HARVARD UNIVERSITY

Library of the
Museum of
Comparative Zoology

umm Amy

'MB lb W

■

imtsr&MAiTY.

THJ1 JOURNAL OF RESEARCH
ON! THE LEPIDOPTERA

published by

The Lepidoptera Research Foundation, Inc.
at

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

EDITOR: William Hovanitz

ASSISTANT EDITOR: Karen Nielson 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.

CLASSES OF
MEMBERSHIP

Regular

$12

year

Family

15

year

Contributing

25

year

Subscribing

50

year

Sponsor

100

year

Life

250

for life

Subscriptions to the Journal are $15.00 per year. Special subscriptions are avail¬
able to students at $10.00 per year.

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.,
Arcadia, California, U.S.A. 91006. The publisher is THE LEPIDOPTERA RESEARCH
FOUNDATION, INC. The general editor is William Hovanitz at the above address.
The secretary-treasurer is Barbara Jean Hovanitz at the same address. The owner is
THE LEPIDOPTERA RESEARCH FOUNDATION, INC. THE LEPIDOPTERA
RESEARCH FOUNDATION, INC. is a non-profit organization incorporated under the
laws of the State of California in 1965. There are no bond holders, mortgages or other
security holders.

Second Class postage paid at Arcadia, California, U.S.A.

MUS. COMP. ZOOL
}.. (! . A R

Journal of Research on the Lepidoptera 16(l):l-67, 1977

E.b,- AUG 10197?

do Santa Barbara Museum of Natural, Rfstpxy
2559 Puesta del Sol Road, Santa Barbara, Calif ornla'6&W5

UNIVERSITY

STUDIES ON NORTH AMERICAN PHILOTES
(LYCAENIDAE)

V. TAXONOMIC AND BIOLOGICAL NOTES, CONTINUED
OAKLEY SHIELDS

Department of Entomology, University of California
Davis, Calif. 95616

INTRODUCTION

This paper continues in the format established in Part IV
(Shields, 1975) for the remainder of the named subspecies of
Nearctic Shijimiaeoides (= Philotes) (except that S. rita rita
and S. rita coloradensis will appear in a separate paper with
Michael Fisher). Parts II- V are intended to be read as a set
and not to be interpreted separately. One more paper in this
series is in preparation, describing several new subspecies and
reporting a unique collection of lohn Lane’s from southern
California. This project required six years to complete, as Shi¬
jimiaeoides ranks with C olios, Euphydryas, Speyeria, and Aga-
thymus in taxonomic complexity. Apologies are extended to
contributors for lengthy delays in returning some of their ma¬
terial, due to having all specimens before me for viewing, rather
than relying on comparisons drawn from memory. Eriogonum
distributions follow Reveal (1969b) for the most part.

NON-HOST ERIOGONUM

Certain Eriogonum species are not used by Shijimiaeoides.
I have never seen Shijimiaeoides flying around E. inf latum Torr.

6 Frem. and E. deflexum Torr. & Ives, two species commonly
encountered on field trips, nor around E. elatum Dough ex Benth.
or E. cernuum Nutt. None were seen on blooms of E. fendlerb
anum ( Benth. in DC. ) Small near Vermejo River, Hwy. 64; and

7 mi. S. Raton; both in Colfax Co., New Mexico ( VIII-16-70). A
visit to the type locality of E. apachense Reveal near Bylas,
Arizona ( see Reveal, 1969a ) on 30 Aug. 1970 produced negative
results. None were seen around the blooms of E. pelinophilum
Reveal at 8.6 mi. W. of Hotchkiss, Delta Co., Colorado (see
Reveal, 1973). No Shijimiaeoides were found on the blooms of
E. hieracifolium Benth. in DC. along Hwy. 73 from near Indian

1

2

OAKLEY SHIELDS

J. Res. Lepid.

Pine to Carrizo, Navajo & Gila Cos., Arizona, VII-27-70 ( Shields
#124). Eriogonum abertianum Torr. in Emory var. abertianum
grows commonly near Ramsey Canyon, Cochise Co., Arizona,
VII-28-70 ( Shields #126) and 13-14 mi. NE of Lordsburg,
Hidalgo Co., New Mexico, VIII-25-70 ( Shields #156) but is not
a host of S. vita rita at these locations.

BEHAVIOUR NOTES

At 6 rd. mi. S. Sherwin Summit, along Hwy. 395, Mono Co.,
V-18-70, S. enoptes langstoni flew commonly around the white-
flowered Eriogonum kennedyi var. purpusii (one oviposition
observed), while S. battoides nr. glaucon avoided this species
and alighted on the scarcer yellow-flowered E. umbellatum var.
chlorothamnus , in a mixed stand of the two.

At the canyon 3/4 mi. SSE Horse Thief Spgs., Kingston
Range, San Bernardino Co., IX-15-70, a fresh male S. enoptes
dammersi investigated a male of the white skipper Heliopet es
ericetorum that was perched with outstretched wings on the
mud. It tried to alight on the skipper (“hovering” over it as if
it were a flower) then flew off after this brief encounter. Per¬
haps significantly, dammersi was associated with the white-
flowered Eriogonum wrightii var. wrightii in this canyon.
Shijimiaeoides rita pallescens (Tilden & Downey, 1955)

Tilden & Downey. 1955. Bull. S. Calif. Acad. Sci. 1955, vol. 54,
pp. 25-29, pi. 11, ff. 1 & 3, pi. 12, ff. 4-6.

This was originally described as a separate species, but
Mattoni (1965, p. 94) noted that in the shape of the valve it is
similar to elvirae and thus considered pallescens to be a sub¬
species of rita. There is little doubt that Mattoni’s decision is
correct, as S. rita pallescens inter grades with rita elvirae in ex¬
treme western Nevada and eastern California and displays
affinities to rita emmeli in western Utah. The holotype male
comes from Little Granite Mountain, Dugway Proving Grounds,
Tooele Co., Utah, VIII-20-53 (H. E. Cott). The paratype series
was taken at this locality, at Dog Area in Dugway Proving
Ground, and in the Stansbury Mountains. Tilden & Downey
adequately describe pallescens and picture the adults and male
valve ( in comparison with rita rita ) . Previously known from only
the type locality, pallescens ranges across northern Nevada in
scattered locations.

Number of teeth on terminal end (cucullus) of valve = 11

16(1): 1-67, 1977

PHILOTES

3

(1), 12 (1), 14 (5), 15 (4), 16 (1), 17 (5), 18 (4), 19 (7),
20 (4), 21 (7), 22 (1), 24 (1); N = 41, average = 17-19, from
Sand Mtn., Currant, and Ralston Valley, Nevada.

DISTRIBUTION (viewed 62 $ gen. from 10 localities):
CALIFORNIA: Mono Co.: Mammoth, 1$, vii (CAS). NE¬
VADA: Lyon-Mineral Co. line: U.S. 95, Co. line, 5$ 7 $ , ix
(AM). Nye Co.: Jett Canyon, Toiyabe Range, 1 $ , viii (AM).
County undetermined: Montello Wells, Is, viii (CM). UTAH:
Tooele Co.: NW Willow Springs, Stansbury Mts., 17 $ 23 $ ,
viii (AM).

HOSTS AND DISTRIBUTION: Eriogonum kearneyi Tid-
estr. var. kearneyi. CALIFORNIA: Mono Co.: ca. 5-6 rd. mi.
WSW Calif.-Nevada border along Cal. State Hwy. 167, 6700',
T3N, R28E, 10 $ 11$, vii (AM), adult assoc. ( Shields #7).
NEVADA: Churchill Co.: SE base of Sand Mtn., 4000', T17N,
R32E, ca. 7 air mi. WNW Frenchman, 78 $ 47$, vii-ix (AM),
oviposition ( Shields #5 & #244). Nye Co.: NW corner T9N,
R58E, sand flats, 7 rd. mi. S Currant, 5000', on dirt rd. to Nyala,
Railroad Valley, 48 $ 60$, vii, viii (AM), oviposition & larvae
reared to adult ( Shields #3); 7 air mi. NW Lunar Crater (1/2
rd. mi. E Sandy Summit), on US Hwy. 6, 6000', T7N, R52E, E of
Hot Creek Range, 2$, viii (AM), adult assoc. ( Shields #11);
Ralston Valley, 5600', from jet. Hwy. 8 A & US Hwy. 6 to V-/i mi.
W. of this jet., E of Tonopah, T3N, R43E, 17 $ 14 $ , vii (AM),
oviposition ( Shields #4); 6 rd. mi. W of Warm Springs on US
Hwy. 6, 6200', E side Stone Cabin Valley & S end of Hot Creek
Range, T4N, R49E, 1# 2$, viii (AM), adult assoc. ( Shields
#12). UTAH: Tooele Co.: sand hills, 3 rd. mi. SW Johnsons
Pass, 5200', E side Skull Valley, T6S, R7W, 36$ 32 $ , viii ( AM ),
adult assoc. ( Shields #14). Eriogonum microthecum Nutt. var.
foliosum (Torr. ) Reveal. NEVADA: Nye Co.: 5 rd. mi. W of
Five Mile Spring on US Hwy. 6, 6400', at S end Monitor Range,
NW 1/4 of T3N, R47E, 33 3$, viii (AM), adult assoc. ( Shields
#13). Eriogonum baileyi S. Wats. var. baileyi. NEVADA:
Washoe Co.: 3 rd. mi. SSE of US Hwy. 395 on Nevada Hwy. 3B,
SE of Little Washoe Lake, 5100', T17N, R20E, 5$ 14 $ , vii
(AM), oviposition ( Shields #8). Eriogonum baileyi S. Wats. var.
divaricatum (Gandg. ) Reveal. NEVADA: Ormsby Co.: 1 mi. S
Carson City, 49007, at base of E slope of Carson Range, 2 $ 3 $ ,
vii (AM), adult assoc. ( Shields #9). Eriogonum heermannii
Dur. & Hilg. var. humilius (S. Stokes) Reveal. NEVADA:
Washoe Co.: 11 rd. mi. NW of Nixon, W shore Pyramid Lake,
3900', 1$, viii (AM), adult assoc. ( Shields #245). See Map 1.

4

OAKLEY SHIELDS

J. Res. Lepid.

Map 1. — Distribution of Shijimiaeoides rita pallescens (above dotted line)
and S. r. elvirae (below).

16(l):l-67, 1977

PHILOTES

5

Ssp. pallescens is in flight from early July to September.

One pair in a series of 17 males, 23 female pallescens from
Willow Springs, Tooele Co., Utah, are like emmeli.

One male from 5-6 mi. WSW Calif.-Nev. border along Hwy.
167, Mono Co., California, is unusual in that the aurora is almost
absent. Populations on E. baileyi in Washoe Co., Nevada, are
small and atypical.

Near Currant, Nye Co., Nevada, pallescens is at least double-
brooded, as larvae collected on 2 Aug. 1969 emerged prior to
30 Aug. 1969 (fresh adults were on wing 6 July 1969 and 2 Aug.
1969).

Eriogonum kearneyi var. kearneyi grows in sandy soils from
extreme east-central California eastward across central and
southern Nevada to western Utah and northwestern Arizona,
flowering from July to September. E. microthecum var. foliosum
is widespread and common from southeastern California in east¬
ern San Bernardino and Inyo Cos. eastward across southern and
central Nevada into northern Arizona and southern Utah, north¬
ward on the Colorado Plateau to Emery and Grand Cos., and in
widely scattered locations as far north as the Wyoming state
line, entering western Colorado in Mesa Co. and proceeding
southward and across southern Colorado to San Luis Valley
and across northern New Mexico to the Sangre de Cristo Mts.,
hence southward to central New Mexico, 4500-7500', flowering
from June to October. E. baileyi var. baileyi occurs in southern
and eastern California and western Nevada northward to south¬
eastern Washington and southwestern Idaho, flowering from May
to September. Var. divaricatum grows in Placer and Nevada
Cos., California and from Washoe to Douglas Cos., Nevada,
flowering from June to September.

Shijimiaeoides rita elvirae (Mattoni, 1965)

Mattoni. 1965 [1966]. Jour. Research Lepid. 1965, vol. 4; pp.
94-99, 84-85 (tables), 96 (map), 99 (type distr. ); f. 5, nos. r-w;
f. 1, col. 4 (4 ff.); f. 7, rows 3-4 (9 ff.).

The holotype male comes from 3.5 miles southwest of Pear-
blossom, Los Angeles Co., California, VIII-20-64. Mattoni’s de¬
tailed verbal description, with black-and-white photographs of
a series of adults and male and female genitalia, and a com¬
parison with the other named rita subspecies, makes repetition
here unnecessary. He recorded the foodplant as Eriogonum
plumatella and included distributional records. Comstock &
Henne (1967) gave a detailed life-history description of elvirae,

6

OAKLEY SHIELDS

J. Res. Lepid.

with a black-and-white painting of the egg, mature larva, and
pupa. They noted that the larvae “did not take normally to our
coastal E. fasciculatum.”

Number of teeth on terminal end (cucullus) of valve = 13
(2), 14 (2), 16 (2), 17 (1), 18 (1), 19 (2); N = 10, average =
16, from Pearblossom mostly, and two other localities.

DISTRIBUTION (viewed 13 $ gen. from 3 different locali¬
ties): CALIFORNIA: Inyo Co.: Big Pine Creek, 6000', 4$ 7?,
viii (KH, PO, CIS); above Bishop, 1 S , vii (LACM); 9 mi. W
Lone Pine, 7$ 5 $ , ix (CIS). Kern Co.: Tehachapi, 1$, viii
(LACM); Walker Pass, 5250', 1 $ , vi (DB); 1 mi. W Walker
Pass, 1 S , ix (CIS). Los Angeles Co.: wash-N boundary Juniper
Hills, Mojave Desert, 3400', 11 £ 11$, vii-ix (AM, KH, KR,
LACM); Juniper Hills, 3400', 2 mi. S Pearblossom, 10 3 24?,
vii, ix (CIS, LACM, KR); Juniper Hills nr. Pearblossom, 7$
13?, viii, ix (JL, PO); Pearblossom, 4^ 5?, viii (GG). Mono
Co. : Sherwin Summit ( old 395 ) , 5 $ 2 ? , ix ( CIS ) .

HOSTS: Eriogonum plumatella Dur. & Hilg. var. jaegeri
(Munz & Johnst.) Stokes ex Munz. CALIFORNIA: San Ber¬
nardino Co.: 1 mi. E Morongo Valley, 2 $ , ix (AM), adult assoc.
( Shields #58). Eriogonum plumatella Dur. & Hilg. var. pluma¬
tella. CALIFORNIA: Los Angeles Co.: Juniper Hills, 3500', 3
mi. SW Pearblossom, 3$ 7 ? , ix (AM), adult assoc. & larvae
reared ( Shields #64). Eriogonum microthecum Nutt. var. ambi-
guum (M. E. Jones) Reveal in Munz. CALIFORNIA: Inyo Co.:
Onion Valley Rd., nr. Tub Sprgs., W of Independence, 4 $ 6 ? ,
viii (LACM, SE), adult assoc. (/. F. Emmel #113 & Shields
#128). Eriogonum kearneyi Tidestr. var. monoense (S. Stokes)
Reveal. CALIFORNIA: Mono Co.: Sherwin Summit, Old Hwy.
395, ix, adult assoc. (J. A. Powell, pers. comm., identified at
UCB Herbarium). See Map 1.

Ssp. elvirae flies from mid June to late September.

There appears to be some gene flow between pallescens and
elvirae based on undersurface characters. Populations from
Walker Pass, Kern Co.; Big Pine Creek, Bishop, and Lone Pine,
Inyo Co.; and Sherwin Summit, Mono Co., in California; and at
Little Washoe Lake, Carson City, and the Lyon-Mineral Co.
line on U.S. 95, in Nevada, display a mixed series, some being
similar to elvirae , others closer to pallescens.

Ssp. elvirae from Juniper Hills, Los Angeles Co., often re¬
semble the small form of battoides ellisii from the eastern Mojave
Desert on the underside. At Walker Pass, Kern Co., elvirae is
sympatric with battoides bernardino, and at Sherwin Summit,

16(1 ):l-67, 1977

PHILOTES

7

Mono Co., it is sympatric (but allochronic) with enoptes enoptes.

Eriogonum plumatella var. plumatella occurs from southern
California to Mohave Co., Arizona (and perhaps southwestern
Utah), flowering from August to November. Var. jaegeri is con¬
fined to the Mojave Desert, California and Mohave Co., Arizona.
On 11 Sept. 1969, J. F. Emmel and I inspected E. plumatella in
full bloom at a wash along Kelbaker Rd., S. of Granite Mts., SW
of Providence Mts.; near Colton Well, 3100', S. end of Colton
Hills; and near Weaver’s Well, Old Woman Mts.; all in San
Bernardino Co., California, without finding any elvirae adults.
Nor did I find elvirae in Sepetmber on E. plumatella near
Scissors Crossing, San Diego Co., California, which supports a
large colony of Apodemia mormo deserti.

Eriogonum microthecum var. ambiguum is found in dry
rocky places from southern Washoe Co., Nevada and adjacent
Nevada Co., California, southward along the eastern flank of the
Sierra Nevada and adjacent desert ranges to Mineral and Esmer¬
alda cos., Nevada, and Mono and Inyo cos., California, 6500-
10,500', flowering from July to September. Eriogonum kearneyi
var. monoense grows on gravelly soils along the eastern flank of
the Sierra Nevada from extreme southern Mono Co. into Inyo
Co., California (endemic), flowering from July to October.

Shijimiaeoides enoptes dammersi (Comstock & Henne, 1933)
Comstock, J. A., & Henne. 1933. Bull. S. Calif. Acad. Sci. 1933,
vol. 32, pp. 23-26.

The holotype male was taken at Snow Creek, Riverside Co.,
California, IX-9-32 (C. Henne). Some 70 paratypes were dis¬
tributed along USNM, Canadian Museum, LACM, and CAS,
collected at Snow Creek (mostly) and Chino Canyon, both
located at the base of the San Jacinto Mts. near Palm Springs.
The type description leaves no doubt as to the identity of this
fall, desert subspecies of enoptes from southeastern California,
extreme southern Nevada, and northwestern Arizona. The only
subspecies that could be confused with dammersi is rita rita ,
which is very similar in appearance but is readily separable
upon examination of the male genitalia ( rita has cristae which
enoptes lacks). Ssp. rita rita flies in central and southeastern
Arizona, southwestern New Mexico, and extreme western Texas.
Foster H. Benjamin examined the male genitalia of dammersi
for Comstock and Henne and stated that “it is allied to enoptes”
(I have examined the large series of paratypes in the LACM

8

OAKLEY SHIELDS

J. Res. Lepid.

collection.) The subspecies is accurately pictured in Emmel &
Emmel (1973, pi. 8, figs. 10-12). A description and figures of
the egg, larvae, and pupa appear in Comstock & Henne ( 1965 ) ,
and the mature larva and pupa are illustrated in Emmel &
Emmel (1973, fig. 56).

Number of teeth on terminal end (cucullus) of valve = 10
(1), 11 (2), 12 (7), 13 (9), 14 (11), 15 (3), 16 (3), 17 (3),
18 (2); N = 41, average 13-14, from Banning, Granite Mts., and
jet. 79 & 179 mostly, plus two other localities.

DISTRIBUTION (viewed 78 3 gen. from 21 localities):
ARIZONA: Gila Co.: East Verde River, 10 mi. N Payson, 1 2 ,
ix ( UCD ) ; Tonto Bridge, 4900', 72 mi. NE of Phoenix, 1 2 , ix
(AM). Maricopa Co.: 1 mi. N Sunflower, 7 3 , ix (KR). Mohave
Co.: Hualapai Mts., 6$ 3 $ , ix (LACM, KH). Yavapai Co.:
Black Hills, 4 3 1 $ , ix (DB); Hillside, 33 3 $ , ix (DB); 1 mi.
S Hillside, 1 3 , ix ( JS ) ; nr. Jerome, 3 3 , ix (DB); Mingus Mtn.,
5500', 6 3 1 $ , ix, (DB, CIS); Sycamore Creek, 3500', 13, x
(DB); nr. Verde Hot Spr., 4 3 5$, ix (DB). CALIFORNIA:
Riverside Co.: Chino Canyon nr. Palm Springs, 3 3 , ix, x (AM,
LACM); Palm Springs, 13, x (LACM); Snow Creek, nr. Palm
Springs, 13 3 13$, ix, x (LACM, GG, JL, YU, CAS); White-
water, 10 3 7 $ , ix, x ( LACM, CAS ) ; Whitewater Canyon, 7 3

5 $ , ix, x (UCD, LACM, AM). San Bernardino Co.: nr. Barnwell,

1 3 1 $ , ix (CIS); Cedar Cyn., Mid Hills, New York Mts., 5 3

6 $ , ix (KH); 5 mi. NW of Cima, 4900', 4 2 , ix (MCZ); King¬
ston Range, 6400', 1 2 , ix (LACM); Mtn. Pass Sta., Ivanpah
Mtns., 12, ix (CIS); New York Mts., 13 3 10 2 , ix (KH,
LACM); Tecopa Pass, 3/4 mi. S of, 4900', 13, ix (AM). San
Diego Co.: nr. Camp Ole, Laguna Mts., 4 3 , viii (CIS); Laguna
Grade, 5000', Laguna Mts., 1 2 , ix (AM); Hwy. 79 nr. Lake
Henshaw, 5 3 4 2 , viii, ix ( KH ) ; Lake Henshaw, Hwy. 79, 4 3
12, ix (CIS, PO); 2 mi. E Lake Henshaw, 2 2 , ix (LACM);

2 mi. W Manzanita, 4 2, x (CIS); Mason Valley, Box Canyon,
1 2, x (LACM); Sentenac Canyon, 1 3 , ix (CM); nr. Warners
Hot Spgs., 1 3 1 2 , ix (LACM). NEVADA: Lincoln Co.:
Meadow Valley Wash, 4200-4400', 8 rd. mi. S turnoff from Cali-
ente, T5S, R66E, 1 3 , ix (AM).

HOSTS AND DISTRIBUTION: Eriogonum wrightii Torr.
ex Benth. in DC. var. wrightii. ARIZONA: Mohave Co.: 1.0 to
1.2 road mi. NW of Hualapai Mountain Park, Hualapai Mts.,

3 2 , ix (AM), adult assoc. ( Shields #180). Yavapai Co.: 4 rd.
mi. E of Cherry, 2 3 1 2 , ix (AM), adult assoc. ( Shields #53);
3 rd. mi. NE of jet. Hwy. 79 & 179, along Hwy. 79, 11 3 7 2 , ix

16(1): 1-67, 1977

PHILOTES

9

(AM), adult assoc. ( Shields #52). CALIFORNIA: San Bernar¬
dino Co.: large canyon just SE of Clark Mtn. (Peak), ca. 5600-
6000', 5 rd. mi. NW of Mountain Pass, 1 $ , ix (AM), adult
assoc. ( Shields #177); vicinity of Cove Spring, 4100', T8N, R13E,
sec. 8 (SE 1/4) & 17 (NE 1/4), Granite Mts., just SW of Provi¬
dence Mts., 29$ 8 $ , ix (AM), adult assoc. ( Shields #59);
canyon 3/4 air mi. SSE of Horse Thief Springs, Kingston Range,
6$ 2 $ , ix (AM), oviposition ( Shields #176); Mid Hills, 5100',
NW exposure of ridge, T13N, R15E, NE 1/4 of sec. 31, along
road, 8 $ 20 $ , ix ( AM ) , adult assoc. ( Shields #60 ) . Eriogonum
wrightii Torr. ex Benth. in DC. var. membranaceum Stokes ex
Jeps. CALIFORNIA: San Diego Co.: Power Station, S of Warn¬
er’s Hot Spgs. at turnoff to Borrego. 1 $ , ix (AM), adult assoc.
( Shields #183). Eriogonum elongatum Benth. var. elongatum.
CALIFORNIA: Riverside Co.: V/2 mi. S Banning, 2300', along
Hwy. Rl, T3S, R1E, sec. 15 (S center of), 42 3 25 $ , ix (AM),
adult assoc. (Shields #57). See Map 2.

Ssp. dammersi is on the wing from late August to late Oc¬
tober.

At Snow Creek, Riverside Co., one female enoptes dammersi
in a series of 13 has blue flecks in a band above the aurora on
the upperside of the secondaries. One female in six from Mid
Hills, San Bernardino Co., has some basal blue, and four females
from Cima, San Bernardino Co., have some basal blue, appar¬
ently lacking in other dammersi populations.

Ssp. dammersi is sympatric (but allochronic) with battoides
martini at Clark Mtn. and Horse Thief Springs, San Bernardino
Co., with enoptes mojave in the Providence Mts., and sympatric
and synchronic with a dwarf battoides ellisii form at Clark
Mtn. At 4 mi. E. Cherry, Yavapai Co., Arizona, dammersi flies,
while at Cherry itself, only rita rita is reported. Likewise, these
two subspecies are sympatric and synchronic or nearly so at
1 mi. S. Hillside, Yavapai Co. Both species use Eriogonum
wrightii var. wrightii at these localities.

Comstock & Henne (1965) observed oviposition and reared
adults of dammersi from Eriogonum elongatum at Snow Creek
and Chino Canyon near Palm Springs, Riverside Co., California.
As they point out, E. elongatum “is more widely distributed in
southern California and Baja California than is the associated
butterfly.” It is also associated with E. elongatum at Banning,
Riverside Co., and at Manzanita, San Diego Co. I have checked
a number of localities for E. elongatum in southern California
during September without locating dammersi.

10

OAKLEY SHIELDS

J. Res. 1

Map 2. — Distribution of S. enoptes dammersi.

16(1): 1-67, 1977

PHILOTES

11

E. elongatum var. elongatum occurs in coastal central and
southern California and northern Baja California, Mexico, flower¬
ing from August to November. E. wrightii var. i vrightii is found
in southeastern California eastward to western Texas and south¬
ward to northern Mexico, flowering from July to September.
Var. membranaceum is distributed in southern California and
northern Baja California, flowering from August to October.

Shijimiaeoides enoptes ancilla (Barnes & McDunnough, 1918)

Barnes & McDunnough. 1916. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, pi. 11, ff. 2 & 5 (as glaucon), p. 152.

Barnes & McDunnough. 1917. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, pi. 17, f. 2 (as glaucon), p. 264.

Barnes & McDunnough. 1918. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 4, pp. 77-79 (O. D. fixed here).

In 1918 Barnes & McDunnough published a correction to
their assertion in 1916 and 1917 that glaucon was the Great
Basin race of enoptes, after viewing the type specimen of
glaucon and finding it had a battoides genitalia (see discussion
of glaucon under spaldingi in Part IV). Thus they erected the
name ancilla for this enoptes subspecies which otherwise so
closely resembles battoides glaucon. Phenotypically ssp. ancilla
differs only slightly from glaucon in having, especially in fe¬
males, “a rather rougher squammation on the underside and a
thickening of the black basal fringe line; the males show only
traces of fulvous on upper side of secondaries.” Ssp. glaucon
and ancilla are separated with assurance only after examining
the male genitalia. “Our type specimens are a series of 6 $ , 4 $
from Eureka, [Juab Co.], Utah (July 1-7).” Two of these were
figured in black-and-white as glaucon in 1916; these leave no
doubt about the proper identity of ancilla. I have examined a
series of 10 8 4 $ ancilla from Eureka, 6500-7500', Juab Co.,
Utah, taken in June and July (CM, gen. det. ). So far, glaucon
has not been found in Utah. Langston’s ( 1969 ) statement that
ancilla is geographically sympatric with rita pallescens in north¬
western Utah is true in the broad sense, but I am unaware of
any specific localities where the two fly together.

Number of teeth on terminal end (cucullus) of valve = 9 (3),
10 (3), 11 (3), 12 (3), 13 (3), 14 (5), 15 (3); N = 23, average
= 13-14. The localities are mostly Kingston Canyon and Bonanza
Gulch, Nevada; others are singletons from seven localities.

12

OAKLEY SHIELDS

/. Res. Lepid.

DISTRIBUTION (viewed 225 # gen. from 105 localities):
COLORADO: Adams Co.: Denver, 1 # , viii (CM). Boulder Co.:
Boulder, 1# 1 2 , vi (AM); Boulder Canyon, 14# 5 2 , v, vii
(YU, AM); Buckingham Pk., 6500', 1 $ , vi (YU); Chataqua
Mesa, 6 3 5$, v (JS); Eldora, 2 $ , vi vii (YU); Flagstaff Mt.,
7$ 6$, vi, vii (AM, YU, LACM, DG); 4-Mile Creek above
Sunset, 7 $ , vi (AM, JS, LACM); 4-Mile Canyon nr. Sunset, 8 $ ,
vi, vii (PO, SE, AM, LACM); Gregory Canyon, Boulder, 36 3
2$, v-vii (YU, LACM, JS, AM); Lefthand Canyon, 4$, vi
(AM, LACM); Magnolia, 1$, vii (YU); Nederland, 1$, vii
(AM); Pennsylvania Gulch, near Sunset, 1 $ 1 $ , vii (JS, AM);
Sawmill Hill., nr. Ward, 1 A , vii (LACM); Six Mile Canyon, 1 $ ,
v (LACM); R. R. Grade, NW Sugarloaf, 2 $ , vi (PO). Clear
Creek Co.: Clear Creek, 4$ (CM); Fall River Rd., 7500', 2 mi.
W Idaho Springs, 7 $ 1 2 , vi, vii (MF); nr. Morrison, 1 $ 1 $ ,

v (BM); Mt. Evans, 13,000', 1 $ , vii (CM); York Gulch, 8000',
Fall River, 1 $ , vi (JS). Delta Co.: Black Ridge, 6500', Smith
Fork Canyon, 14$ 9 2 , vi (SE, LACM). Gunnison River, 5 mi.
S Austin, 6000', 22, vi (SE); S end of Landing Field, E side
Redlands Mesa, NE % of SW %9 sec. 16, T14S, R93W, 6300',
21 $ 2 2, v (AM); Rogers Mesa, 5850', nr. Filter Plant, 3#, v
(SE); Smith Fork Canyon, 5500', 4$ 2 2 , v, vi (SE, LACM);
Smith Fork of Gunnison R., W Hotchkiss, 1 $ , v (SE); 2 mi. S
Smith Fork Canyon, 6500-7000', 4 $ , v (SE). Eagle Co.: 2 mi. N
Basalt, 7500', 1 $9 vi (SE). Garfield Co.: E. Elk Creek, 6800-
7000', 8 $ , vii (SE); No Name Creek, 6400-7400', 1 # , vii (SE).
Gilpin Co.: Golden, 2 $ 1 2 , vi (UCD, CM); 1 mi. into N. Fork
Clear Creek Canyon, 16# 7 2 , vii (JS); Rollinsville, 1#, vii
(AM). Grand Co.: Beaver Creek, 1 $ , vii (JS); Grand Lake, 1 $ ,
vii (CM). Gunnison Co.: 1.5 mi. N Almont, Hwy. 135, 1 2, vii
(AM); Black Mesa, 8500', 1 # , vii (SE); Black Mesa Rd., 8500',
7 $ 1 2 , vii ( SE ) ; Cement Creek, 1 $ , vii ( CIS ) ; Curecanti
Creek, 9000', 1 2 , vi (SE); Gothic, 9500-10,500', 4 $ , vi, vii (YU,
AM); Hwy. 50, 5 mi. W Gunnison, 7000', 9# 6 2, vi (SE);
Vulcan, 7 mi. S Iola, 1#, vi (SE). Jefferson Co.: Bear Crk.
Canyon, 6000', nr. Morrison, 14 $ 1 2 , vii (MF); Castle Rock nr.
Golden, 24 $ 26 2 , v (JS, LACM); Chimney Gulch, 5# 6 2 , v,

vi (CM); gulch % mi. S Chimney Gulch, 11#, vi (JS, GG);
Clear Creek Canyon, 2# 12, vi (SE, LACM); Cub Creek,
Evergreen, 1 # , vi (AM); Dedisse Park, Evergreen, 1 # 3 2 , vii
( AM ) ; Deer Creek Canyon & Hwy. 75, 1 # , vii ( CIS ) ; Golden
Gate Canyon, 1#, vi (JS); Indian Hills, 3# 1 2 , vii (AM);
Lookout Mt., 6500-7200', above Golden, 9# 6 2 , vi, vii (DG,

16(1): 1-67, 1977

PHILOTES

13

BM, LACM, MF); Mother Cabrini Shrine, 3 $ 1 2 , vi (JS, AM);
Phillipsburg, 4$ 4 2 , vii ( JS); Platte Canyon, 2$ 1 $ , v (CM);
Red Rocks Park, 25 $ 8 $ , vi (CIS, JS, LACM); Rooney Ranch,
1 $ , v (AM); Table Mtn., 2 $ , v ( JS); E end of S. Table Mtn.,
7 $ 3 2 , v (JS); Whitehall, 1 $ , vii (CM). Larimer Co.: 3 mi. W
Bellevue, 6700', 3 $ 1 $ , vi ( MF ) ; Copeland Lake, Longs Pk.
Trail, Estes Park, 1 $ , (MCZ); Horsetooth Res. Area, 5400', 1 $
1 $ , vi (MF); Lily Lake, Estes Park, 1$, (MCZ); Long’s Pk.
Inn, Estes Park, 9500', 20 £ 6$ , (MCZ); Soldier Canyon, 6000',
I $ , vi (MF). Moffat Co.: Juniper Mtn. W of Craig, 1 $ 2 $ , vi
(SE); Plug Hat Picnic Area, 7000', Dinosaur Nat’l Mon., 1 $ , vi
(SE). Montezuma Co.: Navajo Canyon, 8000', Mesa Verde Nat’l
Park, 1 $ , vi (AM). Pitkin Co.: Aspen Area, 7500-9500', 3 mi. W.,
I $ , vi (MF). Routt Co.: 4 mi. N Hayden, 10 £ 3 $ , vi (LACM,
SE). IDAHO: Cassia Co.: Bostetter Cmpgd., Rock Cr.-Oakley
Rd., 7000', Sawtooth Nat’l For., 3$ 1 2 , vii (AM); 2 mi. E.
Bostetter Cmpgd., 7000', 1$, vii (AM); Divide N. of Cache
Peak, Oakley-Elba Rd., Sawtooth Nat’l For., 7500', 28 $ , vii
(AM); Rock Creek, S. of the village of Rock Crk., Sawtooth Nat’l
For., 6000', 3$, vii (AM). MONTANA: Madison Co.: Three
Forks, 2$ , vi (CM). Sweet Grass Co.: sage hilltop nr. Big Tim¬
ber Creek, 4400', 1 $ , vi (JS); Little Timber Creek, 5743', Crazy
Mts., 1 $ , vi (YU). County undetermined: Big Timber Canyon,
ca. 6500', (Half Moon Park), Crazy Mts., 1 $ , vi (AM); Rock
Creek & Madison River, Gallatin Nat’l For., 1 S 3$, (CM);
Teepee Creek & Madison River, Gallatin Nat’l For., 2$ 2$,
(CM). NEVADA: Elko Co.: Angel Lake, 8100', Ruby Mts., 13
mi. SW Wells, 1 $ , vii (AM); Horns Gulch, Mountain City, 1 $ ,
vi (CIS); Jarbidge, 6500', 2$, vii (KH); approx. 3 mi. S Jar-
bidge, 6500', 4 $ , vii (JL, KH); Jarbidge Wilderness, Pine Creek
Camp Rd., 1 $ , vii ( AM ); Pine Creek, nr. Jarbidge, 2 $ , vii (GG).
Humboldt Co.: Indian Crk., 6000', Santa Rosa Mts., 4 $ , vii
(AM); Martin Creek Ranger Station, 7000', Santa Rosa Range,
2$, vii (AM). Lander Co.: lower end Birch Creek Canyon,
6200-6500', Toiyabe Range, 2 2 , vi (AM); 4 rd. mi. E. Carroll
Summit, 6700', 9$ 10 $ , vi (AM); Kingston Canyon, Toiyabe
Range, 5 $ 1 $ , vii, viii (JL, AM); S & W-facing slopes of Vic-
torine Canyon, 8000-10,000', S side of Bunker Hill, Toiyabe
Range, 67 $ 419, vii (AM). Nye Co.: Summit Canyon, 7000-
7200', E side Toiyabe Range, 30 <2 1 2 , vi, vii (AM). Pershing
Co.: Buena Vista Creek, 5700-6300', E slope Humboldt Range,
22, vii (AM). Washoe Co.: “T. H. Ranch Ca. Pyramid Lake”,
7$, vi (LACM). OREGON: Harney Co.: l/2 mi. W Burns P.O.,

14

OAKLEY SHIELDS

J. Res. Lepid.

4100', 3 3 5 $ , vi (CIS); Devine Canyon, 12 mi. N Burns,
5036', 13, vi (CIS). UTAH: Cache Co.: Green Canyon, nr.
Logan, 9 3 , vii (AM). Davis Co.: Mill Creek Canyon, 5300', 1 9 ,

vi (CIS); Mueller Park, Wasatch Mts., 1 3 , vi. (AM). Duchesne
Co.: Wolf Crk. Cyn., 7320', 14 mi. NW Tabiona, 4 3 , vi (CIS).
Emery Co.: Big Spring Wash, 2 3 4 9 , v (KT). Juab Co.: Eureka,
6500-7500', 10 3 4 9 , vi, vii (CM). Salt Lake Co.: Alta, 4$ 2 $ ,

vii ( MCZ); City Creek Canyon, 5300', 4 3 , vi (CIS, MCZ, CM);
Lamb’s Canyon, Wasatch Mts., 23, vii (AM); Mill Crk., 6000',
Wasatch Mts., 23 , vi, vii (AM); Mineral Fork, Big Cottonwood
Canyon, 1 3 , vii ( KT ) ; Parleys Canyon, 6000', 1 $ , vi ( CIS ) .
Summit Co.: Co-op Creek, Uintah Mts., 1$, vii (KT); East
Canyon, 1 3 , vi (CIS); 5 mi. E Kamas, 7500', 4 3 , vi (CIS);
Zi mi. S Kimball Jet., Uintah Mts., 4$ 3 $ , vi (LACM, AM);
Park City, 11 3 5 9 , vii (CM); N. Fork Provo River, 7500',
Uintah Mts., 5 3 2 9 , vii (LACM, AM, KT); Shingle Creek,
Uintah Mts., 11 3 5 9 , vi, vii (JS, KT, LACM); Snyderville, 21 3
25 9 , vi (AM, DG, JS, KT, LACM); nr. Snyderville, 3 3 1 9 , vii
(JS, KT); St. Hwy. 105, 7000', 4 3 89, vii (DB). Tooele Co.:
Bennion Crk., Sheeprock Mts., 3 3 , vi (AM); South Willow
Creek, Stansbury Mts., 4 3 , vi, vii (SE, KT, CIS); Stockton, 7 3
2 9 , vi (CM). Uintah Co.: Blue Mtn. Plateau, 8000', Dinosaur
Nat’l Mon., 1 3 , vi (SE); S slope Blue Mt., 8000', 1 3 , vi (CM).
WYOMING: Albany Co.: Johnson Creek, Sybille Canyon, 1 3 ,

vi (DG); Sybille Canyon, 6500', 63 3 9 , vii (DG). Lincoln Co.:
Swift Creek, nr. Afton, 63 1 9 , vi (AM). Park Co.: Roosevelt
Camp, Yellowstone Nat’l Park, 19, vii (CAS). Sublette Co.:
Elkhart Park, 13, vii (DG). Teton Co.: Blacktail Butte, Teton
Mts., 13, vi, vii (MCZ); Grant Teton Nat’l Park, 83 4 9 , vi
(CAS); 4 mi. E. Jackson, 13 3 , vii (AM); 6 mi. N. Jackson, 1 9 ,

vii (DG); Jackson Hole, 1 3 , vi (CAS); Jenny Lake, Teton Mts.,
32 3 13 9 , vi, vii (CM, YU, LACM); Moose, Teton Mts., 3 3 , vi
(LACM); Moran, 3 3 (CM); the Potholes, G. Teton N. P., 8 3 ,
vi (JS, LACM); Signal Mt., Teton Nat’l Park, 2 3 5 9 , vii
(LACM); String Lake, Teton Mts., 11 3 4 9 , vi (LACM); Strong
Lake, 6750', Grand Teton N. P., 1 3 , vii (CIS); Teton Mts., 14 3
1 9 , vii (MCZ, CM); Teton Pass, 8400', 3 3 1 9 , vii ( DG, AM );
Teton Trail, Teton Mts., 8 3 , vi (LACM); Timber Island, Teton
Mts., 13 (CAS). Some additional records appear in De Foliart
(1956) and Klots (1930).

HOSTS: Eriogonum heracleoides Nutt. var. heracleoides.
IDAHO: Bear Lake Co.: 8 Mile Canyon, S of Soda Springs,

16(l):l-67, 1977

PHILOTES

15

Cache Nat’l For., 6500', 2 5 2 9 , vii (AM), adult assoc. (Ellis
#1086). NEVADA: Elko Co.: slope /2 air mi. S of Bonanza
Gulch, 8000-9500', T46N, R58E, sec. 27, 28, 34, Jarbidge Mts.,
14 5 6 $ , vii (AM), oviposition (/. F. Emmel #195). Eriogonum
umbellatum Torr. var. aureum (Gandg. ) Reveal. COLORADO:
Pitkin Co.: Aspen (NE end of airport, hill-slope), U 1? reared
from larvae (AM), ( Shields #137); the ants Tapinoma sessile
(Say) and Formica montana Emery (det. R. R. Snelling) were
tending the larvae. NEVADA: Lander Co.: Kingston Canyon,
6600-6800', Toiyabe Range, 10 5 4 $ , vi (AM), adult assoc.
( Shields #1). Eriogonum umbellatum Torr. var. umbellatum.
NEVADA: Nye Co.: 5 rd. mi. W Five Mile Springs on US Hwy.
6, 6400', NW % T3N, R57E, S end Monitor Range, 2 5 reared
from larvae (AM), (/. F. Emmel #183). UTAH: Grand Co.:
Gateway-Castleton Rd., 2 mi. W Polar Mesa turnoff, N side
LaSal Mts., 8400', 1 5 , vi (AM), adult assoc. (Ellis #1049).
Eriogonum umbellatum Torr. var. nevadense Gand. NEVADA:
Lander Co.: 4.0 rd. mi. E Carroll Summit, 6700', 9 5 10 $ , vi
(AM) adult assoc. (Shields #208). Eriogonum lobbii Torr. &
Gray var. robustius (Greene) M. E. Jones. NEVADA: Lyon Co.:
1 mi. N Silver City, 15 3 2 , vi (AM), adult assoc. (Shields
#186). Eriogonum strictum Benth. ssp. proliferum (Torr. &
Gray) S. Stokes var. proliferum (Torr. & Gray) Reveal. NE¬
VADA: Elko Co.: Wildhorse Reservoir, Hwy. 51, 64 mi. N Elko,
6500', 9 5 1 $ , vi (AM), oviposition (Ellis #1073). Eriogonum
ovalifolium Nutt. var. ovalifolium. COLORADO: Mesa Co.:
Monument Cyn. trailhead, 5000', Colorado Nat’l Mon., 1 5 , iv
(AM), adult assoc. (Ellis #1002). See Map 3.

Ssp. ancilla flies from late April to early August.

A small-sized variety of ancilla with light blue males occurs
at the following localities: Black Ridge, Austin, Redlands Mesa,
Rogers Mesa, and Smith Fork Canyon, Delta Co.; Black Mesa,
Curecanti Creek, and Gunnison, Gunnison Co.; Monument Can¬
yon, Mesa Co.; Juniper Mtn. and Plug Hat, Moffat Co.; and
Hayden, Routt Co., all in Colorado; Big Spring Wash, Emery
Co.; Bennion Creek, Tooele Co.; and Blue Mtn. Plateau, Uintah
Co., all in Utah. Superficially these resemble enoptes mojave
but appear closest to ancilla. They are probably associated with
Eriogonum ovalifolium var. ovalifolium (see host record for
Monument Canyon). One dwarf female from a typical ancilla
population at Snyderville, Summit Co., Utah, is extraordinarily
similar to female mojave on the underside. Perhaps ancilla and
mojave are linked by way of this small, light form from Utah

16

OAKLEY SHIELDS

J. Res. Lepid.

Map 3. — Distribution of S. enoptes ancilla.

16(1): 1-67, 1977

PHILOTES

17

and western Colorado. Both subspecies average 13-14 teeth on
the valve and have the curved edge bearing the teeth upturned
to a 45° angle (also true in dammersi).

Specimens from the vicinity of Burns, Harney Co., Oregon,
are typical ancilla, despite the fact that they are geographically
close to the range of enoptes columbiae. Several specimens in a
series of 26 male columbiae from Mill Creek, Umatilla Co.,
Oregon, approach ancilla. ,

A male from Mt. Charleston, 7200', Clark Co., Nevada, VI-
26-59 (J. C. Downey, CIS) is nearest ancilla although the black
borders are wide (see fig. 3-4); enoptes nr. enoptes also flies
in the vicinity.

Although ancilla is usually readily distinguishable from
battoides centralis, ancilla resembles centralis on the underside
at Grand Lake, Grand Co., and Platte Canyon and Castle Rock,
Jefferson Co., Colorado.

Ssp. ancilla approaches the range of battoides glaucon in
western Nevada, at Pyramid Lake and Silver City. In Elko Co.,
Nevada, the two nearly overlap in range, glaucon occurring
north of Jarbidge on the Nevada-Idaho border and ancilla flying
in the Jarbidge Mts. at Pine Creek near Jarbidge, Jarbidge, 3 mi.
S. Jarbidge and Bonanza Gulch. Based on genitalia, two speci¬
mens from 8 mi. N. of Jarbidge appear to be hybrids between
the two.

Development of the orange auroral band on the lower half
of the underside primaries is present in an occasional ancilla
specimen from Birch Creek Canyon and Kingston Canyon, Lan¬
der Co., Nevada, and about half the females from Black Ridge,
Delta Co., Colorado, have a tinge of this primary aurora. One
male ancilla from Kingston Canyon is unusually large, about
the size of many spaldingi, with some orange on the veins and
a partial orange band on the underside of the primaries.

Eriogonum heracleoides var. heracleoides occurs from south¬
eastern Washington south to northeastern California, eastward
to Montana, Wyoming, and Colorado, flowering from May to
August. E. umbellatum var. umbellatum is common from Wash¬
ington to northern California eastward to Montana and Colorado,
flowering from June to September. Var. aureum is widespread
at lower elevations from Nevada to central Colorado, and in
scattered locations across Idaho and south-central Wyoming,
flowering from June to August. E. lobbii var. robustius is in¬
frequent in extreme west-central Nevada south and west of
Reno, flowering from July to August. E. strictum ssp. proliferum

18

OAKLEY SHIELDS

J. Res. Lepid.

var. proliferum ranges from Washington to central California
and eastward to western Montana, flowering from June to Aug¬
ust. E. ovalifolium var. ovalifolium is widespread and exceed¬
ingly variable, from British Columbia to southern Alberta south¬
ward through Washington and eastern Oregon into California
along the eastern flank of the Sierra Nevada to northern San
Bernardino Co., and southward from Canada along a broad
front through Idaho and Montana into Nevada, Utah, Wyoming,
and Colorado as far south as northwestern New Mexico and
northern Arizona, flowering from April to July.

Shijimiaeoides enoptes enoptes (Boisduval, 1852)

(See discussion in Part IV)

Number of teeth on terminal end (culcullus) of the valve =
13 (2), 14 (5), 15 (3), 16 (3), 17 (6), 18 (3), 19 (2),
20 (3), 21 (1), 23 (1); N = 29, average = 16-17, mostly from
Lake Spalding and Tehachapi Mts., with ten other localities
sampled.

DISTRIBUTION (viewed 112$ gen. from 78 localities):
CALIFORNIA: Alpine Co.: Brightman Station, 4$, vii (KH);
Ebbetts Pass, 1 $ , vi (LACM); Hermit Valley, 2$, vii (JS);
Hope Valley, 2$, vii (CIS); Sonora Pass Ridge, 9700', 4 $ , vi
(CAS); Woodfords, Voight Canyon, 1 $ , ix (DB). Calaveras
Co.: Bloods Crk. at N Fork Stanislaus River, 2 8 2 $ , vii (AM,
LACM); Camp Wolfeboro, 5600-5800', N Fork Stanislaus River,
7 $ 1$, vi, vii (LACM, PO, KH); N Fk. Stanislaus R., S Big
Meadows, 4 $ 1 $ , vi (CIS); N Fk. Stanislaus R., 2 air mi. E of
E end Big Meadow, 1 $ , vii (AM). El Dorado Co.: China Flat,
1$, vii (AM); China Flat Campground from Silver Fork Rd.
from Kyburz, 6 $ 2 2 , vi, vii (LACM, AM, NL); Echo Lake, 5 $ ,
vii ( LACM ) ; Glen Alpine Creek, 1 $ , vii ( DB ) ; 6 mi. N Ice
House Rd. off Hwy. 50, 1 $ , vii (AM); Pilot Creek, 2 $ 1 2 , vii
(CIS); 4 mi. N Riverton on Ice House Rd., 5$ 2 2 , vi (AM);
Twin Lakes, 2 $ 1 2 , vii (PO); Wright’s Lake, 1 $ 2 2 , vii (PO).
Fresno Co.: Huntington Lake, 1$ 1 2 , vi (LACM). Glenn
Co.: Hull Mtn. Lookout & ridge to E, ca. 6600-6900', W edge
Glenn Co., 1 $ 1 2 , vii ( AM ) ; Plaskett Mdws., 4 $ 1 2 , vi ( AM ) .
Humboldt Co.: Orleans, 1$ 1 2 , vi (CIS). Inyo Co.: Argus
Mts., 1 $ , vii (LACM); Brewery Spring, 4880', Surprise Canyon,
Panamint Range, 1 $ , vi (MCZ); Lone Pine Lake, 8500', 1$,
vi (CIS); near Mono Pass, 10,000-11,900', 5$ , viii (MCZ, PO);
Onion Valley Rd., 6800', nr. Tub Springs, 4 $ 2 2, viii (LACM,

16(l):l-67 , 1977

PHILOTES

19

SE); Ruby Lake, 11,500', 1$, viii (PO); above Ruby L., NW
Inyo Co., *19, ix (CIS); Whitney Trail, 9000', 12, vii (CIS).
Kern Co.: Blackburn Canyon, Tehachapi Mts., 1 4 1 $ , vii

(NL); Havilah, 14 1 $ , vi (LACM, CM); 4 mi. SE Keene,
1 $ , ix (CIS); entrance Kern Co. Park, 5200', Tehachapi Mts.,
3 4 , vi (NL); up draw NW of jet. Lebec-Frazier Park Rd.,
1#, vii (NL); Pine Tree Mine, 5000', Tehachapi Mts., 4 4 , vi
(NL). Lake-Glenn Co. lines : 2.0 rd. mi. S Hull Mtn. Lookout
turnoff just S of Hull Mtn. summit, 3 4 6 2 , vii (AM); 3 rd. mi.
SSE of Hull Mtn. Lookout, along Boardman Ridge, 1 4 1 2 ,
vii (AM). Lassen Co.: Bridge Crk. Campground, 1 2 , vii (CIS);
W side Eagle Lake, 1 4 , vi (LACM). Los Angeles Co.: Big
Rock Crk., nr. Valyermo, 5 4 10 2 , viii (GG); dirt rd. to Horse
Flats Campground from Angeles Crest Hwy. (1.0 mi. off hwy. ),
just N of Chilao Rec. Area, ca. 5900', San Gabriel Mtns., 1 4 ,

vi (AM); 0.7 rd. mi. N Newcomb Ranch Inn, wash bottom off

hwy., 5500', San Gabriel Mts., 1 4 , vi (AM); Pacifico Mtn.,
6400', San Gabriel Mts., 24, vii, viii (AM); San Antonio Can¬
yon, 1 <2 (CAS); So. Fork Campgd., So. Fork Big Rock Creek,
San Gabriel Mts., 5 4 5 2 , ix ( JL). Madera Co.: Minaret Ridge,
9265', 1$ 12, viii (AM). Mariposa Co.: Tenaya Canyon,

Yosemite, 12, vii (LACM). Modoc Co.: .8 rd. mi. E jet. of
Hwy. 395 & 299, along Hwy. 299, NE of Alturas, 1 2 , vii (AM);
3 rd. mi. E Pine Creek (town), Warner Mts., 2$, vii (AM);
Fandango Pass, 6100', 2 4 3 2 , vii (CIS); Pine Creek, 4800-
5100', 1-2 mi. E Pine Creek (town), 65 6 2 , vii (AM). Mono
Co.: Lake Lundy, 1 <2 , vii (KH); 1 mi. SE Lake Mary, 14,

vii (LACM); Lundy Lake, 14, vii (PO); Mammoth, 8$ 32,
vii ( LACM ) ; Mammoth Camp, 3 4 1 2 , vii ( LACM ) ; Mam¬
moth Lakes, 5$ 2 2 , vi, vii (LACM, CIS, YU); Minaret Pass,
1 4 , vii (LACM); Minaret Summit, Mammoth Lakes, 13 4 82,
vii (LACM, AM); Saddlebag Lake, 5 4 3 2, vii (AM, LACM);
Sherwin Summit (old 395), 1 4 1 2 , ix (CIS); Sonora Pass, 1 4 ,
vii (LACM); Tioga Pass, 5 4 22, vii, viii (JL, LACM); W
above Tioga Pass, 5 4 4 2 , vii (AM, LACM); 4.9 mi. E Tioga
Pass, Hwy. 120, 9400', 14 1 2 , vii (AM). Nevada Co.: Bear
River nr. Fuller Lake, 3 4 1 2 , vii (CIS); Donner Pass, 14,
vii (CIS); Donner Summit nr. Truckee, 3 4 1 2 , vi, vii (AM);
Truckee, 4 4 6 2 , vii ( CM ) ; entrance to Lake Spalding, 17 4
12 2, vii (AM, CIS). Placer Co.: Big Meadows, 14 2 2 , vi,
vii (CIS); Cisco, 6000', 22, vii (CIS, DB); Donner Summit,
1 4 4 2 , vii (AM); Lake Tahoe, 14, viii (UCD). Plumas Co.:
Blairsden, 12, vii (DB); nr. Chester, 1 4 , vi (AM); Keddie,

20

OAKLEY SHIELDS

J. Res. Lepid.

2 5 1 $ , vi ( YU ) ; 4 mi. W Quincy, 1 5 , vi ( CIS ) . San Bernar¬
dino Co.: Lytle Creek, 7000', San Gabriel Mts, 27 3 8$, vii,
viii (JL, LACM); Upper Lytle Crk., 20 3 15$, viii (KH, PO).
San Luis Obispo Co.: Cottonwood Pass, 2000', 1 5 , ix (AM).
Shasta Co.: 3 mi. N Burney, 1 5 3 $ , vii (CIS); between Caieye
& Ellery Crks., 15-16 rd. mi. NE of Hwy. 5 (McCloud arm
Shasta Res.), 1 5 , v (AM); 2 mi. N Hat Crk, 1 5 1 $ , vii (CIS);
Old Station, 1 $ , vi (CIS). Sierra Co.: 4 mi. N Camptonville,
11 £ 4 $ , vi (PO); 10 mi. W Downieville, 5$ 3 $ , vi (CIS);
Glen Alpine Crk, 6500', 1$, vii (KH); Gold Lake, 3 5 4$,
vii ( AM ) ; Shenanigan Flat, 12 mi. W Downieville, 2 5 3 $ ,
v, vi ( PO ) ; Sierra City, 1 5 , vi ( CAS ) ; W of Sierra City, 4 $ ,

vi (PO); Sierraville, 15, vii (CAS). Siskiyou Co.: Black Butte,

1 $ , viii (CIS); Klamath Lodge, Doggett Creek, 1 5 , vi (DB);
SW entrance to Lava Beds Nat’l Mon. (rd. from McCloud),
nr. Mammoth Crater, 5100', 4$ 2 $ , vii (AM); McBride Spgs,
5000-5500', nr. base Mt. Shasta, 4 5 2 $ , vii (GG); McCloud,
1$, vii (CAS); 5 mi. E McCloud, 2$ 2$, vi, vii (CIS); Jet.
Mt. Shasta Rd. (Hwy. A-10) & McBride Springs Cmpgd. Rd,
5 5 5$, vii (JL, KH); Mt. Shasta City, 3$, vi, vii (CIS);

2 mi. N Mt. Shasta City, 6 5 2 $ , vi (CIS); Pumice Stone Well,

5$, vii, (LACM); So. Fork Sacramento River nr. Mt. Shasta,
3800', 1$ 1 $ , vii (CIS); Soda Sprgs, 1 5 , vi (CM). Tehama
Co.: rocky outcropping on Lassen Trail Rd, 5300', T27N, R3E,
Sll, nr. Onion Butte, 1$ (SM). Trinity Co.: Carrville, 2$,

vii (CIS). Tulare Co.: Monarch Lake, 10,000', 1$, vii (CM).
Tuolumne Co.: Big Oak Flat, 2 $ , vi (CAS); Big Prater Mdws,
N Fork Stanislaus R, 1 5 , vii (CAS); Brightman Flat, 2$ 1 $ ,
vii (PO); Chipmunk Flat nr. Sonora Pass, 1 5 2 $ , vi, vii (CIS);
3% mi. SSW Hetch Hetchy Dam, 4800', 15 1 $ , ix (AM);
Hog Ranch, 4600', 1 mi. E Mather, 1 5 1 $ , vii ( CIS ) ; Relief
Res, nr. Sonora Pass, 1 5 , vi (CIS); Sonora Pass, 1 5 , vii (CIS);
5 mi. W Sonora Pass, 3$ 1 $ , vii (CIS); NW above Tioga
Pass, 1$, viii (CIS); Tuolumne Mdws, Yosemite, 3$ 1$,
vii (CIS, CAS). Ventura Co.: Mt. Pinos, 1 5, viii (LACM).
County undetermined: G. Alpine Cr, Tahoe, 15, vii (CIS);
Sonora Pass, 8000', 2 5 , vi (UCD); nr. Sonora Pass, 1$ 1$
(CIS); Tamarak L, Tahoe, 1 5 , vii (CIS); nr. Vidette Meadows,
2$, vii (CIS). Erroneous (?): Mt. Baldy, 2$, viii (CAS).
NEVADA: Clark Co.: end Kyle Canyon, Mt. Charleston, 1 $ ,
vii (LACM); Mt. Charleston, 1^, vii (AM). Douglas Co.:
W side Carson Valley, 15, ix (DB); SW of Genos, Kingsbury
Grade, 5700', 5 5 1 $ , vii ( CIS ) ; Glenbrook Creek, 2 5 1 $ ,

16(l):l-67, 1977

PHILOTES

21

vii ( DB ) ; Haines Canyon, 5000', 6 3 1 2 , v, viii, ix ( DB ) ;
Kingsbury Grade, 83, viii (NSM); St. Hwy. 19, spring, 5500',
53 , v, viii (DB). Ormsby Co.: Kings Canyon, 3$ , vii (NSM).
Washoe Co.: Verdi, 2$, viii (CIS). OREGON: Gilliam Co.:
4.5 mi. S Condon, 2500', 13, v (AM). Josephine Co.: Grants
Pass, 23 , vi (CM). Lake Co.: Warner Cn. Ski area, 12 mi. NE
Lakeview, 5450', 33, vii (AM). WASHINGTON: Klickitat
Co.: “Klickitat V., W.T.”, 1 3 , vii (MCZ).

HOSTS: Eriogonum nudum Dougl. ex Benth, var. pauci-
florum S. Wats. CALIFORNIA: Los Angeles Co.: South Fork
Big Rock Creek, est. 4500', San Gabriel Mts., 6 3 11 $ , viii-x
(AM, CIS), adult assoc. ( Shields #63). Eriogonum nudum
Dougl. ex Benth. var. deductum (Greene) Jeps. CALIFORNIA:
El Dorado Co.: along W & SW shore Loon Lake, ca. 15 air mi.
N Kyburz, vii (AM), adult assoc. ( Shields #233); 4 mi. N
Riverton on Ice House Rd., 5 3 2 $ , vi (AM), adult assoc.
( Shields #89); South Fork Silver Creek, ca 1 mi. E Chicken
Hawk Spring, 5000', ca. 4 mi. W Ice House Store, “21”, vi (AM),
adult assoc. ( Shields #205). Placer Co.: ridge between Mt.
Judah & Sugar Bowl, 7200-7500', nr. Donner Pass, 8 3 4 2 , vii
(AM), oviposition (/. F. Emmel #190). Eriogonum nudum
Dougl. ex Benth. var pubiflorum Benth. in DC. CALIFORNIA:
Inyo Co.: E. slope 4 Jeffery Campground, Bishop Crk., 7900',
vii (OS), adult assoc. ( Shields #248). Modoc Co.: .8 rd. mi.
E jet. Hwy. 395 & 299, along Hwy. 299, NE of Alturas, 1 2 ,
vii (AM), adult assoc. ( Shields #102); Electric Plant vicinity,
Modoc Nat’l For., 3.0 rd. mi. E Pine Creek (town), Warner
Mts., 23, vii (AM), adult assoc. ( Shields #104). Siskiyou Co.:
SW entrance to Lava Beds Natl Mon. (road from McCloud) nr.
Mammoth Crater, 5100', 4 3 2 2 , vii (AM), adult assoc. ( Shields
#101); ca. 6 rd. mi. NE Weed, on Hwy. 97, 12, vii (AM),
adult assoc. ( Shields #224). Eriogonum nudum Dougl. ex Benth.
var. nudum. CALIFORNIA: Glenn Co.: 1 mi. W Plaskett Ranger
Station, NW Alder Springs, 4 3 1 2 , vi (AM), adult assoc.
( Shields #242); 3 rd. mi. SSE Hull Mtn. Lookout, along Board-
man Ridge, Mendocino Natl For., 13 1 2 , vii (AM), adult
assoc. ( Shields #98). Placer Co.: Hwy. 80, 2 mi. E Emigrant
Gap, vii, oviposition ( Goodpasture #11). Sierra Co.: 3 /2 rd.
mi. SSW Bassetts, on Hwy. 49, est. 4800', “4”, vi (AM), adult
assoc. ( Shields #195). Eriogonum nudum Dougl. ex Benth. var.
scapigerum (Eastw. ) Jeps. CALIFORNIA: Mono Co.: Tioga
Pass, 10,000', vii (OS), adult assoc. ( Shields #251). Eriogonum
wrightii Torr. ex Benth. in DC var. subscaposum S. Wats.

22

OAKLEY SHIELDS

/. Res. Lepid.

CALIFORNIA: Inyo Co.: Onion Valley Rd., 6800', nr. Tub
Springs, 4 <2 2$, viii (LACM, SE), adult assoc. ( Shields #83).
Eriogonum saxatile S. Wats. CALIFORNIA: Los Angeles Co.:
dirt rd. to Horse Flats Campground from Angeles Crest Hwy.
(1.0 mi. off hwy.), just N Chilao Rec. Area, ca 5900', San Gabriel
Mts., 1 $ , vi (AM), adult assoc. ( Shields #91). Eriogonum
davidsonii Greene. CALIFORNIA: Los Angeles Co.: .7 rd. mi.
N Newcomb Ranch Inn, wash bottom off Hwy., 5500', N of
Chilao Rec. Area, San Gabriel Mts., 1 $ , vi (AM), adult assoc.
( Shields #93 ) . See Map 4.

Ssp. enoptes is on wing from early May to early October.

A southern California form of enoptes enoptes has the broad
black borders reduced that are present in males from the Sierra
Nevada and northern California and is a lighter blue. This form
is found in Kern, Ventura, Los Angeles, and southwest San
Rernardino cos. Males from Lytle Creek possess an uppersurface
like enoptes dammersi but an underside like enoptes enoptes.
A few females from Big Rock Creek, Los Angeles Co., and one
from Lytle Creek, have a slight orange band on the underside
of the primaries ( as in hattoides comstocki ) , one with the lower
half predominantly so (in this respect rather similar to rita
spaldingi ) .

Ssp. enoptes enoptes and hattoides intermedia are sympatric
at Echo Lake and Loon Lake, El Dorado Co.; Donner Summit
and Lake Spalding, Nevada Co.; Gold Lake, Sierra Co.; Onion
Butte, Tehama Co.; and Carrville, Trinity Co., all in California.
A strong tendency in enoptes enoptes to resemble b. intermedia
on the underside is manifest at Chester, Plumas Co.; and Donner
Summit, and vice versa at Lake Spalding and Onion Butte. An
occasional specimen in an enoptes enoptes series from Donner
Summit and Truckee, Nevada Co., tends toward enoptes ancilla
on the underside. Two specimens from Kyle Canyon and Mt.
Charleston, Clark Co., Nevada, are like Sierran enoptes enoptes
on the upperside but are smaller and more heavily marked on
the underside.

Ssp. enoptes enoptes and hattoides hattoides are sympatric
at Mono Pass and Ruby Lake in Inyo Co., Mammoth Lakes and
Tioga Pass in Mono Co., and Gold Lake in Sierra Co. A female
from Ruby Lake has the underside heavily spotted as in hattoides
hattoides but the spots are not rectangular.

At Haines Canyon, Douglas Co., Nevada, enoptes enoptes
flies in May and again in August.

16(l):l-67, 197 7

PHILOTES

23

Map 4. — Distribution of S. enoptes enoptes in California.

