• TABLE OF CONTENTS
HIDE
 Front Cover
 Front Matter
 Main






Group Title: Bulletin / Florida Museum of Natural History
Title: A review of Virbia (formerly Holomelina) of America north of Mexico (Arctiidae: Arctiinae: Arctiini) /
CITATION PDF VIEWER THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00101267/00001
 Material Information
Title: A review of Virbia (formerly Holomelina) of America north of Mexico (Arctiidae: Arctiinae: Arctiini) /
Alternate Title: Virbia of America north of Mexico
Physical Description: p. 59-118 : ill. (some col.), map ; 28 cm.
Language: English
Creator: Zaspel, Jennifer M
Weller, Susan J
Cardé, Ring T
Florida Museum of Natural History
Publisher: Florida Museum of Natural History, University of Florida
Place of Publication: Gainesville, FL
Gainesville, FL
Publication Date: 2008
Copyright Date: 2008
 Subjects
Subject: Tiger moths   ( lcsh )
Arctiidae   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Statement of Responsibility: J.M. Zaspel, S.J. Weller, and R.T. Cardé.
Bibliography: Includes bibliographical references (p. 107-110).
General Note: Bulletin - Florida Museum of Natural History ; vol. 48, no. 3, pp. 59-118
 Record Information
Bibliographic ID: UF00101267
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 354973193
issn - 0071-6154 ;

Downloads

This item has the following downloads:

PDF ( 4 MBs ) ( PDF )


Table of Contents
    Front Cover
        Front Cover
    Front Matter
        Front Matter
    Main
        Page 59
        Page 60
        Page 61
        Page 62
        Page 63
        Page 64
        Page 65
        Page 66
        Page 67
        Page 68
        Page 69
        Page 70
        Page 71
        Page 72
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
        Page 78
        Page 79
        Page 80
        Page 81
        Page 82
        Page 83
        Page 84
        Page 85
        Page 86
        Page 87
        Page 88
        Page 89
        Page 90
        Page 91
        Page 92
        Page 93
        Page 94
        Page 95
        Page 96
        Page 97
        Page 98
        Page 99
        Page 100
        Page 101
        Page 102
        Page 103
        Page 104
        Page 105
        Page 106
        Page 107
        Page 108
        Page 109
        Page 110
        Page 111
        Page 112
        Page 113
        Page 114
        Page 115
        Page 116
        Page 117
        Page 118
Full Text





FLORIDA
M MUSEUM.
OF NATURAL HISTORYTM


BULLETIN


A REVIEW OF VIRBIA (FORMERLY HOLOMELINA) OF AMERICA
NORTH OF MEXICO (ARCTIIDAE: ARCTIINAE: ARCTIINI)


J. M. Zaspel, S. J. Weller, and R.T. Carde


Vol. 48, No. 3, pp. 59-118


2008


UNIVERSITY OF FLORIDA


GAINESVILLE







The FLORIDA MUSEUM OF NATURAL HISTORY is Florida's state museum of natural history, dedicated to
understanding, preserving, and interpreting biological diversity and cultural heritage.

The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is a peer-reviewed publication
that publishes the results of original research in zoology, botany, paleontology, and archaeology. Address all inquiries
to the Managing Editor ofthe Bulletin. Numbers of the Bulletin are published at irregular intervals. Specific volumes
are not necessarily completed in any one year. The end of a volume will be noted at the foot of the first page of the
last issue in that volume.



Richard Franz, Managing Editor
Cathleen Bester, Production



Bulletin Committee
Richard Franz, C hijl i n;
Ann Cordell
Sarah Fazenbaker
Richard Hulbert
William Marquardt
Larry Page
Irvy R. Quitmyer
David W. Steadman, Ex officio Member




ISSN: 0071-6154

Publication Date: December 30, 2008







Send communications concerning purchase or exchange
of the publication and manuscript queries to:

Managing Editor of the BULLETIN
Florida Museum of Natural History
University of Florida
PO Box 117800
Gainesville, FL 32611-7800 U.S.A.
Phone: 352-392-1721
Fax: 352-846-0287
e-mail: Irfranz(@&flmnh.ufl.edu









A REVIEW OF VIRBIA (FORMERLY HOLOMELINA) OF AMERICA NORTH
OF MEXICO (ARCTIIDAE: ARCTIINAE: ARCTIINI)





J. M Zaspel', S. J. Weller2,3, and R.T. Card64







ABSTRACT

The tiger moth genus Virbia Walker (Arctiinae: Arctiini) is redescribed and its biology and taxonomic history are reviewed. The recent placement
of Holomelina Herrich-Schaffer as its junior synonym is examined. North of Mexico, 14 Virbia species are recognized, including two new
species, V.fergusoni Zaspel and V. rindgei Carde. Adult habitus photographs, a checklist, diagnoses, and complete descriptions are provided
for all species. Genitalia of the new species are illustrated, and diagnostic character illustrations are provided for separating all previously
described species. Locality information from over 12,000 specimens representing 16 museum collections [in the United States and Canada] is
compiled and available in an electronic database on the Diversity of Life Website (at www.diversitvoflife.org/virbia.html). As some species
appear to be habitat specialists (e.g., V lamae in bogs), photos of selected habitats are provided.



Key Words: Tiger moths, Virbia, Arctiidae, morphology, lepidopteran taxonomy.







TABLE OF CONTENTS

Introduction............................................................... ....... .. ................ 60
Biology of Virbia in Canada and the United States....................................... 62
Taxonomic History of Holomelina and Virbia.............................................. 68
Checklist of Virbia ........................................................................ .............. .. 72
M materials and M ethods .................................................................. ................ 73
Systematic Entomology of Genus Virbia................................... .................... 76
V a u ra n tia ca ............................................ ........................... ................ 7 7
V costa ta ........................................................................ . . ........... 8 1
V. fergusoni n. sp ................................................................... .............. .. 84
V .ferrug in osa ........................................... ........................... ................ 87
V frag ilis. ............................................................................ ................ 8 8
V im m a cula ta ........................................................ ................. . ............ 89
V la e ta ...................................................................................................... .. .. 9 1
V la m a e ................................................... ............................................... 94
V m a rg in a ta ............................................ ............................ ................ 97
V n ig rica n s............................................ ............................. .............. . 98
V op ella .................................................. ............................................... 10 1
V ostenta ............................................................................................ . 102
V rin dg ei n sp ...................................... ........................... . . ........... 104
V rubicundaria ....................................... .......................... . . ............ 106
Acknowledgem ents........................................................................ 107
Literature C ited .. ..................................................................... ............... 107

1 Dept. of Entomology and Nematology, P.O. BOX 110620 Natural Area Drive, University of Florida, Gainesville, FL 32611
2 Dept. of Entomology, 1980 Folwell Ave., 219 Hodson Hall, University of Minnesota, St. Paul, MN 55108
Bell Museum of Natural History, University of Minnesota, 1980 Upper Buford Circle, St. Paul, MN 55108
4 Dept. of Entomology, Univeristy of California-Riverside, Riverside CA 92521
Zaspel, J.M, S.J. Weller, and R.T. Carde. 2008. A Review of Virbia (Formerly Holomehna) of America North of Mexico (Arctiidae: Arctiinae: Arctinii)
Bull. Florida Museum Nat. Hist. 48(3): 59-118.








INTRODUCTION
Colorful and enigmatic, the species of North American
Virbia Walker, formerly Holomelina Herrich-Shaffer,
have challenged lepidopterists' species concepts and
occasionally their civility towards one another, since the
mid-1800s (e.g., scientific exchanges reviewed in Smith,
1889). The phenotypic diversity of adults of some spe-
cies is impressive. For example, within a single reared
brood of V aurantiaca (Huibner), several phenotypes
occur, and several of these were named as species (Fig.
1). Such phenotypic diversity when combined with in-
adequate illustrations and descriptions of many species
has resulted in complicated taxonomic history and con-
fusion. Further, only subtle differences in genitalic mor-
phology exist among the males of some species whose
phenotypes are easily confused (e.g., V aurantiaca and
V nigricans [Reakirt]) (Ferguson 1985; Forbes 1960;
Carde 1968, 1971).
Recently, a phylogenetic review of generic limits
of Virbia proposed that Virbia (Walker 1854) and
Holomelina (Herrich-Schaffer 1855) were congeneric,
and Holomelina was placed as the junior synonym
(Zaspel 2004; Zaspel &Weller 2006). Historically, these
two genera have been associated with each other (e.g.,
Barnes & McDunnough 1917; Forbes 1939, Forbes
1960); Ferguson (1985) suggested that the genera might
be congeneric when he described the Holomelina group.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

Rare in Arctiidae, males possess a true scaphium, and
many species also possess a subscaphium (Ferguson
1985; Zaspel 2004; Zaspel & Weller 2006).
As currently defined, Virbia is comprised of small
to medium sized, slender, brightly colored species (Fig.
2). The genus is distributed from south central Canada
(e.g., northern Manitoba) to the North Atlantic (e.g.,
Nova Scotia), south to central South America (Fig. 3)
(Handfield 1999; Zaspel &Weller 2006). North Ameri-
can species (formerly Holomelina) are slender moths
with red or pink adbasal regions in the hindwings (Figs.
2, 4). Neotropical species typically have the red pig-
ment replaced with yellow hindwing patches (Fig. 5).
Larvae possess homoideous crochets, a secondary loss
of the smaller lateral crochets found in other arctiines
(Kitching & Rawlins 1999; Jacobson & Weller 2002).
Larvae are otherwise typical ofArctiini with clusters of
secondary setae on raised verrucae (Kitching & Rawlins
1999; Jacobson & Weller 2002). Some North Ameri-
can species have been reared on dandelion and other
introduced Plantaginaceae (e.g., Forbes 1960). Typical
for Arctiini, larvae are reported as feeding on unidenti-
fied herbaceous plants; native hosts (pre-European settle-
ment) are unknown (Ferguson 1985; Tietz 1972).
Although little is known about their biology, spe-
cies in this genus have figured prominently in the litera-
ture on chemical ecology and pheromones (Roelofs &


Figure 1. Reared series, Virbia aurantiaca (Top Row: Female parent of brood, second and third rows: Males)






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


Figure 2. Virbia ostenta Edwards, Male.


Card6 1971; Card6 & Roelofs 1973; Card6 1974;
Charlton & Roelofs 1991; Schal & Card6 1985; Schal
& Card6 1986; Schal et al. 1987; Yin et al. 1991; Schal
et al. 1998). Virbia aurantiaca was among the first
non-pest lepidopterans to have its female pheromones
characterized (Roelofs & Card6 1971). Pheromone is
synthesized in the fat body (Schal et al. 1998), trans-
ported through the hemolymph to the pheromone glands,
and then released as a liquid aerosol (Meyer 1984; Schal
et al. 1987).
Here, we provide a faunal treatment of the Virbia
species occurring in North America north of Mexico,
formerly placed in Holomelina Herrich-Schaffer. A
generic diagnosis and description is provided for the ge-


Figure 3. General range distribution of the genus Virbia. Triangles = species originally placed in Virbia; stars
species formerly placed in Holomelina.








nus. We recognize 14 species. One of these, V.
marginata (Druce), is primarily distributed in Central
America with disjunct populations in Colorado, New
Mexico, and Arizona; it has been treated previously as a
synonym of V rubicundaria. Two new species, V.
fergusoni Zaspel and V rindgei Card6, are described.
For all species, we provide diagnoses, descriptions, and
illustrations of diagnostic male and female morphology
(Figs. 11-32). The adult habitus of both sexes are fig-
ured (Figs. 27, Pls. 1-3), and we summarize life history
information (e.g., larval hosts, voltinism, activity periods,
and habitat). A database of over 12,000 specimens is
available on-line through the Diversity of Life website
(www.diversityoflife.org/virbia.html).

BIOLOGY OF VIRBIA IN CANADA AND THE
UNITED STATES
Most of the information concerning the habitats and host
plant use is derived from three sources: field notes of R.
Card6, unpublished sections of theses (Card6 1971), and
the Zaspel database of compiled label data. These data
are supplemented with the few published rearing stud-
ies (Dyar 1897; Forbes 1910, 1960; Wyatt 1939; Ferguson
1953). Data are incomplete for many species, but we
note general trends when possible. Details of species
life histories are provided in the Biology sections for each
species.

HABITAT
Virbia exhibit a high degree of local habitat re-
striction (Figs. 6 and 7), although most species have wide-
spread distributions (e.g., V aurantiaca, V.
ferruginosa). Some phenotypically confusing species
can be sorted simply by either habitat or locality infor-
mation. For example, V lamae Freeman has been col-
lected only diurnally in bogs in the northeastern U.S.
and Canada (Fig. 6) to the western edge of its range,
Marathon County, Wisconsin (Ferguson 1954; Zaspel
database). Virbia nigricans (Reakirt) has been col-
lected only along a narrow corridor of secondary
sucessional habitat comprised of old fields, power line
cuts, and highway edges in western New Jersey
(Hunterdon County) and extreme southeastern
Pennsylavania (Philadelphia and Montgomery Counties;
Card6 field notes, Zaspel database). Virbia nigricans
has been confused historically with the dark form of V
opella, but these species are easily separated using adult
habitus and locality. These species mate diurnally (Card6
1971), and their genitalia are distinctive (see Diagnosis
of V nigricans). Another species with a highly special-
ized distribution is V rindgei Card6 (n. sp.), which is
found in open Ponderosa pine forests in the Black Hills


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

of South Dakota and Wyoming.
The remaining species appear to be more general-
ized in their habitat use, but their distributions are re-
stricted. Virbia rubicundaria (Htibner) has been col-
lected in tidal marshes on St. Catherine's island off the
coast of Georgia and also in moist, inland situations near
the southeastern U.S. ocean shores such as the Florida
scrub community in Highlands Co., Florida (Card6 field
notes; Zaspel database). Additional species with some-
what overlapping distributions can be separated based
on habitat, wing pattern, or voltinism. For example, V.
ferruginosa (Walker) is found in more open situations
such as pitch pine scrub oak barrens (Fig. 7), bogs,
aspen parklands, and montane woods (Schmidt, personal
communication), whereas V opella appears to favor
forested areas comprised of either deciduous woods or
mixed hemlock and deciduous forest with a developed
understory (Card6 1965; Zaspel database). Virbia
aurantiaca ranges from sand dunes and secondary suc-
cession old fields (Massachusetts) to the New Jersey
Pine Barrens, and from the Badlands (North Dakota) to
alfalfa fields (Indiana) (Zaspel database).

IMMATURE MORPHOLOGY AND BIOLOGY
Most larval descriptions provide a brief account of
the number of instars and general appearance. The first
descriptions of immature stages were of V aurantiaca
(Saunders 1869) and V opella (Dyar 1897). In 1960,
Forbes described mature larvae for three species, V
laeta, V aurantiaca, V opella, and the dark form of V
opella that he referred to as nigricans. He noted that
larvae of all species he examined possessed homoideous
crochets, two pairs of transverse verrucae on the pro-
thoracic shield, and abdominal L 1 located below the spi-
racle (Forbes 1960). Forbes (1960) described the larva
of V. aurantiaca as dark, with black setae and verru-
cae, and a contrasting conspicuous broad pale dorsal
band filling the space between verrucae. All the setae
were barbed and increased in length caudally. Larvae
of V laeta are lighter than V aurantiaca, pale gray
with black verrucae and a cream-colored stripe on the
frons; two types of setae were present, dentate, and
feathery (Forbes 1960). The third species, V opella,
was similar in body coloration to V laeta (dull gray) but
had a black head and brown frons; its black setae were
described as short and stiff. Forbes (1960) described a
larva that he attributed to form nigricans (1960: 23)
whose body color was a light orange base heavily dusted
with gray; the setae were gray, not black. This larva
was likely the dark form of V opella and not V. nigricans
based on the locality information provided and the rarity
of V nigricans. Thus, two morphs of V opella larvae






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


Figure 4. Virbia aurantiaca Huibner, Male.


Figure 5. Virbia hypophaea Hampson, Male.

exist: a dull gray morph with black setae (typical V.
opella) and a gray-orange morph with gray setae (dark
form V. opella). Unlike the two previous species, V.
ferruginosa larvae typically possess heteroideous
crochers but have reduced lateral crochets (Schmidt,
personal communication). Several species of Virbia have
been reared on two introduced European plants, dande-
lion (Asteraceae) or plantain (Plantaginaceae), but na-
tive hosts are unknown (Forbes 1960; Tietz 1972).

ADULT BIOLOGY
Most of our detailed information on adult biology is
limited to seven species representing the V aurantiaca
complex. This information is derived from the thesis
work of Card6 (1971), supplemented by continuing field
observations by Card6. Roelofs & Card6 (1971) iso-
lated and characterized female pheromone as 2-
methylheptadecane (2Me- 17c), the first saturated phero-
mone from the Lepidoptera, for seven species in the V.
aurantiaca complex: V aurantiaca, V ferruginosa,
V. fragilis, V immaculate, V lamae, V nigricans, and
V rindgei (as an undescribed species) (Roelofs & Card6
1971). 2Me-17c was used in pheromone traps to study
diel periodicities of attraction flight times. In addition,
chromosome numbers were characterized for seven
species. Methods for those studies are described in


Card6 (1971). A summary of the data is published here
for the first time (Table 1).

COLLECTING ADULTS
Some comments on methods for collecting adults
in the V aurantiaca complex illustrate how adults and
particularly females (for initiation of laboratory colonies)
were obtained. Nearly all of the males and females
used in studies of the pheromone biology were collected
by netting. All species can be collected by daytime flush-
ing (e.g., walking through the habitat), even though spe-
cies differ in whether females call and mate during the
day or night. Afternoons are the most favorable time to
collect; sunny and hot conditions are most conducive for
flight. Flushed males fly in an erratic zigzag pattern,
often for several meters or more, before landing; many
males can be flushed a second or third time. Females
have a reduced propensity to initiate flight, and the pro-
portion of females flushed is low; this more sedentary
behavior of females largely explains the great prepon-
derance of males in collections. When females are
flushed, their flight is readily distinguished: essentially a
straight-line path with reduced velocity, usually lower
altitude, and a much shorter distance than males, occa-
sionally only decimeters. Upon landing in the herbage,
females nearly always drop to ground level-males usu-
ally land on the herbage-and "play dead" (thanotosis).
These females can be difficult to locate unless their pre-
cise trajectory has been tracked, and it is more efficient
to net them on the wing if possible. Rarely, females can
be flushed a second time. Daytime flushing of both sexes
is presumably a defensive, predator avoidance reaction.
Many species also are readily attracted to black light
(males more strongly than females), and this collection
method accounts for an abundance of museum speci-
mens of V aurantiaca, V ferruginosa, V immaculate,
and V rubicundaria. For day-mating species, V lamae
and V nigricans, when black lights are placed in the
precise locations where adults were collected diurnally,
none are attracted to the lights. Oviposition behavior
differs as well. Females of V aurantiaca and V
rubicundaria lay eggs individually, not attached to a
substrate; females of V. ferruginosa, V. fragilis, V
immaculate, V lamae, V nigricans, and V rindgei lay
attached, single-layer masses.

PREMATING REPRODUCTIVE ISOLATION
Among co-occurring species, isolation can be due
to: 1) differences in pheromone, 2) rhythms of female
release of pheromone, hereafter termed "calling," and
male response to pheromone, 3) exclusive seasonal flight
periods, and 4) non-overlapping habitats.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Figure 6. Virbia lamae, habitat, Seawall Bog, Southwest Harbor, Maine (Photograph by R.T. Carde).


Figure 7. Virbiaferruginosa, habitat, xeric outwash pitch pine scrub oak barren in the Montague Plains Wildlife
Management Area, Montague, MA. (Photograph by R.T. Carde).





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


PHEROMONES
Distinctive pheromone channels can be achieved
in moths by a variety of strategies, including varying ra-
tios of the same constituents, or altering chain length,
double bond position, number and/or configuration,
chirality, or functional group (Card6 & Haynes 2004).
Indeed, it is typical to have closely related and sibling
species differ in their pheromone blends, such that these
serve as a primary means of premating reproductive
isolation.
The common pheromone constituent of the V.
aurantiaca complex is 2-methylheptadecane (2Me-17c).
Its presence in the female pheromone gland was con-
firmed for V aurantiaca, V ferruginosa, V fragilis,
V immaculate, V lamae, V nigricans, and V rindgei
(listed as an undescribed species) (Roelofs & Card6
1971). The compound 2Me-17c evoked levels of at-
traction in field trapping trials for all of the above spe-
cies except V. fragilis, which was not tested. In addi-
tion, V rubicundaria and V laeta are attracted to 2Me-
17c, although females have not yet been examined for
the presence of 2Me-17c. In some species, such as V.
aurantiaca and V rubicundaria, the levels of male
attraction relative to the density of field populations indi-
cated that 2Me-17c serves as the primary component.
Conversely, a comparatively poor level of attraction with
2Me-17c for V lamae (Roelofs & Card6 1971) sug-
gests that additional pheromone components could be
involved in communication. Indeed, in V lamae, in addi-
tion to 2Me-17c, several of its analogues are present in
the gland and emitted, including n-heptadecane, n-
nonadecane, 2-methyloctadane, and 2-methylnonadecane
(Schal et al. 1987). Field tests of V lamae with mix-
tures of these constituents in naturally occurring ratios,
however, do not have markedly improved attraction over
2Me-17c alone, leaving open the possibility that some
species specificity awaits documentation of additional
pheromone components. Differences in courtship be-
haviors among the species, another possible factor in
reproductive isolation, have not been examined.

RHYTHMS OF FEMALE CALLING
Pheromone synthesized in oenocytes is transported
via lipophorin to paired, tubular abdominal glands (Schal
et al. 1998), each with a spine-beset opening at the ab-
dominal tip (Yin et al. 1991). A calling female perches
vertically and rhythmically protrudes the tip of her abdo-
men at a rate of ca. once per 1.5 sec (Card6 1971;
Card6 & Roelofs 1973). Pheromone is atomized into
droplets, in pulses coincident with each gland extrusion
(Schal & Card6 1985). This release mechanism is evi-


dently common in arctiids but is not known among other
moth lineages (Meyer 1984; Schal & Card6 1986; Schal
et al. 1987, 1998; review Hallberg & Poppy 2003). The
amount of pheromone released is quite high among moths:
in V lamae up to 350 ng of 2Mel7c in 10 min and 835
ng in 60 min (Schal et al. 1987).
The calling rhythm of V immaculate has been dem-
onstrated to be circadian (Card6 & Roelofs 1973). Pre-
sumably the rhythms of the remaining species in this
complex also have a circadian basis; however, we refer
to them here as diel rhythms, as their endogenous na-
ture has not been confirmed formally. The calling
rhythms illustrated (Figs. 8, 9), all were observed at a
L:D of 16:8 at 240C, but it is important to recognize that,
like most calling and male response rhythms of moths,
the expression of these rhythms is modulated by current
and recent temperature regimes, with lower or decreas-
ing temperatures generally advancing the rhythms of the
night-calling species and delaying rhythms of the day-
active species (Card6 & Roelofs 1973; Schal & Card6
1986). Such temperature-induced modulation of calling
periodicities, however, does not reduce their importance
in isolation of some species pairs.

FLIGHT PERIODS AND HABITAT PREFERENCES
In addition to differences in diel rhythms and po-
tential differences in pheromone blends, distinctive flight
periods and habitat preferences isolate some co-occur-
ring species. In Connecticut and Massachusetts, V
aurantiaca has two broods, with adults occurring mainly
in secondary succession habitats (abandoned hay fields,
power line cuts) from late May to late June and again in
August. In the same areas, V ferruginosa is single-
brooded, with adult emergence in July in open scrub-
oak forests. These habitat types can be contiguous, but
the distinctive emergence patterns alone provide full iso-
lation of the two species in southern New England. The
female calling rhythms of both species from Connecti-
cut overlap broadly (Fig. 8), with calling of V aurantiaca
peaking within the first hour of scotophase (dark seg-
ment of the light-dark cycle) and lasting about 4 hours
(Fig. 8a) and calling of V.ferruginosa commencing af-
ter two hours of scotophase and also lasting for about 4
hours (Fig. 8d). Such differences in female calling
rhythms would appear to provide only weak isolation for
these two species, and, unlike most moths, the diel rhythm
of male attraction in V aurantiaca is considerably
broader than the female's nocturnal calling rhythm: male
attraction to synthetic pheromone occurs from mid-mom-
ing to evening (Card6 1974). Although these temporal
and habitat characteristics in concert provide effective
isolation for V aurantiaca and V ferruginosa in south-







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


(a) 100

90
80
70


50
40
e 40
C 30

20
10
0
10 12 14 16 18 20 22 24 2 4 6
calling time


(c) 100

90
80
0 70
S60
S50
e 40
0
30

20
10
0




(e) 100

90
80
70
= 60
50
S40
30

20
10
0


12 14 16 18 20 22 24 2 4 6
calling tmme


10 12 14 16 18 20 22 24 2 4 6
calling time


(b) 100

90


70
C
60
U
50
0
e 40
a
F 30
20
10
0




(d) 100

90
go
8O
a 70
S60
50
2 40
40
CL 30

20
10
0


lmmcarrat


10 12 14 16 18 20 22 24 2 4 6
calling time


10 12 14 16 18 20 22 24 2 4 6
calling time


) 100
90
80







70
10
0

10 12 14 16 18 20 22 24 2 4 6
calling time


Figure 8. Calling periodicities of 2-day-old females in the Virbia aurantica complex. Observations were conducted
every 30 minutes at 24 C in a 16:8 light:dark regime (see Carde 1971 for methods). Time of scotophase indicated by
the hatched bar. The sources for the lab colonies were: A-V aurantiaca, East Haddam, CT.; B-V immaculate,
Dryden, NY; C-V rubicundaria, Gainesville, FL; D-V ferruginosa Branford, CT.; E-V fragilis, Boulder, CO; and
F-V nigricans, Frenchtown, NJ.






