Volume 63 Number 1
29 Apr 2009
ISSN 0024-0966

Journal of the

Lepidopterists' Society

Published quarterly by The Lepidopterists' Society

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Cover Illustration; Omiodes continuatalis, an endemic Hawaiian crambid moth formerly thought to be extinct, hut recently rediscovered.
Photo by William P. Haines. (See journal article on page 1 1).

Journal of
The Lepidopterists’

CSMIT HSOA^

® 22 2017

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Society

Volume 63 2009 Number 1

Journal of the Lepidopterists’ Society
63(1), 2009, 1-10

A NEW SPECIES OF DRASTERIA HUBNER (NOCTUIDAE: CATOCALINAE: MELIPOTINI) FROM

ARIZONA, WITH COMMENTS ON THE GENUS

Jan Metlevski

Department of Entomology, 123 West Waters Hall, Kansas State University, Manhattan, KS 66506-4004, U.S.A; email: jmetlevs@hsu.edu

AND

Gregory Zolnerowich

Department of Entomology, Kansas State University, Manhattan, KS 66506-4004, U.S.A; email: gregz@ksu.edn

ABSTRACT. Drasteria walshi n. sp. (Noctuidae: Catocalinae) from southeastern Arizona is described and illustrated. 'The new species be¬
longs to the D. fumosa-D. divergens species group. Drasteria walshi is distinguished from related species of Drasteria by wing color and pat¬
tern, and structure of male and female genitalia. Taxonomic problems in Drasteria are discussed.

Additional key words: taxonomy, Drasteria, new species. Arizona, Nearctic

The genus Drasteria Hubner, 1818, with more than
60 described species, is the largest genus in the tribe
Melipotini (sensu Fibiger & Lafontaine 2005). It is
exclusively Holarctic in distribution, with 27 species
known from the Nearctic region and 35 species known
from the Palearctic region. No species are known to
occur in the tropics or Southern hemisphere. The only
comprehensive study of the Palearctic species of the
genus was published by John (1910) almost a hundred
years ago. Since then, a dozen new species and
subspecies of Drasteria have been described from the
Palearctic region. The Palearctic species of Drasteria
are in need of revision because the status of many taxa
remains unresolved, and some widely distributed
species may actually be complexes of closely related
species, as is the case for Drasteria rada (Boisduval,
1848) (Metlevski et al ., unpublished data). Richards
(1939) revised the Nearctic species of Drasteria, and
few changes have occurred in the taxonomy of Nearctic
species since then. Some taxonomic problems pointed
out by Richards, such as the relationship between D.
tejonica (Behr, 1870) andD. howlandii (Grote, 1864), or
the status of D. nichollae (Ilampson, 1926), have not
been resolved yet. Richards' 1939 revision downgraded
to subspecies (race by Richards) a number of taxa

originally described as separate species by other
workers. Poole’s 1989 catalog of the world Noctuidae
(Poole 1989) did not recognize any taxa at the
subspecies level and synonymized all the names
applicable to the taxa treated as subspecies by Richards.
As a result, all Drasteria names synonymized by Poole
have been excluded from the lists of Nearctic fauna,
such as Nomina Insecta Nearctica (Poole & Gentili
1996). Nevertheless, the status of many taxa treated by
Richards as subspecies of other Drasteria species
remains unclear. Drasteria hastingsii (H. Edwards,
1878) is now recognized as a valid species, and Wagner
et al. (in press) provide evidence that Drasteria
graphica atlantica Barnes & McDunnough, 1918
represents a full species. It is likely that others, such as
D. sabulosa abrupta (Barnes & McDunnough, 1918) or
D. adumbrata alleni (Grote, 1877), are also valid
species.

In addition, our current investigations suggest that
some North American species of Drasteria are actually
complexes of closely related species. For example,
populations of D. inepta (Edwards, 1881) from
Colorado and northern New Mexico are not nonspecific
with populations from southeastern Arizona and
southwestern New Mexico. Finally, discoveries of new

2

Journal of the Lepidopterists’ Society

Fig. 1. Medial surface of Drasteria male valve (based on a left valve of D. pallescens (Grote & Robinson, 1866)) (c - costa; co - conus; cp
-costal process; dlcp - dorsal lobe of costal process; hrp - harpe; pulv - pulvinus; sel - sacculus; scle - saccular extension; vlvl - valvula).

Fig. 2. Female genitalia of Drasteria (based on D. sinuosa (Staudinger, 1884)) (aa - apophyses anteriores; an - antrum; ap - apophyses pos-
teriores; be - bursa copulatrix; col - colliculum; db - ductus bursae; la - lamella antevaginalis; Ip - lamella postvagin'alis; pa - papilla analis;
pb - pseudobursa).

Volume 63, Number 1

3

species of Drasterici in North America are still
occurring. Drasteria convergens Mustelin, 2006, was
described recently from California, and below we
describe a new species from southeastern Arizona.

The terminology used here for structures of the male
valva follows the scheme shown in Fig. 1. We use the
term “harpe” in a broad sense since the problem of
homologies of valval structures, to which this term is
applicable in various groups of Noctuidae, remains
unresolved. For this reason we follow with some
modifications the broad definition of the term “harpe”
given by Kuznetzov & Stekolnikov (2001), and define
the harpe as various and not necessarily homologous
sclerotized structures on the inner surface of valva, that
are not derived from the costa or sacculus. Here we use
the term “harpe” for the long and slender process
(processus basalis valvae in John 1910) arising from a
sclerotised, angled ridge on the inner surface of
proximal part of the valva. We also use the term “conus”
for the small process located just below the haq:>e, and

the term “pulvinus” for the shelflike process arising
laterad from the inner side of the basal part of the
saccular extension. The last two terms were introduced
by John (1910) for Drasteria but have not been used
since. Terminology for male genitalia other than valva
follows Goater et al. (2003).

Species of Drasteria have aedeagi with a veiy
complex vesica, consisting of a number of diverticula of
different sizes. The complexity of the vesica of species
of Drasteria was shown only recently (Goater et al.
2003; Mustelin 2006); previous taxonomic works on the
genus have not made mention of the vesica. Our current
investigations show the number and relative position of
diverticula are good characters for grouping members of
the genus into species groups. Thus, we include a
detailed description of the vesica in the species
description.

Terminology for female genitalia follows Kiihne
(2005) and is shown in Fig. 2. Terminology for wing
patterns is shown in Fig. 3.

Fig. 3. Wing pattern of Drasteria. dorsal surface (based on D. christophi (Alpheraky, 1895)) (am - antemedial line; as - apical spot; ash - api¬
cal shadow; b - basal line; ba - basal area: cl - discal spot; in - medial shadow; ma - medial area; pin - postmedial line; pina - postmedial area;
pmb - postmedial band; r - reniform spot; st - subterminal line; sta - subterminal area; t - terminal line; ts - terminal spot; tsh - terminal
shadow).

4

Journal of the Lepidopterists’ Society

AZ Graham Co. 9,000 ft.
Pinaleno Mis, Cunimingham
Campdgr, UV + MV lights
20 May 2002. B. Walsh leg.

wa&ly

WeTLftrsjci

PARATYPE

Onion Saddle
Elv. 7600, ft.
Chirachua Mts AZ
June 14. 2007
Leg. Ray B. Nagle

KSU-MEPAR Collection

<?

Gen. Prep. No
J. Metlevski

,W4

. 2006

Gen. Prep. No. 1068
J. Metlevski 2007

Figs. 4-9. Drasteria toalshi. 4-6, holotype. 4, dorsal surface. 5, ventral surface. 6, pin labels. 7-9, female paratvpe, Chiricahua Mts. 7, dorsal
surface. 8, ventral surface. 9, pin labels.

Volume 63, Number 1

5

Figs. 10-13. D. divergens. 10-11, male, White Pine Co., Nevada. 10. dorsal surface. 11, ventral surface. 12-13, f. socia, female, Santa Bar¬
bara; California. 12, dorsal surface. 13, ventral surface.

Abbreviations for institutions and collections used in
the text are as follows: KSU-MEPAR = Kansas State
University Museum of Entomological Prairie Arthropod
Research; BW = Bruce Walsh collection (Tucson, AZ);
RN = Ray B. Nagle (Tucson, AZ) collection.

Drasteria walshi Metlevski, new species
(Figs. 4-9, 18-20)

Diagnosis. Drasteria walshi is most similar to D.
divergens (Figs. 10-13, 21, 23), especially to the form
with orange hind wings, D. divergens f. socia (Behr,
1870) (Figs. 12, 13). The following features can be used
to differentiate D. walshi from D. divergens: in D.
walshi the underside of the wings is pinkish, while they
are yellow to pale yellow in D. divergens ; in D. walshi

the terminal spots on the underside of the wings are
obscured or completely absent, while in D. divergens
there are strong black terminal spots on the underside
of the wings; in D. walshi the discal spot is a thin dark
comma-like line not connected to the postmedial band,
compared to a strong black discal spot connected to the
postmedial band (at least vein CuA2 is black from the
postmedial band to the discal spot, but usually the
connection is much stronger) in D. divergens.

Among other species of Drasteria occurring in
southeastern Arizona only one, D. tejonica (Behr, 1870),
has forewings with similar pattern and colored hind
wings. Drasteria tejonica however is noticeably smaller
in size and has much stronger black pattern on the hind
wings (Figs. 14-17). Males of D. tejonica (Figs. 14-15)

6

Journal of the Lepidopterists’ Society

Figs. 14-17. Drasteria tejonica, Pima Co., Arizona. 14-15, male. 14, dorsal surface. 15, ventral surface. 16-17, female. 16, dorsal surface. 17,
ventral surface.

can be easily separated from D. walshi by the coloration
of the hind wings, which are white basally in D. tejonica.

In males, the costal processes of the valvae of D.
walshi are robust, elongate, sinuous and pointed at the
tip (Fig. 18). This readily distinguishes D. walshi from
1). divergens (Fig. 23), as well as from any other species
in the genus. Female genitalia of the new species (Fig.
20) could be confused only with those of D. divergens
(Fig. 21) and D. fumosa (Fig. 22). The antevaginal plate
in D. walshi is more than 3 times longer than the
colliculum, while just slightly longer in D. divergens and
about 2 times longer in D. fumosa. The antevaginal
plate of D. walshi is semi-rectangular in shape,
compared to triangular with a blunt tip in D. fumosa.
The shape of the antevaginal plate of D. walshi is similar

to that of D. divergens : both species have a rectangular
anterior portion and a slightly narrower posterior
portion. However, in D. walshi the broader anterior
portion is about half the length of the narrower
posterior portion, while in D. divergens the length of
both parts is subequal.

Description. Head: Light brownish gray; labial palpus light grey,
with some dark brownish gray scales on the external side of first and
second segments; antenna filiform in both sexes.

Thorax: Covered with long flattened and slightly spatulate hairs
dorsally, with long simple hairs ventrally; patagia light brownish gray,
with a dark brown longitudinal stripe; remainder of thorax little darker
and with light pinkish tint; tegulae slightly rimmed with dark brown
hairs on external margin; legs same color as thorax except for tarsi,
which are darker brownish gray with a ring of light gray scales at distal
end of each segment.

Forewing: Length 19-23 mm; males and females similar; ground
color light brownish grav with light pinkish tint; basal line double,
black, enclosed area light brownish gray; basal area brownish gray in
anterior half, dark gray in posterior; antemedial line double, internal

Volume 63, Number 1

and external lines black, enclosed area filled with dark brown, internal
line making a strong long tooth toward base of wing between veins A
and Cu and two small teeth at veins A and Sc, entire antemedial line
almost straight from costa to vein A, abruptly curving inward after vein
A; medial area very light brownish gray, somewhat darker in outer part
and at costa; medial fine brown, not strongly contrasting, double, the
internal line thin, external one wider; both lines strongly expanded
and becoming dark brown or almost black at costa; reniform spot dark
brown, black rimmed, with strong whitish outline on outer margin and
thin whitish outline followed by thin black line on inner margin;
postmedial line double, area enclosed between internal and external
lines dark brown except at costa where pinkish, internal line strong
and black through its entire length, external one weaker, marked with
dark brown or black scales, almost disappearing in the middle;
postmedial line turning abruptly toward the apex at vein Rj and almost
parallel to costal margin until vein R„, then bending again and curving
outward, includes strong tooth pointed outward at vein and
another smaller tooth at vein M3, curving inward in between; making
an abrupt turn at vein CuAp continuing along this vein toward base of
reniform spot then turning toward posterior margin of wing, curving
slightly outward between base of reniform and vein A, turning toward
anal angle of wing at vein A; area between reniform spot and
postmedial line brownish grey to brown, filled with brown and light
brownish grey scales, veins M3 and CuA, within this area dark brown
with strong whitish outlines joined with similar outline on inner
margin of reniform spot; postmedial area dark gray, pinkish along
postmedial line at costa; subterminal line slightly incurved between
costa and vein Rp then slightly waving and subparallel to inner margin
of wing, at vein A, turning toward anal angle of wing, dark brown at
veins, otherwise black in anterior half of wing, mostly brown, with
some black scales between veins in posterior hatf; 4 strong black arrow
marks arising from subterminal line between veins in anterior half of
wing, and pointing toward base of wing; subterminal line with a thin
pale brownish gray outline along outer side, immediately followed by
dark brown line; subterminal area light brownish gray, somewhat
darker in place where terminal shadow is located; apical shadow
obvious, as a dark brown path, connected to dark brown outer outline
of subterminal line; terminal line fine, wavy, obvious, dark brown to
almost black; fringe pinkish at base, otherwise grayish brown,
checkered. Underside pale pinkish to pinkish; reniform spot dark
brown to black; dark brown transversal shadow extending from the
middle of costa through inner edge of reniform toward the anal angle,
ending well before anal angle between veins CuA., and A; subterminal
line greasy pinkish with dark edges at costa, otherwise dark brown to
black, convex, wide at costa, rapidly narrowing toward posterior edge
of wing becoming a thin line at its posterior end, meeting transversal
shadow at vein CuA, and not extending after this point, outer edge of
subterminal line wavy, inner edge evenly convex; terminal and apical
shadows brownish, obscured and sometimes barely visible; terminal
line thin, dark brown, wavy, vanishing at anal angle; fringe pinkish at
base, otherwise grayish brown, lighter toward anal angle.

Hind wing: Ground color orange pink, a few dark brown scales at
anal margin and occasionally a few' brownish scales near base of ufing
and on Cu stem; discal spot a narrow dark grayish brown line along
cross-vein, not connected to postmedial band; postmedial band dark
grayish brown to black, narrow through most of its length but strongly
widened between vein R, and costa, subparallel to outer margin
between anal angle and vein CuA„ approaching vein CuA, curving
abruptly mesad, and at vein CuA, angling toward apex, then turning
toward costa between veins M, and Mp outer margin of postmedial
band turning sharply outward at vein ftp terminal and apical spots
narrow patches, dark grayish brown to black, with obscure inner edge,
almost disappearing in some specimens; terminal line obvious through
most of its length, vanishing near anal angle, dark grayish brown to
black, thin, wavy; basal portion of fringe pink, outer portion pink
orange, greasy orange brown at terminal spot, with some blackish
scales between basal and outer layers at terminal and apical spots.
Underside pale pinkish to pinkish, some brown or brownish scales
along costal margin; discal spot similar to that on upper side but
weaker; postmedial band intermittent, marked by few' brown scales at
costal margin, remaining remnants present only between veins M, and
M„ and between vein CuA, and anal angle; terminal spot a dark
brown to blackish shadow, weak, in some specimens almost
disappearing; apical spot absent; terminal line weaker than on upper
side, otherwise similar; fringe same as upper side.

Abdomen: Ochreous gray, with some scattered dark brown scales.

Male genital capsule (Fig. 18): Uncus strong, curved, pointed at tip,
dorsally with a low' longitudinal crest and dense hairlike setae in third
distal quarter; scaphium long, narrow, articulated at base of uncus;

valvae obovate, slightly asymmetrical, left valve slightly smaller,
narrower in the middle and with more sinuous dorsal margin; valvula
rounded, acute at ventro-distal angle, with a small tuft of setae on
inner surface at ventro-distal end; area occupied by coremata small,
subbasaly on outer side of valva, extending along sclerotized base of
valva from ventral edge of base of costa to ventral side of distal half of
sacculus; sacculus about 4 times as long as wide, its dorsal edge slightly
concave; saccular extension strong, slightly curved, ending just before
tip of valvula in a small free prong-like projection turned inward
toward genital capsule; pulvinus strong, broad, broader on left valva,
tip of pulvinus with robust elongate setae; harpe curved, slender,
smoothly narrowing to tip, apex flattened and rounded; conus present
on both valvae, left one tapered, about 2 times as long as wide, right
one minute, short; tip of conus bearing few microscopic setae; costa
moderately sclerotized, with costal process present on both valvae;
costal process prominent, broad based, strongly elongated, sinuous,
tapered, pointed at tip; costal process on left valva shorter, directed
along longitudinal axis of valva; costal process on right valva longer,
directed slightly dorsad, reaching dorsal margin of valvula; saccus
short, V-shaped: juxta symmetrical, about 2 times as long as wide, with
deep V-shaped incision on dorsal margin and two short, curved, and
divergent ventroanterior arms.

Aedeagus (Fig. 19): strong, straight, tubular, membranous in distal
half dorsally but with a small sclerotized area at base of vesica,
otherwise well sclerotized, its membranous portion finely spiculose;
coecum short, slightly flattened dorsoventrally at tip; ventral plate of
carina strongly sclerotized, long, narrow, slightly expanded at top,
basal half of right margin w'ith a strong longitudinal fold; dorsal plate
of carina absent; inflated vesica short, turned dorsad, with a complex
system of diverticula (insertion on Fig. 19): diverticula arising on
ventral side right of carinal plate (2 large and 4-5 very small): 1.
diverticulus next to the tip of carinal plate, broad, about twice as long
as wade, distally rounded, spiculose on right side, directed caudad, tip
slightly turned left; 2„group of 4-5 aligned small spiculose diverticula,
next right to diverticulus 1, followed by diverticulus 3; 3. the biggest
diverticulus, two times longer than next longest diverticulus, slightly
narrowing to the tip, finely spiculose, directed right laterodorsad,
perpendicular to longitudinal axis of aedeagus; diverticula arising on
ventral side left of carinal plate (1 bigger and 1 veiy small): 4.
diverticulus next to tip of carinal plate, short, bifurcated at tip,
spiculose, directed left laterad. perpendicular to longitudinal axis of
aedeagus, one of tips turned cephalad; 5. very small, lobelike, with no
spicula, between diverticulus 4 and tip of aedeagus; diverticula arising
at left dorsolateral angle of vesica (2 bigger and 1 veiy small): 6.
globular, expanded distally, as long as wide, finely spiculose, directed
dorsocaudad, with a narrow fingerlike projection arising left
ventrolaterally and directed ventrad; 7. second longest, tapering,
finely spiculose, curved, directed cephalad, tip curving right
dorsocephalad; 8. very small, short, narrowing at tip, with no spicula,
between diverticulus 7 and base of vesica; top of vesica exposed
dorsad, bearing 1 small diverticulus and 1 small lobe: 9. minute
diverticulus, with no spicula, arising on right side close to diverticulus
2: 10. small lobe with no spicula, close to the base of diverticulus 6;
posterior side of vesica : 11. bearing a small lobe with no spicula
between diverticulus 1 and top of vesica.

Female genitalia (Fig. 20): Ovipositor telescopic; papillae analis
with moderately sclerotized longitudinal band dorsally, weakly
sclerotized in other parts, narrow, about 5 times as long as wide,
strongly narrowing in distal third, tips rounded, membranous; surface
of papillae with short scattered setae, which are denser on tips, some
long setae at the base and tips of papillae analis; apophyses long and
slender, posterior ones about 1.7 times longer than anterior; lamella
postvaginalis (postvaginal plate) a V-shaped sclerotized band, its ends
not connected to eighth segments sclerotization; antrum wide, with
almost parallel lateral edges, about 2 times as long as wide,
dorsoventrally flattened, with large sclerotized antevaginal plate
ventrally, otherwise membranous; antevaginal plate occupying almost
whole ventral side of antrum, semi-rectangular in shape, slightly
narrowing distally, with slightly convex posterior edge; ductus bursae
short, slightly longer than wide, about 2 times narrower and 2.5-3
times shorter than the antrum; colliculum as long as wide, its left edge
slightly longer than the right one, edges of colliculum not overlapping;
bursa copulatrix asymmetrically ovate, membranous, with some

Fig. 18-22. 18. Male genital capsule of D. walshi (posterior), holotype. 19. Aedeagus of D. walshi with vesica everted (ventral view), holo-
type. Inset: vpc - ventral plate of Carina; 1-7 - numbers of diverticula according to the description. 20, Female genitalia oi D. walshi (ventral
view), paratype, genital prep, by Jan Metlevski No. 945. 21, Female genitalia of D. divergens (ventral view), Tuolumne Co., California, genital
prep. b\ Jan Metlevski No. 774. 22, Female genitalia of D. fumosa(ventr;i\ view), Pima Co., Arizona, genital prep, by Jan Metlevski No. 963.

Volume 63, Number 1

9

Fig. 23. Male genital capsule of D. divergent (posterior), genital prep, by Jan Metlevski No. 794.

longitudinal wrinkles on left side anterad of insertion of ductus bursae,
otherwise smooth; insertion of ductus bursae posteriorly, slightly left;
internal wall of bursa copulatrix in posterior half of bursa with very
minute spines, most visible at wrinkled area, becoming smaller and
barely visible laterad, and disappearing anteriorad; pseudobursa
arising posteriorly, right of insertion of ductus bursae, coiling dorsad
and to right, making 0.5 coil; ductus seminalis arising from the tip of
pseudobursa.

Holotype. d, “AZ, Graham Co. Pinaleno Mts. Cunningham
Campgrd. 9000 ft. UY + MV lights. 20 May 2002. B. Walsh leg.”,
“Genitalia Prep, by J. Metlevski No 766”, deposited in KSU-MEPAR
(Figs. 4-6, 18, 19).

Paratypes. (3 d and 3 9): 1 d, same label data as holotvpe, in BW;
1 9, “AZ, Graham Co. Pinaleno Mts. Cunningham Campgrd. 9000 ft.
Ponderosa pine habitat. 14 June 2005. UV + MV lights. B. Walsh leg.”,
“Genitalia Prep, by J. Metlevski No 945”, in BW (Fig. 20); 1 d, “AZ,
Graham Co. Pinaleno Mts. Cunningham Campgrd. 9000 ft.
Ponderosa pine habitat. 5 June 2007. B. Walsh leg.", in BW; 1 9, “AZ,
Cochise Co., Chiricahua Mts. Onion Saddle, 7700 ft, Ponderosa
Pine/Oak habitat, 14 June 2007. B. Walsh leg.”, “Genitalia Prep, bv |
Metlevski No 1084”, in BW; 1 9, “AZ, Cochise Co., Chiricahua Mts.
Onion Saddle, 7600 ft, June 14 2007. Leg. Ray B. Nagle”, “Genitalia
Prep, by J. Metlevski No 1068”, in RN (Fig. 7-9); 1 d, same label data
as above, “Genitalia Prep, by |. Metlevski No 1069”, in RN.

Variation. Specimens from the Chiricahua
Mountains have the hind wings a slightly brighter
ground color and with a stronger dark pattern compared

to specimens from the Pinaleno Mts.

Biology. Unknown. Specimens have been collected
in the second half of May and first half of June.

Distribution. Pinaleno Mountains (Mt. Graham)
and Chiricahua Mountains in southeastern Arizona.

Etymology. Drasteria walshi is named in honor ol
Bruce Walsh, Department of Ecology and Evolutionary
Biology, University of Arizona, Tucson, in recognition of
his invaluable efforts in studying the lejridopteran fauna
of southeastern Arizona.

Discussion. Female genitalia of Drasteria walshi
have a greatly enlarged antevaginal plate, which
occupies most of the ventral side of the antrum and is
longer than the colliculum. This condition of the
antevaginal plate links D. walshi with D. fumosa
(Strecker, 1898) and D. divergens (Behr, 1870), which
are the only other Nearctic species in the genus sharing
this character state. In the male genitalia, the aedeagus
also provides characters linking D. walshi with the two
species mentioned above in that the general structure of
the vesica, such as the relative position and number of

10

Journal of the Leridopterists’ Society

diverticula, of D. walshi is most similar to that of D.
divergens and D. fumosa.

All I mown specimens of Drasteria walshi have been
collected in the Pinaleno and Chiricahua Mountains at
altitudes between 7500-9000 feet (approximately
2300-2750 m) in ponderosa pine or mixed ponderosa
pine/oak forest habitats. Both the Pinaleno Mts. and
Chiricahua Mts. are isolated high mountain massifs
surrounded by lowland plains grasslands and deserts.

AC KN OWLE DG E ME NTS

We thank P. A. Opler and B. C. Kondratieff (Colorado State
University), J. H. Boone (Field Museum, Chicago), E. H. Met-
zler (Alamogordo, New Mexico), and R. B. Nagle (Tucson, Ari¬
zona) for loans of specimens used in this study. We are grateful
to T. Mustelin (San Diego, California), D. L. Wagner (University
of Connecticut) and two anonymous reviewers for reviewing the
manuscript. This article is Contribution No. 0S-395-J from the
Kansas Agricultural Experiment Station (KAES) and was sup¬
ported in part by KAES Hatch Project No. 583 (Insect System¬
atic^ ).

Literature Cited

Fibiger, M. & [. D. Lafontaine 2005. A review of the higher classifi¬
cation of the Noctuoidea (Lepidoptera) with special reference to
the Holarctic fauna. Esperiana 11: 7-92.

Goater, B., L. Ronkay & M. Fibiger. 2003. Catocalinae and Plusi-
inae. Noctuidae Europaeae, Volume 10, Sor0, 452 pp.

John, O. 1910. Eine revision der gattung Leucanitis Gn. Horae Soci-
etatis Entomologicae Rossicae 39: 585-640.

Kuhne, L. 2005. Revision und Phylogenie der Gattungsgruppe Crvp-
sotidia Rothschild, 1901. Tachosa Walker, 1869, Hypotacha
Hampson, 1913, Audea Walker, [1858] 1857 und Ulotrichopus
Wallengren, 1860 (Lepidoptera, Noctuidae, Catocalinae). Esperi¬
ana Memoir 2: 1-220.

Kuznetzov, V. I.. & A. A. Stekolnikov. 2001. New approaches to the
system of Lepidoptera of world fauna (On the base of the func¬
tional moi-phologv of abdomen). St. Petersburg, Nauka. 462 pp.

Lafontaine, J. D. & M. Fibiger. 2006. Revised higher classification
of the Noctuoidea (Lepidoptera). Canadian Entomologist, 138
(5): 610-635.

Mustelin, T. 2006. Taxonomy of southern California Erebidae and
Noctuidae (Lepidoptera) with descriptions of twenty-one new
species. Zootaxa 1278: 1—47.

Poole, R. W. 1989. Noctuidae 1-3. In Heppner. |. B. (ed.), Lepi-
dopterorum Catologus (New Series) fasc. 118. Leiden, New York,
Kopenhavn, Koln. 1314 pp.

Poole, R. W. & P. Gentili (eds.). 1996. Nomina Insecta Nearctica.
Volume 3. Diptera, Lepidoptera, Siphonaptera. Entomological
Information Sendees, Rockville, Maryland, 1143 pp.