24

OAKLEY SHIELDS

J. Res. Lepid.

Eriogonum nudum is widespread from Washington to Cali¬
fornia and Baja California, eastward into extreme western Ne¬
vada. Var. nudum is found from southern Washington southward
along the coast to near San Francisco and in the mountains
from Washington through Oregon to western Nevada and in
California to Tulare Co., flowering from June to September. Var.
pubiflorum occurs from southern Oregon, southward to Mon¬
terey Co., and in eastern California to Kern and Inyo cos.,
flowering from July to October. Var. deductum ranges in the
Sierra Nevada from Lassen Co., California southward to Inyo
Co., and in adjacent extreme western Nevada, flowering from
June to September. Var. pauciflorum is found in the mountains
of southern California from Los Angeles and San Bernardino
cos. southward to northern Baja California to San Pedro Martir,
flowering from August to October. E. wrightii var. subscaposum
grows in the mountains of eastern and southern California and
extreme west-central Nevada, flowering from July to October.
E. saxatile extends from San Jacinto and San Bernardino mts.
west and north to the San Gabriel, Argus, and Panamint mts.,
and the southern Sierra Nevada north to Fresno Co., in the
Santa Lucia Mts., California, and in Nye and Esmeralda cos.,
Nevada, flowering from May to July. E. davidsonii occurs in the
southern Coast Ranges and mountains of southern California
and northern Baja California eastward into western and northern
Arizona, southern Utah, and extreme southwestern Colorado,
flowering from June to September.

Shijimiaeoides enoptes tildeni (Langston, 1963)

Langston. 1963 [1964]. Jour. Lepid. Soc. 1963, vol. 17; pp. 212-
216; f. 3; fig. 5, no. E.

The holotype male was collected at Del Puerto Canyon, 22
mi. W. of Patterson, Stanislaus Co., California, VIII-11-62 (R. L.
Langston). This is at best a weakly defined subspecies closely
allied to enoptes enoptes and enoptes bayensis. If tildeni de¬
serves a name, no doubt other populations within enoptes
enoptes should be so considered, although I prefer to treat
highly variable subspecies like enoptes enoptes, enoptes ancilla,
battoides glaucon, and battoides ellisii as plastic subspecies for
which infra-population names are superfluous. However, tildeni
is not hereby reduced to synonymy, to help emphasize its pos¬
sible evolutionary position between e. bayensis and enoptes

16(1): 1-67, 1977

PHILOTES

25

enoptes. Ssp. tildeni is pictured in black-and-white by Langston,
who also figures a drawing of the male genitalia, along with
some distribution records.

Number of teeth on terminal end (cucullus) of valve = 12,
17, 18 (N = 3), from Del Puerto Canyon.

DISTRIBUTION (viewed 10 $ gen. from 4 localities):
CALIFORNIA: Monterey Co.: 3 mi. SE Parkfield, 1 2 , viii
(CIS). San Benito Co.: New Idria, 2$, viii (PO); hills above
New Idria, 3200-3600', 22, viii (CIS). San Luis Obispo Co.:
Cottonwood Pass, 6 mi. NE Cholame, 5 $ 1 2 , ix ( CIS ) ; Polonio
Pass, Hwy. 46, E of Cholame, 1 $ , ix (AM); 8 mi. W Simmler,
4$ 4 2 , ix (CIS). Stanislaus Co.: mine, Del Puerto Canyon, ca.
22 mi. W Patterson, 24 $ 2 2 , v (AM, GG); 0.3 rd. mi. SSE of
mouth of Hideout Canyon, 1550-1600', Del Puerto Cyn. Rd., sec.
16, T6S, R5E, 5$, vii (AM).

HOSTS: Eriogonum nudum Doug, ex Benth. var. auriculatum
(Benth.) Tracy ex Jeps. CALIFORNIA: Stanislaus Co.: Del
Puerto Cyn., 21-22 mi. W Patterson, 26 <3 92, vii-ix (LACM,
AM, PO, CIS, GG), adult assoc. (/. F. Emmel #144). Eriogonum
covilleanum Eastw. CALIFORNIA: Stanislaus Co.: ca. 2-3 rd.
mi. W of Frank Raines Park, 1300', Del Puerto Canyon, 7 $ 1 2 ,
v (AM), adult assoc. ( Shields #191). Eriogonum parvifolium
Sm. in Rees. var. paynei (Wolf ex Munz) Reveal in Munz. CALI¬
FORNIA: Ventura Co.: 12.1 rd. mi. E of jet. Hwy. 33 & 150,
1000', 6.1 rd. mi. NW of jet. Hwy. 150 & 126 (N of Santa Paula
along Hwy. 150 ) , 17 S 8 2 , vi ( AM ) , adult assoc. ( Shields #96 ) .

Ssp. tildeni is on wing from mid May to mid September.

In Del Puerto Canyon, Stanislaus Co., California, tildeni uses
the annual E. covilleanum in May when E. nudum is just sprout¬
ing new growth for the year, then in July to September another
flight uses E. nudum blooms. Specimens from the different flights
are nearly indistinguishable in appearance, although these may
represent two reproductively isolated, single-brooded popula¬
tions. The population on E. parvifolium var. paynei near Santa
Paula, Ventura Co., is intermediate between tildeni and smithi.

E. parvifolium var. paynei is endemic to Santa Paula Canyon,
Ventura Co., California. E. covilleanum occurs from Alameda
and western Stanislaus cos. southward to Fresno and Kern cos.,
California (endemic), flowering from April to June.
Shijimiaeoides enoptes smithi (Mattoni, 1954)

Mattoni. 1954. Bull. S. Calif. Acad. Sci. 1954, vol. 53, pp. 160-162
& 165, pi. 43, ff. 8-9.

26

OAKLEY SHIELDS

]. Res. Lepid.

The holotype male comes from Bums Creek, State Highway
1, Monterey Co., California, VIII-20-48 (C. I. Smith & R. H. T.
Mattoni). Ssp. smithi is distinguishable from its nearest sub¬
species in geographic proximity, e. bayensis and e. tildeni, by
its distinctive underside. Mattoni (1954) lists five characters
that in combination distinguish smithi from other subspecies.
He thought that its fall flight (actually late summer) and gross
appearance suggest that it is closely related to enoptes dammersi.
However, in average teeth number on the valve it differs from
dammersi and is closer to enoptes tildeni; also there is an inter¬
mediate population between the two near Santa Paula. (Ssp.
dammersi appears to be more closely related to enoptes mojave
and enoptes langstoni than to enoptes enoptes.) Distribution
records for smithi are given in Mattoni (1954) and Langston
(1963). Ssp. smithi is known only from Monterey Co., chiefly
along the coast. Mattoni (1954) figures the upper and lower
surface of a male smithi in black-and-white. Langston (1963),
after making genitalic preparations from various localities with¬
in its range, noted that Mattoni was correct in placing smithi
as a subspecies of enoptes.

Number of teeth on terminal end (cucullus) of valve = 13 (1),
14 (1), 15 (2), 16 (3), 17 (6), 18 (7), 19 (2), 20 (4),
21 (3); N = 29, average = 17-18, all from Seaside.

DISTRIBUTION ( viewed 45 $ gen. from 6 localities ) :
CALIFORNIA: Monterey Co.: 36 mi. S Big Sur, 6 8 4 $ , vii
( GG ) ; Burns Creek, 1 $ , viii ( KH ) ; /2 mi. N Dolans Creek, 1 $ ,
viii (CIS); 3 mi. S Lucia, 150', U 1$, viii (LACM, CIS); 9 mi.
NW Lucia, 1 <$ , viii (CIS); Marina Beach (dunes), 2$, viii
( CIS ) ; Monterey, 10 $ 10 $ , vi-viii ( CAS ) ; Seaside, 18 $ 22 $ ,
vii (CIS, KH, PO).

HOSTS: Eriogonum parvifolium Sm. in Rees var. parvi-
folium. CALIFORNIA: Monterey Co.: dunes at NW end Olym¬
pia Ave., ca. 25', Seaside, 67 3 47$, viii (AM), adult assoc.
( Shields #133). Eriogonum latifolium Sm. in Rees. CALIFOR¬
NIA: Monterey Co.: dunes W of Marina, 32 $ 20 $ , vi (J.
Emmel), oviposition (/. F. Emmel #366), no E. parvifolium in
the immediate area.

Ssp. smithi flies from June to late August.

Bruce Walsh (in litt.) found a smithi colony at Vasquez
Knob, 1600-1800', Carmel Valley, Monterey Co., in association
with E. parvifolium; one oviposition was observed.

16(1 ):l-67, 1977

PHILOTES

27

Shijimiaeoides enoptes bayensis (Langston, 1963)

Langston. 1963 [1964]. Jour. Lepid. Soc. 1963, vol. 17, pp. 208-
212; f. 2; f. 5, nos. c-d.

The holotype male was taken at China Camp, near Point
San Pedro, Marin Co., California, VI-17-61 (R. L. Langston).
Langston described bayensis from 237 specimens at ten locali¬
ties. At best, it is a weakly-defined subspecies closely allied to
enoptes tildeni although both have significantly different aver¬
ages in valve teeth numbers. Langston adequately pictures
bayensis in black-and-white, figures a drawing of the male
genitalia, and gives distribution records. Langston & Comstock
( 1966 ) give a life history description of bayensis , unusually rich
in ecological details, including the fact that the larvae would not
feed on substitute species of Eriogonum furnished them. They
note that the foodplant in Solano and Contra Costa counties is
Eriogonum latifolium ssp. auriculatum (now placed by Reveal
as E. nudum Dougl. ex Benth. var. auriculatum ( Benth. ) Tracey
ex Jeps.). Early records of “Lycaena enoptes ” from the hills
near Mission Dolores, San Bruno Hills, and Lone Mountain in
the San Francisco region (Williams, 1910, p. 39) are perhaps
referable to this subspecies.

Number of teeth on terminal end (cucullus) of valve = 17
(1), 19 (5), 20 (2), 21 (2), 23 (4), 24 (1); N = 15, average =
20-21, all from Pt. Richmond.

DISTRIBUTION (viewed 17 $ gen. from 1 locality):
CALIFORNIA: Contra Costa Co.: Point Richmond, 36 $ 12$,
v, vi, (GG, PO, KH). Humboldt Co.: Fruitland, 7 $ 14 $ , vi
(CIS, CAS). Sonoma Co.: 2 mi. N Camp Meeker, 1 $ , vii (CIS).
(Langston, 1970, lists a record for Benecia State Park, Solano
Co., vi.)

HOST: Eriogonum nudum Dougl. ex Benth. var. nudum.
CALIFORNIA: Sonoma Co.: several miles N & E of Duncan
Mills (fide La Due) ( Shields #185).

Ssp. bayensis flies from mid May to mid July.

Specimens from Fruitland, Humboldt Co., California, deter¬
mined as bayensis by Langston, show some tendency toward
northern California populations of enoptes enoptes. A close
relative of bayensis is no doubt tildeni , which is often similar
on the underside to the bayensis population from Point Rich¬
mond.

Ssp. bayensis is probably closely related to tildeni but does
resemble columbiae in general appearance. In northwestern

28

OAKLEY SHIELDS

J. Res. Lepid.

California there may be some gene flow in facies towards enoptes
enoptes as well. It is noteworthy that smithi and bayensis show
no intergradation.

E. nudum var. nudum is found from southern Washington
southward along the coast to near San Francisco and in the
mountains from Washington through Oregon to western Nevada
and in California to Tulare Co., flowering from June to Sep¬
tember.

Shijimiaeoides enoptes columbiae (Mattoni, 1954)

Mattoni. 1954. Bull. S. Calif. Acad. Sci. 1954, vol. 53, pp. 162-
164 & 165, pi. 43, ff. 7 & 11.

The holotype male comes from the Columbia River near
Brewster, Okanogan Co., Washington, V-5-47 (D. Frechin). As
Mattoni pointed out, it is closely related to enoptes ancilla (al¬
though readily distinguishable from it in most populations ) . Ssp.
columbiae is found in extreme northern Oregon and eastern
Washington, while ancilla occurs in western Colorado to western
Nevada, northward to western Wyoming, extreme western
Montana, and extreme southcentral Idaho. (Mattoni noted that
a subspecies of battoides was sympatric with columbiae and
was nearly indistinguishable until genitalic separations were
made. He said, “This battoides form appears close to P. battoides
glaucon.”) Mattoni (1954) made some genitalic dissections of
the type series and noted that columbiae was a geographic race
of enoptes , distinguishable from other enoptes subspecies by a
combination of six characters. He adequately illustrates the
upper and lower surface of males from the type series, in black-
and-white.

Number of teeth on terminal end (cucullus) of valve = 13
(3), 14 (2), 15 (1), 16 (1), 17 (1); N = 8, average = 13-14, all
from Kusshi Creek, Washington.

DISTRIBUTION (viewed 51 3 gen. from 26 localities):
OREGON: Baker Co.: Snake River Canyon, 8-9 mi. N Hunting-
ton, 4 3 2 $ , v (DB). Jefferson Co.: Metolius River, N of Lower
Bridge Cmp., 1 $ 2 $ , vi (DM). Linn Co.: Santiam Pass, 1$,
vi (KH). Umatilla Co.: Mill Creek nr. Kooskooskie, Wash., 26$ ,
vi (DB). Wasco Co.: 6. 3-6.5 mi. SW The Dalles, 1750-1800',
17$ 7$, v (AM); 7.3 mi. SW The Dalles, 1850', 1$, v (AM);
Juniper Flat, 10 mi. W Maupin, 1890', 2$ , v (AM); 1.7 mi. SW
Rowena, 525', 1$, v (AM). Yamhill Co.: McMinnville, 4 $ , vi
( LACM ) . WASHINGTON: Douglas Co.: Rock Isl. Dam, 4$
3$, v (DM). Kittitas Co.: 3 mi. W Cle Elum, 1$ 1 $ , vii

16(l):l-67, 1977

PHILOTES

29

(CIS); Easton, 1$ (CAS). Klickitat Co.: Kusshi Creek, 32$,
vi (DB); Satus Creek, 5$ 5?, v (DB); Satus Pass, 3200-3500',
4$ 3$ v-vii (CIS, AM); Satus Pass Hwy., 4 mi. N Kusshie
Creek, 2 $ 1 $ , v ( DM ) ; Satus Pass Ski Area, 1 9 , vi ( LACM ) .
Okanogan Co.: Brewster, 4$ 7$, v (LACM, CM, YU); Black
Canyon, nr. Brewster, 2$, v (LACM); Pateros, 2$ 2 $ , vi
(YU). Yakima Co.: Ahtanum Creek, 1800-2200', 10$ 11 $ , v
(CIS, JS, PO, KH); Bear Canyon off White Pass Hwy., 3000-
3200', 10$ 5 2 , v-vii (JS, PO, CIS, AM, DM); Cottonwood
Creek, 2000', 1 $ 1 9 , vi (CIS); Cowiche Crk. Rr., 1 2 , vii (DM);
Harwood, 1 $ , vi (DM); Kusshi Crk., 2000-3000', 3 $ 4 2 , v, vi
(AM); Kussler Crk., 2200', 1 $ , v (CIS); Mill Creek, 1500-1600',
9 $ 2 2 , iv, v (KH, JS, CIS, MF); Oak Creek off White Pass Hwy.,
3000-3500', 3 $ 1 2 , v-vii (DM, PO, CIS); Satus Pass Hwy., ca. 15
mi. S Topp., 1 $ 2 2 , v (DM); N of Satus Pass, 1 2 , v (LACM);
2 mi. W Tieton, 1 $ , vii (DM); White Pass Hwy. 8 mi. E Tieton
Ranger St., 1 $ 1 2 , vii (DM). County undetermined: “E. of G.
Dalles”, 1 $ , v ( CAS ) .

HOST: Eriogonum compositum Dougl. ex Benth. in Lindl.
var. compositum. OREGON: Malheur Co.: Simmons Gulch at
Malheur River, 23 rd. mi. E Juntura, 2$ 2 2, v (AM), adult
assoc. (D. P. Levin, s. n.). See Map 5.

Ssp. Columbiae flies from late April to mid July.

The female of an in copula pair of enoptes taken 8 mi. E of
Tieton Ranger Station, Yakima Co., Washington, is like colum¬
biae but the male is similar to enoptes enoptes. One male enoptes
(Fig. 12, 13), similar to enoptes enoptes on the underside but
otherwise close to columbiae, was taken at Tumwater Canyon,
Chelan Co., Washington, V-29-57 (D. L. Bauer, DB).

Although columbiae is so far known only from one location
on E. compositum, the closely related ancilla uses E. heracleo-
ides, and battoides glaucon in the vicinity of Brewster, Wash¬
ington, an area of sympatry with columbiae, uses E. heracleo-
ides var. angustifolium. The range of var. angustifolium cor¬
responds to the range of columbiae and thus may be its principal
foodplant.

E. compositum var. compositum is found in southern Wash¬
ington and western Washington from Mt. Rainier southward to
California as far as Lake Co., flowering from April to July.
E. heracleoides var. angustifolium ranges along the Columbia
River drainage in northern Oregon northward to northeastern
Washington and southern British Columbia and eastward into
central Idaho where it is local and sporadic, flowering from

30

OAKLEY SHIELDS

J. Res. Lepid.

Map 5. — Distribution of S. enoptes columbiae.

16(l):l-67 , 1977

PHILOTES

31

May to July.

Shijimiaeoides battoides glaucon (Edwards, 1871)

(See discussion of spaldingi, in Part IV)

DISTRIBUTION (viewed 110 $ gen. from 72 localities):
CALIFORNIA: Alpine Co.: Carson Pass, 8573', 1 $ , vi (CIS).
El Dorado Co.: Meyers, 6300', 2 2 , vi (DB); Mt. Tallac, 1$,
vii (CIS). Inyo Co.: Andrews Camp, Bishop Creek, 3$ 6 $ , vi
( LACM ) ; Bishop Creek, 3 $ 3 $ , v, vi ( LACM ) ; Lone Pine
Canyon, 1 $ , vii (LACM); Mahogony Flat, 7500-8000', Panamint
Mts., 1 $ 1 $ , vi (DB). Modoc Co.: Buck Creek, 1$, vi

(LACM); Fandango Pass, 6100', Hist. Mon. #546, Warner Mts.,
1 $ , vi (CIS). Mono Co.: Bodie, 2$ , vii (CM); Convict Lake,
1 $ , vi (AM); Cottonwood Canyon, 7000', S of Brodie, 5$ 32
+ “10”, vi (LACM, CIS); Grant Lake, 7 air mi. S Lee Vining,

4 2 , vi (CIS); Grant Lake, 7800', 5$ 2 2 , vi, vii (AM); just W
of Grant Lake, 32, vi (JL); 2.5 mi. above Grant Lake, 7800',
2$ 4 2 , vii (AM ) ; June Lake, 1 $ , vi (AM ) ; Lower Rock Creek,
5500', Swall Mead, 1 $ 2 2, v (DB); Lundy, 1$, vii (CM);
Mammoth, 4 $ 1 2 , vi, vii (LACM); 4 mi. E Monitor Pass, 2 $ ,
vi (CIS); Mono Lake, 1# 2 2 , vii (CM, LACM); W shore,
Mono Lake, 12$ 1 2 , vi (LACM, AM); Rush Creek, 1 $ , vi
(LACM); Silver Lake, 1 $ , vi (LACM); l/2 mi. SE Tom’s
Place, 48$ 6 2 , vi (LACM, CIS); Warren Creek, 1$, vii
(LACM). Placer Co.: Ward Peak, 8500', 12, vii (LACM).
Shasta Co.: Hat Creek — 3N, 1 $ , vi (CIS); Old Station, 1 2 ,
vi (CIS). Siskiyou Co.: Mt. Shasta, 7600', 12, viii (AM);
Pumice Stone Well, 1$, vii (LACM). County undetermined:
Glen Alpine, Tahoe, 1 2 , vi (CAS). IDAHO: Ada Co.: Boise,
4500-7500', 8 $ 12 2 , vii (MCZ). Butte Co.: nr. Craters of Moon
Nat. Mon., 7$ 4 2 , vii (JS); Craters of the Moon Nat. Mon.,
4$ , vi (UI). MONTANA: Sula, 4700', 1 $ , vi (UI). NEVADA:
Douglas Co.: Carson Valley, 2$ 5 2 , vi (NSM). Ormsby Co.:
Carson Range, W Stewart, 1 $ 1 2 , vi (DB); Eagle Valley, 9$

5 2 , v, vi (NSM); Graphite Cyn., 1 mi. W Carson City, 9 $ 9 2,
vi (GG); Graphite Cyn., ca. 2 mi. S Carson City, 7$ 5 2 , vi
(JS, PO); Kingston Cyn., 5500', 2$ 8 2 , vi (PO); Lakeview
Hill, 12$ 6 2 , vi (LACM); Voitaire Cyn., 5200', 5$ 6 2 , vi
(DB). Storey Co.: Ophir Grade, 11$ 52, vi (NSM); Virginia
City, 7 mi. Canyon, 5$ 2 2 , v (DB). Washoe Co.: Nellie Spgs.,
S. R. 34, 5$ 2 2, v (NSM); Reno, 1$ 1 2 , vii (CM); Reno,

32

OAKLEY SHIELDS

J. Res. Lepid.

N City Limit 395, 2$ 3 $ , v (DB); 2 mi. N Reno, 5 5 , vi (GG,
PO). Nevada-Idaho border: N of Jarbidge on Nev.-Ida. border,
2$, vi (CIS). OREGON: Baker Co.: Baker, 2$, vii (CM);
Durkee, 2$ , viii (CM); along rd. between Joseph & Hwy. 86,
13.2 rd. mi. N of jet. with Hwy. 86, Duck Creek, at turnoff to
Fish Lake, 2 $, vii (AM). Clackamas Co.: Mt. Hood, 1 <2 2$,
vii (MCZ). Gilliam Co.: 4.5 mi. S Condon, 2500', 3$ 7$,
vi (AM); 5 mi. NE Lonerock, 3 $ , vi (DB). Grant Co.: 1 mi.
S Senaca along Silvies River, 5000', 1 $ , vii (CIS). Harney Co.:
V2 mi. W Burns P.O., 4100', 1$, v (CIS); Cricket Creek, 13
air mi. NW Burns, 4900', 1 $ , vii ( CIS ) ; Devine Canyon, 12
mi. N Burns, 5000', 1 $ , vii ( CIS ) ; Roaring Springs, 4600', 1 $ ,
vi (CIS). Jackson Co.: Mt. Ashland Loop Rd., 1$, vii (CIS).
Klamath Co.: Bly Mt., 5000', 2 $ , vi (CIS); 1.6 mi. E Hwy.
97, 9 mi. N Kirk, 4650', 1 $ , vii (AM). Lake Co.: Drake Peak,
Warner Mts., 1$, vii (AM). Morrow Co.: 2-4 mi. S Hardman,
1$ 1 $ , vi (DB). Umatilla Co.: 12 mi N Ukiah, Carney Bu.
L. O. rd. & Hwy. 395, 14 $ , vi (DM). Wasco Co.: 11.2 mi. SW
The Dalles, 1750', 6^ 8$, v (AM); Juniper Flat, 10 mi. W
Maupin, 1890', 24 $ 21 $ , v, vi ( AM); 2 mi. NNW Mill Crk.
bridge, Warm Springs Indian Res., 2500', 1# 2 $ , vi (AM);

2 mi. SE Tygh Valley, Hwy. 197, 1400', 16$ 28$, vi (AM).
Wheeler Co.: Mountain Creek, 6 mi. E Mitchell, 2$ 1 $ , vii
(AM). WASHINGTON: Adams Co.: 5 mi. SW Sprague, 1$,
vi (PO). Chelan Co.: Cashmere, 3$ 3$, v (DB); nr. Drury,
2$, vi (DB); First Crator, 1$, vi (CM); Leavenworth,
l£, v (DB); Tumwater Canyon, 1$, v (DB). Grant Co.:
Beverly, 3$ 1$, v (DM). Kittitas Co.: Easton, 3$ (CAS);

3 mi. W Vantage, 2$, v (DM); Wenas Crk. Rd., 10 mi. SW
Ellensberg, 3$ , v (DM); Wymer, 2$ , v (DB). Okanogan Co.:
Alta Lake, 1 $ , vi (AM); Alta Lake St. Pk., 1 $ 2 $ , v (DM);
Brewster, 20 $ 11 $ , v, vi (YU, PO, AM, LACM); Pateros, 2$
1$, v (CM, CAS); Similkameen River, 1 <3 1 $ , vi (CIS).
Yakima Co.: Satus Pass Hwy., ca. 15 mi. S Topp, 2$ , v (DM);
Wenas Crk., 2400', 1 $ , vi (PO). County undetermined: Priest
Rapids, 400', 2$, vi (LACM). BRITISH COLUMBIA: Garnett
Valley, Summerland, 3 $ , vi (LACM); Oliver, 1 $ , vi (LACM);
2 mi. N Princeton, 4$ 4 $ , vi (KH, JL, LACM); Shingle Cr.
Road, Keremeos, 2 $ , vi (LACM).

HOSTS: Eriogonum umbellatum Torr. var. nevadense Gand.
CALIFORNIA: Mono Co.: Cottonwood Canyon, 8200-8300',
2 mi. S Bodie, 4 $ , vii ( AM ) , oviposition ( Shields #6 ) ; peak
slope ca. 1 mi. W Hwy. 395 on Hwy. 89, SW of Topaz Lake,

16(l):l-67, 1977

PHILOTES

33

PLATE 1

Fig. 1. — Shijimiaeoides rita, unnamed ssp., $ upperside, Uinta Canyon,
Duchesne Co., Utah, VIII-4-59 (J. C. Downey).

Fig. 2. — Same, underside.

Fig. 3. — S. enoptes nr. ancilla, $ upperside, Mt. Charleston at 7200', Clark
Co., Nev., VI-26-59 (J. C. Downey).

Fig. 4. — Same, underside.

Fig. 5. — S. enoptes nr. enoptes, $ upperside, Kyle Canyon, 7100', Mt.
Charleston, Clark Co., Nev., VII-14-66 (P. H. Amaud, Jr., CAS).

Fig. 6. — Same, underside.

Fig. 7. — S. enoptes nr. enoptes, So. Cal. population, $ upperside, Black¬
burn Canyon, Tehachapi Mts., Kern Co., Calif., VII-10-64 (N. La Due).
Fig. 8. — Same, underside.

Fig. 9. — Same, 9 upperside, So. Fk. Campgrd., So. Fork Big Rock Crk.,
San Gabriel Mts., Los Angeles Co., Calif., IX-9-66 (J. Lane).

Fig. 10. — Same, underside.

Fig. 11. — S. enoptes, unnamed ssp., $ underside, Hemet Ridge, 6000',
San Jacinto Mts., Riverside Co., Calif., V-24-08 (F. Grinnell, Jr., LACM).
Fig. 12. — S. enoptes nr. columbiae, with underside similar to enoptes enop¬
tes, $ upperside, Tumwater Canyon, Chelan Co., Wash., V-29-57 (D. L.
Bauer ) .

Fig. 13. — Same, underside.

Fig. 14. — S. battoides nr. glaucon, $ underside, Uncompahgre Plateau,
8400', near 25 Mesa Ranger Station, Montrose Co., Colo., VI-22-62 (K. B.
Tidwell ) .

Fig. 15. — S. battoides nr. glaucon (?), $ underside, Telescope Mt., Inyo
Co., Calif., VII-9-37 (LACM).

PLATE 2

Fig. 16. — Same, 9 underside.

Fig. 17. — S. battoides near ellisii, $ underside, Kingston Canyon, Toiyabe
Range, Lander Co., Nev., VII-22, 23-64 (J. Lane, LACM).

Fig. 18. — S. battoides unnamed ssp., $ upperside, Hwy. 89A, 8 mi. ESE
of Fredonia, Coconino Co., Ariz., V-18-68 (K. Roever & R. Funk).

Fig. 19. — Same underside.

Fig. 20. — Same, $ upperside.

Fig. 21. — Same, underside.

Fig. 22. — S. battoides glaucon X enoptes ancilla hybrid by genitalia, $
upperside, Pattee Canyon, 3500', Missoula Co., Montana, VI-23-62 (J.
Scott ) .

Fig. 23. — Same, underside.

Fig. 24. — S. enoptes ancilla, $ gynandromorph, Sybille Canyon, 6500',
Albany Co., Wyo., VII-4-66 (R. Hardesty).

Fig. 25. — S. enoptes dammersi, 9 underside, unusual aberrant, Hualapai
Mts., Mojave Co., Ariz., IX-4-58 (K. Hughes).

Fig. 26. — S. battoides nr. allyni, $ somewhat aberrant, Cedros Island,
Mexico, III-18-39 (CAS).

Fig. 27. — S. battoides nr. glaucon, 9 aberration, Mt. Hood, Ore., VII-20-
37 (CM).

Fig. 28. — S. battoides glaucon, 9 aurora aberrant, 2 mi. NNW Mill Cr.
bridge, Warm Spgs. Indian Res., 2500', Wasco Co., Ore., VI-18-66 (B. &
S. Perkins, AM).

Fig. 29. — Habitat of S. rita pallescens 8 mi. S of Currant, W. slope Grant
Range, Nye Co., Nev., with Eriogonum kearneyi var. kearneyi in foreground.
Fig. 30. — Habitat of S. battoides martini 3 mi. SW Beatty, Nye Co., Nev.,
with Eriogonum fasciculatum var. polifolium in foreground.

34

OAKLEY SHIELDS

Plate 1

16(1 ):l-67, 1977

PHILOTES

35

Plate 2

36

OAKLEY SHIELDS

/. Res. Lepid.

vi (AM), adult assoc. ( Shields #197); Warren Creek, 9000',

vii (OS), adult assoc. ( Shields #250). Eriogonum umbellatum
Torr. var. chlorothamnus Reveal. CALIFORNIA: Mono Co.:
1.6 rd. mi. N Mono-Inyo Co. line, along Hwy. 395, ca. 6 rd.
mi. SE Sherwin Summit, ca. 5700', 1$, v (AM), adult assoc.
( Shields #82). Eriogonum umbellatum Torr. var. stellatum
(Benth.) M. E. Jones. OREGON: Grant Co.: 5.0 rd. mi. WSW
jet. Rd. 162 & Rd. 1427, along Rd. 162, T16S, R33l/2E, E of
Seneca, 14, vii (AM), adult assoc. ( Shields #112). Wallowa
Co.: along rd. between Joseph & Hwy. 86, 17.8 rd. mi. N jet.
with Hwy. 86, 1.2 rd. mi. SW Wallowa Mtn. summit, 1 4 1 $ ,
vii (AM), adult assoc. ( Shields #116). Eriogonum heracleoides
Nutt. var. heracleoides. IDAHO: Adams Co.: Seven Devils
Campground Rd., 5 mi. W Hwy. 95, Payette Nat’l For., 4500',
14 2$, vii (AM), adult assoc. ( Ellis #2000). OREGON:
Grant Co.: Summit Prairie, at jet. of Rd. 162 & Rd. 1427, T16S,
R34E, Malheur Nat’l For., SE of Prairie City, 14, vii (AM),
oviposition ( Shields #111). Eriogonum heracleoides Nutt. var.
angustifolium (Nutt.) Torr. & Gray. WASHINGTON: Okanogan
Co.: 4 mi. E Brewster, 10 4 19 9 , v (AM), adult assoc. (J. A.
Justice, s. n. ). Eriogonum sphaerocephalum Dougl. ex. Benth.
var. halimioides (Gand. ) S. Stokes. CALIFORNIA: Modoc Co.:
3.0 rd. mi. NE of jet. Hwy. 395 & 299, along Hwy. 299, Warner
Mts., “10”, vii (AM), oviposition ( Shields #103). Eriogonum
flavum Nutt, in Fras. var. piperi (Greene) M. E. Jones. IDAHO:
Adams Co.: Summit of Heavens Gate, 7 Devils Range, 8407',
Payette Nat’l For., 1$, vii (AM), adult assoc. ( Ellis #2001).
See Map 6.

Ssp. glaucon is on wing from early May to mid August. Popu¬
lations are variable, and distinct segregates may exist.

Ssp. battoides glaucon and enoptes columbiae are sympatric
at Juniper Flat, Wasco Co., Oregon; Tumwater Cyn., Chelan
Co., and Brewster, Okanogan Co., Washington; and have simi¬
lar facies at Pateros, Okanogan Co., Washington. One male near
Kirk, Klamath Co., Oregon, has an upperside as dark as battoides
oregonensis but an underside like glaucon. A female from Fan¬
dango Pass, Modoc Co., California, is intermediate between
glaucon and intermedia; enoptes enoptes is sympatric there.
A ssp. near glaucon is sympatric with enoptes langstoni near
Sherwin Summit, Mono Co., California. Specimens of glaucon
from Mt. Ashland, Jackson Co., Oregon, are atypical.

A singleton male (gen. det. ) from Uncompahgre Plateau,
8400', near 25 Mesa Ranger Station, Montrose Co., Colorado,

16(l):l-67, 1977

PHILOTES

37

Map 6. — Distribution of S. battoides glaucon,

38

OAKLEY SHIELDS

J. Res. Lepid.

VI-22-62 (KT) is unusual in that it is closer to glaucon than
to centralis or ellisii in facies and represents a considerable east¬
ward range extension (See Fig. 14). However, the specimen is
not typical glaucon.

Female glaucon from Bishop Creek, Inyo Co., and Grant
Lake, Mono Co., have some basal blue on the dorsal surface,
lacking in other glaucon populations.

Eriogonum umbellatum var. stellatum extends from south¬
eastern Washington and northern Idaho into northern Utah
southward to Del Norte, Siskiyou, and Trinity cos., California
and along the western flank of the Sierra Nevada to Kern Co.,
flowering from July to September. Var. chlorothamnus is known
from southern Mono Co. s. along the e. side of the Sierra
Nevada to s. Inyo Co., California, flowering from late June to
September. E. heracleoides var. angustifolium ranges along the
Columbia River drainage in northern Oregon northward to
northeastern Washington and southern British Columbia and
eastward into central Idaho where it is local and sporadic,
flowering from May to July. E. sphaerocaphalum var. halimioides
is found in central and eastern Oregon south to northern Cali¬
fornia and northwestern Nevada, then eastward into south¬
western Idaho, flowering from May to July. E. flavum var. piperi
occurs mainly (but not always) west of the Continental Divide,
from south-western Alberta, western Montana, and northwestern
Wyoming westward across northern and central Idaho to north¬
eastern Oregon and eastern Washington and southern British
Columbia, flowering from June to August.

Shijimiaeoides battoides battoides (Behr, 1867)

(See discussion in Part IV)

DISTRIBUTION (viewed 28 $ gen. from 16 localities):
CALIFORNIA: Fresno Co.: Bullfrog Lake, 10,634', 1$, viii
(LACM); Cartridge Creek Basin, 1$, vii (CIS); Huntington
Lake, 4$ 3 $ , vi, vii ( LACM ) ; Kaiser Crest, 14 $ 3 $ , vi, vii
(AM, LACM, KH); Kaiser Crest, 10,000', 7 S 9 $ , vi, vii (KH);
Kaiser Pass, 9800-10,100', 5 S 2 $ , vii (AM, YU); South Fork,
Cartridge Creek, 1 <$ , vii (CIS); Windy Ridge, 1$, vii (CIS).
Inyo Co.: Lone Pine Lake, 8500', 4 $ , vi (CIS); vie. Mono Pass,
11,500-12,500', 17 $ 27$ + “4”, viii, ix (PO, CIS, CAS, MCZ,
JS, AM); above Ruby Lake, 2$ 2$, viii, ix (CIS); Whitney
Trail, 9000', 2 $ , vii (CIS). Madera Co.: Minaret Ridge, 9265'
1$ 4$, viii (AM). Mono Co.: Bud Lake, 1 S, vii (LACM);

16(l):l-67, 1977

PHILOTES

39

W end Dana Plateau, NE Tioga Pass, 11,000-11,400', 1 8 , viii
(AM); McLeod Lake, 1$, vii (AM); Mammoth, 10$ 3 $ , vii
(LACM); Mammoth Camp, 7 8 3 $ , vii (LACM); Mammoth
Crest, 13 $ 10 $ , viii ( CIS, CAS ) ; Mammoth Lakes, 3 8 1 $
+ "10”, vii, viii (LACM, CIS, CAS, YU); Minaret Summit,
Mammoth Lakes, 7 8 5 $ , vii (CIS, LACM, AM); Pilot Knob
nr. Tioga Pass, 1$, viii (JL): Rock Creek, 2 8, viii (LACM);
Tioga Pass, 2 8 29, vii, viii (CAS, CIS, LACM); 4.9 mi. E
Tioga Pass, 9400', 1?, vii (AM); E end Tioga Pass, 10,000',
1$, vii (AM); W above Tioga Pass, 6$ 8$, vi-viii (LACM,
CIS); Virginia Lakes, 1 8 , vii- (LACM); Walker River (error ?),
48, vii (LACM). Sierra Co.: Gold Lake, 2 8 4 $ , vii (CM).
Tulare Co.: Mineral King, 10,500-11,000', 4 8 4 $ , vii (LACM,
CAS, CM); Monarch Lake, 10,000', 1$, vii (CM). Tuolumne
Co.: Cockscomb Pk., Yosemite Nat’l Park, 2 8 2 8 , vii (LACM);
Helen Lake, Yosemite, 1$, vii (LACM); May Lake trail, Yo¬
semite, 1 $ , vii ( PO ) ; Mt. Hoffman, Yosemite, 1 $ , vii ( LACM ) ;
Sonora Peak, 1$, viii (CIS); Tuolumne Meadows, 1$ 2$,
vii (LACM); N & NW above Upper Gaylor Lake, 10,600-
11,000', NW Tioga Pass, 1$ 1$, viii (AM); Vogelsang Pass,
1$, viii (LACM). County undetermined: boundary Hill, Yo¬
semite, 1 $ , vii ( CIS ) . Some additional records appear in Erik-
sen (1962). In mid- August 1963 I took it commonly in French
Canyon (Star Lake, Steelhead Lake, 11,700-11,800') and near
Wahoo Lakes, 11,500', Fresno Co.

HOSTS: Eriogonum lobbii Torr. & Gray var. lobbii. CALI¬
FORNIA: Inyo Co.: trail between Mono Pass & Ruby Lake,
11,500', viii, 3 larvae in flowerheads (/. F. Emmel #109, det. by
D. Breedlove). Eriogonum incanum Torr. & Gray. In the Yo¬
semite region, reported flying over this species (Garth, 1935);
CALIFORNIA: Mono Co.: .2 mi. W Mammoth Mtn., 9000', 1 8 ,
vii (OS), adult assoc. ( Shields #249); crest W. above Tioga
Pass, ca. 10,800', vii (OS) adult assoc. ( Shields #252). Eriogonum
polypodum Small. CALIFORNIA: Inyo-Fresno Co. line: Mono
Pass, 12,100', viii, ca. 20 3rd-4th instar larvae in flowerheads
(/. F. Emmel #108, det. by D. Breedlove). See Map 7.

Ssp. battoides is on wing from mid June to September.

Specimens from Kaiser Crest, Fresno Co., are not as boldly
square-spotted as they are, e.g., at Mammoth or Tioga Pass, but
are closer to battoides battoides than to battoides glaucon. At
Gold Lake, Sierra Co., battoides battoides and battoides inter¬
media are sympatric, with no evidence of intergradation. This
ssp. is often abundant at high elevations in the Sierra Nevada.

40

OAKLEY SHIELDS

J. Res. Lepid.

Map 7. — Distribution of S. battoides hattoides.

16(l):l-67, 1977

PHILOTES

41

E. lobbii var. lobbii is widespread in the Coast Ranges from
Lake Co. to Siskiyou and Humboldt cos., and in the Sierra
Nevada from Plumas Co. southward to Mariposa and Inyo cos.,
California, and on Mt. Rose and Slide Mountains, Washoe Co.,
Nevada, flowering from June to August. E. incanum ranges in
the Sierra Nevada from Tulare and Inyo cos. northward to
Tuolumne and Alpine cos., and infrequently to Nevada Co.,
California, and isolated on Ashland Peak, Jackson Co., Oregon.
E. polypodum occurs in the southern Sierra Nevada in Tulare,
Inyo, and Fresno cos., California (endemic), flowering from
July to September.

Shijimiaeoides battoides intermedia (Rarnes & McDunnough,

1917)

Rarnes & McDunnough. 1917. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, p. 214, pi. 16, ff. 4-6, p. 262.

This subspecies was originally described from “Shasta Co.,
Calif, (presumably the vicinity of Castella in the Upper Sacra¬
mento Valley) captured in the third week in July.” Barnes &
McDunnough (1917) regarded intermedia as so similar to
enoptes enoptes in maculation that the two would be inseparable
were it not for their striking difference in genitalia. They de¬
scribe intermedia as having the spots on the underside of the
secondaries greatly reduced in size, the aurora broken ( ± ) into
lunules, the terminal dark line reduced to a fine hair-line in
width, the black border moderately broad in males, and the
fringes of the secondaries white and uncheckered. They picture
the “types” adequately in black-and-white. Langston (1969)
refers to intermedia specimens from Kern and Los Angeles
counties taken 30 to 60 years ago. These are probably the newly
described battoides comstocki from Tehachapi, Kern Co., and
the typical intermedia (possibly erroneous) from Mt. Baldy,
Los Angeles County.

DISTRIBUTION (viewed 42$ gen. from 14 localities):
CALIFORNIA: Del Norte Co.: Little Grayback, NE Del Norte
Co., 4600', 6$ , vii (CIS); Little Grayback Pass, 21 $ , vii (CIS).
El Dorado Co.: Angora Peak, Tahoe, 1 9 , vii (CIS); Echo Lake,
2$, vii (JS, CIS); N ridge of Echo Lake, 7400-7700', 1$, vii
(LACM); Fallen Leaf Lake, 1$ 1 $ , vi, vii (CAS, LACM);
Tamarak L., 2$, vii (CIS). Nevada Co.: Bear River nr. Fuller
L., 6$ 3$, vii (CIS); Donner Pass, 1$ 19, vii, viii (CIS);
entrance to Lake Spalding, 10 $ 7 $ , vii (AM, CIS). Placer Co.:

42

OAKLEY SHIELDS

J. Res. Lepid.

Cisco, 1$, vii (DB). Plumas-Sierra Co. line: Gold Lake, 13 <2
19 9, vii (KH, AM, LACM, CM). Sierra Co.: Salmon Lake
Lodge, S Gold Lake, 1$, vii (CIS). Siskiyou Co.: Caribou L.
& N slope & summit Sawtooth Ridge, Trinity Alps, 6800-7900',
3$, 4$, vii (AM); Castle Lake, 79$ 42, vi-viii (CIS, CAS,
GG, AM, LACM, DB, PO); Castle L., 5600-6000', 7$ 3 2 , vii
(AM); S shore Castle L., 3 $ 22, viii (AM); along E side
Castle L., 3 $ 22, vii (AM); N slope of peak S of Castle L.,
5600-6400', 1 $ 3 2 , vii ( AM ) ; Klamath Lodge, Doggett Creek,
1$, vi (DB); Little Castle Lake, 5300', 22, vii (CIS, GG);
Mt. Shasta, 3$, vii (DB); Mt. Shasta, 7600', 2$, viii (AM);
Trinity Alps, Caribou Mt., 6300-7000', 1$, vii (CIS). Tehama
Co.: rocky outcropping on Lassen Trail Rd., 5300', T27N, R3E,
Sll, nr. Onion Butte, 10$ 4 9 , vi (SM). Trinity Co.: Carrville,
2$ 1 9 , vi (CAS, CIS). Erroneous record (?): Mt. Baldy, 1$
1 2 (CAS). OREGON: Morrow Co.: 5 $ , vii (NSM).

HOST: Eriogonum marifolium Torr. & Gray. CALIFORNIA:
El Dorado Co.: along W & SW shore Loon Lake, ca. 15 air mi.
N Kyburz, vii (AM), adult assoc. (Shields #232). Siskiyou Co.:
along E side Castle Lake, SW of Mt. Shasta (town), 3$ 22,
vii (AM), adult assoc. ( Shields #100). Eriogonum incanum
Torr. & Gray. CALIFORNIA: El Dorado Co.: NW ridge along
NE end of Echo Lake, 7500', vii (OS), adult assoc. (Shields
#247). See Map 8.

Ssp. intermedia flies from early June to early August.

Ssp. intermedia in facies sometimes resembles enoptes
enoptes (e.g., Lake Spalding, Gold Lake, Castle Lake, and Onion
Butte) and on the underside it sometimes resembles Plehejus
lupini (e.g., Echo Lake and Loon Lake). Both P. lupini and
ssp. intermedia fly over the same E. marifolium flowers at Loon
Lake, Echo Lake, and Donner Summit. Thus there is perhaps
predation pressure selecting for convergence in undersurface
pattern. Both intermedia and oregonensis use E. marifolium and
are thus perhaps closely related; indeed, both show some affinity
to glaucon, their probable common ancestor. Material from
Ft. Klamath, Oregon, is intermediate among glaucon, intermedia,
and oregonensis. Eriogonum jamesii, used by centralis, is placed
in a section next to E. marifolium; both are the most primitive
members of their respective sections. Because of this affinity,
there may be a relationship between centralis and the intermedia-
oregonensis group. Both centralis and intermedia on the upper-
surface have extensive black borders, and populations from
Burns Crk. and Cedar Pass in Modoc Co., California, are nearly

16(1): 1-67, 1977

PHILOTES

43

Map 8. — Distribution of S. battoides intermedia.

44

OAKLEY SHIELDS

J. Res. Lepid.

indistinguishable from centralis. ( E . jamesii does not extend to
California. )

E. marifolium occurs in the Sierra Nevada from Tuolumne
Co. northward to Siskiyou and Modoc cos., California, eastward
to the Pine Forest Range of Humboldt Co., Nevada, and north¬
ward to central and east-central Oregon, flowering from June
to August.

Shijimiaeoides battoides oregonensis (Barnes & McDunnough,
1917)

Barnes & McDunnough. 1917. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, p. 214, pi. 16, ff. 1-3, p. 262.

Barnes and McDunnough originally described oregonensis as
a subspecies of battoides from Crater Lake, Klamath Co., Ore¬
gon, from a long series taken during the last two weeks of July.
They merely point out that oregonensis compared to battoides
battoides has a broad black border in males, undersurface spots
less quadrate and the ground color a “greener hue”, and say
that the figures (1-3) on pi. 16 should provide the identification
of this subspecies, when compared to battoides battoides. Topo-
typical material I have seen from Crater Lake fits the descrip¬
tion and figures, although the underside has a dusky appearance.
This is probably a weakly defined subspecies close to glaucon
and perhaps intermedia (see discussions of glaucon and inter¬
media), confined to Klamath Co., Oregon. Langston (1969,
p. 55) thought that oregonensis was a high elevation subspecies
in southern Oregon that decreased in altitude with latitude in
Washington and British Columbia. However, material I have
examined from outside Klamath Co. is glaucon (except for one
peculiar population from Mt. Hood; see fig. 27).

DISTRIBUTION (viewed 4$ gen. from 4 different lo¬
calities): OREGON: Klamath Co.: 2 mi. W of jet. 230 & 97,
1 $ , vii (JS); Crater Lake, 4$, vii, viii (LACM, CM); E Rim
Crater Lake, 1 $ , vii ( AM ) ; S entrance Crater Lake Nat’l Park,
1 $ , vi (CAS); 2 mi. W Diamond Lake Hwy. & US 97 Jet., 1 $ ,

vi (CIS); Hillman Peak, 7500', Crater Lake Nat’l Park, 1 S
1$, vii (AM); 1.6 mi. E Hwy. 97, 9 mi. N Kirk, 4650', 12,

vii (AM); Miller Creek, 5000', 1$ 1 9 , vi (CIS).

HOST: Eriogonum marifolium Torr. & Gray. OREGON:
Klamath Co.: ca. 5 air mi. NE of Chemult, along rd. to Walker
Mtn. Lookout, Walker Rim, “3”, vii (AM), adult assoc. ( Shields
#107); .7 mi. SE of jet. Crater Lake N.P. boundary (S) and

16(1): 1-67, 1977

PHILOTES

45

Hwy. 62, on Hwy. 62, vii (AM), oviposition ( Shields #223);
5 rd. mi. NE Sun Pass, on Hwy 232, vii (AM), adult assoc.
( Shields #225); 6 rd. mi. NE Sun Pass, on Hwy. 232, “11”, vii
(AM), oviposition ( Shields #227).

Ssp. oregonensis flies from late June to early August.
Shijimiaeoides battoides centralis (Barnes & McDunnough, 1917)
Barnes & McDunnough. 1917. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, pp. 214-215, pi. 16, ff. 7-8, p. 262.

The “types” of centralis were taken at Salida, Chaffee Co.,
Colorado, July 1-7. Barnes and McDunnough (1917) say that
centralis is distinguished by the broad black borders on the
upperside in males, a prominent orange shading at the anal
angle of the secondaries, a brown underside with spots larger
than intermedia and well ringed with white, a continuous aurora,
and primaries subterminally with a diffuse smokey shade above
the outer angle. It is noteworthy that they felt centralis is very
similar to intermedia otherwise (see discussion under inter¬
media). The ssp. centralis male “type” from Salida is clearly
illustrated in black-and-white. Scott, Ellis and Eff (1968) report
the foodplant of centralis as Eriogonum umbellatum, but I have
found it only in association with E. jamesii.

DISTRIBUTION (viewed 74$ gen. from 41 localities):
CALIFORNIA: Modoc Co.: nr. Burns Creek Ranger Station,
3$, vii (LACM); Cedar Pass, 1$, vi (CIS). COLORADO:
Alamosa Co.: Great Sand Dunes Nat’l Mon., 2$, vii (AM).
Chaffee Co.: Maysville, 6$, vii (CIS, JS, YU); Poncha Creek,
2 $ 2 9 , vii (LACM, JS). Conejos Co.: Alamosa Reservoir, 7500',
2$ 19, viii (MF); l/2 mi. N of E end of Alamosa Res., 34 $
19 9, vii (JS); W end Alamosa Res., 49 $ 1 9 , vii (GG, JS);
Gato Creek, 8250' 2 $ 2 9 , vii (JS); Schilling’s Spring, 43$ , vii
(JS); Z2 mi. SE Schilling’s Spring, 85$ 229, vii (JS). Custer
Co.: Ben West Hill, 2$ 1 9 , vii (LACM); Hermit Lake Rd.,
9000', W Westcliffe, 2 $ , vii ( KH ) ; Johnson Gulch, 7700' 3 $ 2 9 ,
vii (JS); Zi mi. W Rosita, 2$ 1 9 , vii (JS); Sand Gulch, 1 mi.
SW Greenwood, 3 $ , vii (JS); 2 mi. N Silver Cliff, 1 $ 1 9 , vii
(JS); Welsh Pony Ranch, W of Hardscrabble Creek, 1$, vii
(JS); Fremont Co.: Box Canyon, 4$ 1 9 , vii (JS); 1 mi. NE
Calcite, 14$ 4 9 , vii (JS, SE, LACM); nr. Cotopaxi, 19, viii
(MF); 3 mi. SW Cotopaxi, 2 9 , viii (JS); 1 mi. W Democratic
Mtn., 1$, viii (JS); Falls Gulch, 6$, vii (LACM, JS); South
Fork Falls Gulch, 2$, viii (JS); Fleming Mtn., 1$, vii (JS);
Hayden Creek Campground, 1$, viii (JS); Zi mi. E. Hillside
Cemetery, 1 $ , viii (JS); 2 mi. NE Hillside, 1 $ , viii (JS); Kiln

46

OAKLEY SHIELDS

/. Res. Lepid.

Gulch, 4$ 1 $ , vii (LACM, JS); Kuntz Gulch, 1 $ , viii (AM);
Oak Creek, nr. Cotopaxi, 6500-7000', 2$ 1 $ , vii (MF); Rouch
Gulch, 1$, vii (JS); Spring Creek, 2 $ 1$, viii (JS); 1 mi. S
Wellsville, 1$, vii (JS). La Plata Co.: 1 mi. S Durango, 7000',
3# 2 $ , vi (SE). Montezuma Co.: Morefield Cyn., Mesa Verde
Nat’l Park, 1 $ , vii (AM; Wetherill Mesa, Mesa Verde Nat’l Park,
1 $ , vii (AM). Rio Grande Co.: “D” Hill, Del Norte, 32$ 4$ ,
vi-viii (SE, MF). Saguache Co.: Cotton Creek, 9000', 11$, viii
(SE); North Crestone Campground, 2$ , viii (SE); Orient Can¬
yon, ESE Villa Grove, 2$, vii (JS); ca. 5 mi. W Villa Grove,
1$, vii (JS); 6 mi. W Villa Grove, 11$ 1 $ , vii (JS). San
Miguel Co.: Hwy. 141, 3.5 mi. S Hwy. 145, 7800', 7$ 5 2 , vi
(SE). NEW MEXICO: Bernalillo Co.: San Embudido Canyon,
SW base Sandia Mts., 14$ , vii (JL, KH). Lincoln Co.: 5 mi. N
Angus, Hwy. 37, 7050', 1 $ , viii (CAS). McKinley Co.: 4 mi. S
Fort Wingate, 1$, vii (CAS). Sandoval Co.: Frijoles Canyon,
Bandelier Nat’l Mon., 1$, viii (AM); Jemez Spgs., 1$, vii
(CM); ca 1 mi. N Sandia Man Cave, Las Huertas Canyon, S
of Placitas, 1 $ 1 9 , viii (AM). Taos Co.: 4 mi. S Questa, 7800',
1$, vii (CM). Torrance Co.: Capillo Peak summit, 9200', Man-
zano Mts., 1$, vii (CM). County undetermined: ridge NE of
Chupidero Canyon, 9100', Sangre del Cristo Range, 1 $ , vii
(CM).

HOST: Eriogonum jamesii Benth. in DC. var. jamesii.
COLORADO: Conejos Co.: W end Alamosa Reservoir, 49 $ 1 9 ,
vii (GG, JS), adult assoc. ( Shields #141). Rio Grande Co.: “D”
Hill, Del Norte, 32 $ 4 9, vi-viii (SE, MF), adult assoc. ( Shields
#139). NEW MEXICO: Sandoval Co.: Frijoles Canyon, Ban¬
delier Nat’l Mon., 19, viii (AM), adult assoc. ( Shields #147).
See Map 9.

Ssp. centralis flies from late June to mid August.

Some centralis from San Embudido Canyon, Bernalillo Co.,
New Mexico, show a likeness to battoides ellisii from north-
central Arizona in having a Harrow aurora and similar markings
on the underside. One male from Fort Wingate, N.M., also dis¬
plays this tendency. Specimens from Hwy, 141, 3.5 mi. S. of Hwy.
145, San Miguel Co., Colorado, are rather atypical centralis and
parallel ancilla cfn the underside; they are not close to ellisii.
A singleton female from Alta, Salt Lake Co., Utah, VII-6-43
(MCZ, V. Nabokov) appears to be this phenotype as well.

A long series of centralis from Schilling’s Spring, Conejos
Co., Colorado, is rather similar on'the dorsal and ventral surface
to rita coloradensis which is not sympatric, and differs consider-

16(l):l-67 , 1977

PH1LOTES

47

Map 9. — Distribution of S. battoides centralis.

48

OAKLEY SHIELDS

J. Res. Lepid.

ably from rita spaldingi which is sympatric! Both centralis and
spaldingi are also sympatric at Alamosa Reservoir, Conejos Co.
Ssp. centralis and r. coloradensis are sympatric at Great Sand
Dunes Nat’l Mon., Alamosa Co.; Ben West Hill and Silver Cliff,
Custer Co.; Hillside, Fremont Co.; “D” Hill, Rio Grande Co.;
and Villa Grove, Saguache Co., in Colorado.

Of considerable importance are centralis records from Modoc
Co., California (Burns Creek and Cedar Pass), a dramatic range
extension from the southern Colorado-northern New Mexico
stronghold. Specimens from Carrville, Trinity Co., California,
are intermediate between centralis and intermedia on the under¬
side.

An unusual male specimen from Oak Creek, Fremont Co.,
Colorado, has exceptionally heavy black borders that encompass
the outer half of the upperside.

Eriogonum jamesii var. jamesii is found from central Arizona,
most of New Mexico, and northern Texas, northward into eastern
Colorado and western Kansas into southern Wyoming, flowering
from June to September.

Shijimiaeoides battoides bernardino ( Barnes & McDunnough,

1916)

Barnes & McDunnough. 1916. Contrib. Nat. Hist. Lepid. N.
Amer., vol. 3, pp 116-117, pi. 11, ff. 9, 11, 13, p. 152.

The type locality of bernardino is Camp Baldy, “San Bernar¬
dino Mts.” [San Gabriel Mts.], San Bernardino Co., California.
Originally described only briefly as “a much smaller race” than
battoides battoides with paler ground and less heavy markings
on the underside, the figures of the type (upper and lower sur¬
faces) definitely establish its identity. The statement, “We have
other specimens from San Diego, Pasadena and Havilah,” also
coincides with the known distribution of bernardino. Langston
(1965, Map 1) gives a distribution map of bernardino in Cali¬
fornia. This is probably the subspecies referred to as “Lycaena
battoides ” from the Argus Mts. by Riley (1893, p. 244). Com¬
stock and Dammers (1934) give a complete life history descrip¬
tion of bernardino. They illustrate the last instar and pupa, as
do Emmel & Emmel (1973).

DISTRIBUTION (viewed 117$ gen. from 94 localities):
CALIFORNIA: Fresno Co.: Parkfield Grade Rd., 4.5 mi. S Hwy.
198, 1 $ , v (AM). Inyo Co.: Argus Mts., 1 S , vi (LACM); Big
Pine, 1$ 2 9, vi (CAS); 4 mi. W Big Pine, 1 9 , vi (UCD);

16(1): 1-67, 1977

PHILOTES

49

Carroll Crk., 9-11 mi. SW Lone Pine, 5 $ , v, vi (CIS); 8 mi. W
Lone Pine, 6200', 3$ 3 9 , vii (AM); Surprise Cyn., Panamint
Mts., 1 $ 1 $ , iv (CIS); Tuttle Crk., 2 mi. SW Lone Pine, 26 $
8$, v (CIS); Wyman Creek Canyon, 6800', White Mts., 19,
vii (AM). Kern Co.: Frazier Park, 4 a 7 $ , vi, vii (CIS, GG,
KH); 1 mi. W Jet. Frazier Park Rd., & U.S. Hwy. 99, la, vi
(KH); Havilah, 14 a 6$, vi (AM, CM); Keene, 1 a , vi (CIS);
Lebec, 4 $ 1 9 , vi (PO, CIS, GG); nr. summit, road up Mesquite
Canyon, 3500', El Paso Mts., 1 a 3 9 , v (AM); Mojave, 6 a 1 9 ,

v (CM); 6 mi. NW Mojave, 19, v (CIS); Mt. Pinos, 3$ 19,

vi (LACM); 4 mi. N Muroc, 1 $ , iv (CIS); Randsburg, 2a,

v (YU); 9 mi. W Randsburg, 1 $ , iv (YU); Tehachapi, 1 $ 1 9 ,

vii (CM, LACM); Walker Pass, 4200-5250', 18 a 4 9 , v, vi
(DB, LACM). Los Angeles Co.: Azusa, 1 $ , iii (LACM); 5 mi.
N Beverly Hills, Benedict Canyon, 1 9 , vi ( LACM ) ; Bouquet
Canyon, 4 $ , vi (LACM, KH); Camp Josepho, Rustic Cyn.,
6.3 3 9 , vi (GG); Cobal Crk. Cyn., Padua Hills, 1$, v (KH);
Cold Creek Cyn., Santa Monica Mts., 3$ 29, v (KH); Cold-
water Canyon, Santa Monica Mts., 1 $ , vi (LACM); Devils
Canyon, 5000', 2$ 3 9 , vi (AM); Eatons Wash, Pasadena,
3$ 1 9 , vi (MCZ, LACM); Encino, 3 <3 5 9, v (UI); Glendale,
5 <2 3 9 , v, vi (DB); Glendora, 7$ 2 9 , v, vi (AM, LACM,
CAS); Griffith Park, 12 a 4 9 , iv, vi (CAS, CIS, JL, CM); Horse
Trail Campground, Angeles Nat. For., 19, vi (AM); Lake
Hughes, la 2 9 , vi (KH); La Sierra Canyon, 1200', Santa
Monica Mts., 1 $ , vi (YU); La Tuna Canyon, 61a 139, v-vii
(AM, YU); Littlerock, la + “4”, iv (KH, CIS); 1 mi. W
Little Rock, 2 a 1 9 , v ( CIS ) ; Long Canyon, 3000', San Gabriel
Mts., 1 9 , vi (CM); Los Angeles, 12 a 7 9 , vi (CAS, LACM);
Malibu Lake, Santa Monica Mts., 2 a 1 9 , v, vi (KH, CIS);
Malibu Rd. to Tapia Park, 2 a 1 9 , viii (JL); Mint Canyon, nr.
Acton, 2 a 1 9 , v, vi (LACM, CM); Mt. Baldy foothills, la,

vi (KH); Mt. Wilson, 1 $ , vi (CM); Pacific Palisades, la, vii
(CAS); Pallatt Creek, la 1 9 , iv (CIS, AM); Pasadena, 9 a
39, vi, ix (CM, AM, LACM); Pearblossom, 3a, v (GG);
1.1 road mi. E jet. Pine Canyon Rd. & Ridge Route Rd., just
NE Sandberg, 2 a 3 9 , vi ( AM ) ; Rancho Santa Ana Botanic
Gardens, 1 9 , vi (LACM); Ridge Route nr. Whitaker Peak Rd.,
Tejon Mts., 3a 19, v (KH); Roscoe, 7a, v (LACM); San
Antonio Canyon, 15 a 3 9 , vi (CAS, CIS); San Antonio Wash,
2a 19, vii (AM); Sandberg, 2a 1 9 , vi (KH); 7 mi. NW
San Fernando, 1 a , vii (CIS); San Gabriel Canyon, San Gabriel
Mts., 2a 2 9 , v, vi (CAS, AM); West Fork, San Gabriel River,

50

OAKLEY SHIELDS

J. Res. Lepid.

1 3 , vi (LACM); San Rafael Hills, Glendale, 1$, vii (DB);
Santa Anita Canyon, San Gabriel Mts., 2 3 , vi (LACM; Santa
Monica Mts., 2 3 1 $ , vi (LACM); Sepulveda Pass, 1$ 1$,
vii (KH); Sierra Pelona Rd., Mint Canyon, 2 3, vi (AM);
Soldiers Home, 2 3 , vi (CM); Stunt Cyn., Santa Monica Mts.,
3$, v (JL); Switzers Camp, 3 3 3$, vii, viii (AM); Switzer’s
Trail, 3500', San Gabriel Mts., 1$ 1 9 , vi (LACM); Tanbark
Flat, 1$, vii (CIS); Temescal Cyn., Santa Monica Mts., 4 3,
v (KH); Topanga, 1500', 25 3 22$, v-vii (LACM); Topanga
Cyn., 13 2$, v (KH); Tujunga, 2$, v (LACM); 2 mi. NW
Valyermo, 33 3 23 $ , v (CIS); 2 mi. E Valyermo, 12 3 3$,

v (CIS). Monterey Co.: Arroyo Seco, 12 3 4 $ , v, vi (CIS, CAS,
LACM); 6 mi. W Greenfield, 1 3 , vi (KH); Jamesburg, 1 3 3 $ ,

vi (KH). Orange Co.: 4 mi. NE Laguna Beach, 1 3 , vii (CIS);
Orange Co. Park, 1 3 1 $ , vii (LACM). Riverside Co.: Aguanga,
3 3 , vi, vii ( CAS ) ; 6 mi. E Anza, 1 3 3 $ , v ( CIS ) ; Chino Can¬
yon, nr. Palm Springs, 1 3 , iv (AM); Citrus Exp. Sta., 1 $ , vi
(CIS); Corona, 1 3 , vi (CM); 1 mi. W Cottonwood Spr., Joshua
Tree Nat. Mon., 13, iv (CIS); Dripping Springs, 2$, vi
(LACM); Esperanza Rd., SE Cabazon, 1$, v (GG); Garnet
Queen Mine, 6000', Santa Rosa Mts., 1 3 , vi (CM); Gavilan
Peak, nr. Lake Matthews, 2 3 , iv (KH); Idyllwild, 53 4$,

vii (YU, CIS); Lee Lake, 7 3 3 $ , v, vi (CAS, LACM); Palm

Springs, 13, iv (UCD); near Palm Springs, 13 1 $ , iii

(LACM); Palms to Pines Hwy., E slope San Jacinto Mts., 3000',
3 3, iv ( JL ) ; Perris, 43 1 $ , vi, vii ( LACM ) ; Pinyon Crest, 12
rd. mi. SW Palm Desert, 1 3 , iv (CIS); 1 mi. E Pinyon Flat,
1$, vii (CIS); 3 mi. E Pinyon Flat, Santa Rosa Mts., 13, v
( CIS ) ; 4 mi. E Pinyon Flat, 4 3 1 $ , iv ( CIS ) ; Railroad Canyon,

3 3 1$, v (LACM); hills 3 mi. SW Redlands, 1 3 , iv (CIS);
Riverside, 3 3 3$, iv-vi (LACM, CAS, CM); 5 mi. W Sage,

4 3 5 $ , vii (CIS); San Jacinto Mt., 13 3 $ , vi, vii (LACM);
Santa Ana Mts., 1 3 , vi (DB); Snow Creek, 33 1$, v (CAS,
LACM); 1-2 mi. S Temecula, 13 3 10$, vii (CIS). San Benito
Co.: 12 mi. S Paicines, 3 3 4 $ , vi (CIS, CAS, LACM); Pinna¬
cles Nat. Mon., 1 3 1 $ , v (KH); 4 mi. E Pinnacles Nat. Mon.,

5 3 6 $ , vi (CIS, CAS, LACM); 3 mi. S Tres Pinos, 4 3 12,
vi (CIS, CAS, LACM). San Bernardino Co.: Alta Loma, 13
1 $ , vii (AM); Baldy Mesa, nr. Cajon Pass, 53 1 $ , vi (LACM);
Barton Flats, 1 3 , vi (CAS); Bowen Ranch Rd., Ord Mts., SE
Hesperia, 4 3 2$, v (GG); Burns Spring, 10 air mi. NE Yucca
Valley, long series, v (JL); Cactus Flats, 6000', San Bernardino
Mts., 2 3 3 $ , vii (MCZ); Cajon Pass, 2 3 3$, vi (JL); Camp

16(1): 1-67, 1977

PHILOTES

51

Baldy, la, vii (MCZ); 4 mi. SE Devora, 5 a 1 $ , vi (CIS);
9 mi. N Fontana, 1 a , vi (CIS); Granite Mts., nr. Apple Valley,
3$ 1 $ , iv, v (KH); Lucerne Valley, 4 a 20 $ , v (LACM,
CM); Hwy. 38 below Marton Pk., 3300', San Bernardino Nat.
For., 39 a 23$, vii (AM); Morongo, 2a, v (LACM); Oak
Glen, 1$, vii (CAS); Phelan, 2a, vi (LACM); 12 mi. ENE
Redlands, 3200', 2 a , vi (AM); 16 mi. E Redlands, la 3$,
vii (GG); Santa Ana Cyn., nr. San Andreas Fault, 3 a , vi (GG);
Santa Ana River, 5000', 1$, vii (AM); 1 mi. SE Sheephole
Summit, Sheephole Mts., 2 a 1$, iii, iv (CIS, JL); Victorville,
la, v (LACM). San Diego Co.: Alpine, 1 S 1 $ , vii (CM);
6 mi. E Banner, 3 S 5 $ , vi (CIS); 2 mi. W Barrett Dam, 1 S ,
vii (CIS); Bedford Canyon, 1 $ , vi (CIS); Black Mtn., 2 s
2$, vi (CIS, KH); 2 mi. SE Bonita, la 2 $ , vi (CIS); Box
Canyon, la, v (CAS); Buckman Spr., 3 a 1 $ , v, vi (CIS);
Calavo Gardens, La Mesa, 19a, 5$, v, vi (AM); nr. Camp
Ole, Mt. Laguna, 9 a 15 $ , vii (AM, CIS); Cardiff, 1 a , vi
(CIS); Challas Hts., 5 a 2 $ , iv (CM); Crest, nr. El Cajon,
5 a 5 $ , vi (GG); Cuyamaca, la, vii (KH); Del Mar,
la 1$, vi (KH); 1 mi. S Del Mar, 8 a 18 $ , vii (CIS);
Dictionary Hill, 1 a , vi (AM); El Cajon, 26a 4$, iv-vi (AM,
LACM, GG, YU); Fairmont Cyn., 4 a 2 $ , v, vi (CIS); 4 mi.
E Fallbrook, la 3 $ , vii (CIS); Garnet Peak, 5400', Laguna
Mts., 3 $ , vi (AM); Harbison Cyn., 1 $ , vi (CIS); 3 mi. NW
Henshaw Dam, 8a 9 $ , vii (CIS); 5-8 mi. NE Jamul, 2a 2$,
vi (CIS); 1 mi. W Julian, 1$, vii (CIS); Jet. St. Hwy. 76-78,
1 $ , vi (KH); Kearney Mesa, 1 a , vi (CM); 2-4 mi. N Laguna
Jet., 6a 4 $ , v, vi (AM, CIS); Laguna Mts., 2a, v, vi (KH,
LACM); La Jolla, 11a 11 $ , vi (CIS); Lakeside, 2$, v, vi
(CIS); 2 mi. NE Lakeside, la 1 $ , vi (CIS); La Mesa, 3 a
1 $ , iv, v (CAS, LACM); 1 mi. E Lyons Pk., 4 a 6 $ , vii (CIS);
Mason Valley, 6a 1 $ , iv, v (LACM); Mission Dam, la 2$,
vi (CIS); Mission Gorge, 10 a 7 $ , vi, vii (KH, PO, CIS);
Monument Pk., 1 a , vi (CIS); Mt. Helix, 5 a , v, vii (CIS, CAS);
Mt. Laguna, 18a 15$, vi, vii (CIS); Mt. Laguna Jet., 6a,
v (CIS); Mt. Palomar, 8 a 3 $ , vi, vii (CIS); Mt. Soledad, La
Jolla, 2 a 2 $ , vi (CIS); 3 mi. NE Poway, 2 a 1 $ , vi (CIS);
Powderhouse Cyn., la 1$, v (CIS); “S” Mtn., la, v (CIS);
San Diego, 7a 6$, v-vii (CM, CIS, LACM); 2 mi. SW Santa
Ysabel, la, vii (CIS); Santee, 1 a , vi (KH); Sentenac Cyn.,
la, hi (UI); Solano Beach, 25 a 19 $ , vi (AM, CIS, UCD);
Sorrento Beach, 2 a 8 $ , vii (AM, CIS); Sunnyside, 1$, vii
(CIS); Tecate Mtn., 14a 6$, vi (KH, PO, CIS): Thing

52

OAKLEY SHIELDS

J. Res. Lepid.

Valley, 1$, vii (CIS); Warners Hot Springs, 1$ 4 $ , vi, ix
(CM); Witch Creek, 2 $ 2 $ , vi (CM). San Luis Obispo Co.:
6 mi. S Creston, 1 $ , vi ( CIS ) ; nr. Pozo, 1 $ , vi ( KH ) ; 5 mi. E
Sta. Margarita, 1 9 , vi (CIS). Santa Barbara Co.: 1 mi. S
Buellton, 9^ 7 $ , v, vi (CIS); Green Cyn., 8 air mi. W New
Cuyama, 1 <$ 1 ?, v (CIS); Mission Cyn., Santa Barbara, 4$
l9? v (AM); 6 mi. NW New Cuyama, 15 $ 5$, v (CIS);
2 mi. E Solvang, 1$, vi (CIS); 10 mi. E Twitchell Dam,
Cuyama River Cyn., 1# 1$, v (CIS). Stanislaus Co.: 7 mi. E
Frank Raines Park, Del Puerto Cyn., 1$, v (J. Mori Coll.)
Ventura Co.: Circle X Ranch, B. S. A. Camp, 1500', 3 $ 1 9 ,

iv (CIS); 1 mi. W jet. Frazier Park Rd. & U.S. 99, 3 $ 49,
vi (KH). County undetermined: Fairview, 1$ 39, v (CAS);
Fredalba, 1 <$ , vii (CM); Nellie, 23 3 139, vii, viii (CM);
Ridge Route, Sierra Madre Mts., 1 3 1 9 , vi (LACM). NE¬
VADA: Churchill Co.: 7 mi. W Frenchman, 4$ 5 9 , vi (JS);
N end Sand Springs Mts., 4$ 1 9 , vi (DB); S end, Stillwater
Mts., 5 mi. W Frenchman, 4000', 11$ 4 9 , v, vi (DB). MEXI¬
CO: BAJA CALIFORNIA DEL NORTE: Arroyo Seco, Sierra
San Pedro Martir, 1 $ , v ( CAS ) ; 6 mi. NE Arroyo Seco, 1 $ ,

v (CIS); 12 mi. S Ensenada, l£, v (CAS); Laguna Hansen,
19, vii (LACM); 4 mi. SW La Zapopita, Valle de Trinidad,

1 $ , iv ( LACM ) ; 9 mi. W Mike’s Sky Ranch, Sierra San Pedro
Martir, 2800', 3 $ , v ( RH ) ; 20 mi. W Mike’s Sky Ranch, Sierra
San Pedro Martir, 1800', 3$, v (RH); 2 mi. W Rancho El
Ciprer, Sierra San Pedro Martir, 1800', 5 $ 3 9 , iv ( RH ) ;

2 mi. E San Telmo, Sierra San Pedro Martir, 1000', 4 $ , iv (RH);
2 mi. W Socorro, Sierra San Pedro Martir, 3000', 4 $ , v, vi ( CIS,
CAS).