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

em New England, both species are single-brooded in
Maine (Brower 1974), Nova Scotia (Ferguson 1954),
and generally throughout southern Canada, sharing the
same flight period from July to mid August. Some par-
titioning by habitat also seems important in Nova Scotia
(Ferguson 1954).
In southeastern Pennsylvania (Philadelphia Co.),
A.M. Shapiro collected V aurantiaca, V nigricans,
and V immaculate in the same old field, secondary suc-
cession habitat. Virbia nigricans is a day-calling spe-
cies (Fig. 8f), whereas, as mentioned, V aurantiaca
calls in early evening (Card6 1974), and V immaculate
(Card6 & Roelofs 1973) calls somewhat later in the
evening (Fig. 8b). Such partitioning of calling times would
afford some, but not complete isolation of V nigricans
from the other two species. The male response win-
dow of V aurantiaca includes attraction in daytime in
southeastern Pennsylvania as it does in the Connecticut
population (Card6 1974).
Similar levels of reproductive isolation by habitat
preference and calling times should isolate V lamae from
V aurantiaca and V ferruginosa where they co-oc-
cur. Virbia lamae is a day-calling species (Schal &
Card6 1986; Schal et al. 1987), and it is exclusively a
denizen of bogs.
Partial temporal (adult emergence, calling times)
and spatial isolation occur for four species of the V
aurantiaca complex (Table 1). Virbia aurantiaca is
bivoltine with adults emerging from May to June and
again in August in southern New England and the mid-
Altlantic United States (Card6 1971); adults of V.


14 16 18 20 22 24 2 4 6
calling time


Figure 9. Calling periodicity of 2-day-old Virbia lamae
females from Manistique, MI conducted hourly at 240 C
in a 16:8 light:dark regime (see Schal and Card6 1986
for methods). Time of scotophase indicated by the
hatched bar.

aurantiaca emerge year-round in the south (Forbes
1960). Virbia immaculate is multivoltine in the north-
central and northeastern United States, with adults
emerging from late May and June and again in August
and early September (Zaspel database). Two species,
Vferruginosa and V lamae, are univoltine with a July
flight period (Forbes 1960, Card6 1971). Virbia
rubicundaria and V nigricans are multivoltine, with
adults emerging year-round in the southeastern United
States (Zaspel database).
For those species that occasionally overlap in emer-
gence and locality, female calling times may be impor-


Table 1. Biological information for nine V, !,i species (Card6 1968 & 1971).
information not available.


n = chromosome number.


Species Chromosome # Female Male Male comes Voltinism
calls attracted to lure

V aurantiaca n = 25 yes yes 10am-10pm bi/multivoltine
V ferruginosa n = 32 yes yes night univoltine
V fragilis n = 25 yes yes -
V immaculate n =21 yes yes night multivoltine
V lamae n = 28 yes weak day univoltine
V nigricans n= 16 yes very weak day multivoltine
V rubicundaria n = 23 yes yes night multivoltine
V rindgei weak day -








tant in species discrimination. Three species are active
and mate diurnally; four are active nocturnally (Table
1). Males of V aurantiaca have the broadest known
male activity period with male attraction to female phero-
mone from midmorning (10:00) to early evening (20:00)
(Card6 1974) (Table 1). Virbia rubicundaria females
call nocturnally (Card6 1971). As yet unidentified com-
ponents of females' pheromone may confer some speci-
ficity in the males' response, resulting in diminished at-
traction of heterospecifics (Carde 1971).
Even if occasional mating of heterospecifics oc-
curs, the range of chromosome numbers among the spe-
cies suggests that most hybrid combinations would be at
a disadvantage (Table 1). Haploid numbers range from


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

16 (V nigricans) to 32 (V.ferruginosa) (Table 1, Card6
1971).

TAXONOMIC HISTORY OF HOLOMELINA
AND VIRBIA
Here we review the taxonomic history of Virbia, in-
cluding Holomelina. We also provide a taxonomic re-
view of species, but limit ourselves to those that occur in
North America north of Mexico, all of which formally
were placed in Holomelina. We first review the exten-
sive taxonomic history of Holomelina, and then review
the literature for Virbia. Last, we present a brief sum-
mary of the recent phylogenetic study and its taxonomic
implications (Zaspel & Weller 2006).


Table 2. Taxonomic History of Virbia. B & B = Barnes & Benjamin, B & MD = Barnes & McDunnough, G-M =
Gu6rin-M6neville, N& D = Neumogen & Dyar.


Family Genus Species Synonyms Genus


Lithosiidae


Geometridae


Geometridae


Lithosia


Lithosina


Crocota


Eubaphe


laeta G-M


opella Grote





costata Stretch.


ferruginosa Walker





ostenta Edwards
immaculate Reakirt
nigricans Reakirt
rubicundaria Huibner


fragilis Strecker
aurantiaca Huibner


treatii Grote
rubropicta Packard

flava B & B
obscura Stretch
belmaria Ehrmann
rubricosta Ehrmann

opelloides Graef
intermedia Graef
parvula N. & D.
cocciniceps Schaus
pallipennis B & MD
quinaria Grote
choriona Reakirt
trimaculosa Reakirt
bucholzi Wyatt

calera Barnes

nigrifera Walker
!. f, ,O.i Stretch
diminutive Graef
rosa French

bimaculata Saunders
brevicornis Walker


Crocota
Lithosia


Crocota
Holomelina
Holomelina
Holomelina

Crocota
Crocota
Crocota
Eubaphe
Eubaphe
Crocota
Holomelina
Crocota
Holomelina

Holomelina

Cothocida
Crocota
Crocota
Crocota


Crocota
Crocota


lamae Freeman
marginata Druce





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

TAXONOMIC REVIEW OF HOLOMELINA
The taxonomic history of Holomelina Herrich-
Schaffer is intertwined with two other genera, Eubaphe
(Hibner 1823) and Crocota (Hibner 1823). These two
genera originally were described among a series of
lithosiine genera, and subsequent authors inferred that
Hibner considered these genera as Arctiidae (Watson
et al. 1980). Eubaphe and Crocota are now placed in
Geometridae (Fletcher 1954; Watson et al. 1980), but
the associated geometrid and arctiid species were treated
as Eubaphe for nearly 120 years (Beutenmiller 1898;
Druce 1885; Schaus 1901; Forbes 1910; Barnes &
McDunnough 1917, 1918; Freeman 1941; Ferguson
1954). Similarly, arctiid and geometrid species were
described in Crocota through the 1800s (Walker 1854;
Grote 1863; Graef 1887; Smith 1889) or treated as such
in checklists (Neum6gen & Dyar 1893a). When Hibner
described Eubaphe, he listed a single species, E. lobula
(Hibner 1823), and thus E. lobula is the type species
by monotypy (Fletcher 1954). Hibner subsequently
described Eubaphe aurantiaca (Hiibner 1827);
Herrich-Schaffer used E. aurantiaca as the type spe-
cies for Holomelina (Herrich-Schaffer 1855). Unlike
Eubaphe and Crocota, Holomelina has always been
placed inArctiidae (e.g., Holland 1903). Hampson (1920:
316) asserted that Eubaphe had priority over
Holomelina, and most taxonomists followed Hampson
until Fletcher (1954) revised Eubaphe. In his revision,
Fletcher (1954) noted that the type species, E. lobula,
was a geometrid, and "the next valid name to replace
Eubaphe in the Arctiinae appears to be Holomelina
Herrich-Schaffer 1855, the type species of which is
Eubaphe aurantiaca Htibner" (Fletcher 1954: 155).
However, as late as 1960, some authors considered
Eubaphe to be the valid generic name for these arctiid
species (e.g., Forbes 1960: 21). The confusion of three
generic names, Eubaphe, Crocota, and Holomelina,
has resulted in congeneric species and subspecies being
described in different genera (Table 2).
Although Holomelina species (regardless of ge-
neric name) have been treated as Arctiidae, the three
genera, Holomelina, Eubaphe, and Crocota have been
placed in various subfamilies, including Lithosiinae
(Walker 1854), Arctiinae (Neum6gen & Dyar 1893a,b),
and Eubaphianae (Smith 1889). Earlier authors associ-
ated Eubaphe and Crocota with Lithosiinae, listing these
genera in sequence with lithosiine genera Cisthene and
Delphyre in checklists (Stretch 1885), species' descrip-
tions (Walker 1854), and faunal treatments (Seitz 1914).
Eubaphe (=Holomelina) was placed inArctiini (Forbes
1960), and Holomelina subsequently has remained in
Arctiini (Ferguson 1985; Franclemont 1983; Watson &


Goodger 1986; Jacobson & Weller 2002).
Discrimination of species was contentious in the
1800s (Smith 1889), and species taxonomy was further
complicated with individual species and subspecies de-
scribed in lithosiine genera (Lithosia author, Brycea
Walker, Boenasa Walker, Ptychoglene Felder), arctiine
genera (Cothocida Walker, Cytorus Grote), or geometrid
genera (Crocota Walker, Eubaphe Walker) (Table 2).
In some cases, subspecies were described in different
genera than the original species (e.g. Lithosia laeta
Guerin-Meneville and current syonoym Crocota treatii
Grote; Table 2). Descriptions were often cursory and
so general, that they applied to several species without
distinguishing among them, for example, Crocota costata
Stretch (Stretch 1885). In phenotypically variable spe-
cies, such as V costata and V.ferruginosa, a prolifera-
tion of names resulted (Table 2).
Complicating matters further, some authors refused
to acknowledge the work of others. In his Preliminary
Catalogue of the Arctiidae of Temperate North
America with notes, Smith (1889) states: "The matter
has been further confused by Mr. Grote's persistent re-
fusal to refer to Mr. Reakirt's species." Even when
authors discussed others' work, agreement on species
names could not be reached, resulting in the application
of several names to the same species (see Smith 1889).
Depending on the author treating the Nearctic fauna,
the number of valid species ranged from five (Walker
1854) to 17 (Smith 1889), and the number of varieties
ranged from none (Walker 1854) to seven (Neum6gen
& Dyar 1893a,b).
In the early 1900s, the species' numbers were re-
vised down to either six species (one variety) (Holland
1903) or eight species for the U.S. and Canadian fauna
(Barnes & McDunnough 1917). Holland commented
that "great confusion" existed concerning the number
of species, and that larval rearing and mating studies
were needed to resolve problematic species (Holland
1903: 115). Contemporary taxonomic treatments that
included species occurring in Central and South America
recognized 28 Eubaphe species and six varieties (Strand
1919) or 26 species, nine subspecies, and three varieties
(Seitz 1914).
Little additional taxonomic activity occurred until
the 1950s and 1960s. Ferguson (1954) reviewed the
Canadian fauna and renewed interest in the genus.
Ferguson (1954) noted that species in Eubaphe had been
difficult for taxonomists for some time; he stated, "The
old question of which is a species and which is not, still
remains paramount" (Ferguson 1954: 212). In treating
the E. aurantiaca species complex, Ferguson discov-
ered that differences in male genitalia between Eubaphe





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Figure 10. Strict consensus of 779 trees: L= 1698, CI= 41, RI= 65, resulting from the analysis of all taxaAT matrix.
Numbers below the branch are decays, and unlabelled branches have a decay of 3. Cl C6 = Clades 1 through 6.
Parallel lines = species formerly placed in Holomelina, Solid lines = species originally placed in Virbia, dotted line =
outgroups.





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

species were scarce, and offspring reared in the labora-
tory were phenotypically diverse, i.e., they did not match
the parents. (Ferguson 1954: 212). Extensive reared
series at the National Museum of Natural History
(NMNH) document the array ofphenotypes. Additional
reared series are housed at the American Museum of
Natural History (AMNH, Joseph Muller), Cornell Uni-
versity (CUIC, J. Franclemont material), and the Uni-
versity of California, Riverside (UCR, R. Card6 mate-
rial).
Forbes (1960) provided a synopsis of the known
biology of the genus and commented on the misuse of
the generic name Eubaphe. He defended its usage as
providing taxonomic stability in general usage (footnote
3a, Forbes 1960: 21). Forbes provided illustrations of
diagnostic male genitalic features and a short key to the
H. opella and the H. aurantiaca complex consisting of
H. aurantiaca, H. laeta, and H. lamae. Other names
were subsumed as varieties.
In 1965, Card6 published redescriptions of H.
aurantica, H. ferruginosa, and H. lamae and provided
a partial key to the Nearctic Holomelina species. Card6
(1965) commented on the sibling species in the H.
aurantiaca complex, listing H. aurantica, H.
ferruginosa, and H. lamae as distinct species. In this
paper, Card6 provided commentary on the H.
aurantiaca and H. opella complexes. He was the first
to realize that H. nigricans was not a synonym of H.
opella based on adult habitus and male genitalia (Card6
1965). Card6 summarized additional preliminary find-
ings stating, "Areas of uncertainty are currently under
investigation." In 1968 and 1971, Card6 completed a
set oftaxonomic, chromosome, behavior, and pheromone
studies on Holomelina for his M.S. and Ph.D. at Cornell
University. His taxonomic work is not available under
the International Code of Zoological Nomenclature
(ICZN 1999), because unpublished theses do not con-
stitute valid publications. He was unsuccessful in locat-
ing Huibner types for H. aurantiaca and H.
rubicundaria (Card6 1968, 1971), which complicated
his attempts to determine their species' limits. Card6's
work included two new species that remained as manu-
script names (Card6 1968) until now. Although Card6's
taxonomic revisions were not published, Franclemont
(1983) followed his treatment of Holomelina in the
Checklist of the Moths of America north of Mexico
(Hodges et al. 1983).

TAXONOMIC REVIEW OF VIBRIA
Virbia (Walker 1854) was described in Arctiidae
and associated with lithosiine moths. Its familial place-
ment has never been controversial (Nye et al. 1980;


Ferguson 1985). It is a typical member of the Arctiini
except some species possess homoideous crochets
(Forbes 1939, 1960; Ferguson 1985; Kitching & Rawlins
1999; Jacobson & Weller 2002). Virbia was erected by
Walker (1854), and he described two species, V mentions
and V luteilinea. Kirby (1892) subsequently designated
V mentions as the type species. Until recently (Zaspel
2004; Zaspel &Weller 2006), the only generic synonyms
were Bepara Walker (described in Melameridae) and
Terna Walker (described in Callimorphidae).
As of 1986, 31 of the 39 species placed in Virbia
were described in f i,', (Watson & Goodger 1986).
The remaining species were described in Bepara Walker,
Brycea Walker, Cothocida Walker, Josioides Felder,
Phaeochlaena Huibner, Ptychoglene Felder, or Terna
Walker. Again, the small size of the moths led some
authors to place these species in lithosiine genera (e.g.,
Lithosia Fabricius, Josioides Felder). A few species
were described in other families: Geometridae (Crocota
Huibner, Eubaphe Huibner), Dioptidae (Phaeoclaena
Huibner), and Noctuidae (Phalaena L.). In contrast to
the turbulent taxonomic history of Holomelina, Virbia
has received little revisionary attention. Specific no-
menclature has been comparatively stable; there were
only four synonyms and two subspecies recognized in
the most recent checklist of neotropical species (Watson
& Goodger 1986).

PHYLOGENETIC PLACEMENT OF VIRBIA AND HOLOMELINA
There has been a long taxonomic history of asso-
ciating these genera. Herrich-Schaffer (1855) was the
first to do so when he described Holomelina. He in-
cluded Phalaena medarda Stoll (Fig. 11) as one of two
species comprising Holomelina. Later authors did not
follow him and placed P medarda in F ',w Seitz (1914:
294) noted that Virbia species were structurally much
like Eubaphe but differed by the branching of the up-
permost median vein of the forewing. For over 90 years,
checklists and faunal treatments have associated
Eubaphe and fn,.,i (e.g., Seitz 1914; Forbes 1939;
Hampson 1920). Forbes (1939: 220) remarked that
Eubaphe and Virbia were closely related and very un-
like other arctiine genera. Ferguson (1985) listed char-
acters uniting T ', ',i and Holomelina in the Holomelina
generic group, and noted that they "might not be dis-
tinct" (Ferguson 1985: 199), in other words, congeneric.
Recent checklists (Hodges et al. 1983; Watson &
Goodger 1986) treated f, b,, and Holomelina as mem-
bers of the Arctiini. Phylogenetic studies have come to
the same conclusion (Kitching & Rawlins 1999; Jacobson
& Weller 2002).
A phylogenetic analysis based on adult morphol-





























Figure 11. Male genital capsule, Virbia medarda. 8S =
eighth sternite, Co = coremata (highly developed), J =
juxta, V = valve sacculuss region).


ogy was conducted to determine whether Holomelina
and f, !,,,,i were monophyletic with regards to each other
(Zaspel 2004; Zaspel & Weller 2006). Those studies
included 46 ingroup taxa (of 70 possible) and six arctiine
species as outgroups. All Holomelina species, and about
half the species of Virbia were included. Fifty-eight
characters (195 states) were described and scored from
non-genitalic and genitalic structures (Zaspel 2004;
Zaspel & Weller 2006).
The maximum parsimony analysis yielded 779 trees
with a length (L) of 1698, a consistency index (CI) of 41
and a retention index (RI) of 65 (Zaspel & Weller 2006;
Fig. 10). The monophyly of the clade comprised of
Holomelina and f ,,i, species was well supported.
The clade had a decay index greater than 10 (Bremer
1992) and was supported by five unreversed
synapomorphies: 1) M2 missing in the hindwing, 2) pres-
ence of a scaphium, 3) presence of a subscaphium, 4)
apical hooks on the uncus, and 5) juxta ornamentation.
The North American species with the Holomelina phe-
notype clustered into two clades, the H. aurantiaca clade
and the H. opella clade. The H. opella clade was
sister to the clade consisting of IVrhbia, the yellow and
black phenotype (Fig. 10). Multiple trees resulted from
conflicting resolutions of the apical clade of neotropical
species, consisting of those species traditionally assigned
to Tf /',1 Different analyses could not recover a recip-
rocally monophyletic f /,i', and Holomelina (Zaspel


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

2004; Zaspel & Weller 2006). That is, the H. opella
and H. aurantiaca clades were never grouped exclu-
sive of f ; i, species. Thus, the former Holomelina
was an artificial assemblage of two different dispersal
events from Mexico and the southwestern U.S. into
Canada and the central and eastern U.S. (Zaspel 2004;
Zaspel & Weller 2006). Holomelina (Herrich-Schaffer
1855) was placed as junior synonym of Tf, '1,1 (Walker
1854) (Zaspel 2004; Zaspel & Weller 2006) to reflect
the results of the study. No subgenera were recognized
because they would be either paraphyletic or typically
treated as Holomelina (e.g., V opella group).

SUMMARY
The taxonomic history of Holomelina is compli-
cated by the confusion with geometrid generic names
and the frequent acrimonious debates over species' lim-
its among 19' century American lepidopterists. These
debates were fueled by the phenotypic diversity of the
moths. As noted in the Biology Section, a single species
encompasses adult morphs found in monomorphic, co-
occurring species. Virbia, in comparison, has had a rela-
tively quiet taxonomic history.
The long-standing, phyletic association of Virbia
and Holomelina was confirmed (Zaspel & Weller 2006),
and in retrospect, the result was not surprising. The red
and yellow hindwing coloration and species' distributions
were the principal characters formally used to distin-
guish the genera. The phylogenetic study that supports
this synonymy is discussed at length in Zaspel and Weller
(2006). Below we provide an abbreviated checklist for
the species. A checklist for the entire genus can be
found in Zaspel and Weller (2006).

CHECKLIST OF VIRBIA FOR NORTH
AMERICA
In this checklist, species are arranged alphabetically.
Valid species are in bold with synonyms in italics. The
genus in which a species was originally described fol-
lows the author name in parentheses, and an asterisk
indicates that the type was examined. When the type is
not from the United States, the country is noted.

Genus Virbia Walker 1854
Bepara Walker, 1865a
Cathocida Hampson, 1901, misspelling
Cothocida Walker, 1865b
Crocata authors, misspelling
Crocota authors, not Hutbner, 1823
Cytorus Grote, 1866
Eubaphe authors, not Htibner, 1823
Holomelina Herrich-Schaffer, 1855





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

Terna Walker, 1865a
V aurantiaca (Hibner, 1827) Eubaphe
bimaculata (Saunders, 1869) Arctia
brevicornis* (Walker, 1854) Crocota
V costata (Stretch, 1885) Crocota
opelloides (Graef, 1887) Crocota
intermedia* (Graef, 1887) Crocota
parvula* (Neum6gen & Dyar, 1893b) Crocota
cocciniceps (Schaus, 1901) Eubaphe
pallipennis* (Barnes & McDonnough, 1918)
Eubaphe
V. fergusoni Zaspel n.sp.
V. f ,1111g11 i,'" (Walker, 1854) Crocota
quinaria (Grote, 1863) Crocota
choriona (Reakirt, 1864) Crocota
trimaculosa (Reakirt, 1864) Crocota
buchholzi* (Wyatt, 1963a,b) Holomelina
V f,,g111,"' (Strecker, 1878) Crocota
V immaculate* (Reakirt, 1864) Crocota
V laeta (Guerin-Meneville, 1832) Lithosia
treatii (Grote, 1865) Crocota
rubropicta (Packard, 1887) Lithosia
V lamae (Freeman, 1941) Eubaphe
V. ,,iiI1ihig0,"' (Druce, 1885) Eubaphe (Guatemala)
V. ig,,,,," (Reakirt, 1864) Crocota
,.,,,t,,'" (Walker, 1865b) Cothocida
V opella (Grote, 1863) Crocota
flava (Barnes & Benjamen, 1925) Eubaphe
obscura (Stretch, 1885) Crocota
belmaria (Ehrmann, 1895) Crocota
rubricosta (Ehrmann, 1895) Crocota
V ostenta* (Edwards, 1881) Crocota
calera* (Barnes, 1907) Holomelina
V rindgei Carde n. sp.
V rubicundaria (Huibner, 1827) Crocota
l!;,, i (Stretch, 1885) Crocota
diminutive (Graef, 1887) Crocota
rosa (French, 1890) Crocota

MATERIAL AND METHODS
SPECIMENS AND PREPARATION
Dissection methodology follows Winter (2000) and
is fully described in Zaspel and Weller (2006). Varia-
tions on standard techniques include using only cold 10%
KOH for 18-24 hours; most wings were not cleared
and slide-mounted. This was done to ensure the small,
fragile structures were not 'over cleared' prior to dis-
section. These moths have very fragile wings; there-
fore, 1-2 drops of 30% ethanol were added to the wings
of pinned specimens for observation ofvenation. Larger,
more robust wings placed briefly in weak bleach (30
seconds), and then standard procedures were followed


Figure 12. Virbia aurantiaca, male forewing, all R veins
present. C = costa, Sc = Subcosta, R'- = radial veins 1 -
5, M- '= medial veins 1 -3, CuA1 CuA2 = cubital anal
veins 1 and 2, A = anal veins.


Sc + Ri
Rs
M1
M3
-CuAl
CuA2


B
Figure 13. Virbia laeta, male, A- Forewing, R2 = R2 +
R3 fused or R3 absent. B- Hindwing, M2 absent, Sc + R'
subcosta fused with radial vein 1, Rs = radial sector
vein, other abbreviations as in Fig. 12.







R4
R1


Mi1
M2
M3
CuAi
CuA2
1A+2A


Figure 14. Virbia cyana, male forewing, R2 and R3 ab-
sent, R4 + R5 = fused other abbreviations as in Fig. 12.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

for staining with eosin Y (1% in distilled water; Fisher
Scientific, Pittsburgh, PA) and slide mounting. All prepa-
rations went through a dehydration series before mount-
ing (Winter 2000). Balsam preparations went through
two additional dehydration treatments of oil of clove (15
minutes), and then xylene (15 minutes) before slide
mounting. Euparol mounts were transferred from the
dehydration series into a final 15-minute treatment in
Euparol essence (Bioquip, Garden City, CA) before slide
mounting. Permanent slide mounts (Canada balsam
[Sigma, St Louis, MO] or Euparol [Bioquip, Garden City,
CA]) were made of abdominal pelts, genitalia, legs,
wings, labial palps and antennae. Slides were placed on
trays and cured in drying ovens for 24 48 hours. Terms
for wing venation follow Scoble (1992). Terms for ab-
dominal and genitalic morphology follow Klots (1970),
Weller et al. (2000), Jacobson and Weller (2002) and
Kristensen (2003). For females, when two bursae exist
(double or connate condition), we could not determine
which one was the appendix busa based on position or


A B


Figure 15. A. Condition of the tarsal claw, Virbia
aurantiaca, simple; B. Condition of the tarsal claw,
Virbia cyana, slightly bifid.


Figure 16. A. Virbia opella, male. An = antenna simple
and ciliate.; B. Virbia laeta, male. An = antenna emar-
ginate, asymmetrical flagellomeres.






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

presence of a signum. Therefore, we made no distinc-
tion between corpus and appendix in the descriptions.

