Title: Bulletin of the Allyn Museum
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00079423/00034
 Material Information
Title: Bulletin of the Allyn Museum
Series Title: Bulletin of the Allyn Museum.
Abbreviated Title: Bull. Allyn Mus.
Physical Description: v. : ill. ; 23 cm.
Language: English
Creator: University of Florida. News Bureau.
Allyn Museum of Entomology
Florida State Museum
Florida Museum of Natural History
Publisher: The Museum
Place of Publication: Sarasota Fla
Subject: Entomology   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
Dates or Sequential Designation: Began in 1971.
Issuing Body: Vols. for <1985>- issued by the Florida State Museum; <1988>- by the Florida Museum of Natural History.
General Note: Separately cataloged in LC before no. 48.
General Note: Description based on: No. 4, published in 1972; title from caption.
General Note: Latest issue consulted: No. 123, published in 1988.
 Record Information
Bibliographic ID: UF00079423
Volume ID: VID00034
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 - 01451276
lccn - 87643372
issn - 0097-3211


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Published By
The McGuire Center for Lepidoptera and Biodiversity
Florida Museum of Natural History
University of Florida
P. O. Box 112710
Gainesville, FL 32611-2710
Number 150 20 November 2007


George T. Austin1, Andrew D. Warren1'2, Carla M. Penz3, Jorge E. Llorente-Bousquets2,
Armando Luis-Martinez2, and Isabel Vargas-Femandez2
'McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History,
University of Florida, P.O. Box 112710, Gainesville, Florida 32611
2Museo de Zoologia "Alfonso L. Herrera", Facultad de Ciencias, Universidad Nacional Aut6noma
de M6xico, Apdo. Postal 70-399, M6xico 04510 D.F., M6xico
Department of Biological Sciences, University of New Orleans, New Orleans, Louisiana 70148

ABSTRACT: A new species of Opsiphanes occurring in western Mexico is described,
illustrated, and compared with congeners and other members of the tribe Brassolini. It is
known from relatively mesic canyons at the interface between the Sonoran Desert and the
Sierra Madre Occidental in Sonora and, p -0ii!. disjunctly, in Colima. Larval foodplants
are identified, the mature larva and pupa are described and illustrated, and biogeographic
attributes are discussed.

KEY WORDS: biogeography, butterflies, early stages, foodplants, genitalia


Opsiphanes Doubleday, [1849] (Nymphalidae: Morphinae: Brassolini) are medium-
sized butterflies distributed over most of the Neotropics from Mexico to Paraguay and
Argentina. Adults of Opsiphanes are fast fliers, and their wings are proportionally small
compared to their large thorax. Anecdotal information suggests that mating is
crepuscular (e.g., Fruhstorfer 1912, DeVries 1987); in some species, groups of males
have been observed displaying on the wing and, in others, single males have been seen
patrolling edges of forests (CMP, pers. obs.) and among larval foodplants (ADW, pers.
obs.). Larvae of Opsiphanes feed on various genera of Arecaceae and Musaceae (Penz et
al. 1999), and some species regularly use coconut palm (Cocos ,,n if~ ra L.) as a larval
foodplant (Young and Muyshondt 1975; Young 1977; DeVries 1986, 1987). The 11

known species of Opsiphanes (Bristow 1991, Casagrande 2004) are dark brown or tawny
on the dorsum usually with a prominent transverse band on the forewing that ranges in
color from yellow-orange to white. Several species have similarly-colored submarginal
bands on the hindwing. The venter is variously striated, especially on the hindwing, and
has prominent eye-spots (ocelli) that are common on brassoline butterflies. Males of
Opsiphanes have large scent organs laterally on the abdomen, in addition to hairpencils
and patches of mealy androconial scales on the wings. During late afternoon, scent
released through these organs by displaying males probably plays a key role in mate
attraction (e.g., Srygley and Penz 1999). Within the Opsiphanes-group (Penz 2007), the
quartet Mielkella Casagrande, 1982, Orobrassolis Casagrande, 1982, Blepolenis Rober,
1906, and Opsiphanes are structurally similar. Among those genera, the male valva
typically displays a cluster of subterminal spines. On Opsiphanes, these spines are fused
to form a flattened process that is a defining character of the genus (Fig. 16a).
Opsiphanes boisduvallii Doubleday, [1849] (Figs. 1-2, 7-8) had been thought to be
unique within the genus in having a completely tawny ground color to both surfaces of
the wings and in lacking any trace of a transverse band on the dorsal forewing. Valval
morphology, nonetheless, places the unusually colored 0. boisduvallii within
Opsiphanes. Perhaps because of the immediately recognizable phenotype of O.
boisduvallii, detailed examination of tawny individuals of Opsiphanes has been delayed,
and all reports of tawny Opsiphanes have been automatically assumed to represent O.
boisduvallii (i.e., Bristow 1991, see also below). However, a striking undescribed
phenotype, similar to 0. boisduvallii, occurs at the northwestern limit of the genus'
distribution in western Mexico and this study provides a description of this new species.
Eight specimens in addition to photographs of living and pinned individuals were
examined and compared with other species of Opsiphanes, as well as Mielkella singularis
(Weymer, 1907), Orobrassolis ornamentalis (Stichel, 1906), and three species of
Blepolenis. These comparisons allowed us to verify the generic placement of the new
species. A description of the early stages is also provided.


The new species was compared with several other brassolines to confirm its generic
affinity and provide evidence that it represented an undescribed species. Pinned
specimens were used to examine general external morphology, and dissections of male
and female genitalia were performed using standard procedures. Comparisons of external
morphology and patterning were made with all species of Opsiphanes except Opsiphanes
camena Staudinger, [1886]; comparisons were also made with Mielkella, Orobrassolis,
and the three species of Blepolenis. Genital morphology was examined for M. singularis;
Orobrassolis ornamentalis; Blepolenis; Opsiphanes boisduvallii; Opsiphanes cassina C.
& R. Felder, 1862; Opsiphanes invirae (Htibner, [1808]); Opsiphanes sallei Doubleday,
[1849]; Opsiphanes quiteria (Stoll, 1780); Opsiphanes bogotanus Distant, 1875; and
Opsiphanes tamarindi C. & R. Felder, 1861. Specimens so examined and dissected are
deposited at the American Museum of Natural History, New York, NY; McGuire Center
for Lepidoptera and Biodiversity, Gainesville, FL; Milwaukee Public Museum,
Milwaukee, WI; Museo de Zoologia "Alfonso L. Herrera", UNAM, Mexico D.F.,
Mexico; and San Diego Natural History Museum, San Diego, CA. Photographs taken in
the field of live individuals were also examined. Field observations of larval biology
were made in Sonora, Mexico; larvae and pupae are described from photographs.