HOSTS: Eriogonum fasciculatum Benth. var. polifolium
(Benth. in DC.) Torr. & Gray. CALIFORNIA: Los Angeles Co.:
Pearblossom, 3$, v (GG), adult assoc. ( Shields #184); 1.1 rd.
mi. E jet. Pine Cyn. Rd. & Ridge Route Rd., just NE Sandberg,
2$ 3 9 , vi (AM), adult assoc. ( Shields #94). Eriogonum fascic¬
ulatum Benth. in DC. var. foliolosum (Nutt.) S. Stokes ex
Abrams. CALIFORNIA: Fresno Co.: Parkfield Grade Rd., 4.5
mi. S Hwy. 198, 1 9 , v (AM), adult assoc. (Shields #85). Santa
Barbara Co.: 1 mi. W La Cumbra Lookout, E of San Marcos
Pass, Santa Ynez Mts., 1 $ , vi (AM), adult assoc. (Shields #97).
Eriogonum heermannii Dur. & Hilg. var. humilius (S. Stokes)
Reveal. NEVADA: Churchill Co.: 7 mi. W Frenchman, 4$
5 9 , vi (JS), adult assoc. (Shields #173 & 199). Eriogonum
heermannii Dur. & Hilg. var. floccosum Munz & Johnst. CALI-

16(1 ):l-67, 1977

PHILOTES

53

FORNIA: San Bernardino Co.: large canyon just SE of Clark
Mtn. Pk., Clark Mtn., 5 rd. mi. NW Mountain Pass, 2 $ , ix
(AM), adult assoc. ( Shields #175). Eriogonum heermannii Dur.
& Hilg. var. sulcatum (S. Wats.) Munz & Reveal in Munz.
CALIFORNIA: San Bernardino Co.: Ronanza King Mine Can¬
yon, E side Providence Mts., 1 $ , ix (AM), adult assoc. ( Shields
#179).

Ssp. bernardino is on wing from mid March to early August
(there are several September records), although not continuously
so at any one locality.

Females of bernardino from Pallatt Creek, Los Angeles Co.,
and Tuttle Creek, Inyo Co., have extensive blue on the upper-
side (as do enoptes langstoni near Lone Pine). A short series
from Big Pine, Inyo Co., are very light in color. Material from
Solano Beach, San Diego Co., shows a slight tendency toward
allyni. One bernardino male (gen. det. ) from Chino Canyon,
Riverside Co., in April is remarkably like enoptes dammersi on
the underside, which flies at this same locality in September and
October. The rare September form from the eastern Mojave
Desert on E. heermannii varieties appears nearest S. battoides
ellisii.

Ssp. bernardino is found in association with Eriogonum
fasciculatum var. foliolosum at Arroyo Seco, Monterey Co.; Sage
and Temecula, Riverside Co.; Paicines, Pinnacles, and Tres Pinos,
San Benito Co.; Bonita, Camp Ole, Fallbrook, Henshaw Dam,
Jamul, Lakeside, and Mt. Palomar, San Diego Co.; Buellton,
Green Canyon, New Cuyama, and Twitchell Dam, Santa Barbara
Co.; and Circle X Ranch, Ventura Co. (Langston records). Var.
foliolosum is found in low mountains in southwestern California
and northern Baja California.

Ssp. bernardino is also reported for E. fasciculatum var.
flavoviride at 1 mi. W Cottonwood Spring, Joshua Tree Nat’l
Mon., Riverside Co. (Langston records). Var. flavoviride extends
along the northwest margins of the Colorado Desert and adjacent
parts of the Mojave Desert, southeastern California.

Eriogonum heermannii var. humilius is found in Inyo and
Mono cos., California northward and eastward to Humboldt and
Nye cos., Nevada, flowering from June to October (see Reveal,
1970). Var. floccosum occurs in the eastern Mojave Desert in
Little San Bernardino, Providence, and Clark Mts., San Bernar¬
dino Co., and in southern Clark Co., Nevada, flowering from
August to October. Var. sulcatum ranges in Inyo and San Ber¬
nardino cos., California, eastward across Clark and southern

54

OAKLEY SHIELDS

/. Res. Lepid.

Nye cos., Nevada into Washington Co., Utah, and in northern
Mohave and Coconino cos., Arizona, flowering from July to
September.

Shijimiaeoides battoides martini (Mattoni, 1954)

Mattoni. 1954. Bull. S. Calif. Acad. Sci. 1954, vol. 53, pp. 157-
160 & 164, pi. 43, ff. 3-4.

The holotype male was described from Oatman, Mohave Co.,
Arizona, IV-17-48 (R. H. T. & D. Mattoni), along with the
allotype female and 27 paratypes. The male genital valve indi¬
cates that martini is a subspecies of battoides , according to
Mattoni (1954). He found it at Oatman in close association
with E. fasciculatum var. polifolium. It is distinguished from
all other subspecies of battoides by a combination of light blue
cyanic overlay, light grey underside, very wide auroral band,
and small macules (?), according to Mattoni (1954). The closest
relative of martini appears to be bernardino, which also uses
E. fasciculatum (see discussion). Mattoni (1954) gives some
distribution records and figures the upper surface of the male
holotype and lower surface of the female allotype, in black-and-
white (see also Emmel & Emmel, 1973, pi. 8, figs. 4-6).

DISTRIBUTION (viewed 25$ gen. from 14 localities):
ARIZONA: Mohave Co.: Goldroads, 1 $ 3 $ , v (AM); Kingman,
4$ 1 $ , iv (DB); Oatman, 24$ 8 $ , iv, v (AM, LACM, KH,
CAS). Pima Co.: between Quijotoa and Sells, 1$, v (LACM).
Yavapai Co.: Black Canyon, nr. Bumble Bee, 2$, iii (DB);
Hwy. 69, 4 mi. N Canyon, 2600', 11$ 20 $ , iv (CIS, CAS,
LACM); nr. Rock Springs, 4$ 6 $ , iv (DB, CIS); Hwy. 69
(now 1-17), 4 mi. N Rock Springs, 2$ 5 $ , iv (KR); Hwy. 79,
2 mi. N Rock Springs, 17$ 13 9, iv (KR). CALIFORNIA:
Inyo Co.: Argus Mts., 4$, v (YU); Homewood Canyon, 3500',
Argus Range, 1 $ , vi (MCZ). San Bernardino Co.: Bonanza
King, Providence Mts., 3 $ , iv (LACM); Bonanza King Mine,
1 $ 1 9 , iv (PO); 2 mi. N Bonanza King Mine, Providence Mts.,
1 $ , iv (PO); Clark Mts., 1 $ 1 9 , iv (LACM); Gilroy Canyon,
Providence Mts., 9$ 6 9 , iv (KH); wash below Gilroy Cyn.,
5$ 69, iv (JL); Sunflower Wash, 3200', Old Woman Mts.,
11$, iv (CIS, LACM). NEVADA: Nye Co.: Amargosa Desert,
6 mi. W Beatty, 3 $ , v ( DB ) .

HOST: Eriogonum fasciculatum Benth. var. polifolium
(Benth. in DC.) Torr. & Gray. CALIFORNIA: San Bernardino
Co.: canyon 3/4 air mi. SE Horse Thief Springs, 4400-4700',

16(l):l-67, 1977

PHILOTES

55

Kingston Range, 11 3 , vi (AM), adult assoc. ( Shields # 2 ).
NEVADA: Nye Co.: 3 mi. SW Beatty, 5 3 1 $ , v (AM), adult
assoc. ( Shields #79). See Map 10.

Ssp. martini is on wing from late March to late June.

Specimens from the vicinity of Beatty, Nevada, appear to be
intermediate between martini and bernardino. Also, both mar¬
tini and bernardino occur in the Argus Mts., California, (al¬
though not typical martini). The specimen from between Qui-
jotoa and Sells, Pima Co., Arizona, is large, with a partial red
band on the underside of the primaries, but is otherwise like
martini.

In the Providence Mts. and at Clark Mtn., San Bernardino
Co., martini flies in late April, while an unnamed form about the
size of bernardino , that is perhaps best described as near ellisii,
associated with two varieties of Eriogonum heermannii , is found
there in September. This latter form also occurs at Kingston
Canyon, Toiyabe Range, Lander Co., Nevada, VII-22 & 23-64,
one male (LACM). In the Stillwater Mts., Churchill Co., Ne¬
vada, in June, specimens resemble both bernardino and this
small form near ellisii , and use a third variety of E. heermannii
( see bernardino records ) .

Material from the Old Woman Mts. is clearly martini , while
a short series from the Sheephole Mts. (both in San Bernardino
Co.), some 40 miles to the southwest, is bernardino.

Although bernardino is sometimes reported associated with
E. fasciculatum var. polifolium, as at Pinyon Crest, Riverside
Co., and at Mt. Laguna and 6 mi. E Banner, San Diego Co.
(Langston records), martini is exclusively on this variety. Var.
polifolium ranges on the deserts of southern California and
northern Baja California, southern Nevada, southwestern Utah,
and western Arizona.

DISTRIBUTION SUMMARY

Subspecies of each Shijimiaeoides> species are mostly allo-
patric, as are their host Eriogonum species. Some, like ancilla,
glaucon, and enoptes enoptes, have a wide range both in eleva¬
tion and latitude. Others are narrowly restricted to certain plant
communities or habitats: spaldingi in ponderosa and pinyon-
juniper habitats; bernardino and dammersi to chaparral (and
desert); emmeli and ellisii to the Colorado Plateau; pallescens
to the Great Basin; battoides battoides to high elevations in the
Sierra Nevada; pallescens and emmeli in sandy desert; and
allyni, smithi, and bayensis to the seacoast.

56 OAKLEY SHIELDS /. Res. Lepid.

Map 10. — Distribution of S. battoides martini.

16(l):l-67, 1977

PHILOTES

57

Several “ring”, or Rassenkreis, distributions (involving a
number of allopatric subspecies of a polytypic species) suggest
themselves (although no breeding or genetic evidence is avail¬
able as yet):

I.

rita emmeli

rita rita

rita coloradensis

II. enoptes bayensis enoptes columbiae

enoptes enoptes “ rita ” spaldingi enoptes ancilla

enoptes langstoni Utah & W. Colo, small

enoptes ancilla

enoptes mojave

These rings of related subspecies could be due to re-contact with
the parent population from range expansion and subsequent
hybridization and suggest that these are still subspecies that have
not as yet attained the reproductive isolation of a species.

There are some striking disjunctions within Shijimiaeoides.
For example, S. battoides comstocki is known only from Te-
hachapi, Kern Co., California, and Park City, Summit Co., Utah.
S. battoides centralis occurs in northern New Mexico and south¬
ern Colorado, with two outlying populations in Modoc Co.,
California. S. battoides allyni is found in the Los Angeles coastal
region and on Cedros Island off Baja California. S. battoides
glaucon occurs in the Northwest from northeastern California,
northwestern (and extreme northeastern) Nevada, eastern Ore¬
gon, eastern Washington, Idaho, extreme southern British Colum¬
bia, and Sula, Montana, with one relict population on the Un-
compahgre Plateau in western Colorado. These disjunctions may
mean there was a previously wide distribution in the past, with
the intervening populations gone due to environmental extinc¬
tion or competitive displacement.

Two subgenera of Eriogonum ( Eucycla and Oligogonum)
are widely used by Shijimiaeoides and no doubt harbour new,
undescribed subspecies. Eucycla is common and widespread
throughout most of the trans-Mississippi West, from northern
Mexico northward into the Great Plains, westward to Baja Cali¬
fornia, Mexico, and the west coast of the United States (includ¬
ing the islands adjacent to the coast) to southwestern Canada.
Oligogonum occurs in western United States from northern
Mexico into western and northern Texas, westward to Califor-

58

OAKLEY SHIELDS

/. Res. Lepid.

nia, and northward to southwestern Canada, with outlying popu¬
lations in east-central Alaska and in Virginia and West Virginia.

ACKNOWLEDGEMENTS

In addition to those people mentioned in Part IV who loaned
material, I would like to thank: (AMNH) Frederick H. Rindge,
American Museum of Natural History; (UCD) R. O. Schuster,
University of California at Davis; (CF) Clifford D. Ferris; (DG)
Dennis Groothuis; (UI) D. P. Levin, University of Idaho; (RK)
Roy O. Kendall; (BM) Bryant Mather; (DM) David V. Mc-
Corkle; (SM) Sterling O. Mattoon; and (CS) Charles A. Seker-
man (OS = Oakley Shields, personal collection). John F. Emmel
and Richard M. Shields took the photographs. Charles R. Crowe,
Ernst J. Dornfeld, and Jon H. Shepard kindly provided many
locality records for Oregon and Washington, but these were not
used in the present study unless I actually examined the speci¬
mens. Drs. Lee D. Miller, J. F. Emmel, N. E. Gary, W. H. Lange,
J. L. Reveal, and R. W. Thorp, and Mr. J. Lane, read the paper
and offered suggestive criticisms. I thank the two reviewers for
their comments. This part completes a series submitted in partial
fulfillment of the degree, Doctor of Philosophy, University of
California, Davis, California.

BIBLIOGRAPHY

COMSTOCK, J. A. & C. M. DAMMERS, 1934. Additional notes on the
early stages of Californian Lepidoptera. Bull. So. Calif. Acad. Sci. 33:
25-34.

COMSTOCK, J. A., & C. HENNE, 1965. Notes on the life history of Phi-
lotes enoptes dammersi. Bull. So. Calif. Acad. Sci. 64: 153-156.

- , 1967. Notes on the life history of Philotes rita elvirae (Lepidop¬
tera; Lycaenidae). Bull. So. Calif. Acad. Sci. 66: 99-102.

DE FOLIART, G. R., 1956. An annotated list of southeastern Wyoming
Rhopalocera. Lepid. News 10: 91-101.

EMMEL, T. C., & J. F. EMMEL, 1973. The butterflies of southern Cali¬
fornia. Nat. Hist. Mus. Los Angeles County, Sci. Ser. 26: 1-148.
ERIKSEN, C. H., 1962. Further evidence of the distribution of some boreal
Lepidoptera in the Sierra Nevada. J. Res. Lepid. 1: 89-93.

GARTH, J. S., 1935. Butterflies of Yosemite National Park. Bull. So. Calif.
Acad. Sci. 34: 37-75.

KLOTS, A. B., 1930. Diurnal Lepidoptera from Wyoming and Colorado.
Distribution, life zones and habitat notes — new subspecies. Bull. Brook¬
lyn Entomol. Soc. 25: 147-170.

LANGSTON, R. L., 1963. Philotes of central coastal California (Lycaeni¬
dae). J. Lepid. Soc. 17: 201-223.

- , 1965. Distribution and hosts of five Philotes in California ( Lycaeni¬
dae). J. Lepid. Soc. 19: 95-102.

- , 1969. Philotes of North America: synonymic list and distribution

(Lycaenidae). J. Lepid. Soc. 23: 49-62.

16(l):l-67, 1977

PHILOTES

59

- , 1970. Philotes enoptes bayensis and tildeni Langston — four new

localities. Pan-Pac. Entomol. 46: 74.

LANGSTON, R. L., & J. A. COMSTOCK, 1966. Life history of Philotes
enoptes bayensis ( Lepidoptera : Lycaenidae). Pan-Pac. Entomol. 42:
102-108.

MATTONI, R. H. T., 1965. Distribution and pattern of variation in Phi¬
lotes rita. J. Res. Lepid. 4: 81-102.

REVEAL, J. L., 1969a. A new perennial buckwheat ( Eriogonum , Poly-
gonaceae) from southeastern Arizona. J. Ariz. Acad. Sci. 5: 222-225.

- , 1969b. A revision of the genus Eriogonum ( Polygonaceae ) . Ph.D.

dissertation, Brigham Young Univ., Univ. Microfilms, Inc., Ann Arbor,
Michigan. 70-4714.

- , 1970. Additional notes on the California buckwheats ( Eriogonum ,

Polygonaceae). Aliso 7: 217-230.

- , 1973. A new subfruticose Eriogonum (Polygonaceae) from western

Colorado. Great Basin Nat. 33: 120-122.

RILEY, C. V., 1893. Report on a small collection of insects made during
the Death Valley Expedition, pp. 235-268. In: U.S.D.A., Division of
Ornithology and Mammology; North American Fauna , no. 7, The
Death Valley Expedition, Part II. 402 pp.

SCOTT, J. A., S. L. ELLIS, & D. EFF, 1968. New records, range exten¬
sions, and field data for Colorado butterflies and skippers. J. Lepid.
Soc. 22: 159-171.

SHIELDS, O., 1975. Studies on North American Philotes (Lycaenidae).
IV. Taxonomic and biological notes, and new subspecies. Bull. Allyn
Mus. no. 28, 36p.

WILLIAMS, F. X., 1910. The butterflies of San Francisco, California. En¬
tomol. News 21: 30-41.

ADDENDA AND ERRATA

Several additional, relevant works have only lately come to
my attention. Hemming (1931) recognized three subspecies of
Shijimiaeoides divina: (a) d. divina Fixs. (Seoul, Pungking, and
Heijo in Korea; July); (b) barine Leech (Oiwake, Hokkaido;
June-July; Nambu, Honshu); and (c) asonis Mats. (Mt. Aso,
Kyushu; August).

Waterhouse (1903, p. 211-212) described a small rare blue,
Zizera delospila, and in 1938 erected a separate genus, Zetona,
for it. In certain particulars, e.g. small size, wing shape, check¬
ered fringe, white underside, extent of purplish on the upper-
side, brown females, etc. it closely approaches Zizeeria speciosa,
though I reserve judgement on their exact affinity until a detailed
morphologic and genitalic comparison can be made. Zetona
delospila is very rare and flies in June at King’s Sound, Queen’s
Islet, and the Ord River, northwestern Australia. “The adults
fly close to the ground in open areas amongst grass and low
shrubs” (Common & Waterhouse, 1972).

COMMON, I. F. B., & D. F. WATERHOUSE, 1972. Butterflies of Aus¬
tralia. Angus & Robertson, Sydney, 498 p., 41 pis.

HEMMING, A. F., 1931. On two little-known species of the genus Scolitan-

60

OAKLEY SHIELDS

J. Res. Lepid.

tides, Hb. ( Lep ., Lycaenidae). Ann. & Mag. Nat. Hist. (Ser. 10) 8:
577-579

WATERHOUSE, G. A., 1903. Notes on Australian Rhopalocera: Lycaeni¬
dae. Part III - Revisional. Proc. Linn. Soc. NSW 28: 132-275, 3 pis.

- , 1938. Notes on Australian butterflies in the Australian Museum.

No. 1. Rec. Australian Mus. 20: 217-222.

Unfortunately in Part IV, figures 29-33 were wrongly numbered. For
the captions to be correct, change:

29 to 33

30 to 32

31 to 30

32 to 31

33 to 29

16(l):l-67, 1977

PHILOTES

61

S.B. ALLYNI

S.B. MARTINI

* *

S.B. BERNARDINO

E. Mojave small pop.
near batto^tes ellisii

S.B. EUJSII

S.B. COMSTOCKI

S.B. CENTRALIS

» *

S.B. INTERMEDIA

s.b. oreSinensis

S.B. BATTOIDES

S. BATTOIDES GLAUCON

S.B.

S.E. BAYENSIS

%

S.E. COLUMBIAE

#

*

%

S.E. ANCILLA

W. Colo. - Utah small pop's
near enop^ ancilla

S.E. MOJAVE

S.E. SMITHI

%

S.E. TILDENI

S. ENOPTES DAMMERSI

S.R. RITA

*

S.E. LANGSTON I

% *

S. ENOPTES ENOPTES

*

So. Cal . pop's near
enoptes enoptes

S.R. SPALDINGI

*

S.R. COLORADENSIS
S.R. EMMELI

S.R. ELVIRAE

S.R. P ALBESCENS
SHIJIMIAEOIDES RITA MATTONII
Z.S. BO^RTORUM
ZIZEERIA SPECIOSA SPECIOSA

BAUERI

Fig. 31. — Schematic representation of the possible evolutionary development
of the Nearctic Shijimiaeoides subspecies, based on comparative adult
morphology and host Eriogonum classification.

62

OAKLEY SHIELDS

/. Res. Lepid

Fig. 32. — S. enoptes ancilla X S. battoides glaucon $ , possible hybrid, 8
mi. N of Jarbidge, Elko Co., Nev., VI-28-67, leg. J. Lane. Note the inter¬
mediate aspect of the valve cucullus. SEM photo, 60X, courtesy R. O.
Schuster.

16(1): 1-67, 1977

PHILOTES

63

Fig. 33. — Same, showing the teeth regularly arranged and the valve strong¬
ly curved though not as bifurcate as in battoides. SEM photo, 120X, cour¬
tesy R. O. Schuster.

64

OAKLEY SHIELDS

/. Res. Lepid.

Fig. 34. — S. enoptes ancilla X S. hattoides glaucon $ hybrid, Pattee Can¬
yon, 3500', Missoula Co., Montana, VI-23-62, leg. J. Scott. The left valve
seen here has a battoides aspect though not strongly bifurcate. SEM photo,
58X, courtesy R. O. Schuster.

16(l):l-67 , 1977

PH1LOTES

65

Fig. 35. — Same, front view of valve showing 10 teeth irregularly arranged
as in battoides. SEM photo, 73X, courtesy R. O. Schuster.

66

OAKLEY SHIELDS

/. Res. Lepid.

Fig. 36. — S. rita spaldingi, $ genitalia, Bee Springs, Kaibab Plateau, Coco¬
nino Co., Arizona, VI-30-52, leg. D. L. Bauer. Front view of valve cucullus,
showing the aspect of the 14 teeth. SEM photo, 176X, courtesy R. O.
Schuster.

16(1 ):l-67, 1977

PHILOTES

67

Fig. 37. — Same, showing the upraised cristae in the inside of both valvae.
SEM photo, 390X, courtesy R. O. Schuster.

Journal of Research on the Lepidoptera

16(1 ):68, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

ABERRANT SPEYERIA ID ALIA

TIM L. McCABE

Department of Entomology, Cornell University, Ithaca, New York 14853

I collected the specimen on a patch of virgin prairie where
Hesperia dacotae flies. Hundreds of S. idalia were present. Males
had been flying for several days, but females were just starting.
This aberrant male was not interested in feeding at the numerous
coneflowers it passed. It was drifting with the wind and occa¬
sionally checking a bunch of grasses in typical female search
behavior. In flight, it looked very much like a normal S. idalia,
but slightly darker. It was taken, on the wing, about two miles
from where it was first sighted.

COVER ILLUSTRATION

Fig. 1. — Male Speyeria idalia. 2 miles NW of McLeod, Ransom Co., North
Dakota. 10 July 1973. Tim L. McCabe. Dorsal view.

Fig. 2. — Same. Ventral view.

NOTICES

ADDRESS CHANGE: After April 1^ 1977
will be: Santa Barbara Museum of Natural
Del Sol Road, Santa Barbara, CA 93105.

the address of the Journal
History, 2559 Puesta

PUBLICATIONS: Moths of Southern Africa. E. C G Pinhey Tafel-
berg Publishers Ltd. , 28 Wale St. , Cape Town, ‘south Africa.
Available' in U. S. A. from Entomological Reprint Specialists, P. O.
Box 77971, Dockweiler Station, Los Angeles, CA 90007. $35.95
Butterflies of West Malaysia and Singapore, Vol. 1 and 2. W.

A. Fleming, E. W. Classey Limited, Park Road, Faringdon Berk¬
shire, England. Available in U. S. A. from E R S (above)

$47. 00 set. *

ADDITIONAL PUBLICATIONS: Revised Catalouge of the African
Sphingidae (Lepidoptera) with Descriptions of the East African
Species. R. H. Carcasson, E. W. Classey Ltd. , Park Road, Faring¬
don, Oxon, SN7 7DR, England. Available in North America from
Entomological Reprint Specialists, P. O. Box 77971, Dockweiler
Station, Los Angeles, CA 90007. $11.95.

SPECIAL NOTICE: The Smithsonian Foreign Currency Program, a
national research grants program, offers opportunities for support
of research in many countries in (among others) Systematic and En¬
vironmental Biology. Awards are determined on the basis of compe¬
titive scientific review. The deadline for submission is November
1 annually. For further information write the Foreign Currency
Program, Office of International Programs, Smithsonian Institu¬
tion, Washington, D. C. 2056Q.

ADDRESS CORRECTION: Kurt Johnson, author of "Post Pleis¬
tocene Environments and Montane Butterfly Relicts on the Wes¬
ter Great Plains" in volume 14, number 4 of the Journal: Dept,
of Biology, City University of New York, City College, Convent
Ave. and 138th Street, New York, NY 10031.

DEFECTIVE COPIES: Some missing pages have been detected
in volume 14, number 4. Please check your copy; correct
copies will be sent to those who received faulty issues and in¬
dicate such to us.

WRITING CONTEST: F or articles on any entomological subject to
be published in Insect World Digest, a bimonthly magazine The
deadline for submission of articles is September 1, 1 977. For de¬
tailed information, write to the editor. Dr. Ross H. Arnett Jr
P. O. Box 505, Kinderhook, NY 12106. * * *

FOR SALE: 1, 500 exotic butterflies, insects, biologicals. 72 page
catalog $1. 00. Complete Scientific, P. O. Box 307, Round Lake
IL 60073.

WANTED: Correspondence with members interested in trading
selling North American Lepidoptera. Send list of offers to: Ma
.Trpro.f 1950 Cottr ill lane, Westland, MI 48185.

or

rc

Volume 16 Number 1 March, 1977

IN THIS ISSUE

Studies on North American Philotes
( Lycaenidae )

V. Taxonomic and Biological Notes, Continued

Oakley Shields 1

Speyeria idalia

Tim L. McCabe

68

THE JOURNAL

P* Ul

COA4P,

UBftABY

im

Volume 16 Number 2 June, 1977

THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA

published by

The Lepidoptera Research Foundation, Inc.
at

25590 Puesta Del Sol Road, Santa Barbara, California 93105
EDITOR: William Hovanitz
ASSISTANT EDITOR: Karen Nielson 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.

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.

CLASSES OF
MEMBERSHIP

Regular

$12

year

Family

15

year

Contributing

25

year

Subscribing

50

year

Sponsor

100

year

Life

250

for life

Subscriptions to the Journal are $15.00 per year. Special subscriptions are avail¬
able to students at $10.00 per year.

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 2559 Puesta Del Sol Road, Santa
Barbara, California 93105. The publisher is THE LEPIDOPTERA RESEARCH FOUN¬
DATION, INC. The general editor is William Hovanitz at the above address. The
secretary-treasurer is Barbara Jean Hovanitz at the same address. The owner is
THE LEPIDOPTERA RESEARCH FOUNDATION, INC. THE LEPIDOPTERA
RESEARCH FOUNDATION, INC. is a non-profit organization incorporated under the
laws of the State of California in 1965. There are no bond holders, mortgages or other
security holders.

WILLIAM HOVANITZ

1915-1977

It is with great sadness that we note the death
of William Hovanitz on September 14th. His passing
is a profound loss to this Journal and to all those
involved with Lepidoptera. His contributions in teach¬
ing, research, writing and collecting will long be re¬
membered. As a tribute to William Hovanitz, The
Lepidoptera Research Foundation will publish a
memorial supplement to Volume 16, Number 4 of
The Journal of Research on the Lepidoptera . Included
will be two of his classic papers, “Ecological Color
Variation in a Butterfly and the Problem of ‘Protec¬
tive Coloration’,” and “Parallel Ecogenotypical Color
Variation in Butterflies,” and a complete biblio¬
graphy of his papers.

The Foundation, through its Board of Trustees,
has determined that the Journal will continue, and is
currently exploring various means of best carrying
on the Journal in the spirit of its founder. Until such
time, the Journal will continue under the direction
of the Board of Trustees.

The Board of Trustees

Barbara Jean Hovanitz
Paddy McHenry
Rudi Mattoni

r-

r

l!

o

U*'iA ,• AJiJ.

ddl $

BO 'ill

v< Jfc£. esahc : t g rftiw «i U.

M ^crv oc no xlinfiVoH iOBiiiiW lo
b oJ I oV ilr oi of hr roloiq ft u

j

.

IT ti ii mfctlliW oi slird- as *A .boio'i ?'■*

6 , ill// ubWhr - V-ll dblEOr- • 1 P’:< <*

lo

•?*-• u*.

h *i> :fl v>; . «wU i\0 sb-wrAofl a Iwtui*1! s#T

noq*sq oi> • -h ftiil - lo owl ad

Lna b ai noilfthftV

*V* b«R SvaimioloD avil

•«'. fK‘ ujti.;o-r? B f f/j -v!r • .*v aoilfchft '■*

.8Tvr q *sf «1 lo \dqBT

CLASSES OF

e«ool8inT lo JyjvK- rftbmjol orfT

•i bcs’.orrif- *:h vmni'v V arlj iodi V miim uobeBii
riiy :£*> i d 1* • m,- at aaohftv ?.fihoiqy 1 yltaoiif4

I ,ff .iii« <*fi lo tbiqg *wb ai W*io\ »dl iw>

aoifooiib odl labau Miailno* ^ a.

,1 V t: .• lo

sm^rr-

o i u

. IT

K%hw- r i n • • • ’VW>^itv&
^•.r. > • b • rSrfe^

r-*oV v ■ ; .

Vt ■ r . , - ; L>*-r«p»r

oil*.: i (!*■ v

>«l So* R r>»d *..<
lESKARCli f<
»tov« tHJi 7

TION

JiC

fl

Imm.

Journal of Research on the Lepidoptera

16(2) :69-74, 1977

FOODPLANT ECOLOGY OF THE BUTTERFLY
CHLOSYNE LAC INI A (GEYER)

( NYMPHALIDAE )

II. ADDITIONAL LARVAL FOODPLANT DATA
RAYMOND W. NECK

Texas Parks and Wildlife Department
4200 Smith School Road
Austin, Texas 78744

Previously ( Neck, 1973 ) I discussed the larval foodplants
of Chlosyne lacinia var. adjutrix (Scudder). Included were ex¬
tensive personal observations in central and south Texas in ad¬
dition to previous literature records. The purpose of this publi¬
cation is to update this previous communication before addi¬
tional detailed studies are reported in subsequent publications.
Several remarks and corrections concerning conclusions of the
first paper can be made at this time. Several new foodplant
records are available from personal observations. Personal ob¬
servations were made in central Texas, centering within Austin,
Travis County, Texas.

REMARKS ON INITIAL REPORT

Foodplants of Chlosyne lacinia var. adjutrix in central Texas
were grouped according to relative importance — major, occa¬
sional and rarely utilized foodplants (see table 1). Quantitative
data were not available at the time to properly rank the relative
importance of the foodplants within each group, however.
During 1972, a large, diverse site in the upper part of the old
floodplain of the Colorado River ( now impounded and re-named
Town Lake) in Austin, Travis County, was monitored for food-
plant utilization throughout the season (March-October). The
total numbers of broods per major foodplant at this site for the
entire season were as follows: Helianthus annuus L. — 171,
Verbesina encelioides (Cav. ) Gray — 119, and Ambrosia trifida

69

70

RAYMOND W. NECK

J. Res. Lepiu.

L. — 71. These three plants are in this same order in the pre¬
vious publication (Neck, 1973).

The order of the occasional foodplants given in the previous
paper appears to represent their relative importance. Actually,
the first three species ( plant #4 - #6 in table 1 ) are of approxi¬
mately equal significance. These plants are of importance under
certain seasonal and local conditions. Utilization of these three
plants will be discussed in a later article in this series.

Helianthus debilis Nutt. var. cumcumerifolius (T. & G. )
Heiser grows in sandy soils which are somewhat limited in
occurrence in the Austin area as much of the alluvial areas have
now been inundated as a result of multiple damming of the
Colorado River. Additionally, this plant is rapidly replaced in
succession by V. encelioides, a plant which is persistent in a
particular locality from year to year. A site at which cucumeri-
folius is abundant during one season will quite likely contain
no individuals of this sunflower the following season. Such suc¬
cession has been observed in both the alluvial sands (Quarter¬
nary) of the Colorado River in Austin and the Carrizo Sands
(Eocene) which outcrop to the east. My personal experience
would classify this plant as rarely utilized. However, Kendall
(1959) reported a large population of larvae of this plant.
Therefore, cucumerifolius is best considered an occasional food-
plant, but it is not utilized nearly as widely as the other three
plants placed in this category.

The foodplants in group III are only rarely utilized. Many
are not known to have become larval foodplants via adult female
oviposition; records of several plants are known only from larvae
which have crossed-over from another foodplant species. During
extensive observations in 1972 only one of these plants were
personally observed as a foodplant for adjutrix. Several larvae
were found on Heterotheca latifolia Buck, on 27 June 1972.
These larvae, however, had crossed-over from H. annuus; ovi¬
position by wild females on this plant is still unknown (see
Neck, 1973).

A single final instar larva of adjutrix was found on Calypto-
carpus vialis Less, by R. O. Kendall (pers. comm. 16 Sept. 1972)
west of San Antonio in Bexar County. This larva probably
crossed-over from H. annuus on which a brood was observed in
the same general area. Later, I found a single larva on C. vialis
in Austin in an area devoid of other suitable foodplants. These
two additional examples (see first in Neck, 1973) indicate that
C. vialis is an acceptable but little utilized foodplant.

16(2):69-74, 1977

CHLOSYNE LACIN1A

71

Previous mention was made of the apparent unsuitability of
commercial monocephalic varieties of sunflowers for proper de¬
velopment of adjutrix larvae. A recent study of cultivated sun¬
flower plots (Phillips et al., 1973) revealed a peak abundance
of 1980 larvae per acre (with 40,000 plants/acre). This figure
is only one larva for each 20.2 plants and would involve no
more than three to seven egg masses. Of all lepidopteran species
collected, adjutrix was the least abundant and was not discussed
within the text of the report. This unsuitability of a cultivated
form when compared to a wild weedy form is opposite that
reported for Papilio polyxenes (Erickson, 1975).

The “rarely utilized foodplant” initially reported as “Silphuim
sp.” (Neck, 1973) has been identified as Silphium asperrimum
Hook, (see table 1).

Several publications were omitted from the “Literature Cited”
section of the previous paper, i.e. Dethier, 1959; Kendall, 1964;
Remington, 1952; Remington and Pease, 1955; Straatman, 1962.
These articles are listed in the bibliography at the end of this
publication.

NEW LARVAL FOODPLANTS

Four additional plant species are now known to be larval
foodplants for adjutrix.

R. O. Kendall (pers. comm.) found larvae on Silphium albi-
florum Gray on 4 April 1968 in Terrell County (14 km W. of
Dryden), Texas.

On 20 August 1973 a larval brood was observed feeding on
garden grown specimens of the native Helianthus maximiliani
Schrad. in downtown Austin (two blocks from the state capitol
building). Previous years of observation of this plant under
natural conditions had revealed no larval infestations. Utiliza¬
tion of these plants apparently resulted from lack of suitable
foodplants in an urban environment. Additionally, other food-
plants in natural habitats at this time were in dessicated condi¬
tion due to summer drought; H. annuus plants were in poor
shape and V. encelioides had not yet become lush as yet due
to lack of rain.

Infestation of Jerusalem artichoke, Helianthus tuherosus L.,
has been observed in a residential area of Austin. Larvae of two
generations (15 June and 14 July 1975) were found on several
plants. Other adjutrix larvae were located on H. annuus con¬
currently with both instances of H. tuherosus infestations (only

72

RAYMOND W. NECK

J. Res. Lepid.

ten meters separated the two sites). H. tuberosus plants, how¬
ever, were downwind and visually screened (by an eight-foot
privacy fence) from the H. annuus plants. Oviposition on H.
tuberosus by adjutrix occurred as the result of “random” flight
by adjutrix females rather than a directed attraction away from
the H. annuus. Of further interest was the absence of any
adjutrix larvae on numerous individuals of H. maximiliani which
were also present in the same yard as the H. tuberosus.

H. tuberosus is not native to the Austin area. The nearest
native occurrence is in north Texas some 400 kilometers NNE
of the Austin area. In a study of Chlosyne gorgone (Huber) in
Kansas, O. R. Taylor (personal communication) found that H.
tuberosus was the third most frequently utilized larvae food-
plant (after H. annuus and A. trifida). Possibly, H. tuberosus
would be a significant larval foodplant of adjutrix if it occurred
naturally within the resident geographical range of adjutrix.
Both tuberosus and maximiliani are members of the section
Divaricati, maximiliani in Gigantei (see Heiser et al 1969).

H. petiolaris is a member of the section Annui which includes
the previously reported foodplants annuus , argophyllous and
cucumerifolius (Heiser et al, 1969). Members of this section
appear to be particularly suitable for adjutrix larvae although
no adult oviposition has been observed on H. argophyllous. The
highly pubescent nature of the leaf surfaces of H. argophyllous
may not provide sufficient ovipositional cues despite the pre¬
sumed phytochemical resemblance to other related species of
Helianthus, especially H. annuus.

ADDITIONAL LITERATURE RECORDS

Several additional literature records of larval foodplants of
adjutrix have been located (references from California to ex¬
treme western Texas refer to the subspecific taxon crocale
Edwards). Larvae were reported on Xanthium strumarium L.
(as X. canadense Mill.) around Blythe, California (Comstock
and Dammers, 1935). Later, Comstock (1946) reported larvae
“may be found in the fall throughout the Imperial and Coa¬
chella Valleys” on H. annuus. Cockerell (1941) reported that
“larvae abound on sunflowers” ( H . annuus) in southern New
Mexico and northern Mexico.

Bauer (1975) reported that larvae feed on Grindelia micro -
cephala DC. This record originated from an observation by
R. O. Kendall (personal communication). Kendall (in litt. ) has

16(2):69-74, 1977

CHLOSYNE LAC1NIA

73

informed me that this report is unverified; thus, this species is
not accepted as a valid foodplant record at this time. G. micro-
cephala is a member of the tribe Astereae (same as Hetero¬
theca which is known as adjutrix foodplant only by cross-over
larvae); therefore, G. microcephala is unlikely to be a widely-
used foodplant of adjutrix.

ACKNOWLEDGEMENTS

For personal communication of information utilized in this
report, I wish to thank P. Chavez, R. O. Kendall and D. H.
Riskind.

LITERATURE CITED

COCKERELL, T. D. A. 1914. The entomology of Helianthus. Ent. 47: 191-
196.

COMSTOCK, J. A. 1946. A few pests of sunflower in California. Bull. So.
Cal Acad. Sci. 45: 141-144.

- and C. M. DAMMERS. 1935. Notes on the early stages of two but¬
terflies and one moth. Bull. So. Cal. Acad. Sci. 34: 81-87.

CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular
plants of Texas. Texas Research Foundation, Renner, Texas, 1881 pp.

DETHIER, V. G. 1959. Egg laying habits of lepidoptera in relation to
available food. Can. Ent. 91: 554-561.

ERICKSON, J. E. 1975. The comparative utilization of cultivates and
weedy unbellifer species by larvae of the black swallow-tail butterfly,
Papilio polyxenes. Psyche 82: 109-130.

HEISER, C. B„ and C. M. SMITH, S. B. CLEVENGER and W. C. MAR¬
TIN, JR. 1949. The North American sunflowers (Helianthus). Mem.
Torr. Bot. Club 22(3): 218 pp.

KENDALL, R. O. 1959. More larval foodplants from Texas. J. Lepid. Soc.
73: 221-228.

- . 1964. Larval foodplants for twenty-six species of Rhopalocera (Pa-

pilionoidea) from Texas. J. Lepid Soc. 18: 129-157.

NECK, R. W. 1973. Foodplant ecology of the butterfly Chlosyne lacinia
(Geyer) (Nymphalidae). I. Larval foodplants. J. Lepid. Soc. 27: 22-
23.

PHILLIPS, R. L., N. M. RANDOLPH and G. L. TEETES. 1973. Seasonal
abundance and nature of damage of insects attaching cultivated flow¬
ers. Misc. Pub. Tex. Agr. Exp. Sta. 1116: 7 pp.

REMINGTON, C. L. 1952. The biology of Nearctic lepidoptera. I. Food-
plants and life-histories of Colorado Papilionoidea. Psyche 59: 61-70.

- and R. W. PEASE. 1955. Studies in foodplant specificity. I. The

suitability of swamp white cedar for Mitoura gryneus (Lycaenidae).
Lepid. News 9: 4-6.

STRAATMAN, R. 1962. Notes on certain lepidoptera ovipositing on plants
which are toxic to their larvae. J. Lepid. Soc. 16: 99-103.

74

RAYMOND W. NECK

/. Res. tepid.

TABLE 1.

Larval foodplants of Chlosyne lacinia var. adjutrix (Scudder) in central
Texas with additional data not reported in Neck (1973); Nomenclature
from Correll and Johnston (1970).

I. Major Foodplants

1. Helianthus annum L.

2. V erbesina encelioides (Cav.) Gray

3. Ambrosia trijida L.

II. Occasional Foodplants

4. V erbesina virginica L.

5. Viguiera dentata (lav.) Spreng.

6. Helianthus debilis Nutt. var.

cucumerifolius ( T. & G. ) Heiser

III. Rarely Utilized Foodplants

8. Ambrosia artemesiifolia L.

9. Parthenium hysterophorous L.

10. Helianthus argophyllous L.

11. Xanthium strumarium L.

12. Simsia calva (E. & G. ) Gray

13. Calyptocarpus vialis Less.

14. Silphium asperrimum Hook.

15. Gaillardia pulchella Foug.

16. Heterotheca latifolia Buck.

17. Helianthus maximiliani Schrad.

18. Helianthus tuberosus L.

Journal of Research on the Lepidoptera

16(2) :75-82, 1977

DESCRIPTIONS OF A NEW SPECIES OF
EUPITHEC1A AND THE MALE OF
E. COCOAT A PEARSALL ( GEOMETRIDAE ) 1

ROGER L. HEITZMAN and WILBUR R. ENNS2

Department of Entomology
University of Missouri-Columbia 65201

ABSTRACT

A new species, Eupithecia peckorum, is described from Missouri be¬
longing to the palpata species group. The male of E. cocoata Pearsall also is
described revealing this species to be the first Nearctic representation of
its species group.

INTRODUCTION

These findings are the result of a thesis reviewing the
Larentiinae of Missouri. Eupithecia cocoata Pearsall (1908) was
described from a single female specimen from Plummer’s Island,
Maryland. This species has not since been recorded until now.
A series of six specimens from Missouri and Arkansas reveals
two males with unusual genitalic structure. The laterally bifur¬
cate uncus is unique among the Nearctic fauna introducing a
new species group for the region. Its closest ally appears to be
E. haworthiata Dbd., a Palearctic species.

The new species is known exclusively from three vernal speci¬
mens taken in Independence, Missouri over an eight year period.
This is apparently a univoltine species, which is typical for
Missouri Eupithecia. It inhabits a sparsely forested environment.
The wings and the genitalia show unique characters and cannot
be confused with any other species.

Eupithecia cocoata Pearsall
Plate I: Figs. 4,5,6. Plate II: Fig. 8

MALE: Head: Eyes black; vertex heavily gray and brown scaled
with slightly raised scales anteriorly, followed by a thin row of
erect bristles ( chaetosemas ) and very small pale scales, and
posteriorly with a collar of erect larger scales; frons with closely
appressed, gray scales, extending a bit and turning brown at the

Contribution from the Missouri Agricultural Experiment Station, Entomol¬
ogy Research Museum, Journal Series No. 7712. Approved by the Director.
Received for publication 22 Nov. 1976.

2Research Assistant and Professor of Entomology, respectively .

75

76

HEITZMAN and ENNS

J. Res. Lepid.

base of the proboscis; pilifers and proboscis finely pubescent
with a few fine hairs, labial palps moderate in length (about
diameter of eye), broad and down-curving with thick gray scal¬
ing; antennal base gray-scaled, segments as wide as long, with
fine ventral and lateral hairs (about one-third length of segment
in size), dorsum with gray scales.

Thorax: Patagia with broad, gray scales; tegulae extending
beyond base of hindwing, with gray and brown scales and
pencils; dorsum covered by small gray and brown scales; venter
thickly scaled with white, gray, and brown scales, predomi¬
nately light colored; forelegs gray with some tarsal banding
with lighter scales, middle and hind legs pale gray.

Abdomen: More thickly scaled and darker, with predomi¬
nately gray and some brown scales.

Wings: Length of fore wing: 11 mm. Dorsal surface: Fore
wings, ground color gray with some faint, brown, wavy, thin
lines; two very obscure premedial lines; medial line leaving
costa obliquely into discal spot, continuing faintly and obliquely
towards body to inner margin; medial line mirrored by three
more identical lines distally; subterminal line thicker but faint.
Hind wings, ground color gray with brown, wavy, thicker lines,
more prominent than those on forewings but still faint; lines
darker toward inner margin; antemedial line present; medial line
passing thru discal dot; three more distally located lines present,
homologies uncertain. Ventral surface: Identical to dorsum but
lines and ground color fainter. Terminal lines of all wings dark;
fringe alternating gray and brown hairs.

Genitalia: Uncus laterally bifurcate and well formed; tegu-
men large and rounded posteriorly, with long setae laterally;
transtilla weakly formed; labidal arms and feet normally de¬
veloped; labidal pads of most common shape, with hairy bases
and tips, rest naked; labidal papillae abnormally long and stout
(at least one-half size of pads), tipped with fine hairs; juxta
normal; saccus normal; valve broad and without processes;
costa with setae on its entire length; valvula rather thickly
haired and naked proximally; sacculus with a few fine hairs;
aedeagus relatively large, thickly spiculate at its opening and
pointed, vesica with numerous sclerotizations; ninth segment
with hair pencils; ventral plate with base broad and weakly
bilobed, tapering at mid-length into two thin arms.
SPECIMENS: Two males from Blue Springs State Park, Wash¬
ington Co., Arkansas: 3 May 1969 and 4 June 1971 at UV light
(R. L. Heitzman); three females from same locality: 29 May

16(2) :7 5-82, 1977

EUPITHECIA

77

1966, 16 April 1967, and 3 May 1969 at UV light (J. R. Heitz-
man); one female from Sarcoxie, Jasper Co., Missouri, 16 May
1972 at UV light (J. R. Heitzman).

LOCATION OF SPECIMENS: The described male will be
deposited in the United States National Museum, Washington,
D. C. where the female type is located, and one female in the
Entomology Museum of the University of Missouri, Columbia.
The remaining specimens will be retained by the senior author.

Eupithecia peckorum sp. nov.

Plate I: Figs. 1,2,3. Plate II: Figs. 7,9,10
MALE: Head: Eyes black; vertex heavily brown-scaled, with
slightly raised scales anteriorly, followed by thin row of erect
bristles (chaetosemas ), and posteriorly with a collar of erect,
larger scales tipped with dark brown; frons with brown, closely
appressed, scales in medial area, scales dark brown and longer
near eyes and base of proboscis; pilifers and proboscis finely
pubescent, thinly arrayed with longer hairs; labial palps ex¬
tremely long and up-curving (about twice diameter of eyes),
thin but heavily scaled with short and long dark brown tipped
scales; antennal base completely scaled, antennae bifasciculate
with segments slightly longer than wide, thick patches of fine
hairs arising most prominently from two ridges on the ventral
surface of each segment (about one-half length of segment in
size), dorso-lateral surface covered by dark and light brown
scales.

Thorax: Patagia with broad, long, brown scales; tegulae
reaching base of hind wing and brown-scaled; dorsum covered
with small scales; venter thickly covered with light brown scales
and some intermittent darker scales; forelegs light with some
dark banding, middle and hind legs evenly light brown.

Abdomen: More darkly and thickly scaled with variations in
shading.

Wings: Length of fore wing: 9 mm. Dorsal surface: fore
wings, ground color red brown, with dark brown lines; basal
line complete and bowing distally; antemedial line arising at
one-third length of costa and jutting perpendicular to body for
a short distance towards discal dot then abruptly angling down
towards body to inner margin; a very obscure medial line
arising from inner margin but incomplete and difficult to follow;
postmedial line prominent, arising at two-thirds length of costa,
curving distally at beginning but quickly straightening and

78

HEITZMAN and ENNS

J. Res. Lepid.

fading as it nears inner margin; subterminal line represented by
weak series of pale dots encased in dark, transverse dashes, final
dash near inner margin enlarged and suffused into postmedial
line. Hind wings, antemedial and postmedial lines continuing
into hind wings but not as dark; basal, medial and subterminal
lines vague; discal dot present; ground color same as fore wing
but lightened at costal and basal areas. Ventral surface: Fore
wings, ground color pale brown; costa dark brown to ante-
medial; antemedial line very weak; discal dot present; medial
line obscure; postmedial line strong and fading before inner
margin; subterminal line weak. Hind wings, ground color same
as fore wing; antemedial weak; medial obscure; discal dot
present; postmedial line strong and complete; subterminal ob¬
scure. Terminal lines of all wings dark brown; fringe alternating
dark and light brown hairs.

Genitalia: Structures typical of palpata group; tegumen with
extremely long setae; transtilla thickened centrally; labidal arms
and feet well developed; labidal pads regularly haired by minute
setae; labidal papillae very small and tipped with small hairs;
saccus well developed; costa straight and only slightly thickened
medially, displaying several very long setae; valvula with scales
on its anterior edge, remainder heavily haired with a long tuft
of hairs proximally; sacculus with patch of setae, some very
long at its base; aedeagus centrally curved and slightly narrowed
in the middle, vesica with an extensive array of complex sclero-
tizations; ninth segment without pencils; ventral plate with
bowed, stout arms recurving distally, tips pointed ventrally and
rounded laterally, anterior portion or base well sclerotized with
its shape typical of the species group.

FEMALE: Same as male, except antennae not bifasciculate and
hairs smaller; fore wing length 10 mm, antemedial line angled
more towards body, discal dot very vague; all lines on venter
of wings more pronounced.

Genitalia: Ovipositor and apophyses typical of group; ostium
narrower than ductus bursae; cervix bursa far removed from
ductus seminalis which is atypical for the group; ductus semin-
alis normally shaped; bursa copulatrix spherical in shape and
almost entirely embedded with spines.

ETYMOLOGY: We take pleasure in naming this species for
Dr. and Mrs. William B. Peck of Central Missouri State Uni¬
versity, Warrensburg. Dr. Peck is a well known araneologist and
friend of the authors.

TYPES: Holotype, male, Independence, Jackson Co., Missouri,

16(2):75-82, 1977

EUPITHECIA

79

19 April 1971 at UV light (R. L. Heitzman); allotype, same
locality, 17 April 1963 at UV light (J. R. Heitzman); paratype,
one male, data and collector same as allotype.

TYPE LOCALITY: City limits of Independence, Jackson Co.,
Missouri, in sparsely forested and disturbed habitat.

LOCATION OF TYPES: The holotype and allotype will be de¬
posited in the type collection of the United States National
Museum, Washington, D.C. The paratype will remain in the
senior author’s collection.

DISCUSSION

Only one other Nearctic form, E. longidens kerrvillaria
Cassino and Swett (1924), approaches peckorum in appearance.
The former is a light brownish-gray, the lines arise further dis-
tally on the fore wings with the subterminal line plain, and the
distal dot prominent on the female. The bursa, aedeagus, valve
and ventral plate, however, vary quite radically from peckorum.
The ventral plate of peckorum most closely resembles that of
E. pseudotsugata MacKay, but with the arms bowing more and
the apices not so hooked and pointed. The bursa is quite unique
and cannot be readily compared with any other species.

ACKNOWLEDGEMENTS

We wish to thank Dr. Douglas C. Ferguson for his compari¬
son of a specimen of E. cocoata series with the type in the United
States National Museum, Washington, D.C., U.S.A. and his
comments on the new species, and also, Dr. Klaus Rolte of the
Biosystematics Research Institute, Ottawa, Ontario, Canada and
Dr. D. S. Fletcher of the British Museum (Natural History),
London, England for comments on and examination of illustra¬
tions of the genitalia. Also, thanks are extended to Mr. Paul
Szopa of the University of Missouri, School of Forestry, Fisher¬
ies and Wildlife for making photographic prints of the adults.

LITERATURE CITED

CASSINO, SAMUEL E., and LOUIS W. SWETT. 1924. Some New Eupi-
thecias. Lepidopterist, vol. 4, p. 27.

KIRKWOOD, CARL W. 1961. A New Eupithecia from Arizona (Lepidop-
tera, Geometridae ) . Bull. S. Calif. Acad. Sci., vol. 60, pp. 45-6, fig. 1.

LEUSCHNER, RONALD H. 1965. California Costal Eupithecias with De¬
scription of a New Species (Geometridae). Jour. Res. Lepid., vol.
4(3): 191-7, fig. 1.

80

HEITZMAN and ENNS

J. Res. Lepid.

MacKAY, MARGARET R. 1951. Species of Eupithecia reared in the Forest
Insect Survey in British Columbia ( Lepidoptera: Geometridae ) . Can.
Ent., vol. 83, pp. 77-91, pis. 1-4.

McDUNNOUGH, JAMES H. 1949. Revision of the North American Species
of the Genus Eupithecia (Lepidoptera, Geometridae). Bull. Amer.
Mus. Nat. Hist., vol. 93, pp. 553-728, figs. 1-20, pis. 1-32.

PEARSALL, RICHARD F. 1908. Eastern and Central Eupithecias. Jour.
New York Ent. Soc., vol. 16, p. 103.

PIERCE, F. N. 1967. Reprint. The Genitalia of the British Geometridae.
E. W. Classey, England, pp. 1-84, pis. 1-48.

RINDGE, FREDERICK H. 1952. Taxonomic and Life History Notes on
North American Eupithecia (Lepidoptera, Geometridae). Amer. Mus.
Novitates, no. 1569, pp. 1-27, figs. 1-8.

- . 1956. Descriptions of and Notes on North American Geometridae

(Lepidoptera). Ibid., no. 1784, pp. 1-23, figs. 1-19.

- . 1963. Notes on and Descriptions of North American Eupithecia

(Lepidoptera, Geometridae). Ibid., no. 2147, pp. 1-23, figs. 1-7.

16(2) -.75-82, 1977

EU PITH EC I A

81

PLATE I

Figs. 1-3. — E. peckorum n.sp. (holotype male). 1, clasping unit; 2, aede-
agus; 3, eighth sternite.

Figs. 4-6. — E. cocoata Pearsall (male). 4, eighth sternite; 5, aedeagus;
6, clasping unit.

82

HEITZMAN and ENNS

J. Res. Lepid.

PLATE II

Fig. 7. — E. peckorum n.sp. (allotype female), bursa, ventral view.
Fig. 8. — E. cocoata Pearsall, male.

Fig. 9. — E. peckorum n.sp., holotype.

Fig. 10. — E. peckorum n.sp., allotype.

Journal of Research on the Lepidoptera

16(2) :83-86 1977

AUTUMNAL FALSE BROODS OF
MULTIVOLTINE BUTTERFLIES

AT DONNER PASS, CALIFORNIA
ARTHUR M. SHAPIRO

Department of Zoology, University of California,
Davis, California 95616

The phenomenon of autumnal “false broods” produced by
multivoltine “weedy” butterflies under warm, settled weather
conditions has been documented in the northeastern United
States (Shapiro 1962, 1967, 1970) and is well known to local
collectors in many parts of North America and Europe. It has
not been previously recorded at high elevations in mountainous
regions, however. Donner Pass, at roughly 7000 feet in the high
montane-subalpine Sierra Nevada of north-central California,
would seem an unlikely place to look for it. The climate of the
Pass has been vividly described by Emmel and Emmel (1963):
“Snowmelt occurs in early June while snow flurries can be ex¬
pected in early September. Sandwiched between nine months
of winter — spring, summer and fall in the Donner Pass area must
pass rapidly in succession . . . flight or growth seasons are
measured in weeks instead of months.” The Donner Pass winter
has earned its place in the history and folklore of the West. The
Donner Party arrived at the Pass on 1 November, 1846 and found
the snow already five feet deep. Storm after storm added to that
total, raising it to eight feet on the level by 13 December
(Stewart, 1959). Neither Emmel and Emmel nor Stewart, how¬
ever, pointed out that Sierran winters vary enormously in their
timing and their severity. Since 1972, I have conducted pheno-
logical studies of the butterfly faunas of several localities along
a transect paralleling Interstate Highway 80, from sea level at
Suisun Bay to 7000 feet in Donner Pass (Shapiro, 1975a). From
these studies, it has become apparent that the native fauna of
the Pass is mostly univoltine and seasonally conservative, but
such is not the case with “weedy” species occurring over a wide
altitudinal range and characterized by high vagility and develop¬
mental flexibility. Under appropriate autumn conditions, these
species will continue to emerge and to breed. It is thus not at

83

84

ARTHUR M. SHAPIRO

/. Res. Lepid.

all unusual to find butterfly species which do not hibernate as
adults flying in Donner Pass in October and in some years into
November.

In 1975, 29 species of butterflies were flying in my Donner
study area on 30 September. This rather large number reflected
the general lateness of the season due to a cold, wet spring and
several cold summer storms. The first snow of the season, an
inch, fell on 7 October with eight more inches on 10 October
and another eighteen inches by the 12th. However, it warmed
rapidly, with afternoon highs in the 60s F and lows near freezing,
and by 19 October the snow was gone except on north-facing
slopes; five species were flying — Colias eurytheme and Pieris
rapae, both “weedy” non-hibernators, and Nymphalis californica,
Vanessa virginiensis, and Polygonia zephyrus, all hibernators.
In addition, I noted three species of grasshoppers, various Dip-
tera (Tachinidae, Muscidae, Syrphidae) and Hymenoptera, spi¬
ders, four species of mammals, three of birds, and two of lizards;
the mid-afternoon temperature was 64 °F. After a much colder
but snowless interval, it warmed again in early November and I
returned on 4 November (temperature again 64° F) and saw one
individual each of C. eurytheme, P. rapae, and Vanessa carye.