MATERIAL EXAMINED
For the study of species limits, 180 specimens were
dissected, representing the 14 species described below;
dates from specimen labels have been standardized. In
addition, a specimen-level database was compiled in
Microsoft Excel that included locality data (label data),
prior identifications, comments, and repository for speci-
mens of Virbia species. The database includes over
12,000 entries from 16 collections (listed below) follow-
ing the protocol of the NMNH (M. Pogue, personal com-
munication). It is searchable and can be found of the
Diversity of Life website at www.diversityoflife.org/
virbia.html.
The following is a list of the institutional and pri-
vate collections consulted during this study. The acro-
nym of the institution or name of private collection is
followed by the name of the individual that prepared the
loan. Acronyms follow Heppner and Lamas (1982):
AMNH American Museum of Natural History, New
York (E. Quinter); CMNH Carnegie Museum of Natural
History, Pennsylvania (J. Rawlins); CUIC -Comell Uni-
versity Insect Collection, New York (J. Liebherr); CSU
- Colorado State University, Colorado (P. Opler); E. H.
Strickland Entomological Museum, University ofAlberta,
Alberta (F. Sperling); EMEC Essig Museum of Ento-
mology, University of California, Berkeley, California (J.
Powell); FMNH Field Museum of Natural History, Il-
linois (P. Goldstein); LACM Los Angeles County Mu-
seum, California (J. Donahue); MSU Mississippi State
Univeristy, Mississippi Entomological Museum, Missis-
sippi (R. Brown); UNSM University of Nebraska State
Museum, Nebraska (B. Ratcliffe); NMNH National
Museum of Natural History, Washington D.C. (M.
Pogue); TAMU -Texas A and M University Insect Col-
lection, Texas (E. Riley); UCONN University of Con-
necticut Insect Collection; Connecticut (D. Wagner);
UMSP University of Minnesota Insect Collection, Min-
nesota (P. Claussen); RTC Personal collection of R.
Carde, University of California-Riverside.

SPECIAL TERMS
Adbasal-This is the space before the basal bands in most
other noctoids. In arctiids it can be present, but is usually
lost, or fragmented and united with the antemedial band.

Basal-This represents the space between the basal and
antemedial band.

Antemedial-This can be complete or fragmented. Be-


cause of this fragmentation, this band is almost always a
composite of the basal and antemedial bands, and it is
uncertain how often a true antemedial band occurs.

Medial-A band near the middle of the wing between the
antemedial and postmedial areas, sometimes incomplete
and apparent as a spot or bar near the costa.

Postmedial-This is the pale space beyond the postme-
dial line. This is often present as a full band.

Subterminal-This is a zigzag band running from a point
on the costa about midway between the postmedial band
and apex toward the tomus-the anal region or anal angle,
merging with the postmedial band between M3 and Cu 1,
then bending sharply outward again. This band may be
complete, fragmentary or absent.

SPECIES DESCRIPTIONS
Species were delimited primarily by differences in
the male genitalia, especially characteristics of the vesica,
phallus, and juxta. Female genitalia are diagnostic for
some. Due to a redundant distribution of character states
between species (e.g., little interspecific variation), di-
agnostic morphological characters typically are illustrated
once and are referred to in multiple descriptions. Some
Virbia species are almost indistinguishable by their geni-
talic characteristics; in addition, a variety of wing color
patterns occur, rendering species determination difficult.
Within the North American fauna, species range from
pale pink and yellow (e.g., V ferruginosa and V
immaculate) to deep red and brown (e.g., V ostenta).
Some species (e.g., V aurantiaca) have wing patterns
that vary within long reared series from a single female
(Fig. 1). Although variable, these species do share com-
mon pattern elements with other arctiines. All speci-
mens were examined under a true color lamp (OTT-
LITE), and colors are described using the Naturalist's
Color Guide (Smithe 1975).
In general, arctiids have greatly modified wing pat-
tern systems compared to their closest relatives, the
Noctuidae (Ferguson 1985, sensu Fibiger & Lafontaine
2005, not Lafontaine & Fibiger 2006). The forewing
pattern typically consists of light stripes and spots against
a darker background, a reverse of the classic noctuid
pattern. The forewing pattern usually is based on six
lightly colored, transverse bands on a dark background,
these are almost always reduced to five or fewer, or
lost, with many variations occurring within a genus
(Ferguson 1985). Ferguson notes:

Two typical pattern systems occur on
the forewings of many Arctiini: a lon-








gitudinal vein pattern and a supplemen-
tary longitudinal pattern consisting of
paracostal, cubital, and postcubital band,
also called the primary and secondary
longitudinal patterns, respectively
(Ferguson 1985:183).

Similarly, the basic hindwing pattern ofArctiini is
reversed from the noctuid one, and consists of a colored
background with either dark spots or dark outer-mar-
ginal borders (Ferguson 1985). A discal spot or spots
may be present, but often are modified or lost (Ferguson
1985).
Nomenclature for pattern elements of Virbia fol-
lows Ferguson (1985). Descriptions of wing regions,
patterns, and colors specific to Virbia species are de-
fined in the "Specialized Terms" section. The described
pattern elements are strictly for species' identification
purposes and are not intended to be interpreted as ho-
mologies of pattern elements across Noctuoidea.
To facilitate the construction of consistent diagnoses
and descriptions, DELTA (Descriptive Language for
Taxonomy) was used (Dallwitz et al. 1993). Species
descriptions were first constructed using DELTA, and
then minor formatting changes were made.

SYSTEMATIC ENTOMOLOGY
Order LEPIDOPTERA
Family ARCTIIDAE
Subfamily ARCTIINAE
Tribe ARCTIINI
Genus Virbia Walker
Virbia Walker, 1854: p. 470. Type species: Virbia
mentions Walker, 1854.
Bepara Walker, 1865a.
Cathocida Hampson, 1901, misspelling.
Cothocida Walker, 1865b.
Crocata authors, misspelling.
Crocota authors, not Hfibner, 1823.
Cytorus Grote, 1866.
Eubaphe authors, not Hfibner, 1823.
Holomelina Herrich-Schaffer, 1855.
Terna Walker, 1865a.
Diagnosis (Figs. 1,2,4,5, Pl. 1-3).-- These are
relatively small arctiids, with forewing length ranging from
8.2 mm to 17.1 mm in males (N= 14) and from 8.7 mm
to 18.5 mm females (N = 11). Antennae are typically
simple and ciliate (Fig. 16A). Forewing with two
predominant pattern types: dark forewings and hindwings
with bright yellow markings (Fig. 5) or yellowish areas
replaced with pink or reddish markings (e.g., V ostenta)


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

(Fig. 4). Male genitalia are unique in Arctiini in the pos-
session of a true scaphium (except for V cyana) (Figs.
25A, B). The corpus bursa can be single, double, or
connate with the corpus (Fig. 17C). Virbia can be sepa-
rated from other Arctiidae by the loss or reduction of
the M2 in the hindwing (Fig. 13B) and the scaphium.
Description.-- Adult habitus (Figs. 1,2,4,5, P1.
1-3): Head color uniform, but variable, from dark black
(Fig. 2) or brown (Pl. 3, Nos. 2-3) to pinkish orange
(Fig. 4). Third segment of labial palpus typically long,
exceeding front by at least half its length; color variable
from yellow to pink, gray, or varying intensities of brown.
Ocelli absent. Thorax with dorsal and ventral scale col-
oration variable, from dark black (Fig. 5) or brown (P1.
3, Nos. 2-3) to pinkish orange (Fig. 4). Forewing typi-
cally with R' R5 all present (e.g., V aurantiaca: Fig.
12), R3 absent or trace in V laeta (Fig. 13A), and R2, R3
absent or trace in V cyana (Fig. 14). Hindwing with
M2 absent (Fig. 13B). Wing color pattern one of two
main types (see diagnosis). Abdomen with color vari-
able from dark black (Fig. 5) or brown (Pl. 3, No. 2) to
pinkish orange (Pl. 2, No. 4). Sclerotization of male 8'
stemite often species specific, shapes including: trian-
gular (e.g., V.phalangia: Fig. 28A), oval (e.g., V. opella:
Fig. 28B), pentagonal (e.g., V marginata: Fig. 28C),
pentagonal with expanded base (e.g., V divisa: Fig.
28D), antecosta reinforced by transverse ridge, rectan-
gular width greater than length (e.g., V luteilinea: Fig.
28E), and M-shaped (e.g., V metazonata: Fig. 28F).
Coremata typically present, but variable developed.
Male genitalia: Phallus typically with coecum penis
rounded, occasionally weakly notched (e.g., V
ferruginosa: Fig. 22A). Vesica directed ventrally in
most species, directed posteriorly in V latus, V cyana,
and V. polyphron. Phallic sclerite typically present, but
variable: narrow, rounded triangular (e.g., V ostenta:
Fig. 22B), wide and nodular (e.g., V aurantiaca: Fig.
22C), triangular, blunt (e.g., V inversia: Fig. 18A), sharp
and triangular (e.g., V endophaea: Fig. 18B), and square
(e.g., V. polyphron: Fig. 18C). Vesica typically rugose
with endophallus ornamentation on right side, under phal-
lic sclerite present, either as ventral comutuslike patch
(e.g., V aurantiaca: Fig. 22C) or peglike patches (e.g.,
V endophaea: Fig. 18B). Genital capsule (Figs. 25A,
25B) dorsad with tegumen either M-shaped or concave
(e.g., V costata: Fig. 23A), with lobes (e.g., V
aurantiaca: Fig. 23B), or entire (e.g., V. laeta: Fig. 23C).
Ventrad with uncus base typically cordate (e.g., V
aurantiaca: Fig. 23B). Ventrally directed hooks present
on apex of uncus with bases fused or separate; usually
two hooks present (e.g., V costata: Fig. 23D), but only
one in V semirosea and lacking in V cyana, V





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

phalangia, and V polyphron. Scaphium present (ex-
cept V cyana), conical (e.g., V laeta: Fig. 23E), or flat-
tened (e.g., V phalangia: Fig. 23F). Costa typically
elongate with apex entire (e.g., V laeta: Fig. 25A), some-
times forked (e.g., V opella: Fig. 27D). Saccus typi-
cally rounded, setose (e.g., V marginata: Fig. 27C) in-
frequently thin, square (e.g., V. feronia: Fig. 27E). Fe-
male genitalia: Papillae anales convex, rounded;
pseudopapillae anales typically present (e.g., V.
nigricans: Fig. 24B), infrequently lacking. Posterior apo-
physes entire, long and slender. 8ttergite unornamented.
Anterior apophyses entire, long and slender. Eighth ster-
nite round, wide and entire. Location of ostium bursa
just above 8' sternite, ornamentation of ventral edge
variable: central broad indentation with small lateral
prominences (e.g., V aurantiaca: Fig. 17A), straight
(e.g., V lamae: Fig. 17B), split in center with lateral promi-
nences (e.g., V opella: Fig 30A), small lateral promi-
nences (e.g., V. rubicundaria: Fig. 30B), or toothed (e.g.,
V satara: Fig. 30C). Shape of ductus bursa variable:
rectangular (e.g., V aurantiaca: Fig. 17A), hourglass-
shaped (e.g., V lamae: Fig. 17B), goblet-shaped (e.g.,
V opella: Fig. 17C), or rectangular, tapering in middle of
base (e.g., V metazonata: Fig. 17D). Sclerotization of
ductus bursa extending into corpus bursa, except in V.
marginata. Corpus bursa with (e.g., V. opella: Fig. 17C)
or without additional connate bursa (e.g., V aurantiaca,
V lamae: Fig. 17A & B), typically with one to three
round signa.
Discussion.- Species with the Holomelina pat-
tern are often polymorphic, and several phenotypes may
occur in a single brood (Fig. 1). Rearing studies are
needed of species exhibiting the yellow-black Virbia
phenotype to determine whether polymorphism is preva-
lent. In Mexico, species with T, ii and Holomelina
patterns co-occur.
Both male and female genitalia are useful for dis-
tinguishing phenotypically confusing species. The lobing
and ornamentation of the vesica (endophallus) is often
species' specific. In several species, females exhibit
species' specific ornamentation of the ostium bursa or
corpus bursa, and several species have pseudopapillae
anales (Fig. 24B).
These moths often are found misplaced in
uncurated Geometridae, Notodontidae (Josia), or
lithosiine arctiids. They can be separated from
Geometridae by the placement of the tympana on the
metathorax (versus on the base of the abdomen in
Geometridae). They can be separated from Josia based
on forewing venation (M2 arising from mid discal cell in
Josia and from near M3 in T, ',i). The hindwing vena-
tion (M2 absent in Virbia) usually will separate these


species from other arctiids.

Virbia aurantiaca Huibner
P1. 1 No. 5 (female), Nos. 3-4 (males)
Eubaphe aurantiaca Hilbner 1827: 206 [type lo-
cality: Pennsylvania, USA].
Crocota bimaculata Saunders 1869: 5 [type lo-
cality: London, Ontario, Canada].
Crocota brevicornis Walker 1854: 2 [type local-
ity: United States].
Crocota choriona Reakirt 1864: 371 [type local-
ity: Philadelphia, Pennsylvania, USA].
Holotype.-- The holotype of Eubaphe
aurantiaca is lost according to Lloyd Martin, Curator
of Lepidoptera at Naturhistorisches Museum, Wien (per-
sonal communication 6/25/2004). A neotype is desig-
nated here and labeled to ensure nomenclatural stability.
The specimen was chosen from the nearest locality to
the original type locality. The neotype, male [CMNH] is
labeled: Virbia aurantiaca (Hbn. 1827) [RED LABEL].
Label data: At light New Lisbon, N.J. Aug. 7 1934, E.P.
Darlington//Exch. A.N.S.P. GM. Acc. 20359.
Material examined.- Type material: Neotype des-
ignated above. All material is from the NMNH unless
otherwise noted: CANADA: Nova Scotia: Lequille,
1.viii.1946, D. C. Ferguson (1 Male, JMZ100). Que-
bec: Temiscouta Co., 30.vi.1985, leg. George Hensel (1
Male, JMZ94). MEXICO: Cordoba: 16.v.1908, Fred
Knab (1 Female, JMZ92). Jalapa: Wm. Schaus Col-
lection (1 Male, JMZ93). UNITED STATES: Arkan-
sas: Washington Co., Springdale Blue Spruce Camp-
ground, 28.v.1966, R. Heitzman, (1 Male, JMZ96).
Connecticut: New Haven, Hamden, 3.ix.1967, D. C.
Ferguson (1 Male, JMZ119). Kansas: Labette Co.,
Oswego, 3.vi.1966, Gary F. Hevel collector, at lights (1
Male, JMZ 115). Kentucky: Oldham Co., Homer Bird
Sanctuary 25.viii. 1977, C. V. Covell Jr. (1 Male, JMZ 113).
Maine: Southwest Harbor, 21.vii.1936, near Colorado,
marginata as in Barnes Collection 1937 (1 Male,
JMZ183); Willimantic, 15.vii.1930, H.Wilhelm (1 Male,
JMZ 101). Maryland: Queen Annes Co., Gransonville
Horsehead Sanctuary, 24.vii.1986, D. C. Ferguson (1
Female, JMZ109). Michigan: July Barnes Collection
(1 Male, JMZ112). Montana: Sweetgrass Co., Big Tim-
ber Creek 7 mi. N. Big Timber, 29.vii.1966, D. C.
Ferguson (1 Male, JMZ99). New York: Long Island,
Greenport, viii (1 Male, JMZ103). New Jersey: Or-
ange Mts., 24.vii. 1937, Otto Bucholz (1 Female, 1 Male,
JMZ110; JMZ111). North Dakota: Slope Co., Bum-
ing Coal Veins Pine Ridge, elev. 2500', T 136N, R 102W,
Section 11, 21.vii.1982, Eric H. Metzler (2 Males,
JMZ 116; JMZ 118). Oklahoma: Sequoia Co., Tenkiller





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Ob


Db



I.___


-'B


B.-Db
SI


Ds


B


~W


Figure 17. A. Bursa complex, Virbia aurantiaca, ventral view. B = bursa (membrane wrinkled), Db = ductus bursa
(rectangular and narrowing toward ventor), Ob = ostium bursa (central, broad indentation and small lateral promi-
nences); B. Bursa complex, Virbia lamae, ventral view. Db = ductus bursa (hourglass), Ob = ostium bursa (straight),
S = signum; C. Bursa complex, Virbia opella, dorsal view. B = bursa connatee and arising from ductus, membrane
of additional lobe wrinkled), Db = ductus bursa (goblet, sclerotization extending from ductus bursa into corpus bursa);
D. Bursa complex, Virbia metazonata, dorsal view. B = bursa connatee, arising from ductus; membrane of additional
lobe smooth), Db = ductus bursa (square).


-0d'





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

Lake, 3 mi. W. Blackgum, 6-9.vii. 1979, D. & M. Davis
(1 Male, JMZ117). South Carolina: McClellanville,
Wedge Plantation, 22.iv.1974, D. C. Ferguson (1 Male,
JMZ102). Tennessee: Nashville, 16-23.vi., Barnes
Collection (1 Male, JMZ95). Texas: Concan, Gamer
Ste. Pk. 1.vi.1964, A. & M. E. Blanchard (1 Male,
JMZ62), Hill Co., Lake Whitney Park, 21.iv.1985 (1
Male, JMZ98); Sinton, Welder Wildlife Foundation,
22.iv.1964, A. & M. E. Blanchard (1 Male, JMZ67);
Cameron Co., Brownsville, 17.xi. 1941, Collection of Fred
Rindge, AMNH (1 Male, JMZ288); SantaAna Wildlife
Refuge, 21.x.1970, A. & M. E. Blanchard (2 Females,
JMZ 194; JMZ 196). Virginia: Leesburg, 4.vii.1970, D.
C. Ferguson, reared ex ovo on dandelion (1 Male,
JMZ108). Wisconsin: Lindsey, 23.vii.1964, J. W.
Barnes (1 Male, JMZ97). Wyoming: New Castle, 6
mi.N.W., 28.vi.1965, R. W. Hodges (1 Male, JMZ114).
Diagnosis. i,.', b aurantiaca may be confused
with V ferruginosa, V lamae, V marginata, V.
nigricans, and V rubicundaria, but it is most com-
monly confused with V. ferruginosa. The two species
are separated using hindwing maculation and potentially
emergence dates. The subterminal hindwing band of V.
aurantiaca is solid, whereas the subterminal area of V.
ferruginosa has a series of blotches and spots, not a
solid band. In New York and neighboring states, V.
ferruginosa emerges in July, whereas broods of V.
aurantiaca emerge in June and August (Cardel965,
1974). Flight period is less reliable in the Maritime Prov-
inces, northern Michigan, and northern Maine, where
there is a single brood and the adult emergence of V.
aurantiaca overlaps with V ferruginosa (Zaspel data-
base). In the southern states (e.g., Georgia, Florida), V.
aurantiaca may be encountered anytime in the sum-
mer (Zaspel database, Adams personal communication).
Virbia aurantiaca can be separated from V.
nigricans and V lamae if detailed label data are avail-
able. Tb 'a'1,1 lamae is a bog endemic and males fly (with-
out flushing) diurnally. f .'i 0 nigricans has been col-
lected only in a narrow corridor of disturbed secondary
forest on the border of western New Jersey (Hunterdon
County) and eastern Pennsylvania (Philadelphia area).
This species is much darker and smaller (wing length
8.2 mm) (Figs. Plate 2, Nos. 8,9) than V aurantiaca
(wing length 9.7 mm) (Pl. 1, Nos. 3-5). The shape of
the juxta also separates the three species. In V.
aurantiaca, the juxta is ovoid with the ventral margin
expanding into a larger, orbicular shape (Fig. 20B),
whereas in V lamae and V nigricans, it is cylindrical
and emarginated as in V marginata (Fig. 27C). In fe-
males, V aurantiaca has pseudopapillae anales; the other
two do not. Lateral prominences surround the ostium


bursa in V aurantiaca (Fig. 26D) and V nigricans but
not V lamae (Fig. 26B).
Virbiafragilis, a western species, occasionally is
misidentified as V aurantiaca, especially specimens
from the western edge of V. aurantiaca's range. Virbia
fragilis is a much larger species (14 mm), and the
hindwing color is flesh ocher and lacks subterminal mark-
ings. The phallic sclerite is wide and nodular in V
aurantiaca (Fig. 22C), whereas it is highly reduced in
V fragilis.
In Central America, V aurantiaca co-occurs with
V marginata (range: Guatemala, Mexico, SW USA).
Diagnostic differences are provided under V marginata.
Occasionally, pale specimens of V aurantiaca lack
the subterminal hindwing band, and these can be con-
fused with the boreal V immaculate or V rubicundaria,
but V immaculate is a larger species (forewing length
12 mm) and has a reduced phallic sclerite compared to
V aurantiaca (Fig. 22C), and V rubicundaria is a
smaller species (forewing length 8.9 mm). Females of
V aurantiaca can be confused with those of V
rubicundaria, but V rubicundaria is restricted to the
southeastern USA and is rarely sympatric with V
aurantica. Virbia rubicundaria females are also much
smaller (forewing length 8.7 mm vs. 9.7 mm, respec-
tively) and brighter than V aurantiaca females (com-
pare Pl. 2. No. 4, Pl. 1, No. 5).
Description.-- Male habitus (Pl. 1, Nos. 3-4):
Head dark clay throughout. Antennal scales dark clay.
Third segment of labial palpus exceeding front of head
by at least half of entire length, first and second seg-
ments salmon, apex of third segment grayish horn. Tho-
rax dorsal and ventral scales clay throughout. Legs clay,
tarsal claw simple (Fig. 15A). Forewing length 10 mm
(N = 10); R'-R all present, m3 present, extending to
wing margin (Fig. 12). Dorsal surface ranging from
clay to cinnamon throughout; ventral surface salmon.
Hindwing dorsal surface peach red, typically with solid
raw umber subterminal band, ventral surface salmon.
Abdomen with dorsal and lateral abdominal scales clay
to salmon, dorsal tergites occasionally with medium size
fawn spot in center, size of spot tapering with each seg-
ment towards posterior, ventral abdominal scales yellow
ocher throughout. Sclerotization of 8th sternite penta-
gon (Fig. 28C). Coremata reduced to small patches of
scales on membranous lobes. Female habitus (Pl. 1,
No. 5): Head clay. Antennal scales clay. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length, all segments clay. Thorax dorsal,
lateral, and ventral scales salmon to clay. Legs clay
with patches of salmon scales on femora and tibiae, tar-
sal claw simple (Fig. 15A). Forewing length 9.7 mm (N





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Dps












DVVe












C

Dps










D




Figure 18. A. Phallus, Virbia inversia left, lateral view. Dps = phallic sclerite (triangular, blunt), Ve = vesica (upper
medial-proximal projection on left side), Cp = patch of comuti (on venter); B. Phallus, Virbia endophaea left, lateral
view. Dps = phallic sclerite (triangular, sharp), PCm = peglike patches of comuti (under phallic sclerite), PI =
basiphallus (single sclerotized lobe-like projection at midpoint), Ve = vesica; C. Phallus, Virbia polyphron left, lateral
view. Dps = phallic sclerite (square, rugose), R = rugose patch (under phallic sclerite), Ve = vesica (directed
laterally), Vel = right dorsal lobe, Ve2 = left lateral lobe; D. Phallus, Virbia costata left, lateral view. CoP = coecum
penis (rounded and orientation straight), Dps = dorsal phallic sclerite (narrow and rounded), PI = basiphallus (inflected
ventrally at base), Ve = directed (ventrally, membrane rugose, left side with membranous rugose projection).






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

= 10), venation same as in male. Forewing dorsal sur-
face salmon to cinnamon with faint olive brown discal
spot present, ventral surface salmon, peach red scales
lining costal margin. Hindwing dorsal surface peach
red with natal brown discal spot, natal brown subtermi-
nal markings as spots and blotches often not reaching
posterior margin, ventral surface identical. Abdomen
with dorsum intermixed with salmon and clay, occasion-
ally with fuscous spots on all segments, lateral scales
salmon with fuscous spots on all segments, ventrally in-
termixed with salmon and clay. Male genitalia: Phallus
(Fig. 22C) with rounded coecum penis, straight,
basiphallus inflected ventrally at midpoint, phallic scler-
ite wide, nodular. Vesica directed ventrally, membrane
rugose, bilobed, elongate, second lobe sclerotized, lower
left side of vesica with small membranous projection,
right side of vesica with patch of comuti under phallic
sclerite. Genital capsule (Fig. 25A) with dorsum of
tegumen lobed, sides oftegumen divergent, conjunctiva
membranous (Fig. 23B). Uncus base cordate (Fig. 23B),
ventrad with uncus apex fingerlike, bifid and with two
fused hooks (Fig. 27A). Scaphium rounded, triangular.
Subscaphium absent. Extensions of tegumen sclero-
tized. Juxta vertically ovoid with ventral margin expand-
ing into larger orbicular plate (Fig. 20B). Costa of valva
elongate and tip entire, 20 or more costal setae. Saccu-
lus horizontally obovate, broadly hooked toward costa.
Saccus rounded, concave in center. Female genitalia
(Figs. 26C & D, 17A): Pseudopapillae anales present.
Pheromone glands membranous, branching before tip
of anterior apophyses, moderately thick, greater than 1/
8 but less than 1/4 width of 7th abdominal sternite. Os-
tium bursa slightly concave with small lateral promi-
nences (Fig. 30). Ductus bursa rectangular, narrowing
toward venter, sclerotization extending into corpus bursa
(Fig. 17A). Corpus bursa with three signa, membrane
of the corpus bursa wrinkled (Fig. 26C).
Variation. -- The most common phenotype of V.
aurantiaca is the 'red form' (Pl. 1, Nos. 3-5); however,
a yellow phenotype resembling V marginata (Pl. 3,
No. 1) also occurs infrequently. Nearly all specimens
of V aurantiaca we examined have a characteristic
round, un-notched coecum penis and a dorsally rounded
tegumen. Rarely, red form individuals have a notched
coecum penis as in V marginata or V.ferruginosa (Fig.
22A). Some males from the South, specifically Texas,
have darker raw umber forewings with a white annellar
spot and a smaller forewing length than specimens from
the northeast.
Biology.-- This species can be reared on dande-
lion and plantain (Forbes 1960, Card6 1968), but its na-
tive host is unknown. Larvae feed readily on artificial


diet (Zaspel database). The chromosome number ap-
pears to be 25 (Card6 1971). In southern Connecticut,
two broods can be reared, with one emerging in June
(peak emergence around June 20) and the other in Au-
gust (Card6 1971). Additional generations can be reared
in the southern U.S, and the species is considered facul-
tatively multivoltine (Forbes 1960). Females lay indi-
vidual eggs unattached, like V rubicundaria (Card6,
rearing notes). Females have been observed releasing
pheromone nocturnally (Fig. 8), but males are attracted
to pheromone from 10:00 to 22:00 hours (Card6 1974).
Distribution.-- This species is among the most
widely distributed in the V aurantiaca complex. In east-
ern NorthAmerica, specimens have been collected from
Manitoba and Nova Scotia, Canada, along the eastern
seaboard, south to Cordoba, Mexico (see database for
records from these localities). The western edge of its
distribution is less clear. The species has been collected
in Texas, Mississippi, Missouri, Tennessee, Louisiana,
Oklahoma, Kansas, and North and South Dakota. We
examined a single specimen from Montana that either
represents a highly disjunct population or is misidentified.