Richards, A. G. 1939. A revision of the North American species of
the Phoberia-Melipotis-Drasferia group of moths (Lepidoptera,
Phalaenidae). Entomologica Americana 19: 1-100

Received for publication 1 July 2008; revised and accepted

9 December 2008

Volume 63, Number 1

11

Journal of the Lepidopterists’ Society
63(1), 2009, 11-20

BIOLOGY AND DISTRIBUTION OF A RECENTLY REDISCOVERED ENDEMIC HAWAIIAN
LEAFROLLER MOTH, OMIODES CONTINUATALIS (CRAMBIDAE)

Cynthia King

University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Made Way, Gilmore Hall 310, I lonolulu, III 96822

e-mail: cbaking@hawaii.edu

Daniel Rubinoff

University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Maile Way, Gilmore I Pall 310, Honolulu. HI 96822

e-mail: rubinoff@hawaii.edu

and

William Haines

University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Maile Way, Gilmore Hall 310, Honolulu, III 96822

e-mail: whaines@hawaii.edu

ABSTRACT. The endemic Hawaiian leafroller moth Omiodes continuatalis was documented by early entomologists to be one of the most
common species encountered during initial collecting trips in the Hawaiian Islands at the end ol the 19th century. The species was declared ex¬
tinct in 1982, and while subsequent surveys have resulted in their “rediscovery”, it is evident that O. continuatalis populations have experienced
declines and localized extinctions throughout their historical range. The objective of our research was to document the life history, biology, gen¬
eral morphology, and behavior of this little studied species. To complete the investigation, we observed both wild caught and laboratory reared
O. continuatalis. We also conducted light trap surveys to document the current distribution of O. continuatalis across the Hawaiian Islands. We
have integrated our findings from this study with all available historical data on O. continuatalis, to create a document which can be referenced
for future research relating to the biology or conservation of this species.

Additional Key words: pyraloid, light trap, fossorial, development, extinction

The genus Omiodes Guenee, 1854 (Lepidoptera:
Crambidae) occurs in tropical and subtropical regions of
Oceania, Southeast Asia and the Americas, and contains
over 100 species, including 23 species endemic to the
Hawaiian Islands. The subject of this research, Omiodes
continuatalis (Wallengren, 1860), is a generalist grass-
feeding species, originally documented from all of the
high Hawaiian Islands, with the exception of Kahoolawe
(Zimmerman 1958). The Reverend Thomas Blackburn
found O. continuatalis to be one of the most common
moths encountered during collecting trips in Hawaii
prior to 1880 (Zimmerman 1958). Swezey reported the
species from “drier, lower” habitats, but does not
provide specific collection localities beyond a single
reference to a population in Olinda, Maui (Swezey
1907; Zimmerman 1958). Despite their early
documented widespread abundance, 100 years later the
species was declared extinct by the U.S. Fish and
Wildlife service, and other agencies (Gagne & Howarth
1982; IUCN 2007). Omiodes continuatalis and 13 other
Omiodes species were listed as “extinct or possibly
extinct” based on reviews of museum collections which
indicated that specimens had not been collected for

several decades. Non-target impacts of the biological
control agents introduced for the coconut and sugar
cane leafroller pest species were blamed for the
extinctions (Gagne & Howarth 1982). However, data
from surveys of more recent museum collections
completed in 2003 revealed extant O. continuatalis
populations (Haines et al. 2004). Specimens were
identified from light trap samples collected at five mesie
forest studv sites on both the leeward and windward

J

sides of Hawaii Island (Haines et al. 2004). In addition,
field collections completed in 2003 confirmed O.
continuatalis to be abundant at several localities on East
Maui (Haines unpublished data).

In the past, O. continuatalis larvae have been
collected from native pili grass Heteropogon contortus
(L.), as well as introduced species such as kikuyu grass
Pennisetum clandestinum (Hochst. ex Chiov.) and Hilo
grass Paspalum conjugation Berg. (Zimmerman 1958).
Like other species in the group, O. continuatalis larvae
are leaf-rollers, and use silk to bind plant foliage
together to create a refuge in which they feed and take
shelter. Early observations by Swezey describe how O.
continuatalis larvae typically feed near the base of their

12

Journal of the Lepidopterists’ Society

host plants, within the grass matrix or in leaf litter at the
soil surface (Zimmerman 1958). This behavior is
uncharacteristic of other Hawaiian Omiodes larvae,
which generally feed higher on host plant foliage,
fastening or rolling the tips of leaves together (Swezey
1907). Omiodes continuatalis larvae were also noted to
be physically distinct from larvae of other Omiodes
species, in that they attain a greater body width and
often exhibit pinkish-green coloration in contrast to the
monochromatic green coloration displayed by larvae of
other Omiodes species (Zimmerman 1958).

Unlike the sugarcane leafroller O. accepta (Butler,
1877), O. continuatalis is not considered a pest and
consequently few data have been collected to document
its life history. Omiodes continuatalis appear to share
similar habitat requirements, feeding habits and
physical characteristics with O. accepta (Zimmerman
1958), and initially seemed likely to have comparable
biology and development. The objective of this chapter
is to describe the complete life history of O.
continuatalis , providing new information on its biology,
ecology, general moqdiology and life history, and to
integrate recent findings with information gathered by
early naturalists, to create a document that can serve as
a reference for future research on the biology and
conservation of O. continuatalis.

Materials and Methods

We collected data for this laboratory and field study
from February 2005 to March 2008.

Insect rearing. We established laboratory colonies
of O. continuatalis using wild female moths captured
from UV light traps set on Kailiili Road in Kokomo,
Maui (484m) and at the University of Hawaii Kula
Agricultural Research Station (975m), Mani. Moths
were collected alive in plastic specimen vials, and held
at ambient temperature until being transferred into
rearing cages. Moths were easily identified using the key
to Omiodes species in Zimmerman (1958), as O.
continuatalis is a veiy distinctive species. Rearing cages
were composed of hand-made 61cmx46cmx31cm PVC-
framed sleeve cages covered with fine insect mesh
sleeves. Moths were fed a 30% honey/70% water
solution, and were provided potted sugarcane, pili grass
and/or Hilo grass, on which to oviposit. We obtained
young sugarcane starts from Hawaii Commercial and
Sugar (IICS) and Hawaii Agricultural Research Center
(I4ARC) fields, while pili grass and Hilo grass were
propagated from seed. These plants provided food and
served as host plants for colony larvae. The substrate in
each host plant pot consisted of a 50/50 blend of
Sunshine Mix 5® potting soil and vermiculite. All moth
colonies and host plants were housed in the University

of Hawaii at Manoa, Gilmore Hall greenhouses, and
University of Hawaii, Kula Agriculture Research Station
greenhouses. In Manoa, temperatures ranged seasonally
from a mean low of 18.7°C in the winter to a mean high
of 24.3°C during the summer (Manoa Lyon Arboretum
weather station), and plants were watered three times
each day. In Kula, temperatures ranged from a mean
low of 11.1°C in the winter 2006, to a mean high of
24.4°C during the summer 2006 (UI4 Kula weather
station), and plants were watered twice daily. We
collected additional wild O. continuatalis moths and
integrated them into laboratory colonies periodically to
replenish adult stock and maintain genetic diversity.

Biology and morphology. To elicit oviposition, we
isolated female moths in vials lined with wax-paper and
temporarily denied them access to a food source. These
stressful conditions commonly stimulate female moths
to oviposit. Eggs deposited in each egg mass were then
counted under a Leica dissecting microscope to
determine the total number present, and the minimum
and maximum number of eggs per mass. In order to
quantify mean egg size, we selected 50 eggs randomly
from 50 O. continuatalis egg masses, and measured
them at the widest point using an ocular micrometer in
a Wild Heerbrugg microscope.

We obtained data on O. continuatalis larval
development and moiphology by observing 100
individual larvae develop from egg to pupal stage.
Larvae were stored in individual 2-oz Solo® cups
(ventilated with 3-6 holes in the lid). Cups were cleaned
anti larvae provided with fresh sugarcane foliage eveiy
2-3 days. We measured the head capsule width of each
larva daily to document larval growth; head capsule
widths were measured at their widest point, as above for
eggs. We also recorded changes in larval moiphology
following each larval molt. Only head capsule widths of
larvae which survived from egg to pupal stage (n = 52)
were used to assess instar duration and size ranges. We
completed a one-way analysis of variance to document
variation in head capsule width by instar.

Behavior. Data relating to O. continuatalis larva!
behavior were collected based on observations of
individual larvae contained in laboratory colonies, or
from colony larvae deployed on potted host plants at
selected Maui field sites. We collected data
opportunistically, observing behaviors in the field during
larval deployment and retrieval, and in the laboratory
during daily colony maintenance and larval feedings.
Adult O. continuatalis were isolated and observed
individually to assess adult lifespan under laboratory
colony conditions.

Distribution. Field sites were selected to be
surveyed based on accessibility and the presence of O.

Volume 63, Number 1

13

continuatalis host plants. We collected Omiodes
continuatalis adults using two types of UV light traps:
sheet light traps and bucket light traps. Sheet light traps
consisted of an 18” UV light bulb placed in front of a
large, white bed sheet suspended vertically. Bucket light
traps were constructed with an 18” UV light bulb
attached to three 10.2cmx25.4cm clear plastic flight
intercept vanes, and secured on top of an 8 liter bucket
with bungee cords. Each bucket trap was baited with a
killing agent such as cyanide or a Hot Shot No-Pest
Strip®. Individual moths landing on sheet light traps
were collected alive, however insects collected from
bucket traps were always dead. We set at least one of
the two trap types at each field site to survey for ().
continuatalis adults. When an initial survey yielded
numerous O. continuatalis specimens, we revisited sites
to replenish and diversify laboratory colonies. Visual
surveys of and sweep netting for adult O. continuatalis
were also completed at field sites to complement data
collected from light trapping. To quantify total survey
effort, the number of nights and the number of traps
that were used, were multiplied to obtain the total
number of “trap nights” completed at each survey
location.

Results and Discussion

Biology. Omiodes continuatalis are highly attracted
to UV light, and both male and female moths were
regularly collected from light traps. In addition, O.
continuatalis appeared to be more attracted to UV light
traps than other Omiodes species. Once drawn to a
light, O. continuatalis adults frequently remained
around the light for long periods of time, in contrast to
the sedge-feeding species O. anastrepta Meyriek, 1899,
which are not as readily attracted to light traps, even
when adults are observed flying in close proximity to a
trap. Over the course of one year, adult O. continuatalis
were collected, on multiple occasions, from light traps at
Kokomo, Makawao, and Kula, on Maui (Table 1). Data
from these surveys indicate that O. continuatalis is
multivoltine, with more than three generations per year.
Considerable climatic seasonal variation exists at mid-
high elevation localities, so if O. continuatalis
populations are mutivoltine at these three mid-high
elevation field sites, it is likely that populations are also
multivoltine and not seasonal at low elevation sites,
where seasonal changes in temperature and rainfall may
be even less restrictive to O. continuatalis development.
Based on mitochondrial genetic data, O. continuatalis
populations do not exhibit much structure, either within
or among islands (Haines, unpublished data), suggesting
that moths are strong dispersers. Because of this, we do
not suspect that populations on islands other than Maui

exhibit very different morphology or life history
characteristics from those observed in our colonies.

Laboratory colonies of O. continuatalis were
maintained continuously from July 2005-December
2005, and from March 2006-March 2008. In laboratory
colonies, the moths were most successfully reared on
sugarcane host plants, as compared with pili grass and
hilo grass, which the moths were previously
documented feeding on in the field. At one point during
rearing of the colony, some larvae escaped from cages
and were discovered feeding on Canavalia pubescens ,
Hook. & Arn. (Fabaceae) an endemic Hawaiian legume
which was being grown in the same greenhouse as food
for another caterpillar species (O. monogona Meyriek,
1888). The O. continuatalis larvae were caged with C.
pubescens, and surprisingly, multiple generations were
reared on the plant. There are no previous records of
this grass-feeding species surviving on a dicot host, yet it
survived through four generations on C. pubescens with
no obvious moi-phologieal or reproductive
disadvantages. The success of the moth on this host
plant over multiple generations indicates that its host
range in the wild might be broader than previously
expected, although opposition preference may be more
important than nutritional suitability as a determinant of
host range. While the utilization of this host plant was
unexpected given the documented life history of the
species, it may be explained by recent molecular
analysis of the genus. Phylogenetic analysis based on
mitochondrial DNA shows that O. continuatalis is the
sister species, and the closest relative, of O. monogona,
the sole Hawaiian Omiodes species which feeds on
fabaceous host plants (Haines unpublished data). Other
tropical Omiodes species are documented to feed on
Fabaceae, thus this dietaiy shift may reflect a reversion
to an ancestral host plant retained in the evolutionary
‘memory’ of some Hawaiian Omiodes. Evolutionary
memory is the idea that larvae may retain an ability to
feed on ancestral hostplants on which they are no longer
found in nature (Janz & Nylin 1998). This discovery has
important implications not only for the conservation of
the species, but also for the evolution of host plant shifts
in the genus.

Adult O. continuatalis (Fig. 1) lived a maximum of 18
days under laboratory conditions. In cages, O.
continuatalis females were observed ovipositing on both
the upper and lower surfaces of host plant vegetation,
and females did not avoid ovipositing on material such
as wood, plastic, wax paper and screen mesh. Egg
masses ranged in size from a single egg, to a maximum
of 73 eggs, with a mean of 7.25 (± 0.81 S.E.) per
individual egg mass (n = 115). A limited survey of O.
continuatalis females (n = 33) indicated that gravid

14

Journal of the Lepidopterists’ Society

Fig. 1. Adult Omiodes continuatalis moth.

females can lay more than 400 eggs over their lifetime,
and up to 122 eggs in a 24 hour period. Egg masses are
composed of even rows or clusters of Battened eggs,
slightly overlapping one another (Fig. 2). Eggs have a
mean width of 1.22mm (± 0.025) (n = 50), are light
yellow in color, and exhibit fine surface reticulations and
an iridescent sheen. Omiodes continuatalis eggs took
four days to hatch at 32°c, and eight days to hatch 22°c.

Morphology. Swezey broadly described the life
history and physiology of O. continuatalis (summarized
by Zimmerman 1958). Zimmerman (1978) later
provided limited descriptions and illustrations of larval
moiphology, including a larval key to most Hawaiian
Omiodes spp. However, no detailed research has been
reported that fully characterizes the life history and
larval stages of O. continuatalis.

Based on head capsule width measurements from 52
individuals, we observed O. continuatalis larvae
transitioning through seven to nine developmental
instars (Fig. 2). In a previous study, O. continuatalis
larvae were observed transitioning through as many as
10 instars prior to pupation (King & Rubinoff 2008).
Analyses of variance of head capsule widths by instar
indicates that mean head capsule widths for instars 1
through 9 are significantly different from one another
(Figure 2; df = 8; F = 1823.68; P = <0.001). Due to
considerable overlap in head capsule widths in each
instar (Fig. 3), it is difficult to distinguish discrete larval
instars at any stage of development based on
measurements alone.

Upon eclosion, larval head capsules are reddish-

brown in color and have a mean width of 0.35mm
(range: 0.32-0.38). The reddish-brown coloration
persists throughout the entire larval stage. Unlike the
larvae of other Hawaiian Omiodes species which
maintain bright green body coloration (Swezey 1907),
O. continuatalis larva! coloration is comparatively less
vivid. The abdomen is a gradient of off-white to light
olive, and once larvae begin feeding they acquire the
approximate pigment of the vegetation they are
digesting. Larvae may molt to the second instar (mean:
0.45mm, range: 0.40-0.50) and the third instar
(0.64mm, 0.50-0. 74mm) after three to five days. In the
third instar, larvae develop a single black head spot on
each of the head capsule lobes, as well as two black
spots on the prothoracic shield. O. continuatalis larvae
continue to molt every three to five days (4th: 0.92mm,
0.66-1. 10mm; 5th: 1.28mm, 0.90-1. 42mm; 6th:
1.78mm, 1.3-1. 96mm), growing in size and developing
more intricate patterns of melanization on the
prothoracic shield, as well as tubercles on the 2nd and
3rd thoracic segments and the 7th-10th abdominal
segments. During the penultimate instar (6th; 7th:
2.32mm, 1.8-2. 52mm; 8th: 2.84mm, 2. 34-3. 56mm), O.
continuatalis larvae acquire a faint pink tint, while the

Fig. 2. Omiodes continuatalis larvae exhibit changing pat¬
terns of melanization on the prothoracic shield during nine de¬
velopmental instars. A. Cluster of eggs. B. First instar. C. Sec¬
ond instar. D. Third instar. E. Fourth instar. F. Fifth instar. G.
Sixth instar. H. Seventh instar. I. Eighth instar. J. Ninth instar.

Volume 63, Number 1

15

head, thoracic and abdominal spots are reduced. Molts
to the final instar (7th; 8th; or 9th: 3.19mm,
2. 86-3. 50mm) also occur in three to five days. At this
stage, head spots disappear completely, and the thoracic
and abdominal segments become veiy pink.
Immediately prior to pupation, O. continuatalis larvae
become pale and begin to shrink in size to form pre¬
pupae. It is common for lepidopterous larvae to change
color prior to pupation, when the procuticle increases in
area in the cuticle column above the epidermal cells
(Chapman 1998). Pre-pupae develop into pupae over
1-2 days, and remain as pupa for 13-17 days. In total,
immature O. continuatalis development was 36-57
davs.

J

Behavior. Early instar larvae feed on the surface and
inner tissue of host plant foliage, often near the midrib
of the leaves. Omiodes continuatalis larvae cannot be
characterized as gregarious, like the larvae of O.
blackburni (Butler, 1877) which share the same shelters
throughout most of their development, however early
instar O. continuatalis larvae often feed in close
proximity to one another. Aggregations of larvae on host
plants especially in the first and second instars are a
natural result of larvae emerging from eggs laid in
clusters. General observations from this research
suggest that first instar O. continuatalis larvae are more
likely to survive when feeding together. First instar
larvae that were separated, and made to feed
independently, frequently failed to feed successfully and
progress to the subsequent instar. The reasons for this
increased mortality are not clear, but may involve the
cumulative benefit of aggregated larvae feeding
cooperatively during early instars. Research indicates
that cooperative feeding behavior has many advantages
for lepidopterous larvae, including increasing feeding
efficiency on physically tough host plant material (Clark
& Faeth 1997). It is possible that O. continuatalis larvae
exhibit this type of cooperative feeding due in part to
the fact that they have evolved on grasses, which are
tougher than many dicot host plants and contain higher
concentrations of silica (Falin 1982). When O.
continuatalis larvae reach the third instar they continue
to consume the surface and inner tissue of host plant
leaves, but fed independently from other larvae more
frequently than in the early stages. Third instar larvae
began to fasten silk threads across the foliage and/or
roll-up foliage to create a protected shelter in which to
retreat. During the fourth instar and/or when larvae
attain a head capsule width of 1mm, they begin to feed
on the entire leaf structure. Some larvae were observed
to feed predominantly inside their shelters, while others
migrated back and forth between their shelter and
feeding areas elsewhere on the host plant. Fate instar

larvae primarily feed outside the shelters they construct.
Feeding patterns were easily discerned based on areas
of feeding damage on field deployed host plants, and
host plant material provided to isolated O. continuatalis
larvae.

In colony, O. continuatalis larvae occupied all
available space on the host plants provided, from the
sheaths at the veiy base of the sugarcane, to the top of
each blade. In contrast, when placed at lower densities
on host plants in the field, larvae generally stayed on the
green sugarcane foliage on the mid to upper portion of
the plant. When foliage was available at the base of the
plant, larvae often constructed shelters there, using both
fresh and dead vegetation as materials. This behavior is
consistent with early observations of larval feeding
habits made by Swezey (Zimmerman 1958). Additional
observations by King & Rubinoff (2008) indicate that
some O. continuatalis larvae display a more unusual
habit, burrowing beneath the soil surface at the base of
their host plants for shelter. In some cases, larvae were
recovered up to 14cm below the soil surface. This
fossorial behavior was observed in early, middle and late
instar larvae, and is not pupation behavior.

Another behavior we observed during laboratory
colony rearing was larval cannibalism. O. continuatalis
larvae, specifically late instars, were observed feeding on
other live O. continuatalis larvae. This behavior was
more common on occasions when larval density was
high and host plant resources were decreasing, but still
available. Most frequently, late instar larvae were
observed feeding on middle instar larvae. In one case a
ninth instar larva was found feeding on another ninth
instar larva. In all cases, the larvae being consumed did
not appear to fight or tiy to avoid the predation, but
remained stationary during the attack. The cannibalistic

Instar

Fig. 3. Boxplot of headcapsule widths of Omiodes continu¬
atalis larvae by instar. Medians and inter-quartile ranges are rep¬
resented for each instar, and box widths are proportional to sam¬
ple size.

16

Journal of the Lepidopterists’ Society

Table 1. Locations surveyed for O. continuatalis on the main Hawaiian Islands.

Elevation

Trap

O. continuatalifi

Island

Location

(m)

Method

Date

nights

present

Surveyorfs)

Kauai

Pu'u Ka Pele Forest Reserve

855

light trap

8/6/08

1

No

W Haines

Koke'e State Park, Camp Sloggett

1060

light trap

5/16/05, 10/6/07

3

No

W Haines, C. King

Mohihi Road Koke'e State Park

1110

light trap

6/2/04

2

No

W. Haines

Koke'e State Park, Koke'e Museum

1130

light trap

8/7/07

2

No

W Haines

Na Pali Kona FR, Pihea trail

1140

light trap

8/7/07

i

No

W Haines

Koke'e State Park, Nualolo Trail

1170

light trap

8/8/07

i

No

W I Lines, M. Medics

Na Pali Kona FR, Alakai Swamp Trail

1180

light trap

8/7/07

3

No

King, Haines, Rubinoff

Na Pali Kona FR, Pihea trail

1200

light trap

5/18/05, 7/23/06,
8/18/06

4

No

Eiben, Haines, Rubinoff

Koke'e State Park,

Near Awaawapuhi Trail

1245

light trap

6/3/04

1

No

W Haines

Na Pali Kona FR, Kalalau Lookout

1255

light trap

8/17/06

2

No

|. Eiben, D. Rubinoff

Oahu

Aina Haina, Niuiki Circle

2

light trap

3/24/05

i

No

C. King

Mokuleia Beach

2

light trap

5/1/05

i

No

W Haines, C. King

Sandy Beach

3

light trap

3/6/05. 6/20/05

2

No

W Haines, C. King

Ewa, open grass lot

15

light trap

5/21/05

i

No

C. King

Diamond Head Crater, wetland

80

light hap

4/3/05, 2/15/08

4

No

W Haines

Palolo Valley

80

light hap

2/25/08, 2/26/08.
2/27/08, 2/28/08

4

No

W Haines

HARC Maunawili

160

light trap

5/26/05, 6/22/05

3

No

C. King

Lyon Arboretum, sugarcane patch

228

light hap

1/27/07

2

No

C. King

Lanipo Trail

240

light hap

5/29/05

i

No

W Haines, C. King

Dole pineapple field

256

light trap

3/12/05

i

No

Haines, King, Vorsino

I Iawaiiloa Ridge Trail

425

light hap

6/14/06

i

No

J. Eiben, W Haines

Wiliwilinui Trail, access

440

light trap

6/6/06

i

No

W Haines

Wa'ahila Ridge

475

light trap

6/8/06

i

No

W Haines

Kahanahaiki Valiev

605

light trap

2/24/07

i

No

C. King

Mt. Tantalus

613

light hap

6/3/05, 6/22/05

2

No

W Haines, C. King

Wiliwilinui Trail, Summit

731

light hap

6/24/06

i

No

W Haines

Ko'olau Moutains, Konaliuanui Trail

762

light trap

6/1/06

i

No

W Haines

Palikea Trail

777

light trap

5/15/06

i

No

W Haines

Mokuleia FR, ;dong Mt, Kaala Road

1100

light trap

6/15/07

2

No

W Haines

Waianae-Kai FR, near Ka'ala NAR boundary

1190

light hap

6/15/07

i

No

W Haines

Ka'ala NAR, hog

1200

light hap

6/15/07

i

No

W Haines

Ka'ala NAR. DOFAWshed

1210

light trap

6/15/07

i

No

W Haines

Maui

Kanaha Beach Park

3

light trap

3/20/05

i

No

C. King

Maliko Gulch sugarcane field

76

light trap

6/1/05, 7/13/06

2

No

C. King

West HCS sugarcane field

137

light trap

6/6/06

1

No

C. King

Iao Vdley

183

light trap

6/5/06

1

No

C. King

Waiehu Valley Trail

183

light hap

7/6/06

1

No

C. King

Haliimaile sugarcane fiekl

274

light hap

6/1/05

1

Yes

C. King

University of Hawaii Agricultural Park

305

light trap

6/1/06

1

No

C. King

Waihe'e Ridge Trail

305

\ismd survey

6/8/05

0

Yes

W Haines

2955B Kaili'ili Rd, Kokomo

484

light trap

3/19/05“

15

Yes

W Haines, C. King

Pu'u Kukui Watershed Kaluanui Valley

485

light hap

45/06

2

No

W Haines

Waihe'e Ridge Trail

670

light hap

6/8/05

i

No

W Haines

“dates =3/19/05, 5/25/05, 6/1/05, 7/19/05, 9/3/05, 9/23/05, 2/8/06. 3/7/06, 4/18/06, 7/8/06, 3/9/07, 4/19/07, 5/18/07, 8/17/07, 2/6/08

Volume 63, Number 1

Table 1. Continued

Island

Location

Elevation

(ni)

Method

Date

Trap

nights

O. continuatalis
present

Surveyor(s)

Maui

Makawao Forest Reserve, Site 1

750

light trap

3/23/05

1

No

W. Haines

Makawao Forest Reserve, banana patch

762

light trap

6/19/06,7/14/06,

7/15/06

8

Yes

W. Haines

Makawao Forest Reserve, Site 5

825

light trap

.3/23/05

1

Yes

W. Haines

Kipahulu Valley Delta Camp, HALE

860

light trap

1/15/04,9/14/04

4

No

W. I Iaines, D. Rubinoff

Makawao Forest Reserve

914

light trap

3/19/05,5/25/05,

6/1/05

4

Yes

W. Haines, C. King

Makawao Forest Reserve, Site 2

915

light trap

3/23/05

1

Yes

W. Haines

Makawao Forest Reserve, Site 6

930

light trap

3/23/05

1

Yes

W. Haines

UH Kula Field Station

975

light trap

'3/9/07°

5

Yes

C. King

Makawao Forest Reserve, Site 4

1050

light trap

3/23/05

i

No

W Haines

Waipoli Road, Kula

1065

light trap

6/20/04

2

Yes

W Haines

Haleakala Ranch

1280

visual survey

623/06

0

Yes

C. King

Makawao Forest Reserve

1280

light trap

63/06, S/17/07

2

Yes

W. Haines, C. King

Makawao Forest Reserve, Site 3

1330

light trap

3/23/05

i

Yes

\ Y. Haines

Kula FR

2073

light trap

64/06

i

No

C. King

Molokai

Moomomi Preserve

10

light trap

7/5/05

i

No

W. Haines

Molokai Forest Reserve, TNC Barracks

915

light trap

7/7/05, 1230/05

3

No

W. Haines

Kamakou Preserve, Kamakou Flats

1105

light trap

5/1 604

2

No

W. Haines

Kamakou Preserve, Puu Kolekole

1130

light trap

.5/19/04,7/7/05

2

No

W. Haines

Kamakou Preserv e, Pepeopae Bog

1240

light trap

5/19/04, 7/605

2

No

W. Haines, D. Rubinoff

Lanai

Lanai Game Management Area

.365

ligjit trap

7/3/05

i

No

W. Haines

Kanepuu Preserve

520

light trap

7/2/05

i

No

W. Haines

Kanepuu Preserve

.544

light trap

3/608

2

Yes

D. Rubinoff, R Schmitz

Munroe Trail

1000

light trap

7/3/05

i

No

W. Haines

Kahoolawe

Base Camp, south of Kanapou Bay

5

light trap

1/14/06

i

Yes

S. Meyers

Big Island

Pahala sugarcane field

292

light trap

610/05

i

No

C. King

Honokaa sugarcane field

303

light trap

69/05

2

Yes

C. King

Glenwood, Omega Rd.