The 1976 season was very advanced; the snow pack was far
below normal and was nearly gone by late April. Most species
emerged and disappeared several weeks earlier than in 1975, and
by 4 October only 12 species were flying. Continued warm
weather produced late emergences of multivoltines and also
encouraged altitudinal dispersants, so that 15 species were flying
on 15 October with a mid-afternoon temperature of 68 °F. On
3 November, in 66° weather, six species were still flying: Pieris
rapae, P. protodice, Colias eurytheme, Pleheius acmon, and
Pyrgus communis, all “weedy” non-hibernators, and Precis
coenia, an immigrant from lower elevations (Shapiro, 1973, 1974).
On 10 November, with 60° and variable cloudiness, six species
were again recorded: P. rapae, P. protodice, C. eurytheme,
P. acmon, P. coenia, and the hibernator Vanessa virginiensis.
The butterflies were visiting widely scattered blossoms of Aster,
Monardella, and dandelion (Taraxacum) . On 14 November a
foot of snow fell in the Pass, but immediately thereafter warm,
stable conditions returned with daily highs in the 60s and lows
in the upper 20s to low 30s. On 20 November, with afternoon
temperatures from 63-66 °F, snow remained only locally, mostly
on north-facing slopes. Five species were recorded: P. protodice
and C. eurytheme (the latter common) and three Nymphalids,

16(2):83-86, 1977

MULTIVOLTINE BUTTERFLIES

85

presumably hibernators: V. virginiensis, V. carye, and Polygonia
zephyrus. Plants recorded in bloom included species of Poten-
tilla, Achillea , Aster, Solidago, Tragopogon, Taraxacum, Matri¬
caria, Cirsium, Erodium, Sidalcea, Wyethia, Phacelia, Castilleja,
and Chrysothamnus . Two species of grasshoppers, Muscid,
Syrphid, and Tachinid flies, bumblebees, spiders, and one species
of bird were seen, along with black bear tracks in the snow.
Colias visited Aster and Taraxacum and were seen flying from
1220 to 1510 hours. Twenty C. eurytheme were collected: 12
males, 4 orange females and 4 white females, of which one
white was very worn and the other specimens all more or less
fresh; one male was soft-winged. The abundance of C. eurytheme
was probably an accident of seasonal timing; the first spring
records (flyups from the east?) were on 14 May, a month
earlier than usual; an entire additional generation may have
been reared in the Pass in 1976, providing a large reservoir of
hatchable autumn pupae. The “rock garden” habitats of char¬
acteristic univoltine spring species such as Incisalia “fotis” windi,
Callophrys lemberti, and Plebeius lupini were visited but were
devoid of butterfly activity.

The probability of successful reproduction by late-emerging
multivoltines at Donner Pass is so low that one must assume
natural selection would operate against such “mistakes,” con¬
stantly perfecting diapause-induction systems. That, however,
presupposes the existence of resident overwintering populations
there. In the case of P. protodice and P. acmon that supposition
is almost certainly false. C. eurytheme seems to overwinter only
sporadically. Of the “weedy” multivoltines, only P. rapae
(Shapiro, 1975b) seems to be a permanent resident, but it also
has a temperature-photoperiod system for induction of pupal
diapause which would limit the amount of wasted fall develop¬
ment. The occurrence of autumnal “false broods” in so difficult
an environment as Donner Pass underscores the poor adaptation
of “weedy” multivoltine species to cold climates, the ephemeral
nature of their populations there, and their contrast with the
native montane, subalpine, and alpine fauna, which is “obli-
gately” univoltine under realistic environmental regimes.

ACKNOWLEDGMENTS

This research was supported in part by grant D-804 from
the Committee on Research, U. C. Davis. It has also benefitted
from remarks made by students and associates to the effect that

86

ARTHUR M. SHAPIRO

J. Res. Lepid.

anyone taking a butterfly net to Donner Pass after Labor Day

must be crazy, thereby obliging me to do it.

LITERATURE CITED

EMMEL, T. C. and J. F. EMMEL. 1963. Ecological studies of Rhopalocera
in a high Sierran community — Donner Pass, California. II. Meteoro-
logic influence on flight activity. J. Lepid. Soc. 17: 7-20.

SHAPIRO, A. M. 1962. Colias activity in November and December in Penn¬
sylvania. J. Lepid. Soc. 16: 129-130.

- . 1967. The origin of autumnal “false broods” in common Pierid

butterflies. /. Res. Lepid. 6: 181-193.

- . 1970. Further observations on “false broods” of Colias eurytheme

in central New York ( Lepidoptera: Pieridae). Ent. News. 81: 73-77.

- . 1973. Altitudinal migration of butterflies in the central Sierra Ne¬
vada. J. Res. Lepid. 12: 231-235.

- . 1974. Altitudinal migration of central California butterflies. J. Res.

Lepid. 13: 157-161.

- . 1975a. The temporal component of butterfly species diversity, in

M. L. Cody and J. M. Diamond, eds., Ecology and Evolution of Com¬
munities. Harvard University Press, Cambridge, Mass. pp. 181-195.

- . 1975b. Ecological and behavioral aspects of coexistence in six Cru¬
cifer feeding Pierid butterflies in the central Sierra Nevada. Amer.
Midi. Nat. 93: 424-433.

STEWART, G. R. 1959. Donner Pass and Those Who Crossed It. Lane
Books, Menlo Park, California. 96 pp.

Journal of Research on the Lepidoptera

16(2) :87-130, 1977

Continued from 15( 4) :225-240, 1976.

10. Pinus-Quercus Associes
INDICATOR SPECIES
Chlosyne definita ( 1 )

Callophrys leucania ( 1 )

Thecla augustula (1)

Theda gabatha ( 1 )

Theda clarina
Theda denarius

CHARACTERISTIC SPECIES
Papilio thoas autocles
Phoebis agarithe maxima
Phoebis philea
Phoebis sennae marcellina
Eurema d. daira
Eurema mexicana
Eurema lisa
Eurema nise nelphe
Euptychia gemma freemani
Euptychia mollina
Euptychia gigas
Actinote guatemalena
veraecruzis
Thessalia t. theona
Junonia evarete
Euptoieta hegesia hoffmanni
Mestra amymone

11. Littoral
INDICATOR SPECIES
Anaea morvus boisduvali
CHARACTERISTIC SPECIES
Papilio thoas autocles
Parides p. polyzelus
Colias cesonia
Phoebis sennae marcellina
Phoebis argante
Phoebis agarithe maxima
Eurema albula
Eurema mexicana
Eurema lisa
Eurema nise nelphe
Eurema proterpia
Eurema nicippe
Eurema dina westwoodi

of the Deciduous Woodland

Strymon melinus
Hemiargus i. isola ( 1 )

Baeotes hisbon zonata
Symmachia tricolor hedemanni
Isapis agyrtus hera ( 1 )

Anatole rossi

Hamadryas februa gudula
Hamadryas feronia farinulenta
Hamadryas g. guatemalena
Limenitis iphicla
Libytheana carinenta mexicana
Eumaeus minyas
Calycopis beon
Electrostrymon cyphara
Strymon yojoa
Theda brescia

Hemiargus ceraunus zachaeina
Hemiargus huntingtoni
hannoides
Everes c. comyntas
Leptotes cassius striata
Mesene croceela
Peplia lamis molpe
Woodland

Alorpho peleides montezuma
Agraulis vanillae incarnata
Dryas julia moderata
Heliconius cleobaea zorcaon
Heliconius ismenius telchinia
Heliconius charitonius
vazquezae

Heliconius petiveranus
Anartia jatrophae luteipicta
Anartia fatima venusta
Biblis hyperia aganisa
Hamadryas februa gudula
Hamadryas ferronia farinulenta

87

135

88

GARY N. ROSS

J. Res. Lepid.

Ascia m. monuste
Anteos clorinde
Anteos maerula
Euptychia hesione
Euptychia hermes sosybius
Caligo memnon
Melinaea lilis imitata
Mechanitis polijmnia lycidice
Mechanitis egaensis doryssus

Marpesia chiron
Limenitis iphicla
Limenitis paraeca
Eumaeus minyas
Calycopis beon

Hemiargus ceraunus zachaeina
Everes c. comyntas
Peplia lamis molpe

12. Swamp Forest
INDICATOR SPECIES

Thecla ant incus (1) Eurybia lycisca (2)

CHARACTERISTIC SPECIES
Dimorphia fortunata
Eurema albula
Eurema dina westwoodi
Tithorea harmonia salvadoris
Melinaea lilis imitata
Mechanitis polymnia lycidice
Mechanitis egaensis doryssus
Mechanitis menapis saturata
Hypothyris lycaste dionaea
Ithomia patilla
Oleria paula
Dircenna klugi
Pteronymia cottyto
Greta oto
Greta nero
Taygetes andromeda
Euptychia hesione

Euptychia metaleuca
Euptychia themis
Euptychia hermes sosybius
Euptychia libye
Caligo memnon
Morpho peleides montezuma
Heliconius cleobaea zorcaon
Heliconius ismenius telchinia
Heliconius petiveranus
Heliconius charitonius
vazquezae

Pyrrhogyra hypensor
Pyrrhogyra otolais neis
Hamadryas februa gudula
Hama dry as feronia farinulenta
Hamadryas g. guatemalena
Calycopis beon
Thecla marysas damo

13. Mangrove Woodland
INDICATOR SPECIES
None.

CHARACTERISTIC SPECIES

The same as those for the Swamp Forest.

14. Recently Abandoned Milpas
INDICATOR SPECIES

Tmolus echion echiolus ( 1 ) Mesene margaretta
Thecla ares ( 1 )

136

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

89

CHARACTERISTIC SPECIES

Papilio thoas autocles
Appias drusilla poeyi
Ascia m. monuste
Colias cesonia
Phoebis sennae marcellina
Phoebis philea
Phoebis argante
Phoebis agarithe maxima
Eurema d. daira
Eurema boisduvaliana
Eurema mexicana
Eurema proterpia
Eurema nicippe
Eurema lisa
Eurema nise nelphe
Danaus gilippus strigosus
Dryas julia moderata
Euptoieta hegesia hoffmanni
Chlosyne janais
Chlosyne l. lacinia
Thessalia t. theona

Phyciodes vesta

Phyciodes claudina guatemalena
Phyciodes a. ardys
Phyciodes myia
Phyciodes griseobasolis
Anartia jatrophae luteipicta
Anartia fatima venusta
Dynamine mylitta
Hamadryas februa gudula
Hamadryas g. guatemalena
Marpesia chiron
Limenitis paraeca
Anaea aidea
Calycopis beon
Strymon yojoa
Thecla marsyas damo
Hemiargus ceraunus zachaeina
Hemiargus huntingtoni
hannoides
Everes c. comyntas
Leptotes cassius striata
Peplia lamis molpe

15.

INDICATOR SPECIES

Opsiphanes boisduvalii ( 1 )
Opsiphanes cassiae castaneus
Phyciodes eranites mejicana ( ]
Hamadryas iphthime ( 1 )
Limenitis erotia ( 1 )

Chlorippe pavon
Prepona laertes pallantias (2)
Anaea marthesia (2)
Callophrys goodsoni ( 1 )
Atlides poly be

Pastures

Evenus regalis ( 1 )

Thecla cypria
) Thecla janais (2)

Thecla vibidia ( 1 )

Thecla hecate (2)

Thecla ligurina ( 1 )
Thecla scopas
Ancylusis jurgensenii ( 1 )
Anatole agave ( 1 )
Theope eleutho

137

90

GARY N. ROSS

J. Res. Lepid.

CHARACTERISTIC SPECIES

Papilio thoas autocles
Appias drusilla poeyi
Ascia m. monuste
Colias cesonia
Phoebis sennae marcellina
Phoebis philea
Phoebis argante
Phoebis agarithe maxima
Eurema boisduvaliana
Eurema mexicana
Eurema proterpia
Eurema lisa
Eurema nise nelphe
Eurema dina westwoodi
Eurema nicippe
Danaus gilippus strigosus
Euptychia hermes sosybius
Dryas julia moderata
Euptoieta hegesia hoffmanni
Chlosyne janais
Chlosyne l. lacinia
Thessalia t. theona
Phyciodes vesta
Phyciodes frisia tulcis

Phyciodes claudina guatemalena
Phyciodes a. ardys
Phyciodes myia
Phyciodes griseobasolis
Anartia jatrophae luteipicta
Anartia fatima venusta
Dynamine mylitta
Hamadryas februa gudula
Hamadryas g. guatemalena
Marpesia chiron
Limenitis paraeca
Anaea aidea
Calycopis beon
Strymon yojoa
Heterosmaitia palegon
Thecla marsyas damo
Thecla meton
Thecla tephraeus
Hemiargus ceraunus zachaeina
Hemiargus huntingtoni
hannoides
Everes c. comyntas
Leptotes cassius striata
Peplia lamis molpe

16. Hedgerows

INDICATOR SPECIES

Callophrys miserabilis (2)
Panthiades ochus ( 1 )

Thecla neora (1)

Thecla barajo (2)

CHARACTERISTIC SPECIES

Graphium e. epidaus

Papilio thoas autocles

Par ides p. polyzelus

Eurema albula

Eurema dina westwoodi

Thecla hassan (1)

Thecla hesperitis ( 1 )

Thecla demonassa (2)

Thecla ambrax ( 1 )

Anartia jatrophae luteipicta
Anartia fatima venusta
Metamorpha stelenes biplagiata
Biblis hyperia aganisa
Dynamine mylitta

138

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

91

Oleria paula
Pteronymia cottyto
Euptychia hesione
Euptychia themis
Euptychia hermes sosybius
Euptychia libye
Caligo memnon
Morpho peleides montezuma
Dryas julia moderata
Heliconius cleobaea zorcaon
Heliconius ismenius telchinia
Heliconius petiveranus
Heliconius charitonius
vazquezae

Phyciodes claudina guatemalena
Phyciodes phillyra

Hamadryas februa gudula
Hamadryas g. guatemalena
Marpesia chiron
Limenitis iphicla
Limenitis paraeca
Anaea aidea
Calycopis beon
Strymon yojoa
Heterosmaitia palegon
Thecla marysas damo
Thecla meton
Thecla tephraeus
Charis velutina
Lymnas p. pixe
Peplia lamis molpe

In addition, the data indicate that a second division — less
pronounced than the first but nonetheless significant — can be
made. A comparison of the formations at relatively low altitudes
indicates that the greatest diversity occurs between the species
in the Lower Montane Rain Forest and the remaining forma¬
tions. Indeed, 24 species (13 indicator, 11 characteristic) occur
commonly only within the Lower Montane Rain Forest. This
“uniqueness” is approached by only one other formation, the
Semi-Evergreen Seasonal Forest, which has 25 species (23 indi¬
cator, two characteristic) found commonly only within its bor¬
ders. Thus, a line separating the Lower Montane Rain Forest
from all other formations at relatively low elevations can be
drawn.

In conclusion, my analysis of the butterfly fauna has led me
to the opinion that the classification system employed by Andrle
( 1964 ) does not reflect accurately the existing relationships ( at
least for butterflies) but that the system originally expressed by
Goldman (1951) is more applicable. Following the system em¬
ployed by the latter, I divide the Sierra into two major zones:
first, a Lower Tropical Zone, subdivisible into a Humid Lower
Tropical Subzone (corresponding to the Humid Tropical Upper
Subzone of Andrle), and an Arid Lower Tropical Subzone
(corresponding to the Arid Tropical Zone of Andrle); and sec¬
ond, an Upper Tropical Zone, nondivisible and corresponding to

139

92

GARY N. ROSS

/. Res. Lepid.

the Humid Tropical Upper Subzone of Andrle. These zones and
subzones are defined and characterized in Table III.

BIOTIC PROVINCE

The concept of the Biotic Province, originated by Vestal
( 1914 ) and developed by Dice ( 1943 ) , by definition dictates
that the entire Sierra de Tuxtla fall within only one category.
This, according to Goldman (1951) and Goldman and Moore
(1945) is the Veracruz Biotic Province, which “embraces the
tropical lowlands from eastern San Luis Potosi, southern Tama-
ulipas, and northeastern Puebla, southwesterly through Veracruz
and Tabasco and small portions of northern Oaxaca and Chia¬
pas.” However, I question the validity of including the Sierra
within the “tropical lowlands” of Veracruz since the majority of
the land surfaces within the range have an average elevation in
excess of 1,000 feet and four volcanoes have maximum eleva¬
tions in excess of 3,000 feet. Furthermore, most of the flora and
fauna found at elevations in excess of 2,500 to 3,000 feet seem
to have their affinities with forms common farther south. I con¬
clude, therefore, that a new biotic province should be erected
for the Sierra de Tuxtla.

TABLE III

LIFE ZONES AND CORRESPONDING PLANT

FORMATIONS IN THE SIERRA DE TUXTLA

I. Lower Tropical Zone. — This zone is located from sea level to
approximately 2,500 to 3,000 feet in altitude and probably re¬
ceives an average annual rainfall of less than 150 inches. The
zone is divisible into two subzones.

A. Humid Lower Tropical Subzone. — This subzone is found
principally on the Gulf slopes of the major volcanoes below
2,500 to 3,000 feet where the average annual rainfall probably
averages between 150 and 110 inches. Only one plant forma¬
tion — Lower Montane Rain Forest — is included in this sub¬
zone.

B. Arid Lower Tropical Subzone. — This subzone is found
principally along the coast and on the leeward slopes of the

140

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

93

major volcanoes below 2,500 to 3,000 feet, areas in which the
average annual rainfall probably is less than 110 to 100
inches. This subzone includes the following plant formations:
Littoral Woodland
Mangrove Woodland
Swamp Forest
Savanna

Deciduous Woodland and the Pinus-Quercus Associes
Semi-Evergreen Seasonal Forest and the Bursera-Sabal-
Orbignya Associes

II. Upper Tropical Zone. — This zone is located from approxi¬
mately 2,500 to 3,000 feet in altitude to the peaks of the princi¬
pal volcanoes (3,750 feet, 5,250 feet, and 5,450 feet for Volcans
San Martin Pajapan, Santa Marta, and San Martin Tuxtla, re¬
spectively) where the average annual rainfall probably is in
excess of 150 inches. This zone includes the following plant
formations:

Montane Rain Forest (the Liquidambar-Quercus Associes
being an ecotone)

Montane Thicket
Elfin Woodland

Because the “Miscellaneous Formations” (Recently Aban¬
doned Milpas, Pastures, and Hedgerows) occur throughout the
Sierra where man has settled, I have not assigned these to any
life zone.

AREAL DISTRIBUTION

Faunal-Floral Relationships

A total of 359 species of butterflies representing 133 genera
and eight families now have been recorded from the Sierra de
Tuxtla. Of these species, 258 (72% ) are found primarily in open
and relatively open plant formations throughout the range, and
for the most part were collected in fields, pastures, and along
hedgerows and the margins of forests. In general, members of
the Lycaenidae and Riodinidae visit the blossoms of Cordia
spinescens and Calliandra grandiflora very frequently; members
of the genera Hamadryas, Historis , Smyrna, Gynaecia, Prepona,
and Anaea (Nymphalidae) are attracted to fermenting sap oozing
from the trunks of citrus trees and to fermenting juices of fallen
fruit (principally mangoes); members of the genera Papilio,
Graphium (Papilionidae), Colias, Anteos, Phoebis, Eurema
(Pieridae), Chlosyne, Phyciodes, Diaethria, Dynamine, Marpesia
(Nymphalidae), Hemiargus, Leptotes, and Everes (Lycaenidae)
visit flowers (and damp earth) indiscriminately.

141

94

GARY N. ROSS

J. Res. Lepid.

The interiors of forests with closed canopies are inhabited by
a minority of butterfly species (101 species, 28% of total spe¬
cies). These species belong to 53 genera, 13 subfamilies and
eight families and are listed in Table IV. From this table, cer¬
tain correlations can be made. First, approximately 41% of the
butterfly species inhabiting the forests of the Sierra de Tuxtla
belong to the families Ithomiidae and Satyridae. Second, all
members of the family Ithomiidae and the subfamily Lycoreinae
(Danaidae) occur within forests. Third, more than half the
members of the Pierinae (five species, 55%) and Dismorphiinae
(four species, 66%), (Pieridae), Satyrinae (19 species, 83%)
and Brassolinae (three species, 60%) (Satyridae), and Ama-
thusiinae (two species, 66%) (Nymphalidae) are found within
forests. Fourth, relatively few species in the Lycaenidae (10
species, 11%) and the Riodinidae (13 species, 27%) occur
within forests.

TABLE IV

GENERA OF BUTTERFLIES COLLECTED IN THE
SIERRA DE TUXTLA WITHIN FORESTS
WITH CLOSED CANOPIES

(Numbers behind genera indicate number of species involved;
numbers behind subfamilies indicate total number of species
collected. )

A. Genera in which all species occur within forests
Catasticta (1) Pieridae: Pierinae (9)

Archonias (1) Pieridae: Pierinae
Itaballia (3) Pieridae: Pierinae
Tithorea (1) Ithomiidae Tthomiinae (20)

Melinaea (1) Ithomiidae Tthomiinae
Mechanitis (3) Ithomiidae Tthomiinae
Hypothyris (1) Ithomiidae Tthomiinae
Napeo genes (1) Ithomiidae Tthomiinae
Ithomia (2) Ithomiidae Tthomiinae
Hyposcada (1) Ithomiidae Tthomiinae
Oleria (2) Ithomiidae Tthomiinae
Aeria (1) Ithomiidae Tthomiinae

142

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

95

Dircenna (1) Ithomiidae:Ithomiinae
Episcada (1) Ithomiidae:Ithomiinae
Pteronymia (1) Ithomiidae:Ithomiinae
Greta (3) Ithomiidae:Ithomiinae
Hypoleria (1) Ithomiidaeilthomiinae
Ly corea (1) Danaidae:Lycoreinae (1)

Pierella (1) Satyridae:Satyrinae (23)

Pedaliodes (1) Satyridae:Satyrinae
Dioriste (1) Satyridae:Satyrinae
Eryphanis (1) Satyridae:Brassolinae (5)

Caligo (2) Satyridae:Brassolinae

Polygonia (1) Nymphalidae:Nymphalinae (90)

Myscelia ( 2 ) Nymphalidae : Nymphalinae
Eumaeus (2) Lycaenidae:Lycaeninae (88)

Theorema (1) Lycaenidae:Lycaeninae
Oenomaeus (1) Lycaenidae:Lycaeninae
Perophthalma (1) Riodinidae:Riodininae (42)
Leucochimona (2) Riodinidae:Riodininae
Mesosemia (2) Riodinidae:Riodininae
Eurybia (1) Riodinidae:Riodininae
Thisbe (1) RiodinidaeiRiodininae
Polystichtis (1) RiodinidaeiRiodininae

B. Genera in which half or more species occur within forests

Parides (5) PapilionidaeiPapilioninae (21)

Dismorphia (4) PieridaeiDismorphiinae (6)

Taygetes (4) SatyridaeiSatyrinae (23)

Euptychia (12) SatyridaeiSatyrinae
Morpho (2) Nymphalidae: Amathusiinae (3)

Epiphile (1) Nymphalidae: Nymphalinae (90)

Catonephele (1) Nymphalidae: Nymphalinae
Calycopis (2) Lycaenidae:Lycaeninae (88)

Euselasia (4) Riodinidae:Euselasiinae (6)

C. Genera in which less than half the species occur within forests

Graphium (2) PapilionidaeiPapilioninae (21)

Eurema (3) Pieridae:Coliadinae (22)

Heliconius (1) Nymphalidae :Heliconiinae (17)

Chlosyne (1) Nymphalidae: Nymphalinae (90)

Phyciodes (1) Nymphalidae: Nymphalinae
Limenitis (1) Nymphalidae: Nymphalinae

143

96

GARY N. ROSS

J. Res. Lepid.

Prepona (2) NymphalidaeiNymphalinae
Anaea (3) NymphalidaeiNymphalinae
Thecla (4) LycaenidaeiLycaeninae (88)

Emesis (1) RiodinidaeiRiodininae (42)

Although the species in the previously mentioned families
and subfamilies share a common habitat type (tropical forest),
they nonetheless have different preferred micro-habitats. The
ithomiids usually are found in dank ravines in nonspecific butter¬
fly assemblages and in the vicinities of four flowering plants:
Tournefortia glabra, Eupatorium macrophyllum, E. pittieri, and
Psychotria padifolia. The satyrids are more randomly distributed
throughout the forests, Taygetes spp., Dioriste tauropolis, Ped-
iliodes pisonia circumducta, Eryphanis aesacus, and Caligo spp.
preferring montane formations and Euptychia spp. preferring
less dense forest formations. The single species of Lycorea
(DanaidaeiDanainae) is found in relatively open areas within
the Semi-Evergreen Seasonal Forest and the Lower Montane
Rain Forest and usually in small butterfly assemblages. The five
species (three genera) of Pierinae are found in the Lower Mon¬
tane Rain Forest and the Semi-Evergreen Seasonal Forest and
usually within nonspecific butterfly assemblages — Catisticta n.
nimbice preferring less dense areas and Archonias tereas and
Itaballia spp. preferring more dense and shaded locales. The four
species of Dismorphia (DismorphiinaeiPieridae) prefer the mon¬
tane formations — D. praxinoe and D. fortunata the Lower Mon¬
tane Rain Forest and D. euryope and D. nemesis the Elfin Wood¬
land and Montane Thicket. The two species of Morpho (Ama-
thusiinaeiNymphalidae) have very dissimilar ecologies — M.
polyphemus luna prefers the montane formations above 3,000
feet and M. peleides montezuma prefers the Lower Montane
Rain Forest and other forests at relatively low elevations. The
ten species (five genera) of lycaenids and 13 species (eight
genera) of riodinids are found in a variety of forests; all seem
to prefer the relatively bright sections.

ALTITUDINAL DISTRIBUTION

The maximum altitude of the Sierra is relatively low (5,450
feet) and thus vertical temperature change is not great. It is
reasonable to assume that the entire Sierra falls within the
normal altitudinal range of most butterfly species. (Indeed,

144

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

97

species that are common in sunny fields along the coast and at
relatively low altitudes frequently were seen sailing over the
peaks of the highest volcanoes on sunny days.) Yet, as stated
previously, the greatest diversity in the butterfly fauna occurs
between the Upper and Lower Tropical Zones and so the obvious
conclusion is that plant formations with their characteristic
plants, some of which probably serve as larval and adult food
plants, seem to be the principal factor governing butterfly dis¬
tributions. In general, the majority of the butterfly species (320
species, 89%) in the Sierra are primarily residents of the plant
formations below 2,500 to 3,000 feet ( the Lower Tropical Zone ) ;
only 39 species (11% ) are commonly found in the forests above
approximately 3,000 feet in elevation (the Upper Tropical Zone).

Climatic Relationships

SEASONAL VARIATION IN
RUTTERFLY POPULATION DENSITIES

Although the climate of the Sierra de Tuxtla is relatively
mild and uniform, enough diversity exists to produce a notice¬
able seasonal fluctuation in butterfly populations. In general,
populations reach maximum densities in August, September,
and October and minimum densities in January, February,
March, and April. In fact, many species virtually disappear dur¬
ing the winter and spring months, even at relatively low eleva¬
tions. Although many species of butterflies are known to migrate
to other areas during the winter months (Williams, 1930, 1958),
relatively few of these species (17 species, approximately 5%)
occur in the Sierra. These are: Graphium philolaus (Papilioni-
dae), Eurema nicippe, E. albula, E. lisa , Phoebis statira, P. trite ,
P. philae, P. argante, P. sennae, Anteos maerula, Ascia monuste
(Pieridae), Agraulis vanillae, Vanessa virginiensis, Junonia
evarete, Marpesia chiron (Nymphalidae), Danaus plexippus,
and D. gilippus (Danaidae). The majority (59%) of these be¬
long to the family Pieridae.

During my residence in the Sierra, I saw butterfly move¬
ments (which possibly could be termed migrations) in July and
August involving principally Phoebis spp. and Marpesia chiron.
Each day hundreds of individuals were observed as they flew
usually between ten and 30 feet above the ground and in a

145

98

GARY N. ROSS

J. Res. Lepid.

northeasterly direction toward the Gulf. In addition, during the
fall months populations of Danaus plexippus and Vanessa vir-
giniensis increased, possibly because of an influx of migrants
from the north. Therefore, because of the relatively few migrant
species (approximately 5%), and because of a population in¬
crease during the fall of at least two species ( Danaus plexippus
and Vanessa virginiensis) , I conclude that migrations are an in¬
significant factor in the reduction of the winter butterfly popu¬
lations in the Sierra de Tuxtla.

An alternative and more plausible explanation is that many
species undergo an egg, a larval, or pupal diapause during the
winter period, possibly as a result of decreased amounts of day¬
light, reduced temperatures, and reduced rainfall, or, any com¬
bination of these. Unfortunately, there is little data to support
this hypothesis for I observed diapause (larval) only in one
species — Anatole rossi (Riodinidae; Ross, 1966).

Although spring-dry season population minima are the gen¬
eral rule, a few exceptions do exist. The seven species of
Graphium (Papilionidae:Papilioninae), Dismorphia jethys (Pier-
idae:Dismorphiinae), and Actinote guatemalena veraecruzis
(Nymphalidae:Acraeinae) have maximum population densities
in spring and early summer. All of these species disappear sub¬
sequent to the commencement of the summer rains. Also, Morpho
theseus justiciae (Nymphalidae: Amathusiinae ) and Actinote
leucomelas ( Nymphalidae :Acraeinae) are common in March and
April but then disappear completely only to reappear in Sep¬
tember, October, and November. Thus, these last two species
seem either to be double brooded and to have relatively lengthy
immature stages or to undergo diapause during some stage in
their life cycles. Philaethria cl. clido is common only in the fall
months!

DAILY VARIATIONS IN
BUTTERFLY POPULATION DENSITIES

During any given season, daily cycles in population densi¬
ties are evident. Generally, most butterfly species are strongly
photopositive and reach maximum activity levels between 10:00
A.M. and noon. However, during the periodic cool spells during
the winter and on days of heavy cloud cover, maximum activity
periods are delayed from one to two hours.

146

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

99

The notable exceptions to the previous generalization are the
two species of Caligo ( Satyridae:Brassolinae), Eurybia lycisca
( Riodinidae:Riodininae), and all members of the Ithomiidae.
The caligos are decidedly crepuscular and frequently wander out
of the forest habitats and into more open areas at dusk; the two
specimens of Eurybia lycisca were collected as they flew in a
very dark, dense thicket at 6:30 P.M.; and members of the
Rhomiidae are active under practically all weather conditions
( including fog and light rain ) in addition to practically all hours
of daylight ( dawn to dusk ) .

Faunal Relationships

AFFINITIES AND ORIGINS

Although the Sierra has been open to animal and plant move¬
ments from the north and the range is sufficiently near the
northern limits of the Neotropical region to permit an influx
of Nearctic forms, the fauna and flora have remained essentially
tropical. In his avi-faunal investigations, Andrle (1964) states
that 43% of the avian species recorded in the Sierra have south¬
ern origins as compared to 26% with more northern affinities
(the remaining percentage represents species with unknown or
uncertain origins). Unfortunately, the science of butterfly zoo¬
geography has not advanced to the state whereby the origins
and affinities of most genera and species groups can be defined
(see Hovanitz, 1958). However, several generalizations, cor¬
relations, and speculations can be made about the Sierra’s butter¬
flies.

First, of the 359 species representing 133 genera, only 77
species (21%) representing 45 genera (34%) ever have been
recorded from within the borders of the United States (refer¬
ences: dos Passos, 1964; Ehrlich and Ehrlich, 1961). These spe¬
cies, which are listed in Table V, for the most part are found in
open, sunny areas throughout the Sierra. Furthermore, of these
77 species, 47 (61% ) are known to be breeding residents (refer¬
ence: Klots, 1951); the remaining species, 30 (39%), enter the
United States only as occasional strays. Thus, the majority of
the species of butterflies in the Sierra (79% ) do not occur even
as strays just 500 miles to the north.

147

100

GARY N. ROSS

/. Res. Lepicl.

TABLE V

SPECIES OF BUTTERFLIES COLLECTED IN THE
SIERRA DE TUXTLA KNOWN TO OCCUR WITHIN
THE BORDERS OF THE UNITED STATES

Papilionidae (5 species, 25%
Battus polydamas
Parides areas mylotes
Papilio polyxenes asterius

Pieridae (20 species, 54% of
Appias drusilla poeyi
Ascia m. monuste
Colias cesonia
Anteos clorinde
Anteos maerula
Phoebis sennae marcellina
Phoebis philea
Phoebis argante
Phoebis agarithe maxima
Phoebis statira jada,

of represented species )
Papilio thoas autocles
Papilio anchisiades idaeus

represented species)
Eurema d. daira
Eurema boisduvaliana
Eurema mexic-ana
Eurema salome
Eurema proterpia
Eurema lisa
Eurema nise nelphe
Eurema dina westwoodi
Eurema nicippe
Nathalis iole

Danaidae (4 species, 100% of represented species)

Danaus p. plexippus Danaus eresimus montezuma

Danaus gilippus strigosus Lycorea ceres atergatis

Satyridae (2 species, 8% of represented species)

Euptychia gemma freemani Euptychia hermes sosybius

Nymphalidae (31 species, 29%
Dryadula phaetusa
Agraulis vanillae incarnata
Dryas julia moderata
Heliconius petiveranus
Heliconius charitonius
vazquezae

Euptoieta hegesia hoffmanni
Chlosyne janais

of represented species)
Chlosyne l. lacinia
Chlosyne definita
Thessalia t. theona
Phyciodes vesta
Phyciodes frisia tulcis
Vanessa virginiensis
Junonia evarete

148

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

101

TABLE V

Nymphalidae ( continued )
Anartia jatrophae luteipicta
Anartia fatima venusta
Metcimorpha stelenes
biplagiata
Hypanartia lethe
Biblis hyperia aganisa
Mestra amiymone
Eunica monima
Dynamine dyonis
Hamadryas febma gudula

Lycaenidae (16 species, 21% c
Eumaeus minyas
Chlorostrymon s. simaethis
Chlorostrymon telea
Calycopis beon
Tmolus echion echiolus
T molus azia
Callophrys miserabilis
Strymon melinus

( continued )

Hamadryas feronia farinulenta

Marpesia chiron

Marpesia petreus

Chlorippe pavon

Hist oris odius

Anaea aidea

Anaea pithyusa

Libytheana carinenta mexicana

of represented species)

Strymon yojoa
Strymon columella istapa
Strymon bazochii
Hemiargus ceraunus zachaeina
Hemiargus i. isola
Everes c. comyntas
Leptotes cassius striata
Celastrina argiolus gozora

Second, of the 133 genera found in the Sierra, 58 (44%)
have been recorded in the United States (reference: dos Passos,
1964). These, in addition to the number of species represented
in the Sierra, the United States, and the relative number found
within the Neotropical regions, are listed in Table VI. An analy¬
sis of the data presented in this table reveals that of the 58
genera, only 19 (33%) are represented by a greater number of
species in the United States than in the Sierra, and that of these
19, only four (7%) — Colias (Pieridae), Polygonia (Nymphali¬
dae), Vanessa (Nymphalidae), and Callophrys (Lycaenidae) —
have a greater representation in the United States (Nearctic
region) than in the entire Neotropical region. Lacking additional
information, one may assume with some justification that the
center of origin of a genus corresponds to the area containing
the greatest number of species (Savage, 1958). Thus, I conclude
that at least four genera of the 133 (3%) represented in the
Sierra probably have Nearctic origins or at least have their great¬
est affinities with Nearctic forms and that the remaining genera
(129, 97%) probably have Neotropical origins or have their
greatest affinities with Neotropical forms.

149

102

GARY N. ROSS

]. Res. Lepid.

Third, an anlysis of the 37 species (excluding new, endemic
species) that have not been recorded from the Sierra according
to Hoffmann (1940) — 10 range extensions within the state of
Veracruz, 18 new state listings, and nine new national listings
(Table VII) — reveals that 76% of the new listings (all but the
10 intrastate range extensions) are species with known distri¬
butions that are farther south than the Sierra. The remaining

TABLE VI

COMPARISON OF BUTTERFLY GENERA COMMON
TO THE SIERRA DE TUXTLA, THE UNITED STATES,
AND THE NEOTROPICAL REGIONS

Genus (and

No. of species

No. of species

Relative no. of

family )

in the Sierra
de Tuxtla

in the United
States

species in the
Neotropical regions
as compared to the
Nearctic regions

Papilionidae

Graphium

7

1

greater

Battus

3

2

greater

Parides

6

1

greater

Papilio

5

21

greater

Pieridae

Dismorphia

6

1

greater

Appias

1

1

greater

Ascia

1

2

greater (4)

Colias

1

16

lesser

Anteos

2

2

same (2)

Phoebis

7

6

greater

Eurema

11

10

greater

Nathalis

1

1

same ( 1 )

Danaidae

Danaus

3

3

same (3)

Lycorea

1

1

greater

Satyridae

Euptychia

15

9

greater

Nymphalidae

Dryadula

1

1

same (1)

Agraulis

1

1

same (1)

Dryas

1

1

same (1)

Heliconius

11

2

greater

Euptoieta

1

2

greater (3)

150

103

16(2):87-130, 1977 BUTTERFLIES OF TUXTLA

Chlosyne 5

Thessalia 1

Phyciodes 10

Polygonia 1

Vanessa 1

Junonia 1

Anartia 2

Hy panart ia 2

Biblis 1

Mestra 1

My s celia 2

Eunica 1

Diaethria 2

Dynamine 2

Hamadryas 6

Marpesia 4

Limenitis 9

Chlorippe 3

Hist oris 1

Smyrna 1

Anaea 11

Libytheana 1

Lycaenidae

Eumaeus 2

Chlorostrymon 2
Calycopis 4

Tmolus 3

Callophrys 7

Atlides 1

Panthiades 1

Strymon 6

Thecla 47

Hemiargus 3

Everes 1

Leptotes 1

Celastrina 1

Riodinidae

Euselasia 6

Calephelis 3

Emesis 4

29

greater

2

greater (?)

11

greater

10

lesser

4

lesser

2

greater

2

greater

1

greater

1

same (1)

2

same (2)

2

greater

2

greater

2

greater

1

greater

4

greater

4

greater

7

greater

1

greater

2

same (2)

1

greater

5

greater

2

greater

2

greater (3)

3

greater

2

greater

2

greater

24

lesser

1

greater

1

greater

12

greater

2

greater

3

greater

1

greater (?)

2

greater

1

same (1)

1

greater

9

greater

2

greater

151

104

GARY N. ROSS

J. Res. Lepid.

TABLE VII

NEW BUTTERFLY RECORDS FOR THE
SIERRA DE TUXTLA
(EXCLUDING NEW ENDEMIC SPECIES)

Species Nearest Previous Associated Plant

Recorded Locale Formations

I. Intrastate range
Graphium branchus
(Papilionidae)
Graphium belesis
(Papilionidae)
Graphium agesilaus
neosilaus
(Papilionidae)
Dismorphia euryope
(Pieridae)
Hamadryas g.
guatemalena
(Nymphalidae)
Prepona brooksiana
(Nymphalidae)
Eumaeus debora
(Lycaenidae)

Chlorostrymon s.
simaethis
(Lycaenidae)
Electrostrymon
cyphara
(Lycaenidae)

extensions

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Coatepec, Veracruz

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Sierra Madre Oriental
(Veracruz)

Thecla hecate
(Lycaenidae)

Sierra Madre Oriental
(Veracruz)

II. New state records

Taygetes kerea
(Satyridae)

Chiapas, Sierra Madre
del Sur (Mexico)

Phyciodes griseo-
basolis

(Nymphalidae)
Catagramma lyca
(Nymphalidae)
Limenitis erotia
(Nymphalidae)

Chiapas (Mexico)

Tabasco, Chiapas,
Oaxaca (Mexico)
Chiapas (Mexico)

Semi-Evergreen
Seasonal Forest
Semi-Evergreen
Seasonal Forest
No specific formation

Montane Thicket,
Elfin Woodland
Unrestricted

Montane Thicket

Montane Rain Forest,
Montane Thicket,
Elfin Woodland
Hedgerows

Lower Montane Rain
Forest, Semi-
Evergreen Seasonal
Forest
Pastures

Deciduous Woodland
and the Pinus-
Quercus Associes
Recently Abandoned
Milpas, Pastures

Lower Montane Rain
Forest
Pastures

152

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

105

Species

Limenitis sentia
(Nymphalidae)
Anaea proserpina
(Nymphalidae)
Strymon melinus
(Lycaenidae)

Thecla thales
( Lycaenidae )

Thecla denarius
(Lycaenidae)

Thecla ter a
(Lycaenidae)
Thecla politus
(Lycaenidae)
Perophthalma tullius
lasius

(Riodinidae)
Leucochimona v.
vestalis
(Riodinidae)
Isapis agyrtus hera
(Riodinidae)

Celephelis sp. 2
(Riodinidae)

Calydna venusta
(Riodinidae)
Ernesia lupina
(Riodinidae)
Polystichtis sudias
(Riodinidae)

III. New national
Epiphile plutonia
(Nymphalidae)
Limenitis oherthuri
(Nymphalidae)

Recorded Locale
Nearest Previous

Yucatan (Mexico)
Chiapas (Mexico)
Oaxaca (Mexico)

Chiapas (Mexico)

Tabasco (Mexico)

Chiapas (Mexico)

Pacific Coast of
Mexico

Chiapas (Mexico)

Chiapas (Mexico)

Chiapas (Mexico)

Tabasco (Mexico)

Oaxaca (Mexico)
Guerrero (Mexico)
Tabasco (Mexico)

records

Guatemala

Guatemala

Associated Plant
Formations

Swamp Forest,

Pastures

Montane Thicket,

Elfin Woodland
Pinus-Quercus Associes
of the Deciduous
Woodland

Liquidambar-Quercus
Associes of the
Montane Rain Forest
Pinus-Quercus Associes
of the Deciduous
Woodland

Lower Montane Rain
Forest

Montane Rain Forest

Liquidambar-Quercus
Associes of the Mon¬
tane Rain Forest
Lower Montane Rain
Forest, Montane
Rain Forest
Pinus-Quercus Associes
of the Deciduous
Woodland

Recently Abandoned
Milpas, Pastures,
Hedgerows
Semi-Evergreen
Seasonal Forest
Semi-Evergreen
Seasonal Forest
Liquidambar-Quercus
Associes of the Mon¬
tane Rain Forest,
Montane Rain Forest

Montane Thicket

Lower Montane Rain
Forest

153

106

GARY N. ROSS

J. Res. Lepid.

Species

Nearest Previous

Associated Plant

Recorded Locale

Formations

Calycopis pisis

Guatemala

Liquidambar-Quercus

(Lycaenidae)

Associes of the Mon¬
tane Rain Forest

Thecla ares

Guatemala

Recently Abandoned

(Lycaenidae)

Milpas

Thecla mulucha

Guatemala

Semi-Evergreen

(Lycaenidae)

Seasonal Forest

Thecla amhrax

Nicaragua

Hedgerows

(Lycaenidae)

Thecla dodava

Panama

Montane Rain Forest

(Lycaenidae)

Thecla tamos

Costa Rica

Deciduous Woodland,

(Lycaenidae)

Montane Rain Forest

The ope eleutho

Panama

Pastures, Pinus-

(Riodinidae)

Quercus Associes of
the Deciduous
Woodland

percentage (24%) of new listings, representing ten species, are
of species with distributions that are only slightly farther north
than the Sierra. These ten species belong to genera that prob¬
ably have had Neotropical origins.

In summary, I conclude that the butterfly fauna of the Sierra
de Tuxtla is essentially tropical.

ENDEMISM

Following the Tertiary uplift of the Sierra de Tuxtla, the
area has been subjected to relatively few physical disturbances.
Other than several volcanic eruptions, ash falls, and lava flows,
the flora has had a considerable span of time in which to develop
and mature. Even during the relatively cool periods of the
Pliocene and Pleistocene the proximity of the Gulf of Mexico
with its warm, moist winds, probably caused a relatively small
decrease in the Sierra’s average yearly temperature ( Dorf , 1959 ) .
However, during that period a slight shift in floral (and faunal)
elements conceivably took place; specifically, an extension of
the previously minor subtropical elements (Griscom, 1932, 1950)
and possibly even the establishment of new immigrants from
more northern areas. Subsequent to the Pleistocene, the climate
has ameliorated, which in turn has caused the retreat of the

154

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

107

subtropical floral elements to relatively high elevations of the
major volcanoes (and possibly the elimination of some forms)
and the expansion once again of the more tropical elements (see
Griscom, 1932, 1950). Today these latter predominate. Thus,
since the Pleistocene relatively stable conditions have existed in
the Sierra and there probably have been no major modifications
in the flora and fauna by physical elements except minor vol¬
canic disturbances and hurricanes — both of which have been
relatively rare in occurrence. This stability in environment com¬
bined with geographic isolation make the area ideally suited
for endemism both in flora and fauna. Firschein and Smith
(1956) state that at least eight endemic forms of amphibians and
reptiles have been reported from the range. Wetmore (1943)
listed five birds and Lowery & Newman (1949) one additional
bird endemic to the Sierra.

My investigations of the butterfly fauna revealed five en¬
demic forms: three species, one subspecies, and one form (two
of the species and the one subspecies still remain undescribed).
These endemics are listed in Table VIII. The majority of these
forms (four species, 89%) belong to the families Lycaenidae
and Riodinidae, many members of which are known to have
fairly restricted distributions. Furthermore, of these five endem¬
ics two occur in the high montane forests (Upper Tropical
Zone) and three in the Lower Tropical Zone (Arid Lower Tropi¬
cal Subzone). All endemics but one (Callophrys nr. longula)
belong to genera that never have been recorded within the Unit¬
ed States and which I conclude have had Neotropical origins.

TABLE VIII

BUTTERFLIES ENDEMIC TO THE SIERRA DE TUXTLA

Associated Plant

Species Endemic Form Formations

Morphotheseus Form (but probably Montane Rain Forest,

justiceae form

good subspecies )

Montane Thicket.

schwezeri

Elfin Woodland

(Nymphalidae)
Callophrys nr.

Species

Elfin Woodland

longula
( Lycaenidae )
Thecla nr. antincus

Species

Swamp Forest

(Lycaenidae)
Anatole rossi

Species

Pinus-Quercus As-

(Riodinidae)

socies of the De-

ciduous Woodland

155

108

GARY N. ROSS

J. Res. Lepid.

Theope eleutho Subspecies Pinus-Quercus As-

n. spp. socies of the De-

(Riodinidae) ciduous Woodland,

Pastures

REFERENCES CITED

AMADON, D. & D. R. ECKELBERRY. 1955. Observations on Mexican
birds. Condor 57:65-80.

ANDRLE, R. F. 1964. A biogeographical investigation of the Sierra de
Tuxtla in Veracruz, Mexico. Unpublished dissertation. Louisiana State
University. 236 pp.

BEARD, J. S. 1944. Climax vegetation in tropical America. Ecology 25:
127-158.

- . 1949. The natural vegetation of the Windward and Leeward Is.

Oxford Forestry Memoir No. 21. 192. pp.

- . 1953. The savanna vegetation of northern tropical America. Ecol.

Mon. 23:149-215.

- . 1955. The classification of tropical American vegetation types.

Ecology 36:89-100.

BUDOWSKI, G. 1959. The ecological status of fire in tropical American
lowlands. In “Actas del XXXIII Con.gr. Intern, de Americanistas” 1:
264-278. San Jose, Costa Rica.

CHERMOCK, R. L. 1950. A generic revision of the Limenitini of the world.
Amer. Midi. Nat. 43:513-569.

CLEMENTS, F. E. 1936. Nature and structure of the climax. Jour. Ecol.
24:252-284.

CLENCH, H. K. 1955. Revised classification of the butterfly family Lyca-
enidae and its allies. Ann. Cam. Mus. 33:261-274.

- . 1964. A new species of Riodinidae from Mexico. Jour. Res. Lepi-

dop. 3:73-80.

COMSTOCK, W. P. 1961. Butterflies of the American tropics, the genus
Anaea, Lepidoptera, Nymphalidae. Amer. Mus. Nat. Hist., New York.
214 pp.

DAVIS, L. I. 1952. Tropical woods. Sixteenth breeding bird census. Aud.
Field Notes 6:314-315.

DICE, L. R. 1943. The biotic provinces of North America. Univ. Mich.
Press. Ann Arbor, Michigan. 78 pp.

DICKEY, D. R. & A. J. VAN ROSSEN. 1938. The birds of El Salvador.

Field Mus. Nat. Hist., Zool. Ser. 23:1-609.
dos PASSOS, C. F. 1964. A synonymic list of the Nearctic Rhopalocera.

Mem. No. 1. The Lepidop. Soc. 145 pp.

EDWARDS, E. P. & R. E. TASHIAN. 1959. Avifauna of the Catemaco basin
of southern Veracruz, Mexico. Condor 61:325-337.

EHRLICH, P. R. & A. N. EHRLICH. 1961. How to know the butterflies.

Wm. C. Brown Co. Pub. Dubuque, Iowa. 262 pp.

EMSLEY, M. 1963. A morphological study of imagine Heliconiinae (Lep.:
Nymphalidae) with a consideration of the evolutionary relationships
within the group. Zoologica 48:85-131.

FIRSCHEIN, I. L. 1950. A new toad from Mexico with a redefinition of
the cristatus group. Copeia 2:81-87.

FIRSCHEIN, I. L. & H. M. Smith. 1956. A new fringe-limbed Hyla ( Am¬
phibia: Anura) from the new faunal district of Mexico. Herpetologica
12:17-21.

156

16(2) -.87-130, 1977

BUTTERFLIES OF TUXTLA

109

FORBES, W. T. M. 1944. The genus Phyciodes ( Lepidoptera, Nymphali-
nae). Ent. Amer. 24:139-207.

FOX, R. M. 1956. A monograph of the Ithomiidae (Lepidoptera). Part I.
Bull. Am. Mus. Nat. Hist. 111:1-76.

FRIEDLAENDER, I. 1923. Uber das Vulkangebiet von San Martin Tuxtla in
Mexiko. Zeitschr. Vulkanologie 7:162-187.

GARCIA, J. A. 1835. Eruptionen des Vulkanes von Tustla in den Jahren
1664 und 1793. Neues Jahrb. Minerologie, Geognosie, Geol., und
Petrefaktenkunde, pp. 40-45.

GARTH, J. S. & J. W. TILDEN. 1963. Yosemite butterflies: an ecological
survey of the butterflies of the Yosemite sector of the Sierra Nevada,
California. Jour. Res. Lepidop. 2:1-96.

GODMAN, F. C. & O. SALVIN. 1879-1901. Biologia Centrali-Americana.

Insecta. Lepidoptera-Rhopalicera. 2 vols. London. 487 pp.; 782 pp.
GOLDMAN. E. A. 1920. The mammals of Panama. Smiths. Mis. Coll. 69:
1-309.

- . 1951. Biological investigations in Mexico. Smiths. Misc. Coll. 115:

1-476.

GOLDMAN, E. A. & R. T. MOORE. 1945. The biotic provinces of Mexico.
Jour. Mamm. 26:347-360.

GOODNIGHT, C. J. & M. L. GOODNIGHT. 1954. The opioionid fauna of
an isolated volcano in southeastern Veracruz. Trans. Am. Microscop.
Soc. 73:344-350.

GRISCOM, L. 1932. The distribution of bird-life in Guatemala. Bull. Am.
Mus. Nat. Hist. 64:1-439.

- . 1950. Distribution and origin of the birds of Mexico. Bull. Mus.

Comp. Zool. 103:341-382.

HEPBURN, H. R. & G. N. ROSS. 1964. Collembola from Mexico. Ent.
News 75:219-220.

HIGGINS, L. G. 1960. A revision of the Melitaeine genus Chlosyne and
allied species (Lepidoptera, Nymphalinae ) . Trans. R. Ent. Soc. Lond.
112:381-467.

HOVANITZ, W. 1958. Distribution of butterflies in the New World. In
“Zoogeography.” Edited by V. L. Hubbs. Am. Ass. Adv. Sci. Pub. No.
51. 509 pp.

KLOTS, A. B. 1931. A generic revision of the Pieridae (Lepidoptera). Ent.
Amer. 12:139-204; 205-242.

- . 1951. A field guide to the butterflies of North America, east of the

Great Plains, Houghton Mifflin Co., Boston, Mass. 349 pp.

KoPPEN, W. P. 1936. Handbuch der klimatologie. Vol. I, Part C, 44 pp.

Verlag. von Gebriider Borntraeger. Berlin.

LEOPOLD, A. S. 1950. Vegetation zones of Mexico. Ecology 31:507-518.

- . 1959. Wildlife of Mexico; the game birds and mammals. Univ.

Calif. Press. Berkeley, Calif. 568 pp.

LOWERY, G. H., JR. & W. W. DALOUEST. 1951. Birds from the state of
Veracruz, Mexico. Univ. Kan. Pub., Mus. Nat. Hist. 3:531-649.
LOWERY, G. H., JR. & R. J. NEWMAN. 1949. New birds from the state
of San Luis Potosi and the Tuxtla Mountains of Veracruz, Mexico.
Occ. Papers Mus. Zool. La. State Unw. 22:1-10.

MEDEL Y ALVARADO, L. 1963. Historia de San Andres Tuxtla, 1532-
1950. Coleccion Suma Veracruzana, Serie Historiografia. 2 vols. Edi¬
torial Citlaltepetl. Mexico D.F., Mexico. 555 pp.; 627 pp.
MALGAREJO VIVANCO, J. L. 1960. Breve historia de Veracruz. Biblio.
Facul. Filos. Letras, Univ. Veracruzana. Xalapa, Mexico. 268 pp.

157

110

GARY N. ROSS

J. Res. Lepid.

MERRIAM, C. H. 1892. The geographic distribution of life in North Amer¬
ica with special reference to the Mammalia. Proc. Biol. Soc. Wash.
7:1-64.

MERRIFIELD, R. G., R. R. FOIL, & T. HANSBROUGH. 1964. The in¬
fluence of fertilization on growth and nutrient content of loblolly pine.
Forestry: Res. Rep. N. La. Hill Farm Exp. Sta. 1964:13-30.

MICHENER, C. D. 1942. A generic revision of the Heliconiinae (Lepidop-
tera, Nymphalidae). Am. Mus. Novit. 1197:1-8.

MOCINO SUAREZ DE FIGUEROA, J. M. 1870. Informe de Don Jose
Moziho sobre la erupcion del Volcan de San Martin, Tuxtla (Vera
Cruz) occurido en el ano 1793. Bol. Soc. Geogr. Estad. Rep. Mex.
2:62-70.

MUNROE, E. & P. R. EHRLICH. 1960. Harmonization of concepts of
higher classification of the Papilionidae. Jour. Lepidop. Soc. 14:169-
175.

MURRAY, G. E. 1961. Geology of the Atlantic and Gulf Coastal Province
of North America. Harper and Brothers. New York, N.Y. 692 pp.

PESSIN, L. J. 1937. The effect of nutrient deficiency on the growth of
longleaf pine seedlings. Occ. Papers Sth. For. Exp. Sta. 65. New Or¬
leans, La. 7 pp.

PYBURN, W. F. 1963. Observations on the life history of the tree-frog,
Phyllomedusa callidryas (Cope). Texas Jour. Sci. 15:155-170.

- . 1964. Breeding behavior of the leaf-frog, Phyllomedusa callidryas,

in southern Veracruz. Year Book Am. Philos. Soc. 1964:291-294.

- . 1966. Breeding activity, larvae and relationship of the treefrog

Hyla phaeota cyanosticta. Southw. Natur. 11:1-18.

ROSS, G. N. 1963. Evidence for lack of territoriality in two species of
Hamadryas (Nymphalidae). Jour. Res. Lepidop. 2:241-246.

- . 1964a. An annotated list of butterflies collected in British Honduras

in 1961. Jour. Lepidop. Soc. 17:11-26.

- . 1964b. Life history studies on Mexican butterflies. I. Notes on the

early stages of four papilionids from Catemaco, Veracruz. Jour. Res.
Lepidop. 3:9-18.

- . 1964c. Life history studies on Mexican butterflies. II. Early stages

of Anatole rossi, a new myrmecophilous metalmark. Jour. Res. Lepidop.
3:81-94.

- . 1964d. Life history studies on Mexican butterflies. III. Nine Rho-

palocera (Papilionidae, Nymphalidae, Lycaenidae) from Ocotal Chico,
Veracruz. Jour. Res. Lepidop. 3:207-229.

- . 1966. Life history studies on Mexican butterflies. IV. The ecology

and ethology of Anatole rossi, a myrmecophilous metalmark (Lepidop-
tera: Riodinidae). Ann. Ent. Soc. Am. 59:985-1004.

SAVAGE, D. E. 1958. Evidence from fossil land mammals on the origin
and affinities of the western Nearctic fauna. In “Zoogeography.” Edited
by C. L. Hubbs. Am. Ass. Adv. Sci. Pub. No. 51. 509 pp.

SCHIEFERDECKER, A. A. G. & J. N. TSCHOPP. 1922. Geological report
on the Veracruz embayment and the Veracruz isthmus region down to
the country of Minatitlan. Unpublished Isthmus Geol. Rep. No. 73. 32
PP-

SCLATER, P. L. 1897. List of additional species of Mexican birds, obtained
by M. Auguste Salle from the environs of Jalapa and San Andres
Tuxtla. On a collection of birds received by M. Salle from southern
Mexico. Proc. Zool. Soc. Lond. Part 25:201-207; 226-230.

SEARS, P. B. 1952. Palynology in southern North America I: archeological
horizons in the basins of Mexico. Bull. Geol. Soc. Am. 63:241-254.

158

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

111

SEITZ, A. 1923. Macrolepidoptera of the world. Vol. 5. The American
Rhopalocera. Stuttgart, Germany. 1139 pp.

STOATE, T. N. 1950. Nutrition of the pine. Bull. For. Bur. Aust. 30. 61 pp.
VESTAL, A. G. 1914. Internal relations of terrestrial associations. Am.
Natur. 48:413-445.

WALLACE, A. R. 1876. The geographical distribution of animals. 2 vols.

Macmillan and Co,. London, England. 503 pp.; 607 pp.

WELLING, E. C. 1966. Flight habits of Morpho theseus justitiae. Jour.
Lepidop. Soc. 20:95-101.

WETMORE, A. 1943. The birds of southern Veracruz, Mexico. Proc. Nat.
Mus. 93:215-340.

WILLIAMS, C. B. 1958. The migration of butterflies. Oliver and Boyd.
London, England. 473 pp.

- . 1958. Insect migrations. Macmillan Co., New York, N.Y. 235 pp.

APPENDICES
Appendix A

CHECK-LIST OF BUTTERFLY SPECIES COLLECTED
IN THE SIERRA DE TUXTLA
FAMILY PAPILIONIDAE
SUBFAMILY Papilioninae

TRIBE Graphiini
SUBTRIBE Graphiiti

1. Graphium phaon ( Boisduval )

2. Graphium branchus ( Doubleday )

3. Graphium belesis ( Bates )

4. Graphium philolaus (Boisduval)

5. Graphium epidaus epidaus (Doubleday, Westwood, &

Hewitson )

6. Graphium agesilaus neosilaus ( Hoffer )

7. Graphium calliste calliste ( Bates )

TRIBE Troidini
SUBTRIBE Battiti

8. Battus polydamas ( Linnaeus )

9. Battus belus varus ( Kollar )

10. Battus laodamas copanae (Reakirt)

159

112

GARY N. ROSS

/. Res. Lepid.

SUBTRIBE Troiditi

11. Parides photinus (Doubleday )

12. Parides montezuma (Westwood)

13. Parides polyzelus poly zelus (Felder)

14. Parides sesostris zestos ( Gray )

15. Parides iphidamas (Fabricius)

16. Parides areas mylotes ( Bates )

TRIBE Papilionini

17. Papilio polyxenes asterius Stoll

18. Papilio thoas autocles Rothschild & Jordan

19. Papilio androgens epidaurus Godman & Salvin

20. Papilio anchisiades idaeus Fabricius

21. Papilio victor inus victorimis Doubleday

FAMIFY PIERIDAE
SUBFAMIFY Dismorphiinae

22. Dismorphia (Dismorphia) praxinoe (Doubleday)

23. Dismorphia (Dismorphia ) fortunata ( Fucas )

24. Dismorphia (Dismorphia) eury ope (Lucas)

25. Dismorphia (Acmepteron) nemesis (Latreille)

26. Dismorphia (Enantia) alb ania (Bates)

27. Dismorphia (Enantia) jethys (Boisduval)

SUBFAMILY Pierinae

28. Catasticta nimbice nimhice (Boisduval)

29. Archonias (Archonias) tereas (Hiibner)

30. Appias (Glutophrissa) drusilla poeyi (Butler)

31. Leptophobia aripa elodia (Boisduval)

32. Itaballia (Itaballia) demophile calydonia (Boisduval)

33. Itaballia (Itaballia) pisonis kicaha (Reakirt)

34. Itaballia (Pieriballia) viardi viardi (Boisduval)

35. Ascia (Ascia) monuste monuste (Linnaeus)

36. Melete isandra ( Boisduval)

SUBFAMILY Coliadinae

37. Colias (Zerene) cesonia (Stoll)

38. Anteos clorinde ( Godart )

39. Anteos maerula ( Fabricius )

160

113

16(2):87-130, 1977 BUTTERFLIES OF TUXTLA

40. Phoebis (Phoebis) sennae marcellina (Cramer)

41. Phoebis (Phoebis) philea (Johansson)

42. Phoebis (Phoebis) argante (Fabricius)

43. Phoebis (Phoebis) agarithe maxima ( Neumoegen )

44. Phoebis (Phoebis) intermedia Butler

45. Phoebis (Rhabdodryas) trite (Linnaeus)

46. Phoebis ( Aphrissa ) statira jada ( Butler )

47. Eurema ( Eurema ) albula ( Cramer )

48. Eurema ( Eurema ) daira daira (Godart)

49. Eurema (Eurema) boisduvaliana Felder & Felder

50. Eurema (Eurema) xanthochlora (Kollar)

51. Eurema (Eurema) mexicana (Boisduval)

52. Eurema (Eurema) salome (Felder)

53. Eurema (Pyrisitia) proterpia (Fabricius)

54. Eurema (Pyrisitia) lisa Boisduval & Le Conte

55. Eurema (Pyrisitia) nise nelphe (Felder)

56. Eurema (Pyrisitia) dina westwoodi (Boisduval)

57. Eurema ( Abaeis ) nicippe ( Cramer )

58. Nathalis iole Boisduval

FAMILY ITHOMIIDAE
SUBFAMILY Ithomiinae

TRIBE Tithoreini

59. Tithorea harmonia salvadoris Staudinger

TRIBE Melinaeini

60. Melinaea lilis imitata Bates

TRIBE Mechanitini

61. Mechanitis polymnia lycidice Bates

62. Mechanitis agaensis doryssus Bates

63. Mechanitis menapis saturata Godman

TRIBE Napeogenini

64. Hypothyris ly caste dionaea Hewitson

65. Napeogenes tolosa ( Hewitson )

TRIBE Ithomiini

66. Ithomia leila Hewitson

67. Ithomia patilla Staudinger

TRIBE Oleriini

68. Hyposcada virginiana virginiana ( Hewitson )

69. Oleria zea ( Hewitson )

161

114

GARY N. ROSS

/. Res. Lepid.