Virbia costata Stretch
P1. 1, No. 10 (female), Pl. 2, No. 1 (male)
Crocota costata Stretch 1885: 103 [type locality:
Texas, USA].
Crocota opelloides Graef 1887: 42 [type locality:
Texas, USA].
Crocota intermedia Graef 1887: 42 [type locality:
Texas, USA].
Crocota parvula Neumogen & Dyar 1893b: 140
[type locality: western Colorado, USA].
Eubaphe cocciniceps Schaus 1901: 269 [type lo-
cality: "USA"].
Eubaphe pallipennis Barnes & McDonnough
1918: (1) 85, pl. 14, Fig. 14 [type locality: "USA"].
Material examined.-- Type material: Holotype
of V costata not examined. Holotype C. intermedia:
Crocota intermedia Texas, E.L. Graef Collector, 41D1/
/Collection Brkyln. Mus.//Type No. 34162 U.S.N.M//
Crocota intermedia Graef, type Tex. Coll. Edw. L.
Graef, NMNH. Holotype of C. parvula: Crocota
intermedia parvula Colo. Bruce//Col. B. Neumogen//
Typicum Specimen//Type No. 33695 U.S .N.M//41D2//
Crocota intermedia v. parvula Type Female
Neumogen, NMNH. Holotype E. pallipennis: E.
costata v. pallipennis Type B & McD.//Glenwood
Spgs., Col. July 16-23//47 B3 [sic.], NMNH. All mate-
rial deposited in NMNH unless otherwise noted:
UNITED STATES: Arizona: Yavapai Co., Mayer,
23.vii.1959, R. F. Sternitzky (1 Female, JZ220). Colo-





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Figure 19. A. Male genital capsule Virbiafergusoni. S = saccus (rounded, concave in center), Sa = sacculus (peanut
shaped), SbSc = subscaphium (completely surrounding anal tube); B. Phallus, Virbiafergusoni, left, lateral view.
CoP = coecum penis (rounded), Cp = cornutal patch, D = dorsal, Dps = dorsal phallic sclerite (reduced), PI =
basiphallus (inflected ventrally at base), Ve = vesica, bilobedd, directed ventrally with rugose membrane, right side
with patch of comuti under phallic sclerite), V = ventral; C. Male genital capsule Virbia opella. SbSc = subscaphium
(lobed and incomplete, e.g., not completely surrounding anal tube), S = saccus (rounded, concave midventrally); D.
Phallus, Virbia opella, left, lateral view. D = dorsal, Dps = phallic sclerite (reduced), Ve = vesica (directed ventrally,
membrane rugose, bilobed, elongate, second lobe rugose), LCn = left side of vesica (with ventral patch of comuti),
RCn = right side of vesica (with patch of comuti under phallic sclerite), V = ventral.





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


DAMDM




VM
A


C


Figure 20. A. Juxta, Virbia opella: vertically ovoid with
ventral margin (VM) expanding into large orbicular shape,
dorsal margin (DM) concave with pointed edges; B.
Juxta, Virbia (H.) aurantiaca: vertically ovoid, concave
on both sides with ventral margin expanding into large
orbicular shape; C. Juxta, Virbiaferruginosa: with trans-
parent patches (P) present, (broadly rounded).


rado: Glenwood Springs, 16-23.vii., compared with type
collection in Brooklyn Museum (1 Female, JZ60); 24-
30.vii., E. costata, pallipennis paratype (1 Male, JZ65);
La Plata Co., Durango, Elev. 6, 500 ft., 2.vii. 1957, F. &
P. Rindge, AMNH (1 Male, JZ273). Oklahoma:
Comanche Co., Wichita Mts. Nat. W. Refuge, The Nar-
rows, 10-18.vii.1984, D. & M. Davis (1 Male, JZ88).
Texas: Alpine, 8-14.viii.1928, Poling collector, Barnes
Collection (1 Male, JZ89); Brewster Co., Big Bend Nat'l
Park, Green Gulch, 6.v.1972, A. & M. E. Blanchard,
AMNH (1 Male, JZ256). New Mexico: Eddy Co.,
Big Canyon, Guadalupe Mts., 24.vii.1959, A. B. Klots,
Holomelina intermedia i. intermedia det. by Graef,


AMNH (1 Female, JZ257).
Diagnosis.-- This species is sexually dimorphic
(Plate 1, No. 10; Plate 2, No. 1). Females superficially
resemble V laeta females and males, but V costata is
restricted to southwestern U.S., whereas V laeta oc-
curs in the north central and southern U.S. Virbia
costata females are larger (wing length 13.5 mm) than
males of V. laeta (wing length 11 mm), and smaller than
females of V laeta (wing length 17 mm).
Males of V costata can be confused with males
of V fragilis, a western species, but the two can be
separated by male genitalic characters. The genitalia of
V costata have a large subscaphium and a phallic scler-
ite that is narrow and rounded (Fig. 18D), whereas V
fragilis lacks a subscaphium and has a highly reduced
phallic sclerite (not shown).
Description. Male habitus (Pl. 2, No. 1): Head flesh
ocher throughout. Antennal scales tawny olive. Third
segment of labial palpus short; first and second segments
flesh ocher, third segment dark tawny olive. Thorax
dorsal and ventral scales tawny olive throughout. Legs
clay, tarsal claw simple (Fig. 15A). Forewing length
11.7 mm (N= 10), R3 absent, M3 present, extending to







Db










S


Figure 21. Corpus bursa, Virbia ferruginosa, dorsal
view. B = bursa connatee and both bursae arising from
ductus bursa), Db = ductus bursa, L = lobe of second
bursa, S = signum (three signa present).








wing margin (Fig. 13), dorsal surface pale pinkish buff
suffused with light salmon, ventral surface clay dusted
with orange rufous. Hindwing dorsal surface flesh ocher
with edges fringed with pale pinkish buff scales, ventral
surface yellow ocher dusted with flesh ocher. Abdo-
men with dorsal, lateral, and ventral scales flesh ocher.
Sclerotization of 8th sternite rectangular, length greater
than width (Fig. 28E). Coremata reduced to small
patches of scales on membranous lobes. Female habi-
tus (Pl. 1, No. 10): Head clay, occiput ranging from
salmon to peach red among specimens. Antennal scales
clay. Third segment of labial palpus exceeding front of
head by at least half of entire length, first and second
segments salmon, third segment clay. Thorax dorsal
and ventral scales clay with tufts of salmon on ventral
surface near wing base. Legs tawny olive with light
salmon scales on femora and tibiae, tarsal claw simple
(Fig. 15A). Forewing length 13.5 mm (N = 10), vena-
tion as in male. Forewing dorsal surface solid clay to
solid olive brown with salmon along costal margin, ta-
pering toward wing tip, ventral side salmon to peach red
with pattern beginning at proximal portion of adbasal
region, expanding transversely to most distal portion of
postmedial region, pattern meeting inner margin, inner
margin with clay to fuscous scales extending from post-
medial region to wing apex, occasionally slightly sinu-
ate. Hindwing dorsal surface salmon, pattern beginning
at proximal portion of adbasal region, pattern expanding
transversely to most distal portion of postmedial region,
extending to margin of clay to fuscous scales, inner
margin extending from postmedial region to wing apex,
occasionally slightly sinuate, in some specimens this post-
medial region is with athin, broken subterminal line, ven-
tral surface identical. Abdomen with dorsal scales
fuscous, lateral scales on first three segments salmon,
lateral abdominal scales tapering toward posterior in
fuscous scales, ventral side fuscous throughout. Male
genitalia: Phallus (Fig. 18D) with rounded coecum pe-
nis, straight; basiphallus inflected ventrally at base, curved
toward venter, phallic sclerite narrow and rounded.
Vesica directed ventrally, membrane rugose, bilobed,
elongate, second lobe membranous, left side of vesica
with a rugose, lobed projection and cornutuslike patch
located ventrally, right side ofvesica with rugose patches
under phallic sclerite. Genital capsule dorsally with
tegumen m-shaped, conjunctiva weakly sclerotized (Fig.
23A). Uncus base cordate (Fig. 23B). Uncus apex
triangular and with two distinct hooks. Scaphium
rounded, triangular. Subscaphium completely surround-
ing anal tube (Fig. 25A), extensions of tegumen sclero-
tized. Juxta ovoid with ventral margin expanding into
larger orbicular shape, dorsal margin concave with


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

pointed edges (Fig. 20A). Costa ofvalva elongate and
tip entire, 20 or more costal setae. Sacculus clawlike,
extending under juxta on both sides. Saccus rounded,
concave in center. Female genitalia: Opening of ovi-
positor glandular, lacking pseudopapillae anales. Phero-
mone glands membranous and unbranched, moderately
wide, greater than 1/8 but less than 1/4 width of 7th ab-
dominal sternite. Ostium bursa slightly concave with
small lateral prominences (Fig. 30B). Ductus bursa rect-
angular, narrowing toward venter, sclerotization extend-
ing into corpus bursa (Figs. 17A, 26C). Corpus bursa
double, connate, with three signa, both bursae round with
wrinkled membrane (Fig. 26C).
Variation.- There is almost no variation in size or
color among males or females.
Biology. Virbia costata has been reared on plan-
tain (Zaspel Database), but its native host is not known.
A single flight period occurs in July in Arizona, New
Mexico, Colorado and Oklahoma. Two flight periods
occur in the Big Bend region (Texas): the first in June
and the second in early August (Zaspel database).
Distribution. This species occurs in the western
United States; the eastern edge of its range is western
Oklahoma; the northernmost record is from Colorado.

Virbiafergusoni Zaspel n. sp
P1. 3 Nos. 4 (female), 5 (male)
Holotype.-- GA: Clinch Co., 7 mi. NE Fargo,
13.vii.1981, D. C. Ferguson//reared on Taraxacum//
Holomelina MEM n. p. det. D. Ferguson//JMZ298,
(Male), NMNH.
Paratypes.-- UNITED STATES: Alabama:
Baldwin Co., Bon Secour, NWR 300 15'46"N, 870 45'
27"W, 12.v.1994, R. E. Seymour//in oak-pine forest//
William Cross Expedition//Holo. n. sp. det. D. C.
Ferguson, MESSU:MEM, (1 Female, JMZ299). Florida:
St. Petersburg, Fla.//Bames Collection, NMNH, (1 Fe-
male, 2 Males); Fla. Collection Wm. Schaus, NMNH,
(1 Male); Putnam Co., Univ. of Florida Preserve, Welaka,
22.iii. 1987//D.C. Ferguson Collector, NMNH, (1 Male);
Cassia Lake Co. Fla., 8.v.1964 Taylor//Collection of
Bryant Mather//16305//genitalic preparation 1865-112
Ring Carde, NMNH (1 Male); Highlands Co., Highlands
Hammock S. P., 12.vi.1987, T. L. Schiefer//Holomelina
n. sp. det. D. C. Ferguson 1994, MESSU:MEM, (1 Fe-
male, JMZ312); Highlands Co., Highlands Hammock S.
P., 12.vi.1987, T. L. Schiefer, MESSU:MEM, (1 Male);
Highlands Co., Highlands Hammock S. P., 12.6.1987, T.
L. Schiefer, MESSU:MEM, (1 Male); 15 mi. W. of Co-
coa Beach, 19.iii.1966, R. H. Leuschner// LACM, (1
Male, JMZ313); 15 mi. W. of Cocoa Beach, 22.iii. 1966,
R. H. Leuschner//Holomelina wyatti Carde det. Ring





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

T. Card6, LACM, (1 Male); 15 mi. W. of Cocoa Beach,
31.iii.1966, R. H. Leuschner//Holomelina wyatti Card6
det. Ring T. Card6, LACM, (1 Male); 15 mi. W. of Co-
coa Beach, 22.iii.1966, R. H. Leuschner//LACM, (IMale,
JMZ214); 25 mi. W. of Cocoa Beach, 19.iii.1966, R. H.
Leuschner//LACM, (1 Male, JMZ213); 15 mi. W. of
Cocoa Beach, 19.iii.1966, R. H. Leuschner//Holomelina
wyatti Card6 det. Ring T. Card6, LACM (1 Male); 25
mi. W. of Cocoa Beach, 6.v.1966, R. H. Leuschner,
LACM, (1 Male); Alachua Co., Gainesville, 15.v.1993,
Jeff Slotten Collector, LACM (1 Male, JMZ215);
Alachua Co., Gainesville, 7.v.1993, Jeff Slotten Collec-
tor, LACM, (1 Male); Alachua Co., Gainesville,
15.ix.1995, Jeff Slotten Collector, LACM (1 Male);
Alachua Co., Gainesville, 5.iv. 1993, Jeff Slotten Collec-
tor, 1 Male (LACM); Alachua Co., Gainesville, 3.iv.1993,
Jeff Slotten Collector, LACM (1 Male); Alachua Co.,
Gainesville, 15.v.1993, Jeff Slotten Collector, LACM (1
Male); Alachua Co., Gainesville, 3.v. 1993, Jeff Slotten
Collector, LACM (1 Male); Alachua Co., Gainesville,
5.iv. 1993, Jeff Slotten Collector, LACM (1 Male); Florida,
Orange Co. Tosohatchee St. Preserve (off Rt. 520 Tay-
lor CrRd), 12.v.2001, Jeff Slotten Collector; Deland, Fla.
16.iii.1953, H. Ramstadt, CNHM 1957 A.K. Wyatt Lepi-
doptera Colln.//male genitalia on slide #938 M.O. Glenn
12.xi. 1962, FMNH (1 Male); Lakeland Fla. 21.vii. 1941,
McElhose Chicago N.H. Mus. (Al McElhose Collec-
tion)//male genitalia on slide #53 M.O. Glenn 24.v. 1964.
Georgia: Charlton Co., Okefenokee Natl. Wildlife Ref-
uge, 19.iv. 1991, D.C. Ferguson//Boardwalk Trail Chesser
Island, NMNH (1 Female); Charlton Co., Okefenokee
Natl. Wildlife Refuge, 1 .x. 1992, D.C. Ferguson//Board-
walk Trail Chesser Island Homestead, NMNH (1 Male);
Charlton Co., Okefenokee Refuge Hdqtrs., near Folkston
4.v.1981, D.C. Ferguson, NMNH (1 Male JMZ64);
Charlton Co., Trader's Hill, Folkston, 27.ix.1992, D.C.
Ferguson, NMNH (1 Female, JMZ59); Clinch Co., 7
mi. NE Fargo, 7.v. 1981, D.C. Ferguson//reared ex ovo
on Taraxacum, NMNH (1 Male, JMZ210); Clinch Co.,
7 mi. NE Fargo, 14.vii.1981, D. C. Ferguson//reared ex
ovo on Taraxacum, NMNH (1 Male, JMZ211); Clinch
Co., 7 mi. NE Fargo, 5.v.1981, D. C. Ferguson, NMNH
(1 Male, JMZ212); Clinch Co., 7 mi. NE Fargo,
13.vii.1984, D.C. Ferguson, NMNH, (1 Male); Clinch
Co., 7 mi. NE Fargo, 13.vii. 1981, D. C. Ferguson//reared
on Taraxacum//Holomelina MEM n. p. det. D. Ferguson
1994, MESSU:MEM (1 Male). Mississippi: Harrison
Co., Long Beach 10.iv.1996, R. Kergosien,
MESSU:MEM (1 Male). All material is from the NMNH
collection unless otherwise noted: UNITED STATES:
Florida: Putnam Co., Welaka, University Reserve,
17.iv.1973, D. C. Ferguson (1 Male, JMZ86). Geor-


gia: Atlanta, 17.v. 1946, P. W. Fattig (1 Male, JMZ127);
Clinch Co., 7 mi. NE Fargo, 5.v. 1981, D. C. Ferguson//
parent of brood (1 Female, JMZ185), Clinch Co., 7 mi.
NE Fargo, 13.vii.1981, D. C. Ferguson//reared ex ovo
on Taraxacum (1 Male, JMZ298). South Carolina:
McClellanville, Wedge Plantation, 24.iv.1974, D. C.
Ferguson (1 Male, JMZ85).
Etymology.-- This species is named in honor of
Dr. Douglas Ferguson. His reared series have been in-
valuable in establishing phenotypic diversity within and
among species. Dr. Ferguson encouraged JMZ's inter-
est in the revision of Holomelina and Virbia. His assis-
tance and informed discussion of these moths is grate-
fully acknowledged.
Diagnosis.-- The coloration is distinctive. This
species is bright, geranium pink with varying amounts of
dark brown scales on the hindwing adbasal and subter-
minal areas (Pl. 3, Nos. 4,5). This species can be con-
fused with lighter V opella males, but V.fergusoni has
much darker forewings and more pronounced contrast
in the hindwings. Paler forms of V. fergusoni lack the
discal spot in the hindwing that occurs in V opella. Two
male and one female genitalic character can be used to
separate this species from V opella. The vesica of V
fergusoni lacks a comutuslike patch located ventrally
on the left side of the vesica (Fig. 19B) that occurs in V
opella (Fig. 19D). The subscaphium is not lobed in V
fergusoni (Fig. 19A) and is lobed in V opella (Fig. 19C).
Females of V.fergusoni have a signum, while V opella
females lack signa (Fig. 17C). This species may be
confused with the much smaller V rubicundaria, but it
can be separated from the latter as determined in the
diagnosis of V rubricundaria.
Description.-- Male habitus (Plate 3, No. 5):
Head raw umber, occiput peach red. Antennal scales
grayish horn color. Third segment of labial palpus ex-
ceeding front of head by at least half of entire length,
first and second segments peach red, third segment raw
umber. Thorax dorsal and ventral scales raw umber.
Legs cinnamon, with a mixture of peach red and clay on
femora and tibiae, tarsal claw slightly bifid (Fig. 15B).
Forewing length 11 mm (N = 10), R3 absent (Fig. 13),
M3 present, extending to wing margin. Dorsal surface
raw umber with dark fuscous on crossveins of discal
spot forming a small transverse band, ventral surface
geranium pink, terminal margins a fringe of raw umber.
Hindwing dorsal surface peach red, subterminal region
raw umber, occasionally meeting postmedial region in
sinuate pattern, most specimens with raw umber scales
shading in region from subterminal band along fold from
discal cell to 3A, fuscous discal spot present or absent,
ventral surface identical. Abdomen with dorsum gera-





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


nium pink with patches of fuscous on most segments,
lateral and ventral scales buff throughout. Sclerotiza-
tion of 8th stemite rectangular, length greater than width
(Fig. 28E). Coremata absent. Female habitus (Pl. 3,
No. 4): Head cinnamon, occiput salmon. Antennal scales
fuscous. Third segment of labial palpus short, color same
as in male. Thorax dorsal and ventral scales cinnamon.
Legs raw umber with patches of geranium pink on
femora, tarsal claw slightly bifid (Fig. 15B). Forewing
length 12 mm (N = 10), venation as in male. Forewing
dorsal surface cinnamon, occasionally with salmon cos-
tal margin, ventral surface peach red throughout.
Hindwing dorsal surface peach red, terminal margins
fringed with tawny scales, ventral surface identical.
Abdomen with color same as in male. Male genitalia:
Phallus (Fig. 19B) with straight rounded coecum penis,
basiphallus inflected ventrally at base, curved toward
venter, phallic sclerite reduced. Vesica directed ven-
trally when everted, membrane rugose, bilobed, elongate,
second lobe of vesica rugose, right side of vesica with
patch of comuti under phallic sclerite (Fig. 19B). Geni-
tal capsule (Fig. 19A) dorsally with tegumen entire, con-
junctiva sclerotized (Fig. 23C). Uncus base cordate (Fig.
23B), ventrad with uncus apex fingerlike, bifid, with two
fused hooks (Fig. 27A). Scaphium conical, pointed.
Subscaphium completely surrounding anal tube (Fig.
19A), extensions of tegumen sclerotized. Juxta ovoid
with ventral margin expanding into larger orbicular shape,
dorsal margin concave with pointed edges (Fig. 20A).
Costa of valva elongate and tip forked; 20 or more cos-
tal setae (Fig. 19A). Sacculus peanut-shaped (Fig. 19A).
Saccus rounded, concave in center. Female genitalia:
Pseudopapillae anales present (Fig. 24B). Pheromone
glands membranous, branching after tip of anterior apo-
physes, moderately thick, greater than 1/8 but less than
1/4 width of 7th abdominal sternite. Ostium bursa
straight, without lateral prominences (Fig. 17B). Duc-
tus bursa hour-glass shaped (Fig. 17B), sclerotization ex-
tending form ductus bursa into corpus. Corpus bursa
bilobed, typically with one signum, membrane smooth
(Fig. 21).
Variation.-- Some male specimens of this spe-
cies are significantly lighter in color with a raw umber
subterminus extending toward the wing base, forming a
blotchy line where the vannus meets the remigium. The
male genitalia of some individuals had costa apices that
were entire (Fig. 19A) rather than bifed. No color varia-
tion in females was observed.
Biology.- Virbia fergusoni was reared on
dendelion (Taraxacum spp.) by Ferguson. Its native
host is not known. This species appears to have mul-
tiple broods with adult flight periods from mid-March


Cp







A

Dps Vel


_,M-ve2




B


Vel


Ve2


Figure 22. A. Phallus, Virbiaferruginosa, left, lateral
view. CoP = coecum penis (slightly knotched, orienta-
tion straight), PI = basiphallus (inflected ventrally at mid-
point), Dps = phallic sclerite (lateral view, wide and nodu-
lar), Ve = vesica (directed ventrally, membrane bilobed,
lobe membranous), Cp = patch of comuti (under phallic
sclerite); B. Phallus, T, i,, ostenta left, lateral view.
Dps = phallic sclerite (narrow and rounded), Ve = vesica
(directed ventrally, membrane smooth, entire), Cp =
patch of comuti (located ventrally), Ve1 = right dorsal
lobe, Ve2 = left lateral lobe; C. Phallus, Virbia
aurantiaca left, lateral view. CoP = coecum penis
(rounded, orientation straight), PI = basiphallus (inflected
ventrally at midpoint), Dps = phallic sclerite (wide and
nodular), Ve = vesica (directed ventrally, with membra-
nous projection = Pr), Cp = patch of comuti (under phal-
lic sclerite), Ve 1 = right dorsal lobe, Ve2 = left lateral
lobe.





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

through mid-July. A single specimen was taken in Sep-
tember (Georgia). One label indicates the habitat was a
mixed oak-pine forest.
Distribution. -- This species is restricted to the
southeastern United States; the most northern specimen
was collected in McClellanville, South Carolina, from
Georgia and northern Florida west to Baldwin Co., Ala-
bama.