700

light trap

2605

2

No

W Haines

Big Island

Glenwood Near Kahaualea NAR

700

light trap

2605, 7/30/06

6

No

W. Haines

Ka'u, Kaiholena

770

light trap

7/31/06

2

No

W Haines, J. Giffin

Ola'aFR

822

light trap

25/05

i

No

W. Haines

Ola'a FR

975

light trap

25/05

i

No

W. Haines

Ka'u, Kaiholena

975

light trap

7/31/06

i

No

W. Haines, J. Giffin

Kahuku Ranch, HAVO

1005

light trap

5/18/06

3

No

W. Haines, D. Rubinoff

Volcano Village

1128

light trap

6605, 11/2605

2

No

C. King

Ola'a Tract, HAVO

1150

light trap

.5/17/06

4

No

W Haines

Escape Rd/Thurston. HAVO

1158

light trap

4/1605,629/05,

5/18/06

3

No

W. Haines, C. King

Kilauea Field Station, HAVO

1158

light trap

25/05

1

No

W. Haines

Kipuka Puaulu, HAVO

1200

ligjit trap

10/15/04

1

Yes

W. Haines

Ola'a Tract, Pu'u Unit

1300

light trap

5/17/06

1

No

W. Haines, D. Rubinoff

Waildi

1432

light trap

6/11/05

1

No

C. King

Mauna Loa Strip Roacl HAVO

1500

light trap

10/14/04

1

Yes

W Haines

Kona Forest Unit. Hakalau Natl Wildlife Refuge

1615

light trap

4/1205

2

No

C. King

Keamoku flow; H AM )

1707

light trap

4/15/05,62605

3

No

C. King

° dates = 3/9/07, 4/7/07, 4/19/07, 5/10/07, 5/18/07

18

Journal of the Lepidopterists’ Society

Fig. 4. Map of current distribution of Omiodes contimiatalis in the Hawaiian Islands, based on recent field survey data.

larvae began their attacks at the tenth abdominal
segment, working forward and consuming the entire
contents of the abdomen. The only structures not
consumed by the cannibalistic larvae were the head
capsule and the cuticle. Thus, the end product of such
an encounter resembled a shed head capsule and cuticle
following a developmental molt. Perhaps the behavior
has never been documented in Omiodes larvae because
Hawaiian Omiodes species have not been reared in
colony prior to this investigation. Cannibalism,
prompted by competition for limited resources such as
host plant resources and over-wintering sites, has been
documented in non- Hawaiian crambid species (Baskauf
2003), so it is possible that O. contimiatalis larvae may
exhibit the behavior in the natural environment under
high density conditions.

Distribution. We completed more than 166 trap
nights of surveys at 85 field sites for populations of O.
contimiatalis across the Hawaiian Islands (Table 1).
While previous collecting records show O. contimiatalis
to be present in mostly diy or mesic habitats
(Zimmerman 1958; Haines et al. 2004), our surveys

show that O. contimiatalis populations are present in
diy, mesic and also wet habitat types (Table 1) (Fig. 4).
On Hawaii Island, O. contimiatalis was collected from
three mesic habitats: Kipuka Puaulu (1195m) and
Manna Loa Strip Road (1492m) on Mauna Loa in
Hawaii Volcanoes National Park, and a remnant
sugarcane field near the town of Honokaa (303m). All of
these sites receive a mean annual rainfall (MAR) of
1500-2000mm. On Maui, O. contimiatalis were
collected from seven distinct field sites: Waihee Ridge
trail (305m). Haliimaile/Haleakala Hwy sugarcane field
(274m), Kokomo (484m), UH Kula field station (975m),
Makawao Forest Reserve (914m, 1280m), and

Haleakala Ranch (1280m). With the exception of
Waihee Ridge, which is located on West Maui, all of the
field sites are located on East Maui. The two Makawao
Forest Reserve sites and the Haleakala Ranch site are
wet habitats (2QQ0-3000mm MAR). Waihee is a mesic
habitat (1500mm MAR), as is Kokomo (1500-2000mm
MAR), while Haliimaile/Haleakala Hwy sugarcane field
and the UH Kula field station are dry habitats, due to
their position in the rain shadow of East Maui (MAR

Volume 63, Number 1

19

<750mm). O. continuatalis were also collected on the
islands of Kahoolawe and Lanai. Both of these islands
are exceedingly dry (630mm MAR) as they are also
located in the rain shadows of Maui and Molokai.

Based on survey data of O. continuatalis , it is clear
that this species can persist in diverse habitat types
ranging from diy to wet. O. continuatalis also occurs in
habitats exhibiting varying levels of alteration and
disturbance. The species is present in predominantly
native forest habitat (e.g. Mauna Loa, Makawao Forest
Reserve), as well as areas where native habitat has been
converted to pastureland (e.g. Haleakala Ranch), and/or
agricultural land (e.g. Honokaa and Haliimaile
sugarcane fields). Their persistence appears possible so
long as the habitat contains sufficient host plant
resources, in the form of native or introduced grasses, to
support larval development. Given their widespread
distribution, their palpable ability to adapt to significant
changes in habitat, and their ability to utilize invasive
grasses as host plants, O. continuatalis would seem an
unlikely candidate to face high risk of declines and/or
extinction. Nevertheless, the current distribution of O.
continuatalis on the islands of Maui, Kahoolawe, Lanai
and Hawaii represents a significant range reduction for
a species which was previously common, but is now
extirpated from, Kauai, Oahu, and Molokai.

It is possible that the reduction in distribution is not
as great as these survey data might indicate. Even small
variations in season, moon phase, air temperature,
precipitation and wind speed can affect light trap catch
on a given day. Therefore, it is possible that O.
continuatalis may have gone undetected at sites where
light trapping occurred on only one occasion. The
Kailiili Road study site in Kokomo, Maui was surveyed
frequently due to the consistent O. continuatalis trap
catch. Even so, there were at least two occasions where
O. continuatalis adults were not observed.
Furthermore, during surveys on Hawaii Island, O.
continuatalis was not collected from light traps at the
Keanroku How on Mauna Loa, where the species has
been previously recorded (Haines et al. 2004). Thus, it
is possible, that O. continuatalis are present at
additional field sites on Maui, Kahoolawe, and Hawaii
Island. Accordingly, the species may remain undetected
on other islands despite extensive negative data from
these survey efforts. This hypothesis seems more
plausible at sites where traps were only set-up for 1-2
trap nights, as opposed to field sites like Kokee State
Park or the Alakai Swamp on Kauai where more than 18
trap nights have cumulatively been completed, and no
O. continuatalis have yet been detected.

Potential factors influencing the decline and
restricted distribution of O. continuatalis were not

quantified under the scope of this research. Given the
persistence of the moth in severely degraded habitats, it
seems unlikely that habitat destruction or habitat
alteration alone could be significant limiting factors for
the species. There are habitats on the islands of Kauai,
Oahu and Molokai, both pristine and disturbed, which
we presume contain suitable resources to support O.
continuatalis , and yet the species has still not been
“rediscovered” on these islands. If habitat destruction
and alteration are not consistently affecting the
persistence of this species across the Hawaiian Islands,
then what other factors may be contributing to their
declines? The intentional and accidental introduction of
natural enemies into the Hawaiian Islands has been
viewed suspiciously by entomologists for over a centuiy
(Perkins 1897; Zimmerman 1958; Howarth 1983;
Henneman & Memmott 2001), and some of these
parasitoids and predators appear to suppress
populations of other Omiodes species (Zimmerman
1958; Bess 1974). Additional research is currently
underway to examine the mortality factors, specifically
parasitism, which may be affecting O. continuatalis
populations. However, the individual and compound
effects of introduced predators, such as ants and
spiders, are also likely mortality factors, and their effects
on Omiodes species remain to be investigated.

Acknowledgements

We thank A. Arcinas (Hawaii Agricultural Research Center)
and M. Nakahata (Hawaii Commercial & Sugar Company) for
providing plant material, and D. Oka and G. Otani (University of
Hawaii Kula Agricultural Research Station) for field preparation
and greenhouse assistance. J. Patu provided essential laboratory
assistance. Access permits were granted by S. Meidel (Haleakala
Ranch), G. Shishido and B. Gagne (Department of Land and
Natural Resources, Division of Forestry and Wildlife), and C.
Haines and G. Haines (Pacific Produce). Special thanks to M.
Wright, C. Hooks, J. Eiben, L. Leblanc and L. Kaufman (Uni¬
versity of Hawaifi, Department of Plant and Environmental Pro¬
tection Sciences) for valuable input on research methods, and
also W. Haines (University of Hawai i, Department of Plant and
Environmental Protection Sciences) for access to unpublished
data. Anonymous reviewers provided valuable feedback on this
publication.

Literature Cited

Baskauf, S. f. 2003. Factors influencing population dynamics of the
southwestern corn borer (Lepidoptera: Crambidae): A reassess¬
ment. Environmental Entomology 32(5): 915-928.

Bess, H. A. 1974. Hedylepta blackbumi, a perennial pest of coconut
on wind-swept sites in Hawaii. Proceedings of the Hawaiian En¬
tomological Society 21: 343-353.

Chapman, R. F. 1998. The insects, structure and function. Cam¬
bridge, Cambridge University Press.

Clark, B. & S. Faeth. 1997. The consequences of larval aggregation
in the butterfly Chlosyne lacinia. Ecological Entomology 22:
408-415.

Fahn, A. 1982. Plant anatomy, 3rd ed. New York, Pergamon Press.
Funasaki, G. Y.. P Y. Lai. L. M. Nakahara, J. W. Beardsley, & A. K.

20

Journal of the Lepidopterists’ Society

Ota. 1988. A review of biological control introductions in Hawaii:
1890 to 1985. Proc. Hawaii. Entomol. Soc. 28: 105-160.

Gag^e, W. C. & F. G. Howarth. 1982. Conservation status of en¬
demic Hawaiian Lepidoptera. Proc. 3rd Congr. eur. Lepid., Cam¬
bridge.

Haines, W. P, J. Giffin, & D. Foote. 2004. Rediscovery of five
species of Omiodes Guenee (Lepidoptera: Crambidae) on
Hawaii Island. Bishop Museum Occasional Papers 79: 45^49.
Henneman, M. L. & J. Memmott. 2001. Infiltration of a Hawaiian
community by introduced biological control agents. Science 293:
1314-1316.

Howarth, F. G. 1983. Classical biocontrol: Panacea or Pandora's box.

Proc. Hawaii. Entomol. Soc. 24(2): 239-244.

1UCN. 2007. IUCN - The World Conservation Union.

Janz, N. & S. Nylin. 1998. Butterflies and plants: A phylogenetic
study " Evolution 52(2): 486-502.

King, C. & D. Rubinoff. 2008. First record of fossorial behavior in
Hawaiian leafroller moth larvae, Omiodes continuatalis (Lepi¬
doptera: Crambidae). Pacific Science 62(1): 147-150.

Perkins, R. C. L. 1897. The introduction of beneficial insects into the
Hawaiian Islands. Nature 55: 499-500.

Swezey, O. II. 1907. The sugar cane leaf-roller ( Omiodes accepta):
With an account of allied species and natural enemies. Honolulu,
Experiment Station of the Hawaiian Sugar Planters Association:
1-59.

Zimmerman, E. C. 1958. Insects of Hawaii, Lepidoptera: Pyraloidea.

Honolulu, University of Hawaii Press.

_ . 1978. Insects of Hawaii, Microlepidoptera I. Honolulu, Uni¬
versity of Hawaii Press.

Received for publication 23 May 2008; revised and accepted 9 Dec
2008.

Volume 63, Number 1

21

journal of the Lepidopterists’ Society
63(1), 2009, 21-26

A NEW SPECIES OF MORRI SONIA (NOCTUIDAE) FROM
SOUTHEASTERN NORTH AMERICA

J. Bolling Sullivan

200 Craven St., Beaufort, North Carolina 28516; e-mail: sullivanl4@earthlink.net

AND

James K. Adams

Department of Biology, Dalton State College, Dalton, Georgia 30720; e-mail: jadams@daltonstate.edu

ABSTRACT. We describe Morrisonia triangula. (Noctuidae: Hadeninae: Orthosini) from material collected from Virginia to Texas. Illus¬
trations of adults and their genitalia are provided. The generic placement is discussed.

Additional key words: Achatia, Egira, Himella, Orthosia

For at least 40 years collectors in the southeastern
United States have been aware of an undescribed
species of hadenine provisionally placed in the genus
Morrisonia bv Jack Franclemont. Apparently,
Franclemont never collected the species but was aware
of it from material sent to him for identification by
Charles Kimball prior to the latter’s publication of the
Lepidoptera of Florida in 1965. Kimball (1965)
mentions 5 specimens collected in Escambia Co.,
Florida, 4 in April of 1961 and one in May of 1962.
Additional material, much of which is in the U. S.
National Museum, was collected in the period from
1960-80 in Virginia, the Carolinas, Louisiana and Texas
by several eastern collectors. Kimball (1965) indicated
that Franclemont would describe the species but a
manuscript was never completed. In this paper we
describe the new species and discuss some of the
current taxonomic problems posed by the “genus”
Morrisonia.

Materials and Methods

Specimens were examined from the National
Museum of Natural History, Washington, DC (USNM)
and Furman University, Greenville, SC (FU). Data
were provided by Steve Roble on material in the
Virginia Department of Natural Resources (VDNR),
Richmond, VA; by Steven Hall on specimens in the
North Carolina Heritage Program (NCIIP), Raleigh,
NC; and by Richard Brown on material in the
Mississippi State Museum (MSM), Starkville, MS. Data
were also provided from the personal collections of
James K. Adams; Vernon Brou, Abita Springs, LA;
Chuck Ely, Nacogdoches, TX; living Finkelstein,
Atlanta, GA; Howard Grisham, Maysville, AL; Ed
Knudson, Houston, TX; Hugo Kons, Gainesville, FL;

Tom Neal, Gainesville, FL; Brian Scholtens,
Charleston, SC; Jeff Slotten, Gainesville, FL; and J.
Bolling Sullivan.

Genitalia were prepared by digestion in 10%
potassium hydroxide, dissected in water, and
photographed with a Nikon Coolpix 995 camera
attached to a Nikon SMZ800 microscope. Additional
photographs were provided by Donald Lafontaine (see
Lafontaine 1998). Recently collected specimens were
sent to Paul Hebert at the University of Guelph for
barcode COl analyses (Ratnasingham & Hebert, 2007).

Morrisonia triangula sp. nov. Sullivan & Adams

(Figs. 1-2)

Description. Head: Antennae fasciculate, setae exceeding
shaft diameter at base in male, shorter than shaft diameter in fe¬
male; scape straw with rusty scales intermixed; antennal dorsum
scaled with alternating straw and rusty bands. Palps upturned,
slightly longer than eye diameter, tan, dark brown and reddish
scaled. Frons straw above tongue, short scaling but dorsal half of
frons protruding with ventral part deep maroon, dorsal part
straw, tan with reddish scales; a distinct dark line visible across
upper part of frons. Area lateral to eye blackish scaled, tan scales
below tongue. Thorax: Neck with a mixture of reddish and straw
scales, largely erect. Collar a mixture of white, straw, tan and
deep maroon scales forming a series of horizontal bands; scaled
tuft behind collar. Tegulae reddish brown with some white
scales; scaling underneath mostly white. Legs with reddish scal¬
ing dorsallv, brown with cream scales ventrallv; single pair of
spurs on L2, two pairs on L3. Abdomen: two reddish dorsal scale
tufts on first two segments, 2nd tuft small. Dorsum of abdomen
tan with scattered reddish scales; underneath more reddish but
similar.

Forewing: Pattern without obvious cross lines, dominated by
a black basal and anal dash and a black triangular wedge at mid
costa; whitish scaling in costal area before wedge, distal to
wedge are 5 whitish costal spots, the most distal the largest;
cream elliptical mark between dashes, marginal line black, scal¬
loped; anal dash with smaller dash on costal side, whitish scaling
on vein along anal dash; wing base above and below basal dash

Journal of the Lepidopterists’ Society

darkened; orbicular and reniform spots lightly outlined, filled
with the surrounding light gray ground color of the forewing; or-
bicular oblong, posterior to triangular black wedge; reniform
kidney shaped, just distal to triangular wedge. Hindwing : Fuscus
with marginal black band and tan marginal scales forming fringe.
Underside: With distinct discal spots; perceptible PM (post-
medial) line. Reddish on marginal half of forewing; black scaling
on veins distal to PM line giving a striated appearance; basal half
of forewing with blackish scaling. Hindwing with similar scaling,
black scales densest on costal edges of both wings.

Male genitalia (Fig. 6): Valvae symmetrical, extending beyond
uncus when viewed laterally, flared toward apex; costal region
sclerotized, saccular region membranous; digitus arising from
costa and curving ventrally well beyond saccular edge; clasper
C-shaped, deriving from well developed medial plate; flared api¬
cal portion of valva setose with corona; uncus gently tapering
and arching ventrally, slender, subbasal setae on dorsal side;
gnathos and socii absent; anal tube slightly sclerotized ventrally;
tegumen arms broad, forming inverted V; penieillum well devel¬
oped and located on pointed flap at posterior end of tegumen;
vinculum V-shaped, well developed; pleurite slender, curving up
to dorsal side of tegumen; juxta flaring posteriorly, with deep V
in middle, posterior edges rounded; aedeagus rounded at base,
sclerotized and tapering to a gentle curv ing apex; vesica with 3
or more spirals, basal spiral with longitudinal row of short cor-
nuti, largest cornuti in short row above long, longitudinal row,
diverticula on first and second spirals, ribbon-like band of em¬
bedded cornuti beginning on second spiral, cornuti patch (6-10
long cornuti) ventral to second diverticulum, distal end of vesica
membranous.

Female genitalia (Fig. 9); Lightly sclerotized anal papillae, an¬
terior and posterior hypophvses equal in length; ostium a simple
flared opening, ductus sclerotized to bursa; accessory bursa aris¬
ing ventrally, spiralled with sperm duct arising near tip: bursa a
well defined sack with 4 evenly spaced longitudinal signa ex¬
tending 2/3 toward anterior end, walls of bursa striated.

Types: Holoti/pe: Male (Fig. 1): U. S. A. North Carolina:
Craven Co., Croatan National Forest, Road 169, 2 iv. 2000, J. B.
Sullivan. Paratypes: (11 males, 8 females) - North Carolina: Col¬
lection data same as holotype (2 males); Craven Co., Croatan
National Forest, Road 147, 1 1 iv. 2000 (2 males, 1 female); 23 iv.
2006 (2 males, 1 female); 12 v. 1996 (1 female); 6 vi. 2006 (1 fe¬
male); Croatan National Forest, Road 3046, 26 vi. 1998 (1 fe¬
male) J. B. Sullivan. Georgia: Lumpkin Co., Blood Mountain,
.25 mi. S of Neel’s Gap, 3100', 27 v. 2006 (1 female). Louisiana:
St. Tammany Parish, sec 24T6SR12E. 4.2 mi. NE of Abita
Springs, 30 iii. 1998 (1 male); 10 iv. 1998 (1 male); 25 iii. 2002 (1
male); 28 iii. 2004 (1 female). Texas: Tyler Co., Big Thicket Na¬
tional Preserve, Turkey Creek Unit, 15—16 iv. 1994 (2 males); 8
iv. 1995 (1 female). The holotvpe will be deposited into the U. S.
National Museum and paratypes into the Canadian National
Collection, British Museum, Florida State Collection of Arthro¬
pods, Texas A&M University, and the Texas Lepidoptera Survey
(with Charles Bordelon and Edward C. Knudson, Houston, TX).

Etymology. The name triangula is an adjective
describing the distinctive triangular mark along the
forewing costa. We suggest Triangle-barred Mor¬
risonia as the common name.

Diagnosis. Morrisonia triangula is abundantly
distinct in maculation and genitalia from other species
of Morrisonia and from other hadenine species
encountered in the Eastern United States. In pattern it

resembles less heavily marked forms of M. evicta (Grt.)
but is larger, has the distinct costal triangle and except
perhaps for a few mountain localities, is allopatric. If the
two species do fly together, one would expect M. evicta
to he on the wing earlier in the spring. M. triangula
more closely resembles M. latex (Gn.) (Figs 3, 4) in size
and ground color, and, to some extent, in the major dark
markings of the forewing. The two species certainly
occur in similar habitats in the mountains and possibly
on the coastal plain.

When the wing pattern is obscured, the genitalia of
either sex can be used to distinguish the species. Males
of M. triangula lack hair pencils at the base of the
abdomen as do M. confusa (Hbn.) and M. latex. The
digitus is almost basal in M. confusa hut crosses the
membranous sacculus more distally in M. latex and
triangula. In M. latex the membranous area is swollen
distally, in triangula it tapers evenly. The everted vesica
of triangula is unique in the genus because it has no
large cornutus nor comb-like concentrations of smaller
cornuti. Instead, it has a ribbon-like band of small
cornuti .

Distribution and Habitat. This species occurs from
the southeastern corner of Virginia along the coastal
plain to all of Florida and west to east Texas. There are
veiy few piedmont records (which may reflect a lack of
collecting in this area). In the mountains it occurs from
Averv Co. in North Carolina to the foothills in NE
Alabama. County records are as follows: VIRGINIA:
City of Suffolk, City of Virginia Beach, Isle of Wright;
NORTH CAROLINA: Bladen, Brunswick, Carteret,
Craven, Cumberland, Hoke, Macon, Martin, Moore,
Onslow, Pender, Union; SOUTH CAROLINA:
Charleston, Greenville, Oconee; TENNESSEE:
Cocke; GEORGIA: Fulton, Long, Lumpkin, Rabun,
Towns, Union; FLORIDA: Alachua, Escambia, Liberty,
Monroe, Okaloosa; ALABAMA: Bibb, Conecuh,
Sumpter; MISSISSIPPI: Forrest, Franklin, George,
Hancock, Harrison, jackson. Pearl River, Pike;
ARKANSAS: Ashley, Union; LOUISIANA (Parishes):
Iberville, St. Tammany, West Feliciana, Winn; TEXAS:
Hardin, Jasper, Polk, Sabine, Tyler.

Collections of this moth are not limited to a specific
habitat. It occurs in swamps, savannahs, pinelands and
mesic hardwood sites throughout the Coastal Plain.
Mountain sites are equally undistinguished.

Flight Season. In the Coastal Plain of North
Carolina the extended flight period runs from late
March into }une and perhaps early July. Most males fly
in mid April and most females are in flight in May-June.
Mountain records (above 4000’) are May through mid
July. For other states with numerous records, the
recorded flight periods are as follows: Mississippi:

Volume 63, Number 1

23

Figs. 1-4. Adult Morrisonia triangula Sullivan and Adams. Male. Little Deep Creek Rd., Croatan National Forest, Craven Co,
N. C. April 2, 2000, J. Bolling Sullivan 2. Morrisonia triangula Sullivan and Adams. Female. Croatan National Forest Rd. 147,
Craven Co., N. C. April 1 1, 2000, }. Bolling Sullivan. 3. Morrisonia latex (Guenee). Male. East side of Carp Ridge, 45°24.63"N, 76°
03.45'W, Ontario, 14 June 2003, j. Troubridge. 4. Morrisonia latex (Guenee). Female. Temiscouata Co., Quebec, 16 June 1993,
Henry Hensel.

March 27-June 2; Louisiana: March 12-June 15; Texas:
April 8-May 22; South Carolina: May 2 1— [uly 25 (the
latest recorded specimen, from the mountains [Oconee
Co.]); Georgia: April 23-May 27; Florida: March
17-May 23; Alabama: June 3- June 12. There is no
evidence for a second brood.

Generic Placement. The genus Morrisonia is
attributed to Grote (1874), with evicta as the type
species, but no characters are given in his work to help
delineate the genus. Hampson (1898-1913) included
taxa from North America and New Zealand but species
from New Zealand have since been removed (Dugdale
1988). Forbes (1954) characterized the genus as follows:
“vestiture always mixed, the division of the tuft on upper
part of face obvious; collar usually with suggestion of a
central ridge, and concave on either side; double dorsal
tufts of thorax distinct; several distinct tufts on
abdomen.” Since Forbes’ treatment of the genus, M.
distincta Hbn. has been moved to the genus Achat ia
(Hodges et. al. 1983) and Mamestra latex (Gn.) has been
moved into Morrisonia (Poole 1989). Other North

American genera placed in the tribe Orthosiini by
Lafontaine (2007) are Acerra, Stretchia , Perigonica,
Orthos ia , Himella, Egira and Achatia. Members of the
tribe are united by characters of the larval hypopharynx
and spinneret (Fibiger & Lafontaine 2005), characters
which we have not evaluated. Genitalic variation in this
tribe and within several of the genera is extreme
(Ronkay et. al. 2001). The genus Morrisonia is no
exception. What characters then seem to place this new
species in Morrisonia? Two external characters other
than wing pattern support placement of the new species
in Morrisonia. First, the vestiture is coarse, a mixture of
scales and hairs, and in most of the other genera only
hair scales are present. Second, there is a well-
differentiated line across the Irons that occurs in the
other Morrisonia species but is extremely faint when
present (rarely) in other genera. Comparison of related
species and genera ( Egira alternans (Wlk.), variabilis
(Sm.), dolosa (Grt), crucialis (Harv.), mbrica (Harv.),
cognata (Sm.); Stretchia inferior (Sm.); Perigonica
angulata (Sm.); Himella fidelis Grt.; Orihosia praeses

24

Journal of the Lepidopterists’ Society

Figs. 5-7. Male genitalia of Morrisonia. 5a,b. Morrisonia evicta (Grote). a. Valves. Dunrobin, Ontario (CNC 13538). b. Aedea-
gus. Same data as male valves. 6a, b. Morrisonia triangula Sullivan and Adams, a. Valves. Wedge Plantation, 7miN McClellanville,
South Carolina (CNC 13964). b. Aedeagus. same data as male valves. 7a,b. Morrisonia latex (Guenee). a. Valves. North Gower,
Ontario (CNC 13539). b. Aedeagus. same data as male valves.

7a

1 mm

Volume 63, Number 1

25

8

lmm

Figs. 8-10. Female genitalia of Morrisonia spp. 8. Morrisonia evicta (Grote). Female. Aweme, Manitoba (CNC 13560). 9. Mor¬
risonia triangula Sullivan and Adams. Female. Key Largo, Florida (CNC 13542). 10. Morrisonia latex (Guenee). Female. Lac Mon-
dor, Ste Flore, Quebec (CNC 13540).

(Grt), rubescens (Wlk.), hibisci (Gn.), alurina (Sm.),
pacifica (Han/.), revicta (Morr.), pulchella (Han/.), mys
(Dyar), erythrolita (Grt.), garmani (Grt.) and segregata
(Sm.)) and all 5 Morrisonia species using COl
barcoding techniques (Hebert et. al. 2006) reveals that,
whereas most species sort with their supposed
congeneric members, there are often inclusions of one
or two additional non congeneric species in each
generic cluster. In the case of Morrisonia both evicta
and triangula sort together but may be separated from
the other Morrisonia species by Orthosia hibisci or
rubescens. Examination of the male and female
genitalia (Figs. 5, 6, 8, 9) shows that M. evicta and
triangula are quite similar in all characters and the pair
is most similar to M. macens (Hbn.). M. confusa and M.
latex have genitalia which are quite divergent from the
macens, evicta, triangula complex. Significant

divergence is seen in the COl data from M. latex but
not M. confusa.

Diverse assemblages of species are characteristic of
Orthosia, Egira, and perhaps other orthosiine genera
with multiple species (Ronkay et al. 2001). Finding good
generic characters is difficult at best. For M. mucens,
evicta (Fig. 5) and triangula (Fig. 6) we could list the
spiral vesica with groups of cornuti but no major spine,
a short diverticulum near the apex of the vesica, and the
valva, diaphanous along its ventral edge, indented from
the ventral side past the midpoint and expanded distallv.
The valve process derives from the costal plate; the
medial plate often produces a small process. In females
(Figs. 8, 9) the appendix bursa is coiled, the ductus
bursa well sclerotized, and the ostium simple. These
same characters are either missing or highly modified in
M. confusa and latex (Fig. 10).

26

Journal of the Lepidopterists’ Society

Morrisonia confusa and latex are polyphagous (Covell
2005; Rings et al. 1992). Most records for Morrisonia
evicta are from Primus , although Kalmia and Cornus
are listed also, perhaps erroneously (Wagner et al 2008).
Jack Franclemont reared Morrisonia mucens from
Water Oak ( Quercus nigra L.) as reported in Godfrey
(1972). While most orthosiines require tender new
foliage, both M. confusa and M. latex develop
comparatively slowly on older, hardened summer
foliage. M. confusa is exceptional among noctuids in
taking as long as 6-7 months to mature (Wagner 2005).
The habits of M. confusa and M. latex are strikingly
different from each other and completely different from
the other spring feeding orthosines. M. confusa begins
the year feeding externally on new growth as do the
other species, but in the last instar it fashions a shelter
on old summer foliage, densely lines it with silk, and
continues to feed for upwards of 5-6 months.
Morrisonia latex also feeds throughout the summer but
rests on bark by day (at least in the last instar), a habit
completely unique to other orthosines (Wood & Butler
1989; Wagner et al 2008). Based on the sum of
differences in larval morphology, behavior, and life
history, M. evicta , confusa and latex could easily be
placed in three separate genera.

Thus, everything points to inclusion of triangula in
the genus Morrisonia , considering that it seems to be
the closest relative to evicta , the type species. In fact it
is a better fit than some of the other species currently
placed there. When the larval biology of triangula is
known, hopefully, it will confirm this placement rather
than add more heterogeneity to yet another genus in the
Orthosiini.

Acknowledgements

YVe would like to acknowledge the following persons who
contributed data, insight or both: Vernon Brou, Richard Brown,
Chuck Ely, Irving Finkelstein, Howard Grisham, Steven Hall,
Paul Hebert, Ed Knudson, Hugo Kons, Don Lafontaine. Tim
McCabe, Tom Neal, Steve Roble, Dale Schweitzer, Brian
Scholtens, Darryl Searcy, Jeff Slotten, John Snyder and Dave
Wagner. Figures were prepared by Jocelyn Gill.

Literature Cited

Coyell, C. V., Jr. 2005. A field guide to the moths of eastern North
America. 2nd Ed. Houghton Mifflin Col, Boston, Mass.

Dugdale, J. S. 1988. Lepidoptera-annotated catalogue, and keys to
family group taxa. Fauna of New Zealand 14: 1-262.

Fibiger, M. & J. D. Lafontaine. 2005. A review of the higher classi¬
fication of the Noctuoidea (Lepidoptera) with special reference
to the Holarctic fauna. Esperiana (Buchreihe zur Entomologie),
Band 11, pp. 7 - 92.

Forbes, W. T. M. 1954. Lepidoptera of New York and neighboring
states. Noctuidae, Part III. Cornell University Agricultural Ex¬
perimental Station. 433 pp.

Godfrey, G. L. 1972. A review and reclassification of larvae of the
subfamily Hadeninae (Lepidoptera, Noctuidae) of America north
of Mexico. Tech. Bull. No. 1450. U. S. Dept. Agriculture, Wash¬
ington, D. C. 265 pp.

Grote, A. R. 1874. List of the Noctuidae of North America. Bull, of
the Buffalo Society of Nat. Hist. 2: 1-77.

Hampson, G. F. 1898-1913. Catalogue of the Lepidoptera Phalaenae
in the British Museum. Volumes 1-13.

Hebert, P. D. N., E. H. Penton, |. Al. Burns, D. H. Janzen & W.
Hallwachs. 2006. Ten species in one: DNA barcoding reveals
cryptic species in the neotropical skipper butterfly Astraptes fttl-
gerator. Proc. Natl. Acad. Sci. 101: 14812-14817.

Hodges, R. W„ T. Dominick, D. R. Davis, D. C. Ferguson, J. G.
Franclemont, E. G. Munroe, J. A. Powell (eds). 1983. Check
list of the Lepidoptera of America north of Mexico (including
Greenland). E. W. Classey Ltd. and The Wedge Entomological
Research Foundation, London. 284 pp.

Kimball, C. P. 1965. The Lepidoptera of Florida: an annotated check¬
list. In Arthropods of Florida and neighboring land areas. Vol. 1.
Fla. Dept. Agile., Gainesville. 363 pp.

Lafontaine, J. D. 1998. Noctuoidea: Noctuidae (part), Noctuinae
(part-Noctuini). Fascicle 27.3. In R. B. Dominick et al. (eds). The
moths of America north of Mexico. The Wedge Entomological
Research Foundation. Washington, D. C. 348 pp.

_ . 2007. Updated electronic checklist of the moths of America.

Poole, R. W. 1989. Lepidoptorum catalogus. Fascicle 118, Noctuidae,
3 volumes.

Ratnasingham, S. & P D. Herbert. 2007. Bold: The barcode of life
data system. Molecular Ecology Notes 7(3): 355-364.

Rings, R. W, E. H. Metzler, F. J. Arnold. & D. H. Harris. 1992.
The owlet moths of Ohio (Order Lepidoptera: (Family Noctu¬
idae). Ohio Biol. Survey Bull., New Series, Vol. 9, College of Bio¬
logical Sciences, Ohio State University, Columbus, Ohio. 219 pp.

Ronkay, L.. J. L. Yela & M. Hreblay. 2001. Noctuidae Europaeae,
Hadeninae II. Soro: Entomological Press. 452 pp.

Wagner, D. L. 2005. Cateipillars of eastern North America: A guide
to identification and natural history, Princeton University Press.
496 pp.

_ , D. F. Schweitzer, J. B. Sullivan, & R. C. Reardon. 2008.

Owlet caterpillars of Eastern North America (Lepidoptera: Noc¬
tuidae). Princeton Universitv Press (in preparation).

Wood, P. S., & L. Butler. 1989. Biology and immature states of Po-
lia latex (Guenee) (Noctuiidae). [. Lepid. Soc. 43: 299-304.

Received for publication 31 January 2008; revised and accepted
20 October 2008

Volume 63, Number 1

27

Journal of the Lepidopterists’ Society
63(1), 2009, 27-30

TINACRUCIS NOROESTA, NEW SPECIES, NORTH AMERICA’S LARGEST TORTRICINE MOTH

(TORTRICIDAE: ATTERIINI)

[erry A. Powell

Essig Museum of Entomology, University of California, Berkeley, 94720, e-mail: poweIlj@nature.berkeley.edu

ABSTRACT. The Neotropical tribe Atteriini is briefly characterized, and Tinacrucis noroesta is newly described in order to make the name
available for a forthcoming book on moths of western North America. This species, the northernmost of the tribe, occurs in the Sierra Madre
Occidental, Mexico, and mountains of southern Arizona, where it was first collected in 1927. The larvae are believed to be polvphagous leaf
rollers on broadleaf trees and shrubs but have not been discovered in the field.

Additional Key words: Egg mass adornment scaling, sexual dimorphism. Chihuahua, Durango, Arizona, leaf-roller.

The Atterini comprises one of the smallest and most
well defined tribes of the Tortricinae. Its monophyly is
convincingly supported by a suite of uniquely derived
morphological and behavioral characters associated with
oviposition (Powell 1986). The adults are among the
largest tortricid moths in the Western Hemisphere, FW
length ranging to 22 mm. Females are conspicuously
larger than males, and there is pronounced sexual
dimorphism in wing shape and color patterns in some
genera (. Anacrusis , Tina, Tinacrucis ), whereas the sexes
are similar in other genera (e.g., Atteria, Templemania).
The eggs are laid in imbricate, overlapping patches,
presumably a synapomorphy with Archipini and
Sparganothini. Females possess obvious patches of
corethrogyne scales covering the venter of A6 and A7; a
thick pad of long, thin, dark scales on A6, which are
spread over the egg mass and its periphery, and an
enormous patch of upright, thicker scales on A7 that are
curled at the tips, which are glued to the substrate in
clumps, like a fence surrounding the egg mass. Most
atteriines are nocturnal, but Atteria are diurnal and
brightly colored, evidently specific Batesian mimics of
Pseudatteria (Tortricinae: Polyorthini). Atteriine larvae
are similar to sparganothines but are much larger leaf
rollers. The distance between V setae on A9 is greater
(1.5-2X) than on A8, the crochets are bi- or triordinal,
and there is a well developed anal fork. I have reared
several species of Templemnaia, Tina, and Tinacucis
from eggs to maturity on synthetic diet, which suggests
atteriines are polyphagous, and Dan Janzen and others
have reared field collected larvae of Costa Rican
Anacusis from several unrelated plant families,
enhancing that assumption. The Atteriini includes about
40 described species assigned to eight genera, in
addition to numerous undescribed taxa in Central and
South America. A few species range into northern
Mexico, one into Arizona.

Tinacrucis Powell, 1986, is distinguished by having
elaborate antennal setulae in males, the costa sinuate in
both sexes (unmodified in male Anacrucis and
Templemania), markedly different size and FW pattern
between the sexes (though not as extreme as in Tina),
greatly enlarged A8 tergum, forming a hood enclosing
the male genitalia. Tinacrucis includes six named
species and several undescribed in Mexico and Central
America. I have reared broods of several of them from
single female egg masses, enabling association of the
markedly differing sexes.

I have been aware of the species described here for
50 years, based on males taken by J.A. Kusehe in the
Chiricahua Mountains, Arizona, in 1927, but until
recent years, we lacked sufficient material to confirm
conspecificity of the Arizona species with specimens
collected in Durango, Mexico. This species is described
now because we discuss it in our forthcoming book.
Moths of Western North America (Powell & Opler in
press), scheduled for publication by the University of
California Press in May, 2009. A treatment of the
Sparganothini and Atteriini, by J. W. Brown and Powell,
is in preparation for Moths of America North of Mexico,
an appropriate place for this description, but it will be
published later. Both of those volumes will include color
images of the new species.

Tinacmcis noroesta Powell, new species
(Figs. 1, 2, male, female from
El Salto, Durango, Mexico)

This is the largest tortricine moth in America north of
Mexico. Males have a relatively narrow FW with sinuate
costa, pale beige to yellowish reticulated with rust lines
and irregularly mottled with rust-brown. Females are
larger with broader wings, pale chocolate- to rust-brown
with faint darker and white banding in costal area when
fresh and with huge masses of modified scales ventrally
on abdominal segments 6 and 7.

28

Journal of the Lepidopterists’ Society

Fig. 1 . Tinacrucis noroesto Powell <$ — Mexico, Durango, YV of El Salto, VII. 5. 1964 (J. Martin) CNC.

Fig. 2. T. noroesta Powell

Mexico, Durango, YV of El Salto, VII. 1.1964 (J. Martin) CNC.

Volume 63, Number 1

29

Fig. 3. Tinacrucis noroesta, male genitalia, ventral aspect, valvae spread; aedeagus, lateral aspect shown to right; a, aedeagus with
vesica extended, revealing the deciduous comuti.

Fig. 4. T. noroesta, female genitalia, ventral aspect; a, detail of translucent ventral plate of A9; b, detail of signum, dorsal aspect.

30

Journal of the Lepidopterists’ Society

Description. MALE. Forewing. Length 14.7-16.3 mm (n =
25), narrower than female, length 2. 6-2. 8 x width, costa sinu¬
ate, concave beyond middle, apex produced and subtended by
concave terminal margin; ground color pale beige to yellow,
reticulate with rust-brown along veins and numerous cross lines
between veins, confined by rust-brown clouding to irregular
blotches, basally, in cell, and subterminal area (resembling
skeletonized spots on a fallen leaf); costal area beyond middle
whitish in fresh specimens. Hindwing. Broad, pale cream col¬
ored, tinged with rust apically; underside variably mottled by
rust in costal and apical areas, which is visible from above. Ab¬
domen. Scaling pale beige to rust tinged caudally, venter paler,
with segmental dark, medial spots. Genitalia . As in fig. 3
(drawn from paratype, El Salto, JAP slides 2893; n = 8); uncus
only slightly flared apically, socii densely covered with elongate
scales; joined tip of gnathos elongate, slightly flared; a pair of
dentate projections on posterior margin of transtilla usually
widely separated but adjacent in one specimen from Jalisco;
basal half of valva with weakly projecting saccular ridge re¬
cessed from anterior margin; vesica with a compact group of 8
bladelike, deciduous cornuti situated in two rows; after dehis¬
cence, the attachment scars are visible at high magnification
(100X) on the withdrawn vescia near the phallobase/aedeagus
junction.

FEMALE. Forewing. Length 17.0-20.0 mm (n = 6); broad,
length 2.45-2.55 x width; costa weakly concave beyond middle;
ground color milk chocolate- to rust-brown, showing faint
transverse, whitish and dark bands on costal half beyond middle
on fresh specimens; underside rust tinged; fringe narrowly rust.
Hindwing. As in male. Abdomen. Scaling pale beige dorsally;
venter paler, with large, medial, black spots on segments 3—5;
A6 witli a thick patch of thin, dark brownish scales; A7 with a
conspicuous patch of upright, pale to dark scales, each bent at
the tip. Genitalia. As in fig 4, a, b (drawn from paratype, Sum-
merhaven, JAP slide 9080; n = 3); papillae anales broad, darkly
pigmented, with raised setiferous nipples; sterigma a broad
bowl subtending A9, which is modified as a translucent plate
covered with tiny spicules, more densely at margins (4a);
signum a densely wrinkled patch at bursa surface, preceded by
a small, blind pouch, projected inwardly as a broad, weakly
wrinkled, hollow blade (4b).

Holotype female: ARIZONA, Pima Co., Summerhaven,
7,800', Santa Catalina Mountains, emerged 10 December 1997,
larva reared on synthetic diet from egg deposited by female col¬
lected 29 August 1997 (R. B. Nagle. JAP lot 97H9); in Essig
Museum of Entomology, University of California, Berkeley.

Paratypes (35 6, 7 9): MEXICO, CHIHUAHUA, Sierra de
la Catarina, 7,900', 18 rd. mi. NE Buenaventura, 1 6
VIII. 21. 1976 (J. & K. Donahue, LACM). DURANGO. 10 mi.
YV of El Salto, 9,000', 1 6 VI. 13.1964, 5 d, 19 VII.5/29.1964 (J.
Martin, CNC), 1 6 VII. 30. 1964 (Chemsak & Powell, EME), 1 6
VIII. 8. 1964 (Powell, EME). JALISCO, 18 km SW Sayula, 1 6
X.18.1992 (J. McCarty, EME). USA; ARIZONA, Apache Co.,
Greer, White Mts., 3 6 VHI.6/8.1962 (E. & I. Munroe, CNC),
2 6 VIII. 5. 1986 (R. Leuschner, RL), 6 Alpine, 8,200',
VIII. 3. 1986 (R. Leuschner, RL). Cochise Co., Barfoot Ridge,
8,500'. Chiricahua Mts., 2 6, VIII.6.1927 (J. Kusche, CAS);
Barfoot Park, 1 9 IX.27.1987 (N. McFarland, EME); Rustler
Park, 3 6 VII.30.1987, 1 6 VII.24.1989, 1 i VIII. 7.1997 (R. &

J. Robertson, EME), 1 d VIII.6.1991 (Robertson & Powell,
EME); Pinery Canyon, 6,500', Chiricahua Mts., 1 d
VIII.6.1991 (Robertson, EME); Pinery Canyon 6800’ 1 9
VIII. 1.2005 (K. Richers, KR); Miller Canyon, 6,200’, Huachuca
Mts., 1 6 VIII. 7. 1974 (Powell, EME). Graham Co., Soldier
Creek Camp, 9,350', Graham Mts., 1 9 VIII. 16.1974 (R. & J.
Wielgus, USNM); Mt. Graham, Cunningham Campgr., 3 d
VII. 20.2007 (C. Ferris, CF, EME). Pima Co., same data as
holotype, 5 d, 2 9 VII.24 to VIII.29, 1991 to 1999 (EME, RN),
1 9 VIII. 1996, eggs infertile (EME), 1 d VII.21.1998, (D. Fer¬
guson, USNM).""

Diagnosis. The geographically nearest congener is
Tinacmcis apertana (Walker), with which T. noroesta
Hies in the mountains of Durango, Mexico. I have
reared T. apertana in Nuevo Leon from eggs,
confirming association of the sexes. The two are similar,
but the FW of apertana males has more extensive
network pattern on a paler, cream yellowish ground,
defining a broad, brown costal triangle; females of
apertana have cream-tan FW, more or less uniformly
strigulate with thin, brownish, transverse lines, with a
brownish bar from mid-costa curving towards the
tornus, and a poorly defined subterminal patch,
contrasted with the more uniform rust tan of noroesta.
T. sebasta (Walsingham), which was described from
Guatemala and occurs in Mexico, has the reticulate
pattern restricted to the basal half of the male FW, anti
the female FW pattern is similar to that of apertana but
darker tan. The uncus of T. apertana has a wide, T-
shaped apex, and that of sebasta is broadly spatulate
subapieally, tapering to the tip, whereas it is narrow,
only weakly flared in noroesta. T. apertana and sebasta
have narrower valvae than does noroesta , with a more
distinct saccular ridge. The female signum is short and
broad in T. apertana , elongate in sebasta , more slender
than in noroesta.

In addition we have two male specimens from
Mexico, Sinaloa, 8 road miles W of El Palmito, 6,400',
X. 12.1975 (Chemsak & Powell, EME), which are
similar but differ in several respects. They are slightly
larger than the other T. noroesta specimens, and their
colors are richer, more contrasting yellow and brown.
The male genitalia have a slender uncus, not flared. The
dentate projections of the transtilla are closely adjacent,
and there are slight differences in the valva. These
specimens are provisionally considered to represent
another population of T. noroesta.

Acknowledgements

Carolyn Mullinex made the drawing of the male genitalia. Sev¬
eral collectors made special efforts in response to my pleas to col¬
lect this remarkable species in Arizona and Mexico; the late John
Chemsak, Ron Leuschner, Ray Nagle, the late Ron Robertson, and
Ron Wielgus. Paratypes are deposited in the California Academy
of Sciences (CAS); Canadian National Collection, Ottawa (CNC);
Essig Museum of Entomology, UC Berkeley (EME); Los Angeles
County Museum of Natural History (LACM); Ron Leuschner,
Manhattan Beach, California (RL); Ray Nagle. Tucson, Arizona
(RN); K. Richers, Bakersfield, California (KR); and U.S. National
Museum of Natural History, Washington, D.C. (USNM).

Literature Cited

PowelI, J. A. 1986. Synopsis of the classification of Neotropical Tor-
tricinae, with descriptions of new genera and species (Lepi-
doptera: Tortricidae). Pan Pacific Entomologist 62:372-398.

Received for publication 27 Aug 2008: revised and accepted 17 Nov 2008.

Volume 63, Number 1

31

Journal of the Lepidopterists' Society
63(1), 2009, 31-36

MELITARA WALKER (PYRALIDAE) IN WESTERN CANADA: THE DOCUMENTATION OF M.
SUBUMBRELLA (DYAR) IN THE PRAIRIE PROVINCES DEMONSTRATES THE VALUE OF
REGIONAL COLLECTIONS AND SPECIES LISTS.

Thomas J. Simonsen

Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada; email: thomas.simonsen@ualberta.ca

Jason J. Dombroskie

Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada.

AND

Gregory R. Poiil

Natural Resources Canada, Northern Forestry Centre, Edmonton. AB, Canada.

ABSTRACT. The moth genus Melitara comprises 8 recognized species of medium sized to large micromoths that all feed on Opuntia cacti
as larvae. One species, M. dentata is widespread in western North America, including the southern parts of the Canadian provinces British Co¬
lumbia, Alberta and Saskatchewan. We report the presence of a second species, M. subumbrella in southern Alberta and Saskatchewan, a species
hitherto widely believed to extend no farther north than southern Idaho and Wyoming. Closer examinations of Melitara collections in major
Canadian museums and public collections revealed several additional M. subumbrella specimens from Alberta, including specimens from the
1940s. Two specimens were actually reported by K. Bowman (1951) under a previous taxonomic arrangement, but were' subsequently over¬
looked by later workers on North American Phycitinae. Combined with subtle differences in mitochondrial DNA, this leads us to conclude that
M. subumbrella has been present in Canada for a long time. This provides an excellent example of the scientific value of continuous insect col¬
lecting, well-curated regional collections, and regional faunal lists.

Additional key words: Disjunctive distribution, mitochondrial DNA, cactus-feeding

The pyralid genus Melitara Walker, a prominent
group of cactus-feeding phycitines, is found throughout
much of the USA, southwestern Canada, and northern
Mexico. One species, M. prodenialis Walker, is
widespread throughout the east, and another species,
M. dentata (Grote), is widespread in the west. The
remaining 6 species are till restricted to Mexico and the
southwestern USA with only M. subumbrella (Dyar)
extending north to Wyoming and southern Idaho, and
west to southern California (Heinrich 1956; Mann 1969;
Neunzig 1997). Bowman (1951) reported M. dentata
and Olycella [ =Melitara ] nephelepasa from Alberta, but
his records were ignored by later workers. The latter is
referable to M. subumbrella, which was considered to
be a synonym of M. nephelepasa at the time
(McDunnough 1939). A full list of currently recognized
species of Melitara and their reported range is given in
Table 1. Neunzig (1997) included the genus Olycella
Dyar in Melitara, with the three species O.
subumbrella, O.junctionella (Hulst) and O. nephelepasa
(Dyar) (Heinrich 1939, 1956). This synonymy has later
been shown to be systematically justified (Simonsen
2008). It is not currently clear whether Melitara and
Olycella (both sensu Heinrich 1956) are reciprocally

monophyletic and thus deserve subgenus status. All
species in the genus feed strictly on prickly pear cactus
in the genus Opuntia (Mann 1969) and are thus
restricted to habitats with Opuntia.

Identifying Melitara subumbrella. Both M.
subumbrella and M. dentata are large micromoths with
wingspans exceeding 50 mm. With their stout bodies
and rectangular wings they have a superficial similarity
to medium sized Noctuoidea. The resting moth with its
large porrect palps, and the wings coiled cigar-like
around the body is fairly easy to identify as a phycitine
(Fig. 1). Fresh specimens can fairly easily be identified
to species without dissection (at least in areas where
only M. subumbrella and M. dentata occur). Both
species have white hind wings and gray forewings
(sometimes with a yellowish cast — especially in older
collection specimens). Melitara dentata has a fairly
elaborate zigzag pattern across the forewing, whereas
M. subumbrella has uniformly gray forewings almost
devoid of any pattern (Fig. 2a, b). Old, faded, and
especially denuded specimens can be much harder to
identify. In this case dissection is often necessary.
Females are easy to identify based on genital ic
dissections as M. subumbrella. The species formerly

32

Journal of the Lepidopterists’ Society

Fig. I . Live Melitara dentata displaying typical Phycitinae
posture with the wings rolled around the abdomen and the an¬
tennae pressed down along the back. CAN: AB: Buffalo:
50.848°N -110.696°W: 16 viii 2006. Photo: JJD.

grouped in Olycella are the only members of Melitara
with a signum on the corpus bursae (Heinrich 1939,
1956; Neunzig 1997). The males are harder to identify,
but the shape of the valves differs slightly between the
two species. In M. dentata the valve has a fairly pointed
apex with curved dorsal and ventral margins. The costa
extends almost to the apex of the valve (Fig. 3a). In M.
subumbrella the valve has a much broader and blunter
apex, and the costa does not extend as far (Fig. 3b).
Finally, the flight period is often a veiy good way to tell
the species apart. All M. subumbrella from Canada were
collected between May 15 ami July 2, whereas all M.
dentata were collected between July 27 and October 25,
so there is only a small risk of an overlap in flight time
between the two species. This pattern where M.
subumbrella is a late spring to early summer flyer, and
M. dentata is a late summer to fall flyer seems to be the
same in the western USA, though some M. dentata are
reported from early Julv (Neunzig 1997).

Herein we report on an examination of M.
subumbrella specimens, to determine if Canadian
populations are distinct from those farther south. To do
so, we examined morphological and genetic characters.

Materials and Methods

Morphology. To search for morphological
differences, two male and one female M. subumbrella
from Grasslands National Park SK, one male and one
female M. subumbrella from central USA, and several
male and female M. dentata were examined. Abdomens
were dissected and macerated in 10% KOH, and
stained in Chlorazol Black in a 70% ethanol solution by
TJS, both for use in this study and others (e.g. Simonsen
2008). Terminology follows Heinrich (1956), Klots
(1970) and Neunzig (1997).

Molecular analysis. To test for genetic differences
between the Canadian and USA populations, we

sequenced 740 bp from the cytochrome oxidase subunit
I (COI) gene from three specimens from Grasslands
National Park and one from Colorado. DNA was
extracted from legs with the QIAgen’s DNEasy
extraction kit (QIAgen Sciences, Maryland, U.S.A).
PCR reactions were performed using the Jerry - Pat
primer pair (Simon et al. 1994), and the PCR cycling
profile was: 95.0°C for 2 minutes followed by 35 cycles
of denaturizing (95.0°C for 30 sec.), annealing (45.0°C
for 30 sec.) and extension (72.0°C for 2 minutes). PCR
products were purified using the QIAgen’s PCR
Purification Kit, and the PCR primers were also used
for direct sequencing. Sequencing was done with an
ABI Prism 377 DNA sequencer using Big Dye®.
Consensus sequences from the two sequencing
directions were constructed using Sequencher 4.1 and
aligned by eye. The sequences are deposited on
GenBank under accession numbers

FJ422995-FJ423000.

Phylogenetic analysis. An exhaustive maximum
parsimony (MP) analysis was carried out in PAUP0
4.10b (Swofford 2002). The haplotype distributions
were analyzed in MacClade 4.0 (Maddison & Maddison
2000). The tree was rooted using a M. dentata specimen
from Alberta as outgroup.

Fig. 2. Melitara dentata (a) and M. subumbrella (b) display¬
ing subtle, yet clear differences in forewing pattern (arrows
point to the zigzag pattern on the forewing of M. dentata).

Volume 63, Number 1

33

Table 1. Currently recognized species of Melitara and their distributions following Heinrich (1956), Mann (1969), and Neunzig (1997).

Species

Distribution

Melitara prodenialis Walker

Southeastern U.S.A. from Texas and Florida to New York

Interior western North America from north western Mexico, Arizona and western Texas to

Melitara dentata (Grote)

southwestern Canada.

Melitara texana Neunzig

Texas

Melitara doddalis Dvar

Southern Arizona, southern New Mexico and western Texas.

Melitara apigramella Blanchard & Knudson

Southwestern Texas

Melitara junctionalla Hulst

Southern Texas and eastern Mexico.

Melitara nephelepasa (Dyar)

Central Mexico

Southwestern USA from southeastern California and western Texas to southern Idaho,

Melitara subumbrella (Dyar)

Wyoming and Nebraska.

Results

Morphology. When dissected, males from
Grasslands National Park were found to be
indistinguishable from males from Colorado, USA. The
dissected female from Grasslands National Park,
showed a very subtle difference. The signum on the
corpus bursae was less developed than the
corresponding structure in specimens from Colorado.

Molecular analysis. The MP analysis resulted in a
single tree 62 steps long (Fig. 4). The four specimens
from Grasslands National Park all shared the same
haplotvpe; whereas the specimen from Colorado
differed in three base pairs, corresponding to a
difference of 0.4%. M. clentata differed in 59 base pairs
compared to all four M. subumbrella specimens,
corresponding to a difference of 8.0%.

Discussion

Melitara in Canada. In 2004 GRP collected five
specimens of M. subumbrella in Grasslands National
Park, Saskatchewan. In his annotated list of Lepidoptera
in Alberta, Bowman actually reported two specimens of
M. subumbrella from the Medicine Hat region
(Bowman 1951), though he listed them as Ohjcella
nephelepasa (following McDunnough 1939). Melitara
subumbrella is technically a new record for Canada.
However, a simple update of Bowman's (1951)
checklist, following the synonymy reported in Munroe
(1983) would yield the report of this species in Alberta.
Both the specimens and Bowman’s paper have been
overlooked by later workers on the genus in North
America (e.g. Heinrich 1956; Mann 1969; Neunzig
1997). One of Bowman's specimens was located in the
University of Alberta Strickland Museum. Four other
specimens were misidentified as M. dentate in the same
collection, including another specimen collected by

Bowman in 1940 near Medicine Hat. The latter is
obviously Bowmans second specimen which later must
have been misplaced in a tray with M. dentate. The
three remaining specimens were collected in 2005 at
two different localities in southeastern Alberta. This
inspired us to examine other public collections that we
felt could hold either overlooked or misidentified
specimens of M. subumbrella from western Canada.
Examinations of the materials in the collections of the
Canadian Forest Service (Edmonton, AB), Royal
Alberta Museum, (Edmonton, AB), Olds College (Olds,
AB), Royal Saskatchewan Museum (Regina, SK) and
the Canadian National Collection, (Ottawa, ON)
revealed one M. subumbrella from Lethbridge, AB
collected by C. E. Lilly in 1949, and one from Onefour,
AB collected by GRP in 1995. The same examinations
also showed that M. dentate occurs from south-central

Fig. 3. Differences in male genitalia between Melitara den-
tata (a) and M. subumbrella (b). Note the more pointed apex of
the valve and that the costa extends to the apex in M. dentata.

34

Journal of the Lepidopterists’ Society

Saskatchewan to the southern interior of British
Columbia (Table 2), a more widespread Canadian
distribution than reported recently by Neunzig (1997),
who only listed the species from southern Alberta. The
distributions of both species based on the collection
data are shown in Fig. 5. These records underscore the
importance of regional collectors, and their published
lists and collections, in determining species ranges.
Local collectors typically have extensive knowledge of
productive collecting localities, and often amass
extensive collections of excellent quality (Ferris 1986;
Hendra 2005, 2007). These collections and publications

are a neglected resource that can yield many exciting
finds to the diligent taxonomist.

Two prickly-pear species, Opuntia fmgilis (Nutt.)
Haw. and O. polyacantha Haw., occur (sometimes
abundantly) throughout the southern parts of Canada’s
Prairie Provinces (personal painful observations).
Though Fragile Prickly-pear Cactus (Opuntia fmgilis), a
possible host plant for M. clentata (Mann 1969; Neunzig
1997), occurs scattered in remnants of the Peace River
grasslands in northwestern Alberta (Royer & Dickinson
2007), there are no records of M. clentata from that
region. It is, however, quite possible that M. clentata

Table 2. Records of Melitara in Canada based on the collections listed in the text. Collection acronyms: CNC: Canadian National
Collection of Insects, Arachnids and Nematodes; NFRC: Northern Forestry Centre Research Collection; OC: Olds College: RSM:
Royal Saskatchewan Museum; UASM: University of Alberta, Strickland Museum.

Species

Locality

Coll, period

Year

No.

Collection

M. clentata

Grasslands National Park, SK

24/7-12/9

2002

6

RSM

M. clentata

Buffalo Pond Provincial Park, SK

7/8-25/8

1974, 1984-85

3

RSM

M. clentata

Wood Mountain Park, SK

4/9

2002

2

RSM

M. clentata

Fort Qu'Appelle, SK

12/8-31/8

1976, 1991

2

RSM

M. clentata

St. Victor, SK

14/8

2002

i

RSM

M. clentata

Matador, SK

7/8

2002

i

RSM

M. clentata

Eastend, SK

reared

1974

i

RSM

M. clentata

Saskatchewan Landing Provincial Park, SK

10/8

1981

i

RSM

M. clentata

Scout Lake, SK

27/8

1968

4

CNC

M. clentata

Swift Current, SK

8/8

1936

1

CNC

M. clentata

Tolinan Bridge, AB

3/8-25/10

2000-2, 2006

13

UASM, NFRC, OC

M. clentata

Taber, AB

14-15/8

1999, 2005

2

OC

M. clentata

Medicine Hat, AB

17/8

1943

1

UASM

M. clentata

Edgerton, AB

24-25/8

2002

1

UASM

M. clentata

Steveville Bridge, AB

13/8

1999

2

UASM. NFRC

M. clentata

Buffalo, AB

16/8

2006

i

UASM

M. clentata

Champion, AB

3/8

1961

12

CNC

M. clentata

Coaldale, AB

4/8

1961

8

CNC

M. clentata

Manyberries, AB

10/8

1951

1

CNC

M. clentata

Lethbridge, AB

26/7-24/8

1921-22, 1949, 1986

6

CNC

M. clentata

Willow Creek Canyon, AB

29/7

1961

4

CNC

M. clentata

Kamloops, BC

5/8-9/8

1937, 1956

4

CNC

M. clentata

Nicola, BC

29/8-6/9

1922-23

4

CNC

M. clentata

Oliver, BC

1/8-15/9

1923, 1953

113

CNC

M. subumbrella

Grasslands National Park, SK

2/6

2004

5

NFRC

M. subumbrella

Onefour, AB

19/6

1995

1

NFRC

M. subumbrella

Lost River Valley w. of Onefour, AB

20-21/5

2005

1

UASM

M. subumbrella

Medicine Hat, AB

31/5

1940

2

UASM

M. subumbrella

Oldman River Hwy. 36, s ofVauxhall, AB

18/5

2005

2

UASM

M. subumbrella

Lethbridge, AB

15/5

1949

i

CNC

Volume 63, Number 1

35

does occur in the Peace River region, as suitable
habitats in the region are fairly poorly collected,
especially during M. dentata’ s main flight period (G. G.
Anweiler pers. com.).

The status of Melitora subumbrella in Canada.

The three oldest specimens of M. subumbrella from
southern Alberta show that the species was present in
the province 60-70 years ago. The presence of recently
collected individuals from different localities in
southeastern Alberta and Saskatchewan clearly
demonstrates that M. subumbrella occurs at multiple
localities in western Canada. It is also not unthinkable
that the species still is overlooked and is more
widespread, not only in southwestern Canada, but also
in Montana, though an examination of the Melitara
specimens in the entomological collection of Montana
State University in Bozman did not result in additional
specimens.

Are Canadian M. subumbrella different from M.
subumbrella in the U.S.A.? Ideally, many more
specimens from both Canada and the U.S.A. should
have been included to strengthen the results of the
molecular analysis. Unfortunately such material was not
available. Nonetheless, the difference reported here is
well within variations that have been found in other

tarn subumbrella in Canada and west central USA. The closed
circle marks the clade comprising the four Canadian specimens.
The six digit code after the species name is the specimen ID.
The numbers indicate the number of unique base pair changes.

Fig. 5. Distribution map for Melitara in western Canada
based on the records listed in Table 2. An open circle marks a M.
subumbrella locality, and a closed triangle marks a M. dentata lo¬
cality.

insect populations considered to be conspecific (e.g.
Simonsen et al. 2008; Cognato 2006; Laffin et al. 2005a,
b; Scheffer & Grissel 2003). The slight difference in
female genitalia may well represent individual variation.
It is not known whether the species is present in
Montana, and the variation thus could be continuous.
Despite the apparent geographical isolation, these
differences do not in our opinion justify separate
taxonomic status (subspecies or otherwise) for the
Canadian M. subumbrella populations. The species
should be sought after in Montana to determine
whether the distribution is continuous or the Canadian
populations are disjunct.

Acknowledgements

We wish to thank Paul Opler, of the Cillette Museum, Col¬
orado State University for loans of specimens from the U.S.A.
and for permission to extract DNA from these. We thank the fol¬
lowing people and institutions for help and assistance: |ean-
F ranyois Landry, Canadian National Collection, Ray G. Poulin,
Roval Saskatchewan Museum, Tyler Cobb, Royal Alberta Mu¬
seum, Ken Fiy, Olds College and Michael A. Ivie, Montana
State University, Bozman. Felix A. II. Sperling, Gary G. An¬
weiler, Danny Shpeley, Marie Djernaes and Lisa Lumley, all of
the University of Alberta, and Brenda Laishley, Canadian Forest
Sendee, Edmonton. AB, are thanked for help, advice and dis¬
cussions. Financial support was provided by The Carlsberg
Foundation, Copenhagen, Denmark (post doc salary forT. J. Si¬
monsen). NSERC (a Discovery Grant to Felix A. H. Sperling
paid for lab work and various expenses), and Natural Resources
Canada (support for faunistics work by G. R. Pohl).

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Ferris, C. D. 1986. Presidential address, 1986: unexplored hori¬
zons — the role of the amateur lepidopterist. J. Lepid. Soc. 40:
247-254.

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Journal of the Lepidopterists’ Society

I Ieinrich, C. 1939. The cactus-feeding Phycitinae: a contribution to¬
wards a revision of the American pyralidoid moths of the family
Phycitidae. Proc. U. S. Nat. Mus. 86: 331^13.

_ . 1956. American moths of the subfamily Phycitinae. U. S. Nat.

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Hendra, L. 2005. Kenneth Bowman, Alberta Lepidopterist. Blue Jay
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_ . 2007. Kenneth Bowman, Alberta Lepidopterist, part 2. Alberta

Lepidopterists' Guild website: http://www.biology.ua]berta.
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07.pdf

Klots, A. B. 1970. Lepidoptera. Pp. 115-130. In S.L. Tuxen (ed.).
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Laffin, R. D., L. M. Dosdall, & F. A. II. Sperling. 2005a. Popula¬
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_ , _ . & _ . 2005b. Population structure and phvlogenetic

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McDunnough, [. II. 1939. Check list of the Lepidoptera of Canada
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Munroe, E. 1983. Pyralidae (except Crambinae). pp. 67-85. In:
Hodges, R. W., Dominick, T., Davis, D. R., Ferguson, D. C.,
Franclemont, J. G. Munroe, E. G., & Powell, J.A. 1983. Check
list of the Lepidoptera of America North of Mexico. E. W.

Classey Ltd. and the Wedge Entomological Research Founda¬
tion, London, UK. 284 pp.

Neunzig, II. H. 1997. Pvraloidea, Pyralidae (part). Fascicle 15.4 In
Dominick, R. B., et al. (eds). The moths of America north of Mex¬
ico. The Wedge Entomological Research Foundation. Washing¬
ton, D.C. 157pp.

Royer, F. & R. Dickinson. 2007. Plants of Alberta. Lone Pine Pub¬
lishing. Edmonton, AB, Canada. 527pp.

Scheffer, S. }. & E. E. Grissell. 2003. Tracing the geographic ori¬
gin of Megastigmus transvaalensis (Hymenoptera: Torymidae):
an African wasp feeding on a South American plant in North
America. Molec. Ecol.12: 405-414.

Simon, C., F. Frati, A. Beckenbach, B. Crespi, H. Liu. & P. Flook.
1994. Evolution, weighting, and phylogenetic utility of mito¬
chondrial gene sequences and a compilation of conserved poly¬
merase chain reaction primers. Ann. Entomol. Soc. Amer. 87:
651-716.

SlMONSEN, T. J. 2008. Phylogeny of the cactus-feeding phycitines and
their relatives (Lepidoptera: Pyralidae) based on adult morphol¬
ogy: evaluation of adult character-systems in phvcitine systemat-
ics and evidence for a single origin of Cactaceae-feeding larvae.
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_ . R. L. Brown, & F. A. H. Sperling. 2008. Tracing an invasion:

phylogeography of the cactus moth, Cactoblastis cactonun (Berg)
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Received for publication 5 June 2008; revised and accepted

27 October 2008.

Volume 63, Number 1

37

Journal of the Lepidopterists’ Society
63(1), 2009,37-47

NOT ALL BILATERAL GYNANDROMORPH BUTTERFLIES ARE INTERSPECIFIC HYBRIDS: NEW

PAPILIO SPECIMENS FROM FIELD POPULATIONS

Mark Scriber

Department of Entomology, Michigan State University, E, Lansing, MI 48824; research associate McGuire Center for Lepidoptera and
Diversity, University of Florida, Gainesville, FL 3261 1-2710; email: scriber@pilot.msu.edu

Rodrigo Mercader

Department of Entomology, Michigan State University, E. Lansing, MI 48824

Howard Romack

28 Irish Lane, Cambridge, NY 12816,

AND

Mark Deering

Lepidoptera Curator, Sophia Sachs Butterfly House, 15193 Olive Blvd., Chesterfield. MO 63017

ABSTRACT. Bilateral gynandromorphs (half male, half female) have previously been reported in interspecific hybrids. However, we report
6 new nearly perfect bilateral gynandromorphs, three (one Papilio canadensis , and two P. glaucus) of which were collected from areas not near
hybrid zones. New interspecific gynandromorphs also are described for apparent P. nitulus x P. multicaudatus hybrids. Bv using hand-paired in¬
terspecific hybrids we have generated both diapausing and direct developing (non-diapausing) gynandromoiphs. While X (or Z)-linked regula¬
tion of diapause in these Papilio may be similar throughout the male and female halves of the individual, the dark melanic color [Y (or W)-
linked] is distinctly different. Wing color regulation is differentially implemented on dorsal and ventral surfaces as seen in sexual mosaics
described here. The developmental irregularities resulting in bilateral gynandromorphs and sexual mosaics are not solely due to interspecific ge¬
netic incompatibilities. However, genetic factors seem to play a major role in their expression. The existence of dark melanic female patchy pat¬
terns supports the suggestion that control of the W (=Y)-linked b* melanic gene is both cell-specific and sex-specific in its ability to suppress the
BAS enzyme activity that normally shunts dopamine to the yellow papiliochrome pathway. Interspecific hybrids involving dark mothers (with
the Y-linked b* gene) mated to P multicaudatus , P. nitulus , and P. canadensis males (all species lacking the dark morph) have also produced hy¬
brid daughters with “intermediate” suppression of melanism giving a dusty colored appearance (differing from color mosaics). This uniform
“dusty” coloration often has been referred to as “cinnamon,” and may reflect incomplete genetic penetrance.

Additional key words: melanic wing color, sex chromosomes, Papilio glaucus, P. canadensis, P. nitulus, P. multicaudatus, P. appalachiensis

Lepidoptera that have developed abnormally to
produce mosaic color patterns and a mixture of other
traits, often representing intersexes (part female and
part male) have frequently been described (Clarke &
Ford 1980; Ayala & Kiger 1984; McCafferty &
Bloodgood 1986). If such developmental errors occur
early in development (perhaps from fertilization of a
binucleate oocyte; Blanchard & Descimon 1988) the
result might be a perfect (or near-perfect) bilateral
gynandromoiph with one half being male and one half
female (Clarke & Clarke, 1983).

A large number of gynandromoiphs have been
reported in Lepidoptera from at least 10 different
species of Papilionidae, eight species of Lycaenidae,
four Hesperiidae, ten Pieridae, four Nymphalidae, and
several families of moths including Saturniidae,

Pyralidae, Geometridae, and Lymantriidae (reviewed in
Scriber & Evans 1988b; see also Sibatani 1980, 1983a,
b; Shuey & Peacock 1985). Cockayne (1935) provides
numerous additional records, and the Milwaukee Public
Museum’s fames R. Neidhoefer Collection has several
hundred mosaics and at least 150 bilateral
gynandromorphs. David and Marc Perlman in Illinois
have also compiled a large collection of mosaics and
gynandromorphs.

A number of suggestions have been advanced to
potentially explain these developmental abnormalities
including: extreme temperatures (Standfuss 1900;
Drescher & Rothenbuhler 1963; Riotte 1978),
ultraviolet light (Nekrutenko 1965), viral infections
(Gardiner 1972; Blau 1978; Sevastopulo 1973),
translocations of parts of the sex-chromosomes and/or

38

Journal of the Lepidopterists’ Society

autosomes (Blanchard & Descimon 1988; Seri her &
Evans 1987; Marec et al. 2001), and background
uranium radiation-induced somatic mutations and the
Chernobyl nuclear power plant disaster (Dantchenko et
al. 1995). Interspecific hybrid crosses have also been
suspected as a potential cause of
mosaic/gynandromorphs (Whicher 1915; Cockayne
1935; Clarke & Sheppard 1953; Clarke et al. 1977; Platt
1983; Blanchard & Descimon 1988; Scriber & Evans
1988a, b). In Papilio such gynandromorphs have been
found to be most often the result of interspecific (or
intersubspecific) crosses (Scriber & Evans 1988b).
Asymmetrical color mosaics may also result from such
interspecific crosses, but are more often seen from the
central parts of a species geographic range, or from
intraspecific pairings, which suggests that mosaic
gynandromorphs could be the result of more complex
and basically different developmental processes than
bilateral gynandromorphs (Clarke & Clarke 1983;
Blanchard & Descimon 1988).

The evidence we had collected from 1979-1986 with
the Papilionidae in eastern North America suggested
that perhaps only interspecific hybridization would
generate bilateral gynandromorphs, and at that time we
had never found a “natural” (i.e. field-collected)
bilateral gynandromoiph (Scriber & Evans 1988b).
However, we have subsequently collected interesting
mosaics and near-perfect bilateral gynandromoiphs
from field populations, some of which were not clearly
near a hybrid zone where introgression from different
species was obvious. We report on some of these
interesting aberrations here.

Tiger Swallowtail Gynandromorphs and
Color/Sexual Mosaics. For an earlier study (Scriber &
Evans 1988b), we reared nearly 30,000 butterflies from
more than 4,000 families of Papilio glaucus and P.
canadensis (previously considered subspecies P. glaucus
canadensis ; Hagen et al. 1991). Of these individuals we
observed 6 of 34 color mosaic adults, all of which
possessed basic dark (mimetic) wing patterns of females
interrupted by yellow patches or blotches on one to
three of the individual wings. These yellow patches
represent either the non-mimetic female, or male type
pattern. Such color mosaics may represent intersexes or
gynandromorphs with part male and part female
asymmetric traits. They may be either gynandromoiphs
or sexual mosaics (Blanchard & Descimon 1988;
Cockayne 1935; Ford 1955; Sevastopulo 1973). In
Papilio glaucus and P. alexiares Hopffer, both
polymoiphic yellow and black forms are found (Scriber
et al. 1996). Only these two Papilio species have the
polymorphic dark and yellow morph females, clearly
showing the color contrast in gyndromoiphism/

mosaicism. One such P. glaucus female (with one yellow
half and one mostly dark half) has recently been figured
and described by Bradshaw (2000).

In addition to these 34 color mosaics, we also
observed three perfect bilateral gynandromoiphs
(including one that is half dark female and half yellow
male) and two “nearly” perfect bilateral
gynandromoiphs basically half dark and half yellow
(Scriber & Evans 1988a, b). These perfect bilateral
gynandromoiphs were all offspring from interspecific
hybrid crosses. Furthermore, while the dark color of the
female half was expressed in the pairing of a dark Ohio
female with a P. alexiares Mexican male (Fig. 1), the
female half of the second gynandromoiph [a dark
female P. glaucus from South Carolina (with the b+
gene) mated with a P. canadensis male from Marinette
Co., Wisconsin] had the dark potential suppressed (Fig.
2; Scriber et al. 1996). The third gynandromoipih (a
yellow P glaucus from Pennsylvania mated with a P.
canadensis from Juneau Co., Wisconsin) had the female
half yellow because it likely lacked the Y-linked dark
gene from the mother and also probably received an X-
linked suppressor gene from the P canadensis father
(Scriber et al. 1996).

Such gynandromoiphs are not only fascinating in
appearance, but also with regard to physiology. Both of
these P. glaucus x P canadensis bilateral
gynandromoiphs emerged in the second year and did
not develop directly as did many of their “normal¬
looking” brothers. This reflects an X-linked obligate
diapause that is hemizygous in the female hybrids (and
the female half of these gynandromoiphs; XY or ZW),
but heterozygous in its sibling male hybrids (XX or
WW), which allows direct development and adult male
eclosion without diapause. Apparently the two halves of
both gynandromoiphs were cued to eclose as adults in
the next season rather than directly (at the 16:8 L:D
photophase, 23°C rearing conditions; Scriber & Evans
1988b).

Of the 9,000 or more field collected specimens
during this study (1979-1987; Scriber & Evans 1988a,
b) we never collected an obvious bilateral or nearly
bilateral gynandromoiph. The closest was one reared
from a field-collected female from Jasper Co. Texas
(Fig. 4). However, we collected many color mosaics
(dark and yellow patterns being the most obvious). In
fact, 33 of the 39 color mosaics were field-captured
individuals (6 were lab-reared from hand-paired
hybrids). While most of these field-collected color
mosaics were from Georgia, Ohio, Illinois, and Texas,
one family from a female captured in Dane Co.,
Wisconsin produced 17 of the color mosaics. It is
possible that some introgression from P canadensis

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39

Figs. 1-3. 1. A bilateral gynandromorph (from brood #4210) from a lab cross of a dark morph P. glaucus female (Adams Co., Ohio) mated
to a wild collected male P. alexiares garcia from Nuevo Leon, Mexico, (a) dorsal, (b) ventral. 2. A bilateral gynandromorph (from brood #628)
from a lab cross of a dark morph P. glaucus female (Pickens Co., South Carolina) mated to wild collected male ( P. canadensis from Marinette
Co., Wisconsin (collected 1-5 July 1983 by Don Caine), (a) dorsal, (b) ventral. 3. A gynandromorph, or sexual mosaic, which appears to be more
than 60% male (from brood #2830) from a dark morph P. glaucus female (reared from eggs obtained from a dark morph female wild collected
in Clay Co., West Virginia) bred to a wild collected P. rutulus male (reared from eggs obtained from a wild female collected in Sacramento Co.,
California), (a) dorsal, (b) ventral.

could have occurred in this geographic area since it is
very near the hybrid zone (Luebke et al. 1988). In fact,
the famous 19th century Strecker collection (Strecker
1878; Walsten 1977; Ehle 1981; Shapiro 1981a, b;
Clarke & Clarke 1983) of P. glaucus color (or sexual)
mosaics (which had been on loan from the Chicago
Field Museum to the Allyn Museum) may reflect such
introgression in those mosaics near the Papilio glaucus/
P. canadensis hybrid zone in the Midwest and
Pennsylvania (Scriber et al. 2003). Here we review our
hypothesis in light of new information.

We reared these hybrid larvae only through to
pupation on black cherry (Primus serotina) leaves, since
this host is used by both P. canadensis and P. glaucus
(and by all four other species in the P. glaucus group;
Scriber 1996). If we had attempted to rear these
gynandromorphs on tulip tree, Liriodendron tulipfera
(this P. glaucus favorite is toxic to P. canadensis; Scriber
1982) or on quaking aspen, Populus trenudoides (this P.
canadensis favorite is toxic to P. glaucus, Lindroth et al.

1988), these insects might have might have survived to
become adults, as normal hybrids, since detoxification
enzymes are inherited autosomally, and are used inside
the gut, they can be expressed throughout the insect
regardless of whether they are produced by one or both
halves (male or female) of the gynandromorphs (Scriber
1986; Scriber et al. 1989).

Results

We collected a female Papilio canadensis from
Cheboygan Co., Michigan during the period 9-12 June
1997. Larvae were reared on black cherry. One of the
resulting adults was a nearly perfect bilateral
gynandromoiph with blue hindwing color and female
genitalia on the right, and with male genitalia and
coloration on the left (Fig. 5).

Another near-perfect bilateral gynandromorph was
actually collected in the field on 15 June 1996 in
southwestern Vermont near the New York border
(II. R.). This specimen has basically female blue

40

Journal of the Lepidopterists’ Society

Figs. 6-8. 6. A bilateral gynandromorph of P. canadensis collected (H.R.) in Bennington Co., Vermont on 15 June 1996 (a) dorsal, (b)
ventral. 7. A bilateral gynandromorph collected (H.R.) on 3 June 1989 in Lincoln Co., Wyoming. Possibly P. rutulus female (left half) and P.
multicaudatus male (right half). 8. A hybrid (lab-paired, lab-reared) of a P. rutulus female x P. multicaudatus male (brood #7806).

5 (enlarged)

Figs. 4-5. 4. A nearly perfect gynandromorph or sexual mosaic (from brood #1091) from a wild collected dark morph ( P. glaucus female
from Jasper Co., Texas, (a) dorsal, (b) ventral. 5. A bilateral gynandromorph lab-reared from a family that was derived from a female P.
canadensis collected in Charlevoix Co., Michigan, 9-12 June 1997 (a) dorsal, (b) close-up (In collection of Mark Deering).

Volume 63, Number 1

41

coloration on the right hindwing and slightly heavier
(wider) black bands (as in females) on the right
forewings (Fig. 6; this specimen appears to possess 2
male claspers and no ovipositor).

The third nearly perfect bilateral gynandromorph is
one collected from Lincoln Co. (Wyoming) on 3 June
1989. This individual appears to be half female P.
rutulus and half male P. multicaudatus (Fig. 7). Both
species lack the dark morph females and probably the Y-
linked b+ gene (Seriber et al. 1990, 1996). The right
hindwing has the extra “tail " and the right forewing
appears somewhat similar to P. multicaudatus in
banding. The left female half looks rutulus- like. For
comparison, we also present a female from an
interspecific lab pairing of P. rutulus female x P.
multicaudatus male (Fig. 8, brood #7806). In this
specimen, the two species traits appear somewhat
intermediate throughout and did not segregate into the
two halves (one female and one male, as in the
interspecific gynandromorph; Fig. 7).

A field-captured, nearly-perfect bilateral
gynandromorph between the nymphalids Limenitis
arthemis arthemis (Drury) and L. a. astyanax (F.) was
captured in the same area of the southwestern
Vermont/New York border as the P. canadensis
gynandromoiph (not figured) and may be part of an
active hybrid zone. These Papilio and Limenitis species
(sub-species) complexes have significant introgression
and exhibit centers within their hybrid zones in New
England, as they do across the Midwest in Michigan and
Wisconsin (Platt & Brower 1968; Seriber 1996).

We have also lab-reared a dark nearly perfect bilateral
gynandromorph from a dark, east Texas P. glaucus
mother (brood #8858, Janice Bossart). A specimen
photo sent to us by Loretta Lutman of Ashboro, North
Carolina (Randolf Co.) also appears to be a wild,
bilateral gynandromoiph of P. glaucus, with the female
half dark (Fig 9).

Perfect bilateral gynandromorphs, with half
expressing male traits and the other female traits, are
extremely rare. Color/sexual mosaics with asymmetrical
blotches or patterns of color (or other mixed gender
traits) are somewhat more common than bilateral
gynandromorphs among these swallowtails, but they are
still quite rare. Earlier studies with nearly 30,000 lab-
reared North American Papilio resulted in 30-40 color
mosaics (approximately 0.001 % of reared adults was an
obvious color and/or sexual mosaic; Seriber & Evans
1988b). It is easiest to find color (or sexual) mosaics in P.
glaucus when the female is of the dark morph, but the
hindwing blue color of females is also helpful in spotting
them (Fig. 10). One thing that is clearly characteristic of
color mosaics is that the dorsal and ventral patterns are

determined independently; a few examples are
illustrated in Figs. 11-16. Another mosaic from Illinois
shows asymmetry (Fig. 17). It is also possible to have
bilateral differentiation as in size or color for the same
sex and single species individual larvae, pupae, and
adults (see Seriber & Hagen 1990).

Discussion

We describe six new and nearly perfect bilateral
gynandromoiphs from field collections in northern
Michigan, southwestern Vermont, Montana and Texas.
Four of these are likely to represent interspecific or
intersubspecific hybrids (Figs. 5-7, and the Limenitis
from Bennington Co. Vermont, collected by HR). While
perfect and near-perfect bilateral gynandromorphs
appear to be often generated by interspecific
hybridization, we report two gynandromoiph specimens
from Texas and North Carolina (Fig. 9) and East Texas
(]. Bossart and JMS, not figured here) that were not
near the P. canadensis/ glaucus hybrid zone.

The somewhat unique ability of individual butterflies
with one half male and one half female to develop as
larvae, pupae, and live adults capable of flight (e.g.
Seriber et al. 1995; Conners 2000) is fascinating, but still
poorly studied and understood. The dramatic
differences in wing color with partial dark morph
females and yellow-patterned males enables us to easily
detect individual bilateral gynandromorphs or
asymmetrical sexual mosaics, but the yellow morph
female gynandromorphs of tiger swallowtail butterflies
are no less fascinating.

Here we have presented one example of interspecific
bilateral gynandromoiphs that were bred from Papilio
glaucus female and P. canadensis male parents (Fig. 2;
see also Seriber et al. 1995). This interspecific pairing
united genes in the resulting gynandromorph that have
segregated into the two gender-differentiated halves
with the heterogametic female [XY (=ZW)] possessing
the Y-linked black (b+) gene of dark morph females plus
the X-linked suppressor (s+; Seriber et al. 1996),
resulting in the yellow (suppressed) tiger pattern for the
female half (Fig. 2). This contrasts with the interspecific
(P. glaucus x P. alexiares) gynandromoiph with the dark
melanic (b+) gene and no suppressor (s', an enabler),
where the female half is black (Fig 1).

The biochemical pathways (temporally and spatially
regulated by enzymes, including dopa carboxylase) are
apparently coordinated by multiple enzyme activities of
the melanin (black) and papilioehrome (yellow) specific
pathways. These enzyme regulations must be conducted
simultaneously in the same gynandromorphs (Koch et
al. 2000a), all of which is coordinated by the single
major Y-linked factor (Seriber et al. 1996) on the female

42

Journal of the Lepidopterists’ Society

Figs. 9-11. 9. A natural bilateral gynandromorph of P. glaucus from Randolph Co., North Carolina (photo from Loretta Lutman). 10. A
sexual mosaic (gynandromoiph) with mostly male characters (except the blue coloration in the left hindwing) brood 2025, from a wild-collected
yellow female, Clark Co., Georgia. 11. A color/sexual mosaic (lab-reared from a female collected in Ohio, 1989 (a) dorsal, (b) ventral.

Figs. 12-13. 12. A gynandromoiph or sexual mosaic, which appears to be more than 60% female (from brood #3622) from a pure P. glaucus
lineage (in which the mother was the daughter of a dark morph female collected in Hocking Co., Ohio and mated to a male from Adams Co.,
Ohio), (a) dorsal, (b) ventral. 13. A gynandromorph or sexual mosaic, which appears to be more than 75% female but with male claspers (from
brood #3935) from a wild collected dark morph P. glaucus female (Scioto Co., Ohio), (a) dorsal, (b) ventral.

Volume 63, Number 1

43

side of the individual; but these genes are either not
expressed, or not present, on the male half oi the
individual. The activity of other sex-linked traits such as
obligate diapause regulators would explain the direct
development (non-diapause) of both halves of the P.
glaucus/P. alexiares gynandromorph and the one year
delay (diapause) of the P. glaucus/P. canadensis
gynandromorph (with both the hemizygous female and
the heterozygous male having the X-linked obligate
diapause gene, od+). Other sex-linked traits such as
opposition preferences could not be analyzed, but in
the future researchers could examine the sex-linked
diagnostic allozymes and mitochondrial DNA variation
with regard to distribution in the two halves of such
gynandromorphs.

In melanic females, black melanin replaces the
background yellow' papilioehrome (Koch et al. 1998).
The key enzyme involved is BAS (n -(5- alanyl-
dopamine-synthase), which shunts (suppresses)
dopamine from the melanin pathway into the
production of the yellow papilioehrome pigment
(probably in response to dosage of the Z (=Y)-linked b+
gene products; Scriber et al. 1996). The mixed mosaics
we figure here appear to reflect the fact that melanism
is both sex-specific and pattern-specific (cell
autonomous), and therefore melanic specimens of P.
glaucus are not just null mutants as in some ebony
Drosophila strains (Koch et al. 2000b). The
suppressor/enabler gene (s'/s+) that permits dark female
expression of the b+ melanin gene has been identified
on the Z (=X) chromosome (Scriber et al 1996), and
may involve tyrosine hydroxylase, or linked factors that
are involved in production of dopa (common to both
melanin and papilioehrome pathways) (Koch et al.
2000a, b). The late incorporation of tyrosine into
melanin may also be involved in the incomplete
penetrance (where yellow scales are brownish)
suggesting that sex-linked control genes and their sex-
linked modifiers exert their effects by differential
interpretation of the patterning signal (e.g. the delay in
development of melanic scales is female-specific)
(Ffrench-Constant & Koch 2003).

The specific melanin (b+) modifier genes (s+/s') on the
Z-chromosome, remain to be identified, but we have
isolated the tyrosine hydroxylase gene from the Z-
chromosome off! glaucus (Putnam et al. 2007). The Z-
chromosome also carries other genes of major
significance in this species, including oviposition
preferences (Scriber 1993; Mercader & Scriber 2007;
Mercader et al. 2009), diapause regulation (Rockey et al
1987a, b), post-diapause emergence delays (Scriber et
al. 2008), and allozymes (lactate dehydrogenase (Ldh),
and phosphoglueonate dehydrogenase (Pgd)), which

appear to be species-diagnostic for P. glaucus, P.
canadensis, and P. appalachiensis (Hagen et al. 1991;
Scriber & Ording 2005; Scriber et al. 2008). The genetic
incompatibilities of some recombinant hybrid genotypes
appear to involve the Haldane effect in the
heterogametic female pupae (Hagen & Scriber 1995).
The delayed emergence of recombinant hybrid swarms
(P. appalachiensis: Pavulaan & Wright 2002; Scriber &
Ording 2005) may be controlled by the Z-linked
obligate diapause gene (od+; Scriber et al. 2008) or a
closely linked post-diapause developmental rate
controlling gene on this Z-chromosome, such as
identified for the delayed mid-July flight of European
corn borer (Dopman et al. 2005). The tpi allozyme locus
appears to correspond closely with this factor in the
corn borer, and we have determined that it is also Z-
linked in P. glaucus/P. canadensis (Hagen & Scriber
1989; Scriber 1994).

Females of P. glaucus that are intermediate in their
expression of melanism (with different degrees of
“dusted” or “cinnamon” appearance; Fig 18) can be a
result of rearing and/or pupal eelosion at higher
temperatures (Ritland 1986). Such intermediate
melanism may reflect variable penetrance, suggesting
that the black melanin is superimposed over the yellow
papilioehrome (Koch et al. 1998), which may appear
brown in color (Ffrench-Constant & Koch 2003).
However, we have also seen such intermediate
melanism in hybrid daughters of dark females mated to
P. multicaudatus (Fig 9), to P. rutulus (Scriber et al.
1995), and to P. canadensis (from Dane Co. WI, not
figured here). Most P. canadensis males have X-linked
suppressors (s+) that completely prevent the expression
of melanism in hybrid daughters of dark mothers
(Scriber et al. 1996). Here we report the partial
expression of melanism (“cinnamon" with striations) in
daughters of a primary F-l hybrid pairing between a
dark P. glaucus mother mated with a male P. canadensis
from central Michigan (in Isabella Co. on the cooler
side of the historical hybrid zone; see below). These are
not color mosaics as described earlier. These
“cinnamon” intermediates of this Michigan P.
canadensis father and dark P. glaucus mother, reared
and emerged at 22° C, were also not due to high
temperatures (28-30° C) as in Fig. 18 (see Ritland
1986).

This result suggests that the wild Isabella Co. male
parent of this pairing (#20011) had at least one Z-linked
enabler (s ), which permitted expression of the Y-linked
dark gene (b+). The introgression of the X-linked
enabler has also been documented in daughters of
another Isabella Co. P. canadensis male mated to a dark
P. glaucus mother (#20015; Scriber et al. in prep.) and

44

Journal of the Lepidopterists’ Society

Figs. 14-15. 14. A gyriandromorph or sexual mosaic which appears to be more than 50% female (from brood #4196) from a lab cross of a
dark morph P. glaucus female (Champaign Co., Illinois) mated to a hybrid male (mother of which was the daughter of a dark morph P. glaucus
female collected in Georgia and whose father was a P. canadensis collected in Lincoln Co., Wisconsin, (a) dorsal, (b) ventral. 15. A female color
mosaic (from brood #3770) from a dark morph P. glaucus female (the daughter of a wild dark morph P. glaucus female, #3104, collected in
Athens Co., Georgia) mated to a wild P. canadensis male (collected in Tompkins Co., New York), (a) dorsal, (b) ventral.

Figs. 16-18. 16. A field-reared P. glaucus from Lancaster Co. PA 2004 (pupa from Win. Houtz). 17. A specimen from David and Marc
Perlman (photo provided, pers. comm, to JMS). 18. An intermediate “dusty” or “cinnamon” type reared from Dane Co., Wisconsin at high
temperatures (28-30° C).

Volume 63, Number 1

45

also in daughters of several males of the hybrid swarm
late flight at the New York/Vermont border where no
dark females (with the W-linked b+ gene for melanism;
Seriber et al 1996) have ever been collected (the nearest
collection of a dark female is roughly 150 miles south;
Seriber et al. in prep). The Z-linked enabler has also
been detected in males captured from the hybrid swarm
of late flight individuals in P. appalachiensis populations
from West Virginia, but here, on the warmer side of the
hybrid zone (Seriber et al. 2009b), the W-linked gene
for melanism (b+) does also occur as evidenced by
recombinant hybrid-like dark females (Pavulaan &
Wright 2004; Seriber & Ording 2005; Seriber et al.
2009a, b).

Individual species diagnostic traits moving northward
across the historical hybrid zone include: 1) autosomal
tulip tree detoxification (Seriber 1986, 2002; Seriber et
al. 1999), 2) several wing color traits (Luebke et al.
1988), 3) some X-linked traits such as the allozyme allele
Pgd-100 (Hagen & Seriber 1989; Hagen et al. 1991;
Seriber et al. 2008), and 4) opposition preferences
(Seriber et al. 1991; Mercader & Seriber 2007;
Mercader et al. 2009). However, other X-linked species
diagnostic traits (including Ldh-100 allozyme alleles and
bivoltine potential (ock); Rockey et al. 1987) as well as
the Y-linked dark gene (b+) and certain mitochondrial
DNA alleles have not moved northward across the
historical hybrid zone (Stump et al. 2003; Seriber et al.
2008). The divergent selection pressures on different
parts of the X-chromosome (and differential movement
of these parts) in the genomes of both P. glaucus and P.
canadensis may be historically old (Putnam et al. 2007).
However, recent origins of interspecific introgression
and recombination combined with strong divergent
selection on different regions of the X-chromosome may
have been catalyzed by recent climate warming,
possibly leading to the Mountain Swallowtail (potential
hybrid species), P. appalachiensis (Pavulaan & Wright
2002; Seriber & Ording 2005). We will continue to
investigate the linkage relationships of the dark morph
enabler gene (s-) with other parts of the X-chromosome
and how it may be involved in the easily detected sexual
color mosaics.

Acknowledgements

This research was supported in part by the College of Natural
Science, and the Michigan Agricultural Experiment Station
(Project MICL #01644), NSF grants (DEB 9220122, DEB
9981608, DEB 0716683), and the Ray and Bernice Hutson En¬
dowment of the Department of Entomology. We thank Matt
Ayres, Janice Bossart, Jessica Beering, Robert Dowell, Cheryl
Frankfater, Robert Hagen, Robert Lederhouse, Matt Lehnert,
Jim Maudsley, James Nitao, Michelle Oberlin, and Patti Tidwell
for assistance in the field and lab. Thanks are extended to David
and Marc Perlman and Joseph Zeligs for their color mosaic pho¬

tos and to Loretta Lutman for the photo of the wild bilateral gy-

nandromorph. We thank Austin P. Platt and Charles V. Covell

for very helpful reviews.

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Received for publication 26 February 2008; revised and accepted 9
December 2008.

48

General Notes

Journal of the Lepidopterists’ Society

Journal of the Lepidopterists’ Society
63(1), 2009, 48-51

PITHING AND MINING BY A PUNKIE: THE UNUSUAL FEEDING STRATEGIES OF NOLA
CILICOIDES (GROTE, 1873) (NOCTUIDAE: NOLINAE)

Additional key words: Lysirnachia ciliata, leafmining, plant defense, hypertrophied tactile setae, leaf shelter

Herbivorous insects have evolved innumerable
strategies for coping with plant defenses. Among these
are efforts made by insects to incapacitate a plants ability
to transport defensive substances through its vascular or
lactifer systems to sites of feeding injury. One means of
doing such is to trench, vein cut, or otherwise breach the
delivery systems of the host so that milky latexes, resins,
and gums cannot be transported to the feeding site
(Dussourd & Eisner 1987; Dussourd 1993). Another
strategy is to girdle the host, either as an adult (e.g.,
ovipositing females) or as a larva (Forcella 1982;
Dussourd 1993; Hanks 1999; Albanese et al. 2007). A
third option is mass assault: bark beetles are a classic
example. Adults attack host trees in such numbers that
individuals under siege cannot individually “pitch out" all
the invading beetles (Wood 1982). Here we report an
example of a fourth option available to herbivores for
thwarting a host’s defensive system: the hollowing
(pithing) of non-woody stem tissues.

In early July 2007 we found young (approximately 3rd
instar) larvae of Nola cilicoides (Grote) (Noctuidae:
Nolinae) pithing upper stems of fringed loosestrife
(. Lysirnachia ciliata L.) (Primulaceae). Larvae tunneled
into the side of new shoots, removed pith and associated
vascular tissues, backed out, then crawled further up the
compromised stem to feed on flowers and young leaves.

Nola cilicoides (Fig. 1) occurs from southern Canada
to northern Florida and west to Arkansas (Forbes I960;
Handheld 1999; Ferge & Balogh 2000; Coveil 2005). In
the Northeast the moth is local and uncommon in
swamps, floodplains, wet meadows, damp woods, and
other mesic to wet community types. Dyar (1899) did
not provide life history information for N. cilicoides in his
study of the early stages of Nearctic nolines. McCabe, in
Rings et al. (1992), reported that the larva of N. cilicoides
feeds on new leaves and flowers of fringed loosestrife
( Lysirnachia ciliata). Collection dates for adults in
Connecticut and New York normally range from mid
June into July, with the occasional specimen taken as
early as late May or as late as August and September —
we are unsure of the number of generations. In Georgia
there are least three full broods (James Adams pers.
comm.).

Study Site and Methods. Observations were made

on a colony of N. cilicoides located in Lebanon,
Connecticut along a powerline cut south of Burnham
Road and north of Route 87 (N41° 39’ 15.5” W72° 14’
07.6”). Rearing was done in plastic bags and 40 dram
plastic vials. Laivae were reared at 21-23° C with
photoperiods set to match those outdoors in central
Connecticut. Larval and adult vouchers and digital
images have been deposited at the University of
Connecticut.

Results. On 6 and 11 July, 2007, young laivae of N.
cilicoides were found feeding on the shoots of fringed
loosestrife plants that were coming into flower. Laivae
were located by surveying for loosestrife plants with
flagging shoots. Each flagged shoot had a small entry
hole and small area of cored pith, 5-10 centimeters
below the shoot apex (Figs. 2, 4). On each flagged plant,
we located 1 or 2 approximately 3rd instar N. cilicioides
feeding externally (Fig. 3), distal from the wound, on
floral buds, flowers, and new leaves. Laivae showed a
preference for buds and flowers (Figs. 6, 7) over leaves.
Some larvae tied together adjacent leaves and flowers,
forming a weak shelter, but silk deposition was sparse
and their shelters were poorly defined.

Laival densities were low; only five caterpillars were
found on roughly 200 plus fringed loosestrife stems that
were searched in early July. In the vicinity of the colony
and at other locations in New London and Tolland
counties, we searched for, but did not find, laivae on
either swamp loosestrife [L. terrestris (L.) Britton,
Stems, & Poggenb.j or whorled loosestrife (L.
quadrifolia L.).

On 9 and 19 August, 2007, we returned to the same
stand of fringed loosestrife in Lebanon, ostensibly to
look for cocoons. While we failed to find cocoons on
more than 60 checked stems, and any evidence on newly
bored shoots, we discovered something surprising:
young Nola cilicoides larvae, 2-3 mm in length mining
leaves of fringed loosestrife. Laivae had entered
through the lower epidermis, often adjacent to a vein,
and then tunneled into the lamina, removing
parenchymatic tissues, leaving a full depth blotch mine
(Fig. 5). All but one mine had a single entrance/exit hole
on the lower leaf surface; the one exception had three
entry holes. Dark green to black frass collected in the

Volume 63, Number 1

49

Figs. 1—9. Nola cilicoides (Grote), all Lebanon, New London Co., Connecticut except adult from Whitefield, New Hampshire.
1. Adult female. 2. Shoot damaged by single larva (arrows point to entry points). Note wilted section of stem above bore hole. 3.
Middle instars feeding on young leaves. 4. Detail of 2. 5. Occupied leafmine. 6. Last instar “coring” flower bud with head fully in¬
serted. 7. Last instar feeding on remains of what was previously an open flower. 8. Last instar. While not fully visible, SD setae ex¬
tend close to all four edges of the image (see text). 9. Cocoon under construction, upper end with abdominal terminus visible. 10.
Finished cocoon. Note discolored area of stem above and below cocoon where bark strips were removed.

50

Journal of the Lepidopterists’ Society

mine, i.e., little had been expelled. The smallest mine
located, was only a few mm in diameter and contained
the cadaver of a larva 1.25 mm in length.

In the wild, we found approximately 3rd instar larvae
grazing on patches of parenchyma from leaf
undersides — scraping away small patches of leaf tissue
without feeding through to the upper leaf surface — as
late as mid September, indicating that at least some N.
cilicoides larvae vacate their mines to feed externally,
before the onset of winter.

Beginning as early as late August, our captive (lab-
reared larvae) spun small, circular to oval, cocoons
between overlapping leaves. Cocooned larvae were
foreshortened, only 2. 6-2. 8 mm in length, and evidently
had entered a non-feeding diapause in which they
would pass the winter. [Meganola spodia Franclemont,
an oak-feeding noline, also overwinters as a partially
grown larva (Butler 1989).]

The full grown larva is 10-12 mm in length (n=8). It
is typical for a noline: the prolegs are absent on A3 and
secondary setae are present, grouped onto five (dorsal,
subdorsal, lateral, subventral, and ventral) warts. Both
the dorsal and subdorsal warts have an incomplete black
ring around their bases (Figs. 6, 7). The ground color is
dirty yellow-orange above, more yellow laterad, and
mostly unpigmented below the subspiracular stripe; the
creamy to pale orange middorsal stripe is weak over Tl.
Snbdorsal warts on T1-T3 and A7-A9 bear one
exceptionally long seta that extends well beyond the
body (see below). The head, prominently marked with
black (Fig. 7), is drawn into the prothorax at rest.

|nly-collected, lab-reared larvae formed their pupal
cocoons along a stem (Figs. 9, 10), but extensive
searches of plants in the wild, including those with
wilted tips, did not yield any cocoons. The central
chamber of cocoons measured roughly 3 mm x 10 mm.
The lower end of the cocoons had an additional 5 nun-
long ‘‘tail’' that tapered downward; the upper end was
higher in profile (n=3) (Figs. 9, 10). Like other nolines,
numerous strips of stem epidermis "bark" were
collected and then woven into the wall of the cocoon.
Our caterpillars positioned more than 40 narrow strips
of bark, removed from adjacent areas of the stem, into
their cocoon walls. Eclosion occurred 1-2 weeks after
pupation.

Discussion. The greatly elongated setae that extend
from the SD warts at either end of the body of Nola
larvae (Fig. 8 and caption) presumably sen e as an "early
warning" system for the larvae. Those issuing from T3
are the longest: in one individual, the longest SD seta
extended forward 5.8 mm on a larva only 10 mm in
overall length. The rearward-projecting SD seta on A9
of this same caterpillar measured 3.6 mm. In addition.

the subdorsal warts on these segments are oriented in
such a way (in living larvae) that the hypertrophied setae
are directed well forward on the thoracic and rearward
on the abdominal segments. Analogous long dorsal or
subdorsal setae also have been reported from
ehoreutids (e.g., Brenthia ), pyralids (e.g., Monoloxis), as
well as other noctuids (e.g., Aboblemma Walker) (Aiello
& Solis 2003; Rota 2003, 2008; Wagner et al. 2008; Rota
& Wagner 2008).

Although Lijsimcichia ciliata has not been
investigated, the tissues of other Lijsimachia species
have been found to contain a variety of bioactive
compounds including flavonoids, triterpenoid saponins,
and phenolic acids (Chang et al. 2004). To what degree
these are transportable or serve in defense remains
unstudied. Our thesis presupposes that N. cilicoides
larvae tunnel out the pith of their hostplant to
incapacitate its vascular tissues — we acknowledge that
the pithing behaviors reported here may have little to do
with circumventing the plant’s ability to translocate
antiherbivory substances. Perhaps pithing and the
subsequent flagging help the larva to construct its leaf
shelter, prevent subsequent growth and maturation of
the shoot, enhance survivorship from attacks by natural
enemies, or in some other way improve or enhance
larval performance or survival. Regardless, the
stereotypical behavior is commonly expressed, and stem
boring and bud coring represent unique traits for
Nearetic nolines, which tend to be rather unspecialized
leaf feeders (Forbes I960; Wagner 2005; Wagner et al.
2010).

As noted above, larvae preferentially sought out
flower buds and flowers over leaf tissues. A preference
for reproductive tissues has been reported for other
nolines. Nola sorghiella Riley feeds on the seed heads
of sorghum and various other grasses and appears to be
completely tied to reproductive tissues (our wild-
collected larvae failed when offered only leaves). In the
Old World, spiny bollworms of the genus Earias are
major pests of cotton flowers (Pinhey 1975; Kitching &
Rawlins 1998).

More smprising was the finding that N. cilicioides
larvae engage in leaf mining. Leafmining is rare among
noctuids. In the East only a handful of apaineines mine,
and most of these only for the first one or two instars
(■Wagner et al. 2010; Eric Quinter pers. com¬
munication). It is exceptional for a "leafminer" to bear
secondary setae; more commonly, setae tend to be
reduced or absent among leafmining clades (cf. Stehr
1987). We did not observe leafmining behaviors prior to
our August visits. Interestingly, life histories with stem
mining in spring generations and leafmining in late
summer generations have been reported for a number

Volume 63, Number 1

51

of European Momphidae (Koster 2002). In North
America, Mompha solomonella Adamski & Brown,
tunnels in shoots of Cephalanthus accident alio L.
(Rubiaceae) in the spring, but second and subsequent
generations mine leaves (Wagner et al. 2004).

Much remains to be elucidated about the biology
Nola cilicoides. As noted, the selective advantages of
pithing and ultimate explanation for this exceptional
behavior have yet to be determined. Both observational
and controlled studies are needed to better understand
the roles of the hypertrophied tactile setae of Nola and
other lepidopterans. We are also in a quandary about
the phenology and broodism of N. cilicoides in New
England. We were only able document the existence of
a single brood — one that produced its pithing larvae in
early July. Either we missed a small spring brood, or the
partially grown larvae that spun overwintering cocoons
in August, waited until early July of the following season
to break their diapause.

Acknowledgments

Tim McCabe, as so many times before, guided DLW’s initial
efforts. Deane Bowers, Dave Dussourd, Laura McLoud, Jessica
McShane, April Rodd, and Bo Sullivan read drafts of the paper.
An anonymous reviewer offered many helpful suggestions. An¬
drea Farr assembled the plates. Both authors were supported
on a grant from the CT-Department of Environmental Protec¬
tion Wildlife Division through the State Wildlife Grants Pro¬
gram; additional support was provided by Northeast Utilities.

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- . D. Adamski, & R. L. Brown. 2004. A new species of Mompha

(Lepidoptera: Coleophoridae: Momphinae) from buttonbush
(Cephalanthus occidentals L.) with descriptions of the early
stages. Proc. Entomol. Soc. Wash. 106: 1-18.

- , J. Rota, & T. L. McCabe. 2008. Larva of Abablemma Nye (Lep¬
idoptera: Noctuidae: Hvpenodinae) with notes on lichenivory and
algivory in Macrolepidoptera. Ann. Entomol. Soc. Amer. 101:
40-52.'

- . D. F. Schweitzer, J. B. Sullivan & R.C. Reardon. 2010.

Owlet cateqiillars of eastern North American (Lepidoptera: Noc¬
tuidae). Princeton Univ. Press. In review.

Wood, D. L. 1982. The role of pheromones, kairomones, and al¬
lomones in the host selection and colonization behavior of bark
beetles. Ann. Rev. Entomol. 27: 411-446.

David L. Wagner, Department of Ecology and
Evolutionary Biology, University of Connecticut, Storrs,
Connecticut 06269, USA, e-mail: clavid. ivagner
@uconn.edu and Bryan Connolly, Department of
Ecology and Evolutionary Biology, University of
Connecticut, Storrs, Connecticut 06269, USA.

Received for publication 6 Nov 2007; revised and accepted
28 May 2008

52

Obituary

Journal of the Lepidopterists’ Society

Journal of the Lepidopterists’ Society
63(1), 2009, 52-57

EUGENE G. MUNROE

(8 September 1919-31 May 2008)

I recently wrote an obituary for Eugene M unroe for
the Bulletin of the Entomogical Society of Canada
[2008. 40 (1): 142-146], Unfortunately there was not
enough space for his list of publications. In 2000, I
wrote a detailed account of Genes life and professional
career for a Festschrift in Tropical Lepidoptera
[11: 1-6]; the list of publications was incomplete. Gene’s
impressive publication list is published in its entirety
here. See also the 2008 Pyraloid Planet [2:2] for
reminisces by colleagues. Michael Shaffer, The Natural
History Museum, London, generously reviewed this
version of Gene’s publications.

To learn more about Gene’s life, accomplishments,
enthusiasm for research, and love of Lepidoptera, refer
to the publications mentioned above. Here, I would
like to briefly share some historical information
regarding the Lepidopterists’ Society and Gene. He
was a Charter and Honorary Life Member of the
Lepidopterists1 Society. In preparation for the First
Annual Lepidopterists’ Society Meeting, Dec.
29th-30th, 1950, held at the American Museum of
Natural History, New York, the President, ]. H.
McDunnough, appointed three committees. Program,
Local Arrangements and Nominating. Gene was
appointed member of the Nominating Committee.
Also, at the first meeting Gene participated (1951e) in a
symposium entitled “Geographic subspeciation in the
Lepidoptera” where Charles Remington also presented
the introduction. Gene was involved in the first annual
Season Summary as a coordinator (1947b, 1949b, c,
1951b) for Area 7, the Northeast, and later as editor-
(1951d) of the Season Summary. He served as
President of the society in 1958. In 2004 Gene received
the Karl Jordan medal from the Lepidopterists’ Society.

Gene Munroe passed away 31 May 2008 at the age of
89 in Ontario, Canada.

List of Publications

Munroe, E. G. 1942. The Caribbean races of Anartia jatrophae Jo¬
hansson (Lepidoptera: Nymphalidae). American Museum Novi-
tates 1179: 1-4.

_ . 1947a. Four new Pieridae from the West Indies. American

Museum Novitates 1362: 1-5.

_ (coordinator). 1947b. The field summary of Lepidoptera - 1947

season - 7, Northeast-Maryland north to Quebec and Newfound¬
land. Lepidopterists' News 1: 95.

_ . 1948a. The geographical distribution of butterflies in the West

Indies. Ph.D. thesis. Cornell University, Ithaca, New York.

_ . 1948b. Further North American records of Acentropus niveus

(Lepidoptera: Pyralidae). Canadian Entomologist 79: 120.

_ . 1949a. Some remarks on the genus concept in Rhopalocera.

Lepidopterists’ News 3: 3-4.

_ (coordinator). 1949b. The field season summary of North

American Lepidoptera for 1949, Area 7, Northeast-Maryland
north to southern Quebec. Lepidopterists' News 3: 97-101.

_ (coordinator). 1949c. Field season summary of North American

Lepidoptera for 1948, Area 7, Northeast. Lepidopterists’ News 8
(supplement): 9-11.

_ . 1949d. An unnoticed character in the Satumioidea. Entomo¬
logical News 60: 60-65.

_ . 1949e. Some remarks on the orithija group of the genus

Junonia (Lepidoptera: Nymphalidae). Entomologist 82: 157-158.

_ . 1949f. A new genus of Nymphalidae and its affinities (Lepi¬
doptera: Rhopalocera). Journal of the New York Entomological
Society 57: 67-78.

_ . 1949g. Notes on fish of the interior of the Labrador Peninsula.

Arctic 2: 165-173.

_ . 1950a. The occurrence of a butterfly in the Pribilof Islands.

Lepidopterists’ News 4: 44.

_ . 1950b. The generic positions of some North American Lepi¬
doptera commonly referred to Pyrausta Schrank. (Lepidoptera:
Pyralidae). Canadian Entomologist 82: 217-231.

_ . 1950c. Subventral tubercles of Saturnoid larvae — a supple¬
mentary note. Entomological News 61: 39-41.

_ . 1950d. The dina group of the genus Eurema in the West Indies

(Lepidoptera: Pieridae). Journal of the New York Entomological
Society 58: 172-191.

_ . 1950e. The systematic^ of Calisto (Lepidoptera: Satyrinae),

with remarks on the evolutionary and zoogeographic significance
of the genus. Journal of the New York Entomological Society 58:
211-240.

_ . 1951a. Field notes on the butterflies of Knob Lake, northern

Quebec. Lepidopterists’ News 5: 7-9.

_ (coordinator). 1951b. Field season summary of North American

Lepidoptera for 1950, Area 7, Northeast-Delaware & Pennsylva¬
nia north to southern Quebec. Lepidopterists’ News 4: 102-106.

_ . 1951c. The De Rabie paintings of Lepidoptera in the Blacker

Library of Zoology, McGill University, with notes on the butter¬
flies represented therein. Lepidopterists' News 5: 55-57.

_ (ed. ). 195 Id. The field season summary of North American

Lepidoptera for 1951. Lepidopterists’ News 5: 81-110.

_ . 1951e. V. Subspeciation in the Microlepidoptera. Lepidopter¬
ists’ News 5: 29-31.

_ . 1951f. A previously unrecognized species of Nymphula (Lepi¬
doptera: Pyralidae). Canadian Entomologist 83: 20-23.

_ . 1951g. The identity and generic position of Chauliodes dis-

junctus Walker (Megaloptera: Corydalidae). Canadian Entomol¬
ogist 83: 33-36.

_ . 1 95 1 h . Pest Trichoptera at Fort Erie, Ontario. Canadian Ento¬
mologist 83: 69-72.

_ . 1 95 1 i . North American Pvraustinae: notes and descriptions

(Lepidoptera: Pyralidae). Canadian Entomologist 83: 161-169.

_ . 195lj. The geographic variations of Dasyuris polata

(Duponchel) in North America. (Lepidoptera: Geometridae).
Canadian Entomologist 83: 290-294.

_ . 1951k. The genus Junonia in the West Indies (Lepidoptera:

Nymphalidae). American Museum Novitates 1498: 1—16.

Miller, M. J. & E. G. Munroe. 1951. Schistosome dermatitis in
Quebec. Canadian Medical Association Journal 65: 571.

Munroe, E. G. 1952a. The illibalis group of the genus Palpita Hiib-
ner (Lepidoptera: Pyralidae). Canadian Entomologist 84: 43-55.

Volume 63, Number 1

53

_ . 1952b. Support for the proposals submitted by Mr John G.

Franclemont in regard to the generic name “Phalaena” Linnaeus,
1758, and the names published by Linnaeus for groups of that
genus (Class Insecta, Order Lepidoptera). Bulletin of Zoological
Nomenclature 6: 317.

_ . 1952c. Support for the proposed use of the plenary powers to

validate the trivial name “adippe” (as published in the combina¬
tion “ Papilio adippe”) as from Denis and Schiffermiiller, 1775
(Class Insecta. Order Lepidoptera). Bulletin of Zoological
Nomenclature 9: 136.

_ . 1952d. Comment on Mr. C. P. dos Passos proposal relating to

the trivial name “ plexippus ” Linnaeus, 1758, as published in the
combination “ Papilio pexippus” (Class Insecta, Order Lepi¬
doptera). Bulletin of Zoological Nomenclature 9: 139-140.

_ . 1953a. The Karanasa butterflies — a critical review. Lepi-

dopterists’ News 7: 16- 23.

_ (ed.). 1953b. Field season summary of North American Lepi¬
doptera for 1953. Lepidopterists' News 7: 66-118.

_ . 1953b. Lines of communication. Division of Entomology

Newsletter 34(4): 1-2.

_ . 1953c. Chaidiodes disjunctus Walker: a correction with de¬
scriptions of a new species and new genus (Megaloptera: Cory-
dalidae). Canadian Entomologist 85: 190-192.

_ . 1953d. Report on field work at Knob Lake, Quebec, 1948. In

Entomological Research in Northern Canada. Progress Report,
1948. Part 1. The Northern Insect Survey. Appendix D. Ottawa:
Canada Dept. Natl. Def.. Def. Res. Bd. (Rep. No. DR18): 1-34.
_ . 1953e. Some aspects of butterfly distribution in the West In¬
dies. (Abstract of invitation paper). Proceedings of the Entomo¬
logical Society of Washington 55: 112.

_. 1953f. The size of island faunas. (Abstract). Proceedings of the
7th Pacific Science Congress, Zoology 4: 52-53.

_ . 1953g. The phylogeny of the Papilionidae. Proceedings of the

7th Pacific Science Congress, Zoology 4: 83-87.

_ . 1954a. Visit to Europe, 1953. Entomology Division Newsletter

32(3): 4.

_ . 1954b. The eastern North American subspecies of the bar¬
berry geometrid (Lepidoptera). Canadian Entomologist 86:
282-283.

_. 1954c. A new Phlyctaenia, with notes on two related species
(Lepidoptera: Pvralidae). Canadian Entomologist 86: 428-431.

_ . 1954d. A new pyralid of Palaearc-tic affinities from Colorado

(Lepidoptera). Canadian Entomologist 86: 499-500.

_ . 1955a. The genus Epipagis Hiibner, nec Hampson, in North

America. (Lepidoptera: Pvralidae). Canadian Entomologist 87:
249-252.

_. 1955b. Figures of type specimens. Lepidopterists’ News 9:
140-141.

_ . 1955c. Karl Jordan, the lepidopterist. Transactions Royal En¬
tomological Society London 107 (Jordan Jubilee volume): 69-75.
_. 1956a. The genus Orenaia in North America, with the descrip¬
tion of a new species (Lepidoptera: Pvralidae). Canadian Ento¬
mologist 88: 74-78.

_ . 1956b. Geshna primordialis Dyar. with descriptions of two

new genera and two new subspecies (Lepidoptera: Pvralidae).
Canadian Entomologist 88: 126-128.

_ . 1956c. Restriction and revision of the genus Diastictis Hiibner

(Lepidoptera: Pvralidae). Canadian Entomologist 88: 208-228.

_ . 1956d. Canada as an environment for insect life. Canadian En¬
tomologist 88: 372-476.

_ . 1956e. The North American species of Diatlirausta Lederer

(Lepidoptera: Pvralidae). Canadian Entomologist 88: 579-583.
_. 1956f. Pvralides recoltes a l'Tle de la Guadeloupe par M. L.
Berland. Revue Franyaise d'Entomologie 23: 121-127.

_ . 1957a. North American subspecies of Pyrausta unifascialis

(Packard) (Lepidoptera: Pyralidae). Canadian Entomologist 89:
91-94.

_ . 1957b. North American components of the genus Diasemia of

authors (Lepidoptera: Pvralidae). Canadian Entomologist 89:
164-170.

_ . 1957c. Comparison of closely related faunas. Science 126:

437-439.

_ . 1957d. Comment on Dr. Elwood C. Zimmerman's proposal to

add to the "Official list of generic names in zoology" the names
“ Oeobia ” Hiibner, [1825], and “Hellula” Guenee, 1845 (Class In¬
secta, Order Lepidoptera). Bulletin of Zoological Nomenclature
13: 183-185.

_ . 1957e. Proposed suppression under the plenary powers of the

generic name Oeobia Hiibner, [1825] (Class Insecta, Order Lep¬
idoptera) for the purposes of the law of priority but not for those
of the law of homonymy. Bulletin of Zoological Nomenclature 13:
186-189.

_ . 19571. Northward extension in recent years of the range of the

lettered sphinx moth. Canadian Field Naturalist 71: 37.

_ . 1958a. Collecting insects in Pacific islands and Southeast Asia.

Division of Entomology Newsletter 36(5): 1-2.

_ . 1958b. A new subspecies of Sparagmia gigantalis Guenee

(Lepidoptera: Pvralidae). Canadian Entomologist 90: 127-128.

_ . 1958c. Pycnarmon jaguaralis (Guenee) and an allied species

(Lepidoptera: Pvralidae). Canadian Entomologist 90: 234—235.

_ . 1958d. Far-eastern Pvralidae (Lepidoptera). Canadian Ento¬
mologist 90: 249-254.

_ . 195Se. Some species of Polygrammodes Guenee (Lepidoptera:

Pyralidae). Canadian Entomologist 90: 257-274.

_ . 1958f. A revision of the genus Epicorsia Hiibner (Lepidoptera:

Pyralidae). Canadian Entomologist 90: 293-301.

_. 1958g. Catalogue of Lederer types of Pvralidae in the British
Museum (Natural History) wfith selections of Iectotvpes (Lepi¬
doptera). Canadian Entomologist 90: 510-517.

_. 1958h. Hampson's Schoenobiinae (Lepidoptera: Pyralidae).
Proceedings of the 10th International Congress of Entomology' 1:
301-302.

_ . 1958i. Systematics as a tool in ecology. Proceedings of the 10th

International Congress of Entomology, Ecology 2: 663-667.

_ . 1958j. The geographic distribution of the Scopariinae (Lepi¬
doptera: Pyralidae). Proceedings of the 10th International Con¬
gress of Entomology, Geographical Distribution 1: 831-837.

_ . 1958k. Ranges of representative insects, ticks and spiders. At¬
las of Canada, Plate 37, 24 maps.

_, A. Diakonoff, & E. L. Martin. 1958. Catalogue of Snellen's
types of Pyralidae, with selections of Iectotvpes. Tijdschrift voor
Entomologie 101: 65-88.

_ . 1959a. Canadian species of Dioryctria Zeller (Lepidoptera:

Pyralidae). Canadian Entomologist 91: 65-72.

_ . 1959b. New Pyralidae from the Papuan Region (Lepidoptera).

Canadian Entomologist 91: 102-112.

_ . 1959c. Pvralidae from the collection of the California Academy

of Sciences (Lepidoptera). Canadian Entomologist 91: 161-167.
_ . 1959d. The phlogosaria complex of the genus Plagodis (Lepi¬
doptera: Geometridae). Canadian Entomologist 91: 193-208.

_ . 1959e. New genera and species of Pyralidae (Lepidoptera).

Canadian Entomologist 91: 359-371.

_ . 1959f. Four new species of Evergestis (Lepidoptera: Pyrali¬
dae). Canadian Entomologist 91: 406-411.

_. 1959g. Revision of the genus Linosta Moschler (Lepidoptera:
Pyralidae) with characterization of the subfamily Linostinae and a
new subfamily. Canadian Entomologist 91: 485—488.

_ . 1959h. New species and new subspecies of Palpita (Lepi¬
doptera: Pvralidae). Canadian Entomologist 91: 641-650.

_ . 1959i. Pyralidae collected in lower California by Mr. J. Powell

(Lepidoptera). Canadian Entomologist 91: 725-727.

_. 1959j. The pyralid fauna of the Western United States. Presi¬
dential address to the Sixth Pacific Slope Meetings of the Lepi¬
dopterists’ Society, August 1959. Journal of the Lepidopterists’
Society 13: 241-244.

_ . 1960a. New tropical Pyraustinae (Lepidoptera: Pyralidae).

Canadian Entomologist 92: 164-173.

_ . 1960b. A new genus of Pyralidae and its species (Lepidoptera).

Canadian Entomologist 92: 188-192.

_ . 1960c. New species of Polygrammodes and a related new

54

Journal of the Lepidopterists’ Society

genus (Lepidoptera: Pvralidae). Canadian Entomologist 92:
279-284.

_ . 1960d. The Meyrick types of Scopariinae (Lepidoptera: Pyral-

idae) in the British Museum (Natural History), exclusive of
Hawaiian species. Canadian Entomologist 92: 891-897.

_ . 1960e. Liocoris , Lygus and ethics. Bulletin of the Brooklyn

Entomological Society 55: 104-108.

_ . 1960f. An assessment of the contribution of experimental tax¬
onomy to the classification of insects. Revue Canadienne de Bi-
ologie 19: 293-319.

_ . 1960g. Professional and amateur research in Lepidoptera.

(President's Annual Address, 1959). Journal of the Lepidopterists’
Society 14: 1-4.

_ & P. R. Ehrlich. 1960. Harmonization of concepts of higher

classification of the Papilionidae. Journal of the Lepidopterists’
Society 14: 169-175.

_ . 1961a. Geography and systematic research. Proceedings of the

9th Pacific Scientific Congress 19: 72-74.

_ . 1961b. The classification of the Papilionidae (Lepidoptera).

Memoirs of the Entomological Society of Canada 17: 1—51 .

_ . 1961c. Synopsis of the North American Odontiinae, with de¬
scriptions of new genera and species (Lepidoptera: Pyralidae).
Memoirs of the Entomological Society Canada 24: 1—93.

_ . 1962. A new species of Linosta (Lepidoptera: Pyralidae). Cana¬
dian Entomologist 94: 922-923.

_ . 1963a. The g ilvarius group of Aspilates Treitschke (Lepi¬
doptera: Geometridae). Canadian Entomologist 95: 259-287.

_ . 1963b. Perspectives in biogeography. Canadian Entomologist

95: 299-308.

_ . 1963c. A new species of Nepytia (Lepidoptera: Geometridae),

of economic importance to Douglas fir in British Colombia.
Canadian Entomologist 95: 407— 113.

_ . 1963d. A new species of Tetralopha (Lepidoptera: Pyralidae)

from Vaccinium. Canadian Entomologist 95: 677-679.

_ . 1963e. New Neotropical Pyraustinae (Lepidoptera: Pyralidae).

Canadian Entomologist 95: 704-712.

_ . 1963f. Collecting insects in the Territory of Papua and New

Guinea, 1957. Proceedings of the 9th Pacific Science Congress,
Entomology 4: 32 — 41 .

_ . 1963g. Systematic Entomology Survey Program of the Canada

Dept, of Agriculture. Proceedings of the 9th Pacific Science Con¬
gress, Entomology 4: 42-43.

_ . 1963h. Entomological research in Canada. Proceedings of the

9th Pacific Science Congress, Entomology 4: 132-135.

_ . 1963i. Characteristics and history of the North American

fauna: Lepidoptera. Proceedings of the 16th International Con¬
gress of Zoology 4: 21-27.

_ . 1964a. Problems and trends in systematics. Canadian Ento¬
mologist 96: 367-377.

_ . 1964b. Four new species of Cliniodes Guenee, with notes on

the genus and some relatives and segregates (Lepidoptera: Pyral¬
idae). Canadian Entomologist 96: 529-538.

_ . 1964c. New genera and species of Glaphyriinae (Lepidoptera:

Pyralidae). Canadian Entomologist 96: 1261-1307.

_ . 1964d. Some Neotropical genera resembling Epicorsia Hiib-

ner (Lepidoptera: Pyralidae). Memoirs of the Entomological So¬
ciety of Canada 33: 3-76.

_ . 1964e. Insects of Campbell Island. Lepidoptera: Pyralidae. Pa¬
cific Insects Monographs 7: 260-271.

_ . 1964f. Biosystematics and dynamic ecology. Royal Ontario Mu¬
seum, University of Toronto Life Sciences Contributions 59: 1-17.

_ . 1965. Zoogeography of insects and allied groups. Annual Re¬
view of Entomology 10: 325-344.

_ . 1966. Revision of North American species of Udea Guenee

(Lepidoptera: Pyralidae). Memoirs of the Entomological Society
of Canada 49: 1-57.

Wier, J. R. & E. G. Munroe. 1966. The interdisciplinary research.
Agricultural Institute Revue. November-December: 9-13.

Munroe, E. G. 1967a. Four new species of Udea from South America
(Lepidoptera: Pyralidae). Canadian Entomologist 99: 392—397.

_ . 1967b. A new species of Thliptoceras from Thailand, with

notes on generic and specific synonymy and placement and with
designations of lectotypes (Lepidoptera: Pyralidae). Canadian
Entomologist 99: 721-727.

_ . 1967c. A new genus and a new species of Pyraustinae (Lepi¬
doptera: Pyralidae). Canadian Entomologist 99: 1104-1110.

_ . 1967d. Parapilocrocis citribasalis, a new genus and species of

Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist 99:
1156-1159.

_ . 1967e. Titles. Nature 214: 1064.

_ . 1968a. New Pvralidae (Lepidoptera) from the Papuan Region.

II. Canadian Entomologist 100: 394—401.

_ . 1968b. New species of Apilocrocis [Amsel], with a key to

known species (Lepidoptera: Pyralidae). Canadian Entomologist
100: 441^448.

_ . 1968c. Insects of Ontario: geographical distribution and post¬
glacial origin. Proceedings of the Entomological Society of On¬
tario 99: 43-50.

_ & A. MUTUURA. 1968a. Contributions to a study of the Pyrausti¬
nae (Lepidoptera: Pyralidae) of temperate East Asia. I. Canadian
Entomologist 100: 847-861.

_ & _ . 1968b. Contributions to a study of the Pyraustinae

(Lepidoptera: Pyralidae) of temperate East Asia. II. Canadian
Entomologist 100: 861-868.

_ & _ . 1968c. Contributions to a study of the Pyraustinae

(Lepidoptera: Pyralidae) of temperate East Asia. III. Canadian
Entomologist 100: 974-985.

_ & _ . 1968d. Contributions to a study of the Pvraustinae

(Lepidoptera: Pyralidae) of temperate East Asia. IV. Canadian
Entomologist 100: 986-1001.

_ . 1969a. Transfer of Cacographis colombiana [Schaus] from

Midilinae to Nymphulinae, with description of Midilambia n.
gen. (Lepidoptera: Pyralidae). Canadian Entomologist 101:
994-996.

_ . 1969b. Scoparia macroplianes Meyrick, 1888 (Insects, Lepi¬
doptera, Pyralidae): request for suppression of neotype designa¬
tion following rediscovery of holotvpe. Bulletin of Zoological
Nomenclature 26: 167-170.

_ . 1969c. Insects of Ontario: geographical distribution and post¬
glacial origin. Proceedings of the Entomological Society of On¬
tario 99: 43-50.

_ . 1969c!. Book review - Ecology and biogeography of high alti¬
tude insects, by M. S. Mani. Canadian Entomologist 101:
1007-1008.

_ & A. Mutuura. 1969a. Contributions to a study of the Pyrausti¬
nae (Lepidoptera: Pyralidae) of temperate East Asia. V. Canadian
Entomologist 101: 299-305.

_ & _ . 1969b. Contributions to a study of the Pyraustinae

(Lepidoptera: Pyralidae) of temperate East Asia. VI. Canadian
Entomologist 101: 897-906.

_ & _ . 1969c. Contributions to a study of the Pyraustinae

(Lepidoptera: Pyralidae) of temperate East Asia. VII. Canadian
Entomologist 101:1069-1077.

_ & _ . 1969d. Contributions to a study of the Pyraustinae

(Lepidoptera: Pvralidae) of temperate East Asia. VIII. Canadian
Entomologist 101: 1239-1248.

_ & P. D. McTaggart-Cowan. 1969. The science council of

Canada. Agricultural Institute Review March-April 1969: 1820.

Mutuura, A., E. G. Munroe, & D. A. Ross. 1969a. American species
of Dioryctria Zeller (Lepidoptera: Pyralidae). I. Western Cana¬
dian species of the zimmermani group. Canadian Entomologist
101: 1009-1023.

_ , _ , & _ . 1969b. American species of Dioryctria Zeller

(Lepidoptera: Pvralidae). II. Western Canadian species of the
baumhoferi and ponderosae groups. Canadian Entomologist 101:
1042-1047.

Munroe, E. G. 1970a. Book review- A short history of the browntail
moth, by William Curtis. Canadian Entomologist 102: 170.

_ . 1970b. A new genus and three new species of Chrysauginae

(Lepidoptera: Pvralidae). Canadian Entomologist 102: 414^120.

Volume 63, Number 1

55

_ . 1970c. A short history of the brown-tail moth. Canadian Ento¬
mologist 102: 640.

_ . 1970d. Types of Nymplnilinae, Scopariinae, Cybalomiinae,

Odontiinae, Evergestinae, and Pyraustinae (Lepidoptera: Pyrali-
dae) in the Zoological Institute, Academy of Sciences of the
U.S.S.R., Leningrad, with selections of lectotvpes. Canadian En¬
tomologist 102: 1025-1035.

_ . 1970e. Revision of the subfamily Midilinae (Lepidoptera:

Pvralidae). Memoirs of the Entomological Society of Canada 74:
1-94.

_ . 1970f. New species of Syllepis Poey ([Lepidoptera]: Pvralidae:

Pyraustinae), with a key to known species. Journal of the Lepi-
dopterists’ Society 24: 287-291.

_ & A. MUTUURA. 1970a. Contributions to a study of the Pyrausti¬
nae (Lepidoptera: Pvralidae) of Temperate East Asia. IX. Cana¬
dian Entomologist 102: 294-304.

_ & _ . 1970b. Contributions to a study of the Pyraustinae

(Lepidoptera: Pvralidae) of Temperate East Asia. X. Canadian
Entomologist 102: 1489-1507.

Mutuura, A. & E. G. Munroe. 1970. Taxonomy and distribution of
the European coni borer and allied species: genus Ostrinia
[Hiibner] (Lepidoptera: Pvralidae). Memoirs of the Entomologi¬
cal Society of Canada 71: 1-112.

Munroe, E. G. 1971a. Eudonia borealis (Tengstrom) and E. alpina
(Curtis) (Lep., Pvralidae). Entomologist's Gazette 22: 96.

_ . 1971b. A new name for Papilio zetes (Westwood, 1847), nec.

Linnaeus, 1758 (Papilionidae). Journal of the Lepidopterists’ So¬
ciety 25: 142.

_ . 1971c. Distribution and geographical differentiation of Marpe-

sia eleuehea Hiibner (Nvmphalidae), with descriptions of two
new subspecies. Journal of the Lepidopterists’ Society 25:
185-193.

_ . 1971d. Status and potential of biological control in Canada.

Part IV, Review of biological control in Canada, 1959-1968.
C.A.B., London 4: 213-255.

_ & A. Mutuura. 1971a. Contributions to a study of the Pyrausti¬
nae (Lepidoptera: Pyralidae) of Temperate East Asia. XI. Cana¬
dian Entomologist 103: 173-181.

_ & _ . 1971b. Contributions to a study of the Pyraustinae

(Lepidoptera: Pvralidae) of Temperate East Asia. XII. Canadian
Entomologist 103: 503-506.