70. Oleriapaula (Weymer)

71. Aeria pacifica Godman & Salvin

TRIBE Dircennini

72. Dircenna klugi ( Geyer )

73. Episcada artena ( Hewitson )

74. Pteronymia cottyto ( Guerin )

TRIBE Godyridini

75. Greta nero ( Hewitson )

76. Greta oto ( Hewitson )

77. Greta anetta ( Guerin )

78. Hypoleria cassotis ( Bates )

FAMILY DANAIDAE
SUBFAMILY Danainae

79. Danaus (Danaus) plexippus plexippus (Linnaeus)

80. Danaus (Tasitia) gilippus strigosus ( Bates )

81. Danaus (Tasitia) eresimus montezuma Talbot

SUBFAMILY Lycoreinae

82. Ly corea ceres atergatis ( Doubleday )

FAMILY SATYRIDAE
SUBFAMILY Satyrinae

83. Pierella luna heracles Boisduval

84. Taygetes mermeria excauata Butler

85. Taygetes virgilia (Cramer)

86. Taygetes andromeda ( Cramer)

87. Taygetes keneza Butler

88. Taygetes kerea Butler

89. Euptychia gemma freemani ( Stalings & Turner )

90. Euptychia hesione ( Sulz )

91. Euptychia met aleuca (Boisduval)

92. Euptychia mollina Hiibner

93. Euptychia lahe Butler

94. Euptychia similis Butler

95. Euptychia themis Butler

96. Euptychia undina Butler

162

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

115

97. Euptychia disaffecta Butler & Druce

98. Euptychia hermes sosybius ( Fabricius )

99. Euptychia gigas Butler

100. Euptychia libye ( Linnaeus )

101. Euptychia glaucina Bates

102. Euptychia sericella Bates

103. Euptychia nr. alcinoe Felder

104. Pedaliodes pisonia circumducta ( Thieme )

105. Dioriste tauropolis (Doubleday & Hewitson)

SUBFAMILY Brassolinae

106. Opsiphanes (Opsiphanes) boisduvalii Westwood &

Hewitson

107. Opsiphanes (Opsiphanes) cassiae castaneus Stichel

108. Eryphanis aesacus (Herrich-Schaeffer)

109. Caligo memnon ( Felder )

110. Caligo uranus Herrich-Schaeffer

FAMILY NYMPHALIDAE
SUBFAMILY Amathusiinae

TRIBE Morphini

111. Morpho theseus justiciae Salvin & Godman

112. Morpho polyphemus luna Butler

113. Morpho peleides montezuma Guenee

SUBFAMILY Acraeinae

TRIBE Acraeini

114. Actinote leucomelas ( Bates)

115. Actinote guatemalena veraecruzis Jordan

SUBFAMILY Heliconiinae

116. Philaethria dido dido (Clerck)

117. Dryadula phaetusa ( Linnaeus )

118. Agraulis vanillae incarnata (Riley)

119. Dione juno huascama (Reakirt)

120. Dione moneta poeyii (Butler)

121. Dryas julia moderata (Stichel)

122. Heliconius (Eueides) cleobaea zorcaon (Reakirt)

163

116

GARY N. ROSS

J. Res. Lepid.

123. Heliconius (Semelia) vibilia vialis (Stichel)

124. Heliconius (Semelia) lineata ( Salvin & Godman)

125. Heliconius (Semelia) aliphera gracilis (Stichel)

126. Heliconius (Heliconius) ismenius telchinia Doubleday

127. Heliconius (Heliconius) doris transiens Staudinger

128. Heliconius (Heliconius) sapho leuce Doubleday

129. Heliconius (Heliconius) sara veraepacis Bates

130. Heliconius (Heliconius) petiveranus Doubleday

131. Heliconius (Heliconius) charitonius vazquezae Comstock

& Brown

132. Heliconius (Heliconius) hortense Guerin

SUBFAMILY Nymphalinae
TRIBE Argynididi

133. Euptoieta hegesia hoffmanni Comstock

134. Chlosyne janais (Drury)

135. Chlosyne hippodrome (Geyer)

136. Chlosyne lacinia lacinia (Geyer)

137. Chlosyne erodyle ( Bates )

138. Chlosyne definita Aaron

139. Thessalia theona theona (Menetries)

140. Phyciodes (Phyciodes) vesta (Edwards)

141. Phyciodes (Eresia) frisia tulcis ( Bates )

142. Phyciodes (Eresia ) claudina guatemalena ( Bates )

143. Phyciodes (Eresia) phillyra (Hewitson)

144. Phyciodes (Tritanassa ) atronia ( Bates )

145. Phyciodes (Tritanassa) ardys ardys Hewitson

146. Phyciodes (Tritanassa ) eranites mejicana ( Roeber )

147. Phyciodes (Tritanassa) my ia (Hewitson)

148. Phyciodes (Tritanassa) griseobasolis Roeber

149. Phyciodes (Tritanassa ) clara ( Bates )

TRIBE Nymphalini

150. Polygonia g-argenteum ( Doubleday & Hewitson )

151. Vanessa virginiensis (Drury)

152. Junonia evarete evarete ( Cramer )

153. Anartia jatrophae luteipicta Fruhstorfer

154. Anartia fatima venusta Fruhstorfer

155. Metamorpha stelenes biplagiata (Frustorfer)

156. Metamorpha epaphus ( Latreille )

157. Hypanartia lethe (Fabricius )

158. Hypanartia dione Latreille

164

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

117

TRIBE Biblini

159. Biblis hyperia aganisa Boisduval
TRIBE Eunicidi

160. Mestra amymone ( Menetries )

161. Pyrrho gyra hypensor Godman & Salvin

162. Pyrrho gyra edocla aenaria Fruhstorfer

163. Pyrrho gyra otolais neis Felder

164. Pseudonica flavilla canthara ( Doubleday )

165. Temenis laothoe liberia (Fabricius)

166. Epiphile adrasta bandusia Fruhstorfer

167. Epiphile plutonia Bates

168. Catonephele nyctimus (Westwood)

169. Catonephele numilia esite (Felder)

170. Nessaea aglaura (Westwood & Hewitson)

171. Myscelia cyaniris Doubleday & Hewitson

172. Myscelia rogenhoferi Felder

173. Eunica monima ( Stoll )

174. Eunica alcmena alcmena Doubleday & Hewitson

175. Catagramma lyca Doubleday & Hewitson

176. Catagramma titania Salvin

177. Catagramma casta Salvin

178. Diaethria anna ( Guerin )

179. Diaethria astala (Guerin)

180. Dynamine mylitta ( Cramer )

181. Dynamine dyonis Geyer

TRIBE Ageroniidi

182. Hamadryas februa gudula (Fruhstorfer)

183. Hamadryas feronia farinulenta (Fruhstorfer)

184. Hamadryas guatemalena guatemalena ( Bates )

185. Hamadryas iphthime ( Bates )

186. Hamadryas amphinome mexicana ( Lucas )

187. Hamadryas laodamia laodamia (Cramer)

TRIBE Marpesiidi

188. Marpesia chiron ( Fabricius )

189. Marpesia harmonia (Klug)

190. Marpesia corita (Westwood)

191. Marpesia petreus (Cramer)

165

118

GARY N. ROSS

J. Res. Lepid.

TRIBE Liminitidi

192. Limenitis (Adelpha) melanthe (Bates)

193. Limenitis (Adelpha) leuceria (Druce)

194. Limenitis (Adelpha) erotia (Hewitson)

195. Limenitis (Adelpha) oberthuri (Boisduval)

196. Limenitis (Adelpha) iphicla (Linnaeus)

197. Limenitis ( Adelpha ) basiloides ( Bates )

198. Limenitis (Adelpha) felderi (Boisduval)

199. Limenitis ( Adelpha ) sentia ( Godman & Salvin )

200. Limenitis ( Adelpha) paraeca ( Bates )

TRIBE Apaturidi

201. Chlorippe cherubina (Felder)

202. Chlorippe paeon ( Latreille )

203. Chlorippe lame (Drury)

204. Historis odius (Fabricius)

205. Smyrna blomfildia datis Fruhstorfer

206. Gynaecia dirce ( Linnaeus )

TRIBE Charaxidi

207. Prepona demophon centralis Fruhstorfer

208. Prepona antimache gulina Fruhstorfer

209. Prepona amphimachus (Fabricius)

210. Prepona laertes pallantias Fruhstorfer

211. Prepona brooksiana Godman & Salvin

212. Anaea (Siderone) marthesia (Cramer)

213. Anaea ( Zaretis ) itys (Cramer)

214. Anaea (Zaretis) callidryas (Felder)

215. Anaea (Anaea) aidea Guerin-Meneville

216. Anaea (Consul) fabius (Cramer)

217. Anaea (Consul) electra (Westwood)

218. Anaea (Memphis) eurypyle confusa Hall

219. Anaea (Memphis) artacaena (Hewitson)

220. Anaea (Memphis) pithyusa (Felder)

221. Anaea (Memphis) proserpina (Salvin)

222. Anaea (Memphis) morvus boisduvali W. P. Comstock

166

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

119

SUBFAMILY Libytheinae

223. Libytheana carinenta mexicana Michener

FAMILY LYCAENIDAE
SUBFAMILY Lycaeninae

TRIBE Theclini
SUBTRIBE Strymoniti

224. Eumaeus minyas Hubner

225. Eumaeus debora Hubner

226. Theorema eumenia Hewitson

227. Chlorostrymon simaethis simaethis ( Drury )

228. Chlorostrymon telea (Hewitson)

229. Calycopis beon (Cramer)

230. Calycopis trebula (Hewitson)

231. Calycopis pisis (Godman & Salvin)

232. Calycopis sp. “C”

233. Tmolus echion echiolus (Draudt)

234. Tmolus crolinus (Butler & Druce)

235. Tmolus azia (Hewitson)

236. Oenomaus orty gnus (Cramer)

237. Callophrys ( Cyanophrys) amyntor distractus Clench

238. Callophrys (Cyanophrys) her odotus (Fabricius)

239. Callophrys (Cyanophrys) leucania (Hewitson)

240. Callophrys (Cyanophrys) miserabilis (Clench)

241. Callophrys (Cyanophrys) goodsoni Clench

242. Callophrys (Cyanophrys) agricolor agricolor (Butler &

Druce )

243. Callophrys (Cyanophrys) nr. longula (Hewitson)

244. Atlides polybe ( Linnaeus )

245. Panthiades ochus ( Godman & Salvin )

246. Strymon melinus Hubner

247. Strymon yojoa ( Reakirt )

248. Strymon columella istapa (Reakirt)

249. Strymon bazochii ( Godart )

250. Strymon albata sedecia ( Hewitson )

251. Strymon serapio (Godman & Salvin)

167

J. Res. Lepid.

120 GARY N. ROSS

252. Electrostrymon cyphara (Hewitson)

253. Cycnus battus jalan (Reakirt)

254. Arawacus aetolus togarna ( Hewitson )

255. Ara wacus sito (Boisduval)

256. Heterosmaitia palegon (Cramer)

257. Allosmaitia pion ( Godman & Salvin )

258. Evenus regalis ( Cramer )

259. Thecla cypria (Ge yer)

260. Thecla mar sy as damo (Druce)

261. Thecla augustala Kirby

262. Thecla lisus Stoll

263. Thecla manors ( Hubner)

264. Thecla inachus carpophora Hewitson

265. Thecla neora Hewitson

266. Thecla laothoe Godman & Salvin

267. Thecla barajo Reakirt

268. Thecla janias ( Cramer)

269. Thecla hassan ( Stoll)

270. Thecla met on (Cramer)

271. Thecla janthina janthodonia Dyar

272. Thecla nr. polibites (Cramer)

273. Thecla vibidia Hewitson

274. Thecla hecate Godman & Salvin

275. Thecla jebus (Godart)

276. Thecla brescia Hewitson

277. Thecla ligarina Hewitson

278. Thecla mycon Godman & Salvin

279. Thecla thales ( Fabricius )

280. Thecla tephraeus (Ge yer)

281. Thecla syncellus syncellus (Cramer)

282. Thecla minthe Godman & Salvin

283. Thecla empusa Hewitson

284. Thecla ares Godman & Salvin

285. Thecla ahola Hewitson

286. Thecla gabatha Hewitson

287. Thecla tarpa Godman & Salvin

288. Thecla maeonis Godman & Salvin

289. Thecla hesperitis ( Butler & Druce )

290. Thecla denarius (Butler & Druce)

291. Thecla plusios Godman & Salvin

292. Thecla clarina Hewitson

168

121

16(2):87-130, 1977 BUTTERFLIES OF TUXTLA

293. Thecla clemonassa Hewitson

294. Theda tera Hewitson

295. Theda coronata Hewitson

296. Theda scopas Godman & Salvin

297. Theda mathewi Hewitson

298. Theda politas Druce

299. Theda basalides (Geyer)

300. Theda mulucha Hewitson

301. Theda amhrax Westwood & Hewitson

302. Theda dodava Hewitson

303. Theda kalikimaka Clench

304. Theda tamos Godman & Salvin

305. Theda nr. antincus Felder

TRIBE Plebejini

306. Hemiargus (Hemiargus ) ceraunus zachaeina ( Butler &

Druce )

307. Hemiargus (Echinargus) huntingtoni hannoides Clench

308. Hemiargus (Echinargus) isola isola (Reakirt)

309. Everes comyntas comyntas ( Godart )

310. Leptotes cassius striata ( Edwards )

311. Celastrina argiolus gozora (Boisduval)

FAMILY RIODINIDAE
SUBFAMILY Euselasiinae

TRIBE Euselasiini

312. Euselasia sergia ( Godman & Salvin )

313. Euselasia hieronymi (Godman & Salvin)

314. Euselasia cheles aurantiaca ( Godman & Salvin )

315. Euselasia cataleuca (Felder)

316. Euselasia pusilla ( Felder )

317. Euselasia eubule (Felder)

169

122

GARY N. ROSS

J. Res. Lepid.

SUBFAMILY Riodininae

TRIBE Riodinini

318. Hades noctula Westwood

319. Perophthalma tullius lasius Stichel

320. Leucochimona philemon nivalis (Godman & Salvin)

321. Leucochimona vestalis vestalis (Bates)

322. Mesosemia tetrica Stichel

323. Mesosemia gaudiolus Bates

324. Euryhia lycisca Westwood

325. Cremna umbra ( Boisduval)

326. Ancylusis jurgensenii ( Saunders )

327. Rhetus arcius thia (Morisse)

328. Isapis agyrtus her a Godman & Salvin

329. Notheme eumeus diadema Stichel

330. Calephelis fulmen (Stichel)

331. Calephelis sp. 1

332. Calephelis sp. 2

333. Charis velutina (Godman & Salvin)

334. Charis myrtea (Godman & Salvin)

335. Charis psaros (Godman & Salvin)

336. Charmona gynaea zama ( Bates )

337. Baeotis hisbon zonata Felder

338. Lymnas pixe pixe Boisduval

339. Mesene margaretta (White)

340. Mesene croceela Bates

341. Symmachia rubina Bates

342. Symmachia accustrix Westwood

343. Symmachia tricolor hedemanni (Felder)

344. Phaenochitonia sagaris tyriotes ( Godman & Salvin )

345. Anteros carausius carausius Westwood

346. Calydna venusta Godman & Salvin

347. Emesis liodes Godman & Salvin

348. Emesis mandana mandana (Cramer)

349. Emesis tenedia Felder

350. Emesis lupina Godman & Salvin

351. Tharops menander isthmiae Godman & Salvin

352. Thisbe irenea belides Stichel

353. Polystichtis sudias ( Hewitson )

354. Anatole agave ( Godman & Salvin )

170

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

123

355. Anatole rossi Clench

356. Peplia lamis molpe (Hiibner)

357. Nymula calice mycone (Hewitson)

358. Calociasma lilina ( Butler )

TRIBE Theopini

359. Theope eleutho Godman & Salvin

Appendix B

PLANTS IDENTIFIED FROM THE SIERRA DE TUXTLA
Acanthaceae

Aphelandra aurantiaca ( Schiedw. ) Lindl.

Odontonema callistachyum (S. & C. ) Kuntze
Ruellia fluviatilis Leonard
Amaryllidaceae
Agave sp.

Anacardiaceae

Spondias mombin L.

Annonaceae

Annona muricata L.

Annona reticulata L.

Annona sp.

Apocynaceae

Stemmadenia galeottiana (A. Rich.) Miers.

Tabernaemontana citrifolia L.

Aquifoliaceae

Ilex belizensis Lundell
Ilex discolor Hemsl.

Ilex nitida (Vahl. ) Maxim.

Araliaceae

Dendropanax arboreus ( L. ) D. & P.

Oreopanax capitatum (Jacq. ) D. & P.

Oreopanax xalapense (H.B.K.) D. & P.

Aristolochiaceae

Aristolochia asclepiadifolia T. S. Brandeg.

Asclepiadaceae

Asclepias woodsoniana Standi. & Steyerm.

Asclepias tuberosa L.

Aspidieae

Didymochlaena truncatula (Swartz) J. Smith

171

124

GARY N. ROSS

J. Res. Lepid.

Bombacaceae

Bernoullia flammea Oliver
Ceiba pentandra (L. ) Gaertn.

Pachira aquatica Aubl.

Boraginaceae

Cordia alliodora (R. & P. ) Cham.

Cordia spinescens L.

Heliotr opium indicum L.

T ournefortia glabra L.

Burseraceae

Bursera simaruba (L.) Sarg. sens. lat.
Campanulaceae

Centropogon affine Mart. & Gal.

Caprifoliaceae

Viburnum acutifolium Benth.

Clethraceae

Clethra macrophylla M. & G.

Clethra suaveolens Turcz.

Combretaceae

Terminalia amazonia (Gmel.) Exell.

Compositae

Ageratum conyzoides L.

Baltimora recta L.

Bidens pilosa L. var. bimucronata (Turcz.) Schultz
Calea cacosmoides Less, (tentative)

Calea longipedicellata R. & G.

Calea zacatechichi Schlecht.

Conyza chilensis Spreng.

Eupatorium incomptum DC.

Eupatorium macrophyllum L.

Eupatorium pittieri Klatt

Eupatorium tuerckheimii Klatt, vel aff. (tentative)

Hidalgoa ternata Llave

Liabum dimidium Blake ( tentative )

Liabum sp.

Melampodium divaricatum (Rich.) DC.
Melampodium kunthianum DC.

Melanthera angustifolia A. Rich.

Polymnia maculata Cav.

Schistocarpha sp.

Senecio sp.

Stevia rhombifolia (H.B.K. )

172

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

125

Vernonia argyropappa Buek.

Vernonia leiocarpa DC.

Convolvulaceae

Ipomoea pes-caprae ( L. ) Sweet
Ipomoea stolonifera ( Cyr. ) Gmel.

Corylaceae

Carpinus caroliniana Walt.

Cucurbitaceae

Anguria tabascensis Donn. Sm., vel aff.

Cycadaceae

Ceratozamia mexicana Brong. sens. lat.

Zamia loddigesii var. angustifolia ( Regel ) Schuster
Cyatheaceae

Alsophila schiedeana Presl.

Cyathea sp.

Cyperaceae

Bulbostylis papillosa Kukenth. (tentative)

Cyperus articulatus L.

Cy perus ligularis L.

Dichromena ciliata Vah.

Rhynchospora globosa (H.B.K. ) R. & S.

Rhynchospora tuerckheimii Clarke
Dilleniaceae

Curatella americana L.

Saurauia sp.

Elaeocarpaceae
Sloanea sp.

Ericaceae

Gaultheria sp. (tentative)

Erythroxylaceae

Erythroxylon tabascense Britton
Euphorbiaceae

Acalypha diversifolia Jacq. var. carpinifolia (Poepp. & Endl.)
Muell.-Arg.

Acalypha unibracteata Muell.-Arg., vel aff.

Alchornea latifolia Sw.

Croton glabellus L.

Croton repens Schlecht.

Croton soliman Schlecht. & Cham.

Gymnanthes actinostemoides Muell.-Arg.

Jatropha curcas L., vel aff.

Rondelitia galeottii Standi., vel aff.

173

126

GARY N. ROSS

/. Res. Lepid.

Fabaceae

Dussia mexicana ( Standi. ) Harms
Erythrina americana Mill.

Gliricidea sepium (Jacq. ) Steud.

Fagaceae

Quercus conspersa Benth.

Quercus ghiesbreghtii Mart. & Gal., vel aff.
Quercus oleoides S. & C.

Quercus peduncularis Nee, vel aff.

Quercus skinneri Benth.

Flacourtiaceae

Casearia nitida (L. ) Jacq.

Casearia sylvestris Sw.

Fleur anthodendr on mexicana (A. Gray) L. Wms.
Xylosma sp.

Gleicheniaceae

Gleichenia palmata (Schaffner) Moore
Gramineae

Aulonemia sp.

Cenchrus incertus M. A. Curtis
Chloris petraea Swartz
Distichlis spicata (L.) Greene (tentative)
Eragrostis sp.

Isachne arundinacea ( Sw. ) Griseb.

Paspalum pectinatum Nees
Paspalum plicatulum Michx.

Sporobolus cubensis Hetchc.

Guttiferae

Clusia salvinii Donn. Sm.

Rheedia edulis (Seem.) Tr. & PI.

Vismia mexicana Schlecht.

Hamamelidaceae

Liquidambar styraciflua L.

Icacinaceae
Calatola sp.

Juglandaceae

Engelliardtia guatemalensis Standi.

Engelhardtia mexicana Standi.

Labiatae

Salvia shannonii Donn. Sm.

Lauraceae

Persea longipes ( Schlecht. ) Meissn.

174

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

127

Phoebe bourgeauviana Mez
Phoebe mexicana Meissn.

Phoebe psychotrioides (Nees) Mez (tentative)
Leguminosae

Albizia idiopoda (Blake) B. & R.

Albizia sp.

Calliandra grandiflora ( L’Her. ) Benth.

Cassia fruticosa Mill, (tentative)

Cassia hispidula Vahl
Cassia occidentalis L.

Cassia spectabilis DC., vel aff.

Crotalaria vitellina Ker
Dalbergia sp. (tentative)

Erythrina mexicana Krukoff
Inga leptoloba Schlecht.

Inga spuria Humb. & Bonpl.

Pithecollobium arbor eum (L.) Urb.

Vigna luteola (Jacq. ) Benth. (tentative)
Magnoliaceae

T alauma mexicana (DC.) G. Don
Malpighiaceae

Byrsonima crassifolia (L.) DC.

Malvaceae

Hibiscus tiliaceus L.

Melastomaceae

Conostegia xalapensis (Bonpl.) DC.

Miconia argentea (Sw. ) DC.

Miconia glaberrima ( Schlecht. ) Naud.
Melioceae

Guarea sp. (tentative)

Meteoriaceae

Pilotrichella flexilis (Hedw. ) Jaeg.
Monimiaceae

Siparuna andina (Tul.) A. DC.

Moraceae

Cecropia mexicana Hemsl.

Ficus cotinifolia H.B.K.

Ficus glaucescens (Liebm.) Miq.

Ficus obtusifolia H.B.K.

Ficus padifolia H.B.K.

Ficus spp.

Pseudolmedia oxyphyllaria Donn. Sm.

175

128

GARY N. ROSS

J. Res. Lepid.

Musaceae

Heliconia latispatha Benth.

Myricaceae

Myrica cerifera L.

Myrica splendens ( Sw. ) DC.

Myristicaceae

Virola guatemalensis (Hemsl.) Warb.
Myrsinaceae

Ardisia sp. (tentative)

Deherainia smaragdina ( Planch. ) Decaisne
Myrtaceae

Eugenia sp. ( tentative )

Orchidaceae

Elleanthus capitatus ( R. Br. ) Reichb.
Oxalidaceae

Oxalis neaei DC. sens lat.

Palmae

Astrocaryum mexicanum Liebm.

Chamaedorea elegans Mart, (tentative)
Chamaedorea ernesti-augustii Wendl. (tentative)
Chamaedorea tepejilote Liebm.

Chamaedorea sp.

Orbignya sp.

Sabal sp.

Passifloriaceae

Passiflora ambigua Hemsl.

Passiflora biflora Lam.

Passiflora coriacea Juss.

Passiflora serratifolia L.

Pinaceae

Pinus oocarpa Schiede
Piperaceae

Piper auritum H.B.K., vel aff.

Piper cordovan C. DC., vel aff.

Piper spp.

Podocarpaceae

Podocarpus oleifolius D. Don
Polypodiaceae
Dryopteris sp.

Pteridium aquilinum var. caudatum ( L. ) Sadele
Proteaceae

Roupala borealis Hemsl.

176

16(2):87-130, 1977

BUTTERFLIES OF TUXTLA

129

Pterobryaceae

Pterobryum densum ( Schwaegr. ) Hornsch.
Rhizophoraceae

Rhizophora mangle L.

Rosaceae

Hirtella racemosa Lam.

Rubiaceae

Borreria suave olens Mey.

Cephaelis elata Sw.

Crusea calcocephala DC.

Deppea excelsa (H.B.K. ) Standi., vel aff.

Hamelia longipes Standi.

H amelia patens Jacq.

Hoffmannia lenticillata Hemsl.

Lindenia rivalis Benth.

Machaonia sp., vel aff. (tentative)

Psychotria padifolia H. & B. (tentative)

Psychotria sp.

Rondeletia strigosa ( Benth. ) Hemsl.

Rudgea cornifolia (H. & B.) Standi., vel aff.
Rutaceae

Zanthoxylum elephantiasis Macf.

Sapindaceae

Thouinidium decandrum (H. & B.) Radik.
Saxifragaceae

Weinmannia pinnata L.

Schrophulariaceae

Escobedia laevis C. & S.

Lamourouxia viscosa H.B.K.

Stemodia durantifolia Sw., vel aff. (tentative)
Simaroubaceae

Picramnia andicola Tub, vel aff. (tentative)
Solanaceae

Solanum ochraceo-ferrugineum (Dunal) Fernald
Solanum schlechtendalianum Walp.

Solanum spp.

Theaceae

Saurauia sp. (tentative)

Tiliaceae

Apeiba tibourbous Aubl.

Belotia sp.

Luehea speciosa Willd.

177

130

GARY N. ROSS

/. Res. Lepid.

Turneraceae

Turner a ulmifolia L.

Ulmaceae

Mirundaceltis monoica (Hemsl. ) Sharp
Urticaceae

Boehmeria sp. (tentative)

Myriocarpa bifurca Liebm. (tentative)
Myriocarpa longipes Liebm.

Trema micrantha (L.) Blume
Urera elata (Sw. ) Griseb.

Verbenaceae

Aegiphila costaricensis Moldenke
Lantana camara L.

Violaceae

Rinorea guatemalensis (Wats.) Bartlett

178

PARATRYTONE MELANE IN SAN LUIS OBISPO
COUNTY, CALIFORNIA (HESPERIIDAE)

SCOTT E. MILLER

Santa Barbara Museum of Natural History
2559 Puesta Del Sol Road
Santa Barbara , California 93105

The distribution of Paratrytone melane (Edwards, 1869) in
California has been treated recently by Heppner (1973). He
noted a large distribution gap in the Coast Ranges from the
Santa Ynez River in Santa Barbara County north to Big Sur in
Monterey County. Heppner suggested that this gap might indi¬
cate an allopatric distribution, with “a northern population
concentrated in the San Francisco Bay area and a southern popu¬
lation in most of coastal southern California.” However, the
present author has located several specimens from San Luis
Obispo County which fill in this prominent distributional gap.

A specimen from Lopez Canyon, collected “2-9- [19] 64” by
Richard Hart, is present in the Santa Barbara Museum of Na¬
tural History collection. Two specimens from Atascadero, col¬
lected “5-10-1932” by Victor L. Clemence, have been located in
the Clemence collection. One of these is now in the author’s
collection, while the other remains in the Clemence collection.

These previously unpublished records show that P. melane is
indeed distributed throughout the Coast Ranges of California.
The records also demonstrate the need for additional study of
the lepidopterous fauna of this region. The author is indebted to
Mrs. Grace Clemence and Mr. Nelson W. Baker for their assist¬
ance.

LITERATURE CITED

HEPPNER, JOHN B. 1973. The distribution of Paratrytone melane and
its spread into San Diego County ( Hesperiidae ) . J. Res. Lepid. 10(4):
287-300.

131

NOTICES

NEW ADDRESS: The address of the Journal is now:
c/o Santa Barbara Museum of Natural History, 2559
Puesta Del Sol Road, Santa Barbara, CA 93105.

PUBLICATIONS: Available from Entomological Reprint
Specialists, P. O. Box 7797 1, Dockweiler Station, Los
Angeles, CA 90007;

Moths of Southern Africa. E. C. G. Pinhey, Tafel-
berg Publishers, Ltd. , 28 Wale Street, Cape Town,

South Africa. $35. 95.

Butterflies of West Malaysia and Singapore, Vol. 1
and 2. W. A. Fleming, E. W. Classey Ltd. , Park Road,
Faringdon, Berkshire, England. $47. 00 set.

Revised Catalouge of the African Sphingidae (Lepi-
doptera) with Descriptions of the East African Species.

R. H. Carcasson, E. W. Classey Ltd. $1 1. 95.

FOR SALE: 1, 500 exotic butterflies, insects, biologicals.

72 page catalog $1. 00. Complete Scientific, P. O. Box
307, Round Lake, IL 60073.

XERCES SOCIETY: An international organization dedi¬
cated to the concervation of terrestrial arthropods and
their habitats. Publications include the journal Atala
and the newsletter Wings. Memberships and subscrip¬
tions US $5. 00 (students $2. 50); contact treasurer, Roger
Pasquier, 235 East 73rd Street, New York, NY 10021.

BUY/SELL/EXCHANGE: Lepidoptera. Send $1.00 for
list of species (worldwide). Specialist in Sphingids, Sa-
turniids and Exotics. Ray Adams, F. R. E. S. , Carretera
Principal 139, Tamaraceite, Las Palmas De Gran Canaria.

WANTED: World moths particularly Geometridae. Will
trade North American or Panamanian moths. Roger L.
Heitzman, Dept, of Ent. , Univ. of Mo. , Agric. Bldg. ,
Columbia, MO 65201.

FOR SALE: 6m by 2. 5m linear Malaise Traps. Proven
design used world wide by military, universities and private
collectors. Additional information available from Dana A.
Focks, P.O.Box 12852, Gainsville, FL 32604, Dept. LRF.

EXCHANGE: Lepidoptera specimens. B. Cech, Vancurova
13, 150 00 Prague, Czechoslovakia.

Volume 16 Number 2 June,

IN THIS ISSUE

Foodplant ecology of the butterfly Chlosyne lacinia
(Geyer) Nymphalidae II Additional larval food
plant data. Raymond W. Neck

Descriptions of a new species of Eupithecia and the male
of E. cocoata Pearsall ( Geometridae )

Roger L. Heitzman and
Wilbur R. Enns

Autumnal false broods of multivoltine butterflies
at Donner Pass, California

Arthur M. Shapiro

An ecological study of the butterflies of the Sierra de
Tuxla in Vera Cruz, Mexico (concluded)

Gary N. Ross

Paratrytone melane in San Luis Obispo County, California
( Hesperiidae ) Scott E. Miller

COL/J9S
/?£*>/& e*to

AS

if+stc r* f/T'crrts

//<*t.&rysS

&*/?: * Z*

*S<*4<jr 4f<\y
*.£V-r.

SO - J/W/ 73
<5x. <:•✓/. G+Atc-s^

HARVARD

coc/^s

/raSaes? o .

£^iir^/njp
<s r^<f £

6+ffr«Z*.

S/,0»*T. C**"*Pst
S2"S"‘7^

£ X . C«// if^vcr

Volume 16 Number 3 September , 1977

« njE, juukinml ur KE9EARCH

THE L E P I DO PTE R A

published by

The Lepidoptera Research Foundation, Inc.
at

2559 Puesta Del Sol Road, Santa Barbara, California 93105
EDITOR: William Hovanitz
ASSISTANT EDITOR: Karen Nielson 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.

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.

CLASSES OF
MEMBERSHIP

Regular

$12

year

Family

15

year

Contributing

25

year

Subscribing

50

year

Sponsor

100

year

Life

250

for life

Subscriptions to the Journal are $15.00 per year. Special subscriptions are avail¬
able to students at $10.00 per year.

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 2559 Puesta Del Sol Road, Santa
Barbara, California 93105. The publisher is THE LEPIDOPTERA RESEARCH FOUN¬
DATION, INC. The general editor is William Hovanitz at the above address. The
secretary-treasurer is Barbara Jean Hovanitz at the same address. The owner is
THE LEPIDOPTERA RESEARCH FOUNDATION, INC. THE LEPIDOPTERA
RESEARCH FOUNDATION, INC. is a non-profit organization incorporated under the
laws of the State of California in 1965. There are no bond holders, mortgages or other
security holders.

Established in 1962

Edited by WILLIAM HOVANITZ

Volume 15

1976

published by

The Lepidoptera Research Foundation, Inc.
at

2559 Puesta Del Sol Road, Santa Barbara, California 93105

Journal of Research on the Lepidoptera

16(3) :133-140, 1977

c/o Santa Barbara Museum, of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

CHARIDRYAS FLAVULA
BARNES AND McDUNNOUGH
( NYMPHALIDAE ) :

A QUESTION OF IDENTITY1

CLIFFORD D. FERRIS2

Bioengineering Program, University of Wyoming,
Laramie, Wyoming 82071

and

MICHAEL S. FISHER

6521 South Logan Street, Littleton, Colorado 80121

INTRODUCTION

The palla boisduval complex of species in the Melitaeinae
has been referred to a number of genera including: Lemonias
Htibner, 1806 [not Felder and Felder, 1859] by Dyar, 1902;
Melitaea Fabricius, 1807 by Barnes and McDunnough, 1918;
Chlosyne Butler, 1870 by Higgins, 1960; Microtia Bates, 1864 by
dos Passos, 1964; Charidryas Scudder, 1872, 1875 by Bauer in
Howe, 1975. The Bauer treatment seems reasonable and recog¬
nizes nine species in Charidryas [Type Species Melitaea nycteis
Doubleday] composed of a six-species western (palla) group
which lacks the dorsal postmedian series of black spots on the
hindwings found in the three-species eastern ( harrisi Scudder)
group. In addition to the maculation, there are differences be¬
tween the two groups in pupal morphology. Because of pheno¬
typic differences in the imagines between the northern Chari¬
dryas and southern Chlosyne, we choose to treat the former as
a full genus, rather than a subgenus as did Bauer.

The taxonomic status of C. flavula Barnes and McDunnough,
1918 has been clouded since its original description. McDun-

ublished with the approval of the Director, Wyoming Agricultural Experi¬
ment Station as Journal Article no. JA 880.

2 Research Associate, Allyn Museum of Entomology, Sarasota, Florida 33580 ;
Museum Associate, Los Angeles County Museum of Natural History, Los
Angeles, California 90007.

133

134

C. D. FERRIS and M. S. FISHER

J. Res. Lepid.

nough (1938), Higgins (1960), and dos Passos (1964, 1969)
offered divergent opinions concerning its position relative to
C. palla (Boisduval) and C. acastus (W. H. Edwards). Appar¬
ently some further study is indicated concerning the affinity of
flavula to these two species. The purpose of this paper is to
review the past treatment of the taxon and to assign it to what
we feel is its proper position for permanent reference in the
nomenclature.

TAXONOMIC CHARACTERS

Barnes and McDunnough (1918) indicated, before formally
describing flavula as a full species, that it might prove to be a
form of palla. McDunnough (1938) retained flavula as a full
species. Higgins (1960) extensively revised the palla group and
placed flavula as a subspecies of gabhii Behr. dos Passos (1964)
listed it under its original status, but later (1969) placed it as
a form of acastus.

The Barnes and McDunnough description does clearly com¬
pare the differences between flavula and acastus , but their study
material was limited and generally incompletely labeled. Based
upon the stated localities from which they had specimens, they
apparently worked with Colorado material from both east and
west of the Continental Divide. The type series contained 13
males, 11 of which were labeled simply “Colorado”, and were
probably collected by David Bruce. Two specimens were labeled
respectively “Glenwood Spgs., [Garfield Co.] Colo.” and “Hall
Valley, [Park Co.] Colo.” Brown (1957) observed on page 79:
“Since no other specimens of flavula have been found on the
eastern slope, I assume that the Hall Valley label was inadvert¬
ently attached to a Glenwood Springs specimen. On this basis
I believe it best to designate Glenwood Springs as the type
locality of flavula”

In the original description, flavula was compared with Cali¬
fornia and Colorado palla , and acastus from Glenwood Springs.
Reference was also made to Utah and Wyoming specimens, but
no conclusions were drawn. An examination of an extensive
series of the flavula phenotype from western Colorado, where
flavula and acastus are sympatric, indicates that there are dis¬
tinct characters which place flavula in the C. palla complex.

There are several characters that reliably separate flavula
from acastus. As noted by Barnes and McDunnough, flavula is
smaller than acastus , but it is also much darker. The brownish-

16(3): 133-140, 1977

CHARIDRYAS FLAVU LA

135

black areas of the wings are more extensive, producing, in most
specimens, a reduction in the checkered maculation. This macu-
lation is distinctly more contrastingly bi-colored, reddish-to-
yellowish dorsally than in acastus. Ventrally, the checkered
maculation on the hindwings is buff-white, separated by fine
black lines along the veins. By comparison, acastus is clearly
flat white to pearly- white, with broader black lines. The basal
area of both wings is more solid reddish in flavula, while in
acastus it is mottled extensively with white and black on the
hindwings and orange-buff on the forewings. On the whole,
C. palla flavula differs from its congener, C. palla calydon (Hol¬
land) by its smaller size, brighter colors above, and larger spot-
bands on the hindwings beneath. The differences are illustrated
in Figures 1-14.

DISTRIBUTION

C. palla flavula occupies a fairly narrow geographic zone in
the plateau country of western Colorado and eastern Utah with
an extension into southern Wyoming. It is locally common in
dry habitats at an altitude range from 6000-8000’ (1830-2440m)
where Sagebrush ( Artemisia tridentata Nutt.) is the dominant
plant species. It does occur, however, in similar habitat up to
9500’ (2895m). The larval foodplant is probably Indian Paint¬
brush ( Castilleja species) as it is for calydon. C. acastus occurs
in semi-arid lowlands in this same region and meets flavula in
and around Glenwood Springs. C. p. flavula is the more montane
of the two insects and replaces acastus completely in similar
habitats at higher elevation.

The flavula phenotype should also occur in northwestern
New Mexico since it occurs in adjacent areas in Colorado. Barnes
and McDunnough stated: “We have it from Utah (Provo, Salt
Lake) and also from the Yellowstone Park region.” The Yellow¬
stone citation undoubtedly refers to calydon based upon speci¬
mens that we have examined from western Wyoming. Holland
did not describe calydon until 1931. We have also examined a
short series of palla collected north of Salt Lake City in Cache
Co.; these do not match the flavula characters defined above.
The same is true of material from Emigration Canyon and
Lamb’s Canyon in Salt Lake Co., Utah. This material is more
unicolorous above, and in terms of maculation, similar to acastus.
Figure 15 indicates the presently known distribution for flavula.

Brown ( 1957 ) discussed the type material of flavula as noted
above. Contrary to Bauer in Howe ( 1975 ) , the specimen figured

136

C. D. FERRIS and M. S. FISHER

/. Res. Lepid.

Fig. 1. — C. p. flavula, male upper, nr. Snowmass, Pitkin Co., Colorado,
13-vi-74.

Fig. 2. — C. p. flavula, female upper, Rabbit Ears Pass area, Routt Co.,
Colorado, 12-vii-74.

Fig. 3. — C. p. calydon, male upper, nr. Nederland, Boulder Co., Colorado,
22- vi-74.

Fig. 4. — C. p. calydon, female upper, same data as male in Fig. 3.

Fig. 5. — C. p. calydon, male upper, South Turkey Creek Canyon, Jefferson
Co., Colorado, 2-vii-68 (Topotype).

Fig. 6. — C. acastus, male upper, Coalmine Point, Black Ridge Breaks, Mesa
Co., Colorado, 26-V-71.

Fig. 7. — C. acastus, female upper, same locality as male, 29-V-70.

16(3) -.133-140, 1977

CHARIDRYAS FLAVULA

Figures 8-14 respectively are the undersides of the specimens shown
Figs. 1-7.

UTAH

138

C. D. FERRIS and M. S. FISHER

J. Res. Lepid.

NEW MEXICO

Fig. 15. — Known distribution of C. palla flavula. Locality data appear in
the Appendix.

16(3):133-140, 1977

CHARIDRYAS FLAVULA

139

as flavula by Brown is good flavula , although his discussion of
the differences between it and calydon is somewhat confused.
Brown did not figure acastus and it was not included in the
Colorado species list, although Barnes and McDunnough (1918,
PL 11, f. 13) illustrated a specimen from Glenwood Springs. It
is possible that some of the material that Brown used for com¬
parison represented acastus from flavula localities, thus further¬
ing the confusion. The male flavula figured in Howe (1975, PI.
72, f. 11) is a poor representation of this subspecies, but does
represent the Utah population previously mentioned.

SUMMARY

The Barnes and McDunnough name flavula is presented
here as a valid subspecies assigned to the Charidryas palla
complex. The distinctiveness of flavula is supported by specific
differences in color, maculation, distribution, and especially by
its sympatric occurrence with acastus at and near the Glenwood
Springs type locality. At Basalt, Eagle Co., Colorado, acastus
and flavula occur sympatrically with no indication of inter¬
gradation between the two species.

LITERATURE CITED

BARNES, W. and J. H. McDUNNOUGH. 1918. Contrib. Nat. Hist. Lepid.
North America. 4(2): 1-208.

BROWN, F. M. 1957. Colorado Butterflies. Denver Museum of Natural
History, Denver, viii + 368 pp.

DYAR, H. A. 1902. A list of North American Lepidoptera and key to the
literature of this order of insects. Bull. U.S.N.M. 52: 1-723 + xix.
HIGGINS, L. C. 1960. A revision of the Melitaeine genus Chlosyne and
allied species (Lepidoptera: Nymphalidae). Trans. Royal Ent. Soc.
London. 112: 381-467.

HOWE, W. H., Contrib. Ed. 1975. The Butterflies of North America.
Doubleday and Co., Inc., Garden City, L.I., New York, xiii + 633 p.
+ 97 plates.

McDUNNOUGH, J. 1938. Check list of the Lepidoptera of Canada and
the United States. Memoirs of the Southern California Academy of
Sciences. Vol. 1.

DOS PASSOS, C. F. 1964. A Synonymic List of the Nearctic Rhopalocera.

The Lepidopterists> Society. Memoir No. 1.

- . 1969. A synonymic list with taxonomic notes concerning the Ne¬
arctic Melitaeinae (Nymphalidae). J. Lepid. Soc. 23(2): 115-125.
SCUDDER, S. H. 1872. 4th Ann. Rep. Peabody Acad. Sci. 1871: 47.

- . 1875. Synonymic list of the butterflies of North America, North of

Mexico. Bull. Buffalo Soc. Nat. Sci. 2: 233-269.

140

C. D. FERRIS and M. S. FISHER

J. Res. Lepid.

APPENDIX

Distribution data for the points indicated on the map (Figure 15) are
given below. We would like to thank Scott Ellis, Ft. Collins, Colorado and
R. E. Stanford, Denver, Colorado for supplying data. The localities listed
on page 79 of Brown (1957) are also included.

COLORADO: Archuleta Co.: nr. Allison, 27-30-v; Arboles, 6100’, v-37.
Delta Co.: Leroux Ck. Rd., 8000’ 8-vi-62; Smith Fk. Ck., 2 mi. E. of Needle
Rock, 29-vi-61. Dolores Co.: Jet. W. Dolores Riv. Rd. and cutoff to Nor-
wood-Dolores Rd., San Juan Nat. For., 7800’, 25-V-72. Eagle Co.: 2 mi. E.
of Basalt, 13-vi-70, 10-vi-76. Garfield Co.: Glenwood Springs (Type Lo¬
cality); No Name Ck., Glenwood Canyon, 6400’, l-vii-67; Grizzly Ck.,
Glenwood Canyon, 6400’, 12-vii-68; French Ck., Glenwood Canyon, 11-vi-
76. Gunnison Co.: Anthracite Ck., vie. Kebler Pass Rd., 14-vi-74. La Plata
Co.: Durango, 12-vi-72; La Posta, 23-V-34; Tiffany, 6200’, 11-14-v. Mesa
Co.: Love Mesa, Uncompahgre Plateau, SW of Escalante Canyon, 8000’,
6-vii-62; Grand Mesa, Lands End Rd., 1 mi. S. of Lookout House, 9600’,
2-vi-72. Montezuma Co.: Taylor Ck., N. of Dolores River, vie. Hwy. 145,
8000’, 15-vii-67; Mesa Verde Nat. Park: Head of Navaho Canyon, 7800’,
26-V-72; Headquarters Canyon, 12-vi-51; Mesa Verde, 27-V-37; Ute Mtn.
Pitkin Co.: vie. Aspen, 7900’, 13-27-vi; Maroon Ck., 29-vii-50; Maroon
Lake Rd., 27-vi-76 (leg. W. B. Wright); Snowmass, 13-vi-73, 13-vi-74.
Routt Co.: Rabbit Ears Pass, 4-vii-48, 24-vii-72, 22-vii-73, 12-vii-74. San
Miguel Co.: Bear Ck., Wilson Peak Rd. nr. Sawpit, 8500’, 18-vii-65; Hwy.
141, 4.5 mi. S. of Hwy. 145 nr. Gurley Reservoir, 8400’, 26-vi-67. WYO¬
MING: Carbon Co.: Battle Ck. Campgd., Medicine Bow Nat. For., 13 mi.
ENE of Slater, 21-vi-73.

Scott Ellis has reported that flavula and acastus are sympatric at Mesa
Verde in Montezuma Co., Colorado. He has collected one specimen that
appears to be a hybrid between the two species.

Journal of Research on the Lepidoptera

16(3) :141-145, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

OCELLUS VARIATION AND WINGSPAN
IN ATTACUS ATLAS LINNAEUS.,

IS THERE A RELATIONSHIP?

( SATURNIIDAE )

MARK HADLEY

7 Beverington Close,

Eastbourne, E. Sussex, BN21.2SB,

England

Considerable variation exists between the size and position
of the elliptical hyaline area of the forewing. It is this area,
when surrounded with darkly pigmented scales that constitutes
the postdiscal ocellus. The discal ocellus is triangular in shape
and situated nearer to the basal area of the forewing. Some
specimens of this species show the complete absence of this
first, elliptical hyaline area (fig. 4), whereas in other specimens
enlargement of this area results in the fusion of the postdiscal
ocellus with the discal ocellus (fig. 3). By chance, the author
when comparing these forms noticed that the form exhibiting
‘fusion’ also had a considerably larger wingspan than the average
for its sex. It was this observation that prompted the author to
investigate further, and try to provide an answer to the question
“. . . is there a relationship between wingspan and postdiscal
ocellus variation? . . .”

A sample of four hundred and ninety set specimens of A.
atlas Linn., were used for the study, all being bred in India.
For each separate individual the length of the elliptical hyaline
area was measured, as well as the distance between the apices
of the forewings. The gross sample was segregated into two di¬
visions, that of male and female specimens. Once this was com¬
pleted scattergrams were prepared illustrating ‘Wingspan vs
Ocellus length’ in millimetres. One for the sample of 146 male
specimens, and a second for the sample of 344 female specimens.

141

142

MARK HADLEY

/. Res. Lepid.

Fig. 1. — Forewing maculation of A. atlas. Typical male.

Fig. 3. — Forewing maculation of A. atlas. Female showing fusion of hyaline
areas, discal and postdiscal.

16(3):141-145, 1977

ATTACUS ATLAS

143

Fig. 2. — Forewing maculation of A. atlas. Typical female.

144

MARK HADLEY

J. Res. Lepid.

RESULTS, (table 1.)

TABLE I

MALES

N=146

Greatest recorded wingspan
= 221 mm

Smallest recorded wingspan
= 175 mm

Mean value for wingspan
= 196 mm

Greatest length of elliptical
hyaline area of forewing
= 14 mm

Smallest length of elliptical
hyaline area of forewing
= 1 mm

Mean value for elliptical hyaline
area of forewing

= 7.56 mm

Correlation was calculated be¬
tween the length of the elliptical
hyaline area and wingspan, the
value obtained was found to be
-0.707

FEMALES

N=344

Greatest recorded wingspan
= 247 mm

Smallest recorded wingspan
= 166 mm

Mean value for wingspan
= 202 mm

Greatest length of elliptical
hyaline area of forewing
= 18 mm

Smallest length of elliptical
hyaline area of forewing
= 0 mm

Mean value for elliptical hyaline
area of forewing

= 10.23 mm

Correlation was calculated be¬
tween the length of the elliptical
hyaline area and wingspan, the
value obtained was found to be
+ 0.252

DISCUSSION

The correlation value between the wingspan of a female and
the length of the elliptical hyaline area was found to be 0.252
which shows that there is a weak, but nevertheless significant
relationship between wingspan and ocellus. Furthermore, as the
figure obtained is positive the relationship is a direct one. Thus,
a specimen with a small wingspan would show a greater tend¬
ency towards the small ocellus size. This relationship is indeed
verified. From the three specimens in the sample exhibiting the
absence of the ocellus the wingspans were 166, 182 and 184
millimetres, these figures are well below the mean for the
sample which was calculated at 202mm. Alternatively, we can
look at the incidence of fusion within the sample. Two examples
of the fused condition were found with wingspans of 221 and
211 mm, both were above the mean for wingspan, this suggests
strongly that the relationship is direct.

We can thus predict that the specimens with unusually small
wingspans will show a reduction in ocellus size with complete
deletion of the area in extreme examples. In specimens showing

16(3):141-145, 1977

ATT AC US ATLAS

145

unusually large wingspans we can predict that the two hyaline
areas of the forewing will increase in size and ultimately fuse.

In the sample of males, the correlation figure was found to
be -0.707, this forcibly shows that there is a strong’ relationship
and good correlation. From the knowledge that the figure is
negative we can arrive at the conclusion, that unlike the case
of the females the relationship within the male population is
indirect. Thus specimens with large wingspans would have
small reduced ocelli on the forewings, while specimens with
smaller than average wingspans would have larger than average
ocelli. This point cannot be illustrated by reference in the sample
to fused or absent areas as none were found. The females there¬
fore in this sample exhibited the extreme variations, within the
male population these extremities were not shown.

ACKNOWLEDGEMENTS

The author would like to express his thanks to Mr. Minet of
Hastings College of Further Education for providing the sta¬
tistical approach to the data and generally helping with the
mathematical aspect. My thanks are also due to Mr. Perrie who
furnished the specimens for the study.

LITERATURE CITED

CONNOLLY, T. G. and SLUCKIN, W., 1969. An introduction to statistics
for the Social Sciences. Macmillan.

CROTCH, W. J. B., 1956. a silkmoth rearers handbook. The Amateur
Entomologist. 12: 1-165.

Journal of Research on the Lepidoptera

16(3) :147-154, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

FOODPLANT ECOLOGY OF THE BUTTERFLY
CHLOSYNE LAC INI A (GEYER)

( NYMPHALIDAE )

III. ADULT RESOURCES
RAYMOND W. NECK

Texas Parks and Wildlife Department
4200 Smith School Road
Austin, Texas 78744

Much data exists for many rhopalocera species concern¬
ing larval foodplants. Such information is of obvious importance
in the study of the ecology and evolution of any particular
species. For many years, however, another important aspect of
rhopaloceran biology has been generally ignored — the import¬
ance of adult energy sources. Some moth species have dispensed
with adult feeding, thus depending entirely upon the larval stage
for acquisition of energy stores. This evolutionary event has not
been observed in rhopalocerans, however; much important data
remain to be collected on various butterfly species. Norris (1936)
produced an early review of the topic but little more was pub¬
lished on this topic until the 1970’s.

Recently, this aspect of rhopaloceran biology has received
more attention by various workers. Shields (1972) reviewed
literature references and added personal records but little syn¬
thesis was attempted. An extreme example of butterfly-plant
mutualism involving the coevolution of Heliconius butterflies and
Anguria/Guranea pollen and nectar plants has been studied by
Gilbert (1975). Gilbert (1972) earlier reported the assimilation
of pollen proteins by Heliconius. Pollen-fed Heliconius ethilla
lay more than five times as many eggs as those without pollen.
Obviously, adult resources are a possible topic of much discus¬
sion and importance. Not all adult resources of lepidoptera are
supplied by plant products; Downes (1973) discussed mud
puddles, dung and carion in this respect.

During the process of collecting data concerning the larval
foodplant ecology of Chlosyne lacinia var. adjutrix Scudder

147

148

RAYMOND W. NECK

J. Res. Lepid.

(Nymphalidae: Melitaeinae), observations were made concern¬
ing adult acquisition of energy and/or water resources. These
data are not as comprehensive as the larval foodplant data, but
what is available is presented here. All observations occurred in
central Texas centering in Austin, Travis County.

ADULT RESOURCES

Verbesina encelioides (Cav. ) Gray. This plant is the major
source of nectar for adjutrix during the fall months when V.
encelioides serves as a primary nectar source for all butterflies.
Quite possibly, adjutrix has some advantage over other butterfly
species in utilization of this plant as a nectar source because
females do not have to leave the area to oviposit on the proper
foodplant. V. encelioides is the second most commonly utilized
foodplant of adjutrix; during the fall months V. encelioides is
by far the major foodplant of this butterfly.

Helianthus annuus L. Nectar of this plant is utilized at times,
but generally it is not very acceptable to adult adjutrix. H. an¬
nuus can be best classified as a “bee plant” in reference to flower
visitation; honey bees and bumble bees are the most frequent
visitors to this plant. This is the primary larval foodplant of
adjutrix , particularly during the spring and early summer
months. The general unsuitability of this plant as a nectar source
for adjutrix does not affect its utilization as a larval foodplant
because various annuals associated with stands of sunflowers are
prime nectar sources.

Heterotheca latifolia Ruckl. This species is a prime nectar
source for adjutrix especially in association with H. annuus over
which H. latifolia is utilized. H. latifolia tends to be ignored in
the presence of V. encelioides.

Verbesina virginica L. This plant does not bloom until the
fall; at this time it may serve as a temporary prime nectar source.
This plant is probably utilized to a lesser extent by adjutrix than
other butterflies because V. virginica occurs in wooded areas of
central Texas; adjutrix tends to avoid such shaded areas.

Zexmenia hispida (H.B.K.) Gray. The small inflorescences of
this species are utilized by adjutrix during the lengthy blooming
period (late spring to> fall). This plant is classified as an “occa¬
sional” larval foodplant (Neck, 1973).

Viguiera dentata (Cav.) Spreng. This plant, also an “occa¬
sional” larval foodplant, is utilized by adult adjutrix during
blooming season which is restricted to the fall months.

16(3):147-154, 1977

CHLOSYNE LAC1NIA

149

Gaillardia pulchella Foug. The colorful inflorescences of this
species may supply nectar to adjutrix adults during spring and
early summer. H. latifolia appears to be preferable to G. pul¬
chella which has “rarely” been utilized as an adjutrix larval
foodplant (Neck, 1973).

Eysenhardtia texana Scheele. This plant is an important peri¬
odic source of nectar. Flowers generally appear in late spring
and fall but react mainly to rainfall.

Lantana horrida H.B.K. and Lantana macropoda Torr. Adult
adjutrix have been observed at the flowers of these species but
the plants generally appear to be ignored. These two species are
major nectar sources for swallowtail and heliconian butterflies.
Possible factors involved in the low utilization of Lantana by
adjutrix will be discussed later in this article.

Rubus trivialis Michx. The blossoms of dewberry are utilized
as nectar sources in early spring, particularly if alternate sources
are uncommon. Such is often the case during a dry early spring
which follows a dry winter; few annuals germinate and produce
blossoms early in the season. When other suitable flowers begin
to appear, R. trivialis is ignored by adjutrix.

Several plants common in areas frequented by adjutrix are
apparently ignored by this species although not necessarily by
other butterflies. Verbena bipinnatifida Nutt, is frequently visited
by Battus philenor ( L. ) , for example. Lupinus texensis Hook, is
primarily visited by bees. Phlox drummondii Hook, is apparently
ignored by adjutrix although the major pollinators of plants of
this genus are butterflies of the genus Colias (see e.g. Levin,
1969).

Non-floral adult resources are also known for adjutrix. Adults
in predominantly male groups may be seen congregated at water
holes ( as previously reported by Bauer, 1953, in Arizona ) , espe¬
cially under high adult density conditions when the nectar
supply may be insufficient. After reviewing the literature of
lepidoptera visits to dung, carrion and puddles, Downes (1973)
concluded that these visits functioned to obtain some unidenti¬
fied nutrient. Studies by Arms et al. (1974) revealed that male
Papilio glaucus were attracted to sand soaked with sodium salts.
These results were interpreted as adult supplement of a larval
diet deficient in sodium. Male adjutrix have been observed to
feed on carrion and feces when sufficiently moist. Several other
melitaeine butterflies have been reported as feeding at mud
puddles and carrion (Voss, 1954, Payne and King, 1969). Dur-

150

RAYMOND W. NECK

/. Res. Lepid.

ing a summer picnic a single male adjutrix was observed to feed
on a piece of fresh watermelon rind for approximately thirty
minutes.

DISCUSSION

Nectar sources heavily utilized by adjutrix are, for the most
part, general ‘‘butterfly flowers” as far as flower visitors are con¬
cerned. Faegri and van der Pijl (1966) remark that butterflies,
generally speaking, “seem to prefer sucking nectar out of narrow
tubes, frequently florets of Compositae.” Nine of the eleven
plants listed above have tubular flowers; seven are Compositae.

Blossom color preference has been shown in several butterfly
species (Dronamraju and Spurway, 1960). The blossom colors
of the most important nectar sources of adjutrix are yellow (V.
encelioides , H. annum, H. latifolia, Z. hispida and V. dentata ),
white (V. virginica, E. texana and R. trivialis ) and red/yellow
(G. pulchella). Blossom color of those common plants which
were noted as being ignored are purple, blue and pink. Detailed
quantitative field data will be required to determine whether
these color differences between utilized (yellow-white) and non-
utilized (blue-pink) are truly significant but a strong tendency
has been demonstrated. Levin ( 1969 ) reported that both corolla
color and outline influence the number of visits by butterflies to
Phlox blossom. Work with Heliconius charitonius has revealed
results that were interpreted as “spontaneous” color preferences
(Swihart and Swihart, 1970) which may be modified by “con¬
ditioning” (Swihart, 1971). These reported “spontaneous” color
preferences may have been due to conditioning, however, be¬
cause wild-caught specimens were utilized in the original ex¬
periments. One would expect a preference for a particular flower
or group of similarly shaped and/or colored flowers to occur
through experience or “conditioning”. Exceptions concerning in¬
stinctual, i.e. genetic, preferences will occur in highly co-evolved
butterfly-plant systems (e.g. Gilbert, 1975).

Studies by Knoll (1926) and Ilse (1928) as reviewed by
Norris (1936) has revealed variation in methods utilized by
rhopalocera to locate nectar sources. Certain butterflies, e.g.
Papilio and various pierids, appear to be attached to blossoms
entirely by visual cues. Other species, e.g. charaxine nymphalid
species, are guided to food sources entirely by olfactory cues.
Ilse (1928) reports that most butterflies, however, utilize both
classes of cues and therefore comprise an intermediate group in

16(3):147-154, 1977

CHLOSYNE LACINIA

151

continuum. Odor is probably largely utilized by adjutrix to
locate feces, carrion and puddles. Flowers may be located by a
combination of the senses. Restriction of the major nectar sources
to blossoms of white or yellow color classes indicates that vision
is important. The near lack of utilization by adjutrix of Lantana,
which is widely used by such visually-oriented butterflies as
Papilio, would tend to indicate that vision is not of prime im¬
portance. Alternately, the pigments present in Lantana corollas
may be quite different from the pigments present in corollas of
the prime nectar sources.

There is no necessarily direct relationship between larval
foodplant and adult nectar plants. However, observations of
adjutrix populations suggest that subtle interacting influences
may exist between these energy sources for the two active phases
of its life cycle.

Previously (Neck, 1973), I stated, “pure, isolated” stands of
A. trifida had never been observed to be infested by adjutrix.
Since that statement was published, a few broods have been
found in isolated stands of giant ragweed. Such utilization is of
uncommon occurrence. The important factor does not appear to
be lack of adult female oviposition stimulation by either H. an-
nuus or V. encelioides as previously suggested (Neck, 1973).
Lack of concentration of adults due to absence of sufficient
nectar sources for adult energy requirements seems to be the
significant factor (See Neck, ms.) V. encelioides is a prime
source for adjutrix and many other insects. H. annuus is only a
minor nectar source for adjutrix , but plants associated with sun¬
flower are important nectar sources. A. trifida produces little, if
any, nectar, being anemophilous (wind-pollinated). Therefore,
adult adjutrix are not as attracted to ragweed as to other larval
foodplants because of lack of nourishment source. Larval infes¬
tations on giant ragweed tend to be heavier in areas where dis¬
turbance of the soil has allowed colonization by other annuals
which are a nectar source for this butterfly.

Both Zermenia hispida (H.B.K. ) Gray and Viguiera dentata
(Cav.) Spreng. are rated as “occasional” larval foodplants of
adjutrix (Neck, 1973, ms.). Z. hispida appears to be preferred
to V. dentata as a larval foodplant. (unpub. data). Such a pref¬
erence is somewhat surprising as the genus Viguiera is very
closely related to Helianthus (Heiser, 1969). The smell of
crushed leaves of V. dentata, to the human nose, is very similar
to the smell of crushed leaves of Helianthus annuus, the primary

152

RAYMOND W. NECK

J. Res. Lepid.

larval foodplant of adjutrix in central Texas. One possible factor
favoring utilization of Z. hispida is the lengthy blooming season
(late spring to fall) of this species in contrast to the restricted
blooming season (fall) of V. dentata. An adult female adjutrix
which is attracted to flowers of Z. hispida is more likely to detect
individuals of this species as acceptable ovipositional substrates
than individuals of V. dentata which supplies no such visual
attraction during much of the year.

Utilization of Verhesina virginica as a larval foodplant is
quite limited but is more likely to occur during the fall months
(unpub. data). As this period of the year is also the blooming
period of this species, adult attraction to flowers for nectar
could increase the utilization of this plant.

Adult resources are of such importance that they represent
a potential limiting factor in the growth of adjutrix populations.
While the presence of various sugars in nectar (Wykes, 1952;
Van Handel et al., 1972) has long been known and/or assumed,
recent work has revealed that nectar of many species contain
amino acids (Baker and Baker, 1973) which could be utilized
by butterfly visitors. Initial investigations have revealed that
butterfly-pollinated blossoms tend to be richer in amino acids
than bee-pollinated flowers (Baker and Baker, 1973b). The lack
of protein supplied by nectar may limit the reproduction of
adjutrix even if moderate amounts of larval foodplant material
are available.

The effect of nectar shortage is not likely to be as dramatic
in adjutrix populations as in populations of Heliconius ethilla
which has intensively co-evolved with its pollen sources ( Ehrlich
and Gilbert, 1973). However, observations of natural popula¬
tions of adjutrix indicate that nectar is extremely important, at
times, as a limiting factor. Early 1970 was a time of abundant
larval foodplant material, but very low population levels of
adjutrix were present. As a result of deficient rainfall, a greatly
reduced amount of nectar was available because few individual
plants were blooming. A similar situation was observed in spring
1976.

SUMMARY

Adult resources (nectar, dung, carrion and mud puddles) of
Chlosyne lacinia var. adjutrix as observed in central Texas are
discussed. Numerous plants supply nectar to adjutrix but flowers
with white or yellow corollas are favored. Primarily males are

16(3):147-154, 1977

CHLOSYNE LACIN1A

153

attracted to dung, carrion and mud puddles. Relative roles of
visual and olfactory cues in location of resources are discussed.
An indirect relationship between larval foodplant utilization and
adult resource utilization is postulated. At times adult resources
are believed to be a limiting factor in the growth of populations
of adjutrix.

LITERATURE CITED

ARMS, K., P. FEENEY and R. C. LEDERHOUSE. 1974. Sodium: Stimulus
for puddling behavior by tiger swallowtail butterflies, Papilio glaucus.
Science 185: 372-374.

BAKER, H. G. and L. BAKER. 1973a. Amino-acids in nectar and their evo¬
lutionary significance. Nature 241: 543-545.

- . 1973b. Some anthecological aspects of the evolution of nectar-pro¬
ducing flowers, particularly amino acid production in nectar. PP. 243-
264 in Taxonomy and ecology (V. H. Heywood, ed.). Systematics As¬
sociation Special Volume #5, 370 pp, Academic Press, London.

BAUER, D. L. 1953. Butterflies at water holes in central Arizona. Lep.
News 7: 146-147.

DOWNES, J. A. 1973. Lepidoptera feeding at puddle-margins, dung and
carrion. J.'Lep. Soc. 27: 89-99.

DRONAMRAJU, K. R. and H. SPURWAY. 1960. Constancy to horticultural
varieties shown by butterflies, and its possible evolutionary significance.
J. Bombay Nat. Hist. Soc. 57: 1-8.

EHRLICH, P. R. and L. E. GILBERT. 1973. Population structure and cy-
namics of the tropical butterfly Heliconius ethilla. Biotropica 5: 69-82.

FAEGRI, K. and L. VANDER PIJL. 1966. The principles of pollination
ecology , Pergamon Press, Oxford.

GILBERT, L. E. 1972. Pollen feeding and reproduction biology of Helicon¬
ius butterflies. Proc. Nat. Acad. Sci. 69: 1403-1407.

- . 1975. Ecological consequences of a coevolved mutualism between

butterflies and plants, pp. 210-240, in Biochemical coevolution between
plants and their insect herbivores (L. E. Gilbert and P. R. Raven,
eds. ) . U. Tex. Press, Austin, 246 pp.

HEISER, C. B. 1969. The North American sunflowers (Helianthus) . Mem.
Tor. Bot. Club 22(3) : 218 pp.

ILSE, D. 1928. Uber den farbensinn der tagfalter. Vergl. Physiol. 8: 658-
692.

KNOLL, F. 1926. Insekten und blumen. Abh. zool. bot. ges. Wien 12: 1-
646.

LEVIN, D. A. 1969. The effect of corolla color on interspecific pollen flow
in Phlox. Evolution 23: 444-455.

NECK, R. W. 1973. Foodplant ecology of the butterfly Chlosyne lacinia
(Geyer) (Nymphalidae). I. Larval foodplants. J. Lep. Soc. 17: 22-33.

- ms. Ibid. II Additional larval foodplant data.

PAYNE, J. A. and E. W. KING. 1969. Lepidoptera associated with pig car¬
rion. J. Lep. Soc. 23: 191-195.

SHIELDS, O. 1972. Flower visitation records for butterflies (Lepidoptera).
Pan-Pac. Ent. 48: 189-203.

154

RAYMOND W. NECK

J. Res. Lepid.

SWIHART, C. A. 1971. Colour discrimination by the butterfly, Heliconius
charitonius Linn. Anim. Behav. 19: 156-164.

- and S. L. SWIHART. 1970. Colour selection and learned feeding

preferences in the butterfly, Heliconius charitonius Linn. Anim. Behav.
18: 60-64.

VAN HANDEL, E., J. S. HAEGEN and C. W. HANSEN. 1972. The sugars
of some Florida nectars. Am. T. Bot. 59: 1030-1032.

VOSS, E. G. 1954. The butterflies of Emmett and Cheboygan Counties,
Michigan with other notes of northern Michigan butterflies. Amer. Mid.
Nat. 51: 87-104.

WYKES, G. R. 1952. An investigation of the sugars present in the nectars
of flowers of various species. New Phytologist 51: 210-215.

Journal of Research on the Lepidoptera

16(3): 155-161, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

A NEW SUBSPECIES OF COLIAS PALAENO
(LINNAEUS) FROM BAFFIN ISLAND, N.W.T.,
CANADA (PIERIDAE)1

JAMES A. EBNER

P. O. Box 556, Okauchee, Wisconsin 53059

and

CLIFFORD D. FERRIS2

College of Engineering, University of Wyoming, Laramie, Wyoming 82071

In 1973, the senior author received a small number of dark
C olios palaeno (Linnaeus) from the vicinity of Frobisher Bay,
Baffin Island, Northwest Territories. In 1975 and again in 1976,
additional specimens were received from the same locality. When
examination specimens were sent to C. F. dos Passos, C. D.
Ferris and the late Colin Wyatt, all agreed that the Frobisher
C olios were indeed a new form of palaeno as described below.

Although dos Passos cited nominate palaeno in his 1964 Check¬
list, it is a European species as noted by Klots 1951, 1975 (in
Howe). To date, all North American palaeno have been referred
to subspecies chippewa Edwards [TL Restricted by F. M.
Brown, 1973 to west end of Great Slave Lake, Northwest Terri¬
tory, Canada], The species is Holarctic and the range of chip¬
pewa in North America extends from Alaska across arctic and
subarctic Canada to Labrador. Baffin Island, in which Frobisher
Bay is situated, is of significant size and is physically removed
from mainland North America. It is not surprising, then, that an
isolated race of palaeno should develop under such conditions.
The new subspecies is now described.

1 Published with the approval of the Director, Wyoming Agricultural Ex¬
periment Station as Journal Article no. JA 896.

2 Research Associate: Allyn Museum of Entomology, Sarasota, Florida ;
Florida State Collection of Arthropods, Div. of Plant Industry, Fla. Dept,
of Agriculture and Consumer Services, Gainesville. Museum Associate,
Los Angeles County Museum of Natural History, Los Angeles, California.

155

156

J. A. EBNER and C. D. FERRIS

/. Res. Lepid.