Virbia ferruginosa Walker
P1. 1, Nos. 6 (female), 7(male)
Crocotaferruginosa Walker 1854: 535 [type lo-
cality, St, Martin's Falls, Albany River, Hudson's Bay,
Ontario, Canada].
Crocota quinaria Grote 1863:30 [type locality,
"Canada"].
Crocota trimaculosa Reakirt 1864: 371 [type lo-
cality, Philadelphia, Pennsylvania, USA].
Holomelina buchholzi Wyatt 1963: 100-102 [type
locality, Suffolk, Virginia, USA].
Material examined. Type material: Syntypes
of V. ferruginosa: Hudson Bay, G Barnston 11.1917//
Arctiidae slide #907//photographed BM negative 43750,
Hudson Bay, G. Bamston 11.1917, also "caterpillar with
fine long brown hairs"//Arctiidae slide #43757//BM nega-
tive 43757; Holomelina ferruginosa data: label 284
Hudson Bay G. Barnston 11.1917, "Crocota
ferruginosa", NHM. Holotype (synonym of V.
ferruginosa): VA: Suffolk, 4.viii. 1941, Otto Buchholz,
C.N.H.M. Type Male Holomelina aurantiaca Wyatt,
Buchholz Collection, ANMH. Paratypes of V.
ferruginosa: VA: Suffolk, 19.vii.1941, Otto Bucholz,
Reared Series//CNHM Paratype//Holomelina, AMNH
(1 Female, 1 Male, JMZ279; JMZ278); Suffolk, Otto
Bucholz, 24.vii.1941//CNHM Paratype//Holomelina,
AMNH (1 Female, JMZ280). All material is from the
NMNH collection unless otherwise noted: CANADA:
British Columbia: Vancouver, 3.vii. 1921, T. A. Moilliet/
/Blackmore Collection (1 Male, JMZ57). Quebec: Lac
ailaTortue, 17.vii.1947, Eugene Munro (1 Male, JMZ90);
Temiscouata Co., 30.vi. 1985, leg. George Hensel (1 Male,
JMZ94). UNITED STATES: Connecticut: New
Haven Co., Guilford, 24.vii.1966, leg. Ring T. Carde,
RTC, (1 Male, JMZ315). Illinois: Edgebrook, 9.x.1947
A. K. Wyatt (1 Female, JMZ162); 11.x.1947, A. K.
Wyatt, Bred (1 Male, JMZ161). Indiana: Tremont,
27.vii.1952, A. K. Wyatt, ex ova (1 Female, JMZ70);
Tremont, 3.vii.1944, A. K. Wyatt (1 Female, JMZ316);
9.vii. 1954,A. K. Wyatt//CNHM 1957 A. K. Wyatt Lepi-
doptera Collection, FMNH (1 Male, JMZ314). Minne-
sota: Olmstead Co. (1 Male, JMZ121). Virginia: Mont-
gomery Co. (1 Female, JMZ195). Wisconsin: Oneida


Co., Lake Katherine, 10.vii.1960, M. M. Bower (1 Male,
JMZ91).
Diagnosis.-- ,, ,i ferruginosa can be confused
with V aurantiaca, V. fragilis, and V immaculate. It
can be separated from V aurantiaca based on colora-
tion and size as described in the diagnosis for V
aurantiaca. This species usually can be separated from
V.fragilis by locality. Virbiafragilis is a western spe-
cies found in the Rocky Mountain region, whereas V
ferruginosa typically occurs in the boreal north and
eastern U.S. deciduous forests. The following male
genitalic features can also be used to distinguish the two
species: males of V. ferruginosa have a bilobed vesica
and a knotched coecum penis (Fig. 22A), whereas those
of V. fragilis have a single-lobed vesica and a rounded
coecum penis (Fig. 22B). Females of V ferruginosa
have a connate, double bursae (Fig. 17C), whereas those
of V.fragilis lack an additional bursa (Fig. 17A).
In the Eastern U.S., the geographic distribution of
Vferruginosa overlaps that of V immaculate, but flight
period is a reliable means of separating the two. The
flight period of V immaculate extends from late spring
to early summer but rarely July, and its second brood
emerges in August; V ferruginosa's flight period is in
July. Abdominal coloration often can be used to dis-
criminate the two. The abdominal tergites of V
ferruginosa often have a central cinnamon brown spot,
whereas V immaculate typically does not. In V
ferruginosa males, the second lobe of the vesica bears
a rugose or sclerotized patch (Fig. 22A) whereas in V
immaculate the second lobe is sclerotized and bears
comuti (Fig. 22C). Unlike V ferruginosa, females of
V immaculate lack an additional bursa.
Description. --Male habitus (Pl. 1, No. 7): Head
cinnamon brown. Antennal scales cinnamon brown.
Third segment of labial palpus exceeding front of head
by at least half of entire length, all segments salmon.
Thorax dorsal and ventral scales cinnamon brown
throughout. Legs clay throughout, tarsal claw simple
(Fig. 15A). Forewing length 13.4 mm (N = 10), R'-R5
all present, M3 present, extending to wing margin (Fig.
12). Dorsal surface cinnamon brown, with a faint olive
brown discal spot present, ventral surface salmon with
peach red scales lining costal margin. Hindwing dorsal
surface pinkish warm buff, cinnamon brown subtermi-
nal markings as spots and blotches often not reaching
posterior margin, exterior posterior margin occasionally
fringed with cinnamon brown scales, discal spot cinna-
mon brown, ventral surfaces salmon. Abdomen light
buff, with first segment on dorsal side a tuft of cinna-
mon brown, tergites often with a medium size cinnamon
brown spot in center, size tapering with each segment







towards posterior. Sclerotization of 8th sternite pen-
tagonal (Fig. 28C). Coremata reduced to small patches
of scales on membranous lobes. Female habitus (Pl. 1,
No. 6): Head clay. Antennal scales clay. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length, all segments clay. Thorax clay.
Legs cinnamon brown with patches of scales on femora
and tibiae from salmon to buff, tarsal claw simple (Fig.
15A). Forewing length 15 mm (N = 10), venation same
as in male. Dorsal surface cinnamon extending from
wingbase through postmedial region, subterminal region
clay color, with a faint natal brown discal spot present,
although rare, some specimens with small white spot or
several large spots in annelar region, ventral surface
dark salmon suffused with peach red scales. Hindwing
dorsal surface peach red, with a natal brown discal spot,
natal brown subterminal markings as spots and blotches
often not reaching posterior margin, ventral surface iden-
tical. Abdomen with dorsum intermixed salmon and clay,
occasionally with fuscous spots on all segments, lateral
sides salmon color with fuscous spots on all segments,
ventral sides a blend of salmon and clay. Male genita-
lia: Phallus (Fig. 22A) with coecum penis weakly notched,
straight, basiphallus inflected ventrally at midpoint, curved
toward venter, phallic wide, nodular. Vesica directed
ventrally when everted, membrane rugose, bilobed, elon-
gate, second lobe membranous; right side of vesica with
patch of cormuti under phallic sclerite. Genital capsule
with dorsum oftegumen lobed, sides oftegumen diver-
gent (Fig. 23B), conjunctiva strongly sclerotized. Un-
cus base cordate (Fig. 23B), ventrad with uncus apex
fingerlike, bifid, with two fused hooks (Fig. 27A).
Scaphium rounded, triangular. Subscaphium absent,
extensions of tegumen sclerotized. Juxta rectangular,
tapering in center, expanding toward ventral base with
backward directed barbs on dorsal margin, broadly
rounded ornamentation present (Fig. 20C). Costa of
valva elongate and apex entire, 20 or more costal setae.
Sacculus horizontally obovate, broadly hooked toward
costa. Saccus rounded, concave in center. Female geni-
talia (Fig. 21): Pseudopapillae anales present (Fig. 24B).
Pheromone glands membranous, branching after tip of
anterior apophyses, moderately thick, greater than 1/8
but less than 1/4 width of 7th abdominal sternite. Os-
tium bursa slightly concave with small lateral promi-
nences (Fig. 30B). Ductus bursa rectangular, narrow-
ing toward venter, sclerotization extending into corpus
bursa (Fig. 21). Corpus bursa double, connate, with
one signum, both bursae round with wrinkled membrane
(Fig. 21).
Variation.-- In some males, the apex of the third
labial palp segment is cinnamon brown or has a patch of


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

cinnamon brown scales. In others, the cinnamon brown
subterminal markings form a large longitudinal band ex-
tending through the posterior margin.
Some males from reared series were curated un-
der the name V buchholzi (AMNH and FMNH).
These males have bright yellow ocher scales through-
out. Card6 (1968) proposed that V buchholzi was a
color variant of V. ferruginosa. Franclemont (1983)
placed it as a junior synonym of V. ferruginosa. We
have examined several V 'bucholzi' specimens and
agree that it is a color variant of V. ferruginosa.
Biology.-- Virbiaferruginosa is univoltine with
adults emerging during July. Larvae have been reared
on dandelion (Taraxacum spp.; Card6, Rearing Notes).
Females release pheromone (exhibit calling behavior)
at night (Fig. 8D).
Distribution.- This species occurs in Canada from
Nova Scotia to British Columbia (Zaspel database). In
the United States, it is sympatric with V aurantiaca in
the Northeast and upper Midwest. Its range extends
south to Virginia, Mississippi, Missouri, and Lousiana.
A few specimens superficially resembling V
ferruginosa have been collected in Colorado; however,
it is likely that these specimens are V. fragilis.

Virbia fragilis Strecker
P1. 2, Nos. 6 (female), 7 (male)
Crocotafragilis Strecker 1878: app: 1859 [type
locality, Pagosa Springs, Colorado, USA].
Material examined.-- Type material: Holotype of
V. fragilis: Crocota fragilis Streck. Pagosa Springs
Col. Orig. Type McCanley//Crocota fragilis Streck.
Colorado, Pagosa Springs McCanley Orig. Type Strecker
Colln. 27530 Field Museum Nat. Hist.//Holotype det.
A. Watson 1986//Genitalic preparation 201 Ring Card6
Genitalic preparation 201 Ring Card6//Lepidoptera type
photograph No. 184 Field Museum, FMNH. All mate-
rial is from the NMNH collection unless otherwise noted:
CANADA: Alberta: Lloydminster, 5.vii.1942, P. F.
Bruggeman (1 Male, JMZ55). British Columbia: Ar-
rowhead Lake//Bames Collection (1 Male, JMZ226).
UNITED STATES: Colorado: Denver, Chimney
Gulch, 16-23.ix (1 Female, JMZ73). New Mexico:
Las Vegas (1 Male, JMZ56). South Dakota:
Pennington Co., Spring Creek Camp, 11 mi. NE Hill
City, 31.vii.1955, F. & P. Rindge, AMNH (1 Male,
JMZ274); Slate Creek, Black Hills, 9 mi. NW of Hill
City, 6000', 20.vii.1964, D. C. Ferguson (1 Female,
JMZ72); 9 mi. NW of Hill City, 6000' July 15, 1964 D.
C. Ferguson (1 Male, JMZ193). Utah: Ephraim Sanpete
Co., Major's Flat, 10.vii. 1981, D. C. Ferguson (1 Male,
JMZ192).





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

Diagnosis. Virbia fragilis is easily confused
with V immaculate, V costata males, and it is some-
times confused with V. ferruginosa. Characters sepa-
rating V. fragilis and V ferruginosa are given above.
Virbia fragilis and V immaculate are geo-
graphically isolated from each other. Virbiafragilis oc-
curs in western Canada and the western and southwest-
ern U.S., whereas V immaculate occurs in north cen-
tral and eastern U.S. Males of V fragilis tend to be
larger (wing length 14 mm) than those of V immaculate
(wing length 12 mm). Male genitalia further distinguish
them: the vesica of V. fragilis has a single lobe (Fig.
22B), whereas the vesica of V immaculate is bilobed
with the second lobe sclerotized.
Males of V fragilis can be separated from V
costata based on genitalia. In V fragilis, the phallic
sclerite is reduced (Fig. 22B), the uncus apex is finger-
like and bifid with two hooks (Fig. 27A). In V costata,
the phallic sclerite is narrow and rounded (Fig. 18D),
the uncus apex is triangular with two hooks (Fig. 23D).
In females, V fragilis possess pseudopapillae anales
(Fig. 24B), and lack an additional bursa (Figs. 24A, 26C),
whereas females of V costata lack pseudopapillae
anales and have a connate, double bursa (Fig. 17D).
Description. --Male habitus (Plate 2, No. 7):
Head clay throughout. Antennal scales clay. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length; all segments clay. Thorax dorsal
and ventral scales clay throughout. Legs clay, with
patches of flesh ocher scales on femora and tibiae; tar-
sal claw simple (Fig. 15A). Forewing length 14 mm (N
10), R'-R5 all present, M3 present, extending to wing
margin (Fig. 12). Dorsal surface clay with cinnamon
scales from wingbase through postmedial region, ven-
tral surface flesh ocher. Hindwing dorsal surface flesh
ocher throughout, ventral surface identical. Abdomen
with dorsal, lateral and ventral scales warm buff. Scle-
rotization of 8th sternite rectangular, length greater than
width (Fig. 28E). Coremata reduced to small patches
of scales on membranous lobes. Female habitus (Plate
2, No. 6): Head color same as in male. Antennal color
same as in male. Third segment of labial palpus ex-
ceeding front of head by at least half of entire length,
color same as in male. Thorax color same as in male.
Leg color same as in male, tarsal claw simple (Fig. 15A).
Forewing length 12 mm (N = 2), venation as in male.
Dorsal surface cinnamon extending from wingbase
through postmedial region, costal margin peach red, ter-
minal wing margins fringed with dark salmon scales, oc-
casionally with a faint natal brown spot between medial
and postmedial regions present. Although rare, some
specimens with a small white spot in annellar region,


ventral surface dark salmon suffused with peach red
scales. Hindwing dorsal surface peach red, with a natal
brown discal spot, natal brown subterminal markings as
spots and blotches often not reaching posterior margin
of wing, ventral surface identical. Abdomen with color
same as in male, but occasionally with patches of
fuscous scales on all dorsal segments tapering towards
posterior. Male Genitialia: Phallus with rounded coecum
penis, straight, basiphallus inflected ventrally at midpoint,
curved toward venter (Fig. 25C), phallic sclerite highly
reduced (Fig. 22B). Vesica directed ventrally when
everted, membrane rugose, bilobed, second lobe sclero-
tized, left side of vesica with a small membranous pro-
jection located ventrally, right side of vesica with a patch
of comuti located under phallic sclerite (Fig. 22C). Geni-
tal capsule dorsum of tegumen concave, bent inward,
conjunctiva weakly sclerotized (Fig. 23A). Uncus base
cordate (Fig. 23B), ventrad with uncus apex fingerlike,
bifid, and with two hooks (Fig. 27A. Scaphium rounded,
triangular. Subscaphium absent, extensions oftegumen
sclerotized. Juxta cylindrical, emarginated, with ends
barbed toward dorsum with broadly rounded ornamen-
tation present (Fig. 20C). Costa of valva elongate and
tip entire, 20 or more costal setae. Sacculus horizontally
obovate, broadly hooked toward costa. Saccus rounded,
concave in center. Female genitalia: Pseudopapillae
anales present (Fig. 24B). Pheromone glands membra-
nous, branching after tip of anterior apophyses, moder-
ately thick, greater than 1/8 but less than 1/4 width of
7th abdominal sternite. Ostium bursa slightly concave
with small lateral prominences (Fig. 30B). Ductus bursa
rectangular, narrowing toward venter, sclerotization ex-
tending into corpus bursa (Fig. 17A). Corpus bursa with
three signa, membrane wrinkled (Fig. 26C).
Variation.- One V fragilis male with white discal
spots on the forewings was found from Alberta, Canada.
Few females were available for examination, but no size
or color variation was observed.
Biology.-- Little is known about the natural his-
tory of Virbiafragilis. It appears to have a single brood,
emerging in July. Peak female pheromone release and
calling behavior occurs in scotophase (e.g., between 20
and 23 hours; Fig 8E), similar to most species in the
complex.
Distribution.-- This species has been collected in
open fields in the Black Hills, South Dakota and in Boul-
der, Colorado. The range extends north to Alberta and
British Columbia, south to New Mexico.

Virbia immaculate Reakirt
P1. 2, Nos. 2 (female), 3 (male)
Crocota immaculate Reakirt 1864: 372 [type lo-





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Cv

Dt
Dt

A B


H











C D




H
Sc






E F





Figure 23. A. Dorsal view of uncus-tegumen complex, Virbia costata. Cv = conjunctiva sclerotizationn weak), Dt =
tegumen (concave, bent inward); B. Dorsal view of uncus-tegumen complex, Virbia aurantiaca. Cv = conjunctiva
sclerotizationn strong), Dt= tegumen (rounded with three-dimensional shapes on both sides with risen node in center),
U = base of uncus (cordate); C. Tegumen, Virbia laeta. Dt = tegumen, entire); D. Uncus, Virbia costata, % view.
H = hooks of the uncus (two hooks present), U = uncus (apex an inverted heart-shape); E. Uncus complex, Virbia
laeta, 3 view. H = hooks of the uncus, Sc = scaphium (rounded, triangular); F. Uncus complex, Virbia phalangia,
ventral view. U = uncus, Sc = scaphium (flattened).





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

cality, Philadelphia, Pennsylvania, USA]
Material examined. Type material: Holotype of
V immaculate: Philadelphia, Pennsylvania//immaculata
Reakirt//C. immaculate Reakirt Coll. Reakirt//Original
Type, [sic.] FMNH. All material is from the NMNH
collection unless otherwise noted: UNITED STATES:
Illinois: Edgebrook, 10.x.1947, A. K. Wyatt, Bred//
1019a (1 Male, JMZ105); 7.x.1947, A. K. Wyatt//CNHM
1957 Alex K. Wyatt Lepidoptera Collection (1 Female,
1 Male, JMZ309; JMZ310); Chicago, 8.x.1948, A. K.
Wyatt//CNHM 1957 Alex K. Wyatt Lepidoptera Col-
lection, FMNH (1 Male, JMZ308). Indiana: Tremont,
25.viii.1952, A. K. Wyatt, ex ova (1 Male, JMZ106).
Michigan: Alpena, 10.vii.1931, Dr. G. W. Rawson//G.
W. Rawson Collection 1962 (1 Male, JMZ126). New
York: Boreas River, 8.vii.1937, L. R. Rupert (1 Male,
JMZ 104). Pennsylvania: New Brighton, 18.vi.1907//
Merrick Museum//Bames Collection (1 Male, JMZ 125);
Warrendale, 21.viii.1946, LACM (1 Male, JMZ191).
Wisconsin: Racine, 0. Meske Collection (1 Male,
JMZ107).
Diagnosis.-- Virbia immaculate can be confused
with V.fragilis, V aurantiaca, and V.ferruginosa. The
characters for separating these are given above under
the relevant species' diagnosis.
Description. Male habitus (Pl. 2, No. 3): Head
clay throughout. Antennal scales scales clay. Third
segment of labial palpus exceeding front of head by at
least half of entire length, all segments clay. Thorax
dorsal and ventral scales clay. Legs clay, with patches
of flesh ocher scales on femora and tibiae, tarsal claw
simple (Fig. 15A). Forewing length 12 mm (N = 10),
R1-R5 all present, M3 present, extending to wing margin
(Fig. 12). Dorsal surface clay with cinnamon extending
from wingbase through postmedial region, ventral sur-
face flesh ocher. Hindwing dorsal surface ocher, ven-
tral surface identical. Abdomen with dorsal, lateral, and
ventral abdominal scales warm buff. Sclerotization of
8th stemite rectangular, length greater than width (Fig.
28E). Coremata reduced to small patches of scales on
membranous lobes. Female habitus (Plate 2, No. 2):
Head color same as in male. Antennal color same as in
male. Third segment of labial palpus exceeding front of
head by at least half of entire length, color same as in
male. Thorax color same as in male. Leg color same
as in male, tarsal claw simple (Fig. 15A). Forewing
length 12 mm (N = 10), venation as in male. Dorsal
surface cinnamon extending from wingbase through post-
medial region, posterior margins fringed with salmon
scales, with a faint natal brown spot between medial
and postmedial regions present, although rare, some
specimens with a small white spot in annellar region,


ventral surface dark salmon suffused with peach red.
Hindwing dorsal surface peach red, with a natal brown
discal spot, subterminal markings as spots and blotches
not reaching posterior margin, ventral surface identical.
Abdomen with color same as in male. Male genitalia:
Phallus with rounded coecum penis, straight, basiphallus
inflected ventrally at midpoint (Fig. 22C), phallic sclerite
reduced (Fig. 22B). Vesica directed ventrally when
everted, membrane rugose, bilobed, second lobe sclero-
tized, left side with small membranous projection located
ventrally (Fig. 22C), right side of vesica with patch of
comuti under phallic sclerite. Genital capsule with dor-
sum concave, bent inward (Fig. 23A), conjunctiva scle-
rotized. Uncus base cordate (Fig. 23B). ventrad with
uncus apex fingerlike, bifid, with two hooks (Fig. 27A).
Scaphium rounded, triangular. Subscaphium absent.
Juxta cylindrical, emarginated, with ends barbed toward
dorsum, broadly rounded (Fig. 20C). Costa of valva
elongate and tip entire, 20 or more costal setae. Saccu-
lus horizontally obovate, broadly hooked toward costa.
Saccus rounded, concave in center. Female genitalia:
Pseudopapillae anales present (Fig. 24B). Pheromone
glands membranous, branching before tip of anterior
apophyses, moderately thick, greater than 1/8 but less
than 1/4 width of 7th abdominal stemrnite. Ostium bursa
slightly concave with small lateral prominences (Fig.
30B). Ductus bursa rectangular, narrowing toward ven-
ter, sclerotization extending into corpus bursa (Fig. 17A).
Corpus bursa with three signa, membrane wrinkled (Fig.
26C).
Variation. Females of V immaculate from some
localities (e.g., Indiana) are typically smaller (wing length
=11 mm) than in others (e.g., Illinois; wing length = 13
mm).
Biology.-- Virbia immaculate has been reared
on both dandelion (Taraxacum) and plantain (Plantago)
(Zaspel database). This species is bivoltine with adults
emerging in June and again in August. Calling occurs in
scotophase or the dark period of the light-dark cycle
(Fig. 8B) and is advanced by decreasing temperatures
(Carde et al. 1973). Six matings have been observed in
an open field in Dryden, New York, all within 30 minutes
of sunset (Carde 1971).
Distribution.-- This species occurs north of lati-
tude 40 degrees, from the Eastern Coast of North
America west to Manitoba. This species can also be
found in Iowa, Illinois, and Indiana.

/, !,,, laeta Guerin-Meneville
P1. 3, Nos. 6 (female), 7 (male)
Lithosia laeta Guerin-Meneville [1832] 1829-
1844: 322 [type locality: "l'Amerique duNord", USA].





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


PsPa


Pg


A B

Figure 24. A. Female genitalia, Virbia laeta, dorsal view. Ap = anterior apophysis, B = bursa (smooth), Db = ductus
bursa, Pa = papillae anales, Pg = pheromone glands, S = signum; B. Ovipositor, Virbia nigricans, dorsal view. Pg =
pheromone glands, PsPa = pseudopapillae anales, internal view.


Crocota treatii Grote 1865: 322 [type localty,
Massachusetts, USA].
Lithosia rubropicta Packard 1887: 52 [type lo-
cality, Rangley Lakes, Maine USA].
Material examined.-- Type material: Holotype
of V laeta: Ex Mus AEO Dris Boisduval//Oberthur Col-
lection//Type TF H6fer//Bd//47BI NMNH. All mate-
rial is from the NMNH unless otherwise noted:
CANADA: New Brunswick: Canequet, 11.vi.1950,
D. C. Ferguson (1 Male, JMZ78). UNITED STATES:
Arkansas: Washington Co., Devil's Den St. Park,
21.vii.1966, R. L. Brown Collector (1 Male, JMZ124).
New Hampshire: Claremont, 20.vi.1908, Collection
Brklyn Museum (1 Male, JMZ80). Florida: Highlands
Co., Archbold Biological Sta. 5.iv.1962, D. C.
Ferguson (1 Male, JMZ79). Maine: Lincoln, 8.vii (1
Male, JMZ123). Minnesota: St. Paul, U. Farm,
22.vii. 1935, A. A. Granovsky//light trap, UMSP (1 Male,
JMZ122). Missouri: Willard, 19.vi., A. E. Brower//
Eubaphe laeta treatii (1 Female, JMZ160). Texas:
Jasper Co., Town Bluff, 6.vi. 1967,A. & M. E. Blanchard
(1 Male, JMZ 159); Montgomery Co., Conroe, 12.iii. 1967,
A. & M. E. Blanchard (1 Male, JMZ77). Wisconsin:


Marathon Co., Township of Stettin, 6.vii.1978, Russell
A. Rahn (1 Male, JMZ81).
Diagnosis.-- Both sexes of this species superfi-
cially resemble V costata females; however, V laeta is
restricted to northcentral and southern U.S. and V.
costata is restricted to southwestern US. See the diag-
nosis of V costata for additional diagnostic differences.
Description.-- Male habitus (Pl. 3, No. 7): Head
fuscous, occiput geranium pink. Antennal scales fuscous.
Third segment of labial palpus short, first and second
segments geranium pink, third segment fuscous. Tho-
rax dorsal and ventral scales fuscous with small tufts of
geranium pink at wing base. Legs fuscous to dark gray-
ish brown, tarsal claw simple (Fig. 15A). Forewing
length 11 mm (N = 10), R3 absent (Fig. 13), M3 present,
extending wing margin. Dorsal surface fuscous to dark
grayish brown with light salmon band running full length
of costal margin, ventral surface dusted in light salmon.
Hindwing dorsal surface geranium pink, pattern begin-
ning at proximal portion of adbasal region, expanding
transversely to distal portion of postmedial region, pat-
tern extending down wing and meets inner margin of
fuscous to dark grayish brown scales extending from





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


- Sc


Sb


H


Dps


Figure 25. Male genitalic capsule of Virbia laeta. A- Ventral view, B- Lateral view; C- Left, lateral view of phallus.
C = costa, CoP = coecum penis, Dps = dorsal phallic sclerite, Dt = dorsal tegumen, H = hook, J = juxta, PI =
basiphallus, Sa = sacculus, S = saccus, Sb = subscaphium, Sc = scaphium, U = uncus, V = vinculum, Ve = vesica.







