annually. F or further infor-
mation write the Foreign Currency Program, Office of International
Programs, Smithsonian Institution, Washington, D. C. 20560

l,5i00 exotic butterflies, insects, biologicals. 72 page catalog ^1. 00
Complete Scientific^ P. O. Box 307 Round Lake, Illinois 60073.

INDEXES: Index to volume 11 is in volume 13, no. 3, to volume 12
is in volume 13, no. 1.

PUBLICATIONS: The Swallowtail Butterflies of North America.
Hamiilton A. Tyler, Naturegraph Publishers Inc. Healdsburg, Calif.
95448. Paper ^5. 95, cloth ^9. 95.

Mites of Moths and Butterflies. Asher E. Treat, Cornell Univ.
Press, 124 Roberts Place, Ithaca, N. Y. 14850. ^35.00

Geographical Variability in Speyeria. Arthur H. Moeck. reprint.
Entomological Reprint Sperialists, P. O. Box 77224, Dockweiler Sta. ,
Los Angeles, CA 90007.

Butterflies of Lebanon. Torben B. Larsen. Nat'l Council for Sci.
Res., Republic of Lebanon. Avail, in U. S. A. from E. R, S. (see above)
^22. 50.

The Swallowtail Butterflies of East Africa, reprint. R, H.

Carcasson, F. R. E. S. E. W. Classay Ltd. in U, S. A, from E. R. S.

(see above). ^4.95.

Macr olepidoptera of Fiji and Rotuma. Gaden S. Robinson. E. W,
Classey, Ltd. in U, S.A. from E, R. S. (see above). ^25.95.

Biology, Ecology and Host Specificity of Microlepidoptera
associated with Quercus Agrifolia, Paul A. Opler. Univ. Calif. Press,
2223 Fulton St. , Berkeley, CA 94720. ^4. 25.

Colour Identification Guide to British Butterflies. T. G. Howarth.
Frederick Waine and Co., Ltd. 40 Bedford Square, London. WCIB.

New York 101 Fifth Ave, :^3. 00 net.

FOR SALE: Mounted, named local moths. Min. order 1000 at ^400.
per thousand in series up to 20 per species. Similar price for 1976
butterflies, S, G. Jewett, Rt. 1, Box 339, West Linn, Oregon, U. S. A.
97068.

WANTED: Correspondence with members interested in trading or selling North
American Lepidoptera. Send list of offers to: Marc Grocoff, 1950 Cottrill Lane,
Westland, Michigan 48185 USA

THE LEFUDOFTERA

Volume 14 Number 4 December, 1975

IN THIS ISSUE

A revision of

the North American Comadia

(Cossidae) Richard M. Brown 189

A note on Oeneis Melissa (Fabricius)

in the western United States (Satyridae)

Clifford D. Ferris 213

Post Pleistocene environments and
montane butterfly relicts
on the western great plains

Kurt Johnson 216

An ecological study of the butterflies of

the Sierra de Tuxtla in Veracruz, Mexico

Gary N. Ross

233

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