Colias palaeno baffinensis Ebner and Ferris, new subspecies
Holotype Male: Frobisher Bay, Baffin Island, N.W.T., Canada,
10-viii-73. Expanse: 18 mm.

Allotype Female (white form): Frobisher Bay, Baffin Island,
N.W.T., Canada, 19-viii-76. Expanse: 21 mm.

The holotype and allotype will be placed in the collection of
the American Museum of Natural History, New York, N.Y.
Paratypes: 17 specimens, 9-12-viii-73; 54 males and 38 females
7-31-vii-75; 13 males and 4 females 26-vii-19-viii-76, all from
Frobisher Bay, Baffin Island, N.W.T., Canada. All material is
from the vicinity of the town of Frobisher Bay.

Paratypes are being distributed to the National Museum of
Natural History, Washington, D.C. and the Canadian National
Collection, Ottawa, Ontario, Canada. Additional paratypes are
placed in the collections of the authors, C. F. dos Passos, Oscar
Dorfmann, Gerald Hilchie and others. The Colin Wyatt collec¬
tion contains examples of the new subspecies.

Figures 1-4

Diagnosis: The new subspecies differs from chippewa and the
European and Japanese subspecies in the very narrow black
borders of the males. This character is consistent and unvarying.
The ratio of the wing width (measurement along the FW costal
margin) to the border width at vein Cu2 is 7.4:1 in the male
holotype of baffinensis; in the specimen of chippewa illustrated
(Fig. 5), this ratio is 6.1:1. These values are typical of the two
subspecies. The white ground color of the females is generally a
more greenish white than in chippewa and the dark markings
tend to be more intense. Ventrally the hindwings in both sexes
exhibit much more dark suffusion than is found in chippewa or
the European and Japanese races, and the HW discocellular
spot is strongly rimmed with dark pink scales not found in the
other subspecies. This melanic condition appears to be genetic
and not related to thermo-regulation, as the melanism exhibited
does not occur in chippewa taken in other localities at the same
or higher latitude. The ventral surfaces are much darker than
the specimen of chippetoa figured by Howe ( 1975, PI. 73, f . 15 ) .
Two subspecies are recorded from western Europe: palaeno
Linnaeus and europome Esper (Higgins & Riley, 1975). The
latter is more brightly colored and the black border in the males
is usually slightly wider than in typical palaeno. Specimens are
shown in Figures 7-8. Nominate palaeno occurs in Fennoscandia,
while europome ranges through portions of France and Germany

16(3): 155-161, 1977

COLIAS PALAENO

157

COL//9S

~/s'*e*j£ /Is
<r /rCs-ris
//ocorysE

/&<*//'"+ Ts.

0.1V T. C /?**&/*
yo - i/+/i. 73

SK. Cm//.

Coc/^S
S*<*Jae./7 o .
4 <*//?* c *sas
£4sf*r-f/&sris
/9ca<%tsSZ

ys *&*'+$/• +r /?*/
Mo»-T. C**"*P/1
y 9- 1////-7 Cm
£ Jr. Cm//- C/rmfcr

EXPLANATION OF FIGURES

Figure 1. — Holotype male, Colias palaeno baffinensis Ebner and Ferris,
Frobisher Bay (town), N.W.T., 10-viii-73, upper side. Label is red im¬
printed with black ink. Figure 2. — Holotype male, underside. Label is
white imprinted with black ink.

Figure 3. — Allotype female, C. p. baffinensis, Frobisher Bay (town),
N.W.T,. 19-xiii-76, upper side. Label is green imprinted with black ink.
Figure 4. — Allotype female, under side. Label is white imprinted with
black ink.

158

J. A. EBNER and C. D. FERRIS

/. Res. Lepid.

Figure 5. — C. p. chippewa, male, Churchill, Manitoba, ll-vii-73, upper
and under sides. Figure 6. — C. p. chippewa, female, same data as male in
Fig. 5, upper and under sides. Fig 7. — C. p. palaeno, male, Les Fonts, H.A.,
France 2000 m, 20-vii-68, upper and under sides. Fig 8. — C. p. palaeno,
female, same data as male in Fig. 7, upper and under sides.

Figure 9. — C. p. aias, male, Asama Mts., Nagano, Japan, 2200 m, 24-vii-74,
upper and under sides. Figure 10. — C. p. aias, female, Mt. Asama, Nagano,
Japan, 2200 m, 23-vii-56, upper and under sides.

16(3):155-161, 1977

COLIAS PALAENO

159

to Rumania. Various forms have been described which have
been elevated to subspecific status by some workers. Petersen
(1947) has discussed the variation that occurs in Scandinavian
populations. Two palaeno subspecies are known from Japan.
The larger and more heavily marked aias Fruhstorfer (Figures
9-10) occurs in the mountains of central Honshu, while sugitani
Esaki is found in the Japanese Alps in the western portion of
central Honshu. The differences between these two subspecies
are slight.

Description: Male. DFW: uniformly dull yellow, discal spot us¬
ually absent, but sometimes faintly indicated; narrow dark bord¬
ers widest at apex and uniformly black to inner margin; variable
in breadth, and sometimes lightly invaded by light streaks along
veins. Fringes pink with costal margin thinly outlined in gray-
black, occasionally pink. Basal area near thorax dark.

DHW : dull yellow, tending to be more greenish than FW.
Dorsal discocellular spot always evident and pronounced as a
cream colored spot; black borders uniformly dark, always less
in breadth than primaries and occasionally interrupted with
lighter streaks along veins. Fringes pink, more pronounced along
inner margin and adjoining outer margin. Basal area darkly suf¬
fused and radiating discally more extensively than in primaries.

VFW: less yellow than above, tending toward white along
tornus to inner margin; dorsal margins apparent. Pink fringes
more contrasted; pink costal margin with inward dark suffusion.
Discal cell spot occasionally present; macular spots rare.

VHW: strongly contrasting with ground color of FW, dark
olive green with intense dark suffusion throughout. Discal cell
spot always well-developed, usually strongly rimmed with dark
pink to brownish-red, occasionally dagger-shaped as in C. hecla
Lefebre. Discocellular spot much larger than in chippewa, more
angular. Fringes pink.

Antennae red-brown above and beneath, pink with club bright
yellow. Eyes brown to brownish-black.

Expanse: base of primary-to-apex: 17-21 mm.

Female. Dimorphic: white form prevalent (3.75:1 ratio) with
some specimens intermediate between white and yellow. Of 43
females examined, 30 were white, 8 were yellow and 5 were in¬
termediate.

DFW: white form: uniformly gray- white. Discal cell spot
absent or inconspicuously present. Dark marginal border always

160

J. A. EBNER and C. D. FERRIS

J. Res. Lepid.

widest at apex, but variable in breadth, sometimes containing
several light markings, either enclosed partially or completely
by a dark band. Fringes pink. Basal area dark brown at thorax.
Costal margin thinly lined with pink edge and sometimes suf¬
fused lightly with dark scales.

DHW: white form: contrasts slightly with FW, usually a more
bluish-gray. Discal spot always evident and cream colored. Outer
marginal black border narrow and always slightly developed.
Fringes pink.

VFW: white form: more glossy white than DFW; apical area
conspicuously tinged with greenish-yellow. Costal margin lined
thinly in pink; fringes pink.

VHW: white form: uniformly dark gray-green. Discal spot
prominent and sometimes silvery. Red-brown rimming usually
strongly evident, but slightly evident in some examples, occasion¬
ally dagger-shaped. Fringes pink and prominent.

Antennae and eyes as in males. Thoracic hairs at base of FW
strongly pink.

Expanse: 19-23 mm.

The yellow form is essentially identical to the white form, but
with all of the white areas replaced by pale yellow.

Variation: Geographic variation cannot be considered, as all of
the specimens have been taken in the same locality. Except for
sexual dimorphism, the subspecies shows remarkably little vari¬
ation. The observed variation relates to the shape of the VHW
discocellular spot as noted above and the dorsal coloration in
the males, which in a very few examples is a dark yellow. A
female Colias specimen from Frobisher Bay appeared to reflect
a hybrid between hecla and palaeno. The ground color was quite
orange, but the dark markings were characteristic of palaeno.

Some specimens of palaeno from both Coppermine and Baker
Lake, N.W.T. appear to be intermediate between chippewa and
baffinensis. The dark rimming of the VHW discocellular spot
is more prominent than in chippewa.

Baffin Island is one of the few areas in North America where
pelidne Boisduval & LeConte and palaeno come into contact.
Hovanitz (1950) discussed Baffin Island pelidne but palaeno
was not recorded from the area. The dot on his distribution map
(Figure 4) occurs in the vicinity of Lake Harbour which is west
of both the town of Frobisher Bay and the body of water bear¬
ing the same name. Rare examples of pelidne phenotypes occur

16(3):155-161, 1977

COLIAS PALAENO

161

in the vicinity of the town of Frobisher Bay and may represent
migrants from a colony elsewhere on the island. In one character,
the strongly rimmed discocellular spot, baffinensis is intermedi¬
ate between the two species. In all other characters, however, it
is clearly palaeno. Perhaps weak introgressive hybridization in
the past has contributed to the discocellular spot character.

Bionomic Notes: The life history of baffinensis is unknown. Be¬
cause of its swift and low flight, the butterflies are difficult to
capture. It appears restricted to the lower elevations of the hills
which surround Frobisher Bay. The butterflies occur in mixed
dry-wet areas on slopes ranging from 300-500 feet (100-160 m)
in elevation. They are absent from the hill tops and the dry lower
regions of the slopes.

Arctic Cotton ( Eriophorum sp., Cyperaceae), Fire Weed and
Willow Herb ( Epilobium sp., Onagraceae) grow in the general
areas where baffinensis flies. It is sympatric with C. nastes Bois-
duval & LeConte and hecla. Five species of Boloria occur in the
region along with Lycaena phlaeas feildeni (M’Lachlan) and
Plebejus aquilo ( Boisduval ) . In the higher and drier areas where
baffinensis does not occur, Erebia rossii ( Curtis ) and two species
of Oeneis may be taken.

LITERATURE CITED

BROWN, F. M. 1973. The types of the pierid butterflies named by William
Henry Edwards. Trans. Amer. Ent. Soc., 99: 29-118.

HIGGINS, L. G. & N. D. RILEY. 1975. A Field Guide to the Butterflies
of Britain and Europe. Collins, London. 381 p.

HOVANITZ, W. 1950. The biology of Colias butterflies. I. The distribu¬
tion of the North American species. Wasman J. Biol., 8(1): 49-75.
HOWE, W. H., Contr. Ed. 1975. The Butterflies of North America.
Doubleday and Co., Inc., Garden City, L. I., New York, xiii -\- 633 p.
4- 97 pi.

KLOTS, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin
Co., Boston, xvi -|- 349 p.

dos PASSOS, C. F. 1964. A Synonymic List of the Nearctic Rhopalocera.

Memoir No. 1, The Lepid. Soc. v -(- 145 p.

PETERSEN, B. 1947. Die geographische Variation einiger fennoskandischer
Lepidopteren. Zool. Bidrag (Uppsala), 26: 329-531.

Journal of Research on the Lepidoptera

16(3) : 162-172, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

DISTRIBUTION OF SHIJIMIAEOIDES RITA ,

ESPECIALLY S. R. RITA AND S. R. COLORADENSIS

(LYCAENIDAE)

OAKLEY SHIELDS

Department of Entomology, University of California,

Davis, California 95616

From a study of Shijimiaeoides ( =Philotes ) recently com¬
pleted (Shields, 1975, and in press), there are a series of fairly
discrete, allopatric S. rita (Barnes & McDunnough) subspecies
that appear to gradually intergrade into one another at the
perimeter of their ranges. This cline is expressed through pheno¬
type, valve teeth counts, and foodplant preferences. I.e., S. rita
emmeli Shields grades into S. r. coloradensis (Mattoni) and
S. r. rita on the east and south and S. r. pallescens (Tilden &
Downey) on the west, and S. r. pallescens grades into S. r. elvirae
(Mattoni) on the west (see Fig. 1). S. r. rita and S. r. elvirae,
although separated by only 200 miles, are at opposite ends of
this spectrum. Likewise, S. r. pallescens and S. r. spaldingi
(Barnes & McDunnough) in the Stansbury Mts., Utah, are very
distinct. Preliminary data on chromosome counts of S. rita are
variable: S. rita emmeli has n = 33, and S. rita pallescens has
n = 34 (2 localities ) and n = 25 ( 2 localities ) ( T. C. Emmel &
O. Shields, unpublished).

In Shields (1975), I move spaldingi to the position of a rita
subspecies since it possesses cristae on the valves. In that paper,
possible ties of spaldingi with r. rita and r. coloradensis are
assumed on morphological and foodplant similarities. S. r.
emmeli flies only in areas of Triassic-Jurassic age sands, while
S. r. spaldingi occupies regions of Upper Cretaceous-Cenozoic
age mostly, at higher, wetter elevations.

S. r. pallescens probably occurs in the little-collected region
of central and southern Nevada to western Utah and north-

162

16(3):162-172, 1977

SHIJIMIAEOIDES RITA

163

Fig. 1.— Distribution of Shijimiaeoides rita subspecies. 1 = S. r. elvirae,
2 = S.r. pallescens, 3 = S.r. mattonii, 4 = S.r. emmeli, 5 = S.r. spal-
dingi (from Shields, 1975, and in press). Black triangles = S.r. rita, black
circles = S.r. coloradensis, half-shaded circles = intergrade populations.
Arrows denote presumed evolutionary trends.

164

OAKLEY SHIELDS

J. Res. Lepid.

western Arizona, where E. kearneyi Tidest. is found on sandy
soils. E. plumatella Dur. & Hilg. grows in southeastern Cali¬
fornia to Mohave Co., Arizona, but so far S. r. elvirae has failed
to turn up there.

Shijimiaeoides rita coloradensis (Mattoni)

Number of teeth on terminal end (cucullus) of valve =
23 (1), 24 (3), 25 (2), 26 (4), 27 (3), 28 (3), 29 (2), 30 (2),
35 (1); N = 21, average = 26-27, from Kendrick and Silver Cliff.
For distribution, see appendix.

HOST: Eriogonum effusum Nutt. var. effusum. COLO¬
RADO: Chaffee Co.: Chalk Creek Trout Farm, viii-11-65, 1$
(J. Scott, JS), adult association ( Shields #138), Scott, Ellis, &
Eff (1968) report oviposition on E. effusum in Fremont and
Lincoln Cos., and adults on this species in Saguache Co. At the
Lowry Bombing Range, Arapahoe Co., Colorado, r. coloraden¬
sis averages small and is in association with an annual Eriogo¬
num species on overgrazed range land (fide Shapiro).

Eriogonum effusum var. effusum (see Fig. 2) is found in
Pennington Co., South Dakota, southward into Converse and
southern Niobrana Cos., Wyoming, southward in southeastern
Wyoming and adjacent extreme western Nebraska to central and
east-central Colorado in the Front Ranges and on the Great
Plains to northern New Mexico from eastern San Juan County
eastward to Union County and with an outlying population in
Socorro County, New Mexico, flowering from June to September
(Reveal, 1971).

Ssp. coloradensis is on wing from early July to late August.

S. r. coloradensis may be found in W. Kansas, SE Wyoming,
and possibly W. Nebraska, as suitable habitats occur there or
once did, although isolated and hard to locate in the farming
areas according to Mike Fisher (in litt.). Scott, Ellis, & Eff
( 1968 ) give some records for coloradensis , repeated here.

Mattoni (1965) described coloradensis in detail for the first
time and compared it with “rather similar” rita rita. From the
results of this study, coloradensis and r. rita are distinguishable
but closely related subspecies that intergrade in north-central
New Mexico. Mattoni (1965) gives some distribution records
for both.

Shijimiaeoides rita rita (Barnes & McDunnough)

Number of teeth on terminal end (cucullus) of valve =
21 (1), 22 (1), 23 (1), 24 (3), 25 (1), 26 (3), 27 (1), 28 (3),
29 ( 1 ) , 31 ( 1 ) ; N = 16, average = 26, all from Ramsey Canyon.

16(3): 162-172, 1977

SHIJIMIAEOIDES RITA

165

tqI'i ?'~Distributi0n °f Eri°g°num eftusum var. effusum (from Reveal,
iy i a / •

166

OAKLEY SHIELDS

J. Res. Lepid.

For distribution and hosts, see appendix.

Kendall (in lift.) says he found larvae of rita rita on E.
wrightii in the flats below Panther Canyon, Big Bend National
Park, Brewster Co., Texas, in 1971. In 1972, he found one larva
on this plant in Green Gulch, and one larva and two adults on
this plant in the approach to Pine Canyon, both in Big Bend
N. P. Comstock (1953) described the egg of rita rita and com¬
pared it with enoptes dammersi (Comstock & Henne).

E. wrightii var. wrightii is found in southeastern California
eastward to western Texas southward to northern Mexico,
flowering from July to September. E. polycladon occurs from
extreme south-central Utah and Arizona across New Mexico to
western Texas southward to northern Mexico, flowering from
July to October. E. rotundifolium ranges from northern New
Mexico southward to Chihuahua, Mexico, eastward into central
Texas and Coahuila, Mexico, flowering from April to October.
(The above information was drawn from Reveal, 1969.) E.
corymbosum var. velutinum is probably only casually used by
rita, as varieties of the species corymbosum are widely used by
battoides ellisii Shields on the Colorado Plateau (Shields, 1975).

Ssp. r. rita is on wing from mid August to late September.

S. r. rita may range into northern Mexico on E. wrightii, but
in southeastern California, extreme southern Nevada, and cen¬
tral Arizona it is replaced by S. enoptes dammersi. In the vi¬
cinity of Cherry and Hillside, Yavapai Co., Arizona, both are
sympatric and synchronic. North, west, and east of the range of
S. r. rita in the vicinity of Prescott, S. enoptes dammersi flies
over E. wrightii at Stanton, Jerome, Mingus Mtn., Sycamore
Canyon, Poland Jet., Montezuma Well, Verde Hot Spgs., Tonto
Natural Bridge, East Verde River nr. Payson, and Sunflower.

Clench (1967) mentions the Silver City, Grant Co., N. M.,
record and the Rio Verde Mts. being NE of Phoenix, for rita
rita.

A specimen of r. rita from the Huachuca Mts., Cochise Co.,
Arizona, no date (AMNH) labelled “P. rita B. & McD., identi¬
fied by Barnes & McDunnough,” one male, bears a red label
“Metatype”; also from this locality at AMNH are 1 $ 1 $ “ex
Barnes Collection,” the female bearing the date VIII-16-23. For
a discussion of the taxonomy of rita rita, see Mattoni (1965)
and Shields (1975).

The “Philotes rita” reported by Garth (1944) from the
southern slopes of the Ajo Mts., Pima Co., Arizona, in April on

16(3) .162-172, 1977

SHIJIMIAEOIDES RITA

167

Eriogonum fasciculatum Benth. is most likely S. battoides mar¬
tini (Mattoni).

Populations in Sandoval and Santa Fe Cos., New Mexico,
are best characterized as clinal between r. rita and r. coloraden¬
sis. At 13 mi. N. Lemitar, Socorro Co., N. M., the majority of
specimens are typical rita rita but some trend toward r. colora-
densis. A short series from SW of Cuba, Sandoval Co., tends
smallish and appears to be a mixture of rita rita , r. coloradensis,
and r. emmeli. A large female S. rita ssp. from Uinta Canyon,
Duchesne Co., Utah, viii-4-59 (J. C. Downey, CIS) is probably
near emmeli or coloradensis although it is not readily classed in
either category.

The S. rita complex is chiefly confined to the primitive species
of the Eriogonum subgenus Eucycla, ranked as the most primi¬
tive subgenus by Reveal ( 1969 ) . The Colorado Plateau in south¬
eastern Utah is likely the place of origin of Eucycla by virtue of
its concentration of endemic species, evolving in the Triassic-
Jurassic sand-dune deserts (Shields, 1975). Following the phylo¬
genetic Eriogonum classification of Reveal (1969, 1971), S. r.
emmeli on primitive E. leptocladon Torr. & Gray would be the
presumed originator of the S. rita complex, giving rise to S. r.
pallescens on E. kearneyi and S. r. coloradensis on E. effusum
(Fig. 1 & Table 1). In turn, S. r. pallescens produced S. r.
elvirae on E. plumatella and S. r. mattonii Shields on E. micro-
thecum Nutt., and S. r. coloradensis gave rise to S. r. rita on
E. wrightii. The derivation of S. r. spaldingi on E. racemosum
Nutt, from S. r. coloradensis would have occurred latest in the
sequence, in Late Cretaceous or Early Tertiary times.

ACKNOWLEDGEMENTS

I wish to thank the following people for loaning material
used in this study: (AM) Allyn Museum of Entomology, Lee
D. Miller; (AMNH) American Museum of Natural History,
Frederick H. Rindge; (RB) Richard Bailowitz; (CAS) Cali¬
fornia Academy of Sciences, Paul Arnaud and Thomas W.
Davies; (CIS) California Insect Survey, Robert L. Langston and
Jerry A. Powell; (CM) Carnegie Museum, Harry K. Clench;
(SE) Scott Ellis; (CF) Clifford Ferris; (MF) Michael Fisher;
Paul Grey; (KH) Keith Hughes; (RK) Roy Kendall; (NL)
Noel La Due; (LACM) Los Angeles County Museum of Natural
History, Julian P. Donahue; (MCZ) Museum of Comparative
Zoology, Harvard, John Burns; (PO) Paul A. Opler; (KR)

168

OAKLEY SHIELDS

/. Res. Lepid.

Kilian Roever; (JS) James A. Scott; (CS) Charles Sekerman;
Arthur M. Shapiro; and (YU) Yale University, Douglas C.
Ferguson and Charles L. Remington. Mike Fisher assisted in
constructing the distribution map of S. r. coloradensis. Dr. James
L. Reveal of the University of Maryland kindly identified all the
Eriogonum species and deposited voucher specimens at the
National Arboretum in Washington, D.C. I thank John Lane
for reviewing the manuscript and offering helpful suggestions.
This work was supported in part by a grant from the Allyn
Museum of Entomology (Arthur Allyn) and an N. S. F. Gradu¬
ate Traineeship.

LITERATURE CITED

CLENCH, H. K. 1967. Further distribution records and taxonomic notes
on Philotes rita ( Lycaenidae ) . J. Lepid. Soc. 21: 141-142.

COMSTOCK, J. A. 1953. Life history notes on four southern Arizona but¬
terflies. Bull. So. Calif. Acad. Sci. 52: 127-136.

GARTH, J. S. 1944. Butterflies of the Organ Pipe Cactus National Monu¬
ment, Arizona. Ent. News 55: 119-124.

MATTONI, R. H. T. 1965. Distribution and pattern of variation in Philotes
rita. J. Res. Lepid. 4: 81-102.

REVEAL, J. L. 1969. A revision of the genus Eriogonum ( Polygonaceae ) .
Ph.D. dissertation. Brigham Young Univ.; Univ. Microfilms, Inc., Ann
Arbor, Michigan. 70-4714.

- . 1971. Notes on Eriogonum— V I. A revision of the Eriogonum micro-

thecum complex (Polygonaceae). Brigham Young Univ. Sci. Bull., Biol.
Ser. 13(1): 1-45.

SCOTT, J. A., S. L. ELLIS, & D. EFF. 1968. New records, range exten¬
sions, and field data for Colorado butterflies and skippers. J. Lepid.
Soc. 22: 159-171.

SHIELDS, O. 1975. Studies on North American Philotes (Lycaenidae).
IV. Taxonomic and biological notes, and new subspecies. Bull. Allyn
Mus. no. 28, 36p.

- . 1977. Studies on North American Philotes (Lycaenidae). V. Taxo¬
nomic and biological notes, continued. J. Res. Lepid. 16: 1-67.

TABLE 1.

Variation in teeth number on terminal end (cucullus) of male valve in

Shijimiaeoides rita subspecies ( Shields,

1975, and

in press).

Average

Range

Sample Size

S. rita coloradensis

26-27

23-35

21

spaldingi

13-15

11-18

35

rita

26

21-31

16

emmeli

20-22

20-26

18

” ” pallescens

17-19

11-24

41

” ” mattonii

—

15-19

3

” ” elvirae

16

13-19

10

16(3): 162-172, 1977

SHIJIMIAEOIDES RITA

169

APPENDIX

Shijimiaeoides rita coloradensis (Mattoni)

DISTRIBUTION (viewed 27c? gen. from 13 localities):

COLORADO: Adams Co . : 1 mi. E Bennett, 5750', viii (MF).

A1 amosa Co . : entrance to Great Sand Dunes Nat'l Mon.,
viii -17-64, 1 cT ( J . F. Emmel , AM). Arapahoe Co . : 0.4 mi.

E Boxelder Crk., 6000', ca. 20 mi. SE Denver, viii (MF);

Lowry Bombing Range, 6000', viii-5-73, 3<?l9 (A. M. Shapiro, AM).
Chaffee Co .: Sal i da, vi i -29-67 , 2c? (J. Scott, JS).

Cheyenne Co . : Aroya (spelled "Arroyo"), viii-21-34, 1?

(C. L. Remington, YU). Custer Co . : 1/4 mi. NW Ben West Hill,

vii- 25-65, 56* (J. Scott, JS, P0); 1/2-1 mi. S Silver Cliff,

viii- 7 & 8-65, 15(3*29 (J. Scott, JS, LACM) ; 2 mi. N Silver

Cliff, vi i - 1 8-65 , 1 ^ 1 9 (J. Scott, JS); 1.5 mi. NW Westcliff,
vii-25-65 , 1 cT 1 9 (j. Scott, JS). El bert Co. : 5-6 mi. E Kiowa,
6300', viii (MF). El Paso Co . : 10 mi. E. Colorado Springs,

Hwy. 94, 6000' (MF); Fountain Valley School & vicinity east
(F. M. Brown, MF); along Hwy. 94 E. of #14, including nr.
Ellicott, Rush, Yoder (MF); locals between Hwy. 94 & north

to Hwy. 24 (Rush to Ramah) along the "Ramah" Hwy. and west
to Peyton and north on county road toward Elbert, slightly
into Elbert Co., distribution fairly continuous from Hwy. 94
north to near Kiowa, first week of August 1973 (MF).

Fremont Co . : nr. Cotopaxi, viii -5-69 • 1 c? 1 9 (MF); 1/2 mi.

NW to 1 mi. W Democratic Mtn., vi i i -20-65 , 1 <T 1 9

(J. Scott, JS); 2 mi. N Hillside, viii-24-68 , 1 c? 39

(J. Scott, JS); Hillside Cemetery, viii-17-65, 50*29

(J. Scott, JS, LACM); 1/2 mi. E Hillside Cemetery, viii-17-65,

170

OAKLEY SHIELDS

/. Res. Lepid.

1? (J. Scott, JS); Hillside School, viii-17 & 23-65 , 1 d* 8?

(J. Scott, JS, LACM); Kuntz Gulch, 1 mi. W Cotopaxi, viii-16-65,
2d*2? (D. Eff, JS, LACM); mouth of Kuntz Gulch, viii-15-65,
12o*17? (J. Scott, JS, PO, LACM); McCoy Gulch, 6800',
viii-16-65, 2<?4?, (J. Scott, JS, PO, LACM). Lincoln Co . :
nr. Jet. 94 & 71 on Hwy. 94 (MF); 2-5 mi. E Arriba & 1-70 east

of there in Kit Carson Co., 5000-5100', viii (MF).

Ri o Grande Co . : "D" Hill, Del Norte, vii-6-68, 1? (S. Ellis, SE).

Saguache Co . : 2 mi. SW Villa Grove, vii-18-66, 5d* 6?

(J. Scott, PO, JS); 4 mi. S Villa Grove, vii-22-69 , 6<?13?,

(J. Scott, JS). Washington Co . : 5 mi. W Last Chance, 5500',
viii (MF). Weld Co . : Hilltop along I-80S (now 1-76) 2 mi.

S. of Empire Resvr., 5000' (MF); IBP site, E. of Nunn, vii,
and vie. of Pawnee Buttes (International Biome Project,

Research Facility, fide Fisher).

WYOMING: Albany Co. : T.15N., R.73W., 7500', vii-9 &

12-72, 2 d* ( C . D. Ferris, CF). Sweetwater Co . : 25 mi.

S Bitter Creek, vii-15 & viii-1-42, 2d* 2? (CM).

Shijimiaeoides rita rita (Barnes and McDunnough)

DISTRIBUTION (viewed 4 3 a* g e n . from 15 localities):

ARIZONA: Cochi se Co . : Huachuca Mts., viii-16-23, 2d*l? ( AMNH ) ;

Ramsey Canyon, Huachuca Mts., viii, ix, 1060*68? (CIS, NL, LACM,
AMNH, AM, P0, JS); Ramsey Canyon, 2 mi. SW of Route 92, 5300',
Huachuca Mts., ix-7-59, 3d*l? (K. Roever, KR). Maricopa Co. :

"Rio Verde Mts.", Phoenix, "Aug. 1893", 2b*(CM). Yavapai Co.:
Cherry, viii-19-53, 2d”l? (J. W. Tilden, AMNH); Cleator, viii-24-74,
ld*(R. Bailowitz, RB); 1 mi. N. Granite Dells, viii-16-76,

10*1? (R. Bailowitz, RB); 1 mi. S Hillside, ix-20-71, 2o*

16(3):162-1 72, 1977

SHIJIMIAEOIDES RITA

171

Peeples Valley, viii-24-76, 1(^1$ (R. Bailowitz, RB); 6 mi. NW Prescott,
viii-21-76, 1$ R. Bailowitz, RB);

(C. Goodpasture, AM); Humboldt, viii-26-53, 3? (J. W. Tilden, AMNH )
Yarnell, viii-21-60, 70*4? (K. C. Hughes, KH, PO, CIS).

County undetermi ned : "So . Arizona. Poling.", 2(^2? (CAS, MCZ).

NEW MEXICO: Dona Ana Co . : San Agustin Pass, Organ Mts. E of
Las Cruces, viii-24-72 , 4(f 4? (L. P. Grey, AM). Hidalgo Co . :
ca. 11 rd. mi. NW Cloverdale, ca. 5500', Coronado Nat'l For.,

Pel one i 1 1 o Mts., viii-24-70, 1 ? (S. L. Ellis, 0. Shields, & M. E.
Toliver, AM). Grant Co . : Silver City, viii-13-?, id*

(J. B. Wallis, CM). TEXAS: Brewster Co .: 21 mi. S Alpine,

ix-20-70, 2? (J. Scott, JS); Big Bend Nat'l Park, ix-25 & 27-71,
ex larvae, 1 cf* 1 ? (R. 0. & C. A. Kendall, RK); Panther Canyon,

Big Bend Nat'l Park, ix-25- 71, 1? (R. 0. & C. A. Kendall, RK);

Pine Canyon, Big Bend Nat'l Park, ix-27-72, 1 d* 1 ? (R. 0. &

C. A. Kendall , RK) .

HOSTS AND DISTRIBUTION: Er i oqonum wr i ght i i Torr. ex
Benth. in DC. var. wri ght i i . ARIZONA: Cochise Co . : jet. of
Ramsey Canyon Rd. & Hwy. 92, 6.0 mi. S of jet. of Hwys. 90 & 92,

SE of Ft. Huachuca, vii -28-70, one larva on flowers, adults
taken commonly here in past (0. Shields) (Shields #125).

Yavapai Co . : Mayer, viii-26-70, 10*2? (C. A. Sekerman, CS),
adult assoc., (area visited with L. Martin) (Shields #181).

NEW MEXICO: Catron Co . : 1/2 mi. S Alma, 5600', ix-11-72,

lo*(0. Shields, AM), adult assoc. (Shields #246). Hidalgo Co.:

22.0 rd. mi. S Animas, ca. 4900', along Hwy. 338, Animas Valley,
viii-24-70, 5<Tl? (S. L. Ellis, 0. Shields, M. E. Toliver, AM),
adult assoc. (Shields #153); ca. 13-14 rd. mi. NE Lordsburg, along

172

OAKLEY SHIELDS

J. Res. Lepid.

Hwy . 90, vi i i -25-70 , 3ld*22? (S. L. Ellis, 0. Shields, M. E. Toliver,
AM), adult assoc. (Shields #155), although most adults at this
locality were roosting on Eriogonum pol ycl adon (Shields #154).

Eri ogonum pol ycl adon Benth. in DC. NEW MEXICO: Grant Co . :

W edge of city limit of Central, along Hwy. 180, viii-26-70,

3? (0. Shields, M. E. Toliver, AM), oviposition (Shields #157);

18 rd. mi. WNW of Silver City, along Hwy. 180, viii-26-70,

2d*l? (0. Shields, M. E. Toliver, AM), adult assoc. (Shields #158).

(Clinal populations between r i ta r i ta and r i ta coloradensis):
Eriogonum ef f usum Nutt. var. eff usum. NEW MEXICO: Sandoval Co . :

2.0 rd. mi. SW of jet. 44 & 85, ca. 5000', along Hwy. 85 (SW of
Bernalillo), v i i i -19-70, 40*2? (S. L. Ellis, 0. Shields, M. E.
Toliver, AM), adult assoc. (Shields #150); 8.0 rd. mi. SW of
jet. of 197 & 44 (SW of Cuba), along Hwy. 197, viii-18-70,

40*2? (S. L. Ellis, 0. Shields, AM), adult assoc. (Shields
#149) ; 18 rd. mi. NW San Ysidro, along Hwy. 44, viii-18-70,

1 d* ( S . L. Ellis, 0. Shields, AM), adult assoc. (Shields #148) .

Santa Fe Co . : 8.4 rd. mi. NE of Cerrillos, ca. 6100', along

Hwy. 10, viii-17-70, 1 <f 1 ? (S. L. Ellis, 0. Shields, AM),
adult assoc. (Shields #144); 10.8 rd. mi. NE of Cerrillos,
ca. 6200', along Hwy. 10, viii-17-70, 1 <? 1 ? (S. L. Ellis,

0. Shields, AM), adult assoc. (Shields #143). Socorro Co . :

13 rd. mi. N of Lemitar, ca. 5000', at La Joya State Game
Refuge exit of Hwy. 25, viii-21 -70 , 25c? 6? (S. L. Ellis,

0. Shields, M. E. Toliver, AM), oviposition (Shields #151) , and
adults also in association with Eriogonum rotundi folium Benth. in
DC. (Shields #152). Eriogonum corymbosum Benth. in DC. var.
vel ut i num Reveal. NEW MEXICO: Santa Fe Co . : 1.1 rd. mi. SW

of Cerrillos, ca. 5800', along Hwy. 10, viii-17-70, 1 d* ( S . L.

Ellis, 0. Shields, AM), adult assoc. (Shields #145).

Journal of Research on the Lepidoptera

16(3) :173-175, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

EVIDENCE FOR TWO ROUTES
OF POST-PLEISTOCENE DISPERSAL IN POANES VIATOR
(HESPERIIDAE)

ARTHUR M. SHAPIRO

Department of Zoology, University of California,

Davis, California 95616

The Sedge-Feeding Marsh Butterfly and skipper fauna
of eastern and central North America shows many interesting
distributional phenomena which were analyzed in an earlier
paper (Shapiro, 1971). That investigation grew out of the ob¬
servation that the Broad-Winged Skipper, Poanes viator (Ed¬
wards), was distributed in a variant of the Coastal Plain —
Great Lakes pattern familiar to plant geographers. At that time
there were no confirmed records in the Mississippi Valley from
northern Iowa, Illinois and Indiana (all within the glaciated
area) to west-central Mississippi (in the Mississippi Embay-
ment, part of the Coastal Plain province). Many experienced
collectors were contacted in the search for such records. In their
absence it was inferred that P. viator, like Euphyes bimacula
( Grote & Robinson ) , had dispersed northward along the Atlantic
Coastal Plain from a glacial refugium in the Deep South and
subsequently colonized the Great Lakes region from the east,
perhaps via the Hudson-Mohawk connective across New York
State. At the conclusion of that study I stated that “new infor¬
mation may well require the re-evaluation of many of the ranges
discussed in this paper.” Such information is now at hand for
P. viator. The critical new records are:

IOWA: Muskrat Slough, Jones County, 5 July 1971, leg. S.
Miller (Miller, 1972). This is at the terminal moraine in east-
central Iowa.

KENTUCKY: Reelfoot National Wildlife Area, Fulton Coun¬
ty, 28 May 1977 and 5 June 1977, leg. L. Gibson and R. Hender¬
son (Coveil, 1977).

173

174

ARTHUR M. SHAPIRO

/. Res. Lepid.

MISSISSIPPI: Bluff Lake, Oktibbeha County, 19 August
1972, and 9.3 mi E Louisville, Winston County, 12 September
1972, both leg. C. T. Bryson ( Bryson, in lift. )

TEXAS: Double Bayou, Chambers County, 13 June 1971,
leg. A. M. Shapiro (in collection of U.C. Davis).

When these records are added to the map of the range of
P. viator given in Shapiro (1971), they provide a strong case for
the northward postglacial dispersal of P. viator on both sides of
the Appalachians (but not in them, as in Speyeria idalia (Dru-

ry)).

The sharp phenotypic and biological disjunction between
viator populations across the well-documented distributional
break in New York State led to the naming of a subspecies,
zizaniae Shapiro — the larger Coastal Plain entity. Now that a
few relict Mississippi Valley populations are known it is possible
that the subspecies will be found to intergrade clinally from
north to south in this part of the range. All of the records re¬
ported here are based on short series, and nothing can yet be
said in any statistical sense. However, the opportunity is now
present to test the alternative hypotheses for the origin of the
Great Lakes populations, both of which are possible. If viator
entered the Great Lakes from the south and then dispersed east¬
ward to its present limit in central New York and north-central
Pennsylvania, the easternmost populations — being the most
recent and farthest removed from the source — might be ex¬
pected to differ the most from zizaniae. If viator entered the
Great Lakes from the east and then spread south — and west¬
ward, the southern-most Great Lakes populations might be ex¬
pected to be the most differentiated from zizaniae.

If the Great Lakes - Coastal Plain pattern is potentially ex¬
plicable in this species by double migration ( with the two groups
of populations never coming into contact across New York), the
same may be true for many or most of the species discussed in
Shapiro, 1971. The best evidence will always be extant relict
populations. Many populations of marsh species might have been
eradicated in the Mississippi Valley during the Xerothermic
interval, giving rise to the present pattern of apparent disjunc¬
tion. All range extensions for the species discussed in the 1971
paper are of potential biogeographic interest and deserve wide
publicity.

16(3): 173-175, 1977

POANES VIATOR

175

LITERATURE CITED

COVELL, C. V. JR. 1977 Reports from the field: big catches. Kentucky
Lepidopterist 3 (2):2.

MILLER, S. 1972. Observations and new records of Iowa Rhopalocera.
J. Lepid. Soc. 26: 229-234.

SHAPIRO, A. M. 1971. Postglacial biogeography and the distribution of
Poanes viator and other marsh butterflies. J. Res. Lepid. 9: 125-155.

Fig. 1. — Distribution of Poanes viator (Shapiro, 1971), with new records
shown by large dots. Dotted line in northeast indicates distributional break
between coastal subspecies zizaniae Shapiro and nominate viator.

Journal of Research on the Lepidoptera

16(3):176-192, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

THE BUTTERFLIES OF
CRATER LAKE NATIONAL PARK, OREGON
J. W. TILDEN1 and DAVID H. HUNTZINGER2

ABSTRACT

Seventy-eight species of Rhopalocera are recorded for Crater Lake
National Park, Oregon. The list is presented in annotated form, with a
discussion of conditions in the Park, and of certain of the records.

INTRODUCTION

Most of the specimens on which this paper is based were
collected in 1957-1962 inclusive. Most of the field work was
done by Huntzinger. Tilden collected in the area in June, 1960,
and in August, 1962, adding a number of previously unrecorded
species. The original intent had been to do considerably more
field work before writing the results, but Huntzinger received
another assignment, and other commitments prevented Tilden
from doing further research in the area. It seems best to present
the results now, so that such work as has been done will not be
lost. It seems certain that this list includes the major number
of butterfly species to be found in the Park. Others will no
doubt be added in time, but it is hoped that the present list
will provide a basis for future work.

In Crater Lake National Park, most of the butterfly species
are concentrated in a relatively few favored areas. Experience
taught that large parts of the Park are depauperate in butterfly
fauna.

The dry area north and west of the lake yielded very few
species or specimens. The peaks (unlike similar areas in many
other mountain masses) support few species not also found at
more moderate levels. Emergence dates, however, are naturally
later for the same species at higher elevations.

The least collected area of Crater Lake National Park is the

1 125 Cedar Lane, San Jose, CA 95127
2 National Park Service

176

16(3):176-192, 1977

CRATER LAKE

177

extreme southwest corner. This lies at a lower elevation than
the rest of the Park, and if new records are sought, this should
be a likely area. One only record is admitted of a species not
actually taken or seen anywhere in the Park. This is Habrodais
grunus (Bdv. ), taken just outside the West Entrance. This
species should occur inside the Park, where oaks are found.

Because of the policy of the National Park Service, large
series of most species were not collected. There are also several
species that were taken only one to a few times. This may
reflect either scarcity of the species, or incomplete collecting.
Relative abundance was estimated from the number of times a
species was seen in the field. Many localities were explored, from
which no specimens were taken. Such localities are not men¬
tioned by name. In those cases in which a species was found
in a number of localities, and seems to be well-distributed in
the Park, this is mentioned under the species annotation.

The senior author has checked all specimens that he could
examine. Only a few were unavailable for study. The physical
basis for this list is housed in four collections, at least for the
present: The Collection of Crater Lake National Park, located
at Park Headquarters; The Collection of San Jose State Uni¬
versity, San Jose, California; the small private collection of
David H. Huntzinger; and the collection of J. W. Tilden. The
most of the Crater Lake specimens now in this latter collection
will later be placed in the California Academy of Science’s
collection.

The sequence of families and species is primarily that of
dos Passos, 1964. The determinations are those of the senior
author, who assumes responsibility for them. Some populations
seem to merit confidence at the subspecific level, others do not.
Mention is made in each case where the population does not
seem to fit any usual named entity.

PREVIOUS WORK

The first specimens retained in the collection of Crater Lake
National Park were taken by H. A. Scullen in 1930-1931. Ten of
these specimens remain. It is not known if there were originally
more. There is also one specimen taken in 1931 by F. Lyle Wynd.

Crater Lake Nature Notes for 1951 has a two-page article
(pp. 10-11) on Crater Lake Butterflies, and lists 18 species,
some of which were not taken by the present authors. No speci¬
mens were found as a basis for this list, which is by Donald C.

178

J. W. TILDEN and D. H. HUNTZINGER

/. Res. Lepid.

Lowrie. Therefore the determinations cannot be checked, but
there seems to be no good reason to doubt them. The article
also describes outbreaks of the California Tortoise-shell ( Nym -
phalis calif ornica) for 1930-1931, and for 1951. The bulk of the
specimens in the collection of Crater Lake National Park were
placed there by the junior author during the years 1957-1960.

BUTTERFLY HABITATS OF THE PARK

Crater Lake National Park is situated in Klamath County,
Oregon. Elevations within the Park range from 4500 ft. at the
South Boundary and 5956 ft. at the North Entrance, to 8938 ft.
at the summit of Mt. Scott, located in the northeast part of the
Park. The high points around the Rim of the Lake exceed
8000 ft. The greater part of the Park lies between 6000 and
7000 ft. in elevation.

In spite of its name, Crater Lake is not in fact a crater, but
a caldera, caused by the collapse inwardly of Mt. Mazama.
Portions of the present Rim have been glaciated. Much of the
entire Park has been covered deeply by pumice and scoria,
ejected at the time of the collapse. On such soils the flora is
sparse and lacking in variety, with few butterfly food plants
present. As a result, the butterfly fauna of such areas is cor¬
respondingly poor. However, certain butterfly species, the food
plants of which are present, reach good populations in these
areas.

It is believed that much of the flora was destroyed by the
glowing avalanche of pumice and scoria at the time of the
collapse. This has been discussed by Tilden (1963b). The flora
that has regrown is uneven in distribution and its density seems
to depend inversely on the depth to which pumice covered
the surface. Areas of light pumice fall recovered more rapidly
and more completely, and now have the densest and most varied
flora, and the greatest variety of butterfly species as well.

The Park is drained by several small streams, which run
down the slope and out of the Park. Sand Creek, running east
out of the Park at the now unused South Entrance, runs through
an area of heavy pumice fall, and supports along its course a
rather meager number of plant species. Annie Creek, running
south out of the Park, flows through an area that has largely
recovered, especially where it reaches the South Boundary. The
canyon of Annie Creek is at times rather spectacular, and there
are small meadows and groves of stream-side trees. This area,
which lies slightly east of the road on which one enters from
the south, and which extends from Annie Creek Station south

16(3):176-192, 1977

CRATER LAKE

179

out of the Park, is the most favored part of the Park for butter¬
flies, and several species were taken only here.

On the South Boundary on the margin of Annie Creek there
are meadows and forest glades, mixed with sagebrush and
Eriogonum, where fly several species that barely enter the Park
at this point. The dirt road that runs east across Annie Creek at
the South Boundary leads into an area of mixed coniferous
forest, meadow, streamside vegetation, aspen grove, and sage¬
brush, offering the greatest number of habitats in the Park, and
showing at present little effect of pumice fall.

Small montane meadows occur at several localities in the
Park. Among these are Pole Creek Meadows, the meadows
below Vidae Falls, and those in the vicinity of Kerr Notch.
These meadows support populations of certain butterflies. Other
favored localities are near Park Headquarters, and on the
glaciated portions of the Rim, where pumice is absent or less
evident.

The Pumice Desert supports very few butterflies, and these
appear to originate elsewhere. Few if any seem to develop there.

The higher points around the Rim of the Lake may be
considered subalpine in nature. The butterfly species in these
localities are few, but some species occur in fair numbers.
Speyeria egleis is found here more commonly than it is else¬
where in the Park.

An annotated list of the butterfly species so far known to
occur in Crater Lake National Park follows. Collectors’ names
are given in parentheses. References to Lowrie’s 1951 paper are
cited as Lowrie ( 1951 ) . As stated above, no specimens are
known to support these records. For the collections, the follow¬
ing abbreviations are used: Crater Lake National Park Collec¬
tion, CLNP; San Jose State University Collection, SJSU; J. W.
Tilden, JWT; David H. Huntzinger, DHH.

Hesperiidae

1. Ochlodes sylvanoides (Bdv. )

South Boundary, 11. VIII. 62 (Tilden) (JWT). Very com¬
mon, both sexes, though apparently not previously reported
in the Park.

2. Polites sabuleti (Bdv.)

South Boundary, 11.VIII.62 (Tilden); Pole Creek, 25.VII.59;
Annie Springs, 16.VII.57, 30.VII.58, 3.VIII.59; Park Head¬
quarters, 28.VII.57; “C. L. N. P.”, 27.VIII.58 (all Hunt-

180

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

Zinger); East Boundary, 11.VIII.62 (Tilden). MacNeill,
who examined all of the skippers for this study, noted that
these specimens represent an unnamed population of Polites
sabuleti. However, considering the variability of this spe¬
cies, it seems unnecessary to add any more names. The
senior author has specimens of three other populations of
P. sabuleti that he has for many years been reluctant to
name.

3. Polites sonora (Scud.)

South Utility Area, 1. VII. 59 (Huntzinger, JWT). So far as
known, these fresh specimens (2 $ $ ) are the only ones
so far examined from the Park. MacNeill notes that they
seem to represent a blend zone between P. sonora sonora ,
and P. sonora siris , being darker below than usual sonora ,
but by no means as dark as siris.

4. Hesperia harpalus oregonia (Edw. )

Well-distributed throughout the Park in the meadows.
Greatly attracted to flowers. All records are for late July
and August. Determination by MacNeill.

5. Hesperia juba (Scud.)

South Boundary, 11.VI.59 ( S ); Annie Springs, 18.VI.59
( $ ), (both Huntzinger, JWT), both much worn. The ap¬
parent scarcity of this species may be an artifact of col¬
lecting. It may fly earlier than most of the collecting was
done.

6. Carterocephalus palaemon (Pallas)

South Boundary, ll.VI. 59 (Huntzinger). Taken within the
Park only the one time. The location of this specimen is
in doubt; it was not seen in any of the collections.

7. Pyrgus ruralis (Bdv. )

South Utility Area, 11.VI.59 ( $ ), 14.VI.59 ( $ ); Vidae
Falls, 25.VI.58 ( $ ) (all Huntzinger) (CLNP); South
Boundary, 22. VI. 60 ( $ ) (Tilden, JWT). Found sitting on
the ground in openings.

8. Erynnis icelus (Scud. & Burg.)

South Boundary, 11.VI.59 (2$ $) (Huntzinger); same,
22-23. VI. 60 (3 $ $ ) (Tilden); Annie Creek Road, 9. VII. 64
( $ ) (Tilden); all JWT. Found streamside, in vicinity of
willows and Populus. Determination by J. M. Burns.

9. Erynnis persius (Scud.)

South Boundary, 11.VI.59 ( $ ) (Huntzinger); same, 22-23.
VI. 60 (4$ $ ) (Tilden) (all JWT). Found together with
Erynnis icelus. In New England, Scudder found that E.

16(3):176-192, 1977

CRATER LAKE

181

persius larvae ate willow and Populus. The food plants for
this Oregon population are unknown. Burns labelled these
as E. persius complex.

10. Erynnis pacuvius lilius (Dyar)

Annie Springs 4. VIII. 58 ( $ ); South Boundary, 11.VI.59
( $ ) (both Huntzinger); same, 22-23. VI. 60 ( $ $ ) (Til-
den) (all JWT). Found in the lower southern part of the
Park where its presumed food plant, Ceanothus velutinus,
is common and forms an understory in the Ponderosa Pine
forest.

11. Thorybes mexicana nevada (Scud.)

Pole Creek, 6.VII.59 ( $ ); Park Headquarters, 17.VII.59
( 9 ) ( both Huntzinger, both JWT ) . This species appears to
be quite scarce in the Park. These specimens were taken
at an intermediate elevation. Why it should seem to be
absent from higher elevations is not clear.

Papilionidae

12. Parnassius clodius claudianus Stichel

Fairly common, especially around the Rim, where in places
may be found quantities of its food plant, Dicentra. Two
species of Dicentra , D. formosa and D. uniflora , occur in
the Park; D. formosa is the larger and more common species.
At the time this work was done in Crater Lake NP, the life
history of P. clodius was not known. In the Sierra Nevada
of California, at intermedite levels, the larvae, which are
purplish black with yellow markings (and look a good deal
like certain millipedes), may be found in the day time
under the leaves at the base of the plants. The subspecies
there is usually considered to be P. c. sol.

Lowrie ( 1951 ) called the Crater Lake Parnassius , baldur,
but they seem closer to claudianus.

13. Papilio zelicaon Lucas

Reported by Lowrie (1951). Seen, but not taken, by Hunt¬
zinger. There is no specimen in the CLNP collection.

14. Papilio rutulus Lucas

South Boundary, 20. VII. 58 (Huntzinger, CLNP); same,
22.VI.60 ( $ ) (Tilden, JWT).

15. Papilio eurymedon Lucas

South Boundary, 20. VII. 58 (Huntzinger, CLNP). The
Papilios are not as scarce as the few records would imply.
Much of the Park is steep and offers few opportunities to
take these soaring butterflies.

182

J. W. TILDEN and D. H. HUNTZINGER

/. Res. Lepid.

Pieridae

16. Neophasia menapia menapia (F. & F. )

Whitehorse Camp, 25.VIII.30 (H. A. Scullen) (CLNP);
“C. L. N. P”, 22.VIII.58 (Huntzinger, CLNP); listed by
Lowrie (1951). This late species is probably more com¬
mon after most of the summer collectors have gone.

17. PierS beckerii beckerii Edw.

Listed by Lowrie ( 1951 ) . The specimen does not seem to be
in the Park collection. However, there seems to be no reason
why the species should not occur in the Park, since it has
been found in the lower Sand Creek Area outside the Park.

18. Pieris sisymbrii Bdv.

The Watchman, 13. VII. 59 (Huntzinger, CLNP). The single
record may be late for this usually early species.

19. Pieris protodice Bdv. & Lee.

Sun Notch, 8.X.53 (B. C. Wood, CLNP); Kerr Notch, 11.
VIII. 62 (Tilden, JWT); Lowrie, 1951, lists Pieris protodice
vernalis Edw.

20. Pieris occidentalis occidentals Reak.

Near Union Peak, 13.VII.59; Park Headquarters, 10.VII.59
(both Huntzinger). Normal-looking P. occidentals , not
form calyce Edw.

21. PierS napi (L. ), probably marginals Scud.

North Junction, 10. VII. 68 (Huntzinger). This species should
be more easily found earlier in the season.

22. Pieris rapae (L).

Park Headquarters, 20. VII. 59 (Huntzinger); Kerr Notch, 11.
VIII. 62 (worn) (Tilden). (both JWT). It is evident that
North Junction, 10. VII. 68 (Huntzinger); Kerr Notch, 11.
this introduced pest has invaded the Park.

23. Colias eury theme Bdv.

Taken by Scullen, Park Headquarters, 2.IX.30 and East
Entrance, 28.VIII.30. Listed also by Lowrie, 1951. Taken
several times and seen frequently during the present study.
Found throughout the Park. All specimens seen were the
summer form amphidusa Bdv.

24. AnthocharS sara flora Wright

Sun Creek, 23.VI.60 ( $ ) (P. A. Fosterla); Park Headquar¬
ters, 15. VI. 60 ( $ ) (Huntzinger) (both JWT). Others were
seen but not taken.

16(3): 176-192, 1977

CRATER LAKE

183

Lycaenidae

25. Habrodais grunus herri Field

Two miles west of Crater Lake National Park, Rogue River
National Forest, 1.IX.59 (1$ 2$ $, worn) (JWT). This
species, the larvae of which feed on oaks, especially Quercus
chryslepis, may occur in the southwest corner of the Park.
For this reason, these specimens from just outside the Park
are included here.

26. Satyrium behrii behrii (Edw.)

South Boundary, 11.VIII.62 ( 5 ) (Tilden, JWT). Much of
the Park seems unsuited to this species and its food plant,
Purshia (Bitter Brush, Antelope Brush).

27. Satyrium saepium (Bdv. )

Park Headquarters, 31.VII.59 (Huntzinger, CLNP). This
species, the larvae of which feed on various species of
Ceanothus, should be more prevalent than this one record
would indicate.

28. Satyrium calif ornicum (Bdv.)

South Boundary, 11. VIII. 62, common but worn (Tilden,
CLNP, JWT). Should be common in its proper season,
along the southern part of the Park.

29. Incisalia augustinus iroides (Bdv.)

South Boundary, 11. VI. 59, South Utility Area, 14. VI. 59
(both Huntzinger, CLNP). These two records, from the
southern limits of the Park, are all for this species. It should
be more widely distributed.

30. Incisalia eryphon eryphon (Bdv.)

This pine-feeding species is the most generally distributed
hairstreak in the Park. It is very unobtrusive, sitting on pine
foliage or visiting flowers of Pussy Paws ( Calyptridium
umbellatum ) .

31. Mitoura spinetorum (Hew.)

South Boundary, 22-23. VI.60 (Huntzinger & Tilden, CLNP,
JWT). Several specimens, at flowers of Calyptridium. All
were worn. The food plants, species of Dwarf Mistletoe
( Arceuthobium ) are common, but the butterfly is quite
irregular in distribution.

32. Mitoura nelsoni (Bdv.)

South Boundary, 23. VI.60 (Tilden, JWT). One only, a $ in
good condition, at flowers of Calyptridium. A close exami¬
nation of the area yielded no others. Associatel with Incense
Cedar, the presumptive food plant.

33. Callophrys lemberti Tilden

184

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

Arent Peak, 3.VII.58 (Huntzinger, CLNP); The Watchman,
12.VII.59 (Huntzinger, JWT); South Boundary, 22. VII. 60
(Tilden, JWT). This recently recognized species seems not
to be abundant anywhere in its known range. It should be
looked for at high elevations. The South Boundary speci¬
men is unexpected, and is badly worn. It may be referable
to some other species, but its condition makes identification
uncertain. It is left here since no other species of the genus
than lemberti is found in the Park.

34. Lycaena heteronea gravenotata Klots

South Boundary, 11.VIII.62 (Tilden, JWT). A single 9,
worn but recognizable. The heavy spotting UNH indicates
that it represents gravenotata Klots, which has been taken
at Diamond Lake, just north of Crater Lake. Collecting in
July should locate fresh material in the Park.

35. Lycaena editha (Mead), near montana Field

South Boundary, 22.VI.60 (Tilden, CLNP); same, 11.VIII.62
(Tilden, JWT). Both males, at flowers of Pussy Paws
(Calyptridium) . Widely distributed in the West, this spe¬
cies should occur more commonly and at higher elevations
than these two specimens indicate.

36. Lycaena mariposa Reak.

The most common copper in the Park, often found at flowers
in the meadows of the Annie Creek drainage, flying from
late June to August. The specimens from the park are not
entirely typical mariposa, but show darkening, especially
of the outer edge both above and below, and are somewhat
intermediate to penrosae Field.

37. Lycaena nivalis nivalis (Bdv. )

Park Headquarters, 31.VII.59 (Huntzinger, DHH); Vidae
Falls, 28.VIII.59 (Huntzinger, JWT); Kerr Notch, 8.VIII.58
(Huntzinger, CLNP); same, 10.VII.59 (Huntzinger, DHH);
same, 11.VIII.62 (Tilden, JWT); also listed by Lowrie,
1951. The few records are clustered on the south and east
shoulders of Mt. Mazama, from Park Headquarters to Kerr
Notch, at median elevations. These localities show less effect
from the pumice fall. These are nearer to nivalis from the
Sierra Nevada, than they are to brotvni dos Passos, which
occurs further north and east.

38. Lycaena helloides helloides (Bdv.)

South Boundary, 20.VII.58 (Huntzinger, CLNP, JWT);
same, 11. VI. 59 (Huntzinger, JWT); same, 22-24. VI.60 (Hunt¬
zinger & Tilden, JWT); Lost Creek, 11.VI.59 (Huntzinger,

16(3) .176-192, 1977

CRATER LAKE

185

DHH); North Entrance, 9.VII.58 (Huntzinger, CLNP).
Common along the southern edge of the Park.

39. Lycaena cupreus cupreus (Edw.)

South Boundary, 22-23. VI. 60 (3$ $ 1$ ) (Tilden, JWT).
The finding of this species, usually associated with high
elevations, at a moderate elevation and in one locality only,
is a bit puzzling. All were found on a small sandy stream-
side flat, visiting flowers of Calyptridium. Others were seen
but not taken.

40. Lycaeides ar gyro gnomon ricei (Cross)

This large showy Blue is one of the characteristic species
of the Park, and apparently of the Cascades in general. It
flies from late June to at least mid-August, and may be
found along roadsides as well as in the meadows. The sub¬
specific name ricei is used, following Nabokov (1949), but
the late F. Chermock informed me (in Litt. ) that he had
seen the type of ricei Cross, and that in his estimation it
was a specimen of some subspecies of Plebejus icarioides,
and for that reason, he had proposed the name fretchini
Chermock for the Oregon population of argyrognomon.

41. Plebejus saepiolus (Bdv. )

Common in the moist meadows on the south and east
shoulders of the mountain. These appear to represent an
unrecognized population. The males are large and brightly
colored; the females are completely brown, and have the
submarginal lunules of the hind wing developed in most
individuals. However, there are so many recognizably dif¬
ferent populations of saepiolus that it seems useless to
propose more names at present.

42. Plebejus icarioides (Bdv.)

In Crater Lake National Park, as in many parts of the wes¬
tern states, this is the commonest and the most ecologically
tolerant of the larger “Blues”. It is always associated with
perennial lupines, and is usually found wherever these
grow. The males of the Crater Lake population have wider
dark borders than most, and look most like the Arizona
subspecies, P. i. buchholzi dos Passos.

43. Plebejus lupini (Bdv.)

Annie Springs, 4-6. VIII. 58 (Huntzinger, CLNP); Kerr Notch,
8.VIII.58 (Huntzinger, JWT); The Watchman, 13.VII.59
(Huntzinger, JWT); South Boundary, 23. VI. 60 (Tilden,
JWT). Lowrie (1951) lists Plebejus acmon (West. & Hew.),
which may occur in the Park, or he may have referred to

186

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

the insect here treated as lupini Bdv. Without specimens,
this point cannot be settled. The females from Crater Lake
are unusually large and dark.

44. Agriades aquilo podarce (F. & F. )

Boundary Springs, 5. VII. 57 (Huntzinger, JWT); Pole Creek,
8.VIII.58 (Huntzinger, CLNP); same, 6.VII.59 (Hunt¬
zinger, JWT, DHH); same, 23.VII.59 ( Hountzinger, DHH);
Annie Springs, 6. VII. 58. This species is found only in very
wet or marshy meadows, and so far has been taken in the
Park only in the Annie Creek drainage. The Crater Lake
population resembles podarce from the Sierra Nevada of
California, much more closely than it does megalo McDun-
nough from further north.

45. Everes amyntula (Bdv.)

South Boundary, 23. VI. 60 (2$ $ ) (Tilden, JWT); South
Utility Area, 23. VI. 60 ( $ ) (Huntzinger, JWT). The male
from the South Utility Area is much larger than the two
females from the South Boundary.

46. Shijimiaeoides battoides oregonensis (B. & McD.)
Generally distributed in the Park, flying in June at the
South Boundary, and in August on the slopes of Mt. Scott.
It is fairly common, but inconspicuous, staying close to
its food plants, which in the Park seem to include several
species of perennial Eriogonum.

47. Glaucopsyche piasus piasus (Bdv.)

Pole Creek, 23.VI.59 (Huntzinger); South Boundary, 23.VI.
60 (2$ $ ) (Tilden, JWT). This is seldom a common
species.

48. Glaucopsyche lygdamus Columbia Skin.

Rather common; a number taken and others seen. It flies
a little earlier than the other large blues, and is worn by
mid- July. Found flying along trails as well as over the
meadows.

49. Celastrina argiolus echo (Edw. )

Fairly common, flying early in the year for this elevation,
the records mostly in early and mid-June; no records after
early July, and these worn. This usually multivoltine spe¬
cies would appear to be one-brooded in the higher moun¬
tains of the west.

Nymphalidae

50. Limentitis lorquini (Bdv.)

16(3):1 76-192, 1977

CRATER LAKE

187

Annie Springs, 19.VI.58 (Huntzinger, CLNP); Vidae Falls,
29.VII.59 (Huntzinger, CLNP); Pole Creek, 11.VIII.62 (Til-
den, JWT). Reported by Lowrie (1951). More common
than these few records indicate. The brick red of the wing
tips is reduced, approaching the subspecies ( or form )
burrisonii Maynard.

51. Adelpha bredowii calif ornica (Butler)

South Boundary, 20. VI. 58 (Huntzinger, CLNP); The Watch¬
man, 14. VIII. 59 (Huntzinger, CLNP). This species would
not be expected to occur commonly in the Park, because of
the absence from much of the region, of the oaks with
which it is associated. However, it is a wide-ranging species
that sometimes occurs as a straggler far from its point of
origin. It should be found regularly in the southwest corner
of the Park.

52. Vanessa atalanta rubria (Fruhst. )

Park Headquarters, 10. VIII. 60 (Huntzinger, CLNP, 2 speci¬
mens). The single record of this conspicuous butterfly may
indicate actual scarcity in the Park.

53. Cynthia virginiensis (Drury)

Listed by Lowrie (1951). No specimens seem to exist for
Crater Lake National Park. Lowrie (p. 10, second para¬
graph, line 4) mentions the “Painted Lady,” but Cynthia
cardui does not appear on Lowrie’s list on p. 11.

54. Cynthia cardui (L. )

North Entrance, 30. VII. 57 (Huntzinger, JWT); same,
17.VII.58 (Huntzinger, CLNP); Annie Springs, 10.VIII.58
(Huntzinger, JWT). Reported by Huntzinger to be very
common in Crater Lake National Park during the summer
of 1958. This was an outbreak year, and a migration year
for this species over the southwestern United States. The
Painted Lady is commoner in the Park than the few records
show.

55. Cynthia anabella Field

Huntzinger reports seeing this species several times when
he was not collecting. He also has seen a colored slide
taken by Richard Brown when Brown was an Assistant
Park Naturalist in 1957. It is also reported by Lowrie (1951).
However, to date no actual specimen for the Park is known
to exist.

56. Junonia coenia (Hbn.)

Huntzinger has seen a colored slide of this species, taken in
the Park. No specimen seems to exist. It is entirely possible

188

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

that this and the two previous doubtful species may occur
within Crater Lake National Park, but evidence in the form
of specimens would be desirable.

57. Nymphalis calif ornica calif ornica (Bdv. )

Huntzinger found this species in outbreak numbers every¬
where in the Park in 1958. Tilden found similar outbreak
numbers in all parts of the Park that he visited in 1962.
The Ceanothus was nearly defoliated in many places.
Lowrie (1951) describes similar outbreaks for 1930 and
for 1951. It appears that Crater Lake National Park is sub¬
ject to outbreaks of the California Tortoise-shell, as are so
many other places in the Pacific States. These outbreaks
are in irregular cycles (gradations) and the number of
years from peak to peak is not predictable.

58. Nymphalis milberti furcillata (Say)

Near Park Headquarters, 22.VIII.30 (Scullen, CLNP);
Sleepy Hollow, 15. VI. 58 (Huntzinger, CLNP); Top of
Wineglass, 5. VII. 57 (Huntzinger, JWT); reported by Lowrie
(1951) (as Aglais milberti Godt. ). N. milberti occurs in all
the western states in the mountains. Overwintering adults
appear as soon as the snow is melted.

59. Nymphalis antiopa (L. )

Annie Springs, 18. VI. 59 (Huntzinger, CLNP). Reported by
Lowrie (1951) (as Aglais antiopa Linn.). This species must
be more common in the Park than records show.