postmedial region to wing tip, occasionally with a slightly
sinuate pattern, ventral surface identical. Abdomen ge-
ranium pink with small dark grayish brown spots in cen-
ter of each abdominal tergite, lateral abdominal scales
geranium pink, ventral abdominal scales fuscous to dark
grayish brown. Sclerotization of 8th stemite rectangu-
lar shape, length greaterthan width (Fig. 20E). Coremata
reduced to small patches of scales on membranous lobes.
Female habitus (Pl. 3, No. 6): Head color same as in
male. Antennal color same as in male. Third segment
of labial palpus exceeding front of head by at least half
of entire length, color same as in male. Thorax color
same as in male. Leg color same as in male, tarsal claw
simple (Fig. 15A). Forewing length 17 mm (N = 10),
venation as in male. Dorsal surface color same as in
male, ventral surface color same as in male. Hindwing
dorsal surface color same as in male, ventral surface
color same as in male. Abdomen with color same as in
male. Male genitalia: Phallus (Fig. 25C) with rounded
coecum penis, inflected ventrally, basiphallus straight,
phallic sclerite absent. Vesica directed ventrally when
everted, membrane smooth, entire, left side with small
membranous projection located ventrally, right side of
vesica with patch of comuti (Fig. 22A). Genital capsule
with dorsum entire (Fig. 23C), conjunctiva sclerotized.
Uncus base cordate (Fig. 23B), ventrad with uncus apex
slightly bifurcate, toothed, with two hooks (Fig. 25A).
Scaphium rounded, triangular. Subscaphium completely
surrounding anal tube, extensions oftegumen sclerotized.
Juxta ovoid with ventral margin expanding into large or-
bicular shape (Fig. 25A). Costa of valva elongate and
tip entire, 20 or more costal setae. Sacculus blunt and
extending under juxta, clawlike. Saccus rounded, con-
cave in center. Female genitalia (Fig. 24A):
Pseudopapillae anales present (Fig. 24B). Pheromone
glands membranous, branching after tip of anterior apo-
physes, moderately thick, greater than 1/8 but less than
1/4 width of 7th abdominal sternite. Ostium bursa
straight, without prominences (Fig. 17B). Ductus bursa
rectangular, narrowing toward venter, sclerotization ex-
tending into corpus bursa (Fig. 17B). Corpus bursa with
three signa, membrane smooth (Fig. 24A).
Biology.-- Larvae have been reared on dande-
lion (Taraxacum) and plantain (Plantago) (Forbes
1960). Adults are typically found in pine woodlands
(Heppner 2003). In Louisiana, V laeta can be collected
every month except December (Brou 2003). In Texas,
two flight periods (March, June) appear to occur, but in
more northern parts of the range, collection dates sug-
gest a univoltine brood emerging in June and July.
Distribution.-- New Brunswick south to Florida,
west to Minnesota south to Texas.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

Virbia lamae Freeman
P1. 1, Nos. 8 (female), 9 (male)
Holomelina lamae Freeman 1941: 123 [type lo-
cality, Baddeck, Nova Scotia, Canada].
Material examined.- Type material: Holotype not
examined. All material is from the NMNH collection
unless otherwise noted: CANADA: Nova Scotia:
Halifax Co., Bog Prospect Road, 21.vii.1852, D. C.
Ferguson (1 Female, JMZ58); Mount Uniake, 19.vii. 1946,
D. C. Ferguson//Eubaphe lamae Freeman//Otto
Bucholz Collection, AMNH (1 Male, JMZ277).
UNITED STATES: Maine: Seawall Bog, Acadia Pk.
Mt. Desert Is., 29.vii.1966, leg. Ring Carde//Female
Parent of Brood (1 Female, 1 Male, JMZ296; JMZ295);
Acadia Pk. Mt. Desert Is, 29.vii. 1966, leg. Ring Carde/
/Female Parent of Brood, RTC (1 Female, JMZ297).
Wisconsin: Marathon Co., Township ofNorrie, Russella
(1 Male, JMZ63).
Diagnosis.-- This species can be confused with
V aurantiaca and is occasionally misidentified as V
marginata. Virbia lamae and V marginata have dis-
junct distributions; V lamae occurs in the Northeastern
U.S. and Canada, and V marginata occurs in northern
Central America with disjunct populations in the south-
western U.S.
Genitalic differences in both sexes separate V
lamae and V aurantiaca. In male V lamae, the juxta
is cylindrical and emarginated as in V marginata (Fig.
27C) whereas in V aurantiaca, the juxta is ovoid with
the ventral margin expanding into a larger, orbicular shape
(Fig. 20B). In females, V aurantiaca has pseudopapillae
anales (Fig. 24B); V lamae does not. The ostium bursa
is unornamented in V lamae (Fig. 26B); in contrast, lat-
eral prominences surround the ostium bursa in V
aurantiaca (Fig. 26D). The dark coloration in the anal
fold of the hindwing can also be used to separate V
lamae from V aurantica and V marginata.
Description. --Male habitus (Pl. 1, No. 9): Head
ranging from clay color to fawn color among specimens.
Antennal scales ranging from clay to fawn among speci-
mens. Third segment of labial palpus exceeding front of
head by at least half of entire length, all segments clay
to fawn. Thorax dorsal and ventral scales from clay to
fawn. Legs clay, tarsal claw simple (Fig. 15A). Forew-
ing length 10 mm (N = 9); R'-R5 all present, M3 present,
extending to wing margin (Fig. 13). Dorsal surface clay
with fawn extending from wingbase through postmedial
region, ventral surface salmon color extending from
wingbase through postmedial region, subterminal region
natal brown often not reaching posterior margin, scales
over veins retaining salmon scales. Hindwing dorsal
surface from yellow ocher to salmon with a natal brown





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


DO









F


Figure 26. A. Female Virbia lamae, dorsal view. Db = ductus bursa (hourglass), Pg = pheromone glands (membra-
nous, branching after tip of anterior apophysis, moderately thick); B. Female Virbia lamae, ventral view. Ob = ostium
bursa (lacking lateral prominences); C. Female Vb,.,ia aurantiaca, dorsal view. B = bursa (with wrinkled mem-
brane), Db = ductus bursa (rectangular, narrowing toward ventor, sclerotization extending into corpus bursa), Pg =
pheromone glands (membranous, branching before tip of anterior apophysis, moderately thick), PsPa = pseudopapillae
anales, internal view, S = signum (three present); D. Female V, .'1, aurantiaca, ventral view. Ob = ostium bursa
(slightly concave with small lateral prominences); E. Female Virbia opella, dorsal view. B = bursa connatee and
membrane smooth), Db = ductus bursa (goblet shape, sclerotization extending into corpus bursa), Pg = pheromone
glands (membranous, branching after tip of anterior apophysis, thin), PsPa = pseudopapillae anales present; F. Fe-
male Virbia opella, ventral view. Ob = ostium bursa (split midventrally with two broad, lateral prominences).


Pg

Do
--- DO

1A
-I

A-


k-bb


r<

B


S---B
-B


V-





Cb
E





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


A B



C



Sa

S

C


C


C
Se
Se


Sa
Sa


D E




Figure 27. A. Uncus complex, Virbia aurantiaca, ventral view. U= uncus (fingerlike, with bifed apex), H= hooks;
B. Uncus in lateral view, Virbia marginata. N = notch (rounded); C. Male genital capsule Virbia marginata. C =
costa (distal tip entire), J =juxta (cylindrical, emarginated with ends barbed toward dorsum), Sa = sacculus (horizon-
tally obovate, broadly hooked toward costa, with setae present), S = saccus (rounded, concave in center); D. Male
genital capsule, V,, ,i opella. C = costa, (distal tip forked), Sa = sacculus (peanut-shaped), Se = setae; E. Male
genital capsule, T, b'iferonia. C = costa, (distal tip entire), Sa = sacculus (thin, square-shaped), Se = setae.





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

discal spot, natal brown subterminal markings a thick
band often not reaching posterior margin, ventral sur-
face identical. Abdomen with dorsal, lateral and ventral
scales light buff with first segment on dorsal side a tuft
of fawn scales, additional tergites occasionally with a
medium size fawn spot in center, size of spot tapering
with each segment towards posterior. Coremata ab-
sent. Female habitus (Pl. 1, No. 8): Head color same as
in male. Antennal color same as in male. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length, color same as in male. Thorax
color same as in male. Legs clay, with patches of buff
scales on femora and tibiae, tarsal claw simple (Fig. 15A).
Forewing length 9.8 mm (N = 4), venation same as in
male. Dorsal clay extending from wingbase through
postmedial region, posterior margin cinnamon brown,
occasionally faint natal brown spot between between
medial and postmedial regions present, large white anellar
spot present, ventral surface clay extending from
wingbase through postmedial region, ocassionally suf-
fused with salmon scales, especially along costal mar-
gin, natal brown subterminal markings as spots and
blotches often not reaching posterior margin, posterior
margin clay with scales over veins retaining salmon
scales. Hindwing dorsal surface salmon, with a large
natal brown discal spot, natal brown subterminal mark-
ings as thick spots and blotches often not reaching pos-
terior margin, ventral surface identical. Abdomen color
same as in male. Male genitalia: Phallus with rounded
coecum penis rounded and weakly notched (Fig. 22A),
inflected ventrally, basiphallus straight, phallic sclerite
wide and nodular (Fig. 22C). Vesica directed ventrally
when everted, membrane rugose, bilobed, elongate, sec-
ond lobe sclerotized, right side of vesica with patch of
comuti under phallic sclerite (Fig. 22A). Genital cap-
sule with dorsum concave, bent inward, conjunctiva
weakly sclerotized (Fig. 23A). Uncus base cordate (Fig.
23B), ventrad with uncus apex fingerlike, bifid, with two
hooks (Fig. 27A). Scaphium rounded, triangular.
Subscaphium absent, extensions oftegumen sclerotized.
Juxta cylindrical, emarginated, with ends barbed toward
dorsum (Fig. 27C). Costa of valva elongate and tip en-
tire, 20 or more costal setae. Sacculus horizontally obo-
vate, broadly hooked toward costa. Saccus rounded,
concave in center. Female genitalia: Lacking
pseudopapillae anales. Pheromone glands membranous,
branching after tip of anterior apophyses, moderately
thick greater than 1/8 but less than 1/4 width of 7th ab-
dominal sternite. Ostium bursa without lateral promi-
nences (Figs. 17B, 26A, 26B). Ductus bursa hourglass
shaped, sclerotization extending into corpus bursa (Fig.
17B). Corpus bursa with three signa, membrane wrinkled


(Fig. 17B).
Variation.-- Of the material examined, V lamae
males rarely with white annellar spots on the forewings;
no additional variation for males or females was observed.
Biology.- Larvae have been reared on plantain
(Plantago) and dandelion (Taraxacum) (Zaspel data-
base). This diurnal species can only be collected with
nets in bogs during July and early August (Carde, Field
notes). Blacklighting in Seawall Bog, Southwest Har-
bor, Maine failed to attract any V lamae, although these
were abundant in daytime (Carde personal observation).
Distribution.- This species can be collected in
Nova Scotia, New Brunswick, Maine, Wisconsin, and
Michigan. Locally, it is restricted to open peat bogs,
usually near large bodies of water (maritime or Great
Lakes).

Virbia marginata Druce
P1. 2, No. 10 (female), Pl. 3, No. 1 (male)
Eubaphe marginata Druce 1885: 136, t. 13, f 8
[type locality, Duefias, Guatemala]
Material examined.-- Type material: Holotype of
V marginata: Type Goodman-Salvin Collection, 97 53'
Duefias Guatemala, G.C. Champion//Type sp. figured
E.C.A. Lep. Het. Eubaphe marginata Druce (NHM).
All material is from the NMNH collection unless other-
wise noted: GUATEMALA: Guatemala City, viii/
Barnes and Schaus Collection (2 Males, JMZ222;
JMZ224). MEXICO: Jalapa, Schaus//ETOwen Do-
nor (1 Female, JMZ69). UNITED STATES: Arizona:
White Mts., Barnes Collection (1 Male, JMZ221). Colo-
rado: Col. remmingtoni det. Ring T. Carde, LACM (1
Male, JMZ182); Denver, Barnes Collection (1 Male,
JMZ184). New Mexico: Jemez Springs, Barnes Col-
lection (1 Male, JMZ223); Sandoval Co., 1 mi. S. Sulfer
Springs, 7800 ft., 31.vii.1961, F., P. & M. Rindge//Col-
lected onNSF grant 69037, AMNH (1 Female, JMZ305).
Diagnosis.-- Virbia marginata (Plate 2, No. 10;
Plate 3, No. 1) has been treated as a synonym of V
rubicundaria (Plate 2, Nos. 4,5); however, these two
species have very different phenotypes. Further, the
distributions of the two species do not overlap. Virbia
marginata occurs in Mexico, Guatemala and the South-
western U.S., whereas V rubicundaria occurs in the
southeastern U.S. Both sexes are morphologically dis-
tinct. Males of V marginata have a tegumen that is
concave (Fig. 23A), whereas those of V. rubicundaria
have a lobed tegumen (Fig. 23B). Females of both spe-
cies have an ostium bursa with lateral prominences (Fig.
30), however, V marginata lack pseudopapillae anales
and typically have one signum (Fig. 17B). Females of V
rubicundaria have pseudopapillae anales and possess








three signa (Fig. 24B). Occasionally, V marginata is
confused with V lamae (Plate 1, Nos. 8,9); however, V.
lamae is restricted to the northern U.S. and Canada,
and the species are not sympatric.
Description.-- Male habitus (Plate 3, No. 1): Head
yellow ocher. Antennal scales yellow ocher. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length, all segments yellow ocher. Thorax
dorsal and ventral scales yellow ocher. Legs clay with
patches of salmon scales on femora and tibiae, tarsal
claw simple (Fig. 15A). Forewing length 10 mm (N =
10), R3 absent, M3 present, extending to edge (Fig. 13).
Dorsal surface clay, ventral surface identical, but occa-
sionally suffused with salmon scales. Hindwing dorsal
surface yellow ocher, pattern beginning at proximal por-
tion of adbasal region, expanding transversely to most
distal portion ofpostmedial region, meeting inner margin
of raw umber scales, extending from postmedial region
to wing apex, occasionally with a slightly sinuate pattern,
small raw umber discal spot present, ventral surface iden-
tical. Abdomen yellow ocher, dorsal tergites occasion-
ally with a medium size fawn color spot in center, size
tapering with each segment towards posterior. Scleroti-
zation of 8th sternite pentagonal (Fig. 28C). Coremata
reduced to small patches of scales on membranous lobes.
Female habitus (Pl. 2, No. 10): Head cinnamon. Anten-
nal scales cinnamon. Third segment of labial palpus short,
cinnamon. Thorax cinnamon. Legs yellow ocher, tarsal
claw simple (Fig. 15A). Forewing length 11 mm (N =
1); venation same as in male. Dorsal surface cinnamon,
with a faint olive brown discal spot, ventral surface iden-
tical. Hindwing dorsal surface flesh ochre, pattern be-
ginning at proximal portion of adbasal region, expanding
transversely to most distal portion ofpostmedial region,
meeting inner margin of dark raw umber, extending from
postmedial region to wing apex, occasionally with a sinu-
ate pattern, large dark raw umber discal spot present;
ventral surface identical. Abdomen flesh ochre, dorsal
tergites occasionally with a medium size dark raw um-
ber spot in center, size tapering with each segment to-
wards posterior. Male genitalia: Phallus rounded coecum
penis rounded weakly notched (Fig. 22A), inflected ven-
trally, basiphallus straight, phallic sclerite wide and nodu-
lar (Fig. 22C). Vesica directed ventrally when everted,
membrane rugose, bilobed, elongate, second lobe scle-
rotized, right side of vesica with patch of comuti under
phallic sclerite (Fig. 22C). Genital capsule (Figs. 27C)
with dorsum oftegumen concave, bent inward (Fig. 23A),
conjunctivamembranous. Uncus base cordate (Fig. 23B),
ventrad with uncus rounded at sides with notch at cen-
ter, with two hooks (Fig. 27B). Scaphium rounded,
pointed. Subscaphium absent, extensions of tegumen


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

sclerotized. Juxta cylindrical, emarginated, with ends
barbed toward dorsum. Costa of valva elongate and tip
entire, 20 or more costal setae. Sacculus horizontally
obovate, broadly hooked toward costa (Fig. 27C). Sac-
cus rounded, concave in center. Female genitalia: Lack-
ing pseudopapillae anales. Pheromone glands membra-
nous, branching after tip of anterior apophyses, thin, less
than 1/8 width of 7th abdominal stemrnite. Ostium bursa
slighty concave with small lateral prominences (Fig.
30B). Ductus rectangular, narrowing toward venter (Fig.
17A). Corpus bursa with one signum, membrane smooth
(Fig. 17B).
Variation.-- Some specimens collected in Colo-
rado, New Mexico and the White mountains of Arizona
are phenotypically identical to V marginata and share
the diagnostic conditions of the coecum penis and
tegumen. Additional V marginata specimens have been
collected from Texas and Mexico, but are rare, making
it difficult to examine population variation. The U.S.
populations appear to have disjunct distributions from
the main species' range. Potentially, these population
isolates are genetically distinct species; however, adult
morphology does not distinguish them.
Biology.-- Unknown.
Distribution.- This species primarily occurs in
Mexico, Guatemala, and Honduras. Specimens from
New Mexico and Arizona are tentatively identified as V
marginata for reasons provided in Variation.

Virbia nigricans Reakirt
P1. 2, nos. 8 (female), 9 (male)
Crocota nigricans Reakirt 1864: 371 [type local-
ity, Philadelphia, Pennsylvania, USA].
Cothocida nigrifera Walker 1865b: 499 [type lo-
cality, "USA"].
Material examined.- Type material: Holotype of
V nigricans: Crocota nigricans Philadelphia, Pa.
Reak. Type Coll. Reak.//Crocota nigricans Reak.
Pennsylvania Philadelphia Type Coll. Reak. Strecker
Colln. Field Museum of Nat. Hist.//genitalic preparation
172 Ring Carde//Lepidoptera Photograph No. 186 Field
Museum [sic.] FMNH. Holotype (synonym of V
nigricans): Holomelina nigricans "D.T." (green la-
bel) nigrifera Walker "Cathocida nigrifera" N.
America, 62 85'//Arctiidae slide 969, BM negative 43760,
NHM. All material is from the NMNH collection un-
less otherwise noted: UNITED STATES: New Jer-
sey: Hunterdon Co., 10.vii. 1964, Joseph Muller, ex ovo
(1 Female, JMZ269); 16.vii. 1964, Joseph Muller, ex ovo
(1 Female, JMZ270); 11.vii.1964, Joseph Muller, ex ovo
(1 Male, JMZ271); 25.vii. 1964, Joseph Muller, ex ovo,
AMNH (1 Male, JMZ272); Hunterdon Co. (1 Female,
JMZ187).





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


a a







A B




A
.......... . ...


C D

a
a






E F








Figure 28. A. Male 8' stemite, Virbia phalangia. A = antecosta triangular, a = anterior, p = posterior; B. Male 8'
stemite, Virbia opella. A= antecosta oval, egg-shaped, a = anterior, p = posterior; C. Male 8' stemite, Virbia
marginata. A = antecosta pentagonal, a = anterior, p = posterior; D. Male 8' stemite, Virbia divisa. A = antecosta
pentagonal with expanded anterior edge, a = anterior, p = posterior; E. Male 8' stemite, Virbia lutelinea. A =
antecosta reinforced by transverse ridge, rectangular, width greater than length, a = anterior, p = posterior; F. Male
8' stemite, Virbia metazonata. A = antecosta M-shaped, a = anterior, p = posterior.







Diagnosis.-- f, ,, nigricans has been confused
with dark V opella specimens. Virbia nigricans is
smaller (forewing length 9.3 mm), and the species re-
stricted to western New Jersey and eastern Pennsylva-
nia. Virbia opella tends to be larger (forewing length
11 mm) and has a wide geographic range. Virbia
nigricans is a daymating species (Fig. 8F, Carde 1971)
and its habitat is secondary old fields. Virbia opella is
active at night. Additionally, dark coloration in the anal
fold of the hindwing can also be used to separate V
nigricans from V opella.
Genitalia of both sexes are diagnostic. Males of V
nigricans lack a subscaphium and the costal tip is en-
tire (Fig. 27C); males of V opella possess a well-devel-
oped subscaphium and the costa tip is forked (Fig. 27D).
Female V nigricans have a corpus bursa with 3 signa
(Fig. 26C), whereas V opella females have a connate,
double corpus bursa (Fig. 17C) and lack signa.
Description.-- Male habitus (Plate 2, No. 9):
Head buff to clay. Antennal scales raw umber. Third
segment of labial palpus exceeding front of head by at
least half of entire length, all segments buff to clay.
Thorax dorsal and ventral scales buff to clay. Legs legs
buff to clay, tarsal claw simple (Fig. 15A). Forewing
length 8.2 mm (N = 10), R'-R5 all present, M3 present,
extending to edge (Fig. 12). Dorsal surface tawny buff
from base, extending toward postmedial region, fading
into clay color through subterminal region to wing apex,
with a faint raw umber discal spot present, ventral sur-
face light to dark flesh ocher, occurring in varibale
amounts. Hindwing dorsal surface natal to army brown
extending from inner margin to outer edge, medial re-


Figure 29. Adult habitus: Virbia opella Grote, Male: 15
miW of Cocoa, Orange County, Florida 19.iii. 1966, R.H.
Leuschner Coll., AMNH.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

gion flesh ocher extending from wingbase along ante-
rior margin to postmedial region, with a natal brown discal
spot, fringes buff to clay color, ventral surface identical.
Abdomen buffto clay. Sclerotization of 8th sternite pen-
tagonal (Fig. 28C). Coremata reduced to small patches
of scales on membranous lobes. Female habitus (Plate
2, No. 8): Head same as in male, but in some specimens
infused with orange-rufous. Antennal color same as in
male. Third segment of labial palpus short, color usually
same as in male or infused with orange-rufous. Thorax
buff to clay. Legs buff or clay, tarsal claw simple (Fig.
15A). Forewing length 9.3 mm (N = 3), venation same
as in male. Dorsal surface same as in male,
occasionally suffused with orange rufous scale, ventral
surface same as in male. Hindwing dorsal surface usu-
ally same as in male, some specimens with large discon-
tinuous terminal bands dark sepia, ventral surface same
as in male. Abdomen usually same as in males, some
specimens suffused with orange rufous. Male genitalia:
Phallus with rounded coecum penis, straight, basiphallus
inflected ventrally at midpoint, phallic sclerite wide and
nodular (Fig. 22C). Vesica directed ventrally when
everted, membrane rugose, bilobed, elongate, second lobe
sclerotized, right side of vesica with patch of cornuti
under phallic sclerite (Fig. 22C). Genital capsule dor-
sally with tegumen concave, bent inward (Fig.23A), con-
junctiva membranous. Uncus base cordate (Fig. 23B).
Ventrad with uncus apex fingerlike, bifid, with two fused
hooks (Fig. 27A). Scaphium conical, pointed.
Subscaphium absent. Juxta cylindrical, emarginated, with
ends barbed toward dorsum (Fig. 27C). Costa elongate
and tip entire, 20 or more costal setae. Sacculus hori-
zontally obovate, broadly hooked toward costa. Saccus
rounded, concave in center. Female genitalia: Lacking
pseudopapillae anales. Pheromone glands membranous,
unbranched, thick, greater than 1/4 width 7th abdominal
sternite (Fig. 24B). Ostium bursa slighty concave with
small lateral prominences (Fig. 30B). Ductus rectangu-
lar, narrowing toward venter; sclerotization extending
into corpus bursa (Fig. 17A). Corpus bursa with three
signa, membrane wrinkled (Fig. 26C).
Variation.-- Little variation exists among V
nigricans males; one specimen examined from Penn-
sylvania has slightly more pinkish coloration in the
hindwing. Similarly, V nigricans females can vary
slightly in size and color.
Biology.-- Larvae have been reared on artificial
diet; the species is bivoltine with adults emerging in late
May to June and again in early to mid August. Males
are diurnal and weakly attracted to 2Me-17c (Carde
1971). Females lay unattached eggs individually like the
species V aurantiaca (Carde 1971).





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico


Figure 30. A. Ostium bursa, ', !,,, opella. Ob = ven-
tral edge of ostium (split), P = lateral prominence; B.
Ostium bursa, T /.' ia rubicundaria. Ob = ventral edge
of ostium bursa (with small, lateral prominences); C.
Ostium bursa, Virbia satara. Ob = ventral edge of os-
tium (toothed, with lateral prominences).


Distribution.- This species has a highly restricted
distribution, occurring only in western New Jersey
(Hunterdon Co.) and Pennsylvania (Philadelphia and
Montgomery Cos.) in secondary secession habitats.