Opsiphanes NI'thl kl irnill, ,11 Austin & A. Warren, new species
(Figs. 3-6, 9-12, 14-17, 19-23)

Diagnosis. Among known species of Opsiphanes, 0. blythekitzmillerae is similar
only to 0. boisduvallii in its tawny wing color. Both males and females can be
distinguished from 0. boisduvallii by the following combination of characters: (1) wings
slightly translucent revealing the ventral eye-spots; (2) large eye-spots on the ventral
hindwing much larger than on 0. boisduvallii; (3) ventral color of the body and wings
pic.i. .!iii, li ll. whitish, in contrast to tawny on 0. boisduvallii.
Description. Male forewing length 45.2 mm, 45.8 mm (paratypes); venation typical
for genus (e.g., Stichel 1904) except distal end of discal cell on forewing less angled from
vein M3 to costa than on any species; forewing with apex produced (extending as far
distad as tornus), termen concave between veins M1 and CuA2 then convex to anal
margin, anal margin straight, hindwing termen convex, slightly undulate, torus rounded;
dorsum deep tawny becoming more tawny-ochreous basad on both wings, appearing
mottled due to ventral markings showing through; forewing brownish along costa
(broadest towards apex) and outer margin cephalad to vein 2A, hindwing narrowly
brownish along outer margin from apex to vein 2A, forewing with suboval blackish
subapical macule in Mi-M2, preceded by three small white subapical macules; hindwing
with yellow-tan or tawny-brown recumbent hairpencil (6 mm) originating in discal cell
just distad of origin of vein CuA2, extending caudad over vein cual-cua2 and overlying
pocket of modified black scales in proximo-caudal portion of CuA,-CuA2; hindwing with
anal area pale yellowish (tan on one paratype) with prominent black cross-striations
(showing through from venter); yellow-tan or tawny semi-erect hairpencil (1.8 mm)
originating just caudad of the middle of vein 2A with its distal end overlapping vein 2A;
narrow shiny area on both sides of vein 2A proximad and distad to hairpencil; remaining
portion of anal area heavily overscaled with pale yellow-tan or tan hair-like scales,
heaviest distad.
Venter with complex pattern; forewing with gray-brown outer margin outlined
proximad with dark brown, followed by band of pale tawny outlined proximad with
scalloped dark brown line defining distal edge of pale (distad) and dark (proximad)
brown postmedial band that grades to pale tawny cephalad and contains a large black
ocellus (6.5 mm, ocelli measured in mid-cell to outer edge of dark outline) in M1-M2, this
with a crescentic white central mark and outlined with pale tawny (extending into
cephalic portion of M2-M3) and then red-brown proximad; fairly prominent (on one
paratype, absent on other, present on male specimen seen in photograph, barely
discernible on photographed live male, Fig. 15) smaller white-pupiled black ocellus in
CuA1-CuA2 outlined with dirty tawny and then black, and two smaller and poorly defined
white-pupiled black macules in M2-M3 and M3-CuA1 (absent on one paratype and
photographed live male except for white dot in M2-M3, anterior supernumerary macule
with pupil on male specimen seen in photograph); postmedial band defined proximad at
its cephalic end by a narrow paler band that is tawny distad and whitish proximad; medial
area largely pale tawny striated with dark brown cephalad of vein CuA2 except distad of
end of discal cell where striations are absent; proximal to this from the end of the discal
cell to the bases of cells between Sc and M3 mottled whitish and dirty tan and crossed by
irregular black lines, this followed proximad in discal cell by a whitish band, a dark tan
band outlined by blackish, whitish, and then tan towards the base; most of wing caudad

of CuA2 pale tawny with an irregular black submedial mark extending caudad from vein
CuA1; oval whitish patch of mealy, i1pp ii !ll,. scent, scales just cephalad of vein 2A near
base; hindwing largely dull whitish, pale tawny in submargin, pale tan in central portion,
heavily striated with blackish striationss mostly 3-4 dark scales in width); large (9.8, 9.1
mm) tan ocellus centered in mid-cell Sc+Ri-Rs but extending into costal cell and
especially Rs-M1, this with crescentic white pupil and heavily outlined by black; another,
but somewhat smaller (8.2, 8.0 mm) and black ocellus centered in cell CuA1-CuA2 with
crescentic white pupil (gray-blue scales within crescent) outlined broadly with tawny
(this extending into cell CuAz-2A and especially into M3-CuA1 as an incomplete satellite
ocellus with a white pupil and a few blackish scales; black absent on one paratype and on
male specimen seen in photograph), the entire marking outlined with blackish proximad
and pale brown distad (white in M3-CuA1 interrupts dark outline on one paratype);
another incomplete ocellus in M2-M3 consisting of a white mark with a tawny patch
distad and a concentration of blackish scaling proximad (a complete ring around ocellus
on one paratype, photographed live male, Fig. 15, and on male specimen seen in
photograph); proximad to ocellus in Sc+R1-Rs is a prominent whitish-tan band, a
blackish margined tan band (this extending into cephalic portion of discal cell) followed
by another whitish band, a blackish smudge, and another tan band.
Dorsal head and anterior thorax pale warm brown, paler behind eyes; eyes dark
(appear tan with dark dots in photograph of living specimen); palpi warm brown on
dorsum, otherwise whitish-tan with a brownish band on outer sides; antennae red-brown;
remainder of dorsal thorax and entire abdomen tawny-ochreous like bases of wings;
ventral thorax and legs gray-tan, forelegs with brownish band on venter (when folded
against body); ventral abdomen pale gray-brown; abdominal scent gland suboval, 2.1, 2.2
mm in length.
Male genitalia (Fig. 16) uncus in lateral view curved downward to sharp point
caudad, about same length as tegumen, with a few setae on ventro-lateral surface, nearly
triangular in dorsal view narrowing to relatively blunt point caudad; tegumen hood-
shaped with two sets of 'hairpencils' (after Penz 2007) on dorsal surface; saccus narrow,
very slightly curved upward, about 2/3 length of uncus and tegumen combined; gnathos
with expanded and smooth proximal region, distal region abruptly narrower, sharply
pointed caudad; valva in lateral view broad cephalad, narrowing caudad to a blunt and
slightly upcurved tip, ventral edge with lobe towards cephalic end, dorsal edge with
unevenly-shaped flattened process directed dorsad and composed of several fused lobes
decreasing in size cephalad, most of ventral surface densely covered with setae; valva in
ventral view broad cephalad, abruptly narrowing caudad where curving slightly inward;
phallus slightly longer than valvae, nearly straight, with numerous pc- -lie- setae
especially near attachment of anellus and on phallobase; juxta heart-shaped.
Female forewing length 51.2 (ex larva, badly damaged), 54.8 i. i.l..pc, 55.7 mm
(paratype), apex rounded, not produced, termen nearly straight with slight ,.. i,,.
centrally, anal margin straight, hindwing termen convex with slight angle at M3, very
slightly undulate, tomus rounded; dorsum tawny (paler than male) becoming more
tawny-ochreous basad on both wings, appearing mottled due to ventral markings showing
through; forewing brownish along costa (broadest towards apex) and outer margin
cephalad to vein 2A, hindwing narrowly brownish along outer margin from apex to vein
2A, forewing with suboval blackish subapical macule in M1-M2, preceded by 3 small
white subapical macules; hindwing with anal area pale tan with prominent black cross-

striations (showing through from venter), heavily overscaled with tan hair-like scales,
heaviest distad.
Venter with complex pattern nearly identical with that of male, but with striations
finer; forewing with black subapical ocellus (6.6, 6.8, 6.9 mm); paratype (and a badly
damaged reared specimen) with white dot in M2-M3 similar to one paratype male (absent
on holotype and photographed live female, Fig. 14); hindwing with ocellus centered in
mid-cell Sc+Ri-Rs larger than on male (11.0, 11.2, 11.5 mm); ocellus centered in cell
CuA1-CuA2 also larger (10.7, 11.0, 11.0 mm), this completely enveloping the satellite
ocellus in M3-CuA1 on holotype, the reared female, and photographed live female and
only partially including a white macule on the female paratype; ocelli represented by
white macules in M1-M2 and M2-M3.
Dorsal head and anterior thorax pale warm brown, paler behind eyes; eyes dark
(appear tan with dark dots in photograph of living specimen); palpi warm brown on
dorsum, otherwise whitish-tan with a brownish band on outer sides; antennae red-brown;
remainder of dorsal thorax and entire abdomen tawny-ochreous like bases of wings;
ventral thorax and legs gray-tan, forelegs with brownish band on venter (when folded
against body); ventral abdomen pale gray-brown.
Female genitalia (Fig. 17) sterigma broader than long, posterior section (Penz 2007)
well-developed with prominent central lobe, anterior section with two very thin arms
curving inward and nearly meeting centrally; lateral section thin, extended dorsally and
fused to edge of tergum 8; ductus bursae long, narrow distally, expanding cephalad to
elongate corpus bursae with parallel signa shorter than corpus bursae; papillae anales
with more-or-less triangular sclerotized base, but with thin projection from dorsal edge
extending caudad, posterior edge of papillae anales curved, this unsclerotized section
heavily adorned with setae.
Types. Holotype female MEXICO: Sonora; Mexican route 16, canyon with palms
near km 196, 30 August 1997, to which is attached a red, printed label / HOLOTYPE /
Opsiphanes blythekitzmillerae / Austin & A. Warren /. Paratypes: male, MEXICO:
Sonora; Cucurpe Road, 18 mi. SE Magdalena, 14 Oct 1979; male, MEXICO: Sonora;
Alamos, 27 August 1994; female, MEXICO: Sonora; Mexican route 16, 'fig-tree' canyon
near km 196, 16 September 1999 (at banana peel). The holotype is deposited at the
Museo de Zoologia "Alfonso L. Herrera", Facultad de Ciencias, Universidad Nacional
Aut6noma de Mexico, Mexico D.F., Mexico.
Additional records (specimens not examined by us unless noted otherwise): all from
MEXICO: SONORA; Cucurpe Road, 18-19 mi. SE of Magdalena (Sierra Baviso), 12
September 1981 (1 female), 3-4 April 1982 (1 male); SONORA; Mexican route 20 (E of
Hermosillo), 13 mi. E of El Novillo (Rio Yaqui), 12 August 1985 (sight), 25-27 April
1987 (sight); SONORA; Mexican Route 16, km 196, E of Rio Yaqui, 2 September 2000
(photograph), 11 November 2002 (photograph); SONORA; junction Aros and Bavispe
rivers, 9 April 2002 (photograph); SONORA; Sierra Madre Mts., Mexican route 16, 9
April 2005 (1 male); SONORA; Tarahumaris Road, ca. 9 mi. NE of Nuri (Arroyo
Honda), 24 September 1988 (2 females); SONORA; near Santa Rosa, 4 mi. E junction
Mexican route 16 and Sonora route 11, 23 September 1988 (1 female); SONORA; Rio
Sahuaripe, 7 miles S of Tacupeto, 27 March 1999 (sight); SONORA; Municipio Ures,
Sierra de Mazatan, riparian canyon, 2908'15"N, 11013'10"W, 1200 m, 9 October 2004
(larva); SONORA; road to Santa Barbara, 2705.745'N, 10845.426'W, 3672', upland
oak area, at sap, 2 October 2006 (sight); SONORA; near El Caj6n, upper Rio Cuchujaqui,

s1)9d ;:av

Figures 1-6. Males of Opsiphanes dorsumm on left, venter on right; line = 2 cm). 1-2) 0.
boisduvallii, MEXICO: Oaxaca; Chiltepec, 26.viii.1969; 3-4) 0. blythekitzmillerae, paratype,
MEXICO: Sonora; Alamos; 5-6) 0. blythekitzmillerae, paratype, MEXICO: Sonora; Cucurpe Rd.

Figures 7-12. Females of Opsiphanes dorsumm on left, venter on right; line = 2 cm). 7-8) 0.
boisduvallii, GUATEMALA: Zacapa; La Union, 20.x.1982; 9-10) 0. blythekitzmillerae, holotype;
11-12) 0. blythekitzmillerae, paratype, MEXICO: Sonora; route 16, 'fig-tree' canyon.

Figures 13-15. Opsiphanes habitat and live adults. 13) Type locality of 0. blythekitzmillerae
(photograph by Robert Behrstock); 14) female 0. blythekit millerae, MEXICO: Sonora; route 16,
km 196, 11 November 2002 (photograph by Douglas Danforth); 15) male 0. blythekitzmillerae,
MEXICO: Sonora; route 16, palm canyon near km 196, 2 September 2000 (photograph by Priscilla


.t~"~;f'~d5~'- j'~


a c d

1 mm


Figures 16-17. Genitalia of Opsiphanes blythekitzmillerae. 16) Male (MEXICO: Sonora, GTA
#13762); a. lateral view of genital capsule, b. ventral view of genital capsule, c. lateral view of
aedeagus, d. dorsal view of tegumen and uncus, e. juxta; 17) female (MEXICO: Colima, CMP #07-
128); a. lateral view of sterigma, ductus bursae, and corpus bursae, b. lateral view of papillae anales,
c. ventral view of sterigma.

10 October 2006 (larva); SONORA; Choquincahui, July 2005 (sight); COLIMA; La
Salada, 1000', 8 May 1967 (1 female examined); COLIMA; Mpio Villa de Alvarez,
Agua Dulce, 250 m, 29 October 1989 (1 male examined), 13 July 1995 (1 female
Type locality. MEXICO: SONORA; Mexico Route 16, canyon with palms near
kilometer 196 (Fig. 13).
Distribution and phenology. The northernmost record for 0. blythekitzmillerae is
from 18 miles southeast of Magdalena de Kino, on the road to Cucurpe. This site is
situated approximately 115 airline kilometers south of the border with Arizona, United
States (at Nogales). This and other records suggest that 0. blythekitzmillerae is a resident
of palm-inhabited canyons to an elevation of 1200 m along the western base of the
northern end of the Sierra Madre Occidental, bordering the Sonoran Desert, at least from
northern to southern Sonora (Fig. 18). Given the few known records, the distribution
given here is tentative. The occurrence of 0. blythekitzmillerae in Colima (Fig. 18)
suggests a broader distribution in western Mexico. Note that all tawny Opsiphanes we
have examined to date from Sinaloa (from the southern half of the state) and Nayarit are
0. boisduvallii. Vargas et al. (in press) did not record Opsiphanes from Sonora.
Available data for 0. blythekitzmillerae suggest it has at least two generations
annually in Sonora. Here, the species has been recorded in March and April and from
August through November. It has been recorded in Colima in May, July and October,
perhaps also representing two generations.



400 0 400
18 Kilometers

Figure 18. Distribution of Opsiphanes blythekitzmillerae (triangles) and 0. boisduvallii (dots) in

Habitat. Areas inhabited by 0. blythekitzmillerae in Sonora host a mosaic of short
tropical deciduous forest and desert scrub, with at least four species of palms in the genus
Brahea Mart. (Arecaceae), including Brahea brandegeei (Purpus) H. E. Moore, Brahea
aculeata i,! ii.lc-cc ., H. E. Moore, Brahea dulcis (Kunth) Mart., and Brahea nitida
Andre, as well as additional palms, Washingtonia robusta H. Wendl. (cultivated) and
Sabal uresana Trel. (Bdrquez et al. 1999, Bdrquez, pers. comm.). Such habitats have
been described and illustrated by various authors (e.g., Bdrquez et al. 1999, Martinez-
Yrizar et al. 2000, also Fig. 13 herein), and frequently occur as isolated montane habitat
patches separated by extensive areas of desert scrub (e.g., Sierra del Aguaje and Sierra
Libre) within the Sonoran Desert, the northern limit of tropical deciduous forests in
western Mexico. These ..cI-i ,ii. tropical habitats host numerous 'micro-endemic' taxa
of neotropical affinities, some of which are at the northern limits of their ranges,
occasionally as highly disjunct populations (Bdrquez 1997, pers. comm.; Bdrquez et al.
1999; Bdrquez & Martinez-Yrizar 2006, in press).
Biogeography. The primary known distribution of 0. blythekitzmillerae, from
northern to southern Sonora, corresponds with the Sonoran area of endemism (which
includes northern Sinaloa) revealed by many groups of plants and animals (Goldman and
Moore 1945, Cabrera and Willinik 1973, Marshall and Liebherr 2000, Morrone 2001,
Morrone et al. 2002, Espinosa et al. 2006). This region includes the lower elevations of
the western slope of the Sierra Madre Occidental, below the pine-oak zone, and most of
the Sonoran Desert west to the Pacific Coast. Other endemic species of butterflies
sharing this primary distribution include Polites norae MacNeill, 1993 (Hesperiidae),
Euchloe guaymasensis Opler, 1987 (Pieridae), and Voltinia ,J.Li..nii; (A. Warren &
Opler, 1999) (Riodinidae). The presence of 0. blythekitzmillerae in Colima, however,
shows that it is not strictly restricted to the Sonoran region, but that it also occurs,
,ppl'Iciill. disjunctly, in the Armerfa-Coahuayana area of endemism as defined by
Espinosa et al. (2006).
Larval foodplants and early stages. In Sonora, larvae were found on Brahea
aculeata (Fig. 22) near Municipio Ures and El Caj6n (one finished feeding on cultivated
Brahea armata S. Watson). Furthermore, 0. blythekitzmillerae co-occurs with Brahea
nitida at the site southeast of Magdalena, the northern known limit of that palm's
distribution r,.i..- R. Felger 2007; see Felger and Joyel 1999, Fleger et al. 2001).
Adjacent to kilometer 196, along Route 16, 0. blythekitzmillerae is associated with a
congener, Brahea brandegeei (Fig. 13). Both of these palms p.'- cli ii,. serve as larval
foodplants. Although known larval foodplants of other Opsiphanes are largely species of
Arecaceae and Musaceae, Brahea has not previously been reported as a larval foodplant
for any member of the Brassolini (Penz et al. 1999).
Mature larvae long and slender, head more-or-less rectangular in dorsal view bearing
two pairs of pointed and granulated scoli caudad; a shorter pair laterad, a longer pair
dorsolaterad. Two known color forms; one (Figs. 19, 21-22) largely reddish brown with
yellow-tan (becoming greenish cephalad) beneath whitish spiracles; entire body covered
with very short white setae emitting from minute white dots (probably chalazae); lateral
surface with two vague whitish lines, one just above spiracles, the other more dorsad.
More prominent and broader dorsolateral white line followed by another vague whitish
line on mid-dorsum. Ground color slightly deeper red-brown along spiracular area and
flanking pale lines. 'Anal fork' (=caudae) tan, intersegmental lines whitish. Head deeper
red-brown (nearly maroon), with broad yellow-green lateral stripe, the yellow-green
continuing onto the lateral scolus. Dorsolateral scolus red with narrowly yellowish tip.

Dorsal head with yellow-green central stripe flanked by narrower lines of a paler and
more yellowish color. Head covered with heavy vestiture of white setae of moderate
length, setae also occurring as pair of blackish tufts ('moustache') laterally from near
ventral portion of anterior head. Second form (Fig. 20) similar, but red-brown replaced
by blue-green with darker green flanking pale lines, mid-dorsal line yellow; and 'anal
fork' concolorous with the body. Green of head more bluish than on body, lateral stripe
yellow-green, narrow grayish stripe between lateral and mid-dorsal stripes, mid-dorsal
stripe also yellow-green becoming greener caudad where flanked by whitish.
Dorsolateral pair of scoli brighter red than on red form with black tips preceded by
Pupae (Fig. 23) stout and somewhat hourglass shaped in dorsal view with slightly
bifurcate head. Dorsum with prominent keel and, in lateral view, convex and somewhat
bulbous cephalad. Ventral edge slightly concave. Pale greenish caudad, but pinkish over
wing pads and head. Abdomen with thin and darker green-brown lateral line below
spiracles, spiracles outlined with pale brown. Less prominent line dorsolaterally, two
thinner lines ventrolaterally. Entire pupa with reticulate pattern of very fine lines, brown
on abdomen and thorax, deeper pink than ground color cephalad. Ochreous line along
edge of wing pad with prominent white macule outlined with brown just caudad of its
Mature larvae of 0. blythekitzmillerae are typical of the genus in their general aspect.
Larvae of Opsiphanes tend to be greenish and often pi. ',.i,,clii. striped with contrasting
colors (Harrison 1963, Young and Muyshondt 1975, Young 1977, Cubero 1985, DeVries
1987), perhaps more so than larvae of 0. blythekitzmillerae; we, however, found no
report of larval dimorphism within the genus. All species studied have prominent
markings on the head with the number of scoli varying from two to four pairs; this may
vary intraspecifically (Young and Muyshondt 1975, Young 1977, Cubero 1985). Pupae
of 0. blythekitzmillerae are also typical; those of the genus are amazingly similar in form
and pattern (Harrison 1963, Young and Muyshondt 1975, Young 1977, Cubero 1985,
DeVries 1987). A pupal case of 0. boisduvallii examined from Colima appears
indistinguishable from those of 0. blythekitzmillerae.
Larvae of 0. blythekitzmillerae fed nocturnally, as ipp .111cil. usual for Brassolini
(e.g., DeVries 1986, 1987) including Opsiphanes (Harrison 1963, Young 1977), and
spend the day in silken creches (Figs. 19-21) as do others of the genus (Harrison 1963,
Young 1977, Cubero 1985). Both solitary larvae wandered away from the foodplant at
night to pupate, one on a vertical surface and the other on a horizontal surface. Pupation
occurred two days after leaving the foodplant as noted for the prepupal stage of other
species in the genus (Harrison 1963, Young and Muyshondt 1975, Cubero 1985). Pupal
duration was 15 and 16 days, within the range previously reported for Opsiphanes
(Harrison 1963, Young and Muyshondt 1975, Cubero 1985).
Etymology. This species is named in honor of Margery Minerva Blythe Kitzmiller.
She was born November 17, 1883 in Malvern, Ohio and died March 10, 1972. She
married Frank Kitzmiller on September 7, 1904 in Cleveland, Ohio and they lived in
Pittsburgh, Pennsylvania. They had three sons, all of whom fought in World War II.
Minerva, or "Bango" to her grandchildren, was a very creative person who wrote poetry,
played piano and sang. This owl butterfly is being named in her honor on behalf of her
five grandchildren, who loved her very much.



Figures 19-23. Larvae and pupae of Opsiphanes blythekitzmillerae (MEXICO: Sonora; Sierra de
Mazatin). 19, 21) Red form of larva on Brahea aculeata, 20) green form of larva on B. aculeata,
22) red form of larva showing distinctive feeding damage on B. aculeata, 23) pupa (artificially
attached to B. aculeata for photography). Images by Michael Wilson.

Discussion. Among butterflies, Nymphalidae is considered one of the best known
families taxonomically and it is thought that few species remain to be discovered (Brown
1996, 1997). This is especially true for large and showy taxa, particularly those that have
obviously distinctive phenotypes. Within Opsiphanes, more than 100 years has elapsed
since the description of a new species (Stichel 1902), making the discovery of
Opsiphanes blythekitzmillerae so close to the border of the United States and the
northernmost known record of a Brassolini (Bristow 1991) all the more remarkable.
It has been four decades since the first known specimen of Opsiphanes
blythekitzmillerae was obtained. A specimen from Sonora, originally identified as O.
boisduvallii, was noted (but not examined) in the revision of the genus by Bristow
(1991), and was assumed to represent the northernmost occurrence of that species, as a
late-season stray. Indeed, 0. blythekitzmillerae had been "hiding" in plain sight for some
time as a photograph on the internet (Warren et al. 2006). Warren first noticed the
unusual appearance of this taxon in December 2005 while examining photos taken by
Doug Danforth. None of the specimens in Warren's series of 0. boisduvallii from
Sinaloa, Nayarit, Jalisco, Colima or Michoacan matched Danforth's image from Sonora,
but the image was tentatively determined as 0. boisduvallii, and posted online, since no
other Opsiphanes had been reported from that far north or had a tawny dorsum. While
sorting collections at the McGuire Center in February, 2007, Austin called Warren's
attention to the unique phenotype of a specimen from Sonora. This led to a detailed
study of the few available images and specimens of Opsiphanes from northwestern
Mexico, leaving no doubt that an undescribed species was involved.
Similarity in size and wing shape between the new species and the Mexican Mielkella
singularis led us to ask whether 0. blythekitzmillerae belonged to that genus.
Accordingly, we compared wing and genital morphology of both taxa. On Mielkella,
wings are pi i-.l mi,,i m,,il, brown dorsally, and the forewing displays an orange postmedial
band that branches into the discal cell. In contrast, in species of Opsiphanes, this band is
yellow or white, and usually does not cross the discal cell (but see below). The region
between veins CuA1 and CuA2 on the ventral hindwing of both Mielkella and Opsiphanes
has enlarged scales with 3-5 long sharp points. Those scales are intermixed with a dense
cover of long and very fine filiform scales on Mielkella, while Opsiphanes have a
variable mix of the pointed scales and more robust filiform scales (see below). The scent
organ originating in the discal cell on the dorsal hindwing of 0. blythekitzmillerae
corresponds with that on other species of Opsiphanes, but is lacking on Mielkella. It
should be noted, however, that the same organ is present on two species of Blepolenis
and even on Opoptera Aurivillius, 1882, and can thus be considered plesiomorphic for
Opsiphanes. The abdominal scent organ occurs throughout the Opsiphanes clade (Penz
2007). Male genitalia of 0. blythekitzmillerae closely resemble those of other
Opsiphanes. On Mielkella (and also Orobrassolis and Blepolenis), the male valva has a
cluster of spines on the distal edge, while on Opsiphanes these are fused to form an
unevenly-shaped flattened process that is characteristic of the genus. Also on Mielkella,
the cluster of spines is located more distally than on Blepolenis, Orobrassolis, or
Opsiphanes (as a flattened process). In short, wing and genitalic morphology indicated
that the new taxon should be included within Opsiphanes. Furthermore, populations of
Mielkella always occur in evergreen forests in Mexico, while populations of Opsiphanes
occur in a wide variety of evergreen and deciduous forests, as well as riparian and
disturbed habitats (JL1, pers. obs.).

Among congeners, Opsiphanes blythekitzmillerae is similar in color to O.
boisduvallii, those two being the only species of Opsiphanes exhibiting a p!c I..i.!!"in, il.
tawny dorsum. The two species, however, differ in several important aspects (Table 1).
Opsiphanes blythekitzmillerae is, on average, notably larger than 0. boisduvallii, with a
proportionally longer forewing having a produced apex and a broader and less truncated
hindwing. The overall shapes of the wings of male 0. blythekitzmillerae, while
incorporating characters seen on other species of the genus, overall do not resemble those
of any individual species. The dorsum of 0. blythekitzmillerae has broader dark margins
on both wings than on 0. boisduvallii and most males of the latter have no dark margin
on the hindwing. The hairpencils of 0. blythekitzmillerae are pale and concolorous,
while these are at least dark distad on 0. boisduvallii; that originating in the discal cell is
longer on 0. blythekitzmillerae than on 0. boisduvallii. The abdominal scent glands of
0. blythekitzmillerae appear similar to those of 0. boisduvallii, although those of the
latter average 2.5 mm in length (2.1-3.1 mm, n=11). The venter on 0. blythekitzmillerae
is whitish compared with tawny on 0. boisduvallii, and more heavily marked with
coarser striations striationss on 0. boisduvallii are mostly 2-3 dark scales in width). The
large sizes of the ocelli on 0. blythekitzmillerae differ from the small ocelli on O.
boisduvallii, but resemble those seen on the majority of other species of Opsiphanes. The
extension of the outline of the ocellus in CuA1-CuA2 cephalad to envelop a satellite
ocellus in M3-CuA1, however, is rarely seen on other Opsiphanes (see below). On O.
blythekitzmillerae, the ocelli on the ventral hindwing are visible dorsally through a subtle
translucency that is considerably more pronounced than the intense and more opaque
colors typical of 0. boisduvallii. As noted above, Opsiphanes have a mix of enlarged
multipointed and filiform scales on the ventral hindwing in cell CuA1-CuA2. These range
from a thick vestiture of hair-like scales [Opsiphanes cassiae (Linnaeus, 1758),
Opsiphanes zelotes Hewitson, 1873, 0. bogotanus, 0. tamarindi], a more even mix of
pointed scales and yet thicker and much shorter rod-like scales (0. invirae, 0. cassina),
to those with very few of those undivided scales (0. sallei, 0. quiteria, Opsiphanes
mutatus Stichel, 1902). Both 0. boisduvallii and 0. blythekitzmillerae are similar to the
latter, but the divided scales have very long points.
Among specimens examined, none were encountered that were intermediate between
0. boisduvallii and 0. blythekitzmillerae; those examined from Sinaloa M I1.I -i I! east
of Mazatlan) are typical of 0. boisduvallii occurring further south. The specimens of O.
blythekitzmillerae from Colima are smaller in size (male forewing length = 40.0 mm,
female = 44.5, 47.2 mm) than are individuals from Sonora. Aside from this, we do not
see differences between material from Sonora and Colima, nor do we detect any patterns
of geographic variation in 0. boisduvallii. Where 0. blythekitzmillerae and 0.
boisduvallii occur in sympatry in Colima, they may be readily separated by wing
characters. We also note one female of Opsiphanes from Baja California Sur, Mexico,
listed by Brown and Faulkner (1982) and illustrated by Brown et al. (1992). This
individual is not 0. blythekitzmillerae and no different from females seen from
populations of 0. boisduvallii in other parts of Mexico. Until additional specimens from
Baja California become available, the range of variation of Opsiphanes there will remain
unknown. The single extant specimen suggests, however, that the population in Baja
California is 0. boisduvallii and not 0. blythekitzmillerae. It is possible that Opsiphanes
may have been introduced into Baja California on ornamental palms; the two records of
0. boisduvallii from Mexico City, at 2250m elevation, most likely represent this type of
human-assisted introduction (Gonzalez-Cota, pers. comm.).

Table 1. Comparative measurements of wing length, hairpencil length, and ocelli width for 0.
blythekitzmillerae and 0. boisduvallii. All specimens of 0. boisduvallii are from Veracruz, Mexico.

0. blythekitzmillerae 0. boisduvallii
male forewing length (mm) 43.7 / 40.0-45.8 /n=3 32.7 / 35.6-38.7 /n=10
average / range / n
female forewing length (mm) 50.7 / 44.5-55.7 / n=5 46.0 / 42.0-50.6 / n=9
average / range / n
hairpencil in HW discal cell (mm) 6.0, 6.0 / n=2 4.4 / 4.0-4.8 / n=10
average / range / n
male ocellus on forewing (mm) 6.5 / n=l 2.9 / 2.4-3.2 / n=10
average / range / n
male anterior ocellus on hindwing 9.1, 9.8 / n=2 4.9 / 4.0-5.4 / n=10
(mm) average / range / n
male posterior ocellus on hindwing 8.0, 8.2/ n=2 3.8 / 3.2-4.2 / n=10
(mm) average / range / n
female ocellus on forewing (mm) 6.8 / 6.6-6.9 / n=3 3.5 / 2.9-4.0 / n=9
average / range / n
female anterior ocellus on hindwing 11.2 / 11.0-11.5 / n=3 6.1 / 4.8-7.2 / n=9
(mm) average / range / n
female posterior ocellus on hindwing 10.9 / 10.7-11.0 /n=3 4.8 / 4.0-5.6 / n=9
(mm) average / range / n

The genitalia of 0. blythekitzmillerae (Figs. 16-17), although establishing its
placement within Opsiphanes, appear uninformative at the species-level. The genitalia of
the small sample of 0. blythekitzmillerae examined (2 males, 1 female from Sonora; 1
male, 1 female from Colima) are consistent with the descriptions given above. We have
noted during our investigations of the genus that there may be considerable intraspecific
variation in the genitalia of both males and females. This variation among males of O.
boisduvallii (n=12) includes the height and breadth of the tegumen, the length and
robustness of the uncus, the length of the saccus, and the breadth and angularity of the
harpe. While these structures tend to be less robust in 0. boisduvallii, as might be
expected due to their smaller average size than 0. blythekitzmillerae, there are males of
0. boisduvallii with genitalia that appear virtually identical to those of O.
blythekitzmillerae. Similarly, the sterigma of females of 0. boisduvallii (n=10) exhibit a
range of variation from having the anterior portion continuous (as illustrated by Penz
2007) to having a wide gap; the two females of 0. blythekitzmillerae examined have a
narrow gap. No character of the genitalia has yet been found that would unequivocally
distinguish the two species. Insufficient samples of most species of Opsiphanes have
been examined to determine if the genital variability seen in 0. boisduvallii is widespread
within the genus or if there are species-specific characters. For example, the shaft of the
phallus of 0. blythekitzmillerae has many pc.--i.k setae, particularly close to the
attachment point of the anellus, and also on the phallobase. Those pC-- i'k.c- setae are also
present on several other Opsiphanes, including 0. boisduvallii, 0. sallei, 0. invirae, 0.
quiteria, and 0. cassina. They are absent on 0. bogotanus and 0. tamarindi (genitalia of
the remaining species of the genus were not examined, see Materials and Methods).
Some elements of the wing pattern are shared among most species of Opsiphanes.
Transverse bands occur on the forewings in variable degrees of development involving
elements of the medial, submedial, and submarginal regions of the wing. The occurrence

of those bands is associated with a nearly uniform dark (generally brownish) region
posteriorly on the forewing with only 0. mutatus and 0. sallei exhibiting immediately
obvious medial patterns caudad of the discal cell. The discal cell on the ventral forewing
of Opsiphanes and the region just distad of it has alternating pale and dark bands. On
many species, the dark (and in some instances, the pale) bands are divided into macules
outlined with black. On 0. blythekitzmillerae along with 0. boisduvallii, 0. camena
(from photograph in Bristow 1991), 0. zelotes, and some 0. invirae, and 0. cassina, the
dark bands are undivided, presenting a more orderly aspect to this region of the wing.
Detailed examination of the pattern elements is likely to provide useful characters for
understanding the relationships within Opsiphanes. While 0. blythekitzmillerae might
have evolved through parapatric speciation from the geographically nearest taxon, 0.
boisduvallii, it is not yet clear which Opsiphanes species is the closest living relative of
0. blythekitzmillerae, due to the distribution and intraspecific variation of morphological
characters among the species of Opsiphanes discussed above.
Supernumerary macules occur widely within Opsiphanes. Individuals of most, if not
all, species have one or more such macules showing varying degrees of development on
the ventral hindwing. Those macules, however, rarely occur on the ventral forewing.
Examination of several hundred specimens of the genus indicated less than one percent
have supernumerary macules on the forewing; these were seen only on 0. boisduvallii, 0.
cassiae, 0. invirae, and 0. cassina. It is thus interesting that all five males and two of six
females of 0. blythekitzmillerae examined from specimens or photographs have
supernumerary macules on the ventral forewing. It is also worthy of note that on O.
blythekitzmillerae the outline of the ocellus in CuA1-CuA2 on the ventral hindwing
encloses or is split by the white of the macule in M3-CuA1. A similar enclosure occurs
very rarely on 0. tamarindi (e.g., Bristow 1991, Figs. 8E, 10B); the white macule more
often splits the dark outline on that species (e.g., Bristow 1991, Figs. 8F, 10C) as it
occasionally also does on 0. cassiae (e.g., Bristow 1991, Figs. 6C, 6F), 0. bogotanus
(e.g., Bristow 1991, Figs. 12D, 13A, 13D), and 0. quiteria (e.g., Bristow 1991, Fig.
In summary, the new brassoline from northwestern Mexico is the northwestern-most
ranging species of the tribe Brassolini. Although it is undoubtedly an Opsiphanes and
variably shares a number of characters with other species of the genus, the unique
combination of these features characterizes the distinct phenotype of O.
blythekitzmillerae. The tawny color of 0. boisduvallii had stood it uniquely apart from
the general dark-colored and pale-banded aspect otherwise universal within Opsiphanes,
and contributed to the delay in discovery of the new species. Opsiphanes
blythekitzmillerae adds a second tawny and unbanded phenotype to the genus, an
apparent enigma that has yet to be elaborated within the current knowledge of the
phylogeny, ecology, and biology of the genus.


Our thanks are extended to the many people who supplied material, data, and images
of or searched collections for the new species of Opsiphanes: R. Bailowitz, B. Behrstock,
J. Brock, H. and P. Brodkin, D. Danforth, J. Mori, D. Mullins, K. Roever, J. Smith, and
M. Wilson. We especially acknowledge M. Wilson who also sent images and extensive
notes on the early stages of the new species. We express our appreciation to R. Felger for
the identification of and information on palms; special thanks to A. Burquez for literature

and information about Sonoran Desert habitats. We also thank M. Wall, San Diego
Natural History Museum, for ..-.pl .Ili the loan of the female from Baja California, J.
Brown for information on that specimen, and curators of other collections for loans. P.
DeVries is thanked for his comments on an earlier draft of the manuscript. We thank J.
and L. Miller and J. Calhoun for critically reviewing the manuscript. We additionally
thank M. Trujano Ortega for producing the map. We are indebted to D. Matthews Lott
for photographing specimens, critically commenting on the manuscript, and expertly
organizing text and figures into this final product and to D.-M. Mintz for assistance in the
production of this publication. Financial support was provided in part by DGAPA-
UNAM (to A. D. Warren), DGAPA-UNAM IN-212006 (to J. Llorente, I. Vargas and A.
Luis), and NSF DEB 0527441 (to C. M. Penz). We also thank an anonymous donor for a
generous financial contribution.


Bristow, C. R. 1991. A revision of the brassoline genus Opsiphanes (Lepidoptera:
Rhopalocera). Zool. J. Linn. Soc. 101:203-293.
Brown, J. W. and D. K. Faulkner. 1982. New Rhopalocera records for Baja California
with the description of a new species of Habrodais Scudder (Lepidoptera: Theclinar
[sic]). Bull. Allyn Mus. 67:1-6.
Brown, J. W., H. G. Real and D. K. Faulkner. 1992. E;B..... ;.. of Baja Califinia.
Faunal Survey, Natural History, Conservation Biology. Beverly Hills, CA: The
Lepidoptera Research Foundation, Inc. iv + 129pp., 8pls.
Brown, K. S., Jr. 1996. The use of insects in the study, inventory, conservation and
monitoring of biological diversity in neotropical habitats in relation to traditional
land use systems. Pp. 128-149 in S. A. Ae, T. Hirowatari, M. Ishii, and L. P.
Brower (eds.), Decline and Conservation of EB ..... i, in Japan. Osaka: Lepid.
Soc. Japan (Yadoriga, special issue).
Brown, K. S., Jr. 1997. Diversity, disturbance, and sustainable use of neotropical
forests: insects as indicators for conservation monitoring. J. Insect Conserv. 1:25-
Bdrquez, A. 1997. Distributional limits of euglossine and meliponine bees
(Hymenoptera: Apidae) in northwestern Mexico. Pan-P.-. ,i'. Ent. 73:137-140.
Bdrquez, A. and A. Martinez-Yrizar. 2006. Conservation and landscape transformation
in northwestern Mexico. Pp. 465-477 in R. S. Felger and B. Broyles (eds.) Dry
Borders: Great Natural Reserves of the Sonoran Desert. Salt Lake City: The
University of Utah Press.
Bdrquez, A. and A. Martinez-Yrizar. (in press). Limites geograficos entire las selvas
bajas caducifolias y los matorrales espinosos y xer6filos: ique conservar? In G.
Ceballos and J. E. Bezaury (eds.) I.i. n...i;-. ....... de Amenazas y Prioridades para
Conservacidn en las Selvas Secas del P.. ,-... Mexicano. Mexico, D.F.: WWF-
Bdrquez, A., A. Martinez-Yrizar, R. S. Felger and D. Yetman. 1999. Vegetation and
habitat diversity at the southern edge of the Sonoran Desert. Pp. 37-67 in R. H.
Robichaux (ed.), Ecology of Sonoran Desert Plants and Plant Communities.
Tucson: The University of Arizona Press.

Cabrera, A. L. and A. Willinik. 1973. E,;. ... ... .r', de America Latina. ... ...i.,,, de
la OEA, Serie de Biologia, no. 13. Washington, DC: Organizaci6n de Estudios
Americanos. 122pp.
Casagrande, M. M. 2004. Tribe Brassolini. Pp. 201-205 in G. Lamas (ed.), Atlas of
Neotropical Lepidoptera. Checklist: Part 4A, Hesperioidea-Papilionoidea.
Gainesville, FL: Scientific Publishers. 439pp.
Cubero, R. 1985. Notes on the life cycle and natural history of Opsiphanes quiteria
quirinus Godman and Eryphanis aesacus buboculus Butler (Brassolidae). J. Lepid.
Soc. 39:33-42.
DeVries, P. J. 1986. Hostplant records and natural history notes on Costa Rican
butterflies (Papilionidae, Pieridae & Nymphalidae). J. Res. Lepid. 24:290-333.
DeVries, P. J. 1987. E;..i..;. of Costa Rica and Their Natural History. I.
Papilionidae, Pieridae and Nymphalidae. Princeton, NJ: Princeton Univ. Press.
Espinosa, D., J. Llorente and J. J. Morrone. 2006. Historical biogeographical patterns of
the species of Bursera (Burseraceae) and their taxonomic implications. J. Biogeogr.
Felger, R. S., M. B. Johnson and M. F. Wilson. 2001. Trees of Sonora, Mexico. New
York: Oxford Univ. Press. 391pp.
Felger, R. S. and E. Joyal. 1999. The palms (Arecaceae) of Sonora, Mexico. Aliso 18:1-
Fruhstorfer, H. 1912. 5 Familie: Brassolidae. Pp. 285-332 in A. Seitz (ed.), Die Gross-
Schmetterlinge der Erde, vol. 5. Stuttgart: Alfred Kernen.
Goldman, E. A. and R. T. Moore. 1945. The biotic provinces of Mexico. J. Mammal.
Harrison, J. O. 1963. On the biology of three banana pests in Costa Rica (Lepidoptera:
Limacodidae, Nymphalidae). Ann. Ent. Soc. Amer. 56:87-94.
Marshall, C. J. and J. K. Leibherr. 2000. Cladistic biogeography of the Mexican
Transition Zone. J. Biogeogr. 27:203-216.
Martinez-Yrizar, A., A. Burquez and M. Maass. 2000. Structure and functioning of
tropical deciduous forest in western Mexico. Pp. 19-35 in R. H. Robichaux and D.
R. Yetman (eds.), The Tropical Deciduous Forest of Alamos: F,...i ;... I, of a
Threatened Ecosystem in Mexico. Tucson: The University of Arizona Press.
Morrone, J. J. 2001. E ;.. .... ,(.i, de America Latina y el Caribe. M & T Manuales y
Tesis, Vol. 3. Zaragoza, Spain: Soc. Entomol. Aragonesa.
Morrone, J. J., D. Espinosa and J. Llorente. 2002. Mexican biogeographic provinces:
preliminary scheme, general characterizations, and synonymies. Acta Zool. Mex.
Penz, C. M. 2007. Evaluating the monophyly and phl1..i.-cicii relationships of
Brassolini genera (Lepidoptera, Nymphalidae). Syst. Ent. 32:668-689.
Penz, C. M., A. Aiello and R. B. Srygley. 1999. Early stages of Caligo illioneus and C.
idomeneus (Nymphalidae, Brassolinae) from Panama, with remarks on larval food
plants for the subfamily. J. Lepid. Soc. 53:142-152.
Srygley, R. B. and C. M. Penz. 1999. Lekking in neotropical owl butterflies, Caligo
ilioneus and C. oileus (Lepidoptera: Brassolinae). J. Ins. Behav. 12:81-103.
Stichel, H. 1902. Aufteilung der Gattung Opsiphanes Westw., Ic-l'.!c-'l.i,, neuer
Brassoliden und synonymische Notizen. Berl. Ent. Zeit. 46:487-524.

Stichel, H. 1904. Lepidoptera, Rhopalocera, Fam. Nymphalidae, Subfam. Brassolinae.
Genera Insectorum 20:1-48.
Vargas-Femandez, I., J. Llorente-Bousquets, A. Luis-Martinez and C. Pozo. (in press).
Nymphalidae de Mexico II i',,i........ Ithomiinae, Morphinae, Charaxinae):
Distribucidn ,.. .. .,.-.., e Ilustracidn. M6xico, D.F.: Facultad de Ciencias, UNAM.
Warren, A. D., J. E. Llorente-Bousquets, A. Luis-Martinez and I. Vargas-Femandez.
2006. Interactive Listing of Mexican E ... i;, ..
http://www.mariposasmexicanas.com/ (ace. 16 Feb. 2007)
Young, A. M. 1977. Notes on the defoliation of coconut palm (Cocos .... .... by the
butterfly Opsiphanes quiteria quirinus in northeastern Costa Rica (Lepidoptera:
Brassolidae). Dtsch. Ent. Z. 24:353-365.
Young, A. M. and A. Muyshondt. 1975. Studies on the natural history of Central
American butterflies in the family cluster Satyridae-Brassolidae-Morphidae
(Lepidoptera: Nymphaloidea). III. Opsiphanes tamarindi and Opsiphanes cassina in
Costa Rica and El Salvador. Studies Neotrop. Fauna 10:19-56.

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