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THE JOURHAL ©F RESEARCH
OH THE LERIJOORTERA

Established in 1962

Founded by William Hovanitz
Edited by Rudolf H. T. Mattoni

Volume 19

1980

published by

The Lepidoptera Research Foundation, Inc.
at

2559 Puesta Del Sol Road, Santa Barbara, California 93105

Volume 19 Number 1 Spring, 1980(81)

IN THIS ISSUE

Date of Publication: September 7, 1981

Butterflies of Clark County, Nevada

George T. & Anna T. Austin 1

Book Review 64

Cover Illustration: Black and white adaptation of a color illustration from A
Flight of Butterflies. Reprint permission given by The Metropolitan Museum of Art,
New York, NY.

Volume 19 Number 2 Summer, 1980(81)

m THIS ISSUE

Date of Publication: September 7, 1981

Nantucket Pine Tip Moth, Rhyacionia frustrana, in Kem
County, California: Integrated Control and Biological

Notes (Lepidoptera: Tortricidae, Olethreutinae)

Dennis M. Poore 65

Moths of North America North of Mexico, Supplemental
literature: 1

J. C. E. Riotte 68

Acknowledgment 7 1

Demonstration of Reproductive Isolating Mechanisms in
Callosamia (Satumiidae) by Artificial Hybridization

Richard S. Peigler 72

Canalization of the Phenotype of Nymphalis antiopa
(Lepdioptera: Nymphalidae) from Subartic and
Montane Climates

Arthur M. Shapiro 82

Aberrant New Mexican Butterflies

Richard Holland 88

Diapause in Various Populations of Pieris napi L. from
Different Parts of the British Isles

E. Lees & D. M. Archer 96

Cover Illustration: Pen and ink rendering of C. securifera cf X C. promethea 9 by
Robin Jarrett of Los Angeles, CA. See R. Peigler article, page 72.

Volume 19 Number 3 Autumn, 1980(81)

IN THIS ISSUE

Date of Publication: November 20, 1981

A Revision of the American Genus Anisota (Saturniidae)

J. C. E. Riotte & Richard S. Peigler 101

Cover Illustration: Scanning electron micrograph of A. peigleri larval integu-
ment. Scale = 22 mm:l mm. See page 170, this issue.

Volume 19 Number 4 Winter, 1980(81)

IN THIS ISSUE

Date of Publication: November 20, 1981

Enzyme Electrophoretic Studies on the Genetic Relation-
ships of Pierid Butterflies (Lepidoptera: Pieridae) I.

European Taxa

H. J. Geiger 181

A New Tortyra from Cocos Island, Costa Rica (Lepidoptera:
Choreutidae)

John B. Heppner 196

Troidine Swallowtails (Lepidoptera: Papilionidae) in South-
eastern Brazil: Natural History and Foodplant
Relationships

Keith S. Brown, Antoni J. Damman & Paul Feeny 199

Karyology of Three Indian Lasiocampid Moths (Lepidoptera)

P. K. Mohanty & B. Nayak 227

Illustrations and Descriptions of some Species of Pyrrhopyginae
from Costa Rica, Panama and Colombia (Hesperiidae)

S. S. Nicolay & G. B. Small, Jr. 230

Errata: Scott 240

Book Review 241

Index: Volume 19 245

Cover Illustration: Left to right’,' pupae of Battus polydamas, Parides bunichus,
P. anchises nephalion, P. agavus and P. proneus, three views.

5'96" 1G05

Published By: The Lepidoptera Research Foundation, Inc,

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

Founder:

William Hovanitz

Editorial Staff: Rudolf H. T. Mattoni, Editor

Lorraine L. Rothman, Managing Editor
Scott E. Miller, Assistant Editor

Associate Editors: Thomas Emmel, U.S.A.

Arthur Shapiro, U.S.A.

Brian O. C. Gardiner, England
Bjorn Petersen, Sweden
Otakar Kudma, Germany
Ichiro Nakamura, U.S.A.

Miguel R. Gomez Bustillo, Spain

Manuscripts may be sent to the Editor at:

9620 Heather Road, Beverly Hills, CA 90210 (213) 274-1052

Notices Material may be sent to the Managing Editor.

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STATEMENT OF OWNERSHIP AND MANAGEMENT

THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a year, Spring, Summer,
Autumn, and Winter, by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the
publication and the general business office are located at 2559 Puesta Del Sol Road, Santa Barbara, California
93105. The publisher is THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The Editor is R. H. T.
Mattoni at the above Beverly Hills address. The Secretary-Treasurer is Barbara Jean Hovanitz at the general
business office. All matters pertaining to membership, dues, and subscriptions should be addressed to her,
including inquiry concerning mailing, missing issues, change of address, and reprints. The owner is THE
LEPIDOPTERA RESEARCH FOUNDATION, INC., a non-profit organization incorporated under the laws of
the State of California in 1965. The President is R. H. T. Mattoni, the Vice President is John Emmel, the
Secretary-Treasurer is Barbara Jean Hovanitz. The Board of Directors is comprised of R. A. Arnold, Barbara
Jean Hovanitz, and R. H. T. Mattoni. There are no bond holders, mortgages, or other security holders.

Journal of Research on the Lepidoptera

19(1): 1-63, 1980(81)

Butterflies of Clark County, Nevada

George T. Austin

and

Anna T. Austin

Nevada State Museum, Capitol Complex, Carson City, Nevada 89710 and

5077 Eugene Avenue, Las Vegas, Nevada 89108

Abstract. Clark County, the southernmost area of Nevada is in the
northern portion of the Mojave Desert. This area is topographically
diverse and with several distinct biotic communities. The butterfly
fauna is also diverse with 125 taxa recorded within the county. Of
these, at least 15 are non-resident strays. Five subspecies of widely
ranging polytypic species are endemic. Over forty percent of the
species have distributions mainly in the North American deserts;
most of the remaining species are more widespread.

Butterflies have been recorded in all months but are most abundant
from May through July. Of the species for which voltinism is known,
more than one -third are univoltine, one -fifth are bivoltine and the
remainder are multivoltine. The number of broods varies somewhat
with habitat and elevations and, in some species, on the occurrence of
summer rainfall.

Introduction

The distribution and occurrence of butterflies in Nevada are poorly
kno^ and there is very little literature pertaining to the state’s butterfly
fauna (Field et al 1974). The only regional list is for the Carson Range in
the western part of the state (Herlan 1962). There is also a recent list of the
species known from the state (Haijes 1980). The butterfly fauna of
adjacent California is known for the Mojave Desert in southeastern
Califomia (Emmel and Emmel 1973). Those of adjacent Arizona and Utah
are known less weU (Haskin 1914, Tidwell and Callaghan 1972). Nevada
records are largely scattered through the literature, listed in the seasonal
summaries published by the Lepidopterists’ Society, in the Nevada State
Museum and in various other collections.

We have resided in Las Vegas continuously since 1961. Our records and
those of others who have lived in or visited the area are now sufficient to
allow publication of an account and analysis of the county’s butterflies.

Our private collection forms the basis for this report. In addition, the
senior author has examined in whole or part several collections and has

2

J. Res. Lepid.

corresponded with others who have collected within the state. Over 13,000

specimens were viewed.

For each taxon, we briefly discuss what is known of its distribution,
phenology and ecology in the county and, where pertinent, taxonomic
comments and comparisons. Following this we list specimens examined,
other reported occurrences, larval foodplants recorded in Clark County
and resources used by adults. Specific records are not listed for many of
the veiy common species except for certain species of interest or for
unusual distributional or seasonal occurrences. Specimens examined are
those actually seen by the senior author in his visits to various collections
or those seen while in the field with other collectors (total number given
first in parentheses). Numbers, distribution among the sexes and collec-
tion are indicated. Additional records include specimens reported to us by
other collectors, sight records Gargely by the senior author) and reports in
the literature or seasonal summaries of the Lepidopterists’ Society.
Specimens in other collections are credited to the person (or institution)
who maintains the collection and, if different, the person who reported the
specimen to us. Sight records are indicated by an “s” following the
observer. Literature citations are presented in the usual form. Records
fi'om the seasonal summaries are cited by the year of the summary (summ.)
which is normally published in the second number of the News of the
Lepidopterists’ Society of the following year with occasional addenda in
later numbers. Specimens reported in the summaries that were later seen
are included under specimens examined. Foodplant records are those of
the authors unless indicated otherwise and are categorized as adult
association (adult assoc.), oviposition (ovip.), larvae and, in a few special
cases, pupae. Adult resource (flowers, unless indicated otherwise) obser-
vations were entirely by the authors. Plant identifications were mostly by
botanists at the University of Nevada, Las Vegas; plant nomenclature
largely follows Munz and Keck (1963) and Beatley (1976). Time of day is
based on the 24 hour clock (PST).

Butterfly nomenclature used herein follows dos Passes (1964) and later
revisionary works where applicable. All comparisons and taxonomic
determinations were made by the senior author. Low elevations generally
pertain to “desert” below the Pmon-Juniper belt {ca. 6000') and high
elevations are above 6000' in montane woodland and forest.

Collections which were examined or from which records were reported
are as follows: D. E. Allen (DEA), R. BaMowitz (RB), A. Bean (AB), J.
Brock (JB), Carnegie Museum (CM), D. Eff (DE), J. F. Emmel (JFE), C. D.
Ferris (CDF), C, F. Gillette (CFG), P. J. Herlan (PJH), H. L. King (HLK),
J. Lane (JL), R. L. Langston (RLL), C. S. Lawson (CSL), J. F. Leser (JFL),
Los Angeles County Museum (LACM), C. D. MacNeill (CDM), D. Mullins
(DM), Nevada State Museum (NSM), F. W. Preston (FWP), K. Roever
(KR), J. A. Scott (JAS), O. Shields (OS), R. E. Stanford (RES), K. B.

19(1): 1-63, 1980(81)

3

Tidwell (KBT), University of Nevada, Las Vegas (UNLV), and R. E. Wells
(REW). Additional Nevada records were supplied by C. Henne and B.
McGuire.

A number of specimens, some important, in the Nevada State Museum
from the Arrow Canyon Range (apparently west slope off U. S. 93) bear no
collection date. The museum card catalog lists the date 13 May 1969 for
some of these. We believe the correct date for all these undated specimens
to be 14 April 1969 based on (1) 2 Heliopetes ericetorum labeled Arrow
Canyon Range, 14 April 1969, (2) the date of 14 April 1969 was reported
for certain of these specimens in the 1969 summaiy, (3) other collections
were made in Clark Co. on 12 April 1969 (Searchlight, McCullough Mts.,
Blue Diamond) and (4) no specimens from Clark Co. (except 2 Euphilotes
battoides haueri which may be mislabeled) with the date 13 May 1969
could be found in the museum. The only other reference to the 13 May date
is of a collection of “P. mojave ” (=P. battoides baueri) in the Delamar Mts.,
Lincoln Co. (1969 summ.).

Description of Clark County

Clark County is located in southernmost Nevada in the northern portion
of the Mojave Desert. Several distinct biotic communities occur in the
county due to great topographic diversity with elevations ranging from
about 500' along the Colorado River to nearly 12,000' on Charleston Peak
in the Spring (Charleston) Mountains. The biotic communities of southern
Nevada were described in some detail by Bradley and Deacon (1965).
Important works dealing with plant distribution in the county include
Clokey (1951) and Beatley (1976). Brief descriptions of the major
vegetative associations follow.

Most of Clark County (78%, Bradley 1967) is dominated by xerophytic
vegetation. The largest association is the creosote bush community which
occurs at elevations below 4200' and is dominated by creosote bush
(JLarrea tndentata) and burrobush (Ambrosia dumosa). Numerous other
shrub species are subdominant and many species of annual plants occur.
At the higher elevations of this association there are often stands of Yucca.
Density and diversity of plants tend to increase along desert washes. Along
some major washes is a distinctive association of greater stature dominated
by mesquites iProsopis spp.) and catclaw {Acacia greggii). At elevations
between 4200' and 6000', the desert vegetation is characterized by
blackbush (Cokogyne ramosissima) with Joshua Tree (Yucca brevifolia) as
a subdominant in some areas.

Elevations between 6000' and 7300' are dominated by a woodland
characterized by Pinon (Hnus monophylla) and juniper (Juniperus osteo-
sperma) often with an understory of sagebrush {Artemkia spp.). A distinct
association including such species as silk-tassel (Garrya flavescem),
manzanita {Arctostaphylos pungens) and oak (Quercus gambelli) develops

4

J. Res. Lepid.

Fig. 1. Map of Clark County, Nevada, indicating major collecting localities
mentioned in the text. Contour lines indicates 6000' (dotted line) and 7500'
(solid line) elevations.

AC - Arrow Canyon Range
CC - Cabin Canyon
CN - Corn Creek
CP - Cottonwood Pass
GC - Grapevine Canyon
GS - Goodsprings area
KC - Kyle Canyon
LC - Lee Canyon
LO - Logandale
MF - Mormon Farm

MM ” McCullough Mountains
MS- Mt. Springs Summit
OV - Overton area
PV - Paradise Valley
RC - Red Cloud Mine
RD - Red Rock area
RR - Railroad Pass
SM - Sheep Mountains
WC - Willow-Cold Creek area
XM - Christmas Tree Pass

19(1): 1-63, 1980(81)

5

in disturbed areas. At elevations above 7300', there is a forest community
composed of Ponderosa Pine (pinus ponderosa) and White Fir (A6ies
concolor) at lower elevations and Limber and Bristlecone pines {Hnus
flexilis and P. aristata) becoming commoner at higher elevations. Under-
story shrubs include current {Ribes spp.), snowberry (Symphoricarpos
hngiflorm) and Mountain Mahogany (Cercocarpus ledifolius).

Other associations include relatively small areas but are important as
butterfly habitat. Cultivated and urban areas are found at low elevations,
principally in Las Vegas, Virgin and Moapa valleys. Marsh vegetation
develops in the presence of permanent water in such areas as Las Vegas
Wash, Tule Springs, Com Creek and in Moapa Valley.

Distinctive riparian associations develop in the vicinity of permanent
water at high elevations, notably at Little Falls in Kyle Canyon and at
Willow and Cold creeks in the northern portions of the Spring Range.
Meadows also occur in some cleared or naturally open areas at high
elevations including the old ski area in Kyle Canyon, along the ridge to
Charleston Peak and in the vicinity of the ski area in Lee Canyon.

In certain areas, especially on the east slope of the Spring Range in the
Red Rocks area, vegetation zonation is depressed in elevation in some
deep, cool canyons with permanent water. This results in a unique mixture
of vegetation not found in other portions of the county.

Annotated List of Species

In the following listing where no initials are given it will be assumed that
the records were made by G. T., A. T. and E. J. Austin. Otherwise, the
record citation will be indicated by the abbreviated collector or collection
name. Males and females in the numerical listing are cited as “m” and “f”
respectively. The locality citations are abbreviated according to the legend
given with Figure 1,

Family Megathymidae
Agathymm alliae (Stallings Turner) ssp.

Locally common only in the vicinity of Agave at middle elevations. The single
brood flies from late September to early November. Clark County material belongs
to the unnamed eastern Mojave Desert population as illustrated in Emmel and
Emmel (1973).

SPECIMENS EXAMINED: (41) 1 mi. W. MS, 28 Sept. 78 (9 m), 30 Sept. 79 (4
m), 10 Oct. 78 (10 m, 1 f), 11 Oct. 77 (1 m, 3 f), 22 Oct. 77 (5 m, 3 f), 2 Nov. 77 (4 0;
CC, 11 Oct. 78 (1 m).

ADDITIONAL RECORDS: 1 mi. W. MS, 1 Oct. 77 (s).

LARVAL FOODPLANTS: Agave utahemis Engelm. var. nevademis Engelm. ex
Greenm. &. Roush. (Agavaceae): 1 mi. W. MS Oarval tubes).

ADULT RESOURCES: m at water.

Megathymus yuccae navajo Skinner.

Uncommon in the higher desert to about 6000' in stands of Yucca. The flight
period is from early March to late May. Clark County material is assigned to the

6

J. Res. Lepid.

broad concept of navajo. In a more strict sense, these insects appear closest to M. y.
maudae. Stallings, Turner & Stallings.

SPECIMENS EXAMINED: (15) Newberry Mts., 15 Mar. 69 (1 f, NSM); CC, 13
April 78 (1 f, CSL), 27 April 78 (1 m; 1 m, CSL); CP Rd., 0.6-1. 8 mi. S. Pahrump Rd.,
16 April 79 (2 m, 1 f), 29 April 79 (1 f), 2 May 78 (1 m), 6 May 78 (1 f, CSL), 28 May
78 (1 m);KC 17.1 mi. W. U.S. 95, 26 Apr. 77 (1 m); 1 mi. W. RD Summit, 28 April 77
(1 m, NSM); 2 mi. N. RC, 4 May 78 (1 m); KC Ski Run, 13 May 78 (1 m).

ADDITIONAL RECORDS: Dead Mts., 7 Mar. 67 (PJH); WiUow Spring, 29
Mar. 78 (s); Wilson Pass, 29 Mar. 78 (s); CP to Pahrump Rd., 29 Mar. 78, 20 Apr. 77,
29 Apr. 78, 11 May 78 (s); 2 mi N. RC, 5 Apr. 78 (s); 2 mi, N. Sandy, 5 Apr. 78 (s);
Potosi Mine Rd., 8 Apr. 71 (REW); CC, 8 Apr. 77 ^B), 10 May 78 (s); XM, 12 Apr.
78, 20 Apr. 78, 3 May 78 (s); GC, 12 Apr. 78 (s); KC, 9-17 mi. W. U.S. 95, 17 Apr. 66,
18 Apr. 77, 1 May 77, 13 May 78 (s), 28 May 73 (JFL); U.S. 93, 8 mi. N. L15, 20 Apr.
75 (s); GS area, 20 Apr. 75 (JB); RD Summit Rd., 1. 5-1.8 mi. E. Lovell Wash, 24
Apr. 77, 28 Apr. 77 (s); WC, 27 Apr. 77 (s); 1 mi. W. WC, 8 May 78 (s).

LARVAL FOODPLANTS: Yucca haccata Ton*. (Agavaceae): Spring Mts., 30
Sept. 68 (larvae, 1968 summ.); RD, 25-27 Mar. 78 (pupae, REW); 1 mi. W. WC, 8
May 78 (ovip.). Yucca schidigera Roezl ex Ortgies (Agavaceae): RD, 25-27 Mar. 75
(pupae, REW); Charleston Blvd. nr. RD, 25 Nov. 78 (3 larvae in feeding tents, DM).

Family Hesperiidae

Lerodea eufala (Edwards).

Common to abundant in agricultural areas of Las Vegas, Moapa and Virgin valleys
with occasional records in the vicinity of desert springs and rarely in open desert
(Railroad Pass). The flight period extends from early May through mid November; it
becomes increasingly common in successive broods suggesting poor overwinter
survival. Four broods were produced at about 45 day intervals in the Logandale area
but only 2 or 3 broods in other areas during 1977.

SPECIMENS EXAMINED: (333) Numerous records for Moapa VaUey with
earliest on 7 May (78, 1 m, CSL) and latest on 11 Nov. (77, 3 m, 5 f). Records away
from Moapa Valley area: Las Vegas, 19 June 78 (1 m, CSL); MF, Las Vegas, 14 Aug.
77 (1 m), 13 Sept. 77 (1 m, 1 f), 25 Sept. 77 (Im, 3 f), 30 Sept. 77 (1 m, 1 f), 2 Oct. 77
(13 m, 8 f), 13 Oct. 77 (1 m), 16 Oct. 77 (Im), 21 Oct. 77 (2 m), 1 Nov. 77 (1 m);
Mesquite, 18 Sept. 72 (3 m, 4 f, NSM); 0.6 mi. W. Blue Diamond, 26 Sept. 77 (1 m);
RR, 3 Oct. 77 (1 m); Virgin Mts., 7 Oct. 73 (1 m, 3 f, NSM); CN, 7 Oct. 77 (If). The
only high elevation record is for WC, 23 Sept. 77 (1 f).

ADULT RESOURCES: Sphaeralcea ambigua QAslvaceae); Polygonum lapathi-
foUwn (Polygonaceae); Convolvulus sp. (Convolvulaceae) ; MecKcn^o satiw (Fabaceae);
HeUanthus annuus, Bebbia juncea, Pluchea sericea (Asteraceae).

Atrytonopsis python (Edwards).

A single record from Pine Creek is the only one for Nevada.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: Pine Creek, 30 June 63 (KR).

Ochlodes yuma (Edwards).

Locally common in or near stands of Phragmites communis flying in 2 broods with
peaks in late June (early June-early August) and in late September (late August
early November). It rarely strays far from these situations although it is found
commonly in alfalfa fields adjacent to Phragmites stands in Moapa Valley.

19(1): 1-63, 1980(81)

7

SPECIMENS EXAMINED: (190) Hidden Valley, 7 June 77 (1 m), 15 June 77
(5 m), 27 June 77 (5 m, 2 f), 4 July 77 (2 m), 12 July 77 (1 m), 25 Sept. 66 (1 m);PV, 9
June 79 (1 m, 1 f); Whitney Mesa, 9 June 79 (1 m), 6 Sept. 77 (3 m), 30 Sept 77 (4 m,
2 f), 12 Oct 77 (1 m), 21 Oct. 77 (1 m); CN, 12 June 66 (4 m, 2 f), 22 June 68 (5 m, 8 f),
18 July 66 (6 m, 7 f), 8 Aug. 77 (1 m, 3 f), 1 Sept. 63 (2 m, LACM), 3 Sept. 75 (14 m,
NSM), 10 Sept 75 (13 m, NSM), 15 Sept. 64 (4 m, 2 f, NSM), 25 Sept 65 (4 m,
NSM), 7 Oct 77 (8 m, 4 f), 14 Oct 77 (3 m, 3 f), 4 Nov. 77 (1 m, 1 f), 12 Nov. 77 (1 f);
LO, 15 June 77 (12 m, 4 f), 27 June 77 (10 m, 1 f), 4 July 77 (5 m, 1 f), 12 July 77 (1 0,
30 July 77 (1 m), 23 Aug. 77 (2 m), 9 Oct. 77 (1 m, 4 f), 23 Oct 77 (2 Bowman’s
Reservoir, 27 June 77 (1 m); Tule Springs, 4 July 62 (3 m); Clark County, 9 Sept. 76
(1 f, UNLV), 15 Sept. 73 (1 f, UNLV); Arden, 15 Sept 64 (1 f, NSM), 25 Sept 65 (1
m, NSM); Roger’s Spring, 15 Sept. 64 (2 m, NSM); OV, 15 Sept 64 (1 f, NSM); 25
Sept 66 (1 f, NSM), 27 Sept 65 (2 m, NSM); Valley of Fire, 27 Sept 65 (1 m, NSM);
Warm Springs, 27 Sept. 65 (1 m, NSM); Las Vegas, 2 Oct 75 (1 m, UNLV), 10 Oct.
70 (1 m, UNLV), 15 Oct 76 (1 f, UNLV).

ADDITIONAL RECORDS: LO, 7 June 77 (s), 10 July 72 (1972 summ.);
Roger’s Spring, 22 June 65, 25 Sept. 65 (PJH); CN, 22 June 68, 29 June 72 (JFL),
29 June 63 (KR), 31 July 65 (RES), 1 Aug. 65 (RES, JL), 1 Sept 63 (KR); Tule
Springs, 29 June 63 (KR); Cold Creek, 29 June 63 (KR); Hidden Valley, 4 Sept 77
(s).

LARVAL FOODPLi^TS: Phragmites communis Trin. (Gramineae): adult
assoc, nearly everywhere found.

ADULT RESOURCES: Heiiotropium curassavicum (Boraginaceae); Prosopis
pubescens, Medicago sativa (Fabaceae); Solidago spectabilis, Pluchea sericea,
Cirsium sp. (Asteraceae).

Atalopedes campestris (Boisduval).

Uncommon in agricultural areas of Moapa Valley (one record for Las Vegas
Valley) flying in 2 broods during July and October. There are no other Nevada
records.

Nevada material is consistently smaller than east coast and Texas specimens and
tends to be more heavily patterned on the hindwing beneath and the females have
considerably more orange above. They appear identical with a small series we have
from the Sacramento Valley, CaMomia and may warrant subspecific recognition.

SPECIMENS EXAMINED: (22) LO, 4 July 77 (1 m), 10 July 68 (3 m), 9 Oct 77
(4 m, 2 f), 16 Oct 77 (3 m, 3 f), 23 Oct 77 (2 m); MF, 2 Oct 77 (1 m); OV, 23 Oct 77
(3 m).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Medicago sativa (Fabaceae).

Polites sabuleti (Boisduval) ssp.

Locally common, especially in fall, in agricultural areas and occasionally in low
alkali areas. One record for higher elevations at Mt Springs Summit. Flies
apparently in three broods in April, June- July and early August to early November.

We refrain from placing southern Nevada material in any named race. Our
material is highly variable ranging from pale specimens with the hindwing markings
beneath nearly lost in the pale ground color which resemble P. s. chusca (Edwards)
to dark individuals from the same location and date with the markings on the lower
hindwing well developed and thus approaching nominate sabuleti.

8

J. Res. Lepid.

SPECIMENS EXAMINED: (96) PV, 14 April 77 (1 m, NSM); 10 mi. W.
Glendale Jet, 17 April 70 (1 f, NSM); MF, 30 April 77 (1 m), 25 June 77 (1 m, 1 f), 14
Aug. 77 (2 f), 3 Sept 77 (8 m, 3 f), 13 Sept 77 (6 m), 25 Sept 77 (4 m, 2 f), 2 Oct 77
(18 m, 1 1 f), 16 Oct 77 (1 f), (1 f); LO, 17 May 78 (1 m), 10 July 68 (1 m), 4 Sept 66 (1
m), 28 Sept. 67 (1 m, 1 f, NSM), 9 Oct 77 (1 m, 3 f), 16 Oct 77 (1 m, 3 OV, 4 Sept
66 (1 f), 26 Sept 65 (3 m); Bowman’s Reservoir, 7 May 78 (1 f, CSL), 27 Sept 78 (1
m), 29 Sept. 66 (1 f, NSM), 29 Sept 77 (2 m), 27 Oct 65 (3 m, NSM); CN, 15 Sept
64 (8 m, 1 f, NSM); OV, 27 Sept. 78 (1 m); Willow Spring, 24 Oct 65 (1 m).

ADDITIONAL RECORDS: CN, 22 June 68 (KR), 10 Sept 75 (CSL), 7 Oct 77
(s); MS, 30 June 63 (KR); Clark County, 8 Aug. 74 (JAS); LO, 23 Aug. 77, 17 Sept
77, 23 Oct. 77 (s); MF, 24 Aug. 77, 30 Sept 77, 21 Oct 77 (s); GC, 14 Sept 77 (s);
Whitney Mesa, 30 Sept 77 (s); OV, 23 Oct. 77, 5 Nov. 77 (s).

ADULT RESOURCES: Polygonum lapathifolium (Polygonaceae); Convolvulus
sp. (Convolvulaceae); Heliotropium curassavicum (Boraginaceae); Medicago sativa
(Fabaceae); Baccharis sp. (Asteraceae).

Polites draco (Edwards).

A male from Kyle Canyon is the only record for the state.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: KC, 16 June 77 (J. W. Tilden, det CDM, RES.).
Hesperia comma harpalus (Edwards).

Not uncommon in the Spring Range from 6000' to 9000'. It flies in apparently two
broods from mid May to early August and late August to late October. Spring Range
material is large and quite orange above and thus differing from Great Basin
populations and possibly showing some influence of H. a susanae Miller. Its double
broodedness appears unique among harpalus.

SPECIMENS EXAMINED: (140) Numerous records especially in KC and the
WC area with records between 12 May (1934, Charleston Mts., 1, LACM) and 24
Oct. (1977, KC Ski Run, 1).

ADDITIONAL RECORDS: LC, 29 May 50 (MacNeill 1964), Charleston Peak
trail, 10,800-11,918', 24 June 68 (JFE, OS); KC, 6768', 30 June 50 (18 m, 12 f,
FWP); Deer Creek Rd., 7000', 1 July 50 (4 m, 8 f, FWP); KC Campground, 2 July 78
(s); KC Ski Run, 2 July 78, 25 July 78, 8 Oct. 77 (s); Charleston Park, 8-15 July 28
(MacNeill 1964); LC Ski area, KC, Deer Creek area, 8 Aug, 74 (JAS).

ADULT RESOURCES: Apocynum androsaemifolium (Apocynaceae); Sarcos-
temma cynachoides (Asclepiadaceae); Penstamon palmeri (Scrophulariaceae);
ChaerMctis douglasii, Chrysothamnus sp., Cirsium sp., Taraxacum officinale
(Asteraceae).

Hesperia nevada (Scudder).

A single specimen of the western Great Basin population of this species is known
from Clark County.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: Charleston Mts., 13 May 31 (MacNeill 1964).
Hesperia pahaska martini MacNeill.

The only records are from the Sheep Range. Its reported foodplant, Tridens
pulchellus, is widespread in the county and the presence of other colonies is
probable.

SPECMENS EXAMINED: (3) Sawmill Canyon, E. side of SM, 6000-6500', 1
July 69 (1 f, LACM), 2 July 69 (1 m, 1 f, LACM).

19(1): 1-63, 1980(81)

9

ADDITIONAL RECORDS: None.

Hesperia juba (Scudder).

Rsffe above 6000' in the Spring Range and in the Virgin Mts. with records
indicating two broods from late May to mid July and late September to late October.

SPECIMENS EXAMINED: (7) WC, 6 June 78 (1 f), 22 June 78 (1 f); CC, 7 June
78 (1 f, CSL); KC Ski Run, 15 July 79 (1 f, CSL); KC, 9 Oct 66 (1 m, 1 f); Echo
Canyon, 23 Oct 66 (1 f).

ADDITIONAL RECORDS: Charleston Park, 29 May 50 (MacNeill 1964); LC,
29 May 50 (MacNeill 1964); Cold Creek, 29 June 63 (KR); Cathedral Rock, 12 July
72 (DEA); KC, 26 Sept 65 (PJH).

Hylephila phyleus (Drury).

Common to abundant in urban and agricultural areas, less common in the vicinity
of desert springs. It is rare away from these situations with occasional records for
desert washes in fall and two records for high elevations. The flight period is from
late March to mid December.

SPECIMENS EXAMINED: (364) Many locations in Las Vegas and Moapa
valleys with a small population in GC. Dates extend from 31 Mar. (78, PV, s) to 15
Dec. (79, Las Vegas, CSL). A sexual mosaic was taken at Hidden Valley on 23 Oct
77.

LARVAL FOODPLANTS: Cynodon dactylon (L.) Pers. (Gramineae): Las
Vegas, 7 June 77 (ovip.).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Sarcostemma hir-
tellum (Asclepiadaceae); Heliotropium curassavicum (Boraginaceae); Prosopis
glandulosa, Medkago sativa (Fabaceae); Viguiera deltoidea, Bebbia juncea, Baileya
multiradiata, Perityle emoryi, Senecio douglasii, Baccharis sp., Cirsium 8p.,Stephan-
omeria sp. (Asteraceae).

Copaeodes aurantiaca (Hewitson).

Fairly common in urban and agricultural areas, less common and somewhat local
in desert washes and around springs and rare at high elevations (Willow Creek, Kyle
Canyon Campground). The flight period extends from early March through early
December in 3 or more broods.

SPECIMENS EXAMINED: (109) Many records for Las Vegas and Moapa
valleys with extreme dates from 3 Mar. (75, Las Vegas, NSM) to 1 Dec. (77, Tule
Springs, s). High elevation records are for WC, 22 June 68 (JFL) and KC
Campground, 25 July 77 (1 f).

ADULT RESOURCES: Sphaeralcea ambigua (Malvaceae); Eriodictyon angus-
tifolium (Hydrophyllaceae); Heliotropium curassavicum (Boraginaceae); Medkago
sativa (Fabaceae); Helianthus annuus, Bebbia juncea, Baileya multiradiata, SoU-
dago spectabilis, Tetradymia axillark, Stephanomeria sp. (Asteraceae).

Pholisora libya libya (Scudder).

Uncommon and very local in (some) stands of Atriplex canescens in Las Vegas and
Moapa valleys. It flies in two broods from mid April to late June and early
September to mid October, The only record away from the valley floors is for Kyle
Canyon at 6800' at an unusual date of 27 July.

Specimens from Moapa Valley are of fhe nominate subspecies; certain

specimens from the Las Vegas Valley (Com Creek) have reduced maculation on the
ventral hind wing thus showing intermediacy towards P. libya lena (Edwards) of the
Great Basin.

10

J. Res. Lepid.

SPECIMENS EXAMINED: (57) Hidden Valley, 4 May 77 (1 m); 1 mi. E. OV, 4
May 77 (4 m); Bowman’s Reservoir, 4 May 77 (1 m, 4 f), 7 May 78 (1 f, CSL), 17 May
78 (1 m, 1 f, NSM; 1 m, 1 f, CSL; 5 m, 2 f), 7 June 77 (1 f), 15 June 77 (2 m, 1 f), 17
Sept. 77 (1 f), 27 Sept. 78 (2 m); LO, 7 June 77 (1 f), 16 June 77 (2 m, 1 f, CSL; 2 m, 1
f), 16 Oct. 77 (1 f); Las Vegas, 18 June 68 (1 f, UNLV); CN, 22 June 68 (Im, 2 f), 3
Sept. 75 (2 m, 2 f, NSM), 15 Sept. 64 (4 m, 2 f, NSM), 15 Sept. 66 (1 m, NSM), 7 Oct.
77 (1 m); MF, 25 June 77 (1 m); KC at Deer Creek Rd., 27 July 61 (1 m); Clark Co.,
17 Sept. 73 (1 m, UNLV); Tule Springs, 25 Sept. 62 (1 m).

ADDITIONAL RECORDS: Bowman’s Reservoir, 13 Apr, 78, 29 Sept. 77 (s);
PV, 24 Apr. 78, 10 May 77 (s); CN, 22 June 68, 29 June 72 (JFL), 29 June 63, 1 Sept.
63 (KR), 25 Sept. 64 (PJH); Las Vegas, 10 Sept. 75 (s), 6 Oct. 62 (JFL); OV, 17
Sept. 77 (s).

ADULT RESOURCES: Heliotropium curassavicum (Boraginaceae); Prosopis
glandulosa, Medicago sativa (Fabaceae); Baileya pleniradiata, Baccharis sp.,
Pluchea sericea (Asteraceae).

Pholisora gracielae MacNeill.

Locally common to abundant in Moapa Valley but only in close association with
Atriplex lentiformis. We have not found this species in stands of this plant in the Las
Vegas Valley or in the Colorado River Valley south of Davis Dam. Its flight period
began sometime before 17 April and continued in 3 nearly non -overlapping broods
through 29 September during 1977. Approximately equal peaks in numbers were at
2 month intervals on 4 May, 4 July and 4 September with numbers sharply lower 1-2
weeks on either side of the peaks. California populations were reported to have but 2
broods (MacNeill 1970, Emmel and Emmel 1973). The first brood has well
developed subapical spots on the forewing; the second and third broods have these
spots much reduced or wanting.

SPECIMENS EXAMINED: (122) Hidden Valley, 13 Apr. 78 (1 m, CSL), 17
Apr. 77 (17 m, 3 f), 4 May 77 (15 m, 7 f), 7 May 78 (1 f, CSL; 1 m, 1 f, NSM), 15 May
77 (5 m), 7 June 77 (1 m), 1 5 June 77 (3 m, 2 f), 27 June 77 (6 m), 4 July 77 (20 m, 7 f),
9 Aug. 77 (1 f), 4 Sept. 77 (7 m, 1 f), 17 Sept. 77 (1 m); Bowanan’s Reservoir, 4 May
77 (2 m, 1 f), 4 July 77 (2 m), 29 Sept. 77 (1 m); LO, 15 June 77 (1 f), 16 June 77 (2 f,
CSL), 17 June 77 (1 f, NSM), 27 June 77 (1 f), 10 July 68 (1 m, 1 f), 12 July 77 (2 m, 1
f), 23 Aug. 77 (3 m, 1 f), 4 Sept. 66 (2 f).

ADDITIONAL RECORDS: Bowman’s Reservoir, 27 June 77, 12 July 77, 23
Aug. 77, 4 Sept. 77 (s); LO, 4 July 77 (s); Hidden VaUey, 12 July 77, 30 July 77, 23
Aug. 77 (s).

LARVAL FOODPLANTS: Atriplex lentiformis fTorr.) Wats. (Chenopodaceae);
Moapa Valley (adult assoc.).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Heliotropium
curassavicum (Boraginaceae); Medicago sativa (Fabaceae).

Pholisora alpheus oricus Edwards.

Locally common, especially along desert washes, above the valley floors. The
single brood flies from mid March to late May. It is invariably found in association
with Atriplex canescens.

SPECIMENS EXAMINED: (57) Ash Springs, 19 Mar. 72 (1 m); KC, 6-17 mi.
W. U.S. 95, 4 Apr. 77 (1 m), 10 Apr. 78 (2 m), 11 Apr. 78 (1 m), 17 Apr. 66 (1 m)^ 18
Apr. 77 (2 m, 1 f), 18 Apr. 78 (2 m), 19 Apr. 78 (5 m, 1 f), 26 Apr. 77 (1 m) 26 Apr. 78
(3 m, 1 f), 30 Apr. 78 (5 m), 1 May 67 (1 m, NSM), 1 May 77 (2 m, 2 f), 2 May 78 (1 m).

19(1): 1-63, 1980(81)

11

11 May 78 (Im), 13 May 78 (1 m); 0.5 mi. W. Blue Diamond, 20 Apr. 77 (If); 0.5 mi.
S. Red Springs, 21 Apr. 79 (3 m, 1 f), 29 Apr. 79 (2 m, 3 f); CP Rd., 0.6-L8 mi. S.
Pahrump Rd., 22 Apr. 78 (4 m), 11 May 78 (1 m, 1 f); LC, 23 Apr. 66 (1 m); WC, 27
Apr. 77 (2 m), 29 Apr. 77 (2 m, 1 f).

ADDITIONAL RECORDS: 9 mi. W. Davis Dam, 23 Mar. 78 (s); Davis Dam, 31
Mar. 62 (R. C. Bechtal,/ide OS); KC, 5000', 3-4 May 75 (1975 summ.); Cold Creek
Ranger Station, 8 May 78 (s); KC, 17.1 mi. W. U.S. 95, 28 May 77 (s).

ADULT RESOURCES: Baileya multiradiata, Accamptopappus shockleyi,
Taraxacum officinale (Asteraceae).

Heliopetes domicella domicella (Erichson).

The single record from the Newberry Mountains is the only one for the state. The
closest known collection site is located near Parker Dam, San Bemadino Co.,
Palifomia (Emmel and Emmel 1973), about 70 mi. to the south.

’ SPECIMENS EXAMINED: (1) GC, 12 Oct. 77 (1 m).

ADDITIONAL RECORDS: None.

Heliopetes ericetorum (Boisduval).

Common, usually near water, at elevations to 8500'; less common along desert
washes and other areas away from water. The species is at least double brooded at
high elevations and probably multivoltine at some low elevation sites. It flies at low
elevations from late March to early November with very few July and August records
and at high elevations from early May to late October with veiy few records from
early August to mid September.

SPECIMENS EXAMINED: (128) The species is found nearly throughout the
county except in the lower desert away from washes or roadsides. Extreme flight
dates are from 21 Mar. (78, Las Vegas, 1 f, CSL) to 4 Nov. (77, Las Vegas, 1 m,
UNLV).

LARVAL FOODPLANTS: Sphaeralcea ambigua Gray (Malvaceae): Lovell
Wash, 8.9 mi. W. Pahrump Rd., 22 June 77 (ovip.). Sphaeralcea grossulariaefoUa
(Hook. & Am.) Rydb. vai. pedata (Torr.) Kearney (Malvaceae): 1 mi. W. WC, 25
May 78 (ovip. lower leaf surface, 12:15 PST).

ADULT RESOURCES: Sphaeralcea ambigua (Malvaceae); Stanleya elata.
Erysimum asperum (Bmssicaceae); Eriogonumfasciculatum, Eriogonum heermannu
(Polygonaceae); Sarcostemma hirtellum (Asclepiadaceae); Eriodictyon angustifol-
ium (Hydrophyllaceae); Verbena gooddingii (Verbenaceae); Monardella sp.
O^amiaceae); Acacia MeliloUts albus, Medicago saliva (Fabaceae); Sambucus
caerulea (Caprifoliaceae); Viguiera deltoidea, Bebbia juncea, Baileya multiradiata,
Chaenactis sp., Semcio ■ douglasii, Senecio multilobatus, Tetradymia axillaris,
Cirsium sp., Pluchea sericea (Asteraceae); males frequently visit water.

Pyrgus scriptura (Boisduval).

Uncommon but widespread mainly along desert washes but also as high as
11,000'. This is a greater elevational range than reported previously (Scott 1975,
MacNeill in Howe 1975). The flight period extends from late March to mid
November. Most records are for April to June and September (no August records)
indicating at least 2 broods. Spring and fall records are largely at lower elevations;
June records are mainly at high elevations,

SPECIMENS EXAMINED: (46) 6 mi. N. Apex, 28 Mar. 71 (2 m, 3 f, NSM); CP
to Pahmmp Rd., 29 Mar. 78 (1 m), 22 Apr. 78 (3 m), 3 May 78 (1 m, CSL), 6 May 78
(1 m, CSL), 11 May 78 (1 m), 28 May 78 (3 m); 2 mi. E. Searchlight, 12 Apr. 69 (1 m,

12

J. Res. LepM.

NSM); 1 mi. N. Dry Lake on L15, 19 Apr. 75 (1 m); WC, 23 Apr. 78 (2 m), 27 Apr. 77
(2 m), 29 Apr. 77 (1 f, NSM; 2 m, CSL), 6 June 78 (1 m); KC, 945 mi. W. U.S. 95, 1
May 77 (1 m), 6 June 78 (1 m), 23 June 77 (1 f), 28 Sept. 77 (1 m); Nipton Pass, 4
May 78 (1 m); WC, 6 June 78 (2 m, 1 f), 15 June 78 (1 m), 22 June 68 (1 f), 22 June 78
(1 m), 23 June 62 (1 m), 20 July 78 (1 m); LC, 8200', 10 June 62 (1 m, 1 f); Lovell
Wash, 4500', 17 June 62 (1 m); KC, 6.6 mi. W., 0.9 mi. N. U.S. 95, 21 June 78 (1 m);
Las Vegas, 22 June 66 (1 f); Hidden Valley, 12 July 77 (1 m); LO, 28 Sept. 67 (1 m,
NSM); Clark Co., 28 Oct. 70 (1 f, UNLV).

ADDITIONAL RECORDS: LC, 0.54 mi. W. U.S. 95, 7 Apr. 71 (REW);Potosi
Ml Rd., 8 Apr. 71 (REW); U.S. 93, 5 mi. N. L15, 17 Apr 77 (s); U.S. 95, 5 mi. N.
KC, 23 Apr. 78 (s); CP to Pahramp Rd., 2 May 78 (s); 1 mi, W. WC, 8 May 78 (s);
Hidden Valley, 15 May 77 (s); WC, 6 June 78 (s); Trout Canyon, 14 June 77 (s); LC
baUfield, 23 June 77 (s); Mt. Charleston, 11,000', 24 June 68 (JFE, OS); MS, 30
June 63 (KR);Las Vegas, 9-10 Sept. 62 (JFL);Moapa VaUey,21 Oct. 67 (JL); 11 mi.
W. Searchlight, 11 Nov. 75 (CDM).

ADULT RESOURCES: Phacelia fremontii, Eriodictyon angustifolium (Hydro-
phyllaceae); Baileya multiradiata, Tetradymia canescem, Taraxacum officinale
(Asteraceae).

Pyrgus communis (Grote).

Common to abundant in many habitats throughout the county to the top of
Charleston Peak. At low elevations, the flight period extends from early February to
mid December and at high elevations from early May to mid November.

SPECMENS EXAMINED: (431) Records from a large number of locations
with extreme dates of 4 Feb. (77, Boulder City, s) to 17 Dec. (70, Calico Basin, 1 f,
UNLV).

LARVAL FOODPLANTS: Sphaeralcea ambigua Gray (Malvaceae): CP, 1.5 mi.
S. Pahramp Rd., 22 Apr. 78 (ovip., lower leaf surface of lower leaves, 12:30 PST).

ADULT RESOURCES: Sphaeralcea ambigua (Malvaceae); Smilacina stellata
(Liliaceae); Tamarix pentandra (Tamariacaceae); Descurmrm pirmata (Brassicaceae);
Eriogonum fasciculatum, Eriogonum wrightii, Eriogonum heermannii, Eriogonum sp.
^olygonaceae); Allionm incamata (Nyctagnaceae); Apocynum androsaemifolium
(Apocynaceae); Sarcostemma hirtellum (Asclepiadaceae); Langlosia setosksima
^olemoniaceae); Phacelia fremontii, Eriodicfyon angustifolium (Hydrophyllaceae);
HeUotropium cura^savicum, Crypatantha sp. (Boraginaceae); Physalis crassifolia
(Solanaceae); Monardella sp. (Lamiaceae); Medicago sativa, Trifolium repem
(Fabaceae); Vigukra multifolora, Helianthus annual, Bebbia juncea, Baileya multi-
radiata, Baileya pleniradmta, Chaenactis fremontii, Pectis papposa, Acamptopappus
shockl^, Chrysothamnus sp., Erigeron sp., Senecio douglmU, Tetradymia canescem,
Cirsium sp., Taraxacum officinale (Asteraceae).

Erynnis brizo burgessi (Skinner).

Locally common in the vicinity of oaks from 4000' to 7000'. The single brood flies
from late March to mid June.

SPECIMENS EXAMINED: (94) WiEow Springs, 23 Mar. 77 (2 m, NSM); Lost
Creek, 23 Mar. 77 (1 m, NSM), 29 Mar. 78 (1 f, CSL), 20 Apr. 77 (2 m); CC, 29 Mar.
71 (2 m, 1 f, NSM), 8 Apr. 79 (6 m), 13 Apr. 78 (1 m, NSM; 2 m), 14 Apr. 77 (5 m, 3 f,
NSM), 27 Apr. 78 (1 m, 1 f, NSM; 2 m), 10 May 78 (4 m, 1 f), 12 May 77 (1 m, CSL);
Icebox Canyon, 21 Apr. 77 (1 f, NSM); WMte Rock Springs, 21 Apr. 79 (1 m); XM,
22 Apr. 79 (1 m); WC, 27 Apr, 77 (12 m, 2 f), 8 May 78 (4 m), 25 May 78 (3 m), 5 June

19(1): 1-63, 1980(81)

13

79 (3 m); WC, 27 Apr. 77 (1 m, NSM; 1 m, 2 f, CSL), 29 Apr. 77 (1 m, NSM), 8 May
78 (1 m), 25 May 78 (2 m), 30 May 66 (1 f), 13 June 77 (2 m, 1 f); Seep Canyon, 28
Apr. 73 (4 m, 1 f, NSM); 1 mi.W. RD Summit, 28 Apr. 77 (1 f, NSM); Pine Creek, 28
Apr. 75 (1 m, NSM); KC, 1 May 7 1 (1 m, 1 f); KC, 20 mi. W. U.S. 95, 2 May 77 (1 m, 1
f); Charleston Mts., 12-14 May 34 (6m, 1 f, LACM); 0.5 mi. E. MS, 13 May 79 (1 m).

ADDITIONAL RECORDS: CC, 8 Apr. 77 (RB), 18 May 78 (s); Pine Creek, 2
May 77 (s); Oak Creek Canyon, 11 May 78 (s); KC Ski Run, 13 May 78 (s); LC, 29
May 50 (LACM);KC Campground, 31 May 77 (s); WC,6 June 78 (s), 11 June 79, 13
June 77 (s).

LARVAL FOODPLANTS: Quercus gambelii Nutt. (Fagaceae): WC, 27 Apr. 77
(ovip. leaf buds). Quercus turbinella Greene (Fagaceae): CC, 10 May 78 (ovip. at
base of leaf).

ADULT RESOURCES: Erodium cicutarium (Geraniaceae); Eriodictyon angus-
tifolium (Hydrophyllaceae); Ribes aureum (Saxifragaceae); adults often visit water.
Erynnis funeralis (Scudder & Burgess).

Uncommon and local especially in desert canyons at elevations below 4400'. The
flight period consists of a relatively large spring brood (mid March to early June), a
small summer brood (late June to late July) and a small fall brood (late August to late
October). The summer and fall broods may not occur in every year and possibly
depend on summer rainfall.

SPECIMENS EXAMINED: (33) 9 mi. W. Davis Dam, 17 Mar. 77 (1, NSM; 1 f,
CSL), 31 Mar. 77 (1 m, 2 f, NSM; 1 m, CSL), 8 Apr. 77 (1 m); Newberry Mts., 23
Mar. 67 (1 m, ex UNLV); GC, 8 Apr. 77 (1 m, NSM; 1 m), 20 Apr. 78 (1 f), 5 May 77 (1
m, CSL; 1 m), 3 July 78 (1 f), 21 July 78 (2 m), 13 Sept. 78 (1 f), 27 Sept. 77 (1 m), 12
Oct. 77 (1 m); XM, 12 Apr. 78 (1 m), 20 Apr. 78 (1 f), 5 May 77 (Im), 23 May 78 (1 f),
20 Sept. 78 (1 m); Nelson, 19 Apr. 64 (1 m, 1 f); CC, 18 May 78 (1 m, CSL), 26 June
78 (2 m); Las Vegas, 27 June 68 (1 m); WC, 20 July 78 (1 m); CN, 10 Sept. 78 (1 f,
CSL); LO, 23 Oct. 77 (1 f).

ADDITIONAL RECORDS: RD, 19 Mar. 72 (s); XM, 25 Mar. 79, 15 Apr. 78, 21
July 78, 27 Sept. 77 (s); PV, 4 Apr. 77 (s); GC, 12 Apr. 78, 15 Apr. 78, 12 May 78, 1
June 78, 21 Aug. 78, 20 Sept. 78, 26 Sept. 78, 4 Oct. 77 (s); Sacatone Wash, 15 Apr.
78, 20 Sept. 78 (s); 9 mi. W. Davis Dam, 15 Apr. 78 (s); CP Rd., 0.6 mi. S. Pahrump
Rd., 22 Apr. 78 (s); CC, 10 May 78, 17-18 May 78, 20 May 78, 7 June 78 (s); Oak
Creek Canyon, 1 1 May 78 (s); Willow Springs, 19 Sept. 78 (s); Pine Creek, 22 Sept.
78 (s).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Amsinc-
kia tessellata (Boraginaceae); Viguiera deltoidea, Bebbia juncea, Chrysothamnus sp.,
Senecio douglasii, Tetradymia axillaris, Tetradymia canescens (Asteraceae).
Erynnis meridianus meridianus Beil.

Locally common in association with oaks above 3000'. It flies from mid March to
early August with two September records. The number of broods apparently varies
with location with possibly 3 at the lower elevations and usually one at higher
elevations. We see no great differences in phenotype between early and late
individuals although the earliest spring specimens tend to be smaller than later
specimens.

SPECIMENS EXAMINED: (143) Numerous records from nearly throughout
the Spring Range in the vicinity of oaks from 19 Mar. (72, Ash Spring, 1 m, 2 f) to 19
Sept. (78, Pine Creek, 2 m). Most specimens are from June. There are several

14

J. Res. Lepid.

records for the Virgin Mts. and one for GC on 13 Sept. 78 (1 m, CSL; 1 f).

ADULT RESOURCES: Ranunculus cymbalaria (Ranunculaceae)/ Rorippa
nasturtium-aquaticum, Erysimum asperum (Brassicaceae); Eriodictyon angustifol-
ium (Hydrophyllaceae); Ribes cereum, Ribes aureum (Saxifragaceae); Cercis occi-
dentalis, Melilotus albus (Fabaceae); Viguiera multiflora, Solidago spectabilis,
Erigeron sp., Senecio douglasii, Senecio multilobatus, Cirsium sp., Taraxacum
officinale (Asteraceae).

Erynnis telemachus Bums.

This species is known at present only from the Virgin Mountains flying in a single
brood in May and early June.

SPECIMENS EXAMINED: (29) CC, 10 May 78 (2 m; 2 m, CSL), 18 May 78 (1
m; 1 m, CSL), 20 May 78 (1 m), 20 May 79 (12 m, 1 f), 29 May 78 (4 m), 7 June 78 (1
m), 8 June 78 (1 m, CSL); Virgin Mts., 30 May 74 (3 m, NSM).

ADDITIONAL RECORDS: None.

Chiomara asychis georgina (Reakirt).

The only record is a stray at Com Creek.

SPECIMENS EXAMINED: (1) CN, 26 Sept. 65 (1 m, NSM).
ADDITIONAL RECORDS: None.

Systasea zampa (Edwards).

A small population occurs in the southern portion of the county in the Newberry
Mountains. There are no other Nevada records.

SPECIMENS EXAMINED: (8) GC, 15 Apr. 78 (1 m), 20 Apr. 78 (1 m, CSL), 21
July 78 (1 m); 9 mi. W. Davis Dam, 20 Apr. 78 (1 m), 26 Sept. 78 (1 m, NSM); XM, 23
May 78 (1 m), 21 July 78 (1 m); Sacatone Wash, 20 Sept. 78 (1, CSL).
ADDITIONAL RECORDS: 9 mi. W. Davis Dam, 3 May 78 (s).

ADULT RESOURCES: Monardella sp. (Lamiaceae); Bebbia juncea (Asteraceae).

Thorybes pylades (Scudder).

This skipper is known for Clark County from 2 records from the Spring Range.
SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: KC, 29 June 63 (KR); Pine Creek, 30 June 63 (KR).
Polygonus leo arizonensis (Skinner).

Three fall records are the only occurrences of this stray in Nevada.
SPECIMENS EXAMINED: (2) 10 mi. W. of Searchlight, 21 Sept. 72 (2 m,
NSM).

ADDITIONAL RECORDS: GC, 13 Sept. 78 (s); LO, 9 Oct. 77 (s).

ADULT RESOURCES: Bebbia juncea (Asteraceae).

Epargyreus clams huachuca (Dixon).

The only state specimens are from the Virgin Mountains where they associate with
Robinia neomexicana from mid May to late June. Individuals of this population
average smaller but are otherwise quite indistinguishable from southern Arizona
huachuca.

SPECIMENS EXAMINED: (14) CC, 20 May 79 (1 m), 29 May 78 (1 m), 7 June
78 (1 m), 26 June 78 (6 m); Virgin Mts., 30 May 74 (5 m, NSM).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Robinia neomexicana (Fabaceae); males visit water.

19(1): 1-63, 1980(81)

15

Family Papilionidae

Battus philenor philenor (Linnaeus).

There are several records of this species from April to October mostly in the low
valleys. Specimens are generally quite worn. It is most likely non-resident but
occurring as a stray from Arizona.

SPECIMENS EXAMINED: (4) CN, 18 July 65 (1 m); Las Vegas, 7 Sept. 68 (1
f); Stewart Springs, 29 Sept. 66 (2 m, NSM).

ADDITIONAL RECORDS: PV, 24 Apr. 78 (s); GC, 5 May 77, 21 Aug. 78 (s);
Las Vegas, 14 June 77, 26 June 77, 23 Aug. 66 (s); KC Ski Run, 25 July 77 (s); CN,
31 July 65 (RES); KC, 7600', 5 Aug. 72 (JFL); Roger’s Spring, 10 Oct. 64 (PJH).

ADULT RESOURCES: Cirsium sp. (Asteraceae).

Papilio bairdii bairdii Edwards.

Rare in the Spring Range; the 4 known specimens are probably from Kyle Canyon
where its probable foodplant {Artemesia dmcunculus) is common. All records are in
mid June.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: (all fide JFE) Mt. Charleston, 11-12 June 35 (2 f,
Rudkin colln.), 15 June 40 (1 m, Calif. Acad. Sci.), KC, no date (T. Davies colln.).
Papilio rudkini Comstock.

Locally common in the upper desert and occasional elsewhere in towns, cultivated
areas and once to 9000'. There are two main broods in March and April and in
September and October with scattered individuals at other times of the year
(records from mid February to late November). The fall brood may be quite small or
lacking, depending on summer rainfall. The dark form “clarki” comprises about
14% of the local population (approximate estimate, no statistical data available).

SPECIMENS EXAMINED: (311 including 45 “clarki”) Records extend from
12 Feb. (77, KC, 1 mi. W. U.S. 95, 1 m) to 21 Nov. (77, Las Vegas, 1 m, UNLV).
'There are very few records (12 specimens) from June through August.

LARVAL FOODPLANTS: Thamnosma montana Torr. & Frem. (Rutaceae):
CC, 27 Apr. 78 (mature larva, dark form, emerged as form “clarki”, CSL); CP Rd.,
1.5 mi. S. Pahmmp Rd., 11 May 78 (ovip. upper leaf surface, 10:00 PST); CC, 20
May 79 (mature larva); RC, 27 May 78 (ovip. on stem at leaf base, 11:00 PST); CP
Rd., 0. 6-1.8 mi. S. Pahrump Rd., 28 May 78 (ovip. on stem at leaf base, 09:45 PST,
11:20 PST); Pahrump Rd., 3 mi. W, Blue Diamond turnoff, 26 Sept. 77 (ovip. on
stem, 09:15 PST, 09:30 PST).

ADULT RESOURCES: Delphinium parishii (Ranunculaceae); Dichlostemma
pulchellum (Amaryllidaceae); Arabis sp. (Brassicaceae); Eriogonum plumatella
(Polygonaceae); Asclepias subulata (Asclepiadaceae); Phacelia vallis-mortae, Erio-
dictyon angustifolium (Hydrophyllaceae); Heliotropium curassavicum, Amsinckia
tessellata, (Boraginaceae); Penstamon palmeri (Scrophulariaceae); Verbena good-
dingii (Verbenaceae); Salazaria mexicana, Salvia dorrii (Lamiaceae); Rihes cereum
(Saxifragaceae); Rosa woodsii (Rosaceae); Acacia greggii, Medicago sativa, Melilotus
albus, Lotus rigidus, Astragalus lentiginosus (Fabaceae); Thamnosma montana
(Rutaceae); Encelia farinosa, Encelia virginensis, Bebbia juncea, Haplopappus sp.,
Chrysothamnus sp., Senecio douglasii, Cirsium sp., Stephanomeria sp., Taraxacum
officinale (Asteraceae).

16

J. Res. Lepid.

Papilio indra martini Emmel & Emmel.

Locally common in the Spring Range where there are two broods from mid March
to late April and late June to mid July. The Spring Range population varies from
individuals with wide postmedian bands resembling P. L nevademis Emmel &
Emmel to those with a near absence of the postmedian band in the hindwing as is
typical of martini (see €dso Emmel & Emmel 197 4). There are additional populations of
indra known from the Newberry, Mormon and Virgin mountains which differ from
these. The records (1963 summ.) of P. indra nearfordi Comstock & Martin refer to
the Spring Range population. Tyler’s (1975) inclusion of fordi in Nevada is also in
error.

SPECIMENS EXAMINED: (6) Ash Springs, 30 June 69 (2 m, 1 f, LACM), 6
July 62 (3 m).

ADDITIONAL RECORDS: Calico Basin, 19 Mar, 62 (JFL); Willow Spring, 19
Apr. 66 (JFL), 17 June 78 (DM); White Rock Springs, 19 Apr, 75 (JB); 21 Apr. 79
(s),^,l mi. E, Lovell Wash, RD Summit Rd., 24 Apr. 77 (s); Ash Springs, 25 June 68
(JFE, OS); north of Deer Creek Campground, 29 June 63 (KR); Lovell Canyon, 10
July 75 (CSL).

LARVAL FOODPLANTS: Lomatium parryi (Wats.) Macbr. (Apiaceae): Calico
Basin, 28 Mar. 72 (larvae, JFL); Ash Spring, 25 June 68 (larvae, JFE, OS).

ADULT RESOURCES: Lomatium parryi (Apiaceae); Cirsium sp. (Asteraceae).

Papilio rutulus rutulus Lucas.

Fairly common is the Spring Range from 6000' to 9000' especially in Kyle Canyon.
It flies from late May to early August in a single brood.

SPECIMENS EXAMINED: (58) KC, 20 mi. W. U.S. 95, 31 May 77 (1 m), 14
June 77 (1 m), 28 June 77 (1 m), 28 June 78 (1 f), 5 July 78 (1 m); above Cathedral
Rock Campground, 17 June 77 (1 m, 1 f); KC Ski Run, 17 June 77 (1 m, NSM); 20
June 79 (1 m, 1 f), 22 June 78 (1 f), 26 June 79 (2 m, 2 f), 2 July 78 (2 m), 5 July 77 (1
f), 7 July 77 (2 m, CSL), 10 July 75 (2 m), 18 July 79 (1 m); KC Campground, 20 June
79 (1 m), 24 June 79 (2 m), 26 June 79 (2 m), 7 July 77 (1 m); KC, 7650', 27 June 65
(1 f); Charleston Park, 30 June 77 (3 m, CSL; 2 m, 2 f, NSM); Little Falls, 5 July 65 (6
m, 3 f), 13 July 65 (2 m, 1 f), 20 July 65 (1 m); KC, 6050', 5 July 65 (1 m); end KC, 13
July 65 (1 m); KC, 7500', 15 July 74 (2 m, 2 f); Rainbow Canyon, 20 July 65 (1 f); KC,
9000', 25 July 65 (1 f).

ADDITIONAL RECORDS: Numerous records for KC, earliest on 29 May 63
(JFL); latest on 1 Aug. 67 (s). One record for west slope in Lovell Wash at Spring
Mountain Youth Camp on 22 June 77 (s).

LARVAL FOODPLANTS: Populus tremuloides Michx. (Salicaceae): KC Ski
Run, 26 June 79 (ovip. lower surface of leaf near midrib, 10:00 PST); 5 July 77
(ovip.); 10 July 79 (ovip. upper leaf surface, 08:50 PST).

ADULT RESOURCES: Ipomopsis aggregata (Polemoniaceae); Penstamon
utahensis (Scrophulariaceae); Erysimum asperum (Brassicaceae); Cirsium sp..
Taraxacum officinale (Asteraceae); visit water on occasion.

Papilio multicaudatus Kirby.

The only known colony of this species in Clark County is in the Virgin Mountains.
Specimens have wide dark borders similar to Arizona material.

SPECIMENS EXAMINED: (18) CC, 27 Apr. 78 (1 m, CSL), 17»18 May 78 (2
m, 1 f; 2 m, CSL; 2 m, NSM), 20 May 79 (3 m), 29 May 78 (2 m, 2 f), 7-8 June 78 (2 m;
1 f, CSL).

19(1): 1~63, 1980(81)

17

ADDITIONAL RECORDS: CC, 10 May 78, 20 May 78, 26 June 78 (s).
ADULT RESOURCES: Males occasionally visit water.

Family Pieridae

Neophasia menapia (Felder & Felder).

Rare but apparently not uncommon in some years in the Spring Range above
7400'. The few records indicate a single brood flying from late July to late August.
SPECIMENS EXAMINED: (1) KC Ski Run, 1 Aug. 77 (1 m).

ADDITIONAL RECORDS: KC, 7400', 29 July 62 (JFL), 31 Aug. 62 (JFL);
Charleston Mts., 6 Aug. 61 (RES); KC, 7800', 8 Aug. 62 (JFL), 12 Aug. 63 (JFL), 18
Aug. 63 (JFL); KC, 8 Aug. 74 (JAS).

ADULT RESOURCES: Argemone minita (Papaveraceae).

Pieris beckeru beckerii Edwards.

Uncommon, but sometimes common, usually at elevations between 2900' and
7200'. Its flight period is from late March to mid November, At least 4 broods
appear to be involved in some years along some upper desert washes; single broods
may be raised in other areas. Summer rainfaU may be regulatory as suggested by
Emm el and Emmei (1973).

SPECIMENS EXAMINED: (121) Numerous records for many locations at
middle elevations in much of the county with dates between 3 Mar. (78, KC, 6.6 mi.
W., 0.9 mi. N. U.S. 95, 1) and 13 Nov. (77, KC, 9.3 mi. W. U.S. 95, 1 m). The species
is most common in the Spring Range and rare in the Newberry and Virgin
mountains. A few records also exist for the Las Vegas and Moapa valleys.

LARVAL FOODPLANTS: Stanleya pinnata (Pursh) Britton (Brassicaceae);
CP Rd., 1.8 mi. S. Pahramp Rd., 29 Apr. 78 (ovip. main stem, 1/2 way up at leaf
base, 11:15 PST); 1 mi. W. WC, 8 May 78 (ovip. at apex of flower bud, 11:30 PST);
Trout Canyon Rd., 0.8 mi. W. Spring Mt. Youth Camp, 14 June 77 (ovip. lower leaf
surface); KC, 15 mi. W. U.S. 95, 23 June 77 (ovip,). Stanleya elata Jones
(Brassicaceae): KC, 9.3 mi. W. U.S. 95, 1 May 77 (ovip. on main stem),

ADULT RESOURCES: Clematis ligusticifolid (Ranunculaceae); Stanleya pin-
nata, Arabis sp. (Brassicaceae); Eriogonum fasckulatum, Eriogonum microthecum
(Polygonaceae); Phacelia fremontii, Eriodictyon angustifolium (Hydrophyliaceae);
Marrubium vulgare, Salvia dorrii (Lamiaceae); Astragalus lentiginosus (Fabaceae);
Teradymia canescens (Asteraceae).

Pieris sisymbru elivata (Barnes & Benjamin).

Uncommon, especially at middle elevations between 4000' and 6500', in most of
the county. The single brood flies from mid March to mid May with a single record in
mid June. All southern Nevada females examined were at least slightly tinged with
yellowish,

SPECIMENS EXAMINED: (100) Records extend from 18 Mar. (72, XM, JFL)
to 17 May (78, CC, s). One record on 14 June 77 (KC, 17.1 mi. W. U.S. 95, s).

ADULT RESOURCES: Arabis sp. (Brassicaceae).

Pieris protodke protodice Boisduval & Le Conte.

Common to abundant at low elevations and less common at the higher elevations
to at least 10,000'. The flight period extends throughout the year. Fresh individuals
are found on warm days in midwinter.

The small spring form “vemalis” flies at low elevations from January to April and
apparent intermediates with the typical form occurring into May. Specimens from

18

J. Res. Lepid.

October to December are large but more heavily marked below than the summer
form.

SPECIMENS EXAMINED: (391) Specimens are from many localities through-
out the county except the highest elevations.

LARVAL FOODPLANTS: Lepidium lasiocarpum Nutt. (Brassicaceae): CP
Rd., 1.5 mi. S. Pahrump Rd., 29 Apr. 78 (ovip., upper leaf surface, 09:30 PST).
Descurainia pinnata (Walt.) Britt, ssp. glabra (Woot. & Standi.) Detl. (Bras-
sicaceae): U.S. 93, 13 mi. N. 1-15, 14 May 78 (ovip., 09:15 PST). Descurainia
pinnata (Walt) Britt, ssp. halictorum (Ckll.) Delt. (Brassicaceae): WC, 8 May 78
(ovip. 2x, lower leaf surface of upper leaves, 13:30 PST).

ADULT RESOURCES: Sphaeralcea grossulariaefoUa (Malvaceae); Lmum lewisii
(Linaceae); Tamarix pentandra (Tamaricaceae); Stanleya pinnata, Sisymbrium
altissimum (Brassicaceae); Eriogonum corymbosum, Eriogonum fasciculatum (Poly-
gonaceae); Menodora spinescens (Oleaceae); Phacelia fremontii, Eriodictyon angus-
dfolium (Hydrophyllaceae); Heliotropium curassavicum (Boraginaceae); Marmfeium
vulgare, Salvia dorrii (Lamiaceae); Fallugia paradoxa (Rosaceae); Acacia greggii,
Prosopis glandulosa, Prosopis pubescens, Medicago sativa, Melilotus albus, Melilotus
officinalis, Astragalus lentiginosus (Fabaceae); Viguiera deltoidea, Viguiera multi-
flora, Helianthus annuus, Encelia virginensis, Bebbia juncea, Baileya multiradiata,
Baileya pleniradiata, Chaenacds fremontii, Haplopappus sp., Chrysothamnus sp.,
Erigeron sp., Senecio douglasii, Senecio multilobatus, Pluchea sericea, Cirsium sp.,
Taraxacum officinale (Asteraceae).

Pieris rapae (Linnaeus).

Common in urban areas, less common in agricultural areas and small, recently
established (since 1968) wild colonies at higher elevations at Willow and Cold
creeks and in Kyle Canyon. A wild, possibly fugitive, colony was found in the upper
desert near Cottonwood Pass in 1978. There are additional records near desert
springs and in the Virgin and Newberry mountains. The flight period is from late
January to mid December at low elevations and from late April to mid October at
high elevations. There is only one record before 1961 (Las Vegas, 12 Apr. 60, PJH).

SPECIMENS EXAMINED: (104) Many records especially in Las Vegas Valley
throughout most of the year.

LARVAL FOODPLANTS: Caulanthus cooperi (S. Wats.) Pays (Brassicaceae);
CP Rd., 1.8 mi. S. Pahrump Rd.,29 Apr. 78 (ovip., 2x, lower leaf surface, just to one
side of mid rib, 12:00 PST). Descurainia sophia (L.) Webb (Brassicaceae): MS, 11
May 78 (ovip,, 3x, lower leaf surface, 12:30 PST). Rorippa nasturtium-aquaticum
(L.) Schinz &Thell (Brassicaceae): WC, 1 July 77 (ovip. lower leaf surface). Brossica
oleracea L. var. botrytis L. (Brassicaceae): Las Vegas (many records of ovip. and
larvae). Brassica oleracea L. var. capitata L. (Brassicaceae): Las Vegas (many
records of ovip. and larvae).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Sisymbrium
altissimum, Rorippa nasturtium-aquadcum (Brassicaceae); Phacelia fremontii,
Eriodictyon angustifolium (Hydrophyllaceae); AmsmcAia tessellata (Boraginaceae);
Medicago sativa, Melilotus albus, Astragalus sp. (Fabaceae); Chaenactis fremontii,
Pluchea sericea, Taraxacum officinale (Asteraceae).

Colias eurytheme Boisduval.

Abundant in agricultural areas and occasional in the desert and near springs at low
elevations and fairly common in open areas at high elevations. Low elevation records

19(1): 1-63, 1980(81)

19

extend from mid February to mid December and those at high elevations from late
April to mid November. Desert records are for spring and fall.

Early spring ^arch, April) and late fall (October, November) specimens are
smaller, paler and usually more heavily dusted with black and have more extensive
pink markings beneath than summer specimens. This indicates that the form
“ariadne” is a valid form in this area and is not a hybrid between this species and C.
philodice Godart as sometimes suggested (Klots 1951, Brown, et al. 1957). Colias
philodice is very rare in southern Nevada. The form “alba” comprises 31% of the
local female population (approximate estimate, no statistical data available).

SPECIMENS EXAMINED: (615) A large number of records from numerous
locations to 1 1 ,000' with records from 1 1 Feb. to 19 Dec. It probably can be found at
sometime as either a resident or stray in all parts of the county.

LARVAL FOODPLANTS: Medicago sativa L. (Fabaceae); LO, 4 July 77
(ovip.). Trifolium repens L. (Fabaceae): WC, 8 May 78 (ovip., lower leaf surface,
12:20 PST).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Linum lewisii
(Linaceae); Tamarix pentandra (Tamaricaceae); Argemone minita (Papaveraceae);
Stanleya pinnata, Sisymbrium altissimum, Erysimum asperum (Brassicaceae);
Eriogonum wrightii, Polygonum lapathifolium (Polygonaceae); Arctostaphylos pun-
gens (Ericaceae); Convolvulus sp. (Convolvulaceae); Apogynum androsaemifolium
(Apocynaceae); Sarcostemma hirtellum (Asclepiadaceae); Eriodictyon angustifolium
(Hydrophyllaceae); Heliotropium curassavicum (Boraginaceae); Lycium sp. (So-
lanaceae); Verbena gooddingU (Verbenaceae); FaUugia paradoxa, Rosa woodsU
(Rosaceae); Lupinus sp. Medicago sativa, Melilotus albus, Melilotus officinalis,
Robima neomexicana (Fabaceae); Viguiera multiflora, Helianthus annuus, Bebbia
juncea, Baikya multiradiata, Eriophyllum wallacei, Pectis papposa, Haplopappus
sp., Chrysothamnus sp., Machaeranthera tortifolia, Baccharis sp., Senecio douglasii,
TetraAymia canescens, Cirsium sp., Taraxacum officinale (Asteraceae).

Colias philodice philodice Godart.

A stray from Las Vegas is the only specimen from the county.

SPECIMENS EXAMINED: (1) Las Vegas, 20 Oct. 1974 (1 f, ex UNLV).

ADDITIONAL RECORDS: None.

Colias alexandra edwardsii Edwards.

A small colony of this specie® was discovered in the southern portion of the Spring
Range in 1978. The only other record for the county is a fall record from Kyle
Canyon.

SPECIMENS EXAMINED: (5) CP Rd., 0.6-1. 8 mi. S. Pahrump Rd., 22 Apr. 78
(1 m), 29 Apr. 78 (1 m), 6 May 78 (1 m, CSL), 11 May 78 (1 m, apparent hybrid x C.
eurytheme or C. philodice); KC, 9.3 mi. W. U.S. 95, 15 Oct. 77 (1 f).

ADDITIONAL RECORDS: CP Rd., 0.6-1.5 mi. S. Pahrump Rd., 16 Apr. 79, 29
Apr. 79, 19 May 78 (s).

Colias cesonia (Stoll).

Usually rare at elevations to 7600' with records from mid February to mid
November. In most years it may be a stray but it was common and produced at least
two broods in the Newberry Mountains in 1978.

SPECIMENS EXAMINED: (47) GC, 15 Apr. 78 (1 f), 21 Apr. 78 (1 f, CSL), ,3
May 78 (1 f), 23 May 78 (5 m, 1 f), 1 June 78 (8 m,NSM; 3 m, CSL); 3 July 78 (1 m, 2

20

J. Res. Lepid.

f), 21 July 78 (2 m, 2 f), 4 Aug. 78 (2 m); CP Rd., 0.6=1, 5 mi. S.Pahmmp Rd., 16 Apr.

77 (1 f), 28 May 78 (1 m); Bowman’s Reservoir, 7 May 78 (1 m, CSL); Bridge
Canyon, 12 May 78 (1 f); XM, 1 June 78 (1 m, CSL), 21 July 78 (2 m); WC, 22 June

78 (2 m, CSL), 1 July 77 (1 m), 4 July 62 (1 m), 20 July 78 (2 m); WC, 4 July 62 (1 f),
20 July 78 (2 m); KC Ski Run, 23 July 78 (1 m), 25 July 78 (1 m, NSM).

ADDITIONAL RECORDS: GC, 20 Feb. 78, 20 Apr. 78, 12 May 78 (s); 1 mi. W.
MS, 26 Mar, 68, 11 May 78 (s); Sacatone Wash, 15 Apr. 78 (s); PV, 19 Apr. 78, 28
Apr. 78 (s); 9 mi. W. Davis Dam, 20 Apr. 78 (s); CP Rd., 0.6-1. 8 mi, S. Pahrump Rd.,
22 Apr. 78, 11 May 78 (s); Hidden Valley, 27 Apr. 78 (s); Pine Creek, 2 May 77, 22
May 78 (s); Willow Springs, 2 May 78 (s); CC, 10 May 78, 29 May 78, 7 June 78 (s);
LO, 17 May 78, 11 Nov. 77 (s); Bunkerville, 17 May 1978 (s);XM, 23May 78,3 July
78 (s); WC, 11 June 79 (s); Lovell Wash, 17 June 62, 18 July 62 (s); Charleston Park,
25 June 68 (JFE, OS); Ash Springs, 6 July 62 (s); Echo Canyon, 23 July 78 (s); KC
Ski Run, 25 July 78 (s); Charleston Mts., 31 July 65 (JL, RES); WC, 30 Aug. 77 (s);
Jean, 2 Sept. 78 (s); RR, 28 Oct. 77 (s).

ADULT RESOURCES; Clematis ligusticifolia (Ranunculaceae); Amsonia
tomentosa (Apocynaceae); Eriodictyon angustifolium (Hydrophyllaceae); Marrub-
ium vulgare, Monardella sp. (Lamiaceae); Bebbia juncea, Tetradymia canescens,
Cirsium sp., (Asteraceae).

Phoebis sennae marcellina (Cramer).

Scattered records of strays are the only ones for the state. A relatively good flight
was noted in the spring of 1978.

SPECIMENS EXAMINED: (5) Willow Spring, 10 May 66 (1, JFL); GC, 1 June
78 (1 m, CSL), 3 July 78 (1 f); OV, 10 July 68 (1, JFL); KC, 7400', 18 Aug. 63 (1,
JFL).

ADDITIONAL RECORDS: PV, 10 Apr. 78, 24 Apr. 78, 19 May 79 (s); XM, 12
Apr. 78 (s); GC, 12 Apr. 78 (s); Hidden Valley, 13 Apr. 78 (CSL, s); CC, 13 Apr. 78
(s); WC, 22 June 78 (s).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Prosopis juliflora
(Fabaceae).

Eurema mexicana (Boisduval).

Four specimens and a few sightings represent the only state records. This species
may occasionally establish itself as a breeding insect in southern Nevada but this
has yet to be shown.

SPECIMENS EXAMINED: (4) Willow Springs, 10 May 66 (3, JFL); 21 May 66
(1, JFL).

ADDITIONAL RECORDS: 9 mi. W. Davis Dam, 23 Mar. 78, 20 Apr. 78 (s); I-
15, 2 mi. N. Dry Lake, 19 Apr. 75 (s).

Eurema nicippe (Cramer).

Common in a variety of habitats to 7600' but mostly below 4000' in desert washes
and the vicinity of springs. Records extend from late Januaiy to mid December
involving possibly 4 broods in some areas. Open desert records indicate that there
are 2 usual broods, one in early spring and one in fall.

Specimens from April through September are mostly yellow beneath; those from
February and October to December represent the reddish winter form.

SPECIMENS EXAMINED: (126) Numerous records from the lower canyons
from 29 Jan. (78, Las Vegas, s) to 11 Dec. (77, GC, 1 m, 3 f). Less common at higher
elevations from 10 Apr. (68, MS, s) to 16 Nov. (77, KC, 20 mi. W. U.S. 95, s).

19(1): 1-63, 1980(81)

21

LARVAL FOODPLANTS: Cassia armata Wats. (Fabaceae): 9 mi. W. Davis
Dam, 3 May 78 (ovip., lower surface of flower bud, 09:00 PST).

ADULT RESOURCES: Sarcostemma hirtellum (Asclepiadaceae); Physalis cras-
sifolia (Solanaceae); Monardella sp. (Lamiaceae); Medicago sativa, Melilotus albus
(Fabaceae); Viguiera deltoidea, Helianthus annum, Bebbia juncea, Senecio doug-
lasii, Cirsium sp. (Asteraceae).

Nathalis iole Boisduval,

Common and widespread in many habitats to 9000', Its flight period extends from
early February to mid December at low elevations and from late April to late
October above 6000'. At least 4 broods appear to be involved in some areas but
possibly only 2 in low desert.

Very early spring (February) specimens represent the winter form. The lower
surface of both sexes of this form is very heavily overscaled with black nearly
obscuring the underlying paler scales. The females are pale yellow with a reduction
of the black markings on the inner margin of the fore wing and on the hind wing. The
yellow hindwing of the winter form contrasting with orangish hindwings of the
summer form indicates, perhaps, that southern Nevada material belongs to the
central segregate of this species (see Clench 197 6). The validity of these color forms
is yet to be demonstrated.

SPECIMENS EXAMINED: (107) Records are for much of the county with the
earliest on 6 Feb. (77, 0.5 mi. S.W. Laughlin, 1 m, 1 f) and latest on 12 Dec. (77,'rule
Springs, s).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Eriogonum fascicu
latum (Polygonaceae); Eriodictyon angustifolium (Hydrophyllaceae); Heliotropium
curassavicum (Boraginaceae); Viguiera multiflora, Chaenactis fremontii, Senecio
douglasii, Taraxacum officinale (Asteraceae).

Anthocaris pima Edwards.

Common in some years, rare or absent in others in desert below 5200'. The flight
period extends from mid Februaiy to early May. There is one main brood and
possibly a partial second brood.

SPECIMENS EXAMINED: (123) Numerous records at low elevations between
11 Feb. (75, U.S. 93, 8 mi. N. 1-15, s) and 4 May (78, Nipton Pass area, 2).

ADULT RESOURCES: Descuriarm pinnata (Brassicaceae); Phacelia fremontii
(Hydrophyllaceae); Amsinckia tessellata, Cryptantha sp. (Boraginaceae).
Anthocaris sara thoosa Scudder.

Uncommon mostly between 4000' and 7000' but it occasionally occurs at lower
elevations. The single brood flies from mid February to late May with a peak in late
March and April.

The material from southern Clark County is identical with California populations.
'The ventral maculation of Virgin Mountains material tends to be heavier as is
southern Utah material. We agree with Emmel & Emmel (1973) on the separation of
thoosa from Arizona A. s. inghami Gunder especially on the basis, especially, of
ventral coloration.

SPECIMENS EXAMINED: (160) Pine Creek, 19 Mar. 64 (1 m, 1 f), 23 Mar. 77
(1 m, NSM); KC, 17.1 mi. W. U. S. 95, 22 Mar. 77 (1 m), 11 Apr. 77 (1 f); XM, 23
Mar. 78 (2 m,NSM; 2 m, 1 f), 25 Mar. 79 (21 m, 15 f), 12 Apr. 78 (1 m); Wilson Pass,
25 Mar. 78 (1 f); CC, 29 Mar. 71 (1 m, 1 f, NSM), 8 Apr. 79 (14 m, 26 f), 13 Apr. 74 (1
f, NSM), 13 Apr. 78 (8 m, 3 f), 14 Apr. 74 (3 m, NSM), 19 Apr. 75 (2 f, NSM), 27 Apr.

22

J. Res. Lepid.

78 (3 m, NSM; 4 m), 10 May 78 (1 m, 1 f, NSM; 1 m); Willow Springs, 29 Mar. 78 (1
m); 2 mi. N. RC, 5 Apr. 78 (1 m, NSM; 1 m), 1 1 Apr. 78 (1 m, NSM), 15 Apr, 79 (6 m,
1 f), 4 May 78 (1 m); Nipton Pass, 12 Apr, 69 (6 m, 5 f, NSM); Lost Creek, 20 Apr. 77
(1 m, NSM); Potosi Mt. Rd. atPahrump Rd., 20 Apr, 77 (1 f); RD Summit Rd., 1.5-
2.1 mi. E. Lovell Wash, 24 Apr. 77 (2 f), 28 Apr. 77 (1 m, 2 f); Lovell Wash, 6.8 mi.N.
Pahrump Rd., 24 Apr. 77 (1 f); WC, 27 Apr. 77 (2 f), 25 May 78 (1 f); Seep Canyon,
28 Apr. 73 (2 f, NSM); 1 mi. W. RD Summit, 28 Apr. 77 (5 m, 1 f, NSM); Cold Creek,
8 May 78 (1 m).

ADDITIONAL RECORDS: Several records from middle elevations with the
earliest record on 16 Feb. (77, Spring Mt. Ranch, s).

ADULT RESOURCES: Arctostaphylos pungens (Eriaceae); Phacelia fremontii
(Hydrophyllaceae); Amsinckia tessellata (Boraginaceae).

Euchloe hy antis lotta (Beutenmueller).

Uncommon but widespread especially at middle elevations to about 6200'. It flies
in one brood from early March to mid May.

SPECIMENS EXAMINED; (134) Numerous records, usually away from the
valley floors from 5 Mar. (78, PV, 2 f) to 19 May (78, 0.5-1. 5 mi. E. MS, 1 m).

ADULT RESOURCES: Phacelia fremontii (Hydrophyllaceae); Rihes cereum
(Saxifragaceae); Rhus trilobata (Anacardiaceae); Baileya multiradiata, Taraxacum
officinale (Asteraceae),

Family Riodinidae

Apodemia mormo mormo (Felder & Felder).

Common and occasionally locally abundant, especially along roadsides and
washes where Eriogonum inflatum is common, to 6200'. It flies in at least 4 broods
from early February to mid November. It is most common and widespread in spring;
summer and fall broods may reflect rainfall patterns and/or the availability of
alternate foodplants.

The common phenotype is of the nominate race with little tendency towards true
A. m. deserti Barnes & McDunnough although the latter name has been applied to
southern Nevada material. This insect commonly associates with Eriogonum
■inflatum and E. fasciculatum. In fall, a larger and darker phenotype associated with
Eriogonum microthecum occurs at higher elevations. This phenotype occurs
similarly in the desert ranges of California {fide JFE). Whether this represents a
distinct unnamed subspecies needs confirmation.

SPECIMENS EXAMINED: (388) Records for many localities throughout the
county between 4 Feb. (77, Boulder City, AB) to 16. Nov. (77, 1.8 mi. E. Blue
Diamond, 1 m). The large, dark phenotype is known from such locations as MS and
WC between 6 Sept. (69, NSM) and 7 Oct, (77).

LARVAL FOODPLANTS: Eriogonum deflexum Torr. (Polygonaceae): 0.5 mi,
E. Boulder City, 3 Oct. 77 (ovip. at base of flower). Eriogonum plumatella Dur. &
Hilg. war. plumatella Dur. & Hilg. (Polygonaceae): 10 mi. W. Davis Dam, 3 Aug. 77
(larvae on flower head). Eriogonum inflatum Torr. & Frem. (Polygonaceae):
Searchlight, 1 Oct. 68 (larvae, 1968 summ.). Adults also associated with. Eriogonum
fasciculatum (RR, GC), Eriogonum microthecum (MS, WC), Eriogonum wrightii
(Pine Creek) and Eriogonum heermannii (2 mi. N.RC),

ADULT RESOURCES: Sphaeralcea ambigua (Malvaceae); Eriogonum fascicu
latum, Eriogonum heermannii, Eriogonum microthecum, Eriogonum plumatella,
Eriogonum umbellatum, Eriogonum wrightii, Eriogonum sp. (Polygonaceae); Cryp-

19(1): 1=63, 1980(81)

23

tantha sp. (Boraginaceae); Bebbia juncea, Baileya multiradiata, Baileya plenirad-
iata, Chaenactis fremontii, Palafoxia linearis, Acamptopappus shockleyi, Solidago
spectabilis, Chrysothamnus sp., Baccharis sp., Senecio douglasii, Tetradymia
canescens, Stephanomeria sp. (Asteraceae).

Apodemia palmeri marginalis (Skinner).

Locally common in mesquite stands in the valleys. It flies from mid April to mid
October with 3 or possibly 4 broods in favorable years.

Southern Nevada material agrees well with marginalis described from nearby
southern California and is paler and has consistently complete orange borders in
contrast to the darker, supposedly nominate race from southern Arizona, with its
often incomplete orange borders. A review of the species seems in order as the
species was described from just north of Clark County at St. George, Utah.

SPECIMENS EXAMINED: (236) Numerous records between 17 Apr. (77,
Hidden Valley, s) and 13 Oct. (77, PV, 1 m).

LARVAL FOODPLANTS: Prosopis glandulosa Torr. (Fabaceae): PV, 31 July
77 (ovip.). Prosopis pubescens Benth. (Fabaceae): PV, 2 Aug. 77 (ovip.).

ADULT RESOURCES: Heliotropium curassavicum (Boraginaceae); Prosopis
glandulosa, Prosopis pubescens (Fabaceae); Baccharis sp. (Asteraceae).

Calephelis nemesis californica McAlpine,

Rare in fall with all records except one (Virgin Mountains) from the Logandale
area of Moapa Valley in association with Baccharis. There are no other records for
the state.

SPECIMENS EXAMINED: (13) Bowman’s Reservoir, 30 July 77 (1 m); LO, 4
Sept. 66 (1 m), 27 Sept. 78 (2 m), 16 Oct. 77 (3 m, 2 f), 28 Oct. 77 (3 m); Virgin Mts.
12 Sept. 74 (1 f, NSM).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Baccharis sp. (Asteraceae).

Calephelis wrighti (Holland).

Small numbers fr<5m the Newberry Mountains are the only Nevada records. They
probably represent a small breeding colony as its known foodplant, Bebbia juncea,
is common there.

SPECIMENS EXAMINED: (20) GC, 21 July 78 (1 m), 13 Sept. 78 (1 m; 1 m,
CSL), 20 Sept. 78 (1 m, CSL), 26 Sept. 78 (2 m, 3 f, CSL; 1 m, 1 f), 29 Oct. 78 (6 m),
3 Nov. 77 (1 m); Sacatone Wash, 20 Sept. 78 (1 m, CSL; 1 m).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Bebbia juncea (Asteraceae).

Family Lycaenidae

Satyrium behrii behrii (Edwards).

Rare with records between early June and mid July. It may be more widespread
than presently known in the Pinbn-Juniper belt where its foodplant, Purshia is not
uncommon.

SPECIMENS EXAMINED; (17) WC, 5 June 79 (2 m), 1 1 June 79 (9 m, 1 f), 22
June 68 (1 m), 22 June 78 (4 m).

ADDITIONAL RECORDS: MS, 23 June 65 (PJH), 30 June 63 (KR); WC, 13
July 72 (DEA).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Mar-
ruhium vulgare (Lamiaceae).

24

J. Res. LepM.

Ministrymon leda (Edwards).

Locally common in association with Prosopis glandulosa in Paradise Valley near
Las Vegas and scattered records elsewhere. There appear to be 2 broods. Most
individuals are typical leda although there are single spring and fall specimens of
form “ines” from Hidden Valley.

SPECIMENS EXAMINED; (30) Hidden Valley, 13 Apr. 78 (1 f “ines”), 12
July 77 (1 f), 16 Oct. 77 (1 m “ines”); GC, 1 June 78 (2 m), 3 July 78 (1 f); Lovell
Wash, 7.5 mi. N. Pahrump Rd., 22 June 77 (1 f); PV, 9 May 78 (1 m), 24 June 78 (1
m), 18 July 77 (4 m, 4 f), 26 July 77 (2 m, 4 f), 31 July 77 (4 f); 27 June 66 (1 m); KC,
17.1 mi. W. U.S. 95 (1 m); WC, 19 Aug. 77 (1 f).

ADDITIONAL RECORDS: PV, 26 May 79, 3 July 77, 2 Aug. 77 (s).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Eriodictyon
angustifolium (Hydrophyllaceae); Acacia greggii, Prosopis glandulosa (Fabaceae);
Helianthus annuus, Bailey a multiradiata (Asteraceae).

Callophrys fotis fotis (Strecker).

Usually uncommon but in some years exceedingly abundant, at least locally, at
middle elevations from 4000' to 6800', always in association with Cowania
mexicana. The flight period is from late March to mid May. Fresh specimens from
Cold Creek on 13 June 1977 are undoubtedly from a partial second brood perhaps
in response to early June rainfall.

SPECIMENS EXAMINED: (230) GS, 28 Mar. 68 (1, NSM); Lost Creek, 29
Mar. 78 (2 m, 1 f); CC, 8 Apr. 79 (1 m, 1 f); 2 mi. N. RC, 9 Apr. 79 (3 m), 15 Apr. 79 (7
m, 8 f); CP Rd., 0.6-1. 8 mi. S. Pahrump Rd., 16 Apr. 77 (4 f); 16 Apr. 79 (5 m, 7 f);
White Rock Spring, 21 Apr. 79 (1 m, 9 f); WC, 23 Apr. 78 (7 m, 6f), 27 Apr. 77 (30 m,
33 f; 12 m, 13 f, NSM), 29 Apr. 77 (12 m, 2 f, NSM), 8 May 78 (2 m, 2 f), 13 June 77 (1
f); RD Summit Rd., 1,5-2. 1 mi. E. Lovell Wash, 24 Apr. 77 (18 m, 22 f), 28 Apr. 77 (3
m); WC, 27 Apr. 77 (5 m, 2 f), 8 May 78 (1 f); 1 mi. W. RD Summit, 28 Apr. 77 (2 m, 1
f, NSM); KC, 15.7 mi. W. U.S. 95, 1 May 77 (2 m, 1 f); KC, 17.1 mi. W. U.S. 95, 2
May 77 (1 f); LC, 6 May 62 (1 m); Charleston Mts., 12 May 34 (1 f, LACM).

ADDITIONAL RECORDS: GS, 23 Mar. 68 (1968 summ.), 20 Apr. 75 (JB);
RD, 23 Mar. 75 (JB, REW); RC, 24 Mar. 74 (REW), 24 Mar. 75 (JB), 7 Apr. 74
(REW); Potosi Mt. Rd., 6-8 Apr. 68 (REW); CP, 7 Apr. 74 (REW); RD near Calico
Hills, 19 Apr, 75 (JB); White Rock Springs, 19 Apr. 75 (JB); CP Rd., 0.6-1. 5 mi. S.
Pahrump Rd., 20 Apr. 77, 29 Apr. 79 (s); KC, 6800', 3-4 May 75 (1975 summ.); 0.5
mi. E. MS, 13 May 79 (s).

LARVAL FOODPLANTS: Cowania mexicana var. stansburiana (Torr.) Jeps.
(Rosaceae): WC, 27 Apr. 77 (ovip.).

ADULT RESOURCES: Rihes aureum {Saxifragace ae); Arctostaphylos pungens
(Ericaceae); Salix lasiolepis (Salicaceae); Rhus trilobata (Anacardiaceae); Taraxa-
cum officinale (Asteraceae).

Callophrys eryphon eryphon (Boisduval).

One relatively fresh male from the Virgin Mountains is the only county record.
The nearest known Nevada population is in the Wilson Creek Range, Lincoln
County (GTA, CSL).

SPECIMENS EXAMINED: (1) CC, 20 May 76 (1 m, NSM).

ADDITIONAL RECORDS: None.

Callophrys spinetorum (Hewitson).

Uncommon at middle elevations from 4400' to 7000'. It flies in 2 broods from early

19(1): 1-63, 1980(81)

25

April to late May and early June to early August. The earlier brood occurs mainly
below 6000'; later brood is mainly above 6000'. Adults are particularly fond of the
flowers of Eriodictyon. Males commonly perch on ridge-top pinons and junipers.

SPECIMENS EXAMINED: (42) White Rock Springs, 21 Apr. 79 (11 m); RD
Summit Rd., 1. 5-2.1 mi, E. Lovell Wash, 24 Apr. 77 (3 f), 28 Apr. 77 (2 f); 0.5 mi, E.
MS, 13 May 79 (8 m); WC, 25 May 78 (1 m), 5 June 79 (1 m), 1 1 June 79 (1 m), 20
July 78 (1 m, 1 f), 28 July 77 (1 m); Deer Creek Rd., 7000', 3 July 61 (2 m); Lovell
Wash, 6100-6600', 14 July 62 (1 m, 3 f), 18 July 62 (3 m, 3 f).

ADDITIONAL RECORDS: Potosi Mt. Rd., 6 Apr. 74 (REW), 8 Apr. 71
(REW); WC, 27 Apr. 77, 30 May 66 (s); Willow Springs, 10 May 66 (JFL); Mt.
Charleston, 15 June 40 (Shields 1965); RD Summit Rd., 1.5 mi. E. Lovell Wash, 22
June 77 (s); SM, 7000', 23 June 61 (KR); LC, 6000', 23 June 61 (KR); Charleston
Peak, 29 June 35 (Howe 1975); WC, 13 July 72 (DEA), 16 July 72 (JFL); Charleston
Mts., 6 Aug. 61 (1961 summ.).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae) ;Arctostap%/os
pungens (Ericaceae); Eriodictyon angustifolium (Hydrophyllaceae); Salix lasiolepis
(Salicaceae); Rhus trilobata (Anacardiaceae); Tetradymia canescens (Asteraceae).
Callophrys siua siva (Edwards).

Fairly common at the middle elevations in the Pinon-Juniper belt. It flies in 2
broods from late March to late May and early June to late July with some variation

depending on elevation.

Clark County material is assigned to nominate siua on the advice of John Lane. All
populations in the county, however, show some intermediacy with the unnamed
Great Basin brown population occurring farther north.

SPECIMENS EXAMINED; (131) CC, 8 Apr. 79 (2 m), 20 May 79 (1 f); MM, 12
Apr. 69 (1 m, NSM); 2 mi. N. RC, 15 Apr. 79 (4 m, 2 f); CP Rd., 1.8 mi. S. Pahramp
Rd., 16 Apr. 77 (1 f); RD Summit Rd., 1.5-2. 1 mi. E. Lovell Wash, 24 Apr. 77 (1 m, 2
f), 28 Apr. 77 (1 m; 1 f, CSL), 2 June 77 (1 m, 1 f), 22 June 77 (4 m, 7 f), 27 July 77 (1
m); WC, 27 Apr. 77 (1 m, 1 f), 29 Apr. 77 (2 m, 1 f, CSL; 1 f, NSM) 30 May 66 (1 m), 6
June 78 (1 m, 1 f), 13 June 77 (2 f), 22 June 78 (1 m,' 3 f), 4 July 62 (1 f), 28 July 77 (1
m); WC, 27 Apr. 77 (1 f), 5 June 79 (6 m, 3 f), 6 June 78 (1 f), 1 1 June 79 (23 m, 10 f),
22 June 68 (1 f), 22 June 78 (1 f), 1 July 77 (1 m, 1 f); KC, 5680', 1 May 66 (1 m); Pine
Creek, 2 May 77 (1 f); 0.5 mi. E. MS, 13 May 79 (2 f); Charleston Mts., 14 May 34 (1
f, LACM); Potosi Mt. Rd. at Pahrump Rd., 2 June 77(1 m); Trout Canyon Rd., 14
June 77 (5 m, 2 f); KC, 14.5 mi. W. U.S. 95, 14 June 77 (1 m), 20 June 77 (1 m), 20
June 79 (1 m, 1 f), 6 July 77 (2 m); Lovell Wash, 22 June 77 (3 m), 14 July 62 (1 m, 1
f), 18 July 62 (1 m, 2 f), 27 July 77 (1 m); LC at Mack’s Canyon Rd., 23 June 77 (1 f);
Mormon Well, 1 July 69 (1, LACM); 26 mi. E. CN, Mormon Well Rd., 1 July 69 (1,
LACM); Sawmill Canyon, 6000-7000', 2 July 69 (5, LACM); KC, 17.1 mi. W. U.S.
95, 5 July 78 (2 f); Cold Creek Ranger Station, 20 July 77 (2 m).

ADDITIONAL RECORDS: Early spring specimens have also been taken near
GS, 23 Mar. 77 (KBT); Pine Creek, 23 Mar. 72 (JFL): RC, 24 Mar. 75 (JB) and
Potosi Mt. Rd., 8 Apr. 71 (REW). Additional records are for locations and span of
dates listed above.

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Stanley a pin-
nata (Brassicaceae); Eriogonum mkrothecum, Eriogonum umhellatum (Poly gonaceae);
Arctostaphylos pungens (Ericaceae); Sarcostemma cynachoides (Asclepiadaceae);
Eriodictyon angustifolium (Hydrophyllaceae); Cryptantha jamesii (Boraginaceae);
Melilotus albus, Trifolium repens (Fabaceae); Salix sp. (Salicaceae); Rhus trilobata

26

J. Res. Lepid.

(Anacardiaceae); Viguiera multiflora, Baccharis sp., Senecio douglasii, Senecio
multilobatus, Tetradymia canescens (Asteraceae).

Callophrys sheridanii comstocki Henne.

Locally common with all records except one (Virgin Mountains) for the higher
desert area between Goodsprings and Mountain Springs Summit. There are two
broods from late March to early May and late August to mid September. The fall
brood is more yellow-green below than the spring brood as found in California
populations (Emmel and Emmel 1973).

SPECIMENS EXAMINED: (152) 0.7 mi. NNW Wilson Pass _(4 m, JFE);
Wilson Pass, 25 Mar. 78(1 m), 29 Mar. 78(1 m); CC, 29 Mar. 71 (1 f, NSM); 6 mi. N.
GS, 4 Apr. 77 (10 m, 3 f, DM); 2-3.3 mi. N. RC, 5 Apr. 78 (9 m, NSM; 30 m, 1 f), 11
Apr. 78 (2 m, NSM), 15 Apr. 79 (39 m), 4 May 78 (1 m), 21 Aug. 79 (11 m), 26 Aug.
79 (2 m, 1 f), 2 Sept. 78 (2 1 m, 9 f), 3 Sept. 78 (2 m, NSM), 8 Sept. 78 (2 m, NSM), 1 1
Sept. 78 (2 m).

ADDITIONAL RECORDS: GS, 23 Mar. 68 (Callaghan 1970); RC, 24 Mar. 75
(JB, RE W), 7 Apr. 74 (RE W); Potosi Mt. Rd., 6 Apr. 7 1 (RE W); GS, 1 1 Apr. 7 1 (s);
N. GS, 20 Apr. 75 (JB).

LARVAL FOODPLANTS: Eriogonum heermannii Dur. & Hilg. var. sulcatum
(S. Wats.) Munz & Reveal (Polygonaceae): 2-3 mi. N. RC, several dates (adult
assoc.).

ADULT RESOURCES: Eriogonum heermannii, Eriogonum microthecum (Poly-
gonaceae); Rhus trilobata (Anacardiaceae); C hrysothamnus sp. (Asteraceae).
Atlides halesus concorani (Clench).

Locally common in the valleys in stands of Prosopis or Acacia on which its
foodplant, Phoradendron californicum, is parasitic. The flight period extends from
late February to mid November involving at least 4 broods with peaks in early April,
early June, late July and October. The last 2 broods appear to be more numerous
than earlier broods. There are a few high elevation specimens which inay represent
strays or small colonies on Phoradendron juniperinum.

SPECIMENS EXAMINED: (106) Records extend from 25 Feb. (78, PV, 1 f) to
11 Dec. (77, GC, 1 f). High elevation specimens are: 0.5 mi. E. MS, 13 May 79 (2 m),
19 May 78 (2 m); WC, 28 July 77 (1 f); WC, 6 June 78 (2 f).

LARVAL FOODPLANTS: Phoradendron californicum Nutt. (Viscaceae) on
Prosopis glandulosa: PV, 18 July 77 (ovip.).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Eriogonum fascicu-
latum, Eriogonum plumatella, Eriogonum sp. (Polygonaceae); Sarcostemma hirtellum
(Asclepiadaceae); Eriodictyon angustifolium (Hydrophyllaceae); Acacia greggii,
Prosopis glandulosa (Fabaceae); Salix sp. (Salicaceae); Sambucus caerulea (Capri-
folicaceae); Bebbia juncea, Solidago spectabilis, Baccharis sp., Senecio douglasii,
Tetradymia canescens, Pluchea sericea (Asteraceae).

Strymon melinus pudica (H. Edwards).

Common, especially in agricultural areas, but with records for a variety of habitats
to 9000'. The flight period extends from late March through mid November
involving several broods. Desert records are for spring and fall. Clark County
material is tentatively assigned to the California race on the advice of Glenn A.
Gorelick pending completion of his study on this species.

"SPECIMENS EXAMINED: (209) Records for numerous locations between 25
Mar. 79, XM, (1 m) and 16 Nov. (77, KC, s); high elevation records extend from 30
May (66, WC, 1 m) to 26 Oct. (77, WC, 1 f).

19(1): 1-63, 1980(81)

27

ADULT RESOURCES: Tamarix pentrandra {Tamaricaceae); Eriogonum fasci-
culatum, Eriogonum heermanni, Eriogonum plumatella, Eriogonum umbellatum,
Eriogonum wrightii (Polygonaceae); Apocynum androsaemifolium (Apocynaceae);
Sarcostemma hirtellum (Asclepiadaeae); Eriodictyon angustifolium (Hydrophyl-
laceae); Marrubium vulgare (Lamiaceae); Fallugia paradoxa (Rosaceae); Acacia
greggii, Prosopis glandulosa, Prosopis pubescens, Medicago saliva, Melilotus albus
(Fabaceae); Helianthus annuus, Bebbia juncea, Bailey a pleniradiata, Hymenoxys
cooperi, Solidago spectabilis, Chrysothamnus sp., Baccharis sp., Senecio douglasii,
Tetradymia axillaris, Tetradymia canescens, Pluchea sericea, Cirsium sp. (Asteraceae).

Lycaena helloides (Boisduval).

Locally common in agricultural areas of Moapa and Virgin valleys. There are at
least 3 broods in a season ranging from mid April to late October. The fall brood is
by far the largest, peaking in early to late October. The larvae undoubtedly feed on
Rumex sp. which commonly borders fields and adults are often taken at Medicago
flowers.

SPECIMENS EXAMINED: (56) LO, 13 Apr. 78 (1 m), 10 July 68 (1 m), 9 Aug.
77 (1 m, 2 f), 23 Aug. 77 (1 f), 17 Sept. 77 (1 f), 28 Sept. 67 (2 m, NSM), 28 Sept. 68
(2 f, NSM), 9 Oct. 77 (1 m, 9 f), 10 Oct. 64 (6 m, 12 f, NSM), 16 Oct. 77 (7 m, 4 f), 2 1
Oct. 79 (1 m); OV, 7 June 77 (1 m); Hidden Valley, 23 Oct. 77 (1 m, 3 f).

ADDITIONAL RECORDS: Bunkerville, 17 May 78 (s); Hidden Valley, 7 June
77 (s); LO, 27 June 77, 30 July 77, 4 Sept. 77, 29 Sept. 77, 23 Oct. 77 (s).

ADULT RESOURCES: Convolvulus sp. (Convolvulaceae); Medicago saliva
(Fabaceae); Helianthus annuus, Baccharis sp. (Asteraceae).

Lycaena dorcas castro (Reakirt).

The single southern Nevada record was reported by Ferris (1977) from Charleston
Park (Kyle Canyon). If a colony once occurred here, it apparently no longer exists as
there are no recent records for this well collected area. The specimen may be
mislabelled.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: Charleston Park, 8000', 26 July 28 (1 m,fide CDF).
Brephidium exilis (Boisduval).

Locally abundant in somewhat alkaline areas of valleys, otherwise rare to common
in a variety of habitats to elevations of 8400'. The flight period at low elevations is
from early March to mid Decertiber and at high elevations from early May to mid
November. The lateness of the first records at high elevations suggests no
overwinter survival there. Reproduction (2-3 broods) occurs at least as high as
6800' where the species is associated with Atriplex canescens. At low elevations, 4 to
5 or more overlapping broods are produced.

SPECIMENS EXAMINED: (513) Many records from virtually throughout the
county. In the lowlands earliest and latest records are 3 Mar. (67,LO,NSM) and 12
Dec. (77, Tule Springs, s). Montane records are from 8 May (78, WC) to 12 Nov. (77,
WC, s).

LARVAL FOODPLANTS: Atriplex canescens (Pursh) Nutt. ssp. linearis (Wats.)
Hall & Clements (Chenopodiaceae): several locations, many dates (adult assoc.).
Suaeda torreyana Wats. (Chenopodiaceae); Hidden Valley, 9 Oct. 77 (larvae);
several locations, many dates (adult assoc.). Salsola spp. (probably including S.
iberica Sennen & Pau. and S. paulsenii Litv., see Beatley 1976) (Chenopodiaceae):
many locations, many dates (adult assoc.).

28

J. Res. LepM.

ADULT RESOURCES: Sphaeralcea ambigua (Malvaceae); Tamarix pentandra
(Tamaricaceae); Lepidium fremontii (Brassicaceae); Eriogonum fasciculatum,
Eriogonum heermannii, Eriogonum wrightii, Polygonum lapathifolium (Poly-
gonaceae); Eriodictyon angustifolium (Hydrophyllaceae); Heliotropium curassaui-
cum (Boraginaceae); Prosopis glandulosa, Medicago satiua, Melilotus albus (Fabaceae);
Rhus trilobata (Anacardiaceae); Viguiera multiflora, Bebbia juncea, Baileya pleni-
radiata, Baileya multiradiata, Chrysothamnus sp., Baccharis sp., Senecio douglasii,
Senecio multilobatus, Cirsium sp., (Asteraceae).

Leptotes marina (Reakirt).

Common to abundant in a variety of habitats to 7t)00' with occasional occurrences
as high as 11,000'. The flight season is from late March to mid November at low
elevations and from late April to late August at high elevations. Desert records are
mainly from March to May. At least 3 broods are produced in mesquite habitat
(May, June, July), continuous broods in agricultural areas (May-October) and a
single brood at high elevations (June-July).

SPECIMENS EXAMINED: (403) Numerous records at many locations at low
elevations with records from 25 Mar. (72, 9 mi. W. Davis Dam, REW) to 12 Nov.
(77, Tule Springs, s). High elevation records are few but widespread from 29 Apr.
(77, WC, CSL) to 26 Oct. (77, WC, 1 m).

LARVAL FOODPLANTS: Acacia greggii Gray (Fabaceae): PV, 8 June 77
(ovip. at base of flower buds). Prosopis glandulosa Tott. (Fabaceae): 0.5 mi. E. Blue
Diamond, 22 May 78 (ovip. on flower buds, 09:30 PST).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae) ; Larrea triden-
tata (Zygophyllaceae); Tamarix pentandra (Tamaricaceae); Sisymbrium altissimum,
Rorippa nasturtium- aquaticum (Brassicaceae); Eriogonum sp., Eriogonum fascicu-
latum {Folygonaceae); Sarcostemma hirtellum {ksclepiadaceae); Eriodictyon angus-
tifolium (Hydrophyllaceae); Heliotropium curassavicum (Boraginaceae); Acacia
greggii, Prosopis glandulosa, Prosopis pubescens, Medicago satiua, Melilotus albus.
Trifolium repens (Fabaceae); Rhus trilobata (Anacardiaceae); Viguiera deltoidea,
Bebbia juncea, Baileya multiradiata, Chaenactis fremontii, Haplopappus sp., Bac-
charis sp., Senecio douglasii, Tetradymia canescens, Pluchea sericea, Cirsium sp.
(Asteraceae).

Hemiargus ceraunus gyas (Edwards).

Fairly common throughout at elevations below 7800' and occasional to 11,000'.
The flight period at low elevations is from early April to mid November. Along small
desert washes, there are broods in April-May and October-November; in mesquite
stands and agricultural areas, there is a large brood flying from early July through
mid September depending on location and with scattered individuals at other times
of the year and at high elevations, there are at least 2 broods peaking in July and
October.

SPECIMENS EXAMINED: (179) Numerous records, especially at low eleva-
tions from 4 Apr. (77, PV, s) to 12 Dec. (77, Tule Springs, s). High elevation records
are from 13 June (77, WC, 1 f) to 12 Nov. (77, WC, 3 m).

LARVAL FOODPLANTS: Psorothamniis fremontii (Torr.) Barneby (Fabaceae):

RR, 14 May 77 (ovip. on new leaf growth, 10:30 PST); U.S. 93, 5 mi. N. 1-15, 14 May
78 (ovip. on leaf buds, 09:15 PST).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Eriogonum de-
flexum, Eriogonum fasciculatum, Eriogonum wrightii (Folygonaceae); Physalis eras-

19(1): 1-63, 1980(81)

29

sifolia {Solanaceae); Acacia greggii, Prosopis glandulosa, Prosopis pubescens, Medi-
cago saliva, Melilotus albus, Psorothamnus fremontii (Fabaceae); Bebbia juncea,
■Chaenactis fremontii, Palafoxia linearis, Erigeron sp., Senecio douglasii, Tetradymia
axillaris (Asteraceae).

Hemiargus isola alee (Edwards).

Commoner than the above with an equally broad ecological range but has a more
restricted season. Records extend from early April to late July at low elevations and
from late April to early August (once late October) at high elevations. Desert
records are for a single brood in April and May, at least 3 broods are produced in
mesquite stands (April, May, July) and a single brood flies in July at high elevations.
This species is outnumbered by ceraunus only in agricultural areas.

SPECIMENS EXAMINED: (212) Numerous records from many locations to
11,000'. Records extend from 8 Apr. (77, several locations) to 31 July (77, PV s) at
low elevations and 18 Apr. (77, KC, 17.1 mi. W. U.S. 95, 1 f) to 5 Aug. (77, WC, 1 f
and Cold Creek, 1 m) at high elevations. One very late specimen on 26 Oct. 77 (WC,
1 m).

ADULT RESOURCES: Sisymbrium altissimum (Brassicaceae); Eriogonum
umbellatum (Polygonaceae); Sarcostemma hirtellum (Asclepiadaceae); Eriodictyon
angustifolium (Hydrophyllaceae); Acacia greggii, Prosopis glandulosa, Prosopis
pubescens, Melilotus albus, Melilotus officinalis. Trifolium repens (Fabaceae); Rhus
trilobata (Anacardiaceae); Viguiera multiflora, Baileya pleniradiata, Tetradymia
axillaris, Senecio douglasii, Pluchea sericea, Cirsium sp. (Asteraceae).

Plebejus melissa melissa (Edwards).

Rare in agricultural areas of Moapa Valley in fall with records from early
September to late October. Clark County material is tentatively assigned to the
nominate race but may represent an undescribed subspecies. The insect is fairly
small, the wings (especially the hindwing) appear more rounded than those of other
populations examined and the females are quite dark with barely a hint of basal blue
overscaling.

SPECIMENS EXAMINED: (7) OV, 4 Sept. 66 (2 m, 1 f); LO, 4 Sept. 66 (1 m),
16 Oct. 77 (1 m, 1 f); Hidden Valley, 23 Oct. 77 (1 f).

ADDITIONAL RECORDS: OV, 29 Sept. 66, 10 Oct. 64 (PJH).

Plebejus icarioides evius (Boisduval).

Common to abundant in stands of Lupinus above 6400' mainly in the Kyle Canyon
and Lee Canyon areas of the Spring Range. The flight period of the single brood is
from mid May to early September. There are records from the Virgin Mountains
from mid May to late June and in early October.

The Spring Range specimens are of large size with relatively strong maculation
and fit closest to the concept of evius. The females are variable and over 60% have
orange scaling on the hindwing above. The few specimens from the Virgin
Mountains are smaller in size, have reduced maculation beneath and the females
have increased black scaling on the forewing above and no orange scaling. They thus
tend towards the Great Basin P. i. ardea (Edwards).

SPECIMENS EXAMINED: (316) Numerous records for the Spring Range to
11,300' from 12 May (34, Charleston Mts. 1 f, LACM) to 1 Sept. (77, KC Ski Run, 6
m, 3 f). Also recorded in the Virgin Mountains: 20 May 79 (1 m), 30 May 74 (2 m, 1 f,
NSM), 7 June 78 (s), 26 June 78 (2 m, 3 f), 7 Oct. 73 (1 f, NSM).

ADULT RESOURCES: Linum lewisii (Linaceae); Eriogonum umbellatum (Poly-

30

J. Res. Lepid.

gonaceae); Potentilla propinqua (Rosaceae); Lupinus sp., Melilotus albus (Fabaceae);
Chaenactis douglasii, Erigeron sp., Senecio douglasii (Asteraceae).

Plebejus shasta charlestonensis Austin.

Usually rare (may be common in some years) in Kyle and Lee canyons above 8200'
and in the Willow Creek area at 6000' to 8000'. The flight period of the single brood
is from early July to mid August. This taxon was recently described by Austin
(1980).

SPECIMENS EXAMINED: (17) LC Ski area, 5 July 76 (1 m, DM); LC, 8250'-
8800', 21 July 63 (7 m, 7 f); KC, 9000', 25 July 65 (1 m); KC, 8500', 27 July 65 (1 m).

ADDITIONAL RECORDS: Deer Creek Rd., 7000', 1 July 50 (FWP, figured in
Howe 1975); Cathedral Rock, 10 July 72 (DEA); WC, 6000-8000', 15 July 28
(Gunder colln. at American Museum of Natural History, fide JFE); LC Ski Area, 1 2-
17 Aug. 63 (JFL).

Plebejus acmon acmon (Westwood & Hewitson).

This taxon is known in the county only from the Newberry Mountains and a single
specimen from Las Vegas. The flight period is from late March to late October. The
Newberry Mountains’ population of P. acmon (referred to as acmon acmon) has a
pinkish-orange submarginal band on the secondaries and a narrow, non-infusive
dark margin on the primaries. All other Clark Co. populations (referred to as acmon
texanus) have an orange (with no pink tinge) submarginal band on the secondaries.
The dark margin on the primaries is broad with an indistinct inner border.

SPECIMENS EXAMINED: (15) XM, 12 May 78 (1 f), 23 May 78 (1 f), 1 June
78 (1 m, 2 f), 3 July 78 (1 f), 21 July 78 (1 m, 1 f), 4 Aug. 78 (1 f); Las Vegas, 29 June
62 (1 m); GC, 3 July 78 (1 m, 1 f) 21 July 78 (1 m), 4 Aug. 78 (1 m), 29 Oct. 78 (1 m).

ADDITIONAL RECORDS: 9 mi. W. Davis Dam, 31 Mar. 77 (CSL); XM, 21
May 73 (PJH); GC, 26 Sept. 78 (s).

ADULT RESOURCES: Eriogonum fasciculatum, Eriogonum plumatella (Poly-
gonaceae); Tetradymia axillaris (Asteraceae).

Plebejus acmon texanus Goodpasture.

This race of acmon constitutes the bulk of records for the species in Clark County,
ft is found throughout except in the Newberry Mountains and at the highest
elevations. It flies in several broods from late March to late October.

SPECIMENS EXAMINED: (230) Many locations and dates between 28 Mar.
(71, 6 mi. N. Apex, 1 m, NSM) and 26 Oct. (77, WC, 1 m).

ADULT RESOURCES: Eriogonum umbellatum (Polygonaceae); Sarcostemma
cynachoides {Asclepiadaceae); Eucrypta micrantha, Phacelia fremontii, Eriodictyon
angustifolium (Hydrophyllaceae); Crypthantha sp. (Boraginaceae); Potentilla pro-
pingua (Rosaceae); Melilotus albus, Astragalus lentiginosus (Fabaceae); Viguiera
multiflora, Eriophyllum wallacei, Tetradymia canescens, Cirsium sp. (Asteraceae).
Everes amyntula (Boisduval).

Uncommon in the Spring Range at elevations between 6000' and 9000'. It appears
to be single brooded with records from mid May to late July and single records in

late August and early October.

SPECIMENS EXAMINED: (28) Charleston Mts., 12 May 34 (5 m, LACM);
WC, 25 May 78 (1 m), 5 June 79 (1 nf), 6 June 78 (1 m), 13 June 77 (1 m), 22 June 68
(2 m), 22 June 78(1 m), 30 Aug. 77 (1 m), 1 Oct. 73 (1 m, UNLV); KC Campground,
31 May 77 (2 m), 8 June 77 (2 m, 3 f), 17 June 77 (1 m); WC, 13 June 77 (1 m), 28
July 77 (2 m); above Cathedral Rock Campground, 17 June 77 (1 m); LC Ski Area,
29 July 77 (1 m, 1 f).

19(1): 1-63, 1980(81)

31

ADDITIONAL RECORDS: WC, 1 July 77 (s), 13 July 72 (DEA), 16 July 72
(JFL), 28 July 77 (s); KC Campground, 22 June 78 (s).

ADULT RESOURCES: Eriogonum umbellatum (Polygonaceae).

Euphilotes battoides nr. ellisii (Shields).

A population with a phenotype similar to that of populations in desert mountains
of eastern California flies during August and September in the southern Spring
Range. We have been unable to find it around its suspected foodplant in other
portions of the Spring Range.

The use of the genus Euphilotes follows Mattoni (1977).

SPECIMENS EXAMINED: (22) 2-3.3 mi. N. RC, 19 Aug. 79 (2 m), 21 Aug. 79
(Im, 2 f), 26 Aug. 79 (1 m, 1 f), 2 Sept. 78 (1 m, 6 f), 3 Sept. 78 (1 f, NSM), 8 Sept. 78
(1 m, 2 f, NSM; 2 f); 11 Sept. 78 (2 f).

ADDITIONAL RECORDS: None.

LARVAL FOODPLANTS: Eriogonum heermannii Dur. & Hilg. var. sulcatum
(S. Wats.) Munz & Reveal (Polygonaceae): 3 mi. N. RC, 2,11 Sept. 78 (adult assoc.).

ADULT RESOURCES: Eriogonum heermanni (Polygonaceae).

Euphilotes battoides martini (Mattoni).

Common in association with Eriogonum fasciculatum especially in the Newberry
Mountains. Smaller populations occur in the Spring, Virgin and Las Vegas ranges.
The flight period is from mid March to late May.

SPECIMENS EXAMINED: (357) Newberry Mts., 8 Apr. 70 (1 m, NSM); 9 mi.
W. Davis Dam, 8 Apr. 77 (5 m, 1 f, NSM; 7 m, 4 f), 20 Apr. 78 (9 m, NSM; 7 m, 3 f), 22
Apr. 79 (12 m, 12 f), 1 May 75 (1 m, 1 f, NSM); 3 May 78 (9 m, 12 f), 12 May 78 (1 m,
1 f), 23 May 78 (1 m); GC, 8 Apr. 78 (2 m, 1 f, NSM; 3 m, 1 f), 12 Apr. 78 (1 m, NSM; 2
m, 2 f), 15 Apr. 78 (5 m, 1 f), 20 Apr. 78"(4 m, NSM; 12 m, 1 f), 3 May 78 (8 m, 5 f);
XM, 22 Apr. 79 (39 m, 2 f), 3 May 78 (25 m, 22 f), 5 May 77 (4 m, 12 f, NSM; 6 m), 6
May 79 (29 m, 30 f), 12 May 78 (5 m, 6 f), 23 May 78 (4 f); Pine Creek, 2 May 77 (1
m), 22 May 78 (3 m, 4 f); Bridge Canyon, 3 May 78 (4 m, 3 f); CC, 10 May 78 (3 m, 1
f), 17 May 78 (5 m, 5 ”f), 18 May 78 (2 m, 1 f), 20 May 79 (1 f); U.S. 93, 5-8 mi. N. 1-15,
14 May 78 (1 m, 4 f).

ADDITIONAL RECORDS: GC, 18 Mar. 72 (JFL), 12 May 78 (s); XM, 18 Mar.
72 (JFL), 20 Apr. 78 (s); 9 mi. W. Davis Dam, 25 Mar. 72 (RE W), 31 Mar. 77 (CSL);
CC, 20 May 78, 29 May 78 (s).

LARVAL FOODPLANTS: Eriogonum fasciculatum Benth. ssp. poliofolium
(Benth.) S. Stokes (Polygonaceae): 9 mi. W. Davis Dam, 8 Apr. 77 (ovip. on flower
heads, 10:00 PST); GC, several dates (adult assoc.); XM, several dates (adult
assoc.); Pine Creek, 2 May 77, 22 May 78 (adult assoc.); CC, several dates (adult
assoc.); U.S. 93, 5-8 mi. N. 1-15, 14 May 78 (adult assoc.).

ADULT RESOURCES: Eriogonum fasciculatum (Polygonaceae); males and,
occasionally females visit mud,

Euphilotes battoides baueri (Shields).

This taxon has been recorded once in Clark County. It was originally reported as
‘‘"Philotes” mojaue collected on 14 April 1969 (1969 summ.); the specimens are the
only ones located in Nevada State Museum bearing the date of 13 May 1969. We
retain the label date but point out the strong possibility that the correct one is in
April (see introduction) or that these specimens are not from Clark County.

SPECIMENS EXAMINED: (2) AC, 13 May 69 (1 m, 1 f, NSM). This record
also was reported by Shields (1975).

ADDITIONAL RECORDS: None.

32

J. Res. Lepid.

Euphilotes enoptes (Boisduval) ssp.

A nearly purple, heavily marked population of enoptes occurs in the Spring Range
between 6100' and 7200'. It flies in association with Eriogonum umbellatum from
early June to mid August. This unnamed population has been variously referred to
as near E. e. enoptes and E. e. ancilla (Barnes & McDunnough).

SPECIMENS EXAMINED: (192) WC, 5 June 79 (60 m), 6 June 78 (53 m), 11
June 79 (33 m), 13 June 77 (9 m), 20 July 77 (7 m, 1 f); end KC, 2 July 36 (1 m,
LACM); KC Campground, 25 July 77 (2 m, 1 f); WC, 20 July 78 (10 m, 5 f), 28 July
77 (5 m, 4 f), 19 Aug. 77 (1 m).

ADDITIONAL RECORDS: WC, 17 June 77 (s), 16 July 72 (JFL), 28 July 77
(s); Mt. Charleston, 7200', 26 June 59 (Shields 1977); KC, 17.1 mi. W. U.S. 95, 6
July 77 (s); KC, 7100', 14 July 66 (Shields 1977).

LARVAL FOODPLANTS: Eriogonum umbellatum Torr. var. subaridum S.
Stokes (Polygonaceae): WC, 28 July 77 (ovip. on flowers, 08:30 PST).

ADULT RESOURCES: Eriogonum umbellatum (Polygonaceae); males fre-
quent mud.

Euphilotes enoptes dammersi (Comstock & Henne).

The only records are from mid September to early October in the Newberry
Mountains and in the Spring Range at Pine Creek. The one previous record for
Nevada is for Lincoln County (Shields 1977).

SPECIMENS EXAMINED: (55) GC, 13 Sept. 78 (2 m, NSM), 20 Sept. 78 (1
m, 2 f, NSM; 4 m, 4 f), 26 Sept. 78 (2 m, 3 f, NSM; 2 m 5 f), 27 Sept. 77 (3 m), 4 Oct.
77 (6 m, 2 f); XM, 13 Sept. 78 (1 m, 2 f); Pine Creek, 19 Sept. 78 (1 m), 19 Sept. 79 (3
m, 5 f), 22 Sept. 78 (2 m, 5 f).

ADDITIONAL RECORDS: Sacatone Wash, 26 Sept. 78 (s).

LARVAL FOODPLANTS: Eriogonum wrightii Torr. ex Benth (Polygonaceae):
GC, 4 Oct. 77 (ovip. on flower buds, 10:00 PST).

ADULT RESOURCES: Eriogonum plumatella, Eriogonum umbellatum, Eriog-
onum wrightii (Polygonaceae); males visit mud.

Euphilotes mojaue (Watson & Comstock).

A population of this species was discovered in the Virgin Mountains in 1978. It
flies from early May to early June at elevations between 3800' and 6000'. Although
this population is undoubtedly mojaue there are certain differences from nominate
mojaue and the population probably deserves a name. There are no previous
Nevada records (those reported in the 1969 summ. are actually battoides, see
above).

We agree with Mattoni (1977) that mojaue deserves specific status and is not a
race of enoptes as suggested by Shields (1975).

SPECIMENS EXAMINED: (84) CC, 10 May 78 (9 m), 18 May 78 (25 m, 9 f), 20
May 78 (6 m, 4 f), 20 May 79 (16 m, 10 f), 29 May 78 (1 m, 2 f), 7 June 78 (2 m).

ADDITIONAL RECORDS: None.

LARVAL FOODPLANTS: Eriogonum nidularium Cov. (Polygonaceae): CC, 20
May 78 (abdominal probing near unopened flower buds, 13:55 PST).

ADULT RESOURCES: Eriogonum uimineum (Polygonaceae); Cryptantha sp.
(Boraginaceae).

Philotiella speciosa speciosa (H. Edwards).

Two specimens are known from Clark County. One bears no date on its label but
was apparently taken on 14 April 1969 (1969 summ., see introduction above;

19(1): 1=63, 1980(81)

33

Shields, 197 4, reports this as a May specimen) and the other was taken in June. The
genus Philotiella was recently proposed by Mattoni (1977),

SPECIMENS EXAMINED: (2) AC, no date (1 m, NSM); Willow Spring, 17
June 78 (1 m, ex DM).

ADDITIONAL RECORDS: None.

Glaucopsyche lygdamus (Doubieday) ssp.

Small numbers of this species fly from early April to mid May in the higher desert
between Goodsprings and the Pahrump Road in the Cottonwood Pass area of the
southern Spring Range. There is also one record for the McCullough Mountains. It
appears to belong to the unnamed eastern Mojave Desert population that is known
from several desert ranges in California (Enimel and Emmel 1973).

SPECIMENS EXAMINED: (113) MM, 12 Apr. 69 (1 m, NSM); CP, 0.6M.8 mi.
S. Pahrump Rd., 16 Apr. 77 (1 m), 16 Apr. 79 (65 m, 5 f), 22 Apr. 78 (5 m, 3 f), 29
Apr. 8 (7 m, 2 f), 29 Apr. 79 (14 m, 7 f), 2 May 78 (2 m), 11 May 78 (1 f).

ADDITIONAL RECORDS: CP and RC, 7 Apr. 74 (REW); 1 mi. N. CP, 12 Apr.
76 (JB); CP Road, 0.6 mi, S. Pahrump Rd., 20 Apr, 77 (s).

LARVAL FOODPLANTS: Astragalus lentiginosus Dough var. fremontii (A.
Gray) S. Wats. (Fabaceae): CP Rd., 1,5 mi. S. Pahrump Rd., 22 Apr. 78 (abdominal
probing among flowers).

ADULT RESOURCES: Dicheiostemma pulchellum (Amaryllidaceae); Arabis
sp, (Brassicaceae); Salvia dorrii (Lamiaceae); Astragalus lentiginosus (Fabaceae).
Glaucopsyche lygdamus oro Scudder,

A few records from Kyle Canyon from mid June to late July indicate the presence
of a small colony of this taxon in the Spring Range. It is associated with Lupinus and
seems best placed in the Rocky Mountain-Great Basin race oro.

SPECIMENS EXAMINED: (4) above Cathedral Rock Campground, 8000b 17
June 77 (1 m, 1 f); KC Ski Run, 2 July 78 (1 f), 23 July 78 (1 m).

ADDITIONAL RECORDS: None.

Celastrina argiolus cinerea (Edwards).

Common to locally abundant at elevations between 3600' and 95007 Records
extend from mid March to late October. Early spring records especially are from the
more mesic lower canyons with the first high elevation records in late April.

Early spring specimens tend to be smaller than those from later broods. Spring
females have relatively narrow black borders above; summer females are heavily
bordered with black, a form to which the name “arizonensis” has been applied.

SPECIMENS EXAMINED: (376) Many records from the Spring Range from
19 Mar. (72, Calico Basin, s) to 24 Oct. (65, Calico Basin, 1 m). There are a few
records for the Virgin Mts. and one for the Newberry Mts. (GC, 20 Sept. 78, 1 m).

LARVAL FOODPLANTS: Petrophytum caespitosum (Nutt.) Rybd. (Rosaceae):
Deer Creek, 25 June 68 (ovip. on flower buds, JFE, OS). Peraphyllum ramosissi-
mum Nutt. (Rosaceae): RD Summit Rd., 1.5 mi. E. Lovell Wash, 24 April 77 (ovip.
on flower buds).

ADULT RESOURCES: Eriogonum plumatelia, Eriogonum umhellatum, Eriog-
onum wrightii (Polygonaceae); Arctostaphylos pungens (Ericaceae); Apocynum
androsaemifolium (Apocynaceae); Sarcostemma cynachoides (Asclepiadaceae);
Eriodictyon angustifolium (Hydrophyllaceae); Potentilla propinqua (Rosaceae);
Lupinus sp. Melilotus alhus (Fabaceae); Acer giabrum (Aceraceae); Rhus triiobata
(Anacardiaceae); Viguiera multiflora, Chrysothamnus sp., Senecio douglasii, Tetra-
dyrnia canescens, Cirsium sp. (Asteraceae),

34

J. Res, LepM.

Family Libytheidae

Lihytheana bachmanii larvata (Strecker).

Usually uncommon to rare, but abundant in some years to 7 600'. It occurs in most
habitats but is most common along desert washes where Chrysothamnus is
flowering. All except three records are from late August to mid December and
probably represent migrants. The May and June records may be of individuals
raised locally on Celtis reticulata which occurs rarely in the Red Rock area and in the
Virgin Mountains. This remains to be documented.

SPECIMENS EXAMINED: (104) GC, 14 Sept. 77(1 m), 4 Oct. 77(1 f), 12 Oct.
77 (2 f), 25 Oct. 77 (7 m, 1 f), 3 Nov. 77 (2 m, 4 f), 11 Dec. 77 (2 m, 1 f); Bowman’s
Reservoir, 17 Sept. 77 (1 m); LO, 28 Sept. 66 (3, NSM), 16 Oct. 77 (1 f); Las Vegas,
3 Oct. 77 (1 m, UNLV); 2.9 mi, W, Searchlight, 4 Oct. 77 (1 m, 1 f); 1.8 mi. E. Blue
Diamond, 1 1 Oct. 77 (1 m), 22 Oct. 77 (6 m, 1 f), 2 Nov. 77 (2 m, 1 f); WC, 14 Oct. 77
(1 f); KC Ski Run, 15 Oct. 77 (1 m); Whitney Mesa, 21 Oct. 77 (1 m, 4 f); Pahrump
Rd., 3 mi. W. Blue Diamond turnoff, 22 Oct. 77 (14 m, 9 f); 0.6 mi, W. Blue
Diamond, 22 Oct. 77 (1 f); Newberry Mts., 23 Oct. 70 (17, NSM); 2 mi. W.
Searchlight, 25 Oct. 77 (3, NSM); Cold CreekRanger Station, 26 Oct. 77 (3 m); WC,
26 Oct. 77 (1 m, 1 f); Tule Springs, 4 Nov. 77 (2 m); CN, 12 Nov. 77 (1 m, 1 f); 1 mi.
W. MS, 16 Nov. 77 (1 f); KC, 20 mi. W.'U.S. 95, 16 Nov. 77 (1 f); RD, 25 Nov. 77 (1 f,
UNLV); PV, 28 Nov. 77 (1 f).

ADDITIONAL RECORDS: GC, 23 Mar. 78 (s); Willow Springs, 10 May 66
(JFL); MS, 30 June 63 (KR); Potosi Mt. Rd., 23 Aug. 75 (JB); CN, 1 Sept. 63 (KR);
14 Oct. 77, 26 Oct. 77, 4 Nov. 77 (s); 15.9 mi. W. Searchlight, 4 Oct, 77 (s); Las
Vegas, 6 Oct. 70 (JFL); LO, 7 Oct. 64 (PJH); KC, 9.3 mi. W. U.S. 95, 15 Oct. 77 (s);
jet. U.S. 93 and 1-15, 23 Oct. 77, 5 Nov, 77 (s); Echo Canyon, 24 Oct. 77 (s); 0.2 mi.
E. Boulder City, 25 Oct. 77 (s); WC, 26 Oct. 77 (s); Tule Springs, 1 Nov. 77, 12 Nov.
77, 12 Dec. 77 (s); 1.0 mi. W. MS, 1 Nov. 77 (s); UNLV campus, 1 Noy. 77 (s); 0.6 mi.
W. Blue Diamond, 1 Nov. 77 (s); RR, 3 Nov. 77 (s); 0.5 mi. E. Boulder City, 3 Nov.

77 (s); 1.8 mi. E. Blue Diamond, 16 Nov. 77 (s).

ADULT RESOURCES: Eriogonum sp. (Polygonaceae) ; Me/i/otus albus (Fabaceae);
Chrysothamnus sp., Baccharis sp., Senecio douglasii, Cirsium sp. (Asteraceae).

Family Nymphalidae

Asterocampa celtis montis (Edwards).

A small population of this taxon occurs in association with Celtis reticulata in the
Virgin Mountains. There are no other records for Nevada.

SPECIMENS EXAMINED: None.

ADDITIONAL RECORDS: 0.4 mi. N. Nay (Whitney on maps) Ranch, 11 Sept.

78 (KR), 11 Oct 78 (s).

Limenitis archippus obsoleta Edwards.

Local in Moapa Valley in close association with stands of Populus and/or Saiix. It
flies in at least 2 broods from early June to late October; an early brood is suggested
by a specimen from Beaver Dam, Mohave Co., Arizona, on 7 April 1963 (1 m). The
June brood appears to be the most numerous.

SPECIMENS EXAMINED: (53) Hidden Valley, 7 June 77 (1 m), 27 June 77 (5
m), 12 July 77 (1 m), 9 Aug. 77 (1 m); Bowman’s Reservoir, 7 June 77 (1 m), 15 Jpne
T7 (7 m, 3 f), 16 June 77 (1 m, NSM), 27 June 77 (4 m, 1 f), 4 July 77 (3 m), 30 July 77
(2 m), 23 Aug. 77 (1 m), 9 Oct. 77 (1 m); LO, 15 June 77 (1 m), 4 July 77 (1 m), 23

19(1): 1-63, 1980(81)

35

Aug. 77 (2 m), 9 Sept. 79 (1 m, 2 f), 27 Sept. 78 (3 m, 1 f), 9 Oct. 77 (2 m, 1 f), 16 Oct.

77 (1 m, 1 f), 21 Oct. 79 (1 m); OV, 16 June 77 (2 m, NSM), 10 Oct. 64 (2 f, NSM).
ADDITIONAL RECORDS: LO, 27 June 77, 9 Aug. 77 (s); Hidden Valley, 4

July 77, 30 July 77 (s); Bowman’s Reservoir, 9 Aug. 77, 4 Sept. 77, 29 Sept. 77 (s);
OV, 27 Sept. 78 (s), 1 Oct. 68 (1968 summ.).

Limenitis weidemeyerii angustifascia (Barnes & McDunnough).

A population assigned to this taxon flies in the Virgin Mountains in May and June.
This material shows no intermediacy with neuadae. It is large with the more rounded
wings of the Arizona race but has slightly wider median white bands. In this feature
it is slightly intermediate towards L. w. latifascia Perkins & Perkins. Only one of the
nine specimens examined has a trace of white in the cell.

SPECIMENS EXAMINED: (9) CC, 4000-6000\ 29 May 78 (1 m), 7 June 78 (4
m), 26 June 78 (4 m).

’additional RECORDS: CC, 20 May 79 (s); Virgin Mts., 7200', 30 June 66
(JFL).

Limenitis weidemeyerii nevadae (Barnes & Benjamin).

This distinctive taxon occurs in the Spring Range from 4400' in mesic canyons to

about 8400'; it is most common in Kyle Canyon (above 6500') and in the Cold-
Willow creeks area. There are also records of this race from the Sheep Range. The
single brood flies from late May to late August with a peak in early July.

SPECIMENS EXAMINED: (183) Many records from 22 May (78, Pine Creek,
1 m) to 22 Aug. (77, KC Ski Run, 1 f).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Eriodictyon
angustifolium (Hydrophyllaceae); Marrubium vulgare (Lamiaceae); Cirsium sp.
(Asteraceae).

Adelpha bredowii eulalia (Doubleday).

Fairly common in certain low, mesic canyons and at high elevations in the vicinity
of oaks. It occasionally strays as far as Las Vegas. The flight period at low elevations
is from mid April to mid July and early September to early November. At high
elevations, it flies from late May to early October. At least two broods are involved.

The Spring Range population apparently represents the westernmost extreme of
this subspecies’ range.

SPECIMENS EXAMINED: (95) RD Springs, 14 Apr. 64 (1 m); Lost Creek, 3
May 77 (1 m, NSM), 2 June 77^3 m, NSM); CC, 12 May 77 (1 m, NSM), 17 May 78
(1 m), 20 May 78 (1 m), 20 May 79 (1 m), 29 May 78 (6 m), 7 June 78 (4 m), 26 June

78 (2 m, 1 f), 28 Sept. 74 (4 m, 5 f, NSM), 1 1 Oct. 78 (1 m, 1 f); Pine Creek, 22 May 78
(3 m), 2 June 77 (1 m, NSM), 1 Sept. 75 (1 f, NSM), 12 Sept. 75 (1 f, NSM), 19 Sept.
77 (1 m, 1 f), UNLV), 24 Sept. 73 (2, UNLV), 29 Sept. 71 (1, UNLV), 5 Oct. 77 (1 m,
1 f); Virgin Mts., 30 May 74 (2 f, NSM), 7 Oct. 73 (1 m, 1 f, NSM); RD Summit Rd.,
1.5-2. 1 mi. E, Lovell Wash, 2 June 77 (1 f), 22 June 77 (1 m, 1 f); Willow Spring, 2
June 77 (2 m, NSM); WC, 22 June 68 (1 f), 13 Oct. 78 (1 m); Deer Creek Rd., 2.8 mi.
N. KC, 26 June 79 (1 m); KC, 20 mi. W. U.S. 95, 28 June 78 (1 m), 5 July 78 (1 f), 25
Aug. 78 (2 m), 26 Sept. 77 (1 m); KC Campground, 24 June 79 (2 m, 1 f), 30 June 77
(1 f, NSM); Charleston Park, 30 June 77 (1 m, 2 f, NSM); KC Ski Run, 20 June 79 (2
m), 2 July 78 (1 f); KC, 7500', 15 July 74 (2 m, 1 f); Clark Co., 9 Sept. 73 (1, UNLV),
9 Sept. 74 (1, UNLV), 12 Sept. 76 (1, UNLV), 24 Sept. 73 (3, UNLV), 29 Sept. 71
(1, UNLV), 30 Sept. 76 (1, UNLV), 13 Oct. 74 (1, UNLV), 15 Oct. 74 (1, UNLV)r
KC, 11 Sept. 74 (2 f, NSM); RD, 23 Sept. 77 (1 m, UNLV), 15 Oct. 73 (1, UNLV);

36

J. Res. Lepid.

Las Vegas, 24 Sept. 74 (1, UNLV), Oct. 74 (1, UNLV), 21 Oct. 71 (1, UNLV), 10
Nov, 76 (1, UNLV); Lovell Canyon, 25 Sept. 75 (1 m, NSM).

ADDITIONAL RE CORDS: Many additional records from locations and within
the dates cited above. Additional collection sites are: MS, LC, Camp Bonanza,
Arden Springs, Ash Springs and Oak Creek Canyon.

LARVAL FOODPLANTS: Quercus gambelii Nutt. (Fagaceae): RD Summit
Rd., 22 June 77 (ovip.).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Chryso-
thamnus sp., Cirsium sp. (Asteraceae).

Vanessa atalanta rubria (Fruhstorfer).

Fairly common locally in fall during some years but rare at other times of the year.
There are a few records of mostly single individuals from March to August and a
large brood from late September to early November. The habitat of V. atalanta is
generally adjacent to riparian areas although it is occasionally found away from
these situations.

SPECIMENS EXAMINED: (49) CC, 10 May 78 (1 m); WC, 5 June 79 (1 f), 22
June 78 (2 m), 14 Oct. 77 (1 m, 1 f), 26 Oct. 77 (1 f); WC, 22 June 78 (1 f), 14 Oct. 77
(5 m, 6 f), 26 Oct. 77 (2 m, 3 f); Lovell Wash, 4475', 17 June 62 (1 m); Clark Co., 15
July 74 (1, UNLV); Deer'Creek, 21 Aug. 71 (1, UNLV); Lovell Wash, 25 Sept. 75 (1
f, NSM); KC, 20 mi. W. U.S. 95, 27 Sept. 78 (2 f); LO, 27 Sept. 78 (2 f); KC, 3998', 9
Oct. 66 (1 f); KC, 6940', 9 Oct. 66 (3 f); Tule Springs, 13 Oct. 77 (7, CSL), 14 Oct. 77
(1 f), 4 Nov. 77 (2 m); KC Ski Run, 13 Oct. 78 (1 m), 15 Oct. 77 (1 m); Echo Canyon,
24 Oct. 77 (1 m).

ADDITIONAL RECORDS: 9 mi. W. Davis Dam, 25 Mar. 79 (s); XM, 12 Apr.
78 (s); Hidden Valley, 13 Apr. 78 (s); White Rock Spring, 21 Apr. 79 (2); CP Rd., 1.5
78 (s); Hidden Valley, 13 Apr. 78 (s); White Rock Spring, 21 Apr. 79 (s); CP Rd., 1.5
mi. S. Pahrump Rd., 29 Apr. 79 (s); Lovell Wash, 5375', 17 June 62 (s); WC, 11 June
79, 22 June 78, 1 July 77, 20 July 77 (s); KC Ski Run, 20 June 79, 2 July 78, 8 Oct. 77
(s); Lost Creek, 23 June 78 (s); LC ballfield, 29 July 77 (s); CN, 14 Oct. 77, 4 Nov. 77
(s); Echo Canyon, 15 Oct. 77 (s); KC Campground, 15 Oct. 77, 24 Oct. 77 (s); 'Tule
Springs, 1 Nov. 77, 29 Nov. 77 (s).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Amsinckia
tessellata (Boraginaceae) ; Marmhiam vulgare (Lamiaceae);Me/^7o^as albus (Fabaceae);
aenactis sp. Haplopappus sp., Chrysothamnus sp., Baccharis sp. (Asteraceae).

Vanessa virginiensis (Drury).

Rare, occurring mostly at high elevations but also occasionally in desert washes and
agricultural areas. It is apparently more common in some years than in others. It flies in
2 broods from mid June to early August and mid September to early November.

SPECIMENS EXAMINED: (19) Cold Creek Ranger Station, 13 June 77 (2 f);
PotosiMt. Rd. at Pahrump Rd., 14 June 77 (1 f); Overton, 15 June 77 (1 f);LC ballfield,
23 June 77 (1 m), 7 July 77 (1 m, NSM; 1 m); CC, 26 June 78 (1 m); KC, 20 mi. W. U.S.
95, 28 June 77 (1 m); ridge above KC, 11,200', 19 July 78 (1 m, 1 f); WC, 20 July 77 (1
m), 23 Sept. 77 (1 m); CN, 15 Sept. 64 (1 m, NSM); Lovell Canyon, 6000', 25 Sept. 75
(1 f, NSM); KC Ski Run, 8 Oct. 77 (1 m), 15 Oct. 77 (1 m); GC, 25 Oct. 77 (1 f); Tule
Springs, 4 Nov. 77 (1 m).

ADDITIONAL RECORDS: WiUow Creek, 17 June 77, 1 July 77 (s); KC Ski Run,
26 June 79, 2 July 78, 18 July 79 (s); GC, 21 July 78 (s); WC, 28 July 77, 14 Oct. 77, 26
Oct. 77 (s); Charleston Mts., 6 Aug. 61 (RES); 1.8 mi. E, Blue Diamond, 22 Oct. 77 (s).

19(1): 1-63, 1980(81)

37

ADULT RESOURCES: Clematis ligustidfolia (Ranunculaceae); Erysimum asperum
(Brassicaceae); Eriodictyon angustifolium (Hydrophyliaceae); Lupinus sp., Medicago
saliva, Melilotus albus (Fabaceae); Viguiera deltoidea, Encelia farinosa, Chryso-
thamnus sp., Senedo douglasii, Tetradymia canescens, Cirsium sp., Taraxacum
officinale (Asteraceae).

Vanessa cardui (Linneaus).

Common to abundant in nearly all habitats. Low elevation records are for every
month with desert records primarily in spring and fall. High elevation records are
from early April to mid November. A large northward migration from 4 to 24 March
1968 with scattered individuals still moving into mid April (similar movements were
seen at this time in Sonora, Mexico, and southern California, 1968 summ.). Large
numbers in April 1973 and June and July 1965 (see also Emmel and Wobus 1966)
showed no migratory tendencies and probably reflected favorable reproduction.
The 1973 numbers were during one of the best local spring blooms of annuals on
record; 1973 was a year of enormous numbers of the butterfly in California north to
the Oregon border (Shaprio 1974). Several dwarfs were obtained in June and July
1965. Smaller scale movements also occur in other years {e.g., 1978) which may go
unnoticed.

SPECIMENS EXAMINED: (202) Records are from every month and for
nearly all locations in Clark County to the top of Charleston Peak.

LARVAL FOODPLANTS: Sphaeralcea ambigua Gray (Malvaceae): Las Vegas,
several records (ovip., larvae). Althaea rosea (L.) Cav. (Malvaceae): Las Vegas,
several records (ovip., larvae). Cirsium neomexicanum A. Gray (Asteraceae): 2 mi.
N. RC, 5 Apr. 78 (ovip., 2x, upper leaf surface, 10:10 PST).

ADULT RESOURCES: Clematis ligustidfolia (Ranunculaceae); Tamarix pen-
tandra (Tamaricaceae); Erysimum asperum (Brassicaceae); Eriogonum fasciculatum,
Eriogonum heermannii, Eriogonum microthecum, Eriogonum sp. (Polygonaceae);
Arctostaphylos pungens (Ericaceae); Menodora spinescens (Oleaceae); Apocynum
androsaemifolium (Apocynaceae); Phacelia vallis-mortae, Eriodictyon angustifolium
(Hydrophyliaceae); Sarcostemma hirtellum (Asclepiadaceae); Heliotropium curas-
savicum, Amsinckia tessellata {Boraginaceae); Mimulus guttatus (Scrophulariaceae)
Verbena gooddingii (Verbenaceae); Marrubium vulgare (Lamiaceae); Medicago
sativa, Melilotus albus, Psorothamnus fremontii (Fabaceae); Rhus trilobata (Ana-
cardiaceae); Lomatium parryi, Angelica scabrida (Apiaceae); Viguiera deltoidea,
Helianthus annuus, Encelia farinosa, Bebbia juncea, Psilostrophe cooperi, Baileya
multiradiata, Chaenactis fremontii, Pectk papposa, Haplopappus sp., Chrysothamnus
sp., Baccharis sp., Senedo douglasii, Pluchea sericea, Cirsium sp., Taraxacum
officinale (Asteraceae).

Vanessa annabella (Field).

Common to abundant in nearly all habitats. Lowland records are from early
February to mid December and high elevation records are from early April to mid
November. Desert records are for spring and fall.

SPECIMENS EXAMINED: (149) Specimens were seen for all months except
January, August and December but there are sight records for the latter two
months.

LARVAL FOODPLANTS: Sphaeralcea ambigua Gray (Malvaceae); many re-
cords (larvae). Althaea rosea (L.) Cav. (Malvaceae); Las Vegas, several records
(ovip., larvae).

38

J. Res. Lepid.

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Lepidium fre-
montii, Erysimum asperum (Brassicaceae); Eriogonum fasciculatum (Poly|onaceae);
Abronia villosa (Nyctaginaceae); Apocynum androsaemifolium (Apocynaceae);
Phacelia fremontii, Eriodictyon angustifolium (Hydrophyllaceae); Heliotropium
curassavicum (Boraginaceae); Marrubium 'Vulgare (Lamiaceae); Ribes aureum
(Saxifragaceae); Medicago saliva, Melilotus albus (Fabaceae); Helianthus annuus,
Encelia farinosa, Bebbia juncea, Baileya multiradiata, Chaenactis fremontii, Solidago
spectabilis, Chrysothamnus sp., Baccharis sp., Cirsium sp., Taraxacum officinale
(Asteraceae).

Precis coenia (Huebner).

Common in agricultural areas and relatively rare away from these occurring
occasionally in desert washes, at springs ind at high elevations. The flight period is
from mid April to mid November. Most specimens are from agricultural areas from
late August to mid November. Wild populations are irregular and suggest that the
species is maintained in these areas by repeated immigrations.

SPECIMENS EXAMINED: (85) CC, 19 Apr. 75 (1 m, NSM), 18 May 78 (1 m),
26 June 78 (1 m); XM, 6 May 79 (1 m), 21 July 78 (1 m); WC, 22 June 78 (1 f), 13
Oct. 78 (1 m, 1 f); Cold Creek Ranger Station, 22 June 78 (1 m), 26 Oct. 77 (1 m);KC
Ski Run, 24 June 79 (1 f); Cold Creek, 20 July 78 (1 m); GC, 21 Aug. 78 (1 m);
Bowman’s Reservoir, 23 Aug. 77 (1 m), 29 Sept. 77 (1 m); LO, 23 Aug. 77 (1 m), 4
Sept. 77 (1 m), 17 Sept. 77 (1 m), 28 Sept. 67 (1 m, 2 f, NSM), 28 Sept. 68 (1 m,
NSM), 29 Sept. 77 (1 m), 9 Oct. 77 (3 m, 3 f), 16 Oct. 77 (1 m); Clark Co., Sept. 73 (1,
UNLV); OV, 4 Sept. 66 (3 m), 15 Sept. 63 (2 m, 1 f, NSM), 25 Sept. 66 (3 m), 27
Sept. 78 (1 m, NSM), 29 Sept. 66 (1 m, 1 f, NSM), 23 Oct. 77 (9 m, 4 f), 5 Nov. 77 (5
m, 6 f), 11 Nov. 77 (9 m, 2 f); 1 mi. W. MS, 30 Sept. 79 (3 m, 1 f); Hidden Valley, 9
Oct. 77 (1 m); Las Vegas, 12 Oct. 70 (1, UNLV); 1.8 mi. E. Blue Diamond, 22 Oct.
77 (1 f).

ADDITIONAL RECORDS: Other records for span of dates above. Additional
locations are CP Rd., Pine Creek, RC area. Sawmill Canyon, RR and Tule Springs.

ADULT RESOURCES: Medicago saliva (Fabaceae); Baccharis sp., Tetra-
dymia axillaris, Tetradymia canescens, Cirsium sp. (Asteraceae).

Nymphalis calif arnica (Boisduval).

Scattered records exist for this species above 6700' in the Spring and Sheep
ranges. There is also one record for the Virgin Mts. In most years it is rare or absent
but it was reported as “very abundant” in 1950 (FWP).

SPECIMENS EXAMINED: (6) CC, 5 June 79 (1, CSL); KC, 12 June 66 (1 m);
Clark Canyon, 8000', 10 July 75 (1 f, NSM); Little Falls, 8000', 31 July 62 (2, JFL);
KC, 78000', 11 Oct. 71 (1, JFL).

ADDITIONAL RECORDS: WC, 22 June 78 (s); Hidden Forest, 23 June 61
(KR); Little Falls, 29 June 63 (KR); KC, 6700-7500', 30 June 50 (FWP); KC Ski
Run, 2 July 78 (s); Charleston Mts., 10-12 Aug. 63 (JFE).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae).
Nymphalis milberti furcillata (Say).

Rare in Kyle Canyon and the Cold-Willow creeks area and usually not far from its
foodplant Urtica holsericea which grows in small patches at these localities. Records
extend from late April to early July, probably reflecting a single brood emerging in
late June and overwintering.

SPECIMENS EXAMINED: (3) WC, 27 Apr. 77 (1 m; 1, CSL), 4 July 62 (1 m).

19(1): 1-63, 1980(81)

39

ADDITIONAL RECORDS: Little Falls, 15 May 72 (JFL); WC, 22 June 68, 22
June 78 (s); Charleston Peak, 24 June 68 (JFE, OS).

LARVAL FOODPLANTS: Urtica holosericea Nutt. (Urticaceae): KC, 23 May

63 (larvae, JL).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae).
Nymphalis antiopa antiopa (Linnaeus).

Uncommon over a wide elevational range to at least 9000'. Low elevation records
are generally from the vicinity of springs or towns. The flight period extends from
early February to mid November. One brood is involved with overwintering adults
occurring as late as 5 July and overlapping with the next generation which has been
caught as early as 14 May.

SPECIMENS EXAMINED: (30) Willow Spring, 23 Mar. 77 (1 m, 1 f, NSM);
Lost Creek, 23 Mar. 77 (1 f, NSM), 20 Apr. 77 (1 f, NSM); Pine Creek, 23 Mar. 77(1
m, NSM), 9 Oct. 71 (1, UNLV); KC, 20 mi. W. U.S. 95, 26 Apr. 77 (2 m), 5 July 78 (1
m, 1 f), 6 July 77 (1 m); 2 mi. N. RC, 4 May 78 (1 f); Clark Co., 14 May 67 (1, UNLV),
9 Sept. 74 (1, UNLV); KC, 14 May 67 (1 f), 6 June 67 (1, UNLV); LO, 17 May 78 (1
f); WC, 25 May 78(1 m), 22 June 78 (1 m), 23 June 62 (1 m), 1 July 77(1 m), 14 Oct.

77 (1 f); WC, 30 May 66 (1 f), 23 June 62 (1 m); RD Summit Rd., 2.1 mi. E. Lovell
Wash, 22 June 77 (1 m);KC Campground, 24 June 79 (1 m);KC, 3340', 5 July 65 (1
m); Little Falls, 20 July 65 (1 m); Las Vegas, 18 Oct. 76 (1, UNLV); 'Pule Springs, 4
Nov. 77 (1 f).

ADDITIONAL RECORDS: Records for all months from 8 Feb. (70, Las Vegas,
in desert area, s) to 18 Nov. (79, Las Vegas, s) mostly for locations cited above.
There are also records for the Sheep, Newberry and Virgin mountains.

Polygonia satyrus satyrus (Edwards).

Less widespread than P. zephyrus reflecting the local distribution of its probable
foodplant, Urtica holosericea. The 2 broods fly from June to August and in October.
The latter brood overwinters and flies again in March and April. The overwintering
brood is darker brown and more yellowish beneath than the summer brood.

SPECIMENS EXAMINED: (29) Lost Creek, 23 Mar. 77 (1 m, CSL); WC, 27
Apr. 77 (1 m, 1 f), 5 June 79 (1 m, 2f), 11 June 79 (1 m, 1 f), 22 June 68 (1 m), 22 June

78 (2 f), 1 July 77 (4 m), 20 July 77 (2 m, 2 f), 20 July 78 (1 m), 8 Aug. 71 (1 f), 13 Oct.
78 (1 m); KC, 20 mi. W. U.S. 95, 14 June 77 (1 f); KC Campground, 24 June 79 (1
m); WC, 1 July 77 (1 m), 14 Oct. 77 (1 m), 26 Oct. 77 (2 m); Deer Creek, 21 Aug. 71
(1, UNLV).

ADDITIONAL RECORDS: WC, 13 June 77, 22 June 78 (s), 29 June 63 (KR),
28 July 77 (s); Little Falls, 22 June 62, 26 June 63, 4 July 72, 5 July 62 (JFL); KC,
9500', 1 July 72 (JFL); Clark Canyon, 10 July 75 (CSL); Charleston Mts., 31 July 65
(RES); Mack’s Canyon, 28 Aug. 75 (CSL).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Pen-
stamon palmeri (Scrophulariaceae); Marrubium vulgare (Lamiaceae); Melilotus
albus (Fabaceae); Salix lasiolepis (Salicaceae); Chrysothamnus sp. (Asteraceae).

Polygonia zephyrus (Edwards).

Locally common above 6000' to as high as 11,200'. It flies in 2 broods, the largest
numbers appearing in September and October representing the overwintering
population which again flies in March and April. A summer brood flies from late
May to August. The latter consists of larger individuals which are more yellow-
orange above and browner beneath than the fall brood.

40

J. Res. Lepid.

SPECIMENS EXAMINED: (65) KC, 17.1-20.0 mi. W. U.S. 95, 22 Mar. 77 (2
m), 28 June 78 (1 m), 25 Aug. 78 (1 m), 26 Sept. 77 (1 m, 2 f), 27 Sept. 78 (4 m, 3 f), 3
Oct. 77 (1 f), 12 Oct. 77 (1 m); Willow Spring, 26 Mar. 78 (1 f, CSL); Deer Creek, 31
May 77(1 m); WC, 22 June 78(1 m); Deer Creek Rd., 2.8 mi. N. KC, 24 June 79(1
m), 26 June 79 (1 f); KC Campground, 24 June 79 (1 m), 8 Oct. 77 (1 m); KC Ski
Run, 26 June 79(1 m), 10 July 79 (2 m), 12 July 78 (1 m), 20 July 65 (1 m), 27 Sept.
78 (4 m), 28 Sept. 77 (4 f), 8 Oct. 77 (2 m); KC, 8000-9000', 5 July 65 (2 m, 1 f), 13
July 69 (1 m), 25 July 65 (1 f); LC, 8000-9000', 5 July 76 (1 m, 1 f, DM), 29 July 77 (1
m); Little Falls, 13 July 65 (1 f), 20 July 65 (1 m, 1 f); ridge above KC, 11,200', 15
July 79 (1 m), 19 July 78 (1 m, 3 f); Echo Canyon, 28 Sept. 77 (1 m, 2 f), 8 Oct. 77 (3
m), 23 Oct. 66 (1 m); KC, 9 Oct. 66 (3 m); RD, 15 Oct. 73 (1 f, UNLV).

ADDITIONAL RECORDS: KC Ski Run, 24 June 79, 18 July 79 (s); Little Falls,
29 June 63 (KR), 8 Aug. 62 (JFL); KC, 1 July 72 (JFL), 8 Aug. 74 (JAS), 26 Sept. 65
(PJH); Deer Creek Rd., 2.8 mi. N. KC, 10 July 79 (s); Cathedral Rock, 13 July 72
(DEA); Charleston Peak Trail, 11,200', 14 July 77 (s); LC, 8 Aug. 74 (JAS), 17 Aug.
63 (JFL), 23 Sept. 75 (s); Charleston Mts., 10-12 Aug. 63 (JFE); Echo Canyon, 9
Oct. 66, 15 Oct. 77 (s).

LARVAL FOODPLANTS: Ribes cereum Dough (Saxifragaceae): KC, 31 Aug.
67 (larvae. Shields et al. 1969); ridge above KC, 11,200', 19 July 78 (ovip., 2x, on
main stem at base of branch, 12:00 PST; plant adjacent to snowbank, leaves
unopened).

ADULT RESOURCES: Chrysothamnus sp. (Asteraceae).

Chlosyne palla vallismortis (Johnson).

Not common in Kyle Canyon between 6700' and 8000'. Records are for mid May
and mid June to mid July. The flight season appears to be very short in some years. I
took specimens on 5 and 7 July 1977 but none were seen by me on 17 June or 15 July
in the same area. Lawson did not see it on 30 June and Mullins took it on 2 July. In
1965, however, specimens were taken over a 5 week period.

We tentatively assign the Spring Range population to vallismortis of the Panamint
Mountains of California. The populations are similar in their large size although that
of the Spring Range is slightly redder in color.

SPECIMENS EXAMINED: (85) Charleston Mts., 14 May 34 (10 m, 1 f,
LACM); WC, 11 June 79 (1 m); KC, 6800', 13 June 65 (1 m); KC Campground, 20
June 79 (17 m), 24 June 79 (15 m, 1 f), 26 June 79 (9 m, 2 f), 5 July 77 (9 m), 7 July 77
(5 m, 1 f), 20 July 65 (1 m); 1 mi. N. KC on Deer Creek Rd., 2 July 77 (6 m, 2 f, DM);
Deer Creek, 11 July 65 (1 m); KC, 7500', 15 July 74 (1 m, 2 f).

ADDITIONAL RECORDS: KC, 6768', 30 June 50 (FWP); KC, 6770', 1 July 50
(Howe 1975); between KC and Deer Creek, 7000', 1 July 50 (FWP); KC, 8000', 5
July 65 (s).

ADULT RESOURCES: Viguiera multiflora (Asteraceae).

Chlosyne neumoegeni neumoegeni (Skinner).

Common to abundant, mostly in desert washes, to 5000'. The flight period
involves a large spring brood from early March (one January and two February
records) to mid May with a peak in mid April and an irregular and sometimes absent
(depending on rainfall) fall brood from early September to early November. Some
colonies may occasionally produce a small second spring brood (1978, Newberry
Mountains).

SPECIMENS EXAMINED: (422) Specimens or other records are for the dates

19(1): 1-63, 1980(81)

41

from 3 Mar. (63, RR, s) to 7 June (78, CC, s) and 3 Sept. (61, Blue Diamond, KR) to
2 Nov. (77, 1,8 mi. E. Blue Diamond, 2 f). There are records for 21 Jan. 64, 6 Feb. 77
and 8 Feb. 70.

LARVAL FOODPLANTS: Acamptopappus shockleyi Gray (Asteraceae): 0.5
mi. S. Red Spring, 27 Apr. 79 (ovip. on flower buds, 09:15 PST); 12 mi. N. 1-15 on
U.S. 93, 15 May 77 (larvae on disc flowers). Macaeranthera tortifolia (Gray) Cronq.
& Keck (Asteraceae); XM, 22 Apr. 79 (ovip. lower surface of lower leaves near
center of plant, 10:30 PST); 0.5 mi. S. Red Spring, 21 Apr. 79 (larvae); 9 mi. W.
Davis Dam, 3 May 78 (small larvae on disc flowers); XM, 12 May 78 (larvae mostly
on disc flowers, also feeding on ray flowers after disc consumed); 12 mi, N. 1-15 on
U.S. 93, 15 May 77 (larvae on disc flowers).

ADULT RESOURCES: Eriogonum fasckulatum (Polygonaceae); Menodora
spinescens (Oleaceae); Cryptantha sp,,Amsinckia tessellata (Boraginaceae) ;L3/ciiim
andersonii (Solanaceae); Encelia farinosa, Behhia juncea, Baileya muitiradiata,
Baileya pleniradiata, Perityle emoryi, Eriophyllum wallacei, Chaeriactis fremontii,
Pectis papposa, Haplopappus sp., Acamptopappus shockleyi, Dyssodia cooperi,
Chrysothamnus sp., Baccharis sp., Senecio douglasii (Asteraceae).

Chlosyne lacinia crocale (Edwards).

Common to abundant in agricultural areas of the Las Vegas and Moapa valleys;
very rare elsewhere with records for Virgin Valley, Corn Creek and Grapevine
Canyon. The flight period extends from mid April to late October involving 2-4
broods depending on availability of the larval foodplant, largely Helianthus annum.
The latter grows commonly along edges of fields and may be cut or burned at times
depending on the particular- farmer’s practices.

All three forms occur in Clark County with the form“rafescens” predominating,
comprising 70% of the population and forms “crocale” and “nigrescens” with 26%
and 4%, respectively.

SPECIMENS EXAMINED: (342) Numerous records from Moapa and Las
Vegas valleys between 13 Apr. (78) and 23 Oct. (77). Records away from here are
Mesquite, 13 Apr. 78 (s), 18 Sept. 72 (1 f, NSM); Bunkerville, 17 May 78 (1 m,
NSM); GC, 21 Aug. 78 (1 f), 27 Sept. 77 (s); CN, 7 Oct. 77 (1 m).

LARVAL FOODPLANTS: Helianthus annuus L. (Asteraceae): Hidden Valley,
23 Aug. 77, 29 Sept. 77 (many larvae). Xanthium strumariumh. (Asteraceae): MF,
13 Sept. 77 (many larvae).

ADULT RESOURCES: Tamarix pentandra (Tamaricaceae); Polygonum lapa-
thifolium (Polygonaceae); Convolvulus sp. (Convolvulaceae); Heliotropium curas-
savicum (Boraginaceae); Medicago satiua (Fabaceae); Viguiera deltoidea, Helianthus
annuus, Baileya pleniradiata, Baileya muitiradiata, Baccharis sp., Pluchea sericea
(Asteraceae).

Chlosyne californica (Wright).

Locally common in some years in the Newberry Mountains; rare elsewhere
occurring as far north as Willow Creek in the Spring Range. It flies in 2 broods from
mid March to early June and early September to late October, The timing and size
of the second brood may depend on summer rainfall.

SPECIMENS EXAMINED: (167) GC, 23 Mar. 78 (2 m, NSM; 4 f), 8 Apr. 77 (1
m), 12 Apr. 78 (3 m, 1 f, NSM; 9 m, 3 f), 15 Apr. 78 (3 m, 5 f), 20 Apr. 78 (1 f, NSM; 1
m, 1 f), 3 May 78 (1 f), 5 May 77 (1 f, NSM; 2 m, 2 f), 12 May 78 (1 f), 1 June 78 (2 f,
NSM), 26 Sept. 78 (1 m, 2 f, NSM; 1 m), 27 Sept. 77 (27 m, 22 f), 4 Oct 77 (4 m, 6 f),

42

J. Res. Lepid.

12 Oct. 77 (2 f), 25 Oct. 77 (1 f); MM, 12 Apr. 69 (2 m, NSM), 26 Sept. 68 (1 f,
UNLV); XM, 15 Apr. 78 (1 m, 1 f), 22 Apr. 79 (10 m, 3 f), 3 May 78 (1 f), 6 May 79 (2
m, 5 f), 23 May 78 (1 m), 1 June 78 (2 m, NSM), 27 Sept. 77 (3 m, 1 f); 9 mi. W. Davis
Dam, 15 Apr. 78 (1 f), 20 Apr. 78 (2 f, NSM), 12 May 78 (1 m), 23 May 78 (1 m);
Sacatone Wash, 15 Apr. 78 (1 m, 2 f); 1 mi. W. Nipton Pass, 4 May 78 (2 m); Bridge
Canyon, 12 May 78 (1 f), 26 Sept. 78 (1 m, NSM); 2 mi. N. RC, 27 May 78 (2 f); CP
Rd., 1.5 mi. S. Pahrump Rd., 28 May 78 (1 f); Crescent Peak, 21 Sept. 72 (3 m, 5 f,
NSM); 2.8 mi. E. Nelson, 27 Sept. 77 (1 m), 25 Oct. 77 (1 f).

ADDITIONAL RECORDS; XM, 18 Mar. 72 (JFL), 12 Apr. 78, 20 Apr. 78, 5
May 77, 12 May 78, 4 Oct. 77 (s); 9 mi. W. Davis Dam, 12 Apr. 78, 22 Apr. 79, 3 May
78 (s); Bridge Canyon, 3 May 78 (s); CP Rd., 0.6 mi. S. Pahrump Rd., 19 May 78 (s);
WC, 6 June 78 (s); Ash Spring, 30 June 69 (JFE, OS); Blue Diamond, 3 Sept. 61
(KR); 0.5 mi. E. Boulder City, 3 Oct. 77 (s).

LARVAL FOODPLANTS: Viguiera deltoidea Gray (Asteraceae): GC, 14 Sept.
77 (larvae, skeletonizing leaves).

ADULT RESOURCES: Eriogonum fasciculatum (Polygonaceae); Amsinckia
tessellata (Boraginaceae); Salvia sp. (Lamiaceae); Viguiera deltoidea, Encelia
farinosa, Bebbia juncea, Perityle emoryi, Eriophyllum wallacei, Chaenactis fremontii,
Haplopappus sp., Baccharis sp., Senecio douglasii, Pluchea sericea (Asteraceae).

Phyciodes texana texana (Edwards).

The only Nevada records are for Grapevine Canyon in April and Cabin Canyon in
May.

SPECIMENS EXAMINED: (2) GC, 12 Apr. 78 (1 m), 20 Apr. 78 (1 m).
ADDITIONAL -RECORDS: GC, 15 Apr. 78 (s); CC, 20 May 79 (s).

Phyciodes tharos (Drury) ssp.

The only Nevada record is from Corn Creek. They belong to the unnamed
southwestern United States-northwestern Mexico population. This taxon was
called distincta by Bauer in Howe (1975). No type was, however, designated. Thus
P. tharos distincta is a nomen nudum. The specimens are identical with material
from Imperial Co., California.

SPECIMENS EXAMINED; (2) CN, 9 Sept. 64 (2 m, NSM).

ADDITIONAL RECORDS: None.

Phyciodes phaon (Edwards).

Very rare with records from mid September to early November for the Las Vegas
area.

SPECIMENS EXAMINED: (9) Las Vegas, Sept. 75 (1 m, ex UNLV), 16 Sept.
77 (1, UNLV), Oct. 75 (1 m, ex UNLV), 8 Oct. 77 (1 f, UNLV), 13 Oct. 76 (1 f), 13
Oct. 77 (1 m, UNLV), 15 Oct. 78 (1 f), 7 Nov. 77 (1 f, UNLV); Clark Co., 1 Oct. 76 (1
f, ex UNLV).

ADDITIONAL RECORDS: None.

Phyciodes pallida barnesi Skinner.

A single specimen for this taxon exists for Clark County. Populations are known
for adjacent Lincoln Co., Nevada, (GT A) and Washington Co., Utah (Tidwell and
CaUaghan 1972).

SPECIMENS EXAMINED: (1) CN, 15 July 64 (1 m, NSM).

ADDITIONAL RECORDS: None.

Phyciodes mylitta mylitta (Edwards),

19(1): 1-63, 1980(81)

43

A single specimen from Kyle Canyon referrable to the nominate race was
examined. Another record from the same area is the only other one known from the
county. These may represent a small breeding colony as its known foodplant,
Cirsium, is quite common here.

SPECIMENS EXAMINED: (1) Charleston Park, 30 June 77 (1 f, CSL).

ADDITIONAL RECORDS: Charleston Park, 29 June 63 (KR).

Thessalia leanira alma (Strecker).

Uncommon locally at middle elevations between 3400' and 6100' especially in the

southern portion of the Spring Range in the Cottonwood Pass area. Records extend
from mid April to mid May and a fresh individual seen on 13 June.

SPECIMENS EXAMINED: (184) Blue Diamond, 12 Apr. 69 (1 m, NSM); CP
Rd., 0.6-1. 8 mi. S. Pahrump Rd., 16 Apr. 77 (3 m, 2 f), 16 Apr. 79 (1 m), 20 Apr. 77
(4 m), 22 Apr. 78 (6 m), 29 Apr. 78 (1 m), 29 Apr. 79 (26 m, 13 f), 2 May 77 (2 m), 2
May 78 (7 m, 2 f), 3 May 78 (3 m, 1 f, NSM), 6 May 78 (1 m, NSM), 1 1 May 78 (4 m, 2
f), 13 May 79 (4 m), 19 May 78 (3 m); 3 mi. W. Blue Diamond, 20 Apr, 77 (2 m, 5 f);
0.5 mi. S. Red Spring, 21 Apr. 79 (17 m, 2 f), 29 Apr. 79 (30 m, 4 f), 2 mi. E. MS, 24
Apr. 77 (1 m, 1 f); WC, 27 Apr. 77 (1 m, NSM), 29 Apr. 77 (1 m, NSM); CC, 27 Apr.
78 (1 m), 10 May 78 (1 m); Pine Creek, 2 May 77 (1 m); KC, 6.6 mi. W. U.S. 95, 2
May 77 (1 f), 2 May 78 (4 m), 1 1 May 78 (3 m); 1 mi. W. WC, 8 May 78 (1 m), 25 May
78 (2 m); KC, 9.3 mi. W. U.S. 95, 13 May 78 (1 m); 0.5 mi. E. MS, 13 May 79 (12 m),
19 May 78 (3 m); 1.5 mi. E. MS, 13 May 79 (1 m); RC, 27 May 78 (2 m); Wilson Pass,
ex larva emerged 21 Apr. 78 (1 f).

ADDITIONAL RECORDS: White Rock Spring, 2 May 78 (s); RC, 4 May 78, 27
May 78 (s); 2 mi. N. RC, 4 May 78, 27 May 78 (s), WC, 13 June 77 (s).

LARVAL FOODPLANTS: Castilleja chromosa A. Nels. (Scrophulariaceae):
RD Canyon, 25-27 Mar. 75 (larvae, JB); Willow Spring, 29 Mar. 78 (larvae feeding
on flowers and leaves); Wilson Pass, 29 Mar. 78 (larvae feeding on flowers).

ADULT RESOURCES: Menodora spinescens (Oleaceae); Baileya multiradiata,
Eriophyllum wallacei, Chaenactis fremontii, Acamptopappus shockleyi, Senecio
douglasii, Senecio multilohatus (Asteraceae).

Poladryas minuta arachne (Edwards).

Common very locally on the west slope of the Spring Range at elevations of 4500'
to 6000'. The flight periods of the 2 broods are late April to late June and early

September to mid October.

Southern Nevada material with its two-toned orange ground color is clearly within
the concept of P. m. arachne, especially like northern Arizona populations. In
comparison with a series from Flagstaff, Coconino Co., Arizona, southern Nevada
material differs in having slightly less black on the basal half of both wings above
and below, the ground color of the primaries below is more even (less two-toned)
and the light areas of the secondaries below are less creamy. In fact, these light
areas are the whitest of all populations of minuta examined except those of southern
Arizona, P. m. gilensis (Holland), to which southern Nevada specimens are
comparable in this aspect. The use of the specific name minuta is based on the
studies of Scott (1974).

SPECIMENS EXAMINED: (51) CP Rd., 0.6-1.8 mi. S. Pahrump Rd., 29 Apr.
78 (1 m), 2 May 78 (3 m), 3 May 78 (3 m, NSM), 1 1 May 78 (2 m), 13 May 79 (1 m),
28 May 78 (1 m); 0.5 mi. E. MS, 13 May 79 (12 m), 19 May 78 (3 m); CC, 20 May 78

(1 f); 5 June 79 (1 f, CSL); Trout Canyon Ranch, 14 June 77 (6 m), 18 Sept. 77 (Im);

44

J. Res. Lepid.

6.1 mi. N. Spring Mountain Youth Camp,rd. to Trout Canyon, 14 June 77 (1 m);RD
Summit Rd., 1.5 mi. E. Lovell Wash, 22 June 77 (1 f); MS, 6 Sept. 69 (2 f, NSM); 1
mi. W. MS, 28 Sept. 78 (1 m), 1 Oct. 77 (9 m, 1 f), 11 Oct. 77 (1 f).

ADDITIONAL RECORDS: 1 mi. W. WC, 8 May 78 (s); 1:5 mi. E. MS, 13 May
79 (s).

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); Viguiera
multiflora, Erigeron sp. (Asteraceae).

Euphydryas chalcedona kingstonensis Emmel & Emmel.

One desert population of this species is known in the Newberry Mountains. It flies
in a large spring brood from mid March to early June and a small brood depending
on summer rains in September. Specimens are within the range of variation of
southern California populations.

SPECIMENS EXAMINED: (269) GC, 8 Apr. 77 (1 f, CSL; 1 m, 1 f), 12 Apr. 78
(lm,NSM), 15 Apr. 78 (2 m), 20 Apr. 78 (5 m, NSM; 12 m, 1 f), 3 May 78 (21 m, 2 f),
12 May 78 (10 m, 5 f), 23 May 78 (1 m), 1 June 78 (1 m, 1 f, NSM), 14 Sept. 77 (3 m,
5 f); XM, 12 Apr. 78 (6 m, NSM; 13 m), 15 Apr. 78 (4 m, 1 f), 20 Apr. 78 (12 m, 3 f),
22 Apr. 79 (35 m, 2 f), 3 May 78 (13 m, 3 f), 5 May 77 (1 m, CSL), 6 May 79 (49 m, 26
f), 12 May 78 (11 m, 13 f), 23 May 78 (4 m).

ADDITIONAL RECORDS: XM, 18 Mar. 72 (JFL), 1 June 78, 19 Aug. 79 (s);
GC, 23 Mar. 78 (CSL, s), 27 Sept. 77 (s); Sacatone Wash, 15 Apr. 78 (s).

LARVAL FOODPLANTS: Keckiella antirrhinoides Benth. ssp. microphyllus
(Gray) Keck (Scrophulariaceae): XM, 12 Apr. 78 (last instar larvae feeding on
leaves).

ADULT RESOURCES: Eriogonum fasciculatum (Polygonaceae); Amsonia
tomentosa (Apocynaceae); Phacelia fremontii, Phacelia uallis-mortae, (Hydrophyl-
laceae); Amsinckia tessellata (Boraginaceae); Salvia sp. (Lamiaceae); Baileya
multiradiata, Senecio douglasii (Asteraceae).

Euphydryas anicia morandi Gunder.

This distinctive taxon is locally common in the Spring Range especially in the
meadows on the ridge to Charleston Peak and above the ski area in Lee Canyon.
Most records are for above 10,000' although it is occasionally taken as low as 6800'.
The flight period of the single brood is usually from late June to mid July; a number
of early (1940 and before) collections, however, were made in early May to early
June.

SPECIMENS EXAMINED: (168) Charleston Mts., 7 May 29 (1 f, LACM), 13
May 34 (1 m, LACM), 14 May 34 (17 m, 1 f, LACM), 19 May 34 (2 m, LACM); Mt.
Charleston, 1 June 33 (4 m, LACM); KC Campground, 30 June 77 (1 m, CSL); LC
above ski area, 2 July 77 (6 m, 1 f, DM), 5 July 76 (6 m, 6 f, NSM), 6 July 66 (8 m, 7 f,
DM); Charleston Peak trail above KC, 10, 400--1 1,300', 14 July 77 (27 m, 5 f), 15
July 79 (43 m, 14 f), 19 July 78 (10 m, 8 f).

ADDITIONAL RECORDS: Mt. Charleston, 7 May 29 (DE), 10 May 36 (DE),
10 May 40 (DE), 26 June 36 (DE); trail to Charleston Peak, 10,800-11,500', (TCE,
OS); Mt. Charleston, 10,500-11,360', (JFL); KC, 10,500-11,000', 10-16 July 28
(type series, Gunder 1928).

ADULT RESOURCES! Erysimum asperum (Brassicaceae); Taraxacum of-
ficinale (Asteraceae).

Euphydryas anicia (Doubleday) ssp.

A population which does not seem to fit any described subspecies of anicia flies in

19(1): 1^63, 1980(81)

45

the Virgin Mountains between 4000' and 5000' in April and May. Its relationships
appear towards Rocky Mountain populations.

SPECIMENS EXAMINED: (17) CC, 8 Apr. 79 (1 m, 1 f), 13 Apr. 74 (1 m,
NSM), 14 Apr. 77 (2 m, 1 f, CSL), 27 Apr. 78 (2 m), 10 May 78 (1 m), 18 May 78 (1
m), 20 May 78 (1 m), 20 May 79 (6 m).

ADDITIONAL RECORDS: None.

Speyeria zerene carolae (dos Passes & Grey).

This endemic is locally common on the east slope of the Spring Range from 6700'
to 10,800', but mostly below 7500' in Kyle Canyon. There is but one west slope
record for the low elevation of 5000' in Lovell Wash. The flight period for the single
brood is from mid June to early September with a peak in mid to late July. Very late
records are in late September and mid October.

SPECIMENS EXAMINED: (179) Numerous records with the earliest date on
17 June (66, LC and Deer Creek, s). Latest record is 1 1 Oct. (77, KC Campground,
s). The only record away from the east slope is Lovell Wash, 5000', 17 June 62 (s).

ADULT RESOURCES: Erysimum aspermn (Brassicaceae); Apocynum andro-
saemifolium (Apocynaceae); Rosa woodsii (Rosaceae); Lupinus sp. (Fabaceae);
Angelica scabrida (Apiaceae); Chaenactis sp., Cirsium sp. (Asteraceae).

Euptoieta claudia (Cramer).

Very rare with four known records for the county.

SPECIMENS EXAMINED: (3) Ash Springs, 3800', 19 Mar. 72 (1, JFL); CN,

22 June 68 (2, JFL).

ADDITIONAL RECORDS: Sawmill Canyon, 1 July 69 (1969 summ.); 1 mi. N.

CP, 23 Aug. 75 (JB).

Family Heliconiidae

Agraulis vanillae incarnata (Riley).

The only state records are a stray caught in the authors’ yard in Las Vegas and two

sight records.

SPECIMENS EXAMINED: (1) Las Vegas, 19 June 68 (1 f).

ADDITIONAL RECORDS: 0.5 mi. N. Red Spring, 29 Apr. 79 (s); XM, 6 May

79 (s).

Family Danaidae

Danaus plexippus plexippus (Linnaeus).

Not uncommon especially in mesic canyons, near springs and, after July, at the
higher elevations. Records are for every month except January but it is most
abundant from mid June to mid November. There are many Las Vegas records from
August to November and a few desert records in early spring and fall. High elevation
records are from early July to mid November. A concentration of about 40
individuals was at Tule Springs on 6 Sept. 75 and over 200 were seen in Pine Creek
on 5 Oct. 77.

SPECIMENS EXAMINED: (124) Records from numerous locations through-
out the county between 6 Feb. (77, 1.5 mi. S. Davis Dam, s) to 25 Dec. (65, CN, s).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Stanleya pin-
nata (Brassicaceae); Sarcostemma hirtellum (Asclepiadaceae); Eriodictyon angusti-
folium (Hydrophyllaceae); Marrubium vulgare (Lamiaceae); Cowania mexicana
(Rosaceae); Medicago satiua, Melilotus albus (Fabaceae); Angelica scabrida
(Apiaceae); Viguiera deltoidea, Helianthus annuus, Solidago spectabilis, Chryso-
thamnus sp., Baccharis sp., Senecio douglasii, Cirsium sp. (Asteraceae).

46

J. Res. Lepid.

Danaus gilippus strigosus (Bates).

Locally common in certain low canyons (Pine Creek, Grapevine Canyon) but
found in most areas of the county to at least 8500'. There are records for every
month except March but most records are from mid June to early November.

SPECIMENS EXAMINED: (229) Numerous records through most of the year
from many locations; desert records are few. High elevation records are from 25
May (78, WC, s) to 22 Aug. (77, KC Ski Run, s).

LARVAL FOODPLANTS: Sarcostemma hirtellum (Gray) R. Holm (Asclepia-
daceae): GC, 20 Apr. 78 (ovip. on stems, 13:30 PST), 3 Aug. 77 (mature larva), 27
Sept. 77 (ovip. single eggs on stems between leaf bases, 10:40 PST, 11:00 PST).
Asclepias subulata Dene, in A. DC. (Asclepiadaceae): 9 mi. W. Davis Dam, 3 May 78
(ovip. beneath flower heads, 10:00 PST).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Tamarix pen-
tandra (Tamaricaceae); Eriogonum fasciculatum (Polygonaceae); Apocynam andro-
saemifolium (Apocynaceae); Sarcostemma hirtellum (Asclepiadaceae); Eriodictyon
angustifolium (Hydrophyllaceae); Acacia greggii, Prosopis glandulosa, Prosopis
pubescens, Melilotus albus (Fabaceae); Viguiera deltoidea, Helianthus annuus,
Bebbia juncea, Solidago spectabilis, Chrysothamnus sp., Baccharis sp., Senecio
douglasii, Tetradymia canescens, Pluchea sericea, Cirsium sp. (Asteraceae).

Family Satyridae

Cyllopsis pertepida dorothea (Nabokov).

The population in the Virgin Mountains is the only known colony for the state
except for one in Meadow Valley Wash in Lincoln Co. (1968 summ.). It flies in 2
broods from late May to late June and in September and October. The taxonomy
follows Miller (1974).

SPECIMENS EXAMINED: (23) CC, 29 May 78 (3 m), 7 June 78 (2 f, NSM; 2
m), 8 June 78 (1 m, 1 f), 26 June 78 (5 m, 4 f), 12 Sept. 74 (2 m, NSM), 20 Sept. 72 (1
f, NSM), 11 Oct. 78 (2 m).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Eriodictyon angustifolium (Hydrophyllaceae); occasional-
ly visit water.

Coenonympha California California Westwood.

A single specimen from the Spring Range is the only southern Nevada record.

SPECIMENS EXAMINED: (1) WC, 22 June 78 (1 m).

ADDITIONAL RECORDS: None.

ADULT RESOURCES: Rorippa nasturtium-aquaticum (Brassicaceae).
Coenonympha ochracea brenda Edwards.

Uncommon in the Spring Range above 6500' but mostly above 7500' in open
areas. The single brood usually flies from mid June to early August although there

are May records.

Spring Range material along with certain other Nevada material (White Pine, Nye,
Lander counties) fall within the concept of Utah brenda although there are minor
differences in pattern (see also Brown 1964).

SPECIMENS EXAMINED: (48) Charleston Mts., 12 May 34 (7 m, 5 f, LACM);
KC, 8000', 12 June 66 (1 m), 5 *July 65 (4 m, 2 f); above Cathedral Rock
Campground, 17 June 77 (3 m, 1 f); KC Campground, 22 June 78 (1 m); KC Ski
Run, 24 June 79 (1 f), 5 July 77 (1 m); KC, 7680', 27 June 65 (1 m); Clark Canyon, 2

19(1): 1-63, 1980(81)

47

July 75 (1 m, NSM); Deer Creek, 11 July 65 (2 m); Little Falls, 13 July 65 (1 m, 1 f);
Charleston Peak trail, 10,900-11,300', 14 July 77 (1 m, 1 f), 15 July 79 (1 m), 19 July
78 (4 m, 3 f); KC, 9000', 25 July 65 (1 m), 26 July 64 (1 m), 27 July 66 (1 m, 2 f); LC,
8700', 26 July 65 (1 m).

ADDITIONAL RECORDS: Charleston Mts., 14 May 34 (CDF), 6 Aug. 61
(RES); KC, 7100', 11 June 65, 17 June 66 (s); LC, 17 June 66 (s); KC Campground,

21 June 66, 13 July 65 (s); KC, 7600', 22 June 63, 26 June 63, 29 June 63, 5 July 62
(JFL); KC Ski Run, 22 June 78 (s); KC, 10,000-10,500', 24 June 68 (JFE, OS); KC,
20 mi. W. U.S. 95, 28 June 77, 6 July 77 (s); Little Falls, 29 June 63 (KR), 4 July 72
(JFL); KC, 10,500-11,200', 1 July 72 (JFL); Deer Creek Rd., 7000', 1 July 50
(FWP); Charleston Peak trail, 9000', 14 July 77 (s).

ADULT RESOURCES: Lupinus sp. (Fabaceae); Senecio multilobatus
(Asteraceae).

Cercyonis sthenele (Boisduval) ssp.

Locally common mostly above 5000' but occasionally as low as 3300' on
sagebrush flats or in chaparral vegetation. The single brood flies from mid June
through late August.

All Nevada material was referred to previously by Emmel (1969) as C. s. paulus
(Edwards). Clark County material and that of adjacent areas do not have the
extensive white overscaling beneath as is typical of paulus and resemble certain
material from the desert mountains of southern California. These populations most
closely resemble the Rocky Mountain C. s. masoni Cross but probably are distinct
and deserve to be named.

SPECIMENS EXAMINED: (60) KC, 15 mi. W. U.S. 95, 20 June 79 (1 m); WC,

22 June 78 (1 f), 20 July 77 (1 m, 3 28 July 77 (1 m, 1 f); WC, 1 July 77 (2 m), 4 July

62 (1 m, 1 f), 20 July 78 (1 m, 2 f), 28 July 77 (2 m, 8 f); KC, 3340', 5 July 65 (3 m);
KC, 5680', 5 July 65 (1 m), 18 July 65 (5 m); Lovell Wash, 6100', 14 July 62 (2 m, 1
f); KC Campground, 15 July 77 (5 m), 25 July 77 (3 m); KC, 17.1 mi. W. U.S. 95, 15
July 77 (2 m, 2 f); Lovell Wash, 18 July 62 (1 f); Cold Creek Ranger Station, 20 July
77 (2 m, 1 f); RD Summit Rd., 1. 5-2.1 mi. E. Lovell Wash, 27 July 77 (3 m, 4 f).

ADDITIONAL RECORDS: KC Campground, 20 June 79, 25 July 65, 1 Aug. 77
(s); RD Summit Rd., 1.5 mi. E. Lovell Wash, 22 June 77 (s); CC, 26 June 78, (s);KC,
14.5 mi. W. U.S. 95, 28 June 77, 27 July 77 (s); KC Ranger Station, 29 June 63
(KR); WC, 29 June 63 (KR); KC, 6770', 30 June 50 (FWP); MS, 30 June 63 (KR),
30 June 69 (JFE, OS); Deer Creek Rd., 1 July 50 (FWP); mouth Sawmill Canyon, 1
July 69 (JFE, OS); Mt. Charleston, 9 July 74 (CDF); LC, 9 July 74 (CDF); Lovell
Wash, 10 July 75 (CSL); KC, 17.1 mi. W. U.S. 95, 1 1 July 77 (s); WC, 20 July 78, 5
Aug. 77 (s); KC, 6800', 18 July 65 (s); KC, 14 mi. W. U.S. 95, 25 July 65 (s);
Charleston Mts., 31 July 65 (JL, RES); KC, 7400', 12 Aug. 63, 18 Aug. 63 (JFL);
lower KC, 24 Aug. 75 (JB).

ADULT RESOURCES: Clematis ligusticifolia (Ranunculaceae); Eriodictyon
angustifolium (Hydrophyllaceae); Tetradymia canescens, Cirsium sp. (Asteraceae).

Dubious Records

At least five species have been attributed to the Clark County butterfly fauna
based on misidentifications or poor knowledge of their distribution.

Amblyscirtes eos (Edwards).

The record for this species (1972 summ.) refers to Pholisora alpheus {fide JFL).
This species should be deleted from the state list.

48

J. Res. Lepid.

Ochlodes sylvanoides (Boisduval).

Herlan reported to us that a pair of this species was taken in the Overton area on
29 Sept. 66 and one of these was indicated in the Nevada State Museum catalogue.
This specimen was located and proved to be a male Polites sabuleti. Ochlodes
sylvanoides is thus apparently unknown in Clark County although it is common
farther north in the state.

Ochlodes agricola (Boisduval).

Herlan reported to us that three males were taken at Warm Springs (Moapa
Valley) on 27 Oct. 65 and these were listed in the Nevada State Museum catalogue.
These specimens could not be located in the museum. We thus consider this species
to be of hypothetical occurrence in Clark County.

Hesperia uncas lasus (Edwards).

This species was reported by Herlan (1972 summ.) for Crescent Peak in the
McCullough Range.

A search of the collection at the Nevada State Museum failed to produce a
specimen. No specimens are listed in their catalogue from Clark County. The
species must thus be considered of hypothetical occurrence in Clark County.
Polites sonora sonora (Scudder).

The taxon is included as hypothetical based on the range “to southern Nevada” by
Howe (1975). We know of no southern Nevada records although the species is
regular in northern Nevada.

Faunal Composition

Clark County’s butterfly fauna totals 125 taxa of 118 species. Of these, one
species appears to have become locally extinct (L. dorcas), if it ever occurred, and at
least 15 are considered non-breeding strays (A. python, H. domicilla, C. asychis, T.
pylades, P. leo, C. philodice, P. sennae, E. mexicana, L. bachmanii, P. texana, P.
tharos, P. pallida, E. claudia, A. vanillae and C. California). We doubt that more than
10 to 15 species will be added to the county list in future years. It is unlikely that
many new resident species will be found although such species as Erynnis afranius
(Lintner) and Hypaurotis crysalus (Edwards) are possibilities in the Virgin Moun-
tains.

The 125 taxa belong to 11 families with Lycaenidae, Nymphalidae and Hesperi-
idae being the most important comprising over 70% of the total fauna (Table 1).
Similar species composition occurs in other areas of western North America {e.g..
Brown et at 1957, Emmel and Emmel 1963, Opler and Langston 1968, Tidwell and
Callaghan 1972).

To further consider the distribution of butterflies in Clark County, four major
habitat groupings were delineated. Desert scrub occurs at low elevations generally
below 4000-5000' and includes high water table associations of mesquite. Riparian
and agricultural habitat includes vegetation which requires a nearly constant water
supply whether it be natural along streams or artificial as urban or cultivated areas.
Middle elevations include areas above the desert scrub to approximately 7000' and
roughly delimited by the Pinon-Juniper woodland. Montane habitat is all areas
above 7000'. For the present section, all species which commonly occur in the
particular habitat are included. In subsequent sections, only those species which
have their distributional center of abundance in a particular habitat will be
considered. Species which commonly range over more than one habitat are termed
widespread.

19(1): 1-63, 1980(81)

49

Family composition in the different habitats shows a varied pattern (Table 1). The
desert scrub habitat is dominated by Pierids and Lycaenids, riparian/agricultural
by Hesperiids and Nymphalids, middle elevations (which contains the most total
species) by Hesperiids and Lycaenids and montane habitat by Hesperiids,
Lycaenids and Nymphaiids. Of the minor families, Papilionids and Satyrids are
most important at the higher elevations and Riodinids at lower elevations. Similar
habitat differences were noted by Emmel and Emmel (1963).

Biogeographical Analysis

The location of Clark County in the northern Mojave Desert and in close proximity
to the Great Basin and Sonoran deserts make a biogeographical analysis of the biota
of interest. Each species was assigned to an element based on the geographical
range of the taxon (Table 2). The butterfly fauna of Clark County exhibits complex
biogeographical relationships (Table 3). Forty -four percent of the species have
distributions mainly in the North American deserts. About one-half are largely
restricted to the Mojave Desert and one-quarter of the others are widespread desert
region species. The remaining species of these have distributions in either the Great
Basin or Sonoran deserts. Almost 41% of the county’s species are widespread in
distribution throughout North America, in western United States or in tropical
regions.

Species with a widespread ecological distribution in the county are mostly of
relatively wide distribution in the west as a whole or in desert regions. Desert scrub
species are largely of Mojave Desert distribution. Mojave Desert and Rocky
Mountain elements dominate at middle elevations. Principally montane species
belong to the western North America and endemic elements.

Phenology and Voltinism

Seasonality and voltinism of the Clark County butterfly fauna were examined on
the basis of number of locality-date records for each 10 day period of each month
(Table 4) and by evaluating wear of specimens. It must be recognized throughout
that annual climatic differences tend to lengthen the apparent flight season and to
smooth out peaks and valleys of abundance.

For the county as a whole, butterflies have been recorded in every month of the
year. Species number and abundance are greatest from May through July although
considerable numbers of species are present in all months from April to October
(Table 4). Approximately equal numbers of species are present in each month from
March to October at low elevations. The number of records, however, is greatest in
April and September-October indicating the peak flight period. The spring peak is
mainly in desert scrub habitats; the peak in fall is of species associated with desert
scrub and cultivated habitats. The flight season at high elevations extends from late
April to October with a peak in June and July.

The Mojave Desert biogeographic element has a strong peak in April and a minor
peak in September. The fall peak may be controlled largely by the occurrence of
summer rainfall. Undoubtedly, the availability of adult and larval food resources
restricts the flight season and number of broods of these, mostly desert scrub
inhabiting, species. Certain species of this group occur in more riparian habitats
and fly throughout the summer or occur at higher elevations and are summer
univoltines.

The southwest desert element shows peaks in summer and fall. This group
contains species which are widespread in the county or occur largely in riparian and

50

«/. Res. LepM.

agricultural habitats and have long flight seasons. The phenology of the widespread
element is similar to that of the southwest desert element but becomes common
somewhat earlier in the year. This group contains species found in desert scrub in
spring and in cultivated areas in summer and fall. The western North America and
toopical elements also show a similar seasonal pattern. They are both composed of a
number of common, widespread species in the county.

The endemic element has a single peak in summer as expected of species
inhabiting montane habitat.

The Great Basin element is most common in spring reflective of its composition of
several spring, univoltines The Rocky Mountain element has a spring and summer
peak. Too few records are available for the other elements to draw conclusions on
them.

Sufficient records exist for 99 species to reach some conclusons on their voltinism
(Table 2). Thirty-nine species appear to be univoltine within the county (Table 5).
Twenty-eight of these occur at middle to high elevations. At middle elevations most
univoltines are vernal with S. behrii, L. weidemeyerii and C. sthenek as summer
univoltines and A. alliae and E. b. ellm as fall univoltines. Montane univoltines
fly in summer except for the spring flying E. brizo. Eight additional univoltines
occur largely m desert scrub. Seven of these are vernal and only E. e. dammersi is
autumnal. There are only two (both autumnal) univoltines in riparian and agricul-
tural habitats. The only widespread univoltine is N. antiopa.

Twenty species appear as bivoltines (Table 5). These include four montane
species {H. comma, H. juba, P. satyrus, P. zephyrus) mth summer and fall broods. At
middle elevations, there are six bivoltines. Four {P. indra, C. spinetorum, C. siva, P.
wdnuta) have spring and summer broods, one (C. shendanii) has spring and fall
broods and one (C. pertepida) has summer and faH'broods. Five desert scrub species
are bivoltine. One (M. leda) occurs in mesquite stands and has summer and fall
broods. The remaining four (P. rudkini, C. neumoegeni, C. California, E. chalcedona)
have regular spring broods and irregular fall broods, apparently dependent upon
summer rains. These species may be univoltine m some years. Papilio rudkini has
occasional emergences during mid summer in some years and C. neumoegeni is
known to have had two successive spring broods. The four bivoltines (0. yuma, A.
campestis, C. nemesis, L. archippus) of riparian and agricultural habitats and the
one widespread bivoltine (VI virginkmis) have summer and fall broods.

Forty Clark County species are considered multivoltine or have continuous
emergences throughout their flight season (Table 5). The number and timing of
broods of some may vaiy from year to year and with elevation. Certain species
included here such as A. rnormo may be single brooded in some colonies but have an
overall pattern which mdicatee several broods. Such patterns need further study
and may be related to the seasonal availability of suitable foodplante.

Nearly 40% of the Clark County butterfly fauna- is univoltine. TMs frequency of
univoltines is intermediate to that reported for several Califomia butterfly faunae
(Shapiro 1975). Analysis of voltinism by habitat (Table 5) presents difficulties due
to the presence of widespread species which either stray to high elevations or have
different breeding strategies under different ecological conditions. For these
reasons, species were separated by principal habitat preference. The overall
pattern is an increase m univoltinism with increasmg elevation (Table 5) ae noted in
jother studies (Shapiro 1974, 1975). Univoltinism is primarily vernal at low (70%)

19(1): 1^63, 1980(81)

51

and middle (62%) ele¥ations and a summer phenomenon (93%) at high elevations.
Vernal univoltinism is the usual situation in long summer faunas (Shapiro 197 5) and
the northern Mojave Desert pattern contrasts sharply with the exception found for a
Sonoran Desert butterfly fauna (Austin 1978). The difference between the two
desert faunas can be accounted for by the lack of a regular summer rainy season in
the Mojave Desert. Multivoliinism in low desert butteiflies is largely limited to
species with desert iiparian affinities where foodplants remain green through the
summer.

Of the important biogeograpMcal entities in Clark County, the Mojave Desert
element has the greatest number of univoltines fTable 6). Species which fly in
spring and/or fall (spring and fall univoltines, spring-fall bivoltines) comprise 68%
of the species in this elemnnt. Thus the p-oup as a whole exhibits adaptation to the
generally favorable springs, often favorable falls and usually unfavorable remainder
of the year that is typical of the Mojave Desert. The Great Basin and Rocky
Mountain elements are also represented by a large proportion of (mainly spring)
univoltines (Table 6).

The southwest desert and widespread elements are largely multivoltine (Table 6).
Most of these species are either widespread m the county or occur m riparian and

Phenology, voltinisni and abundance aie highly variable especially in the lower
desert. These variations, as mentioned above, are ultimately due to various climatic
factors, particularly rainfall and proximateiy to the resultant condition of the
vegetation. Warm temperatures in late winter promote early emergence of a number
of species. Later in spring, this early flight may be interrupted or terminated

Acknowledgements: Numerous people were of great help to us during the course
of this study. We first thank all of those (listed in the introduction) w-ho have sent
records and/or allowed us to examine and/or borrow specimens in their care. The
completeness of coverage herein could not have been attained without their help. In
addition, we would like to extend special thanks to those who have also helped in
other ways. First of all, Scott Miller and Gloria Harjes made the complete collection
of butterflies at the Nevada State Museum in Carson City available to us for study
and extended various othei courtesies to the senior author while he was in Caison
City. Difficult groups were examined by various people who gave their opinion as to
the taxonomy of Clark County specimens. We are grateful for their help. Final
taxonomic decisions were made by the senior author and any errors in such must
thus be his. Those who helped in this regard include: D. Bauer (various groups), J.
Burns {Erynnis), J. F. Emmel (various groups), B. McGuire (Hesperia), J. Masters
(Enpkydiyas), R. Mattoni (Euphilotes), D. Mullins (Euphydryas, Poladryas), K.
Roever (various groups), O. Shields (Euphilotes), and R. Stanford (various
sliippers). W. E. Niles and A. Pinzl made most of the plant determinations. Anna
Stanley typed the final two drafts of the ms and without her patience this may have
never been typed. A final thanks is due to Chuck Lawson for his interest in Nevada
butterflies and his companionship in the field.

52

J. Res. Lepid.

Literature Cited

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no. 45.

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HARJES, G. J. 1980. Checklist of the butterflies of Nevada. Nevada State Museum,
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53

HOWE, W. H. 1975. The butterflies of North America. Doubleday, Garden City, NY
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54

«/. Res. Lepid.

TABLE 1. Composition of the Clark County, Nevada, butterfly fauna
by families.

County

{%)

Desert

Scrub

(%)

Riparian/

Agriculture

(%)

Middle

Elevations

(%)

Montane

(%)

Megathymidae

1.6

2.4

0.0

3.1

0.0

Hesperiidae

22:2

11.9

26.8

21,9

22.7

Papilionidae

4.8

2.4

2.4

4.7

4.5

Pieridae

12.7

26.2

17.1

15.6

13.6

Riodinidae

3.2

4.8

4.9

1.6

0.0

Lycaenidae

24.6

31.0

17.1

26.6

22.7

Libytheidae

0.8

2.4

2.4

1.6

0.0

Nymphalidae

24.6

16.7

22.0

18.8

31.8

Heliconiidae

0.8

0.0

2.4

0.0

0.0

Danaidae

1,6

2.4

4.9

3.1

2.3

Satyridae

3.2

0.0

0.0

3.1

2.3

Total Species

126

42

41

64

44

TABLE 2. Biogeographic
Clark County,

affinities, voltinism and habitat

Nevada, butterflies.

data for

Taxon

Biogeographical
Affinity ^

Voltinism^

Primary,

Habitaf^

Agathymus alliae ssp.

MD

U

(F)

ME

Megathymus yuccae navajo

MD

U

(S)

DS

Lerodea eufala

WS

M

RA

Atrytonopsis python

SWD

-

-

Ochlodes yuma

MD

B

(SU,F)

RA

Atalopedes campestris

WS

B

(SU,F)

RA

Polites sabuleti ssp.

WNA

M

RA

Polites draco

RM

_

_

Hesperia comma harpalus

GB

B

(SU,F)

M

Hesperia nevada

SN

■ -

M

Hesperia pahaska martini

MD

-

-

Hesperia juba

WNA

B

{SU,F)

M

Hylephila phyleus

WS

M

RA

Copaeodes aurantiaca

WNA

M

RA

Pholisora libya libya

MD

M

RA

Pholisora gracielae

MD

M

RA

Pholisora alpheus oricus

GB

U

(S)

ME

19(1): 1^63, 1980(81)

55

Taxon

Heliopetes domicella domicella

Heliopetes ericetoriim

Pyrgus scriptura

Pyrgus communis

Erynnis brizo burgessi

Erynnis funeralis

Erynnis meridianus meridianus

Erynnis telemachus

Chiomara asychis georgina

Systasea zampa

Thorybes pylades

Polygonus leo arizonensis

Epargyreus clarus huachuca

Battus philenor philenor

Papillio bairdii bairdii

Papilio rudkini

Papilio indra martini

Papilio rutulus rutulus

Papilio multicaudatus

Neophasia menapia

Pieris beckerii beckerii

Pieris sisymbrii elivata

Pieris protodice protodice

Pieris rapae

Colias eurytheme

Colias philodice philodice

Colias alexandra edwardsii

Colias cesonia

Phoebis sennae marcellina

Eurema mexicana

Eurema nicippe

Nathalis iole

Anthocaris pima

Anthocaris sara thoosa

Euchloe hyantis lotta

Apodemia mormo mormo

Apodemia palmeri marginalia

Calephelis nemesis californica

Calephelis wrightii

Satyrium behrii behrii

Ministrymon leda

Callophrys fotis fotis

Callophrys eryphon eryphon

Callophrys spinetorum

Callophrys siva siva

Callophrys sheridanii comstocki

Atlides halesus corcorani

Biogeographical j.,- Primary,

1 Voltinism „ ... 13
Affinity Habitat

T

-

-

WNA

M

WS

WNA

M

WS

WS

M

WS

RM

U

(S)

M

T

M

RA

SWD

M

M

RM

U

(S)

ME

T

-

-

SWD

M

RA

WS

-

-

SWD

-

-

SD

-

ME

WS

-

-

WNA

-

M

MD

B

{S,F)

DS

MD

B

(S,SU)

ME

WNA

U

(SU)

M

WNA

-

ME

WNA

u

(SU)

M

WNA

M

ME

RM

U

(S)

ME

WS

M

WS

WS

M

WS

WS

M

WS

WS

-

-

GB

U

(S)

DS

WS

M

WS

T

-

-

T

-

-

WS

M

WS

T

M

WS

SD

U

(S)

DS

MD

U

(S)

ME

MD

u

(S)

ME

GB

M

WS

MD

M

DS

SCM

B

(SU,F)

RA

SCM

M

RA

SN

U

(SU)

ME

SD

B

(SU,F)

DS

GB

U

(S)

ME

WNA

-

-

WNA

B

{S,SU)

ME

RM

B

(S,SU)

ME

MD

B

{S,F)

ME

WNA

M

DS

SWD

M

WS

WS

M

RA

RM

-

-

T

M

WS

Strymon melinus pudica
Lycaena helloides
Lycaena dorcas castro
Brephidium exilis

56

J. Res. Lepid.

Biogeographical Primary^

-j. 1 Voltinism „ ... '3

Affinity ^ Habitat

Leptotes marina

T

M

WS

Hemiarqus ceraunus gyas

SWD

M

WS

Hemiarqus isola alee

ws

M

WS

Plebejus melissa melissa

WNA

U (F)

RA

Plebejus icarioides evius

SCM

U (SU)

M

Plebejus shasta charlestonensis

E

U (SU)

M

Plebejus acmon acmon

SN

M

WS

Plebejus acmon texanus

SWD

M

WS

Everes amyntula

WNA

U (SU)

M

Euphilotes battoides nr. ellisii

MD

U (F)

ME

Euphilotes battoides martini

MD

U (S)

DS

Euphilotes battoides baueri

GB

-

Euphilotes enoptes ssp.

E

U (SU)

M

Euphilotes enoptes dammersi

MD

U (F)

DS

Euphilotes mojave ssp.

MD

U (S)

ME

Philotiella speciosa speciosa

MD

U (S)

DS

Glaucopsyche lyqdamus ssp.

MD

U (S)

DS

Glaucopsyche lyqdamus oro

RM

U (SU)

M

Celastrina arqiolus cinerea

RM

M

WS

Libytheana bachmanii larvata

SWD

-

WS

Asterocampa celtis montis

SD

-

ME

Limenitis archippus absoleta

SWD

B (SU,F)

RA

Limenitis weidemeverii anaustif ascia

SD

U (SU)

ME

Limenitis weidemeverii nevadae

E

U (SU)

M

Adelpha bredowii. eulalia

SWD

M

WS

Vanessa atalanta rubria

WS

M

WS

Vanessa virginiensis

WS

B (SU,F)

WS

Vanessa cardui

WS

M

WS

Vanessa annabella

WNA

M

WS

Precis coenia

WS

M

WS

Nymphalis californica

WNA

U (SU)

M

Nymphalis milberti furcillata

WNA

U (SU)

M

Nymphalis antiopa antiopa

WS

U (SU)

WS

Polygonia satyrus satyrus

WS

B (SU,F)

M

Polygonia zephyrus

WNA

B (SU,F)

M

Chlosyne palla vallismortis

MD

U (SU)

M

Chlosyne neumoegeni neumoegeni

MD

B (S,F)

DS

Chlosyne lacinia crocale

SWD

M

RA

Chlosyne californica

MD

B (S,F)

DS

Anthanassa texana texana

T

~

_

Phyciodes tharos distincta

SWD

-

-

Phyciodes phaon

WS

U (F)

RA

Phyciodes pallida barnesi

RM

_

_

Phyciodes mylitta mylitta

SN

~

M

Thessalia leanira alma

GB

u (S)

DS

Poladryas minuta arachne

RM

B (S,F)

ME

Euphydryas chalcedona kingstonensis

MD

B (S,F)

DS

Euphydryas anicia morandi

E

U (SU)

M

Euphydryas anicia ssp.

RM

U (S)

ME

Speyeria zerene carolae

E

U (SU)

M

19(1): 1^63, 1980(81)

57

Taxon

Biogeographical
Affinity ^

Voltinism^

Primary^

Habitat

Euptoieta claudia

WS

Agraulis vanillae incarnata

SWD

M

WS

Danaus plexippus plexippus

WS

M

WS

Danaus gilippus strigosus

SWD

M

WS

Cyllopsis pertepida dorothea

RM

B

(SU,F}

ME

Coenonympha California California

SCM

_

"

Coenonympha ochracea brenda

GB

U

(SU)

M

Cercyonis sthenele ssp.

E

D

(SO)

ME

= Endemic, MD = Mojave Desert,

GB = Great Basin,

SD =

Sonoran

Desert,

SWD = Southwest Desert, SK = Sierra Nevada, RM = Rocky Mountain, SCM =
Southern California Mediterranean, T = Tropical, WNA = Western North
America, WS = Widespread.

2

U = univoltine, B = bivoltine, M = multivoltine , S = spring, SU = summer,
F = fall, dash indicates insufficient data.

^DS = Desert Scrub, RA = Riparian/Agriculture, ME = Middle Elevations,

M = Montane, WS = Widespread, dash indicates insufficient data.

TABLE 3-. Biogeographic affinities of the Clark County, Nevada butterfly fauna.

Element

Primary Habitat

Entire Widespread Agriculture/ Desert Middle Montane

County Riparian Scrub Elevations

(%) (%) (%) (%) (%) (%)

Mojave Desert

18.3

O

o

17.6

66.7

40.0

5.0

Great Basin

6.3

3.7

0.0

13.3

10.0

10.0

Sonoran Desert

4.0

0.0

0.0

13.3

5.0

O

©

Southwest Desert

11.1

18.5

17.6

0.0

0.0

5.0

Sierra Nevada

3.2

3.7

0.0

0.0

5.0

o

©

Rocky Mountain

9.5

3.7

0.0

O

o

30.0

10.0

So. California Mediterranean

3.2

o

o

11.8

o

o

O

o

5.0

Tropical

7.1

11.1

5.9

0.0

o

o

©

O

Western North America

15.1

11.1

17.6

6.7

10.0

35,0

Widespread

18.3

48.1

29.4

o

O

©

o

5.0

Number of species

126

27

17

15

20

20

Taxon Month

19(1): 1-63, 1980(81)

59

Month

60

J. Res. Lepid.

19(1): 1-63, 1980(81)

61

Tajoon Month

62

J. Res. Lepid.

-K. ]

^ I

63

TABLE 5. Voltinism of the Clark Ctoimty, Nevada, butterfly fauna.

Primary

Habitat

No. of

Species

Univoltines

(%)

Bivoltines

(%)

Multivoltines

(%)

Widespread

27

3.7

3.7

92.6

Agriculture-Riparian

17

11.8

23.5

64.7

Desert Scrub

15

53.3

33.3

13.3

Middle Elevations

20

65.0

30.0

5.0

Montane

20

75.0

20.0

5.0

County

99

39.4

20.2

40.4

TABLE 6. Voltinism of the

to biogeographie

Clark County,

affinities .

, Nevada,

butterfly fauna

Biogeographical

Element

Percent

Univoltine

Bivoltine

Multi voltine

Endemic

100.0

0.0

0.0

Mojave Desert

54.5

31.8

13.6

Great Basin

71.4

14.3

14.3

Sonoran Desert

66.7

33.3

0,0

Southwest Desert

0.0

11.1

88.0

Sierra Nevada

50.0

0.0

50,0

Rocky Mountain

55.6

33.3

11.1

So. California Mediterranean

33.3

33.3

33.3

Tropical

0.0

0.0

100.0

Western North America

37.5

18.8

43.6

Widespread

10,5

15.8

73.8

i.

:^|S.7eo''

m

. \Jcy(i P7

TA-r

19(1): 1-63, 1980(81)

19(1): 1-63, 1980(81)

63

TABLE 5. Voltinism of the Clark County, Nevada, butterfly fauna.

Prirtary No. of Univoltines Bivoltines Multivoltines

Habitat Species (%) (%) (%)

Widespread
Agriculture-Riparian
Etesert Scrub
Middle Elevations
Jtontane

27

3.7

3.7

92.6

17

11.8

23.5

64.7

15

53.3

33.3

13.3

20-

65.0

30.0

5.0

20

75.0

20.0

5.0

99

39.4

20.2

40.4

Qsun-^

Journal of Research on the LepMoptera

19(1): 64, 1980(81)

Book Review

Herbivores: Their interaction with secondary plant metabolites.

Edited by Gerald A. Rosenthal and Daniel H. Janzen. Academic Press,
New York, 1979. 718 pp. $59.50.

The title of this book is misleading. It’s really an updating and attempted
res3mthesis of that part of chemical ecology dealing with chemical
mediation of herbivore-plant interaction. The emphasis is on insect
herbivores and secondary plant compounds. The 20 chapters are divided
into eco-evolutionary and primarily chemical. The latter are very compe-
tent reviews of what ecologists may derisively call “chicken-wire chemistry”
and their bibliographies are absolutely indispensible for anyone working in
this field. The former, where the fun is, display once again that chemical
ecology reduces intellectually to the ploy-counterploy model published by
Fraenkel in 1959, and that its practitioners must constantly do battle
against unjustified assumptions of adaptivity, “function,” and evolutionary
history. The most interesting chapter in the book is by Janzen, who
cautions his colleagues against all these deadly sins and in the process
spins a tall tale about phylogenetic inertia (which might even be true). His
caution that “herbivores do not eat Latin binomials” should apply
especially to Lepidopterists, long plagued by questionable host records
and the assumption that “a rose is a rose is a rose.” Gertrude Stein was not
a biologist. Hard-headed hypothesis testers may well feel that many
chemical ecologists aren’t either. Whether or not one is happy about
chemical ecology as an explanatory or a predictive science, this is an
important book.

Athur M. Shapiro, Department of Zoology, University of California, Davis, CA 95616.

INSTRUCTIONS TO AUTHORS

Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, on 8V2 x 11 inch paper with wide margins. Number all pages
consecutively and put author’s name at top right corner of each page. If your typewriter
does not have italic type, underline all words where italics are intended. Footnotes,
although discouraged, must be typed on a separate sheet. Do not hyphenate words at the
right margin. All measurements must be metric, with the exception of altitudes and
distances which should include metric equivalents in parenthesis. Time must be cited on
a 24-hour basis, standard time. Abbreviations must follow common usage. Dates should
be cited as example: 4. IV. 1979 (day-arabic numberal; month-Roman numeral; year-
arabic numeral). Numerals must be used before measurements (5mm) or otherwise up to
number ten e.g. (nine butterflies, 12 moths).

Title Page: All papers must have the title , author’s name, author’s address, and any
titular reference and institutional approval reference, all on a separate title page. A
family citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.

Abstracts and Short Papers: All papers exceeding two typed pages must be ac-
companied by an abstract of no more than 300 words. An additional summary is not
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Name Citations and Systematic Works: The first mention of any organism should
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References: All citations in the text must be alphabetically listed under Literature Cited
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Tables: Tables should be minimized. Where used, they should be formulated to a size
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Illustrations: Color must be submitted as a transparency (i.e., slide) ONLY, the quality
of which is critical. On request, the editor will supply separate detailed instructions for
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comments of reviewers.

Volume 19

Number 1

Spring, ly>S0(8

EV THIS ISSUE

Date of Publication: September 7, 1981

Butterflies of Clark County, Nevada

George T. & Anna T. Austin

Book Review

Cover Illustration: Black and white adaptation of a color illustration from A
Flight of Butterflies. Reprint permission given by The Metropolitan Museum of Art,
New York, NY.

Volume 19

Number 2

Summer, 1980(81)

Published By: The Lepidoptera Research Foundation, Inc.

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

Founder: William Hovanitz

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Jean Hovanitz, and R. H. T. Mattoni. There are no bond holders, mortgages, or other security holders.

Journal of Research on the Lepidoptera

19(2): 65-67, 1980(81)

Nantucket Pine Tip Moth, Rhyacionia frustrana, in Kem
County, California: Integrated Control and Biological
Notes (Lepidoptera: Tortiicidae, Olethreutinae)

Dennis M. Poore

Kem County Department of Agriculture, Bakersfield, California 93302

Abstract. Rhyacionia frustrana had four generations on Monterey
pine {Pinus radiata) in the San Joaquin Valley of California during
1974 with summer generations of approximately six weeks. In Cali-
fornia, the moth prefers P. radiata, but has been found in association
with eight other pine species. The use of chemical, biological and
cultural control practices has evidently lead to its eradication in a
Kem County pine plantation.

Tlie Nantucket pine tip moth, Rhyacionia frustrana (Comstock), is
known to have a wide geographical distribution. Powell and Miller (1978)
show the following range: Central America (Guatemala, Honduras and
Nicaragua), southern Mexico (Oaxaca State), West Indies (Cuba, Jamaica
and Dominican Republic), eastern United States (eastern Texas and
Florida, north to Missouri and Massachusetts). This pine tip moth was
probably introduced into California on infested nurseiy stock from the
eastern United States sometime in the late 1960’s, according to informa-
tion assimilated by a task force of various State and Federal agencies. In
California five counties (San Bernardino, San Diego, Riverside, Orange
and Kem) have been known to have infestations. The hosts^ recorded for
R. frustrana in California are Pinus attenuata Lemmon (Knobcone pine);P.
canariensis Sweet ex K. Spreng. (Canary Island pine); P. halepensis Mill.
(Aleppo pine); P. jeffreyi Grev. and Bah. (Jeffrey pine); P. nigra Arnold
(Austrian Black pine); P. radiata D. Don. (Monterey pine); P. roxburghii
Sarg. (Chir pine); P. sylvestris L. (Scotch pine); P. thunbergiana Franco
(Japanese Black pine) (personal communication - T. D. Eichhn, Insect
Taxonomy Laboratory, California Department of Food and Agriculture).
Nine species of pines (excluding exotics and ornamentals) in the eastern
United States have been recorded as hosts (Powell and Miller, 1978).

Rhyacionia was first detected in Kem County on November 10, 1971.
Tlie specimens were collected from Monterey pines (which is the preferred
host in California) at a Christmas tree farm, Kem County, Wasco,
California, Section 36, Township 26s. Range 24e. M.D. The damage by
burrowing larvae caused severe dieback to the terminals. The dieback

‘Not all hosts have been confirmed from rearing records

66

J. Res. Lepid.

caused secondary and lateral shoots to form, which resulted in reduced
growth, misshappen and compacted trees.

During 1974, the life history of R. frustrana on Monterey pines at the
infested Christmas tree farm was determined*. Peak flight was determined
by extrapolation from numerical counts of larvae, pupae, pupal cases and
adults. The major flight of adults from overwintering pupae occurred April
15; first generation peak flight was May 30 (T 3 days); second generation
peak flight was July 10 (T 4 days); third generation peak flight was August
28 (T 4 days). The summer generations took approximately six weeks to
complete development. A total of four generations occurred. Brown and
Eads (1975) postulated that in’ San Diego County R. frustrana had four
broods annually. Yates and Beal (1962) stated that R frustrana has two to
possibly five generations annually in the southeastern United States,
depending on latitude.

A chemical and a biological control agent were applied in 1976 in an
attempt at eradication (See Table 1). Surveys were conducted in late
summer through fall to determine efficacy of the materials. In the tree
farm, only three live pupae from a single Monterey pine were found on
October 7, 1976. No insecticides were applied during 1977 and 1978.
During 1977 surveys for the pest, only a single larva was found, which was
observed in September. No other specimens of the pine tip moth have
been collected subsequently.

Table 1

Insecticides Applied During 1976

APPL.

DATE

MATERIAL^

RATE/ACRE

APPL. METHOD

3/30

carbaryl - D

5 lbs. a.i.

Back pack duster

4/14

B. T.

50 lbs.

Fixed wing aircraft

4/27

carbaryl - D

6 lbs. a.i.

Fixed wing aircraft

5/11

B. T.

60 lbs.

Fixed wing aircraft

5/25

carbaryl - D

6 lbs. a.i.

Fixed wing aircraft

6/8

B. T.

37 lbs.**

Fixed wing aircraft

6/22

carbaryl - D

6 lbs. a.i.

Fixed wing aircraft

7/6

carbaryl - D

6 lbs. a.i.

Fixed wing aircraft

a.i. = active ingredient

B.T. = Bacillus thuringiensis (Berl.)

♦Used under S.L.N. registration CA-760011 and CA-760012, Dipel® 1 Dust and Sevin® 10
Dust, respectively.

♦♦Expended all available material for the project.

19(2): 65-67, 1980(81)

67

TYee shaping by terminal tipping done at three to four week intervals in
1977 and 1978 was, in fact, a cultural practice aiding in control of the pest.
At the same time, any chlorotic tips were also removed and crushed.

R. frustrana had four generations on Monterey pines in the San Joaquin
Valley during 1974 with summer generations of approximately six weeks
duration. The application of Bacillus thuringiensis (Berl.) and carbaryl at
bi-weekly intervals suppressed the tip moth’s population to a very low
level. Following continuous terminal tipping at three to four week intervals,
no specimens have been collected since September 1977.

Literature Cited

BROWN, L. R. & c. o. EADS, 1975. Nantucket pine tip moth in Southern California:

Identity and Insecticidal Control. Jour. Econ. Entom. 68(2): 380-382.
POWELL, J. A. & w. E. MILLER. 1978. Nearctic pine tip moths of the Genus Rhyacionia:
Biosystematic Review (Lepidoptera: Tortricidae, Olethreutinae). U.S.D.A.
For. Serv. Agric. Handb. 514, 51 pp.

YATES, H. O. & R. H. BEAL. 1962. Nantucket pine tip moth, U.S.D.A. For. Serv., For.
Pest Leafl. 70, 4 pp.

Journal of Research on the Lepidoptera

19(2): 68-71, 1980(81)

Moths of North America North of Mexico, Supplemental
literature: 1

J. C. E. Riotte

Department of Entomology, Bishop Museum, P. 0. Box 19000-A, Honolulu, HI 96819

The Journal of Research on the Lepidoptera herein initiates a project to
continually update the Moths of North America North of Mexico (MONA)
by citing all relevant papers published subsequent to each fascicle
produced. We feel the service will maintain the value of MONA and urge
all readers to submit citations to Riotte for incorporation into the periodic
bibliographies we will publish.

Fascicle 6.2, Gelechioidea - Oecophoridae

Hodges, R. W, Oecophoridae from West Texas. J. Lep. Soc. 29: 89-94 (1975).
Heppner, J. B. Mathildana newmanella (Oecophoridae) in Arkansas. J. Lep. Soc.
30: 18 (1976).

Fascicle 13, Pyraloidea

Munroe, E. Additional material of Scoparia huachucalis Munroe, with description
of male geniialia (Pyralidae, Scopariinae). J. Lep. Soc, 31: 63-66 (1977).
Fiance, S. B. & R. E. Moeller. Immature stages and ecological observations of
Eoparargyractis plevie (Pyralidae: Nimphulinae). J. Lep. Soc. 31: 81-88 (1977).
McFarland, N. Larval foodplant records for 106 species of North American
moths. J. Lep. Soc. 29: 112-125 (1975). Includes some Pyralidae.

Kendall, R. 0. Larval foodplants and life history notes for eight moths from Texas
and Mexico. J. Lep. Soc. 30: 264-271 (1976). Pyralidae and Sphingidae,
Munroe, E. A new genus and species of Odontiinae from Arkansas (Lepidoptera:

Pyralidae). Can. Ent. 105: 669-671 (1973).

Munroe, E. A new Nocteulopsis from Texas (Lepidoptera: Pyralidae: Odonti-
inae). Can. Ent. 106: 655-658 (1974).

Heppner, J. B. Synopsis of the genus Parargyractis (Lepidoptera: Pyralidae:

Nymphulinae) in Florida. The Florida Entomologist 59: 5-19 (1976).

Munroe, E . Additional Material of Scoparia huachucalis Munroe, with description
of male genitalia (Pyralidae, Scopariinae). J. Lep. Soc. 31: 63-66 (1977).
Munroe, E. A new species related to the garden webworm, with a key to American
members of the genus Achyra Guenee (Lepidoptera: Pyralidae: Pyraustinae).
Can. Ent. 110: 499-505 (1978).

McCafferty, W. P. & M. C. Minno. The aquatic and semiaquatic Lepidoptera of
Indiana and adjacent areas. The Great Lakes Ent. 12: 179-187 (1979).

19(2): 68-71, 1980(81)

69

Fascicle 20.1, Mimallonoidea and Bombycoidea (excl. Satumiidae)

McFarland, N. Larval foodplant records for 106 species of North American
moths. J. Lep. Soc. 29: 112-125 (1975). Includes some Lasiocampidae.

Lajonquiere, Yes de. Revision du Genre Phyllodesma Huebner - n(e) Partie -
Especes nearctiques (suite) - Correction et Adjunction. Bull, de la Soc. entomol.
de France, Tome 81, mars-avril, 108-112 (1976).

Fascicle 20.2A, Bombycoidea - Satumiidae (Part)

McFarland, N. Notes on three species of Hemileuca (Satumiidae) from eastern
Oregon and California. J. Lep. Soc. 28: 136-141 (1974).

Smith, Capt. M. J. Life history notes on some Hemileuca species (Satumiidae). J.
Lep. Soc. 28: 142-145 (1974).

Dominick, R. B. Amphion nessu^ (Sphingidae) attracted to female Anisota
virginiensis pellucida (Citheroniidae). J. Lep. Soc. 28: 176 (1974).

Leeuw, 1. A further note on the acceptability of an alternate foodplant for
Heijiileuca maia (Dmry) (Satumiidae). J. Lep. Soc. 28: 301 (1974).

Worth, C. B., T. F. Williams, A. P. Platt & B. P. Bradley. Differential growth
among larvae of Citheronia regalis (Satumiidae) on three genera of foodplants. J.
Lep. Soc. 33: 162-166 (1979).

Worth, C. B. Doubly overwintering Citheronia regalis Fabricius (Lepidoptera:
Satumiidae). J. Lep. Soc. 33: 166 (1979).

Tuskes, P. M. A new species of Hemileuca from the southwestern United States
(Satumiidae). J. Lep. Soc, 32: 97-102 (1978).

Fascicle 20.2B, Bombydoidea - Satumiidae (Part)

Collins, M. M. Notes on the taxonomic status of Hyalophora Columbia (Satumi-
idae). J. Lep. Soc. 27: 225-235 (1973).

Tuskes, P. M. The distribution and larval foodplant relationships of Saturnia
walterorum (Satumiidae). J. Lep. Soc. 28: 172-174 (1974).

Scarbrough, A. G., G. P. Waldbauer & J. G. Stemburg. Feeding and survival of
Cecropia (Satumiidae) larvae on various plant species. J. Lep. Soc. 28:212-219
(1974).

Peigler, R. The geographical distribution of Callosamia securifera (Satumiidae).
J. Lep. Soc. 29: 188-191 (1975).

Orsak, L. J. The type locality of Saturnia walterordm (Satumiidae). J. Lep. Soc.
29: 191 (1975).

Koehalmi, L. & P. Moens. Evidence for the existence of an intergrade population
between Hyalophora gloveri nokomis and H. Columbia in northwestern Ontario
(Lepidoptera: Satumiidae). Can. Ent. 107: 793-799’ (1975).

McFarland, N. Larval foodplant records for 106 species of North American
moths. J. Lep. Soc. 29: 112-125 (1975). Among other 4 Satumiidae.

Miller, T. A. & W. J. Cooper. Portable outdoor cages for mating female giant silk-
worm moths (Satumiidae). J. Lep. Soc. 30: 95-104 (1976).

Miller, T. A. Observations of Eupackardia calleta in Southern Texas (Satumiidae).
J. Lep. Soc. 30: 127-130 (1976).

Peigler, R. S. Observations on host plant relationships and larval nutrition in
Callosamia (Satumiidae). J. Lep. Soc, 30: 184-187 (1976).

70

J. Res. Lepid.

Prentiss, J. B. Time variations of pupal stage of Eupackardia calleta (Satumiidae).
J. Lep. Soc. 30: 187 (1976).

Tuskes, P. M. A key to the last instar larvae of west coast Satumiidae. J. Lep. Soc.

30: 272-276 (1976). Includes one species of Sphingicampa (fasc. 20. 2A).
Scarbrough, A. G., J. G. Sternburg & G. P. Waldbauer. Selection of the cocoon
spinning site by the larvae of //ya/ophoracecropia (Satumiidae). J. Lep. Soc. 31:
153-166 (1977).

Gillaspy, J. E. Antheraea polyphemus (Satumiidae) and Biblis hyperia (Nymphal-
idae) in Texas. J. Lep. Soc. 31: 166 (1977).

Preston, W. B. & W. B. McKillop. A melanistic specimen of Antheraea polyphemus
polyphemus (Satumiidae). J. Lep. Soc. 33: 147-148 (1979).

Toliver, M. E., J. G. Sternburg & G. P. Waldbauer. Daily flight periods of male
Callosamia promethea (Satumiidae). J. Lep. Soc. 33: 232-238 (1979).

Peigler, R. S. Collecting cocoons of Callosamia securifera (Satumiidae). J. Lep.
Soc. 30: 111-113 (1976).

Peigler, R, S. Wing color variation in Callosamia (Satumiidae). J. Lep. Soc. 30:
114-115 (1976).

Peigler, R. S. Hybridization of Callosamia (Satumiidae). J. Lep. Soc. 31: 23-34
(1977).

Peigler, R. S. Parasitism of Callosamia (Lepidoptera: Satumiidae). J. Georgia
Ent. Soc. 12: 111-114 (1977).

Peigler, R. S. Hybrids between Callosamia and Sarnia (Satumiidae). J. Lep. Soc.
32: 191-197 (1978).

Fascicle 21, Sphingoidea

Dominick, R. B. Some notes on the Sphingidae. J. Lep. Soc. 26: 234 (1972). This
note gave th^ idea for MONA follow-up.

Howe, W. H. The mature larva of Sphinx vashti (Sphingidae). J. Lep. Soc. 26:
247-248 (1972). First larval photo.

Franklin, C. M. New distribution records for Ceratomia hageni (Sphingidae). J.
Lep. Soc. 26: 198 (1972).

Dominick, R. B. Life history of Isoparce cupressi (Sphingidae). J. Lep. Soc. 27: 1-8
(1973).

Blanchard, A. Record and illustration of some interesting moths flying in Texas
(Sphingidae,...). J. Lep. Soc. 27: 103-109 (1973). Addition: Amplypterus donysa
(Dmce).

Beutelspacher, C. R. New Mexican Sphingidae records. J. Lep. Soc. 27: 303-304
(1973).

Dominick, R. B. A further field note on Isoparce cupressi (Sphingidae). J. Lep.
Soc. 28: 157 (1974).

Dominick, R. B. Amphion nessus (Sphingidae) attracted to female Anisota
virginiensis pellucida (Citheroniidae). J. Lep. Soc. 28: 176 (1974).

Lloyd, J. E. Genital Stridulation in Psilogramma menephron (Sphingidae). J. Lep.
Soc. 28: 349-351 (1974).

McFarland, N. Larval foodplant records for 106 species of North American
moths. J. Lep. Soc. 29: 112-125 (1975). Includes 1 sphingid.

Rickard, M. A. Life history notes on three Texas Sphingidae. J. Lep. Soc. 29:
167 (1975).

19(2); 68-71, 1980(81)

71

Brown, C. H. A survey of the Sphingidae of Sanibel Island, Florida. J. Lep. Soc.
30: 230-233 (1976).

Kendall, R. O. Larval foodplants and life history notes for eight moths from Texas
and Mexico. J. Lep. Soc. 30: 264-271 (1976). Pyralidae and Sphingidae.
Tuttle, J. P. Cocytius duponchel (Sphingidae): Second United States capture.
J. Lep. Soc. 31: 34 (1977).

Williams, B. D. Life history observations on Hemaris gracilis (Sphingidae). J. Lep.
Soc. 33: 254-257 (1979).

Riotte, J. C. E. A review of the North American hawk moth genus Lapara
(Lepidoptera: Sphingidae). Life Sci. Contr. 79, R. Ont. Mus., p. 40 (1972).
Riotte, J. C. E. Sphinx poecila - a valid North American hawkmoth species
(Lepidoptera: Sphingidae). The Great Lakes Ent. 13: 115-130 (1980).

Fascicle 22.2, Noctuoidea - Lymantriidae

NeU, K. A new subspecies of Orgyia leucostigma (Lymantriidae) from Sable
Island, Nova Scotia. J. Lep. Soc. 33: 245-247 (1979).

Acknowledgments

The implementation of the referee system, starting with Volume 17, has
been an outstanding success in assuring publication of the highest quality
papers. At this point all holdover papers from pre-referee time have been
used, including a few brief taxonomic papers by recognized authorities
which we felt did not require review.

Referees we wish to thank are:

Ray White
Frances Chew
Art Shapiro
Jerry Powell
John Emmel
Douglas Ferguson
Larry Gilbert
Jon Sheppard
L. P. Grey
F. M. Brown
R. S. Peigler
T. 0. Eichlin
Fred Stehr

Tom Emmel
Michael Collins
Ichiro Nakamura
Paul Ehrlich
Paul Tuskes
Richard Hoffman
Robert Lacy
Peter Brussard
Ashley Morton
Pritam Singh
John Huxley
Richard Vane- Wright

Journal of Research on the Lepidoptera

19(2): 72-81, 1980(81)

Demonstration of Reproductive Isolating Mechanisms
in Callosamia (Saturniidae) by Artificial Hybridization

Richard S. Peigler^

Department of Entomology, Texas A&M University, College Station, Texas 77843, USA

Abstract. The isolating mechanisms in Callosamia form an elaborate
array of complex and numerous ones including premating to postzygotic.
The cross C. securifera cf X C. promethea 9 is described for the first time, as
well as a complex cross involving C. angulifera and C. securifera, these in
addition to ten crosses reported earlier. Some of the results obtained are
comparable to those achieved by other workers hybridizing other genera of
Lepidoptera. Each isolating mechanism is discussed and interpreted. The
relationship between speciation and isolating mechanisms is discussed also,
with a review of some recent literature on this topic.

Introduction

The genus Callosamia Packard is a well-defined North American group
of satumiid moths comprised of only three species: C. angulifera
(Walker), C. promethea (Drury), and C. securifera (Maassen). Although
generally considered to be “well-known”, many questions remain un-
answered about the phylogeny and ecology of these insects. Moreover, the
barriers to hybridization in nature are much more complicated than simple
differences in circadian mating behavior. Obtaining and rearing hybrids in
the laboratory, and careful studies of natural populations can help
elucidate these problems. The purpose of this paper is to figure and
describe briefly artificial hybrids obtained since my report on ten earlier
crosses (Peigler, 1977), and to discuss some of the evolutionary, ecological,
and genetic aspects of the genus, with special reference to isolating
mechanisms.

Descriptions of New Crosses

C. securifera cf X C. promethea 9

Three broods of this cross were reared simultaneously. A brief account
of each brood is given followed by a composite description of the stages
based on all three broods. Table 1 contains numbers involved in the
broods.

Brood 1 was the result of a hand-pairing using a male and female (both
from wild-collected cocoons) from Berkeley County, South Carolina.

Museum Associate in Entomology, Los Angeles County Museum of Natural History

19(2): 72-81, 1980(81)

73

Some larvae were reared on tuliptree {Liriodendron tulipifera L.) and some
on sweetbay {Magnolia virginiana L.). Those fed on the former grew larger.
Many larvae died of disease in the penultimate stadium.

Brood 2 was the result of hand-pairing a male from a wild-collected
cocoon from Berkeley Co., S.C. and a reared female from Pine Grove,
Schuylkill Co., Pennsylvania. Larvae were fed tuliptree.

Brood 3 resulted from a natural pairing in a cage between a male from the
same source as the others and a female from Pine Grove, PA from a wild-
collected cocoon. Mortality in this brood occurred during the larval stadia.

Larva (Fig. 1): Color bluish as in pure C. promethea, but most with pale
lateral stripes. Red and yellow scoli large, either weakly clavate or tapered
toward apex, in a few individuals some of these scoli bifid. Two larvae in
Brood 1 were supertuberculate (see Peigler, 1977).

Cocoon: Most with well-developed attachments to stem. Color brownish-
gray to brilliant gold. Exhibiting considerable variation yet mostly
intermediate between parent species.

Male (Fig. 2): Characters of pure C. promethea very dominant, having
very dark ground color and only minimal gold suffusion beyond postmedian
line. Most with a trace of discal mark in forewing, not in hindwing.
Underside more intermediate but traits of C. promethea still predominating.
Wing apices generally more falcate (pointed) than males of either parent
species. Some specimens with sparse scaling on thorax and abdomen.

Female (Fig. 3): Antennae and spiracular pattern intermediate. Color
in most specimens reddish like C. promethea but lighter, some almost
orange. Very pronounced black along postmedian line and around discal
marks, as in C. promethea, but with even more contrast with adjacent
areas. Underside quite intermediate, not dissimilar to pure C. anguliferal
Some of these are larger than any females, hybrid or purebred, in the genus
which I have seen — the proverbial heterosis.

C. [angulifera cT X {angulifera cf X securifera 9)9]cr X C. securifera 9

All C. angulifera stock originated at Clemson, Pickens Co., S.C. and C.
securifera stock at Berkeley Co., S.C. and after hand-patring, the female
laid 110 ova, of which six hatched. One larva was reared on tuliptree to the
pupal stage and a female emerged in July. She was mated to a male C.
angulifera but the ova did not hatch.

Larva: The yellowish lateral stripe prominent. Colored scoli (red
anterior ones, yellow caudal one) very small. Black scoli so minute as to be
almost invisible.

Cocoon: Large and puffy as in pure C. securifera, having a long, thin
peduncle. Color of silk very golden.

Female: Color light orange. Very close to pure C. securifera on upper-
and undersides. Discal marks in all four wings large, more as in C.
angulifera.

74

J. Res. Lepid.

Fig. 1. Mature larva of Callosamia securifera cf X C. promethea 9 from Brood 2. (Coloration in
figure is too blue).

Fig. 2. Hybrid male of C. securifera cf X C. promethea 9 from Brood 2.

Fig. 3. Hybrid female of C. securifera cf X C. promethea 9 from Brood 1. (Coloration of actual
specimen is more orange).

19(2): 72-81, 1980(81)

75

Conclusions from Above Crosses

The larvae of the three broods of C. securifera cf X C. promethea 9 were
all intermediate between parent species but exhibited pronounced
differences between each brood (larvae of the pure species are rather
constant). Larvae within a single brood showed minimal variation among
themselves. Thus, each time a particular cross is repeated, one can expect
similar but not identical results.

One of the conclusions from my earlier experiments (Peigler, 1977) was
that hybrids involving C. promethea are sterile, i.e., cannot be backcrossed
or make F2 crosses. Due to different emergence dates for males and
females and my being away when they emerged, I was unable to test the
fertility of these moths by additional matings. Earlier crosses of C.
promethea with the two other species failed to show two larval characters,
which appeared in the new cross. One of these traits is the presence of a
light lateral stripe, and the other is the whitish ring on the four red thoracic
scoli just above the black bases. These two features can be seen clearly in
the color photograph (Fig. 1) of one of the larvae, which could otherwise
pass for pure C. promethea.

The three broods of the cross C. securifera cf X C. promethea 9 gave
similar results despite the fact that in one the female was from the South
(from a population sympatric with C. securifera) and in the other two the
females were from Pennsylvania (allopatric to C. securifera). This suggests
that geographical genetic variation may be overshadowed by interspecific
genetic differences. Many additional crosses are needed to develop such
conclusions. Source localities of parental stock in hybridization studies
should always be recorded.

The complex cross described above involving C. angulifera and C.
securifera gave evidence that a small amount of fertility can be retained in
hybrids for at least three generations, notwithstanding the fact that
eclosion becomes very low already in the first backcross or F2 cross (this is
item 7 on Table 2), as also shown in some of my earlier crosses. This
sterility occurs even when the ovipositing female is purebred.

As I pointed out (Peigler, 1977), most mortality in hybrid broods occurs
in very young larvae, suggesting a genetically based physiological dis-
harmony. Because of the ever-present problem of disease, and the fact
that broods of the pure species also suffer from the same, such “negative
data” cannot be very useful. I have no quantitative data available for
percent success in rearing the three pure species from eggs to adults, but
the usual results are normally no better than in the Fi hybrid broods.
Workers using Drosophila, certain plants, etc. enjoy higher numbers of
experimental subjects on which to make statistical analyses. My methods
of recording and tabulating data from the hybridization experiments
follow those prescribed by Robinson (197 1) in the introductory chapter of
his book. A summary of results of all Fi crosses I have made thus far is
given in Table 1.

76

J. Res. Lepid.

Isolating Mechanisms

Collins and Tuskes (1979) provided a definitive study of isolating
mechanisms in another genus of satumiid moths, but these were pre-
dominantly prezygotic, and postzygotic ones such as reported here are
desirable to complete their study. The present study of Callosamia would
be enhanced by prezygotic observations such as theirs. I have attracted
males of C. angulifera and C. securifera to captive females of C. promethea
which were emitting pheromone during the flight times of the other two
species. To determine whether pheromone differences exist between the
three species will require observations such as those of Collins and Tuskes
(1979).

Priesner (1968) gave data to suggest that the pheromone of all three
species of Callosamia may be the same, and demonstrated that Hyalo-
phora cecropia (L.) and C. promethea have different pheromones, but there
is a partial degree of interattractivity . Rau and Rau (1929) reported a male
of C. promethea attracted to a female of H. cecropia which emitted
pheromone at the normal time before dawn.

Harbich (1976) presented an array of postzygotic isolating mechanisms
in sphingid moths remarkably parallel to those of Callosamia. General
classifications of reproductive isolating mechanisms were tabulated by
Dobzhansky (1970) and Littlejohn (1969). My Table 2 roughly follows the
format of the latter, but is modified to fit Callosamia in particular.

In nature each of the isolating mechanisms enumerated in Table 2 are
tested if the previous one fails. In captivity only items la and 2 can be
circumvented by artificial methods, and we may assume that temporal
isolation is probably the most important one operating in nature. Selection
should be expected toward those highest on the list (Littlejohn, 1969). In
Callosamia crosses it is possible to see examples of all isolating mechanisms
in Table 2 excepting item 2. Even this one could be tested by placing
captive females in large cages with a choice of plants for oviposition.

I am not aware of any valid reports of wild hybrids ever being found, but
private and museum collections should be checked nonetheless. The
erroneous report of a C. promethea-C. angulifera wild hybrid in the News of
the Lepidopterists’ Society (June 1975, p. 10) was based on the erroneous
statement that these hybrids occur by Collins and Weast (1961). Although
the artificial hybrids of known parentage are recognizable as having
intermediate or combined traits, wild hybrids could be easily overlooked
unless a search with the explicit intention of finding them was undertaken.
The searcher must be familiar with the normal variation which occurs
within each species. If the three species of Callosamia can be crossed so
easily in captivity, surely the primary isolating mechanisms must oc-
casionally fail in nature. Examples of how this could occur include the
possibility of artificial lights (such as streetlights) causing a female of C.
promethea to continue emitting pheromone after nightfall, thus attracting

19(2): 72-81, 1980(81)

77

Table 1: RESULTS OF BASIC Fi CROSSES IN CALLOSAMIA*

CROSS

No. eggs eclosed/
no. eggs deposited

No.

pupating

No. d

No. 9

fast- slow 99
development

angulifera d X promethea 9

33

19

14

fast 99

angulifera d X promethea 9

6/115

2

1

1

angulifera d X securifera 9

ca. 125/150

56

- 25

28

fast 99

angulifera d X securifera 9

14/17

12

7

4

promethea d X securifera 9

155/175

ca. 20^*

4

0

promethea d X securifera 9

109+/235

31

23

8

slow 99

securifera d X promethea 9

125/134

10

6

4

slow 99

securifera d X promethea 9

almost 100% eclosion

32

19

13

slow 99

securifera d X promethea 9

almost 100% eclosion

15

9

6

slow 99

securifera d X angulifera 9

21/128

12

6

5

slow 99

angulifera d X Sarnia cynthia 9

over 90% eclosion

10

4

0

♦Including results from Peigler (1977 and 1978).

♦♦Less than half pupated within the 55 cocoons which were spun.

Note: The only basic F i cross combination which I have not had the opportunity to make is promethea d'
X angulifera 9.

Table 2: ISOLATING MECHANISMS IN CALLOSAMIA

1. Reduction or elimination of cross -mating

a. Temporal

b. Mechanical

2. Incorrect or poor choice of foodplant for oviposition

3. Zygotic mortality — eggs fail to hatch

4. Weak Fi progeny

a. Larval inviability

b. Disruptive diapause of pupa (see Peigler, 1978)

c. Adults not vigorous or well -formed (see Peigler, 1977)

5. Differing developmental times for female hybrids

6. Incorrect temporal activity of hybrid adults

7. Fi hybrid or backcross sterility — partial or complete

Note: Mechanism la is premating, lb-7 are postmating; 1-2 are
prezygotic, 3-7 are postzygotic

78

J. Res. Lepid.

males of C, angulifera. Even if the situation just proposed were to occur, it
would be additionally necessary for the female to have failed to mate with a
male of C. promethea earlier, which is also unlikely. Another possibility for
cross matings is the emergence of females of two species in close proximity
(within several cm), and males mating with both when attracted to the first
“calling” female.

Circadian temporal isolation (allochronic mating behavior) in Callosamia
was finally clarified by Ferguson (1972). Stated simply, C. securifera flies
(i.e., females emit pheromone and males respond) during midday hours, C.
promethea flies in late afternoon, and C. angulifera flies after sundown but
before midnight. Supposedly a margin of one hour or more falls between
these flight times when no species is flying. Male and female Fi hybrids of
Callosamia are possibly at a disadvantage for timing of correct mating
behavior (item 5 on Table 2). Some data for this are in Peigler (1977).

One area of worthwhile investigation would be comparison of exact flight
times of all three species of Callosamia between sympatric and allopatric
populations. Specifically, would C. promethea fly earlier and/or later in
Cedar Rapids, Iowa where it is common but the other two species do not
occur, than populations of C. promethea living among its congeners?
Indeed Ferguson (1972) stated that C. securifera flies earlier in coastal
South Carolina (sympatric with C. promethea) than in south-central
Florida (allopatric to C. promethea). The flight times of these two diurnal
species probably vary with atmospheric conditions (Collins and Weast,
1961), with population density (low population levels perhaps having a
broader flight time to ensure all females are mated), and with latitude,
since photoperiod varies with the latter. Lepidopterists in several states
could make useful contributions by keeping careful and persistent records
on the circadian behavior of these moths, and noting the aforementioned
parameters. Some data are already in place (RauandRau, 1929; Toliver et
al., 1979). There appears to be no seasonal temporal isolation in
Callosamia, as was given by Ferguson (1972) for two species of Hyalophora
in southeastern Canada.

Mechanical isolation (incompatibility of genitalia) is virtually negligible
between C. securifera and C. angulifera, but C. promethea has considerably
larger genitalia in both sexes than its congeners. Should we then be
surprised that the flight time of C. promethea falls between the other two?
Shapiro (1978) and Dobzhansky (1970) gave some convincing arguments
against the “lock and key” significance, but I believe that mechanical
differences in Callosamia do play a role in reducing natural hybridization. I
have discussed (Peigler, 1977 and 1978) how this difference also hinders
artificial hybridization experiments. Male genitalia of the hybrids are
intermediate in size and shape.

Information on foodplant specificity in this genus can be found
elsewhere (Ferguson, 1972; Peigler, 1976; Feeny and Scriber, 1979).

19(2): 72-81, 1980(81)

79

Since tulip tree is known to be the best foodplant for hybrids, ovipositing
females of C. promethea which had mated to one of the other species would
likely select plants unsuitable for the hybrid larvae, since C. promethea is
polyphagous. A female of C. securifera would oviposit on sweetbay,
although tuliptree would be a better host for her offspring if she had mated
to a nonconspecific male. Brood 1 of the cross C. securifera cf X C.
promethea 9 included a striking difference in size of adults between those
reared on tuliptree and those on sweetbay. Slower growth rates were also
seen when rearing on sweetbay the larvae of C. angulifera cf X C. securifera
9 (Peigler, 1976). It is possible that tuliptree is the best foodplant for pure
C. promethea, as shown by certain growth data given by Feeny and Scriber
(1979). Since C. angulifera is monophagous on tuliptree, the foodplant
difficulty would not occur in wild hybrid broods in which the mother was C.
angulifera. An Fi hybrid female of any given parentage in the genus may be
no more likely to select the best foodplant (tuliptree) than a less optimal
one. Such foodplant differences in closely related species of Lepidoptera
can certainly be interpreted as an isolating mechanism.

Female hybrids showing both heterosis (as I mentioned under the cross
C. securifera cf X C. promethea 9) and reduced viability (as I mentioned
under C. promethea cT X C. securifera 9 in Peigler (1977)) were obtained in
crosses of the genus Colias (Pieridae) by Grula and Taylor (1980). The
latter authors attributed these differences between sexes in hybrid broods
and between reciprocal crosses to the X-chromosome, which apparently
contains most or all genes governing size, developmental rate, wing
pigmentation, and wing color pattern. Callosamia has not been investi-
gated cytogenetically, except for spermatogenesis in C. promethea (Robin-
son, 1971). Moths in the same subfamily (Satumiinae) as Callosamia may
have the XXcf :XY 9 or XXcf :X09 type of sex determination, the presence
of sex chromatin seemingly correlated with the latter (Gupta and Narang,
1980). It would be best to reserve speculation about the genetic
implications of results from crosses within this genus until such basic
cytogenetic information is available. Haldane’s Rule (Robinson, 1971: 24;
Dobzhansky, 1970: 333) that the heterogametic sex is rarer or has reduced
viability in hybrid broods certainly appears to hold for Callosamia. The
pronounced sexual dimorphism of these moths will make such studies all
the more enticing, since the findings of Grula and Taylor (1980) suggest
increased sexual dimorphism connotes increased genetic incompatibility
in hybridization of Lepidoptera.

A conspicuous phenomenon which is well-illustrated by Callosamia
hybrids is that of males emerging before or after females of a hybrid brood,
frequently the females diapause and the males do not (see Table 1). This
was also shown by Cocault et al. (1980) in a hybrid brood of the saturniids
Graellsia isahellae (Graells) cT XActias luna (L.) 9. Oliver (1978 and 1979)
found the phenomenon in his nymphalid hybrids as did Grula and Taylor

80

J. Res. Lepid.

(1980) in their hybrids of Colias. The differing developmental rates of the
sexes in hybrid Lepidoptera are here proposed as an isolating mechanism
(item 5 in Table 2) because such would reduce frequency of F2 and
backcross matings when hybrid broods are produced in nature. Based on
the few F 1 crosses made thus far in Callosamia, it appears that fast female
development occurs in hybrid broods in which the father is C. angulifem,
and slow female development is to be expected in broods fathered by C.
securifera or C. promethea.

Discussion

Some workers have advocated that isolating mechanisms are the result
of speciation, not the cause (Ehrlich and Raven, 1969) while others appear
to take the opposing viewpoint (e.g., Bush, 1969). Lewontin (1974: 161)
felt that isolating mechanisms arise as a result of genetic divergence due to
allopatry, and then become selectively reinforced upon secondary contact.
These matters were also ably reviewed by Futuyma and Mayer (1980) who
concluded that the genetic and selective causes of reproductive isolation
are still largely unknown. In Callosamia I would propose that the large
amount of genetic incompatibility inferred from the hybrid crosses is
strong evidence that the genomes of the three species differ considerably,
but allozyme analysis would be desirable as proof. These and other
isolating mechanisms were probably increased in number and intensity
after speciation was complete. Once allochronic mating behavior became
perfected (whatever the cause) to the point that cross matings were no
longer significant in number, the other mechanisms would increase in
number and intensity due to genetic divergence, the three species then
being on separate paths of evolution. The postzygotic ones cannot be
assumed, in my opinion, to have evolved to be relied upon when the
prezygotic ones failed. They are merely coincidental, but are to be
considered valid isolating mechanisms in that they reduce chances of
hybrids being produced, or when produced will fail to pollute the genomes
of the pure species. Although the evolutionary processes which result in
speciation and in reproductive isolation are still poorly understood, the
two processes must evidently go hand-in-hand to the extent that one is not
necessarily the result of the other.

Acknowledgments: I thank Dr. Joseph E. Eger and two anonymous referees for
the Journal for reviewing the original manuscript and offering helpful suggestions
for its improvement. Dr. G. R. Gamer of Clemson University photographed the
larva (Fig. 1). Special thanks are due to my mother, Paralie Williams Peigler, who
cared for growing larvae and emerging adults of the hybrids described in this paper
during the summer when I was away on extended trips.

Literature Cited

BUSH, G. L. 1969. Sympatric host race formation and speciation in frugivcrous flies
of the genus Rhagoletis (Diptera, Tephritidae). Evolution 23: 237-251.
COCAULT, R., G. LECOCQ & R. VUATTOUX. 1980. Erforg einer Hybridation zwischen

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Graellsia isabellae und Actias luna (Lep.: Attacidae). EntomoL Z. (Stuttgart)
90: 14-16.

COLLINS, M. M. &P. M. TUSKES. 1979. Reproductive isolation in S3unpatric species of
dayflying moths {Hemileuca: Satumiidae). Evolution 33: 728-733.

COLLINS, M. M. & R. D. WEAST. 1961. Wild silk moths of the United States. Collins
Radio Co., Cedar Rapids, Iowa, 138 pp.

DOBZHANSKY, T. 1970. Genetics of the evolutionary process. Columbia Univ. Press,
New York, xiii -f 505 pp.

EHRLICH, P. R. & P. H. RAVEN. 1969. Differentiation of populations. Science 165:
1228-1232.

FEENY, P. & J. M. SCRIBER. 1979. Growth of herbivorous caterpillars in relation to
feeding specialization and to the growth form of their food plants. Ecology 60:
829-850.

FERGUSON, D. c. 1972, Satumiidae (in part), Bombycoidea, in R. B. Dominick et al.,
The moths of America north of Mexico, fasc. 20, 2B: 155-275, col. pis. 12-22.

FUTUYMA, D. J. & G. C. MAYER. 1980. Non-allopatric speciation in animals. Syst. Zool,
29: 254-271.

GRULA, J. w. & 0. R. TAYLOR, Jr. 1980. Some characteristics of hybrids derived from the
sulfur butterflies, Colias eurytheme and C. philodice: phenotypic effects of the
X-chromosome. Evolution 34: 673-687.

GUPTA, M. L. & R. c. NARANG. 1980. Chromosome numbers, sex chromatin and sex
chromosome mechanism in some satumiid moths of India. Entomon. 5: 13-18.

HARBICH,H. 1976. Isolationsmechanismenund Arterhaltung im Genus Celerio (Lep.,
Sphingidae). EntomoL Z. (Stuttgart) 86: 33-42.

LEWONTIN, R. c. 1974, The genetic basis of evolutionary change. Columbia Univ.
Press, New York, xiii + 346 pp.

LITTLEJOHN, M. J. 1969. The systematic significance of isolating mechanisms, pp.
459-482 in International Conference on Systematic Biology, Univ. of Michigan,
1967. Natl. Acad. Sci. Publ. 1692, 632 pp.

OLIVER, c. G. 1978. Experimental hybridization between the nymphalid butterflies
Phyciodes thaws and P. campestris montana. Evolution 32: 594-601.

1979. Experimental hybridization between Phyciodes thaws and P.

batesi (Nymphalidae). J. Lepid. Soc. 33: 6-20.

PEIGLER, R. s. 1976. Observations on host plant relationships and larval nutrition
in Callosamia (Satumiidae). J, Lepid. Soc. 30: 184-187.

1977. Hybridization of Callosamia (Satumiidae). J. Lepid. Soc. 31: 23-

34.

. 1978. Hybrids between Callosamia and Sarnia (Satumiidae). J. Lepid.

Soc. 32: 191-197.

PRIESNER, E. 1968. Die interspezifischen Wirkungen der Sexuallockstoffe der
Satumiidae (Lepidoptera). Z. vergl. Physiol. 61: 263-297, 1 tabl.

RAU, P. & N. L. RAU. 1929. The sex attraction and rhythmic periodicity in giant
satumiid moths. Trans. Acad. Sci. St. Louis 26: 81-221, figs. 1-2.

ROBINSON, R. 1971. Lepidoptera genetics. Pergamon Press, New York, 687 pp,

SHAPIRO, A. M. 1978(1979). The assumption of adaptivity in genital morphology.
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Journal of Research on the Lepidoptera

19(2): 82-87, 1980(81)

Canalization of the Phenotype of Nymphalis antiopa
(Lepidoptera: Nymphalidae) from Subarctic and
Montane Climates

Arthur M. Shapiro

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

Abstract. Nymphalis antiopa from the montane Sierra Nevada of California
respond phenotypically to pupal cold shock in a manner similar to lowland
California antiopa and unlike subspecies hyperborea from Alaska. This could
result from historical factors, gene flow, or canalization of the phenotype in
the Sierran climate, which shows very wide diel temperature fluctuations but
not the prolonged cool spells common in the Alaskan summer.

The concept of canalization of development (Waddington, 1957) has
been thoroughly assimilated into the evolutionary framework of modern
biology. Like many other biological concepts deriving from seemingly
impregnable Darwinian deduction, it has rarely been tested experimentally;
although physiological adjustment in geographic populations is routinely
demonstrated, developmental buffering is not. Shapiro (1981) attempted
to test the hypothesis that canalization is adaptive to the physical
environment, using the phenotypic response of the Mourning Cloak
butterfly, Nymphalis antiopa (Linnaeus) to pupal cold shock. It has been
known for over a century that various species of the tribe Nymphalini
respond to temperature shock by producing predictable, often very
radical, pattern aberrations. In the most dramatic aberration of A^. antiopa,
“hygiaea,” the yellow border is doubled in width, and the blue spots
normally arranged in a row basad of it are entirely absent (Fig. 1).
“Hygiaea” can be induced by a variety of treatments (Fischer, 1907;
Standfuss, 1896). One treatment which is very efficacious at inducing it in
near-sea-level Californian stock is exposure of 8-hr-old pupae to 2°C for 2
weeks. Shapiro (1981) reasoned that the subarctic subspecies N. a.
hyperborea (Seitz), which has a relatively high probability of being chilled
in the pupal stage, should be better buffered developmentally against cold
than lowland Californian N. a. antiopa, which are assured of benign
conditions in a Mediterranean climate. Treatment failed to produce any
“hygiaea” among 53 Alaskan animals, vs. 31 among 157 lowland
Californian ones. This is a highly significant difference (X^ = 12.28, p
<0.005). Limitations on the interpretation of these results are spelled out
by Shapiro (1981).

19(2): 82-87, 1980(81)

83

Fig. 1. Bred, cold-shocked Nymphalisantiopa. Left: Donner Pass, Sierra Nevada,
normal (top) and “hygiaea” (center, brood 3; bottom, brood 4). Right:
Vacaville, Solano County, lowland California, normal (top) and “hygiaea”
(center); ssp. hyperborea, Fairbanks, Alaska, 1979 (bottom).

N. antiopa occurs in California from sea-level to tree-line. Populations at
high altitudes, like Alaskan ones, are likely to be chilled during develop-
ment. The pattern of probable exposure differs radically between the two
areas, however. Alaskan hyperborea are larvae in June-early July and
pupae in mid- July (Fairbanks). They are likely to experience periods of
cool, showery weather with temperatures between 40-50°F), but no frost.
Such weather is almost impossible in the Sierra Nevada of California. At
7000 ft. (2100''’ m), antiopa are larvae in late July-early August, and pupae
in mid-August. Afternoons are clear to partly cloudy with frequent highs of

84

J. Res. Lepid.

80°F (27°C), but nocturnal freezes are very common. Thus, Alaskan
antiopa may develop in prolonged cool but not cold conditions, while
Sierran ones are cold almost every night but warm almost every day.
Summer monthly mean temperatures for Fairbanks, Alaska and Truckee,
California are similar, but the diel ranges are very different (Tables 1,2).
Since short exposures (less than 1 wk) to 2°C have little or no phenotypic
effect on N. antiopa, and the full 2-wk exposure is required to obtain the
full expression of “hygiaea,” it would not be surprising if Sierran antiopa
were less well buffered against sustained low temperatures than Fairbanks
hyperborea. This was tested experimentally in 1980.

Five colonies of larvae were collected in the 3rd instar on willows {Salix
spp.) at two Sierran localities: one from Martis Creek, near Truckee,
Placer Co. (elevation about5600ft. = 1707 m), JulyS, 1980, and four from
the east end of Donner Pass along U. S . Highway 40 east of Norden, Placer
Co. (7000 ft. = 2134 m), August 7, 1980. Martis Creek is a sedgy swale

Table 1. Comparisons of summer temperatures for Fairbanks, Alaska and
Truckee, California for a sample year (1977). Data from National Oceanic and
Atmospheric Administration, Asheville, N. C.

VALUES IN °F AND (IN PARENTHESES) °C

Fairbanks* Truckee^

Variable

June 77

July 77

Aug. 77

June 77 July 77 Aug. 77

68.8

73.7

73.2

77.4

80.7

81.6

mean daily maximum

(20.4)

(23.1)

(22.9)

(25.2)

(27.0)

(27.5)

50.4

51.9

52.0

42.1

41.8

44.5

mean daily minimum

(10.2)

(11.0)

(11.1)

( 5.6)

( 5.4)

( 6.9)

59.6

62.8

62.6

59.8

61.3

63.1

mean daily temperature

(15.3)

(17.1)

(17.0)

(15.4)

(16.3)

(17.3)

departure of mean daily

T from long-range mean

+0.6

+2.1

+7.2

+6.3

0

+3.4

daily T

(+0.3)

(+1.2)

(+4.0)

(+3.5)

0

(+1.9)

number of nights with
temperature below 40°F

(4.4°C)

0

0

2

10

12

6

‘Fairbanks Airport, 64°49' N, 147°52' W, elevation 436 ft. = 182.9 m.
^Truckee Ranger Station, 39°20' N, 120°11' W, elevation 5995 ft. = 1827.3 m.

19(2): 82^7, 1980(81)

85

Table 2. Freeze data for Fairbanks, Alaska and Tmckee, California. Station data
as in Table 1. Source: NOAA.

DATE OF LAST SPRING OCCURRENCE OF:

16°F (-8.9°C) 20° (-6.7) 24° (-4,4) 28° (-2.2) 32° (0)

Fairbanks 1977 16.IV 29.rV 29.IV 29.IV lO.V

Tmckee 1977 19.V 19.V 28.V 29.V ll.VI

DATE OF FIRST FALL OCCURRENCE OF:

32°F (Q°C) 28° (-2.2) 24° (-4.4) 20° (-6.7) 16° (-8.9)

Fairbanks 1977 4.IX LX l.X 13.X 16.X

Tmckee 1977 5.Vn 21.IX 22.IX 7.XI 9.XI

NUMBER OF DAYS BETWEEN LAST SPRING AND FIRST
FALL OCCURRENCE OF:

16° (-8.9) 20° (-6.7) 24° (-4.4) 28° (-2.2) 32° (0)

Fairbanks 1977 183 167 155 155 117

Tmckee 1977 174 172 117 115 24

running through Great Basin shrub desert {Artemisia, Purshia). The
Donner site is mesic-montane and receives a very heavy winter snowpack.
Both populations are univoltine. The two sites are about 12 mi (19.3 km)
apart. Martis Creek receives cold air drainage at night and is both hotter by
day and colder by night than Donner Pass. Tmckee Ranger Station (5995
ft. = 1827 m) provides a weather station intermediate in climate and
elevation between them.

The Martis Creek colony was reared to maturity on English Elm, Ulmus
procera (Ulmaceae). This tree became unavailable by August due to
defoliation by beetle larvae, and the four Donner broods were reared
instead on Hackberry, Celtis sinensis (Ulmaceae). All rearing was under
continuous light at 25°C. Eight-hr pupae were refrigerated in the dark at
2°C, held for 14 days, and returned to 25°C. Larval mortality was
negligible m all broods.

Pupal mortality was very heavy in refrigerated groups from all broods,
and exceeded 75% overall. Such heavy mortality is unusual. Moreover,
chilled animals usually die as pharkte adults, which can be scored for
phenotype; in these broods most mortality occurred before wing pigmen-
tation was laid down. Celtis is an unusual host, but the very poor
survivorship of the Martis Creek brood, reared on Ulmus, argues against it
as a major cause of pupal mortality; it seems that Sierran antiopa are
poorly protected physiologically against long chilling in the pupa, even
compared to lowland Californian ones.

86

J. Res. Lepid.

Results and Discussion

Phenotypic sensitivity paralleled pupal mortality: seven “hygiaea” were
produced among 61 scorable Donner animnals (Table 3). This is not
significantly different from 31/157 from lowland California {Z^ = 1.72,
0.2 50 >p >0.1 00) but is more unlike 0/53 from Alaska = 6.23,
0.025>p>0.010). Most suggestively, all four Donner broods gave at least
one “hygiaea” each, just as 12/13 lowland Californian broods reared in
past years have done. Neither of the 1979 Alaskan broods produced even
one individual transitional to “hygiaea.” The very small Martis Creek
emergence does not significantly affect the statistical comparisons.

Alaskan hyperhorea exaggerate their subspecific characters when chilled.
The same modifications were seen in both Martis and Donner antiopa.
They were often quite marked in animals scored “normal” for purposes of
this analysis. Sierran animalss also resembled Alaskan ones in size (mean
LFW: lowland CA, 37 mm; Sierra Nevada, 30.3 mm; Fairbanks, 29.0 mm).
Unlike hyperhorea, Sierran experimentals often showed slight modifica-
tions in the direction of “hygiaea”; almost all of Donner brood 3 showed
them. Several other odd phenotypes were obtained in the chilled Sierran
antiopa: one animal each in Donner 1 and Martis had the wings thinly
scaled, and the Martis individual also had the blue spots oddly shaped and
arranged, and of a violet color. One individual in Donner 2 had the
hindwings symmetrically falcate.

This variability in chilled Sierran antiopa contrasts with both the
unchilled controJs and with wild-collected material. In a series of about 3
dozen from Donner Pass, Truckee, and nearby Sagehen Creek the
phenotype is quite uniform: the size is larger (mean LFW 34.5 mm), there
is a slight increase in black in the borders relative to lowland specimens,
and the blue spots may be slightly enlarged, especially on the hindwing.
The increased black scaling in the borders is duplicated in both control
and experimental Sierran broods, increased in the latter. Certain other
temperature treatments favor increased blue spotting in both European
(Standfuss, 1896) and lowland Californian (Shapiro, unpublished) antiopa.

Sierran antiopa thus behave like lowland Californian ones with respect to
the major aberration “hygiaea,” but in other responses to chilling they are
intermediate in response between lowland Californian and Alaskan
broods, and more variable than either. The “hygiaea” response can be
accounted for in at least three ways. Two of these imply lack of adaptation
to local climate: phylogenetic inertia (recent common ancestry of lowland
and Sierran antiopa) and gene flow (constant or intermittent). There are no
data bearing directly on either. The species is highly dispersive, and
occurs locally from the coast across the Sierra in riparian habitats.
Differences in the phenotypes of chilled lowland and Sierran animals
argue for some degree of differentiation among the populations. The third
hypothesis specifically invokes local climatic adaptation. The dissimilarity

19(2): 82-87, 1980(81)

87

Table 3. Results of temperature-shock experiments with Nymphalis antiopa from
the Sierra Nevada of California.

Normal

Normal

Hygiaea Hygiaea Unscorable

Brood, source

live

dead

live

dead

dead

Total

Donner Pass #1

experimental

18

0

1

0

53

72

control

10

0

0

0

0

10

Donner Pass #2

experimental

18

1

2

1

45

67

control

10

0

0

0

0

10

Donner Pass #3

experimental

14'

1

2

0

57

74

control

10

0

0

0

0

10

Donner Pass #4

experimental

2

0

1

0

23

26

control

8

0

0

0

2

10

Martis Creek

experimental

22

4

0

0

11

17

control

5

0

0

0

0

5

Totals

experimental

54

6

6

1

189

256

control

Total Reared:

43

0

0

0

2

45

301

^most individuals show definite “hygiaea” tendencies.

^includes one thinly scaled and otherwise aberrant animal (see text).

of Alaskan and Sierran animals could reflect the different temperature
regimes and local adaptation to them, but until considerably more is
known of the ecology and genetics of this butterfly it is not possible to
distinguish among these three hypotheses. Despite its abundance, the
Mourning Cloak is rather poorly known biologically (Young, 1980).

Acknowledgements: I thank Marc Minno, Michael Collins, and James W. Beever
m for assistance and Wanda Reynolds and Carol Erickson for discussion.

Literature Cited

FISCHER, E. 1907. Zur Physiologie der Aberrationen- und Varietaten- Bildung der
Schmetterlmge. Archiv fur Rassen- und Gesellschafts-Biologie 4(6): 761-793.
SHAPIRO, A. M. 1981. Phenotypic plasticity in temperate and subarctic Nymphalis
antiopa (Nymphalidae): evidence for adaptive canalization. J. Lepid. Soc. 34: (in
press).

STANDFUSS, M. 1896. Handbuch der palaarktiscken Gross-Schmetteriinge furForscher
und Sammler. G. Fischer, Jena. 392 pp.

WADDINGTON, c. H. 1957. The Strategy of the Genes. Allen and Unwin, London. 262 pp.
YOUNG, A. M. 1980. Some observations on the natural history and behaviour of the
Camberwell Beauty (Mourning Cloak) Butterfly, Nymphalis antiopa (Linnaeus),
in the United States. Entomol. Gazette 31: 7-18.

Journal of Research on the Lepidoptera

19(2): 88-95, 1980(81)

Aberrant New Mexican Butterflies

Richard Holland

1625 Roma, N.E., Albuquerque, New Mexico 87106, USA

Abstract. Photographs of aberrant Plebejus acmon texanus, Euphilotes
rita rita, Speyria atlantis dorothea, Phyciodes campestris Camillas, Strymon
melinus franki, Thessalia theona thekla, Eyphydryas anicia alena and
Hesperia pahaska pahaska are shown. Limenitis astyanax arizonensis ab.
doudorffi is illustrated and discussed as an arizonensis X L. weidemeyerii
angustifascia hybrid.

Over the 18 seasons (1964-1981) I have collected in New Mexico
(Holland, 1974), it is inevitable I would encounter a number of aberrant
specimens. The purpose of this article is to illustrate the more unusual
New Mexican forms collected.

Figure 1 shows a male Plebejus acmon texanus Goodpasture with
radically abnormal black markings on the ventral wing surfaces. This
specimen was taken at a moist spot in a shady side canyon near Mayhill,
NM, at the confluence of Cox Canyon and Wills Canyon. The dorsal
surface of this specimen is normal. A number of other small blues were
present at this same moist spot; when they were disturbed and took flight,
the illustrated specimen immediately gave an appearance quite different
from anything in my previous experience. A typical specimen is shown for
comparison in Figure 1.

Figure 2 illustrates a normal female Euphilotes rita rita (B. & McD.) and a
specimen with the ventral forewing (YFW) postmedial black spots in cells
Ml and M2 markedly displaced distally so they fuse with the submarginal
spots. The two specimens in Figure 2 were taken in a series of 20 (lOcTcf
and 1099); none of the others exhibited the displaced postmedian spots.
The abnormality of the specimen in Figure 2 was not apparent until after
capture. The dorsal surface of this specimen is normal. Leeuw (1979) has
recently figured a specimen of Satyrium acadica acadica (Edwards) which is
remarkably similar to the aberrant P. acmon texanus shown here in Figure
1. On the basis of three examples in three different species, one could
speculate that lycaenids may have a tendancy towards atypical expression
of the postmedial and submarginal spots ventrally.

Figure 3 illustrates male and female Speyeria atlantis dorothea Moeck
which show grossly enlarged silvering of the basal part of the ventral

Not refereed

19(2): 88-95, 1980(81)

89

Fig. 1. Plebejus acmon texanus ventral surface:

Top, normal 27 Aug. 75, Curtis Canyon Dam, Sacramento Mts., nr.
MayhiU, Otero Co., NM, 6600' (2000 m), leg. R. Holland.

Bottom, aberrant cf, 20 May 75, confluence of Cox Canyon and Wills
Canyon, Sacramento Mts., nr. Mayhill, Otero Co., NM, 6500' (2000 m), leg.
R. Holland.

Fig. 2. Euphilotes rita rita ventral surfaces:

Top, aberrant 9,

Bottom, normal 9, both specimens 1 Sept. 80, 1 mi. E. of Organ, San
Augustin Pass, Organ Mts., Dona Ana Co., NM, 5500' (1650 m), leg. R.
Holland & J. McCaffrey.

hindwing (VHW), This figure also shows normal specimens for comparison.
The aberrant specimens were taken 14 days and five miles (eight km)
apart. It is thus doubtful, but not impossible, they were from the same
brood. Of some 50 other dorothea taken during 1966-68 on Mt. Taylor,
NM, no other aberrant examples appeared, nor have I ever seen a similarly
silvered atlantis (Edwards) of any subspecies from elsewhere. Due to the
double occurrence of this aberration, it is tempting to attribute its cause to
some recessive allele which is present with very low frequency in the Mt.
Taylor population of dorothea. Jn flight, such specimens were not
distinguishably abnormal, nor are their dorsal surfaces.

Figures 4. and -5. show what is probably an extreme melanic of Phyciodes
campestrk camillm Edwards, although the specimen may be Phyciodes
mylitta arizonemis Bauer. (Both species were present in numbers at the
time and place that this specimen was captured.) Note that even the

90

J. Res. Lepid.

forewing shape is abnormal. Instead of having the usual convex or straight
outer margin, this specimen has the margin slightly concave at Cui. The
specimen is a male. A normal P. campestris camillus is also shown in these
figures.

Figures 4 and 5 also show a normal male Strymon melinus franki Field
and a male of the form which has been called meinersi Gunder. In this form,
the normally orange-red ''thecla spot” in cell Cu2 on both surfaces of the
hindwing is pale yellow. The usual orange-red marking at the anal angle is
also pale yellow, as well as reduced in area. (In the specimen illustrated, a
part of the anal angle has been lost from both wings, especially the right.)

Fig. 3. Speyeria atlantis dorothea ventral surfaces:

Top left, normal cT, 19 July 68, 1 mi. S. of La Mosca Outlook, Mt. Taylor,
Valencia Co., NM, 10,300' (3150 m), leg. R. Holland.

Top right, normal 9, 16 July 66, San Mateo Spr., San Mateo Canyon, Mt.
Taylor, Valencia Co., NM, 8800' (2700 m), leg. R. HoUand.

Bottom left, aberrant cT, 30 July 66, Tapia Canyon, Mt. Taylor, Valencia
Co., NM, 9300' (2850 m), leg. R. Holland.

Bottom right, aberrant 9, 16 July 66, La Mosca Canyon, Mt. Taylor,
Valencia Co., NM, 9800' (3000 m), leg. R. Holland.

19(2): 88-95, 1980(81)

91

Fig. 4. Dorsal surfaces, all specimens except as noted leg. R. Holland:

Upper left, normal d‘ , Phyciodes campestris camillus, 18 Aug. 68, 1 mi. W. of
New Canyon C. G., Manzano Mts., Torrance Co., NM, 8200' (2500 m).
Left next to top, extreme aberration cT, Phyciodes campestris camillus (?),
same data as specimen above.

Left next to bottom, normal cT, Strymon melinus franki, 30 July 68, 2 mi. S.
of New Canyon C. G., Manzano Mts., Torrance Co., NM, 7400' (2250 m).
Bottom left, aberration meinersi cf, Strymon melinus franki, 3 mi. W. of
Sanostee, Chuska Mts., San Juan Co., NM, 5500' (1650 m).

(continued on page 95)

92

J. Res. Lepid.

Oddly, the meinersi specimen has the postmedian band inwardly suffused
with the normal melinus (Huebner) orange-red on the VHW, although the
amount of suffusion is greatly reduced. The VFW postmedian line
suffusion on the meinersi specimen is completely absent. Similarly, the
meinersi specimen has the tip of the antenna club a normal melinus orange -
red.

In addition to the figured specimen, I have a female meinersi taken in
poor condition at New Canyon, 7400' (2300 m), Manzano Mts., Torrance
Co., NM, 30 July 68, leg. R. Holland. Normal melinus may have the
abdomen either grey or with the last four segments orange-red. The
figured meinersi has a grey abdomen; the unfigured meinersi has the last
four abdominal segments dorsally white.

In April, 1974, at Alamo Canyon, near Alamogordo, NM, there was a
great population explosion of Thessalia theona thekla (Edwards). Figures
4 and 5 also show a female taken at this time with the normal theona
(Menetries) pattern almost completely obliterated. The normal form is
also illustrated. The complete brown occlusion of the normal VHW basal,
medial and marginal pearlish bands is especially striking. The marginal
pearlish band normally present on the VFW is similarly occluded.
Typically, the dorsal surface of theona thekla is tricolored: chocolate, light
fulvous and dark fulvous, with each color fairly sharply delineated. In the
aberrant specimen, the chocolate and dark fulvous are not at all sharply
delineated, and the light fulvous is completely absent except for a row of
submarginal smudges on the forewing. The usual ventral theona thekla
pattern is also tricolored: chocolate, dark fulvous and pearl, with the pearl
bands sharply set off into rectangular spots by chocolate scaling on the
veins. In this specimen, however, there is almost no pearl ventrally, and
certainly no hint of chocolate scaling on the veins. In normal theona thekla,
the ventral surface is not a particularly close replica of the dorsal surface.
However, the two surfaces are quite alike in this aberration. No other
theona thekla aberrations were seen in this population explosion.

Figures 4 and 5 additionally illustrate a female Euphydryas anicia alena
Barnes & Benjamin with the following abnormalities: the apex of both
surfaces of the forewings are unusually blackened; the usual transverse
basal and discal DFW and DHW black markings are absent; and the
medial and basal markings of the VHW are peculiar in having elongated
pearly spots, especially in cells SC and Cu2. Euphydryas species in general
appear to produce more frequent aberrant individuals than most other
genera, e.g., the forms figured in Comstock (1927). Indeed, I have another
aberrant alena from the same place. Normal alena is given in this figure as
well.

Finally, Figures 4 and 5 illustrate a male of what has been called
Limenitis astyanax arizonensis ab. doudoroffi (Gunder). Perkins and Garth
(1972) assert that the entity is almost certainly a hybrid between L.
weidemeyerii Edwards and L. astyanax (Fabricus).

19(2): 88-95, 1980(81)

93

Neither astyanax nor weidemeyerii are frequently encountered in the
Sacramento Mts. of southeastern New Mexico. Small L. astyanax
arizonensis colonies exist in at least two riparian canyons (Alamo Canyon
and Three Rivers Canyon) on the west slope of this range around 5500 to
6500' (1700 to 2000 m). The Three Rivers location is also the source of the
subject doudoroffi specimen. Limenitis weidemeyerii is even rarer in the
Sacramentos; I have a single specimen taken near the summit of Sierra
Blanca at 11,000' (3300 m), about five miles (eight km) firom where the sub-
ject specimen was captured. The single Sieira Blanca weidemeyerii specimen
appears to be ssp. angustifascia (B. & McD.), although Perkins and
Perkins (1975) indicate the Sacramento Mts. to be closer to the known
range of typical weidemeyerii. (Evidently Perkins and Perkins (1975) had
very scant Sacramento Mts. weidemeyerii available. In any event, my single
weidemeyerii specimen forms an uncertain basis for determining which
subspecies inhabits the Sacramento Mts. so tenaciously.)

Perkins and Garth (1972) illustrate an example of doudoroffi from the
Allen Hancock Foundation (AHF), and mention the existence of three
other specimens, one in the Los Angeles County Museum (LACM), one
taken by Bauer, and the type taken by M. Doudoroff.All four of these
specimens are from Arizona, where the weidemeyerii subspecies would
unquestionably be angustifascia, the same subspecies which appears to
exist in the area where this doudoroffi specimen was taken. The present
specimen differs only in very minor ways from that illustrated by Perkins
and Garth: On the DFW it has only two, not three or four subapical white
spots; and the whitish median band on the VFW anterior to M2 is more
expressed.

Perkins and Garth indicate that their doudoroffi specimen, like the one
illustrated here, has furruginous scales suffusing the apex of the VFW.
Tliey maintain that, in this trait, it resembles astyanax arizonensis, and
that in weidemeyerii angustifascia the corresponding suffusion is black.
However, my Sierra Blanca weidemeyerii has the area suffused with
ferruginous scaling as extensively as the doudoroffi. Indeed, approximately
half the weidemeyerri I have seen from New Mexico have at least some
ferruginous scaling in this area, although the Sierra Blanca specimen is the
most extensively so scaled.

Hybridization among Limenitis species is widely documented; see, for
example, Shapiro and Biggs (1968), Perkins and Gage (1970), Gage
(1970), Hovanitz (1949), Simpson and Pettus (1976), and Platt, Rawson
and Balogh (1978), Due to the extreme scarcity of both weidemeyerii and
astyanax in the Sacramento Mts.', it is easy to postulate a female of one
species accepting courtship from the most nearly eligible male to be
found — even if that male was of the other species. In any event, I am fully
convinced that the question mark of Perkins and Garth (1972) can now be
removed, and ab. doudoroffi can be recognized as a weidemeyerii X

94

J. Res. Lepid.

astyanax arizonensis hybrid. Such recognition is, of course, without
nomenclatorial significance, because aberration and hybrid names do not
have code standing.

After the initial submission of this article, a very baffling female Hesperia
was captured on the south slope of the Capitan Mts. in Lincoln Co., NM.
The entire VHW of this specimen, which is shown in Figure 6, is
immaculate — a condition not characteristic of any Hesperia. At the same
time and place, Hesperia pahaska pahaska Leussler (common) and H.
uncas uncos Edwards (occasional) were flying. Hesperia viridis (Edwards)
had been taken nearby earlier in the season. The aberrant specimen was
immediately recognizable as peculiar in the field. In fact, I first missed a
sitting shot while the insect was nectaring at a thistle. Upon returning to
the same thistle four hours later, I was rewarded by finding the striking

Figure 6

Hesperia pahaska pahaska
ventral surfaces:

Top, aberrant pahaska (?) 9,
26 July 80, Peppin Canyon, S.
slope, Capitan Mts., nr. Capi-
tan, Lincoln Co., NM, 7000'
(2150 m), leg. R. Holland.
Middle, intermediate pahaska
9, 14 June 80, road S. of Capi-
tan Gap at National Forest
boundary, Capitan Mts., Lin-
coln Co., NM, 6500' (2000 m),
leg. R. Holland.

Bottom, normal pahaska 9, 27
July 80, Pierce Canyon, S. slope
Capitan Mts., nr. Capitan, Lin-
coln Co., NM, 6300' (1900 m),
leg. R. Holland.

I

19(2): 88-95, 1980(81)

95

Hesperia had also returned. Later examination of H p. pahaska from this
locality revealed a female specimen with VHW markings greatly reduced,
but not completely missing. It is on the basis of this intermediate specimen
and the general abundance of pahaska in the area that I concluded the
immaculate specimen is probably also pahaska. The intermediate speci-
men and a typical pahaska female are also illustrated in Figure 6.
Literature Cited

COMSTOCK, J. A., 1927. Butterflies of California. Published by the author, Los
Angeles. 334 pp.

GAGE, E. V., 1970. A record of a naturally occurring Limemtis hybrid (Nymphalidae).
J. Lepid. Soc. 24(4): 270.

HOLLAND, R., 1974. Butterflies of six central New Mexico mountains with notes of
Callophrys (Sandia) macfarlandi (Lycaenidae). J. Lepid. Soc. 28(1): 38-52.
HOVANTTZ, w., 1949. Increasing variability in populations following natural hybridi-
zation, pp. 339-355. In Jepson, Mayr, & Simpson (eds.) Genetics, Paleontology,
& Evolution. Princeton Univ. Press, 474 pp.

LEEUW, L, 1979. Aberrant Satyrium a. acadica (Lycaenidae). J. Lepid. Soc. 33(3):
204-205.

PERKINS, E. M. & E. V. GAGE, 1970(1971). On the occurrence of Limenitis archippus X
L. lorquini hybrids (Nymphalidae). J. Res. Lepid. 9(4): 223-226.

PERKINS, E. M. & J. s. GARTH, 1972(1973). Limenitis weidemeyerii angustifascia X L.
astyanax arizonensis = (?) ab. doudoroffi (Gunder), 1934. J. Res. Lepid. 11(4):
229-234.

PERKINS, S. F. & E. M. PERKINS, 1975. The Subfamily Limenitinae. In: Butterflies of
North America, (ed. by Howe, W.). Doubleday & Company, New York, xiii + 633
p. + 97 pi.

PLATT, A. P., G. w. RAWSON & G. BALOGH, 1978. Interspecific hybridization involving
Limenitis archippus and its congeneric species (Nymphalidae). J. Lepid. Soc.
32(4): 289-303.

SHAPIRO, A. M. & J. D. BIGGS, 1968(1970). A Limenitis hybrid from New York. J. Res.
Lepid. 7(3): 149-152.

SIMPSON, R. G. & D. PETTUS, 1976. Records of Limenitis hybrids from Colorado. J.
Res. Lepid. 15(3): 163-168.

( continued from page 91 )

Upper right, Limenitis astyanax arizonensis aberration doudoroffi = L.
weidemeyerii X L. astyanax arizonensis cf, 23 June 75, Three Rivers C. G.,
nr. Carrizozo, Sacramento Mts., Lincoln Co., NM, 6500' (2000m).
Center, normal 9, Thessalia theona thekla, 30 Apr. 74, W. slope of San
Augustin Pass, 18 mi. E. of Las Cruces, Organ Mts., Dona Ana Co., NM, c.
5500' (1650 m), leg. G. S. Forbes.

Bottom middle, aberrant 9, Thessalia theona thekla, 28 Apr. 74, Alamo
Canyon, nr. Alamogordo, Sacramento Mtjs., Otero Co., NM, 5500' (1650
m).

Middle right, normal 9, Euphydryas anicia alena, 16 May 76, Grasshopper
Spr., nr. Ramah, Zuni Mts., McKinley Co., NM, 7500' (2300 m).

Bottom right, aberration 9, Euphydryas anicia alena, same data as
specimen above.

Fig. 5. Ventral surfaces of specimens in Fig. 4.

Journal of Research on the Lepidoptera

19(2): 96-100, 1980(81)

Diapause in Various Populations of Pieris napi L. from
Different Parts of the British Isles

E. Lees

and

D. M. Archer

Postgraduate School of Biological Sciences, University of Bradford,

Bradford, W. Yorks, England

Diapause has been defined (Beck, 1967) as a genetically determined
state of suppressed development, the manifestation of which may be
induced by environmental factors. It follows, therefore, that different,
geographically isolated populations of a butterfly should have different
assemblages of genes determining diapause and hence should show a
somewhat different diapause response to the same environmental condi-
tions. The phase of development in which Pieris napi L. undergoes
diapause is the pupal phase. Whether or not an individual undergoes pupal
diapause depends in part on the photoperiod and temperature en-
countered during late larval life (Danilevskii, 1961). This, in turn,
determines whether a particular population of the species is monovoltine
or polyvoltine. P. napi is usually regarded as being partially bivoltine in the
British Isles (Lees and Archer, 1974), but this statement hardly does
justice to the complex situation which prevails over the country as a whole.

Colonies of P. napi which are strictly monovoltine are known (Lees, 197 0)
and these are usually found at altitudes of 800 feet or more in Northern
England and Scotland. Elsewhere in the British Isles the species is at least
partially bivoltine or even partially trivoltine. It should be noted, however,
that even in those parts of the country where there is a second and third
brood, a significant part of the pupae derived from the spring generation
are diapause pupae and do not develop into butterflies until the following
year.

In order to learn more of the diapause characteristics of Pieris napi in
different parts of the British Isles, we sampled spring populations of the
species from various localities ranging from Teignmouth in S. Devon to
Dollar in Scotland, although the majority of samples were from popula-
tions in N. England (c.f. Fig. 1). Eggs from the captured females were kept
in plastic boxes until they hatched and the resultant larvae were
maintained under controlled conditions of photoperiod and temperature
until they pupated. The larvae were fed on the leaves of Hedge Garlic
iAlliaria petiolata)^ a common foodplant of this species in the field.

19(2): 96-100, 1980(81)

97

The photoperiod chosen for the experiments was 18 hours, which is a
“long-day’' photoperiod, similar to the daylength of the longest day over
much of England. The larvae were reared at two temperatures viz. 18
degrees C and 12 degrees C. The former is similar to the mean maximum of
the warmest month in S. England, the latter is similar to the mean
maximum of the warmest month at altitudes of 1000 feet in N. England and
Scotland. The numbers and percentages of diapause and non-diapause
pupae obtained in the various experiments are shown in Tables 1 and 2.

Table 1

Experiments at 18 Kfr. Photoperiod and 18 De^ees C

Locality

Elevation

(M.)

Offspring
of Female

No. of Diapause
Pupae Produced
Among Progeny

No. of Non-Diapause
Pupae Produced

Among Progeny

Dollar

48

1

5 (19.2%)

21 (80.8%)

Dollar

48

2

6 (16.6%)

30 (83.4%)

Dollar

48

3

8 (23.5%)

26 (76.6%)

Dollar

48

4

6 (14.0%)

42 (86.0%)

Bellingham

122

1

8 (16.6%)

40 (83.4%)

BeUingham

122

2

6 (14.3%)

36 (85.7%)

Bellingham

122

3

6 ( 8.7%)

39 (91.3%)

Bellingham

122

4

4 (18.2%)

18 (81.8%)

High Force

381

1

7 (15.5%)

38 (84.6%)

Ifigh Force

381

2

8 (16.6%)

40 (83.4%)

High Force

381

3

9 (20.9%)

36 (79.1%)

Ihgh Force

381

4

4 (16.0%)

21 (84.0%)

Amside

15

1

6 (17.6%)

38 (82.4%)

Amside

15

2

2 ( 6.9%)

27 (93.1%)

^^side

15

3

3 (10.4%)

25 (89.6%)

SU’ensaU

16

1

6 (14.3%)

36 (85.7%)

Strensall

16

2

3 (10.8%)

25 (89.2%)

Strensall

16

3

4 (11.8%)

30 (88.2%)

SteensaU

16

4

2 ( 6.6%)

28 (93.4%)

Menston

48

1

3 (13.1%)

20 (86.9%)

Menston

48

2

2 ( 7.3%)

25 (92.7%)

Menston

48

3

2 ( 8.7%)

23 (91.3%)

Menston

48

4

4 (11.8%)

30 (88.2%)

Nantmch

30

1

2 ( 8.7%)

22 (91.3%)

Nantwich

30

2

1 ( 4.0%)

24 (96.0%)

Nantwich

30

3

3 ( 7.9%)

36 (92.1%)

Teignmouth

8

1

2 ( 6.7%)

28 (93.3%)

Teignmouth

8

2

4 (11.8%)

30 (88.2%)

Teignmouth

8

3

3 ( 7.7%)

36 (92.3%)

Teignmouth

8

4

2 ( 6.4%)

29 (93.6%)

98

J. Res. Lepid.

Table 2

Experiments at 18 Hr. Photoperiod and 12 Degrees C

Locality

Elevation

(M.)

Offspring
of Female

No. of Diapause
Pupae Produced
Among Progeny

No. of Non-Diapause
Pupae Produced
Among Progeny

Dollar

48

1

31 (100%)

0 ( 0.0%)

Dollar

48

2

43 (100%)

0 ( 0.0%)

Dollar

48

3

19 (100%)

0 ( 0.0%)

Dollar

48

4

25 (100%)

0 ( 0.0%)

Bellingham

122

1

23 (100%)

0 ( 0.0%)

Bellingham

122

2

37 (100%)

0 ( 0.0%)

Bellingham

122

3

24 (100%)

0 ( 0.0%)

Bellingham

122

4

36 (100%)

0 ( 0.0%)

High Force

381

1

20 (100%)

0 ( 0.0%)

High Force

381

2

34 (100%)

0 ( 0.0%)

High Force

381

3

31 (100%)

0 ( 0.0%)

High Force

381

4

28 (100%)

0 ( 0.0%)

Amside

15

1

24 (88.8%)

3 (11.2%)

Amside

15

2

48 (85.7%)

8 (14.3%)

Amside

15

3

30.(83.3%)

6 (16.7%)

Amside

15

4

20 (77.0%)

6 (23.0%)

Strensall

15

1

21 (91.3%)

2 ( 8.7%)

Strensall

15

2

27 (90.0%)

3 (10.0%)

Strensall

15

3

50 (89.4%)

6 (10.6%)

Menston

48

1

22 (84.6%)

4 (15.4%)

Menston

48

2

40 (85.1%)

7 (14.9%)

Menston

48

3

42 (89.4%)

5 (10.6%)

Nantwich

30

1

20 (80.0%)

5 (20.0%)

Nantwich

30

2

37 (83.4%)

7 (16.6%)

Nantwich

30

3

39 (81.4%)

8 (18.6%)

Teignmouth

8

1

29 (82.9%)

6 (17.1%)

Teignmouth

8

2

39 (79.6%)

10 (20.4%)

Teignmouth

8

3

32 (80.0%)

8 (20.0%)

Teignmouth

8

4

21 (81.6%)

5 (18.5%)

From the results given in Table 1 it is obvious that larvae from all
localities, maintained at a temperature of 18 degrees C and 18 hrs
photoperiod, produced some pupae which were diapause and some which
were non-diapause. In all cases the numbers of non-diapause pupae greatly
exceeded the numbers of diapause pupae, but the percentages of non-
diapause pupae were highest in the population from S. England
(Teignmouth) and from locahties in N. England at low altitudes. It is
reasonable to infer that at a temperature of 18 degrees C, populations of P.
napi would be overwhelmingly bivoltine, although in all the localities we

19(2): 96-100, 1980(81)

99

have sampled at least some genotypes for monovoltinism are present. Hus
temperature is above the mean temperature of the wairmest month in any
part of the British Isles, so that populations in nature are likely to be
somewhat less bivoltine than the experimental data suggests.

The results in Table 2 indicate that larvae maintained at the lower
temperature of 12 degrees C behave very differently from those at 18
degrees C, in the extent to which they yield diapause puape. Larvae from
Aree localities (High Force, Bellingham and Dollar) gave no non-diapause
pupae under these conditions and those from other localities did not give
more than 20% non-diapause pupae. Once again the percentages of non-
diapause pupae were highest in the population from S. England
(Teignmouth), and the populations which behave in a strictly monovoltine
fashion are those from high altitudes in N. England and from Scotland. The
experimental data is therefore in agreement with what is observed in
nature viz. the occurrence of monovoltine colonies in localities where the
summer temperatures are low.

Our findings suggest that populations of P. napi show a physiological
cline in relation to the tendency of their larvae to give diapause pupae as
one moves from north to south in the British Isles. Northern populations
are more prone to give diapause pupae than southern ones and there is a
weU-defined gradient wiMi extremes at Dollar in Scotland and Teignmouth
in S. England. This physiological dine is accompanied by a phenotypic
^adient. The female butterflies from Scotland and the extreme north of
England have much darker veining to the wings and a proportion of them
have a ground colour which is some shade of yellow instead of white. These
tendencies, which are more marked in the -spring brood, become prop*es-
sively less evident as one moves from north to south. Warren (1968) has
suggested that the northern forms of P. napi should be regarded as foming
a distinct subspecies P. napi thomsord Warren, and Thomson (1970) has
described the distribution of the latter. It is obvious from Thomson’s
account that the phenotypes he describes constitute a dine.

Literature Cited

ffiCK, s. D. 1968. Insect Photoperiodism. New York and London.

DAMLEVSKn, A. s. 1961. Photoperiodism and Seasonal Development of Insects.
Leningrad.

LEES, E. & D. M. ARCHER. 1974. Ecology of Pieris napi (L.). (Lep. Pieridae) in Britain.
Entomologist’s Gaz. 25: 231-237.

I.BES, E. 1970. A univoltine race of Heris napi in the Yorkshire Pennines, Entomolo-
gist, 103: 260-262.

IHOMSON, G. 1970. The diefeibution and nature of Pieris napi thomsoni WaiTen (Lep.

Pieridae). Entomologist’s Record 82: 255-261.

WARREN, B. c. s. 1968. On an instable race of Pieris adalwinda, located in Scotland.
Entomologist’s Record 80: 299-304.

100

J. Res. Lepid.

Fig. 1. Locations of sampled P. napi populations.

1 - EloUar; 2 - Bellingham; 3 - ffigh Force; 4 - Amside; 5 - StrensaU; 6 -
Menston; 7 - Nantwich; 8 - Teignmouth.

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THE JOURNAL OF RESEARCH
ON THE LEPIJOOPTERA

Volume 19

Number 2

m THIS ISSUE

Date of Publication: September 7, 1981

Nantucket Fine Tip Moth, Rhyacionia frustrana,

Countv, California: Integrated Control and moiogicai
Notes (Lepidoptera: Tortricidae, Olethreutinae)
lennis M. Poore

Moths of I^rth America North of Mexico, Supplemental
literaiure: 1

C. E. Riotte

Acknowledgement

Demonstration of Reproductive Isolating Mechanisms in
Callosapiia (Satumiidae) by Artificial Hybridization
lard S. Peigler

Canalization of ti^e Phenotype of Nymphalis antiopa
(LepidoptergiNNymphalidae) from Subarctic and
Montane Climatel
Arthur M.

Aberrant New Mexic^Butterflies
Richard Holland

Diapause in Variods Populations of Pieris napi L. from
Different Paks of the British Isles

E. Leei

D. M. Archer

Cover Illustration Pen and ink rendering of C. securifera c? X C. promethea ^Ly

Robin Jarrett of Los Angeles, CA. See R. Peigler article, page 72. ’

72

Volume 19

Number 3

1980(81)

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,^'6RAR!ES

Journal of Research on the Lepidoptera

19(3): 10M80, 1980(81)

A Revision of the American Genus Amsoto (Satumiidae)

J. C. E. Riotte

Department of Entomology, Bernice P. Bishop Museum, Honolulu, Hawaii 96819 and Royal
Ontario Museum, Toronto, Canada MSS 2C6

and

Richard S. Peigler

Department of Entomology, Texas A&M University, College Station, Texas 77843

Introduction

Michener (1952) revised the Satumiidae of the Western Hemisphere at the
generic level, but except for occasional mention of species, these were not treated.
Recent discovery of new species of the genus Anisota Huebner in Ontario, Texas,
South Carolina, and Mexico made it desirable to do research into the morphology of
all related species, and as no details have been published about the adults and
immature stages of all the North and Central American species of Anisota since
Packard (1905) and Bouvier (1931), it seemed advisable to summarize our
Imowledge in this paper.

Higher Classification

The genus Anisota belongs to the moth family Satumiidae belonging to the
superfamily Bombycoidea which also includes Lasiocampidae and Bombycidae
(Ferguson, 1971).

The correct subfamily name has been, and continues to be, disputed. Lemaire
(1976) indicated that Adelocephalinae, Boisduval 1868, is the correct name. Earlier
authors used other names such as Diyocampinae, Ceratocampinae, and Syssphing-
inae. The name Citheroniinae has had wide usage, even in recent works (Michener,
1952; Ferguson, 1971). The subfamily has frequently been elevated to family rank
under the various names throughout much of the literature (i.e., Citheroniidae,
Syssphingidae). Whether considered a family or subfamily, the group is clearly a
monophyletic assemblage. It is one of the more primitive subfamilies of Satumiidae
’and is exclusively New World in distribution. Most genera and species are
neotropical; Anisota is in fact, the only predominantly nearctic genus of the
subfamily, excepting the allied Dryocampa Harris. Michener (1952) and Ferguson
(197 1) characterized the subfamily morphologically — it is our purpose from here on
to deal only with the genus Anisota.

As the lists of synonyms indicate, species of Anisota have often appeared in the
literature under several generic names (Adelocephala, Dryocampa, etc.). Fortunately
however, the name Anisota has rarely been applied to species which are not tme
Anisota, an exception being the Brazilian Psigida apollinairei (Dognin) which Grote
(1867) redesciibed as Anisota walkeri.

Michener (1952) subdivided the genus Anisota into two subgenera: Dryocampa
and Anisota, the characters of which may be found in Table 9 of his paper. Ferguson
(197 1), however, pointed out correctly that the one species belonging to Dryocampa

102

J. Res. Lepid.

“seems quite far removed from all the species of Anisota” He therefore elevated
Dryocampa to generic rank in which the present authors follow him, especially
because of the characters shown in the immature stages, the male genitalia and
other morphological characters. In the present paper therefore we only treat the
species of Michener’s subgenus Anisota. Dryocampa represents a separate migra-
tion into North America from the neotropical fauna. The genus is perhaps closest to
Anisota {sensu stricto), an idea supported by the work of Pease (1961). We propose
that Dryocampa and Anisota are sister-groups, diverged in Central America, and
radiated into North America where their respective hostplants {Acer and Quercus)
are more available.

Methods

Genitalic dissections were made of a certain number of male and female
specimens of each species from different populations where applicable. Also the
heads of male and female specimens were detached from the body, cleaned,
measured as to eye-size and head-proportions as well as the form of the rear joint of
the head. Also anterior legs were removed and cleaned for examination of the
epiphysis. Labial palpi were also removed and cleaned and studied.