60. Polygonia faunas rusticus (Edw. )

Near Park Headquarters, 29. VIII. 30 (Scullen, CLNP); South
Boundary, 23. VI. 60 (Tilden, JWT). Reported by Lowrie,
1951. The South Boundary specimen is much worn, and
may be overwintered. The small numbers may reflect the
type of collecting. It should be more common, especially
along the Annie Creek drainage.

61. Polygonia zephyrus (Edw.)

The common Anglewing of the Park, found in forest open¬
ings and along trails, often in the vicinity of its food plants,
Ribes spp. (Currant, Gooseberry). An August specimen is
freshly emerged. Individuals noted in June and July are
worn, and probably overwintered.

62. Chlosyne palla (Bdv.)

South Boundary, 20.VII.58 (Huntzinger, CLNP); same,
23.VI.60 (4$ $ 1$ ) (Tilden, JWT). So far taken in an
area of small extent, in the immediate vicinity of Annie
Creek where it runs out of the Park. The habitat is moist

16(3) :1 76-192, 1977

CRATER LAKE

189

and overgrown. These specimens are not referred to any
subspecies. They resemble whitneyi Behr, from the Sierra
Nevada of California, more than they do nominate palla.

63. Chlosyne hoffmanni segregata (B. & McD.)

Quite generally distributed in the Park, the common checker-
spot, from South Boundary to near the Rim at Kerr Notch.
It flies in meadows and forest glades, and a wide range of
dates indicates a rather long flight period, with later emerg¬
ence at higher elevations. Since segregata was described
from the vicinity of Crater Lake National Park, these speci¬
mens should be quite typical.

64. Phyciodes campestris (Behr)

Park Headquarters, 31. VII. 59 (Huntzinger, CLNP); Meadow
just below Vidae Falls, 13. VII. 59 (Huntzinger, CLNP). This
seems to be an uncommon species in the Park. The few
specimens appear closest to montana Behr.

65. Phyciodes mylitta (Edw.)

Union Peak, 13.VII.59 (Huntzinger, CLNP); Park Head¬
quarters, 17. VII. 59 & 26. VIII. 59 (Huntzinger, CLNP).
Rather oddly, these records are for localities well within
the Park and at fair elevations, not from the lower levels.

66. Euphydryas editha lawrencei Gund.

Arent Peak, 3. VII. 58 (Huntzinger, CLNP); East Rim, near
base of Mt. Scott, 11.VIII.62 (Tilden, JWT, 1$). The
meager records suggest that this is not a common insect in
the Park. Both captures are for high elevations and late in
the season. While these are here referred to lawrencei
Gunder, there is some question as to whether they may be
remingtoni Burdick. There may even be some doubt that
remingtoni Burdick is distinct from lawrencei.

67. Boloria epithore chermocki Perk. & Perk.

South Utility Area, 22. VI. 59 ( $ ) (Huntzinger, JWT); Pole
Creek, 22.VI.59 (2$ $ ) (Huntzinger, JWT); same, 11.
VIII. 62 ( $ ) (Tilden, JWT); South Boundary, 23.VI.60
(2$ 2 ) (Tilden, JWT). So far this species has been taken
in the Park only in the Annie Creek drainage. It has been
found only in lush streamside situations, but is locally
common. A specimen exists in the CLNP collection, data
unknown.

68. Speyeria coronis (Behr)

Pole Creek, 24.VII.57 (?) (Huntzinger, TWT); South
Boundary, 24. VI. 60 ( 2 ) (Tilden, JWT). These are the
only records for the Park, and S. coronis must be considered

190

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

rather uncommon in the area. Outside the Park, along Sand
Creek, specimens were taken by Tilden in 1957 and in 1962
(June). All specimens resemble snyderi Skinner very close¬
ly, and might be referred to this subspecies.

69. Speyeria zerene (Bdv. )

South Utility Area, 12. VII. 59 (Huntzinger, CLNP); Park
Headquarters, 31. VII. 59 (Huntzinger, CLNP); South Bound¬
ary, 23.VI.60 & 11.VIII.62 (2$ $ 2? 2 ) (Tilden, JWT);
Vidae Falls, 10.VIII.62 ( $ ) (Tilden, JWT); Kerr Notch,
11. VIII. 62 ( 2 ) (Tilden, JWT). The several specimens
show great variation. Some closely resemble subspecies
conchyliatus from northern California; others are as light
as some specimens from eastern Oregon. This species typi¬
fies the range of variation seen in certain Speyeria popula¬
tions in the Crater Lake region, as noted by Tilden (1963).

70. Speyeria callippee (Bdv.)

Stewart Falls, 17. VII. 57 ( $ ) (brown disc) (Huntzinger,
JWT); North Gate, 9. VII. 57 ( $ ) (green disc) (Huntzing¬
er, JWT). The only two so far found within the boundaries
of the Park. This species is quite common east out of the
Park in the Sand Creek Basin. The Stewart Falls specimen
would fit a series of Uliana H. Edw. from Lake Co., CA.
The North Gate specimen has a dull green disc with the
ground color showing through, and resembles the dull green
population of the arid Oregon interior. Such variation in
two specimens of S. callippe shows the futility of attempt¬
ing subspecific determinations of Speyeria from regions in¬
habited by unstabilized populations, such as are found in
the volcanic regions of interior Oregon.

71. Speyeria egleis (Behr)

The most common Speyeria in the Park, and the only one
found in numbers at higher elevations. S. egleis from the
Sand Creek basin east out of the Park are quite variable,
but those taken inside the Park are quite uniform in appear¬
ance, and seem referable to oweni (Edw.), closely resem-
ling specimens from Mt. Shasta.

72. Speyeria atlantis dodgei (Gunder)

North Entrance, 30.VII.57 ( 2 ) (Huntzinger, JWT); Vidae
Falls, 28. VII. 59 (Huntzinger, CLNP); same, 11.VIII.62 ( 2 )
(Tilden, JWT); South Boundary, 23. VII. 60 ( $ ) (Tilden,
JWT); same, 11.VIII.62 ( $ 2 ) (Tilden, JWT). Found in
suitable habitats here and there in the Park. Several seen
and not taken.

16(3) :17 6-192, 1977

CRATER LAKE

191

73. Speyeria hydaspe (Bdv.)

Pole Creek, 24.VII.59 ( $ ) (Huntzinger, JWT); Vidae
Falls, 28.VII.59 (Huntzinger, CLNP). These seem to be
the only records for the Park. The specimens are a bit light¬
er than purpurascens H. Edw. from northern California.

Danaidae

74. Danaus plexippus plexippus ( L. )

There is a specimen in the CLNP collection. This butterfly
must occur in the Park. It has most likely been neglected in
the search for less conspicuous species.

Satyridae

75. Coenonympha California eryngii H. Edw.

Annie Springs, 18.VI.59 (Huntzinger, CLNP); South Bound¬
ary, 22.VI.60 (Tilden, SJSU, JWT). The small numbers
probably indicate, not scarcity, but that this insect flies
earlier, before most collectors arrive for the summer. It
seems to be absent from higher elevations.

76. Cercyonis pegala boopis (Behr)

Pole Creek Meadow, 15.VIII.30 (Scullen, CLNP); South
Rim Crater Lake National Park, 6. VIII. 30 (Scullen, CLNP);
reported by Lowrie, 1951. Not taken by Tilden & Huntzing¬
er. Lowrie’s inclusion may be on the basis of Scullen’s speci¬
mens, which are listed in the Crater Lake NP Collection,
and in Lowrie, as alope Fab.

77. Cercyonis oeta oeta (Bdv.)

The most common and widely distributed satyr in the Park.
It flies from late June into August, in almost all of the more
open and arid habitats.

78. Oeneis nevadensis nevadensis (F. & F.)

South Utility Area, 14.VI.59 (Huntzinger, JWT); South
Boundary, 20.VII.58, worn (Huntzinger, CLNP); same,
22-24. VI.60, common (Tilden & Huntzinger, SJSU, JWT);
Vidae Falls, 10. VIII. 62, badly worn (Tilden, JWT). Except
for the one record from Vidae Falls, this species seems to
be found in the Park only along the southern limits, where
it flies in openings and glades in coniferous forest. It is
known to have a two year cycle. In the Crater Lake area
its periods of abundance are on the even-numbered years.

192

J. W. TILDEN and D. H. HUNTZINGER

J. Res. Lepid.

ACKNOWLEDGEMENTS

The senior author wishes to thank the National Park Service
for collecting permits which made a part of this study possible.
He also wishes to thank Bruce W. Black, Chief Park Naturalist
at the time that the last of the field work was done, for his help
and cooperation.

REFEBENCES

BRIGGS, LYMAN J. 1962. When Mt. Mazama lost its top. National Geo¬
graphic, 122(1) : 128-133.

LOWRIE, DONALD C. 1951. Butterflies of Crater Lake National Park.
Crater Lake Nature Notes, Vol. XVII, 1951, pp. 10-11.

TILDEN, J. W. 1963. The Argynnis populations of the Sand Creek Area,
Klamath Co., Oregon. J. Res. Lep. 1(2) :109-113.

WILLIAMS, HOWELL. 1961. Crater Lake— The story of its origin. Uni¬
versity of California Press, Berkeley & Los Angeles, XII + 98 pp., 10
pits., 8 figs.

NOTICES

NEW ADDRESS: The address of the Journal is now: c/o Santa Barbara
Museum of Natural History, 2559 Puesta Del Sol Road, Santa Barbara,
California 93105.

MAILING DATES: As of this issue, the mailing date of each number of
the Journal will be listed in the Notice column. Volume 16, Number 3:
December, 1977.

NEW BOOKS: Available from Entomological Reprint Specialists, P. O.

Box 77224, Dockweiler Station, Los Angeles, California 90007, USA:

Allan, P. B. M. Leaves from a Moth Hunter’s Notebook. 1977. $9.00.
Dennis, R. L. H. The British Butterflies: Their Origin and Establish¬
ment. 1977. $20. 00.

Goater, Barry. The Butterflies and Moths of Hampshire and the Isle
of Wright. 1975. $13.00.

Holloway, J. D. Moths of Borneo, with Special Reference to Mount
Kinabalu. 1976. $14.00.

Pinhey, E. C. G. Moths of Southern Africa. 1975. $35. 95.

FOR SALE: 1, 500 exotic butterflies, insects, biologicals. 72 page catalog
$1.00. Complete Scientific, P.O.Box 307, Round Lake, IL 60073.

XERCES SOCIETY: An international organization dedicated to the conser¬
vation of terrestrial arthropods and their habitats. Publications include
the journal Atala and the newsletter Wings. Memberships and subscriptions
US $5. 00 (students $2. 50); contact treasurer, Roger Pasquier, 235 East
73rd Street, New York, NY 10021.

BUY/SELL/EXCHANGE: Lepidoptera. Send $1.00 for list of species (world¬
wide). Specialist in Sphingids, Saturniids and Exotics. Ray Adams, F. R. E. S
Carretera Principal 139, Tamaraceite, Las Palmas De Gran Canaria.

WANTED: World moths, particularly Geometridae. Will trade North
American or Panamanian moths. Roger L. Heitzman, Dept, of Ent. , Univ.
of MO, Agric. Bldg., Columbia, MO 65201.

FOR SALE: 6m by 2. 5m linear Malaise Traps. Proven design used world¬
wide by military, universities and private collectors. Additional information
available from Dana A. Focks, P. O. Box 12852, Gainsville, FL 3260
Dept. LRF.

v v

EXCHANGE: Lepidoptera specimens. B. Cech, Vancurova 13, 150 00 Prague
Czechoslovakia.

WANTED: Correspondence with members interested in trading or selling
North American Lepidoptera. Send list of offers to: Marc Grocoff, 1950
Cottrill Lane, Westland, MI 48185.

MANUSCRIPTS: May be sent to the Assistant Editor at the Journal address.

Volume 16

Number 3

September , 1977

IN THIS ISSUE

Charidryas flavula Barnes and McDunnough
( Nymphalidae ) : A question of identity

Clifford D. Ferris and Michael S. Fisher

Ocellus variation and wingspan in

Attacus atlas Linnaeus, Is there a relationship?

Mark Hadley

Foodplant ecology of the butterfly

Chlosyne lacinia ( Geyer ) ( Nymphalidae )

III. Adult resources Raymond W. Neck

A new subspecies of Colias palaeno (Linnaeus)
from Baffin Island, N.W.T., Canada (Pieridae)

James A. Ebner and Clifford D. Ferris

Distribution of Shijimiaeoides rita, especially
S.r. rita and S.r. coloradensis (Lycaenidae)

Oakley Shields

Evidence for two routes of post-pleistocene
dispersal in Poanes viator ( Hesperiidae )

Arthur M. Shapiro

The butterflies of Crater Lake National Park,

Oregon

133

141

147

155

162

173

J. W. Tilden and David H. Huntzinger

176

MUS. COMP. ZOOU.
LIBRARY

AU6 3 0 1978

harvard

UNIVERSITY

Number 4

December, 1977

THJE JOURNAL OF RESEARCH
ONI THJE LEPI DORTER A

published by

The Lepidoptera Research Foundation, Inc.
at

2559 Puesta Del Sol Road, Santa Barbara, California 93105
FOUNDER: William Hovanitz
ACTING EDITOR: Rudolf H. T. Mattoni

ASSISTANT EDITORS: Karen Nielson Hovanitz, Scott E. Miller
Associate Editors :

Thomas C. Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601

Maria Etcheverry, Centro de Estudios Entomologicos, Casilla 147, Santiago,
Chile.

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.

CLASSES OF
MEMBERSHIP

Regular

$12

year

Family

15

year

Contributing

25

year

Subscribing

50

year

Sponsor

100

year

Life

250

for life

Subscriptions to the Journal are $15.00 per year. Special subscriptions are avail¬
able to students at $10.00 per year.

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 2559 Puesta Del Sol Road, Santa
Barbara, California 93105. The publisher is THE LEPIDOPTERA RESEARCH FOUN¬
DATION, INC. The general editor is R. H. T. Mattoni at the above address. The
secretary-treasurer is Barbara Jean Hovanitz at the same address. The owner is THE
LEPIDOPTERA RESEARCH FOUNDATION, INC. THE LEPIDOPTERA RESEARCH
FOUNDATION, INC. is a non-profit organization incorporated under the laws of the
State of California in 1965. There are no bond holders, mortgages or other security
holders.

Journal of Research on the Lepidoptera

16(4): 193-200, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

PHOTOPERIOD AND TEMPERATURE
IN PHENOTYPE DETERMINATION OF PACIFIC SLOPE
PIERINI: BIOSYSTEMATIC IMPLICATIONS1
ARTHUR M. SHAPIRO

Department of Zoology, University of California,

Davis, California 95616

For the past dozen years I have been studying seasonal
phenotype determination in the family Pieridae. As you all
know, many butterflies and moths which have several gener¬
ations a year display seasonal “forms” or phenotypes, often so
different in appearance that they were described as different
species; their conspecificity was proven only by breeding them,
one from the other, or preferably by split-brood experiments in
which different rearing conditions elicited different phenotypes
in the progeny of single females. From the earliest days of
evolutionary biology down almost to the present, such seasonal
variation was subsumed by the familiar Victorian word “poly¬
morphism.” In the past decade, however, particularly since the
advent of “electrophoretic genetics” — in which allelic frequen¬
cies are estimated by examining the mobility of enzymes in
an electric field for a suitably large sample of the population in
question — the word “polymorphism” has been increasingly ap¬
propriated by geneticists to refer specifically to variation which
has a genetic, rather than an environmental, basis. This re¬
striction can be traced to E. B. Ford’s now classic definition of
polymorphism as the occurrence in a population of more than
one allele at a locus, such that the rarest allele is too common
to be maintained solely by recurrent mutation. What, then, are
we to call seasonal phenotypic variation which is under en-

1 Read by John H. Lane at the 23rd annual meeting of the Pacific Slope
section of the Lepidopterists’ Society, Nevada State Museum, Carson City,
Nevada, July 2, 1976.

193

194

ARTHUR M. SHAPIRO

J. Res. Lepid.

vironmental control? We can call it just that; or we can use Ernst
Mayr’s convenient word polyphenism, coined in parallel to poly¬
morphism, and defined as “the occurrence of several phenotypes
in a population, the differences between which are not the re¬
sult of genetic differences.” Studies in various laboratories —
H. J. Muller’s and Rolf Reinhardt’s in East Germany, Ward
Watt’s at Stanford, and our own — are establishing how wide¬
spread and complex a phenomenon seasonal polyphenism is in
the Lepidoptera. We are finding, much as the great develop¬
mental geneticist C. H. Waddington foresaw twenty-some years
ago, that developmental plasticity provides an alternative to
genetic polymorphism which allows natural populations to cope
with a variable environment in an economical, adaptive way.
We are also finding that studies of the physiological mechanisms
determining phenotype allow us to interpret the history of a
specific group’s climatic and geographic adaptation — and this
in turn gives us clues bearing on the perennial problems of
species diversity and community stability.

Mayr pointed out in 1961 that any adaptation can be studied
from the standpoint of proximate or ultimate causality. Any
time we ask a “why” question in biology, we are really asking
several questions at different levels all at once. Let us consider
the familiar Gray-Veined White, Pieris napi (sens. lat. — if you
buy B. C. S. Warren’s single-character taxonomy you can leave
the room now). Everyone knows, or thinks he knows, that Cal¬
ifornian napi are seasonally diphenic — they have a spring phen¬
otype with black veins, known as venosa , and a summer one
without black veins, called castoria. I’ll show in a minute or
three that you don’t really know that, but first let’s ask “ Why
does Pieris napi have two seasonal phenotypes?” There are
several possible answers. At the proximate, or immediate, level
we have physiological mechanisms which translate from some
environmental cue to an eventual phenotype by affecting me¬
lanin pigment synthesis and deposition. The most complete
studies of such systems have been done not on Pieris napi but
on the Nymphalid Polygonia c-aureum in Japan by Hidaka,
Aida, Fukuda, Endo, and Takahashi. Their papers, beginning 13
years ago, are not well known in this country; most English-
speakers are still laboring under the misapprehension fostered by
E. B. Ford in a casual remark in Butterflies, that the seasonal
forms of Polygonia are food-determined. Actually P. c-aureum is
broadly representative of seasonally polyphenic butterflies in

16(4):193-200, 1977

PACIFIC SLOPE PIERINI

195

that the primary factor controlling adult phenotype is larval
exposure to daylength (photoperiod), with some input from
temperature. That shouldn’t be too surprising. In middle lati¬
tudes photoperiod is a much more reliable predictor of season
than temperature is: the daylength is essentially the same on
every June 25, but the temperature may be wildly different.
These Japanese workers have shown with painstaking thorough¬
ness that long days interacting with high temperatures stimulate
certain cells in the larval brain to produce a hormone, which
travels through the nerves to the corpora cardiaca of the pupa,
whence it is released into the hemolymph (blood) where it in¬
duces development of the summer adult phenotype. But that is
not the end of the story: the dark, summer animals breed im¬
mediately, but light, winter animals are quite uninterested in
sex — and Endo wanted to know why. It turns out that ovarian
maturation and female pheromone (sex-stimulant chemical)
production are also under neurosecretory control — the relevant
hormone being daylength-dependent. In Pieris napi, unlike Poly-
gonia species, there is no adult hibernation and sexual behavior
is not at issue. But the hormonal regulation of adult phenotype
is probably very similar.

We have now studied about a dozen napi populations from the
Yukon to central New Mexico, including a variety of Californian
ones. It appears that adult phenotype in the Gray-Veined White
is under the control of photoperiod, temperature, and certain
genes, interacting in complex — and immensely interesting —
ways. The basic situation in nature is that pupae which go into
programmed dormancy — “diapause” as it is known to physi¬
ologists — produce dark-veined, i.e. spring phenotypes, and
those which develop directly, without diapause, produce light-
veined, or summer ones. This applies even to populations which
are normally single-brooded in nature, with “obligate” diapause
— we can prevent diapause by appropriate rearing conditions,
which lie outside the range of real environments the animals
would encounter afield. Since the primary control of diapause is
photoperiodic, until quite recently it was assumed that pheno¬
type was physiologically coupled to development — that the
hormonal control of diapause also affected pigment synthesis and
deposition. But I now no longer believe that — at least, not quite.
The blame for my disillusionment lies with August Weismann.

196

ARTHUR M. SHAPIRO

J. Res. Lepid.

You may remember Weismann’s name from your genetics
classes; he first enunciated the dogma of the isolation of the
germ cells from environmental influences — the critical repudi¬
ation of Lamarckian heredity, the inheritance of acquired
characteristics. He was one of the greatest of the early Darwin¬
ians, and until his failing eyesight forced him to become a
theoretician, he worked on Leps, including Pieris napi. Now,
Weismann was a monstrous clever fellow: 70 years before Mayr
he talked about multiple levels of causality, and he even dis¬
cussed seasonal polyphenism from that standpoint. When he
bred napi , biological photoperiodism hadn’t been thought of yet;
the influence of daylength on insect development remained to
be discovered in the 1940s by Danilyevskiy in Russia. So Weis¬
mann figured temperature was the environmental cue, and de¬
signed experiments accordingly; and he got ambiguous results,
at least with Pierids. Since we now know these critters to be
strongly photoperiodic, I felt it necessary to repeat Weismann’s
experiments under controlled daylengths to determine whether
there really might be a temperature effect. Not having central
European napi , I settled for coastal central Californian.

It turned out that certain chilling treatment applied to non¬
diapause pupae quite unambiguously produced the spring phen¬
otype! The identical treatment was much less effective on Inner
Coast Range material. This difference among stocks under¬
scores the rather obvious fact that physiological mechanisms
have a genetic basis — the interface between Mayr’s proximate
and ultimate levels of causality. Rut how is one to account for
the apparent redundancy of photoperiodic and temperature
mechanisms? The fact is that the photoperiodic mechanism has
never been unambiguously demonstrated at all! Diapausing
pupae will never eclose unless chilled for a number of weeks.
Although we know that temperature treatment has no effect on
adult pigmentation when the pupa is in deep diapause, what if
diapause intensity gradually decreases, allowing for increasing
sensitivity to environmental cues? This is precisely the clinical
picture of diapause being advanced by Tauber and Tauber at
Cornell (see their recent review in Annual Review of Ento¬
mology). We are currently testing this hypothesis by monitoring
the day-to-day metabolism of diapausing napi pupae and sub¬
jecting them to various temperature regimes when they begin
to wake up.

16(4) -.193-200, 1977

PACIFIC SLOPE PIERINI

197

Underlying whatever proximate, i.e. physiological, answers to
our “why” question are genetic answers, and these reflect a
history of natural selection. The ecological basis of selection —
the adaptive value — is the ultimate level of causality. Ward
Watt at Stanford, formerly at Yale, followed up on Leigh and
Smith’s work and demonstrated the efficacy of seasonal pheno¬
types in body-temperature regulation of the Orange Sulphur,
Colias eurytheme. Our work points in the same direction for
Pieris napi. One striking aspect is the consistent sexual difference
in napi — in all populations and in both seasonal phenotypes,
males are more heavily marked than females. Field studies sug¬
gest that this confers an adaptive advantage in that the darkest
males can become active at lower temperatures in the morning,
giving them first crack at newly-emerged virgin females. ( Pieris
napi , like most butterflies, has a diel periodicity of emergence,
with most eclosions occuring shortly after sunrise even at low
temperatures. )

I hope the preceding remarks have given you some feel for
the directions in which polyphenism studies are going. But I’d
like to wind up with some comments on their biosystematic
implications for Pieris napi, because I have a petition in to the
International Commission on Zoological Nomenclature that is
apt to raise some eyebrows when it appears in the Bulletin. I
might add, with no snobbery intended, that I commit taxonomy
only when dragged into it kicking and screaming by the biology.

I said several minutes ago that everyone “knew” that Californ¬
ian napi had two seasonal phenotypes, venosa and castoria. Last
year Bob Langston suggested at these meetings that they might
be two different species; our work dispels that notion. Clearly,
they are seasonal phenotypes produced by the same genome
responding to different environmental cues. The problem — as
Langston first pointed out to me — is that what we have been
calling “castoria” isn’t what Reakirt described as castoria in
1867! Here is Reakirt’s O.D.:

Size and form of Pieris oleracea, Harris.

Male, upper side pure white, inner half of costa of
primaries, and base of both wings, strewn with a
few dark atoms; a rounded black spot in the medio-
superior interspace of the fore wings ... no other
markings; fringes white, expanse 2-2.12 inches.

198

ARTHUR M. SHAPIRO

/. Res. Lepid.

Underneath immaculate white; a faint yellowish
tinge on the apex of the primaries, and along the
costa of the secondaries.

Body black, with whitish hairs below; antennae
black, with incomplete white annulations inter¬
rupted above. Club yellowish, or yellowish brown
at tip.

Hob. — California. Coll. Tryon Reakirt.

Herman Strecker, in 1877, was the first to consider castoria
as the second brood phenotype of double-brooded California
napi; in this he was followed by W. H. Edwards in his (1881)
revision of the napi group, and by most authors since. Yet very,
very few specimens collected in the wild from double-brooded
populations come near to this description. Almost all of them
have some dark scaling on the veins beneath, at the apex of the
fore- or hindwing above, or whatever. When it turned out that
the so-called type of castoria at the Field Museum was a pseudo¬
type ( in fact it is a typical second brood specimen from a coastal
population, matching Strecker’s sense but not Reakirt’s descrip¬
tion of castoria!), it was exceedingly difficult to find prospective
neotypes that matched the O.D. Now, as it happens the O.D.
matches perfectly the normal phenotype produced by non¬
diapause pupae of interior, single-brooded California napi. Con¬
fused? Let’s backtrack a bit.

Our experiments concur with field data in suggesting that
there are two subspecies of napi in central California. Sub¬
species venosa is found on and near the coast, in places subject
to summer fog; it is best developed from San Francisco south.
It is at least partially double-brooded, with the two seasonal
phenotypes. In the hot, dry, fogless Inner Coast Range and on
the west slope of the Sierra Nevada occurs a single-brooded
subspecies, showing only a spring phenotype in nature. The old¬
est available name for this population is usually considered to
be microstriata Comstock, 1925. The spring phenotype of micro-
striata is lighter than that of venosa reared under identical con¬
ditions. When diapause is artificially prevented, microstriata
will make a summer phenotype in the lab, and it matches per¬
fectly Reakirt’s O.D. of castoria — being lighter than the sum¬
mer phenotype produced by venosa. Recently we have found
a couple of places, in cold canyons, where microstriata is natur¬
ally double-brooded and makes a summer phenotype just like
our lab one. This raises an uncomfortable taxonomic specter.

16(4):193-200, 1977

PACIFIC SLOPE PIERINI

199

Suppose it could be shown that what Reakirt had was not a sec¬
ond brood coastal specimen at all, as everyone has now assumed
for 99 years, but one of the very rare interior ones? Then the
oldest valid name for the inland subspecies would be castoria,
and we would be obliged to use it; the familiar usage for the
second-brood phenotype of venosa would be scrapped, and the
name microstriata would sink into oblivion.

And everyone would hate me.

Where did Reakirt’s type come from? “California,” says the
O.D. But F. Martin Brown has shown that Reakirt never col¬
lected in California; he got his stuff from Lorquin. Where did
Lorquin get it? Unfortunately there is still too little evidence
at hand to reconstruct his itineraries very well; we know he
collected in both venosa and microstriata country, though. I
was wrestling with this when I noticed that, in the same paper
as he describes castoria , Reakirt also describes Pieris yreka.
Now, this is rather an infamous description since it can only
apply to rapae — and the type, at the Field Museum, is a spring
rapae — thus implying that rapae was in California before 1867,
which is inconsistent with the idea of a single introduction in
southern Canada about 1860 — but we wander. The point is that,
although the type locality of yreka is not specified, it’s a good bet
it was the town of Yreka. I checked this out in that invaluable
resource, Gudde’s California Place Names , and found that Yreka
was named in 1852. Well, if Yreka was the type locality of
Pieris yreka , was Castoria the type locality of Pieris castoria?
Heres what Gudde says about the town of Castoria: Castoria =
French Camp, San Joaquin County . . . known as Castoria (Latin
for beaver) from 1850-59 . . . formerly the headquarters of
French beaver trappers on the San Joaquin River . . . ! Could
Lorquin, a Francophone, stay away from such a place?

Alas, there are to be no topotypes. 125 years ago French Camp
was in a maze of riparian woodland and marshland; today it
has been thoroughly civilized, and there are no napi there. In
fact, there are no napi currently known anywhere on the floor
of the Great Central Valley. Still, on climatic and biogeographic
grounds we can infer that any population at French Camp must
have been of the interior subspecies — thus our taxonomic
nightmare becomes real.

So I have asked the Commission to do something very odd.
I have asked them to suppress the name castoria under the

200

ARTHUR M. SHAPIRO

/. Res. Lepid.

Plenary Powers for synonymy but not homonymy. If they do, it
will be unavailable for use at the subspecific (or specific) level,
but can continue to be used informally, as a seasonal phenotype
name, the way it is now and has been for 99 years. It was neces¬
sary to take this roundabout route because microstriata hasn't
been used often enough in the literature to qualify for conserva¬
tion under Article 79 — yet common sense demands that nearly
a century of usage of castoria not be upset just because some
dingbat at Davis finds that there’s more to napi than meets the
eye. Like I said, I do taxonomy only under compulsion.

Now do you see why I’m not here?

Journal of Research on the Lepidoptera

16(4) :201-207, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

ANTENNAL SENSILLA OF SOME CRAMBINAE1

J. A. KAMM2
Agric. Res. Serv. USD A,

Corvallis, OR

ABSTRACT

When the antennal sensilla of several common species of Crambinae
were studied with the scanning electron microscope, trichoid sensilla were
dominant; the primary function of Crambinae antennae may therefore be
detection of semiochemicals. Sensilla chaetica ( mechanoreceptors ) were
well represented and would be useful to the moths for orienting within
the grass canopy. Sensilla coeloconica and styloconica were present, but
sensilla auricillicum were absent. The sensilla of Crambinae are simple
and functional relative to those of many Lepidoptera and appear to be
designed for orientation within grassland habitat.

Many Lepidoptera explore a large habitat (e. g., a forest)
and make relatively long flights in response to various stimuli
that are perceived by antennal sensilla of various sizes, arrange¬
ments, densities, and sculpturing (Callahan, 1975). Adult Cram¬
binae (the grass-feeding larvae are commonly referred to as sod
webworms ) , in contrast, make short, erratic flights ( 3-5 meters )
within or slightly above the canopy of grass and also explore
the interior of the canopy by walking. The male perceives and
orients to the sex pheromone of females during flight ( Banerjee
and Decker, 1966; Kamm, 1974). Both sexes, particularly females
heavy with eggs, often fall to the ground in response to mechan¬
ical stimuli and feign death rather than attempt escape by flight.
Since information about the antennal sensilla of the Crambinae
would help to understand the mechanism by which adults per¬
ceive their environment, a study was made to determine the
types of sensilla on the antennae of several common species of
Crambinae and to relate the types with known functions to the
behavior of the moths.

1 Contribution of the Agric. Res. Serv. USDA in cooperation with the
Agric. Exp. St., Oregon State Univ. Technical Paper No. 4492 of the
latter.

2 Mailing address: Department of Entomology, Oregon State University,
Corvallis, OR 97331.

201

202

J. A. KAMM

J. Res. Lepid.

MATERIALS AND METHODS

Study moths were reared in the laboratory from insects col¬
lected near Corvallis, Ore. (Kamm 1970). The antennae of five
species of both sexes were excised from newly emerged adults,
cut into pieces of several segments, and mounted on a specimen
stub with fast drying silverpaint. The mounted antennal seg¬
ments were then coated with a thin layer of gold-palladium
alloy and observed with the scanning electron microscope. In
this way, the entire circumference of the flagellum of females
could be scanned by using a single segment, but two segments
had to be used in observing both sides of the male sensilla since
they surround most of the flagellum. Segments from five anten¬
nae of each sex were examined for each species. Measurements
of sensilla were made at random locations on the antennae but
sensilla that arose from or were adjacent to the scale area of
the flagellum were avoided because they were often of atypical
shape and size. The terminology of Schneider (1964) was used
to classify the sensilla described herein. The species studied
were: Crambus tutillus McDunnough, Chrysoteuchia topiaria
(Zeller), Pediasia trisecta (Walker), Crambus leachellus cypri-
dalis Hulst, and Tahama bonifatella (Hulst) all of which are
commonly found in the grasslands in the Willamette Valley of
Oregon.

OBSERVATIONS

The antennae of all five species were typical setiform and had
a scape, pedicel, and variable number (45-72) of flagellar seg¬
ments. Also, the spine-like Bohm bristles were present in groups
on the scape and pedicel of all species. In general, the number of
sensilla on each flagellum decreased toward the distal end of
the antenna. The shape of the flagellum was variable and dif¬
ferent among species and sexes. Males always had more sensilla
per flagellum, and they covered an estimated 75 percent or more
of the surface area of the flagellum; the remaining area on all
species except F. trisecta was covered with scales, but F. trisecta
had some scales among the field of the sensilla. The sensilla on
females covered 50 percent or less of the surface area of a typical
segment. This difference between sexes was just dicernible on
the high power (45 x) of a binocular microscope with good res¬
olution. The net-like sculpturing on the flagellum was similar
among species but lacked the rough textured terrain common
on the antennae of some Lepidoptera (Jefferson et al., 1970).
The range in size ( p) of sensilla is given in Table 1.

Table 1. Range in size (y) of 10 randomly selected sensilla on the antennae of five species of Crambinae.

16(4):201 -207, 1977 ANTENNAL SENSILLA 203

03

+->

o

CM

o

ID

r—

CM

(U

CO

CM

CO

f—

CO

•r—

1

1

1

1

1

i-

4->

CT>

r—

t—

cn

I'-S

CM

1 —

CM

O-’l

ro

r—

r—

<u

+->

O

o

CM

o

■=3-

03

q_

CO

CM

r —

1

l

1

1

1

c

o

n—

CO

r—

-Q

CM

i—

l~

T.

CO

•I—

r—

03

-a

•r—

un

CO

CO

o

i-

CO

r—

Q.

i

1

i

i

1

CJ

, _

CO

•

CM

l —

CM

' '

ol

CO

O

i —

ID

CM

CT)

i —

co

CM

CO

4->

1

1

1

i

1

=3

-4->

ID

ID

ID

ID

r->.

CM

CM

ol

( X3

•r—

ID

s-

( o

' CM

i —

CO

CM

•r—

CO

CM

Cl

o

1

1

1

1

i

+->

r-N

CM

CO

r->.

o

Ol

CM

r—

CM

03

r—

E ^

r—

„ — ^

O E

•r—

+-)

S- ^

CO

•1—

CM 4- n-

C

t—i

03 Q.

03 r—

<u

»— <

l— l

CJ

u C 0J

CO

•r- 4-

•r-OO)

■a

-o

03

c o

C -r- 03

4—

■r—

• r —

CJ

o

O 4-> r-

O

o

o

• !—

CJ sz

CJ 03 4-

_c:

-c

+->

O 4->

o >

QJ

cj

cj

( u

r— -O

i— a> cu

CL

•r—

•r—

03

aj •<-

>-) l — VO

>>

S-

J-

J c

o 5

+-> a> 03

1—

h-

1—

C_)

o ^

to - - JO

Measurements made with millimeter rule from micrographs.
Reliable measurements of the sensory cone not possible.

204

J. A. KAMM

J. Res. Lepid.

Two types of trichoid sensilla were distinguished: type I
(T-I) was curved and sharply hooked at the tip; type II (T-II)
was shorter than type I, was slightly curved though it lacked the
pronounced hook at the tip, and was almost always adjacent to
type I sensilla (Fig. 1-A). Type I was the most abundant sen¬
silla on the antennae of both sexes. Neither type had any dis¬
cernible longitudinal or annular striations on the surface (Fig.
1-B).

Sensilla chaetica (Ch) were present on most flagellar segments
on all species (Fig. 1-C). High magnification revealed some
with longitudinal or annual striations (Fig. 1-D) but most
had a smooth surface. No constant pattern or distribution on the
flagellar segments was apparent, but the terminal segments of
both sexes often had several sensilla chaetica that extended be¬
yond the apices which suggested a tactile function.

Sensilla coeloconica (Co) occurred in clusters of 2-3 sensilla
on most flagella (Fig. 1-E) and were nearly always located away
from the scales of the flagellum. However, the peg is corrugated
longitudinally in C. topiaria and arises from a shallow pit sur¬
rounded by slanting spines (Fig. 1-F).

Sensilla styloconia (St) were present on the distal margin of
most flagellar segments of all species but occasionally were ab¬
sent from some segments (Fig. 1-C). The elevation of the cuticle
that ends in these sensory cones varied among species (Table 1).
In general, only one sensilla styloconica was present on a flagel¬
lar segment but there were exceptions. Females of T. bonifatella
often had a flagellar segment with a single sensilla followed by
a segment with two sensilla. The tip of the terminal segment
had what appeared to be a modification of sensilla styloconica
that appeared in clusters of 2-4 sensilla (Fig. 1-D) and were
probably specialized for close range perception of stimuli.

DISCUSSION

The species investigated had the types of sensilla found com¬
monly on the antennae of many Lepidoptera. However, the
trichoid sensillum was dominant which suggests that the primary
function of the antennae is to detect semiochemicals. Since the
trichoid sensilla of male Lepidoptera are known to respond to
the sex attractant of females (Schneider, 1962), they probably
have a similar function in the Crambinae. The sensilla coelo¬
conica are olfactory in some insects but in Lepidoptera they
also respond to warm humid air (Schneider and Steinbrecht,

16(4) -.201-207, 1977

ANTENNAL SENSILLA

205

Fig. 1. — A. C. topiaria $ , 700x Sensilla trichodea (T-I); Sensilla trichodea
(T-II). B. C. topiaria $ , 3000x (note smooth surface texturing of sensilla).

C. C. topiaria 9, lOOOx Sensilla chaetica (Ch), Sensilla styloconica (St).

D. T. bonifatella 9 , 5000x; cluster of sensory cones on distal end of term¬
inal segment.

206

J. A. KAMM

). Res. Lepid.

E. C. topiaria $ , 700x Sensilla coeloconica (Co). F. C. topiaria
15,000x; grooved peg of sensil coeloconica.

16(4):201-207, 1977

ANTENNAL SENSILLA

207

1968), which would be useful to the insect in identifying ovipo-
sition sites. The sensilla chaetica are known mechanoreceptors
(Schneider, 1964) and are well represented in the Crambinae.
Since these adults explore the leaves and culms of grasses by
walking, these sensilla are probably essential for orientation
within the grass canopy. Females, which have more sensilla
chaetica, probably rely more on these receptors than males since
flight is difficult and impractical for females heavily laden with
eggs.

Relative to many Lepidoptera, the Crambinae have fewer
sensilla and some types are conspicuously absent. For example,
the auricillicum sensilla (shoe horn sensilla) probably provide
directional perception of stimuli as the insect orients to the
source but these sensilla were absent in the study species. The
Crambinae also lacked the various types of surface texturing on
olfactory sensilla that may permit the insect to distinguish
several types of odors with a single sensilla (Callahan, 1975).
Thus, the sensilla of the study species were simpler and had a
more functional design that those of many Lepidoptera. How¬
ever, they are well adapted to orientation within the limited
habitat of the grass canopy that adult Crambinae explore.

LITERATURE CITED

BANERJEE, A. C. and G. C. DECKER. 1966. Studies on sod webworms. I.
Emergence rhythm, mating and oviposition behavior under natural
conditions. J. Econ. Entomol. 59: 1237-44.

CALLAHAN, P. S. 1975. Insect antennae with special reference to the
mechanisms of scent detection and the evolution of the sensilla. Int.
J. Morphol. Embryol. 4: 381-430.

JEFFERSON, R. N., R. E. RUBIN, S. U. McFARLAND, and H. H.
SHOREY. 1970. Sex pheromones of noctuid moths. XXII. The external
morphology of the antennae of Trichoplusia ni, Heliothis zea, Prodenia
ornithogalli and Spodoptera exigua. Ann. Entomol. Soc. Am. 63:
1227-38.

KAMM, J. A. 1970. Effects of photoperiod and temperature on Crambus
trisectus and C. leachellus cypridalis (Lepidoptera: Crambidae). Ann.
Entomol. Soc. Am. 63: 412-6.

KAMM, J. A. 1974. Emergence rhythm in Chrysoteuchia topiaria. Environ.
Entomol. 3: 380-2.

SCHNEIDER, D. 1962. Electrophysiological investigation on the olfactory
specificity of several attacking substances in different species of
moths. J. Insect Physiol. 8: 15-30.

- , 1964. Insect antennae. Annu. Rev. Entomol. 9: 103-22.

SCHNEIDER, D. and R. A. STEINBRECHT. 1968. Checklist of insect
olfactory sensilla. Symp. Zool. Soc. Lond. No. 23, 279-97.

Journal of Research on the Lepidoptera

16(4) :208, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

INFORMATION ON
AVAILABILITY OF HOLOTYPES
OF THE DESCRIBED

TAXONS AT A PUBLIC INSTITUTION ( RHOPALOCERA)

DALIBOR WEISS

Bruselskd 3, 120 00 Praha 2, Czechoslovakia

I Would Like To Announce that holotypes of the described
geographical races Clossiana angarensis sedychi Weiss (1964),
Colias cocandica sidonia Weiss (1968) and Parnassius evers-
manni magadana Weiss (1970) have been transfered from my
private collection to the National Museum collections in Prague
in order to comply with the stipulation of the International
Commission for Zoological Nomenclature that new taxons are
invalid in the case that they are not deposited in a public insti¬
tution.

LITERATURE CITED

WEISS, D. 1964. Eine neu Unterart der Clossiana angarensis Ersch. aus
der arktischen Region der europaischen UdSSR, Reichenbachia 4:
15-17.

- . 1968. Beitrag zur Kenntnis der Tagfalter-Fauna der Mongolei,

Acta. Faun. Ent. Mus. Nat. Frag 13: 109-118.

- . 1970. A New Parnassius eversmanni Race from Nord-East Si¬
beria (USSR) ( Papilionidae ) , J. Res. Lepid. 9: 215-216.

208

Journal of Research on the Lepidoptera

16(4) :209-211, 1977

Santa Barbara Museum of Natural History
2559 Puesta Del Sol Road

GYNANDROMORPHIC POLITES SKIPPERS

( HESPERIIDAE )

M. C. NIELSEN

3415 Overlea Drive, Lansing, Michigan 48917

During the past 30 years of field collecting Michigan lepi¬
doptera, the family Hesperiidae has been one of my favorites.
Therefore, it was unique that during this period only two gynan-
dromorph Hesperiinae skippers were collected: Polites mystic
(Scudder), Otsego County, 17 June 1962, and P. origines (Fa-
bricius), Monroe County, 1 July 1965. The colored plates reveal
most of the unusual characteristics; however, the following com¬
ments will further describe the two specimens.

Polites mystic

The female pattern on the left side of the forewing is slightly
atypical. The costal area, extending from the basal area to just
beyond the apical spots, is tawny, instead of the predominantly
fuscous on typical females. Otherwise, the female portion is
within the range of normal patterns found in this locality.

The male right side of the forewing is slightly deformed with
a pronounced curve in the basal half of the costal edge, and a
more rounded anal angle. There is a slight interruption of the
tawny costal area just basal of the apical spots. The hindwings,
dorsally, are not noticeably different as compared to normal
specimens; nor are there appreciable differences on the ventral
surfaces. Dissection of the genitalia revealed that it is a female.

Polites origines

At first glance, in most respects the specimen appears to be
a male. The left dorsal surface of the forewing exhibits more
fuscous scaling than the right side, especially pronounced in the
costal area. Also, the left stigma is broken (best seen under
magnification) at the CU2 vein, and the stigmal patch is re¬
duced in area. There are no differences on the hindwing dorsal
surfaces, nor on the ventral surfaces. It was the thick, heavy
abdomen that called my attention to the specimen in the field.
Dissection of the genitalia has since indicated that it is a
female, filled with ova, with some abnormal development of
the post-vaginalis as compared to a normal female.

210

M. C. NIELSEN

)■ Res. I epid.

Polites origines. Top— Gynandromorph, Monroe Co., Mich., 1 July 1965;
Bottom— Male, Monroe Co., 1 July 1965. (leg. M. C. Nielsen)

16(4):209-2U, 1977 GYNANDROMORPHIC POLITES

211

A review of the literature discloses that the gynandromorph
phenomenon in lepidoptera is apparently uncommon. Scudder
(1889) cited a report by Dr. Hagen of 31 gynandromorph
species, mostly European, in the Papilionidae, Pieridae, Lycaeni-
dae, Nymphalidae and Satyridae in which most showed com¬
plete bilateral distinction. He suggested that the left side is
usually female, and in a few cases, one side will have mixed
sexual differences. Ford (1945) illustrated two butterfly species,
bilateral gynandromorphs, in which the female is expressed on
the left dorsal surfaces, and two species that had traces of the
other sex. Both Emmel (1964) and Opler (1966) figured a
Colias and Lycaena, respectively, in which the female is on the
right dorsal surfaces. There have been fewer reports of gynan¬
dromorphs in the moths. Bilateral gynandromorph moths have
recently been reported by Hessel (1964), Muller (1966) and
Blanchard (1969) in which the specimens exhibit female char¬
acteristics on the right dorsal surfaces. In most of the above
references, these authors did not report the dissection of the
genitalia to confirm the sex.

It would appear that these two skippers may be the first
reported Hesperiidae gynandromorphs. Unfortunately, amateur
lepidopterists do not usually favor skippers in their field activi¬
ties, which may account of the paucity of records. As one can
see, these skipper gynandromorphs are not as striking in ap¬
pearance as compared to some of the other gynandromorphs
figured in the above references. Hopefully, other collectors will
be alerted by this experience and report their findings.

LITERATURE CITED

BLANCHARD, ANDRE. 1969. A Gynandromorph Phaeoura mexicanaria
( Geometridae ) . J. Lepid. Soc. 23(4): 274-275.

EMMEL, THOMAS C. 1964. New Gynandromorph of Colias philodice
from Colorado. /. Res. Lepid. 3(1): 63-64.

FORD, E. B. 1945. Butterflies. Collins London.

HESSEL, SIDNEY A. 1964. A Bilateral Gynandromorph of Automeris io
( Saturniidae ) Taken at Mercury Vapor Light in Connecticut. J. Lepid.
Soc. 18(1): 27-31.

MULLER, JOSEPH. 1966. Gynandromorphic Eacles imperialis. J. Lepid.
Soc. 20(2): 118.

OPLER, PAUL A. 1966. A Gynandromorph of Lycaena gorgon. J. Res.
Lepid. 5(4) : 230.

SCUDDER, SAMUEL H. 1889. The Butterflies of the Eastern United
States and Canada, with Special Reference to New England. Cam¬
bridge, Mass., pp. 1759-1760.

Polites mystic. Top— male, Otsego Co., Mich., 19 June 1964; Middle—
Gynandromorph, Otsego Co., 17 June 1962; Bottom— Female, Otsego Co.,
6 July 1958. (leg. M. C. Nielsen)

Journal of Research on the Lepidoptera

16(4) :212, 1977

SPECIAL THANKS

I should like to take this opportunity to express deep and
sincere gratitude to Karen Nielson Hovanitz, who will be con¬
tinuing as assistant editor, Rudi Mattoni, acting editor and Scott
Miller, assistant editor (elect). Without the unselfish and un¬
failing support they have given, the journal could not be con¬
tinuing.

Our plans for continuing the journal are, we believe and
hope, innovative and forward looking. We welcome comments
from our subscribers and were greatly encouraged by the large
number of spontaneous requests to continue the journal, as well
as offers to help do so in any way possible.

To the many friends who sent cards and letters expressing
sympathy to myself and family in the loss of Bill, a very heart¬
felt thank you.

Barbara Hovanitz

212

Journal of Research on the Lepidoptera

16(4) :213-221, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

STUDIES ON THE EXCRETORY SYSTEM

OF THE FULLY GROWN LARVA OF

CALOGRAMA1A FESTIVA DONOV.
(NOCTUIDE)

U. V. K. MOHAMED

Department of Zoology, University of Calicut,

Calicut — 67365 India

HUMAYUN MURAD

Department of Zoology, Makerere University,

Kampala, Uganda

ABSTRACT

Fully grown larvae of Calogramma festiva Donov., possesses six malpigh-
ian tubules, three on either side of the gut arising from a common duct.
The proximal portion remains suspended in the haemolymph as a free
tubule while the distal convoluted portion of the nephric tubules in as¬
sociation with the rectum form the nephro-rectal complex. On the basis
of histological details three regions are distinguishable in the free tubule
whereas the nephric tubule is divisible into proximal and distal regions.
The nephro-rectal complex is formed of nephric tubules and components
from the gut wall. The pronephric epithelium and the longitudinal muscles
form a cover for the nephric tubules.

INTRODUCTION

Very little is known about the excretory system of lepidop-
terous larvae. Ishimori (1924) has described the distribution
of malpighian tubules of certain larval form of Lepidoptera.
Except for the study of Mathur (1966) in noctuid caterpil¬
lars, Prodenia litura and Trichoplusia ni, no attempt has been
made to investigate the structural details of malpighian tubules
of lepidopterous larvae. Studies on the development of tubules
in Pieris hrassicae (Henson, 1932), Venessa urticae (Hen¬
son, 1937) and Philosamia ricini (Srivastava and Khare,
1966) give only inadequate informations on the histology of the
malpighian tubules. The present work therefore, has been under¬
taken to describe the anatomical details of the excretory system
of fully grown larva of Calogramma festiva D.

213

214

U. V. K. MOHAMED and H. MURAD

/. Res. Lepid.

MATERIALS AND METHODS

Adults were collected from the field and reared in the labora¬
tory. The fully grown larvae were anaesthesised with ether
vapour and dissected in Ringer’s solution under binocular mi¬
croscope. For histological purposes, malpighian tubules were
fixed in Bouin’s fluid. Sections were obtained at 5-6 jj, and stain¬
ing was done with haematoxylin and eosin.

RESULTS

The excretory system of the larva of Calogramma festiva D.,
is in the form of six malpighian tubules, three on either sides
of the gut. The three tubules on each side actually arise from a
common duct (Fig. 1). Major portion of each tubule remains
suspended in the haemolymph as free tubule while the distal
portion called nephric tubule in association with the rectum
forms a nephro-rectal complex.

A. Free Tubule

Immediately after its origin from the common duct each tubule
runs towards anterior side of the gut as ascending limb (AMal)
and on reaching the stomodaeum, it turns backwards and runs
posteriorly as descending limb (DMal). The ascending limb of
the malpighian tubule is transluscent whereas, the two thirds
of the length of descending limb is dull white in colour. The
remaining portion of the descending limb is characterised by the
presence of small diverticulae on the wall of the tubule.

On the basis of histological details, three regions are distin¬
guished in the free portion of malpighian tubule. In the first
region (Fig. 2) peritoneal layer (PI) is not distinct and the
basement membrane (BMb) holds a ring of epithelial cells. The
infoldings of the cell membrane (FcMb) in the basal zone of
the cell are loosely packed and are fewer in number. In the
central zone, cytoplasm is coarsely fine and contains elongated
nucleus (N) measuring 5-7 ^ x 2 |x, but in certain cells more
elongated nuclei of 26-29 ji in size are observed. Border zone is
formed of short filaments (sb) which are free towards the lu¬
men. The second region (Fig. 3) is distinct from the first region
in that the infoldings of the cell membrane (FcMb) are thickly
packed in the basal zone of the cell and the reticulate cytoplasm
in the central zone lodges variously shaped nucleus (N). The
nucleus may be dumb-bell shaped, branched or semilunar. The
cytoplasm presents scattered granules around nucleus. Border
zone is formed of elogated free filaments (sb). An outer peri-

16(4):213-221, 1977

CALOGRAMMA FESTIVA

215

PLATE I

Fig. 1. — Diagrammatic representation of hind gut of C. f estiva to show the
arrangement of malpighian tubules.

Fig. 2. — Section passing through the first region of malpighian tubule of
C. f estiva.

ABBREVIATIONS

AInt, anterior intestine; AMal, ascending limb of malpighian tubule; ANt,
anterior portion of nephric tubule; BMb, basement membrane; CD, com¬
mon duct; DMal, descending limb of malpighian tubule; FcMb, infoldings
of the cell membrane; GR, granules; In, intima; iPE, inner pronephric
epithelium; Lum, lumen; mcl, muscle layer; MG, midgut; N, nucleus; oPE,
outer pronephric epithelium; PI, peritoneal layer; PNt, posterior portion of
nephric tubule; RE, rectal epithelium; Rect, rectum; sb, striated border;
Tra, tracheole.

PLATE II

216

U. V. K. MOHAMED and H. MURAD

/. Res. Lepid.

Fig. 3. — Section passing through the second region of malpighian tubule of
C. f estiva.

Fig. 4. — Section passing through the third region of malpighian tubule of
C. f estiva.

16(4):213-221, 1977

CALOGRAMMA FESTIVA

217

toneal layer (PI) is present. In the third region (Fig. 4) the
basal zone is highly reduced. The reticulate non-granular cyto¬
plasm contains variously shaped nucleus (N) in the central
zone. The border zone is formed of many long filaments (sb)
which are free towards the lumen ( Lum ) .

The common duct (Fig. 5) differs in histological details from
the tubule. The peritoneal layer is indistinct and a tough base¬
ment membrane holds a ring of epithelial cells whose boundaries
are not clear. The basal zone is highly reduced. The central zone
contains dense cytoplasm. Some of the nuclei (N) present in
this region are much elongated. The border zone shows elong¬
ated free filaments ( sb ) .

B. N ephro-rectal complex

The nephric tubule in association with the rectum forms the
nephro-rectal complex (Fig. 6) which includes the nephric tu¬
bule (Nt), pronephric epithelium (PE) and the muscle layer
(mcl). The pronephric epithelium takes its origin from the gut
epithelium. Immediately after its emergence from the latter, it
divides into two strata, the inner pronephric epithelium (ipE)
and outer pronephric epithelium ( opE ) both of which are form¬
ed of single layer of cells. The outer pronephric epithelium is
very thin and weakly developed. The inner pronephric epithelium
is well developed and formed of elongated cells. Both the layers
form a loose cover over the rectal wall. Due to the provision of
two epithelial layers the pronephric chamber is divided into an
inner and outer chamber. Each nephric tubule after its entry
into the outer chamber runs upto the posterior limit of the rec¬
tum. Thereafter it pierces the inner pronephric epithelium and
enters into the inner chamber. The nephric tubule within the
inner chamber takes an anteriorly directed course upto the mid¬
dle of the rectum where each tubule ends blindly.

The proximal portion of the nephric tubule (ANt) lying with¬
in the outer chamber is very narrow having very thin wall.
Peritoneal layer and the basal zone are indistinct. The dense
cytoplasm of the central zone lodges the nucleus. The border
zone is highly reduced. In the distal portion of the nephric tu¬
bule (PNt) the basal zone is formed of loosely packed infoldings
of the cell membrane (FcMb). Oval nucleus is found in the
reticulate cytoplasm of the central zone. The border zone is
formed of long filaments ( sb ) which are free at their distal ends.
A longitudinal muscle layer taking its origin from the gut wall
surrounds the outer pronephric epithelium.

218

U. V. K. MOHAMED and H. MURAD

J. Res. Lepid.

PL ATE III

Fig. 5. — Section passing through the common duct of malpighian tubule of
C. f estiva.

Fig. 6. — Longitudinal section passing through the nephro-rectal complex of
C. f estiva.

16(4):213-221, 1977

CALOGRAMMA FESTIVA

219

DISCUSSION

In Lepidoptera there are six tubules in both larvae and adults
(Henson, 1931; Mathur, 1966) with perhaps only exception
of Tiniedae in which the six typical tubules of the larvae are
reduced to only one pair in the adults (Snodgrass, 1935).

The malpighian tubules of insects are invested by a peri¬
toneal coat having abundant supply of tracheoles (Wiggles¬
worth, 1965). The tubules of P. litura and T. ni (Mathur,
1966) are surrounded by a peritoneal layer of very thin squam¬
ous cells with widely separated nuclei. Since there are significant
variations in its structure at various levels of the tubules, it
seems that these cells perhaps play a role in the tubule physi¬
ology. The nuclei of malpighian tubule differ in size, location
and shape not only in various insects but also in different regions
of the tubule of the same insects (Mohamed, 1974). Ber-
theau (1963) distinguishes five regions in the anterior part
and seven regions in the posterior part of the tubule of Carausius
morosus based on the relation between nucleus and cytoplasm.
But the well marked differences between the nuclei of upper
and lower segments of Rhodnius observed so clearly under light
microscope ( Wigglesworth, 1931) have not been reported
in electron microscopic study (Wigglesworth and Salpeter,
1962). The size of the nucleus varies greatly in different
regions of the tubule of certain aquatic bugs and many ter¬
restrial plant feeding bugs (Bahadur, 1961; 1964). In For-
ficula, the diameter of the nuclei of the tubule at certain ecdysis
become almost doubled possibly because of incomplete mitosis
(Henson, 1946). The increase in the size of nucleus in Pieris
brassicae (Henson, 1932) is accompanied by lobulation and
ramification as well. So it seems evident that variously shaped
elongated nuclei of C. f estiva are either due to difference in
increase of size or incomplete mitosis.

Ishimori (1924) has described the arrangement of nephric
tubules in many lepidopterous larvae. The arrangement of the
tubules in larval C. festiva is similar to that described by Hen¬
son (1931) in V. urticae except that instead of loops a number
of convolutions are formed. In C. festiva the distal portion of
the nephric tubule which is lodged in the inner chamber of the
nephro-rectal complex extends only upto the middle of the latter
as it is in the case of Bombyx, Gastropacha and Spilosoma
(SAINI, 1964).

220

U. V. K. MOHAMED and H. MURAD

J. Res. Lepid.

Ishimori (1924) described the ‘cryptonephric envelope’ of
lepidopterous larvae as composed of three layers, the outer
muscle layer, middle epithelial layer and inner membranous
layer. The present observations agree with that of Ishimori
(1924) in the number of layers but differ in the nature of
pronephric epithelial layers. In the larval Calo gramma the outer
pronephric epithelial layer is very thin and weakly developed
whereas, the inner pronephric epithelial layer is thick and is
formed of elongated cells. It also extends firm support to the
view of Srivastava and Khare (1966) that the two layers of the
cells of the cryptonephric envelope originate from the epithelial
layer of the gut wall.

ACKNOWLEDGEMENTS

The present work has been carried out at the Aligarh Muslim
University, India and the authors are grateful to the Head of
Zoology Department for providing necessary laboratory fa¬
cilities. One of us (U.V.K.M.) is thankful to Prof. K. J. Joseph of
the University of Calicut for enabling him to take up this work
at Aligarh.

LITERATURE CITED

BAHADUR, J. 1961. The malpighian tubules of certain aquatic Heterop-
tera. Ann. Mag. Nat. Hist. Lond., 4: 431-440.

- , 1964. The malpighian tubules of certain terrestrial plant feeding

bugs. Zool. Anz. Lepiz., 175: 387-395.

BERTHEAU, P. H. 1963. Evolution of mesenteron and malpighian tubules
in Carausius morosus. Bull. De La Soc. Zool. de France, 12: 496-512.
HENSON, H. 1931. The structure and postembryonic development of
Vanessa urticae (Lep.). 1. The larval alimentary canal. Quart. J. mic.
Sci., 75: 283-305.

- , 1932. The development of alimentary canal in Pieris brassicae and

the endodermal origin of malpighian tubules. Quart. J. mic. Sci., 75:
283-305.

- , 1937. The structure and postembryonic development of Vanessa

urticae (Lep.). II. The larval malpighian tubules. Proc. Zool. Soc.
Lond. (B), 107: 161-174.

- , 1946. On the malpighian tubules of Forficula auricularia. Proc. Roy.

ent. Soc. Lond. (A), 23: 29-39.

ISHIMORI, N. 1924. Distribution of the malpighian vessels in the wall of
the rectum of the lepidopterous larvae. Ann. Ent. Soc. Amer., 17:
75-86.

MATHUR, L. M. L. 1966. Morphology of the excretory system of two
noctuid caterpillars, Prodenia litura Fabr., and Trichoplusia ni Hubner.
Ind. J. ent., 28: 61-66.

MOHAMED, U. V. K. 1974. Studies on the malpighian tubules of insects.

Ph.D. Thesis Alig. Musi. Univ. Aligarh.

SAINI, R. S. 1964. Histology and physiology of the cryptonephridial system
of insects. Trans. R. ent. Soc. Lond., 116: 347-392.

SNODGRASS, R. E. 1935. Principles of insect morphology. McGraw-Hill
Book Co., New York.

16(4):213-221, 1977

CALOGRAMMA FESTIVA

221

SRIVASTAVA, U. S. & KHARE, M. K. 1966. The development of mal-
pighian tubules and associated structures in Philosamia ricini. J. Zool.
Lond., 150: 145-163.

WIGGLESWORTH, V. B. 1931. The physiology of excretion in the blood
sucking insect Rhodnius prolixus. J. Exp. Biol., 8: 411-451.

- , 1965. Principles of insect physiology. 6th edition, Methuen, London.

WIGGLESWORTH, V. B. & SALPETER, M. M. 1962. Histology of the
malpighian tubules in Rhodnius prolixus. J. Insect Physiol., 8: 299-307.

Journal of Research on the Lepidoptera

16(4) :222, 1977

do Santa Barbara Museum, of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

RECTIFICATION OF A RECENT PAPER ON

LEUCANELLA MEMUSAE GARDINER I1

RICHARD S. PEIGLER

Department of Entomology, Texas A&M University
College Station, Texas 77843

In Vol. 15 Of This Journal, Gardiner (1976) has described
and figured the early stages of Leucanella memusae gardineri
Lemaire and compared these to what he said is the nominate sub¬
species, L. memusae memusae (Walker). However, Gardiner’s
figures of what he says represent the nominate subspecies are
actually of L. viridescens (Walker.) The two species have long
been confused in the literature. Because the source locality of the
L. viridescens stock is not given, I am unable to say which sub¬
species of L. viridescens it is. The aforementioned remarks re¬
sult from reference to an excellent and recent revision by Le¬
maire (1973).

It is stated in the original description of L. m. gardineri that
the type series consisted of wild specimens supplied to Lemaire
by F. Plaumann of Brazil. Gardiner states that the type series was
reared by himself in England.

Gardiner’s paper is valuable for confirming that L. memusae
and L. viridescens are not conspecific by showing differences be¬
tween the larvae.

LITERATURE CITED

GARDINER, B. O. C. 1976. The early stages of Leucanella memusae ssp.
gardineri Lemaire ( Saturniidae ) . J. Res. Lepid. 15: 201-205.

LEMAIRE, C. 1973. Revision du genre Automeris Huebner et des genres
voisins, biogeographie, ethologie, morphologie, taxonomie (Lep. At-
tacidae). Mem. Mus. Natl. Hist, natur., (Serie A), 79:233-422, pis.
30-49.

1Attacidae=Saturniidae

222

Journal of Research on the Lepidoptera

16(4) : 223-242, 1977

c/o Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

THE SCOLITANTIDINI
I: TWO NEW GENERA AND A GENERIC
REARRANGEMENT. ( LYCAEN1DAE )

RUDOLF H. T. MATTONI

Agri Science Laboratory, Los Angeles, California 90025

Although i have been working with “Philotes” for over
thirty years, it was apparent after the first few months of casual
study that the then available arrangements of circumscribed
species was incorrect. At the time, the McDunnough (1938) list
of North American species included Philotes battoides ( Behr )
1867, P. glaucon (Edw. ) 1871, P. enoptes (Bois.) 1852, P. mojave
(Watson & Comstock) 1920, P. spaldingi B. & McD. 1917, P.
rita B. & McD. 1916, P. speciosa (Henry Edwards) 1876, P.
sonorensis F. & F. 1865. Lycaena regia, Bois. (1869), a synonym
of P. sonorensis, was the type species Scudder (1876) designated
in erecting the genus. The arrangement of species concepts has
been maintained by most recent workers, incorporating the
recognition of glaucon as a subspecies of battoides following my
diagnosis (1945).

Based upon criteria of facies, habit, and male genitalia, P.
sonorensis appeared widely separated from P. speciosa, with
both quite clearly differentiated from the remaining group of
battoides, enoptes, mojave, spaldingi, and rita. In 1945, I mis¬
takenly believed the above group of species were closely related
to the Palearctic vicrama, baton, abencerragus group on the
basis of my incomplete knowledge of the work of Hemming
(1929). This led me to also suggest they were in Turanana, a
massive error (Mattoni, 1954b). Hemming (1932), unknown
to me, had placed the group in Philotes, where they remained
through Forsters (1938) revision until the work of Henry Beuret.
Beuret ( 1958, 1959 ) illuminated several relationships by examin¬
ing most members of the tribe using a taxonomy based on male
genitalia and androconia. He erected the new genus Pseudophi-
lotes, type species baton ( Bergstrasser ) 1779, to include vicrama
(Moore) 1865, abencerragus (Pierret) 1837, and bavius (Evers-

223

224

R. H. T. MATTONI

J. Res. Lepid.

mann) 1832. He also erected Shijimiaeoides , type species divina
(Fixsen) 1887, which included the Tibetan species lanty (Ober-
thur) 1886 and North American enoptes.

Forster (1940) had in the meantime (and unknown to Beuret
in 1958) corrected the earlier misidentification of Shijimia moorei
(leech) ( Everini ) with the new entity Sinia leechi, Forster,
1940 and included lanty and divina in Sinia.

The Beuret diagnosis of enoptes in Shijimiaeoides was un¬
noticed until Shields commenced his comprehensive study on the
biology, distribution and variation of North American Philotes.
Shields (1974) noted Beurets conclusion and applied Shijimia¬
eoides to the North American enoptes and rita, later including
battoides (Shields, 1975).

Heretofore I have hesitated to erect new genera without
exploring possible relationships within the whole tribe Scoli-
tantidini Tutt 1909 (syn. Glaacopsychini Hemming 1932, see
Beuret 1959). Sufficient morphological data are available which
indicate the propriety of doing so now.