_ & _ . 1971c. Geographical distribution of Pyraustinae (Lep¬
idoptera: Pyralidae) of temperate East Asia. Transactions of the
Lepidopterists’ Society of Japan 22: 1-6.

Coulson, R. N., A. Mutuura, & E. G. Munroe. 1972. The Dioryc-
tria species of loblolly pine in East Texas, with comments on the
occurrence of two new species. Journal of Economic Entomology
65: 868-870.

Munroe, E .G. 1972a. A new species of Midila Walker (Lepidoptera:
Pyralidae: Midilinae) from Venezuela. Canadian Entomologist
104: 711-713.

_ . 1972b. An entomologist in Brazil. Tableau: 2.

_ . 1972c. Centime de lepidopteres de file de Cuba, by Ph. Poey.

Reprinted by E. W. Classey Ltd. Journal of the Lepidopterists’
Society 26: 123.

_ . 1972d. Book review — Jamaica and its butterflies, by F.M.

Brown and B. Heineman. Journal of the Lepidopterists’ Society
26: 264-267.

_ . 1972e. Pvraloidea. In The moths of America north of Mexico.

Fasc. 13. 1A. Subfamilies Scopariinae, Nymphulinae. R. B. Do¬
minick, C. R. Edwards, D. C. Ferguson, J. G. Franclemont, R. W.
Hodges, and E. G. Munroe. E. W. Classey, (eds.), London:
1-134.

_ . 1972f. Pyraloidea. In The moths of America north of Mexico.

Fasc. 13. IB. Subfamilies Odontiinae, Glaphyriinae. R. B. Do¬
minick, C. R. Edwards, D. C. Ferguson, J. G. Franclemont, R. W.
Hodges, and E. G. Munroe. E. W. Classey, (eds.), London:
137-250.

Mutuura, A. & E. G. Munroe. 1972. American species of Dionjctria
Zeller (Lepidoptera: Pvralidae). III. Grouping of species: species

of the auranticella group, including the Asian species, with the
description of a new species. Canadian Entomologist 104:
609-625.

Munroe, E. G. 1973a. A supposedly cosmopolitan insect: the celery
webworm and allies, genus Nomopliila Hiibner (Lepidoptera:
Pyralidae: Pyraustinae). Canadian Entomologist 105: 177-216.

_ . 1973b. A new genus and species of Odontiinae from Arkansas

(Lepidoptera: Pyralidae). Canadian Entomologist 105: 669-671.

_ . 1973c. A new species of Trischistognatha from Venezuela

(Lepidoptera: Pyralidae: Evergestinae). Canadian Entomologist
105: 1081-1083.'

_ . 1973d. A new genus for Noctueliopsis radialis (Lepidoptera:

Pvralidae: Odontiinae). Canadian Entomologist 105: 1361-1362.

_ . 1973e. Transfer of Cymoriza abrotalis Walker, 1859, from

Nytnphula Schrank to Disimidila Dvar (Pyralidae: Nymphulinae,
Midilinae). Journal of the Lepidopterists’ Society 27: 290-201.

_ . 1973f. Book review — Die tagfalter Europas und Nordwest

Afrikas, ein Taschenbuch fur Biologen, by L. G. Higgins and N.
D. Riley, illustrated by B. Hargreaves, and translated by W.
Forster. Bulletin of the Entomological Society of Canada 5(3): 30.

_ . 1973g. Book review — An index to the described life histories,

early stages and hosts of the macrolepidoptera of the continental
United States and Canada, by H. M. Tietz. Bulletin of the Ento¬
mological Society of Canada 5(3): 94-95.

_ . 1973h. Book review — Emperor moths of south and central

Africa, by E. Pinhey. Bulletin of the Entomological Society of
Canada 5(3): 96-97.'

Mutuura, A. & E. G. Munroe. 1973. American species of Dionjctria
[Zeller] (Lepidoptera: Pvralidae). IV. The schuetzeeUa group and
the taxonomic status of the spruce cone moth. Canadian Ento¬
mologist 105: 653-668.

Munroe, E. G. 1974a. Pyraloidea. In The moths of America north of
Mexico. Fasc. 13. 1C. Evergestinae. R. B. Dominick, D. C. Fer¬
guson, J. G. Franclemont, R. W. Hodges, and E. G. Munroe. E.
W. Classey, (eds.), London: 253-304.

_ . 1974b. Three new genera and four new species of Pyralidae

(Lepidoptera) from South America. Canadian Entomologist 106:
11-20.

_ . 1974c. New Pvralidae (Lepidoptera) from the Papuan Region.

III. Canadian Entomologist 106: 21-30.

_ . 1974d. Pyralidae of the Third Archbold Expedition. I. Sub¬
family Odontiinae (Lepidoptera). Tijdschrift voor Entomologie
117: 27-37.

_ . 1974e. New genera, species, and generic transfers of Neotrop¬
ical Odontiinae (Lepidoptera: Pvralidae). Canadian Entomologist
106: 127-137.

_ . 1974f. Transfer of Cybalomia angustalis Dognin to Udea

Guenee, with description of a related new species (Lepidoptera:
Pyralidae). Canadian Entomologist 106: 139-142.

_ . 1974g. A new Noctueliopsis from Texas (Lepidoptera: Pyrali¬
dae: Odontiinae). Canadian Entomologist 106: 655-658.

_ . 1974h. Hard decisions ahead for Canada’s biological scientists.

Science Forum 37 7(1).

Mutuura, A. & E. G. Munroe. 1974. A new genus related to Dioryc-
tria Zeller (Lepidoptera: Pyralidae: Phycitinae), with definition of
an additional species group in Dioryctria. Canadian Entomologist
106: 937-940.

Munroe, E. G. 1975a. Biogeography of North American Odontiinae.
Canadian Entomologist 107: 129-154.

_ . 1975b. Butterflies and moths in winter. Nature Canada 4(1):

26-29.

_ . 1975c. Entomology in the N.C.C. Symposium on the ecology

of the Mer Bleue. 2pp.

_ . 1975d. Science policy committee. Bulletin of the Entomologi¬
cal Society of Canada 7: 77.

_ . 1975e. Moths of importance to crops, forests and environmen¬
tal quality in North America, with particular reference to un¬
solved taxonomic problems. The Wedge Entomological Research
Foundation, New York, N.Y. 31pp.

Dominick, R. B. & E. G. Munroe. 1975. Why should individuals, or-

56

Journal of the Lepidopterists’ Society

ganizations or groups support the Wedge Entomological Re¬
search Foundation? The Wedge Entomological Research Foun¬
dation, New York, N.Y. 13pp.

MUNROE, E. G. 1976a. Pvraloidea. In The moths of America north of
Mexico. Ease. 13. 2A , Pvralidae, Subfamily Pvraustinae, Tribe
Pyraustini. B. Pyralidae, (conclusion). R. B. Dominick, T. Do¬
minick, D. C. Ferguson, J. G. Franclemont, R. W. Hodges, and E.
G. Munroe. E. W. Classey, (eds.), London: 1-78.

_ . 1976b. Pvraloidea. In The moths of America north of Mexico,

Fasc. 13. 2B. Pyralidae, Subfamily Pvraustinae, Tribe Pyraustini
(conclusion). R. B. Dominick, D. C. Ferguson, J. G. Fran¬
clemont, R. W. Hodges, and E. G. Munroe. (eds.), E.W. Classey,
London: 81-150.

_ . 1976c. Hapalia nigristriatalis a synonym of Udea angustalis

(Pyalidae: Pvraustinae). Journal of the Lepidopterists' Societv30:
11.

_ . 1976d. New genera and species of Pyraustinae (Lepidoptera:

Pyralidae), mainly from the collection of the British Museum
(Natural History). Canadian Entomologist 108: 873-884.

_ . 1976e. Book Review — Macrolepidoptera of Fiji and Roturna: a

taxonomic and hiogeographic study, by Gaden S. Robinson. Bul¬
letin of the Entomological Society of Canada 8: 13-14.

_ . 1976f. Obituary — Richard Bayard Dominick, 1919-1976. Bul¬
letin of the Entomological Society of Canada 8(4): 19-20.

_ . 1977a. Lepidoptera. In Biota Acuatica de Sudamerica Austral.

S. II. Hurlbert (ed.), San Diego State University, San Diego, Cal¬
ifornia 14: 254-255.

_ . 1977b. Two species of Palpita Hiibner described by Janse, with

descriptions of new species (Lepidoptera: Pyralidae: Pvrausti¬
nae). Canadian Entomologist 109: 67-76.

_ . 1977c. Synopsis of Heortia Lederer, with descriptions of a new

species and a new subspecies (Lepidoptera: Pyralidae: Odonti-
inae). Canadian Entomologist 109: 429-441.

_ . 1977d. A new genus and two new species of Neotropical

Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist
109: 481^185.

_ . 1977e. A new genus and species of Odontiinae, with remarks

on Balaenifrons Hampson and Taurometopa Meyrick (Lepi¬
doptera: Pyralidae). Canadian Entomologist 109: 865-869.

_ . 1977f. New Indo-Malayan Odontiinae (Lepidoptera: Pyrali¬
dae), with taxonomic and synonymic notes. Canadian Entomolo¬
gist 109: 1341-1358.

_ . 1977g. Additional material of Scoparia huachucalis Munroe,

with description of male genitalia (Pyralidae, Scopariinae). Jour¬
nal of the Lepidopterists' Society 31: 63-66.

_ . 1977h. Canada’s endangered species and habitats: Insects.

Why protect insects? Canadian Nature Federation: 144-147.

_ . 1977i. Introduction for Monographic des Phycitides et Gal-

leriides par E. L. llagonot. Facsimile edition. E.W. Classey Ltd.

_ . 1977j. Book review — Insects that feed on trees and shrubs: an

illustrated practical guide, by W. T Johnson & H. II. Lyon. Cana¬
dian Field-Naturalist 91: 105-106. '

_ . 1977k. Book review — Biology of insects, by David J. Horn.

Canadian Field-Naturalist 91(3): 326.

Carde, R. T„ W. L. Roelofs, R. G. Harrison, A. T. Vawter, P. F.
Brussard, A. Mutuura, & E. G. Munroe. 1978. European Com
Borer: pheromone polymorphism or sibling species? Science 199:
555-556.

Munroe, E. G. 1978a. A new species related to the garden webworm.
with a key to American members of the genus Achyra Guenee
(Lepidoptera: Pyralidae: Pyraustinae). Canadian Entomologist
110: 499-505.

_ . 1978b. Epicorsia I Iiibner and allied Neotropical genera: sup¬
plementary notes and descriptions (Lepidoptera: Pyralidae:
Pyraustinae). Canadian Entomologist 110: 673-695.

_ & A. Mutuura. 1978. Meyrick's record of “Mecyna fumacalis,

GN” from Fiji, with a new generic assignment for Pyrausta
homaloxarvtha Meyrick (Pyralidae: Pyraustinae). Journal of the
Lepidopterists’ Society 32:130-134.

Mutuura, A. & E. G. Munroe. 1978. An Old World leaf roller po¬

tentially injurious to Rosaceae, newly discovered in British Co¬
lumbia, Croesia holmiana (Lepidoptera: Tortricidae). Canadian
Entomologist 110: 641-646.

Munroe, E. G. 1979. Lepidoptera. In Canada and its insect fauna.
H.V. Danks (ed). Memoirs of the Entomological Society of
Canada 108: 427-481.

Mutuura, A. & E. G. Munroe. 1979. American species of Dioryctria
(Lepidoptera: Pyralidae). 5. Three new cone-feeding species
from the southeastern U.S. Journal of the Georgia Entomological
Society 14: 290-304.

Bolte, K. B. & E. G. Munroe. 1979. Hemithea aestivaria in British
Columbia (Lepidoptera: Geometridae: Geometrinae). Canadian
Entomologist 111: 1121-1126.

Munroe, E. G. & M. Shaffer. 1980. A revision of Vitessidia Roth¬
schild & Jordan and Vitessa Moore (Lepidoptera: Pyralidae). Bul¬
letin of the British Museum of Natural History, Entomology 39:
241-360.

Inoue, H., E. G. Munroe, & A. Mutuura. 1981. A new species of
Glyphodes from Japan, with biological notes. Tinea 11: 91-97.

Munroe, E. G. 1982. Lepidoptera. In Synopsis and Classification of
Living Organisms. S. P. Parker, (ed.), McGraw-Hill, New York:
612-651. "

_ . 1983a. Pyralidae (except Crambinae). In Check List of the

Lepidoptera of America north of Mexico. R. W. Hodges, T. Do¬
minick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G.
Munroe, and J. A. Powell. E. W. Classey and The Wedge Ento¬
mological Research Foundation (eds.), London : 67-85.

_ . 1983b. Wing Patterns in Lepidoptera (Gold Medallists Ad¬
dress). Bulletin of the Entomological Society of Canada 14:
103-112.

_ . 1986. Obituaiy— George Pearson Holland, 1911-1985. Trans¬
actions of the Royal Society of Canada 1: 303-305.

_ . 1989. Changes in classification and names of Hawaiian

Pvraloidea since the publication of Insects of Hawaii, Volume 8,
by. E. C. Zimmerman (1958) (Lepidoptera). Bishop Museum Oc¬
casional Papers 29: 199-212.

Shaffer, J. C. & E.G. Munroe. 1989a. Type material of four African
species of Notarcha Meyrick, wfith designations of lectotypes and
changes in synonymy (Lepidoptera: Crambidae: Pyraustinae).
Proceedings of the Entomological Society of Washington 91:
248-256.

_ & _ . 1989b. Type material of two African species of Her-

petogramma and one of Pleuroptya (Lepidoptera: Crambidae:
Pyraustinae). Proceedings of the Entomological Society of Wash¬
ington 91: 414-420.

Smith, D. S., E. G. Munroe, & S. J. Ramos. 1989. Biogeographical
affinities of the butterflies of the soutlrwest Caribbean Islands,
San Andres and Providencia. Journal ol the Lepidopterists’ So¬
ciety 43(4): 274-288.

Clavijo, J. A. & E. G. Munroe. 1991. Misidentification of Diaphania
liyalinata (L.) (Lepidoptera: Crambidae: Pyraustinae) in the
British Lepidoptera fauna. British Journal of Entomology and
Natural History 4: 9-10.

Munroe, E. G. 1991. Transfer of Autocodes eupselias Meyrick to
Pyraustinae, with notes on the genus Marasmia Lederer and on
cataclystiform wfing patterns in the family Crambidae (Lepi¬
doptera: Pyraloidea). Bishop Museum Occasional Papers 31:
122-130.

Mather, B. & E. G. Munroe. 1992. Heleithia decostalis (Crambidae:
Pyraustinae) new to the United States. Journal of the Lepi¬
dopterists' Society 46: 159-160.

Munroe, E. G. 1992. Three nominal genera of Crambidae from “The
Generic Names of Moths of the World, Vol. 5, Pvraloidea.” Jour¬
nal of the Lepidopterists’ Society 46: 75-77.

_ . 1995. Usingeriessa onyxalis (Dvar) (Lepidoptera: Crambidae:

Nymphulinae), a moth with presumably aquatic larvae, newly
recorded from I Iawaii, with a synopsis of Hawaiian Nymphulinae.
Bishop Museum Occasional Papers 42: 39—42.

_ , V. O. Becker, J. C. Shaffer, M. Shaffer, & M. A. Solis,

1995. Pyraloidea. In Atlas of Neotropical Lepidoptera. Checklist:

Volume 63, Number 1

57

Part 2. Hyblaeoiclea - Pyraloidea - Tortricoidea. |. B. Heppner
(ed.). Association of Tropical Lepidoptera, Gainesville: 34-105,

Clavijo, J. A., E. G. Munroe, & Q. Arias. 1995. The genus Diapha-
nia Hiibner (Lepidoptera: Crambidae): key to species of eco¬
nomic importance. Agronomia Tropical 45: 347-358.

_ & E. G. Munroe. 1996. A new species of Diaphania from

Tropical America (Lepidoptera: Pyraloidea: Crambidae). Tropical
Lepidoptera 7: 151-153.

Munroe, E. G. 1996. Distributional patterns of Lepidoptera in the
Pacific Islands. In The Origin and Evolution of Pacific Island Bio¬
tas, New Guinea to Eastern Polynesia: Patterns and Processes. A.
Least and S. E. Miller (eds.). Academic Publishers, Amsterdam:
275-295.

_ & M. A. Solis. 1999. Pyraloidea. In Lepidoptera, Moths and

Butterflies, Vol. 1. Arthropoda. Insect, Vol. 4, Part 35. Handbook
of Zoology'. N. Kristensen (ed.). Walter de Gruyter & Co., Berlin:
233-256.'

Peck, S. & E. G. Munroe. 1999. Biogeography and evolutionary his¬
tory: wide-scale and long-term patterns in insects. In Ecological
Entomology. C. B. Huffaker and A. P Gutierrez (eds.). John Wi¬
ley & Sons, New York: 231-261.

Shaffer, J. & E. G. Munroe. 2007. Crambidae of Aldabra Atoll
(Lepidoptera: Pyraloidea). Tropical Lepidoptera 14: 1-109.

M. A. Solis, SEL, USD A, Smithsonian Institution,
P.O. Box 37012, National Museum of Natural History,
E-517, MRC 168, Washington, D.C., 20013-7012; email-
alma. solis@ars.uscla.gov

Received for publication 30 September 2008; revised and accepted 17
January 2009..

58

Book Review

Journal of the Lepidopteuists’ Society

Journal of the Lepidopterists’ Society
63(1), 2009, 58

MOTH CATCHER: AN EVOLUTIONIST’S
JOURNEY THROUGH CANYON AND PASS. By
Michael M. Collins. 176 pages, 13 color plates; ISBN
978-0-87417-725-1 (hardcover), ISBN 978-0-87417-
735-0; US$49. 95 (hardcover), US$18.95 (paperback).
Univ. of Nevada Press. Publication date: 2007.

Two threads run through the stories of most
lepidopterists that contribute to their life-long
obsessive fascination with some aspect of entomology,
whether they are professionals or avid amateurs. The
first is some event of life-changing discovery to light the
fire; the second is the timely connection with one or
more knowledgeable and nurturing mentors to fuel the
fire so that it burns throughout a lifetime. Such is
indeed the case with Mike Collins, who shares with us
the beginnings and many subsequent years of his own
journey as a lepidopterist — in this case pursuing the life
history, behaviors and interspecific relationships of
western North American Saturniidae.

Migrating from his native Iowa to the western
mountains, Mike quickly passed beyond the usual goal
of building a large collection of adult specimens and
focused on questions of biology and evolution,
prompted first by the late Walter R. Sweadners
monograph on the evolution of the genus Hyalophora.
Seminal to his later researches, Sweadners treatise on
natural hybridization begged for substantiating
research. The evolutionary relationships among the 3
Hyalophora species — cecropia, glove ri and euryalus —
became the major focus of Mike’s research. Mike
provided important supporting evidence for Sweadners
ideas in collections and by rearing H. gloveri and H.
euryalus from areas of sympatry and allopatry. The
discovery and elucidation of the hybrid zone where
these two species come together in Monitor Pass in
eastern California proved to be a major discovery. This
stoiy is the centerpiece of the book.

The sibling nature of Hemileuca eglanterina, nuttalli
and hero and their isolating mechanisms was another
problem in Mike’s voyage of research and discoveiy.
His research on these and other saturniids was woven
into his undergraduate years at UC Berkeley, and PhD
work under Art Shapiro at UC Davis.

Friendships with several of our colleagues and
stories of field adventures together spice the narrative.
The account of his first meeting with Art Shapiro gave

me a knowing chuckle. A short biography of Sweadner,
and Mike’s meeting with his widow and daughter, was
of special interest to me, as was a photo of this man
who inherited W. J. Holland’s curatorial mantle at the
Carnegie Museum, but for a tragically short period.

Collins’ style is anecdotal, written in the first person,
and easily readable. He uses it effectively when
describing people and research projects without the
cumbersome data found in his scientific publications.
There are numerous short treatises on topics pertinent
to systematic^ and ecology: western montane ecology
(plant and animal), Neo-Darwinism, industrial
melanism, DNA sequencing, speciation (biological
species concept versus phvlogenetie species concepts),
and pre-mating and post-mating barriers, to name a
few. The book thus serves as a good primer for bright
young people interested in entering the field of
population ecology.

Mike’s lifelong passion for his chosen specialty in
entomology — isolating mechanisms among closely
related saturniid moths in the majestic West — shines
through from beginning to end. Mv minor comments
concern uneven referencing: some of the major
literature cited (i.e. Sweadners monograph, p.24), are
mentioned generally (no complete title and other
citation details), but not included in the Bibliography.
The reader must scout these out from other sources.
Also, some less scientifically trained readers might
benefit from definitions of more of the scientific terms
employed ( genome , p. 44; “relief’, p. 56). Other terms
are well defined or explained.

It is always fascinating to see how our colleagues
began their journeys with Lepidoptera — how and from
whom they drew inspiration, nurturing and training—
and then see a summary of their career-long
contributions to the science. Mike’s story is an
enjoyable and informative addition to this lore.

Charles V. Covell Jr., McGuire Center for
Lepidoptera and Biodiversity, Florida Museum of
Natural History, Hull Road at SW 34th St., Gainesville,
FL 32611. Email: ccovell@flmnh.ufl.edu.

Received for publication 10 Dec 2008; revised and accepted
17 Jan 2009

Volume 61, Number 1

59

Journal of the Lepidopterists' Society
63(1), 2009, 59

MANUSCRIPT REVIEWERS FOR 2008 (VOLUME 62)

Manuscript reviewers are anonymous contributors to the scientific rigor, clarity, and quality of text and
illustrations in the papers published by the Journal of the Lepidopterists’ Society. The reviewers’ input is invaluable
and always welcomed by authors, editors and readers. We hope their careful work continues to allow the Journal
to increase quality and readership. On behalf of all the authors and the editorial staff of the Journal, respectful
acknowledgement is given to the reviewers for contributions published in Volume 62.

John Acorn, Edmonton, Alberta, Canada

Matti Ahola, Koski, Finland

Annette Aiello, Washington, DC

Richard Arnold, Pleasant Hill, California

George Balogh, Portage, Michigan

Susan Rorkin, Milwaukee, Wisconsin

M. Deane Bowers, Boulder, Colorado

Lincoln Brower, Roseland, Virginia

John Brown, Washington, DC

John Bums, Washington, DC

John Calhoun, Palm Harbor, Florida

Steven Cary, Santa Fe, New Mexico

Jeff Crolla, Toronto, Ontario, Canada

Michelle DaCosta, Winston-Salem, North Carolina

Robert Dana, Minneapolis, Minnesota

Phillip DeVries, Metairie, Louisiana

Robert Dirig, Ithaca, New York

John Emmel, Hemet, California

Tom Emmel, Gainesville, Florida

Matthew Forister, Reno, Nevada

Terry Harrison, Charleston, Illinois

David Iftner, Pittsfield, Illinois

Brian Inouye, Tallahassee, Florida

Akito Kawahara, College Park, Maryland

Ian Kitching, London, England

Norbert Kondla, McBride, British Columbia, Canada

David Lawrie, Edmonton, Alberta, Canada

David MacDougall, Albany, New York

Billy McCord, Charleston, South Carolina

David McCorkle, Monmouth, Oregon

William Miller, St. Paul, Minnesota

Mark Minno, Gainesville, Florida

Eugene Morton, Cambridge Springs, Pennsylvania

Karen Oberhauser, St. Paul, Minnesota

Paul Opler, Loveland, Colorado

Jerry Powell, Berkeley, California

Ron Priest, East Lansing, Michigan

Tommaso Raeheli, Rome, Italy

Robert Raguso, Ithaca, New York

Jadranka, Rota, Washington, DC

Ronald Royer, Minot, North Dakota

Daniel Rubinoff, Honolulu, Hawaii

Erik Runquist, Davis, California

Ronald Rutowski, Tempe, Arizona

Emily Saarinen, Gainesville, Florida

James Scott, Lakewood, Colorado

J. Mark Scriber, East Lansing, Michigan

Paul Severns, Philomath, Oregon

Jon Shepard, Nelson, British Columbia, Canada

John Shuey, Indianapolis, Indiana

Felix Sperling, Edmonton, Alberta, Canada

Stephen Spomer, Lincoln, Nebraska

Fred Stehr, East Lansing, Michigan

Kieth Summerville, Des Moines, Iowa

Michael Toliver, Eureka, Illinois

Tonya Van Hook, Sweet Briar, Virginia

Tom Vawter, Aurora, New York

Andrew Warren, Castle Rock, Colorado

Wayne Welding, Silver Spring, Maryland

Martha Weiss, Washington, DC

Donald Wright, Cincinnati, Ohio

Myron Zalucki, Brisbane, Australia

EDITORIAL STAFF OF THE JOURNAL
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Biology Department

College of Charleston, 58 Coining Street,
Charleston, South Carolina 29424-001 1 . USA

scholtensb@cofc.edu

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Associate Editors:

Gerardo Lamas (Peru), Kenelm W. Philip (USA), Robert K. Robbins (USA), Felix Sperling (Canada),
David L. Wagner (USA), Ciirister Wiklund (Sweden), Carla Penz (USA), Andrew Warren (USA),

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes,
Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the president
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Send Journal submissions electronically to: scholtensb@cofc.edu or to the editor at the above address. Contributors should feel free to
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Submit manuscripts in triplicate, typewritten, entirely double-spaced, with wide margins, on one side only of white letter-sized paper. Prepare
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Sheppard. P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp.

- 1961a. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10:165-216.

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CONTENTS

3 9088 01912 3868

A New Species of Drasteria Hubner (Noctuidae: Catocalinae: Melipotini) from Arizona, with

Comments on the Genus Jan Metlevski and Gregory Zolnerowich - 1

Biology and Distribution of a Recently Rediscovered Endemic Hawaiian Leafrolleii Moth,

Omiodes continuatalis (Crambidae) Cynthia King, Daniel Rubinojf and William Haines — 11

A New Species of Morrisonia (Noctuidae) from Southeastern North America

J. Bolling Sullivan and Janies K. Adams - 21

Tinacrucis noroesta , New Species, North America’s Largest Tortricine Moth (Tortricidae:

Atteriini) Jerry A. Powell - 27

Melitara Walker (Pyralidae) in Western Canada: The Documentation of M.

SUBUMBRELLA (DYAll) IN THE PRAIRIE PROVINCES DEMONSTRATES THE VALUE OF REGIONAL

Collections and Species Lists. Thomas J. Simonsen, Jason J. Dombroskie and
Gregory R. Pohl - 31

Not All Bilateral Gynandromoiiph Butterflies are Interspecific Hybrids: New Papilio
Specimens from Lield Populations J. Mark Scriber, Rodrigo Me reader, Howard
Romack and Mark Deering - 37

General Notes

Pithing and Mining by a Punkie: The Unusual Leeding Strategies of Nola cilicoides
(Grote, 1873) (Noctuidae: Nolinae) David L. Wagner and Bryan Connolly - 48

Obituary

Eugene G. Munroe (8 September 1919-31 May 2008) M. A. Solis - 52

Book Review

Moth Catcher: An Evolutionist’s Journey Through Canyon and Pass. By Michael M.
Collins. Charles V. Covell Jr. - 58

List of Reviewers for Volume 62-

59

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