Virbia opella Grote
P1. 3, Nos. 2 (female), 3 (male)
Crocota opella Grote 1863: 345 [type locality,
Pennsylvania, USA]
Crocotaflava Barnes & Benjamen 1925: 127-
129 [type locality, Quincy, USA].
Crocota obscura Stretch 1885: 103 [type locality,
Pennsylvania, USA]
Crocota belmaria Ehrmann 1895: 345 [type lo-
cality, Pittsburg, Pennsylvania, USA].
Crocota rubricosta Ehrmann 1895: 345 [type lo-
cality Jeanette, USA].
Material examined.-- Type material: Holotype not
examined. All material is from the NMNH collection
unless otherwise noted: UNITED STATES: Illinois:
Quincey, Poling //Bames Collection (1 Female, JMZ66);
1900.v., Poling //Bames Collection (1 Female, JMZ 158);
1900.vi., Poling //Bames Collection (1 Male, JMZ 157);
Magnolia, 1951.ii., A. K. Wyatt, ex ova (1 Female,
JMZ61). Massachusetts: Plymouth Co., E. Wareham
Agr. Exp. Sta., 10.vii.1976, W. E. Tomlinson at black-
light trap (1 Female, JMZ258); E. Wareham Agr. Exp.
Sta., 27.vii. 1977, W. E. Tomlinson at blacklight trap,
AMNH (1 Male, JMZ261); Martha's Vineyard, F. M.
Jones (1 Male, JMZ128). New Jersey: Cape May
Co., Eldora, J. Muller//Offspring No. 5, 17.viii. (1 Fe-
male, JMZ268); Eldora, 1977.vi.1923, J. Muller (1 Fe-
male, JZ290); Eldora, Jos. Muller//offspring No. 5, 17.viii.
(1 Male, JMZ267); Orange Mts., 6.ix., Otto Bucholz
Collection, AMNH (1 Male, JMZ289); Lakehurst,
nigricans (1 Male, JMZ83). New York: Horseheads,
30.viii.1943, L. R. Rupert //Bred ex ova Food: Dande-
lion//1022 nigricans (1 Male, JMZ84). Pennsylvania:
New Brighton, 7.vii.1975 //Eubaphe opella (1 Male,
JMZ82).
Diagnosis.-- Virbia opella is a highly phenotypi-
cally variable species ranging from a dark charcoal form
(Pl. 3, Nos. 2,3) to a pinkish form (Fig. 29). Nearly all
have a brown discal spot in the hindwing, although the
spot is obscured in the dark form (Plate 3, Nos. 2,3), and
paler forms may lack the discal spot. This species is
routinely confused with V nigricans, and occasionally
with V aurantiaca and V ferruginosa. Virbia
nigricans is restricted to western New Jersey and east-
ern Pennsylvania, and is a much smaller, darker species
(forewing length 9.3 mm) compared to the larger V
opella (forewing length 11 mm). For genitalic differ-








ences, see the Diagnosis of V nigricans. Virbia opella
is easily separated from V aurantiaca and V.
ferruginosa based on forewing venation; R3 is absent
or just a trace in V opella and present in the other two
species.
The pinkish form of V opella (Fig. 29) can be
confused with V. fergusoni (Fig. 91), but V. fergusoni
has darker forewings and more pronounced color con-
trast in the hindwings. For genitalic differences, see the
Diagnosis of V. fergusoni.
Description. --Male habitus (Fig. 28, Pl. 3, No.
3): Head olive brown. Antennal scales olive brown. Third
segment of labial palpus exceeding front of head by at
least half of entire length, first and second segments
orange rufous, third segment olive brown. Thorax dor-
sal and ventral scales olive brown. Legs cinnamon, oc-
casionally suffused with orange-rufous scales, tarsal claw
slightly bifid (Fig. 15B). Forewing length 11 mm (N =
10), R3 absent, M3 present, extending to edge (Fig. 13).
Dorsal surface olive brown to dark drab with a sepia
discal spot present, ventral surface clay suffused with
salmon extending from adbasal region through medial
region. Hindwing dorsal surface identical to forewing,
ventral surface identical. Abdomen olive brown, edges
of tergites, pleurites and sternites layered with scales
ranging from clay to orange rufous among specimens.
Sclerotization of 8th sternite oval, egg shape (Fig. 28B).
Coremata absent. Female habitus (Pl. 3, No. 2): Head
cinnamon. Antennal scales fuscous. Third segment of
labial palpus short; first and second segments range from
clay to orange-rufous, third segment grayish horn. Tho-
rax dorsal and ventral scales cinnamon. Legs intermixed
with olive brown and clay, tarsal claw slightly bifid (Fig.
15B). Forewing length 12 mm (N = 10), venation as in
male.Forewing dorsal surface cinnamon, faint fuscous
discal spot present, ventral surface peach red, terminal
margins a fringe of fuscous scales. Hindwing dorsal
surface peach red, tufts of faint cinnamon scales ex-
tending out from wingbase tapering towards antemedial
region with a dark fuscous discal spot present, ventral
surface identical. Abdomen dorsal abdominal scales
peach red, occasionally with fuscous spots on all seg-
ments, ventral and lateral scales peach red to clay, pat-
tern of fuscous same as dorsal side. Male genitalia:
Phallus (Fig. 19D) with rounded coecum penis, straight,
basiphallus inflected ventrally at base, curved toward
venter, phallic sclerite reduced. Vesica directed ven-
trally when everted, membrane rugose, bilobed, elon-
gate, second lobe of vesica rugose, left side of vesica
with patch of comuti located ventrally, right side of vesica
with patch ofcomuti under phallic sclerite. Genital cap-
sule (Fig. 19C) dorsum with tegumen entire, conjunctiva


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

membranous (Fig. 23C). Uncus base cordate, (Fig. 23B),
ventrad with uncus apex fingerlike, bifid and with two
hooks (Fig. 27A). Scaphium rounded, triangular.
Subscaphium lobed and incomplete, extensions of
tegumen sclerotized (Fig. 19C). Juxta ovoid with ven-
tral margin expanding into larger orbicular plate, dorsal
margin concave with pointed edges (Figs. 19C, 20A).
Costa of valva elongate and tip forked, 20 or more cos-
tal setae. Sacculus peanut shaped. Saccus rounded,
concave in center. Female genitalia: Pseudopapillae
anales present. Pheromone glands membranous, branch-
ing after tip of anterior apophyses, thin, less than 1/8
width of 7th abdominal sternite. Ostium bursa split with
two, large, lateral prominences (Fig. 30A). Ductus bursa
goblet shaped, sclerotization extending into corpus bursa
(Fig. 17C). Corpus bursa double, connate, signa absent,
both with smooth membrane (Figs. 17C, 26E).
Variation.-- Some females of V opella deviate
greatly from the colors discussed in the description. Many
females have olive brown to dark drab scales as in the
males. Some females have these scales, but the pattern
and color discussed in the description can be seen through
these dark scales. Some females display the wing pat-
tern and color discussed in the description, but are much
brighter; these differences in wing color do not seem to
be correlated with distribution (e.g., no pattern of geo-
graphic variation). This species is most closely related
to and may even be confused with V fergusoni. The
most distinctive male genitalic characters that can be
used to separate this species from V. fergusoni species
are the presence of a comutuslike patch located ven-
trally on left side of vesica and the lobing of the
subscaphium. In addition, V fergusoni females typi-
cally have a signum, while V opella females are lacking
signa. In addition to the morphological differences, wing
color and pattern can be used to separate V opella spe-
cies from V fergusoni.
Biology.-- Larvae have been reared on dande-
lion (Taraxacum) (Forbes 1960; Heppner 2003; Zaspel
database). Virbia opella is multivoltine, and adult males
and females fly at night and can be collected at a black
light in habitats ranging from oak forest to scrub oak
(Martha's Vineyard, Massachusetts). In Louisiana, V
opella have three broods from November-February
(Brou, personal communication).
Distribution.- This species occurs from Maine
west to Illinois and south to Texas.

T/,, ostenta Edwards
P1. 1, Nos. 1 (female), 2 (males)
Crocota ostenta Edwards 1881: 12 [type locality,
Prescott, Arizona, USA].





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

Holomelina calera Barnes 1907: 10 [type local-
ity, Huachuca Mts., Arizona, USA].
Material examined. Type material: Holotype of
V ostenta: Crocota ostenta Type Hy. Ed.//No 8195
Collection Hy. Edwards//Prescott Arizona//7600//Type
No. A.M.N.H., AMNH. Holotype (synonym of V.
ostenta): July 16-23//Huachuca Mts. Ariz.//Holomelina
calera Type Barnes//Photograph Pl. 12 No. 3//
PhotographPl. 26 No. 4//47/BL, NMNH. All material
is from the NMNH collection unless otherwise noted:
UNITED STATES: Arizona: Santa Rita Mts., S. S.
Nicolat Madera Canyon, 20.iii.1946 (1 Male, JMZ87);
Huachuca Mts., 24-30.vii//Bames Collection (1 Female,
JMZ225).
Diagnosis.-- Virbia ostenta is clay colored forew-
ings accented with a costal streak of light salmon scales
and with geranium pink extending from the proximal por-
tion of the adbasal region and meeting with the dark
brownish olive ground color (Pl. 1, Nos. 1, 2). Large for
irhbia, the forewing length averages 17 mm. This spe-
cies resembles V costata, but V costata is smaller
(Forewing length 11 -12 mm) and has a darker forewing
that lacks a costal streak. This species can also be mis-
taken for the Mexican V semirosea Druce, but the
forewing of V semirosea is darker, closer in hue to V.
costata females than V ostenta and the hindwing of V
semirosea is more red than pink like V ostenta.
Description.-- Male habitus (Pl. 1, No. 2): Head
yellow ocher. Antennal scales yellow ocher. Third seg-
ment of labial palpus exceeding front of head by at least
half of entire length, first and second segments a blend
of yellow ocher and light salmon scales, third segment
dark clay infused with cinnamon. Thorax a blend of
yellow ocher and clay. Legs dark brown, sepia, tarsal
claw simple (Fig. 15A). Forewing length 17.1 mm (N =
10); R'-R5 all present, M3 present, extends to edge (Fig.
12). Dorsal surface clay with a thin light salmon band
running full length of costal margin, ventral surface
dusted in light salmon. Hindwing dorsal surface dark
brownish olive, pattern with geranium pink beginning at
proximal portion of adbasal region, expanding trans-
versely to most distal portion of postmedial region, ex-
tending halfway down wing and meets ground color with
a sinuate margin, ventral surface identical. Abdomen
dorsal side dark brownish olive, lateral geranium pink,
ventral yellow ocher. Sclerotization of 8th sternite rect-
angular, length greater than width (Fig. 28E). Coremata
absent. Female habitus (Pl. 1, No. 1): Head sepia. An-
tennal scales dark brown, sepia. Third segment of labial
palpus exceeding front of head by at least half of entire
length, all segments dark brown, sepia. Thorax dark
brown, sepia. Legs yellow ochre with sparse patches


of light salmon on femora and tibiae, tarsal claw simple
(Fig. 15A). Forewing length 18.5 mm (N= 4), venation
same as in male. Dorsal surface antique brown with
thin peach red band running full length of costal margin,
ventral surface dusted in geranium pink. Hindwing dor-
sal surface fuscous, pattern with geranium pink begin-
ning at proximal portion of adbasal region, expanding


.








S



S
A

Dps Cpn



... y. .
i. a





Figure 31. A. Male genital capsule Virbia rindgei. J =
juxta (cylindrical, emarginated, with ends barbed toward
dorsum), Sa = sacculus (horizontally obovate, broadly
hooked toward costa), S = saccus (rounded, concave
midventrally); B. Phallus, Virbia rindgei, left, lateral
view. Dps= phallic sclerite (wide, nodular), Ve = vesica
bilobedd, primary lobe elongate with smooth membrane,
second lobe rugose), Cpn = patch of coruti (right side
of vesica under phallic sclerite).







transversely to most distal portion ofpostmedial region,
extending halfway down wing and meets ground color
with a sinuate margin, ventral surface identical. Abdo-
men fuscous. Male genitalia: Phallus (Fig. 22B) with
rounded coecum penis, straight, basiphallus inflected
ventrally at base, phallic sclerite present, narrow and
rounded. Vesica directed ventrally when everted, mem-
brane smooth, entire, left side with patch of comuti lo-
cated ventrally. Genital capsule with dorsum oftegumen
concave, bent inward, conjunctiva weakly sclerotized
(Fig. 23A). Uncus base cordate (Fig. 23B), ventrad
with uncus ape\ sl ighltl bifurcate, toothed with one hook
(Fig. 23E). Scaphium rounded, triangular. Subscaphium
completely surrounding anal tube (Fig. 19C), extensions
of tegumen sclerotized. Juxta vertically ovoid with ven-
tral margin expanding into larger orbicular shape, dorsal
margin concave with pointed edges (Fig. 20A). Costa
of valva elongate and tip entire, 20 or more costal setae.
Sacculus claw-like, extending under juxta on both sides.
Saccus rounded, concave in center. Female genitalia:
Lacking pseudopapillae anales. Pheromone glands mem-
branous, branching before tip of anterior apophyses, thin,
less than 1/8 width of 7th abdominal sternite. Ostium
bursa slighty concave with small lateral prominences (Fig.
30B). Ductus rectangular, narrowing toward venter (Fig.
17A). Corpus bursa double, connate, three signa present
in corpus, both bursae round with smooth membrane
(Fig. 17D).
Variation.- Female V ostenta specimens are rare
in collections. Almost no variation in color or size was
observed in either sex examined.
Biology.- Unknown.
Distribution.- Virbia ostenta is restricted to the
southwestern U.S. (New Mexico, Arizona) and Mexico,
occurring in mountain ranges.

Virbia rindgei Carde n. sp.
P1. 3, Nos. 8 (female), 9 (male)
Holotype.- Reuter Canyon Camp 5 mi. N. of
Sundance Crook Co., WYO., 5900 ft, 8.vii. 1962., F.,
P. & M. Rindge//Collected on NSF Grant G-25314,
Male, AMNH.
Paratypes.-- UNITED STATES: South Da-
kota: Custer Pk.: S. Dak. Lawrence Co., 6794 ft.,
1.vii.1962, F., P. & M. Rindge//JMZ307, Male,AMNH;
Lead, Lawrence Co., S. Dak., 5200 ft., 25.vii.1953, F.
& P. Rindge, Male, AMNH; Spring Creek Camp 11 mi.
NE Hill City S. Dak., 30.vii.1955, F. &P. Rindge, Male,
AMNH. Wyoming: Reuter Canyon Camp 5 mi. N.
Sundance Crook Co., 5,900 ft., 10.vii.1962, F. P. & M.
Rindge//Collected on NSF Grant G 25134//Genitalic
preparations 1963-41 Ring Card6, Male, AMNH; Reuter


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

Canyon Camp 5 mi. N. of Sundance Crook Co., 6100
ft., 13.vii.1959, F., P. & B. Rindge//JMZ276, Male,
AMNH; Reuter Canyon Camp 5 mi. N. of Sundance
Crook Co., 5900 ft., 8.vii. 1962, F., P. & M. Rindge, Male,
AMNH; Reuter Canyon Camp 5 mi. N. of Sundance
Crook Co., 5900 ft., 8.vii. 1962, F., P. & M. Rindge, Male,
AMNH; Reuter Canyon Camp 5 mi. N. of Sundance
Crook Co., 5900 ft., 9.vii.1962, F., P. & M. Rindge,
Male, AMNH; Reuter Canyon Camp 5 mi. N. of
Sundance Crook Co., 5900 ft., 9.vii.1962, F., P. & M.
Rindge//Collected on NSF Grant G-25314//Genitalic
Preparation 163-42 Ring Card6, Male, AMNH; Reuter
Canyon Camp 5 mi. N. of Sundance Crook Co., 5900
ft., 9.vii.1962, F., P. & M. Rindge//Collected on NSF
Grant G-25 314//Genitalic Preparation 164-65 Ring Card6,
Male, AMNH; Reuter Canyon Camp 5 mi. N. of
Sundance Crook Co., 5900 ft., 3.vii.1962, F., P. & M.
Rindge//Collected on NSF Grant G-25314//Genitalic
Preparation 164-72 Ring Card6, Male, AMNH; Reuter
Canyon Camp 5 mi. N. of Sundance Crook Co., 5900
ft., 9.vii.1962, F., P. & M. Rindge//Collected on NSF
Grant G-25314//Genitalic Preparation 164-82 Ring Card6,
Male,AMNH; 5-6 mi. N. of Sundance Crook Co., 5900-
6300 ft., 4.vii. 1962, F., P. & M. Rindge, Male, AMNH;
Reuter Canyon Camp 5 mi. N. of Sundance Crook Co.,
5900 ft., 8.vii. 1962, F., P. & M. Rindge//166//Collected
on NSF Grant G-25314, Female, AMNH. All material
is from the NMNH collection unless otherwise noted:
UNITED STATES: South Dakota: Gunnison Co., 6
mi. E. of Somerset, 29.vi.1959, leg. C. L. Remington (1
Female, JMZ306); Custer Park, 6500-6794 ft.,
30.vii.1955, F. & P. Rindge (1 Male, JMZ311).
Etymology. --This species is named in honor of
Dr. Fred Rindge. He is among the first to collect this
species and recognize it as distinct from other species in
the V aurantiaca complex. Dr. Rindge has had an
illustrious career as a Lepidopterist and his collections
have greatly increased our knowledge of North Ameri-
can lepidopteran fauna.
Diagnosis.-- Virbia rindgei might be confused
with V aurantiaca. Wing pattern, coloration and geni-
talia readily separate these species. Virbia rindgei males
have much darker forewings with rusty undertones (Fig.
P1. 3, No. 8), while V aurantiaca usually have solid
brown forewings (Pl. 1, No. 4). The hindwings of V
rindgei have dark brown shading from the subterminal
band along the fold from discal cell to 3A (often cover-
ing most if not all of the dark brown discal spot), whereas
the subterminal area of V aurantiaca is a solid band.
Virbia rindgei females (Pl. 3, No. 9) have much brighter
forewings than V aurantiaca females (Pl. 1, No. 5). In
males, the dorsal margin of the juxta of V rindgei is





ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

concave with pointed edges (Fig. 31A) whereas the juxta
of V aurantiaca lacks this elaborate dorsal margin (Fig.
20B). Female genitalia are not diagnostic. Although V
aurantiaca is widespread, it has not been collected in
the Black Hills of South Dakota (Lawrence Co.) or
Wyoming (Crook Co.).
Description. Male habitus (Pl. 3, No. 8): Head
mikado brown. Antennal scales mikado brown. Third
segment of labial palpus exceeding front of head by at
least half of entire length, first and second segments
peach red, third segment mikado brown. Thorax mi-
kado brown. Legs cinnamon, with patches of salmon
color scales on femora and tibiae, tarsal claw simple
(Fig. 15A). Forewing length 12 mm (N = 10); R'-R5 all
present, M3 present, extending to edge (Fig. 12). Dorsal
surface mikado brown, faint sepia discal spot present,
ventral surface a mixture of salmon and peach red scales
throughout, posterior margin a fringe of sepia. Hindwing
dorsal surface peach red, subterminal region a thick band
of sepia meeting postmedial region in sinuate pattern, in
most specimens sepia scales shading in region from sub-
terminal band along fold from discal cell to 3A, large
sepia discal spot present, ventral surface identical. Ab-
domen clay, dorsal tergites occasionally with medium
size sepia spot in center, size tapering with each seg-
ment towards posterior. Sclerotization of 8th sternite
pentagonal (Fig. 28C). Coremata reduced to small
patches of scales on membranous lobes. Female habi-
tus (Pl. 3, No. 9): Head dark cinnamon. Antennal scales
dark cinnamon. Third segment of labial palpus exceed-
ing front of head by at least half of entire length, all
segments dark cinnamon. Thorax mikado brown. Legs
clay, with patches of salmon on femora and tibiae, tarsal
claw simple (Fig. 15A). Forewing length 12 mm (N =
3), venation same as in male. Forewing dorsal surface
mikado brown throughout, faint natal brown discal spot
present, occasionally small white spot in annelar region
present, ventral surface identical, dusted with peach red.
Hindwing dorsal surface peach red, subterminal region
a thick band of sepia scales, meeting postmedial region
in sinuate pattern, in most specimens, sepia scales shad-
ing in region from subterminal band along fold from discal
cell to 3A with a large sepia discal spot, ventral surface
identical. Abdomen dorsal, lateral, and ventral abdomi-
nal scales scales clay to peach red, dorsal tergites with
large sepia spot in center, size tapering with each seg-
ment towards posterior. Male genitalia: Phallus with
rounded coecum penis, inflected ventrally, straight, phallic
sclerite wide and nodular (Fig. 31B). Vesica directed
ventrally when everted, membrane smooth, bilobed, sec-
ond lobe sclerotized (Fig. 22C), right side ofvesica with
patch of comuti under phallic sclerite (Fig. 31B). Geni-


Figure 32. Male genital capsule !,,, rubicundaria.
J =juxta (cylindrical and emarginated with ends barbed
toward dorsum), Sa = sacculus (horizontally obovate and
broadly hooked toward costa), S = saccus (rounded and
concave midventrally).


tal capsule (Fig. 31A) with dorsum of tegumen lobed,
sides divergent, conjunctiva membranous. Uncus base
cordate (Fig. 23B), ventrad with uncus apex fingerlike,
bifid, and with two fused hooks (Fig. 27A). Scaphium
rounded, triangular. Subscaphium absent. Juxta cylin-
drical, emarginated, with ends barbed toward dorsum.
Costa of valva elongate and tip entire, 20 or more costal
setae. Sacculus horizontally obovate, broadly hooked
toward costa. Saccus rounded, concave in center. Fe-
male genitalia: Pseudopapillae anales present. Phero-
mone glands membranous, branching after tip of ante-
rior apophypses, moderately thick, greater than 1/8 but
less than 1/4 width of 7th abdominal sternite. Ostium
bursa slightly concave with small lateral prominences
(Fig. 30B). Ductus bursa rectangular, narrowing toward
venter, sclerotization extending into corpus bursa (Fig.
17B). Corpus bursa with three signa, membrane of the
corpus bursa wrinkled (Fig. 26C).
Variation.-- Virbia rindgei has a highly restricted
distribution and is only known from specific localities in
Colorado, South Dakota and Wyoming. This species is
rare in collections and only represented by small series
of mostly males in some of the larger collections (e.g.,
AMNH & NMNH), rendering further study difficult.
Recent collecting attempts in South Dakota and Wyo-
ming were unsuccessful (R.T. Carde, D. Ferguson, J.
Zaspel). This species has not been collected since the
late 1970's.
Biology.-- The larvae are not known. The moths
were collected in Ponderosa Pine forest with an open








understory (Rindge unpublished field notes). From the
label data, this species appears to be single brooded,
with a flight in mid late July, possibly early August.
This species has been collected at elevations near or
above 6000 feet.
Distribution.-- This species has been collected at
high elevations in Colorado, South Dakota and Wyoming.

Virbia rubicundaria Huibner
P1. 2, Nos. 4 (female), 5 (male)
Crocota rubicundaria Huibner 1827: 256 [type
locality, Georgia, USA].
Crocota .~~ 1,'i Stretch 1885: 103 [type local-
ity, Texas, USA].
Crocota brevicornis Walker 1854b: 536 [type lo-
cality, "USA"].
Crocota diminutive Graef 1887: 42 [type locality,
Texas, USA].
Crocota rosa French 1890: 133 [type locality,
Texas, USA].
The holotype of Crocota rubicundaria is lost ac-
cording to Dr. Lloyd Martin, Curator of Lepidoptera at
Naturhistorisches Museum, Wien (personal communi-
cation 6/25/2004). A neotype is designated here and
labeled to ensure nomenclatural stability. The specimen
was chosen from the nearest locality to the original type
locality. Crocota rubicundaria Huibner, neotype Male
[NMNH] is labeled: Tb /,i rubicundaria (Hbn. 1827)
[RED LABEL]. Label data: GA: Trader's Hill, Folkston,
Charlton County 27.ix.1992, D.C. Ferguson//JMZ59,
Female, NMNH.
Material examined.-- Type Material. Holotype
lost. Syntype (synonym V rubicundaria): Holomelina
brevicornis syntype "140"Arctiidae slide 905 BM nega-
tive 43756, NHM. All material is from the NMNH col-
lection unless otherwise noted: UNITED STATES:
Florida: Gainesville, iv.1969, Dale Hubeck (1 Female,
1 Male; JMZ294, JMZ293); Highlands Co., Archibold
Biological Station, 23.vi.1966, Ring T. Carde (1 Male,
JMZ29 1); Archibold Biological Station, 16.vi. 1966, Ring
T. Carde, RTC (1 Male, JMZ292); Florida City,
20.iv. 1947, Otto Bucholz //Otto Bucholz Collection (1
Male, JMZ252); Lakeland, 5.v.1912 (1 Male, JMZ253);
Monticello, 4-8.x.1914, AMNH (1 Male, JMZ254); Lutz,
F. W., 1.iv. 1916, LACM (1 Male, JMZ 190); Florida City,
23.iv. 1947, Otto Bucholz //Otto Bucholz Collection (2
Females, JMZ248; JZ249); 15.iv.1947, Otto Bucholz //
Otto Bucholz Collection, AMNH (1 Female, JMZ250).
Georgia: Liberty Co., St. Catherine's Island, 7-
12.viii.1991, E. Quinter & A. Sharkov, AMNH (1 Fe-
male, JMZ251). Texas: Anderson Co., Tennessee
Colony, 21.vii.1965, A. & M. E. Blanchard (1 Male,


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

JMZ188); Bosque Co., Laguna Park, 24.ix.1970, A. &
M. E. Blanchard (1 Male, JMZ189).
Diagnosis.-- Virbia rubicundaria females can
be confused with V aurantiaca, however, these spe-
cies are rarely sympatric. See diagnosis of V
aurantiaca.
Virbia rubicundaria (Pl. 2, Nos. 4,5) can be con-
fused with V. fergusoni (Pl. 3, Nos. 4,5), but these spe-
cies can be separated based on size, coloration, and geni-
talia of both sexes. Virbia rubicundaria is typically
smaller (wing length 9 mm) with a lighter forewing (clay
with light salmon highlights) compared to V fergusoni
(wing length 11 mm) with darker raw umber coloration.
The male genitalia of V rubricundaria lack a
subscaphium (Fig. 32), the coecum penis is notched and
the phallic sclerite is wide and nodular (Fig. 22C) whereas
Vfergusoni males have a large subscaphium (Fig. 19A),
a rounded coecum penis and reduced phallic sclerite (Fig.
19B). Females of V rubricundaria have a single bursa
(no additional bursae) ornamented with three signa (Fig.
26C). In contrast, females of V fergusoni have a bi-
lobed corpus bursa with a single signum (Fig. 26E).
Description.-- Male habitus (Pl. 2, No. 5): Head
clay with light orange hues. Antennal scales clay with
light orange hues. Third segment of labial palpus ex-
ceeding front of head by at least half of entire length, all
segments a blend of clay and flesh ocher. Thorax clay
with light orange hues. Legs a blend of clay and flesh
ocher, tarsal claw simple (Fig. 15A). Forewing length
8.9 mm (N = 10); R'-R5 all present, M3 present, extend-
ing to edge (Fig. 12). Dorsal surface clay with light
orange hues, ventral surface dark salmon. Hindwing
dorsal surface flesh ocher, ventral surface ranging from
identical to light salmon. Abdomen flesh ocher to dark.
Sclerotization of 8th sternite pentagonal (Fig. 28C).
Coremata reduced to small patches of scales on mem-
branous lobes. Female habitus (Pl. 2, No. 4): Head
color same as in male. Antennal color same as in male.
Third segment of labial palpus exceeding front of head
by at least half of entire length, color same as in male.
Thorax same as in male. Legs same as in male, tarsal
claw simple (Fig. 15A). Forewing length 8.7 mm (N =
10); venation same as in male. Forewing dorsal surface
clay with light orange hues, ventral surface identical,
dark salmon with a faint natal brown spot between me-
dial and postmedial regions present. Although rare, some
specimens with small white spot in annelar region, ven-
tral surface identical. Hindwing dorsal surface flesh
ocher with a natal brown discal spot present, natal brown
subterminal markings as spots and blotches often not
reaching posterior margin, ventral surface identical.
Abdomen same as in male. Male genitalia: Phallus with






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

rounded coecum penis and weakly knotched (Fig. 22A),
inflected ventrally, basiphallus straight, phallic sclerite
wide and nodular (Fig. 22C). Vesica directed ventrally
when everted, membrane rugose, bilobed, elongate, sec-
ond lobe sclerotized, right side of vesica with patch of
comuti under phallic sclerite (Fig. 22C). Genital cap-
sule (Fig. 31) dorsum with tegumen lobed, sides diver-
gent, conjunctivamembranous. Uncus base cordate (Fig.
23B), ventrad with uncus apex fingerlike, bifid, with two
fused hooks (Fig. 27A). Scaphium rounded, triangular.
Subscaphium absent. Juxta cylindrical, emarginated, with
ends barbed toward dorsum. Costa of valva elongate
and tip entire, 20 or more costal setae. Sacculus hori-
zontally obovate, broadly hooked toward costa. Saccus
rounded, concave in center. Female genitalia:
Pseudopapillae anales present (Fig. 24B). Pheromone
glands membranous, branching before tip of anterior
apophypses, moderately thick, greater than 1/8 but less
than 1/4 width of 7th abdominal stemrnite. Ostium bursa
with small prominences on both sides (Fig. 30B). Duc-
tus bursa rectangular, narrowing toward venter, sclero-
tization extending into corpus bursa (Fig. 26C). Corpus
bursa with three signa, membrane of the corpus bursa
wrinkled (Fig. 26C).
Variation.-- Little variation in color pattern exists
among male V rubicundaria specimens. Although rare,
some males have a faint trace of a subterminal band on
the hindwings. Some V rubicundaria males from the
southwest were slightly larger than those from Florida
and Georgia. Of sixteen males, three had a rounded
coecum penis, not gently notched. Some females have
a brownish discal spot in the forewing and one speci-
men had a faint white spot in the annelar region of the
forewing. Female hindwing patterns were consistent.
Biology.-- Larvae have been reared on dande-
lion (Taraxacum) and wild lettuce (Lactuca floridana
Gaertner) (Zaspel Database). A multivoltine species,
moths are collected year-round throughout its range.
Distribution.- This species is found in Georgia
and Florida, and distributed along the Gulf Coastal States
to east Texas.