Although S. divina shares certain morphological features with
enoptes et al., and may in fact be the closest relative to the North
American group, I believe it quite distinct.

EUPHILOTES MATTONI, NEW GENUS

Type Species: Lycaena enoptes Boisduval 1852, Ann. Ent. Soc.

France 2 ( 10) :298
Adult (Table 1, Plates 1, 2, 5, 9, 12)

Head. Antenna 28 segments in male, 30 in female; Club 14-15
segments, nudum 12. Eyes smooth, round in frontal aspect.
Frontal ratio eyes to head coverage 69%. Labial pulps erect,
densely covered with white scales dorsally and caudally, black
hairscales ventrally. Ratio of segments 1 : 3.4 : 1.1, third segment
with a terminal notch about half the length.

Wing shape and venation commonplace for the tribe. Vena¬
tion of forewing radial vein system ratio unique ( rf . Plate 10 ) :
A/B 0.75, B/C 1.50.

Males with dense androconial scales with average length of
.210 mm, width of .149 mm (Ratio 1.4) with 15 striae. Flavone
present as the underside ground pigment.

Legs robust L/W Ratio 4.2, pro- and mesothoracic legs with
large tibial process and two pairs of short, fine calcaria in both
sexes. Ungual edont lobes absent.

16(4):223-242, 1977

THE SCOLIT ANTIDINI

225

Male Genitalia. Tegumen heavy, vinculum light in lateral
aspect, relatively compressed cephalo-caudally. Saccus very
long. Labides not produced, flat, somewhat broad in caudal as¬
pect (L/W Ratio 2.0). Falces short, relatively fine, tapered, not
touching. Aedeagus simple with well developed pair of basal
lopes, seminal duct entering dorso-cephalad. Zone almost term¬
inal. Cornuti spines short, coarse, numerous, borne on a single
dorsal plate.

Female Genitalia. Papillae anales subtriangular. Posterior
aeophyses short, straight, not tapered, attached to papillae anales
subdorsally. Lodix large subcuboid formed as a single apparent
sclerite. Deeply invaginated into the seventh segment. Corpus
bursa simple, elongate, without cervix or signa.

DIAGNOSIS: Euphilotes can be differentiated at once from all
numbers of the tribe by the single character of venation of R
(Table 1). The combination of robust legs, large tibial process,
paired calcaria, and antennal segment number of both sexes;
and single cornutus plate, absence of Chapmans process on the
aedeagus, and short falces in the male are also of generic value.

The conformation of the valves in the male genitalia and
lodix and ovipositor in the female, though of highly significant
taxonomic value, appear for the most part limited to specific
differentiation. In our projected study these and other internal
characters will be subject to multivariate analysis to more ex¬
actly define phenetic relationships.

The specific sets of characters which serve to define Euphi¬
lotes and differentiate it from related genera are given in Table
I and the cited plates. These data compare E. enoptes and its
congeners which have been examined, to all other genera in the
“red maculed” section of the tribe. In several cases it should be
recognized that the evaluations are based on single specimens.
Accordingly further study may show several individual aberrant
traits. The trends, nevertheless, are fairly clear.

Euphilotes is confined to western North America and includes
the species enoptes , rita, mojave, spaldingi, and hattoides. Con¬
trary to the diagnosis of Shields (1975) there is adequate evi¬
dence regarding E. spaldingi as a distinct species on morpho¬
logical and biological grounds and E. mojave on the basis of
several instances of sympatry and cryptic morphological differ¬
entiation. These points will be expanded in a forthcoming paper.
Pending outcome of a long term project Shields and I are com-

226

R. H. T. MATTONI

J. Res. Lepid.

mencing, more complex relationships may be revealed in the
future.

The assignment of other related species in this section is
fairly straightforward. Both Philotes and Scolitantides are mono-
specific. Pseudophilotes as a homogeneous set of species clearly
defined by the produced labides of the male genitalia. The genus
includes P. baton, P. vicrama, P. abencerragus a'nd the newly
described P. sinaicus Nakamura 1974 plus other possible in¬
cluded cryptic species as panoptes. ( Hbn. ) 1808. I have placed
bavins in Sinia on the basis of many shared morphological traits
in spite of a somewhat divergent facies (rf. Figs. 2 & 3).
Higgins ( 1975 ) clearly perceived bavins did not belong with
Pseudophilotes, placing it in Scolitantides. Shijimiaeoides lanty
has been retained with S. divina. Although there are several im¬
portant divergences, both far eastern species appear more closely
related than to any others. Based upon limited data S. divina
is close to Maculinea and may exemplify the link of the “red
macule’’ species to the Glaucopsyche section.

The relationships of speciosa are not well defined. Although
it is not close to P. sonorensis, it is also not congeneric with
Euphilotes. Shields (1974) recognized this fact and placed
speciosa in Zizeeria (Zizeerini) based upon features of the male
genitalia and early stages. Although the S colit antidini and Zi¬
zeerini appear phenetically close, speciosa clearly belongs in
the former tribe by virtue of gross structure of genitalia, legs,
androconia, and probably other factors.

Since Forster (1938) erected Praephilotes for anthracias and
Paleophilotes for triphysina, claiming these as primitive forms, it
appeared fruitful to include the duo in an analysis as possible
relatives of speciosa. P. arthracias is a common butterfly in the
Karakum Desert, P. triphysina a virtually unknown inhabitant
of the Sikiang steppes. Neither appears close to speciosa, nor,
based on the data given, to any other members of the group,
including one another. P. triphysina is particularly bizarre (Table
I and Figures) with regard to venation; male genitalia with the
laterally bent aedeagus and deeply involuted ductus entry, strong¬
ly produced labides, heavy long bent tegumen, a complex bi¬
furcate juxta, and a definative sclerite surrounding the anal
opening. In particulars of the male genitalia there are some sug¬
gestions of relationship to Pseudophilotes. The facies of both
P. triphysina and P. anthracias are decidedly Zizeerine. P. an¬
thracias otherwise appears closer to the Scolitantidid groups. I
have concluded accordingly, that speciosa requires a new genus.

16(4):223-242, 1977

THE SCOL1T ANTIDINI

227

PHILOTIELLA MATTONI, NEW GENUS

Type Species: Lycaena speciosa Henry Edwards, 1876 Proc. Cal.

Acad. Sci. 7:173

Adult (Table 1, Plates 3, 4, 5, 10, 12)

Head; antenna 29 segments in male, 30 in female; Club 13
segments, nudum 10. Eyes smooth, round in frontal aspect.
Frontal ratio of eyes to head coverage 64%. Labial palps erect,
densely covered with hairscales ventrally and scales dorsally.
Both scale types white basally and black proximally. Ratio of
segments 1 : 2.5 : 0.8, third with small terminal notch.

Forewing. Shape commonplace Scolitantiidid but slightly
more elongate (L/W 1.85 vs 1.75 for. Enphilotes enoptes).
Venation commonplace, except branch of R4 + 5 from R3 more
distad than any other genera in the group excepting P. triphy-
sina.

Hindwing. Shape and venation commonplace for the tribe.

Males possess abundant androconial scales with an average
length of .178 mm, width of .126 mm and 13 striae, ratio 1.4).
Flavones present as underside ground pigment.

Legs unique for the tribe, femur more robust than any of the
tribe with L/W ratio 3.0. Proleg with very large tibial process
and one pair of short fine calcaria in both sexes. Tibial processes
absent on meso- and metathoracic legs. Ungual edont lobes
absent.

Male genitalia unique for the tribe. Tegumen and vinculum
heavy in lateral aspect, saccus absent, labides not produced, flat,
suboval in caudal aspect (L/W Ratio 1.3). Falces short, heavy
except distally terminating in fine apical processes bending pos¬
tered and ventrad and parallel with one another. Juxta straight
in lateral aspect, relatively short. Aedeagus unique in possessing
a quadrilobed base. Seminal duct entering dorso-cephalad. Zone
almost terminal. Vesica slender bearing few small blunt, coarse
cornuti spines.

Female genitalia. Papillae anales subquadrate with the mem¬
branous setal caudal portion almost exactly bisecting the quad¬
rate form. Posterior apophyses short, tapered and bent, jointing
the papillae anales subdorsally. Lodix subcylindrilar in caudal
view, formed of two sclerites, the vental appearing the smaller

228

R. H. T. MATTONI

/. Res. Lepid.

PLATE 1

Uppersides. Row 1 (Males) and 2 (Females) Left to Right. Shijimaeoides
divina asonis, Mt. Aso, Kyushu, 24 May, 1970, leg. Setoya and Takagi;
Shijimiaeoides lanty lanty, Tatsienlu, Szechwan, Cheto-Wahui P. 10 May
1930 (ZSB), Sinia bavins bavins, Male, Kohu Mts., Anatolia, 5 May, 1976,
leg. Gillet; Row 3 (Males) and 4 (Females). Sinia leechi, Sunpanting,
Szetchwan, no date, Stotzner Exp. (ZSB); Scolitantiides orion, Srbeko,
Bohemia, 7 May, 1967, leg. Vanek; Pseudophilotes vicrama astarene, Djrezh
River, Yerevan, Armenia SSR, 1000 M, 4 June, 1972, leg. Weidenhoffer;
Euphilotes enoptes enoptes, Male, Tioga Pass, Mono Co., Calif., 14 July,
1958, leg. Shields, Female west above Tioga Pass, Mono Co., Calif., 27
July, 1960, leg. Shields.

PLATE 2

Undersides of PLATE 1.

16(4):223-242, 1977

THE SCOL1T ANTIDINI

229

PLATE 3

Uppersides. Top Row, Philotiella speciosa speciosa, Male, Randsburgh,
Calif., 4 May, 1954, leg. R. Ford; Female, Mojave Desert, Calif., 12 May,
1924, leg. J. A. Comstock; Philotiella speciosa bohartorum , Briceburg, Calif.,
10 May, 1969, leg. J. Lane. Second Row, Paleophilotes triphysina, Male and
Female, Altyn Tag, China, Turkestan, no date (ZSB). Third Row, Praephi-
lotes anthracias, both males, Repetek, Transcaspian, 4 April, 1966 (ZSB);
Kuldaha, Thien-Shan, 1885 (ZSB).

PLATE 4

Undersides of PLATE 3.

230

R. H. T. MATTONI

J. Res. Lepid.

of the two. The lodix is only slightly invaginated into the seventh
segment. Corpus bursa simple, elongate, without cervix or signa.
DIAGNOSIS: Philotiella can be distinguished from other closely
related genera by the single character of venation of R (Table I).
The extremely heavy proleg femur is also unique. Although the
ratio is given for the male (Table I) the female is virtually
identical. The large tibial process of the prothoracic legs in con¬
trast to a small process on the mesothoracic legs is also unique.
The aedegus likewise exhibits characters found only in this
genus: the relative overall length and the four lobed base.

The pattern of variation in the S colit antidini is complex,
probably because of an evolutionary history with both substantial
radiation and convergence. Monophyletic (monophenetic) lines
are difficult to recognize at this time. With data on more char¬
acters, including early stages, we hope to perform multivariate
analysis in order to quantitate possible phenetic distances.

It is intriguing to speculate that the center of origins of the
tribe lies in the vicinity of the North Eastern Tibetan plateau.
In the area of western Szechwan the following genera are repre¬
sented: Glaucopsyche , Sinia, Shijimiaeoides , Maculinea, Phen-
garis, Caerulia and Scolitantides. Within a distance of 2000 km
to the west and north, all remaining Palearctic genera are to be
found: Paleophilotes , Praepliilotes, Pseudophilotes, Turanana,
and lolana. Included with the latter group is Lycaena panope ,
Eversmann 1851, which cannot be generically assigned at this
time; and a new genus and species from the Koh-I-Baba Moun¬
tains of Afghanistan which appears close to Turanana. Pseudo¬
philotes and Euphilotes appear to represent a case of converg¬
ence in facies following adaptive responses towards the Mediter¬
ranean west and east into North America. Both genera appear
actively evolving in adapting to the conditions of their respective
habitats to this day. Such a situation would account for the
taxonomic dynamism of the genera at this point in the biological
history of the earth.

ACKNOWLEDGEMENTS

My special thanks to Drs. Walter Forster and Wolfgang
Dierl of the Zoologische Sammlung Des Bayerischen Staates
(ZSB) for the loan of key specimens, to Mssrs. Boris Izenbek of
Akmene, Lithuania and J. C. Weiss of Hagondage, France for
generously providing much other Eurasian Material, to Dr.

16(4):223-242, 1977

THE SCOL1T ANTIDIN1

231

Takashi Shirozu of Kyushu University, Japan for donating a
beautiful series of S. divina, to John Lane, University of Cali¬
fornia, Davis, for providing the P. speciosa bohartorum speci¬
men used, to John Sorensen for providing the manuscript of his
invaluable morphological study, to Julian Donahue of the Los
Angeles Museum of Natural History by always making their
collection and library completely available, and last but not
least, to my colleague Oakley Shields for the many discussions
on this group we are so deeply committed to study.

Location of Figured Material: (ZSB) from the Zoologische

Sammlung Des Bayerischen Staates, Munich, Germany. All
others from the Los Angeles Museum of Natural History.
Prepared material will accompany their specimens.

LITERATURE CITED

BEURET, H. 1958. Zur Systematischen Stellung Einiger Wenig Bekannter
Glaucopsychidi (Lep. Lycaenidae). Mitt. Ent. Ges. Basel 8:61-79,
81-100.

- . 1959. Zur Taxonomic Einiger Palearktischer Blaulinge ( Lep. Lyca¬
enidae). Mitt. Ent. Ges. Basel 9:80-84.

FORSTER, W. 1938. Das System der Palearktischen Polyommatini ( Lep.
Lycaenidae). Mitt. Munch. Ent. Ges. 28:97-118.

- . 1940. Neue Lycaeniden-Formen aus China I. Mitt. Munch. Ent.

Ges. 30:870-883.

FORD, E. B. 1941. Studies on the Chemistry of Pigments in the Lepi-
doptera, with Reference to their Bearing on Systematics. ( 1 ) The
Anthoxanthins. Proc. Roy. Ent. Soc. (A) 16:65-90.

HEMMING, F. 1929. Revision of the haton-G roup of the Genus, Turanana
Beth.-Bak., with an Account of an Unrecognized Species, Turanana
vicrama Moore. Entomologist 62:27-34, 60-64, 84-89.

HEMMING, A. F. 1932. The Buterflies of Transjordan. Trans. Ent. Soc.
Land 80:269-299.

HIGGINS, L. G. 1975. The Classification of European Butterflies. Collins,
London.

MATTONI, R. H. T. 1954a. Notes on the Genus, Philotes I. Bull. So.
Calif. Acad. Sci. 53:157-165.

- . 1954b. (Abst. ) Taxonomy and Distribution in the Genus, Philotes.

Lep. News 8:8.

McDUNNOUGH, J. H. 1938. Check List of the Lepidoptera of Canada
and the United States of America. Part 1. Macrolepidoptera. Mem.
So. Cal. Acad. Sci. 1:272.

NAKAMURA, I. 1975. Descriptions of Two New Species of Butterflies
(Lepidoptera, Lycaenidae) from the South Sinai. J. Ent. (B) 44(3)

: 283-295.

SHIELDS, O. 1974. Studies on North American Philotes (Lycaenidae)
III. Generic Reassignments and the Biology of speciosa. Bull. Allyn
Mus. No. 19:10pp.

- . 1975. Studies on the North American Philotes (Lycaenidae) IV.

Taxonomic and Biological Notes, and New Subspecies. Bull. Allyn
Mus. No. 28:36pp.

SORENSON, J. T. 1972. The Integumental Morphology of Adult Glau-
copsycini (Lepidoptera:Lycaenidae). Masters Thesis. University of
North Iowa.

232

R. H. T. MATTONI

J. Res. Lepid.

SHIJIMIAEOIDES DIVINA

EUPHILOTES ENOPTES

PHI LOTI ELLA SPECIOSA
0.5 MM

PRAEPHILOTES ANTHRACIAS

PALEOPHILOTES TRIPHYSINA

PLATE 5

Labial palps of eleven species as labelled, scale indicated.

All illustrations, with exception of the venation diagram, to the
same scale. Execution using a Wild M5 Microscope with the
Wild Drawing Attachment, by R. H. T. Mattoni.

16(4):223-242, 1977

THE SCOLIT ANTIDINI

233

PLATE 6 PHIL°TES S0N0RENSIS

Male genitalia of Scolitantiides orion (above); Philotes sonorensis (below).
Upper left figures: Aedeagus, lateral view (above); dorsal view (below).
Upper right figures: entire genitalia with valves not shown, caudal view
showing relative shape or labides and falces.

Lower left figures: entire genitalia lateral view.

Lower right figures: entire genitalia with valves not shown, dorsal view,
caudally oriented to bottom of plate.

234

R. H. T. MATTONI

J. Res. Lepid.

PSEUDOPHILOTES VICRAMA

SINIA B AVI US

- 1

0.5 MM

PLATE 7

Male genitalia of Pseudophilotes vicrama (above) and Sinia bavius (below)
orientation as PLATE 6.

16(4):223-242, 1977

THE SCOUT ANTIDINI

235

SHI JIMI AEOIDES LAN7Y

PLATE 8

Male genitalia of Shijimiaeoides lanty (above) and Sinia leechi (below)
orientation as PLATE 6.

236

R. H. T. MATTONI

J. Res. Lepid.

SHI JIMIAEOIDES DIVINA

EUPHILOTES ENOPTES Q.5 mm

Male genitalia of Shijimiaeoides divina (above) and Euphilotes enoptes
( below ) .

16(4):223-242, 1977 THE SCOUT ANTID1NI 237

PHILOTIELLA SPECIOSA

PRAEPHILOTES ANTHRACIAS

PLATE 10

Male genitalia of Philotiella speciosa (above), Praephilotes anthracias (sec¬
ond row) and Paleophilotes triphysina (third row). Aedeagus lateral and
dorsal views to left, entire genitalia next, entire genitalia without valves
shown caudal view, and lastly, entire genitalia without valves, dorsal view.
Lower figures giving (left) forewing venation of S. orion to show measure¬
ment for relative distance ratio determinations of R to R3, base of R3 to
margin, and distal length R4 + 5 to margin. Lower figure (right) caudal
view of male genitalia of E. enoptes, not showing valves, indicating areas
measured to describe the labides. In this paper only the ratio of A/B
is given.

238

R. H. T. MATTONI

/. Res. Lepid.

PLATE 11

SCOLITANTIDES ORION PHILOTES SONORENSIS PSEUDOPH I LOTES VICRAMA

SHI JIMI AEOIDES LANTY

0.5 MM

I -

SINIA BAVIUS

PLATE 11

Female genitalia of S. orion, P. sonorensis, P. vicrama, S. bavius, and S.
lanty. Upper figure of each depicts the papilla anale and posterior apo¬
physis in lateral view. Middle figure shows the lodix in lateral view oriented
with the ostium vaginalis to the right. The course of the ductus bursa in¬
dicated by broken lines. The lower figure shows the lodix from the ventral
view. The specimen of S. lanty had lost its lodix.

239

16(4):223-242, 1977 THE SCOUT ANT1DIN1

PLATE 12

EUPHILOTES ENOPTES PH I LOTI ELLA SPECIOSA PALEOPHILOTES TRIPHYSINA B

PLATE 12

Female genitalia of S. leechi, S. clivina, E. enoptes, P. speciosa, and P.
triphysine. Orientation as in PLATE 11.

MORPHOLOGICAL DATA FOR RELATED TYPE SPECIES AND SOME OTHER SPECIES IN THE SCOLITANTIDINI

240

R. H. T. MATTONI

/. Res. Lepid.

CM O O

” a

O W IN

«n + * u,

*£>0 0 1/1

\ \

• 41

< C

•H SO z Z <N W O

O C

• o
•h z
\ \

> o> 4>

a> c w

T3 O U

O S Ok

O O O' r~

^ e

\ O \ \

in •• v£> a)

o o in vo c

o o o

Of X)

% 3

O' 4)
0) rH

o

u

o b

W KL

4) *D

C <N
4) ••

%

4J W

5 o

>

4) «

>1 ^

* ^

>j a

o <
8 o

§ B *.

4) \ <0

fc J ■H

.Q

O' M *H
4) 3 H

0 § P

M b 3
b £

_ O' 4)
•b 4) b

O O O
•H P *H
b -P

< (2 K t

b U

o) to
w u
4)

O <M

O O

• H TJ

U W <

16(4):223-242, 1977

THE SCOL1T ANTID1NI

241

O O rH

O' 0)

•h a
<o (0

H 3 XI
T3 e «
<0

3 £

O’ E->

V) a)

u o

«J <0

V c
m o

>i-H

x; (o
a c
o a>
a e

•H «J (U x

> fi »j\

U O' to J

0) *H O

u w a

v (A

v

to o

shijimiaeoides euphilotes philotiella praephilotes paleophilotes

DIVINA ENOPTES SPECIOSA ANTHRACIAS TRIPHISINA

242

R. H. T. MATTONI

J. Res. Lepid.

• O >w

-H C \

\ \ 44

in a» p

r-~ c 0

• O X!

O Z I-I m

X u
O' at
•H CL

01 73 T>

Li RJ
0 3 0

< •
• o
Z ••

m n o o

•H »h m o *3

•a T3

0 O

fl T5 13

o o o

o o r~

•H \
\ 01
O'

\

4-i m

Ll

0 oo

42

mo r~

M OH
•H 0 RJ
RJ g

a > oi

Li \ Rj

rj e u

m 3 oi

3 C -C

O' 0 4J

C O' >1

10 ox

•H -H o

o j rx m ©

O'

>4

Li t0>

01 %1

O p> D

*H V RJ

• O' o

H J O

\ \

O ►, O'

f~ Li C

• ai o

O > H J +

O' 01 rH

-h a rj

RJ RJ 01

Li 44 Li

© O 2 fX

0 O'
4= 01

a j
o

Of 43

% 3

5 o X

•H

• f 01

0 <6 >1 -H

2 Oi U CL,

oi o
c u
o o

% £

O' CL
« X
01 u

1*

00

oo oi

3 2

8 -s

3 01

& 3

o m

CL

00

« a

01 o

>i-H
43 00
CL C
O

a

« <

II

a

Li

O •

H M

1 K I

a -h

RJ RJ
CL C

jU

O CL

44 oi a
van

< RJ O

Journal of Research on the Lepidoptera

16(4) : 243-244, 1977

EDITORIAL

Since the death of Dr. Hovanitz, I have assumed the role of
acting editor of this Journal as directed by the trustees of the
Lepidoptera Research Foundation. We are seeking someone
who will take the position permanently, but are faced with num¬
erous problems in so doing. No small consideration is the phys¬
ical one of having our entire publishing facility, technical edit¬
ing, and business location in one place. Another is the sheer
responsibility of the job description, an item well expanded upon
by Stan Nicolay in his presidential message to the Lepidopter-
ists’ Society (1977 Jr. Lep. Soc. 31:217-222). In the meantime, the
mechanics of publishing so well established by Bill Hovanitz are
well oiled. The JRL has the financial prospects and will continue
as long as appropriate manuscripts need a home.

Although short papers, reviews, habitat descriptions, etc. will
be accepted, the value of the JRL lies in the longer, well illus¬
trated articles we prefer. These articles should emphasize re¬
search on environmentally and genetically induced variation,
patterns of distribution, population structure, behavior, compar¬
ative morphology, phylogenetic and phenetic systematics and
comparative physiology. In a word, on evolution as the central
theme of biology as applied to Lepidoptera.

The liberal use of color plates has been an important adjunct,
made possible by Bills ability to coordinate a printer in Pasa¬
dena and a color separation firm in Missouri. Thus an apparent
extravagance is permitted on our very limited budget. We are
studying setting up typesetting with our own automatic com¬
posing equipment. This step would be undertaken jointly with
other societies who also must find alternatives to the high costs
from “turn key” publishers. Such a move would also ease editorial
work both by propinquity and elimination of more layout efforts.

The basic policy of the Foundation will continue to be the
publication of the JRL with few minor changes. Bill had in¬
tended to standardize formats of all papers, and this will be done
in Volume 17. Changes will include providing more information
in the space available. Instructions to authors will be specified
on the back inside cover. The notice section will be separately
printed in a “newsletter” form which will permit and encourage
increased communication among members. The notices section

243

244

R. H. T. MATTONI

J. Res. Lepid.

is open to all members; we hope you will feel free to more ex¬
tensively use the service. Lastly, if we receive sufficient favorable
comment we will start a section of abstracts of papers of more
than usual interest which have not appeared in the regular Lep-
idoptera press. Accordingly, we should solicit your aid in pro¬
viding reprints or notices of such papers. Our objective is to in¬
crease both membership and member participation to both better
serve you and ease the load on us. The latter will occur with be¬
ing able to purchase outside help. We will have publication up to
date before the end of 1978 (Volume 17 completed). We invite
your comments, criticisms, and help.

Barbara Hovanitz will continue to give generously of her
time in the position of secretary-treasurer. She will handle all
business matters including subscriptions and memberships.
Karen Nielson Hovanitz as assistant editor is responsible for
formatting, layout, coordination of printing, illustrations, publi¬
cations, and mailing. Scott Miller has become a second com¬
petent assistant editor and colleague. Serendipity has provided
an able team. Bill would be proud of us.

R. H. T. Mattoni

Journal of Research on the Lepidoptera

16(4) : 245-248, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

THE USE OFOCECDYSONE
TO BREAK PERMANENT DIAPAUSE

OF FEMALE HYBRIDS BETWEEN
PAPILIO GLAUCUS L. FEMALE AND
PAPILIO RUTULUS LUCAS MALE
C. A. CLARKE

Department of Genetics, University of Liverpool, England

AXEL WILLIG

Department of Biochemistry, University of Oldenburg, Germany

Previously we have shown that both males and females
can be produced by the cross female P. rutulus X male P. glaucus
(Clarke and Sheppard, 1955) but the reciprocal cross only pro¬
duces males, the females remaining in permanent diapause until
they die.

In order to examine the phenotype of the FI females for
genetic purposes we attempted to use ecdysone (Clarke, Shep¬
pard and Willig, 1972) in order to get the female adults to
eclose. Although a number started to develop only one was
sufficiently advanced to score the phenotype, and that did not
emerge. The present paper reports the successful use of -ecdys¬
one and describes the phenotype of the FI females when the
female glaucus parent was the black form.

METHOD

^-Ecdysone was purchased from Simes, Milan. The hormone
was dissolved in ethanol and diluted with distilled water to give
a final concentration of 7.0 fxglml in 10% ethanol-water. The
pupae (1.4 g) were injected with 20 fx 1 each of this solution
(100 ng ^-ecdysone/ g w.w) through the intersegmental mem¬
brane of the abdomen, avoiding piercing the pupal gut, nervous
system or spiracles. The injections were repeated every second
day, each pupa receiving six in all. The butterflies emerged 24
days after the first injection.

RESULTS

Two female insects which were similar to one another were

245

246

C. A. CLARKE and A. WILLIG

/. Res. Leptd.

produced from the two pupae injected. It will be seen from the
plate (bottom right) that they are intermediate between the
black and yellow forms in that the normal yellow pattern is
present but heavily suffused with black scales. The suffusion is
almost complete in a triangular area at the base of the forewing,
extending out as far as the proximal black bar in normal yellow
rutulas. A corresponding black area is to be found in the hind¬
wing extending from the inner margin to the proximal black
line running through the cell towards the anal angle.

A similar female, together with a number of others showing
less yellow, right through to black glaucus, was obtained by back
crossing males of the Fi to black glaucus.

On the ventral surface the Fa females are also suffused with
black but to a much lesser degree, so that the general appear¬
ance is rather like a sooty yellow form.

DISCUSSION

The use of ^--ecdysone opens up the possibility of studying
the genetics of species differences. In the case of the present
cross this seems particularly likely as the F1 hybrid females
had their full complement of developed ova.

The black and yellow female forms of P. glaucus are con¬
trolled by a locus on the Y chromosome, apparently associated
with a cytological marker (Clarke, Sheppard and Mittwoch,
1976). The inheritance of the black form is therefore maternal
since the female is the heterogametic sex. The presence of an
intermediate Fl3 plus similar insects in the back cross using male
hybrids, demonstrates conclusively that rutulus carries an auto¬
somal or perhaps X-linked gene (or genes) which modifies the
effect of the Y-linked allelomorph controlling black. This find¬
ing has interesting implications with respect to the evolution of
the black form, which mimics Battus philenor. P. rutulus, al¬
though it flies with B. philenor in a number of places, including
the coastal range of California, has never evolved a mimetic
form. It seems possible that in the rutulus gene complex the
Y-linked black allelomorph if it arose would be so suppressed
(the Fas having only half the rutulus gene complex) that it
would not produce a phenotype sufficiently close to Battus to
have any advantage.

With the new technique we hope to get a series of back
crosses to rutulus to test this hypothesis.

16(4):245-248, 1977 THE USE OF << -EC DYSON E

247

TEXT FOR PLATE

Top left: Papilio glaucus female, black. Top right: Papilio glaucus female,
yellow. Bottom left: Papilio rutulus male. Bottom right: FI hybrid female,
ex black P. glaucus female X male P. rutulus.

248

C. A. CLARKE and A. WILLIG

J. Res. Lepid.

LITERATURE CITED

CLARKE, C. A. and SHEPPARD, P. M., 1955, The breeding in captivity
of the hybrids Papilio rutulus female X Papilio glaucus male. Lepid.
News, 9, 46-48.

CLARKE, C. A., SHEPPARD, P. M., and WILLIG, A., 1972, The use of
ecdysone to break a two and a half year pupal diapause in Papilio
glaucus female X Papilio rutulus male hybrids. The Entomologist, 105,
137-138.

CLARKE, C. A., SHEPPARD, P. M. and MITTWOCH, U., 1976, Hetero¬
chromatin polymorphism and colour pattern in the tiger swallowtail
butterfly Papilio glaucus L. Nature (Lond. ). 263, 585-587.

Journal of Research on the Lepidoptera 16 ( 4) : 249-264, 1977

do Santa Barbara Museum of Natural History
2559 Puesta del Sol Road, Santa Barbara, California 93105

THE ROLE OFINTRA-AND
INTERSPECIFIC MALE: MALE INTERACTIONS
IN POLYOMMATUS ICARUS ROTT. AND SOME OTHER
SPECIES OF BLUES (LYCAENIDAE)

LENNART LUNDGREN

Laboratory For Ecological Chemistry
University of Goteborge
S-431 33 Molndal, Sweden

INTRODUCTION

The males of Polyommatus icarus start to emerge and
fly before the females. The males show a greater flight activity
than the females. In addition, they mostly fly higher above the
ground vegetation and are more easily observed. The more or
less zigzag-formed patrol flight of the males has in other butter¬
flies (e.g. Argynnis paphia L., Magnus 1956) been interpreted
as a typical appetite behaviour, a sexual search flight which
brings the male into contact with the stimuli of females. A com¬
plete courtship is seldom observed. It is most often the vigor¬
ous mate refusal behaviour of the female or sometimes a pair
in copula that catches the sight. In fact it was the astonishing
frequency of male: male interactions that first struck me when
I started to study this species. During the patrolling the males
perpetually meet, whirl around and often pursue each other in
a straight fast flight. Such “male chases” of more than 25 meters
are not rare.

The aim of this paper is to discuss the role of the intra-
and interspecific male: male interactions in the light of our pres¬
ent knowledge of natural regulation of number in populations. A
survey of theories in this field is given by Dajoz ( 1974 ) .

The investigation of the approach reaction of the males was
performed in 1963 and in 1964, some complementary experiments
in 1966.

The field observations brought about the following ques¬
tions :

1. Are all butterflies visually equally attractive to the P. icarus
males?

2. Is the approach reaction of the P. icarus males released to
the same extent by conspecific males and females?

3. How attractive are the blue males of other species of blues
compared to P. icarus females and P. icarus males?

249

250

LENNART LUNDGREN

J. Res. Lepid.

MATERIAL AND METHODS

Spanned and dried butterflies were used as dummies to
release the approach reaction of the males. The dummies were
hanging freely close over the vegetation in a thin black thread
from 1 meter long hazelrods obliquely stuck in the ground. The
position of the dummies was randomized by arranging the
“angle rods” in a Latin square. A simultaneous testing of e.g.
5 types of dummies required consequently 5x5 rods placed at
a distance of 1 meter from each other. A moderate clearing of
the area was made to remove tall plants which could conceal
a dummy. Futher details about the experiments are summarized
in Tables I - IV.

The responses of the males were classified as approach-
reactions and fluttering around reactions. The approach reac¬
tions were recorded as a “D”, alluding to the fact that the re¬
sponse often was a fast dip movement against the dummy. It
was in practice hard to distinguish between D and F reactions
(fluttering around) though two observers were always re¬
cording the responses. D was much more frequent than F. The
experimental method was best suited to comparing the ap¬
proach tendency of the males to various visual dummies. At the
compilation of the results the D and F reactions were put to¬
gether.

If the same male when flying through the square responded
to more than one dummy, this was recorded as well as the
order of the approaches. The proportions of the results were
not changed when the repeated responses were excluded.

RESULTS

The field observations showed that butterflies are strikingly
more attractive to the males of Polyommatus icarus than other
insects are. This preference has in other species been explained
by the typical way of movement of butterflies.

1. Are all butterflies visually equally attractive to the P. icarus

males?

In five experiments (Table I) 4 dummies were various blue¬
winged blues but the 5th alternative was another type of butter¬
fly also occurring in the habitat. The choice of species was
partly determined by the supply of spanned and dried butter-

16(4):249-264, 1977

POLYOMMATUS ICARUS

251

flies. It is obvious that the F. icarus males did approach blue
colored blues far more often than other butterflies. The greater
attractiveness of the copper, Palaeochrysophanus hippotlioe L.
(Lycaenidae) and the fewer approaches to the brown lycaenid
females ( Cyaniris semiargus Rott. ) can also be noted.

2. Is the approach reaction of F. icarus males released to the
same extent by conspecific males and females?

The experiments show (Table II) that the males are as easily
attracted to specimens of their own sex. The brown females
of this species are more or less flushed with blue scales. The
dummy females used were intermediate blue. Out of 156 females
collected on Oland in 1965, 10 were classified as blue-shimmer¬
ing at the wing base, 55 were blue almost to submarginal lunules,
and 91 were blue almost to the black marginal line. No com¬
pletely brown females with or without orange lunules were
caught. The blue females are of course over-represented as they
can be more easily seen.

3. How attractive are the blue males of other species of blues
to the P. icarus males as compared to P. icarus females and
P. icarus males?

The answer to that question is that the males of the tested
species ( Cyaniris semiargus Rott., Lysandra icarus Esp. and
.Lysandra argester Rgstr. ) successfully compete with F. icarus
butterflies of both sexes about the approach reactions of the
F icarus males (Table II).

Even if the experimental method only tested the initial ap¬
proach stage of the male chase, the experiments have shown
that intra- and interspecific male: male interactions are two
characteristic ways of behaviour in Polyommatus icarus. Table
III illustrates the same behaviour in Plebejus argus L.

4. What stimuli release the approach responses of the males?
The experimental squares were set up in wind-protected

parts of the habitat. As a rule only the upper side of the dummy
wings was exposed to the males. In the first experiment the
spanned butterflies were first hanged 30 centimeters above
the vegetation. Most of the males flew below the dummies. Be¬
tween 9:20 a.m. and 2 p.m. 19 nonresponses and 8 approaches
were registered. The dummies were then lowered to hang close
over the vegetation. Between 2 and 5:10 p.m. only 2 out of 37
males passed through the area without taking notice of the dum-

252

LENNART LUNDGREN

J. Rea. Lepid.

mies. This behaviour indicates that colour signals from the upper
side of the wings are the active stimuli. An objection is that the
under side of the dummy wings is shaded. This was not the
case when a spanned and dried male was rotated around the
body axis. The rotating dummies were fixed to a thin steel axis
running through a 2 m long hollow spinning rod. The rotational
velocity could be varied with the spinning reel which drives
the axis. The alternating flashes of upperside and underside
colour from a male dummy with a “fluttering speed” which
seems normal to our eyes did not increase the atractiveness of
the male as compared to when presented not rotating.

To examine the reaction of the male to a larger blue wing
surface “giant males” (3 x) were made by covering a butterfly
silhouette of stiff paper with the wings from 3 males. The giant
males got almost as many responses as the normal P. icarus
males (Table II). As similar experiments have already been
made (Tinbergen 1942; Magnus 1956) no further experiments
were made to see if the ordinary size of the male is optimal. The
reactions to deformed males suggest that the shape of the
coloured object is unimportant to the males as in other species
of butterflies which have been investigated in this respect.

In many species of blues e.g. Polyommatus icarus and
Lysandra icarius the ground colour of the females varies from
brown to more or less blue. The readiness of the males to ap¬
proach blue objects ought to lead to a selection in favour of the
blue colour genes. The grey-green mutant colour form valesina
of Argynnis paphia is heriditary dominant expressed only in
females. Magnus (1956) has shown that one explanation of its
rareness is a sexual preference by the males for the normal yel¬
low-brown colour of the female.

The two species Plebejus argus and Lycaeidas idas are very
similar and often hard to distinguish without examination of
the genitalia. The F. argus females are brown but the L. idas
females more or less blue and rarely completely brown. The
square experiments have confirmed (Table III) the existence
of interspecific male approaches between these species.

P. argus and L. idas are consequently suited objects for an
investigation of the female colour preference of the males.
Fortunately, I had in my possession some blue and some brown
spanned female specimens of L. idas. The most eligible locality
found in the neighbourhood was an area in a pine forest clear-
cut some years ago, with Calluna vulgaris as a dominant in the

16(4):249-264, 1977

POLYOMMATUS ICARUS

253

herb layer. 60 males were caught and released again, 28 were
P. idas and 32 were P. argus. As it is impossible to determine
with certainty to which species the approaching males belong
without catching the butterflies a pure population of L. idas
would have been better.

The supply of spanned females permitted only a 3 x 3 square
with: 1 L. idas brown female, 2 L. idas blue female and 3
L. idas male. The result (Table IV A) is very interesting. Males
of both species respond to the dummies which was verified
by catching some of the reacting males. The experiment in¬
dicates a visual preference selection of blue L. idas females.
Even the P. argus males are more easily attracted to the blue
L. idas females than to their own brown females. This remark¬
able fact was further supported by two experiments in a locality
where only Plebejus argus were flying (Table IV B).

A similar visual interspecific female preference has earlier
been found in other species. According to Petersen et al. (1952,
1954, 1963) the white males of Pieris bryoniae Hubner prefer
the white females of P. napi L. to their own yellow females.
This may be a secondary effect of intra- and interspecific
male: male interactions released by the white male colour of
Pieris. The P. bryoniae males do more often approach each
other than their own females. Males of Papilio rutulus Bois-
duval are in the beginning of the courtship more often attracted
to female P. multicaudus Kirby than to their own females
( Brower, 1959 ) . In this case the explanation can be quantitative
differences. To human vision the females have the same yellow
colour, but the yellow area on the wings of P. multicaudus is
larger.

Sign stimuli are mostly qualitatively optimal. One of the
phenomena which the dummy experiments have revealed to the
ethologists is the existence of supernormal stimuli. Various
selection pressures often prevent the realization of quantita¬
tively optimal stimuli in the organism. It must yet be considered
as remarkable that in two sympatric species or two allopatric
species with a contact zone the females of one of the species
have got supernormal sign stimuli to the approach reaction of
the males of the other species. It is necessary to ask for the
compensating isolating mechanisms. Brower found a partial sea¬
son isolation and a difference in altitude preference in Papilio
multicaudus and P. rutulis. Petersen established that where
Pieris bryoniae and P. napi appear as different species habitat

254

LENNART LUNDGREN

J. Res. Lepid.

isolation and a bad hybrid vitality were active. An understanding
of the isolation of P. argus and L. idas needs further investi¬
gations. Although they morphologically are very similar species
the male genitalia are strikingly different (according to Chap¬
man the most different in tribus Plebejini). Sengiin (1944)
showed that curtailed males without the whole copulation ap¬
paratus including aedeagus are capable of fertilizing the females.
As Petersen (unpubl.) has pointed out such experiments are only
indicating. Small quantitative differences in male copulation
success can have a considerable isolating importance. Plebejini
is characterized by female genitalia deviating from those of
the main part of Rhopalocera. The vaginal orifice is situated on
the tip of an extrudable organ (hypostema + heina) behind
the 7th sternite. As I have failed to handpair these butterflies
it seems that the female copulation apparatus is not extruded
reflexably when the valves of the male grip the abdomen tip.
It may function as an effective refusal apparatus against males
which do not present the right stimuli.

5. Are the male: male interactions the defence part of a ter¬
ritorial behaviour?

Gently following males of Polyommatus icarus in various
localities did not indicate that the male: male interactions
are concerned with the defence of an area through chasing or
leading the intruder away from a domain to which the male
returns. Every male was followed until it was lost. The most
frequent reason was interference with another male. The males
were frequently lost during the flight chase or it was impos¬
sible to identify the traced male afterwards.

During observations of another species I spent some whole
days on a 200 meters long rectangular meadow. In this locality
the first icarus males emerged on the 11th June in the year 1966.
The male resided all the day in a well defined area in the east
part of the meadow. The size of the domain was estimated to
1/3 of the length and 1/8 of the width of the meadow. On the
13th June male No. 2 appeared. The first male was still flying
in the same area as before. Male No. 2 spent the whole day
in a less well-defined domain west of the flight area of No. 1.
In the afternoon a sustained “fluttering around” was observed
near the ground in the contact zone between the domains fol¬
lowed by a fast chase to the west end of the meadow. Later on
both males were again observed in their respective areas. On

16(4) : 249-264, 1977

POLYOMMATUS ICARUS

255

the next day several males were flying. The population was in
constant restlessness. The male chases were frequent.

Without individual marking one can not be absolutely sure
that defended territories do not exist but my field observations
indicate that the P. icarus males locate their mates by patrolling
according to the terminology of Scott (1972). Tendencies to
perching are observed at low population densities but male: male
interactions not attributable to the defence of a defined area
to which the defender returns are much more frequent.

A 16 mm colour film on the biology of Cupido minimus L.,
Maculinea arion L., Lycaeides argyrognomon Bgstr., Aricia
allous H-Gbg., Cyaniris semiargus Rott., Polyommatus icarus
Rott., Lysandra icarius Esp. and L. argester Bgstr. has been
made and it is filed in the Department of Entomology at the
Institute of Zoology, at Uppsala university, together with a
scenario in English. In this film (1,5 hrs) male:male inter¬
actions as well as feeding behaviour, courtship and egglaying
are documented.

DISCUSSION

I find it very unsatisfying to explain the male: male inter¬
actions as the result of a poorly characterized releasing mechan¬
ism for the sexual appetite flight. There ought to have been a
selection against unnecessary approaches as indicated by the
few approaches to other butterflies. The selective disadvantage
of “timewaste” during the search for females measured as re¬
duced fertilization rate has been demonstrated (Parker, 1970)
in a dung fly ( Scatophaga stercoraria L. )

What evolutionary advantages have maintained the high
level of intra- and interspecific interactions between blue males?
There must also have been a selection for epigamic recogni¬
tion. That is why the males’ chasing of each other seems too
sustained only to be a mate recognition behaviour. The expla¬
nation can be that the male: male interactions are agression
concerned with the defence of an area. Perching behaviour
is defined by Scott (1972) as a mate-locating method in which
males sit at characteristic sites and dart out at passing objects
in search of females. This type of behaviour is frequent in
butterflies but I know of only two investigations focused on
the return of the male to the same area, the defended territory.
A comparison between these is made in Table V.

According to Baker (1972) the benefit of the territorial be-

256

LENNART LUNDGREN

J. Res. Lepid.

haviour of the nymphalid butterflies, Aglais urticae L. and
Inachis io L. is an increased female quota for a territory-occupy¬
ing male. The optimum territories have an edge or comer site
where the stream of females is more intense. The study area
is not more closely described but the localities seem not to
be isolated. Both species are migrant. Voluntary male displace¬
ment occurs between the territorial periods. The distance cov¬
ered during a fine day is about 1 km in Aglais urticae and
0.5 km in Inachis ino. Without physically injuring the intrud¬
ers the territorial defence is rather elaborate involving in
I. ino defence by the very presence of a male in the territory,
chasing or leading the intruder away, and demonstration of
better manoeuverability and thus as a consequence seemingly
potential superiority in a coming courtship contest about the
same female. A low territory: male ratio, i.e. the ratio of suitable
territories to the number of males, is suggested to select for a
break-down of the territorial system. Tendencies in that direction
are shown by A. urticae through the increasing readiness of the
incoming males to share a territory with males already estab¬
lished rather than to continue to search for an unoccupied
territory.

Douwes (1975 b) has investigated the territorial behaviour
in Heodes virgaureae L. as a part of a detailed study of the
ecology and ethology of the adult stages of this species. The
study area was a well isolated, fairly small locality of the species
with a low migration and a low territory: male ratio. Only a
few males defend a couple of optimal territorial sites (perching
areas). Douwes’ results indicate that territorial H. virgaureae
males are inferior to non-territorial ones in the competition for
females and nectar resources, since females and suitable flowers
occur principally in the open field at some distance from the
male territories. The selective advantages of this territorial be¬
haviour is hard to understand but Douwes suggests it might be
that it makes the males cover the habitat better in their search
for females. Tendencies to territorial behaviour occur also out¬
side the territories preferred. The few males which occupy the
preferred territorial sites due to their superiority in competition
seem to “sacrifice” themselves for the best of the population.
The consequence ought to be a lower female quota and a
negative selection which work to eliminate this behaviour. A
solution of these difficulties might be to look upon the pre¬
ferred territorial sites as located not “at a distance from” but

16(4):249-264, 1977

POLYOMMATUS ICARUS

257

“on the route to” feeding and oviposition sites. According to
the description they can be edge territories similar to the ones
in 1. ino and A. urticae. That only one female was observed in
the territories could perhaps be explained by the more unob¬
trusive behaviour of the females and by the fact that the ter¬
ritorial study was performed before the peak of the female
number. The distribution study (Douwes, 1975 a, Fig. 5) shows
that the territorial sites lie at intense traffic routes. In addition
they lie close to two of the three most loco-climatically favour¬
able sites and two places with abundant Tubuliflorae flowers.
Douwes also studied thoroughly another population which
virtually lacked territorial sites and territorial males at least
with the characteristics defined from the other locality. Such
a habitat-adapted behavioural polymorphism is conceivable.
Chitty (1967) writes about intraspecific competition: “If we can
expect that interference has severe effects on survival and re¬
production it is less reasonable to assume that all genotypes
suffer equally than to assume that they are affected non-ran-
domly”. . Through the work of Gilbert and Singer (1973) we
know that in the butterfly Euphydryas editha “the longstained
selection pressure associated with the detailed ecology of each
population can produce intraspecific genetic differentiation in
dispersal tendency such that even populations with abundant
resources may differ in dispersal behaviour”. Douwes (1976 a)
found that also in the non-territorial population the colour re¬
sponsible for the attraction of the male to the female is less
strongly expressed in the female than in the male. This means
that the male: male interactions must be frequent also in the
non-territorial population.

Breakdown of territorialism at least in the butterfly species
discussed here does not lead to breakdown of the male: male
interactions. The intraspecific competition seems to have dif¬
ferent expressions at different population densities in the same
way as has been found in e.g. Mus musculus (Anderson, 1961)
with the exception that when territorialism breaks down when
the population density rises, no social hierarchy is established
in the butterflies in absence of ability to individual recognizing.

The male: male interaction are truely. density-dependent. If
the variations in behaviour as a function of population density
are summarized (Table VI), it seems highly probable that the
male: male interactions are the feed-back part of the natural
regulation of population density. All females are initially unwill-

258

LENNART LUNDGREN

/. Res. Lepid.

ing. Small differences in persistance are aimed to affect fe¬
cundity.

According to Magnus (1963) performing of “mistake” ap-
proachflights in Argynnis paphia gives a lowered stimulus thresh¬
old and a sexual over-excitement. The males’ chasing of each
other may contribute to maintain the sexual mood especially
in the beginning of the flight period when females are rare as
the males begin to hatch earlier than the females.

Why are the male butterflies more brilliantly coloured?
This problem has been discussed by many authors, e.g. Scott
(1972) and Douwes (1976 a). Is it because sexual selection
is more important in the males whereas in the females there
is a counter selection against predation? On the other hand male
colour is not exposed during the decisive stages of the courtship.
The existence of polymorphism mainly restricted to the females
speaks for sexual differences in selection. One suggested ex¬
planation is genetic (Magnus, 1963). If male: male interactions
have the vital importance for population regulation, which is
suggested in this paper, male colour is meant for other males.
This may also include chemical male signals. Douwes (1975 b)
found a gap between the male response to conspecific males in
H. virgaureae and to other butterflies which cannot only be ex¬
plained by differences in visual stimuli. In the Latin square ex¬
periments with P. icarus odor-males painted with a chloroform
extract of fresh males got more approach responses than non¬
odor males (Table I). These results may indicate that male
odor is a signal in the male: male interactions.

Color similarity and color unimportance at the end of the
courtship mean selection for chemical and tactile specificity.
The main components of the male wing volatiles have been
identified (Lundgren & Bergstrom, 1975) only in one species
of blues (Lycaeides argyrognomon ) . Since then the methods
for isolation and concentration of the odor substances as well
as the sensivity of the GLC-MS instrument have been improved.
A GLC of the volatiles from the male wings of the same species
(Fig. 1) shows a very complicated peak pattern, representing
more than 120 components with rich possibilities of species
specificity of the male scents. Polyommatus icarus , Lysandra
icarius, L. argester and Cyaniris semiargus have partly over¬
lapping spectra of larval host plants. They also share the nectar
resources. Nectar is continually produced but Wynn-Edwards
(1970) emphasizes correctly that all supplies of food are finite

16(4) -.249-264, 1977

POLYOMMATUS ICARUS

259

■*-' 0)

<D 03

bO 2 <D **

o S bo

C3 C) -*-> O

1 ^

2 go > .

1/3 S

bO^

x §T3 oh

M a^

3 6

O O *C

U odooco

260

LENNART LUNDGREN

J. Res. Lepid.

in terms of their yield per unit area and time. Changes in the
population structure of Euphydryas editha (Gilbert and Singer,
1973) indicate that the number of butterflies per unite of volume
of nectar is the operative density index. The distribution of the
lycaenid Heodes virgaureae (Douwes, 1975 a) shows a positive
correlation with the distribution of Tubifloral nectar plants.

It is hard to explain the high frequency of interspecific
male: male interactions between blue males compared to the
number of responses to males of other butterfly species unless
it involves some kind of group advantage. It is difficult to show
that the resource overlap is important enough to select for
interspecific male color similarity and male: male interactions
capable of regulating dispersion and population density within
the group in the same way as this behaviour obviously does
within the species. This remains a fascinating possibility. Such
an ecological relationship between species does not conflict
with the niche concept as long as this is not misinterpreted
as spatial. On the contrary the interspecific male: male inter¬
actions may be an important part of the niching of the re¬
sources.

SUMMARY

An high frequency of male: male interactions was observed in
Polyommatus icarus. The approach reactions of the males were
investigated using spanned and dried butterflies as dummies
arranged in Latin squares. All butterflies are visually not equally
attractive for the P. icarus males. Other butterflies (“non-blues”)
got only a few responses. Males in flight are as easily attracted
to specimens of their own sex as to females. Males of other
species of blues compete successfully with P. icarus females
about the approach reactions of the P. icarus males. The blue
wing colour of the upper side of the wings is an important
stimuli for the approach reaction. A visual preference for blue
females was found in Lycaeides idas where the female colour
varies from blue to brown. Also males of Plejebus argus pre¬
ferred visually the blue L. idas females to their own brown
females. The female genitalia of these species which have to be
extruded before the males succeed in copulation may be a
compensating mate refusal apparatus. Polyommatus icarus was
in the studied habitats found to be a patrolling species with
tendencies to perching. The male . male interactions cannot be
merely “mistake” approaches meant for females. True territorial
behaviour occurs in butterflies but male: male interference per¬
sists after the breakdown of territorial behaviour at low ter-

16(4):249-264, 1977

POLYOMMATUS ICARUS

261

ritory:male ratios. The following suggestions are made: The
male: male interactions are the feedback part of the natural
regulation of population density. The more brilliant male colours
and perhaps also the male scents are primarily directed to
other males. Colour similarity of males and interspecific male:
male interactions may be adaptive within species complexes
by adjusting the fecundity to overlapping resources.

ACKNOWLEDGEMENTS

I am grateful to Dr. Bjorn Petersen for focusing my interest
on the lycaenids, a neglected butterfly group in behaviour re¬
search. I should like to thank Professor Bertil Kullenberg for
providing stimulating working facilities at the Ecological Station
on Oland. My thanks are due to my wife for her help with the
field observations. I am also indepted to Gunnar Stenhagen for
making the GLC-analyses of the wing volatiles. Besearch support
from the National Swedish Enviroment Protection Board and
the Knut and Alice Wallenberg Foundation are gratefully ac¬
knowledged.

262

LENNART LUNDGREN

J. Res. Lepid

Table I

Polyommatus icarus. Male approach responses to some other butterfly
species compared to responses to blues of the species P. icarus (P.i.),
Cyaniris semiargus (C.s.), Lysandra icarius (L.i. ) and L. argester (L.a.).
Dried and spanned butterflies. Latin square 5x5. The odor dummies were
painted with a chloroform extract from 5 fresh male or female specimens.

Dummies

Other butterfly
species

Argynnis paphia $ 2

Brenthis ino £ 2

Palaeochrysopanus 17

hippothoe #

Coenonympha 0

pamphilus #

Cyaniris 2

semiargus 9 ( brown )

Table II

Polyommatus icarus. Interspecific male responses to males of other
species of blues compared to conspecific butterflies of both sexes. Dried
and spanned butterflies. Latin Square 3x3 and 5x5. Species name ab¬
breviations the same as in Table I.

Dummies

Exp.

1

2

3

4

No.

P.i. 9

Blues

P.i. 9 P.i. $

P.i.

1

12

(odor)

27

( odor)
37

44

2

16

20

18

30

3

39

49

57

63

P.i. 9

P.i.

C.s. $

L.i. $

4

20

19

16

14

P.i. 9

P.i. $

C.s. £

L.a. $

5

22

21

33

27

Exp.

1

2

3

4

5

No.

P.i. 9

P.i.

c.s. a

1

8

23

12

P.i. 9

P.i. #

C.s. 9

L.i. $

P.i. $

2

22

49

29

30

28

3

14

14

18

9

16

4

21

20

17

19

18

5

4

4

9

7

8

Sum

(exp.

2-5)

61

87

73

65

70

Table III

(3x)

Plehejus argus. Intra- and interspecific male .-male approaches. Dried
and spanned butterflies. Latin square 3x3.

Exp.

1

Dummies

2

3

No.

Plebejus

Plebejus

Cyaniris

argus $

idas #

semiargus $

1

10

23

19

2

20

42

27

30

65

46

Table IV

Plehejus argus and Lycaeides idas. Male approaches to brown and blue
females respectively. A. experiments with a mixed population of P. argus
and Lycaeides idas. Males of both species respond to the dummies. B. Only
P. argus occurred in the locality. Dried and spanned butterflies. Latin
square 3x3.

A. 1 2 3

Lycaeides idas Lycaeides idas Lycaeides idas
brown female blue female male

17 81 96

B.

Exp.

No.

1

2

Plebejus argus
brown female
0
7

Lycaeides idas
blue female
10
48

Plebejus argus
male
6
18

16(4) -.249-264, 1977

POLYOMMATUS ICARUS

263

Table V

Comparison between the territorial behaviour in Inachios io, Aglais
urticae, and Heodes virgaureae.

Behaviour

characteristics

I. io

(Baker, 1972)

A. urticae
(Baker, 1972)

H. virgaureae
(Douwes, 1975 b)

site of the

en route to

at the oviposition

at some distance

territories

oviposition sites

sites

from feeding and

territory :

high

low

oviposition sites
low

male ratio
number of males

1

4

1 - 2

per territory
mating

at roosting place

at the

at feeding

place

to which the

territory

places

time for holding

female is followed
240 min

90 min

1 - 2 days

a territory
time for setting

midday earlier

midday

morning

up a territory
number of

than A.u.

1

2

1

territories per
male and day
voluntary

before midday

before and

after the

displacement

until settling down

between the

territorial

in a territory

territorial periods

period

types of

deterring through

chasing or

chasing or

male: male

presence, chasing

leading

leading

interactions

(leading) away,

away

away

territorial :

contest of ma-
noeuverability
high

high

low

nonterritorial
male ratio
suggested

increased female

increased female

spacing out the

selective

quota, proximity

quota, containing

males, increased

advantages

of roosting sites

roosting sites

chances for the

Variations in

females to be
fertilized, de¬
creased female
quota for territorial
males

Table VI

behaviour as a function of population density.

Increasing population density

Increasing

Decreasing

male: male interactions
disturbances of egglaying females
physiological stress
net emigration

territorialism
territority:male ratio
undisturbed courtship persistence
fecundity

264

LENNART LUNDGREN

J. Res. Lepid.

LITERATURE CITED

ANDERSON, P. K. 1961. Density, social structure and non-social environ¬
ment in house mouse populations and the implications for regulation of
numbers. Tans. N.Y. Acad. Sc. 23: 447-451.

BAKER, R. R. 1972. Territorial behaviour of the nymphalid butterflies,
Aglais urticae (L.) and Inachis io (L.) J. Anim. Ecol. 41: 453-469.

BROWER, L. P. 1959. Speciation in butterflies of the Papilio glaucus
group. II. Ecological relationships and interspecific sexual behaviour.
Evolution 13: 212-228.

CHITTY, D. 1967. The natural selection of self-regulatory behaviour in
animal populations. Proc. Ecol. Soc. Australia 2: 51-78.

DAJOZ, R. 1974. Dynamique des populations. Masson et Cie, Ed. 301 pp.

DOUWES, P. 1975 a. Distribution of a population of the butterfly Heodes
virgaueae. Oikos 26: 332-340.

- , 1975 b. Territorial behaviour in Heodes virgaureae L. (Lep.,

Lycaenidae) with particular reference to visual stimuli. Noru). J. Ent.
22: 143-154.

DOUWES, P. 1976 a. Mating behaviour in Heodes virgaureae with par¬
ticular reference to the stimuli from the female (Lep.: Lycaenidae).
Ent. Germ. 2(2): in press.

- , 1976 b. Ecology and ethology of the butterfly Heodes virgaureae

(L.) Ph.D. Thesis.

GILBERT, L. E. and M. C. SINGER, 1973. Dispersal and gene flow in
a butterfly species. The American Naturalist 107: 58-72.

LEDERER, G. 1960. Verhaltensweissen der Imagines und der Entwick-
lungsstadien von Limenitis Camilla Camilla L. ( Lepidoptera, Nympha-
lidae). Zeit. fiir Tierpsychologie, 17: 521-546.

LUNDGREN, L. and G. BERGSTROM. 1975. Wing scents and scent-
released phases in the courtship behaviour of Lycaeides argyrognomon
(Lep.: Lycaenidae). J. Chem. Ecol. 1(4): 399-412.

MAGNUS, D. B. E. 1956. Experimental analysis of some “overoptimal”
sign stimuli in the mating behaviour of the fritillary butterfly Argyn-
nis paphia L. (Lepidoptera, Nymphalidae). Proc. Tenth Int. Congr.
Ent. Montreal. 2: 405-418.

MAGNUS, D. B. E. 1963. Sex-limited mimicry II. Visual selection in the
mate choice of butterflies. 16th Int. Congr. Zool. 4: 179-183.

PARKER, G. A. 1970. The reproductive behaviour and the nature of
sexual selection in Scatophaga stercoraria L. (Diptera: Scatophagidae).
J. Anim. Ecol: 205-228.

PETERSEN, B. O. TORNBLOM and N. O. BRODIN. 1952. Verhaltens-
studien am Rapsweissling un Bergweissling ( Pieris napi L and Pieris
hryoniae Ochs.). Behaviour 4: 67-84.

PETERSEN, B. and O. TENOW. 1954. Studien am Rapsweissling und
Bergweissling. ( Pieris napi L. and Pieris hryoniae O.) Zool. Bidrag
fran Uppsala. 30: 169-198.

PETERSEN, B. 1963. Breakdown of differentiation between Pieris napi
L. and Pieris hryoniae Ochs, and its causes. Zool. Bidrag fran Uppsala
35: 205-262.

TINBERGEN, N., B. J. D. MEEUSE, L. K. BOEREMA and W. W.
VARIOSSEAU. 1942. Die Balz des Samtfalters, Eumenis (Satyrus)
semele (L. ). Z. Tierpsychol. 5: 182-226.

SCOTT, J. A. 1972 (1973). Mating of butterflies. Jour. Res. Lepid. 11
(2): 99-127.

SENGiiN VON, A. 1944. Experimente zur Sexuell-mechanischen Isolation.
Rev. del la Faculte des Sciences 9 (4): 239-253.

WYNN-EDWARDS, V. C. 1970. Feedback from food resources to popu¬
lation regulation. In: Watson, A. 1970. Animal population in relation
to their food resources. Blackwell Sci. Publ.

NOTICES

NEW ADDRESS: The address of the Journal
Natural History, 2559 Puesta Del Sol Road

s now: c/o Santa Barbara Museum of
Santa Barbara, California 93105.

MAILING DATES:
will be I i sted

As of this issue, the mailing date of each number of the Journal
in the Notice column. Volume 16, Number 3: December, 1977.

NEW BOOKS: Available from Entomological Reprint Specialists, P.0. Box 77224,
Dockwei ler Station, Los Angeles, California 90007, USA:

Allan, P. B. M. Leaves from a Moth Hunter's Notebook, 1977. $9.00.

Dennis, R. L. H. The British Butterflies: Their Origin and Establishment.
1977. $20.00.

Goater, Barry. The Butterflies and Moths of Hampshire and the Isle of
Wright. 1975. $13.00.

Holloway, J. D. Moths of Borneo, with Special Reference to Mount Kinabalu.
1976. $14.00.

Pinhey, E. C. G. Moths of Southern Africa. 1975. $35.95.

FOR SALE: 1,500 exotic butterflies, insects, biologicals. 72 page catalog
$1.00. Complete Scientific, P.0. Box 307, Round Lake, IL 60073.

XERCES SOCIETY: An international organization dedicated to the conservation of
terrestrial arthropods and their habitats. Publications include the journal
Ata I a and the newsletter Wi nqs. Memberships and subscriptions US $5.00 (students
$2.50); contact treasurer, Roger Pasquier, 235 East 73rd Street, New York, NY
10021 .

BUY/SELL/EXCHANGE: Lepidoptera. Send $1.00 for list of species (worldwide).
Specialist in Sphingids, Saturniids and Exotics. Ray Adams, F.r.e.S., Carretera
Principal 139, Tamaraceite, Las Palmas De Gran Canaria.

WANTED: World moths, particularly Geometridae. Will trade North American or
Panamanian moths. Roger L. Heitzman, Dept, of Ent. , Univ. of M0, Agric. Bldg.,
Columbia, MO 65201.

FOR SALE:
mi I itary,
from Dana

6m by 2.5m linear Malaise Traps. Proven design used worldwide by
universities and private collectors. Additional information available
A. Focks, P.0. Box 12852, Gainsville, FL 3260 Dept. LRF.

v v

EXCHANGE: Lepidoptera specimens. B. Cech, Vancurova 13, 150 00 Prague,
Czechoslovakia.

WANTED: Correspondence with members interested in trading or selling North
American Lepidoptera. Send list of offers to: Marc Grocoff, 1950 Cottri I I
Lane, Westland, Ml 48185.

MANUSCRIPTS: May be sent to the Acting Editior at the Journal address.

NEW PUBLICATION: Lepidopterists'
hardbound history of the first 25
($6.00 to Lep. Soc. members) from
Louisville, KY 40208.

Society Commemorative. Volume: 374 page
years of the Lepidopterists' Society. $10.00
Dr. C.V. Coveil Jr., University of Louisville,

WANTED TO BUY: Comstock's Butterflies of California. Also, any
print books on North American Lepidoptera. J. G. Manesis, M.D.,
Fargo, Norht Dakota 58102.

other out-of-
Fargo Cl inic.

The note "Type locality for Calosaturnia walterorum Johnson (Saturn i i dae)",
which appeared in this journal (15(4): 214, 1977) was published by mistake, having
been formally withdrawn from consideration by the Journal of Research on the
Lepidoptera prior to its acceptance and publication in the Journal of the
Lepidopterists' Society (29(3): 191, 1975). We regret this error.

THE JOURNAL OF RESEARCH
ON THE LEFIJ DORTER A

Volume 16 Number 4 December, 1977

IN THIS ISSUE

Photoperiod and temperature in phenotype
determination of Pacific slope Pierini:
biosystematic implications

Arthur M. Shapiro 193

Antennal sensilla of some Crambinae

J. A. Kamm 201

Information on availability of holotypes of the
described taxons at a public institution
( Rhopalocera )

Dalibor Weiss 208

Gynandromorphic Polites skippers ( Hesperiidae )

M. C. Nielsen 209

Special thanks Barbara Hovanitz 212

Studies on the excretory system of the fully grown
larva of Callograma f estiva Donov. (Noctuidae)

U. V. K. Mohamed and Humayun Murad 213

Rectification of a recent paper on
Leucanella memusae gardineri

Richard S. Peigler 222

The Scolitantidini I: Two new genera and a
generic rearrangement (Lycaenidae)

Rudolf H. T. Mattoni 223

Editorial R. H. T. Mattoni 243

The use of <X-ecdysone to break permanent diapause
of female hybrids between Papilio glaucus
L. female and Papilio rutulus Lucas male

C. A. Clarke and Axel Willig 245

The role of intra- and interspecific male: male
interactions in Polyommatus icarus Rott. and
some other species of blues ( Lycaenidae )

Lennart Lundgren 249

czflcme

Bookbinding Co., Inc.

100 Cambridge St.
Charlestown. MA 02129