Special emphasis was placed on the immature stages. In the larvae, the suranal
plate was studied and figured, which is distinct for each of thespecies. The pupae
were photographed in a position to show their specifically different cremaster. In
two species, scanning electron micrographs (Plate V) were made of the cuticle of
larvae (see Byers & Htnks, 1973).

Localities and flight periods for specimens examined are cited under each species
account. Full specimen data have been deposited in the library of the Royal Ontario
Museum and are accessible for reference. Most of the material collected or received
by the authors during the course of this study has been deposited into major
museums (notably ROM, AMNH and LACM). The abbreviations used in the text
for collections from which material for this study was examined are as follows:

AMNH

BPBM

BMNH

CalAcSci

CLU

CM

CNC

FIS

FMNH

FSCA

Heitzman Coll.
IBM

Kendall Coll.
LACM

Lemaire Coll.

LSU

LM

McM

American Museum of Natural History, New York, New York

Bernice P. Bishop Museum, Honolulu, Hawaii

British Museum (Natural History), London, England

California Academy of Sciences, San Francisco, California

Clemson University, Clemson, South Carolina

Carnegie Museum of Natural History, Pittsburgh, Pennsylvania

Canadian National Collection of Insects, Ottawa, Ontario

Forest Insect Survey, Canada

Field Museum of Natural History, Chicago, Illinois

Florida State Collection of Arthropods, Gainesville, Florida

Collection of J. Richard Heitzman, Independence, Missouri

Institute de Biologia, Mexico City, Mexico

Collection of Roy & C. A. Kendall, San Antonio, Texas

Los Angeles County Museum of Natural History, Los Angeles,

California

Collection of Claude Lemaire, France (now in MNHN)
Louisiana State University, Baton Rouge, Louisiana
L5nnan Entomological Museum, Ste-Anne-de-Bellevue,
Quebec

McMaster University, Hamilton, Ontario

19(3): 10M80, 1980(81)

103

MHNM

MNHN

MSU

Peigler Coll

RIB

ROM

Sieker Coll.

TAMU

UArk

UCal

UFl

UG

UGa

UIl

UM

UMo

USNM

USPM

UWO

YPM

Museo de Historia Natural, Mexico City, Mexico
Cambridge, Massachusetts

Museum National d’Histoire Naturelle, Paris, France

Michigan State University, Lansing, Michigan

Collection of R. S. Peigler, Greenville, South Carolina

Entomology Research Institute, Belleville, Ontario (now closed)

Royal Ontario Museum, Toronto, Ontario

Collection of William E. Sieker, Madison, Wisconsin

Texas A&M University, College Station, Texas

University of Arkansas, Fayetteville, Arkansas

University of California, Berkeley, California

University of Florida, Gainesville, Florida

University of Guelph, Ontario

University of Georgia, Athens, Georgia

University of lUinois/Illinois Natural History Survey

University of Michigan, Ann Arbor, Michigan

University of Missouri, Columbia, Missouri

United States National Museum of Natural History,

Washington, D.C.

University of Minnesota, St. Paul, Minnesota
University of Western Ontario, London, Ontario
Yale University’s Peabody Museum of Natural History, New
Haven, Connecticut

Acknowledgements. Although it would be impossible to name everyone here who
helped us with this revision, we begin by thanking the following persons for making
available material in their care of their respective institutions: F. H. Rindge
(AMNH), P. H. Amaud, Jr. (CalAcSci), J. R. Holman (CLU); the late H. K. Clench
(CM), E. Munroe and D. F. Hardwick (CNC); C. R, Beutelspacher (IBM), A. H.
Rose and P. D. Syme (Environment Canada, FIS), R. L. Wenzel (FMNH), H. V.
Weems, Jr., F. W. Mead and G. W. DeWe (FSCA), J. P. Donahue (LACM), P. E. L.
Viette (MNHN), J. B. Chapin (LSU), V. R. Vickery (LM), D. M. Davies (McM), J.
M. Bums (MCZ), R. L. Fischer (MSU), P. Beime (RIB), G. B. Wiggins (ROM), J. A.
Powell (UCal), D. H. Pengelly (UG), C. L. Smith (UGA), J. G. Stemburg (UIL), T. E.
Moore (UM), W. R. Emis (U. Mississippi), D. C. Fdrguson (USNM), P. J. Clausen
(USPM), W. W. Judd (UWO), and K W. Brown (YPM).

We extend our appreciation to the following persons for help in various ways: S. B.
Atkins, H. D. Baggett, V. A. Brou, Earll M. Brown, G. R. Camer, the late R. B.
Dominick, J. E. Eger, Connie A. Kendall, Bert Kohlmann, Peter Loy, J. Wm.
McCord, B. Mather, R. T. Mitchell, W. E. Sieker, J. D. Solomon, and B. D.
Williams, EH.

Special thanks go to R. A. Crowson, Dept, of Zoology, Univ. of Glasgow, for
providing photographs of A. stigma F.; to Th. Finlayson for donating ail research
material of her late husband to this study and for taking the senior author to the type
locality of A. finlysoni in 1967; for making photographs used in our color plates we
thank Joseph E. Eger, n (Plate I, figs. 2, 7, 12; III, 1; IV, 11), G. R. Camer (1, 6, 10),
Robert R. Murray (I, 9), W. D. Winter, Jr. (1, 8), C. Lemaire (EV, 5); all others were
made at the ROM (L. Warren, Head Photographer). Gratefully acknowledged is the
assistance of E. Munroe, the late H. K. Clench, G. B. Wiggins, and D. Barr (ROM)

104

J. Res. Lepid.

for critically reading early drafts of this manuscripto The excellent drawings in this
paper were all made by artists at the ROM: Anker Odum and Zile Zichmanis.
Acknowledgment is also made for the use of the scanning electron microscope (in
ROM) established through a grant from the National Research Council to the Dept,
of Zoology, Univ, of Toronto, for development of a program in systematic and
evolutionary zoology.

We extend our gratitude for determinations of parasites to the following research
scientists of the U. S. Dept, of Agriculture, Beltsville, Maryland: G. Gordh, R. W.
Carlson, L. Knutson, C. W. Sabrosky, P. M. Marsh, and E. E. Grissell. J. M. Tucker
identified Mexican oaks for us.

The very constructive criticisms and suggestions given by an anonymous reviewer
for the Journal were much appreciated, and we thank this person for helping
improve the overall work in several ways.

Finally we express our deepest appreciation to Roy 0. Kendall and Dr. Claude
Lemaire who have over a period of several years generously and consistently
supplied rearing notes, data and material from their collections, and have helped
directly in numerous other ways.

Generic Diagnosis and Morphology of the Imago

ANISOTA HUEBNER

Anisota Huebner, 1820: 192; Michener, 1952: 409. Type species: Bombyx
stigma Fabricius, designated by Grote, 1874: 260.

A North and Central American genus of small to medium sized moths of brownish
color; always with extremely reduced clypeus in the adult and a white discal spot on
the forewing; the male in many cases diurnal; strong sexual dimorphism in some of
the species, some sexual dimorphism in all species. Morphological features which
all species of the genus have in common as adults are:

a) male antennae doubly bipectinate; female antennae simple and ciliate

b) epiphysis in many females absent, not as Michener (1952: 408) and
Ferguson (1971: 63) say in all females; Michener’s own preparation (AMNH)
of an anterior leg of a female of A. virginiensis has an epiphysis; when present
much reduced in size

c) anterior tibia in male and female dorso-apical with two lateral projections,
the exterior one usually a long spine, the interior one a much shorter and
usually blunt projection; middle and posterior tibiae also with two projections
of this kind, but much shorter and in many cases similar in size; middle and
posterior tibiae in males and females bear one pair each of tibial end-spurs; a
pair of peculiar spines in the females only arises from the apical lobes of the
penultimate anterior tarsal segment. Leg morphology is described and
figured by Oiticica (1940).

d) pulvillus, paronychium (lateral lobe) and empodium (generally only one
tubercle with one long strong seta) well developed

e) free anellus in male genitalia present and valves not fused

f) uncus of male genitalia without protuberance

Michener (1952) says of Anisota: “This subgenus includes most of the species of
Anisota. They are brown, with a small white spot at the apex of the discal cell of the
forewing. It includes two rather well-marked groups of species distinguished
primarily by the size of the eyes. In dissimilis Boisduval and oslari Rothschild the

19(3): 10M80, 1980(81)

105

eyes are reduced in size so that their upper ends do not reach the lower margins of
the antennal sockets, and their lower ends do not reach the lower extremity of the
frons. In this group, moreover, the frontal protuberance is more reduced and the
proboscidial fossa is even shallower than in the other group which contains the
remainder of the species. In dissimilis and oslari also the labial palpi are more
distinctly two-segmented than in the remaining species of the genus. The species of
this subgenus occur in eastern North America, westwaird to Arizona and south into
Mexico.”

The basic premise of the foregoing is valid. The species olAnisota can indeed be
separated into two groups with the aid of the size of the eyes, or of a certain ratio of
the head and the position of the eyes or of the form of the frontal protuberance, or on
the basis of the visibility or obscuration of the laterofrontal suture. All of these
criteria give rise to identical groupings, and it should be taken into account that it is
not so much the size of the eyes as their position and angle to the head which makes
the difference, and which makes the eyes appear to be of quite different size. To
compare eyes of these moths without exact measurements gives very wrong results;
Ferguson (1971: 74) mentions the “very small eyes” of the males of A. senatoria. It
is in fact astonishing that exact measurements of the eyes do not show a great
difference among the species, except for A. virginiensis which was included by
Michener among the “big-eyed” species and actually has the smallest eyes of the
entire genus, as correctly pointed out by Ferguson (1971: 74). On the other hand,
the eyes of A. dissimilis and A. oslari are in no way “very reduced” as Ferguson
(1971: 64) says. Table 1 of the eye measurements will better illustrate these points.

TABLE 1

Eye Measurements (in mm)

Species

a-b in male

a-b in female c-d in male

c-d in

Group A

A. consularis

1.30

1.31

0.70

0.65

A. fuscosa

1.70

1.56

1.09

0.96

A. manitobensis

1.43

....

0.87

....

A. senatoria

1.33

1.24

0.70

0.62

A. stigma

1.37

1.44

0.97

0.95

Group B

A. assimilis

1.23

....

0.67

....

A. discolor

0.98

....

0.50

....

A. dissimilis

1.23

....

0.70

....

A. finlaysoni

1.17

1.17

0.67

0.63

A. oslari

1.14

....

0.55

....

A. peigleri

1.33

1.33

0.60

0.80

A. pellucida

1.04

1.21

0.55

0.72

A. punctata

....

....

....

....

A. virginiensis

0.93

1.08

0.50

0.53

106

J. Res. Lepid.

Fig. 1. The two types of heads in Anisota, with large and small eyes. See Tables 1

and 2.

The table clearly shows group A with the large eyes and group B with the small
eyes. Ingroup A the measurement a-b does not go significantly below 1.33 mm while
in group B it does not go over 1.33 mm. In group A the measurement c-d does not go
below 0.70 mm while in group B it does not exceed 0.70 mm. Females show slight
differences with the males; in group A their measurements tend to be lower than in
the males while in group B they tend to be slightly higher. Regrettably there were
not enough females available to check every species. In group A the eye
measurements separate A. fuscosa from all the other species, especially from the
closely related A. stigma. In group B, A. peigleri shows a slight deviation with a-b the
highest for the group and c-d just in the middle between the highest and lowest
values.

Further examination has shown that in no species the eye reaches the antennal
socket at all. To see this clearly the head should be detached, cleaned and
measured. If this is done, one finds that the distance between the two antennal
sockets (e-f) and the distance between one antennal socket and the upper edge of
the eye (g-h), give a ratio which can be very useful in cormection with the values of
Table 1 . See Table 2 and Figure 1 . The data in Table 2 corroborate the conclusions
from Table 1, namely there are two groups of Anisota species which can be
separated by certain eye and head measurements. It is also remarkable that
visibility of the laterofrontal suture and the size of the frontal protuberance support
the groupings established with our measurements. It is interesting to note that A.
fuscosa also deviates here by far from the normal values while everything otherwise
falls nicely into place. Because of the dissection necessary and the scarcity of some
species in collections, it was possible to measure for some species only one
specimen, but in the other species where a series of measurements was taken, the
variation was very small, indicating that the single ones measured are probably
representative of their species, at least in the present context.

The labial palpi show for the entire second group a more distinctly two -segmented
condition than in the first group.

Many specimens were dissected as to the genitalia and other anatomical features;
also Michener’s dissections (AMNH) were used. The form of the aedoeagus of the
male genitalia can be furthermore used to divide the second group into two
subgroups: A. dissimilis, A. finlaysoni, A. oslari, and A. peigleri have a straight

19(3): 101-180, 1980(81)

107

TABLE 2

Head Morphology
(numbers in mm)

ratio of

e-fig-h

latero frontal

frontal

Species

male

female

suture

protuberance

Group A

A. consularis

3.6:1

3.1:1

slightly obscured very prominent

A. fuscosa

6.7:1

6.0:1

obscured

protruding

A. manitobensis

3.3:1

absolutely

obscured

prominent

A. senatoria

3.0:1

3.2:1

slightly obscured protruding

A. stigma

3.6:1

3.3:1

obscured

protruding

Group B

A. assimilis

2.0:1

visible

reduced

A. discolor

1.7:1

visible

reduced

A. dissimilis

1.8:1

visible

reduced

A. finlaysoni

1.8:1

2.7:1

visible

reduced

A. oslari

1.9:1

visible

reduced

A. peigleri

1.8:1

visible

slightly

protruding

A. pellucida

1.8:1

1.8:1

visible

reduced

A. punctata

A. virginiensis

1.7:1

1.8:1

visible

reduced

aedoeagus, whOe A. assimilis, A. discolor, A. pellucida, and A. virginiensis have the
aedoeagus curved. In the first group the aedoeagus is straight in all species. It is
noteworthy that the female genitalia, which were not studied by Ferguson (1971),
also support the above grouping: only the A. assimilis subgroup has a prominent
antrum and this is of an unusual form when seen in lateral view (Figure 8), and in all
species of the second group the ductus bursae makes an abrupt, almost rectangular
turn where the bursa copulatrix is attached. In all species of the first group the
ductus bursae is straight.

In some species of the genus a special sexual dimorphism occurs as a parallel
development in both groups. This character has been used by previous authors
for the purpose of subdivision of the genus into species groups. However, this seems
to be artificial and untenable and it should be recognized that other morphological
features form a better basis for a natural grouping of similar species within Aniso to.
For a correct interpretation of the somehow striking sexual dimorphism see
Ferguson (1971: 63). Some diurnal males have evolved very transparent forewings
(A. pellucida, A. discolor, A. virginiensis) or tend toward that (A. senatoria, A.
peigleri); other diurnal males (A. oslari) show no such trend but nonetheless
probably also resemble orange wasps (V espidae) in flight.

It does not seem justifiable to correlate reduction in eye size with a trend to
diurnal activity (see Ferguson, 1971: 63). Our Table 1 shows clearly that there are
small eyes among species with sexual dimorphism as well as among those without,

108

J. Res. Lepid.

and the same is true for large eyes. It is also important to consider the statement of
Mazokhin-Porshnyakov (1969: 2): “As a rule, the mobile and fast flying species,
both diurnal and nocturnal, have large complex eyes; male insects often have bigger
eyes than females.”

Morphology of the Immature Stages

Egg. This is, according to Packard (1905): “elliptical, flattened, each end alike, a
little longer than broad, shell very thin, parchment like, the surface with obscurely
marked microscopic irregular hexagonal areas. The eggs are laid in large patches on
leaves”. We add: patches on older leaves with preference at apices of leaves on the
lower perimeter of the tree.

Larva. Larvae, according to Packard (1905) with “body cylindrical, slightly
flattened at the end; armature consisting of two recurved slender smooth meso-
thoracic horns [except for A. finlaysoni]; prothoracic spines reduced to low
tubercles;all other dorsal and lateral spines of the body small, conical, acute; the
two dorsal spines of the eighth abdominal segment, remote, not fused into a caudal
horn. Suranal plate rugose or granulated, with from three to four lateral and two
terminal stout conical spines. Body with conspicuous longitudinal bands or
stripes.” The latter, however, is not always the case. The headcapsules and suranal
plates are specific and figured as far as possible. Here it should be noted that in
Peterson (1959: Fig. L 12) the captions of H and C should be interchanged, i.e., H
should read “A. senatoria” and C should read “A. stigma”.

Pupa. About the pupa Mosher (1916) (see also Mosher, 1914) says: “Without
prominent scattered spines on the thoracic segments, the longest never four times
the length of those covering the segments; antennae with the central axis never
bearing pominent spines, the spines never curving posteriorly; maxillae always one-
fourth the length of wings”. We provide figures of the species-specific cremasters of
species for which these were available. These structures have been shown to be
taxonomically useful (Giehsler, 1965).

Intrageneric Classification and Phylogeny

Different groupings may be derived for the 15 species oiAnisota, depending on
characters used. Eye size and head morphology discussed earlier give one such
arrangement, whereas genitalia give another. However, we prefer the following
groupings based on habitus of larvae and adult moths as well as geographical
distributions:

A. stigma group

A. stigma
A. fuscosa
A. consularis
A. manitobensis

A. senatoria group

A. senatoria
A. finlaysoni
A. peigleri

A. pellucida group

A. pellucida

A. leucostygma

A. virginiensis
A. discolor

Mexican group

A. dissimilis
A. punctata
A. assimilis
A. oslari

19(3): 10M80, 1980(81)

109

Certain features of the zoogeography are clear while others are difficult to
ascertain. The genus evidently originated in Mexico or southward, possibly as
specialized feeders on oaks. The distribution and numbers of species in North
America was likely greatly influenced by glaciation and other climatic changes
during the Pleistocene.

Larger size of the adults, very spiny larvae, south Mexican distribution, and
noctumalism are judged to be plesiomorphic characters (based on outgroup
comparison) in such species as A. dissimilis. Both A. dissimilis and A. punctata show
affinities with the A. stigma group; A. assimilis shows a relationship to the A.
pellucida group through the genitalia. All of the species appear to be still very
closely interrelated and the above four groupings merely indicate still closer
affinities. It is likely that there remain undiscovered species in Mexico or
southward. Lepidopterists have traditionally collected most heavily in lowland
rainforests where the fauna seems richer, and Anisota are more likely encountered in
oak-pine biotope in the mountains. If new species are described from the United
States or Canada in the future, these will likely be ones which are presently “hiding”
under names of widespread and variable species such as A. stigma, A. virginiensis,
and A. senatoria.

The phenomenon of allopatry is quite evident with the most closely related
species. When species are so alike in their ecological requirements (i.e., fill the same
niche), then they are mutually exclusive or ecologically antagonistic, possibly due to
competition or more likely the absence of isolation mechanisms to prevent
hybridization (see Mayr, 1969, p. 193-197; Stebbins, 1969). Such allopatry or
parapatry have led workers to consider that some of these species are merely
subspecies. Our observations indicate that all 15 taxa which we herein recognize are
full, discrete species. We have noted consistent differences in genitalia of both
sexes, adult phenotype and leg morphology, larval color and morphology, pupal
morphology, egg size and embryo development; and in the A. pellucida group we
have observed sterility and inviability in Fi and backcrosses of artificial hybrids.
Furthermore we find no indications of blend zones or dines, despite the immense
amount of material examined. Van Valen (1976) offers an interesting discussion on
these matters.

To elaborate on this important concept of allopatry let us view some specifics. Any
given locality tends to have only one representative from a given group: in New
Jersey one will collect A. stigma, A. senatoria, and A. virginiensis; in upper South
Carolina one collects A. stigma, A. peigleri, and A. pellucida; in East Texas we find A.
fuscosa, A. senatoria, and A. discolor.

Although we do not promote usage of common (vernacular) names for these
insects, a few remarks may be appropriate here. According to one list (Benoit, 1975)
the English and French common names for four species are: A. finlaysoni,
shorthomed oakworm, anisota de Finlayson; A. senatoria, orange-striped oakworm,
anisota a lignes orangees; A. stigma, spiny oakworm, anisote stigma; A. virginiensis,
pink-striped oakworm, anisote rose du chene. Holland (1903) refers to A.
virginiensis as “the Virginian Anisota” and A. stigma as “the Stigma Moth”. Names
such as spiny, pink-striped, and orange -striped oakworms have become less
appropriate with the discoveries of more species in recent years, since each name
might apply to an entire group of species.

The species of Anisota are arranged alphabetically in the text for convenient
reference in their groups: Mexican group - pellucida group - senatoria group -
stigma group.

no

J. Res. Lepid.

Key to Adults

1. antennae doubly bipectinate (males) 2

antennae simple and ciliate (females). 5

2. forewings with hyaline area around discal cell. 3

forewings without hyaline area around discal cell 4

3. medium reddish brown; outer margin of both wings perceptibly convex; anal

angle of hindwings all rounded; aedoeagus of male genitalia rounded

A. virginiensis

deep reddish brown; outer margin of both wings straight or nearly so; anal angle
of hindwings not rounded; aedoeagus of male genitalia more slender and oblong.

..... A. pellucida

deep yellowish brown or soft brown; when fresh with a slight grayish hue; outer
margin of both wings convex; anal angle of hindwings strongly rounded;
aedoeagus of male genitalia rounded, more narrow than in A. virginiensis but a

little wider than in A. pellucida, proximal end straight. A. discolor

medium yellowish brown; outer margin of forewing slightly concave (between
M2 and Cu lb) , of hindwing straight, anal angle of hindwing almost rectangular. . .

A. finlaysoni

light brown or soft brown; sprinkling with dark scales strong; outer margin of
foreing perceptibly convex, of hindwing straight; anal angle of hindwing

rounded; only known from Florida and Georgia A, consularis

medium reddish brown (North) or soft reddish brown (South); outer margin of
both wings almost straight; anal angle of hindwing not at all rounded; aedoeagus

of male genitalia straight A. senatoria

medium reddish brown; sprinkling with dark scales strong; hyaline patch
covered with scales to some extent; discal spot large; margin of hindwing
straight A. peigleri

4. forewing brownish orange; hindwing dark blackish brown, venation of forewing
the color of hindwing; abdomen blackish brown; Mexico only. . . A. dissimilis
male still unknown; probably somewhat similar to the above species, Mexico

only. A. punctata, n. sp.

forewing deep reddish orange; hindwing soft reddish brown; Mexico, Arizona to

western Texas. A. oslari

unicolorously gray reddish orange with a slight tinge of reddish in hindwing;

Mexico only. A. assimilis

color of both wings deep orange to brownish orange; strongly sprinkled with
dark scales; outer margin of both wings entirely rounded, especially apex of fore-
wing A. stigma

color of both wings deep orange yellow to medium yellow brown with little
contrast; strongly sprinkled with dark scales; outer margin of forewing slightly
straightened and not rounded at apex; eyes strikingly large. ...... A. fuscosa

strong medium reddish purple suffusion on both wings, basic color brownish
orange; only Manitoba to Wisconsin. A. manitobensis

5. connection (ductus bursae) between ostium bursae and bursa copulatrix

straight 6

this connection not straight. 7

6. deep orange yellow; wings semitransparent in most specimens; usually only
slightly sprinkled with dark scales; outer margin somewhat straightened

19(3): 101-180, 1980(81)

111

between Ms and apex; postmedian band usually quite visible; ductus bursae
obviously slender. .......................................... A. senatoria

deep orange yellow to brownish orange; wings very strongly sprinkled with dark
scales; outer margin of both wings well rounded. ............... A. consularis

deep orange; wings opaque; strongly sprinkled with dark scales; outer margin of

both wings well rounded. A. stigma

almost uniformly medium orange colored with a slight pinkish tinge on the

hindwing; Mexico only. A. assimilis

7. brownish orange strongly suffused with purplish, especially in basal and
terminal areas; ductus bursae only slightly bent to the left; Manitoba to

Wisconsin. .................. ................ A. manitobensis

deep orangish yellow forewing; hindwing duller, somewhat suffused with
blackish-brown; venation on both wings of same color; abdomen blackish-
brown, joints of segments outlined in ochreous brown; ductus bursae sigmoidal;

Mexico only A. dissimilis

brownish orange on forewing; medium orange hindwing; venation not pro-
nounced; abdomen tawny olive-ochreous; head and thorax same; postmedian
lines very strongly developed; strongly sprinkled with dark scales. . A. punctata
light brown with a light ochreous tinge; hindwing slightly darker; ductus bursae

sharply bent to the left; Mexico, Arizona to western Texas A. oslari

unicolorous deep orange yellow to medium yellow brown; strongly sprinkled
with dark scales; outer margin of wings as in male; ductus bursae twisted twice;

eyes strikingly large. ................. A. fuscosa

soft yellow; outer margin only slightly rounded between Cu lb and apex;
postmedian lines on both wings straight; but weak; purplish suffusion in
terminal area very reduced; ductus bursae twisted upwards. ... A. finlaysoni
deep orange yellow; mostly strongly sprinkled with dark scales, marginal area of
forewing purplish; outer margin almost straight; large (25-30 mm); ductus

bursae twisted twice. .A. peigleri

considerable contrast between gray purplish pink terminal area and remainder
of wing brownish orange or wine colored; rarely with dark scales; ductus bursae

sharply bent to the left. A. virginiensis

wings opaque and occasionally with dark scales and strongly colored medium

reddish brown; ductus bursae bent to the left directly cephalad of antrum

A. pellucida

monotonous light brown to dark orange yellow; sometimes with grayish over-
tones; often sprinkled with dark scales; bursa copulatrix very slightly bent off of

ductus bursae. A. discolor

size large; deep orange yellow; terminal area of forewing slate; ductus bursae
twisted twice; Central America. ...... A. leucostygma

Key to Mature Larvae

1.

2.

not unicolorous. 2

unicolorous 3

in two colors (light gray olive and deep pink) evenly and widely striped; thoracic
horns 6 mm long; head deep yellow. .......................... A. virginiensis

in two colors (dark greenish yellow and deep pink); thoracic horns 6 mm long;
head deep greenish yellow. .................................... A. pellucida

112

J. Res. Lepid.

in two colors (dark olive and very deep red); not quite evenly striped; thoracic
horns 8 mm long; head medium olive. ........................... A. discolor

first three abdominal segments yellow, following ones reddish brown; thoracic
horns long and blunt on tips; black spots between abdominal segments; head
yellow. ........................................................ A. dissimilis

3. unicolorous black. .......................................................... 5

other color. ................................................................ 4

4. color soft red with broken yellow stripes; body very strongly granulated;
thoracic horns 5.5 mm long; head deep orange. ..................... A. oslari

color various shades of brown, densely covered with pinkish white granulations
of uneven size; laterally paler stripes only faint; thoracic horns 7 mm long; head
brownish orange. ................................................. A. stigma

color light brown, on each segment a very wide black saddle; on saddle pinkish
white granulations of uneven size; thoracic horns 5 mm long; head deep orange
yellow. ........................................................ A. assimilis

color medium orange to light brown; body covered with many white spots and
ecrescences; dorsal, lateral, supraventral brownish stripes; gray dorsal band;
thoracic horns clubbed, 5 mm long; head brownish orange. ....... A. ftiscosa

color medium orange with lighter brown stripes; thoracic horns 6 mm long; head

pale tan, ..................................... ......... A. manitobensis

color various shades of tan, with numerous longitudinal thin black stripes;
thoracic horns ca. 6 mm long; head brownish orange. ............ A. consularis

5. with very orange yellow continuous ..stripes; thoracic horns 4 mm long; head
blackish red brown, ........................................... A, senatoria

with sometimes faint, interrupted very orange yellow stripes; subspiracular
“ornaments” much reduced; thoracic horns 6.5 mm long; very spiny; head
black. .......................................................... A. peigleri

with brilliant orange yellow continuous stripes, no thoracic horns except a pair
of short scoli 1 mm long; head black. .......................... A. finlaysoni

Note: The names of colors used in the foregoing keys follow those given in the

Inter-Society Color Council — National Bureau of Standards, Circular 553.

Ecology, Collecting in the Field, and Rearing in Captwity

Rearing and collecting the various species oiAnisota is an integral part of studying
them. These activities can be very profitable because it is generally easy to collect
larvae on trees and adults at lights. The larvae are rather easy to rear in captivity.
The senior author has studied the species in southeastern Ontario; the junior author
has collected from North Carolina to Florida across to Texas and in Mexico and
Ontario.

Searching for ova is generally unproductive unless one is in an area where high
populations exist of one or more of the species. It is then best to search on the
undersides of oak leaves not too high from the ground. The heavy females do not
oviposit in the tops of trees unless the tree is only a couple meters high. Isolated
trees, or those along a forest-field interface are good places to search. Trees near
lighted areas (such as rest areas on major highways) which attracted female moths
are more likely to have ova and larvae. Larvae begin feeding on the leaf which bore
the ova so it is advisable to look for chewed leaves on ends of branches. When larvae
only a few days old are found in this manner, the egg patch is also present nearby.
The remaining ova may contain ovarian parasites.

19(3): 101-180, 1980(81)

113

Larvae of Anisota are gregarious all through life, especially in the early instars.
Young larvae spin invisible silk mats to aid aggregating behavior. Mature larvae wiU
spread out over an entire small tree but on large trees they all are generally on one or
few branches, still relatively near each other. When a collector begins to detach
larvae which he has found, the adjacent ones detect the danger and drop to the
p-ound; even larvae in molting position will drop. Larvae are very easy to remove
from branches, never hanging on tightly as in many Satumiidae. Molting larvae
detached from their silk pads almost always molt successfully, even in the bottom of
a glass jar. Larvae sit on twigs in clusters in the position of sphingid larvae, with the
anterior end raised. Brodie (1929) described and figured this behavior. Larvae also
will sometimes sway rapidly from side to side in unision when disturbed. This
behavior is not understood. To collect the largest larvae among a colony is to collect
mostly females. It is tempting to take large ones because one figures that less
rearing work will remain to be done. Male larvae, being smaller, mature sooner and
leave the tree several days earlier than their female siblings.

Since the defoliation is usually quite noticeable, it can be productive to look for
colonies while driving along a road, but sometimes, in the Southeast at least, the
defoliators turn out to be colonies of Datana spp. (Notodontidae). First-instar
larvae of Datana skeletonize leaves, unlike Anisota. Other times the larvae of
Anisota have left the tree already to pupate. If the defoliated branch has begun to
recover by producing new leaves, one can assume the larvae left weeks earlier. If no
new leaves have yet formed, it is often possible to find a few straggling larvae, or at
least some exuviae, to determine which species had been present. We have
observed that the same individual trees, indeed certain lower branches, are
defoliated year after year by the same species of Anisota.

Larvae of certain species of the genus are not well camouflaged. The coloration of
those species such as A. senatoria (black with orange stripes) is apparently
aposematic. Immunity to avian predation may be due to the tough spiny texture of
the caterpillars and/or to possible toxic substances, such as tannins derived from
oak leaves.

It is incorrect to assume that Anisota are found on any oak. All oaks are accepted
freely in captivity; larvae will feed on a mixed diet of several oak species with no
hesitation. The tough evergreen oaks such as live oak {Quercus virginiana Miller) are
eaten freely as well as the species of Castanea, the chestnuts and chinkapins.
Possibly the nearly extinct American chestnut {Castanea dentata (Marshall)
Borkh.) was a major host in the past. Species of Quercus with tough, pubescent, or
evergreen leaves are generally the least preferred in nature. In an area with several
species of oaks, one will find some species attacked growing among others
unattacked. Ovipositing females are apparently very discriminating. In the South-
east (North Carolina to Florida to East Texas) the two favorites are undoubtedly
water oak (Q. nigra L.) and southern red oak (Q. falcata Michaux). The former is
commonly planted in yards and parks and may be heavily defoliated. Other
favorites are Q. marilandica Muenchh., Q. velutina Lam., Q. phellos L., Q. palustris
Muenchh. and similar species. Q. alba L., Q. stellata Wang,, and Q. virginiana are
not frequently attacked. However, in extreme southern Florida it is probable that Q.
virginiana is a usual host because other oaks are not available. Likewise in Texas,
west of the forested areas of the eastern part of the state, the Anisota are obliged to
live on Q. stellata, the most prevalent oak. In France, Dr. Claude Lemaire has reared

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Fig. 2. Suranal plates of mature larvae: a. A. stigma, b. A. fuscosa, c. A.
consularis, d. A. manitobensis , e. A. oslari, f. A. assimilis.

species of Anisota from the Atlantic states with a little difficulty on Q. pubescens
Willd., a species similar to Q. stellata, but found that larvae of A. disco/or from Texas
did especially well on that host. Tough and/or pubescent leaves are more offensive
to very young larvae. Reports of larvae eating hazel {Corylus) (Packard, 1905;
McGugan et ai, 1958) we believe to be correct.

It is also apparent that the hot, dry, sandy, scrub-oak areas, so suitable for
Hemileuca (Satumiidae), are not suitable biotope loi Anisota. These areas include
turkey oak (Q. laeuis Walter) and myrtle oak {Q. myrtifolia Willd.) growth from
central South Carolina down through Florida and the limestone hills with Q.

19(3): 101-180, 1980(81)

115

fusiformis Small in central Texas. Anisota are sometimes collected in such areas, but
the populations are very low, if at all present.

When larvae have finished feeding they drop to the ground and begin to wander
for a day or so in search of a place to enter the ground to pupate. Collecting of larvae
crawling on the ground is desirable because no further feeding is required by such
larvae, but some parasitism should be expected. It is virtually impossible to locate
pupae in the soil so we recommend the collector not waste his time in such an
endeavor. Nematodes may attack pupae in the soil but we have no records for this.
Pupal mortality may also be caused by occasional flooding or droughts.

The best method for collecting adult moths is to attract them with ultraviolet or
mercury vapor lamps (such as street lights). One can hang an ultraviolet light or
lantern in front of a sheet. White light sources near a brick wall or similar surface
(the lighter colored the substrate is, the better) are very suitable places to find
adults because a suitable resting place is available to the moths after arriving at the
Mght. Adults which have been at rest are veiy lethargic and thus easily captured;
they will drop into a box or killing jar if gently pushed. Females of A. peigleri arrive at
light just at dark whereas males of A. stigma may not arrive until 2:00 AM (EST).
Most adults are available under lights before midnight except, of course, those
species with diurnal males which are rarely taken at lights. The earlier in the day
that the male flies, the fewer at light: we know of only one male A. pellucida taken at
light. The rare cases where diurnal males are found at light may involve individuals
which were brought there by the larger females, the female flying to the light before
copulation had terminated. This is only a theory but often pairs are found near each
other under light sources.

Males can be attracted to females which will emit pheromone in captivity. We
believe that the pheromone is the same, or about the same, for all the species (see
our remarks under “Artificial Hybridization”). Therefore a female of any species
may attract males of any other species, provided that she emits pheromone at the
right time of day or night, during the correct season, and in the right location.
Hybridization of the species in nature is prevented by allopatry, different flight
times of day and night, and/or different flight times through the year. Adjusting of
the photoperiod with artificial light or darkness allows one to induce a female to
emit pheromone at any desired time. Assembling males are rapid fliers, and a net is
usually necessary to capture them.

Several handbooks are available to provide information on rearing caterpillars.
Consequently, we only wish to give here a few remarks peculiar to, or important in,
the rearing otAnisota. Larvae are easy to rear but tend to stunt easily when reared in
containers on cut food and the result is undersized adult moths. Best rearing is done
in large cloth bags on limbs of oaks because such growing food provides more
nutrition than cut food. Also ventilation and sunlight should reduce incidence of
disease in a brood. However, once larvae stop feeding they cannot be left in the bags
to pupate as with cocoon-spinning species. There is no evidence that a diet of
several oak species produces larger or healthier larvae. Humidity may be advantageous.

Although most lepidopterists prefer to let larvae which pupate subterraineously
burrow into a container of soil, this is not at all necessary. Contracted larvae pupate
quite successfully in any container, with no soil, or they can be placed on a layer of
moist sand or peatmoss. Summer pupae destined to produce adults in less than one
month generally do veiy well if simply kept from dessication. However, with

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J. Res. Lepid.

diapausing (over-wintering) pupae one should expect some mortality. One method
which the junior author uses is to keep pupae on a layer of moist-to-wet sand in a
plastic shoebox in the refrigerator (ca. 4-5°C) all winter. A too tight lid results in
molding; too open causes dessication; both will be fatal to the pupae. Misting of
pupae with water triggers emergences.

The diapause is apparently broken by longer day length rather than higher
temperatures. Anisota-pupae kept at room temperature (ca. 25°C) all winter still
produce adults at the normal emergence time in many cases. Pupae kept under
laboratory conditions sometimes yield adults in autumn or spring, while summer is
the normal flight time for the particular species. It is possible that the diapause is
attuned to specific needs in the phenology such as host plant requirements. Even in
the milder climates, larvae of Anisota strongly tend to be late season feeders. In
such areas there is ample time for another generation but this does not occur.
Probably larvae of the genus require mature oak leaves. Other lepidopterous larvae
require spring foilage of oak. The levels of proteins, carbohydrates, and tannins in
oak leaves change through the seasons. The excellent study by Feeny (1970) will
provide the reader with more information.

When adults emerge the rearer may wish to obtain matings or cross-matings.
Males can be kept refrigerated for several days until a female is available. The
container in the refrigerator should have a damp cloth. Such chilled males will
become active within a few minutes after being taken from refrigeration, and will
mate quickly with a female which is emitting pheromone. Females which are chilled
in this manner however, usually fail to emit pheromone afterwards. Hand-pairing
has not been successful with the small adults.

A problem is encountered if one tries to hold adult moths for long periods at room
temperature. Emergences from pupae are in morning or mid-afternoon, depending
on species and geographic location, and the moths of both sexes commence to
flutter at dusk. If specimens are needed for a collection but one is also hoping to
obtain pairings, we recommend that they be kept beside a lamp all night while alive.
Another practice used to prevent wing damage can be tried with females after they
have mated. This is to put them into envelopes with wings folded back while
ovipositing. While this method works well for large satumiids, the females of
Anisota seem to prefer to flutter some and more eggs will probably be obtained from
a female kept in an open container. The presence of oak foliage does not appear to
increase oviposition. It is likely that almost all females of Satumiidae taken at lights
have mated and will thus lay fertile ova. This usually appears to be true for females
of Anisota, but not always. Some individuals oviposit freely; others do not, even if
mated.

The rearer choosing to send out ova to enable others to rear larvae should keep in
mind the remarks of Hitchcock (1961b), i.e., first-instar larvae have higher mortality
in smaller groups. The very young larvae apparently work as a team in cutting the
oak foliage. Therefore, it is best to supply at least 30 ova to another rearer.

Population levels or densities may be estimated in an area by abundance of larval
colonies or by numbers of adults taken at lights. The two do not always appear to
“agree” however. For example, in northwestern South Carolina A. peigleri is
consistently abundant as a larva but adults are taken only occasionaOy at lights. In
the same area larvae of A. stigma are difficult to find but adults are abundant at
lights. The apparent discrepancy is explained by the fact that females are taken at

19(3): 101-180, 1980(81)

117

Fig. 3. Suranal plates of mature larvae: a. A. senatoria, b. A. peigleri, c. A.
finlaysoni, d. A. pellucida, e. A. virginiensis, f. A. discolor.

light less often than males, and males of A. peigleri are diurnal. Therefore, many
males of A. stigma are collected at lights, some females of both species, and rarely
any males of A. peigleri.

Females of some species such as A. senatoria and A. peigleri deposit high numbers
of ova in one place, and this contributes to the ability of such species to become
pests by extensive defoliation. Larval colonies of other species such as A. stigma and
A. pellucida usually contain less than 25 larvae. If fecundity of all species is roughly

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J. Res. Lepid.

Fig. 4. Head capsules of mature larvae: a. A. stigma, b. A. fuscosa, c. A.
consularis, d. A. manitobemis, e. A. oslari, f. A. assimilis.

equal, we can assume females of the latter species oviposit in more sites, but fewer
eggs per site.

In Florida and coastal Georgia, populations of A. pellucida are consistently high.
Anisota peigleri has maintained a very high population in upper South Carolina for
at least 12 years; in East Texas the same appears to hold for A. senatoria. The
sporadic outbreaks of A. senatoria in New England are a very different phenomenon.
In this case we would use the ecologists’ term “fugitive species” for A. senatoria.
Fortunately, Anisota is a group which is not especially harmed by human
interference such as habitat destruction, since the various species live well in yards
and parks where oaks are planted, as well as in the countryside. Conservation of
populations oi Anisota may become necessary however. Hessel (1976) mentions A.
finiaysoni as suffering from habitat destruction. Indeed we have noted an absence of
records for this species between Toronto and the type locality and suspect that
agiicultural practices have destroyed most oaks in that area. Many forest entomolo-
gists consider A. senatoria to be a pest such as Ignoffo et al. (1973) in Missouri and
Hitchcock (1958, 1961a) in Connecticut. Large scale sprayings with chemicals or
pathogens should be used only where absolutely necessary. The aerial application
of spores of Bacillus thuringiensis Berliner is hailed as harmless to the environment
by entomologists attempting to control populations of the gypsy moth {Lymantria
dispar (L.), Lymantriidae). That claim is absurd since the pathogen kills many non-
target species of Lepidoptera.

Collecting of material throughout the range of the genus is needed to augment
what is known about the group. Amateur lepidopterists can make significant

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119

contributions by preserving adults and associated larvae from localities where they
collect. Data are especially needed for circadian flight times of even the common
and widespread species. Some of the aforementioned field observations may only
be applicable to the southern states, creating the need for information on some
northern populations for comparison. The lack of material and information for the
Mexican species is particularly obvious. The mechanisms of diapause in Anisota are
poorly understood. It is hoped that our information in this section will stimulate
additional research by others.

Parasitiem

During the larval stages, the species oi Anisota are much more vulnerable to attack
by parasites than predators. The known parasites all belong to the orders Diptera
and H5m[ienoptera. Records for parasitism in species in the northeastern states (A.
stigma, A. senatoria, A. virginiensis) can be found in Schaffner & Griswold (1934)
and Muesebeck et al. (1951), For tachinid parasitism one can do no better than to
consult Amaud (1978). The junior author has reared parasites from Anisota in the
southern states and a list of these plus notes on their habits follows. Our records
from Maryland were provided by Robert T. Mitchell.

Diptera

Tachinidae: Goniinae

Lespesia datanarum (Townsend) (= Achaetoneura anisotae Webber). In A.
pellucida from Martin, Florida; Ludowici and Statesboro, Georgia; in A. peigleri

Fig. 5, Head capsules of mature larvae: a. A. senatoria, b. A. peigleri, c. A.
finlaysord, d, A. pellucida, e. A. virginiensis, f. A. discolor.

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J. Res. Lepid.

from Clemson, Greenville, and Westminster, South Carolina. The ovipositing flies
deposit their eggs on a larva and the maggots feed inside the host, emerging to
pupate in soil after the host has pupated. Parasite puparia will generally diapause at
the times that the host species does. Usually there is one to three parasites per host.

Eumasicera stemalis (Coquillett). In A. consularis from Statesboro, Georgia and
in A. uirginiensis from Pine Grove, Pennsylvania. These flies have a life-cycle much
like L. datanarum but the adults are lighter gray and smaller.

Belvosia hifasciata (Fabricius). In A. senatoria and A. discolor from Walker Co.,
Texas. (What is apparently this species has been reared from A. assimilis by R. O.
Kendall from Chihuahua.) These large flies have yellow-tipped abdomens and can
be seen flying around defoliated oak trees in the autumn searching for hosts. The
phenology is similar to L. datanarum but these are solitary (gregarious in larger
hosts such as Eacles) parasites and the maggot does not leave the host pupa to
pupate.

An undetermined species of Tachinidae attacks A. finlaysoni in St. Williams,
Ontario (size of Lespesia, puparium obtained only; specimen in Royal Ontario
Museum).

Hymenoptera

Trigonalidae

Poecilogonalos costalis (Cresson). In A. disco/or and A. senatoria from Walker Co.,
Texas. This insect superficially resembles a yellowjacket {Vespula maculifrons
(Buysson)). It occurs solitarily and is probably a hyperparasite on tachinid parasites
in Anisota. Trigonalids represent a small and poorly known family.
Ichneumonidae

Hyposoter fugitiuus (Say). In A. peigleri from Greenville and Clemson, South
Carolina; in A. pellucida from Anderson Co., S. C.; Baton Rouge, Louisiana and
Saint Francis Co., Arkansas; in A. discolor from Giddings, Texas; in A. senatoria
from Walker Co., Texas; in A. consularis from Statesboro, Georgia; in A. fuscosa
from Brazos and Walker counties, Texas. In A. finlaysoni from Shannonville,
Ontario (leg. L. R. Finlayson, in alcohol in Royal Ontario Museum). The species is
reported by Muesebeck et al. (1951) to attack A. stigma, A. uirginiensis, A. senatoria
and numerous other hosts in Satumiidae and other lepidopterous families. In
Anisota very young larvae are attacked. In the second or third instar the host dies
and its dried skin is fastened to a leaf or stem by the parasite larva within which it
spins its cocoon within the host (figured by Felt, 1930). In Baton Rouge we found
the incidence of parasitism to be extremely high and encountered no hyperparasitism.

The following species of Ichneumonidae are hyperparasites of H. fugitivus: Gelis
tenellus (Say) in A. peigleri from Hendersonville, North Carolina; Lymeon orbus
(Say) in A. fuscosa from Walker Co., Texas; Isdromas lycaenae (Howard) in A.
peigleri from Hendersonville, N.C. The last species closely resembles its host, H.
fugitivus, but is smaller. The material from North Carolina was given us by David
Montross, entomologist at Clemson University.

Braconidae

Apanteles anisotae Muesebeck. In A. discolor, A. fuscosa, and A. senatoria from
Walker Co., Texas and A. fuscosa from Brazos Co., Texas. These small black wasps
affect larvae in any instar. When mature, the parasitic larvae emerge and spin light
yellow cocoons on the surface of the host. Larger hosts support more parasites but
the number rarely exceeds ten.

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Chalcidae

Ceratosmicra meteori Burks. This yellow wasp is a solitary hyperparasite of H.
fugitivus in A. fuscosa from Brazos and Walker counties, Texas and in A. peigleri
from Clemson and Seneca, South Carolina and Hendersonville, North Carolina. R.
T. Mitchell has taken both this species and Spilochalcis albifrons (Walsh) as
hyperparasites of H. fugitivus in A. senatoria at Patuxent Wildlife Research Center
at Laurel, Maryland.

Eupelmidae

Anastatus reduvii (Howard) (possibly a synonym of A. semiflavidus Gahan). In A.
pellucida and A. comularis from White Springs, Florida; in A. consulam from
Statesboro, Georgia; and in A. fuscosa, A. discolor, and A. senatoria from Walker
Co., Texas. These tiny wasps are ovarian parasites; usually less than half the eggs of
an egg mass is attacked. The parasites emerge from the host eggs several days after
the larvae of Anisota eclose from the adjacent unaffected ova. This is because the
parasites must develop through a larval and pupal stage within the host egg. There
is only one parasite per host egg. Beal (1952) reported a species of Anastatus in A.
senatoria [A. peigleri ?] in North Carolina. The wasps also attack ova (the
overwintering stage) of satumiids in the genus Hemileuca (Watts & Everett, 1976).
Eulophidae

Horismenus floridanus (Ashmead). This is a solitary hyperparasite in Apanteles
anisotae from Walker Co,, Texas. It emerges from the side of the cocoon, whereas
adults of Apanteles exit from the end of their cocoon. The females of H. floridanus
oviposit in the larva of Anisota, this being clear because the hyperparasites emerged
from Apanteles which had emerged from their host and pupated in the lab
(unexposed).

Tetrastichus sp. or spp. In Anisota from Walker Co., Texas and in A. peigleri from

GreenviUe, South Carolina. The minute wasps are egg parasites much like A.
reduvii. They can be stramineous or black but this color difference may be sexual
rather than taxonomic. Hitchcock (1961b) records these parasites from A. senatoria
in Connecticut.

Perilampidae

Perilampus carolinensis Smulyan. In A. senatoria from Laurel, Maryland (R. T.
Mitchell). This larval parasite is metallic blue and about 4 mm long. Apparently is a
solitaiy endoparasite killing half-grown larvae.

Most parasites attacking Anisota are not host-specific to the genus. Many have
geographical distributions wider than the individual species of Anisota, thus
attacking the different host species in different parts of their (the parasites’) range.
Some species of Cratichneumon (Ichneumonidae) are also recorded as parasites in
Anisota (Hitchcock, 1961c; Heinrich, 1977). Voucher specimens for most of our
above records are in the USNM and other museums.

John Abbot

An early pioneer naturalist and artist of Georgia was John Abbot (1751-1840?)
who sent countless specimens and paintings of these to European museums and
collectors. Excellent biographical treatments are given by Coulboum (1973) and
Harris (1972); the latter also provides a bibliography on sources about Abbot’s life.

Before leaving England at the age of 22, Abbot had made the acquaintance of the

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entomologist Mr. Dm Dmry. After settling in Georgia in 1776 Abbot sent many life-
sized colored illustrations of insects to such persons as Dr. J. A. Boisduval of Paris
and Dr. James Edward Smith of England. Two species of Anisota were figured as
new in the book by Smith: The Rarer Lepidopterous Insects of Georgia published in
1797. Smith used some of Abbot’s drawings in his book with appropriate credit to
the artist, and this is why the authorship of many Lepidoptera (including A.
senatoria and A. pellucida) is sometimes seen in the literature as “Abbot & Smith”.

Although Abbot lived in Georgia in Burke County (a part of which later became
Screven County) and also in Bulloch County, we cannot do more than assume type
localities for taxa described and figured in Smith’s book. There is even the
possibility that some specimens Abbot used as models for his paintings originated
from areas outside of Georgia.

Other taxonomic problems created by this book include the fact that some plates
contain larvae of one species mixed with adults of related species (Riotte, 1972: 11
and Ferguson, 1978: 82), and the fact that the drawings, however elegant, must be
interpreted as diagramatic at best and should not be given the favor that
systematists would afford color photographs. Colors on the plates differ from copy
to copy which was a main reason for creation of the synonymy of A. virginiensis
sinulis, the material not agreeing with some figures of A. pellucida. Also the copy of
Smith’s book in the Carnegie Museum library has very light-colored figures of
''Phalaena stigma'" so that we were at first certain they represented A. consularis
(larva, male, and female). Later we decided they probably indeed represent A.
stigma upon viewing the differently colored plates in the copy at the CNC in Ottawa.
Comments on the correct interpretation of Abbot’s plate of “Phalaena senatoria"
are given under our text of that species. We have attempted to overcome these
taxonomic difficulties by consulting the original plates for the book in the BMNH
and designation of neotypes of the concerned taxa.

Treatment of the Species
MEXICAN GROUP
Anisota assimilis (Druce)

Dryocampa assimilis Druce, 1886: 170, pi. 15, fig. 5.

Anisota assimilis; Bouvier, 1931: 22; Schuessler, 1936: 212; Draudt, 1930: 814;
Hoffman, 1942: 245; Ferguson, 1971: 64; Lemaire, 1976: 47.

Anisota leucostygma (not Boisduval, 1872); Draudt, 1930: pi. 142f; Bouvier,
1931: 18 Ipartim); Hoffman, 1942: 245. [Error in determination.]

ADULT

Male (pL IV, fig. 11): Head, thorax, legs, abdomen reddish brown; fore- and
hindwings uniformly reddish brown, more brown on forewing, hindwing more
reddish; postmedian line only very slightly marked on both wings on upperside, a
little more pronounced on underside; white discal spot small. Underside of wings
colored as upperside but lighter. Outer margin slightly rounded in forewing and
more so in hindwing. Length of fore wing 21-23 mm (4 specimens measured).

Female (pi. IV, fig. 12): Described here for the first time. Head, thorax,
abdomen, legs, fore- and hindwings on upper and underside aU almost uniformly
lighter or darker fawn colored, with a pinkish tinge on the hindwings; less intensely
colored outside postmedian line; no dark scales on wings; postmedian line barely

19(3): 101-180, 1980(81)

123

Fig. 6. Male genitalia, with aedoeagus removed and alongside: a. A. stigma, b. A.
fuscosa, c. A. consularis, d. A. manitobensis, e. A. oslari, f. A. assimilis.

perceptible; a little more conspicuous on underside; white discal spot larger than in
mcde. Outer margins as in male. Length of forewing 30-35 mm (3 specimens
measured).

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J. Res. Lepid.

Head: Laterofrontal suture visible. Frontal protuberance transverse, not promi-
nent. Male antenna with 18 rami. Posterior aspect of head not investigated because
of rarity of specimens.

Legs: Male epiphysis slender, a little longer than half the length of the tibia,
densely covered with very short hairs, a few long ones laterally and at apex;
empodium as usual, with one moderately long seta; epiphysis in female not found.
GENITALIA

Male (fig. 6f): Uncus M-shaped, apices heavily chitinized and broadly triangular;
gnathos narrowly triangular; valves uneven interiorly, more straight than rounded,
ventral part of bifurcation pointed and strongly chitinized; dorsal part of equal
length but much wider and blunter than the ventral pointed end; free anellus heart-
shaped with broadly rounded apices; saccus very much elongated cephalad into two
spoon-shaped structures which are joined together by a bridge. Aedoeagus larger
than in other groups, curved, but not as much as in A. virginiensis although very
similar; carina as in A. virginiensis, but teeth a bit larger; vesica with one well-
developed comutus at the end and a much reduced one in the middle; cephalic
margin of proximal end angular (ROM gen. prep. 3-034).

Female (fig. 8i): Ovipositor valves of a peculiar shape, much wider at the base
than at the apex, apex not rounded but linear; lobuli vaginales high; sterigma of
medium height; antrum showing marked relation to A. virginiensis, appearing in
lateral view as an oversized oval connection between ostium bursae and bursa
copulatrix which is connected by a very short ductus bursae; ostium bursae strongly
circumscribed; ductus seminalis connected laterally in first quarter of antrum;
bursa copulatrix round; signum, if present, very small, round (ROM gen. prep. 3-
084).

EGG

Y ellow when deposited, becoming various shades of brown through development.
Length 1.7 mm, width 1.5 mm, height 1 mm.

LARVA (pi. I, fig. 7)

The following description is based on material from R. O. Kendall. Data on earlier
instars are too inadequate and fragmentary to give at the present time.

Fifth instar: Body, headcapsule, and anal plate beige brown; horns on second
thoracic segment smooth with only a few short, thin setae, only slightly clubbed, 5
mm long; on each segment a black saddle from spiracle to spiracle, the width of a
segment, sometimes so much expanded that only fragments of the beige brown
body color are visible between; on the saddle a pattern of porcelain white smaller
and larger ecrescences, the latter with a small seta at the end of each; a subdorsal,
supraspiracular and subspiracular row of white thorns with a small seta on top, in
some spcimens stronger and more or less blackened; spiracles black; an extended
black ring around the outer half of the legs or prolegs, a similar patch on segments
without prolegs, with small and medium white ecrescences, the latter ones with a
small seta at the end of each; a thin dorsal line; a light beige ventral one; the eighth
segment with one subdorso-lateral more prominent thorn on each side; on the ninth
segment a dorsal one; length 43 to 47 mm.

PUPA (pi. Vn, fig. 10)

Dark reddish brown; 25 mm long; stout; cremaster very short; evenly protruding;
similar to A. oslari but with a wider base. '• ’

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DISTRIBUTION

From the figures in Draudt (1930) (see discussion below) we have the record of
Tamaulipas. Numerous other widespread records show the species to occur from
Mexican states bordering the United States south to Chiapas. Since the latter
locality (Comitan) is only ca. 50 km from the Guatemala border, the species is
probably also present in that country.

TYPE MATERIAL

The male holotype was collected in Santa Ana, Chihuahua, Mexico by Buchan-
Hepbum according to the original description. This specimen was examined by the
junior author at the BMNH. The type locality remains unclear because at least five
places in Chihuahua are named Santa Ana.

BIOLOGY AND REMARKS

According to Draudt (1930) a paper by A. Dampf was to be published describing
the immature stages, but this paper could not be located and presumably was never
published. Although Draudt treated Dampf ’s material as A. leucostygma, the color
figures (pi. 142^ leave no doubt that the species was actually A. assimilis. The
Dampf material in Draudt’s collection was destroyed in World War II (Franz Daniel,
Zool. Samml. Bayerisch. Staates, in litt.).

Although Ferguson (1971: 64) remarked that this species is known only from the
unique male type, we found numerous specimens avaOable for study as listed under
material examined. Inadequate series are available however to clarify apparent
differences between northern and southern populations. A series from Chihuahua
reared from larvae collected on Quercus grisea by Roy and Connie Kendall agrees
well with the holotype, as does the male seen from Durango. Specimens from other
localities are larger with less intensely marked postmedian lines in the males, and
there exists more variation among the series from Chihuahua. The latter also exhibit
more gray clouding in the wings.

The nearly mature larvae collected by the Kendalls were found in late September
and adults emerged the following summer. Adults from Oaxaca were taken in
September, fi’om Chiapas in June, from San Luis Potosi in April and from Veracruz
in July. Apparently these scant data suggest the species is univoltine in the northern
part of its range and bivoltine in the southern part. The immature stages are herein
described and figured for the first time.

MATERIAL EXAMINED

MEXICO. CHIAPAS: Comitan (Lemaire Coll., Peigler Coll, ROM, all ex

Lemaire Coll.). CHIHUAHUA: Santa Ana (BMNH); near Creel, 2200 m (Kendall
Coll., Lemaire Coll, Peigler Coll, all ex Kendall Coll). DURANGO: near
Durango City (BMNH). OAXACA: no additional locality data (Lemaire Coll).
SAN LUIS POTOSI: Posada El Sol, Tamazunchale (IBM), Tamazunchale
(AMNH, UCal). VERACRUZ: Poza Rica (UCal). Examined 20+ males, 30+
females; dissected 3 males, 1 female.

Anisota dissimilis (Boisduval)

Adelocephala dissimilis Boisduval, 1872: 93.

Dryocampa dissimilis; Druce, 1886: 170, pi. 15; Kirby, 1892: 740.

Anisota dissimilis; Packard, 1905: 116; Bouvier, 1927: 273; ibid., 1931: 21;
Schuessler, 1936: 212; Draudt, 1930: 814, pi 142g; Hoffmann, 1942: 245;

126

J. Res. Lepid.

Ferguson, 1971: 64; Lemaire, 1976: 48.

Anisota suprema Hemy Edwards, 1884: 16; Schaus, 1884: 102 (larva); Druce
1897: 4 15; Packard, 1905: 115,pl. 19,figs. 7, 7a; Bouvier, 1927: 273; Schuessler,
1936: 218; s5mon3unized by Draudt, 1930: 814; type locality: Jalapa, Veracruz,
Mexico.

ADULT

Male (pi. ni, fig. 10): Head and thorax mahogany brown; abdomen blackish
brown; legs blackish; basal area of forewings and posterior tuft of abdomen
mahogany brown; terminal area with purplish overtone; venation on forewings
blackish; hindwings dark blackish brown; postmedian line blackish, well-marked on
forewings only; white discal spot medium sized; underside of all four wings alike,
with reddish mahogany basal areas, terminal area with purplish overtone; post-
median line and venation well-pronounced and blackish on all wings. Outer margin
of forewings slightly rounded, of hindwings rounded. Length of forewing 22-27 mm
(12 measured). In some specimens the hindwings are not as deeply colored as usual;
in these the venation shows up in the normal darker coloration.

Female (pi. HI, fig. 11): Much lighter colored; head and thorax ochreous brown;
abdomen blackish brown, the segments outlined in ochreous brown; legs blackish;
forewings leathery brown, or orange to yellowish brown; hindwings suffused slightly
with color of male hindwings in much diluted form or with a pinkish hue; venation on
forewings reddish brown, on hindwings blackish; postmedian Une well-marked on
all wings, blackish; white discal spot larger than in male; underside of both wings
with basal area rosy brown, terminal area like hindwings above, postmedian lines
broad and blackish. Outer margin as in male. Length of forewing 32-37 mm (23
measured).

Head: Laterofrontal suture strongly visible. Frontal protuberance transverse,
insignificant. Male antenna with 18 rami. Posterior aspect of head showing well-
developed short triangular lateral projections, apex straight.

Legs: Epiphysis found only in male, very thin, length % that of tibia, rod-shaped,
covered with extremely short hairs at apex, with some long ones marginally.

GENITALIA

Male (fig. 10a): Uncus distinctly M-shaped, straight laterally, two separate
sclerotized elevations on each arm; gnathos broad, triangular shield with straight
lateral sides cephalad; valves of the usual form with specific peculiarities, ventral
part of bifurcation a sharp, heavily chitinized thorn, dorsal one a bit longer, much
wider; free anellus a wide evenly-shaped heart crescent with very short apices.
Aedoeagus only weakly chitinized, straight and short, carina without comuti; vesica
also short but with comuti, one in middle, the other minutely serrate and at the
apex. (Gen. prep. no. 349 Michener, AMNH).

Female (fig. 10b): Ovipositor valves rounded at apex; lobuli vaginales very low,
sterigma as usual; antmm short, ending in an irregular triangular shape; ductus
bursae bent abruptly laterad at half length, then again cephalad where oblong bursa
copulatrix is attached; no signum; ductus seminalis connected at beginning of
antram, ventro-medially (unusual). (Gen. prep, no. 350 Michener, AMNH).

EGG

Unknown to us.

19(3); 101-180, 1980(81)

127

LARVA

No specimens were available for the present study, nor do any illustrations exist,
Tlie following description is summarized from Schaus (1884). Length ca. 60 mm;
head, mesothoracic segment and anal segment yellow; other segments black
dorsaUy, the metathoracic one having a yellowish subdorsal line; mesothoracic
segment also mrked with four small black spots dorsally and one laterally.
Beginning on the metathoracic segment four dorsal rows and one lateral row of short
pointed black spines occur. The two inner dorsal spines on the metathoracic
segment are, on the contraiy, long and blunt at the tips; small patches of white spots
on thoracic segments; color of first three abdominal segments yellow, the following
ones reddish brown; black spots between abdominal segments; prolegs yellowish;
abdominal segments 2, 3, and 4 yeUow ventrally, the succeeding ones having a
yellow line; “centrally and exteriorly a large black spot”. Anal segment covered with
spiny ecrescences. Young larvae differ in being greenish black dorsally and without
the patches of white spots.

PUPA (pi. Vn, fig. 11)

Two dried pupae were available for study. End of pupa gradually tapering;
cremaster long, bifurcated; bifurcation short and strong; on both sides of thorax a
patch of very heavy sclerotization with irregularly arranged pits; form of patch and
arrangement of holes seem to be specific but more pupae should be studied.
DISTRIBUTION

Aside from the records given below under material we examined, there is the
additional one of Guatemala (Bouvier, 1931: 21) and the remarks of Hoffmann
(1942): “Tierras fria y templada de Veracruz, Montanas de Hidalgo, Puebla y
Oaxaca. Sierra Volcanica Transversal. Guanajuato. Durango.”

TYPE MATERIAL

Type locality: Oaxaca, Mexico (not specified as to Oaxaca State or Oaxaca City).

We have seen the pair of cotypes from the Boisduval collection in the MNHN in
Paris. We herein designate the male as lectotype and the female paralectotype.

The holotype male of A. suprema we have seen in the AMNH.

BIOLOGY AND REMARKS

The food was reported to be oak (Schaus, 1884). The insect is widespread in
central and southern Mexico at moderate to high elevations and probably extends
into Guatemala. It is said to be abundant at Jalapa (Packard, 1905), but the junior
author failed to find it there on a brief trip through that area.

It is not possible to say much about the flight times since the few records are
widely scattered geographically. Records for Chiapas of March, May, June, and
July suggest two broods. Also it has been taken in Chimalapa in September, in
Jalapa in May, and in Zacualpan in June. The species is probably not sympatric
with its ally to the north, A. punctata.

A male from Jalapa (pi. HI, fig. 12) in the MHNM from the Roberto Mueller
collection is unusually suffused with black. It appears more normal underneath.
Good colored figures of both sexes are given by Packard (1905), Druce (1886), and
Draudt (1930).

As we stated under “Intrageneric Classification and Phylogeny” this species
appears to be the most primitive species in the genus, the geographical distribution
supporting this idea.

128

J. Res. Lepid.

The abdomen has a blackish coloring of scales but the intersegmental membranes
are bright orange. Several satumiids in the Hemileucinae (e.g., Dirphia, Molippa,
etc.) show this warning coloration of orange and dark stripes, curling the abdomen
when disturbed to expose the orange.

MATERIAL EXAMINED

MEXICO. CHIAPAS: Rancho San Ramon near Mun Ochuc (= Oxchuc)
(LACM, Lemaire Coll., Peigler Coll). DURANGO: Chilpancingo Palos Colorados
(AMNH). DISTRITO FEDERAL: Mexico City (AMNH). HIDALGO: Zacual-
pan (AMNH, ROM). OAXACA: (AMNH, ROM, MNHN), Chimalapa (Lemaire
CoU.). PUEBLA: “ManzaniUa” (AMNH), Coatepec (USNM). VERACRUZ: Jalapa
(— Xalapa) (AMNH, MHNM), Mirador (ranch of R. Mueller near Cordoba)
(MHNM), Orizaba (USNM). Examined 15 males, 27 females, used pair of
dissections of Michener (AMNH).

Anisota oslari W. Rothschild

Anisota oslari Rothschild, 1907: 432; Jordan, 1908: pi. 10: 13; Bouvier, 1931;
19; Schuessler, 1936: 212; Draudt, 1930: 814; Hoffmann, 1942: 245; Ferguson,
1971: 78; Lemaire, 1976: 47.

Anisota skinneri Biederman, 1908: 77; Barnes and McDunnough, 1910: 400
(biology); - synonymized: Barnes and McDunnough, 1917: 28; type locality:

“Arizona”.

Anisota neomexicana Brehme, 1909: 324; synonymized: Barnes and McDun-
nough, 1917: 28; type locality: Fort Wingate, New Mexico.

ADULT

Male (pi. IV, fig. 9): Head, thorax, abdomen orange -ochreous; legs wine-red;
forewings dark reddish mahogany; no hyaline patch; hindwings deep wine-red;
postmedian line very slightly marked; white discal spot variable in size; forewings
.triangular with an acute apex; underside of both wings wine-red, darker in forewing
basal area; lighter apically. Outer margin of both wings usually rather straight.
Length of forewing 23-29 mm (12 specimens measured).

Female (pi. FV, fig. 10): Head, thorax, abdomen, legs light ochreous brown;
forewings beige; hindwings light wine with mauve tinge; postmedian line broad and
slate colored; white discal spot variable in size; underside of wings beige with mauve
tinge, basal area of forewings with light wine-red shading. Outer margin of wings
rounded. Length of forewing 30-41 mm (11 specimens measured).

Head: Laterofrontal suture visible. Frontal protuberance transverse, not promi-
nent but somewhat more produced than in the other species of the group. Male
antenna with 19 rami. Posterior aspect of head shows a scalloped outline; one well
developed lateral process together with one large rounded process differentiates A.
oslari from all other species of the genus.

Legs: Epiphysis slender, half as long as tibia, densely covered with very short
hairs, laterally and at apex a few longer ones; observed only in the male. Empodium
with one tubercle bearing a medium-sized seta.

GENITALIA

Male (fig. 6a): Uncus quite broadly M-shaped, apices heavily sclerotized;
gnathos broad with rounded apex, margins somewhat sinuate; midsection of lateral
margin of valves curved, upper and lower sections straight; parts of bifurcation of

19(3): 101-180, 1980(81)

129

unequal length, dorsal part not chitinized, much wider than ventral ones; free
annellus a heart shaped crescent, Aedoeagus very short, straight, carina with a small
tooth on ventral edge; vesica with one well-developed comutus at mid-length; the
usually strong comutus at apex mdimentary, a very slight chitinization of edges of
triangular apex so that A. oslari has only one real comutus on vesica. (Gen. prep.
ROM 3-032).

Female (fig. 8h): Ovipositor valves broadly rounded, whole stmcture large,
“angular”; lobuli vaginales as in A. finlaysoni; sterigma more straight; antrum
absent; ductus bursae not chitinized, making an abmpt 90° turn laterally where
bursa connected; ductus seminalis connected in upper fifth of ductus bursae
laterally. (Gen. prep. ROM 3-038).

EGG

Smooth, flatly elliptical, pale yellow, 2 mm by 1.8 mm by 1.2 mm; hatching after
about 14 days.

LARVA (pL I, fig. 8)

The following is taken from Bames and McDunnough (1910):

First instar: Body at hatching yellowish, later greenish-grey, legs black. Length 3
mm.

Second instar: Head red, body olive brown, turning later red-brown; skin
granulated; all tubercles and spines shiny black. Homs on second thoracic segment
2.3 mm long, covered with minute bristles, apex slightly bifurcate. Spiracles black,
legs black. Length 9 mm.

Third instar; Head orange-red with fine network of darker lines, sparsely
covered with very minute setae. Body brick-red. Skin granulated with a well
developed lateral fold. Homs on second thoracic segment slightly recurved, 5.5 mm
long. Spiracles black; legs black. Length 19 mm.

Fourth instar: Head and body as in previous instar. Mesothoracic horns 8 mm.
Spiracles, prolegs black. Length 38-50 mm, presumably according to the sex.

Fifth instar: Head reddish-brown, shiny; body dark brick-red, very strongly
granulated, with broken yellow subdorsal and spiracular stripes, the latter being
chiefly confined to a yellow patch about spiracle. Homs on mesothorax only 5.5 mm.
Spiracles black; legs pale red. Length 50-65 mm. Material in alcohol from Texas
received from R. O. Kendall and from Arizona received from E. M. Brown was used
to verify this description. Headcapsule shown in fig. 4e, suranal plate in fig. 2e.
PUPA (pi. Vm, fig. 3)

Material from Texas and from Arizona was available for study. Dark brown;
posterior end gradually narrowing, margins proceeding smoothly from segment to
segment; cremaster long, bifurcated; parts of bifurcation thin and longer than in A.
dissimilis; the patches on the mesothorax have a different form and pattern of
shallow pits than in A. dissimilis.

DISTRIBUTION

The species is known from southeastern Arizona, New Mexico, west Texas and
Sonora. In fact it seems that A. oslari, as so many other Lepidoptera found in
Arizona, New Mexico and Texas, has its origin in Mexico. Ferguson (197 1) says that
Texas specimens were of a more recent date, however, in AMNH there is a female
specimen from Sunny Glen Ranch, 1500 m, Brewster Co., Texas, collected on 10
June 1926. In the same collection is also a male and a female with the label

130

J. Res. Lepid.

“Janeiro” and no other data from the Frank Johnson collection. This locality could
not be identified.

TYPE MATERIAL

Type locality: Nogales, Santa Cruz County, Arizona. Additional type data: des-
cribed from 2 females obtained by E. J. Oslar in July 1903 “from chrysalids dug
from the roots of the century or mescal plant”. Location of cotypes: British
Museum (Nat. Hist.). We hereby designate one female (dated 10 July 1903) as
lectotype and the other as paralectotype; these specimens were examined by the
junior author.

For A. skinneri a lectotype was designated by Ferguson (1971); it is the specimen
which was labelled as the male type in USNM.

For A. neomexicana a lectotype was selected with F. H. Rindge from among the
cotypes in AMNH and is hereby designated: male. New Mexico, 28 June, no year,
ex coll. J. A. Grossbeck.

BIOLOGY AND REMARKS

Ferguson (1971: 79) suggested that A. oslari might be a junior synonym of A.
assimilis but the true identity of the latter species was uncertain to him. Ample
material of A. assimilis is now available to clarify this uncertainty,

Earll M. Brown of San Diego has reared larvae of A. oslari from Arizona on
Quercus kelloggii in California. According to Biederman (1908) the food is “black
(live) oak”, but we are unsure which oak species was meant. Larvae can probably be
collected on most oaks where the moth occurs. As far as we know, our figure of the
larva is the first one to be published.

The males are diurnal, flying during midday. Although they lack transparent
patches on the forewings, they are undoubtedly effective mimics of wasps with their
rapid flight and reddish orange coloration. The females are nocturnal and attracted
to light. There is one generation annually, with adults flying in late summer or early
faO, and larvae found in late fall. Reared material may emerge in early summer.

The males show several color forms from light mauve (as in females) to dark wine
(as figured) and dark slate gray. Usually, the forewings and hindwings differ slightly
in color, in both sexes. The color forms coincide to those of A. assimilis. The male
and female figured by Ferguson (1971: pi. 5, figs. 25, 26) are very typical
representatives for the species.

MATERIAL EXAMINED

MEXICO. SONORA: not seen, reported by Hoffmann, 1942. UNITED
STATES. ARIZONA: no other data (AMNH, CalAcSci); Garces (CM); Madera
Canyon, Santa Rita Mountains (LACM); Nogales (CalAcSci, LACM); Palmerlee
(CM); Pena Blanca Lake, Santa Cruz County (BPBM, Peigler Coll., ROM); Sonoita
Creek, Santa Cruz County, S. of Patagonia (LACM); Sunnyside, Huachuca
Mountains (LACM); Texas Canyon, Chiricahua Mountains (CNC). NEW MEXICO:
no other data (AMNH, CalAcSci, FMNH, ROM); Fort Wingate (AMNH, CM, CNC,
FMNH, LACM, USNM); Frijoles Canyon (AMNH); Jenny Springs (CM); Jimez
Springs (CNC). TEXAS: Fort Davis (USNM); Musquiz Canyon, Jeff Davis
County (ROM); Nickel Creek, Guadelupe Mountains (USNM); Sunny Glen Ranch
(AMNH). Examined 58 males, 64 females, dissected 1 male and 2 females.

19(3): 10M80, 1980(81)

131

Anisota punctata Riotte & Peigler, New Species

Anisota punctata Riotte & Peigler, 1980: on this page.

ADULT

Male: unknown. It is expected that the male is nocturnal and resembles the male
of A. dissimilis somewhat in appearance, but with many small dark spots on all
wings.

Female (pi. HI, figs. 8-9): Head, thorax, abdomen below ochreous brown, legs
with a purplish hue; abdomen above with an olive hue; forewings leathery brown
with numerous large spots of dark scales; hindwings purplish pink with some dark
scales and a slight marginal border of leathery brown; postmedian line straight, very
well perceptible on both wings, dark mauve; white discal spot very large; underside
of wings rosy brown with strong sprinkling of black spots, especially apically on
forewing and entire hindwings. Outer margin of forewings straight, of hindwings
only slightly rounded. Length of forewing 37 mm. Specimen not dissected because
it is unique.

EGG, LARVA, PUPA

Unknown.

DISTRIBUTION

Known only from the type locality but probably occurring in other northern
Mexican states at appropriate elevations in the same mountain range.

TYPE MATERIAL

Type locality: 27 km west of Linares, Nuevo Leon, Mexico. Known only from the
single holotype female collected at this locality on 23 July 1976 at 2 100 hrs by R. S.
Peigler. The holotype is deposited in the AMNH in New York City.

BIOLOGY AND REMARKS

The holotype was collected on a trip into Mexico sponsored by the Entomology
Department of Texas A&M University, of which the junior author was a member.
On 23 July 1976 an ultraviolet light in conjunction with a Coleman lantern was run
on a sheet and the moth arrived at this light source two hours after sunset. No other
specimens of Anisota were collected on the entire trip. The female was kept aMve in
a jar with oak leaves but died the following evening, unfortunately without
depositing any ova.

The following summer the junior author returned to the same locality and ran the
same lights for three consecutive nights (22-24 July 1977) but no additional
specimens were obtained. That season had been unusually dry; a stream alongside
the collecting site was dry, whereas it had been flowing in July 1976. Several species
of Satumiidae and Sphingidae taken there in 1976 were not seen in 1977. The
habitat of the type locality is forested. It lies in the foothills of the Sierra Madre
Oriental. Pressed specimens of the oaks growing at the type locality were sent to
John M. Tucker, Department of Botany, University of California, Davis, California.
Dr. Tucker, an authority on New World oaks, made the following determinations:

Note Added in press: The LACM has a female of this new species, apparently recently donated,
which is said to agree entirely with our figures of the holotype. The data for the specimen are:
MEXICO, San Luis Potosi, Ei Narzanjo, 4 August 1975, Terry W. Taylor. There is not time to
borrow the specimen and designate it a paratype before publication.

132

J. Res. Lepid.

Quercus canbyi TreL, Q. fusiformis Small, Q. polymorpha Schl. & Cham. The first
species has a high density in the area and with its large, tender, non-pubescent
leaves, may be the primary host of A. punctata.

PELLUCIDA GROUP

Anisota discolor Ferguson New Status

Anisota virginiensis discolor Ferguson, 1971: 83; Lemaire, 1976: 47.

ADULT

Male (pi. IV, fig. 6): Sexual dimorphism strongly developed. Head, abdomen,
fore- and hind wings above sepia brown with a very slight purplish hue apically on
the forewings; head, abdomen, legs, fore- and hindwings beneath predominantly of
a greenish brass color, however, on the forewings apically and on the hindwings
along the postmedian line and from wingbase to tomus purplish shades, on the
forewings the veins are conspicuously overlaid by the greenish brass color;
sprinkling with dark scales not observed in this species of the group; postmedian
line on the forewings well, on the hindwings only slightly perceptible; hyaline patch
not very extended; white discal spot medium sized; outer margin of forewings
rounded, of hindwings almost straight. Length of forewing 18 mm (10 specimens
measured).

Female (pi. IV, fig. 5): Head, abdomen, fore- and hindwings above almost
unicolorous fawn with only very slightly purplish apically (fresh specimens tend to
have a greenish shade over the entire wings); head, abdomen and legs beneath
orange; fore- and hindwings beneath basically light purplish, strongly overlaid with
greenish, ochreous orange; postmedian lines on both wings, above and beneath, well
marked, purplish; white discal spot medium sized; only very few and very small dark
scales on the forewings above costaUy; outer margin of forewings slightly, of
hindwings strongly rounded. Length of forewing 27 mm (10 specimens measured).

Head: Laterofrontal suture very conspicuous, wide; frontal protuberance well
developed and protruding. Male antenna with 14 to 15 rami. Posterior aspect of
head pentagonal, laterally one conspicuous, round process.

Legs: Epiphysis dorsally strongly convex, ventrally straight, ending in point,
covered with very short bristles, length little more than half of tibia. Epiphysis may
be different in one and the same specimen on left and right leg, dorsally and
ventrally straight and of even width, ending in blunt ending, length a little less than
half of tibia; in some specimens epiphysis may be rudimentary, very thin, hardly half
length of tibia; not observed in female; empodium one tubercle with slender thin
seta; at end of anterior tibia spines normally developed, i.e., one strong one and one
lesser.

GENITALIA

Male (fig. 7f): Squarish; uncus wide M; gnathos sometimes with waving sides.
Mostly a very broad round shield; bifurcations of equal length; interior angle of
valve pointedly elongated; free annellus long, round tube with somewhat heart-
formed opening; aedoeagus rounded, more narrow than in A. virginiensis but a little
wider than in A. pellucida; proximal end straight; carina prolongation of main body
of aedoeagus; row of small teeth on carina; vesica with very prominent apical
cornutus with teethed edge; there was no small, rudimentary second sclerite in 6
dissected males from Texas, however, such sclerite was often found in specimens of

19(3): 10M80, 1980(81)

133

A. virginiemis from Pennsylvania. (Gen. prep. ROM 3-164).

Female (fig. 9d): Ovipositor valves tapering off apically, of unusual form; lobuli
postvagmales high, antevaginales very low; antrum like strong triangular funnel,
connecting ostium bursae and strong, short, slightly oblique ductus bursae which
has in its first third laterally strongly sclerotized ecrescenses; ductus seminalis
attached at beginning of ductus bursae posterior-laterally; bursa copulatrix very
slightly bent off ductus bursae, rounded oval, no signum, covered with fine network.
(Gen. prep. ROM 3-061).

EGG (The foUowing descriptions of egg and larvae—first to fourth instar— by C.
Lemaire).

Yellow changing into orange and brown.

LARVA (pL I. fig. 12)

First instar: Body color yellowish green, slightly blackish; very near to second
instar of A. senatona; headcapsule and thoracic legs black.

Second instar: Body color dingy greeiysh-yeUowish; headcapsule and thoracic
legs black.

Third instar: Body color dark olive brown; headcapsule beige-brown; thoracic

legs black; prolegs olive; subdorsal and supraspiracular lines rose brown; two very
slight infraspiracular whitish lines.

Fourth instar: Body color blackish brown; headcapsule beige-brown; thoracic
legs greenish with black apex; prolegs olive black; white granulation not dense over
body; subdorsal and infraspiracular lines purplish brown.

Fifth instar: Body color beige-brown; headcapsule, anal plate, thoracic legs of
same color; white granulation not too dense over body; dorsal line purplish brown;
of same color also wider subdorsal, supraspiracular and subspiracular bands; a
spiracular waved line of same color; spiracles black with no border; horns on
thoracic segment II clubbed, 8 mm long; length of fifth instar larva 42 mm.
Headcapsule shown in fig, 5f, suranai plate in fig. 3f.

PUPA (pL vm, fig. 2)

Blackish brown, similar to that of A. virginiemis but the bifurcation perceptibly
straighter and longer, even more so than in A. pellucida.

DISTRIBUTION

This species was recently described from a series of specimens from several
localities in eastern Texas. We now have records from Oklahoma (from P. Loy) and
Louisiana (from R. 0, Kendall); the species is probably only present in the western
parts of Louisiana since we found it was replaced by A. pellucida at Choudrant and
Baton Rouge. It is recorded in Texas as far northwest as Hamilton and as far
southwest as Giddings. It is commoner than either A. fuscosa or A. senatoria in the
grassland prairies between the Edwards Plateau and the forested areas of eastern
Texas.

TYPE MATERIAL

Type locality: Spring, Harris County, Texas. Location of type: United States
National Museum, type no. 71496. Additional type data: female holotype, labelled
Spring, Harris County, Texas, 27 August 1963, A. and M. E. Blanchard.
BIOLOGY AND REMARKS

There are two generations per year but the main one is the later one. Many
overwintering pupae probably do not produce adults until the flight time of the

134

J. Res. Lepid.

second brood. These flight times occur in late June to July and late August through
aU of September. Larvae can be coUected easily in eastern Texas from late
September to early November, some years more commonly found.

The male fUes at midday. Their coloration is a smoky brown somewhat as males of
A. finlaysoni but with more robust and thickly scaled wings. Some with the wine red
coloring of A. pellucida occur occasionally. One male was attracted to a calling
female in College Station, Texas, on 31 August 1978 at 1240 hrs (CDT). A diurnal
sphingid {Hemaris sp.) also appeared but an attempt to capture it failed. This
attraction was not surprising because Dominick (1974) had similar experiences with
males of the sphingid Amphion nessus (Cramer) coming to females of A. pellucida in
South Carolina.

In the females there is much more color variation than in either A. pellucida or A.
virginkmis. Soine are reddish or orangish as in those two species but most are
“coffee-with-cream” colored. The postmedian area may be very purplish or
scarcely contrasting from the median area.

Records for foodplants include Quercus nigra in Giddings, Q. macrocarpa in
Hamilton, Q. stellata most often in College Station, and in eastern Texas usually the
same oaks selected in the Atlantic States by other Anisota, namely Q. velutina, Q.
nigra, and Q. falcata but also Q. marilandica. Although Q. virginiana grows
commonly in College Station, no larvae of Anisota have been found on it by the
junior author in three seasons.

MATERIAL EXAMINED

UNITED STATES. LOUISIANA: no other data (CalAcSci); Vernon Parish
(Kendall Coll). OKLAHOMA: Claremore, Rogers Co. (AMNH), TEXAS: no
other data (AMNH); Anderson Co. (Larva) (TAMU); Beaumont (LACM, TAMU);
College Station (AMNH, BPBM, LACM, Lemaire Coll., Peigler ColL, ROM,
TAMU, USNM); Giddings (LACM); Hamilton (AMNH); Houston (USNM); Hunts-
ville State Park (LACM); Kamack (MSU); New Waverly (USNM); San Jacinto Co.
(LACM); Spring (USNM); Stubblefield Lake in Walker Co. (AMNH, BMNH,
BPBM, CNC, Peigler Coll., ROM); Town Bluff (USNM). Examined long series of
males and females, dissected 5 males and 4 females.

Anisota leucostygma Boisduval

Adelocephala leucostygma Boisduval, 1872: 85.

Adelocephala leucostigma; Druce, 1886: 171.

Anisota kucostygma; Bouvier, 1927: 273; ibid., 1931: 18; Draudt, 1930: 813;
Hoffmann, 1942: 245; Ferguson, 1971: 64; Lemaire, 1976: 47.

ADULT

Male: unknown. It is expected to be diurnal.

Female (pi. IV, fig. 8): Head and thorax reddish orange; legs and abdomen lighter
orange; forewings reddish orange interiorly of postmedian line, golden brown with a
very light hue of slate color exteriorly of postmedian line; this line very well
developed on both wings; white discal spot large (as name implies); hindwings
reddish orange; only sprinkled slightly with dark scales on apical area of forewings;
underside of wings reddish orange. Outer margin of both wings evenly rounded.
Length of forewing 3 1 mm.

GENITALIA

Female (fig. 8g): Ovipositor valves not too large; sterigma very wide; lamellae

19(3): 101480, 1980(81)

135

postvaginaies not too Hgh; lamellae antevaginales laterally strongly expressed, not
too higti; ostium bursae not very wide, reaching into ductus bursae like hook (see
illustration); antram not too pronounced; ductus bursae caudaliy becoming very
wide, narrowing considerably cephalad before bursa copuiatiix wdiich is round and
attached without turn; signnm small, round; surface of bursa copuiatiix with net-
Hke pattern; apophyses very strongly developed; ductus seminaiis attaching at
dorsal side of antram. (Gen. prep. ROM 3-082, now in Paiis Museum),

EGG, LARVA, PUPA

Unknown.

DISTRIBUTION

Boisduval (1872: 86) says: “Nous avons recu cette espece de Guatemala et
d'Oaxaca”. The specimen described above is the only one extent. It has no locality
label, except BoisduvaFs first identification label stating only “Mexique”.

TYPE MATERIAL

The one extant specimen passed through the collections of Charles Oberthuer
and Philipon before becoming permanent property of the MNHN in Paris. Bouvier
(1931: 18) declares: “Dtepres Charles Oberthuer, la femelle de la collection
Philipon est le type meine de leucostygma; elle porte Fetiquette suivante ecrite de la
main de Boisduval: '’Adelocephala sfygrna-, Mexique.” Dans sa description, en
effet, Boisduval rapproche son espece de stigma.” What Bouvier did we would call
today the “designation of a lectotype”, and with this the case may rest.

The labels on the specimen read: 1) handwriting ol Boisduval: Adelocephala
Stygma B Mexique; 2) handwriting of Oberthuei: Adelocephala Leucostygma,
Boisdiiv. (AimaL Soc. Entom. Belgique, 1871-72, pages 85, 86, femelle, typiciim
specimen Boisduvalianum; 3) handwriting of Bouvier. Anisota leucostygma Boisd.
femelle type + printed: E. L. Bouvier + handwritten: ver.(ifiedj.

REMARKS

If there were not the unmistakable assertion of Oberthuer and the declaration of
Bouvier for the identity of A. leucostygma, and if the genitalia of the unique
specimen were not so impressively different from those of the three other species
belonging to the same group, one could perhaps understand that entomologists are
tempted to consider A. leucostygma as a mislabeUed specimen from somewhere in
the southern United States. How^eveiv one has to be very careful with unique
specimeng, A. punctata is a unique female from Mexico, too, and A. assimilis was
until very recently a unique male from Mexico in the British Museum (N.H.). In the
lattei case various negative suppositions had been aired but finally the species was
abundantly relocated in Chihuahua. Tins may very well happen to „4. leucostygma.
One should also considei the fact that the specimen has on the original Boisduval
label as locality only “Ivfexique”, and that at the time when the specimen came into
the possession of Boisduval, about the middle of the 19th century, Mexico was still
commonly understood by Euiopeans to extend much farther to the north than now
and to covw aU of Spanish speaking North America. We might disregai d the locality
designations in BoisduvaFs original description: Guatemala and O’axaca. Also
here political boundaries in those days weie other than they are now. The southern
states of today Mexico did belong to Guatemala. As fai' as then Oaxaca is concerned
Boisduval may have been influenced by the locality from wRere he received Ms
Adelocephala dissimUis (1871: 93), but one should not doubt his handwTitteii label

136

J. Res. Lepid.

“Mexique” (sensu lato). One should here consider other cases of “unbelievable”
species, like the Hawaiian hawkmoth Tinostoma smaragditis (Meyrick) which,
because it just looked so strange, was green like Eumorpha labruscae from Brazil,
was by later workers suspected not to be Hawaiian at all but was thought to have
been sent in the mail from Brazil to Hawaii and then given to Perkins who, however,
expressly says that he received the specimen from persons who had found it in their
house on Kaua’i. Today, also very recently, several specimens of this indigenous
insect are known and in collections. Therefore we should have some trust into the
solid, early workers, and we can only hope that much better and subsequent
collecting in Mexico will finally also turn up again A. leucostygma.

Specimens of A. assimilis reared from Tamaulipas by Dampf were assumed by
Draudt ( 1 930) to be A. leucostygma but his figures of Dampf’s material (destroyed in
war action) clearly show A. assimilis. Bouvier (1931) and Hoffmann (1942) therefore
erroneously reported A. leucostygma as being from Tamaulipas.

In some of the literature it can be found that Boisduval considered his A.
leucostygma to be systematically close to A. stigma. Maybe so, as his handwritten
label suggests. However, his original description of A. leucostygma shows that he
then changed his mind in the right direction.

Druce (1886) reports only briefly about the species which he has not seen from
Mexico and Guatemala. Bouvier (1927) then does not mention “Guatemala”
anymore.

MATERIAL EXAMINED

The one extant specimen of A. leucostygma was loaned to the senior author by P.
E. C. Viette. It was then dissected and the genitalia show it to be very much distinct.

Anisota pellucida (J. E. Smith)

Phalaenapellucida J.E. Smith, 11 91: H: 115, pi. 58; - s3monymized: Westwood,

1837.

Dryocampa virginiemis; Westwood, 1837: 24, pL 13: 2.

Dryocampa pellucida; Harris, 1841: 293.

Anisota pellucida; Grote, 1864: 93; Packard, 1864: 385; Lemaire, 1976: 47.

Adelocephala pellucida; Boisduval, 1872: 87.

Anisota virginiensis; Packard, 1905: 103.

Anisota virginiensis sinulis Riotte, 1970: 89; - synonymized: Ferguson, 1971;
type locality: Gainesville, Florida.

Anisota virginiensis pellucida; Ferguson, 1971: 82.

ADULT

Male (pi. IV, fig. 3): Sexual dimorphism strongly developed. Head, abdomen
above, fore- and hindwings above very dark brownish wine-red; legs, abdomen
below dark orange; in many specimens apically sprinkled with dark scales;
postmedian line well perceptible on both wings, also on underside; white discal spot
small; an ochreous-brown hue can be found on hindwings, thorax and dorsal part of
abdomen, it is, however, very often very faint. Underside of hindwings very dark,
mostly only at anal angle of hindwings and at costal margin of forewings dark
orange-ochreous suffused. Outer margin of both wings in most specimens straight,
in some even showing some convexity; anal angle of hindwings only very slightly
rounded. Length of forewing 14-16 mm (25 specimens measured).

Female (pL IV, fig. 4): Densely scaled; interior field dark orange brown; exterior

19(3): 101-180, 1980(81)

137

field mauve; head, thorax, abdomen dark orange brown; apically sprinkled with
dark scales in many specimens; postmedian line strongly expressed; white discal
spot not too large. Outer margin of both wings strongly rounded. Length of forewing
21-28 mm (25 specimens measured).

Head: Laterofrontal suture visible. Frontal protuberance reduced. Male anten-
na with 12 rami. Posterior aspect of head a smooth ellipsoid without lateral
processes and in this quite distinct from other Anisota species.

Legs: Epiphysis observed only in males; light colored; slightly less than of
length of tibia; convex dorsally; straight laterally; ending in a point; only slightly
covered with bristles. Empodium one small tubercle with one thin small seta. At end
of anterior tibia only one strong apical spine, no second one.

GENITALIA

Male (fig. 5d): Uncus W-shaped, wide, apices strongly sclerotized; gnathos a
narrow, oblong shield, reminding of a sugar loaf; valves from a very broad base
narrowing into unusual long processes; outer margin of valves in the middle strongly
bent which renders them hexagonal; interior angle of valve more or less pointed, in
some specimens even elongated into a veritable thomlike process. Aedoeagus
rounded, short, carina long and wide; at apex with a row of small teeth; vesica with
one apical comutus; proximal end narrow and pointed. (Gen. prep. ROM 3-099).

Female (fig. 9c): Ovipositor valves not too large, rounded at apex, in dorsal view
broader at base, in lateral view very angular, with blackish pigmentation; sterigma
wide; lamella postvaginalis low, lamella antevaginalis almost only like margin of
ostium bursae; this very wide posteriorly, then narrowing into ductus bursae like
funnel; antrum not too pronounced; ductus bursae only slightly sclerotized,
cephalad of antrum straight to the left ending in oblong bursa copulatrix; no signum
but surface of bursa copulatrix with fine net like pattern; apophyses long and
relatively strong; ductus seminalis attaching on dorsal side of antrum. (Gen. prep.
ROM 3-053).

EGG

Orange; 1.1 x 1.4 x 0.8 mm.

LARVA (pi. 1. vig. 11)

First instar: Body color beige-brown; dark brown broad dorsal stripe; fine
spiracular dark brown lines; head blackish brown; thoracic horns smooth, finely
bifurcated, black; inferiorly to lateral lines a yellowish patch around the scolus on
each segment. Length of first instar larva: 5 mm.

Second instar: Body color olivish- brown; darker brown broad dorsal stripe;
spiracular lines dark brown; supra- and subspiracular lines yellowish, fine; horns
with thorns, finely bifurcated; head very dark reddish cherry brown. Length of
second instar larvae: 7 mm.

Third instar: Body color dark olive brownish; dorsally darker; spiracular space
bordered on both sides by yellowish-whitish broken fine lines; subspiracularly
broad outstanding vermillion line (broken at segments); additional little white
tubercles (also on suranal plate) everywhere; horns long, with secondary thorns,
ends shortly bifurcated; on dorsum in middle thin darker olive line and subdorsally
on each segment thin broken vermillion lines; head as before. Length of third instar
larva: 12 mm.

Fourth instar: No change, except head now light brownish-beige. Length of
fourth instar larva: 20 mm.

138

J. Ifes. LepM.

PLATE n. 1. Anisota stigma Clemson, South Carolina, reared ex ovo 21 July
1975 by J. C. E. Riotte (ROM). 2. A. stigma 9, Clemson, South
Carolina, 7 July 1975, at light, R. S. Peigler and J. W, McCord
^eigler Coll). 3. A. fuscosa d", Leeeville, Louisiana, ex larva 4
August 1960, R. O. & C. A. Kendall (ROM). 4, A. fuscosa 9, same
data as fig. 3 except date: 15 August 1960. 5. A. comuhm d, St.
Johns County, Florida, reared ex ovo 16 August 1974 by J. C. E.
Riotte (ROM). 6. A. consularis 9, Statesboro, Georgia, ex larva on
QuercusfalcataSJuly 1976, R. S.Peiglerand J. W. McCord (ROM).
7 A. rrmnitohensis d, Pembina Valley, Manitoba, ex larva, 17 April
1953, F.LS. (ROM). 8. A. manitohemh 9, Waushara County,
Wisconsin, 3 July 1936, C. Harrington (ROMexSieker CoU.). 9-10.
A. peigleri, d X consularis 9, hybrid pair, Greenville, South Carolina/
Statesboro, Georgia, 22 July 1977 and 17 September 1976, R. S.
Peigler (ROM). 11-12. A. virginiemis d X discolor 9, hybrid pair,
Pine Grove, Pennsylvania/Hamilton, Texas, 28 June and 20 July
1977, R. S. Peigler (Peigler Coll).

PLATE 1. Mature larvae of Anisota. l.A. stigma, Greenville, South Carolina.

2. A. fuscosa, Walker County, Texas. 3. A. consularis, St. Johns
County, Florida. 4. A. senatoria, southeastern Michigan, 5. A.
finlaysoni, Shannonville, Ontario. 6. A. peigleri, Clemson, South
Carolina. 7. A. assimiUs, Creel, Chihuahua. 8. A. oslari, Arizona. 9.
A. peigleri d‘ X consularis 9, hybrid. 10. A, virginiemis, Pine Grove,
Pennsylvania. 11. A. pellucida, McClellanville, South Carolina. 12.
A. discolor, Walker County, Texas.

19(3): 10M8O, 1980(81)

139

N.B. : Plates I and 11 were inadvertantly transposed in the original color printing,
and although out of order, they are properly numbered and labelled.

140

J. Res. Lepid.

Fifth instar: No change, except for kind of silvery shine because of the many
white tubercles aU over body; lateral subdorsal lines a little thicker, vermillion;
horns long (7-8 mm), black, somewhat roundly bent, without remarkable secondary
thorns, ends no longer bifurcated; spiracular space not bordered anymore by
anything. Length of fith instar larva: 35 mm. Headcapsule shown in fig. 5d, suranal
plate in fig. 3d.

PUPA (pi. Vn, fig. 7)

Dark brown; not too spiny; cremaster heavy set, originating from broad base, short
and wide, bifurcation long, almost entirely straight, not too pointed.
DISTRIBUTION

This species ranges from coastal North Carolina deep into the Florida peninsula
and across to Louisiana where it becomes parapatric with A. discolor. It is very
common in coastal South Carolina through Florida, much less common in the
piedmont of the former state. The presumed range into Tennessee, Arkansas and
North Carolina where it probably contacts A. virginiensis needs elucidation.
TYPE MATERIAL

The name is based on Abbot’s painting in J. E. Smith (1797: pi. 58). As this is
only a vague basis for a taxonomic entity in the case concerned because of the
extreme variation in the rendition of the plates in the various existing copies of the
work in question, it was considered right to designate a neotype. As such was chosen
the male holotype of Anisota virginiensis sinulis Riotte and with this the type locality
fixed as Gainesville, Florida. Additional type data: taken 22 September 1955 by C.
N. Patton in Gainesville, Florida. Location of neotype: Florida State Collection of
Arthropods, Division of Plant Industry, Florida Department of Agriculture,
Gainesville, Florida. In the said collection is also the female allotype of Anisota
virginiensis sinulis Riotte, taken 21 September 1964 by J. W. Perry in Gainesville,
Florida.

BIOLOGY AND REMARKS

The junior author has found larvae on Quercus nigra in South Carolina, Georgia,
Florida and Louisiana. That oak, along with Q. falcata, are favorite hosts throughout
much of the range. D. Baggett collected larvae in northeastern Florida on Q.
myrtifolia and Q. imbricaria. J. D. Solomon sent us larvae in alcohol from coastal
Mississippi collected on Q. nuttallu and Q. lyrata. Larval colonies are small,
averaging only about ten individuals by the time the last instar is reached, because
females deposit fewer ova at each site than do those of other species groups.

The moth is double-brooded throughout its range but the flight times vary
considerably with latitude. Records range from 1 1 April at Punta Gorda, Florida, to
3 October at Oneco, Florida, so there might possibly even be a third brood at these
lowest latitudes. During the summer the pupal stage lasts only about two weeks.
Dominick & Edwards (1971) and Solomon (1971) reported on the circadian sexual
activity of the moths.

It should be remarked that the larva shown in J. E. Smith (1797: pi. 58) on Q.
marilandica definitely represents the present species, although the adults are not
well recognizable.

MATERIAL EXAMINED

UNITED STATES. FLORIDA: no other data (AMHN, CNZ); Eglin (Hilton
Coll.; Ft. Meade (FMNH); Gainesville (FSCA, LM, ROM); Indian River (AMNH);

19(3): 101-180, 1980(81)

141

Fig . 7 . Male genitalia, with aedoeagus removed and alongside: a. A. senatoria, b .

A. peigleri, c. A. finlaysoni, d. A. pellucida, e. A. virginiensis, f. A. discolor.

Jupiter (AND^H); Largo (CNC); 5 km S. of Leesburg (FMNH); Oneco (AMNH);
Orlando (AMNH); Parish (LM); Punta Gorda (FMNH); Swannee River State Pai:k
(LM); White Springs (BPBM). GEORGIA: Athens (CLU); Clarke Co. (CLtJ);
Emory University (FSCA); Fish Eating Creek (FSCA); Statesboro (BPBM, LACM,

142

J. Res. Lepid.

PLATE in. 1, A. peiglen Greenville, South Carolina, attracted 4:30 PM EDT
on 7 July 1976 to a female of A. comularis, R. S. Peigler (Peigler
Coll.). 2. A. peiglen 9, Clemson, South Carolina, reared ex ovo 12
July 1975 by J. C. E. Riotte (ROM). 3. A. peiglen 9, Clayton,
Georgia, ex larva 18 May 1975, on Quercus velutina, R. S. Peigler
and J. W. McCord. 4. A. senatoria cf , Neotype, Gibsland, Bienville
Parish, Louisiana, 21 August 1976, at light at Interstate 20 rest
area, R. S. Peigler (AMNH). 5. A. senatoria 9, same data as fig. 4
peigler Coll). 6-7. A. finlaysoni 9, Kingston, Ontario, 6 and 2
May 1970, J. C. E. Riotte (ROM). 8 -9. A. punctata 9, upperside and
underside, holotype, 27 km W Linares, Nuevo Leon, Mexico, 23
July 1976, at light, R. S. Peigler (AMH), 10. A. dissimilis 9",
Zacualpan, Mexico, June 1915 (ROM ex AMNH). 11. A. dissimilis
9, Oaxaca, Mexico, no date (ROM ex AMNH). 12. A. dissimilis
Jalapa, Veracruz, May (MHNN), an unusually melanic male.

19(3): 101-180, 1980(81)

143

PLATE rV. 1. A, virgmiensis cf , Chaff eys Locks, Ontario, reared ex ovo 6 August
1970 by J. C. E. Riotte (ROM). 2. A. virginiensis 9, Rondeau
Provincial Park, Ontario, 22 June 1965 (ROM), 3. A. pellucida cf,
holotype of A. virginiensis sinulis, neotype of A. pellucida, Gaines-
ville, Florida, 22 September 1955, C. N. Patton (FSCA). 4. A.
pellucida 9, McClelianville, South Carolina, reared ex ovo 1
September 1972 by J. C. E. Riotte (ROM). 5-6. A. discolor 9 cf, both
College Station, Texas, reared 1976-1977 on Quercus pubescens in
France by C. Lemaire (Lemaire Coll.), female in natural resting
position. 7. A. discolor 9, Claremore, Rogers County, Oklahoma, 19
August 1977, Peter Loy (AMNH). 8. A. leucostygma 9, lectotype,
Oaxaca, Mexico no date (MNHN). 9. A. oslari cf, Pena Blanca Lake,
Santa Cruz County, Arizona, reared on Quercus kelloggii by Earll M.
Brown and emerged 23 June 1977 (Peigler Coll.). 10. A. oslari 9,
Jeff Davis County, Texas, ex larva 13 August 1968, R. 0. & C. A.
Kendall (ROM). 11. A. assimilis d‘ near Creel, Chihuahua,
Mexico, ca. 2134 m, ex larva on Quercus grisea 26 May and 20 June
1979, R. O. & C. A. Kendall (Kendall Coll.). 12. A. assimilis 9,
Comitan, Chiapas, Mexico, June 1963 (ROM ex Lemaire Coll.).

144

J. Res. Lepid.

ROM); Stonehenge (CLU). LOUISIANA: no other data (CalAcSci); Baton Rouge
(AMNH, LSU, ROM); Edgard (ACM); Harahan (CMZ); L. Charles (AMNH);
Pointe Coupee Parish (LSU); Prairieville (BPBM, DPI, LACM, LSU); Sorrento
(AMNH); Sunshine (AMNH). MISSISSIPPI: Aditon (ROM); Bovina (AMNH,
BMNH, LACM); Clinton (AMNH); Pearl (ROM); Tupelo (LACM); Vicksburg
(AMNH, LACM). NORTH CAROLINA: Carteret Co. (AMNH); Fontana (CM);
Whiteville (Peigler Coll.). SOUTH CAROLINA: Clemson (LACM, Peigler Coll.,
ROM); Greenville (Peigler Coll.); McClellanville (ROM); Manning (AMNH, LACM,
Lemaire Coll., ROM); Orangeburg (LACM, ROM); Pendleton (BMNH); Six Mile
(Lemaire Coll.); Summerville (FSCA). Examined 30 males and 40 females,
dissected 7 males and 7 females.

Anisota virginiensis (Drury)

Bombyx virginiensis Drury, 1793: 23, pi. 13: 2.

Dryocampa virginiensis; Westwood 1837: 24, pi. 13: 2; Walker, 1855: 1496.

Anisota virginiensis; Grote, 1874: 261; Kirby, 1892: 739; Neumoegen and Dyar,
1895: 148; Dyar, 1896: 166; Beutenmueller, 1898: 440; Holland, 1903: 95, pi.
8: 9, 10; Packard, 1905: 102, pi. 20: 1-3 (adults), pi. 4: 1-5, pi. 5: l-5,pL29: 1-
Id, pi. 50: 1-ld, pi. 52: 1-lc (larvae); Forbes, 1923: 668; Bouvier, 1931: 19;
Draudt, 1930: 814, pi. 142g; Schuessler, 1936: 218-220; Lemaire, 1976: 47.

Bombyx astynome Olivier, 1790: 43; synonymized: Westwood, 1837: 24; type
locality: Carolina and Virginia; fixed by Ferguson, 1971: 80, as Virginia.

Phalaena (Bombyx) perspicellina Martyn, 1797: pi. 30: 88. Type locality un-
known. New synonymy.

Dryocampa pellucida, Morris, 1862: 232.

ADULT

Male (pi. rV, fig. 1): Sexual dimorphism strongly developed. Head, thorax,
abdomen brownish wine-red; the abdomen in some specimens with a tendency to be
ochreous; legs ochreous; forewings brownish wine-red, hindwings somewhat more
on the brownish side; postmedian line not very much accentuated; white discal spot
large; underside of wings with an ochreous hue especially on the hindwings; dark
scales very small and scarce, some may be detected almost always when examined
under magnification; hyaline patch on forewing strongly developed. Outer margin of
forewings straight making the wing triangular; of hindwings in most specimens
rounded, however, all transitions to absolutely straight ones may be found in one
and the same population, e.g.. Pine Grove, PA. Length of forewing 18-20 mm (20
specimens measured).

Female (pi. IV, fig. 2): Much larger than male and not very thickly scaled; wings
distinctly rounded; terminal area of wings greyish-purple, remainder brownish-
ochreous; contrast may be strong between the two areas, however, in many
specimens it is only slight; there may be found also some of the wine-red male
coloring in some females; postmedian lines well expressed on both wings; white
discal spot large; about the dark scales what is said under the male applies also here;
underside of wings as upperside; abdomen, head, thorax, legs all ochreous. Length
of forewing 24-29 mm (30 specimens measured).

Head: Laterofrontal suture visible; frontal protuberance transverse, not promi-
nent; male antenna with 15 rami; posterior aspect of head showing two well
developed lateral processes of the margin.

19(3): 101-180, 1980(81)

145

Legs: Epiphysis in male a little more than half as long as the tibia; rounded
dorsally, straight ventrally, ending in a point, covered with very small hairs; in
females only occasionally present, much smaller than in male, originating at
beginning of second quarter of tibia, extending half way into third quarter, same
form as in male. Empodium moderate, consisting of one tubercle with a single small
seta. On anterior tibia only one obtuse process instead of the usual apical spines.
GENITALIA

Male (fig. 7e): Uncus M-shaped, heavily chitinized; gnathos not broad, triangular
shield; lateral margins of valves almost straight; inner margin of valves more uneven
than in A. assimilis; apex bifurcate, ventral part a little shorter, strongly chitinized,
dorsal part twice as wide; free annellus heart-shaped, with high apices, the cleft
strongly chitinized (as m A. consularis); aedoeagus different from all other species,
except those of the group, rounded like a boomerang, carina with serrated ridge
ventrally; vesica with one elongated spine -like, minutely serrate comutus spically
and in some populations with preapical one reduced to a chitinized plate or spot;
proximal end wider in lateral view than in other species of the genus. (Gen. prep.
ROM 3-002).

Female (fig. 8f): Very compact looking, especially the antrum; ostium narrow and
strongly sclerotized; antrum wedgeformed cephalad; ductus bursae abruptly bent
where bursa copulatrix is connected; lobuli vaginales very high; valves of ovipositors
of almost uniform width, large and apically rounded. These last three characters
diagnostic for the species. Sterigma as usual, perhaps not as wide as in other
species. The ductus seminalis is connected almost on the dorsal side of the end of
the antrum. Bursa copulatrix more or less rounded and not too large. Signum
present in some specimens, in other absent. (Gen. prep. ROM 3-008).

EGG

Orange, more rounded than flat; 1.1 x 1.6 x 0.8 mm.

LARVA (pi. I, fig. 10)

First instar: Body color dark olive; headcapsule and anal plate black; subspiracu-
lar faint yellowish line just perceptible; horns on thoracic segment II black ending in
two long setae; spiracles black with veiy indistinct bordering; spines on body not too
strongly developed; length 5 mm.

Second instar: Body color light yellowish olive; headcapsule light beige brown;
anal plate of body color but paler; horns on thoracic se^ent II very long and
slender, setae much shorter than in first instar; some scattered white granulations;
between subdorsal p*eemsh lines a dm*ker dorsal band; spiracular line also greenish;
spiracles blackish with body colored border; length 12 mm.

Third instar: Body color greyish olive; headcapsule as before and still lighter;
anal plate pale amber; horns on thoracic segment n shorter than before, the setae
very reduced; white granulations on body more conspicuous; a very faint dorsal line
(no band); subdorsal lines light pinkish; pinkish also a subspiracular band; spiracles
black, almost no bordering, between two fine whitish lines; length 20 mm.

Fourth instar: Body color as before; headcapsule greenish; anal plate pale
amber; horns on thoracic segment 11 shorter and thinner, no setae at end; white
granulations as before; dorsal line more expressed; between the subdorsal pinkish
lines patches slightly darker than the body; spiracles black with whitish bordering;
conspicuous subspiracular pinkish band; length 25 mm.

146

J. Res. Lepid.

Fifth instar: Body color greyish olive; headcapsule greenish; anal plate dull
amber; horns on thoracic segment 11 almost smooth, clubbed, 6-8 mm long; white
granulation not too dense over entire body; dorsal line fine; space between the pink
subdorsal lines darkened in patches; the subdorsal lines now wider and more
prominent; spiracles black with whitish bordering in a whitish, narrow, wavy band;
subspiracular band also wide, pink; all the colors more saturated than in the instar
before; length 42-55 mm. it should be noted that aU the colors in A. virginiensis are
pastel colors not strong colors as in A. pellucida. Headcapsule shown in fig. 5e,
suranal plate in fig. 3e.

PUPA (pL Vn, fig. 8)

Of the same general type as usual in the genus, but more slender posteriorly, the
lateral segmental thorns stronger, some double; the bifurcation of the cremaster
with an inwardly bent bifurcation.

DISTRIBUTION

The species is very widely distributed; it ranges from Arkansas through Missouri
northward to Minnesota and then east through Wisconsin, Illinois to the eastern
seaboard south to Virginia; in Canada it extends from Manitoba eastward through
Ontario and Quebec to Nova Scotia.

TYPE MATERIAL

Type material for the three names: virginiensis, astynome and perspicellina is not
known to be preserved anywhere.

As type locality for virginiensis Drury names Virginia; as that of astynome Olivier
gives Carolina and Virginia; Martyn does not mention type localities.

It is well known that the correct identity of the species has been questionable since
Drury’s (1773) original figure, which is in color but only certain as being a species of
Anisota. Smith (1797), Westwood (1837) and Boisduval (1872) variously confused
the Druiy taxon with A. pellucida and even A. senatoria. Packard (1905) synonymized
A. virginiensis and A. pellucida, and Morris (1892) even synonymized A. virginiensis
to A. pellucida, making thus the senior synonym the junior one.

To finally solve this problem we are designating herein a neotype for both
names: virginiensis and astynome.

With F. H. Rindge we select and designate as neotype for both names of A.
virginiensis and A. astynome a female from Suffolk, Virginia, collected 19 June 1944
by Otto Buchholz. This neotype is in the American Museum of Natural History in
New York.

BIOLOGY AND REMARKS

A. virginiensis is the widest ranging species in the North. The moths fly in June and
July mostly. TYie species is sympatric with A. manitobensis in some areas. The
preferred foodplant is Q. rubra.

Papers published which supposedly give the flight times of A. virginiensis
(Dominick emd Edwards, 1971; Solomon, 1971) actually refer to A. pellucida.
However, we believe the flight time for both these species is the same time of day.

There is evidence that A. discolor and A. virginiensis share some characters which
are not common to A. pellucida, In the fourth-instar larvae of the first two species
the stripes are light pink, while in A. pellucida these stripes are magenta. The ova of
A. pellucida are smaller and are mostly brown during emb^ogenesis, whereas the
larger eggs of A. discolor and A. virginiensis go through a red phase before hatching.

19(3): 10M80, 1980(81)

147

MATERIAL EXAMINED

CANADA. MANITOBA: Morden (CNC); Thornhill (CNC). NOVA SCOTIA:
Argyle (USNM); L. Kejimkujik (USNM); see also in the Proceedings of the Nova
Scotian Institute of Science, Session of 1952-53 (VoL XXH, Part 3), p. 209 under
no. 846. ONTARIO: Aberdeen (FIS); Acton (CNC); Algonquin Prov. Park (McM);
Banks (CNC); Bear L (CNC); Benny (FIS); Bergland (CNC); Blueberry 1. (ROM);
Brent (FIS); Capreol (ROM); Chaff eys Locks (ROM); Chalk River (FIS); Grand
Bend (CNC); Johns Falls (larva) (ROM); Kenebec 1. (FIS); Latchford (FIS);
Norman Twp. (ROM); Ottawa (CNC); Petawawa (FIS); Port Rowan (CNC); Rainy
River (CNC); Rondeau Prov. Park (ROM); Rosebank (ROM); Sault Ste. Marie
(CNC, McM); Severn Falls (FIS); Silver Water (FIS); Simcoe (CNC); St Thomas
(UWO); Sudbury (FIS); Trenton (CNC); Toronto (ROM); Vittoria (CNC); Washago
(CNC). QUEBEC: Aylmer (CNC); Calumet (LM); Ft. Coulonge (CNC); Laniel
(LM); Meach L. (CNC); Montreal (LM); Norway Bay (CNC); Orleans (LM-
Sheppard Coll.); St. Johns (LM); Ste -Anne -de-Belle-vue (LM); Wakefield (CNC).
UNITED STATES. ARKANSAS: no other data (AMNH). CONNECTICUT:
Putnam (AMNH). ILLINOIS: Lacon (UIl); Oregon (city) (UIl); Palos Park (UR);
Putnam Co. (UR). MAINE: no other data (YPM). MICHIGAN: Agr. College
(MSU); George Reserve (MSU); Gull. L. Biol. Station (MSU); L. Orion (MSU);
Wakelee (UR). MINNESOTA: no other data (AMNH); Itassa Park (USPM);
Ramsey Co. (USPM). MISSOURI: UMo CoU.; Columbia (also larvae); Curryville;
Meramec Highlands; Roaring River State Park (larvae); Round Spring State Park
(larvae); Vichy; Heitzman Coll, the foUowing Counties: Benton; Bollinger; Boone;
Carroll; Crawford; Cole; Jasper; Johnson; Lewis; Maries; Montgomery; Ozark;
Randolf; St. Louis; Taney; Texas; Warren; Washington. NEW HAMPSHIRE: no
other data (AMNH); Franconia (AMNH). NEW JERSEY: no other data (AMNH);
Elizabeth (AMNH); Essex Co. (AMNH); Fort Lee distr. (AMNH); Montclair
(•AMNH); Morris Co. (AMNH); Newark (AMNH, FMNH); Ocean Co.; Orange Mts.
(AMNH); Union Co. (AMNH). NEW YORK: Bear Mts. (AMNH); Bedford
(AMNH, FSCA); Bronx (AMNH); Bronx Park (AMNH); Brooklyn (CNC); Buffalo
(ROM); Clayton (AMNH); Ithaca (MSU); New York (AMNH). OHIO: Millport
(CM); Summitville (UMo). PENNSYLVANIA: Delaware Water Gap (AMNH);
Alleghany Co. (CM); Charleroi (CM); Finleyville (CM); Ingomar (CM); Jeannette
(CM); Pine Grove (AMNH, LACM, Peigler Coll.); Pittsburgh (CM); Scranton
(AMNH). RHODE ISLAND: Elmwood (CM); Rockland (AMNH). TENNES-
SEE: Johnson City (AMNH). VIRGINIA: Amherst (CM); Suffolk (AMNH).
WISCONSIN: Balsam L. (USPM). Examined 100-f males, 175+ females, dis-
sected 9 males and 11 females.

SENATORIA GROUP

Anisota finlaysoni Riotte

Anisota finlaysoni Riotte, 1969: 141; Ferguson, 1971: 76; Lemaire, 1976: 47.
Anisota senatoria (not J. E. Smith, 1797); Forest Lepidoptera of Canada (B. M.
McGugan, Co-ordinator), 1958: 48 (partim), [Error of determination.]

ADULT

Male (pi. Ill, fig. 6): Sexual dimorphism strongly developed. Head, thorax,
abdomen, legs, fore- and hindwings normal sepia brown and not too strongly

148

J. Res. Lepid.

sprinkled with dark scales in the apical area of forewing; hyaline patch on forewing
clear; postmedian line well marked; white discal spot quite large; on the underside
of wings a distinctive brass-yellowish suffusion. Outer margin of forewing very
slightly concave between M2 and Cu lb, of hindwings almost all straight. Length of
forewing 17-20 mm (16 specimens measured).

Female (pi. HI, fig. 7): Thinly scaled. Unicolorous, pronounced tawny ochreous;
only slightly sprinkled with dark scales at apex of forewing; postmedian lines on
both wings straight and unconspicuous; white discal spot quite large; a purplish
suffusion in the terminal area usually much reduced but in some populations
occurring; underside of wings with ochreous -yellow suffusion. Outer margin of both
wings rounded, the hindwings more so than the forewings. Length of forewing 24-30
mm (17 specimens measured).

Head: Laterofrontal Suture strongly visible. Frontal protuberance transverse,
insignificant. Male antenna with 18 rami. Posterior aspect of head showing no
lateral projections, only a low convexity on either side, apex rounded.

Legs: Epiphysis in male well developed, broad, longer than two thirds length of
tibia, width about half of length; one side rounded, the other straight, apex blunt,
covered with very small hairs. In most females epiphysis absent, in others veiy small
but of same form as in male. Empodium formed of two tubercles, the distant one
well developed with a long, strong seta, the proximal one very small with a very small
and thin seta,

GENITALIA

Male (fig. 7c): Uncus V-shaped, apices strongly sclerotized; gnathos a broadly
rounded shield, usually with a pronounced triangular process arising apically;
valves perceptibly curved distally; apex bifid, ends of almost equal length, ventral
part of the bifurcation sclerotized, dorsal part wider; free annellus a ring tapering off
apically. Aedoeagus short, straight, carina bent sharply, blunt, with a row of small
serrations; vesica with two conspicuous triangular comuti, the edge of each serrate;
the one at the middle of the vesica with unusual thin, small triangular process
dorsaOy; coecum of the same general form as in the other species. (Gen. prep. ROM
3-004).

Female (fig. 8e): Ovipositor valves rounded at apex; lobuli vaginales high,
triangular with more or less pointed apex; ostium bursae a wide funnel-like
structure; antrum absent; ductus bursae not sclerotized, short, sharply bent where
bursa copulatrix is attached; bursa copulatrix small, round, signum mostly absent or
very small; ductus seminalis connected at the very beginning of ductus bursae
laterally. (Gen. prep. ROM 3-006).

EGG

The freshly laid eggs are of a bright yellow color, darkening after some days to a
dark reddish brown. Before hatching they turn gray. The eggs are laid preferentially
at the apices of the leaves of oak. They hatch after about 15 days and are
approximately twice as big as the small eggs of A. senatoria.

LARVA (pi. I, fig. 5)

First instar: Light yellowish, body not hairy, longitudinally striped with shadowy
blackish, faintly pigmented lines: a narrow one at the dorsum and the spiracles,
two broader ones in between. The setae are thin and small and the two scoli on the
second thoracic segment are only very slightly thicker than all the other ones.

19(3): 101-180, 1980(81)

149

Length of first instar larva: 5 mm.

Second instar: Coloration as in the mature larva : blackish-brown with yellowish
stripes: two narrow dorsal ones extending only to abdominal segment 7 inclusive, a
straight and somewhat subdorsal line, an undulate supraspiracular one interrupted
between the segments, a straight infraspiracular and much interrupted line; the
base of the infraspiracular spines, inferior to the infraspiracular stripe, is of the
same color. Spiracles black. A light yellowish ventral median stripe. Prongs of the
prolegs of the same color. Main difference with aE other known larvae of Anisota is
the absence of the elongated, thin and slender “horns” on the second thoracic
segment.In their place are just two short scoli (1 mm long) which are only very
slightly larger than all the remainder. Pronotum heavily cMtinized, black with
yellowish borders which coincide with the second lateral yellow stripe. Head very
dark blackish red-brown, elongated, the fi’ontal plate quite large. Length of second
instar larva: 10 mm.

Third to last (fifth) instar: There is no perceptible change between the larva of
the second instar and the following ones except for the length. This is a remarkable
distinction from the larva of A. senatoria. Length of third instar larva: 21 mm; of
fourth instar larva: 32 mm, of fith instar larva: 50 mm. Headcapsule shown in fig.
5c, suranal plate in fig. 3c.

PUPA^L Vn, fig. 6)

BlacMsh brown; spiny and slender; cremaster bifid; the two forks stronger than in

A. senatoria and well pointed.

DISTRIBUTION

Since the original description and the publication of Ferguson (1971) it was found
that the distribution of the species corresponds to that of many other moth species
in southern Ontario which occur at the north shore of Lake Erie and then again
around Belleville and eastward.

TYPE MATERIAL

Type locality: Shannonville, Hastings County, Ontario. Additional type data:
holotype male taken 28 June 1967 sitting at the end of grass stalk at about 1500 hrs;
allotype female at the same date and time ovipositing on a white oak tree. Location
of types: Royal Ontario Museum, Toronto, Ontario. Paratypes in: AMNH,
BMNH, CM, CNC, FIS, FSCA, LACM, MSU, Peigler Coll, RIB, ROM, USNM,
UWO, YPM.

BIOLOGY AND REMARKS

The flight period is from the last third of June to about the middle of July. In the
laboratory pupae may fail to diapause.

The mam foodplant is white oak. Also chestnut oak {Quercus pirn) is found at the

type locality and in some instances larvae of A. finlaysoni were found on same. In
rearing experiments also other species of oak, especially red oak, were taken, and
larvae transferred to red oak trees on the grounds of Queen’s University Biology
Station at Lake Opinicon, Chaffeys Locks, Ontario, completed their development
on these but did not become established. Also the rare Quercus muhlenbergi was
accepted in the laboratory as food by the larvae.

L. R. Finlayson (deceased 1965), Research Officer at the Research Institute,
Canada Department of Agriculture, Belleville, Ontario, discovered the species hi
1946 after a fanner in Shannonville, Ontario, informed the Institute that an

150

J. Res. Lepid.

unknown caterpillar defoliated his oak trees. Finlayson collected over the years,
especially in 1957, considerable material of all stages of the species which he rightly
recognized as new and undescribed. In 1966 finally all specimens accumulated in
Belleville were turned over to the Department of Entomology and Invertebrate
Zoology of ROM (nobody else being interested in it) where careful study supported
Finlayson’s suspicions and provided the stimulus for the present review of the
genus Anisota.

On 28 June 1967 a fieldtrip was made to the original locality in Sbannonville
where the first observations had been made in 1946, One male and one female were
found. The circumstances were the same as already observed by Finlayson: the
male was sitting in the grass, the female was resting on a white oak leaf lajdng eggs.
About 400 eggs were found in three bunches on three white oak trees, in addition to
the eggs of the ovipositing female. AU eggs were reared and material of all stages was
preserved. A few adults emerged from the pupa only a short time after pupating, in
September 1967 which was an especially warm month, and the same happened
again in 1968.

Inspection in 1978 by both authors revealed the colony in Shannonville to be still
strong.

The species has been reared in southern France from Shannonville eggs by C.
Lemaire. Many larvae were dead in the first instar on the European Quercus
pubescem which was easily accepted in the last instar. However, successful rearing
was accomplished in five weeks on Castanea vulgaris. This is interesting and may
perhaps give a clue to the sporadic distribution of the species if it should have
originally preferred Castanea sp. and would after the great chestnut blight not have
been able to adjust so easily everywhere to oak.

The species apparently is sympatric or nearly so with A. senatoria in some areas of
Ontario. Elucidation of these ranges would be a worthwhile undertaking for some
local lepidopterist in the area. We believe that our records for A. senatoria and A.
finlaysoni are free from confusion because the determinations are generally based
on larvae or dissected adults.

MATERIAL EXAMINED

CANADA. ONTARIO: Ameliabourgh Twp. (FIS); Barriefield (FIS); Belleville
(CNC, FIS, ROM, UWO); Campbellford (FIS); CampbellvUle (FIS, ROM); Chip-
pewa (CNC); Deseronto (CNC, ROM); Erinsville (FIS); Frankford (FIS); Glen
Miller (FIS); HagersviUe (ROM); Kingston (ROM); Lindsay (FIS); Mountain View
(CNC); Napanee (CNC, ROM); Odessa (CNC, UWO); Otter Lake (ROM); Palermo
(FIS); Rockton (ROM); St. WiUiams (ROM); Sheffield (CNC, ROM); ShannonviUe
(CNC, ROM); Snyder (CNC, ROM); Thurlow (FIS); Tweed (FIS); TVendinaga
(FIS). UNITED STATES. MINNESOTA: no other data (AMNH); Green Lake
(UCal); Minneapolis (FMNH); Olmstedt Co. (USPM). WISCONSIN: Lake Delton
(Sieker Coll.); Chippewa Co. (CalAcSci). Examined long series of males and
females, dissected 22 males and 15 females.

Anisota peigleri Riotte

Anisota peigleri Riotte, 1975: 105; Lemaire, 1976: 47.

ADULT

Male (pi. ni, fig. 1): Abdomen, fore- and hindwings above strongly mahogany-
brown; head, legs, fore- and hindwings beneath ochreous except for an apical

19(3): 101480, 1980(81)

151

purplish-gray marginal space; forewings above sprinkled with dark scales especialy
apicaUy; beneath dark scales especially on hmdwmgs; postmedian line easily
perceptible on upper forewing and on al wings beneath; hyaline patch on forewings
present but much covered with tMn scales; m'Mte discaJ spot large. Outer margin as
in A. finiaysom, i.e. with forewing slightly concave between Mi and Cu lb, with
Mndwing perceptibly concave. Length of forewing 19-20 mm (12 specimens
measured).

Female HI, figs. 2, 3): LMcolorous lighter and of a clay brown with some
orange hue, except for the marginal area of the forewings which shws on both sides
slight purplish-gray hue, in some specimens (ex-larva, Ciajdon, Georgia) this
purplish strongly expressed over entire whig surface; most specimens with strong
sprinkling of somewhat clustered dark scales on both sides of the wings; some
specimens with only a very lew scales; white cMscal spot large. Outer margm of both
■wings slightly rounded. Length of forewing 25-32 mm (11 specimens measured).

Head: Unique sexual dimorijMsm in certain important characters: laterofrontal
suture visible in male but obsciirred in female; frontal protuberance slightly
protruding in male but reduced in female. Male antennae with 12 rami. Posterior
aspect of head ellipsoid, outline smooth without lateral projections.

Legs: Epiphysis slightly shorter than tibia, bulged unilaterally, densely covered
mth ve^ fine, short hairs, obsen^ed in the male. Empodium very small with a short,
weak seta.

GENITALIA

Male (fig, 7b): Uncus M-shaped, apices steongly sclerotized; gnathos similar to
A, finlaysord, i e. reduction in width from and even narrowing into a sudden
narrowness finally ending in a blunt end; in single specimens the apex of the gnathos
is lip-formed; valves narrow; apex bifid, ends of unequal length, ventral part of
bifurcation sfroiigly sclerotized, dorsal part wider; free anelliis a well-formed
crescent. Aedoeagus long, straight, carina bent sharply, apex like a goose head with
many Mttle teeth on top of “head”, siiU more than in A. finlaysmi; the proximal end
tripartite; dorsally only very slightly chltinized; vesica with a well-developed
comutus on apex, while the preapical one may be present or absent.

Female (fig. 9b): Ovipositor valves rounded at apex, oblong, strong, larg^e; lobuM
vaginales not too Hgh; ostium bursae wide; antrum absent; ductus bursae
unsclerotized not-very long; sharply bent where bursa copuiafrix is attached as in A.
finlaysfjni; bursa copulatrix short and oblong; no signum observed; dpctu^ seminalis
connecting at beginning of ductus bursae.

EGG

Yellow; 1 x 1 x 0.8 mm. Turning reddish brown and then gray as development of
embryo proceeds.

LAEVA (pi. I, fig. 6)

First instar: Body color, anal plate included, ivory; headcapsule black: dorsal line
tlmi, grayish-olive; horns on thoracic segment II long, bifurcate; on thoracic segment
HI shorter hornlike outgrowili, bifurcate, on a thicker base; spines aH over body;
long subspiracular bristles in horizontal row; on abdominal segment 9 bifurcate
bristles on a slightly larger base. ■* ' ■ '

Second instar: Body color yellowish; anal plate brownish-black; tMn dorsal line
^•ayish-olive and pale; a subdorsa! and lateral stripe of same color about three times

152

J. Res. Lepid.

the width as the dorsal one; also a spiracular line and subspiracular rounded scolion
each segment. In some cases the line on dorsum and the staipes are dark blackish-
olive. On both stripes and subspiracular region of each segment is a triangular hom;
on the subdorsal stripes is also a smaller hom latero-caudally from the larger one.
Thoracic horns only slightly bifurcate. Spiracles black. On segment anterior to anal
plate is a dorsal larger hom on a base. Small blackish spots irregularly all over body.

Third instar: Body color jet-black just after molting, later becoming more olive-
gray. Dorsal line black; broken whitish-yellow weak subdorsal lines; slightly thicker
lateral ones of the same type. Spiracular area yellow as in previous instar. Thoracic
horns long and only slightly bifurcate. On abdominal segment 1 1 and anal segment
are single strong dorsal bifurcate spines and two smaller lateral ones, no spiracular
spines. Anal plate black.

Fourth instar: Body color jet-black just after molting, some days later changing
to olive-gray. Very spiny. Three yellowish broken lines on each side; subdorsally
veiy faint broken line, absent in some cases. Homs long, clubbed with very fine
bifurcate setae apically. The spiracular line is unbroken and the subspiracular scoli
surrounded by yellow patches. All lines pale yellow, tending to whitish washed-out
color. Caudal spines as in previous instar. larvae very shiny in this and previous
instars (as in all species in the genus).

Fifth instar: Larvae now velvety jet-black; when viewed at certain angles a trace
of olive-gray color perceptible. Very spiny; much more so than in larvae of A.
finlaysoni and A. senatoria (consult pi. V). Subdorsal yellow lines often almost
absent or only slightly visible as disjunct dashes; lateral lines also broken,
sometimes only remnants visible; supraspiracular line intermittent; subspiracular
line wavy and thicker; a round yellow patch surrounding the spiracle on thoracic
segment 11. On abdomen a narrow yellow line from second abdominal segment to
posterior end, sometimes broken, this line much less evident than in A. senatoria
and A. finlaysoni.

The chaetotaxis of the thoracic legs differs from that of both A. senatoria and A.
finlaysoni.

PUPA (pi. Vn, fig. 5)

Medium spiny; chestnut brown; cremaster bifurcate with lateral ecrescences, the
latter apparently an important species -specific character (which also occurs in
Dryocampa).

DISTRIBUTION

Anisota peigleri, maintains high populations every year in upper South Carolina
and adjacent areas of North Carolina and Georgia. The following South Carolina
counties cover the area where the species appears to be most abundant: Anderson,
Oconee, Pickens, Greenville; in outl5mg areas it becomes rarer and still further
away (Florida and coastal South Carolina) it is difficult to find most years. The
species may ultimately be found in eastern Tennessee, northern Alabama, and
western Virginia, although A. senatoria may also occur in these areas. The
distributional pattern is generally characterized by moderate elevations, being
uncommon in higher mountains but common in Piedmont, with occasional
populations encountered in coastal areas. The populations reported by Beal (1952)
under the name A. senatoria possibly refer to the present species.

19(3): 101-180, 1980(81)

153

TYPE MATERIAL

Type locality: Clemson, Pickens County, South Carolina. Additional type
data: holotype male taken 5 August 1974 at light by R. S. Peigler; allotype female
taken at same locality 11 July 1974 at light by R. S. Peigler. Location of
types: Royal Ontario Museum, Toronto, Ontario. Paratypes: one male and four
females in ROM, one female each in AMNH, Lemaire Coil, Peigler Coll., all
collected at the same locality during the same summer by the same collector.
BIOLOGY AND REMARKS

The larvae of this species are very abundant during September and October in the
counties of South Carolina mentioned above under “Distribution”. Usually a few
lower branches are defoliated on medium-sized trees, but it is not unusual to see
isolated small or medium trees completely defoliated. Non-native oaks such as Q.
paiustris planted in yards and parks are especially attacked. In Greenville County
the larvae have been found on Q. rubra and Q. marilandica; in the Clemson area they
occur on Q. velutina; in all these areas of upper South Carolina the favorites are
however Q. nigra and Q. falcata. Larvae collected near Clayton, Georgia and seen in
Asheville, North Carolina by the junior author were all on Q. velutina.

The junior author and other entomologists at Clemson University (see Clemson
Univ. Coop. Ext. Serv., 1968) always assumed this near-pest to be A. senatoria. A
brood of larvae which the senior author reared in Canada from eggs of the allotype,
quickly demonstrated to him that two species were involved since he was familiar
with larvae of A. senatoria. Once this fact became clear, all of the other differences in
the immature and adult stages between the two species were readily found.

Larvae of A. peigleri are variable in the amount of orange striping, as in A. senatoria,
but these stripes are more often broken and reduced in the former species. The
amount of orange stripes present in A. peigleri appears consistent within colonies,
i.e. sibling larvae resemble each other closely. The spines are longer in A. peigleri,
and this trait is easily seen when a direct comparison is made of mature larvae of
both species. In the adults, the outer edges of the right and left wings diverge more
in A. peigleri, whereas they are closer to being parallel in A. senatoria. The forewings
thus appear more produced in A. peigleri. Although the examples we figure do not
show it, the white dot in the forewing of both sexes tends to be larger in A. peigleri.
Other differences between these two species, including genitalic ones, are given in
the formal descriptions of the two species. Narrower wings and more yellowish on
the underside distinguishes the females of A. peigleri from those of A. stigma.
MATERIAL EXAMINED

UNITED STATES. FLORIDA: Pensacola (LACM); Gainesville (State Agric.
Dept., Honolulu); Interstate-75 Florida Welcome Center (ROM); “Carolina, N.
Amerika” (AMNH). GEORGIA: Athens (UGA); Atlanta (Lemaire Coll.); Clayton
(LACM, Peigler Coll, ROM). NORTH CAROLINA: AsheviUe (LACM); Hender-
sonville (larval parasites with host remains (Peigler ColL, USNM). SOUTH
CAROLINA: Clemson (AMNH, FSCA, LACM, Lemaire Coll., Peigler Coll.,
USNM); Anderson (CNC); Pickens (BMNH); Six Mile (ROM); Seneca (Peigler
Coll.); Florence (CLU, Peigler CoU., UGA); GreenviUe (AMNH, BPBM, LACM,
ROM, Peigler Coll, Lemaire ColL); Westminster (AMNH, ROM); Moncks Comer
(ROM). Examined 254- males and 38 females, dissected 9 males and 8 females.

154

J. Res. Lepid.

Anisota senatoria (J. E. Smith)

Phalaena senatoria J. E. Smith, 1797: 113, pL 57.

Anisota senatoria; Huebner, 1818-1822: 193; Grote, 1864: 93; Packard, 1864:
384; Kirkby, 1892: 739; Neumoegen and Dyar, 1894: 147; Beutenmueller, 1898:
439, pi. 20: 3; Packard, 1905: 107-111, pL 20: 10-12; Holland, 1915: 94; Forbes,
1923: 668; Bouvier, 1931‘ 22; Schuessler, 1936: 213-215; Draudt, 1930: 813;
Ferguson, 1971: 73; Lemaire, 1976: 47.

Dryocampa senatoria; Harris, 1835: 72, 592; 1841: 292; 1862: 405, fig. 200;
Walker, 1855: 1496; Fitch, 1859: 43; Morris, 1862: 231.

Adelocephala senatoria; Boisduval, 1872: 87.

ADULT

Male (pi. ni, fig. 4): Sexual dimorphism strongly developed. Abdomen, thorax,
fore- and hindwings above from dark brown in the south to lighter brown in the north
with slight reddish hue, more expressed in the north; head, legs, fore- and hindwings
beneath dark orange-ochreous except for an apical dark purplish-grey marginal
space in forewings above sprinkled with dark scales especially apically; beneath
dark scales more pronounced; postmedian line strongly perceptible on upper
forewing and on all wings beneath; hyaline patch on forewings somewhat suffused
with thin scales; white discal spot not very large. Outer margin as in A. finlaysoni,
i.e,, of forewing slightly concave between Ma and Cu lb, of hindwing almost all
straight. Length of fore wing 15-19 mm (39 specimens measured).

Female (p. m, fig. 5): Abdomen, thorax, head, legs, fore- and hindwings above
and beneath from normally scaled brownish-yellowish ochreous with some orange
in the south to lighter yellowish ochreous and thinner scaling in the north also in
apparent dine, a more or less pronounced purplish suffusion in the terminal area;
postmedian lines on both wings and on both sides in southern specimens strongly
marked, in northern ones much weaker; white discal spot not very large; the
sprinkling with dark scales may be heavy in the south and nearly absent in the north,
it occurs on both sides of the wings; on the underside of the forewings apically a
more or less pronounced purplish-grey marginal space. Outer margin of all wings
evenly rounded. Length of forewing 26-30 mm (47 specimens measured).

Head: Laterofrontal suture slightly obscured; frontal protuberance well devel-
oped and protruding. Male antenna with 15 rami. Posterior aspect of head with
scalloped margin and perceptible but rounded processes.

Legs: Epiphysiss in male very small, only 3/5 of length of tibia; not observed in
female. Empodium very small with a tiny thin seta.

GENITALIA

Male (fig. 7a): Uncus M-shaped with heavily sclerotized apices; gnathos a
triangular roadly rounded shield; valves slightly rounded, apex bifid, both ends
almost of equal length, ventral part of the bifurcation strongly sclerotized, dorsal
part much broader; free annellus a ring of uniform width; aedoeagus straight and
short, Carina bent ventrally , strongly sclerotized, proximal to apex a low comb with a
few small thorns, apex itself blunt; vesica with comuti, one small one at half-way and
the other at the end spatulate with small thorns; proximal end of aedoeagus almost
straight. (Gen. prep. ROM 3-001).

Female (fig. 8d): Ovipositor valves of medium size and rounded at apex; lobuli
vaginales low; sterigma also low not prominent, but sclerotization more or less

19(3): 101-180, 1980(81)

155

heavy; ostium bursae not too wide, no antrum; ductus bursae not sclerotized at all,
straight, slender; bursa copulatrix elongate, signum absent; ductus seminalis
attached in upper fourth of ductus bursae laterally. (Gen. prep. ROM 3-009).
EGG

Yellow. Eggs from Monroe, MI, 1 x 1 x 0.5 mm; from Connecticut 1.3 x 1.2 x 0,9
mm.

LARVA (pi. I, fig. 4)

First instar: Body color greenish-yellow; headcapsule black; anal plate not
pigmented; thoracic horns on segment II long, smooth, clavate, with two terminal
setae of equal length; thoracic legs dark colored; abdominal prol'egs of body color;
no lines perceptible; length 4 mm.

Second instar: Body color dark greenish, shiny; headcapsule, anal plate and anal
legs jet black; yellowish green lines are now perceptible: a dark dorsal line; pale
yellowish green subdorsal lines; a wider supraspiracular line of the same coloring,
separated by a very narrow dark green line from a broad spiracular one and a row of
conspicuous black tubercles; horns on segment I large, finely spinulated, less
bifurcate than in the first instar; thoracic legs black, abdominal prolegs dark green,
anal legs black; length 10 mm; in the southern populations up to 18 mm.

Third instar: Body color dark yellowish green to olive green; headcapsule and
anal plate black; two narrow yellowish dorsal lines; yellowish subdorsal and supra-
spiracular lines; the spiracular line more narrow; spiracles black with yellowish
border; horns on thoracic segment 11 black, long; length 15 mm; in the southern
populations up to 23 mm.

Fourth instar: Body color now black; headcapsule and anal plate black; two
dorsal yellow (or orange-depending on the population) lines; the subdorsal line
very outspoken, not intermpted at the joints; the supraspiracular line wavy but not
broken; the subspiracular line irregular and broken; on each segment a yellow (or
orange) ornamentation below the subspiracular line on the body segment; the
supraspiracular line is extended on both sides of the anal plate; horns on thoracic
segment II longer and stronger than before; length 20 mm; in the southern
populations up to 34 mm.

Fifth instar: Body color blackish brown; headcapsule and anal plate black; lines
deep yellow ochreous or orange— depending on the population— two dorsal lines
(unbroken); a subdorsal straight one (unbroken); a supraspiracular undulating one;
a subspiracular one is broken by black intermittances and is bearing the subspiracu-
lar row of spines; all spines rather short (as in A. finlaysoni); same subspiracular
ornamentation as before; ventrally a yellow ochreous stripe; spiracles black with
yellowish border; horns on thoracic segment II as before (4 mm long, thin); all legs
and prolegs black; length 35 mm; in southern populations up to 40 mm.

PUPA (pi. Vn, fig. 4)

Spiny and not perceptibly broadened posteriorly. The cremaster bifid, the forks
being slightly turned outwards, sharp and not too long.

DISTRIBUTION

From the Gulf states, especially Texas and Louisiana (also Mississippi according
to Ferguson, 1971: 74), northwards to the Great Lakes, hence through the
southwestemmost part of Ontario to the eastern seaboard and then south to the
Carolinas and sporadically into Florida. The specimens mentioned by Packard,

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J. Res. Lepid.

1905, from Maine may well constitute the northern limit while the specimens
mentioned from Quebec by Winn, 1912, were found to be misidentified A.
virginiemis.

TYPE MATERIAL

With the discovery of A. peigleri it seemed necessary to tie the name A. senatoria to
a neotype. We have selected a specimen from a southern state although none from
Georgia was available.

We herein designate as neotype of A. senatoria the following male (pi. HI, fig.
4): Gibsland, Bienville Parish, Louisiana, 21 August 1976, leg. R. Peigler.
Additional type data; specimen collected at light at a rest area on Interstate 20.
Other males and females collected with this neotype are deposited in several
museums. Location of neotype; American Museum of Natural History, New York.
BIOLOGY AND REMARKS

We agree with Ferguson (1971) that Abbot’s plate leaves no doubt as to the
identity of this species. We do not, however, choose to restrict the type locality to
Screven or Bulloch counties in Georgia as Ferguson (1971) assumed for both A.
senatoria and A. pellucida in his text discussion.

Regarding the diagrammatic aspects of Abbot’s plates see also for fuller
discussion the above chapter. Larval and adult differences between A. senatoria
and A. peigleri are enumerated under the latter species. Additional observations con-
vince us that Abbot figured what we now call A. senatoria rather than what we call A.
peigleri. If the “assumed” type locality of coastal Georgia be accepted, it is
significant that the junior author failed to find A. peigleri in this area during three
years of collecting Anisota there. The data supporting that A. peigleri and A.
consularis share the same flight times imply that the two species could not coexist,
and the latter occurs there.

As pointed out by Kimball (1965) and Ferguson (1971), this species is rare in the
Southeast. In eastern Texas, northern Louisiana up through Missouri the species is
very common and attains pest status (see Ignoffo etal., 1973). This species is much
commoner in Walker Co., Texas, than either A. fuscosa or A. discolor but becomes
rarer westward; we have a record from Bastrop from R. 0. Kendall. The insect is
also abundant (although only periodically) in New England where Hitchcock (1958,
1961 a,b,c) carried out some excellent observations. We refer the interested
lepidopterists to his papers. Additional literature on the pest status include
Clarkson (1883), Claypole (1883), Fitch (1859) and Missouri Forest Pest Report
(1973).

Specimens from Texas and Louisiana appear darker than those from the
Northeast, although we find no larval differences. Benjamin D. Williams III reports,
in litt., that he has attracted males to “caOing” females early afternoon, during the
two hours following noon. He reports Q. palustris as a host record. In the southern
populations it is possible that males fly later in the day since several were collected
at light and the forewings are less transparent. Additional material and data from
central states are needed.

There is an old specimen labelled “Mexico” in the Lemaire collection from the
Hans Fruhstorfer collection which Lemaire kindly verified by dissection. Given
what is said about “Mexico sensu lato” under A. leucostygma this specimen may
well be one of the numerous Texas ones.

19(3): 101-180, 1980(81)

157

MATERIAL EXAMINED

CANADA. ONATRIO: Ailsa Craig (FIS); Amer (CNC); Breslau (FIS); Dere-
ham (ROM); Frome (CNC); Grand Bend (CNC); Kingsville (FIS); Ojibway (44
larvae and 100 pupae ex 1947, original Finlayson research) (ROM); Poplar Hill
(FIS); Ranelagh (ROM); Wallenstein (CNC); Winona (FIS); Yarmouth Centre
(CNC). UNITED STATES. ARKANSAS: Crawford Co. aarvae) (UArk); Hope
(LM). CONNECTICUT: Mansfield (AMNH). FLORIDA: no other data
(AMNH); Archbold Biol. Station, L. Placid (AMNH); Monticello (MCZ). DELA-
WARE: Delaware City (CM); New Castle Co. (AMNH). ILLINOIS: Algonquin
(un); Palos Park (UR). INDIANA: Hessville (FMNH); Knox (AMNH, FMNH,
YPM); N. Judson (UH). IOWA: Decorah (AIVINH). LOUISIANA: Dry Creek
(larvae) (LSU); Gibsland (AMNH, BPBM, LACM, Lemaire Coll,, Peigler Coll.,
ROM, USNM). MASSACHUSETTS: Chatham (MCZ); Martha’s Vineyard
(AMNH, LM, YPM). MICHIGAN: Agric. College (MSU); Joseph Co. (MSU);
Monroe (and larvae) (ROM); Ramona (LACM); Wayne Co. (MSU). MINNESOTA:
no other data (AMNH); Hennepin Co. (USPM); Long Prairie (LACM); St. A. P.
(USPM). MISSOURI: Bagnell (LACM, Sieker ColL, YPM); Phelps Co. (larvae)
(UMo); Pike Co. (Heitzman, in litt); Stone Co. (larvae) (UMo); Texas Co. (larvae)
(Peigler Coll., ROM). NEW JERSEY: no other data (MJCZ); Bay Head (MJCZ);
Clementon (ROM); Cranford (AMNH); Elizabeth (FMNH); Jamesbourgh (AMNH);
Lakehurst (AMNH); Morris Co. (AMNH); Newark (AMNH, CM); Paterson
(AMNH), NEW YORK: Albany (AMNH); Brooklyn (CNC); Buffalo (CM, ROM);
Long Island (AMNH); New Windsor (CalAcSci); New York (AMNH, CM, CalAcSci,
LM); West Farms (CM). NORTH CAROLINA: Southern Pines (LM); Sylva,
Jackson Co. (FMNH). OHIO: Anna (LACM); Millport (CM, ROM, UMo);
Summitville (CM, ROM, UMo). PENNSYLVANIA: no other data (CalAcSci);
Alleghany Co. (CM); Delaware Water Gap (AMNH); Lititz (FMNH); Pittsburgh
(CM); Rock Hill Furnace (CM); Rockpoint (CM); Scranton (AMNH). RHODE
ISLAND: no other data (MCZ); Gienwood (CM); West Kingston (AMNH);
Providence (CM). SOUTH CAROLINA: no other data (CIU); Greenville (larvae)
(Peigler ColL). TENNESSEE: BurrvUle (FMNH); McKenzie (CM). TEXAS:
Bastrop Co. (Kendall ColL, ROM); College Station (Peigler ColL, ROM); Jasper
(larvae) (ROM); Marshall (LACM); Nacogdoches (AMNH); San Jacinto Co.
(ROM); Stubblefield L., Walker Co. (AMNH, BPBM (and larvae), LACM, Lemaire
ColL, ROM). VIRGINIA: Kauhana (MCZ). WISCONSIN: Wausau (LACM).
Examined 114 + males, 114 + females, dissected 16 males and 12 females.

STIGMA GROUP

Anisota consularis Dyar

Anisota consularis Dyar, 1896: 166; Packard, 1905: 106-107, pL V: 8; Bouvier,
1931: 23; Draudt, 1930: 814; Kimball, 1965: 69, pL HI: 23-25; Ferguson, 1971:

72; Lemaire, 1976: 47.

Anisota stigma (not Fabricius, 1775); Ferguson, 1971: 67,72.

ADULT

Male (pi. n, fig. 5): Sexual dimorphism strongly developed. Head, Thorax,
abdomen, legs, fore- and hindwings dark sepia-brown and strongly sprinkled with
dark scales, even on underside; hyaline patch on forewing very thinly covered with
scales; postmedian line well perceptible, especially on hindwing; white discal spot

158

J. Res. Lepid.

well-developed; on underside of wings orange tinge. Outer margin of both wings
almost straight. Length of forewing 19-21 mm (21 specimens measured).

Female (pi. H, fig. 6): Head, thorax, abdomen, legs ochreous brown; forewings
raw sienna to rosy brown (quite variable); hindwings darker and usually with a rosy
tinge; all wings strongly sprinkled with dark scales; postmedian lines weak but
sometimes well expressed; white discal spot quite reduced. Outer margin of both
wings strongly convex. Length of forewing 26-30 mm (19 specimens measured).

Head: Laterofrontal suture slightly obscured. Frontal protuberance perceptibly
protruding. Male antenna with 18 rami. Posterior aspect of head distinctly rounded
at apex, processes slightly rounded.

Legs: Epiphysis observed only in males; % of length of tibia; very convex dorsally
midway along its length; straight ventrally; ending in narrow point; covered with
short bristles; bearing on the ventral margin somewhat longer ones. Empodium one
thin tubercle with one medium-sized seta.

GENITALIA

Male (fig. 6c): Uncus narrowly M-shaped, apices chitinized in a very peculiar
way, the sclerotization forming a ridge of variable intensity; lateral borders of
gnathos sinuate, apically forming a triangular rounded projection; valves basally
and laterally very straight; parts of bifurcation of almost equal length, the dorsal
part twice as wide as the ventral part; free annellus heart-shaped, strongly
chitinized in cleft between the low apices. Aedoeagus very short, carina bent,
straight, its apex with a row of small teeth; vesica very narrow with two small similar
comuti, the apical one with a few small teeth; proximal end not as sharply outlined
as in the other species. (Gen. prep. ROM 3-030).

Female (fig. 8b): Ovipositor valves large, elongate; lobuli vaginales low; sterigma
of medium breadth, not as much lipformed as in the other species; ostium bursae
quite narrow; ductus bursae somewhat bottle-shaped, narrowing towards bursa
.copulatrix which is conspicuously oval and in the specimens inspected had a signum
in the form of around plate with concentric circles of chitinization; ductus seminalis
attached laterally just after the beginning of ductus bursae. (Gen. prep. ROM 3-
031). i
EGG

Yellowish, 1 x 1.1 x 0.6 mm. _ .

LARVA (pi. I. fig. 3)

First instar: Body color pale yellowish with a slight greenish tinge; headcapsule
jet black; anal plate of body color; fine dorsal line olive; subdorsal line same color,
twice as thick as dorsal one; spiracular line same color, waved, interrupted at
segments; long spines on each segment subdorsally, on a thick conelike base, black;
long horns on thoracic segment El bifurcate; short horns on thoracic segment III
bifurcate; length 3 mm.

Second instar: Body color as in first instar, however, some larvae tending to a
more dark coloring; headcapsule light brown; anal plate also; dorsal line narrow
olive-black; subdorsal lines same color washed out, much wider; a washed out olive
supraspiracular line bordered by a whitish line on each side; an olive black
spiracular line with light bordering; spines now short; also horns on thoracic
segment II only of normal length, bifurcate; short, bifurcate horns also on thoracic
segment HI; length 4.5 mm.

19(3): 101-180, 1980(81)

159

Third to fith instars: Body color fleshy to red-brown; headcapsule reddish-beige -
brown; anal plate of body color, however, in flesh colored larvae both, headcapsule
and anal plate are with all the feet red-brown; dorsal, subdorsal, supraspiracular
and spiracular lines olive-blackish with the tendency to the black; often all lines
bordered by a whitish space so that entire larva looks much more white than colored
because of many round white spherish ecrescenses all over the body; larvae with
only few white ecrescenses show the body color between the lines; spines very
strongly expressed; spiracles blackish with whitish border; thoracic horns on
segment II clubbed, 10 mm long; short scoli on segment HI bifurcate; on segments 8
and 9 longer scoli, on segment .9 only one in the middle; length in third instar 10 mm;
in fourth instar 25 mm; in fifth instar 32 mm. Headcapsule shown in fig. 4c, suranal
plate in fig. 2c.

PUPA (pi. Vn, fig. 2)

Compared with pupae of the other species of Anisota quite slender and of even
width; not very spiny; cremaster short and compact; bifurcation straight outwards,
very pointed; color lighter brown.

DISTRIBUTION

Although previously only known from widespread records in Florida, the junior
author collected the species in Long and Bulloch counties of coastal Georgia. The
latter area is very close to the state line of South Carolina so that one might expect
the species in that state (but only near the coast) but it was not found in northern
Charleston County after many years of intensive collecting there by the late R. B.
Dominick. The moth might also be found eventually in other states on the Gulf
Coast. The range is evidently limited by an ecological need for a mild climate.
TYPE MATERIAL

Type-locality: West Palm Beach, Florida. Additional type data: reared from
larvae found in Januaiy 1896 on live oak. Location of types: United States
National Museum, Washington, DC; the male type designated by Ferguson (1971)
as lectotype, August 1896; the female type 15 September 1896. Types were
examined by the junior author.

BIOLOGY AND REMARKS

It is apparent unfortunately that no previous author has had a clear understanding
of the species, including Ferguson (1971) who questioned the color figures in
Kimball (1965, pi. 3, figs. 23, 24 and 25). These do indeed represent A. cpnsularis,
as the senior author verified these specimens (in FSCA) by dissection, 'Ferguson’s
figures for this species (1971, pL 5, figs. 16, 17 and 18) are correct, but his figure 5
on the same plate is also A. consularis, not A. stigma. This latter female was verified
(in USNM) by the junior author, Ferguson (1971: 72) also questioned the correct
identification of the figure in Draudt (1930, pi. 142f), saying it “is an obvious
stigma”. This may be true or the figure is just as likely correct, especially if one
considers the artist may have “corrected” the transparent forewings with the belief
these had been rubbed.

Although it is now abundantly clear that Ferguson’s “Floridian stigma” is actually
A. consularis, he was correct in seeing the relationship of it to the Texan A. fuscosa.
In these two species there is ground color variation from light beige to rust to light
orange to purplish brown in both sexes, while specimens of A. stigma are almost
always rusty colored.

160

J. Res. Lepid.

Fig. 8. Female genitalia, with lateral view of ovipositor alongside: a. A. stigma, b.

A. consularis, c. A. manitobensis, d. A. senatoria, e. A. finlaysoni, f. A.
uirginiensis, g. A. leucostygma, h. A. oslari, i. A. assimilis.

The mature larvae can be greenish cream, or brownish, or-pink fleshcolored, the
latter color form with a black and white lateral stripe on each side. The pink form
was figured by Packard (1905, pi. 5, fig. 8), All the larvae within a brood may be one
of these color forms, or such may not be the case. This explains a remark in Packard

19(3): 10M80, 1980(81)

161

(1905: 107) about larvae of A. consularis being found “in company with those of A.
stigma”, in an area far south of the known range of the latter species. These color
variations occur in mature larvae of A. stigma and A. fuscosa also.

Until now, the only reported host was live oak, Quercus virginiana (Dyar, 1896;
Packard, 1905; Ferguson, 1971). The junior author collected larvae in Bulloch Co.,
Georgia, on Q. nigra and Q. falcata and also on these two trees in central Florida. D,
Baggett collected them in Duval Co., Florida, in dry sandy areas on Q. laevis and Q.
myrtifolia.

In Georgia the species flies in late July through early August judging from larval
finds. In Florida records are mainly July through September depending on latitude
but we believe the species is strictly univoltine.

The relatively high ratio of females over males in collections leaves no doubt that
males are day-fliers. Reared females emitted pheromone in upper South Carolina at
the flight time of A. peigleri. Since these two species are entirely alloptaric, having
the same flight times poses no problem of hybridization in nature. This presumed
flight time is additionally supported by the only partially developed transparentness
of the forewings in the males.

MATERIAL EXAMINED

UNITED STATES. FLORIDA: no other data (AMNH); Cassadega (CNC,
FSCA, HOton Coll.); Crystal River (LACM); Gainesville (FSCA); Jacksonville
(AMNH, LACM); Lake City (USNM); Largo (CNC); Lutz (AMNH, CM, ROM);
Martin in Marion Co. (ROM); Miami (AMNH, CM); Palm Beach Co. (USNM); St.
Johns Co. (ROM); Stamper (CM);'Stuart (USNM); Summer Haven (CM); Suwanee
Springs (Lemaire Coll, Peigler Coll.). GEORGIA: Long Co. (larva) (ROM);
Statesboro, Bulloch Co. (AMNH, LACM, Lemaire Coll, Peigler Coll., ROM).
Examined 40 males, 22 females, dissected 3 males, 3 females.

Anisota fuscosa Ferguson New Status

Anisota stigma fuscosa Ferguson, 1971: 70; Lemaire, 1976: 47.

ADULT

Male (pi. II, fig. 3): Sexual dimorphism not marked in this species. Head, thorax,
legs, abdomen, fore- and hindwings almost' unicolorous reddish brown, only
terminal area of fore wings with a mostly only slight purplish hue, the basal area
without such hue. Forewings sprinkled with dark scales in varying density;
hindwings not or only with very few dark scales in terminal area. No hyaline patch on
forewing; postmedian line on both wings blackish-purplish, not strongly developed;
antemedian line only slightly expressed or even absent; white discal spot prominent;
middle field not contrasting (as it is in A. stigma). Outer margin of somewhat
pointedly triangular forewings more straightened than rounded; of hindwings
rounded. On the underside the unicolorous reddish brown is overlaid with some
ochreous orange; terminal areas wholly or at least apically with purplish hue; only
postmedian lines expressed. Length of forewing 20-21 mm (10 specimens measured).

Female (pi. H, fig. 4): Head, thorax, legs, abdomen, fore- and hindwings
unicolorous clay brown with very slight purplish hue in terminal area. Forewing
strongly sprinkled with dark scales;, hindwings only slightly developed; antemedian
line very slight; white discal spot small. Outer margin of wings as in male. On the
underside unicolorous clay brown, dark scales massed apically on fore wings and
over entire surface of hindwing. Length of forewing 32 mm (4 specimens measured).

162

J. Res. Lepid.

Head: Laterofrontal suture vei^^ much obscured; frontal protuberance very
perceptibly protruding; male antennae with 16 rami; posterior aspect of head
triangular with flat simple or double rounding on each side between base and tip.

Legs: Epiphysis of % length of tibia; ventrally slightly rounded; dorsally highly
rounded in middle; ending in blunt point; covered with fine bristles, ventrally with
somewhat longer fine hairs, Empodium one tubercle with one strong, not too short
seta. • ' :

GENITALIA

Male (fig. 6b): Overall impression square; uncus a strong, broad W; gnathos
evenly proceeding triangular shield, somewhat pointed; longer than in A. stigma;
valves rounded, apex bifid, dorsal bifurcation not much longer than ventral one
which is strongly sclerotized; free annellus a low crescent of even height; aedoeagus
short; carina rounded at apex, strongly sclerotized, with a comb of not too
prominent little teeth, not too large, pointed and thorned comutus apically and a
preapical one which may be a sharp thorn (as in A. consularis) or just a very slightly
sclerotized spot or anything in between, this being observed only in A. fuscosa.
(Gen. prep. ROM 3-159).

Female (fig. 9a): Ovipositor valves large and round apically; the basic part
conspicuousy long, longer than in any other investigated species of Anisota; lobuli
vaginales strong; antrum strong developed, almost as in the A. virginiensis group
with chitinized wedge-form into ductus bursae; ductus bursae twice twisted, first
time after antrum making sharp left turn, then where bursa copulatrix attaches
turning to the right; ductus brusae a wide and strong “hose”; bursa copulatrix quite
elongated with a small round chitinized mark as signum; bursa and ductus showing a
narrow network of fine lines. (Gen. prep. ROM 3-160).

EGG

Light yellow, round; size approximately like that of A. stigma.

LARVA (pi. I, fig. 2)

■ First instar: Body color ivory; headcapsule brown; anal plate brown; thoracic legs
brown; no dorsal line; horns on thoracic segment II brown, bifurcate with thin seta
on each end; 2 rows of subdorsal brown scoli with seta; supra- and subspiracular
scoli very small with seta; spiracles very light brown; length 8 mm.

Second instar: Body color ivory; headcapsule beige; anal plate brown; only short
setae on horns and scoli; subdorsal scoli now already well-developed; spiracular
line, dorsal line and field between subdorsal scoli brownish shadowed; spiracles
brown; scoli on 8th and 9th segments not different; length 13 mm.

Third instar: Body color beige; headcapsule beige; anal plate beige; scoli with
lateral white secondary ones; broad dorsal band brown between rows of scoli;
supraspiracular scoli weak, subspiracular ones stronger developed; spiracles brown
with light yellow frame; brownish band laterally between spiracular and dorsal scoli
(weaker than dorsal band); 2 scoli on 8th segment strong and long; on 9th segment
only 1 scolus also long and strong in the middle of the segment; length 20 mm.

Fourth instar: Body color beige; headcapsule beige; anal plate beige; thoracic
legs beige; scoli as before; 3 well- developed brown bands: dorsal, spiracular and
between subspiracular row of scoli; between the bands the beige body color;
subspiracular scoli on a beige pot; beige also between subdorsal row of scoli and
small supraspiracular scoli; spiracles brown with yellow trimming; scoli on 8th and

19(3): 10M80, 1980(81)

163

Fig. 9. Female genitalia, with lateral view of ovipositor alongside: a. A. /uscosa,
b. A. peiglerii, c. A. pellucida, d. A. discolor.

9th segments prominent; on all brown bands many white spots and round
ecrescenses; thoracic horns 11% mm long, clubbed with 2 obtuse ends with very
small seta each; length 25 mm.

Fifth instar: Body color beige; headcapsule beige; anal plate beige; thoracic legs
beige; body covered with many white spots and round ecrescenses; a very fine
blackish dorsal line in the middle of a blackish brown band between the subdorsal
rows of not too prominent black scoli; horns on thoracic segment 11 blackish,
clubbed, only 5 mm long; a lateral band, a little narrower than the dorsal one,
between the small supraspiracular and the also small spiracular scoli; scoli on 8th
and 9th ssegments only slightly stronger than the subdorsal ones; length 45 mm.
Headcapsule shown in fig. 4b, suranal plate in fig. 2b.

PUPA (pL vm, fig. 1)

Chestnut brown as in other species in the A. stigma group; cremaster very strong
and prominent; arising from a wide and short base; especially the bifurcation very
pronounced.

DISTRIBUTION

This species ranges far into central Texas (Kerrville) beyond the grassland
prairies but is much commoner in the forests of eastern Texas. We figure a pair fi*om

164

J. Res. Lepid.

Leesville, Louisiana, where Roy and Connie Kendall collected the larvae on which
the larval descriptions are based. In western Louisiana A. fuscosa ranges eastward
at least to Choudrant but A. stigma occurs commonly at Vicksburg, Mississippi (B.
Mather, pers. comm.), only ca. 150 km farther east. In southern Louisiana the range
probably does not extend as far east. The species probably occurs in Oklahoma but
we have no records for that state. We do not believe that A. stigma occurs in Texas
unless near the state line with Arkansas.

TYPE MATERIAL

Type-locality: Giddings, Lee Co., Texas. Additional type data: holotype female
taken 15 August 1908 by R. A. Cushman, type no. 71495 United States National
Museum. Type examined by the junior author. Most of the paratypes were collected
in eastern Texas by A. and M. E. Blanchard.

BIOLOGY AND REMARKS

The color forms of the adults and larvae discussed in this section under A.
consularis also exist in the present species, but these variations are even more
striking in both adult ground color and mature larvae. One rare larval color form is
very dark brown (almost black as in A. senatoria). The head, prolegs and anal plate
are orange. It is not dissimilar to the hybrid larva shown in pi, I, fig. 9. No larvae of A.
stigma or A. consularis which we have seen approach this form in darkness, but
larvae of A. fuscosa are usually of the lighter forms.

The most important and obvious trait of the larva of A. fuscosa is the presence of
the broad dark dorsal band which occurs in all instars except the first. It is absent in
some mature individuals of the pink color form.

Larvae found on several species of oak such as Q. stellata in College Station and Q.
nigra, Q. falcata and Q. marilandica in Texas counties and Louisiana parishes to the
east. The junior author searched unsuccessfully for larvae at the type locality
(Giddings) on Q. nigra growing along a stream. The caterpillars are exceedingly
delicate in captivity, and usually die from disease. Of large numbers collected at
various instars during three consecutive autumns in Walker Co., Texas, by the
junior author, very few survived to pupate and no pupae produced adults. R.
Kendall reported similar difficulty. However, C. Lemaire successfully reared a
brood in southern France on Castanea vulgaris.

The flight period extends from 25 May to October but the peak is in early
September of what we believe is one brood per year. The long flight period reflects
the large population level in some areas of its range. Males are commoner at lights
than fenaales, as in A. stigma.

MATERIAL EXAMINED

UNITED STATES., LOUISIANA: Choudrant (AMNH, BMNH, LACM,
Lemaire Coll., Peigler Coll.); Gibsland (AMNH, LACM, Lemaire Coll., Peigler
Coll., ROM); Leesville, Vernon Parish (Kendall Coll., ROM); Many (Kendall Coll.,
LACM); Spring Ridge, Caddo Parish (larvae) (LSU). TEXAS: Biyan (larva)
(TAMU); Cherokee Co. (TAMU); College Station (LACM, Peigler Coll., TAMU);
Fort Worth (CalAcSci); Giddings (USNM); Handley Garva) (USNM); Huntsville
State Park (USNM); Jasper Co. (larvae) (LACM); Kerrville (USNM); Milam,
Sabine Co. (Kendall Coll.); Nacogdoches (BPBM, Peigler Coll.); Shepherd (USNM);
Stubblefield Lake Recreation Area, Walker Co. (larvae) (AMNH, LACM, Peigler
Coll., ROM, USNM); Tennessee Colony (USNM), Tyler (TAMU). Examined long
series of males and females, dissected 5 males, 6 females.

19(3): 10M80, 1980(81)

165

Anisota manitobensis McDunnough

Anisota manitohensis McDunnough, 1921: 73; Brodie, 1929: 98-100, figs. 1-3
(biology); Ferguson, 1971: 71; Lemaire 1976: 47.

ADULT

Male (pi. n, fig. 7): Head, thorax, abdomen, and legs brownish orange; strong
purplish suffusion on both wings; basic color dull orange; no hyaline patch on
forewing;postmedian line dark purple and well-developed; white discal spot
prominent (in one specimen the white spots have a white anterior branch as in
Sphingicampa blanchardi Ferguson); underside of wings as on upperside. Outer
margins of both wings weakly rounded. Dark sprinkling of wings usually lacking.
Length of forewing 19-23 mm (10 measured).

Female (pi. H, fig. 8): Like male but larger. Length of forewing 26-30 mm (4
measured).

Head: Laterofrontal suture totally obscured; a slightly perceptible convexity all
along it; at occiput a small oval eoncave spot. Frontal protuberance broadly
rounded. Male antenna with 15 rami. Posterior aspect of head very scalloped with
conspicuous rounded processes.

Legs: Epiphysis found only in male, about half as long as tibia, dorsally

rounded, ventraUy straight, ending in a ventrally bent point, covered with short
hairs, the ventral margins bearing quite long and straight hairs. Empodium small,
one tubercle with one seta.

GENITALIA

Male (fig. 6d): Uncus V-shaped, apices (% of total length of each arm) strongly set
back and heavily sclerotized; gnathos broadly rounded triangular shield as in A.
stigma, but with blunt apex; valves strongly rounded, giving entire structure a
circular appearance; apex of valves bifid, both ends of almost equal length; ventral
part of bifurcation strongly sclerotized, dorsal part only a little wider; free anellus
crescent-shaped. Aedoeagus very broad and very short, on both sides slightly
convex; carina bent in a smooth curve, its apex rounded with only a few tubercle-like
processes instead of the usual teeth; vesica with two strong comuti at the usual
locations; both with a few uneven teeth, the preapical one with knoblike base (as in
A. stigma); proximal end straight as usual. (Gen. prep. ROM 3-016).

Female (fig. 8c): Ovipositor valves medium sized, strongly rounded at apex;
lobuli vaginales approximately as high as in A. finlaysoni; sterigma strongly
sclerotized, higher than usual, having on either side a peculiar triangular ventrally-
pointed form; ostium bursae quite wide; ductus bursae slightly sclerotized, a very
strong tubular structure, gradually cmwed ventro-laterally; bursa copulatrix usually
small, longitudinal, shaped somewhat like a shoe; signum, if present, a round patch
with an arrowhead-like scaling in the center; ductus seminalis connecting at the very
beginning of ductus bursae. (Gen. prep. ROM 3-017).

EGG

¥

According to Brodie (1929): laid in clusters, “ovate and flattened, 1.51 mm by
1.25 mm, bright sulphur yellow, shiny, quite opaque and perfectly smooth”. After
one week darkening to brown and hatching after 13 days,

LARVA

The following description of each of the instars is taken directly from the paper by
Brodie (1929):

166

J. Res. Lepid.

Fig. 10. Morphological structures, a-b. Male and female genitalia of A. dissimilis.

cA. Male and female genitalia, suranal plate and head capsule of larva of
Dryocampa rubicunda.

19(3): 101-180, 1980(81)

167

First instar: Creamy white, head shiny black; a wide dorsal and a narrow lateral
brownish black line; spiracle brownish; the usual horns on the second thoracic
segment. Changing of color in development to light and dark tan and black.
Length: 3 mm.

Second instar: Shiny black, head dark tan; two dorsal cream colored stripes; a
short light cream colored stripe diagonally through each spiracle; spiracles black;
ventral side dull cream to buff; thoracic horns 2.2 mm, bifurcating at tip. Length: 7
mm.

Third instar: Head light tan; dorsal cream colored stripes present; spines,
previously all black, now black at base and snow white toward tipi spines on anal
segment almost entirely white; short white streak diagonally through each spiracle;
ends of prolegs brown (previously grayish); colors of larva changing rapidly through
development; black parts becoming dark brown, e.g. the body; dorsal cream colored
lines becoming a dark shade of buff. Very shiny in this stage. Length: 10 mm.

Fourth instar: Head light tan, body shiny dark brown; two lighter brown dorsal
stripes; spiracles black; horns on second thoracic segment shiny black with white
branches, 5 mm long; other spines similarly colored; anal segment now dark brown
(previously black) with scattered white spines; prolegs dark brown; white on spines
giving larva appearance of being covered with hoar-frost (similar in A. stigma).
Colors changing during development to light pinkish brown. Length: 22 mm.

Fifth instar: Little change from previous instar; head paler tan; true legs changed
from black to pale tan; body light pinkish brown; many more spines now white;
spiracles black with light pinkish border; horns on second thoracic segment 6 mm
long; the area between the two dorsal lines with a thin central black line and with
dark shading on either side of this line. Length: 50 mm.

PUPA (pi. vn, fig. 3)

Reddish brown; evenly wide leading to a dome -like last segment with a strong
medium sized bifid cremaster; the forks strong and wide but not very pointed.
DISTRIBUTION

This moth was previously only recorded from southern Manitoba until we
obtained specimens from three localities in Wisconsin from Mr. William E. Sieker
(part of these we deposited in ROM and AMNH as indicated under “Material
Examined'*). The species probably also occurs in North Dakota and should
certainly be found in Minnesota.

TYPE MATERIAL

Type locality: Aweme, Manitoba. Additonal type data: male holotype collected
4 July 1907 and female allotype collected 23 June 1904 both by Norman Griddle.
Location of types is CNC with paratypes in CNC and USNM. We examined all of
these. .j

BIOLOGY AND REMARKS

Brodie (1929) fed his larvae on Quercus macrocarpa Michx. He got his larvae from
Mr. T. M. Short. The insect has been taken in recent years by Manitoba
lepidopterist Mr. C. S. Quelch.

The flight period is June and July, the records for Manitoba being for both
months, those of Wisconsin only for July.

As given under our remarks on the distribution, this species can now be listed as
part of the United States fauna. The Wisconsin material required dissection for

168

J. Res. Lepid.

verification of the determination because most were heavily spotted as in A. stigma.
The original description (McDunnough, 1921) and Ferguson (1971) state that A.
manitobensis completely lacks this sprinkling and most of the museum material we
saw bears this out. No problems, however, should arise in distinguishing the two
species since they are apparently allopatric and the latter has more pointed
forewings. Some specimens of A. stigma (in AMNH) from Rhode Island and
Massachusetts appear very much like A. manitobensis with reduced or absent
sprinkling and pink coloring in the postmedian area. The present species is
sympatric with A. virginiensis; see maps 31, 32 in McGugan (1958).

MATERIAL EXAMINED

CANADA. MANITOBA: Aweme (CNC); Darlingford (AMNH, CNC); Deer
Lodge (CNC); Kelwood (CM); McCreary (Peigler Coll.); Pembina Valley (CNC,
ROM); Pine Ridge (Peigler Coll.); Riding Mtns. (CM, USNM); St. Vital (CNC);
Transcona (Lemaire Coll.); Winnipeg (CNC, USNM). UNITED STATES.
WISCONSIN: Arlington (ROM); Madison (one from University of Wisconsin
Arboretum) (Sieker Coll.); Waushara County (AMNH, ROM). Saw 17 males, 8
females, dissected 3 males and 2 females.

Anisota stigma Fabricius

Bombyx stigma Fahricius, 1115: 563; Olivier, 1790: 42; Kerr, 1910: 110 (types).

Phalaena Bombyx stigma-, J. E. Smith, 1797, II: 111, pi. 56.

Dryocampa stigma; Harris, 1841: 292.

Adelocephala stygma; Boisduval, 1871: 86.

Anisota stigma; Huehner, 1818-1822: 193; Beutenmueller, 1898: 430, pi. 20, fig.
4; Holland, 1903: 94, pi. 9, figs. 9, 10; Packard, 1905: 98-102, pi. 20, figs. 4-9;
Forbes, 1923: 667; Draudt, 1930: 813; Bouvier, 1931: 17; Schuessler, 1936: 215-
218; Kimball, 1965: 69.

Anisota stigma stigma; Ferguson, 1971: 67; Lemaire, 1976: 47.

ADULT

Male (pi. n, fig. 1): Head, thorax, abdomen, legs, fore- and hindwingss ochreous
reddish-brown; forewings sprinkled with many dark scales, hindwings less; no
hyaline patch on forewings; postmedian line on both wings dark purplish and well-
developed; antemedian line weak to well expressed; white discal spot prominent;
terminal area of fore wings with more or less outspoken purplish hue; basal area
shows this hue to a lesser degree; middle field never with it. Outer margin of all
wings rounded. On the underside the ochreous ground color more pronounced
against the brownish tones; the purplish in many specimens strong apically on fore-
and hindwing; only postmedian lines strongly expressed. Length of forewing 19-24
mm (25 specimens measured).

Female (pi. 11, fig. 2): Like male, larger, often with more brownish than reddish
overtones and rarely even leaning to yellowish. When marked purplish suffusion
appears with decreased black sprinkling or no purplish suffusion with much
sprinkling indentification can be difficult and genitalic dissection can be helpful.
Also the fact that the anterior wings are broader and more strongly convex in the
outer margin should make the decision easier. The white discal spot is conspicuous.
Length of forewing 27-36 mm (17 specimens measured).

Head: Laterofrontal suture obscured; frontal protuberance very perceptibly
protruding; male antennae with 15 rami; posterior aspect of head with triangular

19(3): 10M80, 1980(81)

169

rounded large thorns.

Legs: Epiphysis in males a little more than half the length of the tibia, dorsally
rounded, ventrally straight, ending in a point, covered — especially ventraliy—with
rather long, dense hairs; in females, if present, only very rudimentary; empodium
small with a very short seat (see: Oiticica, 1940: pi. 12, fig. 2).

GENITALIA

Male (fig. 6a): Overall impression pronouncedly square. Uncus with strongly
sclerotized apices in the form of a broad M; gnathos a triangular shield with rounded
apex, wider than in A. peigleri, A. senatoria and A. virginiensis and not as long as in
these; valves strongly rounded and bent distally, apex bifid, the dorsal bifurcation
longer than the ventral one which is strongly sclerotized; free annellus conspicuously
crescent shaped; aedoeagus short, carina bent and rounded at apex, strongly
sclerotized with a comb of thorns; vesica with pointed and thomed cornutus apically
and another halfway with a knob-like base ending in a trifurcation; proximal end of
the same general form as in the other species. (Gen. prep. ROM 3-045).

Female (fig. 8a): Ovipositor valves large; strongly rounded at apex; lobuli
vaginales low; sterigma compact, strongly sclerotized; ostium bursae of medium
width; ductus bursae not sclerotized, but a straight, strong, tubular structure; bursa
copulatrix sometimes very small, normally, however, large, elongate, ovate; signum
mostly absent, sometimes a round slightly sclerotized patch present; ductus
seminalis attached in upper fifth of ductus brusae, laterally. (Gen. prep. ROM 3-
042).

EGG

Light yellow, more round and not so much flattened; 1.3 x 1.5 x 0.7 mm.
LARVA (pi. 1. fig. 1)

First instar: Body color pastel yellow; headcapsule black; anal plate also black
(note here difference from Packard, 1905: 99, who says that the anal plate is pale,
of the color of the body, not pigmented); thoracic horns on segment 11 very long,
bifmxate; on segment HI bifurcate already at base, shorter, base brownish; spines
on all segments, on a base, this base brownish on segment 1, on the other segments
blackish, on segment 9 one only large scolus on a base, bifurcate with setae on both
ends; dorsal line very thin, olivish, secondaiy spines all over the body in a horizontal
row subspiracularly; spiracular line greyish-olive; length 3 mm.

Second instar: Body color darker into brownish; headcapsule black; anal plate
black; dorsal line narrow, blackish; subdorsal lines wider, blackish; supraspiracuar
and spiracular lines not so wide and fainter in color; spiracles black; length 9 mm.

Third instar: Body color reddish-brown; headcapsule dark mahogany, anal plate
black; horns on thoracic segment 11 black, perceptibly barbed; anal legs with a large,
red, triangular spot; abdominal prolegs dull red with a black spot; the lines showing
in the second instar become faint, reduced to traces; some white granulation begins
to appear over the entire body; first traces of two lateral pinkish lines appear; the
three dorsal lines are blended in one another forming a broad, pale reddish-
brownish band; on each side of the head a pale, whitish line; length 16 mm.

Fourth instar: Body color reddish-brown, headcapsule dark mahogany; anal
plate black; horns on thoracic segment 11 less spinulated than in the third instar,
wth white granulations, 10 mm long; on thoracic segment I six large, strong, forked
scoli, not bifurcate, with a single seta; spiracles black with a thin whitish bordering;

170

J. Res. Lepid.

PLATE V. Scanning electron micrographs of larval integument. 1, 3, 5. A.

senatoria. 2, 4, 6. A. peigleri. Scale of figs. 1-2 is 22 mm:l mm, of figs.
3-4 is 16 mm;300 pm, of figs. 5-6 is 16 mm:30 pm.

color of anal legs verying between dark reddish and black; thoracic legs and
abdominal prolegs dark pitchy; any lines almost not perceptible; length 32 mm.

Fifth instar: Body color brown; headcapsule dark mahogany; anal plate brownish;
supraspiracular lines now better perceptible, greyish-pinkish; the whole larva very
much as the one of the fourth instar otherwise; length 48 mm.

NB: Packard, 1905; 101, mentions an occasional sixth instar that we, however,
were not able to observe in any of our rearings.

Headcapsule shown in fig. 4a, suranal plate in fig. 2a.

PUPA (pi. vn, fig. 1)

Posterior segments decreasing only slightly in diameter; the spines are con-
spicuous; cremaster long and strong, bifurcation weak; in this the pupa is different

19(3): 10M80, 1980(81)

171

from all other known puape of any Anisota; color chestnut brown.
DISTRIBUTION

This species does not occur as far to the north as A. senatoria and A. virginiensis .
We find it, however, widely distributed along the eastern seaboard from Florida to
Massachusetts; westward through extreme southern Ontario to Minnesota, Kansas,

Arkansas. There are no specimens known from Mexico.

TYPE MATERIAL

Type-locality: “America Meridionalis”. Location of the two cotypes: Collection
of insects of the University of Glasgow, Scotland. The female specimen (lower one
on pi. VI, fig. 3) is hereby designated by us as lectotype.

Boisduval, 1871, had expressed some doubt as to the identity of this species
which was figured for the first time by J. E. Smith, 1797, in both larval and adult
stage. Boisduval bases his contention on the fact that Fabricius states as type-
locality “America Meridionalis” and not “America Borealis”, as might be expected
if the species came from the United States. However, when Fabricius named the
species there did not yet exist any United States of America and America just meant
the British Colonies in North America (see Oxford Dicitonary), hence “Am. Merid.”
would be what we call today the “Southern States” including Spanish possessions
like Florida and Mexico. Moreover, the types, preserved in the Hunter collection
and reproduced here after an original photograph made for this purpose in 1967,
show without any reasonable doubt specimens of A. stigma.

BIOLOGY AND REMARKS

This species is so widespread and well-known that very little new information can
be offered here. The moth has been collected with ultraviolet light by the junior
author atop Mt. Mitchell, North Carolina, the highest point in eastern North
America. It is generally common in the northern and southern states of the eastern
half of the continent. The northern range into Canada needs additional records to
delineate it.

One error which keeps creeping up into dealings with A. stigma is the record in
McGugan (1958): “near North Bay in northern Ontario”. The original report shows
as locality Bear Island, Joan Township, Nipissing District, it was made in 1953 from
a larval collection. However, from Bear Island there is an ex larva female of A.
virginiensis in CNC ex FIS from 1956. The identification of the original report was in
the meantime changed to “Anisota sp.” (P. D. Syme, Canadian Forestry Service,
Insect Pathology Research Institute, Sault Ste. Marie, Ontario, in litt.). Also North
Bay is far from “southern Ontario” (Ferguson, 1971: 68) which should have elicited
some suspicion in using this report again.

A. stigma is uncommon in central Florida and absent southward in that state.
Specimens in the AMNH and others sent us by D. Baggett all originating in the
panhandle of Florida average considerably smaller than material from elsewhere.
Additional material, with associated preserved larvae, from the Gulf coast area
would be interesting.

At Clemson, South Carolina, the junior author found larval colonies in the
university arboretum on Q. michauxii and Castanea mollissima. Also at Clemson he
once found larvae on Q. alba, his only observation of this oak species ever being
utilized by Anisota in a southern state.

Seasonal flight times vaiy with latitude, altitude, and weather for the particular
year, but most adults are taken in June, July or August. The circadian flight times

172

J. Res. Lepid.

were mentioned in a previous section. Variation in larval coloring is cited under the
text of A. consularis. Resemblance of adults in New England to those of A.
manitobensis is discussed under the latter species.

The Nova Scotia record given below needs verification due to absence of records
from Maine, New Hampshire, Vermont and Quebec.

MATERIAL EXAMINED

CANADA. NOVA SCOTIA: Truro (CNC). ONTARIO: London (ROM); Port
Colbome (CNC); Queenston (CNC); St. Williams (CNC, FIS); Simcoe (CNC).
UNITED STATES: ALABAMA: 8 km NE Eutaw (AMNH); Ida (CalAcSci);
Loxby (AMNH); Wilmer (AMNH). ARKANSAS: no other data (AIVINH); Blue
Springs Campground (Heitzman Coll.); Bluff City (AMNH); Camden (LACM,
Peigler Coll.); Chidester (Lemaire Coll.); Crawford Co. (larvae) (UArk); Hope (LM,
UMo). CONNECTICUT: Deep River (AMNH); Manfield (AMNH); Old Lynn
(AMNH); Putnam (AMNH). FLORIDA: Alachua Co. (FSCA); Archbold Bio. Sta.,
L. Placid (AMNH); Bonita Springs (AMNH); Gainesville (FSCA); 3 km E. Lamont
(UM); Monticello (MCZ); Quincy (AMNH, FSCA); Shalimar (Hilton Coll.);
Suwannee Springs (pupa ex larva) (ROM); Torreya State Park (AMNH, LACM);
White Springs (larva) (AMNH). GEORGIA: Atlanta (AMNH, CM); Cooper Creek
State Rec. Area, Fannin Co. (AMNH, LACM, UGA); Emory Univ. (FSCA);
Lawrenceville (UM); Macon (UM); Ohoope River (CM); Savannah (UM); Tifton
(UGA). ILLINOIS: Chicago (FMNH); Lacon UIl); Osborn (UM); Palos Park
(FMNH, UH); Putnam Co. (UIl). INDIANA: Bristol (FMNH); Hessville (FMNH,
UIl, UM); Knox (FMNH); Miller (FMNH); Mitchell (FMNH); Tresmont (FMNH,
YPM). KANSAS: Johnson Co. (Heitzman Coll.); Ottawa (AMNH). KENTUCKY:
no other data (MCZ); Morehead (CNC); Rio (UM). LOUISIANA: Baton Rouge
(LSU); Denham Springs (LSU); Indian Mound (FMNH); Tallulah (larva) (UMo).
MASSACHUSETTS: Barnstable (AMNH); Chatham (MCZ); East Warshaw
(AMNH); Edgartown (LM); Lawrence (ROM); Marthas Vineyard (MCZ); Sherborn
(MCZ). MICHIGAN: Agric. College (MSU); Allegan (MSU); Augusta (MSU);
George Reserve (MSU, MU); Highland Park (UM); Lansing (UM); Lenawee Co.
(ROM); Nine Mile Road (UM). MINNESOTA: Minneapolis (YPM). MISSISSIP-
PI: Aditon (ROM); Bovina (AMNH, BPBM, LACM, Lemaire Coll., Peigler Coll.,
ROM); Hattiesburg (AMNH, ROM); Jackson (AMNH, ROM); Johnson State Park
(LACM); Mississippi State (LACM); Oxford (BPBM, LACM); Pearl (AMNH,
ROM); Waynesboro (LACM); Wilmer (LACM). MISSOURI: no other data
(CalAcSci); Adair (Heitzman Coll.); Ashland (larva) (UMo); Bagnell (AMNH,
LACM); Benton (Heitzman Coll.); Boon (dto.); Camden (dto.); Columbia (also
larvae) (UMo); Curryville (UMo); Greene (Heitzman Coll.); Hartsburg (UMo)
Jasper (Heitzman Coll.); Jefferson (dt.); Livingston (dto.); Pike (dto.); Putnam
(dto.); Randolf (dto.); St. Louis (dto.); Ste. Genevieve (dto.); Stone (dto.);
Washington (dto.). NEW JERSEY: no other data (AMNH); Arlington (AMNH);
Cape May Co. (AMNH); Cedar Run (AMNH); East Marion (CNC); Elizabeth (CM);
Green Village (larva) (AMNH); Lakehurst (AMNH); Manahawkin (AMNH); Morris
Co. (AMNH); Newark (AMNH, FMNH); Ocean Co. (AMNH); Paterson (AMNH);
Plainboro (AMNH); Smithville (AMNH); Stanhope (AMNH); Watchung Mts.
(AMNH); Wodersey (AMNH). NEW YORK: no other data (AMNH, LM); Bedford
(AMNH, FSCA); Bronx (AMNH); Brooklyn (CNC); Buffalo (CM); Grand Island
(CM); Leviston (AMNH); Long Island (AMNH, CNC); New York (AMNH);
Northport (AMNH); West Farms (AMNH). NORTH CAROLINA: Bryson City

19(3): 10M80, 1980(81)

173

PLATE VI. 1. Anterior end of larva of A. senatoria showing mesothoracic scoli
typical for the genus. 2. A. finlaysoni which has these horns greatly
reduced. Both original photos by L. R. Finlayson. 3. Cotypes of
"‘Bombyx stigma Fabricius” in Hunter Collection at Univ. of
Glasgow, Scotland.

174

J. Res. Lepid.

(UM); Cedar Mt. (UCal); Crestmont (UM); Fontana (CM, LACM); L. Toxaway
(AmNH); Maxton (AMNH); Mt. Mitchell (Peigler Coll); Raleigh (USPM, larva
LSUO; Southern Pines (LM). OHIO: Columbs (UM). PENNSYLVANIA: Butler
(CM); Delaware Water Gap (AMNH); Mt. Airy (AMNH); Pittsburgh (CM);
Sharpsburg (CM); Shawville (CM); Slippery Rock (CM). RHODE ISLAND: Deardon
(MCZ); Elmwood (CM); North Scituate (AMNH); Rockland (AMNH). SOUTH
CAROLINA: Clemson (AMNH, BPBM, CLU, LACM, Lemaire Coll., Peigler
Coll., ROM); Columbia (LACM); Coosawhatchie (AMNH); Florence (CLU);
Galivants Ferry (AMNH); Myrtle Beach, Arrowhead L. (CNC); Six Mile (AMNH,
LACM); Sleepy Creek (CM). TENNESSEE: Allardt (UM); Burrville (CNC);
Gatlinburg (FMNH); Knoxville (CNC); Monteagle (AMNH, CNC, FMNH); Sun-
bright, Morgan Co. (FMNH). VIRGINIA: Falls Church, Fairfax Co. (UM); Nelson
Co. (CM). WISCONSIN: Chippewa Falls (AMNH). Examined 161 + males, 118+
females, dissected 19 males and 16 females.

ARTIFICIAL HYBRIDIZATION

A few crosses have been made within the genus, aU within the last few years.
Females of any species will apparently attract males of any species providing they
emit pheromone at the right times of day and year for the particular males in the
area.

A. peigleri cf X A. consularis 9

Using a female of A. consularis reared from a larva taken in Statesboro, Georgia,
males of A. peigleri were attracted in Greenville, South Carolina. The percent hatch
was extremely high. Mature larvae (pi. I, fig. 9) were strikingly intermediate , having
reddish heads, black bodies, and yellow-orange stripes. Adults were intermediate,
tending more toward A. consularis. A pair is figured on pi. II, figs. 9-10. Specimens
are in AMNH, LACM and ROM.

A. senatoria cS X A. oslari 9

Using Arizona stock of A. oslari, a male of A. senatoria was attracted at 2:15 PM
(EDT) in Pomfret, Connecticut by Benjamin D. Williams HI. He succeeded in
rearing several to adults. The larvae were not unlike those of the above cross with
light heads and dark bodies, and the adults showed a nice blend of characters. After
overwintering, the females emerged in May and the males in June and July.

A. pellucida d' X A. virginiensis 9

Using adults from caged pupae this cross was made in Toronto by the senior
author with a male from McClellanville, South Carolina and a female from Chaffeys
Locks, Ontario. Larvae and adults were intermediate, even in genital characters.
The series is in the ROM.

A. virginiensis X A. discolor 9

Adults emerged from caged pupae, the father being from Pine Grove, Pennsylvania
and the mother from Hamilton, Texas. Larvae looked more like the father species
and one larva had distinct light green in place of the dark gray stripes. Some
diapaused and others did not. A large series of adults was obtained and these
resembled A. virginiensis mostly and exhibited very little variation among them-
selves. Most of the series is in AMNH with pairs in other collections. A pair is
figured on pi. 11, figs. 11-12.

19(3): 101^180, 1980(81)

175

PLATE Vn, Photographs of pupae showing cremasters. 1. A. stigma. 2. A.

comularis. 3. A. manitobensis. 4. A. senatoria. 5, A. peigleri. 6. A.
finiaysoni. 7. A. pellucida. 8. A. virginiemis. 9. A. pellucida. 10. A.
assimilis. 11. A. dissimilis. 12. D. mbicunda.

176

J. Res. Lepid.

PLATE VIII. Photographs of pupae showing cremasters. 1. A. fuscosa. 2. A.
discolor. 3. A. oslari.

A. virginiensis cT X A. discolor 9 F2

A brother-sister pairing of the above hybrids resulted in ova which gave very little
hatch. Larvae were weak and only three reached the final instar but none pupated.
Possibly a disease killed these larvae.

A. pellucida cf X (A. virginiensis cf X A. discolor 9) 9
A hybrid female attracted a male of A. pellucida in Greenville, South Carolina at
12:28 PM (EDT) and she laid a full complement of ova. All larvae died when very
small which suggests hybrid inviability.

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THE JOURNAL ©F
HJ THE LEFIJ

k\

Volume 19 Numbers Autumn, 1980(81)

IN THIS ISSUE

Date of Publication: December 1, 1981

A Revision of the American Genus Anisota (Satumiidae)

J. C. E. Riotte & Richard S. Peigler

Introduction 101

Higher Classification 101

Methods 102

Acknowledgments 103

Generic Diagnosis and Morphology of the Imago 104

Morphology of the Immature Stages 108

Intrageneric Classification and Phylogeny 108

Key to Adults 110

Key to Mature Larvae 111

Ecology, Collecting in the Field, and Rearing in Captivity 112
Parasitism 119

John Abbot 121

Treatment of the Species 122

Artificial Hybridization 174

Literature Cited 176

Cover Illustration: Scanning electron micrograph of A. peigleri larval integu-
ment. Scale “ 22 mm:l mm. See page 170, this issue.

THE JOU
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Journal of Research on the Lepidoptera

19(4): 181-195, 1980(81)

Enzyme Electrophoretic Studies on the Genetic
Relationships of Pierid Butterflies (Lepidoptera:
Pieridae) I. European Taxa*

H. J. Geiger

Zoologisches Institut der Universitat Bern, Sahlistrasse 8, CH 3012 Bern, Switzerland

Abstract. The phylogenetic relationships of 24 taxa representing the four
European subfamilies of the Pieridae were studied using enzyme electro-
phoretic techniques. The mobilities of 20 enzymes, each determined by a
different locus, were compared. A dendogram is presented indicating the
grouping of the taxa on the basis of biochemical data. Results agree well with
those obtained from conventional systematic studies. In cases where the
relationships between taxa are under discussion, one of the alternatives is
supported. The Pierinae and Anthocharinae are biochemically clearly
characterisable entities although the branching point of these two sub-
families in the dendogram lies only slightly below those of different genera.
These two subfamilies are separated from the Coliadinae by a clear step of
genetic divergence. Within the recently subdivided genus Pieris Shrank s. Itr.
of the Pierinae, three groups of taxa that branch at the same level of
similarity in the dendogram can be recognized. The taxa of the new genus
Artogeia Vrty. are no more closely related to each other than they are to the
taxa of Pieris s. str. Biochemical data support the species rank of the taxa
cheiranthi Hbn., simplonia F. and crameri Btlr., but indicate a remarkably
low level of genetic differentiation between the taxa napi L. and hryoniae
Hbn.

Introduction

The systematics of pierid butterflies have been extensively studied by
several independent taxonomic methods and the Pieridae are now one of
the best known butterfly families. Despite this, diverging views exist on
the relationships and status of several taxa at different hierarchical rank.
Systematic problems at the levels of 1) subfamilies and tribes, 2) genera,
subgenera and species groups and 3) species and subspecies are discussed
below.

1) In a generic revision of the family, Klots (1931/32) grouped the
Pieridae into three subfamilies: Pseudopontiinae, Dismorphiinae and
Pierinae and subdivided the Pierinae into three tribes: Euchloini, Rhodo-
cerini and Pierini. This classification, with or vrithout modifications, has

•This study was supported by the Swiss National Science Foundation Grant No. 3.640.80.

182

J. Res. Lepid.

been followed by virtually all subsequent authors. In a comparative analysis
of the morphology of the Papilionoidea, Ehrlich (1958), following other
authors (e.g. Ford, 1945), elevated the Coliadinae (Rhodocerini sensu
Klots, 1931/32) to subfamily status. This classification reflects the closer
morphological relationship between the Pierini and Euchloini than
between these two tribes and the Coliadinae or Dismorphiinae. Higgins
(1975) adopted this system for the European Pieridae, but gave the
Anthocharinae (Euchloini sensu Klots) subfamily rank. In the same work,
Higgins included the genus Catopsilia Hbn. in the Pierinae in the tribe
Callidryini Kirby but most other authors, including Klots (1931/32), have
regarded this genus as best placed in the Coliadinae.

2) Of the European taxa, species of the genus Pieris Schrank have been
especially problematical. This genus has long been recognized as being
globally distributed and various authors have included a great number of
taxa in it, (e.g. Roeber, 1908). In his revision of the Pieridae, Klots
(1931/32) restricted the number of taxa and subdivided the genus into
four subgenera. Of the European taxa, he arranged brassicae L, in the
subgenus Pieris and the taxa napi L., rapae L., manni Mayer and ergane
Geyer in the subgenus Synchloe Hbn. (together with e.g. callidice Esp.).
Bemardi (1947) also used the system of subgenera, but, in part, arranged
the species in different subgenera than previous authors. In his system, the
subgenus Pieris was divided into three species groups: Pieris 1 s. str. with
brassicae (of the European fauna), Pieris 2 with napi, rapae, manni and
ergane and Pieris 3 which is not represented in Europe. Kudrna (1974),
following Verity (1947), suggested that the species group which includes
napi, bryoniae Ochsenheimer, ergane, manni, rapae and krueperi Staudinger
should be placed in a separate genus, Artogeia Verity, and that Pieris only
houses brassicae, cheiranth Huebner, deota Niceville and brassicoides
Guerin-Meneville. However, Forster and Wohlfahrt (1976) claimed that
the taxa included in Artogeia and Pieris are, in fact, congeneric.

3) Problems with European taxa at the species and subspecies level are
well illustrated by the taxa of the Pieris (Artogeia) napi-group, the status of
which have been widely discussed. In the past, it has not been possible to
determine the phylogenetic distances between many taxa of this group and
it has been necessary to use complicated taxonomic constructions such as
the super- and semispecies concepts of Lorkovic (1962), Varga and Toth
(1968) and Bowden (1972). Theories on the evolution of this group are
cortfused (Petersen, 1949; Varga and Toth, 1978; Bowden, 1972) while
relationships between North American, Asiatic and European taxa remain
uncertain (Bowden, 1962). One reason for this confusion is the great
variability of morphological characters within very similar taxa (for the
variability of wing markings of napi and bryoniae see Mueller and Kautz,
1939). Investigations of closely related taxa indicate that the variability in
coloration seems to be strongly influenced by environmental factors (e.g.
Koyler, 1966; Shapiro, 1975, 1977).

19(4): 181-195, 1980(81)

183

Table 2

Enzyme

Abbreviation

Buffer System

Enzyme Stain

Adenylate kinase
(2 Loci)

AK-1

AK-2

1

Brewer, 1970

Gly c era Id ehyde -phos-
phate dehydrogenase

GAPDH

1

Harris and Hop-
kinson, 1976

Arginine kinase

APK

1

Scholl (1)

Halate dehydrogena-

MDH-l

1

Harris ans Hop-

88 (2 Loci)

MDH-2

kinson, 1976*

Aldolase

ALD

1

ibid

Indophehol oxydase

IPO

1

Brewer, 1970*

Malic enzyfrffe

ME

2

Harris and Hop-

kinson, 1976

Hexokinase

HK

1

ibid

Pyruvate kinase

PK

1

ibid

Glutamate-oxaloace-
tate transaminase

(2 Loci)

GOT-1

GDT-2

1

ibid*

6-Phosphogluconate

dehydrogenase

6-PGD

1

Brewer, 1970*

Fumarase

FUM

1

ibid*

Isocitrate dehydro-

IDH-1

1

Ayala et al . ,

genase (2 Loci)

IDH-2

1972

Glutamate pyruvate
transaminase

GPT

2

Harris and Hop-
kinaon, 1976

Phosphoglucomutase

PGM

1

Brewer, 1970

Phosphoglucose Iso-
marase

PGI

1

Scholl et al.,

1978

Remarks :

Buffer Systems s System Is Tris-citric acid buffer (Ayala et el., 1972)
System 2s Tris borate EDTA buffer (Ayala et al., 1972)

♦With minor modifications, but the original staining medium would also

be sufficient

(1) Scholl (personal communication ) s Agar overlay method. Staining

mixtures Glycin buffer (1.52 M Glycin, 42 mM MgCl2 * 6 ^2^* 9.0) s

5ml, PEP s 10 mg, ATP s 40 mg, NADH^ s 10 mg, LDH s 90 iU, PK s 20 iU

Discussions are also in progress on the Euchloe “aiisonia- complex”,
especially the status of the monovoltine populations flying in the Alps and
Pyrenees and the bivoltine populations of the lower part of France and
Italy (Back, 1979).

Recently biochemical methods have become important in the analysis of
phylogenetic relationships. Enzyme electrophoresis in particular has
proved veiy informative (Ayala et aL, 1974; Avise, 1974; Selander and
Johnson, 1973). In this method, the extent of genetic divergence is
estimated from variation in electrophoretic mobilities of a sample of

184

J. Res. Lepid.

homologous enzymes of the various taxa. This approach is basically a
genetic method but differs from other methods, for example the determin-
ation of interspecific fertility- bridges or sterility-barriers by hybridization
experiments, in that the degree of genetic divergence is estimated from a
defined sample of the genotype. Again this is not the case when evaluation
of phylogenetic relationships is based on morphological data.

It has been demonstrated that phylogenetic independence of a taxon is
correlated with gradual biochemical-genetic divergence (Ayala et al.,
1974; Avise, 1974; Selander and Johnsen, 1973) manifested in an
increasing number of electrophoretically distinguishable enzyme variants.
Enzyme electrophoresis may provide a basis independent of environmental
variation for assessing the evolution of the Pieridae. This paper reports

Tabls 1

T aatm

Number of

Countries

Number of

Number of

animals

of origin

population
samples (1) '

sampling
sites (2)

Aporia crataegi L.

19

CH,F

1

2

Pieris brassicae L.

141

CH,F,D,IRL

9

17

Pieris cheiranthi Hbn.

16

E

1

1

Artogeia rapae L.

154

CH,F, I,D

9

17

Artogeia mannii Mayer

25

F

1

1

Artogeia napi napi L.

276

CH,F, I ,D,GB

16

24

Artogeia napi bryoniae Hbn.

63

CH

4

4

Pontia daplidice L.

5

CH,F,I

0

4

Pontia callidice Hbn.

5

CH

0

1

Euchloe simplonia Freyer

11

CH

0

5

Euchloe crameri Btlr.

11

F

1

1

Euchloe tagis bellezina Bois.

13

F

1

1

Anthocharis cardamines L.

45

CH,F

3

10

Anthocharis euphenoides Staud.

11

F

1

1

Colias phicomone Esp.

78

CH

5

5

Colias palaeno europome Esp.

60

CH

4

4

Colias myrmidone Esp.

6

D

0

1(3)

Colias crocea Geoff.

9

I

0

1

Colias hyale L.

5

D

0

l(3)

Colias australis Vrty

7

CH,D,F

0

4

Catopsilia florella F.

4

E

0

1

Gonepteryx rhamni L.

6

CH

0

1

Gonepteryx cleopatra L.

7

F

0

1

Leptidea sinapis L.

13

CH,F

1

1

Remarks s

Code for countries: CH ^ Switzerland, D ^ Federal Republic of Germany,

E = Spain, F * France, GB = Great Britain, I = Italy
IRL = Republic of Ireland

(1) a population sample consists of a minimum of 10 individuals from
each sampling site

(2) this includes samples with less than 10 individuals

(3) laboratory reared

19(4): 181-195, 1980(81)

185

electrophoretic investigations of European taxa. A more comprehensive
study, incorporating non-European material, is in preparation.

Materials and Methods

The material used in this study is listed in Table 1 . Samples (only adults
were studied) were usually stored at -30°C until electrophoresis. Control
experiments showed that the electrophoretic mobility of the enzymes is
identical in extracts from deep frozen and fresh specimens. There was
however a considerable loss of GAPDH and ALD activity (for abbreviations
used see Table 2) in frozen butterflies.

Supernatant fractions were prepared for electrophoresis from homo-
genates of an individual’s thorax and head or its abdomen. The enzymes
ME and GPT were assayed in supernatants from abdominal homogenates
and other enzymes in supernatants from homogenates of the thorax and
head. All enzymes were electrophoresed on vertical starch gels using
standard procedures of our laboratory which have been described in detail
elsewhere (Scholl et aL, 1978).

The enzymes assayed are listed in Table 2. A total of 20 enzyme loci has
been scored in all taxa, with the exception of Euchloe tagis bellezina and
Leptidea sinapis, for which only 18 enzyme loci were studied.

All taxa have been compared directly with each other. A precise
characterization of all electrophoretic variants (electromorphs) was obtained
by repeated cross-comparisons, with the exception of some rare variants.
For each enzyme the designation of electromorphs (Tables 3.1 and 3.2)
was made by comparison to the most frequent variant of Pieris brassicae,
which defined the index 100. Other electromorphs were designated by
measuring the difference (in millimeters) in their mobility compared to
that of the reference P. brassicae variant (under our conditions of electro-
phoresis) and then adding or subtracting this value from 100. Thus IDH
103 refers to an IDH-electromorph that migrates 3 mm faster than the
most frequent IDH-variant of P. brassicae.

The correct genetic interpretation of electrophoretically detectable
enzyme phenotypes in polymorphic enzyme systems was proved by
mating experiments (for comments see also Scholl et aL, 1978).

To estimate the genetic similarities between the taxa, the statistic I as
defined by Nei (1972) was used. The similarity values I are presented in
comparisons between pairs of taxa (Table 4) and are a measure of the
proportion of electrophoretically identical proteins.

The similarity values I (Tables 4.1, 4.2, 4.3 and 5) were used to generate
a dendrogram (Fig. 2) according to the unweighted pair) group average
clustering method (Ferguson, 1980).

Results

The different electromorphs found at the various enzyme loci are
presented in Tables 3.1 and 3.2. We have observed considerable

186

J. Res. Lepid.

F ioure 1
GOT-1 Zymogram

Intra- and interspecific variability of the enzyme pattern

©

Species

Genotype

Euchlae crameri

112/118

"

103/112

Euchloe simplonia

103/103

"

Pontia daplidice

35/B5

Anthocharis cardamines.

93/93

/ "

Aporia crataegi

/

9,8/98

Pieris brassicae

lOO/lOC

Artogeia napi napi

60/90

"

90/90

I.

»

»

„

GOT -1 heterczygote individuals show a 3-banded phenotype

polymorphism at some loci and this polymorphism was found in most taxa
where a large number of individuals have been investigated. At each locus
only the electromorphs found at the higest frequency are given in Tables
3.1 and 3.2. The rare variants which are not given do not affect the
conclusions of the present work. Enzymes which exhibit a low degree of
interspecific variation, i, e. which migrate the same distance in the majority
of species, are presented in Table 3.1. Enzymes which have greater
interspecific variability are shown in Table 3.2. Since the designation of
the electromorphs is based on the electrophoretic mobility of the enzymes,
it can be seen from Table 3.1 and 3.2 that the relationships obtained from
the enzyme pattern agree well with the taxonomic classification obtained
with classical systematic methods. Electrophoresis patterns are more
similar between congeneric species than between species of different
genera. Electrophoretic identity can only be found in some isolated cases
between species of different subfamilies. It is of interest that three
enzymes (AK-1, AK-2 and GAPDH) are invariant in all taxa investigated.

This correlation between genetic similarity, assessed from enzyme
variants and taxonomic classification becomes clearer if the comparison is
based on coefficients of genetic similarity rather than the enzyme pattern.
These coefficients are listed in Tables 4.1, 4.2 and 4.3 as comparisons
between pairs of the taxa of the subfamilies Pierinae, Anthocharinae and
Coliadinae, respectively. The subfamily Dismorphiinae is not listed, since
only one species {Leptidea sinapis) was studied.

19(4): 181-195, 1980(81)

187

Table 3.1 : Llectrp'norphs found a t hi qhest frequency in each species.

Enzymes with low interspecif ic variation

Taxon

AK-1

AK-2

OAPDH

APK

■^DH-1

ald

IPG

KE

HK

OGT-2

A

crataegi

100

100

100

94

100

100

102

100

106

93

P

brassicae

100

100

100

100

100

100

100

100

100

100

P.

che 1 ranthi

100

100

100

100

100

100

110

100

100

100

A .

rapae

100

100

100

100

100

100

102

98

100

100

A .

ma n n i

100

100

100

100

100

100

102

98

100

100

A .

n . napi

100

100

100

100

100

100

102

103

lie

100

A .

n. br^oniae

100

100

100

100

100

100

102

103

110

100

P.

daplidice

100

100

100

94

100

100

96

104

100

100

P.

callidice

100

100

100

94

89

100

96

100

90

100

E.

simplonia

100

100

100

100

100

100

112

100

98

100

E.

crameri

100

100

100

100

100

100

112

100

98

100

E .

tagis bel .

100

100

94

100

-

112

99

103

100

A.

cardamines

100

100

100

94

89

100

112

100

104

98

A.

euphenoides

100

100

100

100

89

100

112

103

104

98

C.

phicomone

100

100

100

94

100

100

103

103

123

95

c.

palaeno

100

100

100

94

100

100

103

103

116

95

c.

myrmidone

100

100

100

94

100

100

103

103

116

93

c.

crocea

100

100

100

94

lOD

100

103

99

116

97

c.

hyala

100

100

100

94

flDO

Uio

100

103

99

123

95

c.

australis

100

100

100

94

100

100

103

99

116

95

Cal

florella

100

100

100

94

100

100

108

95

112

90

G.

r hamni

100

100

100

94

89

85

103

99

106

97

G.

Cleopatra

100

100

100

94

89

85

103

99

106

92

L.

sinapis

100

100

100

100

92

-

97

106

65

The averages of similarity coefficients and their standard deviations
found between the taxa of the different subfamilies are presented in Table
5 (as indicated in the method section, the calculation of these coefficients
is based on the allele frequencies observed, including the minor alleles at
polymorphic loci, which are not shown in Tables 3.1 and 3.2).

High similarity coefficients were obtained between taxa which are
regarded as closely related from morphological and other data (e.g.
between the members of the genus Colias). These taxa clearly show less
similarity to more distantly related taxa (e.g. comparison of the Colias and
Gonepteryx species). The lowest similarity coefficients are found between
members of the different subfamilies.

The similarity coefficients between the Pierinae and Anthocharinae are
clearly higher than between these two subfamilies and the members of the
Coliadinae and Leptidea sinapis, the only species of the Dismorphiinae

188

J. Res. Lepid.

Table 3.2 : E 1 e c t romor p h s found at highest frequency in each species.

Enzymes with high interspecific variation

Taxon

MDH-2

6-PGD

FUM

IDH-1

IDH-2

PK

GOT-1

GPT

PGI

PGM

A, crataegi

98

1 100

105

93

1 90
[lOO

112

98

102

1 90

1 96

(111

1115

P. brassicae

100

100

100

100

100

100

100

100

100

100

P . cheiranthi

100

93

100

100

100

100

100

100

100

95

A . rapae

90

107

98

95

110

106

88

100

107

100

A . manni

90

100

98

95

100

106

88

IDO

100

100

A . n . napi

90

100

90

98

100

100

90

115

100

102

A. n. bryoniae

90

100

90

98

100

100

90

115

100

102

P . daplidice

90

100

95

86

97

103

93

130

97

110

P . cal lidice

90

89

95

91

97

106

130

81

109

108

E. simplonia

102

87

104

78

90

103

103

126

108

90

E . crameri

102

93

104

78

90

103

112

121

108

100

E. tagis bel.

102

87

105

76

90

103

101

94

97

92

A . cardamines

91

92

105

84

97

103

93

118

91

85

A. euphenoides

91

103

105

84

97

103

93

104

82

1 92
1100

C . phicomone

100

79

106

88

102

102

105

61

90

109

C . palaeno

100

79

106

88

102

102

105

68

80

102

C . myr midone

100

79

106

88

102

100

105

96

96

109

C. crocea

100

79

106

88

102

100

105

68

107

109

C . hyal e

95

79

93

88

102

95

105

81

98

109

C . australis

100

79

106

88

102

95

105

81

98

(102

[109

Cat. floralla

92

79

106

88

108

92

105

81

96

109

G. rhamni

92

79

110

97

142

95

115

89

80

115

G . Cleopatra

92

79

110

1

1 97

140

95

122

91

80

120

L . sinapis

97

96

97

90

120

115

113

113

114

129

investigated. Leptidea sinapis has the lowest similarity coefficients with all
other taxa.

A dendrogram (Fig. 2) has been constructed from the similarity
coefficients (Tables 4.1, 4.2, 4.3 and 5), computed from the primary data
(Tables 3.1 and 3.2). It showss, more clearly, the grouping of the taxa
based on the biochemical-genetic data. The four subfamilies can be
recognized as four branchings, however it is remarkable that the Antho-
charinae branch from the Pierinae. Branch points of the genera are usually
above the level of subfamilies (between 0.36 and 0.59). Colias and
Gonepteryx species branch at a higher level of genetic similarity than
nongeneric species of the subfamilies Pierinae and Anthocharinae.
Discussion

It has become evident from numerous investigations in recent years that
electrophoretic techniques provide a very valuable tool for systematisists.

19(4): 18M95, 1980(81)

189

Table 4.1 ; Coefficients of genetic similarity (l-values)

between Pierinae-speciea .

m

•H

m

m

X

•H

S3

m

C

U

y

C

O

•H

m

a

>1

T3

■D

03

m

CO

U

■H

m

m

c

c

n

<-4

£0

a

c

a

M

m

m

m

m

u

u

B

c

c

*0

u

CL

<

<

<

Q.

a.

A. crataegi

.35

.32

.33

.35

.34

.34

.37

.32

P. brassicae

.88

.53

.64

.56

.57

.42

.32

P. cheiranthi

.52

.58

.51

.51

.37

.32

A. rapae

.90

.51

.56

= 43

.38

h, raanni

.60

.60

.4?

.30

k. n. napi

1.0

.42

.32

A. R. bryoniae

.42

.32

P. daplidicB

.55

L. crameri

t. tagis bell.

A. cardamines

A. euphenoides

E, simpionia

.83

.52

.36

.37

E. cramori

.46

.37

.39

E. tagis bell.

.40

.39

A. cardamines

.74

The application of these techniques in establishing genetic relatedness,
however, appears to be restricted mainly to congeneric species (Ayala et
at, 1974; Avise, 1974; Selander and Johnson, 1973), since the range of
similarity detectable by enzyme-eiecta’ophoresis rapidly decreases beyond
die generic level and reaches the critical point where two species have no
electrophoretically identical enzymes in common. It is of interest therefore
that in the Pieridae it has been possible to clearly establish genetic
affinities even in interspecific comparisons of members of different
subfamilies. It should be emphasized, however, that the sample of
enzymes investigated has some influence on the similarity coefficients,

190

J. Res. Lepid.

since rates of divergence are quite different for individual enzymes. It is
possible that in this study a rather conservative sample of enzymes has
been investigated.

Three enzymes, AK-1, AK-2 and GAPDH, were almost invariant in all
Pierid species investigated and were the main, though not exclusive
contributors, to the level of genetic identity found in inter-subfamilial
comparisons. These three enzymes may prove very valuable for enzymatic
characterization of butterfly taxa beyond the generic level, though further
work will be necessary to establish this point.

Table 4.3 : I - values between Col iadinae-spec ie s

C. palaeno europ

C. myrmidone

C . crocea

C . hy a 1 e

C. australis

Cat. florella

G. rhamni

G. Cleopatra

C. phicomone

.65

.79

.71

.76

.83

.62

.35

.34

C. palaeno europ.

.79

.77

.63

.79

.53

.35

.35

C. myrmidone

.62

.63

.79

.59

.34

.34

C. crocea

.64

.80

.54

.40

.39

C . hy al e

.82

.56

.45

.44

C. australis

.64

.44

. 64

Cat. florella

.33

.33

G. rhamni

.78

19(4): 181-195, 1980(81)

191

PIERIS BRASSICAE
PIERIS CHEIRANTHl
ARTOGEIA RAPAE
ART06EIA MANNI
ARTOGEIA NAPl NAPI
’ ARTOGEIA NAPI BRYONIAE
PONTIA DAPLIDICE
PONTIA CALLIDICE
APORIA CRATAEGI
EUCHLOE SIMPLONIA
EUCHLOE CRAMERI
EUCHLOE TAGIS BELLEZINA
ANTHOCHARIS CARDAMINES
ANTHOCHARIS EUPHENOIDES
COLIAS PHICOMONE
COLIAS PALAENO EUROPOME
COLIAS MYRMIDONE
COLIAS CROCEA
COLIAS AUSTRALIS
COLIAS HYALE
CATOPSILIA FLORELLA
GONEPTERYX RHAMNI
GONEPTERYX CLEOPATRA
LEPTIDEA SINAPIS

1. The genetic divergence of the subfamilies
The subfamilies investigated show different degrees of genetic diver-
gence (Table 5). Leptidea sinapis, the only Dismorphiinae in this study, is
clearly isolated by the biochemical-genetic criteria as it has the lowest
similarity coefficients. The genetic similarity between the Pierinae and
Anthocharinae is greater than between these two subfamilies and the
Coliadinae. The Anthocharinae may be characterized biochemically as a
unit, but their branching point from the Pierini lies at the same level as
those between different genera of the Pierini or that between the genera

192

J. Res. Lepid.

Gonepteryx and Colias (Fig. 2). Therefore this biochemical data support
the decision of Ehrlich (1958) and others to group the Anthocharini and
Pierini as tribes of the Pierinae.

The somewhat closer electrophoretic similarity of Aporia crataegi to the
Anthocharinae than to the Pierinae is probably not significant, but
supports the relatively close biochemical-genetic relationship between the
two subfamilies.

The isolated position of Leptidea sinapis results from the observation
that this species has common electromorphs with other Pieridae only at
the AK-1, AK-2, APK and MDH-1 loci which are all highly conservative,

1. e. have a low number of electromorphs in the material investigated (see
Table 3.1). Lorkovic recently expressed some doubts, based on morpho-
logical observations, as to whether Leptidea is really a Pierid (personal
communication). It would be of interest to investigate whether the low
degree of genetic similarity of Leptidea sinapis is typical for other taxa of
the subfamily Dismorphiinae or the genus Leptidea. At the same time,
however, the genetic divergence of species of other butterfly families from
the Pieridae should be established and compared with the value found for
Leptidea sinapis.

2. The position of Catopsilia florella F. 1775

There is a remarkable high genetic similarity between Catopsilia florella
and species of the genus Colias. This similarity is clearly higher than that
between species of Colias and Gonepteryx. Higgins (1975) classified
Catopsilia florella as a Pierinae species. The biochemical genetic data
support the opinion of most other authors (e.g. Klots, 1931/32), that
Catopsilia florella should be included together with the species of the
genera Colias and Gonepteryx into the subfamily Coliadinae.

3. The division of the genus Pieris Schrank 1801

Verity (1947) and Kudma (1974) maintain that the genus Pirns Schrank
1801 should be subdivided into two genera, Pieris and Artogeia Verity
1947 respectively. The taxa brassicae and cheiranthi, which were thus
isolated in the “new genus Pieris"', show the same biochemical-genetic
similarity with taxa of the “new genus Artogeia" as is found between
Artogeia taxa themselves. Therefore, the electrophoretic data do not
support the proposal that the old genus Pieris should be split in two.

Current investigations of additional taxa of this group may clarify the
phylogenetic relationships.

4. The relationships between the taxa brassicae L. and cheiranthi
Hbn. and between the taxa simplonia F. and crameri Btlr.

Unlike sympatric species where reproductive isolation is directly
demonstrable, the relationships of allopatric populations is often difficult
to determine. Pieris brassicae L. and its isolate P cheiranthi Hbn. in the
Canary Islands, and Euchloe crameri Butler of the Mediterranean region

19(4): 181-195, 1980(81)

193

and its subalpine relative E. simplonia Freyer are examples of such cases.
P. cheiranthi was originally described as a new species, but has since
been regarded by many authors as a subspecies of P. hrassicae (e.g.
Higgins, 1975). Others maintain that they are specifically distinct
(Kudma, 1973; Schurian, 1975). Similarly, jF. simptowa is often considered
as a “glacial” subspecies conspecific with E. crameri, even though many
others have assumed them to be good species on the basis of morphological
differences. In particular, distinctness of their larvae and pupae is well
known (e.g., Catherine, 1920; Verity, 1947; Back, 1979). Neither enzyme
analysis nor morphological studies measure reproductive relationships.
Nevertheless, the magnitude of genetic divergence between related
allopatric forms may provide some insight into such systematic problems
when it is compared with data for the sympatric species pairs. The pairs A.
mpae and A. mannii, A. cardamines and A. euphenoides, G. rhamni and G.
Cleopatra, and those of the genus Colias are at least partially sympatric.
Some are sibling species. Similarity indices for these pairs range from 0.74
to 0.90. Some of these species have wide distribution ranges, but the
genetic divergences in geographically distant populations so far studied
are significantly lower than the range given above; for instance, A. rapae
from Europe and Japan, or A. cardamines from Europe and Japan have the
l value of at least 0.95 (unpublished data). In contrast, the lvalue of 0.88
for E. crameri and E. simplonia, falls within the range of genetic disparity
found for sympatric species pairs. The interspecific genetic similarities of
some sibling species may overlap with intersubspecific similarities in
Drosophila (Ayala et aL, 1974). We have not encountered similar cases in
butterflies, yet future studies may reveal such situations. With this
reservation, therefore, the data suggest specific distinctness of the above
allopatric pairs. Rapid genetic drift and establishment of reproductive
barriers in island populations or experimental stocks founded from small
numbers of individuals have been demonstrated in Drosophila (e.g. Carson,
1968; Templeton, 1980). P. cheiranthi may represent such an example. It
would have been extremely interesting if the genetic composition of the
newly founded colony of P. brassicae in Chile (Gardiner, 1974) were
followed from the beginning and compared with P. cheiranthi and
European P. brassicae.

5. The napi - bryoniae problem

The taxa napi L. and bryoniae Hbn. have been regarded as: different
species (Mueller and Kautz, 1938; Forster and Wohlfahrt, 1976), as
semispecies (Lorkovic, 1962) or as subspecies (Higgins, 1975). Bowden
(1972) describes the holarctic Pieris napi ~ bryoniae complex as a “perfect
example of a superspecies”. The present results on 20 enzyme loci
support previous observations (Geiger, 1978) that biochemical- genetic
differences between these two taxa are extremely low (lvalues between,
population samples of at least 0.992). The same electromorphs occur in

194

J. Res. Lepid.

both taxa, when very infrequent electromorphs are disregarded. The
previous observations (Geiger, 1978) that differences between napi
population samples are sometimes greater than differences between napi
and (Swiss) bryoniae, has been substantiated here with more animals. A
specific report on napi and bryoniae and on some closely related taxa is in
preparation.

Acknowledgments. I am greatly indepted to PD Dr. A. Scholl, Zoological
Institute of the University of Bern, for his generous support of the present work and
for critically reading this manuscript. Comments and suggestions of Dr. I.
Nakamura, New York, considerably helped to improve this paper. I am also grateful
to Dr. Ch. Webb (University of Bristol, England) and Drs. F. and B. Westley for
reading the English versions of the manuscript and to Mrs. L. Frauchiger and Mrs.
V. Siegried (all of the Zoological Institute of the University of Bern) for assistance in
the electrophoretic investigations. This work would not have been possible without
the help of the following persons that sent me living material: Mr. R. Adams,
F.R.E.S., Canai*y Islands (P. Cheiranthi, C. florella)\ Mr. S. R. Bowden, St. Albans,
England {A. napi)\ Mr. G. Flauger, Teublitz, Gennany (C. australis, C. hyale, C.
myrmidone) and Dr. L. Rezbanyai, Museum of Nature Lucerne, Switzerland (A. n.
bryoniae).

Literature Cited

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variability in the Drosophila willistoni group. IV. Genic variation in natural
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AYALA, F. J., M. L. TOACEY, D. HEDGECOOCK & R. C. RICHMOND, 1974. Genetic differentia-
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BACK, w., 1979. Zur Biologie der europaischen und nordwest-afrikanischen
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BREWER, G. J., 1970. An introduction to isozyme techniques. Academic Press, New
York and London.

BOWDEN, S. R., 1972. Pieris napi L. (Pieridae) and the superspecies concept. J.
Lepid. Soc. 26: 170-173.

., 1979. Subspecific variation in butterflies: Adaptation and dissected

polymorphism in Pirn’s (Artogeia) (Pieridae). J. Lepid. Soc. 33: 77-111.
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CATHERINE, G., 1920. Notes sui Anthocharis belia Cr. et ausonia Hb. du departe-
uient du I’Aube. In Oberthuer, C., Etud. Lepid. Comp. 17, pp. 48-53.
Corrigenda by Oberthuer, id. 17 (Planches), p. 46.

EHRLICH, P. R, 1958. The comparative morphology, phylogeny and higher classifi-
cation of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull.
39: 305-370.

FORSTER, w. & T. WOHLFAHRT, 1976. Die Schmetterlinge Mitteleuropas, 2. Tagfalter.
Franckh, Stuttgart.

FORD, E. B., 1945. Butterflies. London. Collins.

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195

FERGUSON, A., 1980. Biochemical Systematics and Evolution. Blackie, Glasgow
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GARDINER, B. 0. C., 1974. Pieris brassicae L. established in Chile; another palearctic
pest crosses the atlantic (Pieridae). J. Lepid. Soc. 28(3): 269-277.

GEIGER, H. J., 1978. Die systematische Stellung von Pieris napi bryoniae: Bio-
chemisch-genetische Untersuchungsbefunde (Lep.: Pieridae). Ent. Z. 88:
229-235.

HARRIS, H. & D. A. HOPKINSON, 1976. Handbook of enzjnme electrophoresis in human
genetics. North-Holland, Amsterdam and Oxford.

HIGGINS, L. G., 1975. The classification of european butterflies. Collins, London and
Glasgow.

KLOTS, A. B., 1931/32. A generic revision of the Pieridae (Lepidoptera). Ent. Am.
12 (n.s.) (3-4): 139-242.

KOYLER, J. M., 1966. The effect of certain environmental factors and chemicals on the
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KUDRNA, o., 1973. On the status of Pieris cheiranthi Huebner (Lep., Pieridae). Ent.
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, 1974. Artogeia Verity, 1947, Gen. rev. for Papilio napi Linnaeus (Lep.

Pieridae). Ent. Gazette 25: 9-12.

LORKOVIC, z., 1962. The genetics and reproductive isolating mechanism of the Pieris
napi bryoniae group. J. Lepid. Soc. 16: 5-19, 105-127.

MUELLER, L. & H. KAUTZ, 1939. Pieris bryoniae O. und Pieris napi L. Osterr. ent. Ver.
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NEL M., 1972. Genetic distance between populations. Amer. Natur., 106: 283-292.
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5: Die Amerikanischen Tagfalter. Kemen, Stuttgart.

SCHOLL, A., B. CORZILLIUS & w. viLLWOCK, 1978. Beitrag zur Verwandtschaftsanalyse
altweltlicher Zahnkarpfen der Tribus Aphaniini (Pices, Cyprinodontidae) mit
Hilfe elektrophoretischer Untersuchungsmethoden. Z. zool. Syst. Evolut.-
forsch. 16: 116-132.

SCHURIAN, K., 1975. Bemerkungen uber Pieris cheiranthi (Lep., Pieridae). Ent. Z.
85: 252-256.

SELANDER, R. K. & w. E. JOHNSON, 1973. Genetic variation among vertebrate species.

Annual Review of Ecology and Scstematics 4: 75-91.

SHAPIRO, A. M., 1975. Photoperiodic control of development and phenotype in a sub-
arctic population of Pieris occidentalis (Lepidoptera: Pieridae). Can. Ent.
107: 775-779.

, 1977. Phenotypic induction in Pieris napi L.: role of temperature and

photoperiod in a coastal California population. Ecological Entomology 2:
217-224.

TEMPLETON, A. R., 1980. Modes of speciation and inferences based on genetic dis-
tances. Evolution 34(4): 719-729.

VARGA, z. & I. TOTH, 1978. Obersicht der taxonomischen und chorologischen
Verhaltnisse in der Pieris (Artogeia) napi-bryordae-Gruppe (Lep.: Pieridae)
mit Ruecksicht auf die Anwendung des Superspezies-Begriffes in der Lepid-
opteren-Taxonomie. Acta biol. Debrecina, 15; 297-322.

VERITY, R., 1947. Le farfalle diume d’ltalia. Vol. EH Firenze.

Journal of Research on the Lepidoptera

19(4): 196-198, 1980(81)

A New Tortyra from Cocos Island, Costa Rica
(Lepidoptera: Choreutidae)

John B. Heppner

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560

The following new species of Tortyra is described here to further
document the unique fauna of Cocos Island, off the western coast of Costa
Rica. Unlike most of the oceanic islands near the west coast of Central
America and northern South America, Cocos Island has a wet tropical
forest. The new Tortyra is almost certainly an endemic of the island and is
here named in honor of Dr. C. L. Hogue of the Los Angeles County
Museum of Natural History, who collected the type series.

Tortyra hoguella Heppner, new species

Size. Forewing length 5. 2-6. 5 mm.

Head. Dark fuscous with white between antennal bases; frons dark
fuscous; labial palpus white with dark fuscous apical segment; antenna
purple iridescent with white area near apex.

Thorax. Lustrous dark grey; patagia fuscous irrorated with white; venter
white; legs banded with dai’k fuscous. Forewing: basal Vs fuscous irrorated
with white; mid Vs of wing with a broad brown band, with a silver and green
iridescent line along basal part of band and a broad white border, distally
irregular; apical Vs of wing fuscous with broad purple iridescent area from
tornus almost to apex; ventral side lustrous bronze fuscous; fringe lustrous
fuscous. Hindwing: uniformly bronze fuscous; ventral side lustrous
bronze fuscous; fringe lustrous fuscous.

Abdomen. Dark fuscous; venter white. Male genitalia: uncus and
gnathos absent; tegumen narrow, broadening dorsally toward extended
anal tube resembling a socius; vinculum strong, rounded; anellus triangular;
valva oblong, narrowing to sharp extended apical thorn-like point;
extensive inwardly directed setal field from dorsal valval margin; aedeagus
with phallobase; comutus a large flat spatula-like shape with a dentaceous
apical edge. Female genitalia: papilla anales setaceous; posterior
apophyses long and slender; anterior apophyses short and very stout;
ostium a simple membranous opening on intersegmental membrane
posterior to rounded posterior edge of 7th sternite; ductus bursae
membranous, long and spiralled, with spirals twisted around bursal
appendage near bursa; ductus seminalis from near ostium; bursa copula-
trix ovate with elongate appendage; signum a simple spicule band on one
side of bursa.

19(4): 196-198, 1980(81)

197

Fig. 1.

Fig. 2.
Fig. 3.

Tortyra hoguella Heppner, new species, genitalia (holotype, JBH slide
1383).

Same, aedeagus (enlarged).

Anellus (enlarged) (paratype, JBH slide 1541).

Fig. 4. Tortyra hoguella Heppner, new species, 9 genitalia (paratype, JBH slide
1555), bursa.

198

J. Res. Lepid.

Fig. 5. Tortyra hoguella Heppner, new species (cf paratype, Cocos Id.).

Holotype cf. Wafer Bay, Cocos Id., 17-22 IV 75, C. L. Hogue (LACM).
Paratypes. Costa Rica: Cocos Id., 17-22 IV 75, C. L. Hogue (5
LACM); Rio Genio, Cocos Id., 17-22 IV 75, C. L. Hogue (1 9, LACM).
(Paratypes to BMNH and USNM].^

Host. Unknown but probably a species of Ficus, as is usual for other
Tortyra species.

Remarks. Tortyra hoguella has one of the most unusual wing patterns in
the genus due to the prominent white forewing fascia. The male genitalia
are similar to several species of Tortyra, with some similarity to Tortyra
spectabilis (Walker) from Brazil, although the wing maculation is unique.
The female genitalia have the ductus bursae spiralled and twisted around
a bursal appendage as is typical for the genus and only this portion is
illustrated. Most female Tortyra genitalia are almost identical.

‘LACM = Los Angeles County Museum of Natural History, Los Angeles, California; BMNH —
British Musuem (Natural History), London, England; USNM = National Museum of Natural
History, Smithsonian Institution, Washington, D.C.

Journal of Research on the Lepidoptera

19(4): 199^226, 1980(81)

Troidine Swallowtails (Lepidoptera: Papilionidae) in
Southeastern Brazil: Natural History and Foodplant
Relationships

Keith S. Brown, Jr.^^ Antoni J. Damman* and Paul Feeny^

‘Section of Ecology and Systematics, Division of Biological Sciences, Cornell University,
Ithaca, New York, 14853, USA and

^Departamento de Zoologia, Institute de Bioiogia, Universidade Estadual de Campinas, C. P.
1170, Campinas, SSo Paulo, BRAZIL 13.100

Abstract. Five species of troidine swallowtails, Parides proneus, P. bunichus,

P. agavus, P. anchises nephalion, and BattuspolydamaSf sympatric in central
S?o Paulo state, southeastern Brazil, differentially utilize the available
Aristolochia hostplants. All early stages differ morphologically, with the
possible exception of eggs of P. bunichus and P. agavus. In two study sites,
the adult populations fluctuated greatly and disharmoniously in approxi-
mate two-month cycles. Individuals moved an average of near 100 m and up
to 1.3 km between successive recaptures, probably in search of flower
resources. Adult lifespans in the field and laboratory averaged less than two
weeks, with a majority of field-recaptured individuals passing through one of
the six major wing wear classes per week. Laboratory cultures, maintained in
a walk-in growth chamber, were used to investigate larval and adult behavior,
pupal diapause, oviposition, and hostplant palatability in the five species of
butterflies, with relation to the six potential hostplants sympatric in the field.

Introduction

Papilionid butterflies in the tribe Troidini have long been known to feed
specifically on plants in the genus Am to/oc/iia, and are commonly referred
to as Aristolochia swallowtails. (Roths child & Jordan, 1906; Munroe, 1960;
Ehrlich & Raven, 1964). Although homopterans, tingid bugs, cassidine
beetles, and larvae of noctuid and pyralid moths have been observed on
Aristolochia, relatively few herbivores other than the Aristolochia swallow-
tails can feed on these plants (D’ Araujo e Silva et aL, 1968; Rausher,
1979a; K. B., pers. obs.). Aristolochia vines characteristically contain
aristolochic acids (nitrophenanthrenes) that have pharmacological effects
on vertebrates (Hoehne, 1942: 14-17; von Euw et al., 1968) and strongly
inhibit feeding by some insects (Bernays & Chapman, 1977; Rausher,
1979a). The plants also contain several benzylisoquinoline alkaloids,
sesquiteipenes, and other secondary components (Fraenkel, 1959;
Hegnauer, 1964).

Aristolochia swallowtails sequester some of these secondary chemicals

from their hostplants, apparently making the butterflies unpalatable to

200

J. Res. Lepid.

many potential predators (von Euw et al., 1968; Rothschild et aL, 1970;
Rothschild, 1972; R. Nishida, pers. comm.). The butterflies advertise
their distastefulness through aposematic coloration, and serve as models
in Batesian and Muellerian mimicry complexes (Rothschild & Jordan,
1906; Brower, 1958; Brower & Brower, 1964; Young, 1971a, 1971b).

Because the biology of the troidine swallowtails is linked so closely to
that of their Aristolochia hostplants, a thorough study of this plant-insect
association could provide insights into the evolution of plant defenses, and
into the use of plant chemicals by insects. The basic biology of the system
has received limited attention until recently. Several studies of the natural
history of various North and Central American troidines have appeared in
the past decade (Young, 1971a, 1971b, 1972a, 1972b, 1973, 1977, 1979;
Muyshondt, 1974; Rausher, 1979a, 1979b; de Vries, 1979). Rausher
(197 9a) studied the interaction of Battus philenor and its two host plants in
southeastern Texas. The South American members of the tribe have been
studied by Moss (1919), D’ Almeida (1922, 1944, 1966), and Cook et al.
(1971).

In this paper we describe an assemblage of five troidine butterflies
sympatric in central Sao Paulo state {Battus polydamas, Parides proneus,
P. bunichus, P. agavus, P. anchises nephalion^), emphasizing the interaction
with their Aristolochia hostplants. The natural histories and laboratory
culture techniques presented here provide the foundation for planned
field and laboratory studies of the biological and chemical aspects of this
Troidini-Aristolochia interaction.

Study Sites and Methods

Wild troidine populations were observed on the grounds of the Horto
Florestal de Sumare, located in the interior of the state of Sao Paulo, Brazil
(22°5TS., 47°16'W., 600 m elevation) (Figure 1, A and B). Between
October and March (summer), temperatures at Sumare average 23°C; the
April to September (winter) average is 18.5°C and daily temperature
ranges often exceed lO'^C. The bulk of the average annual rainfall of 1300
mm falls during the summer. Because Sumarg lies at the intersection of
three major climatic domains, the distribution of the annual precipitation
is highly variable from year to year (Figure 2).

The Horto Florestal (856 ha, of which 50 were intensively covered in the
field work) covers a mosaic of poor, sandy soils and rich soils, originally
supporting cerrado scrub and semi-deciduous open forest, respectively.
The troidines inhabit riverine tangles and the shady middle and under-
story beneath an introduced Eucalyptus canopy of the semi- deciduous
forest (Figure 3). Three species of Aristolochia, A. melastoma, A. arcuata,

^The placement of nephalion and other “species” of Rothschild and Jordan (1906) as subspecies
of anchises, and the species distributions indicated in Figure 5, follow a biosystematic revision
and biogeographical study to be published by K. Brown.

19(4): 199-226, 1980(81)

201

iiiJiiiiWi iiii

iiliMisiiilii

Hiaii!|i«S

Fig. 1 . Map showing study sites for Troidini/ Aristolochia interaction in the region
of Campinas and Sumare, Sao Paulo. Small numbers in detailed maps B
and C indicate subareas for marking of adults (see Fig. 10).

HfiRTO FLOlllSTArDE SOMARE AMARAIS roZENDA SANTA ELISA MONJOLiNHO

202

J. Res. Lepid.

RAINFALL *^EAN TE^ERATURE

Fig. 2. Monthly and yearly fluctuations in rainfall (mm, scale at left) and
temperature (°C, scale at right) in the Horto Florestal de Sumar^ Sao
Paulo Data for 1974-1978 are from Vasconcellos-Neto (1980). Monthly
rainfall data for 1972-1973 are from nearby Nova Odessa (see Fig. 1), other
years are from within the Horto Florestal. Mean monthly temperature is
corrected for altitude difference from data measured in Nova Odessa.
Black areas are very humid months; dotted areas, very dry. The maximum
yearly rainfall (1887 mm in 1976) immediately followed the minimum
yearly rainfall for this decade (1003 in 1975).

19(4): 199-226, 1980(81)

203

Fig. 3. Middle and understory of the Eucalyptus semi-deciduous forest in the
(upper) Horto Florestal de SumarC preferred habitat of the troidines and their

Aristolochia foodplants. Photo at right by J. Vasconcellos-Neto.

Fig. 4. Aristotochia foodplants of troidines, common in central Sao Paulo, a (left),
(lower) A. melastoma; upper, leaves; lower, flower, b (upper center), A. arcuata,
leaf and flower, c (right), A. brasiliensis; upper, leaves; lower, flower, d
(lower center), A. littoralis, flowers, leaves and fruit, from Hoehne (1942).

204

J. Res. Lepid.

and A. brasiliensis (Figure 4), commonly grow in light gaps, along forest
edges, and in mildly disturbed areas within the forest. Two additional
species, A. littoralis (Figure 4d) and A. triangularis, occur infrequently in
the Horto. A. gigantea, cultivated nearby, represents a sixth potential
foodplant for the troidine larvae.

An additional study area near Campinas, SSo Paulo, is the Fazenda
Santa Elisa of the Instituto Agronomico de Campinas. A large troidine
concentration occurs in a Eucalyptus woodlot similar to the Horto Florestal:
Amarais, with abundant A. melastoma and A. arcuata (Figure 1, C). One
km east of Amarais is the 3-ha forest garden Monjolinho (Figure 1 , D), with
a cool, humid understory beneath native trees; here there are patches of A.
esperanzae (near A. brasiliensis) and A. littoralis. A fourth site, visited only
occasionally, is a 250-ha woods north of Campinas (Figure 1, E) with small
numbers of A. melastoma, A. arcuata, and A. drasi/icnsis plants in the more
open understory and along humid edges of a natural semi-deciduous forest
on rich soils. All of the study sites are in the basin of the upper RibeiriTo do
Quilombo (Figure 1).

The five troidines well established at Sumar^ and Campinas have geo-
graphic ranges that differ widely (Figure 5^). P. bunichus and P. proneus
extend into the cool montane and central plateau regions of eastern and
southern Brazil, with the former much more widespread. P. agavus occurs
most abundantly in the mild coastal regions of eastern and southern
Brazil, but also extends far southward and inland. P. anchises and Battus
polydamas are widespread and often prefer warm, dryer climates. At least
six additional troidines are known from central S^o Paulo, including
Parides neophilus eurybates which is occasionally encountered in the study
sites. Observations throughout the Neotropics suggest that the occurrence
of but five or six troidines out of a potential pool of 1 1 to 16 species at a
particular site is a general phenomenon (Moss, 1919; D’Almeida, 1966; K.
B., pers. obs.).

Between November 1978 and September 1979, and again after August
1980, one of us (K.B.) spent two to ten hours each week conducting mark-
recapture studies of the adults in the Horto Florestal, and observing adult
behavior, resource utilization, and oviposition as well as searching for
larvae on the foodplants. Parallel studies were undertaken in the two
Santa Elisa sites starting in the later part of 1980. Captured adults were
sexed, assigned to one of six wing wear classes, numbered on the underside
of each hindwing (P bunichus, P proneus, P. agavus) or forewing (P
anchises, P. neophilus, Battus polydamas) with an indelible marking pen
(“Sharpie”, Sanford Corp.), and immediately released (usually within 30
sec). Butterflies were often captured two to six times within a single
sampling session (Table 1) and were invariably placed in the same wing
wear class all times, attesting to the reproducibility of the wing wear
assignments. Rapid return of marked butterflies to pre-capture activities

19(4): 199-226. 1980(81)

205

FIGURE 5

Geographical Distributions

PARIDES PRONEUS *3^

AMERICA TROPICAL

K§ - ZOQlOGiA - UNICAMP 1171

PARIDES BUNICHUS
PARIDES AGAVUS

PARDES ANCHISES^

\ outer lirbjts

( ) BATTUS POLYDAMAS)

TOTAL NUMBER MARKED

206

J. Res. Lepid.

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19(4): 199-226, 1980(81)

207

was routinely observed, suggesting relative unimportance of marking
trauma.

These studies showed that the Horto Florestal harbored a 6-ha central
concentration of troidines {A in Figure 1, A and B) as well as several
smaller peripheral concentrations (B-G). Amarais had a single colony
covering 25 ha, half of which was heavily overgrown in foodplants (Figure
1, C). Monjolinho showed rather sparse troidines in a few favored light-
gaps (Figure 1, D). Populations of troidines were very sparse in the large
natural woodlot north of Campinas (Figure 1, E). Troidine concentrations
are infrequent within a 20-km radius of the Horto Florestal, the majority of
which is dominated by fields, orange groves, and heavily populated area
(Figure 1).

Laboratory cultures of butterflies were maintained in a walk-in growth
chamber (Figure 6 ab) at Cornell University; stock cultures were also kept
in a greenhouse (Figure 6 cd). The environment in the growth chamber
approximated summer conditions in the middle and lower levels of the
forest at Sumar^. A 13.5 hr “day” period faded into a 10 hr “night” period
through programmed dimming of the lights over a 0.5 hr “dusk” period.
Temperature alternated between a constant 26°C during the day and a
constant 2 1°C at night. Relative humidity was kept between 85% and 90%
throughout the daily cycle, but because of the constant air flow the
atmosphere was a drying one. A diffuse, fine-grained, and relatively low-
intensity light structure, similar to that of the Horto Florestal middle level
(Figure 3), and a dark humid understory, were necessary components for
the adaptation of the butterflies in both the growth chamber and the
greenhouse (Figure 6). Butterflies visited cut or potted flowers as well as
artificial nectar sources (consisting of an orange or yellow scouring sponge
in a plastic petri dish filled with 1:6 honey: water solution). The butterflies
often preferred to rest and sleep on dacron net curtains hung in the culture
areas. Juvenile and adult troidines did not survive as well in the
greenhouse as in the growth chamber, perhaps because of daily and
seasonal fluctuations in temperature, and predation by ants. In both
cultures, however, normal behavior of adults was observed (Figure 6 d-g).
Individual larvae could be followed from day to day by marks on their
tubercles (fourth and fifth instars) made with various colors of Liquid
Paper correction fluid (Liquid Paper Corp.).

Description of Juvenile Stages

The adult stages of most troidines are well known (Rothschild & Jordan,
1906; D'Almeida, 1966); those occurring in the study sites are illustrated
in Figure 7. The juvenile stages are less well known, though larvae and
pupae of most species have been illustrated or preserved. Variations in the
larval characteristics of Parides anchises, P. aeneas andP. lysander (Moss,
1919; Young, 1977; inspection of larvae reared at Cornell University or in
the field or preserved in the British Museum (Natural History) by K. B.)

208

J. Res. Lepid.

Fig. 6. Laboratory cultures of troidines in Cornell University (except for f). a-b,
views of the growth chamber; in a, a P. anchises male is resting on a leaf at
upper left, and two P bunichus are on flowers at the rear, by plastic boxes
used for pupal experiments, c-d, views in the greenhouse; in c, aP. bunichus
is flying at the rear, and in d, another one is approaching an Impatiens
flower. Note the light structure (diffuse in middlestory, dark in understory)
in a-c, and Dacron net curtains in b-d. e, aP bunichus and aP proneus on
Lantana flowers in the growth chamber, f , a courtship of P. bunichus (near
Campinas, S^o Paulo; photo by 1. Sazma); note the exposed white scent
pouch of the male hindwing over the female antennae, g, a copula of P
bunichus in the greenhouse; both male and female had eclosed from the
pupa earlier on the same day.

19(4): 199-226, 1980(81)

209

Fig. 7. Adults of troidines captured in the Horto Florestal de Sumar^. a, Battus
polydamas, male; b, Parides proneus, female; c, P. bunichuss, male; d,
P. agavus, female; e, P. anchises nephalion, male (upper) and female
(lower); f, P. neophilus eurybates, mafe (upper) and female (lower). Note
the white or black androconial scales in the male Parides hindwing anal
folds (c, and e-f upper).

210

J. Res. Lepid.

Fig. 8. Juvenile stages of the five resident troidines in the central SSo Paulo study
sites. Eggs (at left) 8.5x (except for f, egg position under leaf, 1.7x; and n,
egg cluster, 3.4x); first instars (left center) 7x (except for o, group of larvae,
2.3x); fifth instars (right), l.lx; pupae (lower), 0.75x. P. bunichus, a-d, left
center pupae; P. anchises nephalion, e-h, center pupae; P. proneus, i-k, right
pupae; P. agavus, 1-m, right center pupae; B. polydamas, n-p, left pupae.
Pupae, q = dorsal, r = dorsolateral, s = ventrolateral.

19(4): 199-226, 1980(81)

211

suggest that larval color patterns may not be geographically homogeneous.
Color patterns in B. polydamas larvae vary within a population (Moss,
1919). While it is important to briefly describe and distinguish between
juvenile stages encountered in central SSo Paulo, the following larval color
descriptions should be extended to geographically distant populations
with caution.

Measurements are on organisms from laboratory cultures, which usually
but not always gave normal-sized adults. Those with two significant digits
to the right of the decimal were made with an ocular micrometer on
material preserved in Kahle’s fluid; those with only one digit were made
with calipers on living organisms.

Parides bunichus (Huebner, 1822)

Egg (Figure 8 a): irregularly sculptured spheroid, yellow to red, x = 1.32
mm in diameter (max. ™ 1.44 mm, min. = 1.19 mm, n = 89).

First instar larva (Figures 8 be): reddish, last few abdominal segments light
yellow; head capsule 0.79 mm wide (max. = 0.91, min. = 0.71, n = 165).
Second instar: head capsule 1.21 mm wide (max. = 1.36, min. = 1.05, n =
75).

Third instar: head capsule 1.6 mm wide (max. = 1.9, min. = 1.4, n = 17).
Fourth instar: head capsule 2.6 mm wide (max. =3.0, min. = 2.4, n = 31).
Fifth instar: head capsule 3.8 mm wide (max. = 4.1, min. = 3.3, n = 14).
Note non-overlap of the ranges of head capsule widths between instars,
permitting unambiguous identification of larval instar.

All four Parides larvae share a basic pattern, with a deep maroon to dark
brown-black ground color; conical, pointed, often recurved tubercles on all
segments; and a light-colored, oblique side stripe from the base of a light-
colored dorsolateral tubercle on the 4th abdominal segment across a light-
colored subspiracular tubercle on the 3rd abdominal segment to the base
of the proleg on the same. Light-colored tubercles are always present on
the 7th abdominal segment (dorsolateral and subspiracular), and the 1st
(dorsolateral) and 2nd (subspiracular) thoracic segments.

P. bunichus larvae (Figure 8 d) are distinguished by short tubercles (less
than one-half head width), with the light-colored zones strong yellow to
orange, and have additional light-colored dorsolateral and subspiracular
tubercles on the 9th abdominal segment, but not on the 8th.

The characteristic pattern of the light abdominal tubercles (segments 7-9)
of this species and the following three can be recognized in most
individuals by the late first instar, and permits rapid and unambiguous
identification of larvae.

Pupa (Figure 8 qrs, second from left): light brown or green, short cephalic
projections, no abdominal flanges, two dorsolateral maroon dots on the 1st
abdominal segment.

212

J. Res. Lepid.

Parides anchises nephalion (Godart, 1819)

Egg (Figure 8 ef): pinkish-yellow, noticeably lighter in color and larger than
that of P. bunk has (diam. == 1.4 -1.5 mm).

First instar (Figure 8 g): similar to P. 5umc/ms. Head capsule about 0.8 mm
wide (n = 3).

Second instar: head capsule 1.3 mm wide (max. = 1.4, min. = 1.1, n = 20).
Third instar: head capsule 1.8 mm wide (max. = 2.0, min. = 1.6, n = 30).
Fourth instar: head capsule 2.7 mm wide (max. = 3.0, min. = 2.3, n = 8).
Fifth instar: head capsule 4.1 mm wide (max. = 4.9, min. = 3.5, n = 7).
Note the larger width of the head capsule in all stages of P. anchises in
relation to P. bunichus.

Light colored areas pale yellow; distinguished by additional light dorso-
lateral and subspiracular tubercles on the 9th abdominal segment, and
subspiracular tubercles on the 8th and usually all thoracic segments,
producing prominent yellow lateral patches at the front and rear ends of
the larva. All tubercles are nearly as long as the head capsule width, and
twice as long as those in P. bunichus.

Pupa (Figure 8 qrs, center) : green and yellow (brown phase not observed in
southern Brazilian populations), moderate cephalic projections, long
spatulate flanges on 6th and 7th abdominal segments, upper abdomen
widely flaring.

According to Moss (1919), the larva ofP. neophilus eurybates (Gray, 1852),
which should be found occasionally in the study sites, is nearly identical to
that of P. anchises nephalion.

Details of head capsule width for each instar are not available for the
remaining species, described below.

Parides proneus (Huebner, 1825)

Egg (Figure 8 i): grayish-white and small (diameter =1.0 mm).

First instar (Figure 8 j): very yellowish in the posterior abdominal
segments; strongly club-shaped, with an expanded thorax, tapering
posteriorly.

Fifth instar (Figure 8 k): distinguished by additional light yellow dorso-
lateral and subspiracular tubercles on both the 8th and 9th abdominal
segments and on the 3rd thoracic segment. Most tubercles short as in P.
bunichus, but light-colored tubercles twice as long. Ground color strongly
striated with light gray.

Pupa (Figure 8 qrs, right): green or light brown and yellow, moderate
cephalic projections, short flanges on 5th through 7th abdominal segments.

Parides agavus (Drury, 1782)

Egg (Figure 8 1): usually darker red than that of P. bunichus, but similar in
size.

First instar: similar to that of P. proneus, likewise rather club-shaped; less
yellowed posteriorly.

19(4): 199-226, 1980(81)

213

Fifth instar (Figure 8 m): distinguished by additional light- colored
subspiracular tubercles on the 2nd thoracic segment and the 9th
abdominal segment, but not the dorsolateral tubercles on the 9th
abdominal segment; light- colored abdominal tubercles twice as long as
other tubercles. Ground color often mottled with lighter streaks. Some-
what club-shaped as in P. proneus.

Pupa (Figure 8 qrs, second from right): long narrow cephalic projections,
triangular flanges on 6th and 7th abdominal segments, red antenna cases,
otherwise green or beige strongly mottled with yellow.

Battus polydamas (Linnaeus, 1758)

Egg (Figure 8 n): relatively smooth and small (diameter =1.0 mm), and
deposited in clusters (unlike Parides eggs which are always laid singly),
usually 2 to 9.

First instar (Figure 8 o): yellow, turning to brown with lighter tubercles
after one or two days.

Second to fourth instars: tending to dark red or brown with subequal wii'e-
like tubercles, lighter on the 2nd thoracic and fourth and seventh
abdominal segments, thus at times superficially similar to some Parides
larvae, especially in earlier instars.

Fifth instar (Figure 8 p): wire-like subequal tubercles on all segments,
except for dorsolateral tubercles on first thoracic segment and subspiracu-
lar tubercles on 2nd (usually) and 7th abdominal segments which are much
longer (from the second instar, permitting unambiguous identification).
Body maroon, grey, or beige in various proportions, striated with lighter or
darker markings, especially late in the instar.

Tubercles the same color as body or lighter, occasionally bicolored.
Pupa (Figure 8 qrs, left): green or light brown and yellow, with a long
spatulate dorsal process on the thorax (present in all Battus, never seen in
Parides; Moss, 1919).

Adult Biology

In the laboratory, adults of all species eclosed in the morning (see also
Moss, 1919), and were actively flying by midday and for one to two weeks
thereafter. In the field, butterflies were rarely recaptured more than two
weeks after marking, and consistently passed through one of the six major
wing wear classes per week (Table 1, Figure 9). A maximum of 35 days
between first marking and last recapture was recorded for a male of P.
proneus and a male of P. bunichus; two males of P. agavus lasted 49 and 56
days in Monjolinho, a cool, humid and protective environment with
abundant adult resources in the understroy. In the growth chamber,
lifespans of P. bunichus averaged 11.4 days (max. 24 days, n = 47), in
general agreement with the field data. These results approximate those
obtained for troidines in Trinidad forests by Cook et al. (1971) and in
Texas thickets by Rausher (1979a), but contrast markedly with the 2 to 7

214

J. Res. Lepid.

19(4): 199-226, 1980(81)

215

Fig. 9. Aging of recaptured troidines, combined field sites (1980): test of the
hypothesis of passage through one of the six wing wear classes each week.
Recapture lines for individually numbered butterflies always start within
the scored wear class at first marking. Of 61 butterflies which should be in
class 2 (fresh- intermediate) when recaptured, 53 in fact were, with 1 in class
1 and 7 in class 3. For recaptures in the other classes of wing wear, 3 shows
89 in 3, 9 in 2, and 3 in 4; 4 shows 44 in 4, 10 in 3, and 3 in 5; 5 shows 20 in 5,
8 in 4, and 3 in 6; and 6 shows 13 in 6 and 9 in 5. Maximum inferred lifespan
was 49 days in two separate cases. Note that butterflies were placed at
various points within their wing wear class as scored on first capture, in
accord with recapture data. If all individuals are placed only on the
midpoint of their wing- wear class upon fii*st recapture, and seven days are
allotted to each class (except for teneral = 1 day and fresh ” 6 days), the
numbers found in each recapture set, including all data through early
August 1981, are as follows:

Number Found in Each Wing Wear Class,
Predicted Class Total Number As Actually Scored Upon Recapture

upon Recapture of Recaptures 1 2 3 4 5 6

1 3 3

2

101

4

Tl

26

3

139

21

107

11

4

91

T5

7

5

67

3

21

7

6

42

3

20

19

443

It may be noted that aging is somewhat accelerated (with relation to the
hypothesis) in the earlier wing wear classes, and somewhat retarded in the
three later classes, but the overall picture is very coherent.

month lifespans reported by Young (1971a, 1971b, 1972a, 1972b)forwild
and captive adults in Costa Rica.

The mark-recapture studies indicated that butterflies rarely moved
between the main colony of the Horto Florestal {A in Figure 1 AB) and the
outlying concentrations, but often moved around all parts of Area A
(Figure 10). In 1980, males captured in the Horto Florestal colony A
moved an average of 81 m and a maximum of 660 m between successive
captures on the same day (n ~ 158), or 116 m (average) and 970 m
(maximum) on different days (n = 135) (distances were represented by
minimum straight-line paths between epicenters of marking areas; see
Figures 1 and 10). Corresponding values for females were very similar: 77
m (average) and 620 m (maximum) for the same day (n = 55), and 110 m
and 490 m for different days (n = 30). Some evidence for dispersal over
several hundred meters or across major barriers emerged from the data,
especially in periods of sparse or ephemeral adult resources. Between
colony recaptures (A to B or E) numbered only 31 in the total of 825
recaptures, however (Table 1, Figure 10). The occasional appearance in

216

J. Res. Lepid.

COLONY A - SUMARE

Lantana (irregular)

19(4): 199-226, 1980(81)

217

Fig. 10. Movement of adult troidines, and relationship of population concentra-
tions with seasonal nectar sources in Colony A of the Horto Florestal de
Sumare (above) and in the Amarais woods (below) (see Figure 11).
Locations of preferred flowers, which attracted exceptional densities of
Troidines, are indicated on the base maps of each area, along with the
epicenters for captures in each subcoiony (dark circles). At left center is a
summary of all marks and recaptures in the Horto Colony A (as a
schematized heptagon), by subcolony (circles at apices); M = total number
marked in the subcolony, R = total number of first subsequent recaptures,
and numbers by each arrow = first subsequent recaptures in other
subcolonies; external arrows — subsequent recaptures or prior captures
outside Colony A, representing intercolony transfers. At right are graphs
showing movements of populations between different subcolonies through-
out the year (Horto Colony A above, Amarais below), giving respective
dates, number of captures in the indicated subcolony, and total number of
captures for that date; for example, on March 31, 1981, 48 of the 67
troidines captured in Horto Colony A were on Elephantopis flowers in
subcoloies 4-5; on February 19, 1981, half of the 32 troidines captured in
Amarais were on Stachytarpheta flowers in subcolony 5. All subcolonies
were always covered in each day’s sampling, usually several times, but
more time was spent in areas where more butterflies were encountered.
Dotted arrows indicate discontinuities (different years of observation; refer
to Figure 11).

the Horto Florestal of P. neophilus eurybates, apparently not a resident
there, suggests that immigration and emigration of the resident species
could occur, in spite of the unfavorable habitat extending on all sides of the
site (Figure 1). In support of this, a male P. agavus marked in the riverside
thicket!) (Figure 1 A) was captured a week later within the Horto Florestal
colony E, one km to the SW. In the more continuous habitat of the Amarais
forest (Figure 1C), individual movements of over 400 m between the
scattered flower patches were more frequently recorded (23% of all
recaptures). One agavus male flew from Amarais to Monjolinho (about 1.3
km) in an hour and a half; a female bunichus was also captured in
Monjolinho two weeks after being marked in Amarais; and a female agavus
and a male proneus marked in Monjolinho were captured in Amarais, 7 and
32 days later, respectively.

The short life spans of the adults precluded the use of standard methods
(Jolly-Seber, Lincoln-Bailey, Manly-Parr) for estimation of population
sizes from weekly samples; most butterflies were not recaptured, and very
few were recaptured twice (Figure 9). When several samples were made at
2-day intervals, a simple Lincoln analysis suggested that about a third of
the P. proneus population , a fourth of the P. bunichus population, and a
tenth of the B. polydamas population of Horto Florestal colony A was being
marked or sampled in a 4-hr standardized intensive capture period (other
species gave less than five recaptures). In Figure 11, all samples are

218

J. Res. Lepid.

50i

JAI

I97§

PARIDES AGAVUS

1 |«r— J

PARIDES ANCHISES NEPHALiON

^ III, IN..I«i. 1

PARIDES NEQPHILUS EURYBATES

^ ^FEB ' MAR ‘TpR ^MAy“ ^JUN ' JUl""'* " AU6 ' SEP ' OCT

^ ^1980-riELD"— (aai.... »79 FIELD NO TE S — 4u.--^- ■ • •> 1££^

SUMMER FALL HORTO FLORESTAL sprinq

' NOV ' DEC

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AMARAIS

19(4): 199-226, 1980(81)

219

Fig. 11. Variation during the year in relative abundance of troidines in the main
colony A of the Horto Florestal de Sumar^, from field data, 1978-1981. All
numbers are adjusted to a four-hour sampling period. The daily data have
been exaggerated to better define brood emergences - fresh individuals
(wear classes 0-2) are given triple their value, intermediate ones (wear
classes 3-4) double. Dotted lines are based on numbers seen or captured in
weekly visits outside the disciplined mark-recapture periods. Data from
Amarais, November 1980 is presented at right for comparison; uppermost
curves for Amarais populations are also shown as chains of open circles on
the Horto graph, for February to August 1981. Note non- successive broods
(less than two months interval) inB. polydamas, P. bunichus, P. agavus, and
P. a. nephalion; non-overlap of 1978 and 1980 (November), 1979-1981
(February through early August), and 1979 and 1980 (August and
September) data; and desynchronized broods of P. agavus and P. a.
nephalion in relation to the commoner species.

corrected to 4 hr, and more weight is given to younger age classes in daily
samples; in this way, a very approximate picture of brood emergences and
monthly and yearly population fluctuations can be obtained.

Concentrations of butterflies were invariably encountered on the most
abundant flower resources within the colony areas (see Figure 10). Adults
fed exclusively on nectar, in the field and in the laboratory. Pollen feeding
(cf. de Vries, 1979) was not observed in central SKo Paulo or in any other
parts of the Neotropics. “Puddling”, common in the Graphium mimics of
troidines and thus often reported for this tribe, has only been observed
occasionally for Battus, never for Parides. In the Horto Florestal and
Amarais, butterflies visited Lantana camara L. and L. lilacina Desf.
(Verbenaceae) flowers throughout the year. Seasonal nectar sources
included Petrea racemosa Nees et Mart, (late winter) and Stachytarpheta
polyura Schau. (spring and summer) (also Verbenaceae), Psychotria and
Palicourea species in late spring (Rubiaceae), Vochysia tucanorum Mart.
(Vochysiaceae) and iVestoma coalite (Veil.) Woods. (Apocynaceae) in late
summer, and Vernonia rubriramea Mart., V, scorpioides Pers., and other
composites in the fall and winter. The principal nectar source in
Monjolinho was introduced Impatiens (Balsaminaceae), abundant in the
understory, supplemented by Lantana, Ixom, Euphorbia, and other
ornamental flowers in the adjacent sunny garden (exclusive habitat for B.
polydamas). V^en nectar sources dwindled in number, especially in fall
and winter, butterflies were recaptured far from their marking localities,
and males became increasingly protective of the flower patches, attempt-
ing to expel other males which appeared near them. Males often courted
the females near the flower concentrations. During courtship, males
hovered over females, occasionally darting to and fro, and appearing to
brush the white androconial scales on the dorsal surface of the anal fold of
the hindwing against the antennae and head of the female (Figure 6 f).

220 J. Res. Lepid.

Laboratory observations indicated that both males and females could
mate on the day of eclosion (Figure 6 g). Virgin females up to two weeks old
could mate and subsequently laid fertile eggs; before mating, females very
rarely laid (infertile) eggs. The female copulatory pore is internally
plugged by sclerotic bars after mating, so multiple matings are very
unlikely.

When all captures and recaptures (3571 to mid- 1981; Table 1) were
plotted by species and sex against half-hour time intervals from sunrise to
sunset, females of dllParides species were seen to predominate over males
in the earliest and latest hours of the day, while the opposite was true for
Battus (females flew mostly near midday). Parides proneus was active
significantly earlier in the day than the other species (peaking from 9-10
AM; polydamas peaked near 11 and the other Parides species showed
near-constant activity from 10 AM to 2:30 PM).

19(4): 199-226, 1980(81)

221

Oviposition and Larval Biology

Parides females deposited eggs singly, usually on the underside of leaves
(Figure 8 f). Battus, on the other hand, placed eggs in small clusters on the
growing tips of the Aristolochia vines (Figure 8 n). Females of all five
species preferred some species of Aristolochia over others when choosing
oviposition sites, possibly using chemical cues to identify the preferred
species (Table 2). Prior to oviposition the females appeared to inspect the
vines visually before “drumming” the leaf surface with their foretarsi. The
selection of suitable oviposition sites in the field seemed to depend, also,
on the habitat in which the Aristolochia^ grew; P. anchises nephalion often
oviposited in drier portions of the Horto Florestal (C and F, Figure lA),
while Battus polydamas appeared to prefer open habitats (see also
Rausher, 1979b) and P. proneus invariably favored low plants, essentially
always A. melastoma, in dense undergrowth.

Observations of P. bunichus in the growth chamber indicated that
females deposited about 20 eggs per day and could continue to produce
eggs for up to ten days following the first oviposition. Parides females did
not oviposit as exclusively on apical leaves as did Battus females. A.
Uttoralis, A. gigantea, and A. arcuata leaves often formed necrotic zones
where the egg contacted the leaf, similar to that illustrated under a Parides
photinus egg by Ross (1964). These areas occasionally fell from the leaf,
taking the egg with them.

Eggs of Parides had an irregular, hard coating deposited on them by the
female at the time of oviposition. This coating was poorly developed in B.
polydamas. Five to eight days after oviposition the larvae hatched, eating
the egg shell and its coating. Young (1973) proposed that such coatings
contained chemicals that protected the egg. Moss (1919) and L. E. Gilbert
(pers. comm, to K. B.) suggested that the coating might contain energy rich
nutrients to fuel the newly hatched larvae as they sought out the tender
apical leaves on which they prefer to feed. By the late second, or early third
instar, the larvae could move onto older, tougher leaves. The solitary habit
of Parides larvae, coupled with their tendency to eat any unhatched eggs
and attack smaller larvae that they encountered, limited the damage
inflicted on a plant. B. polydamas larvae were gregarious, displaying no
cannibalistic tendencies, and remained several to a plant even after the
third instar. Consequently, they did much damage to the Aristolochia plant
under attack (cf. Moss, 1919). All mature Aristolochias except A.
melastoma, which had little storage root, quickly sent out new shoots when
eaten back to the ground, so that defoliation was generally not fatal to the
plant. Seedlings and small plants, however, might be killed more easily by
extensive herbivory (Rausher and Feeny, 1980).

harvae-Aristolochia Interactions

The laboratory studies showed that larvae of all five species could
develop in 20 to 30 days, agreeing with reports of Moss (1919) and Young

222

J. Res. Lepid.

(1971a, 1973, 1977) for various species of Battus and Parides. Develop-
mental rates of the larvae depended greatly on the species of Aristolochia
on which they fed (Table 3). Most larvae developed rapidly on A. arcuata
and A. me/astoma, and poorly on A. brasiliensis and A. gigantea. A. littoralis
seemed to be of intermediate quality (see also Young, 1973). P. proneus
larvae seemed to be the most specialized, growing rapidly only on A.
melastoma, while B. polydamas could eat and develop well on most
Aristolochias available to it. P anchises nephalion survived better on A.
brasiliensis (which it uses in the field in Sumare) than the other butterfly
species, andP agavus seemed the best adapted to A. littoralis (and even A.
gigantea). The patterns of hostplant palatability roughly paralleled
patterns of preference for oviposition site (Table 2), and helped to explain
the near absence of P proneus and the predominance of P agavus in
Monjolinho (Table 1).

In the field, larvae occurred on a limited subset of the palatable
Aristolochia species available to them (Table 3). The limited use of some
food plants that would support larval growth could have resulted from the
inability of females to locate or recognize these plants in the field, from
unsuitable microclimatic conditions on these plant species linked to
differences in the sites in which the Aristolochias grew, or from physical or
biological conditions external to the tYoidine-Aristolochia system that
influence hostplant palatability.

Pupal Biology

The larvae entered a wandering stage prior to pupation. Even at high
hostplant densities, such as those in the growth chamber, larvae frequent-
ly. selected pupation sites other than Aristolochia plants. Butterflies
normally eclosed after 13 to 17 days as pupae. ManyP. bunichus, and some
P proneus and P. agavus larvae that developed in the drying atmosphere of
the growth chamber went into diapause as pupae. P. anchises and B.
polydamas never entered diapause under these conditions. Diapause inP.

lasted up to 115 days, and was characterized by a loss of 0.05% to
0.5% of total pupal weight per day compared to a loss of 1% to 3% of total
pupal weight per day in non- diapause pupae. Most diapausing bunichus
pupae broke diapause within two weeks of transferral to closed plastic
boxes lined with moist paper toweling, suggesting that humidity plays an
important role in diapause control in this species. D’ Almeida (1966)
reported that B. polydamas, P anchises nephalion, P agavus, and many
other species in Rio de Janeiro usually skipped winter generations.
Perhaps in these species, diapause is cold -induced.

Pupal diapause would enable butterflies to escape the unfavorable
weather and limited flower abundance of cold or dry winters, factors that
appeared to influence the phenology of the troidines at Sumare (Figure
11), All five troidines were present in the spring and summer, when as
many as 250 individuals could be seen in a single day. Some staggering of

19(4): 199-226, 1980(81)

223

brood emergences was verified in this period, perhaps related to the
shared foodplants (Tables 2 and 3), the short adult lifespan (Figure 9), the
long juvenile development time, and varying responses of diapausing
pupae to rainy spells (Figure 11). Only P. bunichus and P, proneus
persisted through the winter months, though both were scarce at times
from March to May (Figure 11), when P. agavus and P. anchises nephalion
were more frequent. B. polydamas was nearly absent during the cooler half
of the year, and increased very slowly in the dry spring of 1980 (Figure 11).
D’Almeida (1966) observed that troidines occurring together in the
vicinity of Rio de Janeiro had staggered peaks of emergence. Foodplant
phenology may also influence the feeding patterns of the troidines. A.
arcuata and A. melastoma initiate most of their new growth in winter when
most butterfly populations are at a low ebb (Figure 11),

Concluding Comments

Several interactions may define the relationship of Parides and Battus to
their Aristolochia hostplants in southeastern Brazil. The chemistry of the
Aristolochias could act to influence the patterns of damage by the
troidines, and also to allow the troidines to identify some of them as
suitable hostplants. Disproportionate levels of parasitism and predation
on any of the Aristolochias could reduce larval survival on those plants,
even were the plants to be intrinsically suitable as food. The spectrum of
Aristolochias used by a troidine species might be limited by competition
with other troidines. The physical environment could restrict the
Aristolochias available to a troidine on a spatial or seasonal basis. The
evolutionary history of each of the five species could influence the pattern
of hostplant utilization directly by limiting the hostplant characteristics to
which they have adjusted and indirectly by limiting environments in which
each troidine can survive.

The larvae of the five troidines at Sumar^ did not grow equally well on all
the Aristolochias available to them when reared under identical conditions
in the laboratory (Table 3). Similarly, the females would not oviposit
equally on the six available Aristolochias (Table 2). In both cases plant
chemistry is implicated. Not all Aristolochias that supported growth of the
larvae of a troidine species were used by that butterfly in the field,
however, indicating that plant chemistry is not the only factor in play.
Because of the seasonal variation in the abundance of the swallowtail
species and in the phenology of growth in some of the Aristolochias, the
presence of tender foliage suitable for consumption by first instar larvae
on an Aristolochia that was otherwise palatable did not always coincide
with the population maxima of a particular butterfly. Only P. bunichus and
P. proneus occurred in numbers during the cool winter; these two species
range into the cooler montane regions, whereas the other three butterflies
range into milder areas. In this way both the physical environment and the

224

J. Res. Lepid.

evolutionary history of the troidines could limit the availability of some
Aristolochias to the larvae. Parasitism and predation of larvae occurred
most frequently on older leaves and on less acceptable foodplants; food
quality may thus affect survival of the larvae indirectly. Competition has
not been demonstrated, and indeed there is a large excess of foodplant
present in Amarais and in parts of the Horto Florestal (colony A) during
much of the year; the destructive feeding of B. polydamas larvae, however,
could very well decimate smaller patches of foodplant and even affect
larvae of Parides feeding on the same plant. Detailed experimental work on
this and similar systems is necessary to answer the many questions posed
by the troidine -Aristolochia interaction.

Acknowledgements. This research was made possible through support of the
Universidade Estadual de Campinas, the Fundac^o de Amparo a Pesquisa do
Estado de SlTo Paulo (FAPESP, Grant 74/544 - Biol6gicas), the Entomological
Club of Oxford, and the Conselho Nacional de Desenvolvimento Cientffico e
Tecnol^gico (CNPq, Fellowship 1112.4339/78 - ZO) to K.S.B., from the National
Science Foundation Grant DEB 79-22130 to P.F., and from Cornell University
(A.D. White Fellowship) to A.J.D. We thank the administration of the FEPASA
railroad and Sr. Pedro Levantese for permission to work in the Horto Florestal de
Sumare and Dr. R. Forster for permission to work in the Fazenda Santa Elisa.
Research on larval survival in Monjolinho was assisted by I. B. Baldassari A. B.
Barros de Morais, and C. T. Teradaira. Facilities at Cornell University were
provided by the Department of Entomology. Lorraine Contardo and Maureen
Carter kindly assisted in tending the laboratory cultures of plants and butterflies.
We thank May Berenbaum, Robert Hagen, Daniel Janzen and Jan Salick for
valuable comment and discussion, and finally an anonymous referee who made
several relevant suggestions.

Literature Cited

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in Locusta migratoria. Ecol. Entomol. 2: 1-18.

BROWER, J. V. z., 1958. Experimental studies of mimiciy in some North American
butterflies. II. Battus philenor and Papilio troilus, P. polyxenes, and P.
glaucus. Evolution 12: 123-136.

BROWER, L. p. & J. V. z. BROWER, 1964. Birds, butterflies, and plant poisons: A study in
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BROWER, L. P., L. M. COOK & H. J. CROZE, 1967 . Predator responses to artificial Batesian
mimics released in a neotropical environment. Evolution 21: 11-23.

COOK, L. M., K. FRANK& L. P. BROWER, 1971. Experiments on the demography of tropical
butterflies. I. Survival rate and density in two species of Parides. Biotropica
3: 17-20.

D’ALMEDIA, R. F., 1921. Notes sur quelques l^pidopteres d’Amerique du Sud. Ann.
Soc. Entomol. Fr., 90: 57-65.

, 1922. Melanges L^pidopt^rologiques. I. Etudes sur les L^pidopteres du

Brasil. Berlin: R. Friedlander & Sohn, ix + 226 pp.

, 1924. Les Papilionides de Rio de Janeiro. Description de deux chenilles.

Ann. Soc. Entomol. Fr. 93: 23-30.

19(4): 199-226, 1980(81)

225

, 1944. Estudos biologicos sobre alguns lepid^pteros do Brasil. Ar^'. Zool.

S. Paulo 4: 33-72, 3 pis.

, 1966. Catalogo dos Papilionidae Americanos. Soc. Bras, de Entomologia,

Paulo, vi + 366 pp.

D’ ARAUJO E SILVA, A. G., C. R. GONCALVES, D. M. GAL V AO, A. J. L. GONCALVES, J. GOMES, M. N.
SILVA &L. de SIM ONI, 1968. Quarto CatSlogo dos insetos que vivem nas plantas do
Brasil - seus parasites e predadores, Parte H. Ministerio da Agriculture, Rio de
Janeiro, xxvi + 662, viii + 265 pp.

DE VRIES, P. J., 1979. Pollen-feeding rainforest Parides and Battus butterflies in Costa
Rica. Biotropica 11: 237-238.

EHRLICH, P. R. & P. H. RAVEN, 1964. Butterflies and plants: A study in coevolution.
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von EUW, J., T. REICHSTEIN & M. ROTHSCHILD, 1968. Aristolochic acid-I in the swallow-
tail butterfly Pach/iopteraaristo/oc/iiae (Fabr.) (Papilionidae). /sraeLJ. Chem. 6:
657-670.

FRAENKEL, G. s., 1959. The raison doetre of secondary plant substances. Science
129: 1466-1470.

HEGNAUER, R., 1964. Chemotaxonomie der Pflanzen, vol. IQ. BirkhMuser Verlag,
Basel, pp. 184-194.

HOEHNE, F. c., 1942. Aristolochiaceas. Flora Brasiliensis, 6 (XV-II). Lanzara/Insti-
tuto de Botanica, Slfo Paulo. 141 pp., 123 pi.

MOSS, A. M., 1919. The papilios of Para. Nov. Zool. 26: 295-319, 3 pi.

MUNROE, E., 1960. The generic classification of the Papilionidae. Can. EntomoL,
Suppl. 17: 1-51.

MUYSHONDT, A., 1974. An unusually long pupal stage of Battus polydamas polydamas
L. (Papilionidae). J. Lep. Soc. 28: 174-175.

RAUSHER.M. D., 1979a. Coevolution in a simple plant-herbivore system. Ph.D. disser-
tation, Cornell University, Ithaca, New York. 240 pp.

, 1979b. Larval habitat suitability and oviposition preference in three

related butterflies. Ecology 60: 503-511.

RAUSHER, M. D. & P. FEENY, 1980. Herbivory, plant density, and plant reproductive
success: The effect of Battus philenor on Aristolochia reticulata. Ecology 61:
905-917.

ROSS, G. N., 1964. Life history studies on Mexican butterflies. 1. Notes on the early
stages of four papilionids from Catemaco, Veracruz. J. Res. Lep. 3: 9-18.
ROTHSCHILD, w. & K. JORDAN, 1906. A revision of the American Papilios. Nov. Zool.
13: 411-752, 6 pi.

VASCONCELLOS-NETO, J., 1980. Dintmica de populac^es de Ithomiinae (Lep.,
Nymphalidae) em Sumare - SP. Master’s dissertation, Institute de Biologia,
Universidade Estadual de Campinas, Sao Paulo, vi + 206 pp.

YOUNG, A. M., 1971a. Mimetic associations in natural populations of tropical butter-
flies. 1. Life history and structure of a tropical dry forest breeding population of
Battus polydamas polydamas. Rev. Biol. Trop. 19: 210-240.

, 1971b. Mimetic associations of tropical papilionid butterflies (Lepidop-

tera: Papilionidae). J. N. Y. EntomoL Soc. 79: 210-224.

, 1972a. Breeding success and suvivorship in some tropical butterflies.
Oikos 23: 318-326.

, 1972b. Mimetic associations in populations of tropical butterflies. 11.
Mimetic interactions of Battus polydamas and B. bellus. Biotropica 4: 17-27.

226

J. Res. Lepid.

, 1973. Notes on the life cycle and natural history of Parides areas mylotes

(Papilionidae) in Costa Rican premontane wet forest. Psyche 80: 1-22.

, 1977. Studies on the biology of Parides iphidamas (Papilionidae: Troi-

dini) in Costa Ricaa. J. Lepid. Soc. 31: 100-108.

, 1979. Oviposition of the butterfly Battus belus varus (Papilionidae). J.

Lep. Soc. 33; 56-57.

Journal of Research on the Lepidoptera

19(4): 227-229, 1980(81)

Karyolo^ of Three Indian Lasiocampid Moths

(Lepidoptera)

P. K. Mohanty and B. Nayak

Department of Zoology, B. J. B. College, Bhubaneswar, Orissa, India

Abstract, Chromosome studies on three species of Lasicocampid moths
(Crinocraspida torrida, Dendrolinus hyrtaca and Taragama siva) revealed n
of 26, 3 1 and 3 1 , respectively. In both size and morphology, the chromosomes
were almost identical. Sex chromosomes, if any, remained unidentifiable.
The details of chromosome morphology and behaviour during meiosis are
discussed.

Introduction

The Lepidoptera constitute a major group of insects of over 200,000
species. Only a small number of these have been cytologically investigated
so far. Most studies have been concerned with the chromosomes of
Macrolepidoptera, particularly butterflies. The Microlepidoptera are
moths which comprise the majority of species, but have been little studied.
To date very little work has been done on Indian species of Lepdioptera
(Gupta, 1964;Rishi, 1973, 1975;Nayak, 1975). Of the family Lasiocampidae,
chromosome counts of only four species have so far been reported (Rishi,
1973; Nayak, 1975). The present paper deals with the chromosome
studies of three species of Lasiocampid moths, two being reported for the
first time.

Materials and Methods

The larvae were collected from their respective host plants (see Table 1)
and reared in cages. Testes of late larvae (5th instar) and early pupae were
found suitable for cytological preparations. Freshly dissected testes were
fixed in aceto-alcohol (3:1) overnight and smears were made on pre-
warmed albuminised slides using 45% acetic acid. The stain used was
Heidenhains’ Iron Haematoxylin.

Observations and Discussion

Crinocraspida torrida: 2n=52. The spermatogonial metaphase chromo-
somes were exceedingly small, homomorphic and spherical, and were not
differentiable into autosomes and sex-chromosomes. The early meiotic
stages were ill-defined due to the diffuse nature of the chromosomes. At
pachytene, they formed shorter and thicker bivalents though still not
countable. At Diplotene, diakinesis revealed chiasma bearing forms like

228

J. Res. Lepid.

Table 1

Collection Data for Study Material

Name of Species

Food Plant

Place of
Collection

iPeriod of

Collection

Crinocraspida torrida Moore

Terminalia

sp.

Bhubaneswar

Dec., 1976

Dendrolinus hyrtaca Cramer

Zizyphus

jujuba

Bhubaneswar

Aug., 1976

Taragama siva Lef.

Eugenia

jambolana

Bhubaneswar

Sept.-Oct.,1975

‘cross’, ‘V’, rod" and dumb-bell-shaped bivalents, many of the chiasmata
being terminal or near terminal. Metaphase I cells invariably showed 26
bivalents. The chromosomes were oval in polar view and dumb-bell-
shaped on an equatorial plane. A few polyploid cells with almost double
the number of metaphase I bivalents were also observed. Metaphase II
showed 26 univalents.

Dendrolinus hyrtaca: 2n=62. Metaphase I cells showed 31 bivalents.
In some plates, however, one of the bivalents appeared to be deeply
stained while in two other plates almost all the bivalents had undergone
early resolution, the,^ partners of each bivalent lying in close proximity to
each other without^ any actual contact. In a good number of plates, twice
the number of normal bivalents were encountered. At anaphase I, a pair of
small deeply stained equal-sized bodies were visible on equatorial region
of the spindle when all others had already reached their respective poles.
Metaphase II showed 31 univalents.

Taragama siva: 2n”62. Metaphase I cells showed 31 bivalents.
Variation in number by one more or one less was marked in a number of
cells. This might be due to precocious separation of the partners of a
bivalent or fusion of two bivalents. In some abnormal metaphase I cells,
many univalent chromosomes were observed along with some bivalents.
The univalents lay a little apart from each other but in homologous pairs.
This configuration much before onset of anaphase I must have been due to
non-pairing or weak pairing of homologues. In anaphase I, the majority of
dividing cells showed homologues of one bivalent to trail behind the rest
during their anaphasic movement towards the respective poles. Such
lagging behaviour even could be followed up to telophase. Metaphase n
showed 3 1 univalents, confirming the haploid complement for this species.

Of the three species chromosomally examined during this study, two of
them, D. hyrtaca and T. siva, have a base number n“31 which is in close
correspondence with the modal haploid number of the family Lasiocampidae

19(4): 227-229, 1980(84)

229

and with that of Lepidoptera in general (Kemewitz, 1915; Beliajeff, 1930;
Saitoh, 1970; Robinson, 1971; White, 1973; Ennis, 1976). The third
species, C. torrida, has aTiaploi^ number n^^B which is quite different
from the modal number. Twelve members of tile twenty species of this
family which have been cytologically examined so far have^the haploid
number 31. The occurrence of low haploid numbers like 28 in Trichiura
crataegi (Federly, 1945), 26 in Trahla vishnu (Nayak, 1975), 25 in
Malacosoma indica (Rishi, 1973) and 26 in C. torrida (present report)
indicate a trend towards evolution of lower chromosome numbers, and
chromosomal fusion rather than dissociation appears to have played the
main role in the process. The lagging anaph^ic movement of a pair of
elements in Dendrolinus hyrtaca and Taragamd siva, perhaps the homo-
logues of a bivalent, points to their sex-chromosome (XX pair) nature.

Acknowledgments. Valuable suggestions by Prof. C. Das of the Department of
Zoology, Berhampur University, are gratefull>| acknowledged. The work was
supported by the assistance of the University Grants Commission.

t-VTP ''

■ Literature Cited

BELIAJEFF, N. K., 1930. Die Chromosomankomplexe und ihre Beziehung zur Phyio-
genie bei den Schmetter lingen. Z. indukt. Abstamm. »u. Vererb. -L. 54: 369-399.
ENNIS, T. J., 1976. Sex chromatin and chromosome numbers in Lepidoptera. Can. J.
Genet. Cytol. 18: 119-130.

GUPTA, Y., 1964. Chromosomal studies in some Indian Lepidoptera. Chromosoma
(BerL), 15: 540-561. ‘

KERNEWITZ, B., 1916. Spermiogenese bei Lepi^pteren mit besonderer Beruck-
sichtigung der Chromosomen. Arch. Naturge^ch. 81: 1-34.

NAYAK, B., 197 5. '^"Studies on the male germinal chromosomes of thirty-one species of
moths and butterflies (Lepidoptera) Prakruti, 12(1 & 2): 141-150.

RISHI, S., 1973. Chromosome numbers of thirty species of Indian Lepidoptera.
GenenPhaenen, 16(3): 119-122.

ROBINSON, R., 1971. ‘Lepidoptera Genetics’. Pergamon Press, New York, Toronto,
Sydney, Braunschweig.

SAITOH, K. & K. KUDOH, 1970. On the chromosomes of two species of moths
(Lepidoptera, L5rmantridae and Lasiocampidae). Kontyu. 38/1: 45-49.

WHITE, M. J. D., 1973. ‘Animal Cytology and Evolution’, 3rd Edn. Cambridge Uni-
versity Press.

;g, 1. Motaphase I ofCrinocraspidatorrulat^ Fig. 2. Metaphase I of Fig. 3. Metaphase I of Taragamcsiw,,

Journal of Research on the Lepidoptera

19(4): 230-239, 1980(81)

niustrations and Descriptions of some Species of
Pyirhopyginae from Costa Rica, Panama and Colombia
(Hesperiidae)

S. S. Nicolay

and

G. B. Small, Jr.

1500 Wakefield Drive, Virginia Beach, VA and
PSC Box 2510, APO Miami, FL 34002

About eleven years ago, the authors (1969) described a new subspecies
of Pyrrhopyge creon Druce from Panama' and remarked that it was not
possible to make a definite statement about the distribution of the species
as a whole. In the intervening years, the junior author has observed and
collected creon in a number of different localities in Panama' and Costa
Rica with some unexpected results. It is the purpose of this paper to bring
our knowledge of this interesting species up-to-date.

Nominate creon is distributed continuously from Northern Costa Rica to
Code' Province, Panama' along both the Atlantic and Pacific slopes of the
series of cordilleras forming the continental divide. The axis of this divide
runs generally in a northwest to southeast direction. More specifically, the
northwestemmost locality at which creon creon has been collected is
Guanacaste Province, Costa Rica, from which it is distributed in a general
southeastward direction through the Cordillera Central and the Cordillera
Talamanca, the last of which enters Panama' in western Chiriqui' province.
The continental divide then continues in an easterly direction as the
Cordillera of Tabasara', which terminates slightly northwest of the town of
Penonome', Code' Province. The southeastemmost record is from near
the town of El Cope', Code', virtually at the terminus of this cordillera. The
Atlantic slope has a year-round wet climate and here the butterfly is found
from an elevation of 750 m up to 2000 m. Most of the Pacific slope has a
pronounced dry season from December to April, and in these areas it
seems to be found only above 1000 m. However, the Pacific slope of the
Cordillera Tabasara' has the same climate as the Atlantic slope, and here
creon can be found again as low as 750 m.

Throughout this range of more than 400 kms. in length, there is
continuous highland with elevations greater than 750 m. However, there
are two distinct regions, in each case separated by less than 40 Ions, from
tile main range, where the species has developed striking and distinctive
subspecies. In each case there is an intervening area of lower elevation

19(4): 230-239, 1980(81)

231

Fig. 1. Upper row, left to right: (A) Pyrrhopyge creon taylori Nicolay & Small d*
holotype, (B) Pyrrhopyge creon liUiana Nicolay Small paratype, (C)
Pyrrhopyge creon Brace c?. Middle row, left to right: (D) Pyrrhopyge creon
taylori Nicolay & SmaU 9 allotype, (E) Pyrrhopyge creon lilliana Nicolay &
Small 9 paratype, (F) Pyrrhopyge creon Bruce 9. Bottom row, left to right:
(G) Mimoniades aerata Godman & Salvin cf, (H) pyrrhopyge sangaris
Skinner d*.

which apparently serves to keep their populations isolated. In neither case
have any specimens that could be called intergrades or hybrids with typical
creon been found.

The first of these regions is an isolated massif with elevations from 750 m
to 1000 m of little more than 30 kms. in length located in extreme eastern
Code' Province and extreme western Panama' Province. This includes the
well known collecting localities of Cerro Campaha and La Mesa (north of El
Valle). It lies about 40 kms. to the east of the eastern terminus of the
Cordillera of Tabasara', and is separated from the latter by a low lying area
of less than 500 m. The climate here is not as wet as in the Cordillera of
Tabasara', and in the months of December to April there is little
precipitation, although the forest remains evergreen due to frequent mists
and cloud cover. Here occurs the very distinctive subspecies lilliana as
described by the authors in 1969.

232

J. Res. Lepid.

The isthmus of Panama' then turns northeastward and altitudes of 750 m
and over are not encountered again until the region of the watershed of the
Chagres River, about 80 kms. to the northeast. Despite intensive collecting
in this region, there has been no sign of creon either here or further east
toward Colombia.

The second region lies in southeastern Costa Rica in the province of
Puntarenas, where south of the Cordillera of Talamancf (Continental
Divide) there occurs an isolated range, the Fila Cruces, of approximately
30 kms. in length, and with elevations of up to 1500 m. Between this range
and the Cordillera of Talamanca is the valley of the Rio Coto Brus, a
relatively low lying area with elevations less than 1000 m and with a strong
dry season of four months with little cloud cover. There are no records of
creon from this valley, which averages about 25 kms. in width. About 6 kms.
south of the town of San Vito de Java is the Las Cruces Tropical Garden,
situated at an elevation of about 1200 m on the northern flanks of the Fda
Cruces. The property includes a forest preserve of over 100 hectares
extending down to the Rio Java at almost 1100 m, and then up the other
side of the valley of this river. Here, and presumably throughout the Fila
Cruces in similar environments, occurs an isolated and distinctive
subspecies of creon which we take pleasure in dedicating to its discoverer,
Mr, Thomas Taylor.

Pyrrhopyge creon taylori Nicolay & Small, new subspecies
Figures lA, ID, 2

Male: Length of forewing, 28 mm ± 1; holotype 28 mm. Upperside:
forewing a dark, shining bronze-purple with a relatively narrow, vaguely
defined dark border on the outer margin and the apex. Hindwing with the
disc shining purple with an even deeper bronze sheen, the dark border of
the wing margin somewhat wider than that of the forewing and more
sharply defined. There is a single orange-red sub-tomal spot in space 1-lb,
the same size and form as in the nominate species. Underside: the purple
coloring of the upper side is repeated but without the shining brilliance, is
more subdued and has a smoky cast. The sub-tomal orange-red spot of the
upper side is repeated in like manner. Fringes concolorous with the dark
outer border.

The abdomen, thorax, tegulae, collar, leg clothing are black; the palpi and
head are covered with intermixed dark reddish-brown and black hairs.

Female: Length of forewing, 33 mm ± 1 mm; allotype 33 mm. Upper and
undersides with all coloring and maculation the same as in the male.

Holotype male, Finca Las Cmces, San Vito (1150 m), Puntarenas, Costa
Rica, 23 VI 1976, G. B. Small, Jr. Collector. Allotype female, same locality
and collector, 24 VI 76. There are 54 male and 20 female parat3TDes
collected by G. B. Small at the type locality with dates ranging from 22 VI
to 3 Vn 1976. Additionally, there are 9 male and 1 female paratypes

19(4): 230-239, 1980(81)

233

collected by Thomas Taylor from the type locality dated 13-20 VI and in
Vin 1971. The holotype is deposited in the Allyn Museum of Entomology,
Sarasota, Florida, the allotype in tihe U. S. National Museum collection,
Washington, D, C. Paratypes are deposited in the British Museum
fl^atural Histoiy), The American Museum of Natural History and the
Carnegie Museum of Natural Histoi^,

As in the case of the subspecies lilliana, the male genitalia clearly
indicates the subspecific relationship of tayloii to nominate creon. And,
unlike lilUana, the facies and general configuration of tayloH is much closer
to creon^ the primary and notable difference being only the basic color. The
shining bronze-purple color is unlike that of any other species of the genus
with which we are familiar. A striking feature of the creon complex of 3
subspecies is that there are no known intergrades, nor have we seen or
taken any specimens that could not be placed at once in the correct
subspecies. Yet, the male genitalia of all three show little or no difference
and, in fact, are absolutely identical, even to the rather subtle asymmetry
of the valvae.

Fig. 2 . Male genitalia - Pyrrhopyge creon taylori - lateral view of uncus, aedeagus in
place, inner surface of both valvae and a ventral view of the uncus.

234

J. Res. Lepid.

Discussion

It is apparent that notwithstanding its considerable capabilities for
strong flight, creon is a very sedentary insect, for gaps of unsuitable
territory of no more than 30 to 40 kms. are sufficient to keep populations
separated. - c

All of the subspecies have the following traits in common:

(1) individuals are numerous, even abundant, at suitable places and
seasons.

(2) males perch with outstretched wings on the tips of leaves from 8 to 40

feet above the ground. ^ ►

(3) the perching behavior of the males commences around 9:30 a.m., is
most prevalent from this time until about 11 a.m. and ceases altogether
about 12:30 p.m.

(4) perching males are quite belligerent toward other passing lepidop-
tera, but are unwary and easy to capture.

(5) females are much more rarely seen,« and then usually searching for
the larval foodplant.

(6) both sexes are partial to a wide variety of flowers, particularly
Eupatroium and are most frequently seen thereon between 8 and 9 a.m.
(later at higher altitudes and/or depending on weather conditions.)

On the other hand, there appear to be certain differences in the behavior
of the various subspecies.

P. creon lilliana perches somewhat lower than the other subspecies,
usually from 8 to 16 feet above the ground. P. creon creon often perches
extremely high and is therefore usually quite difficult to net except when at
flowers. P. creon taylori appears to perch somewhat higher than lilliana but
lower than creon. Since lilliana and taylori have been taken only at two
localities and one locality respectively, the data on them are perhaps
influenced by the nature of the terrain where they were taken, and may
therefore be inconclusive.

At Santa Fe, Veraguas, males of creon creon were seen by both authors to
fly vitually ceaselessly under the canopy of a mature wet forest at heights of
12 to 20 feet above the ground evidently looking for females. They settled
very infrequently, and did not return to a given perch as in normal perching
behavior, making it almost impossible to catch them. This behavior
pattern is perhaps limited to the interior of forests. In any event, at this
locale no individual was seen to engage in “normal” perching behavior,
either in the interior of the forest or along its borders. Such behavior has
never been observed in lilliana and seems to be only occasional in taylori.

In view of the considerably different appearance of lilliana from typical
creon, and the differences in habits mentioned above, it may be that lilliana
should be considered a full species. On the other hand, the genitalia of
creon and lilliana are essentially identical, and lilliana is clearly derived

19(4): 230-239, 1980(81)

235

from creon stock. So, it appears that until some evidence of breeding
incompatibility between them may be produced, the relationship between
these allopatric taxa is best expressed by the subspecific concept.

Unfortunately, nothing is known at present about the foodplants or life
histories of creon and its subspecies.

Specific records:

Recorded below are the data on material studied in the institutional
collections as indicated: (BM) British Museum (Natural History); (AM)
American Museum of Natural History; (USNM) U. S. National Collection,
Smithsonian Institution; (AME) Allyn Museum of Entomology; localities
without an institutional designation are those collected by the authors.
Pyrrhopyge creon creon: No data 6 cT 2 9 (BM), (USNM)

Costa Rica: 6 2 9 (BM)

Tres Rios 2 cf (BM)

Cartago 3 cf 2 9 (BM)

Moravia de Chirippo (1100m) - August
Guanacaste 1 cf (BM)

San Jose 1 cf (AM) 1 cf (USNM)

San Geronimo 4 cf (USNM)

Heredia

Volcan Barba (2000 m) - August
Puntarenas

Monteverde (sight record - GBS)

Panama':

Chiriqui 4 cf 6 9 (BM); 1 cf 1 9 (USNM)

Sta. Clara (1200 m) July 3 cf (GBS)

Cerro Colorado (Continental Divide) 1450 m July-Oct.
40 cf 10 9 (GBS)

Veraguas 1 cf (BM)

Sta. Fe (800 m) Sept. 1 cf (GBS) 2 cf 1 9 (SSN)

Code'

El Cope (600 m) Dec. 1 cf (GBS)

Colombia 1 cf (BM) (?)

Pyrrhopyge creon lilliana
Panama':

Panama'

Cerro Campana (800 m) July-Dee. 50 cf 8 9 (GBS)

Cole'

La Mesa (800 m) Dec. 1 cf (GBS)

Pyrrhopyge creon taylori
Costa Rica:

Puntarenas

Finca Las Cruces, San Vito (1150 m) July- Aug.

50 d- 17 9 (GBS) 6 d* (AME)

236

J. Res. Lepid.

Fig. 3. Male genitalia - Pyrrhopyge sangaris - lateral view of uncus, aedeagus in
place, inner surface of both valvae and a venta*al view of the uncus.

The concept of this paper, begun more than three years ago, was
originally limited to a discussion of the species creon and a description of
the new subspecies tayloii. Subsequent discovery of two male specimens
of Pyrrhopyge sangaris Skinner, and more notably, two male specimens of
Mimoniades (Pyrrhopyge) aerata Godman & Salvin in the Carnegie
Museum collection provided us with the incentive and material to expand
this study to include all three species listed by Evans (1951) at the
beginning of his VIH. Hygieia Group.

P^^hopyge sangaris Skinner
Figures: IH, 3

Pyrrhopyge sangaris Skinner, 1921, 32: 236-237. BeU, 1931, 39: 467.
Evans, 1951, Part I, p 31.

Sangaris has never been illustrated. Superficially it is similar to creon,
but all wing shapes are more sharply angular and there is no mistftoig the

19(4): 230-239, 1980(81)

237

lustrous bronze-green color of sangaris as contrasted to the deep, shining
blue of creon. Skinner’s original description is quoted below. The male is
figured together with an accurate line drawing of the genitalia.

“ .-"Palpi crimson with tips black. Abdomen and legs dark green-black.

Upperside. Primaries shining green-black. Secondaries shining green-
black, somewhat darker than the primaries, with a blood-red spot near the
anal angle. This spot is quadrate, 4 mm. wide and is 3.5 mm. from the inner
margin and about the same distance from the outer margin.

Underside. Primaries as above but lighter in color with the crimson spot
repeated but somewhat smaller and rounder. Expanse (one wing) 23 mm.
Inner margin of hind wing 21 mm.

Type one male in the collection of The Academy of Natural Sciences of
Philadelphia, taken at Hacienda Cincinnati, Sierra San Lorenzo, Magda-
lena, Colombia, July 23rd, 1920, Academy Colombia Expedition, Rehn
and Hebard,

This handsome species has a superficial resemblance to creon Druce but
has the shape of phidias Linn.”

A total of 5 specimens of sangaris were examined in the course of this
study. Three in the British Museum (Natural History), 2 males and a
female from Onaca, Santa Maria, Colombia. Two males in the Carnegie
Museum are from Valpariso (4500 ft.), Dept, of Magdalena, Colombia,
December.

Mimomades aerata (Godman & Salvin) new combination
Figures: IG, 4

Pyrrhopyge aerata Godman & Salvin, 1879: 152, pi 14, fig 3. Mabille &
Boullet, 1908: 177, 182. Draudt in Seitz, 1922, 5: 839, pi 166a. Bell, 1931,
39: 468. Evans, 1951, Part I, p 31.

As far as can be determined, the two male specimens of aerata m the
Carnegie Museum of Natural History are the only two known: the type, a
female in the British Museum (Natural History) is still unique in that
collection. Upon examination of the male genitalia, it became readily
apparent that aerata, belongs to the genus Mimoniades and not Pyr-
rhopyge. The antennal apiculus and wing venation are also characters
compatible with these features found in Mimoniades. A description of the
male follows:

Male: Length of forewing, 28 mm. Upperside: forewing shining bronze-
green with darker shading on the outer third of the wing. Hindwing bronze-
green with the basal half covered with back hairs of nuxed lengths and
density, and all vems somewhat darkened. Underside: bronze-green, the
forewing unmarked, the hindwing with 5 orange-red spots, one each in
interspaces 1 c and 2, then across the cell (the largest), interspace 6 and a
small adjoining spot outside the ceU. Cilia concolorous dark ^een. The

238

J. Res. Lepid.

abdomen with a full-length narrow, dorsal black stripe, laterally with 6
orange-red striped alternating with narrow black stripes. The anal tuft,
vental surface of the abdomen and leg clothing black. The thoreix, palpi,
head, collar and tegulae, black. There are two inconspicuous pale yellow
tufts of hair in the collar located just behind the very minute and
inconspicuous yellowish tufts of hair at the base of the antennae. The
antennal club shaped as for the genus, rather stout and tapered to a blunt
point only at or near the tip.

''fhe male genitalia, figured here for the first time is of a similar form to
that of the species Mimoniades montra Evans. The undivided uncus is
long, without side flanges and in lateral view appears like a long, slender
bird’s neck and head. The valvae are asymmetrical and heavily spined, the
aedeagus slender, bent ventrally and without spines.

All specimens known £u-e from the Dept, of Magdalena, Colombia with
specific data as follows:

Female type, Puebla Viejo, Sierra Nevada de Santa Marta (BM)

Males (2) El Libano (6000 ft.) Dept. Magdalena, Colombia (May)

Fig. 4. Male genitalia - Mimoniades aerata - lateral view of uncus, aedeagus in
place, inner surface of both valvae and a ventral view of the uncus.

19(4): 230-239, 1980(81)

239

Acknowledgments. We wish to express our appreciation to all who have helped in
the preparation of this paper. To those individuals entrusted with the collections in
their care — Harry Clench, the Carnegie Museum of Natural History, R. I. Vane-
Wright, the British Museum (Natural History), Dr. F. H. Rindge, the American
Museum of Natural History, Dr. Lee D. Miller, the Allyn Museum of Entomology
and Wm. D. Field, the U. S. National Museum — go our thanks for allowing us access
to study the material in these collections. And' our grateful thanks to Robert
Robbins for reading the manuscript with a critical ey^e and offering suggestions and
constructive comments for its improvement. .it-

Literature Cited

BELL, E. L, 1931. Studies in the Pyrrhopyginae, with Descriptions of Several New
Species (Lepidoptera, Rhopalocera, Hesperiidae). Jour. N. Y. Ent. Soc. 39:
417-491.

DRAUDT, M., 1922. American Rhopalocera in Seitz, A. Macrolepidoptera of the
World. Vol. 5. Stuttgart.

DRUCE, H. H., 1874. Cist. Entomol. Vol. I.

EVANS, W.H., 1951. A catalogue of the American Hesperiidae. Part I, Pyrrhopyginae.

British Museum (Natural History). London.

GODMAN, F. c. & 0. SALVIN, 1879. Proceedings Zoological Society of London.

, 1879-1901. Biologia Centrali- Americana. Insecta. Lepidoptera-

Rhopalocera. London.

MABILLE, P. & E. BOULLET, 1908. Ann. des Sciences Nat. Zool. Paris. 9th Series.
NICOLAY, s. S. & G. B. SMALL, Jr., 1969. A new subspecies of Pyrrhopyge creon (hesperi-
idae) from Panama. J. Lep. Soc. 23: 127-131.

SKINNER, H., 1921. Ent. News. Vol. 32.

Journal of Research on the Lepidoptera

19(4): 240, 1980(81)

Errata: Scott 14:2; Scott 17:50; Scott & Scott 17(2)

Some unfortunate printer’s errors were made in my recent papers. In vol.
14, p. 2 the sentence “In patrolling, species, interactions occurred
predominantly when resting males investigated moving objects.” is wrong
and must be deleted. In vol. 17, p. 50 the upside down 1 and 3 after “Wet
Mtn. Valley and Salida” should be deleted and a 1 placed in column D, a 3
in column E. The remaining corrections are for vol. 17, no. 2. On p. 73
our address is Estes not Eses St. P. 75, paragraph 2, a comma after
timberline, not after hermodur. Paragraph 3, a period after “here”.
Paragraph 4, delete period after polyxenes. P. 76, line 8, “is”, not “in”. P.
76, paragraph 3, Rutaceae is correct spelling. P. 77 add Prunus to P.
eurymedon hosts. P. 80 Melilotus is correct spelling on line 13. P. 81,
paragraph 7, line 3, should read “plants. Several broods; there are...”.
Delete last two lines on p. 81. P. 82, line 7 and p. 83, line 8, poplar, not
popular. P. 82, paragraph 2, lines 4 and 5 should read “and late Aug.).
Common; the second brood is less common than the first but is sometimes
common in Denver,...”. Same paragraph, line 7, parvifolia is correct
spelling. P. 84, paragraph 4, insert “all” after “12 loc.:”. P. 85, last
paragraph, a comma instead of a parenthesis after “Sept. 13”, and
“period” not “perio.” in last line. P. 87, line 1, should read “two or three
broods in...”. P. 88, paragraph 2, 21 not 2 localities, in first line. P.91, first
two paragraphs should be moved to bottom of page, and the end of the C.
oetus paragraph should read “flowers of all colors, frequently composites,
apparently preferring blue over yellow ones.”, and C. meadii should start a
paragraph. The last three paragraphs on p. 92, and the first two
paragraphs on p, 93, should be deleted. P. 95, paragraph 2 and p, 99,
paragraph 3 should read “dos P.” not “Dosp.” or “dosp.”. The last
paragraph on p. 99 and the first nine lines on p. 100 should be deleted.
P. 102, line 4, should read “early June, late June-late July, Aug. 16-Sept.
13”. P. 106, paragraph 3, line 3, insert “present on” before “the San Luis
Valley...”. P. 107, paragraph 5, line 6, should read “Larvae may feed on
Bouteloua'\ P. 118, paragraph 5, line 7, should read “(6 only northeast-
ward, 3 only southeastward)”. Paragraph 6, line 7, add “P. acmon'' after
shasta. P. 119, paragraph 4, change E. dorothea to C. pertepida. Last
paragraph, line 3, rhesus not urhesus. P. 120, paragraph 3, Phyciodes is
correct spelling, and insert “& Convolvulus'^ after Aster. P. 127, paragraph

I, insert “possibly” after milberti. On back cover, the satellite photo refers
to J. Scott and G. Scott, not Emmel & Shields. Note: if your copy of vol.
17, no. 2 lacks pp. 99-114, write to Scott Miller of the Santa Barbara
Museum for a correct copy (address on inside front cover).

J. & G. Scott, 60 Estes Street, Lakewood, Colorado 80226

Journal of Research on the Lepidoptera

19(4): 241-244, 1980(81)

Book Review

A catalogue/checklist of the butterflies of America north of Mexico.

Lee D. Miller and F. Martin Brown, Lepidopterists’ Society Memoir No. 2, 1981.
280 pp. (Lep. Soc. members $5.00 cloth, $10.00 hai’dbound; non-members
$8.50/17.00).

The long awaited Miller and Brown catalogue is now at hand. This volume was well
worth the wait as it is a far more useful work than dos Bassos’ (Lepid. Soc. Memoir,
no. 1, 1964) synonymic list. The authors are to be highly commended in giving the
systematicist his most useful single book on the nomenclature of North American
butterflies.

Full synonymies are given for each taxon from the generic level down. For each
genus (and synonymic genera) the literature citation of its original proposal and the
type species are presented. Under each species, besides a listing of aU named
subspecies (and synonomies), forms and aberrations, we are treated to the genus
under which it was originally described, the citation of the original description, the
type locality and the location and kind of type specimen when known. This
alleviates us of the boring and time consuming task of pouring through the
Zoological Record to find where something was described. Finally, and probably of
most importance, there are over 650 footnotes which justify, explain or expand
upon the main body of the text.

Overall, Miller and Brown are splitters, especially at the generic level. For
example, they use 21 genera for Pieridae (13 in dos Bassos), seven for Papilionidae
(5), seven for Lycaeninae (l)and 16 for Polyommatinae (11). This is largely a result
of raising dos Bassos’ subgenera to the generic level (e.g., Falcapica, Zerene,
Aphrissa, Pyrisitia, Abaeis). Others are new genera proposed in recent reviews (e.g.,
Hyllolycaenaa, Hermelycaena, Euphilotes, Philotiella). Still others result from
finally recognizing that some of our taxa are congeneric with, especially, Palearctic
ones (e.g., Pontia, Artogeia, Aglais). On the other hand, there is a realization that
some Nearctic genera are sufficiently distinct to warrant recognition {e.g.,Basilarchia).

In any work of this sort, there are bound to be decisions not agreed upon by
everyone, typographical errors not edited out and omissions. This book is no
exception. My following comments point out some of these and are largely directed
towards taxa that occur in Nevada and adjacent areas.

In a number of places (e.g., p. 60, 66, 116, 117, 121, 142) we are referred to a
footnote as “see Note 000”. These are perhaps the most annoying typographical
errors as I wonder if I am missing some important bit of information. Footnote 557
(p. 177) is really note 571. The genera are sequentially numbered yet Nat/iafo and
Feniseca are CXVn and Enantia and Thersalea are CXVIII; genus CXXXII
{Satyrium) is followed by CXXXV {Ocaria).

A couple of other remarks are pertinent before turning to individual species. The
first concerns taxa described from some of Lorquin’s specimens. Difficulties with
Lorquin’s Utah and “Lac Sal” localities have been known for some time (Brown, J.
Lepid. Soc. 21: 271-274, 1967). Miller and Brown restricted the type localities of
Neophasia menapia, Apodemia mormo and Speyeria mormonia to the general area
near or to the west of Pyramid Lake, Washoe Co., Nevada. I have no particular
quarrel with the first two. The menapia could have come from nearly an3rwhere on
the east slope of the Sierras. Likewise, mormo occurs in scattered pockets on the

242

J. Res. Lepid.

valley floor and in lower canyons in the same general area. The mormonia presents a
special problem. East slope Sierran populations in the Reno-Carson City area seem
to fall into the concept of arge especially in their variable silvering. If this is where
Lorquin took the types then there is a nomenclatural problem. We know that
Lorquin was in the Carson City area from Behr’s description of leto, but there are
other lakes that could have been “Lac Sal” including Mono Lake and the Alkali
Lakes east of the Warner Mountains.

Then there are a number of taxa that were proposed by Bauer {in Howe, The
Butterflies of North America, Doubleday, Garden City, 1975). These are right on the
fuzzy edges of validity under the “Code”. All are treated adequately except
Thessalia leanira oregonensis which is not mentioned. Miller and Brown’s com-
ments that apply to the other Bauer taxa {in Howe) apply to this taxon also.

Now to a number of comments on various species.

Erynnis persius - the four “subspecies” are listed although Bums (Univ. Calif, Publ.
Ent. 37, 1964) suggested against recognizing any names in this complex until they
were better known biologically.

Pyrgus communis/albescens - these are presented as separate species “on the advice
of H. A. Freeman” but we are given no hint of why.

Anthocharis cethura/pima - retained as separate species although it is recognized
by collectors in the southwest that there are intermediate populations.

Atlides halesus - the western populations are still jumping around from name to
name: estesi Clench, corcorani Gunder, corcorani Clench, corcorani dos Passos.
Callophrys comstockiAemberti - these are retained as separate species although
there are several intermediate populations. These should, at least, be as sub-
species {comstocki has priority). I prefer treating them as taxa under sheridanii.
Incisalia fotis - schryveri, fotis, mossii, bayensis, doudoroffi and windi are treated as
subspecies. The Sedum feeding taxa are undoubtedly separate from nominate fotis.
Euristrymon - Clench (J. Lepid. Soc. 32: 277-281, 1978) synonymized this with
Fixsenia Tutt. The latter is not even mentioned.

Parrhasius m-album - why this genus, did I miss something after Panthiadesl
Euphilotes - Mattoni’s (J. Res. Lepid., 16: 223-242, 1977) generic revision is fol-
lowed yet there is some inconsistency within the species. Mattoni recognized
battoides, enoptes, mojave, rita and spaldingi as good species. Shields (Bull., AUyn
Mus. 28, 1975; J. Res. Lepid. 16: 1-67, 1977) recognized battoides, enoptes (with
mojave as a subspecies and rita (with spaldingi as a subspecies). Here Miller and
Brown follow Shields except that rita is split into two species, rita and pallescens.
Lycaeides melissa - fridayi is recognized and paradoxa takes precedence over
inyoensis.

Icaricia acmon texana - a slight error here as this was originally described as a
Plebejus.

Agriades franklinii - a totally new face for the species we have known so long as
aquilo or glandon. The latter are apparently Palearctic.

Speyeria coronis gunderi - it has been known for some time that gunderi refers to a
zerene population. Unfortunately it was described from an area where red Sierran
and blond Great Basin zerene mix. However, the holot3q)e is quite indistinguishable
from populations going under the name of cynna. Thus gunderi should be moved
to zerene with cynna as a synonym.

Phyciodes pratensis - takes precedence over campestris under the “First Reviser”
tenet.

19(4): 241-244, 1980(81)

243

Euphydryas anicia - I’ll bet that ab= “duncani” (TL Sta. Catalina Mtns., Arizona)
refers to hermosa rather than wheeleri.

Basilarchia lorquini - in a footnote it is stated that “fridayi” is a hybrid of this species
and B. weidemeyerii nevadae. The weidemeyerii taxon involved is latifascia. Be-
fore latifascia was described much of western Great Basin weidemeyerii were
considered nevadae, a concept is now restricted to populations in the Spring and
Sheep ranges in southern Nevada. Thus, the older literature (correctly at the time)
reported nevadae as one of the parent taxa.

Asterocampa - Fm still amazed that we maintain so many mono ty pic species.
Coenonympha - retention kodiak, inomata, ochracea, ampelos and California as
separate species under the concept of a tullia superspecies. This appears to be a
reasonable means of handling this difficult group.

Coenonympha ochracea brenda - recent studies indicate that brenda does not apply
to Great Basin populations or, for that matter, to ochracea. This is being worked
out by R. E, Gray.

Cercyonis pegala - some changes were made in the synonymy of western members
which were badly needed, especially recognition of certain western Great Basin
populations under the name stephensi.

Cercyonis oetus - the taxon pallescens Emmel and Emmel {Pan-Pacific Entomol. 47 :
155-157, 1971) was omitted.

These are, for the most part, minor points that can be cleared up at a later date in
an “Addenda et Corrigenda’’ once there is sufficient input from others. Again, the
volume is an extremely valuable contribution to systematic Lepidopterology and
should be on all our shelves.

George T. Austin, Nevada State Museum, Capitol Complex, Carson City, NV 89710.

The catalogue is an absolutely necessary work for anyone with the slightest
systematic concerns with the Holarctic butterfly fauna. Although there will be many
arguments based on the arrangement of taxa, the meat is here.

Since I feel only competent to comment on my own group, the Scolitantidini, I have
tried to extrapolate errors of fact and typography to those of the whole book. Thus
there are two such errors in four pages: 1) citation of bohartorum as a synonym of
Philotiella speciosa (it is clearly a valid subspecies and perhaps a species with no
published report to the contrary), and 2) a 000 note on p. 12 1. At this rate 100 errors
can be expected in the some 200 pages.

For the Polyommatinae it is unfortunate the authors did not follow Eliot’s
classification [1973, Bull. Brit. Mus. N. H. 28(6)]. Recognizing Eliot’s sections as
tribes would place Brephidium, Leptotes, and Zizula each in its own tribe. Their
placement here in Lampidini is misleading. Most serious, Hemiargus is also placed
in Lampidini, where it is clearly in Polyommatini, surely one of the best demarked
groups of Blues.

Interpretatively, I would like to point out that Euphilotes spaldingi (511 c) is
clearly specific on morphological, developmental, and behavioral grounds. E. rita
and E. pallescens may also be treated as one species based upon presumptive

244

J. Res. Lepid.

intermediates found in Washington Co., Utah (Shields, pers. comm.). There are
data, both biological and morphological, to indicate E. mojaue and E. enoptes can be
considered distinct. These points, however, have not been formally published, and I
can in no way fault the authors concerning their use.

No doubt every specialist will offer his quarrels. Miller and Brown should be
pleased thereby that such arguments in reality serve to point out the lack of
meaningful systematic revisions for many groups. The authors, in their introduction,
take pains to point out the classification used represents their joint opinion, and
that they were not in agreement on some matters. Such recognition makes the
message that no taxonomy is cast in stone.

We now can look forward to the day there is a similar catalogue of the Palearctic
butterfly fauna.

R. H. T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210.

THE JOURNAL OF RESEARCH
ON THE LEFIDOFTERA

Established in 1 962

Founded by William Hovanitz
Edited by Rudolf H. T. Mattoni

Volume 20

1981

published by

The Lepidoptera Research Foundation, Inc.
at

2559 Puesta Del Sol Road, Santa Barbara, California 93105

Volume 20 Number 1 Spring, 1981(82)

IN THIS ISSUE

Date of Publication: September 20, 1982

Editorial 1

Butterfly Nomenclature: A Critique

Paul R. Ehrlich & Dennis D. Murphy 1

Early Stages of Speyeria nokomis (Nymphalidae)

James A. Scott & Sterling 0. Mattoon 12

Supernumerary Chromosomes in the Domesticated Eri-Silkmoth,

Philosamia ricini (Saturnidae: Lepidoptera)

K. B. Padhy & B, Nayak 16

Geographic Variation and Ecology of Hesperia leonardus (Hesperiidae)

James A. Scott & Ray E. Stanford 18

Notes on the Immature Biology of Two Myrmecophilous Lycaenidae:

Juditha molpe (Riodininae) and Panthiades bitias (Lycaeninae)

Curtis J. Callaghan 36

Immature Stages of Odonna passiflorae Clarke (Lepidoptera:

Oecophoridae): Biology and Morphology

Patricia Chacon & Marta de Hernandez 43

A New Genus and two New Species of Oecophoridae from
Colombia (Lepidoptera)

J. F. Clarke 46

Notes 50

Book Reviews 55

Cover Illustration: Speyeria nokomis pupa from Chihuahua State, Mexico. See
Scott & Mattoon article, page 13, this issue.

Volume 20 Number 2 Summer, 1981(82)

IN THIS ISSUE

Date of Publication: September 20, 1982

International Nepal Himalaya Expedition for Lepidoptera Palaearctica
(ENHELP) 1977, Report No. 1: Introduction and Lycaenidae

Oakley Shields 65

Intergradation between Callophrys dumetorum oregonensis and

Callophrys dumetorum affinis in Northwestern U.S. (Lycaenidae)

James A. Scott & John A. Justice 81

Chromosome Studies in Sixteen Species of Indian Pyralid
Moths (Pyralidae)

P. K. Mohanty & B. Nayak

86

The Biological and Systematic Significance of Red Fecal and Meconial
Pigments in Butterflies: A Review with Special Reference to

the Pieridae

Arthur M. Shapiro 97

On the Nomenclature of Colias alfacariensis Berger 1948
(Lepidoptera: Pieridae)

Otakar Kudrna 103

Some Little-Known U. S. Publications on Lepidoptera I

Cyril F. dos Passos 111

Errata 122

A New Species of Adelpha (Nymphalidae) from Parque Nacional
Braulio Carrillo, Costa Rica

Philip J. DeVries & Isidro Chacon Gamboa 123

Japanese Literature 127

Cover Illustration: The Marsyandi River Valley of Manang, Nepal. See 0.
Shields’ article, page 71, this issue.

Volume 20 Number 3 Autumn, 1981(82)

IN THIS ISSUE

Date of Publication: April 1, 1983

Role of an Ornamental Plant Species in Extending the Breeding
Range of a Tropical Skipper to Subtropical Southern Texas

(Hesperiidae)

Raymond W. Neck 129

Japanese Literature 134

Hipparchia azorina (Strecker, 1899)(Satyridae) Biology, Ecology
and Distribution on the Azores Islands

Steffen Oehmig 136

Courtship Behavior of the Dainty Sulfur Butterfly, Nathalis iole

with a Description of a New, Faculative Male Display (Pieridae)

Ronald L. Rutowski 161

On the Supernumerary Chromosomes of Tarache tropica Guen.

(Lepidoptera: Noctuidae)

P. K. Mohanty & B. Nayak 170

An Apparent Interspecific Fi Hybrid Speyeria (Nymphalidae)

James A. Scott 174

A New Record of Vanessa virginiensis ‘‘ab. ahwashtee'’ from Northern
California (Lepidoptera: Nymphalidae)

Arthur M. Shapiro 176

The Colonization of Violets and Speyeria Butterflies on the Ash-
Pumice Fields Deposited by Cascadian Volcanoes
Paul C. Hammond

179

Book Review

192

Cover Illustration: Artist’s rendition of courtship behavior in the dainty sulfur,
Nathalis iole. Figure was drawn from a 35 mm photograph by Patricia Rutowski. See
page 33, R. Rutowski,

Volume 20 Number 4 Winter, 1981(83)

IN THIS ISSUE

Date of Publication: April 1, 1983

Butterfly Taxonomy: A Reply

Lee D. Miller & F. Martin Brown 193

Nomenclature, Taxonomy and Evolution

Paul R. Ehrlich & Dennis D. Murphy 198

Editorial Note 205

Acknowledgment 206

Notes on the Life History and Baja California Distribution of
Chlorostrymon simaethis sarita (Skinner) (Lepidoptera:

Lycaenidae)

John W. Brown 207

Biology and Immature Stages of Australian Ethmiid Moths
(Gelechioidea)

Jerry A. Powell 214

Field Study of Phyciodes batesii (Reakirt) and P. tharos (Drury) from

a Site in the Black Hills, South Dakota (Lepidoptera: Nymphalidae:
Melitaeinae)

Clifford D. Ferris 235

Notes: Afi Aberration of Glaucopsyche lygdamus (Lycaenidae) with
a Complete Scolitantidine Dorsal Pattern
A Reared Gynandromorph of Tatochila (Pieridae)

Two Homoeotic Pieris rapae of Mexican Origin (Pieridae)
An Apparent “Intersexual” Colias eurytheme (Pieridae)

Arthur M. Shapiro 240

Two New California Catocala Subspecies (Noctuidae)

John W. Johnson 245

Notes 249

Book Review 251

Index for Volume 20 253

Cover Illustration: Scanning electron micrograph of Australian E. postica
eggshell on nylon mesh (x42) by Barry Filshie. See J. Powell article, page 217, this

issue.

INSTRUCTIONS TO AUTHORS

Manuscript Format; Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, on 8V2 x 11 inch paper with wide margins. Number all pages
consecutively and put author’s name at top right corner of each page. If your typewriter
does not have italic type, underline all words where italics are intended. Footnotes,
although discouraged, must be typed on a separate sheet. Do not hyphenate words at the
right margin. All measurements must be metric, with the exception of altitudes and
distances which should include metric equivalents in parenthesis. Time must be cited on
a 24-hour basis, standard time. Abbreviations must follow common usage. Dates should
be cited as example; 4. IV. 1979 (day-arabic numberal; month-Roman numeral; year-
arabic numeral). Numerals must be used before measurements (5mm) or otherwise up to
number ten e.g. (nine butterflies, 12 moths).

Title Page: All papers must have the title , author’s name, author’s address, and any
titular reference and institutional approval reference, all on a separate title page. A
family citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.

Abstracts and Short Papers: All papers exceeding two typed pages must be ac-
companied by an abstract of no more than 300 words. An additional summary is not
required.

Name Citations and Systematic Works: The first mention of any organism should
include the full scientific name with author (not abbreviated) and year of description. New
descriptions should conform to the format: male: female, type data, diagnosis, distribu-
tion, discussion. There must be conformity to the current International Code of
Zoological Nomenclature. We strongly urge deposition of types in major museums, all
type depositions must be cited.

References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbrevations must conform to the World List of
Scientific Periodicals. Do not underline periodicals. If four or less references are cited,
please cite in body of text not in Literature Cited.

Tables: Tables should be minimized. Where used, they should be formulated to a size
which will reduce to 4 x 6V2 inches. Each table should be prepared as a line drawing or
typed with heading and explanation on top and footnotes below. Number with Arabic
numerals. Both horizontal and vertical rules may be indicated. Complex tables may be
reproduced from typescript.

Illustrations: Color must be submitted as a transparency (i.e., slide) ONLY, the quality
of which is critical. On request, the editor will supply separate detailed instructions for
making the most suitable photographic ilustrations. Black and white photographs should
be submitted on glossy paper, and, as with line drawings, must be mounted on stiff white
cardboard. Authors must plan on illustrations for reduction to the 4 x 6V2" page.
Allowance should be made for legends beneath, unless many consecutive pages are used.
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Legends should be separately typed on pages entitled “Explanation of Figures”.
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Review: All papers will be read by the editor(s) & submitted for formal review to two
referees. Authors are welcome to suggest reviewers, and if received, submit name &
comments of reviewers.

Volume 19 Number 4 Winter, 1980(81)

IN THIS ISSUE

Date of Publication: December 1, 1981

Enzyme Electrophoretic Studies on the Genetic Relation-
ships of Pierid Butterflies (Lepidoptera: Pieridae) 1.

European Taxa

H. J. Geiger 181

A New Tortyra from Cocos Island, Costa Rica (Lepidoptera:
Choreutidae)

John B. Heppner 196

Troidine Swallowtails (Lepidoptera: Papilionidae) in South-
eastern Brazil: Natural History and Foodplant
Relationships

Keith S. Brown, Antoni J. Damman & Paul Feeny 199
Kaiyology of Three Indian Lasiocampid Moths (Lepidoptera)

P. K. Mohanty & B. Nayak 227

Illustrations and Descriptions of some Species of Pyrrhopyginae
from Costa Rica, Panama and Colombia (Hesperiidae)

S. S. Nicolay & G. B. Small, Jr. 230

Errata: Scott 240

Book Review 241

Index: Volume 19

Cover Illustration: Left to right, pupae of Battus polydamas, Parides bunichus,
P. anchises nephalion, P. agavus and P. proneus, three views.

j 9'5'‘ 7 ^

J8&

JOURNAL
OF RESEARCH
ON THE LEPIDOPTERA

Volume 20
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Spnng. 1981(82)

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Journal of Research on the Lepidoptera

20(1): 1-11, 1981(82)

Editorial

The recent appearance of the Miller and Brown A Catalog/Checklist of the
Butterflies of America North of Mexico, published by the Lepidopte lists’ Society,
was a well anticipated event, an event which hopefully would have established a
stable nomenclatorial framework for North American butterfly taxonomy for a long
time to come. There has been an immediate reaction to this work from several
quarters. The critique published below, by Ehrlich and Murphy, has already been
widely circulated and formally supported by a significant number of recognized
authorities. For reasons which are unclear, the editors of The Journal of the
Lepidopterists’ Society refused publication of the critique, although it was properly
first submitted there. We strongly assert that forward movement in any scientific
endeavor cannot be achieved without intelligent consideration and discussion. It
should, consequently, be made quite clear that this journal, as a scientific
publication, can take no position in the matter except to strongly express the desire
to have the issue thoroughly aired. We have invited Miller and Brown to rebut the
criticism, and they have agreed to do so presently. The basic issue is substantive to
all Lepidopterists, not only in this region, but internationally. We cordially extend
an invitation for all relevant considered opinion on the matter.

We take this opportunity to emphasize our support of and good relationship with
the Lepidopterists’ Society. That organization has had a great historical impact on
aU workers in the field in North America, and continues as the organization binding
all Lepidopterists in the region together. We in no way wish to convey any sense of
competition between publications of the Society and this journal. We are all
working toward the same objective, have an enormous body of information and
opinion to disseminate, and will continue to do so in a cooperative fashion.

RHTM

Butterfly Nomenclature: A Critique

Paul R. Ehrlich

and

Dennis D. Murphy

Department of Biological Sciences, Stanford University, Stanford, California
94305

Introduction

There are two main goals of formal, latinized nomenclature. The first is
permitting unambiguous communication about what organisms are being

2

J. Res. Lepid.

discussed. The second is expressing, to whatever degree possible, the
evolutionary (phenetic or cladistic) relationships among those organisms.
These goals are not antithetical, but we contend here that the first has
been almost totally ignored lately by many butterfly taxonomists. In part
this is because of a confusion of the nomenclature of butterflies (the
system of names applied to them) with the taxonomy of butterflies (their
classification into groups). And in part it is due to a nearly automatic trend
for specialists to inflate the taxonomic rank of the group on which they
work.

The basic element of the scientific system of nomenclature is the
latinized hinomen. The binomen is made up of two parts, the generic name
followed by the specific name. The binomen is supposed to be a standard,
stable label, understandable to workers in all locations-”Unlike “common”
names that vary from place to place. Thus while the Americans call a
certain butterfly the “Mourning Cloak” the English call it the “Camber-
well Beauty”, the Germans, “Trauermantel”; Swiss, “Sorgmantel”;
French, “Le Morio”; and Spanish, “Antiopa”; scientists everywhere
should call it Nymphalis antiopa.

The preamble to the International code of Zoological Nomenclature
states: “The object of the Code is to promote stability and universality in
the scientific names of animals.” It is instructive, however, to examine how
well the formal nomenclature of butterflies has achieved stability of
names.

Trends in Generic Nomenclature

We have examined the latinized names in seven standard works on
Nearctic butterflies: Holland (1898, 1931); McDunnough (1938); Ehrlich
and Ehrlich (1961); dos Bassos (1964); Howe (1975); and Miller and
Brown (1981). In what follows we will often refer to these works by the
dates only. We traced the history of only those 1898 names in the
Papilionoidea that were still considered to represent valid species in 1961,
and were not of questionable residence (“strays” picked up occasionally
along the southern border of the U.S.). For example, the 1898 names
Lycaena icarioides, Neonympha phocion, and Melitaea arachne (in 1961
Plebejus icarioides, Euptychia areolata, and Poladryas pola respectively)
were included, while Lycaena ardea (in 1961 considered a subspecies of P.
icarioides), Chlosyne chinatiensis (not described until 1944), and Dircenna
klugii (doubtful resident) were not.

Changes in the generic names of the 262 species that fit these criteria in
1898 were followed through the subsequent six publications. The average
species changed generic name 1.8 times, that is, it has had almost three
different generic names in seven standard works. There have, of course,
also been a large number of changes in the specific name or switches
between specific and subspecific status. Thus three different names in 84

20(1): Ml, 1981(82)

3

years— a new name eve^ 28 years— is a very conservative estimate of the
amount of name changing. Nomenclatural instability, rather than stabiliiy ,
has been the rale.

Consider some examples. Although in the United States the common
name Mourning Cloak has remained stable for the entire eighty-four years,
Holland first called it Vanessa antiopa and then Aglais antiopa, whereas
everyone from McDunnough onward has called it Nymphalk antiopa.
Conversely, while everyone from Holland through Howe called the Spice-
bush Swallowtail Papilio troilus, Miller and Brown suddenly declare it to
be Pterourus troilus. But these have been, relatively stable names.
Satyrodes canthus (1898) became S. eurydice (1931, 1938), then Lethe
eurydice (1961, 1964, 1975), and finally Satyrodes eurydice (1981).

Some species have weathered nearly as many generic names as
publication appearances. Thecla m-album (1898, 1931) has been in
Strymon (1938), Panthiades (1961, 1975), Eupsyche (1964), and Parrhasius
(1981). Others have simply bounced back and forth between genera.
Thecla augustus (1902, 1931) moved to Incisalia (1938), to Callophrys
(1961) back to incisalia (1964), back to Callophrys (1975), and finally back
to Incisalia (1981).

As the example of 'Tnckalia-Callophrys'' augustus indicates, continuing
disagreement on whether or not to split a genus may inflate the number of
generic name shifts. But such cases are only a minor part of a general trend
of fractioning genera. Within the sample set of species there has been a
100 percent increase in the number of genera, and therefore a halving of
the number of species per genus, during the past 50 years.

This generic splitting occurred in two waves. The 262 species were in 46
genera in 1898, 49 in 1931, 69 in 1938, 71 in 1961, 67 in 1964, 72 in 1975
and 100 in 1981. Thus from 1938 to 1975 the number of genera remained
more or less stable. The McDunnough list increased the number of genera
some 40 percent over Holland; Miller and Brown ended that period of
relative stability with a similar increase. Our contention is that the
McDunnough changes were largely justifiable and those by Miller and
Brown largely unjustifiable.

In general the name changes made between 1931 and 1938 reflect the
fractioning of a few very large, sometimes polyphyletic genera like
Chlorippe, Melitaea, Satyrus, Thecla, and Lycaena, and the recognition of
clear cases of priority that did not involve splitting (e.g. Anaea for
Pyrrhanaea, Polygonia for Grapta). The large increase in the number of
genera between 1931 and 1938 is almost entirely accounted tor by the
splitting of two genera, the blues (then Lycaena) and hairstreaks (Thecla).
T^e changes between 1975 and 1981, in contrast, reflect in large part, a
refusal to recognize subgenera in disregard of both basic goals of
nomenclature outlined in the introduction.

Some name changes are almost entirely due to changing “styles” in

4

J. Res, Lepid.

splitting and lumping. Consider the case of Euphydryas editha. No one
doubts that Melitaea cinxia is more closely related evolutionarily to
Euphydryas phaeton than to Boloria pales (all are the type species of their
genera). But Melitaea in the broad sense (including phaeton) was
considered too large to be a single genus and Euphydryas was generally
recognized as having generic status by the late 1930s. Thus Melitaea
editha became Euphydryas editha. The latter name remained stable until
Higgins (1978), recognizing several evolutionary groups witlun Euphydrym,
split up that genus into four genera, concluding that the name of
Euphydryas editha should be changed to Occidryas editha.

Note that splitting Euphydryas from Melitaea does not make the
slightest difference in the amount of relationship communicated by the
specific name. For example the first split focussed attention on the
relatively close relationship hetv^eenEuphydryas phaeton and Euphydryas
editha but obscured the somewhat more distant, but no less important,
relationship between Euphydryas editha and Melitaea (— Chlosyne ^
Charidryas) palla. Whether or not the old Melitaea should have been split
is a matter of taste, especially since relationships within the Melitaeini
(sensu Ehrlich and Ehrlich, 1961) are still not well understood. But we
tend to think it was useful from the point of view of the communication goal
of nomenclature. The advantage to lepidopterists who must frequently
discuss species groups within the oldMelitaea is probably greater than the
inconvenience for non-specialists for whom the large, easily recognized
Melitaea was more useful. As we will discuss below, however, there is no
conceivable justification for splitting Euphydryas.

The change of Graphium marcellus from the genus Papilio to the genus
Graphium was justifiable because the old genus Papilio was polyphyletic.
Graphium marcellus is much more closely related to Lamproptera curius
than to Papilio machaon (or Battus philenor). Only by placing all of the
Graphiini and Papilionini (sensu Munroe and Ehrlich, 1960) into a single
genus could marcellus be made congeneric with machaon without recreat-
ing a polyphyletic entity.

The Rule of Obligatory Categories

In the case of Graphium marcellus the need to change the generic name
was clear— it could not reasonably remain in the genus Papilio (whether it
should be in the genus Eurytides, a subset of the genus Graphium as used
here, is a more difficult question). But how does one evaluate the case of
the proposed splitting of Euphydryas? Here one must somehow balance
the needs of different “user” groups.

There is, fortunately, a taxonomic guideline that is veiy useful in
determining where to draw the line. The key point to remember is that
taxonomists should not be creating nomenclature primarily for their own
use, but as a general tool useful to all biologists. The rule is: obligatory

20(1): 1-11, 1981(82)

5

categories above the species level should be kept conservative. What are
“obligatoiy categories”? Categories are ranks in a hierarchic classification.
Obligatory categories (e.g., Mayr, 1969, p. 89) are those that every animal
must be placed in when it is described: species, genus, family, order,
class, and phylum. It is especially important to follow the rule with respect
to genera. As Ernst Mayr wrote (1969, p. 239): “Splitting is particularly
deleterious on the generic level. The generic name is part of the scientific
name of an organism and can therefore be employed more advantageously
to indicate affinity than can the name of any of the other higher
categories.”

If this rule is followed then there can be the best of both worlds.
Communication with non-specialists is facilitated, because it is normally
the obligatory categories that are used for this purpose. But there remains
a wealth of non-obligatory categories with which a taxonomist can
communicate finer points of difference -"Superfamilies, subfamilies, tribes,
subtribes, subgenera and species groups, to name the ones most used by
Lepidopterists.

Effects of ignoring the Rule

As we have seen, it is primarily generic splitting that has destroyed the
stability of the latinized names of butterflies. For example Higgins’ (1978)
splitting of Euphydryas is an attempt to raise what would be reasonable
species groups (or weak subgenera) to generic status. He provided no
discussion of the basis on which he decided that the relationship between
E. editha and E. chalcedona was more important to communicate in the
binomen than that between E. chalcedona and E. phaeton. Higgins also
promoted the weak genus Euphydryas to full tribal level, without
consideration of what that categoric inflation meant for the taxonomy of
the checkerspots (and nymphalids) in general. Presumably the old tribe
Melitaeini (and other tribes of the Nymphalidae such as the Argynnini)
would have to be raised to subfamily level— implying that the checker-
spots are a group as distinct from, say, the fritillaries as the Papilioninae
are from the Pamassiinae. It would further require that the Nymphalidae
(sensu 1961) be divided into eight or so poorly defined famifies.

Miller and Brown (1981) appear to have seen the absurdity of tribal
status for Euphydryas, but nonetheless recognized Higgins’ daughter
genera Occidryas, Eurodryas, and Hypodryas. This, however, leads to
something equally absurd within the Melitaeini: considering the dif-
ferences between Phyciodcs and Occidryas to be of the same order as those
between Occidryas and Hypodryas. In fact Euphydryas is an extremely
cohesive group, not just morphologically but in its behavior, reproductive
biology, chemistry of host plant choice, and allozyme genetics.

Perhaps even more important, Euphydryas are now widely used in the
research of population biologists. It would be as ill-advised to change the

6

J. Res. Lepid.

scope of that generic name today as it would have been to accept the
proposal (made some years ago) that the generic name of Drosophila
melanogaster be changed. It is precisely that sort of nonsense that
frequently leads evolutionists, ecologists and others to ignore the impor-
tant contributions of taxonomists and damages the reputation of taxonomy
as a discipline.

Again, Euphydryas is hardly an isolated instance. Pieris rapae, a name
stable for more than a century and enshrined in thousands of papers in the
economic literature is now supposed to be changed to Artogeia rapae. It is
true that the type of Pieris, P brassicae, is morphologically and chromo-
somally a rather unusual species, but the difference between the two could
have remained expressed, as it has for decades, by subgenera. It is
doubtful that most scientists will accept this change anyway, any more
than they will use Occidryas editha. The problem then becomes the
acceptance of the change by some scientists and the confusion that ensues.

Other examples abound in Miller and Brown (1981). Zerene was
considered a subgenus of Colias in the careful revision of the Pieridae by
Klots (1933) and in his treatment of Colias in Ehrlich and Ehrlich (1961).
Should not the grounds on which that judgment was reversed be
published? Similarly what is the biological justification for spHtting up
Eurema. It seems a most uniform assemblage, and again Klots found no
reason for fragmenting it into several genera. Taxonomies should not be
modified by fiat, but only with the publication of thorough analyses backed
by data.

Among the least warranted changes in the Miller and Brown list is the
resurrection of a series of antique generic names, mostly proposed by
Huebner (1819), within Papilio (sensu 1975). Are we to assume that the
judgment of Jacob Huebner, based on the ve^ limited material and
information of 150 years ago, should take precedence over that of Eugene
Munroe, a modern taxonomist with access to virtually aU papilionid
species? Huebner was a giant among his contemporaries, but in his time
the concept of a genus was far from its modem state. In Munroe ’s classic
paper on the Papilionidae (1961) he states: “...I have failed to find simple
and reliable differentiating characters for what appear to be the natural
groups of Papilionini. I therefore include all the species in a single genus.”
And yet everyone is now expected to drop names used since childhood
and, for example, call the tiger swallowtail Pterourus glaucus.

Huebner’s approach to differentiation at the generic level is particularly
evident where he (1819) proposed the generic name Heraclides for Papilio
thoas. The three species available to Huebner for consideration and
placed by Miller and Brown in Heraclides (thoas, cresphontes, and
androgens) were actually divided by Huebner into two genera. The super-
ficially almost indistinguishable thoas and cresphontes were placed in
Heraclides and androgens was split off in Calaides. Furthermore, in the

20(1): 1-11, 1981(82)

7

same publication the generic names Jasoniades and Euphoeades were
proposed for Papilio turnm andP. glaucus respectively —two color morphs
of the same swallowtail! Both names were buried by the priority of
Pterourus, which had appeared 40 years previously and been rightfully
ignored*

It seems unnecessary to discuss in detail the pointless fragmentation of
genera Antkocaris, Lycaena (semu 1938), Boloiia, Chlosyne, Nymphalk,
and Precis, and more complex cases where certain splitting may well have
been justified (e.g., Philotes, Plebejus, some Theclini, Euptychia). One
justification for the recent generic splitting has been that Europeans have
done it (see Miller and Brown, 1979, also pp. ix and xvii of Ferris and
Brown, 1981 and the nearly identical nomenclature in Miller and Brown,
1981). It is true that there has been a trend in Europe toward having a
single species in each genus (which at completion will completely destroy
the utility of binomial nomenclature). There was once a trend for
European amateurs to name every individual, too (the “aberration’’
craze). But there is not the slightest reason for American lepidopterists to
follow in their footsteps.

Another apparent justification for the recent ultra-splitting seems to
have been the mistaken notion (Miller, 1981, p. 54) that the nomenclatural
principle of priority is more rigid than it actually is (see Mayr et al, 1971)
and that “the primary law of taxonomy involves the concept of binominal
nomenclature” ^iUer, 1981, p. 53). The primary laws of taxonomy have
to do with how one arranges organisms into groups, not how one chooses to
assign names (or numbers, or symbols) to those groups in order to
communicate about the groups and their arrangement. An unfortunate
emphasis on names rather than organisms seemingly has led to attempts to
recognize the maximum number of genera— a sort of bizarre “conservation
of generic names. ” lire resultant trend toward all genera being monotypic,
with the concomitant ignoring of subgenera, makes nomenclatural expres-
sion of relationships below the tribal level increasingly difficult.

In summary, the Miller- Brown catalogue is a superb historical review
and bibliographic tool, and all who study butterflies owe them a great debt
for their enormous effort. But their choice of names is simply unacceptable.
Their names will not be used by most scientists working with butterflies,
including taxonomists, and it mU simply make communication even more
difficult if some butterfly taxonomists persist in using them on the
assumption that to do so is somehow “scientific” or “modem,” or that the
names are in some way “official.” Remember there is no mle that, just
because someone has proposed a new genus or resurrected an old one, the
judgment must be accepted.

Recommendations

Unfortunately the Miller and Brown (1981) names have already been

8

J. Res. Lepid.

used in two otherwise excellent books directed to laypersons (Pyle, 1981;
Ferris and Brown, 1981), and our personal contacts and correspondence
indicate widespread distress with the numerous name changes. To avoid
further confusion we would like to make some recommendations for
stablilizing the nomenclature of North American butterflies:

1) . The generic nomenclature in Howe should be adopted, and no
changes accepted in it except where required because of clear polyphyly or
highly distorted “balance” (Mayr, 1969, p. 241). We recommend this not
because we think that the nomenclature in Howe is perfect. It is, however,
the widely available major compendium on Nearctic butterflies. It further-
more has a reasonable nomenclature that does not constitute a major
departure from other post- 1950 works. We are not sure that the splitting of
Speyeria from Argynnis or Euphydryas from Melitaea was originally
justified, but we are certain that to change such widely accepted names
now would be foolish. It might be wiser to recognize Eurytides as a full
genus as Munroe and Ehrlich (1960) did, but Graphium can be mono-
phyletic and is widely used. Let it be!

2) . Editors should routinely rej ect any work that suggests generic name
changes from those in Howe if it does not contain a thorough biological
justification for the change — polyphyly or imbalance. Papers should also
be rejected unless all changes in rank are accompanied by a discussion of
their consequences for the balance of the system as a whole. “Inclusion of
all. ..species within a single genus. ..fails to recognize their wide generic and
specific differences” (Higgins, 1978) and “something is going on with the
coppers” (Miller and Brown, 1979) do not meet these criteria. Munroe’s
1949 discussion “Some remarks on the genus concept in Rhopalocera”
can still provide excellent guidance in this area.

The problem of generic splitting is now so serious that we believe the
butterflies are about to go the way of the birds, where it is the common
names that are used by virtually everyone for communication and the
binomens, in addition to being unstable, give few cues to relationships. Do
we all reaUy want to use names (Pyle, 1981) like “immaculate green
hairstreak,” “Cuban crescentspot,” and “western black swallowtail”
instead of the shorter Callophrys affinus, Eresia frisia, and Papilio bairdUP.
Or names hke “goatweed butterfly,” “question mark,” “waiter,” and
“crimson-banded black,” that give no clues to affinity? In an attempt to
arrest this trend we are going to petition the Lepidopterists' Society
officially to adopt a list of approved generic names based on Howe (1975)
to be used in its publications, and to appoint a diverse board to oversee its
(hopefully rare!) revision.

Regarding species-level taxonomy, matters are more complicated. We
won’t go into the evolutionary problems here, except to say that they are
much more complex than usually indicated in popular works on butterflies.
Those interested in more details can get access to the literature through

20(1): 1-11, 1981(82)

9

Ehrlich (1961), Mayr (1963), Ehrlich and Raven (1969), and Grant (1981).
Suffice it to say that splitting at the species level can be justifiable because
lumping can conceal important biological differences (e.g., Lycaena
phlaeas may really be very genetically different from L. hypophlaeas). Here
again, though, we would tend to be conservative for purposes of
communication and not deviate from the treatment in Howe without
substantal evidence to justify the change.

We also feel strongly that revisions at the species level of Nearctic
butterflies should be accompanied by exhaustive numerical taxonomic
analysis and/or careful work on the biology of the organisms. The recent
description of Boloria acrocnema (Gall and Sperling, 1980) could serve as
a model for the sort of detailed analysis that should support any proposed
new specific names in the Nearctic fauna. Whether or not one accepts their
judgment, the basis for that judgment is clearly and unambiguously laid
out. Our fauna is now so well known that little is to be gained by reshuffling
names, or creating new ones, on the basis of genitalic dissections alone.
Contraiy to mythology, the shape of the genitalia is no magic indicator of
taxonomic status (Shapiro, 1978).

On the other hand, little damage is done by splitting in the description of
new subspecies. Since the classic work of Wilson and Brown (1953) it has
been clear that most subspecies are not biological entities, and they
relatively rarely figure in the evolutionary literature. But subspecies
names do call attention to certain patterns of phenetic variation, do
communicate information about those patterns among specialists, are
useful politically in attempting to prevent the extinction of genetically
distinct populations, and give pleasure to butterfly collectors. In short
they can be conveniently used by specialists and equally conveniently
ignored by others.

At the family- subfamily level we contend that the basic treatment in
Ehrlich (1968) and Ehrlich and Ehrlich (1967) should be retained. It is the
only recent work that considers the entire breadth of the butteiflies and
the balance of the groups within them in a context of other Lepidoptera
and the insects as a whole. The butterflies are an evolutionary uniform
group as insects go, and there is no reason for them to be divided into more
than 5-6 families. People who revise one group invmably become
impressed with the diversity within that group and want to raise the rank of
the taxon they are revising. This urge leads to serious imbalances. For
example, if the Satyrinae are considered to be a family with Euptychia and
Lethe in separate subfamilies an imbalance is created. That is because
there is much more difference between Papilio and Graphium (only in
different tribes) than between Euptychia and Lethe.

This is not to say that the Ehrlich treatment necessarily should be
permanent. Perhaps, for example, the Riodininae should be considered a
family or the Libytheidae dropped to subfamily status. But suggested

10

J. Res. Lepid.

changes at that level must be accompanied by a consideration of the
relationships of all the major groups of the butterflies worldwide and the
balance of insect taxonomy as a whole. Any changes should be based on
more evidence or better techniques than the Ehrlichs used. Interestingly,
a recent cladistic reanalysis of Ehrlichs’ data (Kristensen, 1976) found no
reason to alter the higher classification beyond dropping the libytheids to
subfamily status. ^

Finally, we would like to reiterate a plea that has been made before
(Remington, 1948; Munroe, 1960). It is that research on butterflies be
focused much more strongly on studying the biology of these fascinating
creatures and much less on continual shuffling of names. Butterflies are
increasingly the subject of important research in ecology, evolution,
animal behavior, and conservation biology. These are areas in which both
professional biologists and amateur lepidopterists have made and can
continue to make substantial contributions. Let’s keep our nomenclature
stable so that we can continue conveniently to talk about it among
ourselves and tell the world about it unambiguously.

Acknowledgments. The following people have been kind enough to read and
criticize this manuscript and, though not necessarily agreeing with all the points
made, to let us state that they support the general principles put forth in
it: Richard A. Arnold, George T. Austin, Carol Boggs, M. Deane Bowers, John M.
Bums, Peter F. Bmssard, Frances S. Chew, Philip J. DeVries, Ernst J. Domfeld, J.
Donald Eff, Thomas C. Emmel, David Faulkner, Hugh A. Freeman, La^ence F.
Gall, Lawrence E. Gilbert, L. Paul Grey, John E. Hafemik, Jane L. Hayes, Cheryl E .
Holdren, Richard W. Holm, Nelson D. Johnson, Roy O. Kendall, Alexander B.
Wots, Robert L. Langston, Ralph W. Macy, John H. Masters, Paul A. Opler, Larry
J. Orsak, Naomi Pierce, Austin P. Platt, Jerry A. Powell, Robert M. Pyle, Mm-k D.
Rausher, Charles L. Remington, James A. Scott, Arthur M. Shapiro, Oakley
Shields, Michael C. Singer, Don B. Stallings, Maureen Stanton, Bmce Tabashnik,
John H. Thomas, Fred T. Thome, Ward B. Watt, David A. West, Raymond R.
White, Bmce A. Wilcox, and Ernest H. Williams.

TTiis work has been supported in part by a grant from the Koret Foundation of San
Francisco.

Literature Cited

DOS PASSOS, c. F., 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc.
Mem. no. 1.

EffliLiCH.P. R., 1958. The comparative morphology, phylogeny and higher classifica-
tion of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. BuU.
39: 306-370.

__ — _ — , 1961. Has the biological species concept outlived its usefulness? Syst.
Zool. 10: 167-175.

EHRLICH, P. R. & A. H. EHRLICH, 1961. How to Know the Butterflies. W. C. Brown,
Dubuque.

_, 1967, The phenetic relationships of the butterflies. 1. Adult taxonomy

and the non-specificity hypothesis. Syst. Zool. 16: 301-317.

20(1): 141, 1981(82)

11

EHRUCH, P. R. & P. H. RAVEN, 1969. Differentiation of populations. Science 165:
12284232.

FERRIS, c. D, &F. M. BROWN, eds,, 1981. Butterflies of the Rocky Mountain States. Univ.
i of Oklahoma Press, Noman.

I GALL, L. F. & F. A. H. SPERLmG, 1980. A new high altitude species of Boloria from south-
western Colorado (Nymphalidae) with a discussion of phenetics and hierarchical
j decisions. J. Lepid. Soc. 34: 230-262.

OTANT, VERNE, 1981. Plant Speciation. Second Edition. Columbia Univ. Press,
I New York.

mCGiNS, L. G., 1978. A revision of the genus Euphydryas Scudder (Lepidoptera:

Nymphalidae). EntomoL Gaz. 29: 109-115.

HOLLAND, W. J., 1898. The Butterfly Book. Doubleday, New York.

. 1931. Tlie Butterfly Book. Revised edition. Doubleday, New York.
HOWE, w. E., ed., 1975. The Butterflies of North America. Doubleday, New York.
HUEBNER, JACOB, 1819. Verzeichness bekannter Schmetterlinge. Augsburg.

KLOTS, A. B., 1933. A generic revision of the Pieridae (Lepidoptera) together with a
study of the male genitalia. Ent. Americana 12: 139-242.

I®ISTENSEN, N. P., 1976. Remarlw on the family-level phylogeny of the butterflies
(Insecta, Lepidoptera, Rhopalocera). Z. Zool. Syst. Evolut.-forsch. 14: 25-33.
MAYR, ERNST, 1963. Animal Species and Evolution. Harvard Univ. l^ess, Cambridge.

- , 1969. Principles of Systematic Zoology. McGraw-Hill, New York.

MAYR, E., G. G. SIMPSON & E. EISENMANN, 1971. Stability in zoological nomenclature.
Science 174: 1041-1042.

McDUNNOUGH, J., 1938. Check list of the Lepdioptera of Canada and the United
States of America. Part 1, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1: 1-272.
MILLER, L. D., 1981. Taxonomy. Ch. 4 in C. D. Ferris and F. M. Brown, eds., Butter-
flies of the Rocl^r Mountain States. Univ. of Oklahoma Press, Norman.

MILLER, L. D. & F. M. BROWN, 1979. Studies in the Lycaeninae (Lycaenidae). 4. The
higher classification of the American coppers. Bull. Allyn Mus., no. 51.

, 198 1. A catalogue/checklist of the butterflies of America north of Mexico.
Lepid. Soc. Mem. no. 2.

MUNROE, E. G., 1949. Some remarks on the genus concept in Rhopalocera. Lep.
News 3: 3-4,

, 1960. Professional and amateur reseaich in Lepidoptera. J. Lepid. Soc.
14: 1-4.

1961. TTie classification of the Papilionidae (Lepidoptera). Can. Ent.

Suppl. 17.

PYLE, R. M., 1981. The Audubon Society Field Guide to North American Butterflies.
Knopf, New York.

REMINGTON, c. L., 1948. Lepidoptera biology. ..open for study. Lep. News 2: 37.
SHAPIRO, A. M., 1978. The assumption of adaptivity in genital morphology. J. Res.
Lepid. 17: 68-72.

wn^ON, E. o. & W. L. BROWN, JR., 1953. The subspecies concept and its taxonomic
application. Syst. Zool. 2: 97-111.

Journal of Research on the Lepidoptera

20(1): 12-16, 1981(82)

Early Stages of Speyeria nokomis (Nymphalidae)

James A. Scott and
Sterling O. Mattoon

60 Estes Street, Lakewood, Colorado 80226 and

2109 Holly Avenue, Chico, California 95926

Abstract. The egg, larval stages, pupa, and developmental period of S.
nokomis from the United States and Mexico are described and illustrated.

Introduction

Comstock (1928, 1940) and Skinner (1907) briefly described the egg,
and young larva of nokomis, but not older larvae or pupae. This paper
describes and illustrates the egg, larval, and pupal stages.

Early stages are based on several hundred eggs, larvae, and pupae
reared from eggs laid by females from Elko County, Nevada {S. n.
apacheana (Skinner)), Taos and San Juan Counties, New Mexico, the
White Mountains of Arizona (S. n. nokomis (Edwards)), and Durango and
Chihuahua states Mexico (S. n. coerulescens (Holland)). About 100 larvae
were preserved from San Juan County, New Mexico (in J. Scott coll.), 10
or less from each of the other sites (in S. Mattoon coll.). Larvae were reared
on Viola, including V. nephrophylla.

Early Stages Description

Egg: Cream colored when laid, becoming tan after a few days. Strongly
ribbed vertically, with ribs rising to several peaks surrounding the
micropyle (Fig. 11). Numerous horizontal crossbars connect the vertical
ribs. Incubation period is about ten days in the lab at about 20®C and
constant light. Larva, Figs. 6-11: There are six instars. Head capsule
widths average approximately 0.35, 0.6, 1.0, 1.4, 2.4 and 3.5 mmfor the six
instars, an average of 60% growth at each moult, based on measurements
of about 50 New Mexico head capsules. The head capsule has a dark area
after the first instar (Fig. 1 1); the dark area is black in later instars. Instars
1 and 2 are cream-colored mottled with brown, with a light dorsal band and
a light lateral band running through the spiracles. Brown mottling occurs
elsewhere especially on intersegmental membranes. Body darker around
sclerotized areas. First instar brown mottling of S. n. coerulescens is very
similar to that of ssp. nokomis. Setal pattern of all instars appears identical
in these two ssp. (Fig. 10). Instars 3 and 4 are orangish cream (head pale
orangish brown), with black spots and lines like instars 5 and 6, except

20(1): 12-16, 1981(82)

13

Figs. 1-6. Pupae. Figs. 6-9. Mature larvae. Figs. 1-4, 6-7, Chihuahua State,
Mexico. Fig. 8, Taos County, New Mexico. Figs. 5, 9-11, San Juan
County, New Mexico.

middorsal pale band whitish in color in instars 3-4. Mature (5-6) larva
orangish ochre (head pale reddish brown) with black spots and lines (Figs.
6-9); middorsal pale band also orangish ochre. Mature larva has a light
dorsal band on abdomen, a lateral light band just ventral to spiracles. Scoli
brown in color with black tips, although dorsal side of two lateral rows of
scoli cream in color, subdorsal scoli cream on prothorax and 9th abdominal
segment scoli dark brown. Dark brown patches occur around scoli. Two
transverse black bands occur behind scoli on dorsum of most segments.

A difference was noted between populations in the color pattern of later
instar larvae. Instars 3-6 of ssp. nokomis (from New Mexico and Arizona),
and apacheana have ground color orangish ochre, whereas ground color of
coerulescem larvae is light yellow.

Pupa: Pupa orangish ochre with black markings (Figs. 1-5). The extent of
black varies especially on the wing cases. Pupae of ssp. nokomis are fairly
dark, with wing cases mostly black (Fig. 5). Pupae of White Mountains,
Arizona nokomis, and apacheana, are lighter, with lighter wing cases.
Pupae of coerulescens (Fig. 1) are still lighter, with predominantly light
wing cases (although some individuals are dark, Fig. 2) and the abdomen is
somewhat lighter also.

14

J. ^s. Lepid.

IT 2T 3T lA 2A 3A 4A 5A 6A 7A 8A 9A

Fig. 10. Setal maps of larvae. Solid lines surround legs (L), prolegs (P), scoli (S),
or sclerotized areas. Dash lines surround less well sclerotized areas.
Small ovals on IT (T ” thorax) and 1-8A (A ~ abdomen) are spiracles.
Stippling on first and second instar shows dark pattern of a typical
segment.

Fig. 1 1 . Head capsules of first, second, third, and mature larva, including view of

left side of first, second, mature larva; terminal segments of first lateral
and posterior view), second, and mature larva; and egg (side view and
dorsal view of micropyle). Stippling indicates dark areas; setae and color
pattern of mature larva head is drawn on opposite sides of the head
capsule.

Developmental Period and Male-Female Emergence Lag

Developmental period from oviposition to emergence of adults is 61 days
for males, 69 days for females, indoors at about 20°C for ssp. nokomis, a
difference of 8 days. However, the lab is much warmer on average than
nature, so development is probably longer in nature. Because first stage
larvae overwinter and adults fly mainly from late July to September, it is
reasonable to estimate a 4 month developmental period in nature for
females. With a 4 month or 122 day developmental time, the 8 days
increases to a 14 day difference between male and female emergence in
nature (8/69 ™ 14/122). Males precede females in emergence in most
insects. S. nokomis males may appear in late July or early in August, but
females normally appear much later in mid or. late August. Scott (1977)

20(1): 12-16, 1981(82)

16

demonstrated mathematically that male butterflies (and most inverte-
brates) should precede females in emergence; he showed that females
should emerge when males are most abundant (which is later than when
most males emerge) in order to maximize the number of matings for males
smd minimize the time mjuired for females to find a mate. This is the
evolutionary explanation for males preceding females in emergence. The
lag is implemented physiologically by the longer developmental time for
females just noted. Scott (1977) noted that the most important factors
influencing the optimum length of the lag in emergence are lifespan of
males and standard deviation (spread) of emergence time of males, and
that the lag should be small only if females mate often. A reviewer suggests
fliat emergence lags occur because “females are larger than males necessi-
tating a longer feeding period. Also, females must accumulate the proteins
and lipids that will be used for egg production.’* Actually, the large size of
females is a consequence of their longer feeding period, not vice versa.
Also, Ronald Rutowski (pers. comm.) found that in Pieris protodice Bd. &
LeC., larger males produce larger spermatophores (which are digested by
females and used for producing eggs) and females prefer to mate with
larger males. So, according to these findings there are quite valid reasons
why males should feed longer to grow to large size (they mate more often
when larger and their mates produce more eggs). Furthermore, it is not
clear that females should be larger, because a small female would use less
energy in flight so would have more energy available for producing eggs,
mid a given size of female could produce more offspring merely by
producing smaller eggs. Plus, larger males might fly farther and mate with
more females, and the larger spermatophores produced by larger males
stretch the female’s bursa copulatrix more and are digested slower so the
female will remate later. Of course, thousands of species of Lycaenidae
exist very well with small sized females. The essential point of this
discussion is that there are clear and obvious reasons why males precede
females in emergence, which have been independently verified by
Wiklund and Fagerstrom (1977), whereas it is not at all clear whether
females should be smaller or larger than males.

literature Cited

COMSTOCK, J. A., 1928. Studies in Pacific coast lepidoptera. Bull. So. Cal. Acad.
Sci. 27: 80-89.

, 1940. Argynnid notes. Bull. So. Cal. Acad. Sci. 39: 75-77.

SCOTT, d. A., 1977. Competitive exclusion due to mate-searching behaviour, male-
female emergence lags, and fluctuation in number of progeny in model inverte-
brate populations. J. Anim. Ecol. 46: 909-924.

SKINNER, H., 1907. Note. Entom. News. 18: 318.

WKLUND, c. & T. FAGERSTROM, 1977. \yhy do males emerge before females? A
hypothesis to explain the incidence of protandry in butterflies. Oecologia 31:
153-158.

Journal of Research on the Lepidoptera

20(1): 16-17, 1981(82)

Supernumerary Chromosomes in the Domesticated
Eri-SiUcmoth, Philosamia ricini (Satumidae: Lepidoptera)

K. B. Padhy and B. Nayak

Department of Zoology, Bonaigarh College, Sundergarh, Orissa, INDIA

Abstract. Supernumerary chromosomes were detected in spermatocytes
of the domesticated silkmoth, P. ricini. The supernumeraries varied from
one to four in number and occupied an extranuclear position. In cases of
single supernumeraries in a cell, it passes to one pole during anaphase I.
Where two supernumeraries were present they form a bivalent in metaphase
I and pass to two different poles during the next anaphase. All super-
numeraries are eliminated from the meiotic cell population forming
“micronucleus -like bodies” during interphase. Their “parasitic” way of
chromatin elimination is discussed.

Microchromosomes occurring as supernumerary elements in the karyo-
type of lepidopteran species have been reported by many workers
(DeLesse, 1960; Bigger, 1977; Nayal, 1974, 1978). Philosamia ricini
possess a haploid chromosome number (n ™ 14) in the germ line. While
examining the chromosome preparations of the testes from a population of
domesticated P. ricini from Orissa, it was observed that in addition to the
normal 14 bivalents in the diplotene and metaphase I, about 4% of the
primaty spermatocytes revealed 1-4 minute chromosomes which are
supernumerary elements. The spermatocytes possessing supernumeraries
showed their number as 1, 2 or 4. When they were more than one in
number, pairing could occur between two such supernumerary chromo-
somes. A third supernumerary, if present, remained an univalent. All
supernumeraries observed appear to be negatively heteropycnotic in
metaphase I and positively heteropycnotic in the interphase. Therefore,
such elements must not have produced by precocious resolution of any of
the bivalents into univatents, nor to breakage of the normal complements,
since they occur along with the normal chromosome complement. They
have been observed to occupy a position outside the metaphase plate (Fig.
1), similar to such elements described by DeLesse (1960). Furthermore,
missegregation of supernumeraries were observed in anaphase I when
they numbered more than one, and when one supernumerary occurred it
passed unresolved to one pole only. At the telophase I and the interphase
they occupied an extranuclear position, forming a micronucleus -like body.
They may be eliminated during germ cell development, therefore, without

20(1): 16^17, 1981(82)

17

Fig. 1. First metaphase spermatocyte of P. ricini indicating 14 bivalents and the
supernumerary chromosome bivalent placed in an extranuclear position
(arrow).

playing any significant role in influencing the development of the
organism.

Although the exact nature of such supernumeraries is not yet known,
these elements may represent a case of chromatinic elimination through
heterochromatinisation of specific segments of normal bivalents. White
(1977) argues that the presence of supernumerary chromosomes in many
natural populations of insects may be produced due to metabolic
disorders induced by agriculture chemicals. Such an explanation is not
apparent here, since the species is maintained under laboratory conditions.
Rather their both heteropycnotic and possibly deleterious nature more
probably permits their classification as a “deleterious parasite” as
suggested by Ostergreen (1945).

Acknowledgments: We are thankful to the U. G. C. India for providing financial
assistance for the present work. Our grateful thanks are also due to Professor C. C.
Das, Berhampur University.

literature Cited

BIGGER, T. R. L., 1977. Karyotypes of three species of Lepidoptera including an inves-
tigation of B-chromosomes in Pieris. Cytologia.

DeLESSE, H., 1960. Speciation et variation chromosomique chez les Lepidopteres
Rhapaloceres. Ann. Sci. Nat Zool. 12, ser. 2: 1-223.

NAYAK, B., 1974, Ph.D. Thesis, Utkal University, Orissa, India.

, 1978. On the supernumerary m-chromosomes in two species of Lepid-
optera. J. Entom. Res. 2: 112-114.

NUR, u., 1977. Maintenance of parasitic B-chromosomes in the grasshopper,
Melanoplus femur ruhrum. Genetics, 87: 458-499.

OSTERGREEN, G., 1946. Parasitic nature of extra chromosomal fragments. Bot.
Notiser. pp. 157-163.

WHITE, J. J. D., 1973. Animal Cytology and Evolution, Cambridge University press,
London, pp. 329.

Journal of Research on the Lepidoptera

20(1): 18-35, 1981(82)

Geographic Variation and Ecology of
Hesperia leonardus (Hesperiidae)

James A, Scott

and

Ray E. Stanford

60 Estes Street, Lakewood, Colorado 80226 and
720 Fairfax Street, Denver, Colorado 80220

Abstract. The systematics of the eastern members of the Hesperia
leonardus complex are studied. H. 1. leonardus and H. 1. pawnee are
conspecific and intergrade in Minnesota, Wisconsin and elsewhere, where
extremely variable populations exist. The name H. 1. montana is restricted to
a recently rediscovered third subspecies occupying a small area in the
Colorado mountains. H. I montana is also extremely variable and is similar in
several characteristics to Minnesota intergrade specimens. We summarize
what is known about distribution, flight periods, habitat, adult behavior and
foodplants of the leonardus complex. Adults have one brood (mostly August-
September), and western populations feed mainly on Liatris flowers. Larvae
eat various grasses, and hibernate in the first stage.

Introduction

MacNeill (1964) defined the H. leonardus group to include two western
species (Columbia Scudder and pahaska Leussler) which he studied in
detail, and two eastern entities not studied in detail, pawnee and
leonardus. The purpose of this paper is to study the taxonomic relation-
ships of the latter two entities and to summarize what is known of their
variation, distribution, behavior and early stages. The most interesting
feature of the complex is that the eastern members pawnee and leonardus ,
appear to belong to one species, despite the gross color difference between
them. Because of the importance of this conspecificity, the evidence for it
is presented first.

Conspecificity of leonardus and pawnee

There are several reasons why we combine leonardus and pawnee into
one species. Pupae are the same (Scott, 1975b; Dethier, 1948). Larvae are
the same although H. 1. pawnee heads are lighter than those of H. 1.
leonardus and if. 1. montana (Scott, 1975b). Scudder’s (1889) drawings of
first instar leonardus leave out many setae which occur in all known
Hesperia species, and the long lateral setae on the ninth abdominal

20(1): 18-35, 1981(82)

19

segment in his drawing is probably a short spatulate seta as in all other
known Hesperia (Dethier, 1939; Scott, 1975b). Eggs are very similar
(Scott, 1975b) (the egg figured by Scudder (1889) is again poorly drawn).
There are no differences in male or femal^ genitalia which we can detect.
Genitalia are too variable individually to detect differences; the genitalia
drawings of pawnee and leonardus by MacNeill (1964) are not “typical”.
There may be slight interpopulational differences in antennal shaft length,
the number of segments of the antennal shaft, and the length of the male
penultimate tarsal segment, but these characteristics are also variable.
Flight periods and adult behavior are very similar.

The main differences between populations of leonardus involve color
pattern of palpi, body and wings. Superficially populations and individuals
look very different from one another (Figs. 2-5).

H. I leonardus and H. I pawnee apparently intergrade over a broad area
from Minnesota and Wisconsin to Manitoba and perhaps Iowa (Fig. 1). In
Spruce Woods Forest in southern Manitoba individuals resemble pawnee
but often have a rust tinge to the ventral yellow color, especially in females
(Figs. 2-3). Four males from Sandilands in southeastern Manitoba are like
Ontario H. I leonardus except the dorsal surface is a little lighter, and the
ventral hindwing spots are small. Samples of H. I leonardus from Crivitz,
Wisconsin are odd in several respects and several characters tend in the
direction of pawnee (Figs. 2-3). Northwestern Wisconsin samples are
intermediate toH. I pawnee andH. I leonardus. Of the two males from Des
Moines, Iowa (Carnegie Museum), “one of these is quite like eastern
leonardus, the other is somewhat paler, more fulvous, suggests transition
to pawnee, but still mostly like leonardus” (H. K. Clench, pers. comm.).

The central Minnesota population is extremely variable in every wing
character, varying from nearly “typical” pawnee to nearly “typical”
leonardus in every character. The few structural characters such as
antennal shaft length to head width ratio are also variable and are inter-
mediate between the two subspecies.

If there are no barriers to hybridization of H. I leonardus and H. I
pawnee, mating should be random and after several generations non-
hnked genes should be independently combined in offspring. If major
barriers to hybridization occur, however, or hybrids are largely sterile, the
population should consist of a majority of individuals recognizable as
either H. L leonardus or K I pawnee plus a minority of individuals with
hybrid traits. The evidence strongly suggests the first interpretation for
the central Minnesota population.

In an effort to analyze differences between H. I leonardus and H. 1.
pawnee, four male characters and five female characters were chosen for
quantitative analysis. These characters were: (1) Ventral hindwing color
(rated from 0 to 7 using eight standard reference specimens varying from
light yellow to deep red brown). This color varies from yellow as in H. 1.

20

J. Res. Lepid.

Fig. 1. Map of H. leonardus populations.

pawnee to red brown as in if. I leonardus in both sexes (Figs. 2-3). (2) The
width of the ventral hindwing postmedian band in cell This band varies
from 0 as in if. L pawnee to 1 .4 mm as in if. /. leonardus in both sexes (Figs.
2-3). (3) Dorsal lightness varies from dark reddish as in if. I leonardus to
Kght fulvous as in if. 1. pawnee in both sexes. This character was quantified
by use of five reference specimens. (4) Darkness of ventral forewing
tornus. This varies from the ground color as in H. I pawnee to completely
black as in if. I leonardus in both sexes. Five reference specimens were
also used. (5) Transparency of the dorsal forewing hyaline spot. This
character was used for females only. This spot varies from the ground color
as in if. I leonardus, to transparent as in if /. pawnee. Four reference
specimens were used to quantify this character. Characters 1-5 are very
variable in the leonardus X pawnee blend zone, much more than in either
H. 1. leonardus or H. I pawnee.

Characters 1-5 were plotted against each other (two of the 14 plots are
shown, Figs. 2-5) in an effort to discover whether there was any
reproductive isolation between leonardus and pawnee that would show up
in wing pattern. No correlations were found except that there are very
slight correlations in both sexes between dorsal lightness and darkness of
the ventral forewing tornus, and in females (but not males) slight
correlations appeared between ventral hindwing color and dorsal lightness
and between ventral hindwing color and darkness of ventral forewing

WIDTH VEMTRAL HINDWING POSTMEDSAN BAND IN CELL M3 (mm)

20(1): 18-35, 1981(82)

21

VENTRAL HINDWING COLOR

0.6

0.4-

0.2

Males

H.I. montana

-r

• •

I ••

• m

> •

•• •, •

% %>

0 1 2 3 4 5 6

VENTRAL HINDWING COLOR GRADE

Figs. 2-5, Scatter diagrams of width of ventral hindwing postmedian spot in cell

versus ventral hindwing color (0-pale yellow, to 6-dark mst red
(for ssp. leonardm) or 6-dark brown (for ssp. montana)). Most ssp. are
on Figs. 2-3, but ssp. montana is on Figs. 4-5.

22

J. Res. Lepid.

tornus. These correlations were very small however, and merely indicate
that in some cases, especially in females, if one part of the wing is light
another part is slightly likely to be light as well.

In general, however, these five characters which distinguish H. I
konardus from H. I pawnee are not correlated with each other. A typical
Minnesota individual may have one character tending toward if. 1.
konardus, another tendig toward if. I pawnee, and others intermediate. It
is our opinion that recombinations have occurred resulting in individuals
with characteristics not observed in if. I pawnee and if. L konardus
populations. For example, some individuals have the ventral hind wing
rust-red but with very small or no spots, and other individuals have this
area light yellow with large spots (Figs. 2-3). This lack of correlation of
characters and the lack of two clusters of individuals corresponding to the
two parental types in any character suggests that the central Minnesota
population is a freely interbreeding population. Due to fragmentation of
natural habitat by farmland the central Minnesota colonies probably now
receive little gene flow from either if. 1. konardus or ff. /. pawnee, and
variability is maintained as in ff. 1. montana through unknown mechanisms.

Males from Wabasha County, Minnesota in this intergradation area
show variation similar to that of specimens from Anoka and Sherburne
counties. Females, however, are slightly more similar to ff. L konardus
than are females from the latter two counties. Of 10 females examined, five
are in the konardus part of the Minnesota population scattergram (Fig. 3),
while the other five are scattered but not in the pawnee comer of the plot.

Flight Period, Habitat, Behavior and Larval Biology

The flight period of ff. konardus is very similar throughout the range:
one brood, usually August to September (Figs. 6-7). Figures 6-7 clearly
show that flight period is earlier at higher latitude and higher altitude.
Ozark populations fly six weeks later than Canada populations, and plains
ff. L pawnee flies later than mountain H. L montana. Pittsburgh,
Pennsylvania, inland populations fly several weeks earlier than those on
the coast at Philadelphia.

ff. konardus populations all occur in meadows or grasslands, commonly
old fields and moist meadows for H. 1. konardus, sandy prairie (near
wooded areas) for the central Minnesota intergrade populations, prairie
(sometimes sandy) for ff. /. pawnee, and open grassy pine forest for H. I
montana.

Adult behavior of konardus populations is very similar. All three
subspecies and the intergrade populations are common BiLiatris punctata
flowers in Colorado, Nebraska, Minnesota, Michigan and New Jersey
(occasionally on other flowers such as Cirsium, Vemonia angustifolia,
Eupatorium purpureum, Solidago, Clematis, Aster, other Liatris species,
“bonehead”). H. I konardus is usually found on flowers other than Liatris,

20(1): 18-35, 1981(82)

23

South Dakota

□ Moles
O F^oies

\

Colorado
(H.l. montono)

100-
80 -

Missouri,

Arkansas rl itr. n 60

Figs. 6-7. Flight period for males and females, in three-day intervals.

commonly on high flowers. Adults are not often seen away from flowers.

I leonardus males commonly choose high perches (4-5 ft.), whereas
western populations usually choose lower perches. H. I pawnee males
when not feeding often perch on small to large hilltops throughout the day.
H. 1. Montana males are usually found in concentrations with females,
often on small hills; males have not been found away from these
concentrations on nearby hilltops used earlier in the summer by H.
pahaska males. Unsuccessful courtship of unwilling females appears
identical in H. I Montana, H. I pawnee and H. I leonardus. It occurs
throughout the day, often on flowers, as males attempt to mate with
feeding females, who flutter their wings until courtship terminates.

H. I leonardus foodplants are Agrostis (Scudder, 1893), PanicuM
virgatuM and Eragrostis alba (Shapiro, 1966). Tietz (1972) lists a
dicotyledon which cannot be a foodplant. Dethier (1939) and Scudder
(1889) raised H. 1. leonardus in the laboratory on common grasses. Many
ovipositions of H. I Montana were seen on Bouteloua gracilis. We have not
discovered oviposition substrate or foodplant for L pawnee. A record of
larval host of pawnee (1970 season’s summary of the Lepidopterists’
Society and Ferris, 1971) is erroneous because it is based on laboratory
feeding of a dying larva. We raised ff. 1. Montana and if. I pawnee to adults
(Scott, 1975b) on Poa pratensis, Cynodon dactylon and other unidentified

24

J. Res. Lepid.

grasses, and believe that laboratory hosts have little relevance to the plant
used in nature by Hesperiinae, which may be host-specific in nature in
spite of broad larval tolerance for laboratory grasses. Some Hesperia
oviposit rather haphazardly, however (Scott, 1975a), so larvae may not be
host-specific.

All H. leonardus populations must overwinter as young larvae, because
eggs hatch immediately and there is not enough time before winter for the
larvae to grow to large size. Overwintering occurs as first stage larva (ssp.
montana), young larva (Scudder, 1893), first stage larva (Scudder, 1889),
most first and some second stage larva (Dethier, 1939), second instar
(Laurent, 1908) (all ssp. leonardus). WhenH. I leonardus, H. 1. pawnee and
H I montana are raised indoors, no diapause occurs and adults emerge
from November to January, taking only about three months to develop
(Scott, 1975b).

Taxonomy— L leonardus

leonardus Harris, 1862. Ihs. Mj. Veget., p. 314. Type locality: Boston,

Mass.

Mia Plotz, 1883 (nomen dubium, dos Passos, 1964).

Uberia Plotz, 1883 (synonym or nomen dubium; see dos Passos, 1960).
stallingsi H. A. Freeman, 1943. Bull. Brooklyn Ent. Soc. 38: 153. TVpe

locality: Blendon, Franklin Co., Ohio.

This subspecies occurs usually in open fields and damp meadows but
occurs in permanently wet meadows and bogs in Virginia (Clark & Clark,
1951).

Description

Apical FW spots distinct, ochre above and below. DFW basal color of females
brown with a very slight orange flush. DFW spots of females distinct, fulvous, one or
rarely two. VHW color light to dark rust red. VHW spots usually large (very rarely
absent), ochre to white (more ochreous in males), median spot in cell Sc + mid
median cell spot almost always present and round, spots larger in females. Fulvous
DHW spots fairly distinct, moderate fulvous suffusion in males, little in females.
VFW tomus almost always black, rarely little lighter than ground color.

Last instar larval head with a light V-shaped genal area and a lateral light area.

Variation within H. leonardus leonardus is mostly clinal, with different characters
showing different dines or patterns of variation (Table 1). There is essentially a
north -south dine in wing size, with a peculiar small sample from Wisconsin. Dark
dorsal phenotypes are prevalent on the Ozark Plateau and in the southeast; the
most fulvous populations occur in Maine, Pennsylvania, Indiana and Wisconsin.
The ventral hindwing band is smallest in the Ozark Plateau. This band is light in the
Great Lakes region, and dark (ochraceous) in soufliem popdations. Ventral
hindwing color is dark in southern Great Lakes and middle Atlantic regions, lighter
(ochreous) northward and westward. The Maine and Ozark samples, greatly
different in other characters, are similar in ventral hindwing color. The indepen-
dent variation of these characters in L leonardus makes desi^afion of additional

20(1): 18-35, 1981(82)

25

subspecies futile.

The Ozark sample is the most distinct. The Ozark material is larger, darker and
tends to have the ventral hindwing spots reduced. The synonym stallingsi
(Freeman, 1 943) named from Franklin County, Ohio, falls about in the middle of the
range of variation we have observed in H. L leonardus.

H. L leonardus X H. 1. pawnee

In c'entral Minnesota there are extremely variable populations inter-
mediate between the subspecies leonardus and pawnee. Its variation has
^eady been discussed.

Description

Apical spots distinct, sometimes indistinct and usually ochre ventrally, sometimes
white in females. DFW basal color of females moderately ochre to brown. DFW
spots of females distinct, most fulvous, some hyaline, usually 1-3, rarely 4 spots.
VHW color ochre yellow to light rust red. VHW spots large to obsolete, usually
ochreous, median spot cell Sc + often absent and median cell spot round but
often small when macular band is present. Fulvous DHW spots in males sometimes
fairly distinct but usually vague due to fulvous suffusion; in females with none to
some fulvous suffusion. VFW tomus yellow to black, weakly correlated with ground
color, slightly darker in females. Larvae undescribed.

H. L pawnee (New combination)

Hesperia pawnee Dodge, 1874, Canad. Ent. 6:44. Type locality: Glencoe,
Dodge County, Nebraska (typ© destroyed).

This light colored subspecies has evolved on the Great Plains. Another
unrelated species, Hesperia ottoe Edwards, has convergently evolved
similar appearance with light yellow underside, possibly for camouflage
against a background of dried grasses. H I pawnee differs from H I
leonardus mainly in the much lighter ochre color, the reduction of the
ventral spots, and in having hyaline forewing spots in females; H I
montana and the central Minnesota population are variable in the hyaline
spot character perhaps due to hydbridization with pawnee. The hyaline
spots apparently also evolve convergently on the plains. Of the Hesperia
species which possess hyaline spots., H. ottoe, H uncos Edwards (only
plains and eastern Great Basin uncos have hyaline spots), H. dacotae
Skinner, H 1. pawnee, H. metea Scudder, H attains Edwards, and some H
meskei Edwards, the first four are plains species.

There is considerable individual variation in some wing characters such
as general dorsal color (very light to dark) and the number and size of faint
ventral hindwing spots (variation noted also by Leussler, 1923). There is
somewhat less variation in ventral color and in color of the female forewing
spots. There is very little geographic variation throughout the range. A
series from Custer County, Colorado, has slightly larger ventral hindwing
ochreous spots (Figs. 2-3). Size decreases clinally from south to north as in
H. I leonardus (Table 2). Specimens from Montana are very small.

Ontario- Massachusetts- Missouri-

Quebec Maine Rhode Island New York Michigan Wisconsin Pennsylvania New Jersey Ohio Indiana Virginia Arkansas

26

J. Res. Lepid.

Lf\ ir\

LA LA

LA

CM r^

fA

fA

CO -J- LA

vo*— cA-d" '<o<A

^ CM csj LA vO

LA vO »-

1— LA
LA \D

CO \D ^

-d- vD »— CM

S I

u i_ ^ I 3

0 0 3^ n:

-Q -C

3 -o'
X —
> 2

Of 04-

O -ST

'o ^

CM

e

m §
(S S
CO

Forewing- length 6 15.3 l6,2 15.0 U.5 15.1 15-5 15.5 15.8 16.0 15.3

0 16.6 17.6 17.0 — 16.7 17-3 17.0 17.5 17.^ 16.8

20(1): 18-35, 1981(82)

27

Table 1. Geographic variation of six characters for H. leonardus leonardus: (1)
forewing length in mm; (2) dorsal forewing fulvous (1-very little fulvous,
4-very fulvous, 2 and 3 -intermediate); (3) dorsal hindwing postmedian
band suffusion (1-narrow, discrete; 2-broad, still discrete; 3-inter-
mediate; 4-suffused); (4) ventral hindwing postmedian band width in cell
Ms (mm); (5) ventral hindwing postmedian band color (1 -white; 2 -very
light ochre; 3-medium; 4-dark ochre_j; and (6) ventral hindwing ground
* color (0-very light yellow; 7-very dark rust red; 1 to 6-intermediate).
Numbers (except for sample size) are averages.

Table 2. Forewing length (mm) of H. leonardus samples not included in Table 1.

Description

Apical spots rather indistinct and fulvous in males, distinct and hyaline in females.
DFW basal color of females moderately ochre to brown. DFW spots of females
distinct, most hyaline, usually 3-4, occasionally 2 spots. VHW color of males
orangish ochre to ochre yellow, of females light ochre to ochre yellow. VHW spots
same as in montana but spots usually obsolete, occasionally moderate size and
ochreous, little lighter than rest of wing. Fulvous DHW spots same as in montana.
VFW tomus usually yellow, rarely brown in males and black in females, slightly
darker in females.

Larvae as in leonardus but heads are lighter laterally.

H. 1. montana (Nev^ combination)

Pamphila pawnee montana Skinner, 1911, Ent. News 22: 413. Type
locality: we restrict to Buffalo Creek, Jefferson County, Colorado (the
town).

H. I montana differs from H. I pawnee mainly in darker (more brown)
color and the presence of ventral hindwing spots (Figs. 2-5). It is almost as
variable as the Minnesota intergrade population.

Description

Apical spots usually distinct, mostly fulvous, often hyaline in females. DFW basal
color of females moderately ochre to brown. DFW spots of females distinct, fulvous
to hyaline, most somewhat hyaline, usually 2-3, occasionally 1 or 4 spots. VHW
color variable, ochre yellow to dark brown, rarely russet brown or greenish brown.
VHW spots moderate size to obsolete, ochreous white, medial spot cell Sc + R*
rarely present, medial cell spot rounded, often absent, spots larger in females.
Fulvous DHW spots fairly distinct, in females with slight to moderately fulvous
suffusion, usually lost in fulvous suffusion in males. VFW tornus yellow to black,
usually brown, weakly correlated with ground color, slightly darker in females.
Larvae similar to H. I leonardus.

We designate a lectotype male in the Carnegie Museum, Pittsburgh, Pennsylvania
(labels include “type no. 7086, Colorado Bruce, Chaffee Co. 7500 ft.”), which fits
the original description and our characterization of H. L montana. Only two of the
eleven cotypes had locality data: one says Chaffee County, Colorado 7500 ft. alt.,
and the other says Salida (Chaffee County, Colorado), May 21, 7500 ft. alt. These

28

J. Res. Lepid.

data are erroneous because H. leonardus adults almost never occur before August,
the habitat at Salida differs from known habitats, and we have not found the species
in Chaffee County in any month despite heavy collecting by James Scott and Glenn
Scott (1978). F. M. Brown (pers. comm.) has determined the itinerary of David
Bruce, based on letters from Bruce to Herman Strecker. The Chaffee County
specimens are obviously mislabeled, because at the time they were collected,
Bruce, a painter, was recuperating for three weeks from a fall from a scaffold, in a
hospital in Redcloud, Nebraska! F. M. Brown found that Bruce had left a collecting
net with the children of Mr. William W. G. Smith at Buffalo Creek, Jefferson
County, Colorado, that August, who on 7 September sent Bruce 7 boxes of
butterflies from the “Platte Canyon Valley.” It seems probable that the Smith
children caught the types of montana at or near Buffalo Creek. We correct and
restrict the type locality to the vicinity of the town of Buffalo Creek, Jefferson
County, Colorado, because of these historical records, and because we have found
montana within several miles of there in August. D. Bruce mislabeled other
butterflies (see Ferris & Fisher, 1977), and E. Oslar is similarly noted for
mislabeling of material (Oslar specimens in the American Museum of Natural
History).

H. 1. montana occurs only in a small area in the South Platte River Canyon system
in the mountains of Colorado. All the records of H. I montana are in the Pikes Peak
Granite, which occurs in the South Platte Canyon southeastward to Pikes Peak. H. 1.
pawnee does not occur on this granite and the granite boundary coincides with the
boundary between montana and pawnee. H. 1. montana is extremely variable (Figs.
4-5); some specimens are identical to plains pawnee and others are brown and have
large cream spots on VHW. The ventral hindwing color ranges from yellow to dark
brown, usually light brown, with occasional individuals greenish brown or slightly
rust brown. In VHW color it is somewhat less variable than the central Minnesota
population, but other wing characters are nearly as variable, and show the same lack
of correlation as in the central Minnesota population. The variability seems to
indicate gene flow from plains H. I pawnee, although we could not find any
intervening population between pawnee from Waterton and montana from the
abandoned South Platte Hotel.

The wing pattern differences between H. 1. montana and H. 1. pawnee are genetic,
because the differences were maintained in individuals raised under identical
laboratory conditions.

There are three hypotheses for the origin of H. 1. montana: 1) introgression of a
pawnee population with Hesperia pahaska; 2) a relict population from a time in the
Pleistocene when H. 1. leonardus occurred in forested areas across what is now the
Great Plains and when H. 1. pawnee was farther south or had not yet evolved; 3) a
population which evolved in its present location from H. 1. pawnee founders. The
first hypothesis, introgression, is supported by the several characters in which F7. 1.
montana is more similar to H. pahaska than to H. 1. pawnee, and by the flight
periods. On the plains the flight periods of H. pahaska and H. I pawnee are well
separated, but in the mountains H. 1. montana flies about a week earlier, so that late
female pahaska might mate with early male montana. On July 2 we found H.
pahaska males hilltopping on hills 200 feet from a later large concentration of H. 1.
montana, but the H. pahaska did not differ from specimens found elsewhere in
eastern Colorado. In view of the difficulty in proving introgression, hypothesis one
must be considered unsupported speculation. In morphology, H. leonardus is most

20(1): 18-35, 1981(82)

29

closely related to H. Columbia which occurs on the pacific Coast. It is less closely
related to H. pahaska, which is sympfitric with H. 1. montana and H. L pawnee in
numerous localities from Colorado to the Dakotas and Montana, but flies in June-
July. There is no evidence to suggest that H. leonardus and K pahaska hybridize
anywhere that they meet, as we previously speculated (MacNeill, 1975). We believe
that some combination of hypotheses two and three represents the probable history
of the population. Hypothesis two is supported by the lack of dark ventral hindwing
phenotypes in the present H. I pawnee population to act as a starting point for
selection of a darker population. However, K I leonardus has rust-red ventral
coloration whereas in L montana color variation involves various shades of brown
and light rust individuals are rare. K h^othesis two is correct, since if. I pawnee
invervened between 1. leonardus and 1. montana the latter two populations may have
diverged in ventral coloration. Hypothesis three is supported by the fact that both
forest subspecies {leonardus and montana) are dark, whereas the plains subspecies
{pawnee) is light. The variability of the color pattern of the wings of if. 1. montana is
hard to explain, but could be accounted for by selection for dark spotted
phenotypes along with occasional immigration of H. 1. pawnee phenotypes.

Distribution

Distribution is plotted on Figure 1 . Flight periods are shown in Figures 6-
7. Localities are listed below. H. 1. leonardus undoubtedly occurs in
Delaware and may be found in eastern Oklahoma. H. L leonardus x pawnee
may be found in central Iowa, southeastern Nebraska and northeastern
Kansas. H. I pawnee may be found in southeastern Alberta. The absence
of H. I pawnee records in the center of its range is probably due to lack of
collecting.

Several records appear erroneous and are not listed. These include many
for if. 1. montana (see text above); if. 1. leonardus from Florida (Scudder,
1889) which, judging from the flight period given, refers to Hesperia
attalus (Edwards); and “Texas” (Evans, 1956). The records for Louisiana
and Alabama are so isolated from other records that they require
confirmation.

if. leonardus leonardus (counties only for U. S.)

164 males, 107 females examined

Nova Scotia: Digby.

Quebec: Montreal, Terrebonne, Aylmer, Lakefield, Norway Bay, Shawbridge,
Lanoraie, St. Anne de Bellevue, Rawdon, St. Maurice, Rigaud, Papineau Co.,
Gatineau Co., Pontiac Co.

New Brunswick: southern part.

Ontario: Perth Rd., Frontenac Co.; Bruce Co.; Clarendon; Chaffey’s Locks,
Leeds Co.; Kent Bridge, Kent Co.; Kahshee Lake, Muskoka Distr.; Asperitos Id.,
Parry Sound Distr.; Gravenhorst, Muskoka Distr.; Georgian Bay; Ottawa; London;
Dublin; Toronto; Simcoe; Bruce Co.; Algonquin Park, Parry Sound Distr.

Maine: Penobscot, Cumberland, Lincoln, York, Hancock, Washington, Somerset,
Franklin.

New Hampshire: Coos; White Mts.; Mast Yard.

20(1): 18-35, 1981(82)

31

Figs. 8-25. H. leonardus pawnee (Figs. 8-18) and H. L leonardus (Figs, 19-25).

Ssp. pawnee: left column (top to bottom): m ups (upperside) Green
Mtn., Jefferson Co., CO, 25 Aug, 1971, J. Scott; mups same data; m
ups Green Mtn., 4 Sept 1969, J. Scott; f ups 3 mi. S. Sedalia, Douglas
Co., CO, 3 Sept. 1967, R. Stanford; f ups Lakewood, CO, 3 Sept.
1960, J. Scott; f ups Little Missouri River, BUlings Co., ND, 22 July

1970, J. Nordin; middle column: m und (underside) (ventral hind-
wing color no. 1) Green Mtn., 25 Aug. 1971, J. Scott; m und 3 mi. E.
Wetmore, Custer Co., CO, 2 Sept. 1971, J. Scott; f und 3 mi. S.
Sedalia, 3 Sept. 1967, R. Stanford; f und 1 mi. E. Parker, Douglas
Co., CO, 11 Sept. 1968, R. Stanford; f und % mi. W. Bitter Lake, Day
Co., SD, 25 Aug. 1971, J. Nordin; ssp. leonardus: m ups near N.
Manchester, Kosciusko Co., IN, 27 Aug. 1970, E. M. Shull; right
column: m ups Eggleston, VA, 26 Aug. 1964, G. Straley; m und
Enfield, ME, 26 Aug. 1964, L. Grey; f ups near N. Manchester, 2
Sept. 1970, E. Shull; f ups Ida Center Rd., Monroe Co., MI, 28 Aug.

197 1, L. Melton; fund near Woodbine, Cape May, NJ, 12 Sept. 1970,
R. Stanford; f und Eggleston, 14 Sept. 1964, G. Straley.

Figs. 26-44. H. leonardus leonardus x leonardus pawnee, all from Sand Dunes
State Forest, Sherburne Co., Minnesota. Left column: m ups 31
Aug. 1966, C. Hansen; m ups, 18 Aug. 1967, P. Nordin; f ups 29 Aug.
1971, J. Masters; f ups 19 Aug. 1967, P. Nordin; m und 19 Aug. 1967,
P. Nordin; m und (ventral hindwing color rating no. 2), 19 Aug. 1967,
P. Nordin; m und 31 Aug. 1966, C. Hansen; middle column: m und
(color rating no. 4), 16 Aug. 1970, J. Nordin; m und (color rating no.
3), 29 Aug. 1971, J. Masters; m und 19 Aug. 1967, P. Nordin; m und
19 Aug. 1967, P. Nordin; f und 29 Aug. 1971, J. Masters; f und 25
Aug. 1969, J. Sorensen; right column: f und 25 Aug. 1969, J.
Sorensen; f und 19 Aug. 1967, P. Nordin; f und 25 Aug. 1969, J.
Sorensen; f und 25 Aug. 1969, J. Sorensen; f und 25 Aug. 1969, J.
Sorensen; f und (color rating no. 6), 19 Aug. 1967, P. Nordin.

Figs. 45-61. H. leonardus montana, all J. Scott. Left column (top to bottom): m
ups N. of Cheesman Lake, Jefferson Co., CO, 3 Sept. 1971; m ups
Nighthawk, Douglas Co., CO, 28 Aug. 1969; f ups Nighthawk, 1 Sept.
1970; f ups Nighthawk, 1 Sept. 1970; m und Nighthawk, 12 Aug.
1971; m und Nighthawk, 29 Aug. 1971; middle column: m und
Nighthawk, 12 Aug. 1971; m und Nighthawk, 28 Aug. 1969; m und
Nighthawk, 1 Sept. 1970; f und (ventral hindwing color rating no. 2),
Nighthawk, 28 Aug. 1969; f und Nighthawk, 29 Aug. 1971; f und
Nighthawk, 28 Aug. 1969; right column: f und Nighthawk, 1 Sept.
1970; f und Nighthawk, 28 Aug. 1969; f und Nighthawk, 28 Aug.
1969; f und Nighthawk, 1 Sept. 1970; f und (color rating no. 5), N. of
Cheesman Lake, 3 Sept. 1971.

Vermont: Windham.

Massachusetts: Barnstable, Nantucket, Essex, Middlesex, Bristol, Dukes,
Worcester; Harwich Point; Pelham Hills; Hallowell (in Mass.?).

New Jersey: Camden, Ocean, Cape May, Middlesex, Burlington, Mercer,

32

J. Res. Lepid.

Bergen, Union.

Rhode Island: Providence, Washington.

Connecticut: Hartford.

New York: Richmond, Nassau, Suffolk, St. Lawrence, Clinton, Jefferson, Essex,
Lewis, Oswego, Hamilton, Warren, Washington, Fulton, Oneida, Albany, Schoharie,
Columbia, Onondaga, Genesee, Erie, Livingston, Chautauqua, Cattaraugus, Al-
legany, Steuben, Yates, Cayuga, Seneca, Schuyler, Chemung, Tompkins, Tioga,
Broome, Chenango, Delaware, Sullivan, Ulster, Orange, Dutchess, Rockland,
Westchester, Greene.

Pennsylvania: Allegheny, Bucks, Butler, Chester, Clarion, Crawford, Delaware,
Fayette, Montgomery, Montour, Lancaster, Somerset, Warren, Westmoreland,
Bedford, Indiana, Schuylkill, Tioga, Susquehanna, Berks, Lackawanna, Monroe,
Pike, Clinton, Potter, Centre.

Maryland: Montgomery, Garrett, Baltimore, Allegheny, Prince Georges, Charles,
Ann Arundel.

District of Columbia: Washington.

Virginia: Arlington, Patrick, Madison, Giles, Montgomeiy, Prince William,
Stafford, Powhatan.

West Virginia: Kanawha, Randolph, Nicholas, Upshur, Lewis.

North Carolina: Durham, Richmond, Buncombe, Avery, Guilford, Clay, Tran-
sylvania.

South Carolina: Greenville.

Georgia: Rabun.

Alabama: Cherokee, Tuscaloosa.

Louisiana: Madison Parish.

Tennessee: Morgan, Marion.

Kentucky: Larue, Jefferson, Meade.

Ohio: Franklin, Lorain, Athens, Lucas, Williams, Hocking, Jackson, Wayne,
Ashland, Vinton.

Indiana: Perry, Brown, Lake, Randolph, Wabash, Kosciusko, LaGrange, Porter,
Steuben, Pulaski.

Illinois: Vermilion, Mason, Cook, Peoria, McDonough, Jackson, Mason, Han-
cock, Schuyler.

Michigan: Monroe, Newaygo, Iosco, Allegan, Montcalm, Kalamazoo, Chippewa,
Livingston, Houghton, Otsego, Oakland, Macomb, Grand Traverse, Presque Isle,
Schoolcraft, Emmet, Cheboygan, Kalkaska, Crawford, Oscoda, Manistee, Roscom-
mon, Ogemaw, Lake, Osceola, Mecosta, Huron, Ottawa, Kent, Clinton, St. Claire,
Ingham, Jackson, Washtenaw, Wayne, Berrian, Brown, Van Buren, Dickinson.

Arkansas: Benton, Washington, Carroll, Ovachita.

Missouri: St. Louis, Iron, Camden, Barry, Greene, Franklin, Jefferson, St.
Francois.

Kansas: Douglas.

Iowa: Scott, Des Moines, Polk, Audubon?, Winneshiek.

Wisconsin: Marinette, Eau Claire, Grant, Langlade, Milwaukee, Green, Sauk,
Dane, Columbia.

Manitoba: Sandilands.

H. I leonardus x K I pawnee (counties and localities)

121 males, 55 females examined

20(1): 18-35, 1981(82)

33

Wisconsin: Douglas (Dairy land, Jackson L. Boughner); Burnett (Crex Meadows
near Grantsburg, J. L. Boughner).

Minnesota: Sherburne (Sand Dunes State Forest SSE Orrock and near Zim-
merman, J. S. Nordin, P. D. Nordin, C. Hansen, J. Masters, J. T. Sorenson, W.
Bergman); Anoka (Section 11, Coon Rapids Township, J. S. Nordin; Bunker
Prairie, R. L. Huber); Wabasha (Kellogg, Allison Bolduc, Gajy Korsmo; Kellogg
Prairie, R. L. Huber); Goodhue (Eggleston, E. M, Brackney); Chisago (2 mi, N.
Chisago City); Crow Wing (Brainard); Ohnstead; Scott; Dakota.

H. leonardus pawnee (counties only for U. S.)

409 males, 139 females examined

Minnesota: Lac Qui Parle, Murray, Yellow Medicine, Chippewa, Pipestone,
Qay, Lincoln, Norman, Swift.

Iowa: Poweshiek?, Woodbury.

Kansas: Smith.

Nebraska: Stanton, Boone, Dodge, Nemaha, Douglas.

South Dakota: Day, Meade, Harding, Pennington, Custer, Lawrence, Brookings,
Marshall, Roberts.

North Dakota: Ransom, Bottineau, Williams, McKenzie, Morton, Grand Forks,
Billings, Ramsey, Slope.

Manitoba: Aweme, Cartwright, Spruce Woods Forest Reserve near Hwy. 258.
Saskatchewan: Redvers.

Montana: Dawson, Custer, Big Horn, Prairie, Chouteau.

Wyoming: Platte, Sheridan, Laramie.

Colorado: Logan, Larimer, Weld, Boulder, Denver, Arapahoe, Jefferson,
Douglas, El Paso, Custer, Pueblo, Morgan.

H. leonardus montana (counties and localities)

208 males, 239 females examined

Colorado: Douglas Co.: Sugar Creek 5 mi. NE Deckers (R. E. Stanford),
Nighthawk 10 mi. NE Deckers, 4.3 mi. SE Deckers, 10 mi. SSE Deckers (all three J.
A. Scott); Teller Co.: junction of Teller, Douglas, Jefferson, and Park Counties (R.
E. Stanford); Jefferson Co.: 1 mi. N Cheesman Reservoir near Wigwam Creek (J. A.
Scott), 6.9 rd. mi. up North Fork of South Platte River from town of South Platte (J.
A. Scott), South Platte Hotel (R. E. Stanford & J. A. Scott), Deckers (R. E.
Stanford).

Summary

Hesperia leonardus Harris, 1862

a. leonardus leonardus Harris, 1862

"? liberia Plotz, 1883 (nomen dubium, dos Passos, 1960)

? lidia Plotz, 1883 (nomen dubium, dos Passes, 1964)
stallingsi Freeman, 1943 (subjective synonym)

b. leonardus leonardus x leonardus pawnee (intergrade populations)

c. leonardus pawnee Dodge, 1874 (new combination)

d. leonardus montana (Skinner) 1911 (new combination)

Type locality correction and restriction: town of Buffalo Creek,

34

J. Res. Lepid.

Jefferson County, Colorado.

Lectotype designated Carnegie Museum,

Acknowledgments. Many people helped us in this project. We especially thank
Charles A. Triplehom (Ohio State University Museum), Julian P. Donahue (Los
Angeles Natural History Museum), Charles Remington (Peabody Museum, Yale
University), Howard V. Weems, Jr. (Florida Department of Agriculture, Gainesville),
Frederick H. Rindge (American Museum of Natural Histoiy), Wilbur R. Eims
(University of Missouri, Entomology Museum), Richard Heitzman, John T.
Sorensen, John Masters, Roderick R. Irwin (niinois Natural History Survey), Lutz
J. Bayer (University of Wisconsin), John Hafemik, John S. Nordin, Philip D.
Nordin, Clark Hansen, and C. Don MacNeill for information and loan of specimens.
We also thank Arthur M. Shapiro, Steven J. Kohler, Charles V. Covell, Jr., Tim L.
McCabe, Richard E. Gray, J. Bolling Sullivan, Frances McAlister, F. M. Brown,
Bryant Mather, Roger M. Kuehn, Arthur C. Sheppard (Lyman Museum, McGiU
University), C. S. Quelch, Warren J. Kiel, Jack L. Harry, William Bergman, Robert
Price, Mogens C. Nielsen, Walfried J. Reinthal, Clifford D. Ferris, Mike Fisher,
Hany K. Clench (Carnegie Museum), William H. Howe, George Lewis (Canadian
National collection), Gerald B. Straley, Lee J. Melton HI, Lucien Harris, Jr.,
Richard B. Dominick, Ronald L. Huber, Jackson L. Boughner, F. Martin Brown,
Ronald R. Gatrelle, WiUiam E. Sieker, H. Townes, Jr., Kurt Johnson, William
McGuire, and Ernest M. Shull for helpful information.

literature Cited

CLARK, A. H. & c. F. CLARK, 1951. The butterflies of Virginia. Smithsonian Inst., Wash-
ington, D. C. 239 p.

DETfflER, V. G., 1939. Early Stages of some Hesperiinae. Canad. Ent. 71: 117-118,
, 1948. Life history of Hesperia leonardus. Bull. S. Calif. Acad. Sci, 47: 1-2.
dos PASSOS, C. F., 1960. Taxonomic notes on some nearctic Rhopalocera. 1. Hes-
perioidea. 14: 24-36.

, 1964. A synonymic list of the nearctic Rhopalocera. Mem, Lepidopterists’
Society No. 1.

EVANS, w. H., 1955. A catalogue of the American Hesperiidae in the British Museum.

Vol. 4, Hesperiinae. London: British Museum of Natural History.

FERRIS, c. D., 1971. An annotated checklist of the Rhopalocera (butterflies) of
Wyoming. Agric. Exp. Sta. Univ. Wyoming, Laramie, Science Monograph No.
23, p. 1-75.

FERRIS, C. D. & M. s. FISHER, 1977. Charidryos flavula B. & McD.: a question of
identity. J. Res. Lepid. 16: 133-140.

FREEMAN, H. A., 1943. A new form of Hesperia leonardus (Harris) from the middle
western United States (Lepid., Hesperiidae). Bull. Brooklyn Ent. Soc. 38:
153-154.

LAURENT, P., 1908. Notes on some early stages of some Pamphila. Ent. News 19:
408-417.

LEUSSLER, R. A., 1923. Notes on variation in 53 specimens Pamphila pawnee
collected at Pilger, Nebraska, Sept. 2, 1922. Ent. News 34: 28.

MacNEiLL, C. D., 1964. The skippers of the genus Hesperia in western North America

20(1): 18-35, 1981(82)

35

with special reference to California. Univ. Calif. PubL Entom. 35: 230 p.

, 1976. Hesperiidae, In W. Howe (ed.) The butterflies of North America.
Doubleday & Co., New York.

SCOTT, J. A., 1975a. Clinical intergradation of Hesperia comma Colorado (Hesperi-
idae). J. Lepid. Soc. 29: 156-161.

, 1975b. Early stages of seven Colorado if espma (Hesperiidae). J. Lepid.
Soc. 24: 163-167.

SCOTT, J. A. & G. R. SCOTT, 1978. Ecology & distribution of the butterflies of southern
central Colorado. J. Res. Lepid. 17: 73-128.

SCUDDER, s. H., 1889. Butterflies of the eastern United States and Canada with
special reference to New England. 3 voL Cambridge, Mass. 1958 p.

, 1893. Brief guide to the commoner butterflies of the northern United
States and Canada. Henry Holt & Co., New York. 206 p.

SHAPIRO, A. M., 1966. Butterflies of the Delaware Valley. Amer. Ent. Soc. Special
PubL 79 p.

SKINNER, H., 1911. New species or subspecies of North American butterflies. Ent.
News 22: 412-413.

TTETZ, H. M., 1972. An index to the described life histories, early stages, and hosts
of the macrolepidoptera of the continental United States and Canada. AUyn
Museum of Entomology, Sarasota, Florida. 1041 p.

Journal of Research on the Lepidoptera

20(1): 36-42, 1981(82)

Notes on the Immature Biology of Two Myrmecophilous

Lycaenidae: Juditha molpe (Riodininae) and Panthiades
bitias (Lycaeninae)

Curtis J. Callaghan^

Apartado Aereo 2184, Cali, Colombia

Abstract. The early stages, habitat, and biology are described for two
sympatric lycaenid butterflies, Juditha molpe (Riodininae) and Panthiades
bitias (Lycaeninae). The larvae of both species use the same foodplant and
are tended by ants of the genus Campanotus.

Introduction

Juditha molpe (Huebner) is a common riodinid butterfly of the neotropi-
cal region ranging from Mexico to Argentina and inhabiting a wide variety
of habitats from rainforest to dry tropical scrub. Guppy (1904) recorded
molpe larvae as feeding on Cassia plants and found in association with
'Targe solitary ants” in Trinidad. His description of the larvae and their
interaction with the ants is very sketchy, however. He merely mentions
that the ants attend and milk the larvae.

Panthiades bitias (Cramer) is a common hairstreak butterfly of the
neotropics, ranging from northwestern and central Mexico to central
Brazil. In spite of its being found commonly throughout its range, its
biology has not been described. A foodplant records is given in Muyshondt
(1973).

The purpose of this paper is to describe the immatures, habitat, and
biology of J. molpe and P. bitias as observed at El Boqueron, Colombia.
This paper represents part of a continuing effort to study and record the
biology of neotropical Lycaenidae.

Description of Immature Stages

J. molpe

Egg: About 0.5 mm in diameter, slightly flattened above and below.
Micropyle small and round, with a dot in the center. Color light green when
laid, then turning to tan. Surface covered with a network of thin lines.
Duration six days. First instar larva, Fig. A. Newly hatched larva 0.9
mm long. Head capsule width 0.2 mm. Head and thorax black, rest light
green. Prothorax with eight small dorsal protrusions with a small seta

‘The author is “Pesquisador Associado” at the Museu Nacional, Rio de Janeior.

Figure A.
Figure B.
Figure C.
Figure D.
Figure E.

First instar larva of J. molpe.
Fourth instar larva of J. molpe.
Cephalad part of pupa of J. molpe.
Final instar of P. bitias.

Pupa of P. bitias.

extending cephalad out of each. Abdomen with small yellow spiracles on
the second through eighth segments, and a carapace on the last segment
with numerous long setae extending from around its edge. All abdominal
and the second and third thoracic segments with small setae bordering the
lower edge. Third instar larva. Length 5 mm, head capsule width 0.8
mm. Head and prothorax black, rest pale yellow. Prothorax with two
deeply bisected horns cephalad with one long seta on each lobe. Also, a
smaller protrusion found on each side of the prothorax with two setae
extending laterally out of each. Another seta extends laterally from the
mesothorax. Abdomen as in first instar with the added presence of two
raised myrmecphilous organs on the eighth segment. The setae at the end
of the last abdominal segment shorter. Duration six days. Fourth instar
larva: Fig. B. Length 7.5 to 14 mm. Head capsule width 1.5 mm. Head
black. Prothorax light brown, rest olive green. Two rows of small yellow
spots dorsally on the abdominal segments. Two myrmecophilous glands

38

J. Res. Lepid.

G

Figure F.
Figure G.
Figure H.
Figure 1.
Figure J.
Figure K.

Adult Juditha molpe.

Adult Panthiades bitias.

Habitat at El Boqueron, Colombia.

Final instar Panthiades bitias attended by ants on foodplant.
Foodplant Calliondra globerrima.

Juditha molpe final instar attended by a Campanotus ant.

light brown. Prothoracic horns blunter and with two setae each. Second
smaller horn on each side with one seta each. One additional seta on the
mesothorax. Spiracles and myrmecophilous glands light brown. Duration
eight days. Fifth instar larva: Fig. K. Length 1 5 to 1 9 mm. Head capsule

20(1): 36-42, 1981(82)

39

width 2 mm. Color light green and brown mottled. Morphology as in fourth
instar. Duration eight days. Prepupal larva: Length 19 mm. Color light
mottled brown. Duration three days. Pupa: Fig. C. Color light mottled
brown with striated darker brown marks. Wing cases darker brown than
abdomen. Attached by a thin girdle passing over the middle of the body
and a silk pad under the last segment. Imago: Fig. F.

P. bitias

Third (?) instar larva: Length 5 mm, head capsule width 0.8 mm. Head
black, rest light brown and green mottled. Head covered by a hood formed
by the first thoracic segment. Second and third segments form a fleshy
bifurcated proturbance extending over the first segment. Abdominal
segments flat and slightly concave dorsally and convex laterally, giving the
larva a hexagonal shape when viewed head on. Yellow spiracles found on
segments one through eight dorsally. Last abdominal segments with a
broad dorsal plate which is slightly concave at the end. Body covered with
small cilia. Final instar larva: Figs. D & 1. Head black, thorax and
abdomen light brown and green. Abdomen with darker green elongated
marks dorsally and two yellow marks on the sixth and seventh segments.
Morphology similar to the third instar, only with the proturbance of the
second thoracic segment and the plate of the abdominal segment being
more deeply bifurcated. Duration nine days . Prepupal larva: Length 20
mm. Color light mottled brown. Morpholo©^ as in final instar. Duration
two days. Pupa: Fig. E. Wing cases dark green, abdomen light green with
three rows of small light brown dots on each side. Attached by silk
pad. Imago: Fig. G.

Discussion

El Boqueron is a small town located on the border of the Departments of
Cundinamarca and Tolima, some 80 km southwest of Bogota at an
elevation of 300 m. Here, the boulder strewn Rio Chocho cuts through
some high ridges on its way to the Magdalena River, forming a deep and
rather picturesque canyon. The area is classified as Dry Tropical Forest
(IGAC, 1977, after Holdridge, 1947), with an annual rainfall of between
one and two meters. The rains usually fall in March through May, and
again in September through November. The dry climate combined with
repeated burnings have practically denuded the area of forest, except for
thin lines of trees along the streams. The general aspect of the countryside
is not unlike that of northern Mexico, with thorny scrub, cacti, and
herbacious plants growing on the higher rocky ground (Fig. H). The
butterfly fauna is likewise similar to that of the northern neotropics with
species such as Heliconius charitonia, Zeonia cesonia and Libythea
bachmanni found commonly following the rains.

The principal foodplant of both J. molpe and P. bitias at El Boqueron is
Calliondra globerrima (Benth.) Britton & Killip, family Mimosacae. It

40

J. Res. Lepid.

grows commonly along the river bed and is sometimes covered by water
during flash floods. Its appearance is that of a small bushy tree with smaU
leathery pinnate leaves and clumps of red and white flowers with long
stamens, which bloom at various times during the year (Fig. J). The fruits
are elongated bean-like pods about 13 cm long with up to a dozen black
seeds inside. This plant does not have extra-floral nectaries, but is often
infested with coccids which attract ants of the genus CampanoUis.

A second plant on which female J. molpe were observed ovipositing was
Bauhinia pediolata (Mutis ex DC) Trioma ex Hook, family Caesolpinaceae.
Tliis shrub has large rounded, pointed leaves, clumps of yellow flowers,
and long bean like fruits. Extra-floral nectaries, located on the stems
between the fruits and flowers, attract the same Campanotm ants as those
on C. globerrima. B. pediolata is a forest species, growing in more shaded
localities than C. globerrima.

lobserved J. mo/pe females ovipositing on both plants between 1200 and
1400 hours. The eggs were laid wherever there were ants, and not on any
particular part of the plant. Strangely, I did not observe oviposition on the
flowers of C. globerrima which is where the larvae feed. Eggs were laid
singly, the female feeling the substrate with her abdomen before oviposit-
ing. Nearby ants would make agressive movements in the form of short
rushes towards the female, at which time she would fly off. Following one
female, I observed six eggs placed on different widely spaced parts of C.
globerrima in about ten minutes. I was unsuccessful in finding larvae on
other nearby C. globerrima plants which were in bloom, but mthout ants.
Also, ovipositing females showed little interest in these plants, sometimes
investigating them, but always returning to the infested plant to oviposit.
This behavior suggests that the main attractant to the ovipositing female
molpe was the presence of ants, and not mere foodplant availability. This
possibility is reiirforced by my observations of oviposition on a completely
unrelated plant, B. pediolata^ but which harbored the same ant species.
Future searches of B. pediolata failed to turn up larvae. In fact, B. pediolata
leaves presented to the larvae in the lab were ignored in favor of C.
globerrima flowers. No B. pediolata flowers were available during the time
of the study to test the larvae preference for this part of the plant.

Upon hatching the smaU J. molpe larvae on C. globerrima move to nearby
clumps of ripe flower buds where they begin to feed. Immature green buds,
flowers and leaves are not eaten, the latter being especiaUy tough and
leathery. The young molpe larvae pass at least the first two instars inside
the bud clumps in the company of smaU coccids and other insect larvae,
possibly dipterids. The larvae lead a solitary existence, and although two
or more were found on the same bud clump, they showed little interest in
one another.

Upon reaching the third instar, both J. molpe andP. bitias leave the buds
and move to nearby stems or leaves where they may remain motionless for

20(1): 36-42, 1981(82)

41

as long as 24 hours before returning to feed on the buds. When moving
about the foodplant, both species weave the head back and forth, spinning
a silk web with which they maintain their grip on the plant surface. Their
mottled green color and flat profiles make both molpe and bitias difficult to
locate on the foodplant. Their presence was usually betrayed by the ants
which showed great interest in the larvae of both species at this stage (Figs.
I & K). At least three ants were always observed in attendance, stroking or
“drumming” on the larvae, which in turn would secrete honeydew which
the ants would eat. The interactions observed between the ants and larvae
were similar to those of Menander felsina and Campanotus ants studied
near Rio de Janeiro (Callaghan, 1977).

The myrmecopWlous organs of J. molpe are located on the eighth
abdominal segment and take the form of two slightly raised lumps with a
hole in the middle, looking like the nectaries on many plants. Unlike M
felsina larvae, there was no observed physiological change in the shape of
the glands when stimulated by ants. Nor were there any organs similar to
the lateral tubercles of Lemonias caliginea (Ross, 1966) or various species
of Audre (Pers. Observation). The “horns” on the prothorax of J. molpe
were observed to be passive organs, though the long setae protruding from
each would suggest that they had a sensory role. When disturbed, the J.
molpe larvae raise the front half of the body and flop it around, as do
various species of Actinote (Pers. Observation).

The P. bitias larvae were equally attractive to ants. However, observa-
tion under magnification failed to reveal any distinct myrmecophilous
organs. The interest shown by the ants in the dorsal plate covering the last
abdominal segments suggests that the organs, perhaps very small, are
located in that area (Fig. The P. bitias larvae did not respond vigorously
when disturbed; instead, they would simply move slowly away. Malicky
(1971) has described the presence of microscopic epidermal glandular
organs which appear to secrete an ant attracting pheromone. These may
not be associated with Newcomer's organ and the obvious “honeydew”
secretions.

As in the case of felsina, the ants protect the larvae against predation.
When I touched a larvae of either species with a stick, the ants became very
excited, running about and exuding great quantities of formic acid which I
could smell on my hands hours later. To eject the acid, however, these
Campantus do not rise up on their hind legs as was observed in Brazil
(Callaghan, 1977). The formic acid had no apparent effect on the larvae.

The only aggressive behaviour noted on the part of the larvae was the
eating of a J. molpe pupa by two molpe larvae. The larvae had been without
food for two days which I suspect was the reason for their action. This
suggests that the molpe larvae may resort to cannibalism if their food
supply fails.

Both molpe and bitias have a prepupal stage lasting two and three days

42

J. Res. Lepid.

respectively in which they stop feeding and remain motionless on the stem
of the foodplant. They turn a mottled brown color, and, in the case of bitias,
release a large amount of honeydew, no doubt attracting ants which may
provide protection during this vulnerable stage. The larval skin is then
shed and the pupal case hardens. In the case of bitias, the imago emerges
about a month later.

Acknowledgments. I wish to thank Dr. Woodruff Benson for reading and making
helpful comments on the manuscript; Mr. Stanley Nicolay for determining and
providing information onP. bitias; and Dr. Gustavo Lozamo C. and Dra. Isabel S. de
Arevalo of the Institute de Ciencias Naturales, Bogota, for securing identification of
the plant species. Finally, my thanks to Dr. Ernesto Schmitt-Mumm of Bogota for
caring for the larvae during my absence.

Literature Cited

CALLAGHAN, c. J., 1977. Studies on Restinga butterflies I. Life cycle and immature
biology of Menander felsina (Riodinidae), a myrmecophilous metalmark. J. Lep.
Soc. 31(3): 173-182.

GUPPY, J., 1904. Some notes on the early stages of Trinidad butterflies. Trans. Ent.
Soc. Lond. p. 227.

Institute Geografico “Agustin Codazzi”, 1977. Zonas de vida o formaciones vege-
tales de Colombia. Bogota, 237 pp.

MALICKY, H. 1971. New aspects on the association between Lycaenid laNae
(Lycaenidae) and ants (Formicidae, Hymenoptera). Jr. Lep. Soc. 24: 190-202.
MUYSHONDT, 1973. Jr, Lep. Soc. 27: 210-219.

ROSS, G. N., 1966. Studies on Mexican butterflies IV, The ecology and ethology of
Anatole rossi, a myrmecophilous metalmark. (Lep. Riodinidae) Ann, Ent. Soc.
Amer. 59: 985-1004.

Journal of Research on the Lepidoptera

20(1): 43-45, 1981(82)

Immature Stages of Odonna passiflome Clarke
(Lepidoptera: Oecophoridae): Biology and Morphology

Patricia Chacon

and

Marta de Hernandez

Departamento de Biologia, Universidad del Valle, Cali, Colombia

Abstract. This paper describes the larva and pupa of Odonna passiflorae
Clarke (Lepidoptera: Oecophoridae), its host and natural enemies.

The feeding habits of Oecophoridae larvae are extremely varied. In
Colombia, Maesara gallegoi Clarke was found boring into stumps and
limbs of apple trees (Malus sp.), and/?i^a sp, dind Borkhausenia sp. were
found chewing on citrus and wheat seeds respectively (Posada et aL,
1976).

Odonna passiflorae Clarke was found on curuba {Passiflora moUissima
Bailey) vines and has become an economically important plague for this
crop.

Methods and Materials

This research was carried out at Tenerife, a small Colombian town
located in the Andes at 2600 m. It has a temperature of 13.8°C and 81%
Relative Humidity (RH).

Curuba was established as a crop in approximately 1960 in this area of
Colombia and has become the most important source of income for many
peasants. Extensive field observations were made from March 1980 to
July 1981.

Adult individuals were obtained by rearing immature stages in the
laboratory (in Cali, 1000 m, 20°C and 67% RH). Larvae were placed in
plastic boxes (10 x 7 x 3.5 cm) containing 8-10 stem pieces (8 cm long) of
curuba on top of a wet layer of sterile soil-sawdust mixture.

Descriptions of larval stages and nomenclature of setae are according to
Peterson (1962).

Results

Description of the larva:

Bodyi 18-21 mm long, cream white with skin smooth; center of
spiracles creamish, peritremes brown (Fig, la). Head: about 2.1 mm
wide, yellowish brown, height of frons greater than length of epicranial

44

J. Res. Lepid.

Fig. 1. a. Dorsal view of caterpillar; b. Frontal view of head capsule; c. Mandible;
d. Prothorax; e. Mesothorax; f. Fourth abdominal segment; g. Eight
and ninth abdominal segment; h. Crochets biordinal circle.

suture; frons width less than its height; the fourth ocellus is much closer to
the third than to the fifth, and the second is always farther from the first
than the third; setae AdP and Pd^ at the same level; seta Ad^ closer to Ad^
than to Ad^ (Fig. lb). Spinneret narrow and rounded apically; mandibles
with five teeth (Fig. Ic). Thorax: Prespiracular group on the prothorax
trisetose, distinctly separated from the cervical shield which is slightly
dark dorsally; seta rho below seta delta; epsilon posteroventral to gamma;
Pi group (subventral) bisetose (Fig. Id). Mesothorax and metathorax with
the seta beta directly below alpha; eta below kappa and Pi group unisetose
(Fig. le). Pro thoracic spiracle as large as 8th abdominal spiracle and larger
than the others. Abdomen: setae eta and kappa adjacent, located below
the spiracle (Fig. If). On 8th and 9th segments, the seta beta is closer to the
dorsomeson than seta alpha (Fig. Ig). The prolegs are short and possess
circles of biordinal crochets present on segments 3 to 6 (Fig. Ih).

20(1): 43-45, 1981(82)

45

Description of the pupa:

Obtected type; dark yellow, from 9.5 to 13.0 mm long; labmm simple;
labial palpi concealed; without functional mandibles; fronto-clypeal
suture obsolete in middle; antennae diverging at apex and reaching almost
to the tip of wings; fore wings not extending beyond the 4th abdominal
segment; first 4 abdominal segments longer than the rest; epicranial
suture not visible.

Host plant and feeding habits:

The larvae are gregarious. They feed on the main stem and limbs of
curuba plants. The young larva bores into the inner bark of the vine and
continues to the heartwood where it makes long tunnels and irregular
galleries (Fig. 2). The larva maintains contact with the outside and expels
large amounts of sawdust and frass that cling in masses on the bark.

Fig. 2. Injury to the trunk of Passiflora moUissima Bailey caused by the borer
Odonna passiflorae Clarke.

Pupation occurs inside the trunk and the new adult emerges through the
bark, leaving a shot-hole effect. Infestation may be recognized by wilted,
off-color foliage and longtitudinal scars in the bark. The borer prefers old
curub vines (6 years of age), but once the old plants are destroyed, they
attack younger plants (+ 1 year of age).

Natural enemies:

The larval stage is attacked by a number of parasites, including wasps
(Icheumonidae) and flies (Tachinidae). Also, disease causing micro-
organisms sometimes result in death of large numbers of borers. The
fungus Beauveria bassiana has been isolated from dead and dying larvae.
This pathogen has been recognized as an effective control agent.

Acknowledgments. We thank Drs. J. F. Gates Clarke and Goro Kuno for
identif3dng the insect and pathogen respectively. Dr. Benjamin Jimenez criticized
the manuscript. This work was supported by the Comite de Investigaciones of the
Universidad del Valle,
literature Cited

PETERSON, A., 1962. Larvae of Insects. Part 1. Edwards Brothers, Inc. Ann Arbor,
Michigan. 315 pp.

POSADA, L., I. Z. de POLANIA, I. S. de AREVALO, A. SALDARRIAGA, F. GARCIA & R. CARDENAS,
1976. Lista de insectos daninos y otras plagas en Colombia. Ihstituto Colombiano
Agropecuario, ICA. Programa de Entomologia. Publicacion Miscelraea No. 17,
484 pp.

Journal of Research on the Lepidoptera

20(1): 46-49, 1981(82)

A New Genus and two New Species of
Oecophoridae from Colombia (Lepidoptera)

J. F. Gates Clarke

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

Abstract. The genus Odonna and the two species, Odonna passiflorae and
0. xenodora are described as new and figured.

The first species, Odonna passiflorae, is described to provide a name for
this moth which is a pest of Passiflora mollissima Bailey in Colombia.
Because the second species, 0. xenodora, collected over 20 years ago, was
found to be congeneric with the first, it is included for completeness’ sake.

The life history and descriptions of the early stages of 0. passiflorae follow.
Patricia Chacon de Ulloa and Marta Rojas Hernandez submitted the
specimens of the adults to me for identification.

Odonna Clarke new genus.

Type Species: Odonna passiflorae, new species (by present designation).
The gender of the generic name is feminine.

Labial palpus slender, recurved; third segment shorter than second, acute.
Tongue well developed; maxillary palpus minute, appressed to base of tongue.
Head rough, side tufts spreading; ocellus absent. Antenna simple, shorter than
forewing; scape with pecten. Thorax smooth. Forewing smooth, costa nearly
straight, termen oblique, 12 veins; lb furcate; Ic strongly preserved; 2, 3, and 4
equidistant; 5 nearer to 4 than to 6; 7 and 8 stalked, 7 to apex; 10 much nearer to 9
than to 11; 11 fi-om one-third of cell; upper internal vein absent; hindwing with 8
veins; 2 remote from 3; 3 and 4 short-stalked, 5 about equidistant from 4 and 6; 6 to
termen slightly below apex; 6 and 7 subparallel. Hind tibia roughened with long
scales. Abdominal terga not setose.

Male genitalia. Uncus and socius present; gnathos absent.

Female genitalia. Signum absent.

I am unable to reconcile Odonna with any previously described genus. In
my key to the Chilean genera (Clarke, 197 1, p. 2) the present genus keys to
Talitha Clarke, but in Odonna 3 and 4 of the hindwing are short- stalked; in
Talitha 3 and 4 are widely separated. Furthermore, the female genitalia of
the two genera are of quite different types, hi Talitha the ductus bursae
is long and coiled and the signum is present; in Odonna the ductus bursae
is short, uncoiled and the signum is absent. In Meyrick’s Key (1922, p. 3)
this genus keys to the African Ceranthes Meyrick. In Ceranthes 3 and 4 of

20(1): 46-49, 1981(82)

47

Fig. 1. Odonna passiflorae, n. sp. Holotype female.

Fig. 2. Odonna xenodora, n. sp. Holotype male.

forewing are approximate and 7 is to costa but in Odonna 3 and 4 are
separate and 7 goes to apex. I have not seen the genitalia of Ceranthes.

Odonna passiflorae Clarke new species.

Figures 1, 3, 5

Alar expanse 24-30 mm.

Labial palpus white profusely irrorate with fuscous. Antenna buff with grayish
fuscous bars; scape fuscous. Head grayish fuscous. Thorax grayish fuscous, with
some whitish scales posteriorly; tegula grayish fuscous, tipped with whitish scales.
Forewing ground color buff but so heavily overlaid and streaked with fuscous that
the ground is largely obscured; costa grayish fuscous mixed with white scales; veins
streaked with fuscous; at basal fourth, in cell, a blackish-fuscous discal spot; at end
of cell a similar spot preceded by whitish scales; cilia grayish fuscous and buff
mixed. Hindwing buff basally shading to grayish fuscous distally; cilia buff with
slight mixture of grayish fuscous. Foreleg buff heavily overlaid fuscous; midleg
similar but with well-defined fuscous bar on outer side of tibia; hindleg buff strongly
overlaid and suffused fuscous. Abdomen olive buff dorsoanteriorly, fuscous
dorsoposteriorly; ventrally with buff scaling.

Male genitalia slides USNM 25128. 25129. Harpe costa terminating in a short,
sharp point; sacculus produced as a dull-pointed process; cucullus long, slender,
spatulate. Uncus bulbous basally; distally pointed. Vinculum U-shaped; saccus
short, weak. Tegumen more than twice as long as broad. Anellus a small, oval plate
with lateral sclerotized extensions. Aedeagus with slender, sclerotized rod dorsally.

Female genitalia slide USNM 25051, Ostium small, oval in a deep V-shaped cleft

48

J. Res. Lepid.

Fig. 3. Odorma passifhrae, n. sp. a, ventral view of male genitalia mth aedeagus
removed; b, aedeagus; c, lateral aspect of male genitalia with aedeagus in
situ.

Fig. 4. Odonna xenodora, n. sp. a, ventral view of male genitalia with aedeagus
removed; b, aedeagus.

Fig. 5. Odonna passiflorae, n. sp. a, ventral view of female genitalia (papillae
anales missing).

Fig. 6. Odonna xenodora, n. sp. ^g venation.

20(l)i 46-49, 1981(82)

49

m 7th sternum; lamella postvaginaMs granular. Inception of ductus seminalis
slightly anterior to ostium. Ductus bursae membranous. Bursa copulafaix membranus.

Holotype. USNM 100175.

TVpe locality. Colombia, Valle, Tenerife.

Distribution. Colombia.

Foodplant. Passiflora mollissima Bailey.

Described from the 9 holotype (May 1980) and 2 cfcf paratypes as follows: Colom-
bia, Valle, Tenerife, June 1981; Colombia, Valle, July 1981. No collector is
indicated on the labels, but the specimens were submitted to me by Pafricia Chacon
de UUoa and Marta Rojas de Hernandez who may be the collectors.

Although generically distinct, pmsiflorae is similar in appearance to the North
American Apachea barberella (Busck), but is lighter in color and lacks the brush of
the second segment of the labial palpus.

Odonna xenodora Clarke new species.

Figures 2, 4, 6

Alar expanse 38 mm.

Labial palpus missing. Antenna pinkish buff with very faint darker annulations.
Head pinkish buff with fuscous scales posteriorly. Thorax pinWsh buff with sparse
fuscous irroration. Forewing ground color buff with fuscous irroration; in cell a
conspicuous longitudinal streak; at end of cell a white discal spot surrounded by
ftiscous scales; veins streaked with fuscous; around tomus and along termen
fuscous sfreaks; cilia mixed buff and fuscous. Hindwing very pale gray (or sordid
white) with veins indicated by pale gra3dsh fuscous; cilia mixed gra5dsh fuscous and
buff. Foreleg and midleg buff overlaid fuscous; hindleg buff with sparse fuscous
irroration. Abdomen first three segmente golden dorsally, third segment irrorate
with slender fuscous scales posteriorly; remainder sordid white dorsally slightly
suffused and faintly urorate grayish fuscous; ventrally sordid white irrorate with
fuscous; abdominal terga not setose.

Male genitalia slide USNM 25135. Harpe costa short, terminating in a digitate
process preceded by two small cones; sacculus with distal extention; cucuUus long,
straplike. Socius pendant; fingerlike. Uncus broad basally with a basal point on each
side; apex pointed, hooked. Vinculum narrow; saccus slender. Tegumentwo-and-a-
half times as long as wide. AneUus with basal plate and lateral sclerotized
extensions. Aedeagus pointed, with strongly sclerotized rod on one side.

Holotype. USNM 100176.

Ttype locality. Colombia, Cauca, Pm*amo de Purace, Lake San Rafael, 3570 m.

Distribution. Colombia.

Foodplant. Unknown.

Described from the unique d* holotype (29 Jan. 1959, J.F.G. Clarke).

Similar to passiflorae, new species, but larger and easily distinguished by
genitalia.

This species exhibits the phenomenon of the increase in wing area, in proportion
to body weight, induced by altitude, commonly found in Andean species.

Acknowledgments. The photographs were taken by Mr. Victor Krantz, Smith-
sonian Institution. The drawing of wing venation was done by Mrs. Elsie
Froeschner.

Journal of Research on the Lepidoptera

20(1): 50^54, 1981(82)

Literature Cited

CLARKE, J. F. GATES, 1978. Neotropical Microlepidoptera, XXI: New Genera and
Species of Oecophoridae from Chile. Smithsonian Contributions to Zoology,
No. 273: l=-80, figs. 1=54, pis. 1=6.

MEYRICK, E., 1922. Lepidoptera - Heterocera: Family Oecophoridae. In Wytsman,
Genera Insectomm 180: 1-224, 6 plates (color).

Notes

A Recondite Breeding Site for the Monarch {Danaus plexippus, Danaidae) in the
Montane Sierra Nevada

The Monarch butterfly {Danaus plexippus L.) traverses enormous distances in its
annual migrations. These movements must carry it through or over extensive areas
of habitat unfavorable for breeding, such as continuous forest and high-altitude
montane regions, but data concerning its behavior in crossing such potential
barriers are scanty. The Monarch as a breeder is characteristically associated with
disturbed habitats in which species of milkweeds (Asclepiadaceae) behave as
ruderals or range weeds, and little has been documented on its ability to find native
milkweeds in their natural habitats, especially in mountainous regions. It seems
certain that disturbance by man and overgrazing by his livestock have made host-
finding easier for female Monarchs; in the pristine state stands of the hosts must
have been much more scattered in the west, at least.

The Asclepiadaceae do not usually reach high elevations in California. Asclepias
speciosa Torrey reaches 3030 m in the Convict Creek Basin on the east slope of the
Sierra Nevada, but records above 2000 m are generally rare. The Monarch crosses
the mountains in its seasonal migrations (it is observed as a migrant in Donner Pass
in both spring and fall but does not have any host plants there, at 2100 m). Shapiro,
Palm, and Wcislo (1981, J. Res. Lepid. 18(2): 92) found it breeding on A. cordifolia
(Benth.) Jeps. above 2000 m on Packer’s Peak; this is a very isolated stand of less
than a dozen shoots, and required a traverse of at least 1.5 km of continuous forest
from the nearest road. A much more isolated stand colonized by Monarchs was
found on 18 July 1981 in the northern Sierra Nevada by Mr. William Overton and
the author. This stand covers more than 1 ha on both sides of USFS road 19N14,
about 1.5 km NNE of English Mountain Ranch, Sierra County, near Damfine
Sprmg (ca. 1940 m). Asclepias speciosa is the most conspicuous plant, and on the
date of our visit was in full bloom. About 30 adult and 50 larval Monarchs were
observed. The site is located in R13E, T18N, mapped on the USFS “Foresthill
and-Big Bend Districts, Tahoe National Forest” sheet (1966) and in the NE corner
of the USGS Emigrant Gap topographic 15' quadrangle. It is accessible from the
south by USFS road 18N18 from Highway 20 and from the north (Hermess Pass) by
road 19N03.

This breeding site is surrounded by dense, continuous coniferous forest,
completely free of Asclepiadaceae, for at least 9 km in all directions, except for a

20(1): 50-54, 1981(82)

51

small and equally isolated boggy meadow at English Mountain Ranch (no
mdkweeds). The nearest Imown Asclepiads are about six plants of A. cordifolm at
1700 m near the Bear River on USES 18N18, an airhne distance of 17.5 km from
Damfine Spring. There are few flowers and almost no butterflies along these roads,
except at English Mountain Meadow, where no Monarchs were seen. In fact, no
Monarchs were seen from the junction of 18N18 with Highway 20, via 19N14 and
19N03 to Henness Pass Road to Highway 89, a 65 km trip, except at Damfine
Spring. The only other butterflies seen at Damfine Spring were male Speyeria
zerene ssp., despite the superabundant nectar resources.

There is no question of the Monarch’s ability to toaverse the distances from its
more usual agricultural-ruderal breeding sites to Damfine Spring, but what is
striking is that it was able to find and colonize so remote an island of milkweed in a
sea of conifers. One may be forgiven for speculating that the probability of such
sites being found by parasites of the Monarch is much lower than more conventional
ones, and that they may contribute disproportionately to seasonal reproduction.
We will watch this site in future years to determine whether it is routinely used and
whether parasitism is high or low compared to more open, weedy sites.

Arthur M. Shapiro, Dept, of Zoology, University of California, Davis, CA 95616

Mate-Locating Behavior of Gnophaela latipennis vermiculata G. & R. (Pericopidae)

Gnophaela latipennis vermiculata is one of the most conspicuous day -flying moths
in the Rocky Mountains. It is figured by W. J. Holland (1903 reprinted 1968. The
Moth Book. Dover Publ., N.Y.). Adults are typically found in moist vdley bottoms
in the Canadian Zone. The strangest feature of the moths is that nearly half of the
adults seen are copulating, on yellow (sometimes white) flowered Compositae
growing in meadows or near streams. In their frequency of copulation they rival only
the bluish Epicauta beetles (Meloidae), popularly termed love bugs, that mate and
feed on various legumes in Colorado. The moths are nearly absent in the morning,
and start to patrol the valley bottoms starting about 1300 (24-hour standard time),
and patrol conspicuously the rest of the afternoon. I found many copulating pairs at
1310, 1350, 1400, 1420 and 1830, usually resting on the composite flowers. Adults
sip nectar from the composites also. Adults are probably distasteful to most
predators, because they fly slowly and are very conspicuous, I have collected ssp.
vermiculata at Taos Ski Basin, 10400', Taos County, New Mexico, August 22-23,
1979; Toll Ranch, Gilpin County, Colorado, July 28, 1977; 3 miles NW Nederland,
Boulder County, Colorado, July 24, 1977; Jim Creek, Grand County, Colorado,
August 9, 1977; Diamond Peak, Moffat County, Colorado, July 8, 1972; Willow
Springs Guard Station, Sublette County, Wyoming, August 8, 1980; and ssp.
latipennis Bdv. at Cedar Pass, Modoc County, California, August 4, 1974; and 2
miles W. Old Mill Campground, Colusa County, California, June 8, 1974 (mating
was not observed in ssp. latipennis).

James A. Scott, 60 Estes Street, Lakewood, Colorado 80226

52

J. Res. Lepid.

On Colias hecla Lefebvre re a recent paper by Oosting & Parshall (Lepidoptera:
Pieridae)

In a recent paper on the butterflies found in the region of Churchill, Manitoba
(Oosting & Parshall, 1978(80), Ecological notes on the butterflies of the Churchill
region of northern Manitoba, J, Res. Lepid. 17(3): 188-203), the authors misquoted
a statement that I made about the behavior of Colias hecla Lefebvre in an earlier
paper (Ferris, 1974, Notes on arctic and subarctic collecting, J. Res. Lepid. 13(4):
249-264). I would like to correct this situation, and offer some additional comments
about the behavior of this butterfly.

With reference to the paper cited above, Oosting and Parshall have stated: “The
authors have found no records of this butterfly from the taiga zone. Ferris (1974, p.
257) misquotes Masters (1971, p. 8) when reporting the occurrence of hecla below
treeline.” The statement that I made concerning the habitat of hecla was based
upon my own collecting experience at Churchill in 1973. Specifically, I stated:
“When they first appear on the wing, Colias nastes and hecla can be found a mile or
more into the Taiga, along the railroad right-of-way (cf. Masters, 1971, p. 8).”
Apparently Oosting and Parshall misinterpreted the abbreviation cf., meaning
compare. In this case, compare with Masters (1971) who stated: “Colias hecla is
quite rare at Churchill, but is apparently a breeding resident there. As far as I know
(I have not personally taken the species) all of the examples taken have been taken
over climax high tundra.” (Masters, 1971, The butterflies of Churchill, Manitoba,
The Mid-con. Lepid. Ser. 2(25): 1-13.)

In 1973 at Churchill, I collected C. nastes moina Strecker first in the taiga well
south of Dene village, and later, as the season progressed, in the climax tundra. My
only captures of C. hecla hela Strecker were in the taiga in open clearings. I departed
Churchill in mid-July, apparently before hecla appeared over the climax tundra. In
addition to apparent habitat preferences, there are phenotypic differences in C.
hecla at Churchill which I have reported in another paper that discusses the
taxonomy of C. hecla (Ferris, 1981, Revision of North American Colias hecla
Lefebvre (Pieridae: Coliadinae), Bull. Allyn Mus. Entomol. In press.)

I have also collected C. hecla in British Columbia and Alaska, and have accurate
records from the Yukon Territory and other arctic regions. In British Columbia,
hecla can be taken in open clearings in forested areas (taiga) along the Alaska
Highway. It shows a definite preference for the World War 11 emergency airstrips,
now overgrown with low vegetation. It occurs in similar areas in the vicinity of
Kluane Lake and Haines Junction, Yukon Territory.

My experience with C. hecla in Alaska has been at Murphy Dome (NW of
Fairbanks), McKinley Park, and Eagle Summit (ca. 110 miles (68 km) N. of
Fairbanks on the Steese Highway). Murphy Dome is above treeline and the “alba”
female form of hecla may be taken there. The only hecla that I observed in McKinley
Park were flying over tundra (above treeline). At Eagle Summit, hecla was collected
in the forested area below and to the north of the Summit proper. Above-and-below
treeline habitats are common in Alaska for several butterfly species, including:
Oeneis jutta ssp., Clossiana chariclea ssp., Clossiana polaris ssp. and Clossiana
freija ssp.

From my experience with many species of arctic and arctic-alpine butterflies,
flight patterns sometimes vary annually with certain species. This perhaps accounts
for the reason that I found C. hecla in the taiga at Churchill, while Oosting and

20(1): 50-54, 1981(82)

53

Parshall did not.

Clifford D. Ferris, P. O. Box 3351, University Station, Laramie, WY 82071

Further migrations of Hipparchia semele (L.) in 1976 and 1980

A substantial migration of the Grayling, Hipparchia semele (L.) was reported in
France in the autumn of 1975 {J. Res. Lepid. 15(2): 83-91, 1976). Since then two
other migrations of H. semele have been seen, one in the autunm of 1976 and the
second in 1980. There were no movements of butterflies during the other years.

No quantitative results are available for the migration of 1976 which happened in
early September in the same place, that is at the French address above. The
migration of 1980 occurred a little earlier in the same locality and was seen to be in
motion a few days before the following results were collected by one of us (P.D.) on
the 29th of August. The following day an outbreak of stormy weather prevailed and
persisted until the 7th of September when there was a feeble resumption of the
migratory stream.

Table I

Number of Butterflies Passing a Transect

Time Number of Butterflies Seen Eveiy 6 Minutes Total

1000 - 1100 7, 12, 14, (-), 8, 19, 17, 22, 23, 20, 24, (-) 166

1100 - 1200 21, 22, 25, 19, 15, 24, 27, 26, 29, 19, (-), (-) 227

1230 - 1330 20, 22, 19, 24, 27, 24, 23, 26, 24, 27, 20, (-) 256

1330 - 1430 22, 19, 20, 18, 16, 20, 15, 13, 10, 12, 7, 5 177

1430 - 1530 9, 6, 5, 8, 10, 4, 6, 3, 2, 0, 1, (-) 54

Grand Total 880

^ediods as described in 1976 paper, (-) ^ not recorded)

The results show that there was a peak of flight activity between 1100-1300 with a
maximum of 29 butterflies recorded during one five minute period. In 1975 no such
clear maximum was apparent.

The numbers of butterflies on migration in 1975 and 1980 were equally
impressive, 596 seen during four hours in 1975 and 880 seen in five hours in 1980.

It is interesting to note that H. semele was migrating with dragonflies which
apparently move in this southeast direction every year at the end of August.

hi conclusion we would like to say that the migration of H. semele appears to be a
fairly regular occurrence in this region and has been recorded in three out of six
years.

John Feltwell, Marlham, Henley Down, Catsfield, East Sussex, TO33 9BN,
ENGLAND and Patrick Ducros, Cabanevielle, St. Martial, Sumene, 30440,
FRANCE

54

J. Res. Lepid.

An Interfamilial Courtship (Lycaenidae, Pieridae)

On 6 April 1982 a fresh male Glaucopsyche lygdamus cf. incognitas Tilden was
observed pursuing an unidentified larger butterfly near Davis, Yolo County,
California. The larger butterfly was unidentifiable in flight: it appeared silvery-
^ay with a bluish cast and was reminiscent of the Argentine “silver Satyrid,”
Argyrophorus argenteus (Blanch.). The pursuit, which was first noted at 1255 hours,
continued for about three minutes on a steep highway embankment. During this
time the unidentified butterfly lit momentarily twice. I was able to overtake the pair
as it lit a third time and determined it to be a white female Colias eurytheme Bdv., of
the vernal phenotype, exceptionally small and very heavily infuscated beneath. The
male lygdamus lit on the forewing of the Colias as it positioned itself for
thermoregulatory lateral basking (the ambient temperature was only about 13°C)
and then attempted repeatedly to make genital contact fi'om a position centered
roughly over the hindwing discoceUular spot. The pair was collected at 1303 after
the female rose to take flight. Her left forewing measured only 20 mm from base to
apex, quite small for a Colias but considerably larger than the male’s 14 mm. She is
among the most melanic vernal examples in our very long series at Davis.

Females of the Davis population of lygdamus are partially blue dorsaUy and
lustrous pale gray-brown ventrally. The distinctly silvery-gray-bluish cast of the
female Colias in sunlight, which was baffling to me, very probably fooled the male
lygdamus as well. Fresh female lygdamus and mating pairs were observed in the
immediate vicinity.

Arthur M. Shapiro, Dept, of Zoolo^, University of California, Davis, CA 95616

Celastrina ladon (Lycaenidae) Females Ovipositing on Sambucus canadenis, a
Plant Unsuitable for Larval Development

From 8 to 16 June 1981 1 repeatedly observed female Celastrina ladon (Cramer)
oviposit on the flower buds of Sambucus canadensis L. (Caprifoliaceae) in Upper
Tyrone Township, Fayette Co., Pennsylvania. A close examination of the six
flowering bushes in the general area revealed that each flower head contained from
one to two up to several dozen eggs. Wild eggs taken from the flower heads and ones
laid on flower buds by captive females hatched normally in the laboratory, and the
larvae began to feed. Within two days, however, all larvae left on Sambucus had
died. About 50 larvae were transferred to offlcinaUs QL.,) Lam. (Leguminosae),

the dominant foodplant of second generation C. ladon in this area, after feeding on
Sambucus for only a few hours. All but two or three of these larvae died during the
next day. A further examination of Sambucus in the area of oviposition observations
and in Preston Co., West Virginia, revealed many additional unhatched eggs and
shells of hatched eggs, but no sign of larvae. All parts of this plant contain
hydrocyanic acid (G. A. Petrides, 1968, A Field Guide to Trees and Shrubs,
Houghton Mifflin, Boston, p. 32), and the flower buds are apparently toxic to young
C. ladon larvae. In light of this, it is surprising that females should oviposit so readily
on this plant in nature.

Charles G. Oliver, R. D. 1, Box 78, Scottdale, Pennsylvania 16683

Journal of Research on the Lepidoptera

20(1): 55-64, 1981(82)

Book Reviews

The Audubon Society Field Guide to North American Butterflies.

Robert Michael Pyle. 1981. 916 pages, 759 colored figures. Alfred A. Knopf, New
York. Price: $11.95, paperback.

The best feature of this book is the superb color photos of butterflies in liatural
settings. Generally these show true colors and picture all North American species.
However, a few figures depict rubbed, out-of-focus, or even “nature faked”
specimens. One exasperating aspect is that only common names (some newly-
invented) are given for the figure captions and text discussions. The photos alone
are not always adequate to identify certain species of Speyeria, Euphydryas,
Euphilotes, Thoryhes, Erynnis, Hesperia, Agathymus and other taxonomic problem-
genera.

The introduction stresses watching, photographing, and rearing butterflies and
cautions against over-coUecting. The text emphasizes description, similar species,
life cycle, flight times, habits, habitats, and range. Entirely too much space is
devoted to the descriptions and similar species. Because of the identification
format, the organization is unorthodox. There is little mention of subspeciation
which is rampant in the West. Hostplant data is accurate but skimpy. The style is
variable, often captivating and drawing on a wealth of information, but sometimes
verbose and redundant. It is not clear how much of the information is from Pyle’s
personal experience, the literature, or from others, as facts are not backed up by
referencing. The habits, habitat, and conservation discussions are positive features.
Special attention is given to migrant species and weedy species of disturbed
habitats. The new Brown & Miller check list is followed rather disconcertingly in its
generic splitting, and the commoner species’ synonyms are mentioned. In the back
there is a glossary of terms, picture credits, host plant index, and butterfly name
index (common and scientific names combined).

Some important biogeographic information is scattered in the text. For example,
Neohasia is related to the South American Catasticta, and Neominois, Hahrodais,
and Hypaurotis are related to East Asian species. Strymon avalona and Boloria
acrocnema have the most restricted ranges of any Nearctic species, and Plebejus
neurona, Lycaerm hermes, and Stymon avalona are peculiarly endemic to southern
California. Vanessa tameamea (Hawaii) is probably derived fi*om the Asian V.
mdica, and another Hawaiian endemic, Vaga blackbumii, is related to a Bonin
Islands species.

Ranges are generally accurate, but Euptoieta claudia and Eurema mexicana are
not residents of Southern California, Libytheana bachmanii is not mentioned for
Arizona and Southern California, and Speyeria callippe is not mentioned for
Southern California. Under Speyeria edwardsii, “Of the western fritillaries, this is
the only one which varies constantly throughout its range,” is a misstatement.
Oakley Shields, 4890 Old Highway, Mariposa, CA 95338.

The color photos in tMs book are superb, and make the book worth the price. The
photos are mostly of living butterflies in natural surroundings. The text is suitable

56

20(1): 65-64, 1981(82)

for the novice butterfly enthusiast, and has the virtue of giving a description, life
cycle, flight, habitat, and range for most species.

The book also has shortcomings and numerous errors. Identification is made more
difficult by four drawbacks: a) at least 27 photos are misidentified (correct names
are: 326 & 337 P. polyxenes, 89 C. interior, 108 P. agarithe, 127 K. lyside, 516 L.
helloides, 467 I rdphon, 430 C. apama homoperplexa, 509 C. argiolus, 543 C.
nemesis, 553 E. ares female, 365 L. bachmannU larvata, 607 left S. coronis halcyone,
613 left C. bellona, 590 P. campestns female, 368 leftP zephyrus, 368 right iV. vau-
album, 654 A. iphicla, 759 H. guatemalena, 758 H. febnm, 686 S. blomfildia, 663
right A. leiUa, 662 right A. celtis, 732 0. taygete, 735 O.juttal, 285 T. con^is female,
165 0. sylvanoides—the black pages make correcting these errors difficult— one
must use “Liquid Paper” to whiten a space for inking); b) many photos are of
butterflies in postures that make them unidentifiable (upperside instead of
underside, or vice versa, or a view showing only the body such as photos 205-210,
the diagnostic spot of A. belli is hidden ^the HW on photo 199, etc.) so individuals
of these species cannot be identified; c) many species have no photo, and are
illustrated by line drawings in the text, which are generally small, rather faded, and
poorly drawn; d) many species have no photo or drawing, and are mentioned only in
1-2 lines appended to another species. Photos 1 16, 323, 380, 381 and 475 seem to
represent dead butterflies. It will evidently take many more years to accumulate
good photos of all the North American butterflies in a living state. The photos are
sorted into 14 groups based on rough appearance (the white butterflies are mostly
grouped together for instance, although the white fonns of Colias are shown with the
yellow butterflies). The grouping of many species is debatable, although the
amateur may find this feature of the book useful. There are only three photos per
page, so considerable flipping through the small book is necessary to find the right
photo. These deficiencies in the photos mean that only the visually distinct species
will be able to be identified using the book, and species that are similar to others
cannot be identified to species.

The photos give only the common names, many of which are new, so to detemodne
the identity of them one must turn to the text. Most traditional common names are
retained, although many are new, and some new ones are unfortunate (for instance
die “Sheep Skipper”, A. ovinia, is named forthe Navajo Indians’ sheep, butactually
A. ovinia inhabits extreme southern Arizona; the Navajos inhabit northern
Arizona).

Another limitation is the coverage of the book. Five hundred seventeen species
are illustrated in color (excluding those misidentified), 61 in line drawings, and the
remaining about 122 of the 700 North American species are not illustrated at all.
Only about 600 of the 700 species have full treatment in the text. Two widespread
native species, Euphilotes spaldingi and Cogia outis, and about 30 strays to the
United States from Mexico or the Antilles, are not mentioned at all. Pseudolycaena
marsyas does not occur in the United States.

The text is adequate, although numerous errors detract from its value. The
generic names used in the book are highly split, many familiar names such as PapUio
glaucus and Pieris rapae falling by the wayside. Some species names are doubtful
{Calephelis “guadeloupe” a synonym of nemesis, Euphyes “ruricola” a nomen
dubium, Celastrina ebema a synonym of C, nigra, Agriades *‘frankUm’'isA. glandon,
L. astyanax a ssp. of L. arthemis, etc.).

20(1): 55-64, 1981(82)

57

Many larval foodplants are errors. Some plants are lab hosts only, and are
doubtfully used in the field {Heracleum for Papilio joanae, Trifolium for Phoebis
sennae, Eurema lisa and Zerene eurydice, Muhlenbergia for Lethe anthedon, Althaea
for Celotes nessus, Stenotaphrum for Nastra Julia, Prosopis juliflora for A. palmeri,
Poa, Avena and Cynodon all for Amblyscirtes vialis). Thlaspi is refused hy Pieris napi
larvae. Horkelia and Potentilla are doubtful for Lycaena xanthoides editha.
Polygonum douglasii applies to Lycaena nivalis, not L. mariposa. The S. macfarlandi
host is Nolina texana, not microcarpa. Ceanothus macrocarpus is now megacarpus
for S. saepium. Myrica is refused by Calycopis cecrops females and larvae. Trema
floridana is now micrantha for S. martialis. Salicomia ambigua is a synonym of
virginica for B. exilis. Hosackia purshiana is dubious for Plebejus emigdionis.
Castilleja is an error for C. palla. Ulmus is an error for P. zephyrus, Rhododendron an
error for P. oreas. Sorghastrum nutans is dubious for N. areolata. Asclepias is
unverified for Danaus eresimus. The E. zestos host is a herb, not woody. Canna is an
error for 0. proteus. The A. anaphus host is “wild bean,” not Phaseolus. Malva,
Abutilon and Althaea plants are based on H. Tietz’ “syrichtus” and apply to P.
communis, notP. oileus. Malva is an error for P. catullus. The host ofEuphyes dion,
Scirpus, is a bulrush (Cyperaceae), not a rush (Juncaceae). Sorghastrum nutans and
secundum for Amblyscirtes hegon are somewhat doubtful, based on Abbot.
Agathymus neumoegeni larvae do not feed in roots as stated. Certain plants are
misspelled throughout: Dryapetes, Saccharum, Gaultheria humifusa, Porlieria,
Rumex triangulivalvis Ctriangularis), Quercus gambellii, Cuscuta, Rivina, Lippia
graveolens, Prunus havardii, Stenandrium.

Few hibernation stage records are given, and nearly half of those given are wrong.
The correct stages are: Colias philodice and eurytheme 3-4 stage larvae, Lycaena
phlaeas and cupreus half grown larvae, Lycaeides melissa half grown larva, S.
hydaspe newly hatched larva, Erynnis baptisiae probably mature larva, Hesperia
comma egg, H. metea mature larva, Polites mystic 4th stage larva (see J. Res. Lepid.
18: 171-200).

The larva and pupa descriptions are mostly adequate, although brief. The
Panoquina panoquinoides larva and pupa descriptions refer to P. errans. The
descriptions of larvae and pupae of Erynnis are poor and are repeated almost
verbatim for most of the Erynnis species, including several species that have never
been reared.

The ranges are mostly correct although often underestimated. However, some
regions stated are errors: P. pilumnus, Z. eurydice, C. definita, C. perditalis and C.
mossii do not occur in Arizona, H. charitonius is not a resident in South Carolina, P.
orseis is not in NE Nevada, T. leanira is not in E , Colorado or New Mexico, Z. cyna is
not in New Mexico, E. favonius is not in West Virginia or Louisiana, C. sheridani is
not in Saskatchewan, C. hesseli is not in Mississippi, H. grunus is not in Idaho or
“coastal Arizona”, L. xanthoides is not in the Great Basin, L. phlaeas is not in
Florida, C. occidentalis is not in Alaska, A. pima and P. rudkini are not in SE Utah, P.
beckeri is not in Black Hills, P. hylas is not in W. Nebraska, the P. zephyrus record
from Manitoba was a misdetermination. P. progne, P oreas is not in NE Wyoming,
C. henshawi is not in W. Texas, N. areolatus is not in Kentucky, P araxes is not in S.
Texas, P manueli is not now in Florida, P. mercurius records from Arizona-New
Mexico are errors, U. teleus and A. toxeus and G. gesta records from Arizona are
errors, E. persius is not in Maritimes, P. alpheus is not in Oregon, N. Julia is not in

58

J. Res. Lepid.

Alabama, K dacotae is not in Illinois, H. sassacus is not in Tennessee, H. attains is
not in Nebraska or Wisconsin, P. origines is not in Montana, P. zabulon is not in
Wisconsin, P taxiles is not in Nevada, P. viator is not in Maine, P. hobomok is not in
the Sangre de Cristo Mts., A. aesculapius is not in Connecticut or New Mexico, M
cofaqui is not in N. Texas.

Some sedentary species are wrongly listed as strays (C. goodsoni, P. erram,
Agathymus (“Brown BuUet”) estelleae, Pholisora mejicana), and Heliconius erato is
treated as a native with no mention that it is actually a very rare stray.

The flight periods listed are mostly correct (although iV. terlooti flies in June-July
also, Colias alexandra has two broods on the plains, Boloria eunomia has a several
week flight period (not 4-5 days as stated), etc.). The number of broods for
multivoltine southern species are underestimated throughout the book {Eurytides
marcellus has 3-4 broods from April-October, not two, for instance); in many cases
when only two broods are listed the number of generations per year is probably four
or more.

The introductoiy material gives very elementary information for the beginner on
anatomy and the life cycle, etc. Concepts such as photoperiod do not appear in the
book. Flowers visited by adults, and adult migrations, are listed. There is nothing on
mate -locating behavior in the book, although anecdotal reference to territoriality
are given throughout the book, although the application of this concept to
butterflies is disputed by research workers. Pheromones are attributed to T. elada,
S. hydaspe and L. creola, when really they have never been studied. The white
female form of Ascia monuste also migrates. Colias behrii is a subalpine, not an
alpine, species, and the definition of alpine in the glossary (“pertaining to or
inhabiting mountains”) is peculiar.

OveraD, the photos are excellent, and the text is suitable for the beginner, but the
numerous errors mean that the book must be used with caution by lepidopterists.

James A. Scott, 60 Estes Street, Lakewood, Colorado 80226

Butterflies of the Rocky Mountain States

Ferris, C. & F. M. Brown, eds., with 8 contributors (F. M. Brown, D. Eff, S. L. Ellis,
C. D. Ferris, M. S. Fisher, L. D. Miller, J. A. Scott and R. E. Stanford). 1981 . 442 p.
Univ. Oklahoma Press.

This book should serve a useful purpose in allowing the identification of Rocky
Mountain (exclusive of Canada and Alaska) butterflies and skippers, and present-
ing their distributions, and a little about their foodplants and habits. Overall, it is a
good book, especially for distribution and identification. It does contain some
errors, however. I have studied Rocky Mountain butterflies for 23 years, and hope
these comments will be useful to Lepidopterists who use the book as a factual
source.

The maps are the best feature of the book, and represent a considerable amount of
effort by Stanford (and the reviewer); the one drawback of the maps is that the dots
are placed in the middle of each county rather than at the exact collection site, which
makes the ranges in mountainous and other non-uniform areas appear too large in
many cases. The county lines usually run along the top of mountain ranges, so two

20(1): 55=64, 1981(82)

59

specimens from the top of a peak may be widely separated on the maps. The
boundaries of the area covered as shown on the maps are not rigidly adhered to in
the text: Poladryas minuta minuta from northeastern New Mexico is missing from
the book (more on it later), as is Euchloe creusa which will probably be found in
Glacier National Park, while Atrytonopsis deva, Thymelicus lineola, Ancyloxypha
arene, Precis evarete nigrosuffusa, and Ascia josephina (a possible stray) do not
occur near the region but are fully treated in the book, and the following are given
full treatment even though they are almost certainly mislabeled from the region:
Thoryhes diversus, Poanes zabulon, Pompeius vema, Wallengrenia egeremet, Piruna
poUngii, Emesia zela, Parrhasius m-album, Cyllopsis henshawi It should be noted
that timberline is about 7500' on the Montana-U.S. border, not 9000' on p. 15 or
11000' on p. 289.

The photos are black and white (except for several color plates), but are well done
for the most part, although some are a bit faded (especially Oeneis jutta), the first
photo of Oeneis chryxus is actually 0. uhlen, the photos of Cercyonis pegala damei
are C. p. boopis, the first photos of Pieris sisymbrii and Cyllopsis henshawi are
females not males, the scale of the ‘‘Pyrisitia” proterpia and Nathalis photos is not
IX, and the Everes comyntas photos are of specimens that unfortunately resemble
E. amyntula.

The authors are not properly credited in the text in many cases; one must consult
the Table of Contents to find who authored what.

Butterflies are made to appear ferocious in the book. “Attacks”, “mock combat”,
“pugnacity”, “combative” and “territories” on pp. 60, 226, 228, 232, 346 and 348
are actually misinterpretations of male mate-locating behavior (see Amer. Midi.
Nat. 91: 103). Actually, butterfly adults have no offensive weapons with which to
fight, and it is the larvae which are often territorial (see J. New York Ent. Soc. 81:
214 and Oecologia 52: 415 for descriptions of larval battles) or even cannibalistic.

Considerable space in the book is devoted to purely taxonomic matters (type
localities and dates of publication, type species of genera, etc. are given), which is
surprising in a regional work. Numerous taxonomic changes are made,, and
comparing the scientific names with such books as Klots (1951) Field Guide,
Ehrlich & Ehrlich’s (1961) How to Know the Butterflies, and Howe’s (1975)
Butterflies of North America, the book is an orgy of splitting. The cabbage butterfly
is no longer Hens, the Tiger Swallowtail no longer is Papilio, etc., etc. A. Klots (Bull.
Brooklyn Ent Soc. 31: 154) and T. N. Freeman (Can. Ent. 68: 277) both published
generic revisions of Lycaena, but their work is scrapped and numerous genera used
which differ very slightly (L. rubMus and L. xanthoides in particular have very
similar genitalia contrary to p. 229, and hybrids between them are known, see J.
Res. Lepid. 8: 51 & 18: 50). Apparently the authors used the genera found in the
new “A Catalogue/Checklist of the Butterflies of America North of Mexico’' by L.
MOler & F. Brown (Lepid. Soc. Memoir no. 2). That work, however, abolishes
subgenera completely (not a single one is used), which is contrary to the opinions of
most zoologists and is certainly not warranted by any rale of nomenclature or logic;
ffierefore, these generic changes should be ignored by aU zoologists except those
few who share a disdain of subgenera. Some names in the book do differ from the
Miller/Brown catalogue: the Catalogue uses Polites coras for peckius*, Euphyes
ruricola for vestris*, Agriades franklini for rustica (glandon*), Atrytonopsis cestus
margarita for python margarita*, EupMlotes pallescens for rita palkscens*, Celastrina

60

J. Res. Lepid.

ladon for argiolus (argiolus ladon*), Incisalia fotis mossii for mossii*, Coenonympha
ochracea, ampelos and inomata as three separate species instead of ssp. of C.
tidlia*, and Erebia epipsodea rhodia* for epipsodea (the asterisks show the names
that are probably correct). The studies of higher classification by P. Ehrlich, almost
the only person this century who has scraped off the body scales to see what is
beneath (Univ. Kansas Sci. Bull. 39: 305-370) are ignored, and doubtful categories
such as Anthocharinae and Marpesiinae are used. Anthocharis andEuchloe actually
belong in the Herinae close to Pieris; all have the same foodplants and body
enzymes (see J. Res. Lepid. 19: 181).

The following are detailed comments, arranged in taxonomic sequence, proceea-
ing through the book from start to finish. Hesperiidae. The white fringed ssp. of
Thorybes pylades does not occur in Idaho as stated on p. 71. Erynnis properdus and
meridianus have the same genitalia, as doE. zarucco and funeralis, and appear to be
conspecific (and see p. 79). The foodplant anserina totPyrgus xanthus is a
misidentification of P. hippiana. Amblyscirtes simius does not perch late in the day,
only in early morning (see J. Anim. Ecol. 42: 663); the observations reported were
made after a dark thunderstorm, and are highly exceptional, if indeed they
represent mate-locating behavior. Amblyscirtes aenus has been reared from eggs
laid by A. “ema” (J. Res. Lepid. 15: 92), so ema is a form of aenus, not a species as
implied on p. 94. Amblyscirtes fimbriata has orange fringes. The type locality of
Atrytone logan lagus is Oak Creek, Fremont (not Custer Co., CO, which is probably
an error anyway because the species does not occur in either county now. Ochlodes
sylvanoides napa is a weak ssp. only in the Colorado Front Range, distinguished
only by larger size (see Papilio (New Series) no. 1, available from the reviewer).
Polites sabuleti is not alpine as stated in the figure caption on p. 118. P. draco and
sabuleti actually have identical genitalia and no structural differences (contrary to p.
119), and apparent intermediates occur on Grand Mesa, Colorado. The Festuca
idahoensis foodplant of Polites sonora is only a guess by E. J. Newcomer (J. Res.
Lepid. 5: 243) (actually some moist meadow grass is more probable), and Distichlis
is only a guess tor Atalopedes campestris (by L. Orsak, Butterflies of Orange County,
Univ. Calif. Press). Hesperia comma Colorado is actually annual in appearance, and
has adapted to the subalpine zone with a shorter developmental period (J. Lepid.
Soc. 29: 156). It is not biennial as stated on p. 125. Ssp. assiniboia has an ochre
VHW. Hesperia woodgatei males actually perch on hilltops (J. Res. Lep. 14: 1) and
not in gullies as stated on p. 128. Hesperia nevada males usually perch on hilltops
also. H pahaska flies 1% months (not 3) earlier than H leonardus montana. The
foodplant Digitaria of Copaeodes aurantiaca is an error by H. Tietz (An Index to the
Described Life Histories, Hosts...; AUyn Museum). Megathymus coloradensis is
treated as a species even though the latest revision by K. Roever in 1975 treated it
as a ssp. of M. yuccae (p. 144 correctly notes the limited utility of chromosome
counts). M. streckeri texanus is characterized by the large DHW postmedian spots in
females, rather than the characters given.

Pieridae, Neophasia menapia tau is a synonym of menapia (the California coast
ssp. was recently named ssp. melanica, Papilio no. 1). Pieris napCa range decreased
in eastern United States due to deforestation rather than due to competition withP.
rapae as stated on p. 19 (see Amer. Nat. 1 18: 655). P. napi and Anthocharis sara are
not semicrepuscular, they merely prefer shade. Pieris chloridice is misspelled twice
on p. 150. L. Higgins’ studies combining Pieris chloridice beckerii, P. callidice

20(1): 55-64, 1981(82)

61

occidentalis and Euchloe ausonia ausonides are ignored, and the combination
Anthocharis cethura pima of T. Emmel and J. Emmel is ignored. E. ausonia
coloradensis occurs only in the southern Rockies according to P. Opler’s revision (J.
Res. Lep. 7: 65), and ssp. ausonides occurs from central Wyoming northward. Colias
meadii elis occurs in Glacier National Park, and possibly in NW Wyoming, and fMes
as strongly as ssp. meadii in the reviewer’s experienc.e. C. eurytheme has a spring
form which is mostly yellow and is far from meaningless, being darker on VHW for
better thermoregulation (see Proc. Nat. Acad. Sci. 63: 768). The appearance,
habitat and behavior of C. scudderii and gigantea suggest they are conspecific, and
the combination C. scudderii gigantea was proposed by John Masters. Trifolium is a
lab host only for ''Pyrisitia” lisa. Stellaria media (Caryophyllaceae) is probably an
error for Nathalis.

Papilionidae. Pamassius phoebus actually hibernates as eggs. Ssp. pseudorotgeri
occurs only in the San Juan Mountains, and not in the Sangre de Cristo Mountains.
The reviewer does not know of any evidence that Pamassius clodius contains
cyanide as stated. The yellow tegulae of Papilio bairdii are very useful for
identification; they are much yellower than inP. polyxenes. Ssp. dodi appears to be a
synonym of P. bairdii brucei. Artemisia is not a foodplant of Papilio indra. Papilio
rutulus is probably a subspecies of P glaucus, because they intergrade in southern
British Columbia in male and female genitalia and wing pattern (Pan Pacific Ent.
52: 23), they intergrade in the Black Hills (Evolution 13: 40). Ssp. canadenis does
not occur in the Black Hills as stated on p. 187, they merely resemble canadensis
there because of the intergradation. Ssp. arcticus is an Alaskan form of P. glaucus
canadensis that resembles rutulus, and not a ssp. of rutulus as stated on p. 188;
rutulus does not occur north of SW British Columbia. Magnolia is not eaten by
Papilio palamedes (J. Lepid, Soc. 16: 198).

Lycaenidae. Apodemia mormo ssp. mejicanus actually occurs in the region, not
ssp. duryi which has a greater extent of more orange color and occurs in S. New
Mexico. The report of Lycaeides idas from North America is confusing (p. 201).
Actually, V. Nabokov’s systematic studies (Bull. Mus. Comp. Zool. 101: 479-541)
are perfectly correct; however nomenclatorial changes in Europe now mean that
Nabokov’s Palearctic species “ismemas” is a synonym of argyrognomon, and
Nabokov’s ‘"argyrognomon” must be called idas (Forster, 1936, Mitt. Munich Ent.
Ges. 26: 41-150). So all references to “argyrognomon” in North America must be
changed to idas. The only two species ot Lycaeides in North America are melissa and
idas. Plebejus acmon lutzi does not eat Lupinus or Astragalus in the region; they are
eaten in California and Arizona only. There are many reasons for using the species
combination Agriadesglandonrustica instead of A. rustica (J. Res. Lepid. 17: 101).
0. Shields treated pallescens as a ssp. of Euphilotes rita (J. Res. Lepid. 16: 2).
Alfalfa is not known to be eaten by Everes comyntas (the record referred to is
actually Medica^o lupuLina, Black Medic). Ssp. herrii belongs toE. comyntas, not to
E. amyntula. Trifolium is not eaten by E. amyntula as far as known. The Rocky
Mountain ssp. of Celastrina argiolus is ladon, or possibly sidara, but definitely not
cinerea, which is related to ssp. echo of California. The forms of argiolus in most of
the Rockies are those that occur in ssp. ladon. Amaranthus is not a valid foodplant
for Brephidium exilis. Lycaena arota males perch only in the morning, and males
seldom patrol to find females (see J. Lepid. Soc. 28: 64). The arctic/alpine North
American Lycaena phlaeas are very closely related to Scandinavian L. phlaeas

62

J. Res. Lepid.

polaiis, even to the extent of having the same form caeruleopunctata with blue spots
on DHW; a relationship ignored in .^nerica. Lycaena cupreus in Montana, NW
Wyoming, and Utah occurs in the Canadian Zone and represents a separate ssp.
(ssp. artemisia, see Papilio no. 1). Lycaena xanthoides editha and L. x. dione are the
proper combinations, not those used (see J. Res. Lepid. 18: 50). L. xanthoides is a
perching species, contrary to p. 228 (see J. Lepid. Soc. 29: 63). Lycaena heteronea
gravenotata is a veiy distinct ssp. (the book is correct in treating klotsi as a form). All
references to Lycaena dorcas in the book actually refer to L. helloides, based on
extensive foodplant and morphological data (see J. Res. Lepid. 17: 40). The
specimens of ''dorcas” illustrated are particularly similar to the helloides il-
lustrated. Rocky Mountain foodplants of "dorcas” are Polygonum and Rumex, not
Potentilla as stated on p. 233. Furthermore, Rocky Mountain "dorcas” oviposits on
trash at the plant base as does helloides, whereas true eastern bog dorcas oviposits
on the tops of shrubs (see Can. Ent. 41: 222). L. helloides hibernates as eggs.
Polygonum douglasii is a valid foodplant of Lycaena nivalis, but an error for L.
mariposa (this mixup is explained by E. Newcomer, see J. Res. Lepid. 2: 276).
Harkenclenus titus mopsus does not occur in the region. Quercus foodplant otH. titus
is an error by H. Tietz. Satyrium behrii has been in Satyrium since the work of
Clench in 1961 (in How to Know the Butterflies), not since 1979, Cercocarpus
montanus is not a valid Satyrium saepium foodplant. Eriogonum is an erroneous
foodplant of Satyrium califomica. Incisalia augustinus feeds on a variety of
foodplants in California, but not in the east. Mitoura spinetorum has one brood only
in the region (two broods occur in S. Arizona and New Mexico, see J. Res. Lep. 4:
233). The W. Colorado and SE Utah low altitude Callophrys populations actually
seem to be somewhat intermediate between C. sheridanii sheridanii and C.
sheridanii comstocki, the VHW line varying from straight, to kinked as in comstocki,
and this line varies from a complete row of spots to nearly absent. On p. 260, 2nd
line from bottom, “the region” shouldread “ Colorado. ’’Fixsemo is now used for the
genus Euristrymon (see J. Lepid. Soc. 32: 279).

Satyrinae. Xyris torta (Xyridaceae) is an erroneous foodplant of Megisto cymela,
another error by H. Tietz. M. cymela actually diapauses in the 4th instar (see J. Res.
Lep. 18: 171). Cyllopsis henshawi seems to be the May-June brood of C. pyracmon,
and C. nabokovi is the August-September brood; the same seasonal forms occur in
C. gemma. To prove this, eggs of the first brood should be reared. Cercyonis sthenele
from Salt Lake City actually resemble ssp. silvestris of California. Poa is only a lab
foodplant of sthenele. C. sthenele and meadii interbreed extensively on the Kaibab
Plateau just NW of the Grand Canyon, and actually ssp. damei is a ssp. of sthenele
(not pegala) that has hybridized with meadii (the reviewer’s 1980 research). Page
274 also mentions hybridization between the two in the Chuska Mountains
(intermediate populations). These two do not appear to be completely distinct
species, and may be allopatric subspecies, although the Kaibab hybridization does
not appear to be completely random, perhaps because sthenele flies up from the
Canyon and meadii prevails on the plateau. The dorsal stripes of Cercyonis oetus
larvae are the same as those of meadii (see the detailed descriptions in W. Edwards’
Butterflies of North America). Sedges are undocumented foodplants of C. pegala.
Neominois ridingsi is always single brooded (the second broods reported for Oeneis
uhleri, alberta and polixenes are also errors). Actually, N. ridingsi and 0. polixenes
are known to be biennial in some places (J. Res. Lep. 18: 171). 0. alberta doubtfully

20(1): 65-64, 1981(82)

63

Mbemates as a pupa (ibid.). 0. melksa beam is characterized by its darker smoky
black color, not the characters given. Arctic workers (K. Philip, C. dos Passes, pers.
comm.) have found no essential difference between Oeneis bore and taygete, so bore
should be the species’ name.

Nymphalinae. Boloria eunomia dawsoni occurs only farther north than Wyoming,
and only ''ursadentis” occurs in Montana (see the map). Ssp. ursadentis and laddi
are actually very similar to caelestis. The Polygonum bistorta foodplant listed for
eunomia almost certainly refers to Polygonum (Bistorta) instead. The Bistorta
vivipara foodplant listed for eunomia is a European host. Viola “papilionacea”
foodplant of Boloria selene is a misidentification of V. nephrophylla (M. Epstein,
pers, comm.), and papilionacea is not native to the region. Boloria bellona has only
one brood in the Rockies. Boloria frigga saga is the ssp. in Alberta, and it probably
occurs in N. Montana also. Boloria epithore borealis is a homonym of European B.
thore borealis. The Dryas food of Boloria alberta is based on field association and lab
oviposition only, although it is probably the field food. The new Boloria acrocnema
is believed by the current revisers of Boloria (and by the reviewer, who has a mss.
describing the complete life history and ecology) to be a subspecies of B. improba.
The Viola tricolor (cultivated pansy) food of Speyeria idalia is only used in the lab.
The neotype of Speyeria nokomis was caught by Mrs. Cockerell at Beulah, New
Mexico, and sold to E. Oslar, based on correspondence from T. D. A. Cockerell to F.
Benjamin of the Smithsonian. Therefore nigrocaerulea falls as a synonym of
nokomis. Theoretically only the endpoints of a dine should receive names, so only
the Calif omia-Nevada nokomis apacheana and the Arizona-New Mexico-southern
Colorado nokomis nokomis should receive names and the intervening material
should be left as dinal forms. Speyeria hydaspe conquista and Speyeria zerene were
supposedly collected by A. Klots from the same two localities in new Mexico but
have never been found since (M. Toliver mss. on the butterflies of New Mexico).
Both are undoubtedly mislabeled specimens from Wyoming where Klots also
collected. Neither species has been found south of northern Colorado. The
probability that both occurred at the same two locations, then both became extinct
at the same time in still-natural habitat, is infinitesimal. Speyeria zerene cynna is
now treated as a synonym of S. z. gunderi (see J. Res. Lepid. 19: 242). Speyeria egleis
Unda also occurs in western Montana and in the Stansbury Mountains of Utah. The
Viola canadensis food of Speyeria cybele is an association record only (by S. Ellis).
Euptoieta claudia may actually lack a true diapause. TTie Siphonoglossa and Ruellia
foodplants of Phyciodes texana are lab hosts only. Early stages of Phyciodes mylitta
and P. pallida were published (see J. Res. Lepid. 14: 84, which gave all the
differences that occur between these species and P. orseis). Phyciodes mylitta
callina is an available name for the SW Colorado-New Mexico- Arizona ssp, of
mylitta. The Helianthus scaberrimus foodplant of Chlosyne gorgone is a synon3rm of
H. laetiflorus. Rudbeckia laciniata is the only Rocky Mountain host for Chlosyne
nycteis, not the plants stated. Chlosyne damoetas is not strictly biennial, but rather
“irregular” in life cycle length, because the half grown larvae diapause for a variable
number of years, often two years but probably also 1-3 years or longer. Interestingly,
the larva and pupa of damoetas are identical to those of C. gabbii. Lowland
checkerspot larvae can diapause for several years (J. Res. Lepid. 18: 171), which
seems to preadapt them for life in the alpine zone, so the alpine damoetas may be
more closely related to lowland Chlosyne than current taxonomy suggests. Female

64

J. Res. Lepid.

damoetas are variable also in Colorado; Alberta populations are somewhat
intermediate to California populations. Chlosyne leanira alma occurs in west cental
Colorado (Mesa and Montrose Counties) and central Utah, and C. leanira fulvia
occurs in SW Colorado (Archuleta, La Plata and Montezuma Counties). The two
ssp. appear to intergrade in Kane County, Utah. The treatment of Poladryas is very
poor. P. minuta minuta (missing from the book) occurs on the plains of Colfax
County, New Mexico, where larvae have the typical red ground color. P. minuta
arachne has whiter larvae and occurs only in the mountains (several thousand feet
higher in altitude in Colfax County). These two subspecies have been hybridized
and backcrossed in nature by releasing lab-raised females in front of wild males (the
stocks from north (not west) Texas ssp. minuta and Colorado ssp. arachne) (see Pan
Pacific Ent. 50: 9, 1974 not 1973), proving that there are no barriers in either
courtship or development between them. This paper also mentioned series with
intermediate tendencies. Ssp. minuta is not extinct as stated, being widespread in
north Texas and E. New Mexico, although it is true that the most extreme
phenotype is in Mexico (which is more extreme than the Kerrville Texas types of
minuta). Recent studies published too late for the book place Euphydryas anicia as
a subspecies of E. chalcedona (J.'Res. Lepid. 17: 245). Page 331 supports this
conclusion, mentioning intergrades between chalcedona and “anicia” bemadetta in
N. Nevada and S. Idaho. The maps show that chalcedona wallacensis and “anicia”
overlap considerably in range, but S. Kohler has found that the anicia are all at
higher altitude, the chalcedona at lower altitude, and there are no known localities
where they are sympatric, although they come close at St. Ignatius in Lake County.
Ssp. bernadetta occurs locally in Madison County, Montana. E. editha gunnisonensk
and alebarki and E. Nevada lehmani in the opinion of the reviewer are synonyms of
hutchinsL It is the adults of Nymphalis califomica and not the pupae which
hibernate (J. Res. Lepid. 18: 171). Salix and Helianthus are erroneous foodplantsof
Nymphalis milberti. TUia is a very dubious foodplant of Polygonia interrogatioms.
Celds is eaten unwillingly by Polygonia comma (see W. Edwards, Butterflies of
North America), and the plants Althaea, Ambrosia and Amboris are errors of H.
Tietz and are not eaten by comma. Polygonia zephyrus is a subspecies of P. gracilis
(research of the reviewer); they have the same male genitalia, and are parapatric and
intergrade in the Canadian Rockies. Polygonia areas and P. oreas silenus have a
second different genitalic form. Ulmus is a dubious foodplant of Polygonia progne.
Ludwigia is an erroneous foodplant of Precis coenia. Nigrosuffusa is the Mexican
ssp. of Precis evarete according to J. Hafemik’s studies,'and at any rate it does not
occur in or near the region. Western Colorado records of Limenitis lorquini are
errors, contrary to p. 351 (see the map). Crataegus is another H. Hetz error, rather
than a valid lorquini foodplant.

Libytheidae. All Libytheana bachmardi in the region are probably migrants of ssp .
larvata.

The length of this review does not imply that this is not a good book. It is a good
book and contains more information than the average butterfly book. Its presenta-
tion of distribution information is excellent. Serious Lepidopterists should note the
points presented above, and in the opinion of the reviewer the generic names used in
W. Howe’s book {Butterflies of North America) should be used rather than the
genera used in this book.

James A. Scott, 60 Estes Sti'eet, Lakewood, Colorado 80226

INSTRUCTIONS TO AUTHORS

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Volume 20

Number 1

Spring, 1981(82)

m THIS ISSUE

Date of Publication: September 20, 1982

Editorial 1

Butterfly Nomenclature: A Critique

Paul R. Ehrlich & Dennis D. Murphy 1

Early Stages of Speyeria nokomis (Nymphalidae)

James A. Scott & Sterling 0. Mattoon 12

Supernumerary Chromosomes in the Domesticated Eri-
Silkmoth, Philosamia ricini (Satumidae: Lepidoptera)

K. B. Padhy & B. Nayak 16

Geographic Variation and Ecology of Hesperia leonardus -
(Hesperiidae)

James A. Scott & Ray E. Stanford 18

Notes on the Immature Biology of Two Myrmecophilous

Lycaenidae: Juditha molpe (Riodininae) and Panthiades

bitias (Lycaeninae)

Curtis J. Callaghan 36

Immature Stages of Odonna passiflorae Clarke (Lepidoptera:
Oecophoridae): Biology and Morphology

Patricia Chacon & Marta de Hernandez 43

A New Genus and two New Species of Oecophoridae from
Colombia (Lepidoptera)

J. F. Gates Clarke 46

Notes 50

Book Reviews 55

Cover Illustration; Speyeria nokomis pupa from Chihuahua State, Mexico. See
Scott & Mattoon article, page 13, this issue.

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Journal of Research on the Lepidoptera

20(2): 65-80, 1981(82)

International Nepal Himalaya Expedition for
Lepidoptera Palaearctica (INHELP) 1977, Report No.
1: Introduction and Lycaenidae

Oakley Shields^

4890 Old Highway, Mariposa, California 95338

Abstract. Seven species of steppe and alpine Lycaenidae were collected
during a 1977 summer expedition which visited 13 localities in the
Thakkhola region of Central Nepal Himalayas. Climatic and geologic
characteristics of the Thakkhola region plus habitat profiles of the collecting
localities are described. Alhulina orbitulus asiatica (Elwes), A. lehana
^oore), Polyommatus stoliczkana arene Fawcett, P. nepalensis Forster, and
Lycaena phlaeas baralacha Moore were residents of the Palaearctic realm,
while Lampides boeticus (L.) and .Rapa/a selira (Moore) barely enter this zone
from lower elevations. L. phlaeas and P. nepalensis extend down into the
Oriental realm as well. The butterfly fauna of the Tibetan frontier adjacent to
Sikkim and Bhutan to the east is very similar to the Thakkhola region, while
the high mountain fauna further west and northwest of Nepal is quite
dissimilar except for the Kumaon Himalayan-Tibet frontier. The Oriental
realm occupies the subtropical zone below Thakkhola to the south.

Introduction

During the summer of 1977, Mr. Hans J. Epstein, the expedition leader,
his sons Mark and Larry, and I explored steppe and alpine regions of
Thakkhola, a high valley of the Kali Gandaki River, and the adjacent
Manang and Sangda regions in Himalayan Central Nepal for Palaearctic
butterflies. From June 1st to August 7th we collected between 2745 and
5395 m, covering about 965 km of main and minor trails in all. We were
ably assisted by Sherpa guides (Danu, head Sherpa) and porters provided
by Lt. Col. James Roberts of Mountain Travel, Khatmandu. An historical
account of this rarely collected area, along with our papilionid and pierid
results, has recently been reported by Epstein (1979a, b). The present
report covers the Palaearctic Lycaenidae I collected on the expedition.
(Epsteins’ collections were for the same species and localities as reported
here.) Although two species of Riodinidae and six species of Hesperiidae
were encountered in the Oriental-subtropical zones during the expedition,
none were seen in the Palearctic zone (Shields, 1981). Climatic, geologic,
and biogeographic background information will be discussed.

‘Research Associate in Entomology, Los Angeles County Museum of Natural Histoiy

66

J. Res. Lepid.

Climatic Regime of Thakkhola

Thakkhola is the high valley (>3050 m) of the Kali Gandaki River in
Central Nepal, a glacial-fed river that originates north of the Himalayan
Ridge in the semi-arid region of the Mustang Kingdom. The high, open
basin cuts into the Tibetan Plateau and is bordered by two north-south
ridges, nearly 6100 m in elevation. A series of tributary valleys, extending
about 16 km, drains the ridges. Below this valley, the Kali Gandaki crosses
the High Ridge from Dhumu to Dana and Tatopani in a narrow S -shaped
gorge cut deeply between Dhaulagiri (8300 m) to the west and Nilgiri
(7530 m) to the east. It then descends 1980 m in about 16 km, with the
climate changing from Tibetan semi-arid (slopes north of the High Ridge)
to Nepali monsoon tropical (south slope). Thereafter, the valley opens and
the river flows southernly for about 50 km. Its tributaries drain all the
glaciated south slope of the High Ridge (above from Krummenacher,
1971).

During the monsoon season (July to September), the moist air which
reaches Nepal from the southwest is forced to rise when it meets the
mountains, resulting in heavy precipitation on the southern sides of the
Himalayas. Upper slopes around 2440 m are almost perpetually covered
with drizzling mist and clouds. When the monsoonal air passes over the
northern sides of the Himalayas, there is a pronounced rain-shadow effect.
Early October to mid-December is the post-Monsoon or retreating
southwest monsoon period. Winter lasts from December to March, when
heavy snowfall occurs in the higher Himalayas. Spring extends from April
to June (above from Banerji, 1952; Critchfield, 1966; Stainton, 1972).

A strong wind blasts through the gap between Daulagiri and Annapurna
and on up the upper Kali Gandaki valley for most of the day, starting
around 10 AM during the monsoon season. This wind clears the rainclouds
from the center of the valley, while the sides of the valley above 4420 m are
usually covered in mist.

In Central Asia, including the Himalayas, there were four glacial periods
corresponding to the Mindel, Riss, Wurm, and post-Wurm stages in the
Alps (TVinkler, 1930; Mani, 1968). Glaciation was extensive on the
northern side of the Himalaya during the Pleistocene, where huge
moraines encumbered all the northern valleys. In the Mt. Everest area,
these north-side glaciers completely blocked most of the valleys and often
were Imked (Trinkler, 1930). On the Tibetan Plateau, glaciers were
sporadic and small except in southeastern Tibet, and the highland was not
covered with a cap of ice (Trinkler, 1930). Today, p^etual snowline in
the Himalayas is 4875 m (5180 m in Nepal), and on tne Tibetan Plateau it
rises to 6100 m due to lack of precipitation. The lower limit of glaciers on
Annapurna (south cirque of Annapurna I) is 3600 m, while their upper limit
there is 5200 m (Vivian, 1970).

20(2): 65-80, 1981(82)

67

Geologic Setting of Thakkhola

Nine of the 14 world’s tallest mountains, >8000 m high in elevation,
occur in the Higher Himalayas of Nepal. Daulagiri (8172 m) is the sixth
highest peak in the world. Tibet is the highest of the world’s plateaus,
averaging 4875 m. The Tibetan slab is an immense monoclinal structure
dipping north. Based on indications from submarine volcanism and
paleomagnetism, Middle Cretaceous-Eocene was the time when the
Himalayan orogeny formed (Verma, 1973; Blow & Hamilton, 1975).

The Inner Himalayas, the region between the Great Himalayas and the
Tibetan Plateau, are composed of Lower Paleozoic to Middle Cretaceous
marine formations of the Tibetan Zone (Colchen, 1975). Towards the
north this zone grades into the Tibetan Plateau across the Nepalese
border. Formations of Thakkhola range in age from Precambrian to mid-
Cretaceous (Krummenacher, 1971).

Most great rivers of the Himalayas have their source in the Tibetan Zone
and cut across the Great Himalayan Range in deep gorges. Below Tukche,
the walls of the Kali Gandaki gorge rise from about 2440 m to over 7925 m,
making it perhaps the greatest canyon in the world.

A major N-S directed fault zone borders the west side of the Kali
Gandaki valley. This fault has a maximum vertical throw of >2700 m; it
sharply cuts all existing structures and is therefore post-orogenic.

Biogeographical AMnities

Elwes et al. (1906) describe the butterfly fauna from the Tibetan Plateau
just north of Sikkim and Bhutan (mostly from Gyantze) which appears
quite similar to that of the Thakkhola region. Indeed, both areas fall within
Ward’s (1935) “Outer Plateau” gravel lands botanical division of south-
central Tibet. The Chumbi Valley on the Tibetan frontier of Sikkim
(Elwes, 1882), about 480 km east of Thakkhola, exhibits a similar fauna to
the Thakkhola region. Butterflies from the Kumaon-Tibet border NW of
Nepal show close affinities to the Nepal Himalayas (see Champion & Riley,
1926). Other early papers on the Tibetan butterfly fauna include Fawcett
(1904), South (1913), and Evans (1915).

In contrast (except for the adjacent Kumaon Himalaya), the butterfly
faunas of the high mountains further to the northwest and west of Nepal,
though having some species in common with Thakkhola, are fundamental-
ly dissimilar, i.e. the north-west Himalaya (Man! & Singh, 1961-1962),
western Karakorum (Evans, 1927), Alai-Pamir (Forster & Rosen, 1940),
Baluchistan (Evans, 1932), Afghanistan (Clench & Shoumatoff, 1956;
Wyatt, 1961), and West Pakistan and Iran (Shirozu & Saigusa, 1963).
Most species of Lepidoptera from Northwest Himalaya are also found in
the Pamirs and other Middle Asiatic mountains (Mani, 1968, p. 2 20). This
biogeographic division is termed the Turkmenian subregion of the
Palaearctic realm and extends over Transcaspia, Turkestan, the Higher

68

J. Res. Lepid.

Fig. 1. Butterfly faunal provinces of Southeast Asia. Arrows indicate the initial
spread from an Upper Burma-Yunnan-Szechwan center of origin.

Himalayas along the south slopes (above treeline), Tibet, Sinkiang,
Mongolia, and southernmost West Siberia (Mani, 1962, 1974). However,
the Mongolian butterfly fauna differs markedly from the central Himalayas
(see Elwes, 1899; Forster, 1965). Mani (1968, p. 228) sets off Northwest
Himalaya, Karakorum, and the Alai-Pamirs as a distinct biogeographic
subunit of the Turkmenian. The bulk of the Indian butterfly fauna proper
is derived from the Oriental tropics (Holloway, 1969, 1974) (see Figure 1).

The southern boundaiy of the Palearctic extends to the southern slopes
of the Himalayas, between 2440-3660 m in the N.W. Ifimalayas, gradually
rising to 3660-4575 m in the east (Seitz, 1923; Riley, 1927). Tree-line or
slightly below tree -line is probably a more realistic boundary in the
Thakkhola region.

The alpine flora of the Himalaya is closely related to the alpine flora of
eastern Tibet and western China (Kitamura, 1955). In Nepal, the East
Himalayan floral elements merge with the West Himalayan elements
(Baneiji, 1962, 1973), at least in the Oriental realm (Bhatt, 1964). The
TTiakkhola-Manang regions lie in the Eastern Himalaya in Baneiji *s
scheme but close to this 83° Longitude merging line. We found some basic
differences in various butterfly subspecies on the east and west sides of
the Kali Gandaki in the Thakkhola region, also in support of this boundary.

The highest known angiosperms collected in Nepal are from just below
6000 m (Webster, 1961).

20(2): 65-80, 1981(82)

69

Lycaenid Localities (Table 1 and Figure 2)

1. Just north of Jhomosom, 2800 m (Camp 1)

Sophora moorcraftiana Benth. ex Baker var. nepalensis, a pioneer on
the steep slopes and new soil along the Kali Gandaki, is the dominant
steppe shrub. Also prominent here was Ephedra gerardiana Wall, ex
Stapf. This was a very dry, windy valley, like a desert. Jhomosom averages
only 270-296 mm rainfall.

2. Between Jhomosom and just south of Kagheni to Dangarjong, 2760-
3140 m (Camp 2 at Dangarjong) (Figure 3)

The trail between Jhomosom and Kagbeni passes along present-day
alluvium. S. moorcraftiana and E. gerardiana occur along the Kali
Gandaki, and Juniperus wallkiana Hook. f. et Thoms, ex Pari, in DC grows
sparsely on the hillsides. Thymus linearis Benth. was abundant south of
Dangaijong, but we found no lycaenids on it. The Jhomosom, Kagbeni and
Dangaijong villages themselves are under cultivation.

3. Pass region 8 km NNW Dangarjong, 4000-4400 m (Camp 3)

We extensively collected in the basin area mth rolling hills 1% km
west of this pass. Alpine meadows are found above treeline, above a zone
of Betula utilis D. Don and conifer forest in Sangda Valley. Plants here
were grasses, Lonicera spinosa Jacq., Sedum, Polygonum, small purple
Aster, and many other alpine flowers. The area has been extensively
grazed by goats and yaks but was a rich collecting site nevertheless.

4. IV2-3 km SE Sangda, 4100-4300 m (Camp 5)

We collected in a broad canyon below a limestone ridge and on grassy
ridge slopes, south above Sangda Valley. Alpine flowers included pink and
white Polygonum, yellow Potentilla bushes, white edelweiss, and white

Table 1, Species Collected vs. Localities

SPECIES LOCALITIES

1 2 3 4 6 6 7 8 9 10 11 12 13

Albulma orbitulus asiatica

Albulma lehana

Polyommatus stoliezkana arene
Polyommatus nepalensis
Lampides boeticus
Lycaena pUaeas baralacha
Rapala selira

X X X X

X XXX

X X

XXX XX

X X

X X X X X

X

X

X

70

J. Res. Lepid.

Fig. 2. Map showing our 13 collecting localities in the Thakkhola, Sangda, and
Manang regions. Glaciers are stippled.

>

Fig. 3. Kali Gandaki River Valley with Annapurna in background.

20(2): 65-80, 1981(82)

71

Saxifraga, Upper talus slopes were nearly devoid of vegetation except for
Saxifraga. The area was ve^ windy at times and heavily grimed by goats,
yaks and horses.

5. 1% km SW Sangda, 3600 m (Camp 4)

Ubetan steppe vegetation of Caraganagerardiana Royle mdArtenmia
grows here on shale.

6, Just below sanctuary of Muktinath, 3600-3700 m (Camp 6)

Tibetan steppe vegetation near cultivation was characterised by

Ranunculus, Potentilla, Geranium, Stellera, Primula, Thymus, Populus
ciliata Wall, ex Royle and thickets of Spiraea and Lonicera.

Fig. 4. Marsyandi River Valley of Manang.

7. 5-10% km SE Thorong Pass, 4200-4900 m (Camps 8 and 9)

We collected mostly in Jargeng Valley, which consists of limestone.
Lush habitat occurred in places along the river, with some alpine meadows
and bare rockslides above timberline. Alpine plants included grasses,
sedges Gentiana, Primula, Pedicularis, Campanula, Lactura, Brassica,
Convolvulus, Rumex nepalensis Spreng., Heracleum, Iris, Ribes, Oxytropis.
Thorong Pass is at 5346 m.

8. Manang ( 3500 m) to valley and slopes 5 km ESE Manang (3370-3700
m) (Camp 11) (Figure 4)

The valley of the upper Marsyandi River is covered by an open
xerophile forest of pines (Pinus griffithi McClelland in Griffith) and
junipers (Junipems squamata D, Don,e/. communis L., J. wallkhiana). On

72

J. Res. Lepid.

Annapurna III above Braga, this open forest creeps up to 4100 m where it
changes into a dense Abies forest, with alpine vegetation above the Abies
forest. The extensively cultivated valley consists of alluvial deposits and
alluvial fans. Other plants in the region include Ranunculus hyperboreus
Rottb., Populus suaveolens, Lonicera myrtillus Hook. f. et Thoms., I^rus,
Vibernum, Oxytropis williamsi Vass., Crataegus, Potentilla and Rosa
sericea. Vegetation of the Marsyandi basin is lusher than the arid KaU
Gandaki basin. In the Marsyandi, the winds are not nearly as strong, and
rainfall is much more frequent during the summer monsoon than in the
Kali Gandaki.

9. 6V2 km W. Khangsar, upper end Khangsar Valley, 4500 m (Camp 14)
We camped here for 6 days in early July, in rolling alpine pastures
on limestone filled with flowers and grasses above timberline. Flowers
included a small purple Aster, legumes, purple Viola, Sedum, Rumex,

Rheum spiciforme Royle, Ranunculus, strawberry, and pink Polygonum.
Usually there was very little sun here in early July.

10. 8 km W. Khangsar, 4000 m (Camp 15)

We camped where three rivers converge. Many different alpine plants
grew here on limestone, including Heracleum, Polygonum, Betula, bush
Lonicera, yeUow sunflower, grasses, purple Azalea, orchid, purple Aster,
chives and Berberis.

11. SE slopes above Tilicho Lake, 4860 m (Camp 16) (Figure 5)

This isolated lake is located on the N. side of Nilgiri and is sur-
rounded on the south by glaciers. Sparse alpine vegetation of Sedum, pink
Polygonum, purple Aster, purple Gentiana and some grasses was noted on
limestone.

12. ''High Camp", NW below Tilicho Pass, 4420 m (Camp 1 7)

This camp had alpine meadows and dells filled with pink Polygonum
and purple Aster. We followed the North Thini Valley between here and
Jhomosom.

13. 5~5V2 air km W. Marpha, 3850 m (Camp 22)

We camped and collected at treeline at the NW comer of old, man-
made terraced fields surrounded by grassy-juniper slopes. A complex
assortment of alpine and temperate forest plants grew here: Juniperus,
Sedum, Thymus linearis, Nepeta leucophylla Benth., white Heracleum,
orange cmcifer, small and bush purple Asters, two yellow composites,
Artemisia, yellow Trifolium, pink scroph, grasses, Salix, red thistle, various
field weeds, Rosa, conifer and hsudwood groves and beech. This locality
was rich in butterfly species and flowers with recent glaciations.

Along the Kali Gandaki between Marpha and Lete, there is a Pinus
excelsa Wall, ex D. Don forest, then 4 km S. of Ghasa starts a broadleaved
trees-pine forest, and then the humid, subtropical monsoon forest (Figure
6). Thus in the apace of 24 air km S. of Jhomosom to the entrance of the
Dhauagiri- Annapurna gap, only about a 915 m drop in elevation, one

20(2): 65-80, 1981(82)

73

Fig. 5. The Tilicho Lake locality.

Fig, 6. Subtropical monsoon forest, 24-32 km SW of Marpha.

74

J. Res. Lepid.

passes abruptly from Tibetan steppe (Palaearctic realm) into the sub-
tropics (Oriental realm). The transition is much less marked on the valley
sides. The butterflies we encountered in the subtropical forest habitat and
hardwood trees-pine forest habitat from Kalopani (2440 m) southeast to
Pokhara were endemic to the Oriental realm (Shields, 1981). Rapala selira
(Moore) appears to be a resident just within the Palaearctic realm, while
Lycaena phlaeas (L.) and Polyommatus nepalensis Forster extend down
into the Oriental realm.

One additional, Oriental species, Celastrina dilectus (Moore) was taken
as a fair condition male 4 miles west of Khangsar, 4500 m, July 9. This is
doubtless a stray from lower elevations where it is known from the
Himalayan foothills from Simla to Nepal.

Lycaenidae Collected

1. ) Albulina orbitulus asiatica (Elwes), 69cfcf 1999; localities 3, 4, 5,
6, 12, 13; flight in mid June, July.

Males sometimes formed mud puddle clubs of 4-6 individuals. We
found a light green pupa under a rock on June 15th that emerged on July
4th, 8-9^2 km NNW Dangaijong. Elevation range: 3600-4420 m.

A. orbitulus is found in the Pyrenees and the European Alps at high
altitudes, southern mountains of Norway, Lapland, Scandanavia, Asia
Minor, the Balkan, Persia, Altai Mts., and Himalaya Mts. to Tibet, north-
central Mongolia, Turkestan, China, and south Siberia as far as Kamt-
chatka as subspecies or possibly closely related species (Elwes, 1899;
Seitz, 1906; Forster, 1965; Higgins, 1975). It is also found in the Amur
region (Mani, 1968), A. pheretes Hubn, is a synonym (see Beuret, 1933)
and occurs in Tarbagatai, Central Asia. A. o. asiatica (Elwes) is found at
high elevations in the northeast comer of Nepal (Riley, 1923; Fujioka,
1970) and Sikkim (Mani, 1968). Elwes et al. (1906) consider ssp. pharis
Fawcett to be a synonym of ssp. asiatica and record it from Tungu,
Khamba Jong, and Lhanak Valley, Tibet. A. o.pharis (TL “ Khamba Jong,
4575 m, Tibet, Fawcett, 1904) is recorded from Phari, 4875 m, and Tinki
La, 4725 m (Riley, 1927). Unspecified subspecies of A. orbitulus are found
at Girthi Valley, Shibchilan, Chojan and Lai Pahar (Champion & Riley,
1926), Po Chu Valley and Gyala (Evans, 1915), and Tiong la, Zhasha-la,
Drowa Gompa, Pugo and Di Chu (South, 1913), all in Tibet. Forster
described ssp. lobbichleri from only two worn specimens from Thakkhola;
fresh specimens have the greenish gloss to the underside characteristic of
asiatica, so lobbichleri is likely a synonym.

2. ) Albulina lehana (Moore), IGcfcf 399 plus 23 others; localities 7, 9,
10, 11; flight in late June to mid July,

Adults flew in meadows, came to moisture, and three were found 10y2
km SSE Thorong Pass on a large purple Aster sp. in meadows in the early
morning before it rained. Elevation range: 4000-4860 m.

20(2): 65-80, 1981(82)

75

A. lehana was described from Leh, 3515 m, Ladak (Moore, 1878, p.

230) and is found in Chitral-Kumaon over 3660 m (Cantlie, 1963),
Kashmir (Mani & Singh, 1962) and the Pamirs and Kighizia (Korshunov,
1972). Elwes et al. (1906) claim that the West-Tibetan form is lehana,
Lowndes (1953) records it {''m.pheretes'') from Manangbot, 3810 to 4875
m, Nepal, July. As orbitulus and lehana occur allopatrically in the
Thakkhola region, they are probably separate species.

3. ) Polyommatus stoliczkana nr. arene Fawcett, 53d'd' 2299; locali-
ties 8, 10; flight in late June to mid July, early August.

40 adults were taken in late June, all flying around Oxytropis williamsi
Vass. (det. by A. O. Chater, BMNH) with rose-purple flowers and light,
hairy leaves. Two females oviposited on the leaf venter at 8:30 and 10:30
AM. One in copulo pair (male carried female) was taken at 9:30 AM.
Specimens were fresh to worn and did not visit mud; most alighted on the
flowers and leaves of the foodplant. Oxytropis grew mostly on old stream
alluvium, i.e. on flats. At 5 km ESE Manang, a dozen adults were taken in
close association with 0. williamsi growing on flats near pines, some
feeding on the flower-heads.

We observed this species generally between Manang and Khangsar.
One male was taken 16-19 km SW Marpha (est. 2560 m) in a Thyme-
legume area of a dry river bed. Elevation range: 2560-4000 m.

This is the ''Eumedonia chiron jermynV of Forster (1961). Nominate
jermyni is found from Chitral to Gilgit (Cantlie, 1963). P. stoliczkana was
originally named from Ladak. The species is quite variable in size and
undersurface markings. According to Mani (1968), it extends from the
Northwest Himalaya to the Sikkim-Himalaya, and is represented by the
ssp. hunza Gr.-Gr. on the southeast Pamir and in the Great Pamir, at
elevations of 3870-4725 m. It is also found in the Western Karakorum
(Evans, 1927). P. s. janetae Evans occurs in Karakorum, Khupjerah,
Gujerab and Baturu (Wu, 1938), and ssp.ariana Moore is found in North
India and southern Kashmir (Seitz, 1906). P. stoliczkana has been taken at
Sanga Chu Dzong, southeastern Tibet, 3660 m, in June (South, 1913).
Riley (1923) reports it common between June 18 and August 12 from
3960-4570 m on Mt. Everest. Elwes et al. (1906) give Gyantze and
Khamba Jong, Tibet, as localities. Evans (1915) records it from Po Chu
Valley and Tsang Po, Tibet, from July to September, In Nepal, Lowndes
(1953) says it was common in grassy places and among junipers in
Manangbot, 3500-4570 m, in July. Shirozu (1955) records it from
Annapurna Base Camp to Chame in late May and early October, above
2500 m. The Nepal material we collected agrees with the description and
figure of ssp. arene Fawcett (TL ™ Khamba-Jong, 4570 m, Tibet)
(Fawcett, 1904; Seitz, 1906).

4. ) Polyommatus nepalensis Forster, common (no exact tally); localities
1, 2, 3, 5, 6, 13; flight in early to mid June, late July to mid August.

76

J. Res. Lepid.

Between Jhomosom and 1% km S. Kagbeni, many were taken in early
June, always around the blue-flowering Sophora moorcraftiana var.
mpalemis bushes. One in copulo pair was on S. moorcraftiana, male
carried female, at 8:00 AM. Just below the sanctuary of Muktinah (3600
m), one female displayed preoviposition behavior on a young, flowerless
Lonicera spinosa bush (no Sophora here). At 5V2 km W. Marpha (3700-
3850 m), adults were in association with Thymus linearis Benth. and
Nepeta leucophylla Benth. (both det. by J, R. Press, BMNH), in a Rosa-
Juniperus- Artemisia area. Sometimes freshly emerged males were found
at mud. We collected a few P. nepalensis in the hardwood-pine zone
between Kalopani and Lethe, and between Kalopani (2440 m) and Ghasa
(2010 m), in August. Elevation range: 2010-3900 m.

Shirozu’s (1955) Polyommatus eros ariana Moore from Tukucha-
Jhomosom and nr. Muktinath is probably this species (he doubted the
original identification). Wadhi and Parshad’s (1968) P. icarus fugitiva
Butl. is also suspect, from Tukcha, Marpha and Dana in April. Forster
(1961) saysP. nepalensis stands near the species everesti Riley (1923) and
may be only a subspecies of it.

5. ) Lampides boeticus (L.), 3^c?; localities 1, 6; flight in June, late
July.

This species is strongly migratory in the west, a resident of S. Europe
and N. Africa, Ceylon, India, Pakistan, Nepal, SE Tibet, Szechuan,
Andamans, Nicobars, Burma, Australia, Tasmania, Lord Howe Island,
etc. It was scarce in the Thakkhola region. There are no recognizable sub-
species within its vast range. It is known to use many legumes as
foodplants.

6. ) Lycaena phlaeas baralacha Moore, 62^cf 2299; localities 2, 6, 7,
8, 9, 13; flight in mid June to mid August.

Adults were sometimes found along streams. Two males QV2 km SE
Thorong Pass, at a bridge, were very territorial. Adults were also taken in
the hardwood-pine zone ca. 24-32 km SW Marpha (2550 m), between
Kalopani and Lethe (2530 m), and between Kalopani (2440 m) and Ghasa
(2010 m). Elevation range: 2010-4500 m.

According to Ford (1923), baralacha (TL ^ Baralacha Pass, 4875 m,
Ladak) is perhaps a race of stygianus Butler. Distinguishing features
between these two are slight and depend upon the amount of FW brown
suffusion (more intense in baralacha than in stygianus). L. p. baralacha
occurs in the Outer Himalayas (Kashmir-Kumaon), and Nepal, where it is
common (in Evans as phlaeas indicus) (Cantlie, 1963). L. p. stygianus
inhabits Baluchistan to Chitral and Ladak (= eleus F.; timeus Cr.) (Cantlie,
1963). L. p. ftavens Ford is found in the Interior Himalayas to Sikkim and
SE Tibet; it is large, with scanty brown suffusion (Cantlie, 1963). Wadhi
and Parshad (1968) report L. p. baralacha indicus) from Lethe,
Tukcha, and nr. Muktinath, Nepal. H. J. Epstein {in litt.) compared the

20(2): 65-80, 1981(82)

77

Thakkhola material with the British Museum collection and concludes it is
best placed as baralacha.

7.) Rapala selira (Moore), 9 specimens, cal. 1% km N. Dangarjong,
3140 m, 3 June 1977.

Mostly fresh specimens were collected in a thicket at the outskirts
of cultivation. This locality is Quaternary alluvia (loess and broken rock).

Nominate selira was described from Kashmir (Moore, 1874). It is
found in the Western Himalayas, Chitral to Kumaon, Nepal, and Tibet,
and is common (~ roana Fruh. In Evans it is listed as micans selira, but
micans is now regarded as a Chinese ssp. of nissa) (Seitz, 1906; Cantlie,
1963). It flies in April-June, between 1220-2745 m (W5mter-Bl3rth, 1957).
The Dangaijong locality may be a new altitude record (3140 m). Wadhi
and Parshad (1968) found this species at Marpha and Jhomosom in late
April and early May. The foodplant is Indigofera purpurea (Leguminosae)
(Sevastopulo, 1973).

Acknowledgments: I thank Dr. Walter Forster (Zoologischen Staatssammlung
Munchen, West Germany) for determining many of the lycaenids. Dr. R. H. T.
Mattoni kindly gave financial assistance and encouragement (specimens housed in
his collection). Hans J. Epstein provided invaluable assistance both in the field and
through correspondence. Dr. Richard A. Arnold’s most useful criticisms of an early
draft are incorporated. Some plant identifications were arranged through the
mediation of Oleg Polunin (British Museum of Natural History) and Hans J.
Epstein. The Sherpas helped collect for us at most sites.

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Journal of Research on the Lepidoptera

20(2): 81-85, 1981(82)

Intergradation between Callophrys dumetorum oregonensis
and Callophrys dumetorum affinis in Northwestern U. S.
(Lycaenidae)

James A. Scott and John A. Justice

60 Estes Street, Lakewood, Colorado 80226 and

1700 Seventh Street NW, Minot, North Dakota 58701

Abstract. Populations of Callophrys dumetorum from the Washington
state area are analyzed using seven wing characters. Samples from the Blue
Mountains of Columbia County Washington and Wallowa County Oregon
are almost exactly intermediate between C. d. oregonensis and C. d. affinis
(new combination). C. d. washingtonia (new combination) from the type
locality in Okanogan County in northern Washington is also intermediate,
but closest to affinis. Eastern Washington specimens from Spokane and
Lincoln Counties are intermediate in several traits but are mostly referable
to C. d. affinis. These populations and C. d. oregonensis seem to be stages in
the intergradation of C. d. dumetorum with C. d. affinis.

Introduction

The status of names within Callophrys ( Callophrys) has been problemat-
ical. Tilden (1963) and Clench (1963) made careful studies of Callophrys,
but they lacked significant samples of dumetorum (Bdv.) and affinis (Edw.)
from the northwest. Scott (1975a), based on small samples collected by
Jon Shepard, predicted that C. dumetorum oregonensis Gorelick and C. d.
affinis might be found to intergrade in Washington and that C. d.
washingtonia Clench is an intergrade population. Adequate samples from
Washington have now been obtained by John Justice from four areas of the
state. Analysis of these series show a pattern of intergradation between
oregonensis and affinis.

Methods

Individuals were studied from southwestern, northern and eastern
Washington, from the Blue Mountains of SE Washington and NE Oregon,
and (for C. d. affinis) Montana, Wyoming, Nevada, Utah and Colorado.
Exact localities and numbers are given in the Appendix. Callophrys
sheridanii (Carpenter) was caught at most of the localities, but we carefully
removed them from the samples before study.

The genitalia do not differ among Callophrys (Callophrys) species. The
following wing pattern characters were studied because they are the only
characters that differ among the samples: 1) dorsal color (gray to
completely fulvous); 2) number of cells with white spots on ventral

82

J. Res. Lepid.

hindwing (from 0 to 8); 3) shade of green on ventral hindwing (including
several shades of green, bluish green, and yellowish green); 4) ventral
hindwing fringe (the fringe base varies from brown to nearly white); 5)
number of cells with dark (gray to tan) color on anal margin and disc of
ventral fore wing (from 2 to 4 cells) (the rest of the wing is some shade of
green); 6) the shade of color (varying from gray to tan) at the most anterior
extension of the non-green part of the ventral forewing; 7) color of the
scales on the costal margin of the ventral forewing (from cream colored to
light brown). The character states and cross references to a color manual
are given in the explanation for Figure 1. If a specimen had some scale loss,
it was compared to the reference specimen for each character state with a
microscope to compare the intact scale color; if scale loss was extensive the
specimen was ignored. Several other characters mentioned by Tilden
(1963) that are useful for distinguishing other Callophrys (sheridanii,
viridis (Edw.), etc.) were found to not vary among (and therefore are not
useful for distinguishing) the samples studied in this paper: shape of fore
and hindwing, color of the frons, number of white rings on the antennae,
and antennal color.

Results

Population phenotypes. Results are shown by histograms (Figure 1).
C. d. oregonensis from Klickitat and Yakima Counties Washington (type
locality Kusshi Creek, Yakima Co. Washington) and C, d. affinis from
Montana south to Colorado and Nevada (type locality vicinity of Fort
Bridger, Wyoming, Brown & Opler, 1970) represent extreme populations
in most traits, so the other samples will be compared to them. The eastern
Washington (Spokane and Lincoln Counties) sample is intermediate
between oregonensis and affinis in dorsal color and slightly intermediate in
ventral hindwing spotting, it is like affinis in the ventral hindwing color and
fringe, and is more extreme than affinis in the three ventral forewing
characters; it can be treated as C. d. affinis. The Blue Mountains sample
(Columbia Co. Washington and Wallowa Co. Oregon), however, is
intermediate between oregonensis and affinis in ventral hindwing fringe, in
ventral forewing costa color, and somewhat intermediate in ventral
forewing dark color, it is like oregonensis in dorsal color and ventral
hindwing color, and it is like affinis in ventral hindwing spotting and
number of ventral forewing dark cells. The Blue Mountains population
therefore appears intermediate between oregonensis and affinis, and
cannot be assigned an available name. The sample from Alta Lake in
northern Washington (the type locality of washingtonia Clench) is
intermediate between oregonensis and affinis in dorsal color, in ventral
hindwing spotting, and in ventral hindwing fringe, it is somewhat
intermediate but closest to affinis in the ventral forewing dark color, and it
is like affinis in ventral hindwing color, number of dark ventral forewing
cells, and ventral forewing costa color. The Alta Lake sample therefore is

20(2): 81=85, 1982(82)

83

20-

16-

12-

8-

4-

O'

c? 9

DUMETORUM AFFINIS

DUMETORUM
OREGONENSIS

12 345 0 I 2345678 SHADE OF

dorsal NUMBER OF GREEN VHW

COLOR CELLS WITH

WHITE SPOTS
VENTRAL HIND
WING

FRINGE NUMBER DARK
VHW OF DARK COLOR
CELLS VFW
VFW

I 23
COSTA
COLOR
VFW

Fig. 1, Histograms showing the number of individuals of each sex from each
sample which have each character trait. Colors are from the Color Harmony
Manual 1958 (chip number and formal name of color given in parentheses).
Character states are 1) dorsal color (1-gray; 2-slightly orange; 3-half
orange; 4-mostly orange; 5-completely orange except the margins (chip #5
la, orange)); 2) number of cells with white spots on ventral hindwing (from
0 to 8 cells have white spots); 3)shade of green on ventral hindwing (G1
green (chip #24 1 c, parrot green); G2-dark green (chip #24 ne, pea green);
B1 -slightly bluish green (chip #23 ia, light paris green); B2-bluish green
(chip #22 ia, brite mint green); Yl-slightly yellowish green (chip #24 ia,
light lime green); Y2-yellowish green (chip #24y2 ia, brite chartreuse)); 4)
fringe of ventral hindwing (1-very light, nearly white; 2-dark band at base of
fringe thin, tan in color; 3-dark band thicker, browner; 4-dark band thick
and brown); 5) number of cells in anal angle and disc of ventral forewing
which have dark non -green color (varying from gray to orangish) (from 2 to 4
cells have dark color); 6) color of this dark area on ventral fore wing (1-gray
(chip #5 ca, pale peach); 2-very slightly tan (between chips #5 ca& 5 ea); 3-
slightly tan (chip #5 ea, peach pink)) — (ssp. dumetorum not treated in this
paper also varies to chip #5 ga, peach, and chip #5 nc, burnt orange); 7)
color of the costal margin on ventral forewing (1-cream; 2-ochre; 3-brown).

84

J. Res. Lepid.

intermediate as well but appears closer to affinis than to oregonensis. A few
individuals from Alta Lake are bluish green ventrally similar to individuals
of virkiis from coastal California.

Behavior. At Alta Lake C. sheridanii flies about three weeks earlier than
C. dumetorum washingtonia. C. sheridanii is found mostly in gullies and
hillsides at Alta Lake, whereas C. dumetorum washingtonia males perch on
prominent shrubs on hilltops, where they evidently wait for females; two
copulating pairs of washingtonia were found on the hilltops. At Alta Lake
female washingtonia were found to be rather generally distributed. West of
Davenport C. dumetorum nr. affinis perched on low (1-2 m tall) mounds in
fairly level sagebrush habitat. In the Blue Mountains of Oregon C.
dumetorum occurred on a mountain ridge with sagebrush and Eriogonum.
C. d. affinis males commonly perch on sagebrush shrubs on hilltops.

Discussion

In this paper we treat affinis as a subspecies of dumetorum because the
two are allopatric entities that seem to be connected by various inter-
mediate populations, thus qualifying as subspecies. The biology and adult
behavior of both are similar as well (Scott, 1975a). C. d. washingtonia is
one of the intergrade populations, but it is most similar to affinis. The Blue
Mountains sample is almost exactly intermediate between oregonensis and
affinis, although two characters are closest to oregonensis and two others
are closest to affinis. The simplest explanation of these findings is that
these populations are not reproductively isolated and represent sub-
species rather than species. We have studied only intergradation between
C. d. oregonensis and C. d. affinis. C. d. oregonensis is an intermediate of
sorts itself, however, since it too is intermediate between C. d. dumetorum
and C. d. affinis in several characters including dorsal color, shade of green
on ventral hindwing, number of dark cells on ventral fore wing, and the
shade of gray to brown on ventral forewing (Gorelick, 1968; Scott, 1975a).

A population tentatively assignable to C. d. dumetorum, because it
appears identical to California dumetorum, occurs in the Puget Sound area
of Washington (we examined 3 from Belfair, 200', Mason Co. Washington,
12 May 1970, coll. Jon Pelham).

To be complete we should mention that C. d. affinis appears to blend
with ‘"apama” (Edw.) homoperplexa Barnes & Benj., so that homoperplexa
and apama are almost certainly subspecies of dumetorum also. C.
dumetorum homoperplexa is nearly identical to California C. dumetorum in
wing pattern {homoperplexa was not named until 1923, over fifty years
after its discovery, because lepidopterists labeled it dumetorum), which
suggests that homoperplexa and apama are subspecies of dumetorum,
because most Lepidoptera species are based on morphological similarity.
Another point of similarity is that dumetorum and homoperplexa are the
only Callophrys (Callophrys) that are polyphagous: dumetorum feeds on
Eriogonum (Polygonaceae) and Lotus (Leguminosae), homoperplexa on

20(2): 81^85, 1981(82)

85

Eriogonum and Ceanothus (Rhamnaceae) (many ovipositions seen in 1980
by J. Scott). Furthermore, intergrade populations between homoperplexa
and affinis seem to occur in south central Wyoming (specimens in the
American Museum), in the Gunnison-Delta-Mesa County area of western
Colorado, and in southern Utah. Larger samples are needed from these
areas to better document the intergradation. C. d. washingtonia, C. d.
affinis and some C, d. dumetomm populations tend to mate on hilltops,
whereas some C. d. dumetomm populations (especially in southern
California) and C. dumetomm homoperplexa tend to mate in gully bottoms
(Scott, 1975b).

literature Cited

BROWN, F. M. & P. A. OPLER, 1970. The types of the Lycaenid butterflies described by
William Henry Edwards. Part H— Theclinae and Strymoninae. Trans. Amer.
Ent. Soc. 96: 19-77.

CLENCH, H. K., 1944. Notes on Lycaenid butterflies: A. the genus Callophrys in North
America. Bull. Mus. Comp. ZooL 94: 217-299.

, 1963. Callophrys (Lycaenidae) from the Pacific Northwest. J. Res.Lepid.

2: 151-160.

COLOR HARMONY MANUAL, 1958. 949 different color chips in removable form ar-
ranged and notated according to the original system of Wilhelm Ostwald, 4th ed.
1958. Container Corp. of America, 38 So. Dearborn St., Chicago, Illinois.
GORELICK, G. A., 1968. A new subspecies of Callophrys ( Callophrys) dumetorum from
Washington and Oregon. J. Res, Lepid. 7: 99-104.

, 1971. A biosystematic study of two species of Callophrys (Callophrys)

m California (Lycaenidae). J. Lepid. Soc. vol. 25, suppl. 2, pp. 1-41.

SCOTT, J. A., 1975a. Subgenus Callophrys In: W. H. Howe, ed. The butterflies of
North America. Doubleday & Co., New York. 633 pp.

, 1975b. Mate-locating behavior of western North American butterflies.

J. Res. Lepid. 14: 1-40.

TBLDEN, J. w., 1963. An analysis of the North American species of the genus Cal-
lophrys. J. Res. Lepid. 1: 281-300.

Appendix; Localities Studied

C. d. oregonensis individuals were from the type locality and vicinity in southern Washington:
SatusPass, Klickitat Co., 17 May 1970, 17 m, 18 f; Mill Creek, Yakima Co., 17 April 1970, 2 m, 2
f;Kusshi Creek, Yakima Co., 17 Mayl970, 1 f; Fort Simcoe, Yakima Co., 17 May 1970, 1 f(allJ.
A Justice). C. d washingtonia individuals were from the type locality in northern Washington:
Alta Lake, Okanogan Co., 2 May 1970, 1 f , 6 May 1970, 4 m, 1 f, 7 May 1970, 6 m, 1 f, 23 May
1970, 5 m, 1 f, 11 June 1967, 5 m, 3 f (all J. A. Justice). A third sample was from eastern
Washington: 4 mi. NW Fairchild Air Force Base, Spokane Co., 31 May 1970, 3 m, 1 f; 6.4 mi. N.
Davenport, Lincoln Co., 3 1 May 1970, 2 1 m, 2 f (all J. A. Justice). A fourth sample was from the
Blue Mountains of southeastern Washington and adjacent northeastern Oregon: Trail 2138
fixjm Godman Springs, Columbia Co. Washington, 8 July 1970, 2 m; Skyline Road S. of Godman
Springs, Columbia Co. Washington, 8 July 1970, 2 m; 1-5 mi. N. of Bear Canyon Campground,
Wallowa Co. Oregon, 9 July 1970, 3 m, 1 f; Forest Service Road N-50, 15 mi. SW Troy, Wallowa
Co. Oregon, 21 June 1970, 12 m, 2 f (all J. A. Justice). The sample of C. d. affinis was from
southwestern Montana (1 m, 1 f), northwestern Wyoming (3 m, 2 f), central Nevada (4 m),
northern Utah (1 m, 3 f), and western Colorado (14 m, 3 f), all collected by J. A. Scott and J. A.
Justice.

Journal of Research on the Lepidoptera

20(2): 86-96, 1981(82)

Chromosomal Studies in Sixteen Species of Indian
Pyralid Moths (Pyralidae)

P. K. Mohanty and B, Nayak

Department of Zoology, F. M. College, P. 0. Balasore, Orissa, BSIDIA 756001

Abstract. The chromosomes of sixteen Indian Pyralid moths collected
from different localities of Bhubaneswar were investigated in male germ line
cells using smear preparations, with two species studied using squashes.
Among the species investigated, eight had n = 3 1 i^ntigastra catalaunalis,
Crocidolomia binotalis, Dichocrocis punctiferalis, Lamprosema indicata,
Lepyrodes neptis, Margorina indica, Maruca testulalis, Pyrausta sanguinalis) ,
two had n ~ 30 {Dichocrocis nilusalis, Galleria mellonella), three had n = 29
{Chilotraea infuscatellus, Cnaphalocrocis medinalis, Orthaga exvinacea) and
one species each had n = 27 {Corcyra cephalonica) , n = 22 {Tryporyza
incertulas) and n == 12 {Chilo supressalis) chromosomes. The trend of
evolution in chromosome number tends toward the lower numbers from
a modal haploid number n — 31 of the family as well as of the order. All
circumstantial evidence indicates the holocentric nature of the chromo-
somes. The sex-chromosomes in the males are not differentiable, but
delayed anaphasic movement of a pair might represent the sex-chromosomes.

Introduction

In spite of intense work carried out during recent years, chromosome
studies in Lepidoptera have not made much headway. Of the 130,000
described species of Lepidoptera only 1,300 species have been chromo-
somally examined, mostly by de Lesse, Suomalainen and Saitoh (c. f.
Robinson, 1971). Most work has been concerned with the chromosomes of
butterflies, which constitute only 10% of the total species. Moths, forming
the majority of species, have been but less attended. Information on
chromosome counts of Indian moths is very meagre at present (Gupta,
1964; RisW, 1973; Nayak, 1975), though some data on a number of
Himalayan butterflies are available (Maeki & Ae, 1966; Saitoh & Abe,
1969, 1970 a, b). Of the family Pyralidae, chromosome numbers of 56
species have been reported. The present note deals with the chromosome
counts of 16 species of Pyralid moths, of which 12 are reported for the first
time.

Materials and Methods

The present investigations were carried out on testes material from late
larvae (5th instar) and early pupae collected from their respective

20(2): 86^96, 1981(82)

87

hostplants. Table I gives the account of hostplants and time and place of
collection of the different species of Pyralids. The cytologicai preparations
were made by the following procedure: hypotonic treatment of testes (in
0.9% sodium citrate) for about 10 minutes; fixation in aceto-alcohol (1:3)
overnight; preparation of smears on pre-warmed albuminised slides using
a drop of 45% acetic acid and finally stained in Heidenheins iron
haematoxylin.

Table I

List of species examined, with foodplant and date of collection given. All
were taken in Bhubaneswar.

Species

Foodplant/Resource

Period of
Collection

Antigastra catalaunalis Dup.

Sesamum indicum

August, 1976

Chilotraea infuscatellus Sn.

Saccharum officinarum

October, 1977

(Sugarcane stem borer)

Chilo suppressalis Walk.

Oryza sativa

March, 1978

Cnaphalocrocis medinalis

Oryza sativa

October, 1977

Guen.

Corcyra cephalonica Staint.

Flour

March, 1976

(Rice moth)

Crocidolomia binotalis Zell.

Brassica sp.

December, 1977

Dichocrocis nilusallis Walk.

Michelia sp.

August, 1976

Dichocrocis punctiferalis

Ricinus communis

September, 1977

Guen.

Galleria mellonella Linn.

Bee -hive

October, 1977

(Wax moth)

Lamprosema indie ata Fabr.

Glycin max

November, 1978

Lepyrodes neptis Cram.

Nyctanthes sp.

September, 1978

Margorina indica Saund.

Trichosanthes anquina

August, 1977

(Pumpkin caterpillar)

Maruca testulalis Gey.

Unidentified

July, 1977

Orthaga exvinacea Hmps.

Mangifera indica

January, 1976

(Mango shoot-webber)

Pyrausta sanguinalis Lirni.

Ocimum basilicum

July, 1976

Tryporyza incertulas Walk.

Oryza sativa

November, 1978

(Rice stem borer)

Observations

Antigastra catalaunalis Dup. (Figs. 1 to 3). A chromosome count in
spermatogonial metaphase cells revealed the diploid chromosome number
as 2n = 62 . Metaphase I normally showed 3 1 bivalents. The data are based
on 70 metaphase I cells in four specimens. Quite often, one bivalent,
presumably the sex-bivalent, appeared to be still undercondensed and
was in the form of a ring while others were at their maximal condensation.

88

J. Res. Lepid.

A number of cells were observed in which almost all the bivalents had been
resolved into univalents except a few which exhibited dumbbell-shape
with a distinct notch along the median line. In late anaphase I, all except
one bivalent separated into their homologues and passed to their
respective poles while components of the laggard still remained on the
equatorial region. Metaphase II showed 31 univalents. The haploid
chromosome number was, thus, confirmed to be n = 31.

Chilotraea infuscatellus Sn. (Figs. 4 to 6) 2n = 58. Examination of 101
metaphase I cells in 10 specimens showed 29 bivalents in each. The
majority of bivalents separated into univalents at anaphase I and passed to
their respective poles synchronously. However, in some cases homologues
remained still attached to each other by one or two interzonal fibres at
their ends and were pulled apart in characteristic V-shapes. As the
chromosomes reached the poles of the spindle, they became split into their
chromatids before they formed daughter nuclei and much before telophase
I. Metaphase II confirmed the haploid number as n ^ 29.

Chilo suppressalis Walk. 2n = 24. In counts of 46 cells in six specimens,
metaphase I cells showed 12 bivalents each. In certain anaphase I cells, a
pair of small deeply stained equal sized bodies were visible on the
equatorial region of the spindle when all other partners had nearly reached
their respective poles. Metaphase II showed 12 univalents (Figs. 7 to 9).

Cnaphalocrocis medinalis Guen. (Figs. 10 to 12) 2n “ 58. Metaphase I
cells showed 29 bivalents, out of which one very often resolved early into
its homologues showing 30 chromosomal elements instead of 29. Meta-
phase n cells showed 29 univalents. Seventeen cells in two specimens
were examined.

Corcyra cephalonica Staint. (Figs. 13 to 15) 2n = 54. Metaphase I cells
showed 27 bivalents each. This has been determined by scoring 66 cells in
nine specimens. In some cells, early resolution of a number of bivalents
into univalents occurred much before the onset of anaphase. A good
number of polyploid cells with double the number of bivalents were also
noticed. In two such cells almost all the bivalents showed early separation,
but the partners of each bivalent remained in close proximity to each other
without any actual contact between them. At anaphase I, all bivalents
separated into their equal sized homologues simultaneously. Metaphase II
cells showed 27 univalents.

Crocidolomia binotalis Zell. (Figs. 16 to 18) 2n ^ 62. A chromosome
count in 59 cells in five specimens established the haploid chromosome
number as n = 31. However, a large number of these cells showed 30
bivalents and two univalents, while few others also contained an admixture
of bivalents and univalents. Anaphase I cells showed the normal disjunc-
tion and synchronous separation of the univalent chromosomes along the
spindle. Occasionally, however, one of the bivalents lagged on the
equatorial plate. Preparations from one specimen contained a faintly

20(2): 86-96, 1981(82)

89

stained chromosome (smaller than the usual sized elements), which
remained off the plate. In anaphase I, this element showed precocious
pole-ward movement to only one of the poles and most likely is the
supernumerary m-chromosome. Metaphase 11 showed 31 univalents.

Dichocrocis nilusallis Walk. (Figs. 19 to 21) 2n = 60. Metaphase I cells
showed 30 bivalents, one of them being distinctly smaller. This has been
determined after examination of 30 cells in four specimens. Certain cells of
the cyst showed 31 chromosomal bodies including the two smaller
elements lying in pair, these may represent resolved homologues of a
bivalent. Some early anaphase I cells were also observed with incomplete
separation of the components of some bivalents. Again, in a good number
of anaphase I cells of a cyst, components of a bivalent trailed behind in
their pole-ward movement. Metaphase II showed 30 spherical univalents.

Dichocrocis punctiferalis Guen. (Figs. 22 to 24) 2n = 62. Examination of
126 cells in seven specimens showed metaphase I cells to contain 31
bivalents in each. Anaphase I indicated lagging movement of a bivalent
which still remained on the equatorial plate when all others had their
homologues passed to their respective poles. Many late anaphase plates
also exhibited the laggard on the bridge of the spindle fibres joining the
daughter nuclei under formation. Metaphase 11 cells showed 31 bivalents.

Galleria mellonella Linn. (Figs. 25 to 27) 2n ~ 60. Examination of 71
cells in nine specimens showed metaphase I cells to contain 30 bivalents
each. Eleven tetraploid cells with double the number of bivalents also were
observed. Anaphase I was normal, but occasionally presented the atypical
lagging behaviour of two of the separating elements. Metaphase II showed
30 univalents.

Lamprosema indicata Fabr. (Figs. 28 to 30) 2n = 62. Metaphase I, as
determined by examination of 152 cells in 13 specimens, showed 31
bivalents. One isolated giant pupal cell was observed that was a tetraploid
in showing 62 bivalents. Some anaphase I cells showed the lagging
behaviour of a pair of small deeply stained equal-sized chromosomal
bodies. Metaphase 11 cells showed 31 univalents.

Lepyrodes neptis Cram. (Fig. 31 to 33) 2n ^ 62. Examination of 55
metaphase I cells of ten specimens revealed 31 bivalents in each. In a
number of cells almost all the bivalents were found to be resolved into their
homologues. Several tetraploid cells with double the number of bivalents
were also encountered. Some anaphase I cells showed a pair of deeply
stained equal-sized bodies (the homologues of a bivalent) on the equatorial
region of the spindle when all other chromosomes had almost reached the
poles. Metaphase II cells showed 31 univalents.

Margorina indica Saund. (Fig. 34 to 36) 2n == 62. Metaphase I cells
showed 31 bivalents as determined by counting 17 cells in three
specimens. Preparation from late pupal testes showed the homologues of
many bivalents remained in closely placed pairs along with a few intact

i;

90

J. Res. Lepid.

91

20(2): 86-96, 1981(82)

Figs.

1 to 3.

Figs,

4 to 6.

Figs.

7 to 9.

Figs.

10

to 12.

Figs.

13

to 15.

Figs.

16

to 18.

Figs.

19

to

21.

Figs.

22

to

24.

Figs.

25

to

27.

Figs.

28

to

30.

Figs.

31

to

33.

Figs.

34

to

36.

Figs.

37

to

39.

Figs.

40

to

42.

Figs.

43

to

45.

Figs.

46

to

48.

Spermatogoniai metaphase, metaphase I and late anaphase I of
Antigastra catalaunalis.

Spermatogoniai metaphase, metaphase I and anaphase I of
Chilotraea infuscatellus .

Spermatogoniai metaphase, metaphase I and anaphase I of Chilo
suppressalis.

Spermatogoniai metaphase, metaphase I and 11 of Cnaphalocrocis
medinalis.

Spermatogoniai metaphase, metaphase I and II of Corcyra
cephalonica.

Spermatogoniai metaphase, metaphase I with m-chromosome and
anaphase I with m-chromosome passing to one pole forming an
extra plate of Crocidolomia binotalis.

Spermatogoniai metaphase, metaphase I and anaphase I of
Dichocrocis nilusallis.

Spermatogoniai metaphase, metaphase I and El of Dichocrocis
punctiferalis.

Spermatogoniai metaphase, metaphase I and II of Galleria

mellonella.

Spermatogoniai metaphase, metaphase I and 11 Lamprosema
indicata.

Spermatogoniai metaphase, metaphase I and II of Lepyrodes neptis.

Spermatogoniai metaphase, metaphase I and 11 of Margorina
indica.

Spermatogoniai metaphase, metaphase I and anaphase I of Maraca

testulalis.

Spermatogoniai metaphase, late abnormal metaphase I cell and II
of Orthaga exvinacea.

Chromosomes of Pyrausta sanguinalis.

Spermatogoniai metaphase, metaphase I and early anaphase I of
Tryporyza incertulas.

bivalents. Several anaphase I ceils showed homologues of a bivalent right
on the equatorial region when all others had their separated homologues
moved towards poles. Metaphase II showed 31 univalents.

Maruca testulalis Gey. (Figs. 37 to 39) 2n = 62. Examination of 60
metaphase I ceils showed 31 bivalents in each. Early separation of a
bivalent into its homologues was noticed in some ceils. Some anaphase I
cells had two of the homologues of a bivalent on the equatorial position
when all others had their separated homologues on the march to the poles.
Metaphase II cells showed 31 univalents.

92

J. Res. Lepid.

Orthaga exvinacea Hmps. (Figs. 40 to 42) 2n = 58. Metaphase I cells
showed 29 bivalents as revealed from examination of 85 cells in six
specimens. Some of anaphase I cells showed the lagging behaviour of two
separating elements of a bivalent. Even at this stage, prior to metaphase II,
which followed without interphase, dissociation of separate homologues
into chromatids had occurred. Metaphase 11 ceils showed 29 univalents.

Pyramta sanguinalis Linn. (Figs. 43 to 45) 2n = 62. Metaphase I cells
showed 31 bivalents, as determined from 161 cells in 18 specimens. Early
separation of a bivalent to its homologues was marked in several cells.
Anaphase I was normal. Metaphase II cells showed 31 univalents.

Tryporyza incertulas Walk. (Fig. 46 to 48) 2n 44. Metaphase I cells
showed 22 bivalents in each of the 106 cells counted in six specimens. In
one cyst, majority of metaphase I cells showed early separation of a
number of bivalents into their homologues. Some cells had all bivalents
resolved much before the onset of anaphase 1. In early anaphase I cells, the
majority of bivalents separated into univalents and migrated to each pole
of the spindle synchronously, although some remained attached to each
other by interzonal fibres and were pulled apart in characteristic V-
shapes, showing bend between arms, giving the impression of the
occurrence of a localised centromere. Late movement of homologues of a
bivalent was also marked in late anaphase I cells. The chromosomes
divided into their respective chromatids as they reached the poles.
Metaphase n cells showed 22 univalents.

Discussion

The Pyralid karyotype is typical of Lepidoptera in general. Numerous
small chromosomes, ill defined meiotic stges, anomalous nature of the
centromere and sex chromosomes render kaiyological analysis obviously
difficult. The chromosomes at spermatogonial mitoses are all homomor-
phic and isodiametric bodies without well defined morphological details.
The early meiotic prophase is diffuse, and a clear delineation of leptotene
and zygotene is not possible due to the appearance of a characteristic
‘synizesis’ stage rendering morphology and exact nature of pairing of the
chromosomes almost obscure. However, the manifestation of the normal
process of lengthwise pairing and parallel conjugation of chromosomes
cannot be doubted since in the following pachytene a haploid number of
thick paired chromosomes forming bivalents became evident. The diplo-
tene for the most part is ‘diffuse' and typical diplotene configuration
cannot be seen due to decondensation of the chromosomes which
practically recede back to the interphase condition. Thus chiasma analysis
is difficult although chiasmata do exist. Chiasma become clear only when
some condensation takes place towards late diplotene and diakinesis,
when mostly they are terminal or near terminal. Interstitial chiasmata with
varying distances from ends producing typical cross configuration are also

20(2): 86-96, 1981(82)

93

observed. Ring configurations with double chiasmata are seen occasional-
ly. In reduction metaphase, the bivalents are evenly distributed and lie
well separated from each other. They are dumbbell-shape in a side view
and oval in polar view.

Although uniformity in chromosome size is a well-defined feature of
Lepidopteran karyotype, slight gradation in size among the chromosomes
are nevertheless evident. Direct correlation between chromosome size
and number is indicated since species with low chromosome number have
larger chromosomes than those with higher number.

While the majority of authors furnish experimental evidence that
Lepidopteran chromosomes are holocentric, with diffuse centromeric
activity along the entire length of the chromosome (Bauer, 1967;
Suomalainen, 1969; Murakami & Imai, 1974), Bigger (1975, 1976) and
Rishi (1978), through the use of improved cytological techniques, claim to
have demonstrated the presence of a localised centromere in such
chromosomes. The current study, however, lends much support to the
holocentric nature of Lepidopteran chromosomes. This conclusion is
based on behaviour during metaphase I when the spindle fibres are
attached to many points on the conjugated chromosomes and no
constriction is marked on any of the chromosomes during their anaphasic
movement in the majority of species investigated. Indeed during no part of
the meiotic cycle is there any evidence of the presence of a localised
centromere. The observation is particularly evident in the meiotic
prophase where usually the chromosomes are less spiralised and are
elongated providing a better display of their linear structure. However, the
appearance of a few ‘V’ shaped chromosomes, similar to centric chromo-
somes in anaphase I of C. infuscatellus and T. incertulas, is an exception
and the occurrence of a localised centromere, at least on those chromo-
somes showing the bend, is indicated. Bigger (1975, 1976) demonstrated
definite constriction on ail the mitotic chromosomes of certain Lepidop-
teran species—Hem brassicae, Pieris napi, Polyommatus icarus and
Pyronia tithonus—hy employing a new air drying technique combined with
ASG banding. According to him at early metaphase the chromosomes of
Lepidoptera exhibit a monocentric type of organisation, or at least a part
of the diffuse centromere (the primary centromere) which exerts dominant
influence over the rest. As metaphase progresses the influence of the
primary centromere is either lost or superseded by the combined influence
of the rest of the diffuse centromere. Thus, according to Bigger, a primary
centromere functions only for a limited part of the meiotic cycle later to be
superseded by diffuse centromeric activity along the whole length of the
chromosome in Lepidoptera. The primary centromere performs the
function of holding the chromatids together at prometaphase and its
disappearance at metaphase reflects the evolutionary change from
monocentric to holocentric chromosomes. Here we may record Schrader's

94

J. Res. Lepid.

(1936, 1939) view of cyclic alternation of centromere from diffuse
organisation in late prophase to a strictly localised one at metaphase in
Amphiuma. Rishi (1978) demonstrated occurrence of a localised centromere
in normal diplotene and diakinesis and metaphase I cells of Trabla vishnu.
Bauer (1967) expressed the opinion that cumulative evidence swung in
favour of evolution of holocentric chromosomes from monocentric ones. If
so, the few bent chromosomes indicating primary centromeric organisa-
tion during present work are the ‘relict’ providing further supporting
evidence in this regard.

Out of the 16 species reported herein, eight species show the haploid
number as n = 31 {A. catalaunalis, C. binotalis, D. punctiferalis, L.
indicata, L. neptis, M. indica, M. testulalis andP. sanguinalis), two species
have n = 30 {D. nilusallis and G. mellonella), three species with n “ 29 (C.
infuscatellus, C. medinalis and 0. exvinacea), one species with n “ 27 (C.
cephalonica), one with n — 22 (T. incertulas) and one has n = 12 (C.
suppressalis). Thus the observed trend appears toward the evolution of
lower chromosome numbers which must have been achieved through
chromosomal fusion. The familiar range of variation, however, lies
between n = 10 as in Perinephala coronata (Bigger, 1960) to n = 41
Haritala ruralis (Bigger, 1960). Polyploidy as a means of numerical
increase of chromosome number in some Lepidopterans, hypothesized by
Lorkovic (1941), appears to have been excluded here since the established
chromosome numbers do not fall into any of a common number. We share
the same view as held by White (1973) and Suomalainen (1969), that

Fig, 49. Histogram showing haploid chromosome numbers in family Pyralidae.

20(2): 86-96, 1981(82)

95

fusion and dissociation are responsible for evolution of chromosome
number in Lepidoptera rather than polyploidy, and that fusion is more
frequent than fission since there are more species with numbers below 31
than there are ones with numbers above 3 1 . Other factors responsible for
an apparent increast in chromosome number include failure of synapsis,
early resolution of one or more bivalents to univalents, and the occurrence
of one or more supernumerary chromosomes as seen in D. punctiferalis.
Out of the 56 species of Pyralids cytologically examined so far, the
majority (26) of species show a haploid chromosome number n ” 31. This
is in accord with the standard Lepidopteran karyotype with the modal
haploid number n 3 1 . Since the sex-chromosomes are not differentiable,
most of the conclusions in this regard are inferential. In the anaphase I of
some species, a pair of chromosomes which show delayed poleward
segregation may represent the XX sex-chromosome pair of the male
Lepidoptera.

Acknowledgments: The authors are grateful to Prof. C. C. Das, Zoology Depart-
ment of Berhampur University, for his helpful suggestions. Thanks are due the
Director of the Zoological Survey of India for identification of the species and to U.
G. C., New Delhi for the financial assistance.

Literature Cited

BIGGER, T. R. L., 1960. Chromosome numbers of Lepidoptera. Pt. 1. Entomol. Gaz.
11: 149-152.

, 1961. Chromosome numbers of Lepidoptera. Pt. IL Entomol. Gaz.

12: 85-89.

, 1975. Karyotypes of some Lepidoptera chromosomes and changes in

their holokinetic organisation as revealed by new cytological techniques.
Cytologia 40: 713-726.

, 1976. Karyotypes of three species of Lepidoptera including an inves-
tigation of B-chromosomes in Pieris. Cytologia 41: 216-282.

BAUER, H., 1967. Die kinetische Organisation der Lepidopteren chromosomen.
Chromosoma 22: 101-125.

GUPTA, Y., 1964. Chromosomal studies in some Indian Lepdioptera. Chromosoma
15: 540-561.

LORKOVic, z., 1941. Die Chromosomenzahlen in der Spermatogenese der Tagfalter.
Chromosoma 2: 155-191.

MAEKI.K. & S. A. AE, 1966. A chromosome study of twenty-eight species of Himalayan
butterflies (Papilionidae, Pieridae); Spec. Bull. Lepid. Soc. Jap. 2: 107-119.
MURAKAMI, A. &H. T. IMM, 1974. Cytological evidence for holocentric chromosomes
of the silk- worms, Bombyx mori and B. nmndarina, (Bombycidae, Lepidoptera).
Chromosoma (Berl.) 47: 167-177.

NAYAK, B., 1975. Studies on the male germinal chromosomes of thirty-one species
of moths and butterflies (Lepidoptera). Prakrati-Utkal University Journal-
Science: Vol. 12 (1 & 2): 141-150.

RISHI, s., 1973. Chromosome numbers of thirty species of Indian Lepidoptera.
Genen Phaenen 16(3): 119-122.

RISHI, s. & K. K. RISHI, 1978. Nature of kinetochore in Trabla vishnu Lef. (Lasio-

96

J. Res. Lepid.

campidae: Lepidoptera). Abst. 3rd. All India Cong, of Cytology and Genetics,
pp. 164.

SAITOH, K. & A. aBE, 1969. Chromosome numbers of some Himalayan butterflies
(Lepidoptera: Danaidae, Nymphalidae and Lycaenidae). Kontyu 37(2): 257-
258.

, 1970a. Chromosome studies in sixteen species of Himalayan butter-
flies— Nymphalidae and Lycaenidae. Spec. Bull. Lepid. Soc. Jap. 4: 141-152.

, 1970b. On the chromosomes of five species of Danaidae from Nepal

Himalaya (Lepidoptera, Rhopalocera). Spec. Bull. Lepid. Soc. Jap. 4: 153-158.

SUOMAI.AINEN, E., 1969. Chromosome evolution in the Lepidoptera. Chromosoma
(Berl.) 47: 167-177.

WHITE, M. J. D., 1973. Animal Cytology and Evolution. 3rd ed., London: Cambridge
University Press, 1973.

Journal of Research on the Lepidoptera

20(2): 97-102, 1981(82)

The Biological and Systematic Significance of Red
Fecal and Meconial Pigments in Butterflies: A Review
with Special Reference to the Pieridae

Arthur M. Shapiro

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

Abstract. Red fecal pellets produced just before pupation and red
meconium voided by teneral adults are distributed in specific butterfly
lineages and may be useful in taxonomy. The red pigments are om-
mochromes; a brief review of their biosynthesis and physiology is presented.
Abnormal production of red ommochromes may follow various disease
processes and injuries in Pierid early stages.

The topic of “red rain” was introduced to several generations of
American Lepidopterists by W. J. Holland, who reproduced on pages 299-
303 of The Butterfly Book (1898) an essay on the subject by Frank Cowan.
Those who rear Nymphalini are familiar with the bright red meconium
which inspired superstitious terror among the European peasantry, but
the literature treating the chemistry and physiology of this phenomenon is
generally unknown to Lepidopterists, and the potential significance of red
excretory pigments in butterfly systematics remains virtually unremarked.
This paper presents a brief survey of these subjects, with some hitherto
unpubhshed information on the phenomenon as it occurs in various
Pieridae.

Meconium — the material voided upon or shortly after eclosion by a
teneral (immature adult) insect — is often pigmented; the color may be
characteristic of the taxon. Red meconia are best known in the Nymphalini,
in the “tortoiseshell” group of genera variously known as Nymphalis,
Aglais, Inachis, Vanessa, etc. (In this paper American rather than
European taxonomic conventions will be followed.) They also occur in
some other Nymphalid genera which are not closely related, such as
Euphydryas and Charaxes. There appear to be very few mentions of fecal
or meconial colors in the literature of butterfly breeding. According to one
anonymous reviewer of this paper, red meconia occur in the Papilionid
genera Pamassius (Tadumia) and Allancastria; P. clodius sol Bryk and
Eisner from Nevada County, California, has pink meconium (Shapiro,
unpublished). In the Pieridae they are widespread but not universal in the
Pierini and Anthocharini (™ Euchloini).

98

J. Res. Lepid.

Chemistry and Normal Biosynthesis

Meconium is regarded as the waste products of metamorphosis. There
are two phases of meconium production in the teneral adult (Lafont and
Pennetier, 1975). In the first, red pigments — ommochromes—and uric
acid are excreted; the second, which is essentially colorless, contains
allantoic acid. Many other compounds have been identified in meconium,
but we are concerned primarily with the ommochromes, a group of
pigments best known from insect eyes, whence the name. They were
reviewed in depth by Linzen (1974), from which much of this section is
abstracted. Ommochromes (Fig. 1) are degradation products of the amino

Oq

NHj

|-CH,-C-COOH

COOH
HjNCH
CH,

6-0 °

COOH
H,N CH

COOH

Cc:)6::

COOH

NH

Glucose

Fig. 1. Structural formulae of the precursor amino acid, tryptophan (A) and its
derivatives ommatin D (B) and rhodommatin (C), the two common red
excreory pigments found in Pierid and Nymphalid meconia. The bio-
synthetic pathways are given in detail by Linzen, p. 123.

acid tryptophan. In the Nymphalini two ommochromes, ommatin D and
rhodommatin, occur as wing pigments as well as in meconium. The same
pigments have been recovered from the meconia of the Palearctic Pierids
Aporia crataegi L. and Pieris brassicae L., in the latter only in veiy small
amounts. Ommochromes seem unreported from Pierid wings. They were
first obtained in large quantities by Becker (1942) and Butenandt et aL
(1954) from the meconium of the European Small Tortoiseshell, Nymphalis
(Aglais) urticae L. In the Red Admiral, Vanessa atalanta L,, up to 130 /xg of
rhodommatin and 115 pg of ommatin D have been recovered from the
meconium of a single animal; in A. crataegi these values are 244 pg and
225 pg respectively. Linzen summarizes the history of these compounds in
the insect as follows (pages 176-177):

These ommochromes are formed early in metamorphosis. They
make their first appearance at the time when the larvae leave the
food and crawl about to find a suitable place for pupation. In
{Nymphalis) urticae, orange pigment appears in the midgut wall at
the time when the larva is spinning the small web to hang itself up.
Twelve hours later, at the time of pupation, the gfit is filled with red
fluid containing about 50 pg each of rhodommatin and ommatin D,
More is known in certain moths. In Ptychopoda seriata Schrk. (Geo-
metridae) and Cerura vinula L. (Notodontidae), studied by authors cited

20(2): 97=102, 1981(82)

99

in Linzen’s review, ommochromes appear twice during development, as
they do in Pieridae. First they appear in the Malpighian tubules, which
excrete them— contributing to a red color in the last few fecal pellets prior
to pupation. Later (after pupation), during the reconstruction of the
tissues, ommochromes appear in vacuoles of the new midgut epithelium
and are released into the lumen of the gut, where they contribute to the
first meconium of the teneral adult.

The only definite adaptive function of these releases is the removal of
excess tryptophan, but they may serve subsidiary functions as well. In
many Lepidoptera meconium is retained by the teneral adult until it is
ready for flight, or even longer, but it may be released early if the animal is
disturbed. Such releases are often forceful, with a definite potential for
defense against predation. Meconium has not, to my knowledge, been
examined for repellent or noxious properties. It is perhaps not too far-
fetched that the blood-red color of first meconium forcibly voided in such
circumstances by the otherwise defenseless teneral adult may be alarming
or at least distracting to vertebrate predators having color vision.

Distribution in the Pieridae

In the Holarctic pierine fauna the last few (1-4) fecal pellets produced
before pupation, and the first meconium of the adult, are red in certain
lineages and not in others; the two seem always to be coupled. These facts
suggest that they may be useful characters for phylogenetic reconstruction
and higher classification. The only previous use of meconium m taxonomy
appears to be by DeBach et al. (1978), working on Aphytis spp.
(Hymenoptera, Aphelinidae).

Ommochrome-excreting Holarctic pierines belong to the Pieris (Synchloe)
callidice Hbn. species-group, all of whose taxa thus far examined {callidice,
protodice Bdv. & LeC., occidentalis Reak., nelsoni Edw.) show the
phenomenon. Meconia are unreported in the formal literature for the
closely related subgenus Pontia and for the enigmatic P. (Pontia?) sisymbrii
Bdv., but one of the anonymous reviewers of this paper reports red
meconia for the Palearctic P. (Pontia) daplidice L. and glauconome Klug.
No red ommochromes are found in feces or meconium of the P. rapae L.
and P. napi L. species -groups (subgenus Artogeia) except under patho-
logical circumstances (see below), and they are present but scarce in P.
(Pieris) brassicae. The highest levels ever measured occur in Aporia
crataegi and in some of the South American taxa discussed below.

In the Andean-Patagonian group of Pierid genera, red fecal pellets and
meconia seem generally distributed in the Tatochila-Phulia series. Taxa
reared by me, all of which show the phenomenon, are Reliquia santamarta
Ackeiy and the following entities in Tatochila: sterodice species-group:
sterodice Stgr., arctodice Stghr., macrodice Stgr., vanvolxemii Capr.,
mercedis Esch.; xanthodice species -group: xanthodice Lucas; autodice

100

J. Res. Lepid.

species-group: autodice Won., hlanchardiiBntl. Quantities are especially
large in the sterodice group.

In the Anthocharini, red feces and meconia occur in the Holarctic
Euchloe ausonia species -group: ausonia ausonia Hbn., a. crameri ButL,
and ausonides Lucas (several segregates). They are pink in Anthocharis
Sara Lucas and Euchloe hyantis Edw. Red excreta are recorded formally in
only one Coliadine species', Eurema hecabe L. (Fukuda et al., 1972), but
are reported by the aforementioned reviewer of this paper to occur in both
Catopsilia and Gonepteryx.

The distribution of red feces and meconia in Pierini and Anthocharini
corresponds to the distribution of “red” eggs (Shapiro, 1981). This at least
suggests the possibility that the red pigment developed by maturing Pierid
eggs might be an ommochrome, since Horowitz (1940) and Horowitz and
Baumberger (1941) found that eggs of the marine Echiuroid worm Urechis
undergo color changes similar to those of Pierids. In this case, the
ommochromes function as respiratory pigments; the color change indicates
a shift from the oxidized to the reduced state. The well-known color
changes of Euchloe prepupae (from gray-violet and yellow to pinkish gray
in E. ausonides, from green to lurid purple in E. hyantis) may also be
mediated by ommochromes.

Tlie taxonomic distribution of ommochromes as excretory pigments in
Pieridae corroborates the conventional phylogenetic interpretation of the
Andean-Patagonian fauna (Shapiro, 1980): post-Pliocene derivation
from a Holarctic ancestor of the Synchloe type. It also supports the
electrophoretic evidence of Courtney (1980) and Geiger (1981) indicating
a close relationship between Pontia and Synchloe on one hand and the
Anthocharini on the other. This raises the serious question of whether the
bizarre Andean Eroessa is really an orange-tip at all. It has been
considered the most primitive member of that group, but that interpreta-
tion would remain tenable only by deriving the Pierini from Euchloe or
something near it, an unlikely scenario on various grounds.

Phylogenetic interpretation in the Pieridae is complicated by an obvious
predisposition to parallelisms and convergences which makes even
cladistic reasoning highly unsatisfying. The distribution of excretoiy
ommochromes parallels phenotypic resemblances which have usually
been interpreted as superficial and convergent. Both types of characters
could have evolved as correlates of dietary specialization (on inflorescences
and infractescences); if tryptophan loads are heavier in the diets of these
groups than in those of leaf-feeders, physiological parallelism or con-
vergence would be implied. It must be stressed that there is no a priori
reason why biochemical characters should be less prone to convergence
than morphological ones.

Ommochromes in Pathological Contexts

Linzen (1974, p. 177) observes that ommochromes may be produced

20(2): 97-102, 1981(82)

101

when insects or their tissues “are maltreated so as to disrupt nomial
metabolic function... it is assumed that under these circumstances the
animals draw on their protein reserves as an energy source, thus leading to
an excess of tryptophan.” Phenomena of this sort are readily observed in
Pieridae.

Pierid breeders eventually make the acquaintance of “red dribble
disease,” a condition of unknown etiology which attacks fourth- and fifth-
mstar larvae with invariably fatal results. The afflicted larva suddenly
stops feeding and begins producing a more or less constant dribble of
bright red diarrhea. It never feeds again, but may live for several weeks,
shrinking all the while as it consumes its reserves. Most afflicted larvae
spin a great deal of silk and may appear to be prepared to pupate, but fail
to do so. The condition is associated with etiolated food, with certain
Crucifers which are edible but apparently mimical to survival (such as
flowering tops, but not rosettes, of the weed Barbarea vulgaris R.Br.), and
with inbreeding depression in long-term cultures; it does not seem to be
infectious or contagious. I have encountered it in cultures of Heris
(Synchloe) and of Tatochila and — interestingly— of P, rapae and P. napi,
which do not ordinarily produce visible quantities of excretory om-
mochromes.

Prepupae of Tatochila placed at 37 °C or higher become ssuffused with
pink, which carries over into the pupa but fades gradually over several days
if the pupa is returned to 25°C. Diseased pupae of Pierk (Synchloe) and
Tatochila, but not Pieris (Artogeia), often develop a red “saddle” in the
mid-dorsum which may spread over the entire body, but not the wing-
cases; such pupae never recover and are found to contain a bacterial broth.
Pupae of Euchloe ausonides and hyantis develop a characteristic rose-
carmine flush if diseased or attacked by parasitoids. The flush starts in one
area and gradually spreads over the abdomen as more tissues become
involved. Heat-shocked pupae of Nymphalini which are killed by the
treatment generally turn purple-pink. All of these phenomena are
suspected to involve abnormal ommochrome synthesis.

Abnormal failure to produce excretory ommochromes is also observed in
from less than one percent to five percent of Pierini in mass culture; only
the second (colorless) meconium is excreted. This phenomenon is
observed both as scattered individuals in large broods and as clusters of
sibs from particular families. The implication of genetic variation in the
ommochrome pathway suggests caution in using evidence from one or a
few reared examples in a typological way for taxonomic purposes. Nonethe-
less, the usefulness of these pigments in classification and the understand-
ing of their function would benefit if more breeders would record the color,
if any, of the final larval feces and meconia of whatever species they have in
culture, and publish this information.

Acknowledgments. I thank Mr. Marc Tatar for help with the literature. South

102

J. Res. Lepid.

American work contributing to this paper was funded by the National Science
Foundation (DEB-76- 186 11), the National Geographic Society, and the UCD
Institute of Ecology. Two anonymous reviewers of the paper provided useful
suggestions, the Fukuda reference (which is in Japanese), and unpublished
information on the distribution of red meconia in some Palearctic genera.

Literature Cited

BECKER, E., 1942. Uber Eigenschaften, Verbreitung, und die genetischentwick-
lungs — physiologische Bedeutung der Pigmente der Ommatin — und Om-
mingruppe (Ommochrome) bei Arthropoden. Z. Vererb. Lehre 80: 157-204.
BUTENANDT, A., u. SCHIEDT, E. BIEKERT, & P. KORNMANN, 1954. Uber Ommochrome. I.
Mitteilung: Isolierung von Xanthommatin, Rhodommatin, und Ommatin C aus
den Schlupfsekreten von Vanessa urticae. Liebigs Ann. Chem. 586: 217-228.
COURTNEY, s. P., 1980. Studies on the biology of the butterflies Anthocharis carda-
mines (L.) and Pieris napi (L.) in relation to speciation in the Pierinae. Unpub-
lished Ph.D. thesis. University of Durham, U.K.

DeBACH, P., M. ROSE & D. ROSEN, 1978. Biological and systematic studies of develop-
mental stages in Aphytis. IQ. Meconia as a possible systematic tool in Aphytis.
Hilgardia 46(3): 102-112.

FUKUDA, H. et aL, 1972. lusects' Life in Japan. Vol. 3, Butterflies, (in Japanese.)
Hoikusha, Osaka, p. 71.

GEIGER, H. J., 1981. Enzyme electrophoretic studies on the genetic relationships of
Pierid butterflies. I. European taxa. J. Res. Lepid. 19: 181-195.

HOLLAND, w. J., 1898. The Butterfly Book. Doubleday, New York. 382 pp.
HOROWITZ, N. H., 1940. A respiratory pigment from the eggs of a marine worm. Proc.
Nat. Acad. Sci. USA 26: 161-163.

HOROWITZ, N. H. & J. P. BAUMBERGER, 1941. Studies on the respiratory pigment of
Urechis eggs. J. Biol. Chem. 141: 407-415.

LAFONT, R. & J. L. PENNETIER, 1975. Uric acid metabolism during pupal-adult
development of Pieris brassicae L. J. Ins. Physiol. 21: 1323-1336.

LINZEN, B., 1974. The tryptophan-ommochrome pathway in insects. Adv. Ins.
Physiol. 10: 117-246.

SHAPIRO, A. M., 1980. Convergence in pierine polyphenisms. J. Nat. Hist. 14: 781-
802.

, 1981. The Pierid red-egg syndrome. Amer. Nat. 117: 276-294,

Journal of Research on the Lepidoptera

20(2): 103-110, 1981(82)

On the Nomenclature of Colias alfacariensis Berger 1948
(Lepidoptera: Pieridae)

Otakar Kudma

Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerailee
150-164, D-5300 Bonn 1, Germany

Abstract. The nomenclature of the butterfly usually called Colias australis
is discussed in chronological order, with comments regarding the relevant
papers published by other authors. The valid name for the species, its author
and date of publication are established, this being Colias alfacariensis
Berger 1948.

“Berger’s Clouded Yellow” has usually been called ''Colias australis”
and its authorship has usually been attributed to “Verity 1911.” However,
two other names— “Co/ias alfacariensis” and "Colias ca/ida”— have also
been used for the species, and their authorship attributed mainly to
“Ribbe, 1905” and “Verity, 1916”, although some other authors have also
been occasionally quoted, e. g. Berger, 1948; Berger & Fontaine, 1948 and
Verity, 1923. Kocak (1981) in his painstaking examination of the
nomenclature of the European butterfly species, was already fully aware of
the unavailability of the name australis in connection with Verity’s
authorship dated 1911 (Verity, 1905-11); Kocak apparently presumed
that the name australis was made available later, elevated to the species-
group rank by an unspecified author at an unspecified date to become the
senior subjective synonym, and consequently the valid name, for the
species. Dutreix (1981) retained the name and authorship as "Colias
australis Verity 1911” but his useful compilation of the literature on the
species helped in the search for the correct name of the species. Blab &
Kudrna (1982) already employed the correct name, author and date of
publication Colias alfacariensis Berger 1948; they did not explain the
reasons behind their action as to do so was beyond the scope of their work.

A chronological order has been chosen as being the most opportune
method to elucidate the complex taxonomic and nomenclatural history of
“Berger’s Clouded Yellow.” Throughout the paper on several occasions a
reference is given to the International Commission on Zoological Nomen-
clature and to the International Code of Zoological Nomenclature; to save
space the former is abbreviated ICZN and the latter simply “Code”. I have
the pleasure of thanking R. V. Melville (ICZN, London), G. Bernard!
(Museum National d’Histoire Naturelle, Paris), A. 0. Kocak (Ankara
University, Ankara) and Ichiro Nakamura (USA) who at various times

104

J. Res. Lepid.

discussed with me some aspects of zoological nomenclature and/or
provided valuable advice which in part concerned also some problems
related to this paper.

Ribbe (1905) described an aberration of Colias hyale Linnaeus 1758 and
named it alfacariensis. His type-material came from the mountains Sierra
de la Yedra (ca. 1800 m) just north of the village Alfacar situated north of
Granada, in southern Spain (i.e. Andalusia); he called this mountain range
incorrectly ‘Sierra de Alfacar’ after the above mentioned village which he
used as his base and this name became subsequently very widespread in
entomological literature, particularly in the German language. The
taxonomic category ‘aberration’ has been used either for some kind of
individual form (s.l.) or for various extreme forms ranging often as far as
pathologial and teratological individuals. “Aberration” is not among the
taxonomic categories recognized by the ICZN. The rank of any aberration
is therefore to be interpreted as infrasubspecific and, consequently, the
names originally proposed for aberrations are unavailable. Should a name
originally proposed for an aberration, or any other infrasubspecific
category, be elevated by one of the subsequent authors to the species-
group rank (i.e. the species or the subspecies), it takes the date and
authorship of its elevation (cf. Code, Art 10b) and its previous infrasub -
specific application remains irrelevant and immaterial. To avoid any
subsequent confusion (cf. Reissinger, 1971) it should be pointed out here
that Ribbe (1905) in the same paper distinguished clearly between
infrasubspecific names for individual form (i.e. aberrations) and available
names interpreted now as of subspecies -rank, which he in accordance with
the predominating custom of the period called “varietas”; Satyrus actaea
nevadensis Ribbe 1905 and Chrysephanus [sic] gordius granadensis Ribbe
1905 were both originally designated as “var[ietas|.”

Although the interpretation of the rank and status of the species -group
and lower category names treated in the works of Seitz (1906-09, 1929-32)
often presents considerable difficulties owing to ambiguity, it may be
noted that Roeber (1907) retained unequivocally the rank of aberration for
the taxon: “Ribbe found in Sierra d’Alfacar [sic] (Andalusia) an aber-
ration which he calls alfacariensis: cf lighter yellow, the underside of the
hindwing being more greyish yellow, 9 above greenish white, similar to
edusa ab. helice but with smaller black markings, the underside also being
very similar to that of helice.'’ (Quotation cited here is taken from the
English edition of the book.)

Verity (1905-11) described a “race” of Colias hyale hyale Linnaeus,
1758 from southern Spain (i.e. “Andalusia”) and named it australis. The
original description was given on page 347, the full original combination on
page XXXV of the same publication, and both were published at the same
time (Verity, 1914); the necessity to consult the systematic index to
establish the original combination and rank of all taxa named in Verity’s

20(2): 103410, 1981(82)

105

work cited above is given elsewhere (Kudma, 1983, in print). Verity (1905-
1 1) also defined the “race” as a infrasub specific category, i.e. below the
subspecies-rank, and the quadrieommal original combination Coiias hyale
hyale australis makes all abundantly clear. It may be noted that also races
of monotypic species remain infrasubspeciilc and their names unavailable.

Verity (1916) described a seasonal form of Coiias hyale (treated this time
as a monotypic species) characteristic of the summer generation of the
populations inhabiting southern Europe and named it calMa; at the same
time he designated syiitypes (1 1 9) from Toscana (C. Italy). Names

proposed for seasonal fonns are always treated as unavailable names as
they are excluded from the Code; they are infrasubspecific names as they
apply only to a part of the population(s) of the species, being exclusive of a
certain season of the year. It should be mentioned here that also names
applied to a selected population(s) as the whole cannot be treated as
species-group names (cl Code, draft, 3rd edition).

Verity Querci (1923-24) used the name calida for the southern
Emopean race of Coiias hyale characterized by the presence of the
seasonal fomi calida among the summer generation(s); the paper was
serialized and the page concerned was published in 1923. Verity & Querci
(1923-24) piovided no description of the race calida, however, a biblio-
graphical reference is given to the original description of the seasonal form
of the same name. A tririom.inal combination, Coiias hyale calida, seems
implied. The race calida includes three seasonal forms: mrnalis Verity
1908 in the first generation, calida in the second and third generation and
“extraord. gen. hyale, L.” in the fourth generation. Verity’s taxonomic
category “race” has given much cause for concern owing to its ambiquity;
it was“-and still often is— quite thoughtlessly interpreted as an equivalent
of the subspecies. The names originally proposed by Verity for races have
therefore been usually treated as if they were available names of
subspecies-rank as from the date of their first publication (cf. Cockayne,
1952), Verity made it abundantly clear on several occasions that Ms
“rate” had little, if anything, in common with the subspecies (Verity,
1905“11, 1912, 1929, 1940); he more or less replaced the subspecies with
the “exerge” (Verity, 1925-26, 1929) in all but his early publications (e.g.
Verity, 1905-1 1, also Turati& Verity, 1911, 1912) and placed the race one
rank below it, i.e. as an infrasubspecific and consequently unavailable
name; this is true even about all monotypic species where the race-name
comes placed third in the implied trinominal original combination
(Kudma, 1983). The significance of the intentions of the original author
has not been adequately appreciated in the present Code, but this has
already been corrected by ICZN (cf. Code, draft, 3rd edition). Verity’s
“exerge” (i.e. subspecies) was determined by its hereditary characters (i.e.
genetically fixed), Ms “}*ace” was determined by its somatic characters (i.e.
phenotypic) caused directly by environmental pressures. This means that

106

J. Res. Lepid.

the “race” differed from the “exerge” (i.e. the higher category) in its
phenotypic features while their genetic characters remained identical. The
race was often denoted by the name of a form thought to be predominant or
characteristic (though not necessarily exclusive) of it, or of one of its
distinct broods in case of polyvoltine species. Such is the case of the name
calida used for a seasonal form and applied by Verity & Querci (1923-24)
to the race characterized by the presence, or predominance, of that
seasonal form in its summer brood(s). The race does not exclude examples
of other conspecific forms from its populations (i.e. so called “mixed”
races of Verity), the “exerge” (i.e. subspecies) does precisely the opposite,
it being exclusive to a definite geographical area; only where two such
“exerges” of the same species meet they form a “contact zone” and can
interbreed. The race has no definite range. In case of all monotypic
species, the genetic characters are specific, the phenotypic features
“racial.” Before his indirect replacement of subspecies by the new fanciful
term “exerge” Verity (1920: 146 in 1919-22) stated: “the term ‘sub-
species’ I should restrict to particular groups of races which only just fall
short of the definite group we call ‘species’.”

BoUow (1930) made the following interpretation of the names calida and
australis: “For the very fine and large summer generation of southern
Europe Verity chooses as typical representative the hyale flying in
Tuscany and names it calida'' and “Further large brightly coloured cfcf
and for a great part also similarly coloured 99 are shown by the race
australis Vrty. from Andalusia. The black markings are often reduced.”
(Quotation from the English edition of the work.) The infrasubspecific
status was retained.

Berger (1945) was first to recognize that Colias hyale consisted of two
distinct species the adults of which differed only slightly but which had
very distinct early stages, particularly the larvae, and one of them was a
monophag feeding on Hippocrepis comosa. In his preliminary com-
munication Berger (1945) cautiously avoided to propose a new name for
the “new species” and called it simply after its foodplant to distinguish it
for the oligophagous Colias hyale; he provided no description but
suggested its identity with Colias alfacariensis of Ribbe (1905).

Berger (1948) published another preliminary paper on the two Colias-
species which contained a brief but adequate description of adults and the
early stages of the “new species”, with notes on its biology and distribution
chiefly in England; he also identified it as Colias alfacariensis. He elevated
to the species-rank the aberration alfacariensis and, erroneously, retained
the authorship and date of publication as: Ribbe 1905^ Berger (1948) was
apparently unaware of the consequences of the elevation of an unavailable
name to the rank of species-group, i.e. that he, not Ribbe, became the
author of the name, and that also the date of publication changed
accordingly: Colias alfacariensis Berger 1948. In the circumstances, he

20(2): 103-110, 1981(82)

107

gave no information regarding the type-material (i.e. syntypes) of the new
species and made no statement with regard to the type-locality; nonethe-
less, as he reported in his paper specifically alfacariemis from England, he
used and listed English specimens when writing up his description of the
species. It is important to state here that Berger’s (1948) paper containing
the original description of alfacariensis was published in August 1948.

A little earlier Berger & Fontaine (1947-48) started to publish a more
detailed paper on Colias alfacariensis and dealt particularly with the
biology and the early stages of both closely related species. The paper was
serialized and the name alfacariemis was applied to the earlier published
description, including also a detailed comparative study of adults, as late
as in the last installment which appeared in December 1948. Berger &
Fontaine (1947-48) attributed the authorship again to Ribbe 1905; they
were apparently both unaware of the nomenclatorial consequences of their
action. Also the second description satisfied the Code.

Hemming & Berger (1950) discussed the nomenclature of the new Colias
and correctly observed that the name Colias alfacariemis became first
available under the Code in 1948 and the author of that name was Berger.
However, they were in error as they placed Colias alfacariemis Berger
1948 in synonymy and replaced it with Colias australis, the authorship and
date of publication they gave to Verity 1911. This replacement was
apparently suggested by F. Hemming who must have overlooked both the
original combination stated by Verity (1905-11) and his definition of the
race, either of which were alone adequate to show the infrasubspecific
nature and the subsequent unavailability of the name australis. In fact,
Hemming & Berger (1950) were the first authors to apply the name
amtralis to the species identical with Colias alfacariensis Berger 1948.

Cockayne (1952) rejected both previously used names, Le. Colias
alfacariensis (regardless of authorship) and Colias australis Verity 1911
and pointed out that the oldest name for the species was ca/ida Verity , with
the date of publication 1923 when this was utilized for a race. He failed to
observe that the authors of the paper were Verity & Querci and the infra-
subspecific nature of the race in Verity’s works. Cockayne’s (1952)
conclusions received little support and attention of subsequent authors,
though the name calida was occasionally used chiefly for the “subspecies”
of australis inhabiting central Europe, that is quite contradictory to both
its original description and the type-locality given by Verity (1916).

Following the conclusions of Hemming & Berger (1950) the name Colias
amtralis Verity 1911 became generally accepted by most of subsequent
authors. There appeared a distinct need to provide an objective definition
of the species by selection and designation of the name-bearing type. Riley
(1954) located in the British Museum (Natural Histoiy) the rest of the type
series that served Verity (1905-11) for the description of Colias hyale hyale
amtralis V erity 19 1 1 at his brief visit to the Museum while in London, after

108

J. Res. Lepid.

only perfunctoiy examination of the unknown number of specimens. Riley
(1954) pointed out the somewhat uncertain origin of the types and
selected the lectotype (male). Warren (1954) aware of small differences
usually present in male genitalia of the two species examined the genitalia
of the lectotype; he pointed out that the selection of the specimen was
unfortunate, if not questionable, as its genitalia were somewhat “transi-
tional.” There is no more need for this lectotype, perhaps a lucky solution
in the circumstances.

Reissinger (1971) discussed the nomenclature of the species. at con-
siderable length and concluded that it should be named Colias alfacariensis
Ribe 1905. His argument was based chiefly on facts irrelevant and/or
immaterial from the nomenclatorial point of view and indicated his relative
unawareness of both the principles of systematic zoology and the rules set
out by the ICZN in the Code. Part of Reissinger’s (1971) argument
consists of misinterpretation of the known facts, such as the distinction
Ribbe (1905) made between individual forms (“aberratio”) and sub-
species (“varietas”).

The above perspective completes a taxonomic history full of errors
which resulted in part from some unfortunate oversights of otherwise
acknowledged entomologists. It is to be hoped that the name Colias
alfacariensis Berger 1948 will soon replace the invalid names discussed
here. It would seem most opportune to conclude this paper with the
designation of the name-bearing type, i.e. the lectotype, of this species
from the material that served Berger (1948) for his description. This would
be best done in cooperation with the author himself, L. A. Berger.
Unfortunately, all my earlier attempts to communicate with him failed to
produce an answer. Should he decide to designate the lectotype from his
syntypes, the selection of the right specimens is of utmost importance. It
would seem that southern England should be the type-locality of Colias
alfacariensis for the time being, to be later restricted according to lecto-
type (to be designated).

It may be useful to summarize here the original combinations of both
infrasubspecific and available names cited in this paper:

A) Index of unavailable (infrasubspecific) names:

Colias hyale alfacariensis Ribbe 1905
Colias hyale hyale australis Verity 1911
Colias hyale calida Verity 1916

B) List of available names for species (synonymy):

Colias alfacariensis Berger 1948 (nec Ribbe 1905)

Colias australis Hemming & Berger 1950 (nec Verity 1911)

Colias calida Cockayne 1952 (nec Verity 1923)

Literature Cited

BERGER, L. A. 1945. Espece nouvelle pour la science. Lambillionea 44(1944): 9-10.

20(2): 103410, 1981(82)

109

. 1948. A Colias new to Britain, Entomologist 81: 129-131.

BERGER, L. A. & M. FONTAINE, 1947-48. Une espece meconnue du genre Colias F.

LambilUonea 47(1947): 91-98, 48(1948): 12-15, 21-24, 91-110.

BLAB, J. & 0. KUDRNA. 1982. Grundlagen fur ein Schmetterlings-Hilfsprogramm:

Oekologie und Schutz der Tagfalter und Widderchen. Natursch.

HOLLOW, c. 1930. 2. Familie: Pieridae, Weisslinge. Gross-Schmetterl. Erde (Suppl.)
1: 93-125.

COCKAYNE, E. A. 1952, CoUos calida Verity: the correct name for the butterfly lately
added to the British list. Entomologists ’s Rec. J. Var. 64: 166-168.

DUTREDC, c. 1980. Etude des deux especes affines Colias hyale Lirme et Colias
australis Verity. Alexanor 11: 297-316.

iffiMMiNG, A. F. & L. A. BERGER. 1950. Nouvelles regies de nomenclature. Application

au cas Colias hyale et Colias australis. Lambillionea 50: 2-9.

KOCAK, A. o. 1981. Critical check-list of European Papilionoidea. [1.] Priamus

1: 46-90.

KUDRNA, 0. 1983. An annotated catalogue of the butterflies named by Roger Verity.

Memorie Soc. ent. ital. (Ms ca. 250 pp., in print).

REissiNGER, E. 1971. Die geographisch-subspecifische Gliederung von Colias
alfacariensis Ribbe unter Beracksichtigung der Migrationsverhaltnisse.
[1]. Atalanta, Munnerstadt 3: 145-176.

RIBBE, c. 1905. Einige neue Formen von Schmetterlingen aus Andalusien. Societas
ent. 20: 137-138.

RILEY, N. D. 1954, The lectotype of Colias australis Verity. Entomologist’s Rec. J,
Var. 66: 35-36.

ROEBER, J. 1907. 2. Familie: Pieridae, Weisslinge. Gross-Schmetterl. Erde
1: 39-74.

SEITZ, A. (Ed.). 1906-09. Die palaearktischen Tagfalter. Gross-Schmetterl. Erde
1: 1-379.

1929-32. Die palaearktischen Tagfalter. Supplement. Gross-Schmet-

terl. Erde (Suppl.) 1: 1-399.

TURATI, E. & R. VERITY. 1911. Faunula Valderiensis nell’alta valle del Gesso (Alpi
Maritime). Boll. Soc. ent. ital. 42(1910): 171-265.

. 1912. Faunula Valderiensis nell’alta valle del Gesso (Atpi Maritime).

BoU. Soc. ent. ital. 43(1911): 168-236.

VERITY, R. 1905-11. Rhopalocera Palaearctica. Papilionidae et Pieridae. 86 + 368
pp., 2 + 12+72 pis.; Roger Verity, Firenze.

1912, Considerazione sulla classificazione dei Lepidotteri e loro

applicazione alio studio di alcuni problemi della evoluzione. Monitore zoU.
ital. 23: 45-56.

- 1914. Dates of publication of “Rhopalocera Palaearctica. IconograpMe

et description des papillons diumes de la region palearctique par Roger
Verity (Papilionidae et Pieridae)”. Novit. zool. 21: 426.

1916. The British races of butterflies: their relationship and nomen-
clature. Entomologist’s Rec. J. Var. 28: 73-80, 97-102, 128-133, 165-174.

. 1919-22. Seasonal polymorphism and races of some European

Grypocera and Rhopalocera. Entomologist’s Rec. J. Var. 31(1919): 26-31, 43-
48, 87-89, 121-129, 178-184, 193-201; 32(1920): 3-8, 140452; 33(1921): 170-
176, 190-193, 210-214; 34(1922): 12-15, 68-73, 89-93, 124-142.

no

J. Res. Lepid.

1925-26. Remarks on the evolution of the Zygaenae and attempts

to analyze and classify the variation of Z. lonicerae, Scheven, and of Z. trifolii,
Esp., and other species. Entomologist’s Rec. J. Var. 37(1925): 101-104, 118-
121, 135-138, 154-158; 38(1926): 9-12, 22-26, 57-62, 69-74.

1929. On the necessity of a revision of the rales of entomological

nomenclature concerning groups of lower rank than specific one. Int. Cong.
Ent. 4(1928)(2): 479-480.

1940. Le farfalle diurne d’ltalia. 1. Considerazioni generali. Super-

famiglia Hesperides. 34 + 131 pp., 4 + 2 pis.; Marzocco, Firenze.

VERITY, R. & O. QUERCI. 1923-24. An annotated list of the races and seasonal poly-
morphism of the Grypocera and of the Rhopalocera of peninsular Italy.
Entomologist’s Rec. J. Var. (Suppl.) 35(1923): 1-4, 5-8, 9-12, 13-16, 17-20,
36(1924): 21-24, 25-28, 29-32, 33-36, 37-40, 41-44, 45-46.

WARREN, B. c. S. 1954. A note on the genitalia of Colias australis Verity. Entomolo-
gist’s Rec. J. var. 66: 36.

Journal of Research on the Lepidoptera

20(2): 11M22, 1981(82)

Some Little-Known U* S. Publications on
Lepidoptera

Cyril F. dos Passes

Research Associate, Department of Entomology, American Museum of Natural History,
New York, New York 10024

Abstract. A confusing history is reviewed, namely, that of the bulletins put
out by the Boston Entomological Society under two titles, viz, “The
Lepidopterist” (1916-17) and “Lepidoptera” (1918-21). An additional
publication, by Samuel E. Cassino, which also went by the name “The
Lepidopterist” (1918-31) is given due notice. All known numbers of these
serials are listed, with dates of issue, pagination and inclusions.

Foreword

This is the first of a two-part series, drawing upon manuscripts which have been
entrusted to me by Dr. dos Passes since failing health has curtailed his research
activities. Among his unpublished papers are dissertations on a number of obscure
publications devoted to Lepidoptera. The serials described hereinafter are
probably the most important of those short-lived bulletins and occasional papers.
Others will be made the subject of a later article. The collations are verbatim from
the dos Passes manuscripts, as is also the historical account aside from some minor
emendation and rearrangement. An included letter from John D. Sherman, the
bookseller, has been omitted since its gist was to corroborate the sequence of plate
figures as listed. The latter topic is touched upon further in the text. Aside from a
trifle of editing, and preparation of the abstracts, I have merely dipped into the
wealth of bibliographic data and notes which Dr. dos Passos had assembled. I have
little claim to co-authorship.

Historical Summaiy

The early 20th century was a period of great activity among amateur lepidopterists
in the United States. Holland’s Butterfly Book had been pubhshed in 1898 and his
Moth Book appeared in 1903. They were most successful and gave great
encouragement to amateur collectors besides enlisting many new members into the
hobby. This from time to time resulted in the formation of clubs and societies and
the publication of amateur journals. Outstanding among these was the Boston
Entomological Club which flourished during this era and sponsored a bulletin
which, along with an offshoot therefrom, is described herein. It should be noted that
this club was separate from the Cambridge Entomological Club which published
Psyche during the same period.

As “Official Bulletin of the Boston Entomological Club” (thusly proclaimed on

‘Abstract and foreword by L. P. Grey, Rte. 1, Box 216, Lincoln, Maine 04457

112

J. Res. Lepid.

the masthead of each issue) numbers 1-10 of volume 1 were published under the
name “The Lepidopterist”, with Rudolph C. B. Bartsch designated as editor. So
far, so good. But then dissention appears to have seized upon the club. Bartsch
resigned as editor (under pressure, one gathers). Beginning with volume 1, number
11, the masthead cites Samuel E. Cassino, publisher, continued through numbers
12 and 13 which concluded the volume. The changeover was explained (rather
lamely) as being due in part to trouble with postal service in the Boston area and
Cassino’s ability to make deliveries more promptly from his printing establishment
in Salem.

One does not have to read altogether between the lines to learn of disagreements
at this time; some of them were aired publicly, in the bulletins. In return for his
services as publisher, Cassino began to demand full editorial powers. These, the
club refused to grant, whereupon Cassino retaliated by copyrighting the title and
continuing with volume 2, number 1, of “The Lepidopterist”. This action was
promptly disclaimed, and so henceforth Cassino’s volume 2, number 2, through to
the final issue, volume 5, number 3, are no longer mastheaded as “Official” and are
to be regarded as a private enterprise sponsored and conducted by Cassino
(although still being issued under the original title, “The Lepidopterist”).

Meanwhile, back at the Club, and beginning likewise as volume 2, number 1, the
“Official Bulletin” was continued under a new title, viz, “Lepidoptera”. The editor
was Nathaniel Stowers, with Robert Swift as scientific editor and E. F. Knight as
advertising manager. This publication was more regular than Cassino’s but did not
last so long. Both, however, published about the same number of issues. For
collation purposes the issues sponsored officially by the Boston Entomological
Club are given in order, under the two titles but treated as one serial, while
Cassino’s schismatic bulletin is collated separately.

Most of the needed data were available in the dos Passos library (which has been
donated to Wittenburg University). Photostats of two lacking numbers were
supplied by Miss Nina Root, of the American Museum of Natural History library;
she aided with other services also. Mr. John D. Sherman assisted, sharing his
formidable knowledge in this field, as did some other colleagues who were queried
in an effort to make reasonably certain that the collations are complete and
accurate.

There remains a bit of a question if all of the plates are accounted for properly,
since these were not always numbered or even bound into the individual numbers.
In fact, the plates sometimes were issued separately and after the number in which
they should have appeared had been published. This, if nothing more, may have
elicited Sherman’s expressed judgment that both publications were “criminal”!

Regardless of this and despite whatever minor flaws in collations may yet be
discovered, the following resume should be of abiding utility for a variety of reasons.
These serials are replete with original descriptions of butterflies and especially of
moths (rich in Catocala and Geometridae). Most of the important lepidopterists of
the period were contributors, e.g., John A. Comstock, William Beutenmueller,
Samuel E. Cassino, L. W. Swett and many others whose proposed taxa must be
dated correctly. Apparently there is no flaw in validity of publication, either with the
“Official” bulletins or with Cassino’s separate project, that is, with reference to
Article 8 of the International Code. Both papers were printed and distributed as
required, mailed to subscribers as well as being advertised for sale separately —
starting out at 35<P annually or 4C per issue!

20(2): 111-122, 1981(82)

113

The Lepidopterist 1916-17

(Rudolph C. B. Bartsch, Editor)

VoL I

(1)

Nov. 15, 1916

Pages

1-8

PL [I]

VoLI

(2)

Dec. 15, 1916

9-18

PL II colored

VoL I

(3)

Jan. 15, 1917

19-36

PL in colored

VoL I

(4)

Feb. 15, 1917

27-34

VoL I

(5)

Mar. 15, 1917

35-42

VoLI

(6)

May 1, 1917

43-50

VoL I

(7)

May 15, 1917

51-58

VoLI

(8)

June 1, 1917

59-66

PL IV

VoLI

(9)

June 15, 1917

67-74

PL V

VoLI

(10)

July 15, 1917

75-82

PL VI

VoL I

(11)

(Samuel E. Cassino, Publisher)

Aug. 15, 1917 83-90 PL VII

VoL I

(12)

Nov. 15, 1917

91-98

VoL I

(13)

Dec. 15, 1917

99-106

PL [Vin] colo]

PP=

Lepidoptera 1918-21

(Robert Swift and Nathaniel Stowers, Editors)

Voi.n

(1)

Jan. 15, 1918

Pages

107-114

[sic] [1-8]

VoL n

(2)

Feb. 15, 1918

9-16

VoL n

(3)

Mar. 15, 1918

17-24

voi.n

(4)

Apr. 15, 1918

25-32

VoL n

(5)

May 15, 1918

33-40

PL I

VoL II

(6)

June 15, 1918

41-48

VoL II

(7)

July 15, 1918

49-56

Voi.n

(8)

Aug. 15, 1918

57-64

VoL n

(9)

Sept. 15, 1918

65-72

Voi.n

(10)

Oct. 15, 1918

73-80

VoL n

(11)

Nov. 15, 1918

81-88

VoL n

(12)

Dec. 15, 1918

89-96

Index 4 pp.

VoL m

(1)

Jan. 15, 1919

1-8

voi.ni

(2)

Feb. 15, 1919

9-16

Directory 2 pp.

VoL m

(3)

Mar. 15, 1919

17-24

voi.m

(4)

Apr. 15, 1919

25-32

VoL m

(5)

May 15, 1919

33-40

Directory 3 pp.

VoL m

(6)

June 15, 1919

41-48

PL I

VoL ni

(7)

July 15, 1919

49-56

PL 11

VoL m

(8)

Aug. 15, 1919

57-64

PL in

VoL m

(9)

Sept. 15, 1919

65-72

PL IV

VoL m

(10)

Oct. 15, 1919

73-80

Voi.m

(11)

Nov. 15, 1919

81-88

PL V

VoL ni

(12)

Dec. 15, 1919

89-96

Index 4 pp.

J. Res. Lepid.

114

Vol. IV

(1)

Jan. 1920

1-8

PI. I cover

Vol. IV

(2)

Feb. 1920

9-16

PL n cover

Vol. IV

(3)

Mar. 1920

17-24

cover

Vol. IV

(4)

Apr. 1920

25-32

cover

Vol. IV

(5)

May 1920

33-40

PI. EH, IV cover

Vol. IV

(6)

June 1920

41-48

PL V cover

Vol. IV

(7)

July 1920

49-56

cover

Vol. IV

(8)

Aug. 1920

57-64

cover

Vol. IV

(9)

Sept. 1920

65-72

cover

Vol. IV

(10)

Oct. 1920

73-80

cover

Vol. IV

(11)

Nov. 1920

81-88

cover

Vol. IV

(12)

Dec. 1920

89-96

Index, 3 pp. cover

Vol. V

(1)

March 1921

1-12

Vol. V

(2)

Sept. 1921

13-24

Information supplied by Dr. A. E. Brower, of Augusta, Maine, indicates that at the
Sixth Annual Meeting of the Boston Entomological Club the following arrangement
was made concerning Lepidoptera:

“The Magazine Lepidoptera is discontinued. Henceforth the Official Bulletin of
the Boston Entomological Club will be ‘The Clansman’ which is published every
month by Mr. Harry G. Haskell, of Hyde Park, Mass.”

The last entry Dr. Brower found in The Clansman concerning the activities of the
Boston Entomological Club was a few miscellaneous notes on insect collectors in
volume 1, numbers 5-6, July-August 1922, on page one. Sic Transit...

The Lepidopterist 1918-31

(Samuel E. Cassino, Publisher)

Pages

Vol. n

(1)

Jan. 15, 1918

1-8

PL [1] colored

Vol. n

(2)

Feb. 25, 1918

9-16

PL 3, 4 PL [2| colored

Vol. n

(3)

Mar. 25, 1918

17-24

Vol. n

(4)

Apr. 25, 1918

25-32

PL “6” [5] colored

Vol. n

(5)

May 25, 1918

33-40

PL 6 [sic]

Vol. n

(6)

June 25, 1918

41-48

PL 7, 8 colored

Voi.n

(7)

July 25, 1918

49-56

PL 9

Vol. n

(8)

Aug. 25, 1918

57-64

Vol. n

(9&10)Oct. 25, 1918

65-80

PL [10] [11]

voi.n

(11 &

Dec. 25, 1918

81-96

12)

Voi.m

(1)

Mar. 15, 1919

97-104

Voi.m

(2)

July 15, 1919

105-112

Vol. m

(3)

Jan. 1, 1920

113-120

Voi.m

(4 & 5) Feb. 16, 1920

121-136

Vol. m

(6 & 7) Feb. 15, 1922

137-150

VoL m

(8)

Mar. 15, 1922

151-158

Voi.m

(9)

Apr. 15, 1922

159-166

PL [12]

Vol. m

(10)

Aug. 15, 1922

167-174

Voi.m

(11)

Sept, 15, 1922

175-182

20(2): 111-122, 1981(82)

VoL m

(12)

Nov. 1, 1922

183-190

Vol.IV

(1)

Apr. 1, 1923

1-8

VoL IV

(2)

June 1, 1923

9-16

Vol.IV

(3)

Dec. 25, 1923

17-24

VoL IV

(4)

Jan. 1, 1924

25-32

Vol.IV

(5)

May 10, 1925

33-40

VoL IV

(6 & 7) June 10, 1925

41-56

Vol.IV

(8 & 9) Feb. 25, 1927

57-72

VoL IV

(10)

Apr. 15, 1927

73-80

voi.rv

(11)

June 1, 1927

81-88

VoL IV

(12)

Mar. 15, 1928

89-96

VoL V

(1)

Aug. 1, 1928

1-8

VoL V

(2)

May 1, 1931

9-16

VoL V

(3)

June 15, 1931

17-24

[corrected in ink]

115

Commencing with volume 2, number 5, Cassino, together with Louis W. Swift,
were described as editors, but with volume 3, number 11, “Editors” was dropped
and thenceforth Cassino was designated as publisher.

There are many line drawings throughout this journal. So far as is known, no
indexes to the volumes were published and no covers have been seen.

The letter previously mentioned, from Sherman, the book dealer, indicated that
even in 1939, when the Boston Entomological Club publications and Cassino’s
bulletin both had not been long out of existence, they were rare items and hard to
come by. The set in the American Museum of Natural History library is believed to
be complete for both.

Some Little-Known U. S. Publications on

Lepidoptera II^

Cyril F. dos Passes

Research Associate, Department of Entomology, American Museum of Natural History,
New York, New York 10024

Abstract. Notes and collations are given for the following publications
which are of historical interest to lepidopterists: The Sierra Club Bulletin
1913; The Butterfly Farmer 1913-14; Lorquinia 1916-19; Southwest Science
Bulletin 1920; Butterfly Park Nature Club News 1929-31; The Lepidopterists'
News 1933 (not to be confused with any publication of the present
Lepidopterists’ Society); Hobbies — The Magazine for Collectors 1936, also
The Entomologists’ Exchange Association 1936, also The Entomologists’
Exchange News 1937-42; The Butterfly Club 1946-47; Club Notes, Moth and
Butterfly Club [?1947]"53, also Notes on Moths and Butterflies 1953-55.

116

J. Res. Lepid.

Sierra Club Bulletin 1913

While decidedly not a poorly known publication, it does not seem to be widely
known that volume 9, number 2 of the Sierra Club Bulletin contains important
papers on Lepidoptera by Vernon L. Kellogg and Fordyce Grinnell, Jr. This number
of the Bulletin is publication no. 48, June 1913, pp. 81-128. It deserves a place in
entomological libraries.

The Butterfly Farmer 1913-14

The Butterfly Farmer, a monthly magazine for amateur entomologists, was
published by Miss Ximena McGlashan, of Truckee, California, commencing with
the September number in 1913 and ending with the August number in 1914. This
publication included, as a leading feature in each number, parts of a comprehensive
correspondence course in entomology conducted under the auspices of The Agassiz
Association, and also included much current news in the entomological world,
especially that part thereof devoted to butterflies and moths. It was copyrighted by
Miss McGlashan. It may be collated as follows:

Vol. 1

(1)

Sept. 1913

Pages

1-16

Vol. 1

(2)

Oct. 1913

17-32

Vol. 1

(3)

Nov. 1913

33-48

Vol. 1

(4)

Dec. 1913

49-64

Vol. 1

(5)

Jan, 1914

65-80

Vol. 1

(6)

Feb. 1914

81-96

Vol. 1

(7)

Mar. 1914

97-112

Vol. 1

(8)

April 1914

113-132

Vol. 1

(9)

May 1914

133-148

Vol. 1

(10)

June 1914

149-168

Vol. 1

(11)

July 1914

169-184

Vol. 1

(12)

Aug. 1914

185-208 including index

This was a useful little publication and still is of interest, particularly for the
biographies and pictures of old-time entomologists. With the second number, C. F.
McGlashan, the father of Ximena, was listed on the cover as associate editor and
business manager, and on page 67 their picture, taken together, was given. Always
on the cover the picture of Miss McGlashan appeared. The subscription price was
$5.00 per year, single copies 60<t.

In addition to The Butterfly Farmer, Miss McGlashan published two large
foodplant lists for the benefit of her students, one based on 'HoW&ndLB Butterfly Book
and the other on the same author’s Moth Book. The foodplant list for butterflies was
Circular No. 6, March 1913, and measured approximately 0.48 m by 0.70 m. The
foodplant list for moths was Circular No. 8, May 1913, measuring approximately
0.48m by 0.78 m. The other circulars have not been seen.

Lorquinia 1916-19 ^

Lorquinia was published by the Lorquin Natural History Club, of Los Angeles,
California, between August 1916 and January 1919. Its officers when the
publication commenced were: Luther Little, President; Rutherford D, Moore,
Vice-President; F. Grinnell, Jr., Secretaiy; and E. P. Chace, Treasurer. The editor

20(2): 111-122, 1981(82)

117

was Paul D. R, Ruthling, He was succeeded by George L. Moxley. At that time the
Club had 20 active, 3 honorary and 10 charter members., Lorguima published many
botanical and entomological papers and notes. It collates as follows:

Vol. 1

(1)

Aug. 1916

Pages

1-8

Vol. 1

(2)

Sept. 1916

9-11

Vol. 1

(3)

Oct. 1916

17-24

Vol. 1

(4)

Nov. 1916

25-32

Vol. 1

(5)

Dec. 1916

33-40

Vol. 1

(6)

Jan. 1917

41-48

Vol. 1

(7)

Feb. 1917

49-56

Vol. 1

(8)

Mar. 1917

57-64

Vol. 1

(9)

Apr. 1917

65-72

Vol. 1

(10)

May 1917

73-80

Vol. 1

(11)

June 1917

81-88

Vol. 1

(12)

July 1917

89-96

Vol. 2

(1)

Aug. 1917

1-8

Vol. 1

(2)

Sept. 1917

9-16

Vol. 2

(3)

Oct. 1917

17-24

Vol. 2

(4)

Nov. 1917

25-32

Vol. 2

(5)

Dec. 1917

33-40

Vol 2

(6)

Jan. 1919

1-12

1+ “4” (12), “5” (13), 14, 15,
“18” (16)]

The Lorquin Natural History Club was affiliated with the Southwest Museum and
quarterly issues of Lorquinia were contemplated. A set of these papers obtained
from the library of Fordyce Grinnell, Jr., has a note in pencil on the first page of the
January 1919 issue, viz, “Last issue.” There is no index to any volume nor have any
covers been seen. Mrs. Dorothy E. Martin, former Librarian of the Los Angeles
County Museum, has confirmed also that volume 2, number 6 was the last number
issued. The organization name was changed to the Lorquin Entomological Society,
which is still quite active in Los Angeles with over 100 members. The society has
published a newsletter (10 issues per year, unnumbered) since about 1956.

Southwest Science Bulletin 1920

The Southwest Science Bulletin was published by the Southwest Museum under
the supervision of the Council of the Southwest Science Association, Marmon Way
and Avenue 46, Los Angeles, California. Only one copy has been seen, which was
issued May 5, 1920 and consists of 32 pages, one colored plate and four black-and-
white plates. It contains articles on butterflies and moths. Nothing further has been
learned about this publication. Mrs. Martin kindly confirmed the fact that only one
number ever was issued.

Butterfly Park Nature Club News 1929-3 1

Butterfly Park Nature Club News commenced publication in March 1929. It was
printed at Roscoe, California. Albert Carter appears to have been the leading spirit.
He had the publication copyrighted. Mr. William D. Field, formerly of the National
Museum of Natural History, supplied the following collation:

118

J. Res. Lepid.

Pages

Vol. 1

(1)

Mar. 1929

4

Vol. 1

(2)

Apr. 1929

4

Vol. 1

(3)

May 1929

4

Vol. 1

(4)

June 1929

4

Vol. 1

(5)

July 1929

4

Vol. 1

(6)

Aug. 1929

4

Vol. 1

(V)

Sept. 1929

4

Vol. 1

(8)

Oct. 1929

4

Vol. 1

(9-10)

Nov. -Dec. 1929

4

Vol. 1

(11-12)

Jan.-Feb. 1930

4

Vol. 2

(1-2)

Mar.-Apr. 1930

4

Vol. 2

(3-4)

May- June 1930

4

Vol. 2

(5)

Oct. 1930

4

Vol. 3

(6)

Jan. 1931

8

Vol. 3

(7)

Apr. 1931

4

Vol. 3

(8)

July 1931

4

Vol. 4

(1)

Jan. 1932

8

Whether anything further was published has not been ascertained.

The Lepidopterists’ News 1933

This publication is not to be confused with any publication of the present
Lepidopterists’ Society, with which it had nothing whatsoever to do.

The Lepidopterists’ News was published by the Florida Society of Lepidopterists
of the Florida Society of Natural History. Only two numbers have been seen, both
mimeographed, edited by Mrs. Elizabeth 0. Groves and J. Harold Matteson, as
follows:

Pages

Vol. 1 (1) June 1932 8

Vol. 1 (2) Dec. 8, 1933 6

Number 1 was letterhead size, number 2 was octavo.

HobbieS“”The Magazine for Collectors 1936

In July 1936 Hobbies ran a department entitled “Natural History” in which Frank
Clay Cross published the “Entomologists’ Exchange Association”. This July issue
is the only one which has been inspected; it was the 9th number and 4 1st year of that
magazine. It catered to many tastes and in 1936 it ran several articles on
Lepidoptera written by Cross. These were:

March 1936 A Hobby on Wings. Vol. 41, no. 1, pp. 112-113, 119
April 1936 Reminiscences of a Butterfly Bungler. Vol. 41, no. 2, pp. 112-113

July 1936 Entomologists’ Exchange Association. Vol. 41, no. 5, pp. 112-
113

Also in the March number of Cross announced (p. 113) organization of the

Entomologists’ Exchange Association, which commenced publication that month.

20(2): 111422, 1981(82)

119

The Entomologists’ Exchange Association 1936
The Entomologists* Exchange News 1937-1942

Information concerning these publications has been supplied by Mr. William D.
Field, formerly of the National Museum of Natural History, it appears that Frank C.
Cross, then of Denver, Colorado, initiated a paper the first numbers of which were
typewritten, viz:

Pages

[VoL Ij [Ij circa March 1936 1

[¥oL 1] [2| circa May 1936 1 2 p. list tentative members

[VoL 1] 3 circa June 1936 1 Bulletin added to title

The second publication was mimeographed and the name was changed to “The
Entomologists’ Exchange News”. The first two numbers have not been seen.

Pages

VoL 2

(3)

July 1, 1937

2

VoL 2

(4)

Aug. 1, 1937

2

Vol. 2

(5)

Sept 1, 1937

6

Voi. 2

(6)

Oct 1, 1937

3

Vol. 2

(7)

Nov. 1, 1937

2

VoL 2

(8)

Dec. 1, 1937

4

VoL 2

(9)

Jan. 1, 1938

3

Vol. 2

(10)

Feb. 1, 1938

2

Vol. 2

(11)

March 1, 1938

2

Vol. 2

(12)

April 1, 1938

2

Vol. 3

(1)

May 1, 1938

3

VoL 3

(24)

June-Aug. 1938

3

VoL 3

(5)

Sept 30, 1938

3

Vol. 3

(6)

Oct 30, 1938

3

Vol. 3

(7)

Nov. 30, 1938

7

Vol. 4

(1)

Jan. 5, 1939

3

VoL 4

(2)

Feb. 4, 1939

6

VoL 4

(34**^*)

April 10, 1939

2

VoL 4

(5*)

May 18, 1939

5

Vol. 4*

(64)

July 25, 1939

5

Vol. 4*

(8)

Sept 5, 1939

5

VoL 4*

(940)

Nov. 20, 1939

5

Vol. 5*

(1)

Jan. 30, 1940

5

Vol. 6*

(24)

March 5, 1940

6

Vol. 5

(44)

June 5, 1940

8

VoL 5

(74)

Aug. 5, 1940

6

Vol. 5

(941)

Nov. 27, 1940

2

Voi. 6

(14)

Feb. 15, 1941

4

Vol. 6

(3)

March 25, 1941

5

VoL 6

(4)

April 17, 1941

6

Vol. 6

. (54)

Aug. 1941

3

Vol. 6

(940)

Oct 1941

3

Vol. 7

(1)

Jan. 10, 1942

4

120

J. Res. Lepid.

Vol. 7 (2) Feb. 13, 1942 3

Volume 7, number 2 is the last number seen. It is not known whether anything
further was published.

♦Asterisks indicate errors in numbering issues, a plague of which infested these
runs. To wit: for 1939: April 10, number “4 & 5” was 3-4 in proper sequence; May
18 was number 5, not “6”, The July issue was correctly numbered but cited volume
number incorrectly as “3”, continued through the two issues following. In 1940,
beginning volume 5 the first two issues were numbered “vol. 4”. Finally, the June
issue, 1940, got back on track as volume 5, numbers 4-6.

This paper started out and ended on letter-size sheets, with an interim of legal-
size issues from volume 2, number 12 through volume 4, number 1.

Inclusions

Lists of members in volume 2, number 5; volume 4, number 2; volume 6, numbers
1-2 ahd 3 and 9-10. Volume 3, number 7 had a list of “species in the Cheyenne
Mountain Museum”. With volume 3, number 2-4 Robert Potts took over as editor.
Volume 5, number 9-11 contains a statement that F. Martin Brown was editor
before Potts. Brown appears also to have been treasurer, at least from volume 2,
number 12 on. A field collecting form was included in volume 4, number “4 & 5” (3 &
4]. Beginning with the issue of July 25, 1939, the News was published by the
Colorado Biological Survey and the Cheyenne Mountain Museum. A “preliminary
list of the butterflies and skippers of Santa Clara and Santa Cruz” was featured in
volume 6, number 4.

The Butterfly Club 1946-1947

These were mimeographed letters, written and distributed by Mrs. Evelyn G.
Williams on behalf of the Howell Mountain Butterfly Club, for a period of about one
year, from her residence in Sanitarium, California. Acquisition of Xeroxed copies
which were kindly provided by Mrs. Bernice Neimi, of Aberdeen, Washington, has
made possible a collation, not given here in full because so easily summarized,
viz: The issues ran in sequence, monthly, from volume 1, number 3, September
1946, through volume 1, number 11, July 1947. Then, volume 2, number 1, dated
September 1947, which is the last number. These were all short papers (Nov.-
March, 1 page; April-Sept., 2 pp). Numbers 1 and 2 of volume 1 have not been seen
although doubtless they exist, as also the missing number 12. It is not known how
many numbers, if any, were published after volume 2, number 1. Perhaps somebody
knows and will enlighten us?

Mr. Dominick J. Pirone, of Yonkers, New York, called attention to this bulletin
and suggested that Mrs. Neimi might have copies.

It appears that Mrs. Williams also wrote a booklet entitled “Starting Your
Butterfly Collection” which she sold for IOC a copy, later increased to 25C. This
booklet has not been seen.

Club Notes, Moth and Butterfly Club [? 1947/ 1953]

Notes on Moths and Butterflies 1953-1955

This serial was published by the Moth and Butterfly Club under slightly different
titles. In 1953 James M. Unseld, Jr. was the editor. Publication apparently ceased
in 1955.

20(2): 111-122, 1981(82)

121

No issues prior to January 1953, volume 7, number 1, have been located, so it is
impossible to say when publication began. After writing to all of the Club members
whose names could be ascertained, an answer finally came from Mr. Richard T.
Arbogast, Savannah, Georgia, that he had volume 7 to 9 almost complete. Later, he
kindly furnished Xeroxed copies of the numbers which were lacking in the dos
Passos library, and also volunteered leads to run down in the effort to obtain
volumes 1 to 6, which were pursued but unfortunately without success.

With the June 1953 issue, “Club” was dropped from the title, Unseld stepped
down as editor and was succeeded by Clifford Verhoeff, of Bristol, Colorado.
Unseld’s predecessor, if any, is not known. Whether volume VIII, numbers 5 and 6
were issued also remains in question; they have not been seen.

Volumes 7 and 8 have been collated as follows:

Pages

Voi. “vni”

(1) [Vol. VII]

Jan. 1953

4

voi. “vni”

(2) [Vol. VII]

Feb. 1953

3

voi. “vni”

(3) [Vol. VII]

March 1953

3

voi. “vni”

(4) [Vol. VII]

April 1953

6

Voi. “vni”

(5) [Vol. VII]

May 1953

3

Vol. VII

(6)

June 1953

2

Voi. VII

(“6”) [7]

July 1953

2

Vol. VII

(8)

Aug. 1953

2

Vol. VII

(9)

Sept. 1953

2

Vol. VII

(10)

Oct. 1953

3

Vol. VII

(11)

Nov. 1953

4

Vol. VII

(12)

Dec. 1953

3

Vol. vm

(1)

Jan.-Feb. 1954

5

Vol. vm

(2)

March 1954

3

Vol. VIII

(3)

April 1954

3

Vol. vm

(4)

June 1954

3

Vol. vm

(“7 & 8”) [5 & 6]

July-Aug. 1954

4

Vol. vm

(“9 & 10”) [7 & 8]

Sept.-Oct. 1954

2

Vol. vm

([11]&(12)

Nov. -Dec. 1954

2

Vol. vm

(1 [& 2]) [Vol. IX?]

Jan.-Feb. 1955

1

Vol. vm

(3 & 4)

March-Apr. 1955

2

Inclusions: January 1953 list of 42 members; January-February 1954 list of 35
members.

A note regarding this Club occurs in the 1953 Annals of the Entomological Society
of America (46: 220). It was by “M. T. J.” [Maurice T. James, the managing editor}
as follows: “According to James M. Unseld, Jr., Gravel Switch, Kentucky, the
Moth and Butterfly Club is an organization open to anyone interested in the study of
Lepidoptera. At present there are fifty-five members. Annual dues are $1.50. The
Club Notes are issued monthly in mimeographed form. Mr. Unseld is the editor.
The April 1953 issue is indicated as Volume VIII, number 4,”

Acknowledgments. Generous assistance was provided by many individuals in the
course of assembling foregoing data. Special thanks go to Mrs. Dorothy E. Martin,
former Librarian of the Los Angeles County Museum, and to Miss Nina Root,
Librarian of The American Museum of Natural History. Mr. Richard T. Arbogast, of

122

J. Res. Lepid.

Savannah, Georgia, aided considerably as noted earlier. Inquiries made during
preparation of the collations were sent to nearly 100 individuals, many of whom
responded helpfully. Unfortunately, some sections are incomplete, but they would
have been far more so without the cooperation of many individuals, especially Mr,
William D. Field, of Washington, D. C.

Quite surely the present treatment could be made more definitive and it is to be
hoped that further information, if available, will be published. It is especially
desirable to consolidate the histories of earlier club and society bulletins now that a
veritable spate of such have begun to come out in recent years.

Errata

In my article “Description and taxonomic implications of an unusual Arizona
population of Apodemia mormo (Riodinidae)” (J. Res. Lepid. 78(3): 201-207), two
errors have been noted which require correction:

p. 204: the plate has been printed backwards so that label information is
transposed for the specimens at each end of a row.
p. 205, line 5: sympatric should read allopatric.

Gregory S. Forbes, Box 3AF, Dept, of Biology, New Mexico State University,
Las Cruces, NM 88003

In the issue “A Revision of the American Genus Anisota” {J. Res. Lepid. 19(3)),
three errors have been noted which require correction:

p. 133: Fifth instar: Body color hlackish-hrowa (not he^^e-brown) as the figure
also shows.

p. 134: Boisduval should be in brackets.

p. 175: Plate Vn. 9. should read: A. pellucida x virginiensis.

Journal of Research on the Lepidoptera

20(2): 123-126, 1981(82)

A New Species of Adelpha (Nymphalidae) from Parqiie
Nacional Braulio Carrillo, Costa Rica

Philip J. DeVries
and

Isidro Chacon Gamboa

Department of Zoology, University of Texas, Austin, Texas 78712 and Museo Nacional,

Apartado 749, San Jose, Costa Rica

Abstract. The new butterfly species Adelpha stilesiana (Nymphalidae) is
described from Costa Rica. This new species apparently represents another
Costa Rican butterfly endemic to the Carrillo Belt. A few rudimentary field
observations are also presented.

In Costa Rica, there are 26 species in the gemis Adelpha (Nymphalidae)
that fall roughly into two groups; the typical white banded species, and a
smaller group of species that are brown dorsally with a broad orange band
across the forewings and without white on either wing. In this latter group
there are four species recorded from Costa Rica: Adelpha melanthe
Bates, A. zalmona sophax Godman & Salvin, A. boreas tizona Felder, and
A. salmoneus salmonides Hall. The interested reader is referred to either
Fmhstorfer (1910-1912) or Godman & Salvin (1879-1901) where these
species are illustrated. During the course of a long term study on the
butterfly fauna of Parque Nacional Braulio Carrillo, a fifth and undes-
cribed species of the orange-banded group has come to our attention. This
species is not found in any world museum studied by one of us (P JD), and it
is quite distinct from its congeners. We here describe this new species of
Adelpha and give some general notes on its distribution and behavior in the
field.

Adelpha stOesiana DeVries & Chacon new species

Male: Eyes. Sparsely hairy except for a dense pathc of hairs on the dorsum of
the eyes near the base of the antenae. Antennae. Dark brown and naked except
for a small patch of white scales on the ventrum near the base.

FW upperside: Dark brown from base to distal end of cell; submarginal area
with a wide orange-yellow band which has a somewhat jagged inner and outer
margin; the band runs from costa at Ri-Ri and extends broadly to distal wing margin
at Mato 2 A. HW upperside: Dark brown as in the FW but bears a faint subcostal
line of lighter brown. FW underside: Similar to A. salmoneus and A. zalmona but
differing distinctly as follows: cell area greyish-violet bearing two wide rufous

124

J. Res. Lepid.

bands, one in mid-cell, the other at the end of the cell; both bands are thinly
bordered by dark brown; are distad of cell end washed with pale ochre, being
brightest on distal margin between M2-2A; a series of six prominent whitish
postmedial spots between veins R3-R4, R4"Mi, Mi"M2, M2-M3, Ms-Cui, and Cui-Cu2;
a series of small spots run distad to those just described but beginning in Rs-Mi,
running posteriorly between all veins and terminating in a double spot in cell Cu2
near the tornus. HW undersides Similar to that of A. salmoneus but with a dingy
violet overcast to the ground color; a dark brown postmedial line runs from the
costal margin, inflating in the discal area, and terminating in the anal angle as a
rufous patch; marginal lobes of wing with spots of ochre centered on veins Mi, M2,
Cui, and Cu2; fringe dark brown.

Genitalia: See figure (3) for details.

Female: The solitary female is virtually identical in pattern to the male.

Length of FW: male: 32 mm; female: 34 mm.

TVpes: Holotype: male, Costa Rica, Provincia de San Jose, Parque Nacional
Braulio Carrillo, Estacion La Montura, May 24, 1981, 1100 m altitude, leg. Ruben
Canet M.. Paratype: female, Costa Rica, San Jose, Carrillo, La Montura, 1100 m,
June 11, 1980, leg. F. G. Stiles. Both in the British Museum (Natural History).

Diagnosis. See figures 1 and 2 for details. This species can be distinguished
from all other Costa Rican congeners by the wide orange forewing band that ends
broadly along the distal margin. This character can be used to distinguish this
species in the field.

Etymology; We name this species for F. Gary Stiles who, incredibly enough, let
us talk him into carrying a butterfly net in addition to his bird collecting gear and
who successfully mist-netted the first specimen.

Discussion

Adelpha stilesiana is known only from two specimens (the types) and
collected in the same locality. This locality is part of the Carrillo Belt (see
DeVries, 1980) which is known for its unusual faunal and floral charac-
teristics. The Carrillo Belt harbors a number of endemic Costa Rican

Fig. 1. Dorsal aspect of holotype.

Fig. 2. Ventral aspect of holotype.

20(2): 123-126, 1981(82)

126

butterflies and is also the habitat where some South American species
terminate their northern range (e. g. Morpho granademis Felder, Heli-
conius ekuchia Hewitson, Eunica nonca Hewitson, and Epiphile eriopk
Hewitson). Due to the total absence of specimens of A. stilesiana in
the British Museum (Natural History), Smithsonian Institution, Carnegie
Museum, Allyn Museum of Entomology, and numerous private collections
viewed by us, we suggest that it is a species endemic to the Carrillo Belt.

From our work in Parque Carrillo, we have observed a number of
individuals of A. stilesiaria from 1100 to 800 meters elevation. Individuals
perch on foliage along the insides of ravines or in the forest canopy from
0800 to about 1100 hours. As most other Adelphcf species in Costa Rica,
individuals perch from 5 to 10 minutes at the same spot and make sorties
out from the perch, returning repeatedly to the same spot. They then move
to a different area that is usually 50 or more meters away.

On the wing, A. stilesiana looks very similar to A. salmoneus and A.
zalmona sophax which also occur in the same habitat. Positive field
identification of A. stilesiana is easy when the insect is perched. The
distinctive ochreaceous forewing band can be seen with the naked eye or
with binoculars. We have seen individuals of this species during May, June
and July.

Acknowledgments: We thank Ruben Canet for supplying us with the holot3?pe
and F. G. Stiles for the paratype and the loan of his jeep. A special thanks to R.
Mattoni for illustrating the genitalia and two anonymous reviewers. This study was
supported in part from a grgint to DeVries by the Society of the Sigma Xi and from
the University of Texas. Field assistance came from the Museo Nacional, Servicioe

0.5 mm

Fig. 3. Genitalia top to bottom: a. Aedeagus dorsal view, b. Ibid, lateral view, c.
Lateral view entire genitaMa, left valve removed. J ^ Juxta, tJ ~ uncus.

Scale: 0.5 mm.

126

J. Res. Lepid.

de Parques Nacionales, and L. E. Gilbert. This paper is dedicated to the memory of
Bill Evans and T. Monk.

Resumen: Se describe una especie nueva del genero Adelpha (Nymphalidae) del
Parque Nacional Braulio Carrillo, Costa Rica. La especie es conocida por solamente
dos ejemplares. Se destinga A. stilesiana del A. zalmona sophax, A. melanthe, y A.
solmoneus por la banda muy ancha en la ala primaria. Los congeneros de A.
stilesiana estan iUustradas en Fruhstorfer (1910-1912) y Godman & Salvin (1879-
1901). Se comunican algunas observaciones sobre la distribucion y la conducta en
su habitat endemica en Carrillo.

literature Cited

DeVRiES, P. J., 1980. Description, natural history, and distribution of a new species
of Eretris (Satyridae) from Costa Rica. J. Lep. Soc. 34: 146-151.
FRUHSTORFER, H., 1910-1912. In: A. Seitz, ed.Macrolepidopteraofthe World, vol. 5.
Alfred Keman, Stuttgart.

GODMAN F. D. & O. SALVIN, 1879-1901. Biologia Centrali- Americana. Lepidoptera:
Rhopalocera, vol. 1, 1879-1886, I-XLV & pp. 1-487; Vol. 2, 1887-1901, pp.
1-782.

Journal of Research on the Lepidoptera

20(2): 127-128, 1981(82)

Japanese Literature

The following cited literature is the beginning of a regular listing of
relevant papers published in Japan (and usually Japanese) for the
convenience of our readership in having available information frequently
not readily accessible. Authors’ addresses are given for purposes of
reprint requests.

“Butterflies and Moths ”(Tyo to Ga), volume 32 (1/2), published September 20, 1981.

MASANAO NAKAMURA. Key to the classification of the Japanese iepidopterous pupae,
pp. 1-12, in English with a Japanese summary.

Pupae of 75 families comprising 20 superfamilies are keyed and grouped into
6 sections based on external morphology.

KODO MAEKI. The chromosome of the Lepidoptera. pp. 13-28, iUusts., in Japanese
with an extensive English synopsis.

Reviews the author’s evidence for holokinetic, rather than monokinetic, organi-
zation of lepidopteran chromosomes.

MAYUMI TAKAHASHI & JUNZO AOYAMA. On Neope niphonica Butler (Lepidoptera:
Satyridae) in the Boso Peninsula, Central Japan, with description of a new sub-
species. I pp. 29-47, illusts., in Japanese with the description and a summary in
English.

Distribution and morphology of N. niphonica kiyosumiensis ssp. nov. are
described. Formal description appears on p. 46.

MAYUMI TAKAHASHI. A record of Tirumala limniace Umniace Cramer (Danaidae)
from Shizuoka City, Central Japan, p. 48, illust., in Japanese.

An additional record of this rare stray from the Oriental tropics.

TAKASHI SHIROZU & OSAMU YATA. Ten new subspecies of the genus Eurema (Lepidop-
tera, Pieridae) from the Indo- Australian Region, pp. 49-62, illusts., in English
with a Japanese summary.

Describes E. brigitta papuana (NE Papua New Guinea), E. andersoni inouei
(Cambodia), E. andersoni bomeensis (Sarawak), E. andersoni nishiyamai (Nias),
E. andersoni kashiwaii (Sumba), E. sarilata luzonensis (Luzon), E. sarilata
bazilana (Bazilan),E. alithagunjii (Ceram), E. alitha halmaherana (Halmahera),
and E. alitha papuana (W. Irian), sspp. nov. Four forms previously associated
with E. hecabe (bidens Butler, sankapura Fruhst., chemys Frahst., and jalendra
Fruhst.) are given new status as sspp. of E. alitha.

HISAKAZU HAYASHI. New lycaenid butterflies from the Philippines, pp. 63-82,
illusts., in English.

Describes Deramas treadawayi (Mindanao), D. kawazoei (Marinduque), D.
philippinensis (Marinduque), D. toshikoae (Leyte), Narathura schroederi,
(Palawan), N. cleander sugimotoi (Mindanao and Leyte), N. philippina (Min-
danao), N. nishiyamai (Mindanao and Leyte), N. hollow ayi (Mindanao, Leyte
and Samar), N. pseudovihara (Mindanao), N. sakaguchii (Negros), N. aronya
natsumiae (Negros), N. staudingeri negrosiana (negros), N. alesia mio (Negros),
Flos setsuroi (Marinduque), Horaga albimacula katoi (Marinduque), Pratapa
tyotaroi (Marinduque), Neocheritra kurosawai (Mindanao), Chliaria shirozui

128

J. Res. Lepid.

(Mindanao), and Sinthusa mindanensis yoshiae (Negros), spp. nov. and sspp.
nov.

SIGERU MORIUCHI. A new KessleHa (Lepidoptera: Yponomeutidae) from New
Guinea, pp. 83-84, illusts., in English with a Japanese summary.

Describes K. neuguineae sp. nov. from Wareng, New Guinea, the first sp. of
the genus reported from the Island.

RIKIO SATO. Notes on Buzura (Amraica) recursaria (Walker) and its allies from Japan
and adjacent countries, with description of a new subspecies (Lepdioptera:
Geometridae). pp. 85-93, illusts., in English with a Japanese summary.
Defines B. recursaria (Walker), gives B. superans superans (Butler), B. superans
confusa (Stdgr.), and B. asahinai (Inoue) stat. nov., and describes from
Formosa B. superans taiwana, ssp. nov,

MAYUMI TAKAHASHI. Inter-specific crossing between Mycalesis gotama fulginia
Fruhstorfer and M. madjicosa amamiana Fujioka. pp. 94-100, illusts., in
Japanese with an English summary.

Reports the results of reciprocal crosses in captivity (Satyridae).

MAYUMI TAKAHASHI. A gynandromorph of Pieris (Artogeia) melete (Menetries). p.
100, illust., in Japanese.

KENJI KISANUKI. Rearing records of the jacintha- form individuals of the Great Egg-
fly, Hypolimnas bolina L. pp. 101-107, illusts., in Japanese with an English
summary.

Reports the phenotypes, either f. jacintha or f. kezia, of the offspring of f.
jacintha females caught in the wild,

MAYUMI TAKAHASHI. A list of the butterflies of Haeterinae and Biinae (Lepidoptera:
Satyridae) collected by two Japanese expeditions in Colombia and Peru,
South America, pp. 108-116, illusts., in English with a Japanese summary.
Thirteen spp. and sspp. are listed with a note on an undescribed form of Pierella
hortona Hewitson from South Colombia.

Authors’ addresses:

Junzo Aoyama

Hisakazu Hayashi

Kenji Hisanuki
Kodo Maeki

Sigeru Moriuchi

Masanao Nakamura
Rikio Sato
Takashi Shirozu

Mayumi Takahashi

Osamu Yata

I- 137 Nishinota, Shioya-cho, Tarumi-ku, Kobe-shi, 655
Japan

9-1 Tamatsukuri-honmachi, Tennoji-ku, Osaka-shi, 543
Japan

707 Ogata, Komae-shi, Tokyo, 201 Japan

Biological Laboratory, Kanseigakuin University, Nishino-

miya-shi, Hyogo-ken, Japan

Entomological Laboratory, College of Agriculture, Univer-
sity of Osaka Prefecture, Sakai-shi, Osaka, 591 Japan
14-12 Miyamae 3-chome, Suginami-ku, Tokyo, 168 Japan
472-2 Makio, Niigata-ken, 950-21 Japan
Biological Laboratory, College of General Education,
Kyushu University, Ropponmatsu, Fukuoka, 810 Japan

II- 13 Kita-Ando 5-chome, Shizuoka-shi, Shizuoka-ken,
420 Japan

Biological Laboratory, College of General Education,
Kyushu University, Ropponmatsu, Fukuoka, 810 Japan

Ichiro Nakamura, compiler, 41 Sunrise Blvd,, Williamsville, New York 14221

INSTRUCTIONS TO AUTHORS

Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
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arabic numeral). Numerals must be used before measurements (5mm) or otherwise up to
number ten e.g. (nine butterflies, 12 moths).

Title Page: All papers must have the title , author’s name, author’s address, and any
titular reference and institutional approval reference, all on a separate title page. A
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comments of reviewers.

Volume 20

Number 2

Summer, 1981(82)

m THIS ISSUE

Date of Publication: September 20, 1982

International Nepal Himalaya Expedition for Lepidoptera

Palaearctica (INHELP) 1977, Report No. 1: Introduction

and Lycaenidae

Oakley Shields 65

hitergradation between Callophrys dumetorum oregonensis

and Callophrys dumetorum affinis in Northwestern U. S.
(Lycaenidae)

James A. Scott & John A. Justice 81

Chromosome Studies in Sixteen Species of Indian Pyralid
Moths (Pyralidae)

P. K. Mohanty & B. Nayak 86

The Biological and Systematic Significance of Red Fecal and
Meconial Pigments in Butterflies: A Review with Special
Reference to the Pieridae

Arthur M, Shapiro 97

On the Nomenclature of Colias alfacariensis Berger 1948
(Lepidoptera: Pieridae)

Otakar Kudma 103

Some Little-Known U. S. Publications on Lepidoptera I

Cyril F. dos Passes 111

Errata 122

A New Species of Adelpha (Nymphalidae) from Parque
Nacional Braulio Carrillo, Costa Rica

Philip J. DeVries & Isidro Chacon Gamboa 123

Japanese Literature 127

Cover Illustration: The Marsyandi River Valley of Manang, Nepal. See O.
Shields’ article, page 71, this issue.

Volume 20

Number 3

Autumn, 1981(83)

THE JOURHA.L OF RESEARCH
OH THE LEF« DORTER A

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Journal of Research on the Lepidoptera

20(3):129-132 1981(83)

Role of an Ornamental Plant Species in Extending the
Breeding Range of a Tropical Skipper to Subtropical
Southern Texas (Hesperiidae)

Raymond W. Neck

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

Human activities have had manifold effects upon populations of various
species of lepidoptera. The southern tip of Texas is largely ranching and
agricultural with little acreage remaining in a natural state (see Foscue,
1932, 1934). Many agricultural pests affect the growth of these agricultural
crops. Several lepidopteran species have apparently increased their
natural range as a result of ornamental plants suitable as larval foodplants
now occurring in this southern Texas area. Previously, native plants of the
proper family or genera were not available.

The blood spot skipper, Phocides palemon lilea (Reakirt), was reported
to occur “as a straggler in southern Texas to Arizona” by Holland in the
second edition (193 1) of The Butterfly Book. No mention was made of this
species in his first edition (1898), probably due to lack of collectors along
the southern border of the United States at this early date. However,
Scudder (1872: 68) had described Erycides sanguinea from “Texas (Capt.
Pope, Mexican Boundary Survey).” The specimen(s) may have been
collected during the initial boundary survey in the 1850’s. Several early
collectors failed to find this species, e.g. Lintner (1884). Godman and
Salvin (1887-1901: IE, p. 296) listed Texas as a questionable locality for
Dysennius albicilla (now synonymized with lilea, see Miller and Brown,
1981). A group from the University of Kansas spent 6 Jime to 8 July 1905
in the Brownsville area collecting insects but did not find lilea (Snow,
1906) Skinner (1911) listed this species and credited Captain Pope. This
record was repeated without further remarks by Lindsey, et aL, (1931).
Klots (1951) did not mention this species, possibly as a result of the
negative report of Freeman (1949) who was unable to find this species in
field work on both sides of the Rio Grande in the 1930’s and 1940’ s. Nor
did Freeman find specimens in several local collections which he
examined. Freeman (1951) stated that lilea had been “recorded for Texas,
but I have never seen a specimen that actually came from the state.”
Kendall and Freeman (1963) listed this species in their compilation of
Texas butterfly and skipper species. MacNeill (1975:576) stated that this
skipper “has long been reported to occur in Texas, but its occurrence there

130

J. Res. Lepid.

has only recently been verified.” In a recent letter to the author, Freeman
stated “soon after I left the Valley (in 1948). . .collectors began collecting
this species in goodly numbers.” Tilden (1974) reported adults from
Brownsville (Cameron Co.) and Santa Ana Wildlife Refuge (Hidalgo Co.).

Larval Foodplants

Common guava (Myrtaceae: Psidium guayava L.) was reported as a
larval foodplant at Brownsville by Lipes (1961). This same species is the
only recorded foodplant reported for lilea in Mexico (at Puerto Vallarta,
Jalisco, by Comstock and Vazquez, 1961; at or near Cd. Mante,
Tamaulipas, by Kendall and Maguire, 1975). Recently, Neck (1978, 1982)
reported larval utilization of strawberry guava {Psidium cattleianum
Sabine), also in Brownsville. A Brazilian subspecies (phanias Burmeister)
has been reported to feed onPsidium spp. (Miles Moss, 1949) andEugenia
uniflora L. (Biezanko, 1963).

P. guayava is “indigenous to the American tropics but has been
distributed to practically all tropical and subtropical areas throughout the
world” (Ruchle, 1948). This plant occurs throughout much of Mexico but
is native only in the “southern part” (Standley, 1920-1926). Guava has
been grown on both sides of the Rio Grande at least since the late 1800’sas
both Townsend (1897) and Bailey (1916 but from observations during
1900 trip) mentioned this plant. Plantings may have occurred much earlier
as it was spread “early” by the Spanish (Ruchle, 1948). P. cattleianum is
native to Brazil but has long been grown in the United States and Mexico
(Standley, 1920-1926).

The lack of a native foodplant for lilea in Texas is almost assured because
no members of the Myrtaceae are native to any part of Texas (Gould, 1969;
Correll and Johnston, 1970). The four alternate plants (papaya, bougain-
villea, hibiscus and banana) which Lipes (1961) offered to lilea as
oviposition sites (unsuccessfully) are not native to this area, despite his
statement to that effect.

Status of Phocides in Texas

As a result of the lack of a suitable larval foodplant for lilea in the native
flora of the lower Rio Grande Valley and the periodicity of the scarce
records for lilea in this area, I conclude that lilea is not a member of the
native resident skipper fauna of the United States in the strictest sense.
This species has extended its breeding range from (probably southern)
Mexico into southern Texas. Note that Hoffman (1941: 241-242) reported
Ulea only as far north as southern Tamaulipas. Following the introduction
of guava into Texas, establishment of breeding populations of lilea
occurred during the nineteenth century. The paucity of records and
absence during the 1930’s and 1940’s indicate extinction subsequent to
initial establishment followed by re-establishment. These earlier popula-

20(3): 129-132, 1981(83)

131

lions could have perished during particularly cold winters, e.g. 1866, 1886
and 1899.

Discussion

A similar dynamic system of establishment- extinction- establishment
has been observed in Melanis pixe (Boisduval) (Riodinidae) (Neck,
1976b). In the case of Melanis, however, native species related to the
introduced foodplant occur in south Texas (two in the same genus,
Pithecellobium, and numerous species in same family, Leguminosae). M.
pixe is probably a very recent addition to the fauna of south Texas as no
records are known prior to the 1950’s.

The dynamic systems of population establishment of these species are
indicative of the tropical element of the flora and fauna of the lower Rio
Grande Valley. The native tropical element is only a small proportion of
the flora and fauna of this region. Many tropical frms are able to maintain
populations once they are introduced, e.g. snails and slugs (Neck, 1976a).
Exclusion of tropical forms is due to a combination of occasional severe
cold winter weather (e.g. 19°F at Brownsville in January 1962) and
frequent dry periods which are even more significant since the construction
of dams and levees to contain floods of the Rio Grande. In the case of
phytophagous insects which are restricted to certain plants as suitable
foodplants, the presence of suitable plants is also important.

A dispersal of Dione moneta poeyii (Butler) from central Tamaulipas into
south Texas in 1964 was discussed by Gilbert (1969). This species moved
far north of its normal range due to unusual weather conditions.
Permanent populations were not established because of inherent unsuit-
ability of the south Texas environment, d. m. poeyii is normally found in
montane forests. Subsequent reports of this butterfly in south Texas since
1964 are known (R. O. Kendall, pers. comm.). Another exceptional case of
long-distance dispersal was witnessed by Freeman (1959) who observed
but could not collect a Mo/pho near Hidalgo, Hidalgo County, on 25 March
1945 (date given by Stallings and Turner, 1946). Similarly, lilea is
assumed to have dispersed to south Texas during favorable climatic
conditions. Permanent or semi-permanent populations have become
established because a suitable larval foodplant is available and other
significant environmental factors (which are largely unknown) are not
hostile to its continued presence.

Both lilea and M. pixe were found by collectors in the 1950’ s. The
occurrence of these tropical species at about the same time in the lower
Rio Grande Valley of Texas may have resulted from peculiar climatic
conditions of the 1950’ s. This period included the worst drought in Texas
on record (1950-1957) which was accompanied by very warm winter
weather (Orton, et a!., 1967). Although severe freezes occurred at
Brownsville in late January 1949 and 1951, temperatures fell no lower

132

J. Res. Lepid.

than 29°F ("1.7°C) from 1952-1961 inclusive with no freezing tempera-
tures during 1954-1958 inclusive. Although this was a dry period in
general, heavy rains occurred in late June 1954 as a result of Hurricane
Alice which moved inland south of Brownsville. This period of warm
winters and at least occasional sufficient moisture may have been a
significant aid to the establishment of these two species in the Brownsville
area, hi 1967 tremendous rains accompanying Hurrican Beulah apparently
allowed the establishment of several butterfly species previously unknown
to Texas (Kendall, 1970a, 1970b, 1972). Several hurricanes which
affected the Brownsville area in 1933 had varied impacts on the butterfly
fauna of south and central Texas (Neck, 1977).

Literature Cited

BAILEY, F. M., 1916. Meeting spring halfway. Condor 18: 151-155, 183-190, 214-219.
BIEZANKO, C.M., 1963. HesperUdae da Zona suesta do Rio Grande do Sur. Pelotas,
OFIGRAF, Brazil.

CORRELL, D. s. & M. c. JOHNSTON, 1970. Manual of the vascular plants of Texas. Tex.
Res. Found., Renner, Tex., 1881 pp.

POSCUE, E. J., 1932. Land utilization in the lower Rio Grande Valley of Texas. Econ.
Geog. 8:1-11.

, 1934. Agricultural history of the lower Rio Grande Valley region. Agric.

Hist. 8:124-137.

FREEMAN, H. A., 1949. Notes on some tropical American skippers (Lepidoptera,
Phopalocera, Hesperiidae). Field & Lab. 17:75-81.

, 1951. Ecological and systematic study of the Hesperioidea of Texas. SMU

Press, Dallas, Tex., 67 pp.

, 1959. Butterfly collecting in Texas and New Mexico. J. Lepid. Soc. 13:

89-93.

OLBERT, L. E., 1969. On the ecology of natural dispersal: Dione moneta in Texas
(Nymphalidae). J. Lepid. Soc. 23:177-185.

GODMAN, F. D. & o. SALVIN, 1887-1901. Lepidoptera-Rhopalocera (3 voL). Biologia
Centrali" Americana, voL 36-38.

GOULD, F. w., 1969. Texas plants — A checklist and ecological summary. Tex. Agr.
Exper. Sta. MP-585.

HOFFMAN, c. C., 1941. Catalogo sistematico y zoogeografico de los lepidopteros
mexicanos. Segunda parte. Hesperoidea. An. Inst Biol. Mex. U. Nac. 12:
237-294.

HOLLAND, w. J., 1898, 1931. The butterfly book. Doubleday & Co., Garden City, New
York.

KENDALL, R. O., 1970a. Three hahstreaks (Lycaenidae) new to Texas and the United
States. J. Lepid. Soc. 24:59-61.

, 1970b. Lerema ancillaris (Hesperiidae) new to Texass and the United

States. J. Lepid. Soc. 24:266.

, 1972. Three butterfly species (Lycaenidae, Nymphalidae and HeHconi-

idae) new to Texas and the United States. J. Lepid. Soc. 26:49-56.

& H. A FREEMAN, 1963. The butterflies and skippers of Texas, a tentative
list. Rob & Bessie Welder Wildlife Foundation, Sinton, Texas, 5 pp. mimeo.

20(3): 129-132, 1981(83)

133

& w. w. McGUiRE, 1975. Larval foodplants for twenty-one species of

skippers (Lepidoptera: Hesperiidae) from Mexico. Bull. AllynMus. 27:7 pp.
KLOTS, A. B., 1951. A field guide to the butterflies. Houghton Mifflin, Cambridge,
Massachusetts.

LINDSEY, A. w., E. L. BELL & R. C. WE^LIAMS, 193 1 . The Hespeiioidea of North America.

Denison U. Bull J. Sci. Lab. 25:1-142.

LINTNER, J. A., 1884. On some Rio Grande lepidoptera. Papilio 4:135-147.

LffES, J. E., 1961. More butterfly records from Brownsville, Texas, including a food-
plant of Phocides polybius (Hesp.) J. Lepid. Soc. 15:114.

MACNEILL, C.D., 1975. Family Hesperiidae. The skippers. InW. H. Howe (Coard. ed.
and illus.) The butterflies of North America, pp. 423-578. Doubleday & Co.,
Garden City, New York, 633 pp.

MILES MOSS, A, 1949. Biological notes on some Hesperiidae. Acta ZooL Lilloana 7:
27-42.

MILLER, L. D. & F. M. BROWN, 1981. A catalogue/checkUst of the butterflies of America
north of Mexico. Lepidopterists Society Memoir 2:280 pp.

NECK, R. w., 1976a. Adventive land snails in the Browns ville, Texas area. South-
western Nat 21:133-135.

, 1976b. Factors affecting the occurrence of Melanis pixe (Riodinidae) in

extreme south Texas. J. Lepid. Soc. 30:69-70.

, 1977. Effects of 1933 hurricanes on butterflies of central and southern

Texas. J. Lepid. Soc. 31:67-68.

, 1978. Bionomic notes on the blood-spot skipper (Hesperiidae: Phocides

lilea sanguinea (Scudder)). J. Lepid. Soc. 32:107-110.

, 1982. Leaf selection for oviposition sites by a tropical skipper butterfly.

J. Lepid. Soc. 35:240-242.

ORTON, R., D. J. HADDOCK, E. G. BICE & A c. WEBB, 1967. Climatic guide. The lower Rio
Grande of Texas. Tex. Agri. Exp. Sta. Misc. Pub. 841:108 pp.

RUCHLE, G. D., 1948. The common guava — A neglected fruit with a promising future.
Econ. Bot 2:306-325.

SCUDDER, S. H., 1872. A systematic revision of the American butterflies; with brief
notes on those known to occur in Essex County, Mass. Peabody Acad. Sci.
(Salem, Mass.), 4th Ann. Rpt., Append. 2:24-83.

SKINNER, H., 1911. The larger Boreal American Hesperiidae, including Eudamus,
Erycides, Pyrropyge mid Megathymus. Trans. Amer. Ent Soc. 37:169-209.
SNOW, F. H., 1906. Some results of the University of Kansas entomological expedi-
tions. Trans. Kan. Acad. Set 20:136-154.

STALLINGS, D. B. & J. P. TURNER, 1946. Texas lepidoptera (Rhopalocera: Papilionoida).
Ent News 57:44-49.

STANDLEY, P. c., 1920-1926. Trees and shrubs of mexico. Contrib. U. S. Nat Herb.
23:1-1721.

TILDEN, J. w., 1974. Unusual and interesting butterfly records from Texas. J. Lepid.
Soc. 28:22-25.

TOWNSEND, c. H T., 1897. On the biogeography of Mexico and the southwestern
United States. H. Trans. Tex. Acad. ScL 2(l):33-86.

Journal of Research on the Lepidoptera

20(3): 134-135, 1981(83)

Japanese Literature

“Butterflies and Moths” (Tyo to Ga), volume 32(3/4), published March 25, 1982.

fflROSHi YOSfflMOTO. Notes on the genus Epipsestis, with descriptions of three new
species from Nepal (Lepidoptera: Thyatiridae). pp. 117-137, illusts., in English
with Japanese summary.

Redescribes or describes 10 spp. of the genus here redefined, including^, dubia
(Warren) sp. rev. and comb, nov.; E. albidisca (Warren), E. bilineata (Warren),
E. renalis (Moore) combs, nov.; E. longipennis, E. medialis, E. mediofusca spp.
nov. Nepalese E. nikkoensis (Matsumura) represents the first record of the sp.
outside Japan.

fflROSHi YOSHIMOTO. Notes on the genus Chandata, with descriptions of three new
species from Nepal and Taiwan (Lepidoptera: Noctuidae). pp. 138-146,
illusts., in English with Japanese summary.

The monotypic genus is redefined to include two groups of 2 and 4 spp.: The
partita group, C. partita Moore and C. c-nigrum sp. nov.; the bella group, C.
aglaja (Kishida & Yoshimoto) comb, nov., C, tridentata sp. nov., C. taiwana
sp. nov., and C. bella (Butler) comb. nov.

SHIGERO SUGI. Illustrations of the Taiwanese Catocala, with descriptions of two new
species. Noctuidae of Taiwan, I. (Lepidoptera). pp. 147-159, illusts., in English
with Japanese summary.

Reviews and illustrates all 1 1 spp. of the genus frpm Taiwan, including descrip-
tions of C. naganoi and C. shirozui, spp. nov.

AKIRA YAMAMOTO & SHIN TAKEI. A new species of the genus Delias from Mindanao, the
Philippines (Lepidoptera, Pieridae). pp. 160-163, illusts., in English. Delias
mandaya, sp. nov., is described from Tagubud Mts., Southeastern Mindanao.
HIROSHI INOUE. A new species of the genus Hypochrosis Guenee from Southeast
Asia (Geometridae: Ennominae). pp. 164-167, illusts., in English with Japanese
summary.

Hypochrosis baenzigeri sp. nov. is described from Thailand (type locality),
Assam, and Taiwan. According to Banziger, the sp. in North Thailand is
lachryphagous (feeds on mammalian lachrymal secretion).

SHIGERO SUGI. A new species in the Euteliinae from Japan (Lepidoptera: Noctuidae).
pp. 168-170, illusts., in English with Japanese summary.

Describes Eutelia clarirena sp. nov. which occurs from central Japan south to
Taiwan and has been erroneously known in Japan as E. sinuosa (Moore), an
Indian sp.

CHRIS SAMSON & ATUHIRO SIBATANI. On the discovery of Mycalesis sara Mathew
female (Lepidoptera: Satyridae) from San Cristobal Island, Solomon Islands,
pp. 171-178, illusts., in English with Japanese summary.

Describes previously unknown M. sara female and proposes a solution to the
long-standing confusion among related taxa of the Solomons.

ATUHIRO SIBATANI & TAKASHI NISHIZAWA. Delias rileyi Joicey et Talbot, a little known
pierid from Irian Jaya. pp. 179-181, illusts., in English with Japanese summary.
The specific status of this taxon, originally described from unique male, is con-

20(3): 134^135, 1981(83)

135

firmed by a series of males recently obtained. Female remains unknown.
KAZUHIKO MORISHTTA & KAZUHISA OHTSUKA. On the male of Lethe kinabalensis Okubo

(Lepidoptera: Satyridae). pp. 182-184, illusts., in English with Japanese
summary.

Describes the first male specimen of the taxon originally described from two
females from Mt. Kinabalu, Sabah, E. Malaysia.

ISAMU HIURA & FUSAO NAKASUJI. A questionnaire on the migration of the rice skipper,

Parnara guttata (Lepidoptera: Hesperiidae). pp. 185-189, map, in Japanese
with English summary.

Discusses the species’ migration patterns and geographic limits of movements
in Japan, 1979, based on the results of survey through mailed questionnaires.
KAZUO SAITOH, AZUMA ABE, YOSHINORI KUMAGAI, & SATOSHI KOIWAYA. Karyological
notes on Sasakia funebris Leech and Sasakia charonda Hewitson (Lepidoptera:
Nymphalidae). pp. 190-192, illusts., in English with Japanese summary.
Number {n ~ 30) and morphology of the chromosomes of S. funebris from
Szechwan, China, are compared with those of the Japanese S. charonda (n = 29).
SHIN-ICHIRO KATO. Notes on the Lycaenidae from Ambon Island, Indonesia, pp.
193-202, illusts., in Japanese with English summary.

Gives an annotated list of 16 genera 22 spp. of lycaenids collected by the author,
including 3 spp. previously unknown from the Island.

Authors’ addresses:

Azuma Abe
Isamu Hiura

Hiroshi Inoue
Shin-ichiro Kato

Satoshi Koiwaya
Yoshinori Kumagai

Kazuhiko Morishita
Fusao Nakasuji

Takashi Nishizawa
Kazuhisa Ohtsuka
Kazuo Saitoh

Chris Samson

Atuhiro Sibatani
Shigero Sugi
Shin Takei

Akira Yamamoto
Hiroshi Yoshimoto

Hirosaki High School, Hirosaki-shi, Aomori-ken 036, Japan
Osaka City Natural History Museum, Sumiyoshi-ku, Osaka
546, Japan

311-2 Bushi, Iruma-shi, Saitama-ken 358, Japan
4-13-20 Kiyoshikojin, Takarazuka-shi, Hyogo-ken 665,
Japan

3- 14-5 Shimotakaido, Suginami-ku, Tokyo 168, Japan
Namioka Weak Children’s School, Namioka-cho, Aomori-
ken 038-13, Japan

2-2-16 Shinjuku, Zushi-shi, Kanagawa-ken 249, Japan
Entomology Laboratory, Faculty of Agriculture, Kyoyo
University, Sakyo-ku, Kyoto 606, Japan

4- 6-8 Asakusabashi, Taito-ku, Tokyo 111, Japan
2-4-2 Shinmachi, Hoya-shi, Tokyo 202, Japan
Department of Biology, Hirosaki University, Hirosaki-shi,
Aomori-ken 036, Japan

The National Butterfly Museum, St. Mary’s, Bramber,
West Sussex BN4 3 WE, England

30 Owen Street, Lindfield, New South Wales 207 0, Australia
41-3 Akadutumi 5-chome, Setagaya-ku, Tokyo 156, Japan
1561 Heathdale Drive, Burnaby, British Columbia V5B
3N9, Canada

1-18 Nogata 2-chome, Nakano-ku, Tokyo 165, Japan
Tokyo High School, 39-1 Unoki 2-chome, Ota-ku, Tokyo,
Japan

Ichiro Nakamura, compiler, 41 Sunrise Blvd., Williamsville, New York 14221

Journal of Research on the Lepidoptera

20(3): 136-160, 1981(83)

Hipparchia azorina (Strecker, 1899) (Satyridae) Biolo^,
Ecology and Distribution on the Azores Islands

Steffen Oehmig

D-5090 Leverkusen 3, Akazienweg 5, West Germany

Abstract. The present paper deals with the speciation and distribution of
Hipparchia azorina (Strecker, 1899). The type locality of H. azorina is
restricted to the central island group of the Azores. The eastern island, Sao
Miguel, is inhabited by Hipparchia miguelensis (LeCerf, 1935) stat. rev..
Hipparchia caldeirense sp. n. is restricted to the island Flores in the western
group. The larvae on Faial, Sao Miguel and Flores feed on Festuca jubata
Lowe. Morphology of adults and early stages is described. The habitat of
Hipparchia azorina, H. caldeirense and H. miguelensis is described and
suggestions regarding their conservation are given.

Introduction

Hipparchia azorina (Strecker, 1899) was described from a single male
specimen, given to Strecker by E. T. Owen, who brought it back from the
Azores. Strecker gave “Azores” as the type locality, wdthout any exact
data. The synonomy follows:

Satyrus azorinus

STRECKER

1899

Satyrus semele maderensis Bethune Baker;

REBEL

1917:17

Satyrus azorinus Strecker;

WALKER

1931:77

Satyrus azorinus Strecker;

GAEDE

1931:157

Satyrus azorinus Strecker;

LE CERF

1935:206

Satyrus azorinus picoensis

LE CERF

1935:208

Satyrus azorinus miguelensis

LE CERF

1935:208

Oeneis ohshimai

ESAKI

1936:483

Satyrus semele azorinus Strecker;

REBEL

1939:47

Satyrus semele azorinus Strecker;

REBEL

1940a:9

Satyrus semele azorinus Strecker;

REBEL

1940b:16

Hipparchia azorinus Strecker;

LESSE

1952:80

Satyrus semele azorinus Strecker;

CARTHY

1957:210

Hipparchia azorina Strecker;

MARSDEN & WRIGHT

1971:180

Hipparchia azorina Strecker;

KUDRNA

1975:205

Hipparchia aristeus azorina Strecker;

HIGGINS

1975:226

Hipparchia azorina Strecker;

KUDRNA

1977:97

Hipparchia azorina Strecker;

HIGGINS & RILEY

1978:122

In order to study the relationship of the populations inhabiting the
Azores in their natural environment, I visited the Azores from June 26th to

20(3): 136-160, 1981(83)

137

July 10th, 1980. The results of this visit and subsequent studies are given
below.

Original descriptions Satyrus azorinus Strecker 1899

Body, head and antennae black. Wings dark brown. Primaries somewhat
dull ochreous on the disk. A small round subapical spot between vein 5 and
6. Secondaries with a strongly sinuate dull ochreous mesial band, this has a
deep sinus inwardly between veins 2 and 3, and another at vein 6. Fringe of
all wings white, with black termination of veins. The disk and mesial band
are not decided or well defined, but dull and suffused, as if showing
through from the underside. Under surface, primaries dull pale ochre.
Costa brown. At end of and beyond the discoidal cell a brown mark
extends from the costa to vein 4. The subapical spot of upper side is
repeated, beyond this spot to the costa brown. A brown marginal band, two
small white spots interior to this band between veins 6 and 8. Secondaries
dark brown, somewhat striated. A mesial band as above but pure white and
sharply defined, interior to this band are two white marks, one near the
base, is irregular and extends from the costa to within the discoidal cell.
The other nearly square is below this in the cell. Fringe as above. Expands
iy2 inches. Type, one cf received from prof. E. T. Owen, who informs me it
came from the Azores, The place for this most interesting species I think
would be with or near Satyrus (Chionobas) pumilus, lama, etc. In a remote
way it also reminds one of S. neomiris.

0 Corvo
i^/Ftores

^ Graciosa

Sao Jorge '''\

"^Terceira

\

/'Sao Miguel

100 km

Santo Maria

Fig. 1. Distribution of the Hipparchia taxa on the Azores Island. - - - Hipparcfua
miguelensis, Hipparchia azorina, ..... Hipparchia caldeireme.

138

J. Res. Lepid.

Distribution

The Hipparchia azorina complex is confined to the Azores. The species
complex has been recorded from five of the total nine islands. Their
distribution is given in Figure 1. The Azores Archipeligo extends between
36°55' and S9°44' north latitude and 25° and 31°15' west longitude. The
distance from Sao Miguel in eastern group to Pico Island in the central
group is approximately 250 km. Pico island is another 250 km from Flores
island in the western group. These distances clearly demonstrate the
dispersal required for the insects to settle upon these islands. It is
possible, according to the hypothesis of Gatter (1981), that the drifting
form of migration (the carrying of the insects by the northeast trade winds)
is responsible for establishment of Hipparchia on the various islands. A
recent good example of this form of migration is the colonisation of
Madeira by Pieris rapae as documented by Wolff (1975). A few years later
Higgins (1977) and Oehmig (1977) reported the reestablishment of
Pararge aegeria on Madeira.

Populations of the Islands
PICO

Hipparchia azorina (Strecker 1899) stat, rev.

Lepidoptera Rhopalocera and Heterocera indigenous and exotic. Suppl.
3:3.

Satyrus azorinus picoensis LeCerf, 1935, Bull. Soc. ent. Fr., 40:206-209.
Syntypes 2 cfcf, 1 9, [Azores], Pico, Museum National d'Histoire Naturelle,
Paris.

Description: cf : upperside forewing dark brown, but never as dark as the
males from the Faial populations. Discal and postdiscal area with a pale
ochre yellow color. Occasional specimens appear with a lightened basal
area. In cell 6 a dark brown spot, sometimes as eye spot. upperside
hindwing dark brown postdiscal area light ochre. In cell 2 sometimes a dark
brown spot, cf: underside forewing range from a pale to a strong ochre,
always lighter than the males from the Sao Miguel populations. The dark
brown eyespot in cell 6 or the spot in cell 2 often absent. Outer margin
more or less sharply defined, cf: underside hindwing basal and discal
area dark brown and project pointed into the white postdiscal area.
Females were not found on the Pico island during this study, therefore
they shall not be described here.

Androconial scales: length: 0.14-0.16 mm, width: 0.015 mm. Andro-
conial patch is not always present. Although small, the androconial form is
similar to that of males of Sao Miguel population.

Male genitalia: Valva length: 1.8 mm, width: 0.3 mm. The valva are
distinctly narrower than in Hipparchia miguelensis LeCerf, 1935. The
dorsal process on the valva is mostly rounded off and not as pointed as that

20(3): 136-160, 1981(83)

139

of the Sao Miguel specimens.

Distribution: Pico, Azores, from 600 m on, upwards to 2000 m. Cabeco
do Encalvado 900 m. Walker (1931) Serra Gorda, Cabeco do Afonso, O
Pico. On Pico up to an altitude of 2000 m; LeCerf, 1935. Males only were
observed on the northern high plateau of the island. Vegetation on the
northern slope of Pico predominantly consists of Calluna vulgaris (L.)
Hull, amongst Pteridium aquilinum (L.) Kuhn andErica azorica Hochst. ex
Seub., which grow up to 3 meters tall. Walker (1931) states that specimens
from Cabeco do Afonso and Serra Gorda were found only singly “on the S.
and SE. of the mountains we found it much commoner”. Rebel (1940b)
cites Silveira and Lagoa do Caiado on Pico island. The main habitat,
however, is found on the southern slopes of Pico island, protected against
the northeast tradewinds. The early stages of H. azorina on the Pico island
remain unknown.

Discussion: Since Strecker (1899) gave as type-locality “Azores”, it is
necessary to identify and restrict the exact place for taxonomic clarity. The
holotype was not inspected directly. However, black and white and color
slides of the holotype were available for comparison. The genitalic
preparation of the holotype showed great similarity indeed with the
genitalia of males from Pico. The holotype was examined by Kudma
(1977), who also examined my material from Pico. In addition the original
description of H. azorina (Strecker, 1899) agrees very closely with males
from Pico, “wings dark brown, primaries somewhat dull ochreous on the
disk, under surface, primaries dull pale ochre”. Males from Flores as well
as males fom Sao Jorge are lightly ochreous colored. The Faial males
certainly have a stronger ochre coloring on the underside of the fore wings,
while their upper surfaces are always dark brown. I propose to restrict the
type locality of H. azorina to Pico.

Material examined: 6 cfcf, forewing length: 21-22 mm, leg. et coll. S.
Oehmig, Azores; Pico: Cabeco do Encalvado 9000 m: 2 July 1980.

Appearance: June-September, possible longer,

FAIAL

Hipparchia azorina ohshimai (Esaki 1936) comb, nov., stat. nov.
Oeneis ohshimai Esaki 1936, Annotnes. zool. jap. 15(4):483-485 [Azores]:
Faial; Caldeira, Pico Gorda: 1021 m; 24 Aug. 1935 and 30 Oct. 1935, leg.
H. Ohshima. Holotype Allotype 9, Paratypes 2 d’d', 3 99, Entomological
Laboratory, Kyushu Imperial University, Fukuoka, Japan.

Description: d': upperside forewing color dark blackish-brown, in ceils 6

and 2 dark brown spots, some without these spots, d: upperside
hindwing color dark blackish-brown. Postdical area from the underside
weakly translucent, d: underside forewing pale to middle ochreous
color, but not so strong ochreous as the forewing underside from the Sao

140

J. Res. Lepid.

Miguel males, with a fine dark brown zigzag postdiscal line. Outer margin
dark brown, project into the postdiscal region, and is not sharply defined.
Some with a narrow outer margin band, d*: underside hindwing, basal
and discal area blackish brown, always taper off along the discoidal and
median veins projecting into the white postdiscal area. The basal area has
white spots. 9: upperside forewing dark brown. Discal and postdiscal
area lighter than the males. In cell 6 a dark brown spot or eyespot, in cell 2 a
dark brown spot. 9: upperside hindwing dark brown, but lighter than the
males. 9: underside forewing light ochre with a dark brown zigzag band.
Outer margin dark brown, sharply defined. Spots from the upperside
sometimes absent. 9: underside hindwing basal and discal area dark
brown, lighter than the males. Outer margin light brown to dark brown.

Androconial scales: length: 0.13-0.14 mm, width: 0.02 mm. Andro-
conial form is somewhat stunted. The androconial patch is often divided
into two patches along the median veins in cell 2 and cell lb. Many
specimens appear without an androconial patch. Spatulate androconial
scales have been observed, such forms being unique to males from Faial.

Male genitalia: Valva length: 1.8 mm; width: 0.4 mm. The valves differ
from those of specimens from Flores and Pico in their greater width; the
dorsal process terminates in a point.

Female genitalia: Signum length: 1.2 mm; width: 0.15 mm. Smaller than
the signa from the Sao Miguel females.

Distribution: Azores, confined to Faial from 700 m to over 1000 ni,
(Walker, 1931) Caldeira southern inclines 900 m, (Esaki, 1936) Caldeira,
Pico Gorda 1021 m, (Rebel, 1940b) above Horta, Caldeira. I found this
subspecies very common on the southern slopes of the Caldeira at 900 m.
The butterflies fly frequently on the south eastern slope of the Caldeira on
Pico Gorda. In the Caldeira there are only single specimens in the cliffs of
the upper crater wall. The biotope here gives the impression of a mountain
meadow. Festuca jubata Lowe, is abundant, as is Calluna vulgaris.
Daboecia azorica Tutin & Warb. grows round in nest form with red
blossoms in summer. The cliffs are often extremely rugged from water
erosion. The grass vegetation also includes occasional plants of Potentilla
sp. The biotopes of Faial island differ clearly from the biotopes of the other
islands in special vegetation.

Early Stages: Ovum: Diameter: 1.1 mm; height: 1.1 mm. The termin-
ology follows Doring (1955). The egg form is a half elliptical barrel shape
with a convex egg base. The top view is circular with longitudinal ribs,
extending from the base to just below the micropylar area, although some
ribs do reach the micropylar area. The ribs are in longitudinal ridges with
two rows in binded aeropyles. The chorion is provided with fine hardly
visible cross lines. Number of ribs, 24. The taxonomic importance of the
micropyles is indicated by Hinton (1981). In many species the number may
be constant, and could therefore be used as a reliable taxonomic character.

20(3): 136-160, 1981(83)

141

However, I have also noticed some species in which the number of
micropyles varies within all the eggs of a female. Four micropyles have
been counted in the Faial ova, the micropyle rosettes are seven leaved.
The ova are individually attached by the females to the foodplants. The
base of the ovum is somewhat adhesive. Eggs are always deposited on
plants protected from the wind. Fifteen days elapsed to hatching under
handling conditions. The egg is white at first, then becomes dappled with
light red-brown, and shortly before hatching it is grey-brown. In the field
females cease oviposition when the sun disappears behind a cloud. Egg
deposition stretched into evening under artificial light. In the laboratoiy
Faial females lay 60-70 eggs.

(LI) first instar: Body length: 3-4 mm; final LI: 6 mm. Duration: 21
days. The larva hatches through a circular gnawing of the chorion on the
micropylar area. Usually a cross-piece remains hinged, forming an
emergence lid, although sometimes the chorion is entirely gnawed through
forming a hole. The chorion itself is not consumed. Emergence of the larva
can only occur under humid conditions. Arid conditions inhibit and may
even prevent the emerging. The description of the line patterns of the larva
follows Shirozu & Hara (1979), Fig. 2, and the description of the head is
according to Beck (1974), The coloring of the larva is that of a Hght sand.
Dorsal, subdorsal, subspiracular and spiracular lines are all light brown.
The end of LI is light brown as well. The head capsule has four primary
setae on either side of the genae, head diameter 0.8 mm. The dorsal side of
the body has four rows of setae, and on the fifth segment of the ventral side
there are 6 setae, which are turned caudally. The anal segment is forked,
with two setae on each side. The LI head capsule of Hipparchia semele L.
has the same number of primary setae as the Hipparchia taxa from the
Faial, Flores and Sao Miguel islands. In complete contrast, the genus
Pseudochazara exhibits primary fine hairs which are very strongly
pronounced (Aussem & Hesselbarth, 1980).

(L2) second instar: Body length: 5 mm; final L2: 8-9 mm. Duration: 17
days . The coloring and line patterns of the body are as in L 1 , although now
there is an additional pale white basal line. The head is light brown,
diameter 1 mm. The density of pubescence increased. Genae are marked
on either side with light brown coronal lines, supraocellar lines and ocellar
Mnes. Anal segment is forked.

(L3) third instar: Body length: 9 mm; final L3: 14 mm. Duration: 17
days. Coloring and marking of the body as in L2. Head capsule diameter,
1.5-1. 6 mm. The dorsal line often interrupted or dotted. Head capsule
dark brown and more pubescent than L2. The lines of the head capsule as
in L2. Abdominal fine hair no longer present. Anal segment is forked.

(L4) fourth instar: Body length: 13 mm; final L4: 18 mm. Duration: 16
days. The color is usually dark brown with a few light brown individuals.
Dark brown dorsal line often interrupted. Subdorsal line light brown;

142

J. Res. Lepid.

subspiracular line dark brown, as broad as dorsal line. The spiracular line
light brown, narrow white basal line distinct. Head capsule dark brown.
The genae lined patterns as in L3 densely pubescent. The fine hair on the
body equally abundant. The anal segment forked. From L4 on, the sexes
differ in diameter of head capsule: cfcf 2.1 mm, 99 2.5 mm.

(L5) fifth instar: Body length: 25 mm; final L5: cSd" 27 mm, 99 30-32
mm. Duration: 26 days. Head capsule diameter: cfd* 3.5 mm, 99 5 nun.
Dimorphic, both dark and pale brown forms present. Dorsal line dark
brown bordered by two thin white lines occasionally dotted and inter-
rupted. Subdorsal line dark brown, supraspiracular line also dark brown
with two fine white border lines. White basal line bordered by two thin
dark brown lines. Thin, short dark brown marks present between dorsal
line and spiracular line. Body thickly covered with short bristly hair. Both
dark and pale brown head capsules observed. On either side of the genae
there is a coronal line, supraocellar line and ocellar line. These markings
are dark brown, and in many specimens are so wide and so dark that the
head appears to be entirely black-brown. The head has a dense covering
short red-brown hair. The bristles of the Faial specimen, however, are
somewhat longer than those of the L5 instar of the Sao Miguel and Flores
populations. Males and the females are easily distinguishable by the width
of the head.

Prepupa and Pupa: During the prepupa stage the markings of the larva
become lighter and more translucent as the body of the larva contracts and
thickens. Under laboratory conditions the prepupa lasts 10 days. In
comparison with the pupae from Sao Miguel, the Faial pupae are dark
brown in coloring, although there is some variation. On the dorsal and
lateral sides of the abdominal segments the Faial pupae have dark brown
pigment markings. On each segment there are up to 10 marks in a double
row. The abdomen of newly formed pupa is capable of movement.
Pupation occurs without the preparation of a web, between the leaves of
Festuca jubata Lowe. Occasionally pupation takes place in open ground. In
most cases in the field a grassy area sheltered from the wind is selected for
pupation. In the beginning of July 1980 I found L5 larvae capable of
pupation and pupae at an elevation of 900 m on the southeast slopes of the
Caldeira. Under laboratory conditions, at temperature of approximately
20°C, the pupal state lasts 21-30 days. The pupae require a humid
environment. If they become too diy, the butterflies cannot eclose
properly, or emerge crippled. This could be the result of adaptation to the
moist often damp ground vegetation of the Azores mountain regions . Pupa
length/width (mm): 15/6; 99 17/7.

Adult Material Examined: 45 cfd", forewing length: 20-22 mm. 25 99
forewing length: 21-24 mm. Leg. et coll. S. Oehmig, Azores, Faial,
Caldeira, southeast slope 900 m, 1 July 1980.

Discussion: The dark color of the male upperside is peculiar to Faial

20(3): 136-160, 1981(83)

143

specimens. The stunted form and small size of androconial scales is so
different compared with androconial scales of the Hipparchia taxa of the
islands of Pico and Sao Jorge, that separation of this taxon from the others
is justified.

Foodplant: Festuca juhata Lowe. (Gramineae).

Appearance: June-October.

SAO JORGE

Hipparchia azorina jorgense Oehmig new subspecies

Holotype cf; Paratypes cf, 8 99; [Azores]: Sao Jorge, Coroas 600 m, leg. D.
T. Pombo, 2 Aug. 1981, Nord Biscoitos Transversal, leg. D. S. Furtado, 26
Aug. 1981. All types in my collection.

Description: cf: upperside forewing dark brown, discal region ranges
from a light ochre to a sand color. In cells 6 and 2 a dark brown spot. The
light sandy colored discal region is typical of the specimen from Sao
Jorge, cf: upperside hindwing dark brown, postdiscal band is a striking
light sandy color, cf: underside forewing very pale ochre, with an eyespot
in cell 6 and a dark brown spot in cell 2. Outer margin dark brown is alwyas
narrow and sharply outlined, cf: underside hindwing basal and discal
area dark brown. A characteristic which only occurs amongst the Sao Jorge
specimens is that the pale white postdiscal band is laid very broadly. The
discal area runs along the discoidal and median veins tapering off into the
white postdiscal band. 9: upperside forewing light ochre to sandy
colored discal and postdiscal area. In cells 6 and eyespots or spots in
brown color. 9: upperside hindwing, a pale postdiscal area is typical as
well for these specimens. 9: underside fore wing quite pale ochre to
sandy color. In cell 6 and 2 brown eyespots or spots. Outer margin brown
sharply defined. 9: underside hindwing basal and discal area dark
brown. The discal area project pointed into the white postdiscal area.

Androconial scales: length: 0.14-0.15 mm; width: 0.018 mm. Andro-
conial patch small not always present. The form and size is similar to that
of the Faial males. At the base however, the form is never spatulate round,
rather it is always pointed.

Male genitalia: Valva length: 1.3 mm; width: 0.35 mm.

Female genitalia: Signum length: 1.3 mm; width: 0.2 mm.

Distribution: Azores, Sao Jorge, from 480 m to over 700 m. Marsden &
Wright (1971) between Urzelina and Ouvidor, Mr. Pombo, Coroas 600 m
leg. 2 Aug. 1981, Mr. Furtado, Nord Biscoitos Transversal, leg. 26 Aug.
1981. The only information concerning the Hipparchia biotopes of the
island of Sao Jorge are based on the results of Marsden & Wright (1971),
who reported the habitat of Hipparchia azorina is to be found from the
upper Callunetum-Ericetum pasture to the Agrostis pasture zone. The
lower boundary was specified by the authors as being at 480 m on the
northern slopes, and 540 m altitude on the southern slopes. The number of

144

J. Res. Lepid.

PLATE L

20(3): 136-160, 1981(83)

145

PLATE L Imagines of the Azores Hipparchia taxa.

a. H. azorina jorgense, cf upperside

b. H. azorina jorgense, d* underside

c. H. azorina jorgense, 9 upperside

d. H. azorina jorgense, 9 underside

e. H. azorina, cf upperside

f. H. azorina, (S underside

g. H. miguelensis, cf upperside

h. H. miguelensis, cf underside

i. H. miguelensis, 9 upperside

j. H. miguelensis, 9 underside

k. H. azorina ohshimai, cf upperside

l. H. azorina ohshimai, cf underside

m. H. azorina ohshimai, 9 upperside

n. H. azorina ohshimai, 9 underside

o. K caldeirense, (f upperside

p. H. caldeirense, d' underside

q. H. caldeirense, 9 upperside

r. H. caldeirense, 9 underside

s. H. azorina ohshimai, 5th instar

t. H. miguelensis, 5th instar

u. H. miguelensis, 5th instar

Figures a-r: 0.6 X natural size.

Figures s-u: 1.05 X natural size.

flying adults observed per 3 minute at varying altitudes taken from
Marsden & Wright (1971) is as follows: Between 480-540 m, fewer than
10; at 720 m, on the southern side, approximately 30; and on the northern
side, 85 adults.

Based on these results, it is apparent that the Hipparchia azorina
population of Sao Jorge occur most frequently at an altitude of approxi-
mately 700 m. The above authors also provided data concerning the
temperatures at the high altitude of 700 m, during the month of
September. Under clear skies the temperature may drop as low as 7°C
with 24°C as the daytime high temperature. One may assume, because
larval feeding occurs exclusively during the night, that by September the
larva have already gone into diapause. Under laboratory conditions at a
temperature of 20°C, 10 weeks are necessary for the development to the
third larval stage. An egg, deposited at the beginning of July, would thus be
expected to reach a maximum third larval stage by mid September.

Material Examined: Holotype cf, forewing length: 22 mm, [Azores], Sao
Jorge, Coroas 600 m, 2 Aug. 1981, leg. Mr. D. T. Pombo, Santa Maria,
Azores. Paratypes cf and 9 leg. Pombo, 7 99 leg. Mr. D. S. Furtado, Sao
Miguel, Azores, Nord Biscoitos Transversal, 26 Aug. 1981. Male forewing
length: 22 mm, female 23-26 mm. All types in my collection.

Discussion: Specimens of Sao Jorge are, with respect to the pale

146

J. Res. Lepid.

forewing upperside and the wide postdiscal area on the hindwing
underside, clearly different from the Hipparchia taxa of Pico and Faial.
Both androconial scales and male genitalia also show differentiation
between the taxa of Pico and Faial. These characters justify a distinct
subspecies.

Foodplant: Not known, probably Festuca jubata (Gramineae).

Appearance: June-September.

SAG MIGUEL

Hipparchia miguelensis (LeCerf, 1935) stat. rev.

Satyrus azorinus miguelensis LeCerf, 1935, Bull. Soc. ent. Fr., 40:206-209
[Azores]: Sao Miguel.

Holotype cf, Allotype 9, Paratypes 2 cfd*, Museum National d'Histoire
Naturelle, Paris.

Description: d': upperside forewing dark brown, but not so dark as
males from Faial and Pico, in cell 6 a dark brown eyespot or spot, in cell 2 a
dark brown spot. The ocelli of some of the specimens have a strong,
underlying ochre yellow color. Some males are rather similar to the
females due to the distinct ochre yellow markings which they exhibit. Most
males, however, are without this strong ochre yellow marking, cf : upper-
side hindwing dark brown, some with a dark ochreous postdiscal area, in
cell 2 a dark brown spot. underside forewing dark ochre yellow,
typical//, miguelensis. In cell 6 a eyespot, in cell 2 sometimes a dark brown
spot. Outer margin dark brown and sharply defined. Costa dark brown, cf :
underside hindwing basal and discal area dark brown, but lighter than the
Faial taxa. The discal area always terminates round into the white or pale
ochre postdiscal area. 9: upperside forewing dark brown, as in the males .
In cell 6 an eyespot, in cell 2 a brown spot. The spots of the specimens have
a strong underlying ochre color. The ochraeous colored markings of the
females are more pronounced than those of the males. 9: upperside
hindwing dark brown with more or less strong ochre postdiscal area. In cell
2 a dark brown eyespot. 9: underside fore wing strong ochre yellow with a
finely marked ochre zigzag pattern in the postdiscal area. In cell 6 an
eyespot, in cell 2 a dark brown spot. Outer margin dark brown, sharply
defined. 9: underside hindwing basal and discal area dark brown. The
discal area between cells 3 and 4 along the discoidal and median veins are
always bluntly rounded off, reaching into the light cream colored
postdiscal area.

Androconial scales: length: 0.17-0.19 mm; width: 0.015 mm. These
are the largest androconial scales of all the taxa from the Azores.
Androconial scales wide and strongly tapered off toward the apex. The
androconial patch is most clearly pronounced of all taxa, but not very large.
There are two along the median vein of the cell.

Male genitalia: Valva length: 2.25 mm; width: 0.45 mm. The valva is

20(3): 136-160, 1981(83)

147

wider than those of the other taxa from the Azores. The terminal
extension is always short in form. The dorsal process is well pronounced,
and tapers off to a point. The uncus is always distinctly longer than in the
taxa of the other island.

Female genitalia: Signum length: 1.6 mm, width: 0.2 mm. Longer and
broader than those of the other taxa.

Distribution: Sao Miguel, Azores, from 600 m to over 1000 m. Gafanhoto
715 m, Vista do Rei 600 m (Rebel, 1940b):17, Lagoa do Fogo 700 m, Pico
da Vara 1103 m. The important environment requirement of H, miguel-
ensis populations is the presence of Festuca jubata. This plant shows
substantial variation in composition and aspect in the habitats where it is
found amongst the islands; from the grassy meadows in the mountains to
plant associations which form thickets, and where Festuca jubata itself
only plays a minor role (Lupnitz, 1975a). In the Vale de Furnas, Gafanhoto
700 m, southern exposure, the habitat is in a densely wooded zone. Laams
azorica (Seub.), which stands approximately 2 meters tall, grows in wide
intervals together with Vaccinium cylindraceum Sm., and Rubus sp. In
between grows Calluna vulgaris (L.) Hull, which intertwines to practically
form a surface. Festuca jubata prospers in thickets amongst Blechnum
spicant (L.) Roth., Osmunda regalis L., Woodwardia radicans (L.) Sm.
individual Heydichium gardnerianum Roscoe and Potentilla sp. Close to
the ground are often Selaginalla sp., Sphagnum sp. and Lycopodium sp. In
most places the ground is heavily covered with foliage. On account of
frequent precipitation the vegetation is often dripping wet for the entire
day. These plant communities can be found near Pico da Vara. The
butterflies, although fewer in number, fly as well on the Vista do Rei. The
habitat is small, perhaps a vestigial environment. Many Lotus uliginosus
Schuhr and Lotas parviflorus Desf. are in bloom, and Festuca jubata is also
present. Single butterflies have also been observed at the inaccessible
western crater- wall of the Sete de Cidades.

Ovum: diameter: 1.1-1. 2 mm; height: 1.1 mm. Micropyles 3. Micro-
pyle rosettes five leaved. There are 26 ribs in longitudinal ridges. The form
and coloring is as in the Faial specimens.

Larva: The development of the larvae from the Hipparchia miguelensis
populations corresponds to that of the population from Faial. The
markings of the larva also coincide with those of the larvae from Faial.
However, in contrast to Faial larvae, the 5th instar of the Sao Miguel larvae
are always a light sandy color with a pattern of light brown lines. The head
capsule is always a pale red-brown. The fine hair of the head capsule is
shorter in the 5th instar than in the Faial 5th instar. In most larvae, the
coronal line and the supraocellar line do not meet in the height of the ocelli,
as is the case of the Faial population.

Pupa: The Sao Miguel pupa is always light brown in color. In contrast to
the Faial pupa, the dorsal abdominal markings are not present, or are at

148

J. Res. Lepid.

least very indistinct. Pupa, length/width (mm): cfcf 15/16; 99 17/7.

Material Examined: 90 d'cf, forewing length: 22-25 mm; 8 99 forewing
length: 24“28mm. Leg. et coll. S. Oehmig, Azores, Sao Miguel, Gafanhoto
700 m, 27 June 1980.

Discussion: Typical for H. miguelensis is the large signum of the females,
and the large uncus and the wide valva of the males. The androconial
scales are the largest of all the taxa of the islands. The ovum shows 26 ribs
and 3 micropyle openings. The micropyle rosette is five leaved. The 5th
instar larva, including its head capsule, is a great deal lighter in color than
the other taxa. The fine hairs of the head capsule of the last instar are
shorter than hairs of the head capsule from the Faial larvae. It is clear that a
distinction is in order differentiating this taxon from H, azorina based
upon the divergent coloring and markings of the larva, as well as the
completely different appearance of the imagines.

Foodplant: Festuca jubata (Gramineae)

Appearance: June-September.

It can surely be assumed that the Hipparchia appearing on the Azores
islands are of allopatric distribution. The concept of superspecies (Mayr,
1967) should be applied here. Mayr has indicated that the superspecies
concept be especially applied to the pattern of variation associated with
insular distribution patterns.

FLORES

Hipparchia caldeirense Oehmig new species

Holotype Paratypes 33 ^cf, 6 99; [Azores], Flores, Caldeira Seca, 700
m, 30 June 1980, leg. et coU. S. Oehmig.

Description: upperside forewing dark brown, basal and discal area

lighter ochre. In cell 6 a dark spot, sometimes with underlying of light ochre
color. In cell 2 a small dark brown spot, however the spot is not present in
all specimens, cf: upperside hindwing dark brown color, the postdiscal
area is only poorly visible. Some specimens with a dark brown spot or
eyespot in cell 2. underside forewing light ochre yellow color in the
basal and discal region. The outer margin borders on a dark brown area,
which always broadly reaches into the postdiscal area, and is not sharply
defined. The brown spot in cell 2 is never present. cS: underside
hindwing basal and discal region dark brown with distinct white markings
in the basal region. This is a typical characteristic of the specimens from
Flores, and appear only rarely in specimens from Pico island. The dark
brown discal area runs along the discoidal and median veins, tapering to a
point and project into the white postdiscal area. 9: upperside forewing
color like the males, but somewhat lighter ochreous in color; dark brown
spots in cells 6 and 2 larger than the males. 9: upperside hindwing Hke
the males. 9: underside forewing like the males but the outer margin is

20(3); 136-160, 1981(83)

149

not as broad and dark as the males. 9: underside hindwing like the males
but somewhat hghter in color.

Androconial scales: Not present. One may assume that they were lost
secondarily.

Male genitalia: Valva length: 0.17 mm; width: 0.2 mm. The Flores
specimens are distinguished by their valves which are the narrowest of all
the Azores taxa. The terminal extension is especially long.

Female genitalia: Signum length: 1.3 mm; width: 0.1 mm.

Distribution: Flores, Azores, Caldeira Seca 700 m and above. The
butterflies fly along the inclines of the Caldeira Seca and the Pico dos Sete
Pes, from altitudes of 700 m and more. The environment is pervaded with
valleys formed by erosion. The vegetation is rich in grass and Festuca
jubata, which grows in dense thickets, is quite common. In the eroded
valleys by water one can find the following shrubs: Laurus azorica,
Viburnum tinus L., Rhammnus gladulosa Ait., Rubus sp., and occasionally
Vaccinium cylindraceaum. Dwarfed Juniperus brevifolia (Seub.) Ant. and
Erica azorica can be found in the back of the valley. Calluna vulgaris
appears as well and occasionally one finds Potentilla sp. In some areas the
environment resembles a fen. A thick cushioning is often built on the
ground by Spaghnum sp. With exception of the protected valleys the
environment gives a barren impression, and is barely protected from the
tradewinds. As a result the butterflies remain primarily in the protected
valleys, however when they do fly out of the valley, then they are often
carried quite some distance by the wind.

Ovum: diameter: 1.0 mm; height: 1.0 mm. The form and coloring is
similar to that of the specimens of Faial and Sao Miguel. There are 24 ribs
in longitudinal ridges, two micropyl openings, micropyle rosettes are 4
leaved.

Larva: The development of the Flores larva corresponds to the larval
development of the taxa from Faial and Sao Miguel. The Flores larva are
much lighter in coloring than the larvae from Sao Miguel. In this respect
the Flores larvae are similar to those of Sao Miguel.

Pupa: The pupa is always light brown in color. The distinctly pro-
nounced abdominal markings of the Faial pupae are barely visible or are
not present. Pupa length/ width (mm): rfd" 13/6.

Material Examined: 34 rfcf, forewing length: 19-22 mm; 6 99, forewing
length: 23-24 mm. Holotype cf, forewing length 21 mm, and paratypes.
Azores, Flores, Caldeira Seca, 700 m, 30 June 1980, leg. et coll. S.
Oehmig.

Discussion: On account of the morphological differences between the
imagines, it is reasonable to separate H. caldeirense from Hipparchia
azorina. In particular the extremely narrow valves of the males and the
small signum of the females justify separation. The complete absence of
the androconial scales can be seen as a barrier to the mating of the Flores

150

J. Res. Lepid.

taxon with the Hipparchia populations of the other Azores islands.
Tinbergen (1941) has shown that H. semele L. males, from whom the
androconial scales have been removed, are still capable of mating.
However, in comparison with the males which still possess the androconial
scales, the former are clearly at a disadvantage in mating. A further feature
supporting the separation of Hipparchia caldeirense from H. azorina is the
different structural character of the ovum upper surface.

Foodplant: Festuca jubata Lowe (Gramineae).

Appearance: June-September.

Foodplants— Adults and Larvae

The imagines of all Hipparchia populations from all the islands primarily
visit the blossoms of Rubus ulmifolius Schott and Rubus hochstetteranum
Seub., as well as Potentilla erecta (L.) Raesch and Potentilla anglica
Laicharding as their nectar sources. On Sao Miguel I observed individual
butterflies as they fed from the blossoms of Calluna vulgaris. All of the
habitats are quite poor in the flowering plants, which give rise to questions
of nutrition for the adult butterflies. Present in the environment are also
the following blossoming plants, which by observation are not visited by
the butterflies for feeding: Vaccinium cylindraceum on Flores and Sao
Miguel islands. Thymus caespititius Brot. on Pico, and Daboecia azorica on
Faial and Pico islands. The females prefer grass cushioned areas, which
are compact and not too large, for oviposition. Oviposition in all observed
cases is confined to Festuca jubata.

The feeding of the larvae always begins at the tip of the leaf and
continues down approximately Vs of the length to the plant base. If there
are several larva in the plant, then the surface is eaten away quite evenly.
The larvae of Hipparchia azorina were found on Faial island at the
southeastern slopes of Caldeira at 900 m altitude on July 1, 1980, in the
grass cushioning of Festuca jubata Lowe. Up to five larvae were found on a
single plant, but usually one finds only one or two larvae per plant. The
larvae appear to favor plants which grow in wind protected rivulets or
valleys. Marsden & Wright (1971) have already pointed out the strong
feeding activity of the larvae upon the grass vegetation. Economic damage
by Hipparchia larvae to the grass vegetation is not cited. It appears likely
that the larvae of all the Hipparchia populations on the various Azores
islands feed monophagously on Festuca jubata, which grows only in
montane regions. On the coast it is replaced by Festuca petraea Seub, On
Sao Miguel, F aial and Flores several females were taken for egg laying, the
eclosed larvae being available during the author’s visit to the Azores. In
breeding these larvae, first on Madeira and later in Germany, the problem
of foodplants presented itself. Festuca jubata was no longer available, and
Festuca ovina L., which occurs in Germany, was not accepted as nutriment.

Table 1. Summaiy of characteristics of the Azore Island taxa of Hipparchia.

20(3): 136T60, 1981(83)

151

8 J
a ®

S S
S 1

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8 a

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Q OJ

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£ «
M p

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3 O ^ §

03 03

S

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> a

X

S ^
•S ®
^ o

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« ^

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® a
-a

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f X o
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5-4 d

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152

J. Res. Lepid.

Festuca scoparia Kerner et Hack, from the Pyrenees, was the first plant the
larvae accepted. All the bred specimens have been reared with this plant.
Behavior of Larvae

The larvae are nocturnal. On July 1, 1980, toward 1800 hours on Faial
island I found L5 instars of K azorina hidden in the cushioning grass.
Sometime after dusk the larvae wander out of the middle of the plants in
order to eat from the tip of the leaves. In such fashion they work their way
down towards the base of the plant, minimizing visible evidence of damage
and their presence. During the day the larvae hide in the grass with their
heads facing upward. In cases of heavy feeding by the larvae, up to Vs of the
total height of the cushioning grass can be eaten away. While still dark,
after feeding, the larvae wander back inside the grass cushion.

Behavior of Adults

The flight of the butterfly begins as soon as the sun breaks through in the
early morning hours. The butterflies interrupt their flight during sudden
rainstorms, but continue to fly during Ught drizzles. When adults rest they
prefer to settle down on plants. The butterflies always fly on the regions of
the mountains which are protected from the wind. The flight is short and
after a short flight the butterflies land again. The butterflies take a slanting
position of 45° with their wings shut as they alight. They fly always more or
less close above the ground vegetation. Oviposition was not observed in
nature, but I assume females deposit eggs on the plant. On the islands of
Sao Miguel, Faial and Flores, I observed the flight of the butterflies by and
large ceased after 1800 hours.

dorsal line

20(3): 136-160, 1981(83)

153

Parasitoides and Predators

No evidence of parasitoides were found from the approximately thirty
larvae collected on Faial island. It is not known whether the eggs are
subject to parasitism. Walker (1931) described some imagines from the
Pico island as having parts of the wings bitten off, and assumed that the
bites were caused by Serinus canaria (Linnaeus, 1738). It would appear
that Motacillacinereapatriciae (Vavrie, 1957) could use if. azorina as their
diet includes insects. Predators as Lacerta spec, do not occur in the
habitats of the H. azorina complex. There are not sufficient data at present
to make judgments on the role of either predation or parasitization in
regulating the populations of these butterflies.

Protection of the Species

On all of the Azores islands the intensive use of land for pasture is of
great economic importance to local agricultural production. As a result
large areas of naturally growing fescue grass vegetation are often broken
up by bulldozers in order to be replanted with pasture grasses for higher
productivity. The Azores Hipparchia species cannot, however, live in
these cultivated pastures. It is reasonable to assume that in the middle of
the 15th century, when the Azores were first colonized, that the islands
were covered to a great extent with woods. The present environment of the
Hipparchia populations on the islands can be seen as secondarily arising
after the initial clearing of the woods by the early settlers. As a result,
Festuca jubata was probably able to strongly expand, and the Hipparchia
populations during that period found a still better environment. Through
today’s methods of agriculture, this development is regressing, and the
pasture environment is now being supplanted by artificially selected
grasses for agricultural reasons.

An extensive use of pasture of the natural fescue grasses, as has always
been done in the past, did not harm the prospering of the Hipparchia
population. The method best suited for the conservation of the butterflies
is the conservation of initially produced native pasture habitats, most
appropriate biotopes being those with a southern to southeastern
exposure. The most suitable habitats on Sao Miguel are Gafanhoto, 715 m,
and the regions near Pico da Vara. On Faial island the slopes of Pico Gorda
and the Caldeira fit the described conditions, and on Flores the best areas
are the slopes of Pico dos Sete Pes. For the islands of Sao Jorge and Pico
there are no data available yet that species habitats require protection. In
any case, care should be taken that such habitats are not allowed to
become afforested with Cryptomeria japonica (L.F.) D. Don., which is
employed in the Azores.

Conclusions

1. Comparative investigations of morphology of adults and particularly.

154

J. Res. L^pid.

of the early stages, provided the basis for a revision of the taxa of
Hipparchia azorina complex.

2. H. azorina Strecker 1899 is present only on the central group of the
Azores. The type locality is restricted to Pico island. H. azorina ohshimai
comb. nov. stat. nov. inhabit Faial island, on Sao Jorge one finds H.
azorina jorgense ssp. n.

3. H. miguelensis LeCerf 1935 stat. rev. comb. nov. inhabits Sao
Miguel island.

4. H. caldeirense sp. n. inhabit Flores island.

5. The habitat of the populations from Sao Miguel, Faial and Flores
identified of the Festuca jubata zone. The foodplant of the larva is Festuca
jubata Lowe., although record of the foodplants on Sao Jorge and Pico has
yet to be brought forth.

6. All populations are monovoltine, hibernation taking place during the
larva stage.

7. The continuing expansion of the area devoted to modem agricultural
production necessitates protection of the species.

Acknowledgments: F or their support of this work I would like especially to express
my thanks to the following persons. Dr. 0. Kudma (Forschungsinstitut und
Museum Alexander Koenig, Bonn) for the examination of the genitalia of the H.
azorina complex and the reading and critical comment of the manuscript. Prof. Dr.
D. Lupnitz (Botanisches Institut der Universitat Mainz) for the identification of
Festuca jubata and Laurus azorica. Dr. L. D. Miller (Allyn Museum of Entomology,
Sarasota, Florida, U.S.A.) for the photographs and loan of the abdomen of the
holotype from H. azorina. Prof. Dr. C. Naumann (Universitat Bielefeld) who
provided for the stereoscan pictures of the eggs of H. azorina. To Mr. Duarte Soares
Furtado, Vila Franco do Campo, Sao Miguel, Azores, I am grateful for six H. azorina
females from Sao Jorge. Mr. Dalberto Teixeira Pombo, Santa Maria airport, who
along with the members of the Centro dos Jovens Naturalistas collected specimens
of H. azorina for me on Sao Jorge under difficult circumstances. In addition to
kindly making one female and two male specimens available to me, Mr. P. Ackery
(British Museum of Natural History London) made slides of the wing scales and the
color slides of the specimens from Sao Jorge. My appreciation goes as well to Mr.
Gomez Vieira, Flores, Azores, for information. I would also like to thank Miss
Maijorie Sonia Mawby for her efforts in translating the manuscript from German
into English.

After the completion of this work in May 1982, Hipparchia specimens from the
Azores were sent to the following addresses: Allyn Museum of Entomology,
Sarasota, Florida, U.S.A. ; Mr. W. L. Blom f , Groningen, NL; Dr. O. Kudma, Bonn;
Prof. Dr. C. Naumann, Bielefeld; and W. Schmidt-Koehl, Saarbmcken, the
remainder are in my collection.

20(3): 136-160, 1981(83)

155

Y

V

V

Y

Sao Jorge

Faial

Pico

Sao Miguel

Fig. 3. Androconial scales of the Azores Hipparchia taxa: Sao Jorge: Hipparchia
azorina jorgense; Faial: Hipparchia azorina ohshimai; Pico: Hipparchia
azorina; Sao Miguel: Hipparchia miguelensis.

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(Azoren). Bot. Jahrb. Syst., 95(2):149-173.

, 1975c. Die vertikale Vegetationsglie derung auf der Insel Pico, Azoren.

Cuad. Bot. Canar., 23/24:15-24. Las Palmas.

20(3): 136-160, 1981(83)

157

MARSEND, c. A. & P. L. WRIGHT, 197 1 . A note on the distribution of the Rhopalocera on
the Island Sao Jorge, the Azores. Entomologist’s Rec. J. Var., 83:177-186.

MAYR, E., 1967. Artbegriff und Evolution. Parey. Hamburg.

OEHMIG, s., 1977. Die Tagfalter Madeiras. Ent. Z., Frankf. a.M., 87:169-176,
189-199.

, 1982. Schattenfrei Insekten Makroaufnahmen mit nur einem Elek-

tronenblitz. Ent. Z., Frankf. a.M., 92:182-184, 199-200.

REBEL, H., 1917. Siebenter Beitrag zur Lepidopterenfauna der Canaren. Ann. K. K.
nat. Hist. Hofmus., 31 (1-4): 1-58.

, 1939. Achter Beitrag zur Lepidopterenfauna der Canaren. Ann. K. K.

nat. Hist. Hofmus., 49:43-68.

, 1940a. Die Arthropodenfauna von Madeira nach den Ergebnissen der

Reise von Prof. Dr. 0. Lundblad Juli-August 1935. XXni. Die Lepidopteren-
fauna Madeiras. Ark. ZooL, 23A(5):1-13.

, 1940b. Die Lepidopterenfauna des Azorischen Archipels. Im Anhang:

Eine Lepidopteren Ausbeute von Madeira. Soc. Sci. Fenn., Comm. Biol., 8(1):
1-59.

SHIROUZU, T. & A. KARA, 1979. Early stages of the Japanese Butterflies in colour.
2 Vols., (11th reprint). Osaka.

STRECKER, H., 1899. Lepidoptera Rhopalocera and Heterocera Indigenous and
Exotic. Suppl. 2:1-3, Reading, Pa. U.S.A. Published by author.

TINBERGEN, N., 1941. Ethologische Beobachtungen am Samtfalter, Satyrus semele
L. J. Orn. Lpz., 89 (Sonderheft):132-144.

WALKER, J. J., 1931. Notes on a Satyrine Butterfly, Satyrus azorinus Strecker
from the Azores. Proc. ent. Soc Lond., 5:77-81.

WOLFF, N., 1975. On the sudden mass occurrence in 1974 of PierisrapaeL. (Lepidop-
tera, Pieridae) in Madeira. Bohn. Mus. munic. Funchal, 29:26-32.

Fig. 5. Head diameter of the larva.

158

J. Res. Lepid.

Fig. 6. Male genitalia. 1. Hipparchia caldeirense, 2. Hipparchia miguelensis, S.Hipparchia
azorina ohshimai, 4. Hipparchia azorina, 5. Hipparchia azorina jorgense, 6. Holotype.

20(3): 136-160, 1981(83)

159

Fig. 7. Female genitalia. 1. Hipparchiacaldeirense, 2. Hipparchiamiguelensis, S.Hipparchia
azorina ohshimai, 4. Hipparchia azorina jorgense.

160

J. Res. Lepid.

Fig. 8. Pico island, north side, view from Lagoa do Capitao 800 m, to the Pico 2351 m, only single males ofH. azorina were observed on
this high plateau.

Journal of Research on the Lepidoptera

20(3): 161-169, 1981(83)

Courtship Behavior of the Dainty Sulfur Butterfly,
Nathalis iole with a Description of a New, Faculative
Male Display (Pieridae)

Ronald L. Rutowski

Department of Zoology, Arizona State University, Tempe, AZ 85287

Abstract. The components and temporal structure of courtship leading to
copulation are described for the dainty sulfur, Nathalis iole (Boisduval).
Most successful courtships were similar to those described in the literature
for several other pierids. However, in 22% of the successful courtships the
male performed a previously undescribed wing spread display in which he
alit in front of and facing away from a perched female and assumed a
stationary posture with his wings fully spread. This display is elicited by the
performance of initial rejection responses by the female. The proximate and
ultimate causes of this faculative male display are discussed.

Introduction

In butterflies, male courtship behavior is viewed as having evolved in
response to mate choice by females to insure sex and species identity, and
quality of a potential mate (Scott, 1972; Silberglied, 1977; Rutowski,
1982). Because errors in the selection of mates can have severe negative
effects on a female’s reproductive success, it is expected that all males
successful in courtship should be required by females to produce basically
similar performances. While collecting data for another study, observa-
tions on the courtship of the dainty sulfur, Nathalis iole Boisduval,
revealed the existence of a male display that had heretofore been
undescribed and surprisingly was not performed in all successful court-
ships. This report describes the courtship of N. iole with an emphasis on
this new display and the contexts in which the display occurs in the hope of
discovering something of its proximate and ultimate functions.

Methods

The dainty sulfur was studied at the Archbold Biological Station 13 south
of Lake Placid, Florida, from July to November, 1981. There it flies all
year and is most common where Bidem pilosa Powell and Turner, its larval
foodplant and an adult nectar source, is abundant.

Virgin females were obtained by rearing from eggs. The eggs were
collected on sprigs of B. pilosa that had been placed in small (8x8x8 cm)
wire cages with field-caught females. The cages were placed outside in full

162

J. Res. Lepid.

sun to induce females to oviposit. The larvae were fed on fresh cuttings of
B. pilosa in a laboratory where the light-dark regimen and humidity were
variable and the temperature ranged from 27-29°C.

To observe courtship behavior, particularly that which preceded copula-
tion, virgin females that varied in age from freshly emerged to no more than
3 days of age were released near free-flying males in the field. (Females not
used on day of emergence were stored at 4°C after their wings hardened
until use.) Durations of courtships leading to copulation were measured
with a stopwatch and represent the time from when the male first arrived
(within 2 cm of the female) until the pair had coupled and the male stopped
moving. Written records of the sequence of events observed were also
made. I especially noted (1) whether the female was flying or perched when
the male first approached, (2) whether the female performed a flutter
response or mate refusal posture when alighted, (3) whether the male
performed a wing spread display, (4) any perch changes by the female after
alighting, and (5) whether the male had his wings spread when attempting
copulation. When the wing spread display was observed the relative
positions of the male and female were noted. Details on the criteria used to
judge the occurrence of these behavior patterns are given in “Results.”

Courtships staged within outdoor cages were recorded on film using
techniques described by Rutowski (1978 and 1979). In spite of numerous
attempts, I was not successful at recording a wing spread display on
motion picture film, hence detailed information on its temporal structure
was not acquired.

In both the field and the cages, mating pairs were separated within a
minute of coupling so that the female could be used again in observations
of successful courtship. Normal copulation lasts about 15 to 20 min.
Throughout this report “virgin female” refers, in addition to the obvious,
to females that have been previously but briefly coupled with a male.

Throughout the paper parametric summary statistics will be presented
as: mean ± one standard deviation (sample size). The 0.05 level was used
in making all decisions regarding statistical significance.

Results

A. Field Observations: Components of Successful Courtship

Fifty-four successful courtships (= leading to copulation) were observed
in the course of releasing 33 virgin females near males in the field. In the
descriptions and data that follow no female accounts for more than 3
successful courtships. The most typical sequence of events was as follows.

After release and while still flying, the female was approached by the
male. The female then dropped quickly to the ground and alit on a leaf,
grass blade, or the soil. While the female sought out a perch the male either
followed her closely or, in some cases, hovered about 10 to 20 cm over the
female. Once the female had perched, she did not move except to extend

20(3): 161-169, 1981(83)

163

her abdomen out from between the hindwings. The male quickly alit next
to the female and oriented head-to-head and tail-to-tail with the female.
He then curled his abdomen toward the female’s, and brought his genitalia
into contact with hers which marked the beginning of copulation. This
sequence of events was observed in 24% of the successful courtships
observed (Table 1).

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164

J. Res. Lepid.

Variations in this basic pattern arose when: (1) the female was already
perched when the male approached to within 2 cm, (2) the female
performed a flutter response or a mate refusal posture when perched, (3)
the male performed a display hereafter referred to as the wing spread
display, and/or (4) the male attempted copulation with his wings
noticeably spread to an angle of about 120-150 degrees. The frequency of
occurrence of courtships with these components is given in Table 1. A
flutter response (Obara and Hidaka, 1964) was recorded any time a
perched female performed at least one wing flick [” a rapid opening and
closing of the wings (Rutowski, 1978)] while the male courted her. The
mate refusal posture (Obara, 1964) was recorded when, in response to the
male’s presence, a female assumed a posture with her wings spread and
her abdomen raised so that its long axis stood perpendicular to the plane of
the female’s wings. Rarely (n ^ 7), females changed their perch after
alighting by resuming flight. In subsequent analyses these perch changes
were grouped with the flutter response and mate refusal postures as initial
rejection responses. The data reveal that perched females were no more
likely to perform rejection responses than flying females (3^ ^ = 2.68, ldf,p
< 0.1).

The wing spread display of the male (Fig. 1) was recorded when, after the
female had perched, he alit facing away from the female but no more than 3
cm in front of her. Once on the substrate, the male spread his wings and
held them fully opened for several seconds during which time the wings
quivered slightly. Also, during the time the wings were spread the
forewings were held a little forward of and above the hindwings, enough to
clearly expose the red-orange sex brand on the dorsal surface of the male’s
hindwing (Klots, 1951). Wing spread displays were performed in 22% of
the observed courtships. They were also observed in some unsuccessful
courtships. In some but not all courtships without the wing spread display,
the female alit on a vertical grass blade, in dense vegetation, or in some
other location that made it impossible for the male to perform a wing
spread display due to the lack of suitable substrate. However, it was not
possible to quantify the frequency of this occurrence because of difficulty
in establishing exactly what constitutes suitable substrate for a wing
spread display.

Whether the female is initially flying or perched has no effect on the
likelihood that the male will perform a wing spread display (Table 1).
Thirty percent of the perched females and 12.5% of those flying when
intially approached by males, were courted with wing spread displays. The
difference was not significant (3^^ == 2.36, 1 df , p = 0.12). However, if the
female performed a rejection response (includes perch changes by female
during courtship), a male was significantly and almost 4 times more likely
to perform a wing spread display. Of the rejecting females, 40% elicited
wing spread displays from the courting males; of females that did not, only

20(3): 16M69, 1981(83)

165

F%. 1 . Courtship behavior in the dainty sulfur, Nathalis iole. The male (wing span
'^2.3 cm) has landed in front of and facing away from the female and is
performing the wing spread display. This figure was drawn from a 35 rmn
photograph.

10.8% elicited the display (3|[^ — 5.81, 1 df, p ” 0.02). The performance of a
wing spread display had no effect on the likelihood that a male would
subsequently attempt copulation with hie wings spread ^ ^ ^ 1.24, p =
0.26). Males that were accepted in copulation by females were examined
and assessed with respect to forewing length and wing wear. Males that
performed wing spread displays were not different from those that did
with respect to these two characters. Virtually all males were in fresh
condition (more than 80% in both groups) and had forewing lengths (base
to wing tip) of 14 mm [males that performed wing spread display: 14 ± 1
mm (9); those that did not: 13.8 + 0.62 mm (34)].

166

J. Res. Lepid.

B. Field Observations: Temporal Structure of Successful Courtship

Table 1 shows that the duration of successful courtship in N. iole is

affected by its form. Initial observations indicated that the position of the
male’s wings (spread or not spread) during copulation attempt had no
effect on the duration of the courtship. To test this hypothesis, I compared
courtships that varied with respect to the male’s wing position during
copulation attempt but in which the female did not perform a rejection
response, was initially perched and remained so, and was not courted with
awing spread display. The duration of courtships in which the male did not
spread his wings during the copulation attempt [5.33 + 1.63 sec (6)] was
not significantly longer than those in which the male did spread his wings
[5.2 ± 1.48 sec (5); t ^ 0.14, 9 df, p = 0.45). In all analyses presented
below, data for courtship with and without this male behavior pattern were
pooled.

As a baseline duration I used the mean duration of successful courthsips
in which the female (1) was intially perched, (2) performed no rejection
responses, and (3) was not courted with a wing spread display. This
baseline duration was 5.27 + 1.49 sec (11) and was used to study the effect
of variation in form on the duration of courtship. When the female was
initially flying but did not perform a rejection response and was not
courted with a wing spread display, the duration of courtship [11 ± 5.71
sec (14)] was significantly longer than the baseline duration (t — 3.23, 23
df , p “ 0.002). Females flew for about 5 to 6 sec on the average before
alighting if not perched when the male first approached.

K an initially perched female performed a rejection response but was not
courted with a wing spread display, the duration of courtship [9.13 + 4,05
sec (8)1 was significantly longer than the baseline (t = 2.92, 17 df, p™
0.005). This was also true if one eliminates from successful courthsips with
a rejection response those in which the female changed perch after the
male’s approach. Although the mean duration of this group was a little
shorter [7.8 ± 4.27 sec (5)] these courtships were still significantly longer
than those without a flutter response or mate refusal posture (t ^ 1.8, 14
df, p = 0.047). Hence, rejection responses typically increased the duration
of courtship by a factor of IV2 to 2.

Male wing spread display also increased the duration of successful
courtship. Courtships identical to the baseline courtships, but including
male wing spread display had a mean duration ofl3.6±6.11 sec (3), which
was significantly longer than the baseline duration (t=4.5, 12df, p =
0,001). These data indicate that the wing spread display has an average
duration of about 8 sec.

C. Film Records

Using 7 lab-reared virgin females, I recorded 16 successful courtships on
motion picture film. In the film records, all females were initially perched

20(3): 161-169, 1981(83)

167

(an artifact of the techniques used) but none were courted with wing
spread displays. Females performed flutter responses or mate refusal
postures in half of the successful courtships recorded. Restrictions on
camera movement prohibited the recording of courtships in which the
female flew to a new perch. Only 14 of the courtships recorded were
complete; all of the analyses that follow are based on information from
these 14.

The mean duration of all courtships was 5.03 ± 3.64 sec (14). However,
unlike the situation in the field, the duration of courtship without rejection
responses [3.93 ± 3.13 sec (8)] was not significantly shorter than that for
courtships with rejection responses [6.07 ±4.01 sec (6); t = -1.125, 12 df,
p = 0.14]. In spite of this difference between field and cage courtships the
film records generally confirm the accuracy of the field measurements.
Courtships recorded on film without rejection responses by females were
identical in form to the baseline courtships and were not significantly
different from them in duration (t = 1.24, 17 df, p ™ 0.12). Similarly,
courtships recorded on film with rejection responses by females were not
significantly different in duration from those of identical form measured in
the field (t = 0.425, 8 df, p = 0.34). However, in both comparisons the
difference was in the direction of the film records yielding shorter
durations than the field records. This is to be expected in light of the
inaccuracies inherent in timing such rapid events with a stopwatch.

In these 16 courtships males showed no preference in the side of the
female from which they attempted and achieved copulation (male to
female’s right: 6 courtships; male to female’s left: 10 courtship s;*^^ =
1.0, 1 f, p = 0.32).

Discussion

Over 70% of the successful iV. iok courtships observed during the field
portion of this study and all courtships recorded on film were basically
similar in temporal and sequential structure to that described for several
other pierids (Peterson and Tenow, 1954; Rutowski, 1978, 1979; Silberglied
and Taylor, 1978; Suzuki, 1977). These courtships were characterized as
rapid affairs, lasting a few seconds, in which the male buffets the female
with his wings and body, the female extends her abdomen in response, and
the male then alights and attempts copulation. However, in the other 22%
of the successful courtships reported here, the male behaved in a way that
has not been previously described for any pierid; he assumed a stationary
posture in front of the female with his wings spread. Because such a
display is so different from any prior observation and because it is a
facultative part of successful courtship, questions immediately arise
concerning its proximate and ultimate functions.

The proximate function of the display appears to be to provide the
female with information about the male not garnered by her during the

168

J. Res. Lepid.

intial phases of courtship. An initial rejection response by a female
increases the likelihood that the male will perform a wing spread display.
Both the rejection response and the display almost double the duration of
courtship over what it is without them. Hence, females that perform
rejection responses gain time and a display with which to make a more
complete assessment of a potential mate. The exact nature of the signals
involved in this assessment are not known but could involve visual signals
including the lack of ultraviolet reflectance and the black bar on the dorsal
forewing (Rutowski, 1977), and/or chemical signals that probably arise
from the male’s sex brand on the dorsal hindwing (Vetter and Rutowski,
1978).

What information is the female gathering about the male during the
display? It seems most likely that the female is assessing the male’s (1)
sexual identity, (2) species identity, and/or (3) quality as a mate relative to
other conspecific males. Both female and male pierids are known to
approach and chase conspecifics on occasion (Rutowski, 1980; Rutowski
et al., 1981). Several observations were made during this study of N. iole
females engaging in such behavior. The wing spread display may be a way
for females to confirm the sex of the courting animal by the performance of
the display and by visual and chemical signals enhanced by the display.
Information about the species identity of a courting male may also be
important to the female in that N. iole is sympatric over a large part of its
range with Eurema daira Godart and E. lisa Boisduval and LeConte, two
species of small sulfurs closely related and visually and behaviorally
similar to N. iole (Rutowski, 1977, 1978). Again, the display may provide
visual, chemical, and behavioral confirmation of a male’s species identity.
Finally, the display may provide the female with information about the
male’s age, size, persistence, or other characteristics that could be
indicative of his overall genetic quaUty and ability to invest in the female’s
offspring. Male butterflies are known to pass nutrients to the female
during copulation that she may use in oogenesis (Boggs, 1981; Boggs and
Gilbert, 1979; Boggs and Watt, 1981). Rutowski (1982) has reviewed the
Lepidoptera for some of the characteristics of males that may be
important in selection among conspecific males by females.

In summary, the male display appears to have evolved as a way of
delivering information to females who are initially unreceptive. Exactly
which portion of the information presented to the female by the display is
most important to her is as yet unclear. However, by comparing the
behavior of N. iole with that of its sympatric and very similar relatives E.
lisa (Rutowski, 1978) and E. daira (unpubl. data) it may be possible to
evaluate the three hypotheses about its ultimate function proposed in the
previous paragraph.

Acknowledgments. My sincere thanks to: the Director and staff of the Archbold
Biological Station for the use of their facilities and equipment, David Smith for his

20(3): 161469, 1981(83)

169

invaluable assistance in the field, Patricia Rutowski for Fig. 1, and the National

Science Foundation for financial support during the study (Grant No. BNS 80-

14120).

Literature Cited

BOGGS, C. L., 1981. Selection pressures affecting male nutrient investment at mating
in heliconiine butterflies. Evolution. 35: 931-940.

BOGGS, c. L. & L. GILBERT, 1979. Male contribution to egg production in butterflies:
evidence for transfer of nutrients at mating. Science. 206:83-84.

BOGGS, c. L. & w. B. WATT, 1981. Population structure of pierid butterflies. IV. Genetic
and physiological investment in offspring by male Colias. Oecologia. 50: 320-
324.

KLOTS, A., 1951. A field guide to butterflies. Houghton Mifflin Co., Boston.

OBARA, Y., 1964. Mating behavior of the cabbage white, Heris rapae crucivora. H.
The ‘mate-refusal posture’ of the female. Zool. Mag. (Dobutsugaku Zasshi)
73:175-178.

OBARA, Y. & T. fflDAKA, 1964. Mating behavior of the cabbage white, Pieris rapae
crucivora. I. The ‘flutter response’ of resting male to flying males. Zool. Mag.
(Dobutsugaku Zasshi) 73:131-135.

PETERSON, B. & o. TENOW, 1954-. Studien am Rapsweissling und Bergweissling {Pieris
napi L. und Pieris bryoniae 0.). Isolation und Paamngsbiologie. Zool. Bids.
Uppsala 30:169-198.

RUTOWSKI, R. L., 1977. The use of visual cues in sexual and species discrimination by
males of the small sulphur butterfly, Eurema lisa (Lepidoptera: Pieridae). J.
Comp. Physiol. 115:61-74.

, 1978. The courtship behaviour of the small sulphur butterfly, Eurema
lisa (Lepidoptera: Pieridae). Anim. Behav. 26:892-903.

, 1979. Courtship behavior of the checkered white, Pieris protodice
(Pieridae). J. Lepid. Soc. 33:42-49.

, 1980. Courtship solicitation by females of the checkered white butterfly,
Pieris protodice. Behav. Ecol. Sociobiol. 7:113-117.

1982. Mate choice and lepidopteran mating behavior. Florida Ent. 75:

72-82.

RUTOWSKI, R. L., c. E. LONG, L. D. MARSHALL, & R. s. VETTER, 1981. Courtship solicita-
tion by Colias females (Lepidoptera: Pieriedae). Amer. Midi. Nat. 105:334-340.

SILBERGLIED, R. E. & o. R. TAYLOR, JR., 1978. Ultraviolet reflection and its role in the
courtship of the sulphur butterflies Colias eurytheme and C. philodice (Lepidop-
tera: Pieridae). Behav. Ecol. Sociobiol. 3:203-243.

SUZUKI, Y., A. NAKANISHI, H. SHIMA, 0. YATA & T. SAIGUSA, 1977. Mating behaviour of
four Japanese species of the genus Pieris (Lepidoptera, Pieridae). Kontyu
(Tokyo). 45: 300-313.

VETTER, R. s. &R. L. RUTOWSKI, 1978. External sex brand morphology of three sulphur
butterflies (Lepidoptera: Pieridae). Psyche. 86:383-393.

Journal of Research on the Lepidoptera

20(3): 170-173, 1981(83)

On the Supernumerary Chromosomes of
Tarache tropica Guen. (Lepidoptera: Noctuidae)

P. K. Mohanty
and

B. Nayak

Department of Zoology, F. M. College, Balasore 756 001, Orissa, INDIA

Abstract. The karyotype of Tarache tropica Guen, consists of 1 to 7 minute
dot-shaped supernumerary chromosomes in addition to the normal haploid
complement, n— 31. The exact nature of the accessory chromosomes,
whether they are true supernumeraries or are the unpaired nonhomologous
elements produced through interspecific hybridization, is not clear, though
the latter explanation appears to be more probable.

Introduction

Many species of plants and animals show one or more supernumerary
chromosomes, characteristically different from ordinary chromosomes in
their karyotype (Wilson, 1925; White, 1973; Jones, 1975). In Lepidoptera,
however, reports on the number of species showing such elements are
rather scanty (Dederer, 1928; Beliajeff, 1930; Federley, 1938; Lorkovic,
1941; Maeki, 1953; de Lesse, 1960a; Bigger, 1976; Rao and Murty, 1976;
Nayak, 1978). Indeed, it is difficult to distinguish genuine B-chromosomes
since the lepidopteran karyotype often includes a large number of small
dot-like compact chromosomes which yield very little information about
their exact morphology. An attempt has been made here to analyze the
nature of such supernumerary elements in the karyotype of Tarache
tropica, a species cytologically investigated for the first time.

Material and Methods

The larvae of Tarache tropica were collected from their host plant
(unidentified) in the close vicinity of Bhubaneswar during July-August,
1976. These were reared in cages in the laboratory. The mature male
larvae (5th instar) and early pupae provided suitable testes material for
the study of spermatocytic chromosomes. Temporary preparation of the
material using Bellings’ acetocaramine was tried. For permanent prepara-
tion, the larval sexual organs were dissected in a colchicine-hypotonic
solution (0.01% Colchicine in 0.45% of Sodium Citrate solution), followed
by washing in fresh hypotonic solution for 30-40 minutes at room
temperature. The testis was then transferred to a slide which was slightly

20(3): 170-173, 1981(83)

171

tipped to remove the hypotonic solution completely. The material was
then torn into pieces in a drop of 60% aceto-ethanol (1:3) fixative and then
squashed. The slide was next placed in a freezing chamber of a freezer for
2-3 minutes and the cover glass removed with care using a razor blade. The
frozen preparation was then subjected to gently blowing air over it. The
slide was then transferred to absolute glacial acetic acid for % to 1 minute
to remove cytoplasmic staining and the slide was again air dried. The
preparation was stained with Giemsa, diluted about 50 times in Sorensens’
phosphate buffer (pH 7) for 8-10 minutes at room temperature.

Observations

The diploid chromsomes, as revealed in some of the spermatogonial
metaphase stages of T. tropica, were 2n=62. The individual chromosomes
were dot-like, almost isodiametric and compact bodies yielding no
information regarding the position of the centromere. Morphologically,
the gonial chromosomes were not differentiable into sex-chromosomes
and autosomes. The haploid chromosome number as established from a
count of pro-metaphase or metaphase I chromosomes was n— 31. Meta-
phase I bivalents appeared oval in a polar view and dumbbell-shaped in a
equatorial view. Late diakinesis or pro-metaphase plates and metaphase I
cells of different specimens when compared showed variation in chromo-
some numbers from specimen to specimen and from follicle to follicle of
the same individual. In addition to the normal karyotype of n— 31
bivalents, a variable number of minute chromosomal elements, much
smaller than the normal bivalents, was encountered. The uneven distri-
bution, irregular number, and minute size all indicated that they were not
homologous to any of the members of the karyotype. They appeared to be
univalents without any homologous partner. In 6 of the 21 individuals
investigated, only one supernumerary element was observed in the
karyotype. In five of these, the supernumerary chromosome was stable
and was observed in every meiocyte, while in the sixth, some (possibly of
one cyst) had single m-chromosome each which was not seen in the
remainder. Metaphase II plates were rare. The m-chromosome frequency
in other specimens was high, 3 to 7 elements were encountered at
metaphase 1. They were smaller than the regular chromosomes and
showed irregular segregation at first anaphase. The exact nature of these
minute elements, whether they are supernumeraries or are unpaired
elements due to interspecific hybridization, is not clear. However, their
uneven distribution indicates their supernumerary nature.

Discussion

In the absence of direct evidence, the origin of supernumeraries is highly
enigmatic. White (197 3) assumes their origin to be through fragmentation
of heterochromatic blocks of normal chromosomes. Robinson (1971), as

172

J. Res. Lepid.

• • •

m • • .

1

3

Fig. 1. Spermatogomal mitosis showing 62 dot-like elements.

Fig. 2. Metaphase I showing 31 bivalents and 7 extra elements of supernumerary
nature.

Fig. 3. Metaphase I with 3 m- chromosomes.

well, holds them to be small segments of normal chromosomes undergoing
accidental breakage and showing erratic meiotic behaviour due to their
acentric nature. In Lepidoptera, where fragmentation is a common aspect
of chromosome evolution, Robinson speculates that the occurrence of
supernumeraries is no doubt a special case where an element is produced
which behaves differently from a chromosome which has fragmented into
two stable bodies, each of which behaves sufficiently normal to become a
part of the karyotype. Jones (1975) is of the opinion that sex-chromosomes,
particularly the X-chromosomes, are very likely the ancestors of micro-
chromosomes. Bigger (1976) observed the karyotype of two British Pirns
species to be identical in cells with and without B-chromosomes. Based on
this observation, he concludes that supernumeraries are true additional
chromosomes and not small fragments of the normal karyotype, as
hypothesized by White (1973) and Robinson (1971). We agree with the
last two authors that numerical variation may arise due either to accidental
breakage of normal chromosomes or mixing of chromosomes from
different nuclei. In the present case of Tarache tropica, the occurrence of 1
to 7 supernumeraries may well be due to interspecific hybridization in
nature and may therefore present non-homologous unpaired univalents.
Rao and Murty (1976) hypothesize that B-chromosomes confer an
advantage on the population. This may not be true at least in the case of
Tarache tropica, since a variable number of such supernumeraries occur in
different follicles of the same individual, as well as in follicles of different
individuals at the same time.

Acknowledgments. We thank Professor G. C. Das, Department of Zoolo^,
Berhampur University, for valuable suggestions and one of us (P.K.M.) thanks
V.G.C., New Delhi, for the financial assistance.

20(3): 170-173, 1981(83)

173

Literature Cited

BELIAJEFF, N. K, 1930. Die Chromosomenkomplexe und ihre Bezeihung zur Phylo-
genie bei Lepidopteren. Z. Ind. Abst. Vererbgsl. 54:369-399.

BIGGER, T. R. L., 197 6. Karyotypes of three species of Lepidoptera including an inves-
tigation of B-chromosomes in Pieris. Cytologia 41:261-282.

DEDERER, P. H., 1928. Variation in the chromosome number in the spermatogenesis
of Philosamia cynthia. Jour. Morph. 45:599-613.

FEDERLEY,H., 1938. Chromosomenzahlen finnlandischer Lepidopteren. I Rhopalo-
cera. Hereditas 24:221-269.

JONES, R. N., 1975. B-chromosome systems in flowering plants and animal species.
Int. Rev. Cytol. Vol. 40:1-100.

LESSE, H. de, 1960a. Speciation et variation chromosomique chez les Lepidopteres
Rhopaloceres. Ann. Sci. Nat. Zool. XII. Ser. 2:1-223.

LORKOVic, z., 1941. Die Chromosomenzahlen in der Spermatogenese der Tagfalter.
Chromosoma 2:155-191.

MAEKI, K., 1953. A chromosome study of Japanese butterflies. Kwansei Gakuin
Univ., Annual Stud. 1:67-70.

NAYAK, B., 1978. Supernumerary m-chromosomes in two species of Lepidoptera.

Joum. of Entomology Research, Vol. 2, No. 1:112-114.

RAO, N. N. & A. S. MURTY, 1976. B-chromosomes in Danaus limniace (Lepidoptera:

Danaidae). Microbios Letters, Vol. 3, No. 10.

ROBINSON, R., 1971. Lepidoptera Genetics, Pregamon Press, Oxford.

WHITE, M. J. D., 1973. Animal Cytology and Evolution, Cambridge University Press,
London.

WILSON, E. B., 1925. The Cell in Development and Heredity, 3rd Ed. New York:
MacMillan.

Journal of Research on the Lepidoptera

20(3): 174-175, 1981(83)

An Apparent Interspecific Fi Hybrid Speyeria
(Nymphalidae)

James A. Scott

60 Estes Street, Lakewood, Colorado 80226

An apparent male Fi hybrid (Figs. 3-4) was captured in a wet meadow
near Bridgeport, Mono County, California, August 25, 1974. It is
intermediate between Speyeria (Speyeria) nokomis apacheana (Skinner)
(Fgs. 5-6) and S. (Semnopsyche) cybele leto (Behr) (Figs. 1-2). Pure
populations of both species were caught there August 7, and the hybrid
and nokomis were caught August 25.

The hybrid male may be characterized phenotypically as follows:
dorsally, l)postmedian HW black spots in cells Ms and Cui are inter-
mediate in shape (nokomis has the spots shaped like Va moons, leto's are
much thicker), and 2) there is a normal sized spot and a small spot at end of
DHW cell (nokomis has two normal sized spots, leto only one spot).
Ventrally, 3) hindwing disk and margin is light brown (the disc and margin
are yelow in apacheana, dark brown in leto), 4) silver spots are inter-
mediate in shape (round in nokomis, more crescentic in leto), 5) submar-
ginal cones of dark scales on HW and FW are partly brown and partly black
(cones are black in nokomis, brown in leto), 6) FW red flush is intermediate
in extent (flush covers most of the wing in nokomis, only half of wing in leto),
and 7) postmedian dots on FW near apex are partly black and partly brown
(dots are black often edged with green in nokomis, brown in leto).

These characters (except 1 and 4) are very different in the two species;
they are clearly intermediate in the hybrid, suggesting that it is an Fi. It is
the first hybrid described in the genus, although Paul Hammond (pers.
comm.) has a natural female hybrid S. cybele pugetensis C. & F. X S.
hydaspe rhodope (Edw.). Both hybrids are between different subgenera.
The subgenus Semnopsyche is based on only one character, the female
bursa copulatrix (dos Passos & Grey, 1945, 1947). Modem systematic
theory emphasizes the use of many characters delineating taxa (Wiley,
1981). S. cybele leto and nokomis are similar in wing pattern, although they
are somewhat different in gene frequencies of some adult body enzymes
(Brittnacher et al., 1978). The heterogametic (XY) sex (the female in
Lepidoptera) is less often represented in interspecific crosses in Lepidop-
tera in general. The occurrence of both male and female hybrids, and their
occurrence between species previously thought to be in separate subgenera,

20(3): 174=175, 1981(83)

175

Figs. 1-6. S. cybele leto, top row (Fig. 1 left, Fig. 2 right);

S. cybele leto X S. nokomis apacheana, middle row (Fig. 3 left, Fig. 4
right);

5. nokomis apacheana, bottom row (Fig. 5 left. Fig. 6 right).

Left column upperside, right column underside.

suggests that hybridization may be fairly frequent in Speyeria, but hidden

by the difficulty of identifying many similar species.

Literature Cited

BRITTNACHER, J. G., s. R. SIMS, & F. J. AYALA, 1978. Genetic differentiation between
species of the genus Speyeria (Lepidoptera: Nymphalidae). Evolution 32: 199“
210.

dos PASSOS, C. F. & L. P. GREY, 1945. A genitalic survey of Argynninae. American
Museum Novitates #1296.

, 1947. Systematic Catalogue of Speyeria (Lepidoptera, Nymphalidae)

with designations of types and fixations of type localities. American Museum
Novitates #1370,

WILEY, E. 0., 1981. Phylogenetics. The theory and practice of phylogenetic system-
atics. Wiley-Interscience. New York. 440 p.

Journal of Research on the Lepidoptera

20(3): 176-177, 1981(83)

A New Record of Vanessa virginiensis ‘‘ab, ahwashtee''

from Northern California (Lepidoptera: Nymphalidae)

Arthur M. Shapiro

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

Abstract. A new record of Vanessa virginiensis “ab. ahwashtee” is pre-
sented from Nevada County, California. The occurrence and biological
significance of this and related aberrations in Vanessa are reviewed.

The study of aberrant wing-pattems in butterflies is currently under-
going a mild renaissance, after being cast into disrepute for decades by the
nomenclatorial excesses of some European workers. Although treated by
Field (1971) with disdain, the “e/ymi” series of aberrations in the genus
Vanessa hold great biological interest because parallel and probably
homologous aberrant phenotypes occur repeatedly in at least five species,
belonging to two subgenera. They probably represent the classic pheno-
copy situation, in which the same aberrant phenotype may be produced by
a mutant gene in normal environments, or by abnormal environments
acting on a normal genotype (Shapiro, 1976). Aberrations of this series
have received names (of no standing in formal zoological nomenclature) in
V. cardui L., V. atalanta L., V. annabella Field, and V. virginiensis Drury.
They also occur in the South American V. carye Hbn. sens. sir. (J. Herrera,
personal communication).

The 'Jetcheri - muellerV' aberrations of the ''elymV series are readily
inducible in V. annabella by cold shock applied to the pupa (Dimock,
1968; Shapiro, 1976 and unpubhshed). “E/ymi” itself is more difficult to
induce in V. cardui, and most specimens are only moderately modified by
cold (Shapiro, 1975). The corresponding phenotype ''ahwashtee’' has not
been induced in V. virginiensis in over 100 trials with cold shock since 1972
(Shapiro, unpublished). Both this species and V. atalanta seem better
buffered against cold shock than the others, though European workers
have successfully manipulated V. atalanta with both heat and cold.

On 28 June 1981 a fresh, perfect male V. virginiensis “ab. ahwashtee”
was collected by the author at Lang Crossing of the South Yuba River,
Nevada County, California, on the west slope of the Sierra Nevada at
about 1500 m. Two normal individuals were seen in the same clump of
flowering A^asiac/ie nepetoides (Labiatae). The capture was verified at the
scene by Mr. Marc Mirmo and Ms. Jennie Dusheck. The specimen (Figs. 1,
2) is very similar to the original "ahwashtee” figured by Comstock (1927,

20(3): 176-177, 1981(83)

177

Fig. 1. Dorsal surfaces of Vanessa. Upper right: bred V. virginiensis , normal.
Upper left: V. virginiensis ab. '' ahwashtee'\ Sierra Nevada, CA, 28 June
198. Lower right: V. cardui, “el3nni”-like aberration induced by pupal
chilling. Lower left: V. annabella, “muelleri”-like aberration induced by
pupal chilling.

5%. 2. Same, ventral surfaces.

178

J. Res. Lepid.

pL 42, fig. 6). The bibliographic history of this name is traced by Field
(1971, p. 48). I have been unable to locate any new records ofahwashtee'’
in the past 40 years, and it is not included in recent reviews of the ''elymV
series (Shapiro, 1973, 1975; Phillips, 1971). The apparent rarity of
aberrations in V. virginiensis parallels the strong pattern canalization
observed in the laboratory and does not seem to be an artifact of its more
restricted distribution and localized abundance as compared to V.
annahella and V. carye. It is often the commonest Vanessa at high
elevations and I have seen several thousand California examples in the
past decade without ever noting a major aberration before.

The cause — genetic or environmental — of ''ahwashtee^' remains prob-
lematical, but the Nevada County example coincides with unusual weather
events. From 1 1 through 13 June 1981 there were nightly severe freezes at
Lang Crossing, with lows down to -5°C (and highs of 15°C). These freezes
were sufficient to kill much of the year’s growth of two particularly
sensitive plants, bracken {Pteridium aquilinum, Polypodiaceae) and tall
knotweed {Polygonum phytolaccaefolium, Polygonaceae) at Lang. The
weather then warmed rapidly to daily highs and lows of about 25-21° and
7°. If the 28 June animal was indeed fresh, it would probably have been a
young pupa about 11 June, and the young (8-12 hr old) pupa is known to
display maximum temperature sensitivity in laboratory experiments. An
attempt to duplicate these conditions will be made when material becomes
available.

Because the “e/ymi” series of aberrations holds such interest for both
developmental and evolutionary biologists, new records should be pub-
lished, preferably with a photograph, and female examples should be bred
from whenever possible. Perhaps the study of aberrations can be resumed
without the absurd multiplication of names, misrepresentation of reared
as wild examples, and stamp-collector competition which removed it from
biology earlier in this century; one may hope so.

Literature Cited

COMSTOCK, J. A., 1927. Butterflies of California. Author, Los Angeles, Calif. 334 pp.
DIMOCK.T., 1968. An extreme experimental aberration of Vanessa cardui. J. Lepid.
Soc. 22: 146.

FIELD, w. D., 1971. Butterflies of the genus Vanessa and of the resurrected genera
Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smithson. Contrib. Zool.
84: 1-105.

PHILLIPS, w. L., 1 97 1 . Rare aberrant forms of Utah Cynthia, the Painted Lady (Lepid-
optera: Nymphalidae). Great Basin Natur. 31: 256-260.

SHAPIRO, A. M., 1973. Recurrent aberration in Cynthia annabella: a review with four
new records (Lepidoptera: Nymphalidae), Pan-Pac. Entomol. 49: 289-293.

, 1975. Natural and laboratory occurrence of "‘elymV’ phenotypes in

Cynthia cardui (Lepidoptera: Nymphalidae). J. Res. Lepid. 13: 57-62.
, 1976. Seasonal polyphenism. Evol. Biol. 9: 259-333.

Journal of Research on the Lepidoptera

20(3): 179^191, 1981(83)

The Colonization of Violets emdSpeyeria Butterflies on
the Ash-Pumice Fields Deposited by Cascadian
Volcanoes

Paul C. Hammond

2435 East Applegate, Philomath, Oregon 97370

Abstract. Violently explosive eruptions from the volcanoes of the Cascade
Range have deposited extensive ash-pumice fields that constitute an
extremely xeric habitat for most types of plants and animals. This study
examines the colonization of this habitat by n5?mphalid butterflies of the
genus Speyeria together with their larval foodplants of the genus Viola
(Violaceae). Study areas include ash fields associated with Mt. Washington,
the Three Sisters volcano system, Mt. Newberry, Mt. Mazama, Mt.
McLoughlin, and Mt. Shasta in Oregon and California. Some species of
Speyeria and Viola have proven to be vigorous colonizers of the ash-pumice
habitat, while other species have been completely excluded from this habitat
due to restrictive limitations in their physiology and ecological adaptations.
Successful colonizing species have invaded the ash fields from several
different directions: (1 .) adjacent regions of the high Cascades, (2.) the arid
lowlands east of the Cascade Range, and (3.) the Sierra Nevada and Siskiyou
Mountains south and west of the Cascades. These organisms demonstrate
how plants and associated insects may colonize and adapt to new habitats
created by volcanic eruptions in the Cascade Range.

Mtroduction

The nymphalid butterflies of the genus Speyeria are intimate associates
of the herbaceous plant genus Viola (Violaceae), since the larvae of
Speyeria feed exclusively upon the leaves and flowers of Viola. Together
these plants and animals are widely distributed throughout much of North
America, and are among the most ecologically versatile of organisms in
terms of the diversity of habitats that they occupy. These habitats range
from dense forests to open prairies; permanently wet bogs to arid deserts.
In addition, violets and Speyeria butterflies are usually very sensitive
mdicator organisms to human-caused disturbances of natural com-
munities. Indeed, many populations of these organisms have become
extinct or are threatened with extinction due to human disturbances such
as agriculture and suburban development. A good example are the
extinctions of Viola hallii Gray and Speyeria callippe Boisduval in the
Willamette Valley grasslands of western Oregon about 50 years ago
(unpublished data).

180

J. Res. LepM.

One of the most unusual habitats occupied by Viola and Speyeria are the
extensive volcanic ash-pumice fields associated with the strato volcanoes
of the Cascade Range. These volcanoes extend from Mt. Garibaldi in
southern British Columbia south to Mt. Lassen in northern California.
Most of these volcanoes have experienced many violently explosive
eruptions of the recent Mt. St. Helens type during their long histories.
During such eruptions, vast quantities of finely divided silicaceous ash and
pumice are ejected, so that adjacent areas downwind from the volcano are
buried under deep layers of this material. The most extensive ash-pumice
field was produced by the cataclysmic destruction of Mt. Mazama about
6600 years ago, and covers hundreds of square kilometers in central
Oregon extending from Crater Lake in northern Klamath County to the
Paulina-East Lakes in southern Deschutes County. Harris (1980) has
prepared a general review of the history and geological characteristics of
the Cascadian volcanoes.

As the ash-pumice fields are deposited during eruptions, most biological
life forms are completely exterminated as a result of burial and suffocation.
For example, David V. McCorkle (personal communication) is currently
monitoring the recovery of Speyeria populations near Yakima, Washington,
following the recent eruption of Mt. St. Helens in May, 1980. Prior to this
eruption, large populations of S. caLlippe and S. coronis Behr were found in
the eastern foothills of the Cascades west of Yakima. The Mt. St. Helens
eruption was relatively mild compared to many past historical eruptions of
other Cascadian volcanoes, and only 6-20 cm. of ash was deposited in the
region near Yakima in 1980. Nevertheless, almost no Speyeria were
observed in this region during 1981 except for a single specimen of S.
coronis. In dramatic contrast, the ash-pumice fields examined in this paper
were created by the deposition of several meters of volcanic material (up to
75 meters near Crater lake), which must have virtually sterilized these
areas of all pre-existing Viola and Speyeria populations. This paper wiU
examine the probable origins of Viola and Speyeria populations that were
able to invade and re-colonize these newly created volcanic habitats, plus
the ecological characteristics and adaptations of the organisms to such
habitats.

These ash-pumice fields constitute an extremely xeric habitat for plants
and animals, since this finely divided volcanic material fails to retain
moisture. In areas where the depth of ash exceeds 30 cm., there is very
little moisture available to herbaceous plants in the surface soil during the
summer growing season, with the exception of the moist areas adjacent to
creeks, lake shores, and boggy seepage areas. Most deep ash fields only
support a forest of small, shrubby Lodgepole Pine (Pinus contorta Dough).
The forest floor is mostly bairen except for scattered Bitterbrush {Purshia
tridentata Pursh) and small tufts of grasses. However, areas with only a
shallow layer of surface ash and pumice will support a more substantial

20(3): 179-191, 1981(83)

181

forest of Ponderosa Pine {Hnus ponderosa Dougi.) or White Fir (Abies
concolor Lindl. & (jord.). At the higher elevations, melting snowbanks are
an important source of moisture in the spring and early summer, and
permit the growth of some herbaceous plants such as lupines (Lupinus sp.)
and Yellow Prairie Violet (Viola nuttallii Pursh.).

Methods and Materials

During the present study, field investigations were conducted on the
volcanic ash fields of the Cascades from Mt* Washington in Linn County,
Oregon, to Mt. Shasta in Siskiyou County, California, during 1974, 1975,
1980, and 1981. Specific study areas included: ^ (1.) ash fields near Big
Lake just north of Mt. Washington in Linn County, (2.) ash fields east of
the Three Sisters volcanoes in Deschutes County, (3.) the Mazama-
Newberry ash fields extending from Crater Lake eastward to Yamsey
Mountain in Klamath County and northward to the Paulina-East Lakes in
southern Deschutes County, (4.) ash fields at the summit of the Cascades
south of Mt. McLoughlin and Lake -of -the -Woods in southern Jackson and
Klamath Counties, and (5.) ash fields around Mt. Shasta in Siskiyou
County.

In each study m-ea, the habitat, vegetation type, sun exposure, and the
consistency of the ash-pumice soil were noted (Table 1.). hidividual violet
plants were measured and observed during the course of their growing
season, and samples of the associated Speyeria species were collected.
Measurements included leaf length, total diameter of plants, and butterfly
forewing length. The butterflies were also examined with regm*d to
variation in the coloration of the ventral hindwings. In addition to samples
of butterflies collected during the present study, older samples collected
between 1950 and 1970 were examined in the collections of Ernst J.
Domfeld of Corvallis, Oregon, Oregon State University, and the National
Museum of Natural History in Washington, D. C.

Origin of the Speyeria Butterflies

In order to understand how the violets and butterflies have colonized the
ash-pumice fields of the Cascade Range, it is necessary to examine the
origin and ecology of these organisms in adjacent non-volcanic regions of
Oregon and northern California that have not been exposed to significant
volanic activity during the past 50,000 years. In a previous study,
Hammond (1974) examined the ecology of Speyeria butterflies across
North America, and a brief summary of this ecolo©^ for the Pacific
Northwest Speyeria is useful for the present study (see Table 2).

Eight different species of Speyeria are widely sympatric in the mountains
through much of the Pacific Northwest from western Montana and
Wyoming to the Cascade Range and southward through the Sierra Nevada
of California. Although adult butterflies frequently fly together, there is

182

J. Res. Lepid.

Table 1. Populations of Viola and Speyeria that have colonized the
volcanic ash-pumice fields of the southern Cascade Range.

Volcanoes

Habitat

Viola

Speyeria

Mt. Shasta

Red Fir forest

purpurea

egleis

Mt. McLoughlin

high elevation Lodgepole
Pine forest

purpurea,

nuttallii

atlan tis

lower elevation mixed
conifer forest

purpurea,

mittallii

egleis

Mt. Mazama, Mt.
Newberry, Three
Sisters

Lodgepole Pine forest

purpurea

egleis, zerene

moist, open areas

purpurea,

nuttallii

callippe,

coronis

wet, boggy meadows

palustris,

adunca

mormonia

Mt. Washington

mixed conifer forest

nuttallii,

orbiculata

atlantis,

hydaspe

Table 2. Topical habitats of Viola and Speyeria species in non-volcanic
regions of the Pacific Northwest.

Habitat

Viola

Speyeria

exposed wet meadows

palustris, adunca

mormonia

sheltered wet meadows with tYnckets glabella, palustris,
of willows, alders, etc. adunca

cybele

dry meadows and grassy prairies

nuttallii

atlantis

at high elevations

dense, moist spmce-fir and pine
forests

glabella, orbiculata,
adunca, nuttallii

hydaspe

open, diy pine forests

adunca, nuttallii,
purpurea

zerene

exposed, dry rocky ridges and talus

purpurea

egleis

slopes at high elevations

sagebrush-juniper communities

nuttallii, purpurea.

callippe, coronis

douglasii, beckwithii

20(3): 179-191, 1981(83)

183

usually fairly sharp segregation among species in the types of habitat
utilized by the larvae. Each species appears to be adapted for utilizing a
particular type of habitat, although Speyeria larvae are not adapted to any
particular species of violet and will feed upon any violet that happens to be
growing in the appropriate habitat (Hammond, 1974). At the same time,
most Speyeria species exhibit considerable ecological plasticity, and
frequently invade the habitats of related species when the latter are absent
for one reason or another (see contrasts between West Coast and Rocky
Mountain S. mormonia, S. atiantis, and S. hydaspe cited below). These
observations of such ‘‘ecological release” provide indirect evidence that
the ecological segregation of Speyeria species is largely the result of
interspecific competition for the larval foodplant in the various habitats.

Two species, S. mormonia Boisduval and S. cybele leto Behr, occupy wet,
boggy meadow habitats. Speyeria mormonia favors completely open,
exposed meadows that are free of shrubs and small trees, while S. cybele
kto favors more sheltered meadows with thickets of willows {Salhc sp.) and
small trees. Another species, S. atlantis dodgei Gunder, also occupies open
meadows at the higher elevations in the mountains, but only dry, well-
drained meadows in contrast to the boggy meadows occupied by S.
mormonia. It should be noted that these relationships change in the Rocky
Mountains, where S. mormonia occupies both the dry and wet meadow
habitats, while Rocky Mountain subspecies of S. atlantis occupy spruce-fir
and aspen forest habitats.

However, the primary forest species in the Pacific Northwest is S.
hydaspe Boisduval, which occupies spruce-fir forests at high elevations
and dense, moist Ponderosa Pine forests at middle elevations. Speyeria
zerene Boisduval is adapted to more xeric conditions and replaces S.
hydaspe in very dry, open pine forests. The species also extends out into
the highly xeric sagebrush-juniper habitat to some extent as well. Speyeria
callippe Boisduval is adapted to the most extreme of xeric habitats, and
occupies sagebrush-juniper communities throughout much of western
North America at the lower elevations. West of the Cascade and Sierra
Nevada Ranges, S. callippe occupies dry, open grassland and oak-pine
shrubland at low elevations. At high elevations in the mountains, S.
callippe is often replaced in the xeric habitat by S. egleis Behr, which
usually occupies barren, exposed rocky ridgetops and talus slopes within
the spmce-fir zone. The last species, S. coronis Behr, is another
semidesert species like S. callippe, and usually occupies sagebrush-
juniper habitats or open, dry pine forests. However, S. coronis is a much
larger species than the others, and appears to require a much larger
biomass of the larval foodplant (based partly upon observations of reared
larvae). Thus, S. coronis is commonly associated with moist areas near
creeks, lakes, and seepage areas in these arid regions where the violets
produce a more vigorous growth.

184

J. Res. Lepid.

Origin of the Violets

Seven different species of violets provide important larval foodplants for
Speyeria butterflies in the Cascade Range (Table 2). Viola palustns L.
occupies wet, boggy meadows near creeks, lakes, and seepage areas
throughout much of western North America, but only meadows with a
permanent source of water. In contrast, V. adunca J. E. Smith var.
bellidifolia Greene occupies wet, boggy meadows that dry out during July
and August. Both of these violets are very common in wet meadows at high
elevations throughout the Oregon Cascades, and are the primaiy larval
foodplants of S. mormonia and S. cybele leto. A third violet, V. nuttallii
Pursh. var. linguaefolia Nutt., occupies dry, well-drained meadows and
open grassy prairies at high elevations throughout much of western North
America, and is the primary larval foodplant of S. atlantis dodgei in the
Cascade Range.

Two species of mesic forest violets, V. glabella Nutt, and V. orbiculata
Geyer, occupy the moist spruce-fir forests at high elevations and are the
primary foodplants of S. hydaspe in the Cascade Range. At lower
elevations, the drier Ponderosa Pine forests are occupied by V. nuttallii, V.
purpurea KelL, and the typical form of V. adunca J, E. Smith. These
violets are utilized by the larvae of both S. hydaspe and S. zerene. Both V.
nuttallii and V. purpurea also extend into the sagebrush-juniper zone
where they are utilized by S. callippe and S. coronis. In addition, the
xerophytic V purpurea var. venosa S. Wats, occupies exposed, rocky
ridgetop and mountainsides at high elevations throughout much of
western North America, and is the primary larval foodplant of S. egleis.

However, the most extreme xerophytes among the North American
violets belong to the V. douglasii-beckwithii species complex. Viola
douglasii Steudel occupies dry, open grasslands and oak-pine brushland at
low elevations in California, extending from northern Baja California north
to Medford and Klamath Falls, Oregon. There is also an isolated relict
population of V douglasii along the Metolius River in Jefferson County,
Oregon. East of the Sierra Nevada and Cascade Ranges, V douglasii is
replaced by the closely related V beckwithii Torr. & Gray in the sagebrush-
juniper community. Both violets are primary larval foodplants for S.
callippe and S. coronis.

Colonization of Mt. Shasta Ash Fields

According to Harris (1980), Mt. Shasta represents one of the largest
stratovolcanoes in the world, reaching 4317 meters in elevation. The
volcano actually consists of a series of eruptive cones that have been built
upon each other. The most recent major cone, Shastina, has only
developed during the past 10,000 years and now reaches 3759 meters in
elevation. Violently explosive eruptions of massive proportions apparently
took place during this period, and pyroclastic flows covered much of the

20(3): 179-191, 1981(83)

185

territory near the base of the volcano. Today the steep upper slopes of Mt.
Shasta are covered with glaciers and snow during much of the year. The
lower slopes of the mountain are covered with a thick layer of finely divided
volcanic ash and pumice that supports a well developed forest of Shasta
Red Fir (^bies magnifica A. Murr. var. shastensis Lemmon). A particularly
deep ash-pumice field extends to the northeast of Mt. Shasta and only
supports a sparse forest of Lodgepole Pine.

The primary violet and Speyeria butterfly found on the Mt. Shasta ash
fields are V. purpurea and S. egleis respectively. Both species are very
common in the Red Fir forests at the higher elevations on the volcano. The
violets grow in the loose ash soil of the forest floor, and even form a thick
carpet of plants in some areas of the forest. Adult butterflies fly mostly in
sunny glades and clearings within the forest near the violets, and only
occasionally stray out into the more open, non-forested areas of the
mountain.

This last point is important when considering the origins of S. egleis in
the non -volcanic habitats of the Sierra Nevada, Warner, and Salmon-
Siskiyou Mountains. In these mountains, V. purpurea is very common in
both dry, open Ponderosa Pine forests and on barren, rocky ridgetops and
exposed mountainsides. Speyeria zerene usually occupies the pine forest
habitat while S. egleis occupies the open, rocky habitat throughout these
mountains. Likewise, S. zerene is the dominant species in the open, dry
pine forests surrounding Mt. Shasta at the lower elevations. However, the
occupation by S. egleis of the Red Fir forests at higher elevations on Mt.
Shasta clearly represents a marked ecological shift from the open, rocky
habitat of S. egleis in the Sierra Nevada, Warner, and Salmon- Siskiyou
Mountains. The reasons for this sharp ecological shift may have the
following possible explanation.

Speyeria zerene exhibits a definite preference for relatively warm forests
at lower elevations, while S. hydaspe usually occupies cool, moist spruce-
fir forests at higher elevations. The failure of S. hydaspe to occupy the Red
Fir forests on Mt. Shasta may be due to the extremely xeric nature of the
loose ash-pumice soil found on most of Mt. Shasta. Speyeria hydaspe does
occur in limited numbers in small stands of particularly dense, moist forest
on the lower slopes of Mt. Shasta, so the species does have the opportunity
to invade the upper Red Fir forests. Therefore, it would appear that the
failure of S. hydaspe to successfully occupy this forest habitat has allowed
S. egleis to make the ecological shift into the forest habitat as an
opportunistic colonizer. The major part of this colonization probably took
place during the past few thousand years following the completion of Mt.
Shasta’s massive explosive eruptions from the recently built Shastina
cone. These S. egleis populations also extend eastward from Mt. Shasta
into the Medicine Lake volcano system.

Colonization of Ash Fields Near Mt. McLoughlin

186

J. Res. Lepid.

Extensive ash-pumice fields are also found along the summit of the
Cascade Range south of Mt. McLoughhn in southern Jackson County,
Oregon, and south of Lake-of-the-Woods in southern Klamath County.
These ash fields support either pure forests of Lodgepole Pine or mixed
forests of Lodgepole Pine, Ponderosa Pine, and White Fir (Abies concolor).
Viola purpurea is still common in this area, but is not so abundant as on Mt.
Shasta. Instead, the dominant violet on these ash-pumice fields is V.
nuttalli var. linguaefolia, a species that occupies dry open meadows and
^assy prairies at high elevations throughout the Cascade Range. In the
ash fields near Mt. McLoughlin, both V. purpurea and V. nuttallii often
form a dense carpet of plants in the ash soil of open clearings within the
pine forest, but the individual plants are usually very small rosettes
measuring only 2-6 cm. in diameter.

The primary Speyeria that utilizes V. nuttallii in the high mountain
meadows and prairies of the Cascade Range is S. atlantis dodgei, and this
species has followed V. nuttallii into the ash-pumice fields south of Mt.
McLoughlin. Speyeria atlantis is the dominant species on the ash fields
along the summit of the Cascades in Jackson County, although a few S.
egleis are also present in this area. However, there is a rapid shift in
dominance over to S. egleis on the ash fields south of Lake-of~the- Woods in
Klamath County. The reasons for these replacements between S. egleis
and S. atlantis are not really known, but are thought to be the result of
subtle differences in temperature and moisture in the two areas. For
example, the ash fields along the summit of the Cascades are usually
covered with a deep snowpack during the winter, while the lower
elevations along the eastern slope of the Cascades in Klamath County
usually receive much less snowfall.

A very similar situation exists in the central Oregon Cascades. Both V.
purpurea and S. egleis are absent from the summit of the Cascades in the
Mt. Washington ash fields of Linn County, but both species are present on
the eastern slope near the Metolius River and east of the Three Sisters
volcanoes in Jefferson and Deschutes Counties. The ash fields north of
Mt. Washington near Big Lake support large populations of V. nuttallii
just as in the Mt. McLoughlin area, and some S. atlantis dodgei are also
associated with these violet populations. However, both V. orbiculata and
S. hydaspe have also penetrated into the Lodgepole Pine forests around
Mt. Washington from the adjacent spruce-fir forests on Santiam Pass.
This is the only instance where a mesic species of violet and Speyeria
butterfly have been observed to occupy a xeric ash-pumice habitat, and is
probably due to the moisture received from the heavy snowpack that
accumulates during the winter along the Cascade summit.

Colonization of the Mt. Mazama-Mt. Newberry Ash Fields

The deepest and most extensive ash-pumice fields in the Oregon

20(3): 179-191, 1981(83)

187

Cascade Range were deposited by massive, violently explosive eruptions
from Mt. Mazama and Mt. Newberry in northern Klamath County and
southern Deschutes County. These ash fields also extend northward in
Deschutes County along the eastern slope of the Three Sisters volcanic
system; a complex of numerous volcanoes that have also produced
significant eruptions of ash and pumice. The most important eruption was
the cataclysmic explosion that destroyed Mt. Mazama about 6600 years
ago, creating the modem Crater Lake caldera. Likewise, Mt. Newberry has
also been replaced with a deep caldera crater that is occupied by Paulina
Lake and East Lake today. The deepest ash fields extend from the south
end of Crater Lake National Park eastward to Yamsey Mountain in
Klamath County and northward to the Paulina-East Lakes in Deschutes
County, an area covering more than 3500 square kilometers. According to
Harris (1980), the depth of these ash-pumice fields ranges from 7-15
meters, and reaches a depth of 75 meters in some of the creek valleys near
Crater Lake. Before this cataclysmic emption, Mt. Mazama was one of the
largest Cascadian stratovolcanoes, reaching an estimated height of 3660
meters. Afterwards, an estimated 65 cubic kilometers of volcanic material
were expelled from the mountain to produce these ash-pumice fields. The
Newberry caldera is thought to have been created through a series of
violently explosive emptions according to Harris, rather than in a single
cataclysmic emption as with Mt. Mazama. The most recent emptions are
thought to have taken place less than 1000 years ago. As Harris has
pointed out, Mt. Newberry is a huge shield volcano extending 32
kilometers in diameter with some 200 parasitic cinder cones on the slopes
of the volcano. In addition to explosive emptions of ash and pumice, the
volcano has also produced numerous thicker flows of basalt and obsidian
during the course of its long history.

The ash fields from Crater Lake to the Paulina-East Lakes are extremely
dry, and only support a sparse, scmbby forest of Lodgepole Pine in most
areas, although Ponderosa Pine is found in some areas where the ash soil is
shallower. As a result, the forest floor is mostly barren and devoid of
vegetation except for scattered shmbs of Bitterbmsh {Purshia tridentata)
and small tufts of grasses. However, there are many places in these ash
fields where ground water does come to the surface, producing small
creeks, seepage areas, and wet, boggy meadows. Stands of willows {Salix
sp.) and Quaking Aspen (Populus tremuloides Michx.) occupy these wet
areas together with a large diversity of herbaceous plants including wild
strawbeny {Fragaria sp.), columbine {Aquilegiaformosa Fisch.), delphinium
{Delphinium menziesii D.C.), penstemons {Penstemon sp.), Indian paint-
bmsh {Castilleja sp.), and various composites.

Of the three species of xerophytic violets found along the eastern slope of
die Oregon Cascades, only V. nuttallii var. linguaefolia and V. purpurea
have been able to colonize the Mazama ash fields. Viola douglasii is found

188

J. Res. Lepid.

on the juniper-sagebrush hillsides around Klamath Falls in southern
Klamath County and again along the Metolius River in Jefferson County.
The region between these two widely disjunct populations was buried
under the Mt. Mazama ash fall, and the violet has never been able to
successfully recolonize the Mazama ash fields. The thick rootstalk of V.
douglasii appears to be adapted to a firm rocky or hard clay substrate, and
it does not grow well in a loose, finely divided ash-pumice soil.

Although V. nuttallii grows well in the ash-pumice soU, it apparently
requires considerable moisture in the spring, and is mostly restricted to
the higher elevations where there is a heavy snowpack in the winter, for
example around the Paulina-East Lakes. Only V. purpurea is common and
widely distributed in the pine forests of the Mazama ash field, and it is
largely restricted to the moist zones of pine forest near the edges of creeks
and boggy seepage areas. Even in this fairly moist habitat, the individual
plants of V. purpurea are very small, measuring only 2-5 cm. in diameter.

Two species of mesophytic violets have been able to colonize the wet
areas in the Mazama ash fields from the high mountains to the west. Viola
palustris is strongly restricted to the edges of creeks and lakes that remain
permanently wet through the year, while V. adunca var. bellidifolia is
extremely abundant in wet, boggy meadows that dry out for part of the
summer. The violets often form a dense carpet of plants in many parts of
these meadows, but the individual plants are very small as in V. purpurea,
measuring only 1-5 cm. in diameter.

Five different species of Speyeria have been able to colonize the Mazama
ash fields, including S. mormonia, S. egleis, S. zerene, S. coronis, and S.
callippe. Moeck (1957) and Tilden (1963) previously studied the Speyeria
populations in this region. Speyeria mormonia occupies the wet, boggy
meadows, utilizing V. palustris and V. adunca as larval foodplants. The
butterfly has almost certainly invaded and colonized the Mazama ash
fields together with these violets from the adjacent high Cascades to the
west. However, S. callippe and S. coronis occupy dry, open habitats, and
have probably invaded the Mazama ash fields from the adjacent sagebrush-
juniper lowlands to the east, which constitute the usual habitat of these
xeric species. Because the Mazama ash fields are mostly covered with
Lodgepole Pine forests, S. callippe and S. coronis are usually rather rare in
this region due to the scarcity of open habitats, but particularly large
populations breed in the open, moist areas around the Paulina-East
Lakes. Speyeria callippe is the dominant species around these lakes, while
S. coronis is less common and S. mormonia is restricted to the wet, bog^
habitats. Thus, mesic S. mormonia and xeric S. callippe appear to have
invaded and colonized the Mazama ash fields from opposite directions.

Speyeria zerene and S. egleis are the dominant species in the Lodgepole
Pine forests of this region, but they are mostly restricted to the moist zones
of pine forest near the creeks and boggy seepage areas like their V.

20(3): 179-191, 1981(83)

189

purpurea foodplant. While S. zerene is the usual Speyeria to occupy dry,
open pine forests throughout western North America, it is often rather
scarce on the Mazama ash fields, and is often replaced in this habitat by
large populations of S. egleis. This latter species has invaded and
colonized the Mazama ash fields from the Mt. McLougWin and Lake-of-
the-Woods ash fields in southern Klamath County. Evidence of this
invasion is suggested by the physical characteristics of butterflies from
clinal populations between Lake-of-the-Woods and Crater Lake. In
addition to occupying pine forests at low elevations east of Crater Lake,
egleis also occupies barren pumice fields around the rim of Crater Lake
itself.

The unique feature of these Mazama S. egleis is their very small size,
which appears to represent an adaptation to the small, dwarfed size of
their larval foodplant. For comparison, a sample of 64 S. egleis mal^from
near Lake-of-the-Woods had a forewing length range of 23-28 mm. (X = 26
mm.), while a sample of 71 males collected on the Mazama ash fields east
of Crater Lake had a range of 21-25 mm. (X — 23 mm.). As such, the
Mazama S. egleis rank among the smallest of all Speyeria butterflies in
existence. These samples are illustrated in Figure 1., and the difference
between the sample means is highly significant p<0.0001). Moreover,

male forewing length mm.

Fig. 1. Comparison of male forewing length in Speyeria egleis samples from the
Mazama ash fields of northern Klamath Co., Oregon, and near Lake-of-the-
Woods in southern Klamath Co.

190

J. Res. Lepid.

laboratory rearing experiments conducted by D. V. McCorkle of Monmouth,
Oregon, and O. D. Spencer of Lincoln, Nebraska, have shown that this
dwarfed size is under strong genetic control, and is not merely an
environmentally induced phenotype. McCorkle reared samples of S. egleis
from several different areas under approximately identical conditions, and
the largest specimens from the Mazama region of Klamath County
reached 25 mm. while the largest from the Siskiyou Mountains of
Josephine County reached 27 mm. Thus, the size of re^ed specimens
from different areas closely corresponds to the size range of wild
specimens, suggesting a strong genetic component to the determination of
forewing length.

This Mazama form of S. egleis also differs from the Mt. Shasta-Mt.
McLoughlin populations in coloration, exhibiting relatively little dark
basal suffusion on the dorsal wing surfaces and often a reddish brown
ventral hindwing. In contrast, the Mt. Shasta-Mt. McLoughlin S. egleis
usually exhibit more extensive dark basal suffusion on the dorsal wing
surfaces and a black-brown or umber brown ventral hindwing.

The Mazama form of S. egleis has a wide distribution in the central
Oregon Cascades that corresponds very closely with the distribution of the
ash-pumice fields in this region, extending from Crater Lake eastward to
Yamsey Mountain and northward to the Paulina-East Lakes and the
Three Sisters volcanoes in Deschutes County. Since this habitat did not
even exist until after the destruction of Mt. Mazama about 6600 years ago,
it would appear that this distinctive Mt. Mazama form of S. egleis has only
evolved from the Mt. Shasta form during the past 6000 years or so.
Considering the fact that the Mazama ash fields represent a particularly
harsh, xerophytic environment, it is interesting to observe that this
organism has been able to colonize and evolve in this new environment in
such a short time period.

While S. egleis has successfully colonized the Mazama ash fields, S.
cybele leto has been completely excluded from this region, and displays the
same relict, disjunct distribution as Viola douglasii. Colonies of S. cybele
are found around Klamath Lake in southern Klamath County and again
along the Metolius River in Jefferson County, but the species has not been
able to recolonize the intervening Mazama ash fields. Speyeria cybele leto
is a large butterfly and apparently requires a large biomass of the larval
foodplant, but the violets that grow in volcanic ash habitats are usually too
small to support such a large butterfly (based partly upon observations of
larvae reared by D. V. McCorkle).

Conclusions

The colonization of volcanic ash-pumice fields in the southern Cascade
Range by violets and Speyeria butterflies is summarized in Table 1. For
comparison, the normal habitats of the plants and animals in non-volcanic

20(3): 179-191, 1981(83)

191

areas are summarized in Table 2. These organisms demonstrate how
plants and associated insects may colonize and adapt to new environments
recently created by the eruptions of Cascade volcanoes. It is interesting
that some violet species such as V. purpurea and V. nuttallii have been
vigorous colonizers of the ash-pumice fields, while other violets such as V.
dbuglasii have been conspicuously unsuccessful. Speyeria egleis has
followed V. purpurea into the ash fields of Mt. Shasta, Mt. McLoughlin,
and Mt. Mazama from the exposed, rocky habitats of the Sierra Nevada,
Siskiyou, and Warner Mountains. In contrast, S. atlantis dodgei has
followed V. nuttallii into the Mt. McLoughlin ash fields from the dry, open
meadows and prairies of the high Cascades. Likewise, S. mormonia has
followed V. palustris and V. adunca into the Mt. Mazama ash fields from
the wet, boggy meadows of the high Cascades. However, both S. callippe
and S. coronis have invaded the Mt. Mazama ash fields from the
sagebrush-juniper lowlands east of the Cascade Range. Speyeria zerene is
also a typical resident of the Mt. Shasta, Mt. McLoughlin, and Mt.
Mazama regions where it occupies open dry pine forests; the same habitat
the species occupies throughout much of western North America.
However, S. cybele leto has been effectively excluded from ash-pumice
habitats due to its requirement for a large biomass of the larval foodplant.

Acknowledgments. I am particularly indebted to O. D. Spencer, Ernst J.
Domfeld, and David V. McCorkle for their kind contributions of much assistance
and data to this paper. I also want to thank L. Paul Grey for many long and
stimulating discussions regarding these various Speyeria populations, and to
several reviewers for suggesting numerous improvements in the manuscript.

literature Cited

HAMMOND, P. c., 1974. An ecological survey of the nymphalid butterfly genus
Speyeria. M. S. Thesis, University of Nebraska, Lincoln, Nebraska, U.S.A.
HARRIS, s. L., 1980. Fire and ice, the Cascade volcanoes. The Mountaineers-
Pacific Search Press, Seattle, Washington, U.S.A.

MOECK, A, H., 1957. Geographic variability in Speyeria. Milwaukee Entomological
Society, Milwaukee, Wisconsin, U.S.A.

TTLDEN, j. w., 1963. The Argynnis populations of the Sand Creek area, Klamath Co.,
Oregon. J. Res. Lepid. 1: 109-113.

Journal of Research on the Lepidoptera

20(3): 192, 1981(83)

The Life of the Meadow Brown.

W. H. Dowdeswell, 1981. ca. 14 x 21 cm; 8 + 165 pp., 19 pis., 27 tabs., 29 figs.
Heinemann Educational Books, London. Price: £ 5.95, paperback.

In this fascinating little book the author deals with the ecology oiManiola jurtina,
one of the “most successful” of all European butterfly species. Professor Dowdes-
well used this widespread and usually abundant satyrid both as the object of his
field investigations and as a convenient research tool to answer some fundamental
biological questions, mostly related to ecological genetics. His research programme
began shortly after the end of World War 11 and extended over a period of some 30
years. It was conducted in cooperation with the acknowledged group of Oxford
scientists centered around Professor E. B. Ford. The book is concisely written,
describing both results and the thought processes directing the research that led
from the formulation of the enquiry to its usually successful conclusion. Choice of
methodology is discussed in some detail, making the book a very useful reading for
less experienced students and amateur lepidopterists. It also clearly shows what
can be achieved with relatively simple means. The text of the book is divided into
seven chapters and provided with a list of the bibliographical references and
comprehensive index. The following aspects of the life oiManiola jurtina are among
those studied by Dowdeswell (the order follows approximately that used in the
book): description of the adults and early stages, life cycle, related species,
variation and its significance, population size and its fluctuation, island populations
and aspects of isolation, spotting and its significance, ‘boundary phenomenon’,
spot-placing as an index of variation, variation-types and their relationship,
geography of variation, sex ratio, evolution of island stabilization, selection,
behaviour, selective agents, ecological bacteriology, enzymes and adaptation,
electrophoresis, inheritance of spotting, spot- variation. Field work, supplemented
by laboratory experiments, took place in Great Britain (including some adjacent
islands) and on the Continent, mainly in Holland and Italy.

A few unfortunate errors entered the text. The genus Maniola was described by
Schrank and not by Linnaeus, as stated on page 6. References are not arranged in
alphabetic order and they are cited in the text by numbers instead of the usual and
significantly more convenient author’s name; some papers are not cited in full
and/or correctly (e.g. Atti Accad. naz. Lincei Rc. instead of incorrect Accademia
Nazionale Dei Lincei on p. 159 and elsewhere). The taxonomic account oiManiola
jurtina is perhaps far too brief, and, to my surprise, the only comprehensive paper
on the systematics of the species is not to be found listed in the references: G.
Tliomson in Tijdschr. Ent. 116(1973): 185-227, 1974. If the reason for the
exclusion of Thomson’s paper was its abundant use of inconvenient multinominal
combinations, denoting not only the superfluous subspecies, but also many
individual and local forms, it should have been explained. In any case, it would have
been very useful to have provided a comprehensive bibliography instead of the list
of only 52 references. Thus the book and especially its readers would have gained
much more. The species of the genus Hyponephele Muschamp which inhabit
Europe should have been included among the British relatives of Maniola jurtina,
as they are closer to it than many of the related species listed.

The book is well written, inexpensive and, above aU, it shows new ways of taking
interest in butterflies. Every lepidopterist should read it.

Otakar Kudma, Rhenusallee 30, D-5300 Bonn 3, West Germany

INSTRUCTIONS TO AUTHORS

Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, on SVt x 11 inch paper with wide margins. Number all pages
consecutively and put author’s name at top right corner of each page. If your typewriter
does not have italic type, underline all words where italics are intended. Footnotes,
although discouraged, must be typed on a separate sheet. Do not hyphenate words at the
right margin. All measurements must be metric, with the exception of altitudes and
distances which should include metric equivalents in parenthesis. Time must be cited on
a 24-hour basis, standard time. Abbreviations must follow common usage. Dates should
be cited as example: 4. IV. 1979 (day-arabic numberal; month-Roman numeral; year-
arabic numeral). Numerals must be used before measurements (5mm) or otherwise up to
number ten e.g. (nine butterflies, 12 moths).

Title Page: All papers must have the title , author’s name, author’s address, and any
titular reference and institutional approval reference, all on a separate title page. A
family citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.

Abstracts and Short Papers: All papers exceeding two typed pages must be ac-
companied by an abstract of no more than 300 words. An additional summary is not
required.

Name Citations and Systematic Works: The first mention of any organism should
include the full scientific name with author (not abbreviated) and year of description. New
descriptions should conform to the format: male: female, type data, diagnosis, distribu-
tion, discussion. There must be conformity to the current International Code of
Zoological Nomenclature. We strongly urge deposition of types in major museums, all
type depositions must be cited.

References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbrevations must conform to the World List of
Scientific Periodicals. Do not underline periodicals. If four or less references are cited,
please cite in body of text not in Literature Cited.

Tables: Tables should be minimized. Where used, they should be formulated to a size
which will reduce to 4 x 6V2 inches. Each table should be prepared as a line drawing or
typed with heading and explanation on top and footnotes below. Number with Arabic
numerals. Both horizontal and vertical rules may be indicated. Complex tables may be

reproduced from typescript.

Illustrations: Color must be submitted as a transparency (i.e., slide) ONLY, the quality
of which is critical. On request, the editor will supply separate detailed instructions for
making the most suitable photographic ilustrations. Black and white photographs should
be submitted on glossy paper, and, as with line drawings, must be mounted on stiff white
cardboard. Authors must plan on illustrations for reduction to the 4 x 6y2” page.
Allowance should be made for legends beneath, unless many consecutive pages are used.
Drawings should be in India ink at least twice the final size. Include a metric scale or
calculate and state the actual magnification of each illustration as printed. Each figure
should be cited and explained as such. The term “plate” should not be used. Each
illustration should be identified as to author and title on the back, and should indicate
whether the illustration be returned.

Legends should be separately typed on pages entitled “Explanation of Figures”.
Number legends consecutively with separate paragraph'for each page of illustrations. Do
not attach to illustrations. Retain original illustrations until paper finally accepted.

Review: All papers wiU be read by the editor(8) & submitted for formal review to two
referees. Authors are welcome to suggest reviewers, and if received, submit name &
comments of reviewers.

©I>« THE LEFIJD0PTERA

Volume 20 Numbers Autumn, 1981(83)

m THIS ISSUE

Date of Publication: April 15, 1983

Role of an Ornamental Plant Species in Extending the

Breeding Range of a Tropical Skipper to Subtropical
Southern Texas (Hesperiidae)

Raymond W. Neck 129

Japanese Literature

Ichiro Nakamura 134

Hipparchia azorina (Strecker, 1899)(Satyridae) Biology,

Ecology and Distribution on the Azores Islands

Steffen Oehmig 136

Courtship Behavior of the Dainty Sulfur Butterfly, Nathalis
iole with a Description of a New, Faculative Male
Display (Pieridae)

Ronald L. Rutowski 161

On the Supernumerary Chromosomes of Tarache tropica
Guen. (Lepidoptera: Noctuidae)

P. K. Mohanty & B. Nayak 170

An Apparent Interspecific Fi Hybrid Speyeria (Nymphalidae)

James A. Scott 174

A New Record of Vanessa virginiensis “ab. ahwashtee'' from
Northern California (Lepidoptera: Nymphalidae)

Arthur M. Shapiro 176

The Colonization of Violets and Speyeria Butterflies on the
Ash-Pumice Fields Deposited by Cascadian Volcanoes

Paul C. Hammond 179

Book Review 192

Cover Illustration: Artist’s rendition of courtship behavior in the dainty sulfur,
Nathalis iole. Figure was drawn from a 35 mm photograph by Patricia Rutowski. See
page 33, R. Rutowski.

Volume 20

Number 4

THE JOURNAL OF RESEARCH
ON THE LEFIOOFTERA

Published By: The Lepidoptera Research Foundation, Inc.

c/o Santa Barbara Museum of Natural History
2569 Puesta Del Sol Road
Santa Barbara, California 93105

Founder:

William Hovanitz

Editorial Staff: Rudolf H. T. Mattoni, Editor

Lorraine L. Rothman, Managing Editor
Scott E. Miller, Assistant Editor

Associate Editors: Thomas Emmel, U.S.A.

Arthur Shapiro, U.S.A.

Brian O. C. Gardiner, England
Bjorn Petersen, Sweden
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Emilio Balletto, Italy
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Journal of Research on the Lepidoptera

20(4): 193-198, 1981(83)

Butterfly Taxonomy: A Reply

Lee D. Miller
and

F. Martin Brown

Allyn Museum of Entomology of the Florida State Museum, 3701 Bay Shore Road,
Sarasota, Florida 33580

Our Catalogue/Checklist (Miller and Brown, 1981) is an updating of that
prepared by Dyar (1903), wliich in turn was taken generally from that by
Skinner (1898). The promised catalogue that was to follow dos Passes
(1964) was not forthcoming, and our publication attempted to fill this gap.
It is an up-to-date (through 1979) revision of the nomenclature which 1)
follows an order acceptable to taxonomists of worldwide background (this
often differs from the arrangements in popular books and more parochial
works); 2) follows the best and most recent revisions of genera (including
the recognition as genera of taxa previously recognized as subgenera); 3)
considers higher taxa on their worldwide complement of species, not just
those from the Nearctic; 4) takes into account biological and ecological
advances in knowledge, such as different foodplant preferences that
separate distinct, but previously overlooked genera; and 5) attempts to
follow the International Code of Zoological Nomenclature. The Cata-
logue/Checklist does not take into consideration political constraints or the
aspects of continued usage of incorrect, frequently employed names.
Should the conservation of these names be desired, the place to apply for
their reinstatement is the International Commission on Zoological Nomen-
clature.

It should be clearly stated here that no new names, at the generic or other
level, have been employed by Miller and Brown (1981). In every case, the
generic names are ones proposed validly by earlier authors. There are 669
notes which provide detailed information (often in telegraphic form
because of space constraints) on authorities followed and, at least by
implication, reasons for changes from pre-existing nomenclature.

Such a treatment as ours is not made without objections, and those
complaints made by Ehrlich and Murphy (1982) require some objective
replies. Their points are by no means new—similar reactions have
followed every reclassification that has appeared. The catalogue format
precludes detailed explanation of every change (though such changes are
documented elsewhere in the literature); Ehrlich and Murphy’s com-

194

J. Res. Lepid.

plaints likewise are unsupported by specific explanations. The usual
objections to nomenclatorial changes are: old, reliable, comprehensive
genera are being split; the literature will be adversely affected by
“wholesale name changes’’; the new names do not reflect relationships,
obscuring the affinities of species that “everyone knows belong together”;
and that in general taxonomists are meddlesome creatures who keep
themselves busy changing names for the sake of changing them for no
biological purpose.

Attacks like those mentioned may be seen in the pages of Strecker (1876:
118-120) berating Scudder (1875), of Holland (1931: 325) and Forbes
(1960) in their not accepting the work of Lindsey, Bell and Williams
(1931), by Hovanitz (1962: 95-96) in his criticism of dos Passos and Grey
(1947) and again by Hovanitz (1965: 18) regarding dos Passos (1964). In
the latter critique, Hovanitz states, “Names proposed by splitters do not
have to be used merely because they have been proposed.” He did not,
however, go so far as to suggest that these names be expunged from the
literature.

We are being criticised essentially for being “out of touch” with modern
systematic work, but are we? We had expected a certain amount of
controversy to arise when the classical treatment of the Papilionidae by
Munroe (1961) was expanded. This expansion was not done solely on the
basis of the genitalia, as suggested by Ehrlich and Murphy (1982); rather
the morphology of all life stages was considered along with biological
parameters such as foodplant preferences. Munroe never claimed to have
all of the answers for the Papilionini, and the passage from his work quoted
by Ehrlich and Murphy indicates that he had questions about his ability to
find differences within “Pqpi/io”, differences which have become apparent
with more life history data. Our treatment, hke that of Eliot (1978),
generally follows the outline of Munroe ’s classification, but at a different
level.

Each of the papilionid genera that we accepted is either a lauraceous,
rutaceous or umbelliferous feeder (the main exception being Papilio, s.
str., which contains some rutaceous feeders, apparently as a response to
competition by congenors). Ferris and Emmel (1982) studied the rutaceous
feeding P. polyxenes color W. G. Wright and showed that while it feeds in
the wild on Rutaceae, its larvae actually do better on Umbelliferae, but it is
out-competed for the umbellifers by the very successful P. zelicaon Lucas.
This switching of foodplant groups within a phyletic line is interesting and
may be of fundamental biological and systematic importance. While there
are some biochemical similarities between a number of Umbelliferae and
some Rutaceae, they are not biochemically identical! Berenbaum (1981)
shows how narrowly Papi/io, s. str., is tied to plants (chiefly Umbelliferae)
with concentrations of furanocoumarins, and he further shows that
rutaceous feeders within that genus also will utilize Umbelliferae that

20(4): 193-198, 1981(83)

195

contain these substances.

Perhaps the most characteristic furanocoumarin-bearing rutaceous
plant is Citrus, the host of Heraclides and Priamides. This strongly
indicates that these genera are sister groups of Papilio (perhaps Priamides,
despite its structural dissimilarities, is a good subgenus of Heraclides), and
that this cluster of genera is evolutionarily very different from the
lauraceous feeders. Among Nearctic swallowtails, the lauraceous feeders
are the ones Miller and Brown (1981) include in Pterourus.

These genera are not so restricted in numbers (“one species per genus”)
as a parochial examination of them would indicate. Papilio is found
throughout the northern hemisphere and contains at least twice as many
species as are represented in the Nearctic. Heraclides is a genus of at least
15 Nearctic and Neotropical species, Priamides contains at least ten
species from the Neotropics and Pterourus has eight or more Nearctic and
Mexican representatives. Further, while all are bird-lime mimics in early
larval instars, only Heraclides and Priamides retain this character in later
stadia. Pterourus larvae in later instars are green with thoracic “eye-
spots”, presumably as a sham defense, and mature Papilio larvae are
conspicuously banded with green and black totally unlike other groups.
The osmeterium of Papilio is shorter and stubbier than that of Heraclides
ajnd Pterourus. Thus, the statements by Ehrlich and Murphy (1982) about
the division of ''Papilio'' by Huebner [1819] are merely their way of setting
up “straw men” so that they could knock them down. In a much quieter
way, we did the synonymization of these names to one another, and the fact
that perhaps Huebner established genera for the wrong reasons or
otherwise faultily cannot diminish that he did establish them.

Other examples could be cited, such as the objections to the division of
"Lycaena" into constituent genera (Sibitani, 1974; Miller and Brown,
1979), but some of the comments made by Ehrlich and Murphy (1982)
suggest that they did not really read these papers for content, only to find
grounds for criticism.

Further criticism is rendered because we have “ignored perfectly good
subgenera” and raised them to generic standing. This may be true,
perhaps, but the steps were taken for reasons entirely different from the
capriciousness attributed to us. Subgeneric names, if consistently applied,
can indeed carry great taxonomic and evolutionary information. The
difficulty is that most advocates of subgenera are not consistent. Thus, we
have a situation analagous to that demonstrated by Eliot (1978: 121)
where members of the genus Euploea Fabricius are tabularly divided into
subgenera, which taxa are not mentioned again in the text. The “use” of
subgenera in this way conveys little or no information about specific
groupings, and Euploea is left as an apparently homogeneous assemblage
of related species. Division of Euploea into separate genera (note that we
are not advocating it), elevated from subgeneric standing and placed into a

196

J. Res. Lepid.

well-defined hierarchical classification, does not obfuscate relationships,
but rather, it strengthens them.

In North American butterflies, good candidates for demotion to sub-
generic standing would include Occidryas, Hypodryas, Abaeis, Pyrisitia,
Falcapica and Priamides, and we would then recommend reinstatement of
at least the subgenera Semnopsyche and Erynnides. Nevertheless, while
Ehrlich and Murphy admit the at least “weak subgeneric” status of
Occidryas, they steadfastly refuse to refer to ''Euphydryas (Occidryas)
editha (Boisduval)”, preferring a strict binominal designation. We could
certainly accept consistent use of subgenera, but not the sporadic usage of
taxonomic names at whatever level.

The arguments put forth on scientific vs. vernacular names are, we feel,
specious and clearly irrelevant in the context of the rest of their paper.
Much the same must be said about the thoughts expressed on species-
level taxonomy. We shall comment neither on the correctness nor the
political morality of the thoughts expressed by Ehrlich and Murphy (1982)
on the reasons for naming subspecies, save to state that there are other
thoughts on the matter.

It cannot be questioned that evolutionary problems are more complex
than are reflected in the pages of popular texts: if they were not more
complex, they would have been solved long ago. Perhaps Ehrlich and
Murphy are correct when they accept the concept of the evolutionary
unimportance of subspecies (Wilson and W. Brown, 1953), but acceptance
of that idea also suggests that the detailed study of even smaller demes (for
example, the Occidryas editha studies of Ehrlich and his coworkers) may
not be entirely or even mostly evolutionary in nature. Gould (1982: 104)
states unequivocally, “We cannot learn everything we need to know about
evolutionary trends by studying what happens within demes, if only
because species can act as units of selection.” The key word in this
quotation is “everything”, and the key thought is that no one study
contains all of the information needed to pass evolutionary judgments.

Another point that Ehrlich and Murphy (1982) seem to have forgotten is
that evolution is a dynamic process, so it is natural that some taxa will be
“better” (i. e., phyletically more divergent from other taxa) than others.
Were evolution a static process, phenetic measures might show similarities
and gaps, but since taxa are in vaiying stages of divergence, phenetic
analysis does not always show relationships correctly. Within taxonomic
lines, some taxa are diverging more rapidly, others less, than their closely
related counterparts. Whether one accepts classical gradualism, the
punctuated equilibrium theoiy of Eldredge and Gould (1972) and Gould
and Eldredge (1977) or something that embodies parts of both theories, it
is clear that taxon “A” might well evolve, by one means or another, faster
or slower than taxon “B”.

We object to formalization of the generic nomenclature in Howe (1975)

20(4); 193^198, 1981(83)

197

chiefly because that book is replete with errors (Ferris, 1976) and because
the nomenclature is at odds with that recognized by specialists throughout
the world. Similarly, we must also reject the attempt to formalize the
higher classification of Ehrlich (1958) and Ehrlich and Ehrlich (1967): that
particular proposal is self-serving. Our argument with the Ehrlich schema
is not necessarily with the arrangement of taxa (we more or less agree), but
rather, with the taxonomic levels in which the categories are placed. The
statement (Ehrlich, 1958) that his classification was somehow more in line
with those adopted in other insect orders was not convincingly defended;
nor did that classification take into account what is known about fossil
butterflies simply because so many of the relevant examples have been
discovered in the last two decades. Many of these fossils indicate slower, a
few faster, rates of evolution than would be indicated by the five to eleven
million years’ duration for invertebrate species given by Raup (1978) and
Stanley (1979). Gould (1982:95) suggests, however, . .some species may
survive longer than others because they inhabit a certain kind of
environment, not because their morphologies are ‘better’ in any conven-
tional sense.” Again, we must reiterate that the last word has not been
written on butterly higher classification or phylogeny.

This reply is essentially a plea for additional research, unfettered by
concerns about “sacred cows”. We were somewhat loath to write this
article, but the mere suggestion of the suppression of Miller and Brown
(1981) by Ehrlich and Murphy (1982) left us with no alternative. We reject
their call for censorship, wondering to what other papers it ultimately
might be applied: science is the censor, not individual scientists or
editors.

In the Editor’s Note preceding Miller and Brown (1981), C. V. Covell,
Jr., states, “No arrangement of taxa has yet proved to be the ‘right’ one,
and we expect the years and the gristmill of scientific discourse to bring us
closer to a true phylogenetic classification of the Lepidoptera.” Our work
was written in that spirit, and we welcome the renewed interest in
taxonomic investigation that we hope it spurs. It is only through such
research that progress can be made.

Literature Cited

BERENBAUM, M., 1981. Effects of linear furanocoumarins on an adapted specialist
insect {Papilio polyxenes). EcoL Ent., 6:345-351.

DYAR, H. G., 1903. A list of North American Lepidoptera and key to the literature of
this order of insects. Bull. U. S. Natl. Mus., (52): xix + 723 pp.

EHRLICH,?. R., 1958.The comparative morphology, phylogeny and higher classifica-
tion of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull.,
39(8):305-370.

, & A. H. EHRLICH, 1967. The phenetic relationships of the butterflies. I.

Adult taxonomy and the non-specificity hypothesis. Syst. ZooL, 16:301-317.

, & D. D. MURPHY, 1982. Butterfly nomenclature: a critique. J. Res. Lepid.,

20:1-11.

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EUOT, J. M., ed., 1978. The butterflies of the Malay Peninsula, 3rd ed. Kuala Lampur,
Malaysia, Malayan Nature Society: xiv + 578 pp., 35 pis.

FERRIS, C. D., 1976. Book review. J. Lepid. Soc., 30:138-143.

, & J. F. EMMEL, 1982. Discussion of Papilio coloro W. G. Wright {—Papilio

rudkini F. & R. Chermock) and Papilio polyxenes Fabricius. Bull. Allyn Mus.,
(76): 13 pp.

FORBES, w. T. M., 1960. Lepidoptera of New York and neighboring states. Agaristidae
through Nymphalidae including butterflies. Part 4. Ithaca, N. Y., Cornell Univ.
Agr, Expt. Sta., Mem. 371: 188 pp.

GOULD, s. J., 1982. The meaning of punctuated equilibrium and its role in validating
a hierarchical approach to macroevolution, in Milkman, R. (ed.). Perspectives
on evolution: pp. 83-104.

, & N. ELDREDGE, 1977. Punctuated equilibrium: the tempo and mode of

evolution reconsidered. Paleobiology, 3:115-151.

HOLLAND, w. J., 1931. The butterfly book (rev. ed.). Garden City, N. Y., Doubleday
Doran & Co.: xii + 424 pp., 77 pis.

HOVANITZ, w., 1962. Argynnis and Speyeria. J. Res. Lepid., 1:94-95.

, 1965 [“1964”]. Book review. J. Res. Lepid., 3:18.

HOWE, w. H. (ed.), 1975. The butterflies of North America. Garden City, N. Y.,
Doubleday & Co.: xiii + 633 pp., 97 pis.

HUEBNER, J., [1819]. Verzeichniss bekannter Schmettlinge [sic.], [part 7]. Augsburg,
Priv. publ.

LINDSEY, A. w., JR., E. L. BELL & R. c. WILLIAMS, JR., 1931. The Hesperioidea of North
America. Denison Univ. Bull. Sci. Lab., 26:1-142.

MILLER, L. D., & F. M. BROWN, 1979. Studies in the Lycaeninae (Lycaenidae). 4, The
higher classification of the American coppers. Bull. Allyn Mus., (51); 30 pp.

, 1981. A catalogue/checklist of the butterflies of America north

of Mexico, Mem. Lepid. Soc., (2): vii + 280 pp.

MUNROE, E. G., 1961. A classification of the Papilionidae (Lepidoptera). Canadian
Ent., suppl. 17:1-51.

doe PASSOS, C. F., 1964. A Synonymic list of the Neactic Rhopalocera. Mem. Lepid.
Soc., (1): V + 145 pp.

, & L. P. GREY, 1947. Systematic catalogue of Speyeria (Lepidoptera:

Nymphalidae) with designations of types and fixations of type localities.
American Mus. Novitates, (1370): 30 pp.

RAUP, D. M., 1978. Cohort analysis of generic survivorship. Paleobiology, 4:1-15.

SCUDDER, S. H., 1875. Synonymic list of the butterflies of North America, north of
Mexico. Part 1, Nymphales. Bull. Buffalo Soc. Nat. Sci., 2:233-269.

SIBATANI,A., 1974. A new genus for two new species of Lycaeninae (s. sir.) (Lepidop-
tera: Lycaenidae) from Papua-New Guinea. J. Australian Ent. Soc., 13:95-110.

SKINNER, H., 1898. A synonymic catalogue of the North American Rhopalocera.
Philadelphia, American Ent. Soc.: xvi + 99 + xiv pp.

CTANLEY, s. M., 1979. Macroevolution: pattern and process. San Francisco, W, H.
Freeman Co.: xii + 332 pp.

STRECKER, F. H. H., 1876 [1872-1878]. Lepidoptera, rhopaloceres and heteroceres,
indigenous and exotic; with descriptions and colored illustrations. Reading,
Pennsylvania, priv. pubL: 143 pp., 15 pis.

WILSON, E. 0., & w. L. BROWN, JR., 1953. The subspecies concept and its taxonomic ap-
plication. Syst. Zool., 2:97-111.

Journal of Research on the Lepidoptera

20(4): 199-204, 1981(83)

Nomenelature, Taxonomy and E¥olution

Paul R. Ehrlich
and

Dennis D, Murphy

Department of Biological Sciences, Stanford University, Stanford, CA 94305

This response to Miller and Brown (1983) will have to be relatively brief,
since we do not have room here to teai h elemental^ courses in systematics
oi evolution. Miller and Brown seem to be undei the impression that we
mwe critical of their taxonomy, as the title of their paper— “Butterfly
Taxonomy, A Reply”— indicates. On. the contrary, as the title and content
of our original paper (Eiiriicli and Murphy, 1982) show, we were critical of
the numemlature they used, not the taxonomy. They thus miss the entire
point of our critique, because nomenclature is not taxonomy, and certainly
not biolo©^! In his basic text Principies of Systematic Zoology, Ernst Mayr
(1969, pp. 407 and 413) defines nomenclature as “a system of names” and
taxonomy as “the theory and practice of classifying organisms.” He
further points out (p. 297):

Tt is the role oJ nomenclature to provide labels for taxa at all levels,
ill order to facilitate communication among biologists. The scientific
names for species oC organisms and for the higher taxa in which they
are placed form a system of communication, a language; they must
fulfill the same basic requirements as any other language.”

Mayr’s three outstanding attributes for scientific nomenclature .are
uniqueness, universality, and stability. Of the latter he says (p. 298):
“As recognition symbols the names of objects would lose much of
their usefulness if they were changed frequently and arbitrarily. It
would surely create confusion if we were to call an object a spoon
today but an apple next week. Yet this basic principle of com-
iminication lias been constantly violated by zoologists. Altogether
too mucli name changing has occurred in zoological taxonomy
during the past 200 yeai“s.”

Similar seiitiiiients are found in the other standard source, George
Gaylord Simpson's Principles of Animal Taxonomy (1961). For example,
he m.akes the tollowing point with emphasis (p. 112):

“A published classification in current use should be changed when it is
definitely incomislent with known facts and accepted principles, but
only so far as necessary to bring it into consistency. ”

200

J. Res. Lepid.

A nomenclature can remain relatively conservative to facilitate com-
munication while the underlying classificatory system can be more fluid to
represent, at best partially, better understanding of relationships. But, as
Simpson admonishes, there should be strong reasons even for changing
the taxonomy, let alone the nomenclature.

In this context, the notion that we were recommending “censorship” in
proposing to stick to names in Howe unless there were reasons (clear
polyphyly, highly distorted balance) for not so doing is seen as preposterous.
Standardization is routine in scientific discourse. If Miller and Brown
submitted to Science or any other refereed scientific journal an article that
contained measurements in chains, rods, bushels, or pecks it would be
rejected until those measurements were replaced with their metric
equivalents. Would that be censorship??

We cannot help but note in passing, in terms of “censorship” that the
Journal of the Lepidopterists’ Society refused to publish our original
manuscript on the novel grounds that the Society had published Miller-
Brown. This refusal was surely a “first” for purportedly scientific journals,
the rest of which routinely publish critiques of their previous articles.
Furthermore the society is now going to require that all contributors to its
season summary follow Miller-Brown nomenclature {News of the Lepidop-
terists’ Society, Sept/Oct 1982, p. 62), even though a major segment of the
society (see acknowledgments to Ehrlich and Murphy, 1982) — in fact, if
our sample is representative, the vast majority of its members — think that
nomenclature a disaster.

Some specific replies:

1. Catalogues are not the place to put unexplained new taxonomic
arrangements.

2. There are no “overlooked” genera; most of the ones resurrected in
Miller-Brown were correctly long-ignored.

3. The “no new names” issue begs the question. Names dredged out of
synonymy where they have properly resided for a century and a half are
operationally “new.”

4. Generally the notes in Miller-Brown are utterly inadequate to justify
the nomenclatural changes, since they do not deal with crucial issues of
polyphyly or balance. For many of the more egregious choices, such as the
resurrection of Pterourus, no explanation is given there at all. In others
they follow nomenclaturally incompetent “revisions,” for instance, ac-
cepting ''Occidryas” even though Higgins (1978) gives no valid reason for
proposing it as a genus.

5. Not using a name does not mean it will be “expunged from the
literature.”

6. Miller and Brown nicely summarize aU of the reasons that their
nomenclature should be rejected in the section “The usual objections...
biological purpose.” Hovanitz’ (1964) objection to the splitting of Speyeria

20(4): 199-204, 1981(83)

201

bom Argynnis is perfectly valid == we have not suggested going back simply
because Speyeria has now become widely accepted. Hovanitz' other
comments show how prescient he was about the downhill slide of butterfly
nomenclature.

7. Nowhere did we “suggest” that the division of Papilio was based on
genitalia— our only reference to genitalia was to the dependence upon
them at the specific level. Miller and Brown now justify the nomenclatural
inflation primarily on the basis of food plant differences. Their main
criterion is that '^Papilio’' feeds on Umbelliferae, ''Heraclides’' on
Rutaceae, and '*Pterourus'’ on lauraceous plants. But ''Heraclides'' is also
on Piperaceae, ''Papilio” is commonly also on Rutaceae and Compositae
in addition to Umbelliferaes and "Pterourus” feed on many families
{Papilio giaucus and P. rutulus are recorded from at least 15 families)
mciuding Rutaceae! The correct interpretation of Berenbaum (a she not a
“he”) is not that natural groupings of swallowtails feed on natural
p-oupings of plants, but that certain swallowtails are highly catholic in
their choice of oviposition plants, as long as the plants share a similar
chemical stimulant.

But even if different subgenera or species groups did feed exclusively on
different groups of plants, that, in itself, is not an excuse for raising them to
generic status. The subtleties of the factors governing larval host plant use
by butterflies are Just beginning to be elucidated (e.g., Chew, 1977;
Holdren and Ehrlich, 1982; Lincoln, et aL, 1982; Murphy, 1983; Rausher,
1982; Singer, 1972; Wiklund, 1982). Taxonomic affinity of the plants is
just one such factor, and in many cases a minor one to boot (Janzen, 1979).
Celastrina argiolus is known to feed on at least 18 plant families and
Strymon melinus on 28. Will the next nomenclatural epic fraction them into
18 and 28 genera respectively?

The basic point, of course, recognized by all well Trained taxonomists, is
that levels of genera, subgenera, etc. are biologically purely arbitrary, and
that the use of genera should therefore be conservative to aid in
communication (Mayr, 1969, p. 239). In addition, reclassification of
Papilio does not remotely meet Simpson’s criterion— there are no “known
facts and accepted principles” that are violated by retaining Papilio in the
sense that it has been used for the past few decades. The bottom line is
that even the discovery of natural groups within Papilio would not in itself y
be a reason to split the genus and change hundreds of names. Polyphyly or
severe imbalance would be such a reason, but unless one or the other can
be clearly demonstrated, taxonomic structure within Papilio should be
recognized by subgenera or species groups.

Additionally, we must note Miller and Brown’s story about Papilio
polyxenes coloro. Although it has no direct relevance to the issue under
discussion, it highlights how casual speculation in the literature gets
ta-anslated into fact. Ferris and Emmel (1982) present not one shred of

202

J. Res. Lepid.

evidence bearing on whether P. zelicaon did, does, or can “outcompete” P
p. coloro — nor do they claim to. Such evidence would be of enormous
interest to the ecological and evolutionary communities as a whole, as
competitive exclusion has never been demonstrated in herbivorous
insects.

8. Large, uniform genera should not be broken up just because they are
large — quite the opposite. And to fractionate small, uniform genera such

as Euphydryas (12 species) is absurd. What possibly could be gained by
dividing it in four?

9. Consistent application of subgenera does not mean burdening
communication with them if there is no need. The genius of the Linnean
system of binomial nomenclature is its parsimony — it avoids the older
system of using an entire phrase to denote an organism. The idea is to
maximize communication while restricting oneself to a two-part name.

10. Nomenclature that is “recognized by specialists” is, especially in
groups like the butterflies where many specialists have extremely narrow
training (or none at all), almost invariably oversplit. Rare is the specialist
who considers broad balance in his or her application of names. One result
of this is a continual shifting of names, often accompanied — as in the
Miller-Brown catalogue^-by no significant advance in understanding of
the organisms. Continual name -changing, we repeat, is the major reason
why most biologists consider taxonomy a non-science. Taxonomy is too
important to evolutionary and ecological biology to destroy its reputation
to please those who confuse manipulating names with science.

11. We are glad to be told that evolution is a dynamic process. Miller and
Brown might like to be informed that everything they say in the three
paragraphs that start “It cannot be questioned that evolutionary prob-
lems. . and end “. . .classification or phylogeny.” is gibberish, irrelevant
to the debate at hand, or both. Those familiar with taxonomic and
evolutionary theory will see that by simply reading them. Others can get
the flavor by considering the phrase “Were evolution a static process. .
(our emphasis). Presumably what Miller and Brown mean is that evolution
in different lines can proceed at different rates (a textbook discussion can
be found in Ehrlich et al., 1974). If there have been significant rate
differences in the lines leading to different groups of butterfly species (it is
not known if there have been), this would not make one iota of difference in
whether conservative nomenclature could be applied to the products of
butterfly evolution.

It might be noted that the question of whether nomenclature should be
conservative is not only independent of evolutionary rates but also of
notions of what kinds of relationship should be the basis of taxonomic
schemes. For example, Ehrlich and Ernst Mayr were on opposite sides of
the phenetics vs. phyletics arguments of a quarter of a century ago, but
they are in close agreement on keeping obligatoiy categories conservative.

20(4): 199^204, 1981(83)

203

12. From the comment on “political morality” we can only assume that
Miller and Brown are more interested in conserving generic names than in
conserving butterflies. We would claim that the only moral course (indeed
the only sane one) is to use every available scientific and political tool at
our disposal in attempts to save Earth's dwindling biological resources.
This issue is explored in depfh elsewhere (Soule and Wilcox, 1980; Ehrlich
and Ehrlich, 1981).

13. Splitting and unsplitting genera, or the publication of catalogue/
checklists, will not bring us any closer to a “true phylogenetic classification
of the Lepidoptera.” Even if “phylogenetic” is rigorously defined, a
phylogenetic classification is not even necessarily the most desirable goal.

One might view this whole argument as scientifically trivial, but it is not.
Sound nomenclature is important to evolutionists, ecologists, and other
biologists as well as systematists. Butterflies are prominent organisms,
fast becoming one of the most important groups of experimental animals.
Confusing and senseless changes now will only impede scientific investiga-
tion, confuse serious amateur lepidopterists, and unnecessarily further
lower the esteem of taxonomists in their colleague's eyes.

In summary, we state again that the Miller- Brown catalogue/checklist is,
as a bibliographic tool, one of the most useful publications on North
American butterflies ever to appear. In its introduction (p. v) we find that
of the two authors “the elder [Brown]. . .favors the use of subgenera, the
younger does not.” It is too bad that Brown’s mature taxonomic judgment
did not prevail. It would be a shame if the resultant misuse of generic
names were to be widely followed and thus cause the work to have an
overall negative impact on science as a whole and the study of butterflies in
particular.

Acknowledgments. We thank Carol L. Boggs, Richard W. Holm, John H. Thomas,
Ward B. Watt, and Bruce A. Wilcox for reading and criticizing this manuscript.

Literature Cited

BERENBAUM, M., 1978. Effects of linear furanocoumarins on an adapted specialist
insect {Papilio polyxenes). Ecol. Entomol. 6:345-351.

CHEW, F. s., 1977. Coevolution of pierid butterflies and their cruciferous foodplants
11. The distribution of eggs on potential foodplants. Evolution 31:568-579.
EHRLICH, P. R., & A. H. EHRLICH, 1981. Extinction: The Causes and Consequences of
the Disappearance of Species. Random House, New York.

EHRLICH, P. R., R. w. HOLM, & D. R. PARNELL, 1974. The Process of Evolution, Second
Edition. McGraw-Hill, New York.

EHRLICH, P. R. & D. D. MURPHY, 1982(1981]. Butterfly nomenclature: a critique. J. Res.
Lepid. 20:1-11.

HIGGINS, L. G., 1978. Aa revision of the genus Euphydryas Scudder (Lepidoptera:

Nymphalidae). Entomol. Gaz. 29:109-115.

HOLDREN, c. E., &p. R. EHRLICH, 1982. Ecological determinants of food plant choice in
the checkerspot butterfly Euphydryas editha in Colorado. Oecologia (Berl.)
52:417-423.

204

J. Res. Lepid.

JANZEN, D. R, 1979. New horizons in the biology of plant defenses, hi G. A. Rosenthal
and D. H. Janzen, eds. Herbivores: Their Interaction with Secondary Plant
Metabolites. Academic Press, New York.

mcoLN, D. E., T. s. NEWTON, p. R. EHRLICH, & K. s. WILLIAMS, 1982. Coevolution of the
checkerspot hnttev^y Euphydryas chalcedona and its larval food plant Diplacus
aurantiacus: larval response to protein and leaf resin. Oecologia (Berl.) 52:
216-223.

MAYR, E., 1969. Principles of Systematic Zoology. McGraw-Hill, New York.

MILLER, L. D. & F. M. BROWN, 1981(82). Butterfly taxonomy: a reply. J. Res. Lepid.
20(4):193-198.

MURPHY, D. D., 1983. Nectar sources as constraints on the distribution of egg masses
by the checkerspot butterfly Euphydryas chalcedona (Lepidoptera: Nymphali-
dae). In press, Envir. Ent.

RAUSHER, M. D., 1982. Population differentiation in Euphydryas editha butterflies:
adaptation to different hosts. Evolution 36:581-591.

SIMPSON, G.G., 1961. Principles of Animal Taxonomy. Columbia, New York.

SINGER, M. c., 1972. Complex components of habitat suitability within a butterfly
colony. Science 176:75-77.

SOULE, M. c. & B. A. WILCOX, 1980. Conservation Biology. Sinauer and Co., Sunder-
land, Mass.

WKLUND, C., 1982. Generalist versus specialist utilization of host plants among
butterflies. In press, Proc. 5th Int. Symp. on Insect-Plant Relationships,
Wageningen.

Journal of Research on the Lepidoptera

20(4): 205, 1981(83)

Editor's Note

The above papers raise an issue which cannot be ignored, with regard to
publication of and use of butterfly nomenclature. The crux of the issue is
stability, with its corollary of conservatism. Under the present circum-
stances the nomenclature of the holarctic butterflies is anything but
stable, not only by virtue of the new catalogue/checklist and substantial
supportfor a different view of both taxonomic concepts and nomenclature,
but by the proliferation of generic names in the Palearctic region.
Pragmatically, the situation leaves this journal with the problem of what
should be the accepted nomenclature for submitted papers. It is our
position to accept the judgment of our reviewers in cases of controversy.
Selection of reviewers will be made, insofar as possible, to provide
balanced judgments. Thus, where options are available, the final decision
will be between author and reviewers. Our bias, as was the strong bias of
the founding editor, William Hovanitz, is for the conservative position. We
wish to make very clear that despite this bias, we will not impose any
nomenclature as an editorial fiat. After all, the nomenclature is largely the
result of taxonomic opinion, which, in absence of convincing supportive
data for most groups, is just that.

We do emphasize, though, that a consensus on nomenclature, at least of
the higher classification, be arrived at as a matter of the highest priority for
Lepidopterists. The reasons for such consensus are clearly stated by
Ehrlich and Murphy. This Journal will continue to publish relevant
considered judgments on the subject with the objective of reaching that
goal.

It should be reiterated that we do not wish in any way to impugn
publications of the Lepidopterists Society or create the impression of any
relationship other than one of respect and support for that organization.

Editorial Board: R. H, T. Mattoni, John Emmel, Arthur Shapiro, Bjorn Petersen,
Otakar Kudma, Miguel R. Gomez Bustillo, Emilio Balletto,

Atuhiro Sibatani, Ichiro Nakamura.

Journal of Research on the Lepidoptera

20(4): 206, 1981(83)

Acknowledgment

We wish to acknowledge our depth of gratitude to many individuals who
so generously gave of their time in serving as referees for manuscripts
submitted to The Journal of Research on the Lepidoptera. The quality of
work which now appears in this journal has shown consistently higher
standards as a consequence. We particularly appreciate the promptness
with which reviews are received. Although publication may be delayed for
various reasons, the first courtesy to authors is return of comments on
their works in a timely manner. This is a courtesy virtually all reviewers
have graciously extended. The following listing cites those individuals who
have served as referees to date, covering papers which have appeared and
will yet appear into volume 22.

Philip Ackery

Glenn Gorelick

Austin Platt

Richard Arnold

L. P. Grey

Jerry Powell

Emilio Balletto

John Hafernik

Robert Robbins

Michael Bentzien

Reink de Jong

Ronald Rutowski

Lincoln Brower

John Heppner

Klaus Schurian

F. Martin Brown

Ron Hodges

James Scott

Frances Chew

Otakar Kudrna

Arthur Shapiro

Julian Donahue

Jorge Llorente

Oakley Shields

Paul Ehrlich

Don MacNeill

Steve Sims

John Emmel

William McGuire

Ray Stanford

Tom Emmel

R. H. T. Mattoni

Paul Tuskes

Larry Eng

Lee Miller

Richard Vane- Wright

Cliff F erris

Scott Miller

Ward Watt

John Franclemont
Lawrence Gall
Hanjurgen Geiger

Dennis Murphy
Ichiro Nakamura
Charles Oliver

Ray White

In addition we wish to express our gratitude to the Santa Barbara
Museum of Natural History for generously providing space and facility for
storage of our inventory of back issues and related publishing materials. In
particular we wish to thank Drs. Dennis Power and F. G. Hochberg of that
institution for their assistance and many courtesies.

Journal of Research on the Lepidoptera

20(4): 207-213, 1981(83)

Notes on the Life History and Baja California
Distribution of Chlorostrymon simaethis sarita (Skinner)
(Lepdioptera: Lycaenidae)

John W. Brown*

791 My Way, San Diego, California 92154

Abstract. The early stages of Chlorostrymon simaethis sarita (Skinner) are
herein described. Larval behavior in association with the host plant
Cardiospermum corindum Linnaeus (Sapindaceae) is briefly discussed.
Additionally, data regarding the insect’s temporal and spatial distribution in
Baja California, Mexico, are presented.

Introduction

Chlorostrymon simaethis (Drury) is an exquisite little neotropical hair-
streak that occurs from Florida and Texas south through the Antillies,
Mexico, Central and South America (Nicolay, 1980). Although commonly
associated with balloon vine {Cardiospermum Linnaeus) (Sapindaceae)
throughout its entire range, the early stages of C. simaethis are seldom
encountered. Zikan (1956) briefly described the last instar larva and pupa
from specimens collected and reared near Itatiaia, Brasil.

In Baja California, Mexico, the subspecies C. simaethis sarita (Skinner)
is occasionally moderately abundant. Additionally, balloon vine has a
rather widespread distribution throughout the peninsula of Baja California
and on the adjacent islands (Wiggins, 1980), with 3 species present, all in
the genus Cardiospermum. The plant’s fruiting bodies are round or
angular, inflated, papery, three-chambered pods 2-4 cm long, 3-5 cm wide,
and strongly veined (Coyle and Roberts, 1975). Each chamber usually
bears a single pea-like seed, although in some cases only one or two of the
three seeds develop to maturity.

Larvae of C. simaethis sarita were discovered inside the pods of a large
Cardiospermum corindum Linnaeus at a locality approximately 10 km
north of Rosarito, Baja California Norte, 8 April 1982, by David Faulkner
and the author. Almost all stages of larval development were present.

Early Stages

First instar. Length 2-4 mm. Body eruciform, light translucent green
with a whitish overcast. No conspicuous markings. Second instar. Length

'Departmental Assistant, Entomology Department, San Diego Natural History Museum, P. 0.
Box 1390, San Diego, CA 92112

208

J. Res. Lepid.

Fig, 1. Last instar larva of Chlorostrymon simaethis sarita inside pod of Cardio-
spermum corindum.

5-6 mm. Body light green to gray-green with a faint mid-dorsal darker
stripe. No additional markings. Third instar. Length 7-8 mm. Body slug-
shaped, color quite variable from light olive green to light brown, a darker
green longitudinal stripe running the entire mid-dorsal length. Body
occasionally with minute reddish speckling. At mid-dorsal crest of each
segment 2-8 is a wedge-shaped reddish brown mark and a slight ovaloid
indentation. Marks usually well defined but may be quite faint. Mark on
third segment enlarged, extending laterally almost over entire dorsal
surface of that segment. Mark on second segment extremely hght. A faint
blackish gray longitudinal stripe on each side of the body. Head
concolorous with body. Final instar. Length 9.0-11.5 mm; width 4.5 mm.
Color and markings variable. Body color from light green to light brown,
becoming powdery brick red as larva approaches pupation. Most of body
covered with fine, short black hairs. Each segment with a black semi-
longitudinal dash midway between the apex and the prolegs; the dashes
together forming an irregular, wavy, longitudinal stripe on each side of the
larva. A mid-dorsal greenish black stripe runs the length of the larva, and a

20(4): 207-213, 1981(83)

209

small, ¥ariable faint-to- dark reddish brown oval-shaped marking occurs on
each side of the stripe at the posterior crest of each segment 2—8. The
markings together superficially appear as a reddish brown longitudinal
stripe down the mid-dorsal apex. Mark on the third segment enlarged,
extending laterally. Spiracles unmarked. Head, when extraded, reveals
two conspicuous, small black ocelli. Mandibles brownish black. Pupa*
Length 7. 5-8.5 mm; width 4.5 mm. Short, stout, and rounded. Color
variable, light browm to dark grayish brown, mottled with black and brown
splotches. Ventral surface much lighter. Dark black markings inconsis-
tently border the wing cases. A black comma-shaped spot on each side of
the body near the region of the head. A faint black mid-dorsal longitudinal
stripe; and a tMn, black lateral stripe where abdominal segments six and
seven join. Entire body sparsely covered with fine light brown hairs, wing
cases bare.

Adults emerged as follows: 299, 4 May 1982; 19, 5 May 1982; 299, 8
May 1982; and 19, 10 May 1982. Five specimens are deposited in the
collection of the San Diego Natural History Museum, and one specimen
was given to Glenn Gorelick, Sierra Madre, California.

The molts were determined by observation of distinct changes in size of
the larvae accompanied by subtle changes in coloration and markings. No
head capsules were obser\^ed or recovered, and no immature stages were
preserved.

Behavioral Notes

Eggs are apparently deposited singly on the developing flower heads.
Larval development takes place 'entirely within the fruiting pods. First
instar larvae bore into a developing gi'een seed, leaving behind a small
entrance hole. Feeding initially takes place inside the seed and the frass is
generally deposited on the exterior of the seed. The larva eventually’
consumes aU of the seeds within a single pod, moving between the
chambers of the pod by boring through the membranous structures
dividing them. Occasionally even the membrane is consumed by the larva.
There is never more than one larva within a single pod. Cannibalism might
account for tlue fact since many lycaenid larvae, under crowded conditions,
are known to devour their siblings (Downey, 1962). However, no can-
nibalism was observed. A single egg per bud is suggested by the
occuirence of a single larva witMn a pod. Externally there is no difference
m the appearance of inhabited and uninhabited pods.

After al the seeds within the pod have been eaten, the mature larva
bores out of the pod and crawls away to pupate. Pupation occurs in debris
and loose soil beneath the host wiihout a giirfle -string attachment. Under
laboratoiy conditions, pupation of progeny of the spring brood lasted from
14 to 17 days.

It is not faiown whether a larva can undergo complete development

210

J. Res. Lepid.

within a single pod, and the data available are inconclusive. Pods which
had an exterior hole contained no larvae and were full of frass, indicating
that the hole is an “exit” hole only, implying the use of a single pod. This
was, however, contradicted by the fact that under laboratory conditions,
mature -appearing larva continued to feed if offered a second opened pod.
Also, Zikan (1956) reported the use of multiple pods of Cariospermum
halicacabum Linnaeus. The contents of two pods seemed to be the
absolute maximum required for complete larval development.

Observations by Fred T. Thome (personal communications) indicate
that newly hatched first instar larvae refused to feed on foliage of
Cardiospermum halicacabum even when offered tender young growth. As
C. halicacabum has been documented by Zikan to be a suitable host, this
further suggests that the larval diet is indeed limited to the developing
seeds. Although many lycaenid larva feed on vegetative parts of their
respective hosts, several feed exclusively on plant reproductive stmctures
such as flowers and seeds (Downey, 1962). The latter is the case with C.
simaethis sarita.

Distributional Notes

Chlorostrymon simaethis sarita occurs the entire length of the peninsula
of Baja California, Mexico (Fig. 4). Captures have been recorded from the
cape region of Baja California Sur to the Sierra San Pedro Martir in Baja
California Norte. Strays are known from as far north as the Colorado
Desert in San Diego County and Palm Springs in Riverside County,
California, much to the north of the known range of the host. The single
confirmed host in Baja California is the widely ranging introduced Cardio-

Fig. 2. Last instar larva of C. simaethis sarita, lateral view.
Fig. 3. Last instar larva of C. simaethis sarita, dorsal view.

20(4): 207^213, 1981(83)

211

Kg. 4. Spatial distribution of C. simaethis sarita in Baja California, Mexico.

spermum corindum. It is not known whether C. simaethis sarita utilizes the
other two species of Cardiospermum, both of which are endemic to Baja
California.

In the northern arid region of the peninsula C. simaethis sarita appears to
be capable of three broods, with the adult emergence probably dictated by
rainfall. Capture records indicate flights in late March to mid- April mid-
June to mid- July, and again in September through November. In the cape

212

J. Res. LepM.

region of Baja California Sur specimens have been recorded in ever}.^
month from October to March representing at least two broods »

Specimens Examined

Depositories abbreviated in the text are as follows: CAS, California
Academy of Sciences, San Francisco, California; Cl, College of Idaho,
Caldwell, Idaho; HF, Allan Hancock Foundation, University of Southern
California, Los Angeles; SDNHM, San Diego Natural Histoiy Museum,
San Diego, Califorma, and RB, Richard Breedlove, San Diego, CaMfomia,

MEXICO: BAJA CALIFORNIA NORTE: Punta Prieta, 2 F, 28 March 1935
(CAS), 1 F, 23 Dec. 1976 (D. Lindeley, SDNHM); Sulfur mine S San Felipe, 2 F, 1
M, 12 Nov. 1967 (D. Patterson, CAS); 7 mi W Las Arrastrae, 3 F, 18 M, 4 Nov. 1967
(D. Patterson, CAS); 14.9 mi N Laguna Chapala, 1 F, 1 M, 3 April 1973 (D.
Patterson, CAS); 7 mi SW Mission San Borja, 1 M, 31 March 1973 (Donohoe &
Patterson, CAS), 1 F, 3 M, 30 March 1973 (J. Powell, CAS); 7 mi E Rancho
Rosarito, 16 F, 11 M, 30 March 1973 (Donohoe & Patterson, CAS); 9 fan NW
Rancho Santa Inez, 1 M, 12 July 1979 (A. Chu, Cl); Rancho Santa Inez, 1 F, 14 June
1979 (J. Miles, Cl); Las Encinae, Sierra San Pedro Martir, 1 M, 17 June 1980 (J.
ftuwn, SDNHM), 1 M, 13 July 1980 (J. Brown, SDNIM); vie. ent. Parque
Nacional Sierra San Pedro Matir, 1 F, 22 June 1979 (J. Brown, SDNHM); Bahia
San Francisquito, 1 F, 3 April 1947 (C. Harbieon,' SDNHM); IslaTiburon, Golfo de
CaMfomia, 2 F, 3 M, 19 March 1962 (C. Harbison, SDNHM); 2.1 mi S Rosarito, 1 M,
23 March 1981 (Faulkner & Andrews, SDNHM); 10 km N Rosarito, 2 F, ex-larvae,
em: 4 May 1982, 1 F, ex~larva, em: 5 May 1982, 2 F, ex-larvae, em: 8 May 1982, 1 F,
ex-larva, em: 10 May 1982 (J. Brown, SDNHM).

BAIA CALIFORNIA SUR: Espiritu Santo Island, 2 F, 7 March 1928 (Craig,
CAS), 2 F, 7 M, 22 Feb. 1936 (J. Garth, HF), 1 F, 11 Feb. 1940 (J. Garth, HF), 1 F, 9
^ 19 Feb. 1932 (J. Garth, HF); 10 mi N Todos Santos, 4 F, 9 M, 26 Dec. 1958
(Leech, CAS); Coyote Cove, Bahia Concepcion, 1 F, 1 Oct. 1941 (CAS); 13.1 mi
NW La Pai, 2 M, 3 Jan. 1959 (Leech, CAS); Cabo San Lucas, 1 F, 17 Jan, 1959
(Leech, CAS), 2 F, 31 Dec. 1978 (Rude, CAS); 5.6 mi SE San Perdito, 1 F, 6 Oct.
1981 (D, Faulfaier, SDNHM); 12.2 mi SE San perdito, nr Rancho Saucito, 2 F, 8
Oct. 1981 (Faulfaier & Andrews, SDNHM); 14 mi N Todos Santos, 1 M,4 Oct. 1981
(Brown & Faulfaier, SDNHM); 29.9 mi S Loreto, 1 F, 11 Oct. 1981 (Faulkner &
Andrews, SDNHM); 5 mi W Loreto, 8 F, 27 Dec. 1976 (D. Lindsley, SDNHM);
BaMa Concepcion, 1 F, 29 Dec. 1976 (D. Lindsley, SDNHM); 2 mi W Loreto, 1 F, 4
12 Dec. 1976, 2 mi SW Loreto, 1 F, 3 M, 4 Dec, 1977, 2-3 km SW Loreto, 1 F, 29
Nov. 1977, 1 F, 30 Nov. 1977 (aU G. Forbes, SDNKM); Hotel Mulege, 1 F, 23 March
1974 (G. Forbes, SDNHM); LaPaz, 1 F, 9 Nov. 1952 (SDNHM), 1 F, 1 M, 29 Nov.
1979 (Brown & Faulkner, SDNIM); Cabo San Lucas, 1 F, 11 Nov. 1952 (SDNHM),
Hotel Finisterra, Cabo San Lucas, 3 F, 1 M, 30 Nov. 1979 (Brown & Faulkner,
SDNIM), 4 F, 1 M, 28 Nov. 1980 (Brown & Brown, SDNHM); San Bartolo, 3 F, 1
M 30 Nov. 1979 (Brown & Faulfaier, SDNHM).

CALIFORNIA: RIVERSIDE CO.: Palm Springs, 1 F, 9927 (SDNHM).

SAN DIEGO CO.: Mason Valley, Anza Desert, 1 M, 8 Oct. 1967 (G. Forbes,

20(4): 207=213, 1981(83)

213

SDNHM); Anza Desert, Vallecitos, 1 M, 11 Nov. 1967 (R. Breedlove, RB); Dexter
Peak, Descanso, 1 F, 21 May 1966 (R. Breedlove, RB).

Acknowledgment. I am sincerely grateful to David K. Faulkner, Chairman,
Entomology Department, San Diego Natural History Museum, for making possible
our numerous trips to Baja California, Mexico. I wish to thank the following for
reviewing the manuscript: Fred T. Hiome, Curator Emeritus, San Diego Natural
History Museum; Roy Kendall, San Antonio, Texas; Dr. John Emmel, Hemet,
California; Rudolf Mattoni, Beverly Hills, California; and especially an anonymous
referee for his many helpful suggestions. Thanks are also extended to Paul Amaud,
Califomia Academy of Sciences, San Francisco, California; William Clark, CoUege
of Idaho, Caldwell, Idaho; and John Garth, Allan Hancock Foundation, University
of Southern Califomia, Los Angeles, for allowing me to transcribe data from
specimens at their respective institutions. Additionally, I am grateful to Poody
Brown for assistance in rearing and photographing the specimens, Shirley Latislaw
for the distributional map, and Rosemarie Fiebig for translating the Zikan (1956)
paper.

literature Cited

COYLE, J. & N. ROBERTS, 1975. A field guide to the common and interesting plants of
Baja Califomia, pg. 118. Nat. Hist. Pub. Co., La Jolla, CA.

DOWNEY, J., 1962. Host-plant relations as data for butterfly classification. Systematic
Zoology 11(4):150-159.

MCOLAY, s., 1980. Tlie genus Chlorostrymon and a new subspecies of C. simaethis.

Journal of the Lep. Soc. 34(2):263-256.

WIGGINS, L, 1980. Flora of Baja Califomia, pp. 803-804. Stanford Univ. Press.
ZIKAN, J., 1956. Beitoag zur biologic von 12 Theclinen-arten. Dusenia 7(3):139-148.

Journal of Research on the Lepidoptera

20(4): 214-234, 1981(83)

Biology and Immature Stages of Australian
Ethmiid Moths (Gelechioidea)

Jerry A. Powell

Department of Entomological Sciences, University of California, Berkeley, CA 94720

Abstract. Biological data for 5 species of Ethmia are reported, E. sphaero-
sticha, E. postica, E. thoraea, E. heptasema, and E. heliomela, based on
observations in New South Wales and Queensland in 1980-81. Their eggs
are characterized, along with larvae and pupae of the first and last species.
The larva and pupa of E. hemadelpha from Western Australia and the pupa
of E. nigroapicella from Hawaii are described. Known or suspected food-
plants of all seven are Boraginaceae.

Ethmiids are small, often brightly colored moths sharing a general
similarity in superficial appearance throughout all faunal regions of the
world. There are remarkable differences, however, in less obvious
morphological characteristics of the adults, in biological traits, and in the
eggs, larvae, and pupae. The group is distinct taxonOmically and has been
regarded as a family (Battler, 1967; Common, 1970; Powell, 1973;
Kuznetsov & Stekobnikov, 1979) or a subfamily of the large cosmopolitan
family Oecophoridae (Hodges, 1978). In species diversity, ethmiids are
best represented in areas of seasonal drought, such as the thorn scrub of
the northern Neotropical Region. Each species is restricted in larval
foodplant preference, and in general Ethmia are dependent upon Bora-
ginaceae and the closely related, primarily Nearctic family Hydrophyl-
laceae (Battler, 1967; Powell, 1973, 1980).

For its size, Australia has a depauperate ethmiid fauna, with only 14
species, but most of them are endemic, including some specialized forms
(Powell, 1982). Among these, three species groups with differing male
genitalia types exhibit striking similarity in these structures to Neotropical
species groups that are known or believed to possess uniquely derived
larval and pupal traits {Ethmia, Bection II of Powell, 1973). Thus it was of
considerable interest from a biogeographic viewpoint to discover whether
the derived features of the immature stages in the New World are shared
by Australian species.

During residence at the Division of Entomology, Commonwealth
Bcientific and Industrial Research Organization (CBIRO), Canberra, in
1980-81, 1 was able to obtain some information on the biology of 5 species
and received material collected by K. T. Richards of Perth, W.A.,

20(4): 214-234, 1981(83)

215

representing a sixth species. As a result, some data on oviposition
behavior are summarized, and larval and pupal descriptions are given for 3
species. With existing descriptions of Ethmia nigroapicella (Saalmueller)
(Moriuti, 1963; MacKay, 1972; Zimmerman, 1978), these data provide a
representative picture for 4 of 5 Australian species groups outlined
elsewhere (Powell, 1982).

Techniques

Methods of handling living material were generally similar to those given
previously (Powell, 1971). Moths were confined for oviposition either in
plastic tubs ca. 25 x 25 x 13 cm with a cardboard floor and nylon stocking
mesh ceiling, or in 10 x 1.5 cm glass or plastic petri dishes with nylon lining
the lid. Newly hatched larvae were reared on branchlets of foodplant in
plastic pill boxes or small refrigerator boxes; larger larvae were housed in
plastic tubs or polyethylene bags lined with paper towels. Full grown
larvae were offered folded paper towels and layers of soft corrugated
cardboard, owing to the preference among many Holarctic Ethmia for
burrowing into «oft material for cocoon construction.

Rearing was conducted in uncontrolled laboratory temperatures, which
varied widely. During hot periods larval lots were housed in better
insulated, cooler rooms to deter disease. Nonetheless two larval groups
became diseased, a common problem with communal rearing of Ethmia in
North America (Powell, 1971).

Hostplant associations have been determined for three species, E.
hemadelpha, E. sphaerosticha, and E. heliomela, by discovery of larvae in
the field. For others, attempts were made to elicit oviposition response by
providing Boraginaceae suspected to be hostplants, by correlation of the
geographic distribution of moths and plants according to records in the
Australian National Insect Collection (ANIC) and Herbarium Australiense,
CSIRO, Canberra. Plant samples presented to moths or caterpillars were
obtained from the National Botanic Garden, Canberra, or commercial
nurseries.

Scanning electron micrographs were executed by Barry Filshie, using
living eggs on leaf substrate. Ethmiid eggs seem to resist distortion due to
the vacuum when processed in situ on leaves and need not be metal coated,
hi preservation, larvae were killed in hot water just below the boiling point.
After sufficient distension occurred, they were transferred to Kahle’s
solution and ultimately to 95% EtOH. Measurements in larval and pupal
descriptions were made by a micrometer disc at 6.3 to 20X magnification,
of eggs at 20X to 40X. The number of individuals measured or
observations upon which statements are based is indicated (n).

Consistent ' biological features of ethmiids have been summarized
elsewhere (Powell, 1971, 1973) and are not repeated here. An exception to
the oviposition pattern shown by Nearctic Ethmia occurs in two Australian

216

J. Res. Lepid.

species, E. heptasema and E. heliomela, which usually deposit eggs in
small clusters. In all previous records for the genus, and the 3 other
Australian species I observed, oviposition is single. The eggs of E.
sphaerosticha are flat, unlike any others known in the genus. None of the 4
species for which late instar larvae and pupae are known possess any of the
uniquely derived features characteristic of Ethmia Section II of the New
World.

Ethmia sphaerosticha (Meyrick, 1886)

This species is unique among all known ethmiids for the enlarged
antennae with expanding scale brushes in the male (Common, 1970: 814,
Fig. 36.27k). The egg and pupal anchoring mechanism also differ from any
previously described in the genus. E. sphaerosticha occurs in the rain
forests of eastern coastal mountains, disjunctly, in northern Queensland
and from southern Queensland to southern New South Wales (Powell,
1982).

Adult behavior. — Although they possess relatively small eyes (eye index
0.85; see Powell, 1973:8), the moths are nocturnal, and both sexes are
attracted to lights. One male was taken flying at 1715 hrs, before sunset,
but in laboratory confinement moths were diumally inactive. Field-
collected females (80M25, 81A105, A113, A121) survived 2-9+ days
(x=6, 7n). Two failed to oviposit, but the others deposited 25-30 eggs
each. One female laid 10 eggs on day 7 of confinement, although only water
was provided as nourishment. All eggs were deposited singly.

Females were housed in a plastic bag and plastic tub with a bouquet of
Ehretia acuminata R. Br. in bloom (80M25) or in petri dishes vdihEhretia
leaves. About 50% of the eggs were placed on leaves, almost always on the
undersides, irrespective of whether leaves were presented to the moths
upsidedown or topside up, or as an upright bouquet. Only two eggs were
laid on the upperside of a leaf in crevices. The remainder were placed on
plastic or glass surfaces above the plant. None was laid on flowers, fruit or
stems, or on the nylon mesh that is often selected by Ethmia (Powell,
1971).

Egg. — The eggs are flat, like those of tortricids, with both overall shape
and surface sculpture (Figs. 1, 2) unlike those of any other described
ethmiid. They ranged 1.17-1.40 X 0.67-0.80 mm in length and width and
were weakly convex, less than 0.25 mm thick. Eggs took the form of the
substrate, and their outline was highly variable. To the unaided eye the
eggs appeared milky, and the color did not change during development,
which was faster than in any other microlepidopteran I have observed.
Only 5-6 days (80M25) or 4-5 days (81A113) were required until hatching.

Larva.— 'Newly hatched larvae were whitish, semi- translucent, including
the head capsule (HC). They fed only on undersides of leaves, skeletonizing
the surface and cutting tiny holes through the full thickness. After feeding

20(4): 214^234, 1981(83)

217

Mgs. 1-6. Eggs of Australian Ethmia: 1, egg E. sphaerosticha on underside of
Ehretia acuminata leaf (x30); 2, E. sphaerosticha, closeup of chorion
sculpture (x235); 3, eggshell E. postica on nylon mesh (x42); 4,
eggshell E. §ioraea on leaf of Cynogiossum amtrak (x45), 5, egg
clutch E. heptasema on underside of Ehretia acMminata leaf (x33); 6,
E. Iwptasema closeup of cliorion sculpture (x200). Magnitications
approximated to published size.

218

J. Res. Lepid.

2-3 days, larvae appeared slightly greenish due to the gut contents and
were extremely well camouflaged on the leaf surface. By the eighth day all
larvae were in the 3rd instar, continuing to feed on the lower surfaces of
leaves. Later instars also fed from beneath leaves, eating large holes but
not feeding at the margins. They rested under flat webs on older leaves,
migrating out to feed on newer foliage (in laboratory conditions). The flat
webs were used for moulting by all instars, and seemed to be constructed
only for this purpose. In the field (81A107) and lab (81A113) most
moulting webs of antepenultimate and penultimate instars were con-
structed on uppersides of leaves no longer used for feeding or on container
walls.

There were 6 discrete instars according to HC measurements (mm):

I -= 0.31 (unfed) to 0.33 (shed HC); H = 0.41 to 0.51 (shed HC’s); HI =
0.98 to 1.02 (shed HC’s); IV — 1.19 to 1.23 (pres, larvae); V “ 1.42 (pres,
larva) to 1.47 (shed HC); VI — 1.76 (starved) and 1.96 (preserved) to 2.10

(living).

The first 3 instars were whitish with no integumental markings. A small
black spot appeared on each frontal lobe in IV, along with tiny spots on D1
pinacula. In instar V larvae were cream colored, their spots increased in
size, and sometimes small crescents were present below D2 and L
pinacula. Extensive, variable black markings characterized the yellowish
final instar (Fig, 7).

Growth was more rapid than that recorded for any North American
ethmiid (Powell, 1971). The final instar was reached on day 15-17
following egg hatch, and full grown larvae appeared ready to pupate by day
19-24 (81A113), despite having fed on which had been refrigerated

25-35 days. Those in lot 80M25 were not given additional leaves after day
7, owing to my absence on a field trip, yet nearly all reached the final instar
and were starved or diseased by day 18. None was reared from egg through
pupation, but those collected as last instar larvae produced adults 18-24
days following collection (80L47.1, 81A107). Thus the entire development
from egg deposition to emergence can occur in 40-46 days.

Final instar larva. — Length 15.8 to 18.3 mm (starved individuals). Head:
HC width 1.76 (starved) to 2.10 mm. Setation typical for the genus (e.g.
MacKay, 1972); Ah A^, A^ and L^ nearly on a straight line, with A^ only
slightly displaced posteriorly. Adfrontal sutures extending to cervical
angle but abruptly approximate well below, adfrontal areas extremely
narrow above P^. Cream colored, broadly blotched with black across
middle and narrowly along ventral border. Body: Primary setal arrange-
ments of thorax and abdomen as in fig. 7. Pinacula of D, SD strongly
raised. LI and L2 on abdomen widely separated, especially on Al, and
nearly on the same plane. Sclerotized depressions associated with
spiracles or thoracic leg setae absent. No secondary setae except the usual
gelechioid lower series of anal proleg. Integument yellowish or cream

20(4): 214-234, 1981(83)

219

7

Figs. 7, 8. Setal maps of Ethmia larvae, Roman numerals refer to thoracic
segments, arabic to abdominal: 7,E. sphmorsUcha; 8, E. henmdelpha.

220

J. Res. Lepid.

colored, variably mottled with black, often more extensively than indicated
m fig. 7. Crotchets of abdominal prolegs irregularly biordinal, in a
mesoseries, 26-28; anal leg crotchets similar, 24-26.

This is the only species of Ethmia known to lack secondary setae of the
abdominal SV groups, including both the prolegs and A9. Fully fed larvae
normally exceed the size range given.

Pupa (fig. 10). -“Length 10.3-13.0 mm (4n). Head without projections,
with a deep groove mesad of each antennal base, lined mesally and
anterolaterally with dense patches of golden setae (fig. 11). Antennae and
wings extended to posterior Fs of A5. A5 dorsally, A6-A7 dorsaUy and
ventraUy movable by deep intersegmental clefts. Dense, mst-orange,
setaceus areas around spiracles of A2-A6, on sides and posterior edge of
A5-A6, all exposed surfaces of A7-A9, and dorsum of AlO (fig. 12).
Spiracles small, of uniform size on A2-A8, closed on A8. Anal legs
represent by raised areas, not exceeding anterior edge of A9, densely
covered with hooked setae.

The setaceous grooves of the pupal head (fig. 11), which have not been
observed in any other ethmiid, are well developed in both sexes. Evidently
they are not homologues of the modified male antennae. The anchoring
mechanism restricted to A9, without free anal legs, is a unique feature
among described Ethmia. It resembles that of certain Stenomatinae
(Pracker, 1915), and it may represent the ancestral state for ethmiids.

In the laboratoiy cocoons were constructed in folds of paper toweling or
in lumens of corrugated cardboard. They were moderately dense, opaque,
about 18 mm in outside length and had a smooth interior cell about 14x4
ram, without the loose mesh known in some Ethmia. Normally Ehretia
acuminata is a rain forest tree with its canopy high above the ground. At
the Bunya Mountains, however, I found one tree in sparse natural woods
with penetrating sunlight, so that the lowest limbs could be examined. The
lower canopy contained many dead branches, most of which had hollow
centers, probably the result of beetle and aculeate Hymenoptera burrow-
ing. Splitting of a random sample of 20 branches 5-10 mm m diameter
produced 2 old cocoons of E. sphaerosticha (determined by larval exuvium
color and pattern of pupal shell setaceous areas). Both had appropriated
abandoned borings in 1 cm sticks.

Nearctic ethmiids are known to wander from hostplant foliage to seek
shelters or burrow into soft substances for pupation (Powell, 1971, 1973:
39). Considering the intemittently wet conditions of rain forest floor
litter, pupation above the forest floor would seem to be selectively
advantageous, and appropriation of abandoned holes in the canopy
branches may be the prefewed cocoon site for E. sphaerosticha and other
teopica! Ethmia.

The duration of pupation was relatively long, in contrast to the rapid
larval life. One male emerged 16 days foUowing initiation of cocoon

20(4): 214^234, 1981(83)

221

consteuction (80L47.1); another emerged 19 days after the larval collection,

Natiiral enemies.— Most of my rearinge were from eggs laid in the lab,
piecluding parasitoids, but among 6 larvae collected from Ehretia, 2 were
paiasitked by Braconidae and Tachinidae (81A107). An undersized
penultimate larva collected 6 January in an old shelter failed to feed, and 6
days later a braconid larva emerged and spun a cocoon, leaving a hole in
the mtersegmental area of abdominal segments 4”5. The Ethmia larva
remained active another day without feeding, before it was preserved. It is
interesting that larvae of E. arctostaphylella (Wlsm.) in California dis-
played this same behavior In response to p,arasitoid feeding and emergence
by Apanteles (Braconidae), which emerged from the side of the 3rd
abdominal segment (Powell, 1971: 46).

The tacHnid developed its piiparium within the pupal shell of E.
sphaerosticha and emerged 6 days later than a male Ethmia that had been
collected as a last instar larva at the same time. One of the two cocoons
discovered in hollow Ehretia twigs at the Bunya Mountains also had an
abandoned tachinid puparium inside the Ethmia pupal shell.

Voltinism— Collection records of adults suggest two or more generations
per season, with most records m November and Janiiaiy to March (Powell,
19cS2). FuD grown larvae were present at Mt. Keira in late November,
adults in late December, 1980, Final instar Fi larvae from the latter
(80M25) were preserved in mid-Januaiy^ and presumably would have
produced adults by early February. Ethmia sphaerosticha has been
collected at Mt. Keira as late as early April (1953 and 1964, V. J.
Robinson); thus, it is possible that 4 flights occur: late October, mid- to
late December, late January to mid -February and late March to early
April. Overwintering presumably occurs as a pupa in diapause, as is Imown
for multivoltine Nearctic species (Powell, 1971, 1974).

Colkction data.— Mt. Keira, Wollongong, N.S.W., larvae 25/26 Nov.
1980 (JAP80L47.1), females (MV light) 20 Dec. 1980 f80M25); Bunya
Mts., Q., females 5 Jan. 1981 (bl) (80A105), larvae 6/7 Jan. 1981
(81A107), female 7 Jan. 1981 (H) (80A113); Whkn Whian State Forest,
NE of Lismore, N. S. W., female (bl) 12 Jan. 1981 (81A121).

Ethmia postica (Zeller, 1877)

This is the most widespread ethmiid on the Australian continent,
occurring in interior areas from, northwestern and south central W. A. to
western Queensland, N. S, W., and Victoria (Powell, 1982).

Adult behamor.— The moths are nocturnal. Virtually all collections have
been made at lights, and females confined in the lab were inactive during
daylight hours. Eight females were caged in late September and October
1980, but dissections showed 5 of 6 salvaged to have been unmated. The
other was mated and deposited 90 fertile eggs (80J112.3) during a 12 day
period beginning 3 days following confinement. Water was supplied, and

222

J. Res. Lepid.

9

Fig. 9. Setalmap of Ethmia heliomela. Shading depicts heavily marked form.

potential foodplants had flowers and aphids rendering copious honeydew,
which may have been used by the females as nourishment. Unmated
females survived 6-21 days (x = 13.5, 7n) in captivity and the gravid
female 23 days, the first 4 in refrigeration.

Various plant samples were presented including Boraginaceae: native
Cynoglossum austmle R. Br., Ehretia acuminata) and exotic {Lithosper-
mum prostratum LoiseL, garden MyosoUs, Echium vulgare L.); Scro-
phulariaceae: (Hebe diosmifolia R. Cunn., Veronica formosa R. Br. and V.
calycina R. Br.); and Fabaceae {Westringia fruticosa Druce). None elicited
oviposition response except by the one female, which was offered
Lithospermum and Myosotis, to which Ehretia (drying) was added on day 2,
and Veronica on day 5. The first 6 eggs, deposited on day 3 or 4, were
placed on Myosotis and Lithospermum nested in plant hairs in the fashion
characteristic of many Nearctic Ethmia (Powell, 1971). Nearly all subse-
quent eggs were poked into the nylon mesh, a substrate also preferred by
captive ethmiids in California.

Only 19 eggs were produced up to day 11 of confinement at room

20(4): 214^234, 1981(83)

223

temperature (16 days following collection); 79% of the eggs were
deposited in days 12-18, 22 of them (24% of the total) during the last night
the female was alive.

E^^,““The eggs were cylindrical with rounded ends and had conspicuous
chorion ridges arranged in parallel rows, similar to those of Hokrctic
Etiimia (fig, 3). Those deposited in nylon mesh ranged 0.69 X 0.36 mm to
0.73 X 0.45 mm in length and width and wwe about as thick as wide,
varying in details of fonn with irregularities of their enmeshment. Eggs
were white with a pearly sheen when fresh, turning yellowish during
development, which required 10-11 days at Intermittently cool room
temperatures.

Larva.— Newly hatched larvae were offered various combinations of
foliage terminals of Myosotis, Lithospermum, Echium, Cymglossum, and
Veronica, during a 15 day period. Several fed for a few days on leaves of
Myosotis, where they were positioned against container sides, but
available plant material was deteriorating, and none of the larvae reached
the second instar. Buds and leaves of Lithospermum, Echium, and
Cynoglossum were densely hirsute and appeared to present insurmount-
able physical barriers to unfed larvae. Repeated attempts to elicit feeding
m open flowers or artificially opened buds, failed; this is the feeding site of
several Near ctic Ethmia that depend upon hirsute borages (Powell, 197 1).

Possible Hostplant Association.— Ethmia postka females were collected
near Queanbeyan, N. S. W., on the southern tablelands near the A.C.T.
border. Although LF.B. Common had never taken this species during 9
season sampling at the site, 18 individuals (11 d", 7 9), mostly in fresh
appearing condition, were collected during a 4 night sequence. As a result
it was assumed that some locally growing native or introduced Boraginaceae
must have been the source; Cynoglossum australe and Echium vuigare
were the most likely suspects according to the A.C.T. Flora. Later,
however, when I had summarized the distribution of E. postica (Powell,
1982) and compared records of Boraginaceae in the Herbarium Aus-
foaliense, Halgania cyanea Lindl. emerged as a possible hostplant. This
species, a low shrub of sand dunes and mallee scrub on sand, has a general
distribution comparable to that of E. postica but is not known to extend
east of the Great Dividing Range. Thus several peripheral southeastern
records for E. postica are outside the range of this plant. Because these
records are temporally sporadic, such as at Queanbeyan, at Black
Mountain, A.C.T. (6 moths in Sept.-Oct., 1962, 1965, 1968 during a 15
year light trapping survey), and in southern Victoria (one specimen in the
Gooding collection from Moe, where Gooding collected for many years), it
is possible that they originate from passive movement of individual moths
to cismoetane areas from the interior via storm front airstreams. The
likelihood of 18 individuals appearing at one MV light following such
movement seems low, yet the fact that 5 of 6 females were unmated lends

224

J. Res. Lepid.

credence to such a hypothesis. Normally female ethmiids are mated when
attracted to lights, but it is well known that migrations of noctuids involve
sexually immature adults (e.g. Common, 1954; Fox, 1978). Moreover, Fox
has documented migration of many Lepidoptera across the Tasman Sea
via storm front airstream movements to the southeast, particularly in
October. While most of his records are for larger, strong flying moths and
butterflies, the arctiid, Utethesia pulchelloides Hamps., seems to be weak
flying to the human observer and also feeds on low growing Boraginaceae
(McFarland, 1979). Utethesia was taken in numbers in New Zealand (9
specimens in one light trap, others at stations along a 400 km band of the
west coast) during a 2 night period following passage of a front. Back
trajectories calculated on meteorological data showed the likely origin of
such moths to have been coastal southern Queensland or northern New
South Wales 60 hours previously, a far greater distance than Halgania-
associated insects would need to migrate to reach the Canberra area.

Collection data. — 3 km NE Queanbeyan, N. S. W., 28 Sept, to 1 Oct.
1980 (MV light I.F.B. Common and J. Powell) (JAP80J112); same data 12
Oct. 1980 (80K124).

Ethmia hemadelpha (Lower, 1903)

A northern species, E. hemadelpha is widespread from west central
Western Australia to the coastal mountains of southeastern Queensland
(Powell, 1982). It was reared in 1962 by K. T. Richards at the Kimberley
Research Station, W. A., from Ehretia saligna. The foodplant record was
reported by Common (1970), at that time the only known host of an
Australian ethmiid.

Final instar larva. — Length 18.6-22.2 mm (2n). Head: HC width 1.57-
1.62 mm. Setation as in other Ethmia (e.g., Mackay, 1972). Adfrontal
sutures extending to cervical angle, adfrontal areas abruptly attenuated
posteriorly. Pale amber colored, mottled with rust except in stemmatal
area. Primary setal arrangements of thorax and abdomen as in fig. 8. SDl
directly dorsad and well separated from the spiracle on Al, anterodorsad
and approximate to spiracle on A2-8. SD2 lacking on abdomen. Sclerotized
depressions associated with spiracles or with thoracic leg setae lacking.
Secondary setae lacking except a cluster of 4-6 translucent, fine setae
anteroventrad of SV on A9 (apparently missing on one individual). D area
white; SD mostly rust colored (on preserved specimens) with round, white
spots. The raised pinacula darker; L area whitish below spiracle, which is
included in rust blotch of SD pinaculum on A2 -8 ; S V area lightly tinged with
rust, V pale. Crotchets of abdominal prolegs in a mesal “penellipse” (ca.
half circle), irregularly biordinal, 22-28; anal prolegs crotchets irregularly
biordinal, 24-26.

Pupa (fig. 13). — Length 10,2 mm (In). Head without projecting structure.
Antennae and wings extending to posterior margin of A4. A5-7 movable by

20(4): 214-234, 1981(83)

225

Figs. 10-13. Pupal characteristics of Australian Ethmia: 10, E. sphaerosticha
pupa, lateral (upper) and ventral (lower) aspects; 11, E. sphaerosticha,
lateroventral portion of frons (dorsum at top), showing detail of
antennal pit; 12, E. sphaerosticha, abdominal setation; 13, E. hema-
depha pupa, lateral (upper) and ventral (lower) aspects.

226

J. Res. Lepid.

deep intersegmental clefts and lateral condyles. Raised areas posterior to
spiracles on A3-A6 bearing dense patches of short setae which extend
below spiracles in slight depressions. Anal legs well developed, boot-
shaped, with dense clusters of hooked setae distally. AlO with a group of 6
blunt, nonhooked setae.

Collection data. — Kimberley Research Sta., W. A., 7 Oct. 1962, ex
Ehretia saligna (K. T. Richards); associated reared adults examined.

Ethmia nigroapicella (Saalmueller, 1880)

Originally described from Madagascar, nigroapicella is widespread in the
Indo Australian and Oriental Regions, from Madagascar to the Seychelles,
India, Burma and the islands of New Guinea, Philippines, Fiji, Samoa,
Ryukyu, Formosa, and it is well known (as E. colonella Walsingham) in
Hawaii, where it is presumed to be introduced. The species reaches the
northernmost extremities of Australia, on the Wessel Islands and Cobourg
Peninsula, N. T. and Sue Island in the Torres Strait, Queensland (Powell,
1982). The Australian specimens are smaller and display minor morpho-
logical differences from that reported elsewhere for nigroapicella, and
confirmation of the assignment of populations on the northern islands to
nigroapicella must await study of further material.

Foodplants. — A notable defoliator of Cordia subcordata Lam. and
occasionally feeding on C. sebestena L. in Hawaii (Swezey, 1944; Zim-
merman, 1978), nigroapicella is also recorded from Ehretia sp., deduced to
be E. dicksoni var.japonica Nakai, by Sattler (1967), and from E. buxifolia
Roxb. and Cordia in Japan (Moriuti, 1963). Fletcher (1933) recorded E.
laevis Roxb. in India, and Cordia subcordata was listed as the host in the
Seychelles (LeGrand, 1965). Among these Boraginaceae, Cordia subcordata
occurs in the Australian areas where nigroapicella is known.

Larva. — The last instar larva has been described from Hawaii by
MacKay (1972) and Zimmerman (1978). The full grown caterpillar is
brightly colored, black with light yellow DL lines and irregular light yellow
spots including the lateral row above the spiracles, and is characterized by
numerous secondary setae in the D, DL, L and SV groups, particularly on
the thorax, differing in this respect from the other Australian Ethmia
described here.

Pupa (figs. 14, 15). — General aspects described by Zimmerman (1978),
who included an outline drawing by MacKay of the ventral aspect. In order
to compare more subtle features of setation and spiracles with the other
species, the pupa is further characterized as follows:

Length 9.6-10.1 mm (2n). Head without projecting structures. Antennae
and wings extended to proximal one third of A5. A5-7 movable by deep
intersegmental clefts and lateral condyles. Spiracles of A3-A7 situated on
irregular raised ridges, followed and subtended by dense patches of
golden setae; A5-A7 sparsely setate on anterior half below the spiracular

20(4): 214-234, 1981(83)

227

patches, spiculate posteriorly (fig. 15). Anal legs well differentiated,
tapering, with distal margin slightly broadened, bearing dense clusters of
hooked setae. AlO without true cremaster setae.

Collection data.— Hawaii: Volcanos Natl. Park, Kamoamoa, 30 March

1981, r.f. Cordia subcordata (F. G. Howarth); associated reared adults
examined.

Ethmia thoraea Meyrick, 1910

A member of the Nigroapicella Group (Sattler, 1967; Powell, 1982),
thoraea is related to several tropical rain forest species to the north. Its
range, however, is broader than that of rain forests, from the north coast of
Queensland to coastal N.S.W. in the Hlawarra District, and inland at Cun-
namulla, Q., and Mt. Kaputar, N.S.W. There are discrete spring and fall
flights. Larvae of a related species, E. nigroapicella (Saal.) feed on Cordia
and Ehretia in India, Japan and Hawaii (Sattler, 1967).

Adult behavior. — This and related species are believed to be strictly
nocturnal, despite their brightly colored, aposematic-appearing hindwings
and abdomen. One female confined in a petri dish lived 12 days during
mid- January. She was fresh appearing and was suspected to be unmated
after producing no eggs during days 1-4 in the presence of Ehretia
(acuminata?) leaves from the Bunya Mts., Q. On day 5 a bouquet of
Cynoglossum australe was added. During the next 4 nights she produced
38 eggs, all laid singly on the Cynoglossum, none on Ehretia. Most were
deposited on unopened flower buds or in sepal cups containing young
nutlets (60%), although an appreciable number were placed on undersides
of leaves (29%); the remaining few were on leaf uppersides and stems.
Most were nested in crevices around buds, where they were sometimes
piled up, or among plant hairs, in the fashion characteristic of many
Nearctic Ethmia (Powell, 1971).

Egg.— Eggs (fig. 4) were similar in shape and surface sculpture to
Holarctic Ethmia, resembling those of E. postica in size, shape, and color.
Chorion characteristics indicate closer relationship to E. heptasema, with
a system of irregular cuneiform, rather than oval or rectangular segments.
Eggs ranged 0.63-0.75 X 0.33-0.41 mm in length and diameter, varying
with placement. Development was rapid, hatching occurring in fewer than
6 days.

Larva.— Newly emerged larvae failed to feed on buds, nutlets, or leaves
of 6-day old Cynoglossum australe and escaped from a faulty container
before other plant material could be provided.

Failure of Ehretia to elicit oviposition, together with the precise selection
of hirsute Cynoglossum buds suggests some Boraginaceae other than
Ehretia is the normal host plant, despite the fact that most collection
records are from rain forest localities.

Collection data.— Mt. Tamborine, Q., 10 Jan. 1981 (MV) (JAP 81 A1 14).

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J. Res. Lepid.

Ethmia heptasema (Turner, 1898)

This species differs markedly in genital characters from any other in the
Australian fauna. According to genital structures, it appears to be most
closely related to E. heliomela, a supposition enhanced by the oviposition
behavior of the two. E. heptasema occurs in rain forests of coastal
mountains from central Queensland to the Illawarra District, N. S. W., and
there are possibly conspecific specimens from New Guinea (Powell, 1982).

Adult hheavior.~The moths are nocturnal. Most collections have been
made at lights, and E. heptasema has the largest eyes of any Australian
Ethmia, with an eye index above 1.0, indicative of nocturnal behavior in
Nearctic species (Powell, 1973: 8). Four females were confined in 2 petri
dishes during a 15 day period in mid -January. Leaves of Ehretia
(acuminatal) from the Bunya Mts., Q. were provided. About 250 eggs were
produced, and females exhibited different preferences for oviposition
sites. In one dish one or both moths deposited 28 eggs, 26 of them in axils
of the raised underside leaf veins where they nested against small tangles
of plant hairs (fig. 5). In the second dish the females deposited 222 eggs,
mostly on leaf edges (57%) or in the damp cotton wick (36%). A few were
placed on glass sides of the container, none on leaf vein axils.

Although a few eggs were isolated singly in each dish, most were
deposited in small, irregularly assembled clusters of 3-9 eggs. Egg
clutches placed in leaf vein axils ranged 1-4 per site (x = 2.4, lln) (fig. 5);
those on leaf margins often were aggregated into larger clusters, often
composed of eggs of different ages, indicating retrn visits to the sites on
successive days. Isolated clutches of uniform age on leaf margins and
container walls consisted of 2-9 eggs (x ~ 4.5, 8n). Females displayed
selection preference for aggregation of eggs once a cluster was started,
with most of the available leaf margins remaining unused. As many as 10-
13 eggs may have been deposited at once in the larger batches, judging
from age groups.

—Individual eggs were cylindrical, resembling typical Holarctic
Ethmia eggs in shape but not chorion sculpture, which was an intricate
system of cuneiform segments (fig. 6). Those in clusters varied in form with
placement. Eggs of E. heptasema were unusually small; isolated eggs
ranged 0.57-0.62 X 0.35-0.38 mm in length and diameter. Differing stages
of development were easily detected because the white eggs turned yellow
within 24 hrs, then reddish orange by 48-72 hrs and began to show
mandibular spots within 110 hrs. Hatching occurred in fewer than 6 days
despite intermittent transport in a field ice box.

Lorua.— Newly hatched larvae were provided with mature Ehretia leaves
which were beginning to deteriorate after storage 5-8 days without
consistent refrigeration. The final larvae to emerge were placed on fresh
Cynoglossum australe, but no feeding took place on either plant.

Nonetheless, the geographical distribution of Ethmia heptasema, the

20(4): 214-234, 1981(83)

229

strong oviposition response to Ehretia, and the precise selection of sites on
the leaves, suggests that Ehretia acuminata is the normal hostplant.
Possibly mature leaves pose physical barriers to feeding by first instar
larvae, and buds or new leaves are requisite to larval establishment.

Collection data.— Mi. Tamborine, Q., 4 Jan. 1981 (MV, bl) (JAP
81A103).

Ethmia heliomela Lower, 1923

Ethmia heliomela is a small, brightly colored moth unlike any other
Australian ethmiid. By structural features it appears to be related to E.
lapidella (Walsingham) of India and the Orient, a species that superficially
resembles E. heptasema. Ethmia lapidella has been reported to feed on
Ehretia dicksoni var japonica Nakai and “soongroo (wild Salvia)''
(Fabaceae?) (Fletcher, 1920: 133; Sattler, 1967). The latter record may
have been based on a misidentified plant or a wandering fuU grown larva.
E. heliomela is known only from eastern montane and coastal rain forests,
from southern Queensland to southern N. S. W.

Adult behavior.— The orange and black hindwings and abdomen* and
relatively small eyes (eye index 0.90-0.95), give this species the appear-
ance of a diurnal moth. The adults commonly fly to lights, however, so it is
presumed to be primarily nocturnal. I found adults perched on Ehretia
foliage in late afternoon and early morning. Under laboratory conditions,
activity consistently took place during afternoon hours, but neither mating
nor oviposition was observed. A freshly emerged male and female were
caged in mid-November, but they survived only two days in unseasonably
warm weather. Dissection showed the female unmated (80L38). Five
females collected from foliage of Ehretia acuminata were individually
confined in 42 X 60 mm plastic vials with branchlets of Ehretia
inflorescences in bud to early bloom. These moths survived only 2-5 days.
Two females deposited 11 and 23 eggs, either singly (62%) or in pairs or
clusters of 3 or 4; usually laid side by side. All were placed on walls of the
containers.

Egg.— The eggs resembled those of E. heptasema (fig. 6) in shape and
chorion sculpture viewed at 50X magnification. They measured 0.61-0.65
X 0.35 mm at the rectangulate-oval base of attachment and were ca. 0.25-
0.30 mm thick. During development the eggs turned pale yellow within 12
hrs, then pink within 48 hrs. Hatching occurred in ca. 10 days.

Larva. — No attempt was made to feed newly hatched larvae, but
caterpillars of the penultimate and at least 2 preceding instars were found
in conspicuous webs in inflorescences of Ehretia."' Quite small larvae
produced copious silk networks encompassing branchlets several cm
apart. Sometimes 2 larvae occupied one web, but they were not colonial,
living individually except where abundant. At Mt. Keira only in the final 2
instars were Imwae found to feed on leaves, usually by extending webbing

230

J. Res. LepM.

from wilted inflorescences onto subtending leaves where they skeletonized
the upper surface. Timing of oviposition seemed to be related to that of
flowering; young larvae were collected November 26 from trees in bud to
early bloom, but another tree with buds just starting to develop had none.
Three weeks later David Walsh observed webbing and larvae on the latter
tree and on Dec, 221 found larvae of the final 3 instars on its inflorescences
in late bloom. In the laboratory late instar larvae fed on leaves after
inflorescence material had been consumed.

Final instar /arua.—Length 15.8-16.8 mm (5n), Head: HC width 1.22-
1.26 mm. Setation as typical for genus (e.g. MacKay, 1972). Adfrontal
sutures reaching cervical angle, adfrontal areas abruptly narrowed poster-
iorly, Black to dark amber brown, conspicuously contrasting pale bands
flanking adfrontals and longitudinally on venter, beneath stemmatal area.
Primary setal arrangements of thorax and abdomen as in fig. 9. Prothorax
with strongly delineated shield, black narrowly edged with amber to
mostly amber blotched with blackish; 5 secondary setae along its anterior
margin on each side; D1 and D2 widely separated. Sclerotized depressions
associated with spiracles or with prothoracic legs absent. L2 tiny on Al-8.
Secondary setae absent except a row of 6 or 7 anterior to SV on A9. Color
pattern variable, mostly dark olive green (appearing black on living
caterpillars), as in fig. 9, with pale bands narrowly at mid-dorsum and of
irregular spots longitudinally in SD, L areas (tan or pale greenish in life); a
separate color phase has the dark markings restricted, lacking from
posterior V2 or Va of each segment so that the larva has segmental,
transverse pale bands. Pinacula sclerotized, dark brown, darker above
spiracles. Crochets of abdominal prolegs uniordinal but varying in size, 30-
34 in a circle, of anal prolegs, 34.

The larval color phases appeared to be discrete, without a gradation of
intermediates, in the last instar, but they could not be separated in early
instars. Penultimate and antepenultimate instars were similarly patterned
to the fgl but were paler. The banded form, segregated as a sublot
(80M30.1), produced normal E. heliomela adults. One larva, perhaps
teneral, was observed that appeared orange with pale cream longitudinal
bands to the unaided eye. Ittoo was separated (80M30.2) and produced^.
heliomela.

Evidently this species is capable of developing dense population levels,
during which larval feeding is not restricted to inflorescences. Larval
webbing encompassed an entire Ehretia tree near Jamberoo, N. S. W. in
May 1980 according to D. S. Stevens {in litt. to I. F. B. Common). His
photographs show strings of larvae festooning the defoliated tree. It is
possible that other Lepidoptera were involved, but only E. heliomela was
reared, from cocoons in the bark of a silk-covered piece of a large branch.

I did not discover a pupation site in the field but in or under loose bark
may be normal, hi the laboratory cocoons were plastered onto paper

20(4): 214-234, 1981(83)

231

Figs. 14-16. Pupal characters of Australian Ethmia: 14, E. nigroapicella pupa,
lateral (upper) and ventral (lower) aspects; 15, E. nigroapicella,
detail of abdominal sculpture and setation; 16, E. heliomela pupa,
lateral (upper) and ventral (lower) aspects.

232

J. Res. Lepid.

towelling incorporating debris, on paper under flat cardboard or in lumens
of corrugated cardboard, and once in a curled dry leaf.

Pupa (fig. 16). — Length 6.8-7. 6 mm (5n). Head truncate, square-
margined ventrally. Antennae and wings extending beyond A4, nearly
across A5. Intersegmental areas preceding A6 and A7 deeply cleft dorsally
and ventrally, movable by lateral condyles. No setaceous areas. Spiracles
of A5, A6 huge, ca. 2X diameter of those on A2-4 and 7; A8 sp absent. Anal
legs well developed, variable in shape, separation, and angle of projection;
bearig a series of short hooked spurs distally. AlO without setae
representing the true cremaster.

The enlarged spiracles of segments 5-6 and loss of spiracle on 8 are
unique among described Ethmiidae and perhaps all Lepidoptera. Accord-
ing to Mosher (1916: 29) spiracles are always present on abdominal
segments 1-8 in Lepidoptera, although those of the 8th are never
functional and show no distinct opening.

Under laboratory conditions development time of pupae varied. Al-
though feeding ceased within 10 days, emergences occurred 18-30 days
following collection of larvae in both November and December, and
continued sporadically up to 80 days. Material collected at Jamberoo in
May produced adults in August and again in November, indicating that
diapause occurs in the pupa and termination is facultative in response to
environmental cues, as is known in Holarctic Ethmia (Powell, 1973: 40;
1974).

Voltinism. — Collections of adults indicate a flight period from October to
February, but 80% of the records are in spring, primarily November and
early December. In New South Wales, only 3 (12%) of the collections were
made in January and February, suggesting that a summer generation is
facultative and relatively rare, as indicated by sporadic emergences in
laboratory conditions. At Mt. Keira, in 1980 I collected both adults and
larvae in late November and again 4 weeks later but could find neither in
early February. Thus it is likely that E. heliomela normally undergoes two
generations in spring and early summer and occasionally emerges later,
particularly if Ehretia blooms sporadically and the moth’s ovipositon is
cued to the same factors that influence inflorescence development.

Natural enemies. —Although many larvae were collected, most of those
not preserved became diseased prior to maturing, probably the result of
suboptimal conditions in the lab. Among 17 individuals reared to maturity,
only one was parasitized. A tachinid emerged within 23 days of the larval
collection after pupation inside a heliomela cocoon, before pupation of the
host larva.

Collection data. — Jamberoo, N. S. W. May 1980 (D. S. Stevens), adults
emgd. Nov. 1980 (JAP80L38); Mt. Keira, Wollongong, N. S. W., 25/26
Nov. 1980, adults (80L46), larvae (80L47) on Ehretia acuminata', same
data 22 Dec. 1980 (80M30).

20(4): 214-234, 1981(83)

233

Acknowledgments. In addition to helping with some of the collections, I F. B.
Common provided much of the background knowledge necessary for the study, as
well as hospitality and facilities during my visit. Grateful thanks are also extended to
D. F. Waterhouse, former Chief, and to other staff members of the Division of
Entomology for assistance with transportation and facilities, making possible the
fieldwork and other aspects of the study. SEM photos of the eggs were made by
Barry Filshie, of eggshells by J. A. DeBenedictis, U. C. Berkeley. Carolyn Millinex
Tibbetts, U. C. Berkeley, made the drawings of pupal characters and inked my
drafts of setal maps of the larvae.

Assembly of specimen data and mapping of distributions were carried out by
Elizabeth Randal. Cooperation by Lesley Lockwood and authorities of the National
Botanic Garden and Herbarium Australiense, CSIRO enabled use of living plant
samples and of collections in their care.

David Walsh, caretaker of the Mt. Keira Boy Scout Camp, Wollongong, N. S. W.,
provided facilities and botanic expertise making possible several collections that
were critical to the study. Discoveries of ethmiids at Mt. Keira and elsewhere by
Victor J. Robinson provided much of the background information for my taxonomic
and biological work.

K T. Richards, Dept, of Agriculture, Perth, W. A., loaned larval and pupal
specimens of Ethmia hemadelphn, and F. G. Howarth and G. M. Nishida, Dept.
Entomology, Bernice P. Bishop Museum, Honolulu, provided pupal shells of E.
nigroapicella, enabling inclusion of their descriptions.

Literature Cited

COMMON,!. F. B., 1954. A study of the ecology of the adult bogong moth, Agrotis infusa
(Boisd.) (Lepidoptera: Noctuidae), with special reference to its behaviour
during migration and aestivation. Austral. J. ZooL, 2:223-263.

, 1970. Lepidoptera (Moths and Butterflies) in Insects of Australia; pp.
765-866. Melbourne; Melbourne U. Press.

DIAKONOFF, A., 1967. Microlepidoptera of the Philippine Islands. U. S. Natl. Mus.,
BuU. 257; 284 pp.

MTCHER, T. B., 1933. Life histories of Indian Microlepidoptera (Second Series).
Cosmopterygidae to Neopseustidae. Imp. Counc. Agric. Research, Sci. Monogr.,
No. 4; 85 pp.

POX, K. J., 1978. The transoceanic migration of Lepidoptera to New Zealand — A
history and a hypothesis on colonization. N. Z. EntomoL, 6:368-379.

HODGES, R. w., 1978. Gelechioidea. Cosmopterigidae. in: Moths N. A. North of
Mexico; fasc. 6.1; 166 pp. + 6 pi. E. W. Classey and Wedge EntomoL Research
Found.; London.

KUZNETSOV, V. I. & A. A. STEKOL'NIKOV, 1979. The systematic position and phylo-
genetic relationships of the superfamily Coleophoroidea (Lepidoptera:
Oecophoridae, Coleophoridae, Ethmiidae) as revealed by the functional
morphology of the male genitalia. EntomoL Review, 57(1):91-103.

LeORAND, H., 1965. Lepidopteres des lies Seychelles et de’ Aldabra. Mem. Mus. Nat.

Hist. NatL, Paris Ser. A. (ZooL), 37:1-210.

MACKAY, M. R., 1972. Larval sketches of some microlepidoptera, chiefly North
American. EntomoL Soc. Canad., Mem. 88; 83 pp. 1978 in Zimmerman, bisects

234

J. Res. Lepid.

of Hawaii. VoL 9(2) Microlepidoptera; fig. 643. U. Press Hawaii; Honolulu.

McFarland, n., 1979. Annotated list of larval foodplant records for 280 species of
Australian moths. J. Lepid. Soc., 33, suppl; 72 pp.

MORlun, S., 1963. Ethmiidae from the Amami-Gunto Islands, Southern Frontier
of Japan, collected by Mr. T. Kodama in 1960. Butterfl. Moths, 14:35-39.

MOSHER, E., 1916. A classification of the Lepidoptera based on characters of the

' pupa. Bull. m. St. Lab. Nat. Hist., 12:17-159, pi. 19-27.

POWELL, J. A., 1971. Biological studies on moths of the genus Ethmia in California
(Gelechioidea). J. Lepid. Soc., 25, Suppl. 3; 67 pp.

, 1973. A systematic monograph of New World ethmiid moths (Lepidop-
tera: Gelechioidea). Smithson. Contr. ZooL, 120; 302 pp.

, 1974. Occmrence of prolonged diapause in ethmiid moths (Lepidop-
tera: Gelechioidea). Pan-Pacific EntomoL, 50:220-225.

, 1980. Evolution of larval food preferences in microlepidoptera. Ann.

Rev. EntomoL, 25:133-159.

, 1982. Taxonomy and geographical relationships of Australian ethmiid

moths (Lepidoptera: Gelechioidea). in review for Austral. J. Zool.

SATTLER,K., 1967. Ethmiidae. Microlep. Palaearct., 2; Textband 185 pp.; Tafelband,
106 plates. Verlag. G. Fromme & Co., Vienna.

SWEZEY, 0. H., 1944. The Kou moth, Ethmia colonella Walsm., in Hawaii, Proc.
Hawaiian Ent. Soc., 12(1943):133-135.

ZIMMERMAN, E. c., 1978. Insects of Hawaii. Vol. 9, 1936 pp. U. Hawaii Press,
Honolulu.

Journal of Research on the Lepidoptera

20(4): 235-239, 1981(83)

Field Study of Phyciodes batesii (Reakirt) and P. thaws
(Drury) from a Site in the Black Hills, South Dakota
(Lepidoptera: Nymphalidae: Melitaeinae)^

Clifford D. Ferris^

Bioengineering Program, University of Wyoming, Laramie, Wyoming 82071

Abstract. Maculation and male genitalic structure are discussed for two
sympatric populations of Phyciodes at a study site in the Black Hills, South
Dakota. Evidence suggests naturally occurring hybrids.

t ft

Introduction

In a recent publication (Fbrris & Brown, 1981), Ferris alluded to a
problem in identifying certain species of Phyciodes from the Black Hills,
South Dakota. Data presented by Oliver (1979, 1980) concerning his
studies of Phyciodes batesii (Reakirt) and P. tharos (Drury) now make
possible relatively easy separation of these two species.

To summarize Oliver’s findings (1980), two separate forms of P. tharos
exist, which he has designated as types A and B. Type A is mutivoltine;
type B is univoltme. According to Oliver, in the western United States,
type B, generally designated as P. tharos pascoensis Wright, is found above
8000' (2440 m), while type A is found in moist canyons below 7000' (2135
m). He reported that both types are sympatric in the Black Hills.

Oliver has also indicated {in litt.) that the ventral color of the antennal
clubs can, be used in some instances to separate tharos types A and B.
Where applicable, the color in A is black or dark gray, while it is orange or
yellowish in B. In northern plains populations, however, light-colored
antennal clubs are a fi^ed^ character. Here types A and B are best
separated by voltinism andthe size and extent of the dorsal dark markings
in the males. Type B specimens manifest broad dark wing borders.

My studies were conducted in a side canyon perpendicular to Little
Spearfish Creek Canyon (shown on some maps as Tinton Creek Canyon),
just behind the Timon Campground, Custer National Forest, Lawrence
Co., South Dakota, 5700' (1740 m). Phyciodes populations in this area

‘Published with the approval of the Director, Wyoming Agricultural Experiment Station as
Journal Article No. JA 1176. , .

^Research Associate: Allyn Museum of Entomology /Florida State Museum, Sarasota, Florida;
Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of
Agriculture and Consumer Services, Gainesville, Florida; Dep^tment of Entomology, Los
Angeles County Museum of Natural History, Los Angeles, California.

236

J. Res. Lepid.

F%. 1. P. batesU (male) from Black Hills study site, 22 July 1973, dorsal left,
ventral right.

Fig. 2. P. batesU (female) same data, dorsal left, ventral right.

Fig. 3. P. tharostype B (male) from Black Hills study site, 22 July 1973, dorsal left,
ventral right.

Fig. 4. P. thaws type B (female) same data, dorsal left, ventral right.

Fig. 5. Melanie females of P. batesU (left) and P. thaws type B (right) from Black
Hills study site; thaws collected on 22 July 1973; batesU collected on 27
July 1977.

20(4): 235-239, 1981(83)

237

Fig. 6. Possible hybrid male P, batesii X thaws type B from Black Hills study site,
22 July 1973, dorsal left, ventral right.

were sampled in 1969, 1970, 1973 and 1977.

At this study site, I have found only thaws type B and batesii on the wing
during the month of July, the time of year I have visited this area. Based
upon my field experience, thaws type B appears to predominate in the
Rocky Mountain region, from northern New Mexico northward into
Alberta and British Columbia.

The antennae of P. batesii have ventrally black tips. This permits rapid
separation of the two species when batesii is sympatric with type B thaws.
Males of the two species are relatively easily separated based upon dorsal
maculation. P. batesii is generally much darker overall than thaws, with
heavy black scaling. It also has a two-tone appearance owing to the light
color of the postdiscal band (indicated by the pointer in Figs. 1 & 2). The
females are not so easily separated based upon maculation, particularly
when they are somewhat melanic, as shown in Fig. 5. The females of both
species have a two-tone aspect dorsally, which in thaws sometimes
approximates P. pratensis (Behr). Generally, batesii females have a bright
aspect with sharply defined maculation, while thaws tends to have a
blurred appearance.

As noted by Oliver (1979), thaws and batesii have achieved similar
phenotypes, but through different genetic means, as his hybridizational
studies have shown. Natural hybrids are apparently rare. Laboratory
studies indicate strong behavioral isolating mechanisms during courtship.

Five field-collected specimens, three males and two females, from the
Black Hills exhibit hybrid characteristics. All five have type B thaws
antennae (orange tip color ventrally). The three males resemble batesii
dorsally. Ventrally one resembles thaws; the two others batesii. These
latter two bear a fairly close resemblance to the hybrid specimen
illustrated by Oliver (1979, Fig. Ic). One of these specimens is shown in
Fig. 6. Black Hills batesii differ significantly in phenotypic appearance
from the New York state population used by Oliver in his breeding studies
(pp. cit). Consequently, one would expect possible batesii X thaws

238

J. Res. Lepid.

hybrids from the Black Hills to differ somewhat in appearance from
crosses between individuals from eastern populations.

The two females dorsally are intermediate with regard to the Black Hills
populations studied. Ventrally they are pale, and not nearly so heavily
marked, as is typical thaws. Four of the specimens show no structural
abnormalities of the abdomen as described by Oliver for laboratory
hybrids. The remaining specimen, the first male described above, has a
very contorted abdomen, but under the microscope it cannot be ascer-
tained clearly if this results from a structural defect, or is simply an artifact
of field handling and subsequent desiccation.

Genitalic Studies

There are differences in the male genitalia between thaws and batesii.
The key structures are the hom-like processes extending from the
posterior of the last tegumen (uncus). Fig. 7 A depicts typical type B thaws
from the Black Hills. Note the squarish aspect of the overall structure. Fig.
7B illustrates batesii from the Black Hills. The overall structure here
presents a rounded aspect with a more acute angle at the base of the
processes than occurs in type B thaws.

The three possibly hybrid males noted above have genitalia of the form
shown in Fig. 7C.‘This structure is intermediate between those shown in
Figs. 7 A, B.

Fig. 7 . Diagnostic portion of male genitalia of P. tharos type B and P. batesii A. P.
tharos type B. B. P. batesii. C. Apparent P. tharos X batesii. Genitalia
from specimen shown in Fig. 6.

Results and Conclusions

On the basis of genitalic characters, it appears that the three males
described above represent natural hybrids batesii X tharos type B. The
two females remain enigmatic since the genital structures in that sex are
not diagnostic.

The apparent hybrid specimens were collected accidentally and at
random. They were not discovered until recently when the author revised

20(4): 235-239, 1981(83)

239

the arrangement of thaws and batesii in his collection based upon Oliver’s
studies.

Analysis of collecting records indicates that 90 males and 27 females of
thaws, and 9 males and 6 females of batesii, with the additional five
apparent hybrids were collected in the study region. Of this total number,
44 males and 16 females of thaws type B, and 8 males and 5 females of
batesii were collected on 22 July 1973. Type B thaws is clearly the
predominant Phyciodes at the study site, and perhaps accounts for the
apparent hybrids. Not all Phyciodes observed, however, were collected.
Hence the true ratio of thaws to batesii may not be represented by the
numbers given.

The study site reported in this paper is on the eastern slope of the Black
Hills. In July 1982, type B thaws and batesii were taken sympatrically on
the western slope in the Black Hills National Forest, Crook Co., Wyoming.

Also in July 1982, these two species were found flying together in
Monroe Canyon, 4400’ (1340 m), Sioux Co., Nebraska (Pine Ridge
region) . Males of both species were taken at mud along the banks of a small
stream. To date, no natural hybrids have been found in this area, but only
limited numbers of the two species have been collected.

Acknowledgments. I would like to thank Dr. Charles G. Oliver for reading and
commenting on two preliminary drafts of this paper, and for providing information
on the genitalic structures in Phyciodes. Dr. Robert J. Lavigne of the University of
Wyoming provided some of the specimens from the western slope of the Black Hills.
An anonymous reviewer also commented on the manuscript.

Literature Cited

FERRIS, C. D. & F. M. BROWN, 1981. Butterflies of the Rocky Mountain States. Univer-
sity of Oklahoma Press, Norman, Okla. 442 pp. .

OLIVER, c. G., 1979. Experimental hybridization between Phyciodes thaws and P.
batesii (Nymphalidae). J. Lepid. Soc. 33(l):6-20.

, 1980. Phenotypic differentiation and hybrid breakdown within Phyciodes

'‘thaws” (Lepidoptera: Nymphalidae) in the northeastern United States. Ann.
Ent. Soc. Amer. 73(6):715-721.

Journal of Research on the Lepidoptera

20(4): 240-244, 1981(83)

Notes

An Aberration of Glaucopsyche lygdamus (Lycaenidae) with a
Complete Scolitantidine Dorsal Pattern

On 8 April 1982 I took a female Glaucopsyche lygdamus incognitus Tilden at
Rancho Cordova, Sacramento Co., California, displaying an extensive dorsal
pattern of black spotting on a blue ground, reminiscent of the normal pattern of
several Palaearctic genera such as Maculinea. Females of this population are
normally black with more or less blue basally, and the aberration (Fig. 1) is much
more extensively blue than is commonly seen. Ventrally, the postmedian spot-band
is enlarged and slightly distorted and the ground-color somewhat darker than
average. No abnormalities were observed in a sample of about 20 animals examined
and released at the site.

The resemblance to Maculinea and other Scolitantidines is extremely suggestive
of homology. Mattoni (1981, The Scolitantidini. 11. The world’s smallest butterfly?
Notes on Turanana, and a new genus and species from Afghanistan (Lycaenidae). J.
Res. Lepid. 18:256-264) gives the occurrence of the dorsal pattern in several taxa of
the genera Micropsyche, Turanana, and Glaucopsyche. According to Mattoni {in
Utt.) it occurs in some Turanana, Micropsyche, Phengaris, Shijimiaoides, and
Philotes (sonorensis); in Shijimiaoides it is limited to females, as in Maculinea alcon.
The genus Caerulea, which seems more closely related to Glaucopsyche than is
Maculinea according to Mattoni, lacks the pattern (but cf. Higgins, 1975, The
Classification of European Butterflies. Collins, London. 320 pp. [pp. 131-133]).
Whatever the proper relationships within this group, the genetic control of the
pattern is apparently labile, with more than one reversal or parallelism necessary in
their history.

A similar aberration in female Glaucopsyche melanops has been reported by
Chapman (1905, Trans. Ent. Soc. Lond. 53:ii-iii), who also noted the similarity to
Maculinea. G. melanops is quite a distinct species of Glaucopsyche, and morphologi-
cally stands apart from the other six or so species in that genus. Chapman named the
aberration var. “wheeleri,” on the basis of two specimens from Digne.

Arthur M. Shapiro, Dept of Zoology, University of California, Davis, CA 95616

A Reared Gynandromorph of Tatochila (Pieridae)

On 2 April 1981 a mosaic gjmandromorph emerged in a laboratory culture of the
Tatochila sterodice Stgr. species-group, derived from Argentina (Pieridae). The
specimen (Fig. 2) is an Fi hybrid of T. s. sterodice male, ex San Martin de los Andes,
Province of Neuquen, and T. vanvotcemii Capr. female, ex Bahia Blanca, province of
Buenos Aires. It was reared without diapause under 10 hours light, 14 dark, day and
night temperatures 75°F^23.9®C and 55°F=^12.8°C respectively.

20(4): 240-244, 1981(83)

241

Fig. 1. Aberration of Glaucopsyche
lygdamus.

Fig. 2. Reared gynandromorph of
Tatochila sterodice x van-
volxemii hybrid, upper and
under surface. See text.
NOTE: Fig. 2 (only) is re-
versed due to a printing
error.

Fig. 3. Reared homoeotic male
Pieris rapae, stock ex
Xochimilco, Mexico. Note
FW tissue on one ventral
HW (see text).

Fig. 4. Wild “intersexual” Colias
eurytheme, upper and under
surfaces; note “feminiza-
tion” of dorsal FW pattern.

242

J. Res. Lepid.

The body and three wings are completely male. The left fore wing is mosaic above
and entirely male below. Its upper surface is about 85% female; the male area
occupies the costal margin down to vein Mi, with a mixture of male and female
scales in interspace M1-M2 marginad of the dark submarginal band. The outer
margin is slightly convex on the mosaic wing, a female character. The shape of the
submarginal band on the female surface is slightly abnormal.

This is the only sexual mosaic produced in a complex hybrid culture of some 7000
butterflies reared over 18 months. Another F 1 brood produced a bilateral male-male
mosaic, presumably resulting from a doubly -fertilized binucleate egg (Shapiro,
1982, Experimental studies of the evolution of seasonal polyphenism. In R. Vane-
Wright and P. Ackery, eds.. The Biology of Butterflies, Symp. Royal Ent. Soc. of
London, in press).

Sibatani’s (1980, Wing homoeosis in Lepidoptera: a survey. Devel. Biol. 79:1-18)
analysis of 44 published gynandromorphs shows that sexual mosaicism is com-
moner on the fore- than the hindwings and on the dorsal than the ventral surface;
there is no right-left difference. Of 19 Pierid gynandromorphs, 13 were confined to
the dorsal surface, 1 to the ventral, 5 mixed. Thus, this individual is of the
statistically commonest type of mosaic gynandromorph. It is very close to a % male-
Vi female individual, but the presence of male tissue on the aberrant wing suggests
the chromosome accident occurred early in the history of the wing primordium but
subsequent to the second zygotic division, which would give a neat Vi picture.

Arthur M. Shapiro, Dept, of Zoology, University of California, Davis, CA 95616

Two Homoeotic Pieris rapae of Mexican Origin (Pieridae)

“Homeosis” was defined by Pennak (1964, Collegiate Dictionary of Zoology.
Ronald Press, New York. 583 pp.) as “transformation of one organ into another by
mutation. , .occurrence of an homologous appendage on a segment where it does not
normally occur.” In Lepidoptera, transformations of entire organs are virtually
unknown, and the term, now usually spelled “homoeosis,” refers to the patchy
conversion of areas on the wings into areas appropriately placed on a different wing
or on the other wing surface. Recognition of this condition depends on pattern
differences which render the inappropriately -placed areas conspicuous. It seems to
be very rare; most recognized examples have probably been published, and a
majority figured. Sibatani (1980, Wing homoeosis in Lepidoptera: a survey. Devel.
Biol. 79:1-18) reviewed 164 cases and presented statistics on the topology of the
transformations.

A homoeotic male Pieris rapae L. (Pieridae) eclosed 19 June 1982 in the first
generation of a laboratory stock established 5 March 1982 from several females
collected at Xochimilco, Distrito Federal, Mexico, the southernmost known
population of P. rapae in the Americas, for photoperiodism studies. The aberration
was reared under 10L:14D with day and night temperatures of 70°F=21.1°C and
40°F=4.4°C, and did not diapause. It is shown in Fig. 3. It is similar to a homoeotic
male P. brassicae L. figured by Gardiner (1963, Genetic and environmental
variation in Pieris brassicae. J. Res, Lepid. 2:127-136 [p. 134]) in having part of one
ventral hindwing converted into ventral forewing. Because of the rearing regime, the

20(4): 240^244, 1981(83)

243

ventral ground color is bright yellow and the white converted area is easy to delimit.
It contains both of the black ventral forewing discal spots in their proper
interspaces. The entire homoeotic area is posterior to Sibatani’sMi-Ma barrier. All
other wings and surfaces are apparently normal.

On 17 August 1982 a homoeotic female emerged in the same culture. This
individual, shown alive in Fig. 5 (black and white) as it was to be bred, was reared
under 10L:14Dwithday and night temperatures of 75°F=23.9°C and 55°F™12.8°C,
and diapaused. It was from the same generation as the first; as noted, the culture
was started by pooling the ova of several wild females, but it seems very likely that
the two homoeotic individuals are full sibs. The converted area is nearly identical to
that on the male, and like it the abnormality is confined to the ventral left hindwing.
To date (early September 1982) no further aberrations have appeared in a total of
some 570 animals examined from this culture, reared on several regimes.

Gardiner {loc. cit.) reported a cluster of three homoeotics deriving from a single
mating involving hybridization of typical brassicae with subspecies cheiranthi Hbn.
from the Canary Islands. All were males. One was sib mated and the line inbred for
three generations, but no additional homoeotics appeared in a total of some 900
progeny. As our Mexican rapae line is to be continued for some time and diapausing
members of the first generation are still in cold storage, every effort will be made to
investigate the presumptive genetic predisposition to homoeosis. The extraordinary

Fig. 5. Reared homoeotic female P. rapae from Xochimilco, photographed alive by
S. W. Woo.

244

J. Res. Lepid.

rarity of two homoeotics appearing in the same brood is underscored by their being
the first and second homoeotic butterflies I have observed in over 20 years of
collecting and breeding, including the rearing of perhaps a quarter of a million
Pieridae for physiological and ecological studies.

Arthur M. Shapiro, Dept of Zoology, University of California, Davis, CA 95616

An Apparent “Intersexual^* C olios eurytheme (Pieridae)

In most Nearctic Colias species, the female forewing has a broad dark marginal
band enclosing spots of the ground-color while the band on the male is continuous,
narrow, and interrupted only by ground-color along the veins, if at all. This
difference does not hold in some Palearctic species, in the circumpolar C. nastes
Bdv., and in some South American species.

A C. eurytheme Bdv. showing a partial “feminization” of a male pattern was taken
18 February 1982 at Rancho Cordova, Sacramento Co., California. The specimen
(Fig. 4) is genitalically male. The dorsal forewings have a double dark band
enclosing the light ground color, in this regard resembling late winter-early spring
females. Ventrally there are non-sexual anomalies: the forewing discal spots are
divided or doubled like those on the hindwing, and there is a rudimentary accessory
discal spot basad of the actual, doubled one on the right hindwing. The shape of the
forewings is abnormal, and the flight of the animal was very low and direct, just
above the ground.

Well-developed male examples of the winter-spring form “ariadne” of C.
eurytheme occasionally show a very slight development of the inner dark band just
below the dorsal forewing apex, enclosing one small “spot” of ground-color anterior
to Rs + 4. The Rancho Cordova specimen seems to represent an extreme manifesta-
tion of this tendency in which the developmental control mechanism which normally
inhibits the inner band in males failed to do so. This may or may not be attributable
to abnormalities in the sex-determination mechanism. Although the relationship
between the male and female patterns in Colias could be interpreted in two ways,
the general rule that reduction is “easier” than complexification in evolution sug-
gests that the normal eurytheme male pattern is a reduction from the female, and
thus evolutionarily derivative. The present aberration would then be considered
reversional, or atavistic. If the Andean Colias (Colias) are monophyletic, either at
least one reversal or one convergence or parallelism for this character has occurred,
since the species flaveola Butl. and weberbaueri Stgr. are monomorphic or nearly so,
the others sexually dimorphic for the forewing border.

Arthur M. Shapiro, Dept of Zoology, University of California, Davis, CA 95616

Journal of Research on the Lepidoptera

20(4): 245=248, 1981(83)

Two New California Catocala Subspecies (Noctuidae)

John W. Johnson

Assistant Research Biologist, Museum of Systematic Biology, University of California,
Irvine, CA 92717

Abstract. Catocala andromache subsp. wellsi Johnson, with nearly black
body and forewings, from Jackson, Amador Co,, CA, and Catocala mcdun-
nought subsp. browerarum Johnson, with dark gray-brown body and fore-
wings, the wings suffused with green scales, from Moore Creek Campground,
Calaveras Co., CA, both localities in the northern Sierra Nevada Range,
collected by R. E. WeDs, are described, the types being deposited in the A.
E. Brower collection, Augusta, Maine.

Catocala andromache (Hy. Edwards 1885) was described from a
specimen from “near San Bernardino/' California. Southern California
specimens from Kem River Canyon in the southern Sierra Nevadas, the
Santa Ynez Range in Santa Barbara County, to San Diego County are
quite uniform with brownish-gray primaries, the maculation even, pattern
lines distinct, and patches of green scales suffusing the wings in green,
varying in the same locality from slightly green to nearly leaf green in fresh
specimens, the green fading in aging outdoor specimens and in storage in
light-proof cabinets. Catocala benjamini (Brower, 1937), described as an
andromache subspecies, elevated to species rank (Brower, 1982), differs
in smaller size (Johnson and Walter, 1978), paler coloration, absence of
green scaling (Brower, 1937), in ovum scupturing, and in a larva unique
from andromache from the first instar onward (Johnson, 1981).

Ralph E. Wells took two males and a female at Jackson, Amador County,
California, that clearly represent a distinct northern California population
of andromache of subspecific rank deserving recognition and description.

Catocala andromache wellsi Johnson new subspecies

Holotype female: antennae dark gray, head, thorax grayish black with scattered
yellowish-brown scales, fore and middle legs greyish black, rear legs lighter,
abdomen proximally lighter, caudal two-thirds dark gray-brown; primaries above
dark gray, nearly black, green scales suffusing the wings, line patterns obscured and
in low contrast, t.a. line double, black, a black shade through the reniform spot to
the posterior part of the t.p. line, t.p. line black, silhouetted outwardly by lighter
scales, s.t. line black, subreniform spot pale gray, very conspicuous (not so in
allotype). Lower surfaces much as in type subsp. but orange color more vivid, darker
shades blacker, contrasts greater. Secondaries above yellow-orange, more vivid,

246

J. Res. Lepid.

median black band strong, reaching inner margin black shading, inner margin black,
fringe darkened, wing bases more heavily black scaled; lower surfaces more
contrasting, orange brighter, dark areas blacker than in subsp. andromache.
Forewing costal margin length 24 mm. Collected 19 June 1979 by Ralph E. Wells at
ultraviolet light, Jackson, Amador County, California, 318 m elevation.

Allotype male: much as in holotype, antennae, legs, body grayish-black;
primaries above grayish-black, patterns obscured, green scales suffusing the wings,
subreniform spot inconspicuous; beneath darker, more contrasting than in type
subsp. Secondaries above and below as in holotype, orange more vivid, dark areas
blacker than in subsp. andromache. Length of forewing costal margin 23 mm.
Collected 22 June 1977 at ultraviolet light, by R. E. Wells, Jackson, Amador
County, California, 318 m elevation.

Holotype and allotype to be deposited in the A. E. Brower collection, Augusta,
Maine. One paratype male, 16 June 1977, Jackson, Amador County, California, 318
m elevation, by R. E. Wells, in R. E. Wells collection.

Catocala andromache wellsi (Figure 1) is readily separable from subspecies
andromache by the grayish black antennae, body, and nearly black forewings with
low contests and obscuring of patterns, and the darker tones of the hindwings and
under surfaces. AU three specimens are much alike, the female holotype being
blackest. Efforts to secure more specimens are continuing. Further search should
discover the distribution of the subspecies in northern California. It is a pleasure to
name the subspecies for its discoverer, Ralph E. Wells, an active and able field
observer and collector of Lepidoptera for many years.

Fig. 1 . Above: Catocala andromache andromache Hy. Edw., cf left, 9 right. Below:
Catocala andromache wellsi Johnson, cf allotype left, 9 holotvoe rivht

20(4): 245=248, 1981(83)

247

Catocala mcdunnoughi broweraram Johnson new subspecies

Holotype female: antennae brownish black, head, thorax, fore and middle legs
very dark brown, with long, black, hoary -tipped setae, these forming a tuft at thorax
dorsal caudal margin, and smaller tufts on abdominal segments 1 and 2 dorsa.
Abdomen dark brown, paler anteriorly, ventrally dark gray brown. Pimaries above
dark brown, contrasts in pattern reduced, the wings suffused with patches of green
scales; wing bases dark brown, t.a. line strong, black, edged by light gray scales
proximally, pale area between line and reniform spot less distinct, subreniform
obvious, but darker; t.p. Une strong, black, in less contrast with background, s.t. line
narrow, medium gray, black bordered, wing submargin dark brown, intervein
lunules dark, subdued; lower surfaces much as in type subspecies, contrasts in
orange bands and dark areas greater. Secondaries much as in type subspecies, but
orange more vivid and black bands, black scaling at wing bases and along inner
margins stronger, outer margin lunules darker, apical white spot smaller; lower
surfaces much as in species, somewhat darker and more contrasting. Forewing
costal margin length 26 mm. Collected 15 July 1979, Moore Creek Forest Service
Campground, Amador- Calaveras Cos., California, elevation 975 m, at ultraviolet
Ught by R. E. WeUs.

Allotype male: much as in holotype, antennae nearly black, head, thorax, fore
and middle legs dark brown with long, black; hoary-tipped setae, tufted at thorax
dorsal caudal edge and on dorsa of abdominal segments 1 and 2; abdomen dark
brown above, paler anteriorly, gray-brown below; primaries above as in holotype,
dark gray-brown, pattern lines black, in low contrast to background, reniform spot
less obvious, subreniform reduced, marginal lunules heavily black outlined, fringes
darker, wings with obvious green scaling; below darker, more contrasting than in
type subspecies. Secondaries as in holotype, orange vivid, dark areas blacker,
greater contrasts above and below than in types subspecies, Forewing costal margin
length 25 mm. Collected 14 July 1979, at ultraviolet light, Moore Creek Forest
Service Campground, Amador- Calaveras Cos., California, elevation 975 m, by R.
E. WeUs.

Holotype and allotype to be deposited in the A. E. Brower collection, Augusta,
Maine, Two paratypes, males, 20 June 1981, 4 July 1981, Moore Creek Forest
Service Campground, Amador- Calaveras Cos., California, elevation 975 m, by R.
E. Wells, in R, E. WeUs collection.

Catocala mcdummoughi mcdunnoughi (Brower, 1937) was named from a type
series from Mount Lowe and Mount Wilson, Los Angeles Co., California. Through-
out the southern California coastal mountains above 1524 m the species may be
taken in woodlands containing Quercus chrysohpis Liebm. (Johnson, 1981). The
southern California population is highly uniform, forewings predominantly brown,
patterns of moderate contrast, and green sclamg never present. The four specimens
taken by R. E. Wells differ strikingly from those of southern California in the
^ayish-black darkening of the brown coloration, reduced pattern contrasts, and the
green scaling suffusing the wings, hi northern California is present a previously
uncollected and unrecognized population of subspecific rank (Figure 2). Efforts
continue to secure more specimens and to explore the distribution of subspecies
browerarum. The author is pleased to name this distinctive subspecies for the
species’ author, Dr. A. E. Brower, lifelong student of American Catocala, and Mrs.
Lurana Brower, able assistant to her husband.

248

J. Res. Lepid.

Fig. 2. Above: Catocala mcdunnoughi mcdunnoughi Brower, c? left, 9 right.
Below: Catocala mcdunnoughi browerarum Johnson, cf allotype left, 9
holotype right.

Acknowledgments: Thanks are expressed to Mr. Ralph E. Wells for his generosity
in permitting study of the specimens of the two subspecies and donation of the
types to the A. E. Brower collection of the Smithsonian Institution, and to Mr.
Gordon Marsh, Curator, Museum of Systematic Biolo^, University of California,
Irvine, and his staff for assistance in securing references and in the typing of the
manuscript.

Literature Cited

BROWER, A. E., 1937. Descriptions of a New Species and a New Race of Catocala
(Lepidoptera, Noctuidae). Bull. Brookl. Ent. Soc. 32(5):184-186.

, 1982. Change in status of Catocala andromache race “benjamini”

(Noctuidae). J. Lepid. Soc. 36(2): 159.

EDWARDS, H., 1885. New Species of Californian Moths. Entom. Amer. 1:49-50.
JOHNSON, J. w., 1981. The immature Stages of six California Catocala, parts 1,2,6.
The Pan Pac. Ent., submitted.

, & E. WALTER, 1980. Similarities and Differences in Fore wing Shape of Six

California Catocala Species (Noctuidae). Joum. Res. Lepid. 17(4):231-239,
2 tabs., 2 figs.

Journal of Research on the Lepidoptera

20(4): 249-250, 1981(83)

Notes

Notes on Maryland No. 10: Three New Butterfly Records for the
State of Maryland

Three new butterfly species have been recorded for the state of Maryland as
follows:

1. Atrytone palatka (Skipper), August 7, 1980, near Bucktown, Worcester

County, Maryland.

The specimen was a worn male. It was collected near the town of Bucktown, on
DeCourseys Road. The specimen was found in a sedge-like area which is typical of
the habitat where we collect Atrytone dion alahamae Lindsey. The specimen was
resting on one of the sedges. This area was further thoroughly investigated without
success for other specimens. Along with the A. palatka were flying A. d. alabamae.

2. Lycaena epixanthe (Bog Copper), July 19, 1981, Garrett County, near Cherry
Creek, near Bittinger.

For many years Robert Simmons has been seeking the Bog Copper, Lycaena
epixanthe Boisduval and Leconte, in the cranberry bogs of western Maryland
without success. On July 19, 1981, William A. Andersen and Philip Kean made a
joint field trip to the mountains of western Maryland. North of Bittinger, Garrett
County, they discovered a cranberry bog near Cherry Creek, Upon investigation of
the cranberry plants, eight Bog Coppers were collected. This is the first record of
the species in Maryland. The butterfly will undoubtedly be found in other cranberry
bogs in Garrett County.

3. Ascia monuste (Great Southern White), September 3, 1980, near Newbridge,
Worcester County, Maryland.

Bill Grooms and John Fales made a joint field trip to the Maryland eastern shore
on September 3, 1980. Near Newbridge, Bill Grooms pulled his usual collecting
stunt by netting a new species for the state of Maryland. Bill observed the specimen
zig-zagging down the road and cruising off the road on both sides attempting to find
a place to perch. The butterfly selected a large, fresh green leaf to alight and rest
upon. Bill approached the specimen very carefully so as not to excite it. As he
approached the specimen, he realized it was a worn male of the Great Southern
White, Ascia monuste Linnaeus. Realizing this was a new species record for
Maryland, he carefully secured the specimen. In his usual kind way he gave the
specimen to John Fales for his Maryland studies of Lepidoptera.

We would like to thank Bill Grooms and John Fales for permission to report their
records. A detailed citation of other Maryland records is given in the recent paper
by Simmons and Andersen, 1978(1980), Notes on Maryland Lepidoptera No.
9: Seven new butterfly records for the state of Maryland. J. Res. Lep. 17(4): 257-
259.

Robert S. Simmons, 8150 Loch Raven Blvd., Baltimore, MD 21204
William A. Anderson, 220 Melanchton Road, Lutherville, MD 21 093
Philip J. Kean, 1215 Stella Drive, Baltimore, MD 21207

250

J. Res. Lepid.

Further Notes Regarding Colias hecla Lefebvre (Lepidopterai

Pieridae) at Churchill Manitoba

Ferris’ recent comments concerning the occurrence of Colias hecla at Churchill
Manitoba (J. Res. Lepid. 20(l):50-54, 1981(82)) are well taken. However, some
additional information may be of interest to Ferris and other readers.

Since the 1974 season I have spent four additional seasons at Churchill
conducting systematic studies of the butterfly populations in the Churchill region.
With help from college students from the Churchill Northern Studies Center and
others, population centers were mapped, and ecological and behavioral data
recorded. This study covered additional areas not visited by Parshall and Costing in
1974 or Ferris in 1973. The results of the study will be reported in a second paper
now in preparation. With regard to the occurrence of C. hecla within the taiga areas,
the following notes may present a clearer picture.

During the entire study only 35 adults were recorded. The year 1977 was by far its
best with the author collecting 12 adults and fellow researchers together collecting
an equal number, the greatest number ever recorded for any single season at
Churchill.

C. hecla was always observed in association with tundra communities. It was never
recorded closer than ±75 meters to a forest-tundra ecotone. The species’
ovipositing choices at Churchill are found in a few locations within the open-spruce
forest ecotone referred to as “taiga” by some researchers. The open-spruce forest
ecotone is really a mixture of both tundra and forest communites. Ferris’ taiga
reference may therefore be considered a tundra observation. Such biotope
classification may help to clear the picture a little; in any case, Ferris’ observations
must be regarded as unique and not a data base for a theory for a taiga population of
eastern C. hecla. Other researchers in the Eastern Arctic should systematically
record flight patterns of C. hecla in their areas.

This author, as Ferris, has also collected hecla in the Western Arctic and the High
Eastern Arctic. The Churchill population of C. hecla is more closely related to
eastern populations than it is to western populations in terms of ecology, behavior,
and total biology. Thus great caution should be exercised when eastern and western
populations are compared, for the Western Arctic populations appear to represent
a less stable genetic entity. Many factors may be altering the biology of the species
in the Western Arctic. The possibility of sibling species within the Colias complex is
just one of the several matters requiring careful research.

I look forward to reading Ferris’ revision and hope that it will reflect careful
biological research which will help answer some problems that exist with hecla. A
literature review based on pinned specimens and the author’s opinion will be of
interest, but not nearly as useful as any biological insights.

I would like to suggest that terms such as taiga, climax and sub-climax, be dropped
from use by lepidopterists when referring to Arctic and Sub-arctic biotopes. These
terms do not reflect what is currently known about botanical communities in the
arctic ecosystem. Besides not representing current ecological thought, the terms
have a far too general meaning and do not help clarify a multi-dimensional research
approach.

David K. Parshall, 4424 Rosemary, Columbus, Ohio 4321 4

Journal of Research on the Lepidoptera

20(4): 251-252, 1981(83)

Book Review

Large White butterfly. The biology, biochemistry and physiology of Pieris brassicae

(Linneus).

Feltwell, J. [et aL], 1981. Series ent. 18(1982): i-XXVI, 1-535. Dr. W. Junk
Publishers, The Hague-Boston-London. Price: Hfl. 225.00/U.S. $98.00

The scope of this monograph is so broad that to attempt a comprehensive critical
review briefly could hardly result in a success. I therefore prefer a combination of a
descriptive “book notice” with a critical “book review” of some selected parts of the
book. J. Feltwell felt it necessary to engage three specialists to deal (alone or with
himself) with certain topics: R. I. Vane-Wright (taxonomy), M. R. Shaw (para-
sitology) and H. D. Burges (pathology).

The work is divided into 18 main sections which are split further into numerous
chapters. The main sections are: Nomenclature (1), Distribution (2), Life history
(3), Foodplants (4), Breeding (5), Development (6), Morphology and Anatomy (7),
Physiology (8), Hormones (9), Biochemistry (10), Migration (11), Senses (12),
Economic importance (13), Parasitic control (14), Pathogenic control (15), Preda-
tors (16), Chemical control (17) and Integrated control (18). Preface (by M.
Rothschild), Foreward, Acknowledgments, List of plates. List of tables and figures.
Botanical specific names index, Zoological specific names index and Subject index
complete the book. Comprehensive lists of references are given at end of each
section, and there are 10 plates, 80 tables and 49 figures in text.

The book is very well bound in cloth (size ca. 16 x 24 cm) and printed on
(unfortunately) glossy art paper which owing to its reflective properties makes
reading especially in artificial light unpleasant. Matt paper, as used for the early
volumes of Series entomologica would have been a significantly better choice, and
perhaps cheaper, too. High technical quality of production has always been
characteristic of the books published in this series, and the price usually expressed
abundantly this aspect. In case of the volume under review, the price has surely
overtaken the earlier standards. It surely is a great pity if a book of this kind is,
because of its prohibitive price, eliminated from reaching many of its potential
readers; or are the publishers particularly keen to provide good business for the
manufacturers of photocopying equipment?

The first section deals with the taxonomy of Pieris brassicae, but is therefore
somewhat inaccurately called just ‘Nomenclaure’. It is written in part by Feltwell
and Vane-Wright, partly by Feltwell alone. Some of the statements made in the
section are both difficult to justify and to understand. The authors deal with
‘subraces’ and with numerous aberrations and other infrasubspecific forms. They
follow B. O. C. Gardiner in their treatment of P. cheiranthi as subspecies of P.
brassicae and do not seem to realize that they contradict their own concept by the
inclusion of P. brassicae azorensis as a ‘subrace’ of their P. brassicae cheiranthi. In
spite of the flood of infrasubspecific names discussed, there is no mention of an
available name proposed for subspecies Pieris cheiranthi benchoavensis Pinker
(1968) and Tinker’s paper is not listed in the bibliography. The statements
regarding the adult morphology seem to be based entirely on observations made by
other authors and are incorrect in some applications (e.g. valva).

The section on ‘Distribution’ shows some gaps of considerable significance.
Although countless reports of English collectors are cited on the absence or

252

J. Res. Lepid.

occurrence of P. brassicae in countries they visited in the course of collecting of
butterflies, many significant indigenous standard works dealing with the distribu-
tion of the species are left out; I name here just a few authors that I easily
recall: Buresch & Tuleschkow, 1929-1943; Gozmany, 1968; Hruby, 1964; Korshu-
nov, 1972; Krzywicki, 1962; Verity, 1947. Local faunistic monographs are only
exceptionally cited. This results in avoidable inaccuracies, e. g. brassicae is stated
to be absent from Crimea although listed by Korshunov (1972).

Sections on ‘Life history’ and ‘Foodplants’ are considerably better than the first
two, though only a small proportion of data applies to wild populations. It would
have been interesting to try to define the ecological factors that are decisive for the
occurrence of the species and limit its distribution. Adult foodplants and preferences
are not dealt with.

Sections on ‘Development’ and ‘Breeding’ deal almost exclusively with observa-
tions relevant to brassicae in captivity. The section on ‘Morphology and Anatomy’
lacks adequate illustrations of genitalia of both sexes and of androconia; the
descriptions provided also leave much to be desired.

The section ‘Predators’ is probably the most complete account so far compiled on
any butterfly species.

Bibliographical references and citation selected for a reference book of this type
are particularly important for the reader. It is therefore most unfortunate that the
bibliographies probably contain a much high proportion of errors than should be
tolerated. About thirty minutes spent comparing some references selected at
random produced the following list of major errors (small errors are not included
here).

Page Original

Feltwell

Schurian, K., 1975. Notizen an Pieris 24
cheiranthi. Ent. Z., Frankf.a.M. 85:
252-256.

Kurentzov, N. J., 1929 53

Naumann, C. L., 1974. Immigrations 55
in Afghanistan 1972. Atalanta B 5:
82-88.

Schurian, K., 1975. Bemerkungen
uber Pirn's cheiranthi. Ent. Z.,
Frankf.a.M. 85: 252-256.

Kusnezov, N. J., 1929

Naumann, C. M., 1974. Beobachtun-
gen uber den Verlauf einer Lepid-
opteren-Immigration in Afghanis-
tan 1972. Atalanta, Muermerstadt
5: 82-86.

Schreiber,H. in Mueller, P. 1976. Der 56
Katalog der Ortlichkeiten im
Bundesrepublik Deutschland. Vol.

2. Lepidoptera.

Schreiber, H. 1976. Fundortkataster
der Bundesrepublik Deutschland.
2. Lepidoptera.

To summarize the observations made in this very brief, and necessarily perfunc-
tory, review, it can be said that the publication of the compilation on Pieris brassicae
was in principle a very good idea. The same implies to the choice made by the
author. That the author had not asked selected specialists to read final drafts of
each section resulted in the book being only a qualified success. The amount of work
Feltwell put into this book would have been worth the ‘last touch of perfection’.

INSTRUCTIONS TO AUTHORS

Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, on 8^2 x 11 inch paper with wide margins. Number all pages
consecutively and put author’s name at top right corner of each page. If your typewriter
does not have italic type, underline all words where italics are intended. Footnotes,
although discouraged, must be typed on a separate sheet. Do not hyphenate words at the
right margin. All measurements must be metric, with the exception of altitudes and
distances which should include metric equivalents in parenthesis. Time must be cited on
a 24-hour basis, standard time. Abbreviations must follow common usage. Dates should
be cited as example: 4. IV. 1979 (day-arabic numberal; month-Roman numeral; year-
arabic numeral). Numerals must be used before measurements (5mm) or otherwise up to
number ten e.g. (nine butterflies, 12 moths).

Title Page: All papers must have the title , author’s name, author’s address, and any
titular reference and institutional approval reference, all on a separate title page. A
family citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.

Abstracts and Short Papers: All papers exceeding two typed pages must be ac-
companied by an abstract of no more than 300 words. An additional summary is not
required.

Name Citations and Systematic Works: The first mention of any organism should
include the full scientific name with author (not abbreviated) and year of description. New
descriptions should conform to the format: male: female, type data, diagnosis, distribu-
tion, discussion. There must be conformity to the current International Code of
Zoological Nomenclature. We strongly urge deposition of types in major museums, all
type depositions must be cited.

References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbrevations must conform to the World List of
Scientific Periodicals. Do not underline periodicals. If four or less references are cited,
please cite in body of text not in Literature Cited.

Tables: Tables should be minimized. Where used, they should be formulated to a size
which will reduce to 4 x 6% inches. Each table should be prepared as a line drawing or
typed with heading and explanation on top and footnotes below. Number with Arabic
numerals. Both horizontal and vertical rules may be indicated. Complex tables may be
reproduced from typescript.

Illustrations: Color must be submitted as a transparency (i.e., slide) ONLY, the quality
of which is critical. On request, the editor will supply separate detailed instructions for
making the most suitable photographic ilustrations. Black and white photographs should
be submitted on glossy paper, and, as with line drawings, must be mounted on stiff white
cardboard. Authors must plan on illustrations for reduction to the 4 x 6y2” page.
Allowance should be made for legends beneath, unless many consecutive pages are used.
Drawings should be in India ink at least twice the final size. Include a metric scale or
calculate and state the actual magnification of each illustration as printed. Each figure
should be cited and explained as such. The term “plate” should not be used. Each
illustration should be identified as to author and title on the back, and should indicate
whether the illustration be returned.

Legends should be separately typed on pages entitled “Explanation of Figures”. ^
Number legends consecutively with separate paragraph'for each page of illustrations. Do
not attach to illustrations. Retain original illustrations until paper finally accepted.

Review: All papers will be read by the editor(s) & submitted for formal review to two
referees. Authors are welcome to suggest reviewers, and if received, submit name &
comments of reviewers.

Volume 20

Number 4

Winter, 1981(83)

IN THIS ISSUE

Date of Publication: April 15, 1983

Butterfly Taxonomy: A Reply

Lee D. Miller & F. Martin Brown 193

Nomenclature, Taxonomy and Evolution

Paul R. Ehrlich & Dennis D. Murphy 198

Editorial Note 205

Acknowledgment 206

Notes on the Life History and Baja California Distribution of
Chlorostrymon simaethis sarita (Skinner) (Lepidoptera:
Lycaenidae)

John W. Brown 207

Biology and Immature Stages of Australian Ethmiid Moths
(Gelechioidea)

Jerry A. Powell 214

Field Study of Phyciodes batesii (Reakirt) and P. tharos (Drury)
from a Site in the Black Hills, South Dakota (Lepidoptera:
Nymphalidae: Melitaeinae)

Clifford D. Ferris 235

Notes: An Aberration of Glaucopsyche lygdamus (Lycaenidae)
with a Complete Scolitantidine Dorsal Pattern
A Reared Gynandromorph of Tatochila (Pieridae)

Two Homoeotic Pieris rapae of Mexican Origin
(Pieridae)

An Apparent “Intersexual” Colias eurytheme (Pieridae)

Arthur M. Shapiro 240

Two New California Catocala Subspecies (Noctuidae)

John W. Johnson 245

Notes 249

Book Review 251

Index for Volume 20 253

Cover Illustration: Scanning electron micrograph of Australian E. postica
eggshell on nylon mesh (x42) by Barry FOshie. See J. Powell article, page 217, this

issue.

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