ACKNOWLEDGMENTS
We thank Michelle DaCosta, Howard Frank, Ralph
Holzenthal, William Miller, and three anonymous
reviwers for helpful comments. We also thank Demetrios
Vital for assistance with illustrations and Kevin Nixon
for assistance with the website. This work was sup-
ported SJW funding sources: NSF-DEB-9981416, NSF-
REU supplement, and a Minnesota Experiment Station
Project, MN-17-022. The project was also supported by
JZ's funding: Bell Museum curatorial assistantships, the


Dayton Wilke Fund (Bell Museum), and a Collections
Study Grant from the American Museum of Natural
History.


LITERATURE CITED
Barnes, W. 1907. New species ofNorth American Lepi-
doptera. Canadian Entomologist, 39: 10-15.
Barnes W., & F. H. Benjamin. 1925. Notes on some
species of Lepidoptera in relation to the Stretch
Collection. Pan-Pacific Entomologist, 1: 127-138.
Barnes W., & J. B. McDunnough. 1917. Checklist of
the Lepidoptera of Boreal America. Herald Press,
Decatur, IL, U.S. 4 (1): 84-85.
Barnes W., & J. B. McDunnough. 1918. Notes and
new species. Contributions to the natural history
of the Lepidoptera of North America, 4 (2): 61-
208.
Bremer, K. 1992. Ancestral areas: a cladistic reinter-
pretation of the center of origin concept. System-
atic Biology, 41: 436-445.
Brou, V. A. 2003. Holomelina laeta (Gr.-Men.) in
Lousiana. Southern Lepidoptera Newsletter 25: 81.
Brower, A. E. 1974. A list of the Lepidoptera of Maine-
Part 1, the macrolepidoptera. Life Sciences and
Agriculture Experiment Station Tech. Bull. 66: 1-
136.
Buetenmiller, W. 1898. Bombycine moths of the vicin-
ity of New York. Bulletin American Museum of
Natural History, 10: 365-366.
Card6, R. T. 1965. Some taxonomic notes on the nearctic
Holomelina with a partial key to species. Journal
of the Lepidopterists' Society, 19: 69-76.
Card6, R. T. 1968. A revision of the Holomelina
aurantiaca species group (Lepidoptera: Arctiidae).
Master's thesis. Cornell University, Ithaca, N.Y.
Card6, R. T. 1971. Aspects of reproductive isolation in
the Holomelina aurantiaca complex (Lepidoptera:
Arctiidae). Ph.D. thesis. Cornell University, Ithaca,
N.Y.
Card6, R. T. 1974. Diel periodicities of female calling
and male pheromone attraction in Holomelina
aurantiaca (Lepidoptera: Arctiidae). Canadian
Entomologist, 106: 933-934.
Card6, R. T. & W. L. Roelofs. 1973. Temperature
modification of male sex pheromone response and
factors affecting female calling Holomelina
immaculate (Lepidoptera: Arctiidae). Canadian
Entomologist, 105: 1505-1512.
Card6 & K. F. Haynes. 2004. Chapter 8. Structure of
the pheromone communication channel in moths.
In Advances in Insect Chemical Ecology, Edited








by R. T. Card6 & J. C. Millar. Cambridge Univer-
sity Press p. 283 332.
Charlton, R. E. &W. L. Roelofs. 1991. Biosynthesis of
a volatile, methyl-brached hydrocarbon sex
phermonefrom leucine by arctiid moths
(Holomelina spp.) Archives of Insect Biochemis-
try and Physiology, 18: 81-97.
Dallwitz, M. J., T. A. Paine, & E.J. Zurcher. 1993.
User's guide to the DELTA system: A general sys-
tem for producing taxonomic descriptions. Edition
4. CSIRO Division of Entomology, Canberra, Aus-
tralia.
Druce,H. 1885. Biologia Centrali-Americana (Zoologia)
Insecta, 1: 41-160. London.
Dyar, H. G 1897. The larva of Crocota opella Grote.
Psyche, 8: 119.
Edwards, H. 1881. Descriptions of two new species of
Lithosiidae. Papilio, 1: 12.
Ehrmann, G A. 1895. A few remarkable variations in
Lepidoptera. Canadian Entomologist, 26: 292-293.
Ferguson, D. C. 1953. On the identity and status of
Eubaphe lamae Freeman (Lepidoptera: Arctiidae).
Canadian Entomologist, 85: 371-373.
Ferguson, D. C. 1954. The Lepidoptera of Nova Scotia,
Part 1 (Macrolepidoptera). Bulletin No. 2. Nova
Scotia Museum of Science, Nova Scotia, Canada.
Ferguson, D. C. 1985. Contributions toward reclassifi-
cation of the world genera the tribe Arctiini, part 1:
introduction and a revision of the Neoarctia-
Grammia group (Lepidoptera: Arctiinae).
Entomography, 3: 181-275.
Fibiger, M., & J. D. Lafontaine. 2005. Systematic list of
the Noctuoidea of Europe (Notodontidae, Nolidae,
Arctiidae, Lymantriidae, Erebidae, Micronoctuidae,
and Noctuidae). Esperiana. Buchreihe zur
Entomologie 11: 7-92.
Fletcher, D. S. 1954. A revision of the genus Eubaphe
(Lepidoptera: Geometridae). Zoologica, 39: 153-
166.
Forbes, W. T. M. 1910. New England Caterpillars, No.2.
Eubaphe nigricans Reakirt. Journal of the New
York Entomological Society, 18: 163-164.
Forbes, W. T. M. 1939. The Lepidoptera of Barro Colo-
rado Island, Panama. Bulletin of the Museum of
Comparative Zoology, Harvard, 85: 234-321.
Forbes, W. T. M. 1960. Lepidoptera of New York and
neighboring states. Part 4. Agaristidae through
Nymphalidae including the butterflies. Agricultural
Experimental Station, Cornell University, Ithaca,
New York. Memoir p. 371.
Franclemont, J. G 1983. Arctiidae. In Check List of the
Lepidoptera of America North of Mexico, includ-


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)

ing Greenland. Edited by R.W. Hodges, T.
Dominick, D.R. Davis, D.C. Ferguson, J.G.
Franclemont, E.G. Munroe, and J.A. Powell. E.W.
Classey Ltd. and Wedge Entomological Research
Foundation, London, England. p. 114-119.
Freeman, T. N. 1941. New species of Canadian Lepi-
doptera. Canadian Entomologist, 73: 123-127.
French G H. 1890. Some new moths. Canadian Ento-
mologist, 22: 133-135.
Graef, E. L. 1887. Some new Bombycidae.
EntomologicaAmericana, 3: 41-43.
Grote, A. R. 1863. Additions to the catalogue of U.S
Lepidoptera. No. 2. Proceedings of the Entomo-
logical Society of Philadelphia, 2: 334-344.
Grote, A. R. 1865. Descriptions of North American Lepi-
doptera. VI. Proceedings of the Entomological So-
ciety of Philadelphia, 4: 315-330.
Gu6rin-M6neville, F. E. 1832. Iconographie du Regne
animal de G. Cuvier. Insectes: Paris and London,
plates 1-110, p. 6-576.
Hallberg, E., & G Poppy. 2003. Exocrine glands: chemi-
cal communication and chemical defense. In Hand-
book of Zoology, Lepidoptera, Volume 2: morphol-
ogy, physiology and development. Edited by N.P.
Kristensen. W. de Gruyter, Berlin, Germany, p.
361-376.
Hampson, G F. 1901. Catalogue of the Lepidoptera
Phalaenae in the British Museum. Volume 3. Trust-
ees of the British Museum, London, England.
Hampson, G F. 1920. Catalogue of the Lepidoptera
Phalaenae in the British Museum. Supplement 2.
Trustees of the British Museum, London, England.
Handfield, L. 1999. Le guide des papillons de
Quebec. Boucherville: Volume 1: 1-982, 123 color
plates. Broquet, Boucherville, Qu6bec.
Heppner, J. B., and G Lamas. 1982. Acronyms for
world museum collections of insects, with an em-
phasis on neotropical Lepidoptera. Bulletin of the
Entomological Society of America, 28: 305-315.
Heppner, J. B. 2003. Arthropods of Florida and neigh-
boring land areas, vol. 17: Lepidoptera of Florida,
Div. Plant Industry, Fla. Dept. Agr. & Comsum.
Serv., Gainesville. x + 670 pp., 55 plates.
Herrich-Schaffer, G A. W. 1850-1869. Sammlung neuer
oder wenig bekannter aussereuropaischer
Schmetterlinge. 6 Volumes. Regensburg, G J.
Manz.
Hodges, R. W., T. Dominick, D. R. Davis, D. C.
Ferguson, J. G Franclemont, E. G Munroe, & J.
A. Powell. 1983. Check List of the Lepidoptera of
America North of Mexico, including Greenland. E.
W. Classey Limited and Wedge Entomological Re-






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico

search Foundation. London, England.
Holland, W. J. 1903. The Moth Book. Doubleday and
Page. New York, U.S.
Huibner, J. 1823. Zutrdge zur Sammlung exotischer
Schmettlinge, Volume 2. Augsburg.
Hiubner, J. 1827. Zutrdge zur Sammlung exotischer
Schmettlinge, Volumes 3. Augsburg, Austria.
ICZN: International Code of Zoological Nomenclature.
1999. Fourth Edition. The International Trust for
Zoological Nomenclature xxix+305 pp. London.
Jacobson, N. L., & S. J. Weller. 2002. A cladistic study
of the Arctiidae (Lepidoptera) and a review of their
biology. Thomas Say monograph series, Entomo-
logical Society of America. Lanham, MD, U.S.
Kirby, W. F. 1892. A synomic catalogue of Lepidoptera
Heterocera, Volume 1, Sphinges and Bombyces.
Gurney and Jackson. London. England.
Kitching, I. J., & J. E. Rawlins. 1999. Noctuoidea. In
Handbook of Zoology, Lepidoptera, volume 1, Sys-
tematics and evolution. Edited by N.P. Kristensen.
W. de Gruyter, Berlin, Germany, p. 355-401.
Klots, A. B. 1970. Lepidoptera. In Taxonomist's glos-
sary of genitalia in insects. Edited by S. L. Tuxon.
Copenhagen, Munksgaard, p. 115-130.
Kristensen, N. P. 2003. Chapter 2 In Handbook of Zo-
ology, Lepidoptera, Volume 2: morphology, physi-
ology and development. Edited by N.P. Kristensen.
W. de Gruyter, Berlin, Germany, p. 361-376.
Lafontaine, J. D. and M. Fibiger. 2006. Revised higher
classification of the Noctuoidea (Lepidoptera).
Canadian Entomologist 138: 610-635.
Meyer, W. L. 1984. Sex pheromone chemistry and biol-
ogy of some arctiid moths (Lepidoptera: Arctiidae):
enantiomeric differences in pheromone perception.
Master's thesis, Cornell University, Ithaca, New
York.
Neumogen, B., & H. G Dyar. 1893a. A preliminary re-
vision ofthe Bombyces ofAmerica north of Mexico.
Journal of the New York Entomological Society,
1:153 -158.
Neum6gen, B., & H. G. Dyar. 1893b. Notes on
Lithosiidae andArctiidae with descriptions of new
varieties. Entomological News, 4: 138-143.
Packard, A. S. 1887. Notes on certain Psychidae, with
descriptions of two new Bombycidae.
EntomologicaAmericana, 3: 51-52.
Reakirt, T. 1864. Contributions towards a monograph
of the genus Crocota. Proceedings of the Ento-
mological Society of Philadelphia, 2: 371-373.
Roelofs, W. & R. T. Card6. 1971. Hydrocarbon sex
pheromone in tiger moths (Arctiidae). Science, 171:
684-686.


Saunders, W. 1869. On a supposed new Arctian. Ca-
nadian Entomologist, 2: 4-5.
Scoble, M. J. 1992. The Lepidoptera: form, function,
and diversity. Oxford University Press, New York.
Schal, C. & R. T. Card6. 1985. Rhythmic extrusion of
pheromone gland elevates pheromone release rate.
Experentia, 41: 1617-1619.
Schal, C. & R. T. Card6. 1986. Effects of temperature
and light on calling behavior in the tiger moth
Holomelina lamae (Freeman) (Lepidoptera:
Arctiidae). Physiological Entomology, 11: 75-87.
Schal, C., R. E. Charlton, & R. T. Card6. 1987. Tempo-
ral patterns of sex pheromone titers and release
rates in Holomelina lamae (Lepidoptera:
Arctiidae). Journal of Chemical Ecology, 13: 1115-
1129.
Schal, C., V. Sevala, & R. T. Card6. 1998. Novel and
highly specific transport of a volatile sex pheromone
by hemolymph lipophorin in moths.
Naturwissenschaften, 85: 339-342.
Schaus, W. 1901. Descriptions of new species of
Heterocera. Annals and magazine of Natural His-
tory 7: 265-270.
Seitz, A. 1914. The Macrolepidoptera of the American
Region, the American Bombyces and Sphinges. The
macrolepidoptera of the world, volume 6 (English
edition). Alfred Kemen, Stuttgart, Germany.
Smith, J. B. 1889. Preliminary catalogue of the Arctiidae
of temperate North America, with notes. Canadian
Entomologist, 21: 194-197.
Smithe, F. B. 1975. Naturalist's color guide. The Ameri-
can Museum of Natural History, New York. Color
work by Hale Color Consultants, Baltimore.
Strand, E. 1919. Arctiidae, subfamily Arctiinae.
Lepidopterorum Catalogus, 22: 1-416. Junk, Ber-
lin, Germany.
Strecker, H. P. 1878. Section II. Lepidoptera. Family
Bombycidae, In Annual report upon the explora-
tions and surveys in the Department of the Mis-
souri, being Appendix SS of the annual report of
the chief of engineers for 1878. Edited by E.H.
Ruffner. US Government Documents. Washing-
ton, D.C., p. 1859-1861.
Stretch, R. H. 1885. Descriptions of new species of
Heterocera. Entomologica America, 1: 101-107.
Tietz, H. M. 1972. An index to the described life histo-
ries, early stages, and hosts ofthe Macrolepidoptera
of the Continental U.S. and Canada. Volume 1.
Allyn Museum of Entomology, Sarasota, FL, U.S.
Walker, F. 1854. List of the specimens of Lepidopter-
ous insects in the collection of the British Museum,
2: 279-581. Trustees of the British Museum, Lon-





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


don, England.
Walker, F. 1865a. List of the specimens of Lepidopter-
ous insects in the collection of the British Museum,
31: 1-322. Trustees of the British Museum, Lon-
don, England.
Walker, F. 1865b. List of the specimens of Lepidopter-
ous insects in the collection of the British Museum,
32: 323-706. Trustees of the British Museum, Lon-
don, England.
Watson, A., & D. T. Goodger. 1986. Catalogue of the
neotropical tiger-moths. Occasional Papers on
Systematic Entomology, 1: 1-71.
Watson, A., D. S. Fletcher, & I. W. B. Nye. 1980. In
The generic names of Moths of the World, Volume
2, Noctuoidea (part). Edited by I.W.B. Nye. Trust-
ees of the British Museum (Natural History), Lon-
don, England.
Weller, S. J., R. B. Simmons, R. Boada, &W. E. Conner.
2000. Abdominal modifications occurring in wasp
mimics of the Ctenuchine-Euchromiine clade (Lepi-
doptera: Arctiidae). Annals Entomological Society
of America, 93: 920-928.
Winter, W. D. Jr. 2000. Basic Techniques for Observ-
ing and Studying Moths and Butterflies. Memoirs
of the Lepidopterists' Society, 5: 1-444.
Wyatt, A. K. 1939. Notes on the identity of certain
forms of Eubaphe. Canadian Entomologist, 71: 96-
99.
Wyatt, A. K. 1963a. A new subspecies of Holomelina
aurantiaca from Virginia (Arctiidae). Journal of
the Lepidopterists' Society, 17: 100-102.
Wyatt, A. K. 1963b. Holomelina aurantiaca
buchholzi, a correction. Journal of the Lepidop-
terists' Society, 18: 118.
Yin, L. R. S., C. Schal, & R. T. Carde. 1991. Sex phero-
mone gland of the female tiger moth Holomelina
lamae (Lepidoptera: Arctiidae). Canadian Journal
Zoology, 69: 1916-1921.
Zaspel, J. M. 2004. Chapter 1: Review of generic limits
of the tiger moth genera Virbia Walker and
Holomelina Herrich-Schaffer (Lepidoptera:
Arctiidae: Arctiinae) and their biogeography. Mas-
ters Thesis, University of Minnesota.
Zaspel, J. M. & S. J. Weller, 2006. Review of generic
limits of the tiger moth genera Virbia Walker and
Holomelina Herrich-Schaffer (Lepidoptera:
Arctiidae: Arctiinae) and their biogeography.
Zootaxa. 1159: 1-68.






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico 111





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Plate 1: 1-10

1. Adult habitus: Virbia ostenta Edwards, Female: Huachuca Mountains., Arizona//Barnes Collection//24-30.vii.,
USNM.
2. Adult habitus: T ',/,,u ostenta Edwards, Male: Williams, Coco-Nino Co., Arizona, 10.viii. 1955, N. McFarland at
light, LACM.
3. Adult habitus: Virbia aurantiaca Htubner, Male: Hartford Co., Farmington, Connecticut 9.vi.1961, leg. Ring T.
Card6, RTC.
4. Adult habitus: Virbia aurantiaca, Htubner, Male: Guilford Co., New Haven, Connecticut, 17.viii. 1966, leg. Ring T.
Card6, RTC.
5. Adult habitus: Virbia aurantiaca Htibner, Male: Guilford Co., New Haven, Connecticut, 19.viii. 1966, leg. Ring T.
Card6, RTC.
6. Adult habitus: Virbiaferruginosa Walker, Female: Edgebrook, Illinois 1-11.ix., A. Kwiat Bred//CNHM 1957
Alex. K. Wyatt Lepidoptera Colln, FMNH.
7. Adult habitus: Virbia ferruginosa Walker, Male: Guilford, New Haven CONN. 3.vii.1965 leg. Ring T. Card6,
RTC.
8. Adult habitus: Virbia lamae Freeman, Female: Seawall Bog, Acadia Pk., Mt. Desert Is., Maine 25.vii.1964, leg.
Ring T. Card6//Female parent of Brood, RTC.
9. Adult habitus: V ,ia lamae Freeman, Male: Seawall Bog, Acadia Pk., Mt. Desert Is., Maine 29.vii. 1966, leg. Ring
T. Card6, RTC.
10. Adult habitus: Virbia costata Stretch, Female: Prescott, Yavapai County Arizona 28.vii. 1970, Collected by Lloyd
Martin//Holomelina intermedia (Graef) det. Ring T. Card6, RTC.




ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico 113









1-A





3 4





5 6





7 8





9 10





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Plate 2: 1-10
1. Adult habitus: f ,.' ,,,costata Stretch, Male: Otero Co. 11.v.1926, N. Mexico//Harold M. Bower Coll. Access'd
C.A.C.M. 1964, FMNH.
2. Adult habitus: f ,i,,, immaculate Reakirt, Female: Edgebrook Illinois, Bred 7.x. 1947, A.K. Wyatt Coll.//CNHM
1957 Alex K. Wyatt Lepidoptera Colln., FMNH.
3. Adult habitus: Virbia immaculate Reakirt, Male: Edgebrook Illinois, Bred 7.x. 1947, A.K. Wyatt Coll.//CNHM
1957 Alex K. Wyatt Lepidoptera Colln., FMNH.
4. Adult habitus: Virbia rubicundaria Hutbner, Female: FLA., Highlands Co. Archbold Biol. Sta. 20.vi. 1966, Ring T.
Card6, RTC.
5. Adult habitus: Virbia rubicundaria Htibner, Male: FLA., Highlands Co. Archbold Biol. Sta. 20.vi. 1966, Ring T.
Card6, RTC.
6. Adult habitus: Virbiafragilis Strecker, Female: Spring Creek Camp 11 mi. NE Hill City Pennington County
S.Dak., 1.viii.1955, F. and P. Rindge, AMNH.
7.Adult habitus: Virbiafragilis Strecker, Male: Custer Park Lawrence County, S. Dakota, 6500-6794 feet, 30.vii.1955,
F. and P. Rindge, AMNH.
8. Adult habitus: Virbia nigricans Reakirt, Female: Hunterdon Co., New Jersey 11 -16.v. 1964, Joseph Muller//ex ovo,
AMNH.
9. Adult habitus: Virbia nigricans Reakirt, Male: Frenchtown, New Jersey, Hunterdon Co. Rte.29/10 mi. N Rte. 519,
4.viii. 1967, R. Card6, RTC.
10. Figure 86. Adult habitus: Virbia marginata Druce, Female: 1 mile S. Sulfer Springs, Sandoval Co., New Mexico,
7800 ft., 31 .vii. 1961, F. and P. Rindge.//Collected on NSF grant G 9037, AMNH.




ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico 115









1 2




3 4











7 8





9 10





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)


Plate 3: 1-9
1. Adult habitus: T; b1,a marginata Druce, Male: Guatemala City Guatemala//Schaus and Barnes coll//Dognin Col-
lection//July//Holomelina marginata, USNM.
2. Adult habitus: Virbia opella Grote, Female: Magnolia Ill. ex ova ii. 1951, A.K. Wyatt, USNM.
3. Adult habitus: Virbia opella Grote, Male: Magnolia Ill. ex ova 9.vii. 1950, A.K. Wyatt//Otto Buchholz Collection,
AMNH.
4. Adult habitus: Virbiafergusoni Zaspel, Female: FLA, Highlands Co., Highlands Hammock State Park, 12.vi. 1987,
T.L. Schiefer//Holomelina n. sp. det. D.C. Ferguson, MSU.
5. Adult habitus: Virbiafergusoni Zaspel, Male: GA: 7 mi. NE Fargo, Clinch Co., 13.vii.1981, D.C. Ferguson//
Reared ex ovo on Taraxacum, USNM.
6. Adult habitus: Virbia laeta Gu6rin-M6neville, Female: Willard Missouri 19.vi., A.E. Brower//Eubaphe laeta Gu6rin-
M6neville Treatii Grote, USNM.
7. Adult habitus: Virbia laeta Gu6rin-M6neville, Male: Marathon County Wisconsin, Township of Stettin 6.vii. 1978,
Russell A. Rahn, USNM.
8. Adult habitus: V, .',a rindgei Card6, Male: Custer Park South Dakota, Lawrence County, 1.vii.1962, F., P., & M.
Rindge, AMNH.
9. Adult habitus: Virbia rindgei Card6, Female: Spearfish Caynon, Black Hills South Dakota, 1.vii.1939,A.C. Frederick,
USNM.






ZASPEL, WELLER and CARDE : Virbia ofAmerica north of Mexico






118 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)




University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs