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The Journal of Research

ON THE LEPIDOPTERA

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Evolution of locomotion i
Zygaenoidea: Limacodid

Journal of Research on the Lepidoptera

H ® S4: 1 - 1 3 , 1 995 ( 1 997 )

JUN 2 9 1998 )

Marc E. Epstein '

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560

Abstract. Larval locomotion of species in the limacodid-group families
Limacodidae, Dalceridae, Megalopygidae, Aididae, and Somabrachyidae
is described in phylogenetic context. Function of structures involved in
locomotion reported include thoracic legs, abdominal prolegs or suck-
ers, and spinnerets. Additional segments with prolegs or suckers in the
limacodid-group families increase their ventral surface contact with the
substrate. The limacodid + dalcerid clade has the most fluid waves of lo-
comotion because of a highly flexible ventrum, tactile lateral setae and
size reduction of prolegs and thoracic legs. On flat surfaces aidids have a
similar locomotion to limacodids due to short prolegs and smooth lat-
eral and subventral warts, which contact the substrate, whereas in
megalopygids the motion of each proleg segment is more apparent be-
cause contact of the substrate is primarily with membranous pads on their
prolegs. Ventral adhesion in the limacodid + dalcerid clade is increased
by the spinneret both in laying down wet silk and in cleaning debris off
the ventrum. Evolution of locomotion and its adaptive significance in the
limacodid group are discussed.

Keywords. Limacodid group, larval locomotion, prolegs, crochets, suck-
ers, spinneret, silk, smooth hostplants

Introduction

External feeding larvae in the moth family Limacodidae are often referred
to as “slug caterpillars” because their sticky ventrum and locomotion are
superficially similar to those of slugs. Dyar (1899:69) referred to the wave-
like motion of their ventral abdominal segments during locomotion as “the
creeping disk.” Hinton (1955:516) noted that when limacodids crawl, “a
liquid is secreted over the cuticle... if not sticky... may function... by increas-
ing the efficiency of the suckers or by surface tension binding the abdo-
men to the leaf surface.” Epstein (1996) found semifluid silk to be a source
of this liquid.

Limacodidae is part of a monophyletic assemblage that includes Mega-
lopygidae, Dalceridae, Aididae, and Somabrachyidae that is referred to as
the limacodid group. A summary of relationships of the limacodid group,
based on cladistic analysis found in Epstein (1996) is as follows: megalopygid
subfamilies Trosiinae and Megalopyginae form a clade at the base of the
limacodid group, and Aididae, often considered a subfamily of Mega-

Paper submitted 25 March 1996; revised manuscript accepted LS June 1996.

/. Res. Lepid.

lopygidae, is a family and sister group to the Limacodidae + Dalceridae.
Somabrachyidae are thought to be a sister group of Megalopygidae, Aididae
or to the remaining families in the limacodid group.

Larv ae of each family in the limacodid group have prolegs on the second
and seventh abdominal segments (A2 and A7), unique in Lepidoptera fami-
lies with external feeding caterpillars. Megalopygidae are the only family
in the group with species that possess well developed membranous pads on
proleg segments A2-7. Dalcerids and limacodids have ventral abdominal
suckers that are considered to be derived from proleg bases on A2-7, and
from warts on A1 and A8 (Epstein 1996).

In this study I present observations on the locomotion found in all
limacodid-group families. This is followed by discussion of phylogenetic
trends in locomotion in these families as they relate to plants and defense.

Materials and Methods

Locomotion was ohseiwed with larvae of a variety of instars crawling at all angles
on clear glass or plastic, and on wires or stems. Laiwae were filmed using a 16 mm
movie camera with a macro lens or videotaped, either directly or through a micro-
scope, using an 8 mm camcorder. For laiwae crawling on glass, locomotion of the
ventrum was recorded from below by using an inverted phase-contrast microscope.
Species observed are included in Table 1.

Locomotion

Caterpillars crawl by serial muscle contractions surrounding a fluid skel-
eton (Casey 1991). Fonv^ard motion begins as the anal prolegs, or claspers
(AlO), are lifted and planted, and continues sequentially with each segment
by contraction of dorsal longitudinal muscles of the segment to the ante-
rior; this lifts several trailing segments while dorsoventral muscles retract
the prolegs. The prolegs are then set down, beyond their original position,
by a contraction of the ventral longitudinal muscle of its segment (Hughes
Sc Mill 1974). Whether in motion or at rest, the amount of ventral surface
contact with the substrate of free-feeding caterpillars can be viewed as a

Table 1 . Larvae of species in the limacodid group on which observations of
locomotion are based in this study.

Limacodidae:

Dalceridae:

Megalopygidae:

Aididae:

Somabrachyidae:

Phobetron pithecium (Abbott Sc Smith), Isochaetes
beutenmuelleri (Hy. Edwards), Tortricidea pallida (H.-S.),
Snnyra coarctata complex, Prolmiacodes badia (Huebner),
Isa textula (H.-S.), Acharia stimulea (Clemens), Parasa
indetermina (Bdv.), Euclea delphmii (Bdv.)

Dalcerides ingenita (Hy. Edwards)

Megalopyge sp. from Belize, Megalopyge crispata (Pack-
ard), M. basalts (Walker), Norape cretala (Grote)

Aidos amanda (Stoll)

P.syrharium sp.

Table 2. Morphological characters of the limacodid group and Zygaenidae that relate to locomotion (from Epstein 1996).

34:1-=13, 1995(1997)

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Limacodidae bmshlike, often broad small to suckers Al-8, crochets crochets usually " lateral setae

at apex (often narrowing minute usually absent absent tactile, below

after first instars) spiracle

J. Res. Lepid.

Figs. 1-3. Use of silk in larvae of Somabrachyidae, Aididae, and Megalopygidae.
1) Haphazard laying down of silk of a first instar Psycharium sp.
(Somabrachyidae), viewed from beneath through glass; 2) Middle instar
Aidos amanda clinging to glass, viewed from above through glass (photo
courtesy of Max and Eileen Price); 3) Figure-8 silk on glass from
Megalopyge sp. (photo by Kjell Sandved) (scale is in mm).

continuum. Limacodids and dalcerids are at one extreme, with most of the
ventrum minus the AlO segment in contact, while the condition commonly
found in geometrids, with only thoracic legs and A6 and AlO prolegs in
contact, is at the other extreme. Other free-feeding caterpillars fall in be-
tween by having a maximum of five prolegs in contact.

Caterpillar locomotion involves a complex of structures, behaviors, and
positions in relation to the contact surface. Morphological characters re-
lating to locomotion in each of the limacodid-group families is given in Table
2. These include external aspects of the spinneret, thoracic legs, abdomi-
nal prolegs or suckers, anal prolegs, the texture of the ventral surface, and
lateral structures.

The descriptions of locomotion in the limacodid-group families are or-
dered from plesiomorphic to derived taxa based on the phylogeny from
Epstein (1996). Somabrachyidae, of uncertain relationship to the other fami-
lies, is placed after the Megalopygidae. The descriptions include informa-
tion from these categories: 1) locomotion on narrow surfaces; 2) locomo-
tion on flat surfaces; 3) use of silk and the spinneret; 4) feeding and rest-

34:1-13, 1995(1997)

ing positions. For the families for which I have the most information, Lima-
codidae and Megalopygidae, I will use these categories as subheadings.

Megalopygidae. 1 ) Narrow surfaces:. Larvae use thoracic legs and all pro-
legs, including the anal pair, to grasp (Fig. 4). 2) Flat surfaces (Fig. 5): Mem-
branous pads (Fig. 6) are the primary contact of the prolegs on A2-7 (as
indicated through glass) ; a pair of tactile, subventral setae positioned at the
anterior margin of each pad touches the substrate. Lateral and subventral
verrucae have little contact with the substrate except in certain instances
(e.g., Mesoscia pusilla) . The anterior, non-crochet portion of the anal pro-
legs contacts substrate at the beginning of a wave of locomotion, while at
least two adjoining proleg pairs are retracted (and compressed) from the
substrate as the wave progresses. 3) Silk: Early instars spin silk on horizon-
tal surfaces in a linear or haphazard manner (as in Somabrachyidae, Fig.

1 ) , and can dangle from silk, while later instars issue silk on the substrate
in a figure-8 (Fig. 3), especially when they are at an angle of > 45 degrees.
4) Feeding and resting. Larvae cling to silk they deposit on the substrate with
their thoracic legs and crochets.

Discussion: Packard (1893) noted that the prolegs on A2 and A7 in
Megalopyge {Lagoa) crispata functioned like the others, even though they
lacked the crochets. Dyar (1899) observed that the membranous pads
(=disks) of Megalopyge opercularis were in exclusive contact with a smooth
glass surface during locomotion. Some species of megalopygids have sucker-
like pads below subventral verrucae in addition to the membranous pads
on the prolegs (e.g., Mesoscia pusilla; Fig. 6). These presumably contact the
substrate much the same way as the proleg pads. The size of the membra-
nous pads relative to the prolegs varies between species, and on A2 and A7
depending on whether crochets are present or absent (Epstein 1996).

Somabrachyidae {Psycharium sp., first and second instars, only). These
larvae prefer to crawl on narrow surfaces such as found on its hostplants
(Restionaceae and Pinus, H. Geertsema pers. comm.). When viewed from
underneath on a glass surface the prolegs sometimes clasped together, as
if to grip a narrow substrate, rather than push directly on it. Silk is depos-
ited in the same, haphazard way as in early instar megalopygids (Eig. 1).

Aididae. Movement on flat surfaces of proleg segments has a fluid wave-
like appearance similar to limacodid and dalcerid species, because they
closely contact the substrate with short, broad prolegs and smooth lateral
and subventral warts below the spiracles (Fig. 2). Most megalopygids, in
contrast, have only the membranous pads contact flat surfaces, whereas the
proleg base and crochets and the spiny and plumose setae on lateral and
subventral verrucae have less contact with flat surfaces. The major differ-
ence between locomotion found in Aidos amanda and in limacodid and
dalcerid species relates to the presence of a flexible ventral cuticle found
in the latter two families. Larvae of A. amanda are difficult to dislodge at
rest because they have a large number of crochets hooked onto silk (Fig.

2) (Epstein 1996); they issue silk while crawling onto a leaf or on smooth

J. Res. Lepid.

34:1-13, 1995(1997)

Figs. 4-6. Locomotion and ventral surface of larval Megalopygidae. 4-5: Late in-
star Megalopyge basalis (head on right end) (from 16 mm film by Kjell
Sandved). 4) Clasping a wire with prolegs and thoracic legs; 5) Viewed
through a horizontal piece of plexiglass; 6) Scanning electron micrograph
of abdominal prolegs and subventral pads on A2 (top) to A3 of Mesoscia
pusilla (scale bar = 0.5 mm).

surfaces, as in species of megalopygids and caterpillars in other families
(Epstein 1995).

Limacodidae. 1) narrow surfaces: Larvae ventrally grasp leaf edges, stems,
or narrow vines along the midline (Figs. 11, 19), or from anterior to poste-
rior (Fig. 10); anal prolegs do not grasp. Sticky silk can also help proximal
abdominal segments stick together while wrapping around a stem. Reduced
thoracic legs have tactile function, while the pretarsal claw assists in grasp-
ing a leaf or petiole or in clutching silk applied to the posterior ventrum.
2) flat surfaces: The locomotory surface, which consists of a highly flexible
cuticle with fungiform tactile setae, moves in fluid waves that expand later-
ally along the leading edge, progressively retracting from the substrate (Figs.
16, 17). Tactile lateral setae, located between the margins of the ventral
surface and the spiracles, contact the substrate during locomotion. Waves
can move in oblique angles when the larva shifts its head and thorax to ei-
ther side. Larvae readily reverse motion either in straight or oblique waves.
The vestigial anal prolegs, spinulose with less elastic cuticle, are raised off
the substrate during locomotion. This assists in expelling frass while in
motion or while feeding. 3) silk: Semifluid silk, or a fluid along with the
silk, is laid down in hgure-8 fashion by the spinneret on substrates during
or prior to locomotion. The fluid can spread from the thoracic region to
the 9th abdominal segment and aids in the adhesion of the suckers. The
silk is also applied directly to the anterior ventrum by rearing up the head
or to the entire ventrum on narrow surfaces when clasping from anterior
to posterior. The apparently sticky silk strands on the ventrum help the
suckers grip to flat or narrow substrates (Fig. 10). Unlike megalopygids,
aidids and numerous other lepidopterans, the larvae often do not leave
discrete strands of silk behind until the onset of cocoon construction. 4)
feeding and resting. Larvae have heads retracted beneath the prothorax while
the ventrum is laterally expanded and is suckered down to the substrate
(assisted by surface film) (Fig. 18) . When the ventral surface gets dirty, lar-
vae will raise their anterior off the substrate and brush the thorax and first
few abdominal segments with their spinneret from side to side.

Discussion: The tight adhesion of the ventrum to the substrate, during
locomotion or at rest, requires only small amounts of the liquid silk to pro-
vide surface tension. Use of scanning electron microscopy revealed no pores
for fluid secretion on the ventral and lateral surfaces (Epstein 1996) , as sug-
gested by Holloway (1986). The presence of a liquid silk droplet was ob-
served in Prolimacodes badia during egg eclosion (Figs. 7-9), although not
during this stage in other species (e.g., Isa textula, Tortricidia pallida) . The

/. Res. Lepid.

Figs. 7-13. Use of semifluid silk in larval Limacodidae (from 8 mm video). 7) Egg
eclosion of ProUmacodes badia; 8, 9) Detail of silk droplet on end of spin-
neret during egg eclosion of P. badia (arrows point to droplet); 10) Late
instar larva of Tortriddia pallida clasping stem from anterior to posterior
while applying silk to posteroventral segments; 1 1 ) Silk strand of T. pallida
at the end of a twig (arrow points to silk strand) (note medial clasping of
ventral surface on the left); 12) Detail of spinneret and ventral thorax of
late instar P. badia (viewed through clear plastic from above); 13) Late
instar T. pallida obliquely grasping stem while applying fluid silk to it.

34:1=13, 1995(1997)

Figs. 14-15. Spinnerets of first and last instar Limacodidae in related genera
Prolimacodes and Semyra (scale bar length in parentheses). 14) First
instar Prolimacodes badia with silk debris on distal margin (50 pm);
15) Last instar Semyra coarctata complex (27 pm).

J. Res. Lepid.

Figs. 16-21 . Locomotion in larval Limacodidae and Dalceridae (viewed from above
through glass). 1 6-1 9: Semyra coarctata complex (photos by Chip Clark).
1 6) Near middle of locomotion sequence; 1 7) At the end of a locomotion
sequence: 18) Larva at rest with ventrum laterally expanded and head
retracted; 19) Lateroventral view of larva medially clasping the edge of
the glass. 20, 21 : Dalcerides ingenita (photos by Laurie Minor-Penland).
20) Near middle of locomotion sequence; 21 ) At the end of a locomotion
sequence.

34:1-13, 1995(1997)

Zygaenidae Megalopygidae Aididae Dalceridae Limacodidae

Fig. 22. Evolution of the ventral surface in the limacodid-group families viewed
in cross section of proleg segment (after Epstein 1996).

spinneret, whether broad at the apex in early instars (Fig. 14) or narrow in
late instars (Figs. 12, 15), functions similarly in both the use of the silk and
in cleaning the ventral surface. Although the prolegs are highly reduced,
their gripping of narrow substrates appears to be aided by a dense pad of
ventral muscles revealed by dissection.

Dalceridae. Locomotion (Stehr Sc McFarland 1985; Figs. 20, 21) and spin-
neret function in dalcerid larvae are similar to those found in limacodids.
Tactile lateral setae used to touch the substrate during locomotion are
shifted dorsad, above or near the spiracles, compared to those in limacodids.
This probably is due to the relative closeness of spiracles to the locomotory
surface. Semifluid silk was observed in Dalcerides ingenita, though not at egg
eclosion. Reports of a “shiny path” trailing behind dalcerid larvae (Genty
et al. 1978) may have been from broad ribbons of silk laid on the substrate
or the result of cleaning with the spinnerets. Fluid debris has been observed
following “brushing” on the anteroventral abdominal segments. Small cro-
chets, present only in mid- to late instars (Stehr Sc McFarland 1985, Epstein
1996), appear not to have any function on smooth leaf surfaces. However,
they may be used by the prepupa when crawling inside the diffuse cocoon.

Evolution of Locomotion in the Limacodid Group

The most noticeable trend when viewing locomotion of the limacodid
group in phylogenetic sequence is the increased proportion of the ventral
surface in direct contact with the substrate (Fig. 22). In megalopygids, at
the base of the limacodid group, this is suggested by the shortness of pro-
legs relative to presumed zygaenid ancestors and the addition of prolegs
on A2 and A7. Moving from megalopygids to aidids, contact increases as a
result of the reduction of lateral and subventral verrucae to smooth warts.
In the limacodid + dalcerid clade, further contact results from reduction

/. Res. Lepid.

of the prolegs on A2-7 to suckers, formation of suckers on A1 and A8, and
the flexibility to the ventral cuticle (Epstein 1996) . The relatively large tho-
racic legs and grasping of narrow surfaces in Somabrachyidae, in addition
to other plesiomorphies (Epstein 1996), suggest that this family may be a
primitive lineage of the limacodid group.

The increased contact of the larval ventral surface suggests a specializa-
tion toward smoother host plants. Features of megalopygids that are effec-
tive in clinging to smooth surfaces include the membranous pads on the
prolegs, and presumably, in some species, pads on subventral verrucae. The
presence of smooth subventral and lateral warts in aidids, which contact
the substrate during locomotion (versus setose verrucae), are also indica-
tive of this type of host plant specialization. Absences of crochets on A2 and
A7 proleg segments in many megalopygids and in aidids, apparently inde-
pendent losses (Epstein 1996), are also suggestive of this trend. The smooth
and flexible ventrum in the limacodid + dalcerid clade, in conjunction with
the spinneret and silk, assists in sticking to smooth host plant surfaces (see
further discussion below) .

Species in the limacodid group are often polyphagous, with the ability to
switch host plants even in later instars (Dyar 1905, 1909, Epstein 1995).
Perhaps predator selection influenced evolution in this direction, since a
combination of slow growth (found throughout the group) and increased
foraging time in seeking a specific host plant could lead to heavy losses of
larval populations from parasitoids and predators. Switching to other host
plant species with similar smooth-leaf textures and suitable chemical make-
ups would theoretically decrease foraging time.

Caterpillar adaptations to predators have been thought to relate to de-
fenses, such as group feeding in spiny caterpillars or crypsis, with locomo-
tion not playing a role (Casey 1991). The majority of caterpillars of the
limacodid group employ these defenses against predators and parasitoids.
However, in limacodids and dalcerids especially, ventral adhesion to the
hostplant and locomotion are so closely linked that locomotion can indeed
be considered an adaptive strategy to avoid predation. Species in the two
families show a marked specialization for cryptic behavior in their ability to
crawl beneath smooth plant surfaces. This is further enhanced by having
less visible mouthparts at leaf edges due to retractile feeding beneath the
thorax, as in other members of the limacodid group (Epstein 1996). These
larvae may also be less easily detected by parasitoids from their less appar-
ent silk trails, perhaps gaining in survival from cryptic silk-use what they
lose in not having the ability to dangle on silk to reach new host plants.

Acknowledgments. I wish to acknowledge the assistance of I^ell B. Sandved (Smithson-
ian Inst.) for 16mm filming andjeff Norris (Univ. Florida) for video of Aidos amanda.
Laurie Minor-Penland and Carl Hansen (Smithsonian Inst.) provided both support
in photography and scanning slide images and making plates in Adobe Photoshop,
and Dane Penland (Smithsonian Inst.) taught me how to use a frame-grabber to
capture video images. Chip Clark (Smithsonian Inst.), and Max and Eileen Price

34:1=^13, 1995(1997)

(Denver, Colo.) provided additional photographic support. Scott E. Miller (Bishop
Museum) reviewed an early draft of the manuscript. Michael Ma and Joanne
Ballerino (Univ. Maryland) graciously allowed me to use their inverted compound
microscope. I had beneficial discussions about caterpillar locomotion with John E.
Rawlins (Carnegie Museum of Nat. Hist.) and John Dodge (Annandale, Virginia).
I was assisted in the use of the scanning electron microscope by Susanne Braden,
Walter Brown and Victoria Godwin (SEM Laboratory, Smithsonian Inst.). Noel
McFarland (Sierra Vista, Ariz.) and Hendrik Geertsema (Univ. Stellenbosch) pro-
vided eggs that became the dalcerid and somabrachyid larvae used in the study.
Thanks also go to Don R. Davis and Ronald McGinley (Smithsonian Inst.), and
Jerome C. Regier (Univ. Maryland) for their interest and support.

Literature Cited

Casey, T.M. 1991. Energetics of caterpillar locomotion: biomechanical constraints
of a hydraulic skeleton. Science 252:112"”113.

Dyar, H.G. 1899. Note on the secondary abdominal legs in the Megalopyg-
idae. Journal of the New York Entomological Society 7:69-70, pi. 2.

— . 1905. The life-history of a cochlidian moth — Adoneta bicaudata Dyar.
Biological Studies by the pupils of William Thompson Sedgwick. Chicago, pp.
11-18.

. 1909. Description of a new species of Euclea\nth its larva [Lepidoptera, Coch-
lidiidae]. Proceedings of the Entomological Society of Washington 1 1:156-158.
Epstein, M.E. 1995. False-parasitized cocoons and the biology of Aididae (Lepi-
doptera: Zygaenoidea) . Proceedings of the Entomological Society of Washington

97:750-756.

“ — 1996. Revision and phylogeny of the limacodid-group families, with evolu-
tionary studies on slug caterpillars (Lepidoptera: Zygaenoidea). Smithsonian
Contributions to Zoology 582. 101 pp., 409 figs.

Genty, P., R. Desmier,J.P. Morin & C.A. Korytkowski. 1978. Les ravaguers du palmier
a huile en Amerique Latine. Oleagineux 33:325-419.

Hinton, H.E. 1955. On the structure, function, and distribution of the prolegs of
the Panorpoidea, with a criticism of the Berlese-Imms theory. Transactions of
the Royal Entomological Society of London 106:455-540.

Hughes, G.M. & P.J. Mill. 1974. Locomotion: terrestrial. Pp. 335-379 in M.

Rockstein, ed. The Physiology of Insecta, Vol. 3. Academic Press, New York.
Packard, A.S. 1894. A study of the transformations and anatomy of Lagoa crispata, a
Bombycine moth. Proceedings of the American Philosophical Society 32:275-
292, plates 1-7.

Stehr, F.W. & N. McFarland. 1985. Crochets on abdominal segments 2 and 7 of
dalcerid caterpillars: “missing link” or anomaly? Bulletin of the Entomological
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Journal of Research on the Lepidoptera

34:14-20, 1995(1997)

Territoriality by the dawn’s early light: the Neotropical owl
butterfly Caligo idomenaeus (Nymphahdae: Brassolinae)

Andre V. L. Freitas, Woodruff W. Benson, Onildo J. MarinFFilho, and
Roberta M. de Carvalho

Curso de Pos-Graduagao em Ecologia, Institute de Biologia, Universidade Estadual de
Campinas, C. P.6109, 13083-970 Campinas, Sao Paulo, Brazil

Abstract. Males of the Neotropical owl butterfly Caligo idomenaeus defend
unusual dawn territories along dirt roads in the Linhares Forest Reserve,
Espfrito Santo, Brazil. The territories are notable for their wide spacing
and the brief period in which owners are present. During mid-winter in-
sects arrived on the territories shortly after 0550 h, as the last bright stars
disappeared from the sky, and remained approximately 15 min before
flying back into the forest. Dawn perching seemed unaffected by substrate
temperatures as low as 12.5° C. Perches were about 100 m apart and resi-
dent butterflies returned to and seemingly repelled invaders from their
territories on consecutive mornings. Territories contained no material
resources. The brief dawn occupancy may be related to the activity pe-
riod of receptive females and to predator risk in these large, palatable
insects.

Keywords. Brassolinae, Brazil, Caligo, crepuscular, mating behavior, ter-
ritoriality

Introduction

Male defense of encounter sites is a common mate-locating tactic in
butterflies (Baker 1983, Rutowski 1991). Mating territories have been re-
corded for many taxa and geographic regions, but are especially well docu-
mented for temperate zone species, especially papilionids (Lederhouse,
1982), lycaenids (Douwes 1975, Alcock 1983a, Alcock Sc O’Neill 1986) and
the nymphalid subfamilies Nymphalinae (Baker 1972, Bitzer & Shaw 1980,
1983, Alcock 1983b, Alcock & Gwynne 1988, Rosenberg Sc Enquist 1991)
and Satyrinae (Davies 1978, Knapton 1985, Wickman 1985). The few tropi-
cal studies to date have reported territorial behavior in typical tropical taxa
(Riodininae, Alcock 1988; Heliconiinae, Benson et ah 1989) as well as in
taxa already studied in temperate areas (Papilionidae, Pinheiro 1990;
Nymphalinae, Rutowski 1991b, 1992, Lederhouse et al, 1992).

Independent of region, territorial behavior, like flight activity in general
(Srygley Sc Chai 1990), is characteristic of sunny habitats with mild thermal
environments (Alcock 1983b, Wickman 1985a, Alcock Sc O’Neill 1986) . This
rule is not universal, and several species of Vanessa defend near sundown
(Alcock Sc Gwynne 1988, Brown Sc Alcock 1991). The exclusively Neotropi-

Paper submitted 2 June 1994; revised manuscript accepted 24 October 1996.

34:14-20, 1995(1997)

cal Brassolinae may provide other exceptions. In Panama, Caligo memnon
Felder engages in territorialdike mating behavior at dusk, and Opsiphanes
cassina (Brassolinae) behaves similarly (Srygley 1994). In the state

of Espirito Santo in southeastern Brazil Caligo Cramer and Catoblepia

amphiwhoellxxhnQT perch along roadsides at dusk (W.W. Benson, pers. obs.) ,
whereas Caligo idomenaeus rhoetus Staudinger does this shortly before dawn,
even during cool winter weather. In the winter of 1992 we studied C.
idomenaeus with the intent of clarifying the significance of dawn perching
behavior in this insect and gain insights into the possible influence of light
and temperature.

Methods

The study was carried out from 27.VII to 6.VIII.1992 (no observations were made
on 4.VIII) along an east-west stretch of 4 m wide dirt road passing through mature
subtropical moist forest in the Linhares Forest Reserve (Reserva Florestal de
Linhares) at Linhares, Espirito Santo, Brazil (19° 10' S, 40° 00' W). Mean winter
temperatures at the reserve are near 20° C, with extremes for the months of July
and August (mid-winter) approximately 10° and 30° C (Companhia Vale do Rio
Doce, unpub. data).

The study area was selected based on the confirmed presence of Caligo idomenaeus.
A 450 m long area was marked off in 50 m segments to aid mapping of butterfly
perches and behavioral events. We conducted observations daily from about 0545
to 0620 h (early dawn to shortly after sunrise) . The owl butterflies were very difficult
to see in the dim light at the beginning of their activity period, especially where
trees arched over the road, and we were only able to follow insects at this time by
spacing ourselves along the road and calling to each other as butterflies passed.
When possible, butterflies were netted and marked by cutting distinguishing notches
along wing margins. The owl butterflies are large and robust and apparently not
impaired by this procedure. Some uncaptured individuals could be individually
recognized by distinctive wing damage. Road-surface temperature was measured
daily at the beginning of butterfly activity using a mercury thermometer. On one
morning during the Caligo idomenaeus activity period we measured incident light at
road level near the perch site at the widest and least obstructed part of the road
using a digital luximeter with 1 lux sensitivity (Extech Instruments). Civil twilight
period and sunrise were obtained from the computer program Earthsun 4.5 (© W.
Scott Thoman, Dryden, NY, 1995).

Results

The only large owl butterfly observed at dawn was Caligo idomenaeus. We
observed three of the four individually recognizable insects on more than
one day. Two individuals marked on perches at dawn were males, and oth-
ers observed in the study area were inferred to be males by their behavior.
Other Caligo captured during twilight hours while perched along reserve
roadways have always proved to be males of C. illioneus and C. idomenaeus
(W.W. Benson, pers. obs.).

J. Res. Lepid.

On most mornings we observed 3-5 Caligo perching in the area and as
many as four non-residents making “fly-throughs.” Perching Caligo were
punctual, with the first individual arriving between 0550 and 0556 h(x =
0552.7 h, s.d. = 2.3 min , n = 7; six observations on marked individual #3)
and the last Caligo departing around the time of sunrise between 0609 and
0612 h (x = 0610.1 h, s.d. = 0.9 min, n = 8; seven observations on individual
#3). Between 27.VII and 2.VIII, when the bulk of the observations were
made, civil twilight began between 0547 and 0545 h and the sun rose be-
tween 0609 and 0607 h. Brighter stars remained visible until about 0550 h,
and the planet Venus could be seen until 0553 h.

Butterflies occasionally arrived in the area and patrolled back and forth
as much as 3 min before perching. The time span over which one or more
individuals were present in the area on a given morning varied between 13
and 20 min (x = 16.7 min, s.d. = 2.5 min, n = 6). The time of arrival and
departure did not seem to be strongly influenced by cloud conditions (clear
to cloudy) or temperature (12.5-18.0° C), and even with a soil tempera-
ture of 12.5° C, four butterflies were active. On 2.VIII under an almost cloud-
less sky, the light intensity increased approximately exponentially from about
2 lux at 0551 h when the first Caligo arrived to 180 lux when the last one
departed at 0610 h.

When arriving in the area, an owl butterfly often patrolled back and forth
several (maximum of nine) times along 10-50 m of road before landing.
The flight was swift and erratic about 1-2 m above the ground. After ar-
rival, butterflies perched near the center of the patrolled area on the ground
in the roadway or on low (< 1 m high) roadside vegetation. Although most
arriving (and departing) butterflies that we followed left (or entered) the
forest within 25 m of the perch, one was observed to fly approximately 240
m before entering the forest.

Interactions occurred when flying butterflies met or a presumed male was
chased when it flew over a perched resident. Interacting owl butterflies flew
in tight circles about each other in level or ascending flight approximately
1-2 m (up to 5 m) above the ground while batting their wings together.
Most interactions terminated after a few 10s of seconds. In six of the seven
observed encounters involving marked resident males, the original male
returned to its perch after the intruder had left. In the remaining case ob-
served near the end of the territorial period neither butterfly returned. On
two occasions, two C. idomenaeus were observed to perch 10-20 m apart,
apparently without seeing one another.

During nine days of observations we identified five sites preferred by Caligo
idomenaeus ioY perching: 10 m (used on 2 d), 100/120 m (3 d), 160 m (7
d), 220/280 m (3d), and 380 m (9 d) from the west boundary of the study
area. The three marked males that returned to the area showed perch
fidelity on successive days. On five consecutive days (27-31. VII) male #1
landed at the 160 m perch (and once at the 10 m perch as it was leaving the
area). Unmarked individual (s) occupied this perch on the two days follow-
ing the disappearance of #1. Male #2, seen in the area 27-28.VII, occupied

34:14-20, 1995(1997)

perches at 100 and 120 m, from which it was expelled by male #1. Male #3
perched in the road at 380 m on seven consecutive days (28.VII~-3.VIII) and
subsequently on 5- 6.VIIL Males #1 and #3 usually rested in the road near
fallen, dead Cecropia leaves. Otherwise, there was no indication that the
butterflies selected perching sites with respect to specific habitat features.

Discussion

Patrolling behavior, interactions between individuals and spacing in male
Caligo idomenaeus are almost certainly related to territorial defense. Indi-
viduals returned daily to specific perches spaced about 100 m apart. These
residents seemed to patrol road segments around their perches and inter-
act by expelling intruders. Although we saw neither courtship nor mating,
defense of mating territories is common in butterflies (see Introduction),
and other Caligo mate during crepuscular encounters (Srygley 1994). Al-
though only three individually recognizible butterflies were monitored, we
believe that our observations on these are representative of the study popu-
lation. On the other hand, our unsuccessful attempts to capture unmarked
individuals may have frightened some butterflies from the area and dimin-
ished perch occupation.

Territorial Caligo idomenaeus p2Liro\ corridors up to 50 m long around their
perches. In contrast, territorial Heliconius p^irol corridors about 15 m long
(Benson et al. 1989), and similar territory sizes seem to exist in tropical
Heraclides 3.nd Battus (Pinheiro 1990). C. idomenaeus is large for a butterfly
(wing length 80 mm), and for this reason territory size may be less con-
strained than in smaller species. The wide spacing between perches may be
advantageous in reducing mate competition between neighbors. Although
C. idomenaeus occurs spottily along roadsides in the Linhares Forest Reserve,
males do not seem to lek around conspicuous landmarks as has sometimes
been reported for other butterflies (DeVries 1980, Lederhouse 1982, Alcock
1983a, Knapton 1985, Alcock & Gwynne 1987) and population distribution
may be more related to habitat favorability than classical lek formation.

Low temperature can prevent butterfly flight, and the ability of C.
idomenaeus to maintain full activity before sunrise with substrate tempera-
tures below 13° C is probably aided by its large size and Caligo' s ability to
increase body temperature by shivering (Srygley, 1994). Our study site is
subtropical with cool winters, and it is interesting to note that C. idomenaeus
was active at temperatures (x = 16.2° C, s.d. = 1.9° C, n = 8) uniformly lower
than Srygley’s (1994) estimate of 19-20° C for the lower critical tempera-
ture for flight in Panamanian C. eurilochus.

Two possibly unique characteristics of territoriality in Caligo idomenaeus
and other brassolids (see Introduction; Srygley 1994) are its occurrence
during twilight hours and the extreme brevity of the territorial bouts. Be-
cause owl butterflies are palatable (Chai 1986) and presumably especially
profitable prey items due to their large body mass and high visibility when
in movement, birds may select strongly against late-flying C. idomenaeus and
thereby constrain activity to situations where their visiblity to predators is

/. Res. Lepid.

hampered. Restricted activity of receptive females could have the same
cause, and same effect on mating conventions.

Published studies suggest that tropical forest butterflies usually spend less
time in territorial defense than butterflies of other environments. Exclud-
ing desert butterflies such as Chlosyne calif ornica (Wright) (Alcock 1983b)
and Strymon melinus Hiibner (Alcock & O’Neill 1986) whose activity is ap-
parently limited by high midday temperatures, and species of the cosmo-
politan genus Vanessa that defend territories just before sundown (Bitzer &
Shaw 1980, 1983, Alcock & Gwynne 1988; Brown & Alcock 1991), 64% of
the 14 temperate-zone butterflies for which data are available typically de-
fend territories 3-6 h daily (Powell 1968, Baker 1972, Douwes 1975, Davies
1978, Wickman & Wicklund 1983, Alcock 1983a, Bitzer & Shaw 1983,
Rutowski & Gilchrist 1988) and an additional 29% defend 6 h or more (see
Lederhouse 1982, Wickman 1985b, Knapton 1985, Rosenberg & Enquist
1991). Similarly long shifts of territorial defense have been reported for
Hypolimnas 3.nd Junoniain tropical savanna (Rutowski 1991b, 1992). Exclud-
ing the desert species and Vanessa mentioned above, Polygonia comma Har-
ris is to our knowledge the only temperate butterfly reported to be territo-
rially active for three or fewer hours a day (Bitzer & Shaw 1983).

The seven tropical forest butterflies for which data have been published
defend territories over relatively shorter time spans, five for 3 h or less daily
and the two remaining for up to 6 h. Besides the 0.25 h period reported
here for C. idomenaeus, the heliconiines Heliconius sara (Fabr.), H. leucadia
(Bates) and Eueides tales (Cramer) defend for 1-2.5 h daily in Brazil, and E.
aliphera (Godart) is territorial for about 5 h daily in Costa Rica (Benson et
ah 1989). Alcock (1988) reports that the Costa Rican forest hesperiids
Celaenorrhinus approximatus William & Bell and Astraptes galesus cassius Evans
defend for about 2 and 3 h respectively, whereas males of the riodine
Mesosemia a. asa Hewitson spend about 4 h per day on territories. Although
each species is no doubt adapted to a unique set of ecological conditions,
we believe the general phenomenon of shorter defense shifts in tropical
forest butteflies may be related to fine-tuning in mate search resulting from
the greater temporal structuring of this environment. However, because of
the small number and limited taxonomic distribution of species studied to
date, and the general lack of information on temporal variation in the costs
and benefits of territorial defense, our purpose here is more to draw atten-
tion to the phenomenon than to provide explanations.

Acknowledgments. Permission to work in the Linhares Forest Reserve and basic lo-
gistic support were provided by the Companhia Vale do Rio Doce and their subsid-
iary Florestas Rio Doce S/A through the reserve manager Renato M. de Jesus. Ad-
ditional financial support came from the Funda^ao MB. AVLF and OJMF received
graduate fellowships from CAPES and RMC from CNPq. The study was carried out
under the auspices of a graduate ecology field course of the Universidade Estadual
de Campinas. We are grateful to Keith S. Brown Jr. and two anonymous reviewers
for their helpful comments on the manuscript We also thank Jose Roberto Trigo

34:14-20, 1995(1997)

for installing the computer program Earthsun 4.5 and K,S. Brown Jr. identifying

the butterflies.

Literature Cited

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— . 1983b. Hilltopping in the nymphalid butterfly Chlosyne californica (Lepi-
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— — 1987. The mating system of three territorial butterflies in Costa Rica. J. Res.
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Alcock, J. 8c D. Gwynne. 1988. The mating system of Vanessa kershawi: males defend
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Baker, R.R. 1972. Territorial behaviour of the nymphalid butterflies, Aglais urticae
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— — . 1983. Territorial behavior of Nymphalis antiopa2Lnd Polygonia comma (Nymph-
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Lederhouse, R.C., S.G. Codella, D.W. Grossmueller & A.D. Maccarone. 1992. Host
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PiNHEiRO, C.E.G. 1990. Territorial hilltopping behavior of three swallowtail butterflies
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/ R£s. Lepid.

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— . 1991b. Temporal and spatial overlap in the matedocting behavior of two
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— — 1992. Male mate-locating behavior in the common eggfly, Hypolimnas bolina
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■ — — . 1985b. Territorial defense and mating success in males of the small heath
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Journal of Research on the Lepidoptera

34:21-38, 1995(1997)

A review of the genus Panara Doubleday, 1847 (Riodinidae)
in southeast Brazil, with a description of two new subspecies

Curtis J. Callaghan

Av. Suba 130-25 Casa 6, Bogota, Colombia

Abstract. The species of the riodinid genus Panara Doubleday, 1847 in
southern Brazil are reviewed and corrections are made in the nomencla-
ture of the the extra-Amazonian species. Two new subspecies, Panara soana
bacana and Panara soana ruschii are described. The characteristics of the
genus Panara are reviewed, and two taxa are removed: Pterographium
elegans (Schaus) , new combination; and Phaenochitonia brevilinea (Schaus) ,
new combination, new synonomy. Separate keys are presented for adult
males and females, as well as comments on the range, adult behavior,
and habitats.

Key Words. Neotropical South America, Panara

Introduction

The genus Panara consists of five species, four of which are confined to
extra-Amazonian Brazil and one distributed throughout the Amazon and
Orinoco drainage. All species have black ground color with a diagonal yel-
low band from the costa to the distal margin, and some of the southern
Brazilian phenotypes also have an orange band on the hindwing from the
apex to the inner margin, a characteristic which varies within species. This
has led to much confusion in the literature (e.g., D’Abrera 1994), such as
the mixing of the species and subspecies as well as the inclusion in the ge-
nus of unrelated riodinid taxa with orange forewing bands. The lack of a
basis for the proper identification of these species has hindered potential
research in the biodiversity of southern Brazil.

The purpose of this review is to 1) present biological and morphological
information on each species of extra-Amazonian Panara, 2) provide a key
to both males and females which will help in their rapid determination, 3)
correct the nomenclature of the extra-Amazonian species of Panara, and
4) review the characteristics of the genus Panara with the removal of those
taxa which do not belong there.

Materials and Methods

During the study I examined numerous museum collections in addition to my
own (CJC), which includes material on loan by Dr. Keith Brown: the Museu
Nacional, Rio de Janeiro (MN); the Universidade Federal de Parana (UFP); the
Smithsonian Institution, Washington, DC (NMNH), the Museum National
d’Histoire Naturelle in Paris (MNHN); the Senckenburg Museum, Frankfurt (SM);

Paper submitted 24 October 1994; revised manuscript accepted 16 November 1995.

J. Res. Lepid.

Fig. 1 . Venation of Panara

the Humboldt Museum, Berlin (MNK); and the Natural History Museum (London)
(BMNH). I examined 334 specimens and made 52 genitalia preparations. Measure-
ments were made with an ocular micrometer and calipers. References to wing cells
and veins follow the Comstock-Needham system in Miller (1969) and the genitalia
terminology follows Klots (1970). In addition, my field trips over a 25-year period
provided data on habitats and adult habits of Panara.

The Genus Panara Doubleday

Harvey (1987) followed Stichel (1910) in pl^Lcing Panara in the Ancyluris
section of the tribe Riodinini. This tribe is characterized by a deeply in-
dented notch in the posterior margin of the tegumen of the male genita-
lia. A true saccus is also absent, the vinculum being ribbon-like ventrally,
and not fused to the valvae.

Within the tribe Riodinini, Panara is related to Lyropteryx Westwood,
[1851], Necryia Westwood, [1851], Cyrenia Westwood, [1851], Ancyluris
Hubner, [1819], NirodiaWes\^NOod, [1851], Swainson, [1829],

Chorinea Grdiy, 1832, NahidaYdrhy, 1871, and Bates, 1862, all of which

have 1) a normal strap-like pedicel in the male genitalia, 2) forewing vein
R2 originating beyond the end of the cell, stalked with R3 and R4, and 3)
the ostium bursa located in the middle of the ventral surface of the abdo-
men, not displaced to the right as in the Riodina section of the tribe.

Panara may be separated from related genera by 1 ) the black ground color
of the wings and the transverse orange band on the forewing, 2) the male
genitalia which are broad and triangular shaped laterally, 3) the wing vena-
tion. At the end of the forewing cell, M2-M3 forms a junction with M3 and

34:21-38, 1995(1997)

GUI, whereas in related genera this junction is considerably more basad of
the cell (Fig. 1).

The genitalia of the southern Brazilian Panam species show considerable
individual variation which makes classification on this basis tenuous. The
most constant character in the male genitalia is the height of the valvae
relative to the transtilla. However, I found an individual of Panara iarbas
with the left valva higher and the right valva lower.

Synonyms and New Combinations

The last revisor of the genus Pawarawas Stichel (1930) who defined six
species and five subspecies, as follows:

P. phereclus (Linne, 1758)

^)barsacus Westwood, [1851]
h)elegans Schdius, 1920

c) episatnius Prittwitz, 1865

Hewitson, 1865
=arctifascia Sutler, 1874

d) lemniscataThieme, 1907

= comes Stichel, 1909
P. aureizona Antler, 1874

=ornata Stichel, 1909
P. thisbe (Fabricius, 1782)

=iarbus (Drury, 1782)

=perditus (Fabricius, 1783)

=ovif era Seitz, 1913

a) eclypsis Seitz, 1913

b) Hewitson, 1875
P. brevilinea Sch?i\xs, 1920

P thy mele Stichel, 1909
P. trabalis Stichel, 1916

As constituted by Stichel, the genus Panara is polyphyletic. Three taxa have
been included erroneousely in the genus: ''Panara' elegans Schaus, 1920,
"Panara" brevilinea Schaus, 1928, and "Panara" sicora Hewitson, 1875. The
removal of P brevilinea was facilitated by its being a junior synonym of an
existing taxon. P. was removed to the genus PterographiumStichel, 1910
by Hall and Willmott (1996).

Phaenochitonia brevilinea (Schaus, 1920) , new combination, new synonomy.
My examination of the type of P. brevilinea at the NMNH suggests that it is a
synonym of Phaenochitonia iasis Godman, 1903, the type of which is in the
BMNH.

Pterographium elegans SchdiUS, new combination. Harvey (1987) discovered
that Panara elegans has the androconia on the anterior margins of the ab-
dominal sclerites, characteristic of the tribe Symmachiini. However, he did
not assign this species to a genus. My examination of P. elegans suggests that
it is near to Pterographium based on the presence of erectile scent hairs in
cell CU2-2A of the dorsal hindwing, the principal character for this genus

J. Res. Lepid.

(Zikan 1949). Therefore, I provisionally place it in Pterographium until the
limits of the genera of the tribe Symmachiini can be better defined. The
remaining group of species is monophyletic, sharing the characteristics
described for the genus Panara above.

With the changes proposed in this review, the following synonymic list
summarizes the classification of Panara:

P. phereclus (Linne, 1758)

ssp. barsacusWQ%t}NOod, [1851]
ssp. lemniscataThiGmc:, 1907
= comc5 Stichel, 1909

P. iarbas (Drury, 1782), replacement name

=thisbe (Fabricius, 1782), preocc. (thysbe Linne, 1764)

=perditus (Fabricius, 1793)
ssp. episatnius Frittwitz, 1865
=arctifascia Fuller, 1874
=eclypsis Seitz, 1913, new synonymy
ssp. thymele Stichel, 1909, new status

P. aureizona Fuller, 1874
=ornata Slichel, 1909

P. soana Hewitson, 1875, reinstated status
=trabalis Stichel, 1916, new synonymy
-dilata Lathy, 1932, new combination, new synonymy
ssp. bacana, new subspecies
ssp. ruschii, new subspecies

P. ovifera Seitz, 1913, new status

Ecology and Behavior

Habitat. The genus Panara is distributed in tropical South America to the
east of the Andes. One species, Panara phereclus (Linn.) ranges from the
Guianas throughout the Amazon and Orinoco drainages to Peru and Bo-
livia at elevations less than 200 m (Fig. 37) , a region characterized by Tropi-
cal Moist Forest habitats (Tosi 1983, Holdridge 1947). The other species
are concentrated in northeastern (Pernambuco) to southeastern Brazil
(Parana, Santa Catarina), reaching the central Planalto (Goias). In south-
ern Brazil, the species distributions are correlated with climatic zones.
Panara soana and its subspecies inhabit the Subtropical Wet Forest and Warm
Temperate Moist Forest zones, north through subtropical lower montane
moist forest in the Serra da Mantiqueira, and the montane formations of
the Subtropical Moist Forest habitat which reach their northern limit at
Santa Teresa, Espirito Santo (Tosi 1983). This distribution more or less
parallels that of the Parana pine tree Araucaria. Panara iarbas and its sub-
species inhabit the lower elevations of Subtropical Moist Forest in Rio de
Janeiro State north to Pernambuco, then west along gallery forests penetrat-
ing the Planalto Central to central Goias, and south to western Parana. The
two other species have limited distribution: Panara ovifera inhabits the cloud
forests of the Serra do Mar above 1300 m and Panara aureizona the coastal

34:21-38, 1995(1997)

Subtropical Moist Forest areas of Santa Catarina and Parana, and occasion-
ally to 900 m.

All species inhabit secondary as well as primary forest habitats. I have never
observed them flying outside the forest, except when visiting nectar sources.

Seasonality. Those species inhabiting lowland areas fly all year round,
whereas P. soana flies from September to May, and P. ovifera February and
March.

Biology. Early stage biology of the genus is unknown.

Adult Nectar Sources. I have observed Panara feeding during the morn-
ing on flowers, especially Eupatorium, and on one occasion on bird drop-
pings in the forest.

Wing Pattern and Predation. All Panara species are black with a yellow-
orange band on the forewing, a pattern shared by many other riodinids
(Melanis, Pterographium, Stichelia, Riodina) and day flying moths (Pericop-
inae) . It is not known whether Panara is distasteful, however males rest on
the dorsal surfaces of leaves with wings spread advertising of their color
pattern. I have never observed attacks by birds, nor have I captured speci-
mens with beak marks, which suggests that vertebrate predation is minimal.

Mating Behavior. All Panara use perching as mate locating behavior. Lo-
calities for perching are small clearings in the forest, such as along roads
and trails. Forested hilltops are frequented, but so are clearings in the same
area which suggests that the forest opening is more important than the
physical summit. From 1100-1500 h males rest on the edge of the dorsal
surface of sunlit leaves with wings outspread and antennae apart (Fig. 4)
awaiting females. When disturbed, they fly with a rapid, gliding flight, re-
turning shortly to their original perching site. When not perching, they rest
on the ventral leaf surfaces with a wingtip protruding beyond the edge.
Females are rarer, but are encountered at nectar sources or in the forest.

Key to the Males of Panara in Southeast Brazil

la. Ventral surface of both wings with dull dark purple scaling

at apex of forewing and base of hindwing..... 2

lb. Ventral surface with strong, iridescent light blue scaling at

apex of forewing and base of hindwing 4

2a. Male hindwing band present P. iarbas

2b. Male hindwing band absent P. aureizona

3a. Male forewing band reduced to an elongated, oval spot;
hindwing band wide and rounded towards costa, tapering

to inner margin......... P. ovifera

3b. Male forewing and hindwing bands narrow (1.5 mm)

elongated; hindwing band straight P. soana

Key to the Females of Panara in Southeast Brazil

la. Veins on both wings outlined with lighter scaling. P. soana

lb. Veins not outlined with lighter scaling 2

2a. Band on forewing reaches distal margin 3

J. Res. Lepid.

2b. Band on forewing does not reach distal margin P. ovifera

3a. Forewing band wide (>3 mm) P. aureizona

3b. Forewing band narrow (<3 mm) P. iarbas

Species Accounts

Panara iarbas (Drury, 1782) (Papilio), replacement name
=Papilio thisbe (Fabricius, 1782), preocc. {Papilio thysbelAnn^, 1764)

= Hesperia perditusYdbricivi^, 1793

Nomenclature. Panara iarbus (Drury 1782): Papilio thisbe\^?iS described by
Fabricius as having yellow bands on both fore- and hindwings, and as com-
ing from “Brazil.” However, Fabricius’ name is a primary homonym of Papilio
thysbelAnne, which refers to a South African lycaenid butterfly currently in
the genus Poecilmitis , 1899. Drury (1782) essentially repeated Fabri-

cius’ description in describing Papilio iarbas, which becomes the next avail-
able replacement name for P. thisbe. No type of either taxa has been located;
however, as no strong ventral surface blue reflections are mentioned in ei-
ther description, the name probably refers to the coastal populations of P.
iarbus from central-southeast Brazil, the males of which consistently have
yellow bands on both the fore- and hindwings.

Hesperia per ditus Y2d^r\c\\xs, 1793: This taxon, described from “French
Guiana” is identical to P. iarbas. However, information supplied by C.
Brevignon, a resident collector (pers. comm.), suggests that this taxon is
not found there. Scudder (1875) designated P. iarbus diS the type species of
the genus Panara Doubleday, 1847.

In view of the inadequacy of the original descriptions, the species is re-
described as follows:

Male. Medium sized (forewing length average 20 mm) , robust riodinid
butterfly with black appendages and a yellow lateral line on the abdomen.
Palpae (Fig. 2) and male foreleg (Fig. 3) as illustrated. Vein R1 on the fore-
wing rises before the discal cell with R2 following it (Fig. 1). Wings black
with an orange-yellow 2-3 mm wide transverse band on the forewing be-
tween the costa to less than 1 mm from the distal margin, and with a sec-
ond transverse band of variable width (0.3-3 mm) on hindwing between
0.5 mm from the apex and the inner margin. Ventral wing surface with same
transverse bands as on dorsal surface, ground color black with a faint purple
reflection, stronger at the forewing apex. Fringe black.

Female. Fore- and hindwing more rounded than male; forewing trans-
verse band width variable (1-3 mm), reaching from costa to distal margin,
curving to anal angle; hindwing band dorsal surface when present extends
from costa to inner margin, convex to base and is duplicated on the ven-
tral surface; when absent dorsally, is reflected by a band of lighter scaling
on the ventral surface.

Genitalia. Male genitalia (Fig. 29) with pedicel a strap-like band connect-
ing aedeagus to base of valvae; tergum deeply indented caudad; vinculum
ribbon-like ventrally, not fused to valvae; valvae sickle-shaped, serrated

34:21-38, 1995(1997)

Fig. 2. Panara male palpus
Fig. 3. Panara male foreleg

caudad; tips of valvae reaching to transtilla; transtilla with two small projec-
tions caudad; saccus reduced.

Female genitalia (Fig. 33) with blade-like papillae anales fused dorsad;
ostium bursae squared, sclerotized; corpus bursae without signa.

Geographical Distribution. Panara ranges from Rio de Janeiro State

north to Pernambuco, then west across the Planalto to Goias, from sea level
to 1000 m, then south through western Sao Paulo State to western Parana.

Geographical Variation. I recognize three distinct populations, repre-
sented by the nominate subspecies, P. iarbas episatnius, and P. iarhus thymele.

Panara iarbas iarbas (Drury, 1782) (Papilio), replacement name

Identification. The nominate subspecies has wide hindwing bands on both
sexes. The male can be separated easily from Panara soana by the lack of
strong blue reflections on the ventral surface at the apex and base of the
wings.

Geographic Distribution. The distribution of P. iarbas iarbas is disjunct,
from the Serra da Carioca and Serra do Mar in western Rio de Janeiro State
to southeastern Minas Gerais, and again in the Zona da Mata from north-
ern coastal Espirito Santo State north to Pernambuco.

Brazil. Rio de Janeiro: Rio de Janeiro, 0-600 m, 25 d, 2$, MN; 5d, 19,
NMNH; 4d , 6 9, UFP; Novo Friburgo, 2 d , 1 9 , NHML; Jacarepagua, 3 d ,

J. Res. Lepid.

Fig. 4. P. iarbus perching, Barra de Sao Joao, R.J. Brazil

UFP; Angra dos Reis, 8d, 1$, MN; Paineiras, R.J., 3d, 1$, MN. Minas
Gerais: Passa Quatro, MG, 1$, NHML. Bahia: “Bahia,” 1$, SM; Itamaraju,
Id, MN; Ilheus, 2d, MN; Pernambuco: 10 km E.Joao Pessoa, 7d, 2$; Sao
Lourengo, Id, NMNH; Tiuma, PE, 19, CJC.

Ecology and Behavior. In coastal Brazil, P. iarbas iarbas inhabits primary
and disturbed humid subtropical forest. Males perch in the late morning
to early afternoon in light gaps and other small clearings and on hilltops,
resting on dorsal leaf surfaces with their wings outspread and head and body
raised at a 30° angle from the leaf surface (Eig. 4). Eemales are encoun-
tered less frequently, flying near the ground in the forests. It is local and
uncommon.

P. iarbas episatniusYriXiMitz, 1865, new combination (Eigs. 8-10)

= P.artifascia Butler, 1874
= P. eclypsis Seitz, 1913, new synonymy
Nomenclature. Panara episatniusFTittwitz,1865: Prittwitz described Panara
episatnius ^rov[\ a female from Rio de Janeiro, currently in the Natural His-
tory Museum, London. Stichel (1930) subsequently designated P. episatnius
a subspecies of phereclus based on the absence of the hindwing band. This
was in error, as P phereclus is limited to the Amazonian drainage.

Panara arctifascia Butler, 1874: Butler described this taxon from a female

34:21-38, 1995(1997)

Fig. 5. P. iarbas iarbas, male dorsal surface
Fig. 6. P. iarbas iarbas, male ventral surface
Fig. 7. P. iarbas iarbas, female dorsal surface
Fig. 8. P. iarbas episatnius, male dorsal surface
Fig. 9. P. iarbas episatnius, male ventral surface
Fig. 10. P. iarbas episatnius, female dorsal surface

from Espirito Santo presently in the Natural History Museum, London,
which is identical to P. episatnius. The two were synonymized by Stichel
(1930).

Panara eclypsis Seitz, 1913: Seitz based his description of Panara eclypsis on
a male from Espirito Santo, designating it as a form of Panara thisbe, which
was subsequently raised to a subspecies by Stichel (1926). P. eclypsis is in
fact the male of P. episatnius, thus becoming a junior synonym of that taxon.
The locality of the type of P. eclypsis is unknown; however the butterfly is
distinct enough as to make the designation of a neotype unnecessary.

Identification. The males P. i. episatnius differ from the nominate subspe-
cies in the reduced width of the band on the dorsal hindwing to 0.5 mm
and ventrally to 1 mm. Eemales differ in the absence of the hindwing band.
P i. episatnius intevgrdide?> to the east of Rio de Janeiro State with P. i. iarbus,
some individuals showing characteristics of both phenotypes. The male
genitalia of material from central Espirito Santo have long points on the
valvae, whereas those from eastern Rio de Janeiro State are identical to nomi-
nate P. i. iarbas.

Geographic Distribution. P. i. episatnius is found throughout Espirito Santo
and adjoining eastern Minas Gerais State below 800 m. This suggests that

J. Res. Lepid.

P. i. episatnius is an isolated population of P. i iarbas which has recently come
into secondary contact.

Brazil. Minas Gerais: Parque Estadual de Rio Doce, 2d , 2 $ , CJC; Espirito
Santo: Linhares, Id, CJC: B. Guapemirim, Id, MN; Boitacazes, Id, MN;
“Espiritu Santo,” 3d, MN; Colatina, 6$, MN; Conceic^ao da Barra, Id, 1$,
UEP; Linhares, 1$, UFP; Baixu Guandu, ES, 2d,UFP.

Intergrades to P. iarbas iarbas: Rio de Janeiro: km 27, Rio-Teresopolis, Id,
CJC; Barra de Saojoao, 2d, CJC; Fazenda Uniao, 4d, CJC.

Ecology and Behavior. These are the same as the nominate subspecies.

P. iarbas thy mele Stichci, 1909, new status (Figs. 11-13)

Nomenclature. Panara thymele was described from a male from Casa
Blanca, Sao Paulo, currently in the Museum fur Naturkunde, Humboldt
Universitat, Berlin. P. i. intergrades in central Bahia with P. i. thymele,

suggesting that they are conspecific.

Identification. The male can be distinguished by the slight S-shaped fore-
wing band and both males and females by the hindwing band concave to
the margin. Specimens from Bahia have thinner bands than those from
Goias.

Geographic Distribution. P. i. thymele is found from western Bahia south
to western Parana, then across the Planalto Central to Goias State.

Brazil. Federal District: Sobradinho, 3d, CJC; Parque da Gama, DF, 2d,
19 , UFP; Agua Limpa, DP, 3d, UFP. Goias: Goias Velho, 2d, 1 9 , CJC; 19 ,
UFP Bahia: km 997, Rio-Bahia, 19, CJC; “Bahia,” 9d, 19, BMNH; Campo
Formosa, Juazeiro, Id, UFP. Parana: Guarapuava, 1000 m. Id, UFP.

Ecology and Behavior. On the Planalto, the subspecies inhabits the gal-
lery forests along streams and cabeceira (headwater) woods where it flies
during the early afternoon hours, frequenting the edges of clearings where
the males perch on the dorsal leaf surfaces with wings open.

Panara aureizonaViuhcr , 1874 (Figs. 14-16)

=P. aureizona f omata Stichel, 1909

Nomenclature. Butler described P. aureizona from a female from “Minas
Gerais,” currently in the Natural History Museum (London). It is sympat-
ric at Joinville, Santa Catarina with P. soana and allopatric with P. i. thymele
to the west. The genitalia are intermediate between P iarbas and P. soana.

Geographical Variation. P. aureizona is very rare in collections, so its dis-
tribution and variation are not well known. An occasional male has a spot
of orange where the band should be on the hindwing.

Identification. Males may be separated by the 3.5 mm wide band from
the costa to the outer margin, and the females by the 4 mm wide forewing
band which extends basad and distad along the costa, and is curved towards
the anal angle on the distal margin. Both sexes lack the hindwing band,
but on the VHW is a faint transverse line of lighter scaling in the normal
position of orange band. In the male genitalia (Fig. 30) , the valvae do not
reach the transtilla; the ostium bursae in the female genitalia (Fig. 34) has
V-shaped sides.

34:21-38, 1995(1997)

Geographic Distribution* P. aureizona ranges from coastal Santa Catarina
and Parana north and west to eastern Minas Gerais (?) . As no other records
have been found between Parana and Minas Gerais, the locality of Butler’s
type is suspect.

Brazil. Santa Catarina: Itaiopolis, 900 m, Id, SM; Garcia, 60 m, 3d, SM;
Blumenau, 50 m, 3d, SM; Macaranduba, 130 m, Id, SM;Joinville, 3d,UFP;
10 d, 10$, NM; Id, 1$, CJC; Jaragua, 200 m, Id, UFP. Parana: Marumbi,
500 m, Id, UFP; 5d, 4$, no locality, BMNH.

17 !a

Fig. 1 1 . P. iarbas thymele, male dorsal surface
Fig. 12. P. iarbas thymele, male ventral surface
Fig. 13. P. iarbas thymele, female dorsal surface
Fig. 14. P. aureizona, male dorsal surface
Fig. 15. P. aureizona, male ventral surface
Fig. 16. P. aureizona, female dorsal surface
Fig. 17. P. soana soana, male dorsal surface
Fig. 18. P. soana soana, male ventral surface
Fig. 19. P. soana soana, female dorsal surface

J. Res. Lepid.

Ecology and Behavior. P. aureizona inhabits disturbed tropical forest from
sea level to about 900 m. The males are found hilltopping atjoinville, Santa
Catarina (H.W. Miers, pers. comm.). The females are encountered more
often beside roads and in the forest.

Panara soana Hewitson, 1875

Identification. Panara soana males may be separated from other Panara
by the blue sheen at the apex of the forewing, at the base and along the
margin of the hindwing combined with a straight, narrow, band on the fore-
and hindwings; and the females by a dusting of lighter scaling along the
veins.

Geographical Variation. I recognize three distinct geographical popula-
tions of P. soana, two of which are new. All three are allopatric with no known
intergrades, which future investigations may show to be separate species.

Panara soana soana Hewitson, 1875, reinstated status (Figs. 17-19)

= P. trabalis Stichel, 1916, new synonymy
= P. dilata Lathy, 1932, new combination, new synonymy

Nomenclature. Panara soana Hewitson, 1875: P. soana was described by
Hewitson from a male labeled “Brazil.” Comparison of the type with mate-
rial from Santa Catarina, Parana, and Sao Paulo suggests that the specimen
originated from this region. Stichel (1909) designated P. soana^s, a subspe-
cies of P. thisbe {iarbas). Examination of these two taxa suggests that this
was in error, as they maintain consistent morphological differences, even
when sympatric (Santa Teresa, ES; Novo Friburgo, Rio de Janeiro) . The type
of P. soana soana is in the Natural History Museum (London).

Panara trabalis Stichel, 1916. Panara trabalis wsis described as a species by
Stichel from a female from Santa Catarina, Brazil, located in the Natural
History Museum, London. The type represents the female of P. soana soana,
the white scaling along the veins and no band on the hindwing being typi-
cal of southern Brazilian populations.

Panara dilata Lathy, 1932. P. dilata^A^ described by Lathy from a female
from Ponto Grosso, Parana, and who assigned it to the genus Lymnas,
Blanchard (currently Melanis Hubner). The type in the Natural History
Museum (London) is the female of P. soana.

Identification. The male of the nominate subspecies is distinguished by
strong light blue reflections on the ventral forewing at the apex and ven-
tral hindwing at the base and around the margin, when viewed at an angle.
The female lacks the band on the hindwing and has light scaling along the
veins of both wings.

The valvae in the male genitalia (Fig. 31) extend above the transtilla, and
the ostium bursae in the female genitalia (Fig. 35) has V-shaped sides.

Geographical Variation. The nominate subspecies Panara soana soana
(Figs. 12-13) ranges from northern Rio Grande do Sul State north along
the Serra do Mar to Sao Paulo State.

34:21-38, 1995(1997)

Fig. 20. P. soana bacana, male dorsal surface
Fig. 21 . P. soana bacana, male ventral surface
Fig. 22. P. soana bacana, female dorsal surface
Fig. 23. P. soana ruschii, male dorsal surface
Fig. 24. P. soana ruschii, male ventral surface
Fig. 25. P. soana ruschii, female dorsal surface
Fig. 26. P. ovifera, male dorsal surface
Fig. 27. P. ovifera, male ventral surface
Fig. 28. P. ovifera, female dorsal surface

Brazil. Minas Gerais: Virginia, 900 MN; Parque Nacional Itatiaia,

900 m, M, MN; \6 , 19, UFP; Itajuba, 4c^; Sao Paulo: “Sao Paulo,” 1 9 , MN;
M, UFP; 26, NHML; Amparo, 19, MN; Cantareira, \6 , 19, MN, 19, UFP,
4 9 , SM; Sa. Japi, 1 9 , CJC. Parana: Ponta Grossa, 46,49 , UFP; Rio Vermelho,
26, CJC; Curitiba, 16, CJC; 76,59, UFP; Sao Luiz, Puruna, 2c?, UFP;
Vossoroca, 3c?, UFP; Santa Catarina: Joinville, 2c?, CJC; Campo Alegre, 7c?,
NMNH; Sao Bento do Sul, 2 c? , 19, CJC: Sao Luis de Parana, 2c? , 19, CJC;
Blumenau, 3 c?, SM; Massaranduba, Ic?, SM.

J. E£s. Lepid.

Fig. 29. P. iarbas iarbas, male genitalia
Fig. 30. P. aureizona, male genitalia
Fig. 31 . P. soana, male genitalia
Fig. 32. P. ovifera, male genitalia
Fig. 33. P. iarbas iarbas, female genitalia
Fig. 34. P. aureizona, female genitalia
Fig. 35. P. soana, female genitalia
Fig. 36. P. ovifera, female genitalia

Ecology and Behavior. P. soana soana inhabits montane subtropical fon
est above 600 m. Males perch on the forest edges during the early after-
noon, resting on dorsal leaf surfaces with wings spread. At some localities
they are common.

Panara soana bacana Callaghan, new subspecies (Figs. 20-22)
Description, Male differs from the nominate subspecies in having a wider
band on the dorsal hindwing and a reduction of the blue iridescence at the
apex and base of hindwing. Cara^a specimens have less blue than those to
the southeast. Female differs in a marked reduction in the white scaling
along the veins and the presence of a yellow transverse band on the hindwing

34:21-38, 1995(1997)

Fig. 37. Distribution of the species of Panara. ■ P. phereclus • P. iarbas
A P. auerizona, □ P. soana, A P. ovifera

from the inner margin near the anal angle narrowing to the costa, where it
turns slightly basad.

Holotype Male. With label “BRAZIL, Minas Gerais, Caraga, 2500 m,
264v“1975, C. Callaghan,” a genitalia label #424 and a red holotype label.
The holotype is deposited in the Museu Nacional, Rio de Janeiro, Brazil.

Paratypes. Passa Quatro, MG, M , MN; Caraga, MG, 1500 m, 3d , 3 $ , CJC;
15d, 8$, NHML; Barbacena, 1200 m, MG, 4d, 2$, CJC; Pogo de Caldas,
600 m, 9d, 1$, MN; Caxambu, MG, 5d, MN; Novo Friburgo, R.J., 3$,
NHML.

Etymology. “Bacana” means “nice” in Portuguese.

Ecology and Behavior. Panara soana bacana inhabits subtropical humid
forest patches in the Serra de Mantiquera and Serra do Mar at 600-1800 m

J. Res. Lepid.

Fig. 38. Distribution of Panara in southeast Brazil. • P. iarbas iarbas, I P. iarbas
thymele, O P. iarbas episatnius, A P. aureizona, □ P. soana, ■ P. soana
bacana, T P. soana ruschii, A P. ovifera

from southeastern Minas Gerais to the Serra de Caraga. It is sympatric with
P. iarbas in Novo Friburgo, 900 m, Rio de Janeiro.

Panara soana ruschii Callaghan, new subspecies (Figs. 23-25)

Description. Male differs from the nominate subspecies in having a longer,
more pointed forewing; forewing band narrower, tapering from costa to 2
mm from distal margin above anal angle; ventral surface blue reflections at
forewing apex and base of hindwing stronger and more extensive, that on
apex of forewing extending along distal margin to 3 mm above anal angle.
Female with very light white dusting along veins, band on forewing 4 mm
wide at costa, tapering to a rounded point 1 mm from distal margin above
anal angle, hindwing without transverse band. Genitalia as in nominate sub-
species.

Holotype Male. With label “BRAZIL E. Santo Santa Teresa 800 m, 5-iv-1973
C. Callaghan,” a genitalia label #420, and a red holotype label.

Paratypes. Santa Teresa, Espirito Santo, 900 m 2d, 29, CJC. The holo-

34:21-38, 1995(1997)

type and a female paratype are deposited in the Museu Nacional, Rio de
Janeiro.

Etymology. This taxon is named in memory of the famous Brazilian con-
servationist, Augusto Ruschi, who I got to know during my visits to Santa
Teresa where he lived.

Ecology and Behavior. The subspecies is currently known only from the
type locality where the males frequent hilltops in the early afternoon, perch-
ing on dorsal leaf surfaces with wings spread.

Panara ovifera Seitz, 1913, new status (Figs. 26-28)
Nomenclature. Panara oviferaw2iS described by Seitz (1913) from a male
from Petropolis, Rio de Janeiro as a form of P. thisbe. The phenotype is rep-
resentative of a unique isolated Panara population. The truncated bands,
extensive blue sheen on the ventral surface, high mountain habitat and
absence of intergrades separate it from P. iarbas. Panara ovifera is allopatric
with P soana. There are no dines and it is consistently distinct morpho-
logically. The type is in the Natural History Museum (London).

Identification. The males of P. ovifera can be separated by the triangular
orange spot tapering below the cell on the forewing, and the wide, short
band on the hindwing. The ventral wing surface has the same pattern of
shiny blue scaling at the apex of the forewing and the base and margin of
the hindwing as P soana soana, but more extensive. In the male genitalia
(Fig. 32), the tips of valvae extend beyond transtilla and the female ostium
bursae (Fig. 36) has V-shaped sides, and a wide sinus vaginalis.

Ecology and Behavior. Panara ovifera is restricted to the pygmy chusquea
cloud forests above 1300 m in the Serra do Mar, Rio de Janeiro State. The
males rest on the upper leaf surfaces with wings outspread beside roads and
other openings in the forest between 1100-1500 h. It is rare.

Material examined. Petropolis, Estrada Imperial, 1300 m, 3d, 39, CJC;
Petropolis, RJ, Id, MN.

Acknowledgements. I am indebted to the curators of the museums visited for access
to the collections under their care, to Drs. Robert Robbins and Donald Harvey of
the Smithsonian and Dr. Keith Brown of the Universidade Estadual de Campinas,
Sao Paulo. Two anonymous reviewers provided helpful comments on the manu-
script.

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Ins-Biol. 12:168.

. 1926. Beitrage zur Kenntnis der Riodiniden Fauna Sudamerikas: Nord Brasil.

Z. Insekten Biol. 20:14-23.

. 1930. In W. Junk, ed. Lepidoptorum catalogus, v. 30 Berlin, 795 pp.

Thieme, O. 1907. Familiae Lemoniidarum supplementa cum nods (Lepidoptera
Rhop.). Berk ent. Z. 52:1-16.

Tosi, J. 1983. Provisional Life Zone Map of Brazil at 1:5,000,000 scale. Tropical
Science Center, San Jose, Costa Rica.

Westwood, J.O. [1851]. In E. Doubleday & W.C. Hewitson. Genera diurnal
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ZiKAN, J.F. 1949. Observagoes sobre os componentes dos generos Phaenochitonia
Stichel e Pterographium Stichel, com a descrigao de um novo genero (Riodinidae,
Lepidoptera). Rev. de Ent. 20:1-3, 535-539.

34:39-47, 1995(1997)

Journal of Research on the Lepidoptera

Lepidoptera of different grassland types across the Morava
floodplain

Miroslav Kulfan, Peter Degma, and Henrik Kalivoda

Department of Zoology, Comenius University, Mlynska dolina B-1, 842 15 Bratislava, The
Slovak Republic

Abstract. The occurrence of the diurnal and readily disturbed Lepi-
doptera species were studied during the 1992-1994 flight seasons on dif-
ferent types of grassland resulting from management practice: mowing,
cattle grazing, and application of liquid manure. Observations were made
across the southern part of the Morava River alluvia on 9 study sites. The
transect method was used with 111 lepidopterous species in 15 families
recorded. Five of these species are vulnerable and two species endan-
gered. Zerynthia polyxena, Iphiclides podalirius, Colias chrysotheme, Melanargia
galathea, Minois dry as, Erynnis tages, Agriphila inquinatella, Loxostege sticticalis,
Zygaena loti, Zygaena angelicas, Scopula virgulata, S. immutata, Idaea spp.,
and Euclidia glyphica were associated with relatively well preserved xero-
thermic grassland habitats (no cutting history). Heteropterus morpheus,
Colias hyale, Lycaena dispar, and Maculinea teleim associated with semi-an-
nually cut and regularly flooded extensive wet meadows. Psammotis
pulveralis, Elophila nymphaeata, Coenonympha glycerion, Lycaena tityrus, and
Phlogophora meticulosa were found in bog habitat. The Lepidoptera com-
munity from the non-mown undisturbed areas with Crataegus sp. (forest-
steppe vegetation) showed the highest diversity (H' = 2.944) and rela-
tively high evenness (e = 0.760). The community from cut wet meadow
that is heavily fertilized with liquid manure has both the lowest diversity
(H' = 1.036) and evenness (e = 0.383).

Introduction

The Morava River forms the border between the Slovak Republic and
Austria and partly between the Slovak and Czech Republics. The total length
of the river is 353 km. In Slovakia the Morava River flows across the Borska
Nizina lowland which forms part of the Pannonian region (Kulfan Sc Kulfan
1992). The lower reach of the Morava floods annually, usually in spring and
often in summer after rainstorms. The territory where the field study was
done, on the border near Austria, is relatively well preserved because it was
inaccessible before 1990 (see map. Fig. 1). The study region is now part of
the protected landscape territory Zahorie, where 1000 ha of inundation
meadows are covered with the subcontinental plant association Cnidion dubii
(Ruzickova, 1994). J. Kulfan (1989, 1990a, b) studied the lowland butterfly
communities of Borska Nizina on irregularly cut meadows near the village
of Rohoznik.

Paper submitted 28 April 1995; revised manuscript accepted 15 May 1996.

J. Res. Lepid.

Fig. 1 . Map of the lower Morava River showing approximate location of the sites
used in this study.

The only data on the grassland Lepidoptera of the lower Morava River
alluvia deal with the distribution of heliophilous species across Borska Nizina
where these species were evaluated by trophic relations to host plants and
from a conservation perspective (Kulfan & Kalivoda 1994).

The purpose of this paper is to compare the Lepidoptera communities
across different grassland management regimes that vary according to cut=
ting intensity, cattle grazing, and the application of liquid manure.

34:39-47, 1995(1997)

Methods and Study Areas

Our investigations were carried out on the lower Morava River near a border
between Slovakia and Austria during the flight seasons of 1992-1994 on sunny days
at about 2 week intervals. Day flying and easily disturbed (roused) Lepidoptera were
caught by net following the transect method ofErhardt (1985). The length of each
transect was 200 m with 414 samples taken.

Nine sites were sampled:

Site 1: Alluvial meadow near Marchegg village cut twice and partly flooded dur-
ing the annual cycle {Carici praecoci-Alopecuretum pratensis dissociation, Spanikova
1975, subassociation typicum Bdl.-Tul. 1963, with occurrence of Iris sibirica).

Site 2: Partly cut narrow area near Marchegg village along the field path, between
Prunus and Crataegus shrub grove and the Morava river {Carici praecoci-Alopecuretum
pratensis, subassociation with Filipendula vulgaris ba\.-Tu\. 1974 with occurrence of
Aristolochia clematitis) .

Site 3: Alluvial meadow near Marchegg village cut twice annually and flooded
regularly in spring and summer following heavy rains {Carici praecoci-Alopecuretum
pratensis, subassociation typicumv^ith occurrence of Lychnis flos-cuculi, Iris sibirica,
Iris pseudacorus, Clematis integrifolia) .

Site 4: Non-mown narrow area near the Devinske Jazero railway station between
the path and the Phragmites australis stand {Carici praecoci-Alopecuretum pratensis,
subassociation typicum^Nith occurrence of Aster lanceolatus and Clematis integrifolia).

Site 5: Alluvial meadow near the Devinske Jazero railway station, usually partly
mown twice annually with spring flooding, edged with Phragmites australis growth
{Lathyrus paluster-Gratiola officinalis Bal.-Tul. 1963 and Carici praecoci-Alopecuretum
pratensis associations, latter subassociation typicum with occurrence of Thalictrum
flavum and Leucojum aestivum ).

Site 6: Alluvial meadow cut twice annually, but rarely flooded ( Carici praecoci-
Alopecuretum pratensis association, subassociation typicum With Colchicum autumnale,
Galium verum, Symphytum officinale, Sanguisorba officinalis. Inula salicina) .

Site 7: Uncut area near Devinska Nova Ves (suburb of Bratislava) with the forest-
steppe vegetation, Crataegus surrounded by agricultural phytocoenoses and by al-
luvial meadows (Carici praecoci-Alopecuretum pratensis, subassociation with Filipendula
vulgaris and Serratulo-Festucetum commutatae Bal.-Tul. 1963 association on gravelly
outcrops and sandy alluvial sediments with occurrence of Galium verum, Sanguisorba
officinalis. Inula salicina, Rumex acetosa, Centaurea jacea, Fragaria viridis, Aristolochia
clematitis) .

Site 8: Alluvial meadow near Devinska Nova Ves, cut twice annually and partly
flooded. Highly modified by application of liquid manure. There is a depauperate
community of plant species ( Carici praecoci-Alopecuretum pratensis, subassociation
typicum) .

Site 9: Alluvial meadow near Devinska Nova Ves modified by cattle grazing with a
depauperate community of plants, partly flooded annually {Carici praecoci-
Alopecuretum pratensis, subassociation typicum with Galium verum, Rumex crispus,
Cirsium arvense) .

The lower Morava floodplain was entirely flooded at the beginning of August
1991, with water level reaching about 2.5 m above ground elevation.

J. Res. Lepid.

Table 1 . General survey of Lepidoptera species found on the Morava River
alluvia (EC = Ecological Characteristics: X = xerothermophil, M = mesophil, H =

hygrophil, U = ubiquist).

Taxon

Incurvaroidea

Adelidae

Adela reaumurella (L., 1758)

Toriricoidea

Tortricidae

Aphelia viburnana (D. et. S., 1775)
Aphelia paleana (Hb., 1793)

Agapeta zoegana (L., 1767)

Olethreutes rivulana (Sc., 1763)

Epiblema uddmanniana (L., 1758)
Dichrorampha gueneeana (Obr., 1953)
Pterophoroidea
Pterophoridae

Pterophorus pentadactyla (L., 1758)

Pyraloidea

Pyralidae

Hypochalcia ahenella (D. et S., 1775)
Tachycera advenella (Gm. et Znk., 1818)
Elophila nymphaeata (L., 1758)

* Chrysoteuchia culmella (L., 1758)
Crambus pascuella (L., 1758)

Crambus lathoniella (Znk., 1817)
Crambus perlella (Sc., 1763)

Agriphila tristella (D. et S., 1775)

* Agriphila inquinatella (D. et S., 1775)
Platytes cerussella (D. et S., 1775)

* Evergestis aenealis (D. etS., 1775)

* Pyrausta despicata (Sc., 1763)

* Loxostege sticticalis (L., 1761)
Ecpyrrhorrhoe rubiginalis (Hb., 1796)

* Sitochroa verticalis (L., 1758)

* Psammotis pulveralis (Hb., 1796)
Pleuroptya ruralis (Sc., 1763)

Zygaenoidea

Zygaenidae

* Adscita statices (L., 1758)

* Zygaena loti (D. et S., 1775)

Zygaena viciae (D. et S., 1775)

* Zygaena filipendulae (L., 1758)

* Zygaena angelicaeO., 1808

Hesperoidea

Hesperiidae

* Erynnis tages (L., 1758)

* Pyrgus malvae (L., 1758)

Carterocephalus palaemon (Pallas, 1771)
Heteropterus morpheus (Pallas, 1771)

* Thymelicus sylvestris (Poda, 1761)

* Thymelicus lineolus (O., 1808)

Hesperia comma (L., 1758)

* Ochlodes venatus (Br. et Grey, 1853)

EC 1

X, M
M
M

X, M
X
X
X
X
X
X
X
H

X, M

X, M
X, M
M, H
H
M
M
M
M

Study Area
4 5 6

34:39-=47, 1995(1997)

Papilionoidea

Papilionidae

* Zerynthia polyxena (D. et S., 1775) X

* Papilio machaon L., 1758 X, M

* Iphiclides podalirius (L., 1758) X

Pieridae

Leptidea sinapis (L., 1758)

* Pieris brassicae (L., 1758)

* Pieris rapae / napi (L., 1758)

* Pontia daplidice (L., 1758)

* Anthocharis cardamines (L., 1758)

* Colias hyale (L., 1758)

* Colias alfacariensis^hhe, 1905
Colias chrysotheme (Esper, 1781)

* Colias crocea (Fourcr., 1785)

* Colias erate (Esp., 1804)

Gonepteryx rhamni (L., 1758)

Nymphalidae

* Apatura ilia (D. et S., 1775)

* Inachis io (L., 1758)

* Vanessa atalanta (L., 1758)

* Cynthia cardui (L., 1758)

* Aglais urticae (L., 1758)

* Polygonia c-album (L., 1758)

Araschnia lev ana (L., 1758)

* Issoria lathonia (L., 1758)

* Clossiana selene (D. et S., 1775)

Clossiana dia (L., 1767)

* Mellicta athalia (Rott., 1775)

Satyridae

* Melanargia galathea (L., 1758)

Minois dryas (Sc., 1763)

* Maniola jurtina (L., 1758)

* Aphantopus hyperanthus (L., 1758)

* Coenonympha pamphilus (L., 1758)

* Coenonympha glycerion (Bkh., 1788)

Pararge aegeria (L., 1758)

* Lasiommata megera (L., 1767)

Lycaenidae

Fixsenia pruni (L., 1758)

* Lycaena phlaeas (L., 1761)

* Lycaena dispar (Haw., 1803)

* Lycaena tityrus (Poda, 1761)

* Everes argiades (Pallas, 1771)

Everes decoloratus (Stdgr., 1886)

* Celastrina argiolus (L., 1758)

Maculinea teleius (Brgstr., 1779)

* Lycaeides argyrognomon (Brgstr., 1779)

Aricia agestis (D. et S., 1775)

* Polyommatus icarus (Rott., 1775)

Geometroidea
Geometridae

* Timandra griseataW. Pet., 1902 X, M

* Scopula immorata (L., 1758) X

* Scopula virgulata (D. et S., 1775) X, M

* Scopula immutata (L., 1758) M

* Idaea serpentata (Hufn., 1767) X, M

4 11

4 4

4
7
7

U 80
X 9
X, M 4
M
X
X

X, M
X
M

U 2

X, M
M

X, M
X
U
M

U 67
M, H 22
M

X, M

M, H
X, M
X

X, M
H
X
X

X, M 7

J. Res. Lepid.

Idaea subsericeata (Haw., 1809)

Idaea aversata (L., 1758)

Lythria purpuraria (L., 1758)

Lythria rotaria (F., 1798)

Catarhoe cuculata (Hufn., 1767)
Epirrhoe alternata (Muller, 1764)
Minoa murinata (Sc., 1763)

Lomaspilis marginata (L., 1758)
Semiothisa clathrata (L., 1758)
Ematurga atomaria (L., 1758)

Siona lineata (Sc., 1763)

Sphingoidea

Sphingidae

Agrius convolvuli (L., 1758)
Macroglossum stellatarum (L., 1758)
Hyles euphorbiae (L., 1758)

Noctuoidea

Arctiidae

Diacrisia sanio (L., 1758)

Syntomis phegea (L., 1758)

Noctuidae

Polypogon tentacularia (L., 1758)
Euclidia glyphica (L., 1758)

Deltote bankiana (F., 1775)

Emmelia trabealis (Sc., 1763)
Macdunnoughia confusa (Stph., 1850)
Autographa gamma (L., 1758)
Phlogophora meticulosa (L., 1758)

13 7

+ 2

X, M

M, H

+ 2

7 4

M, H

Results and Discussion

Table 1 presents the survey results of the Lepidoptera species found on
the Morava floodplain. We have found 111 species in the region, but spe-
cies not recorded during transect counts marked by a + . All the found spe-
cies were used to construct the dendrogram of species similarity, using
Soerensen’s index following the Complete linkage clustering method of
Podani (1993). The summary numbers of individuals of different species
(68 species marked with an asterisk in Table 1) in the course of the periods
1992-1994 (27 samples from each site, 9 samples each year) were used for
the Shannon and Weaver diversity and the Pielou equability (evenness)
indices that are given in Fig. 2. The samples of Lepidoptera were from the
same or similar date of each year with the differences between triplets of
corresponding dates not exceeding 8 days.

Numerical classification according to species similarity shows that the Lepi-
doptera communities from non-flooded or rarely flooded sites with the plant
community at drier elevations (sites 2, 6, and 7) form a separate group (Fig.
2). The community from the driest site (site 7) with forest-steppe vegeta-
tion has the highest diversity (H' = 2.944) and high evenness (equability)
(e = 0.760) . This corresponds with the high species richness on this site, 78
species (Table 1). The Lepidoptera communities of the damp sites (1, 3, 4,
5, 8, and 9) form a single group. Within this group the communities of sites
8 and 9, however, differ considerably from the others. They inhabit sites
affected by negative anthropogenic factors, this is, intensively managed

34:39-47, 1995(1997)

0.511 0 712 0.760 0.735 0.383 0.717 0.736 0.767 0.760

Fig. 2. Dendrograms of classification of Lepidoptera communities in individual
study sites according to species presence/absence (to the left) and ac-
cording to frequency of species (to the right). Diversity and equability
values of these communities are given under the left dendrogram.

meadows. The community of Lepidoptera from site 8 with a depauperate
plant community and application of liquid manure shows the lowest diver-
sity (H' = 1 .036) and evenness (e = 0.383) . In contrast the community from
site 9 with many ruderal flowering plants {Asteracae) for food sources for
adult butterflies has relatively high diversity (H' = 2.148) and evenness (e =
0.717).

The Lepidoptera communities of the more flooded sites, 3 and 5, form a
distinct group. These communities have high evenness, e = 0.760 and 0.735,
respectively. Communities from sites with the same plant association (1 and
4) form another separate group, but are similar to each other at a low level
(Fig. 2).

The hierarchical classification (Podani 1993; Fig, 2) used the Complete
linkage clustering method and Similarity ratio index for making the den-
drogram based on the frequency of 78 species (given in Table 1) which were
derived from all 46 samples from each site.

The classification shows that the Lepidoptera communities of the driest
sites, 6 and 7, form a conspicuous grouping (Fig. 2). The communities of
the intensively managed meadows, sites 8 and 9, form a separate group to-
gether with the community of regularly flooded meadow, site 3. All these
sites, 3, 8, and 9, are extensive open wet meadows without shrubs and trees.
Lepidoptera communities of the narrow areas (1, 2, 4, and 5) surrounded
by zones of a transitional character (e.g., shrubland) or by different habi-

J. Res. Lepid.

tat type (e.g., banks, high Phmgmites australis st3.nds, etc.) form yet another
separate group.

According to the lUCN classification, five of the species found are listed
as vulnerable: Heteropterus morpheus, Zerynthia polyxena, Iphidides podalirius,
Fixsenia pruni, Lycaena dispar. In addition two species are listed as endan-
gered: Colias chrysotheme and Maculinea teleius. Zerynthia polyxena, Papilio
machaon and Iphiclides podalirius are protected by law. The greatest number
of Lepidoptera occurred on site 7 where not only the xerothermic species
occur, but many hygrophilous species immigrate from the nearby wet habi-
tats as well.

According to an ecological classification of lepidopterous species given
by Blab and Kudrna (1982) , Koch (1984) and original data, the species spec-
trum shows that xerothermophilous species predominate in the investigated
region (40.5%) (Table 1). This appears to be a result of the study region,
altitude 138-145 m, with poor sandy soil, being located in the warmest re-
gion of Slovakia (Kulfan &c Kulfan 1992). Some Lepidoptera found are
known to prefer sandy soil, e.g., Lythria rotaria. Many butterflies, as Minois
dryas and Colias chrysotheme fly to these sites from nearby habitats of xero-
thermic character with the forest-steppe vegetation, especially from the
adjacent slopes of Devinska Kobyla, a part of Little Carpathian mountains.

The following species found in this study are useful indicators: Zerynthia
polyxena, Iphiclides podalirius, Colias chrysotheme, Melanargia galathea, Minois
dryas, Erynnis tages, Agriphila inquinatella, Loxostege sticticalis, Zygaena loti, Z.
angelicae, Scopula virgulata, S. immutata, Idaeas^^. 3.nd Euclidia glyphica. These
species indicate relatively well preserved xerothermic habitats with forest-
steppe vegetation characterized by Crataegus over uncut grassland on slightly
elevated places as gravelly and sandy alluvial sedimentary outcrops, exem-
plified by site 7.

Heteropterus morpheus, Colias hyale, Lycaena dispar, and Maculinea teleius in-
dicate extensive wet meadows, cut twice annually and frequently flooded,
as site 3.

Psammotis pulveralis, Elophila nymphaeata, Coenonympha glycerion, Lycaena
tityrus, and Phlogophora meticulosa indicate boggy habitats at sites 1, 3, 4, 5,
and 9.

Two species from the family Nymphalidae, Cynthia carduidccid Aglais urticae,
reached their highest abundance on sites 8 and 9 that were modified by
extensive application of liquid manure and by cattle grazing. The abundance
of flowering weeds on site 9 especially attracted adults of these butterflies.

Acknowdgements. This research was supported by the Ministry of Education and
Science, grant No. 1/990709/93 “Zoocoenoses of characteristic biotopes in sur-
roundings of the rivers Danube and Morava” and grant No. 1/1141/94 “Conserva-
tion and Use of the Gene Pool of the Fauna of Slovakia.” We wish to thank two
reviewers for valuable comments and Rudi Mattoni for final editing of the manu-
script.

34:39-47, 1995(1997)

Literature cited

Blab, J. & O. Kudrna. 1982. Hilfsprogramm fur Schmetterlinge. Kilda-Verlag,
Greven, 136 pp.

Erhardt, a. 1985. Wiesen und Brachland als Lebensraum fur Schmetterlinge.
Birkhauser Verlag, Basel, Boston, Stuttgart, 154 pp.

Koch, M. 1984. Wir bestimmen Schmetterlinge. Neumann Verlag, Leipzig,
Radebeul, 792 pp.

Kulfan, J. 1989. Zur Bionomie des Blaulings Everes decoloratus (Stgr.) (Lep.,
Lycaenidae) bei nordwestlicher Grenze ihrer Verbreitung. Biologia (Bratislava)
4:177-184.

. 1990a. Die Struktur der Taxozonosen von heliophilen Faltern (Lepidoptera)
an manchen Biotopen der Westslowakei. Biologia (Bratislava) 45:117-126.

. 1990b. Sezonna dynamika spolocenstiev heliofilnych motylov na troch lokal-
itach zapadneho Slovenska (Lepidotera) . Entomol. Probl. 20:177.

Kulfan, M. & H. Kalfvoda. 1994; Zygaenoidea, Hesperoidea, and Papilionoidea of the
Morava river alluvia (downstream region). Ecology (Bratislava) 13-Supplement
1:165-173.

Kulfan, M. &J. Kulfan. 1992. Changes of distribution of thermophilous butterflies
in Slovakia. J. Res. Lepid. 29:254-266.

PoDANi, J. 1993. SYN-TAX - pc. Computer Programs for Multivariate Data Analysis
in Ecology and Systematics. User’s Guide, Scientia Publishing, Budapest, 104

pp.

Ruzickova, H. 1994. Wiesenvegetation des Inundationsgebietes des Unterlaufes des
March-Flusses sudlich von Vysoka pri Morave. Ecology (Bratislava) 13-Supple-
ment 1:89-98.

Journal of Research on the Lepidoptera

34:48-68, 1995(1997)

Effectiveness of caterpillar defenses against three species of
invertebrate predators

Lee A. Dyer^

Department of EPOB, Campus Box 334, University of Colorado, Boulder, Colorado 80309

Abstract. The efficacies of larval defenses against invertebrate predators
representing different (but overlapping) foraging guilds were compared
by offering 34 species (287 individuals) of lepidopteran larvae to
Paraponera clavata ants, Apiomerus pictipeshug^, and Polistes wasps.

Overall, the ants were the most likely to eat caterpillar prey, and the wasps
were the most cautious. Larval chemistry and diet breadth were signifi-
cant predictors of rejection by the group of predators; chemically de-
fended specialist herbivores were better protected than generalist herbi-
vores without known chemical defenses. These results provide evidence
for the potential importance of predators in maintaining diet breadth of
phytophagous insects, and they suggest that plant chemistry is part of a
mechanism for restricting diet breadth. Other important larval defenses
included size, morphology, and coloration. Large prey (heavier than 1
g) were less acceptable than smaller prey (lighter than 200 mg) for the
wasps and bugs but not for the ants; hairs deterred predation by the ants
and bugs but not by the wasps; and brightly colored caterpillars were fre-
quently rejected by the wasps but not by the ants and bugs.

Key Words, Caterpillars, defenses, diet breadth, predation, Apiomerus
pictipes, Polistes instabilis, Paraponera clavata, plant secondary compounds,
Lepidoptera, specialization, tropics

Introduction

Faced with a deluge of special cases that often appear to be a morass of
contradictions and confusion, ecologists frequently attempt to generalize
about predominant forces or patterns that are manifested by specific ex-
periments and observations. A few prominent examples in research on Lepi-
doptera include: attempts to use specific studies of oviposition patterns
(particularly those of lepidopteran pest species) to construct a diet-choice
theory for all phytophagous insects (e.g., Courtney & Kibota 1990) ; attempts
to identify the most important processes which organize communities by
examining particular systems of predators, herbivores and plants (e.g.,
Karban 1989); and attempts to explain the high incidence of dietary spe-
cialization in lepidopteran larvae by examining known feeding patterns
(e.g., Ehrlich & Raven 1964). These generalizations, which arose from

'Current address: La Selva Biological Station and Biology Dept., Mesa State College, Grand
Junction, CO 81502

Paper submitted 31 October 1995; revised manuscript accepted 10 May 1996.

34:48-68, 1995(1997)

multiple-species pattern analyses as well as from reductionist (single-spe-
cies or single-system) approaches, have contributed significantly to a theo-
retical framework for a large number of studies on Lepidoptera.

In light of these and many other studies which provide good data for gen-
eralizations, it is surprising that there is a dearth of attempts to character-
ize important components of lepidopteran larval defenses, either by con-
ducting multiple-species experiments or through literature reviews (Witz
1990, Dyer Sc Floyd 1993, Dyer 1995; also for adult Lepidoptera see Maclean
et al. 1989). While multiple-species approaches are generally not as thor-
ough as experiments examining the effectiveness of a particular defense in
one species, they allow for different generalizations on insect defenses which
can ultimately provide a framework for both basic and applied research
questions with specific systems. For example, Bernays and Cornelius (1989)
demonstrated that a number of species of leaf rollers were extremely palat-
able to ants; their generalization that trade-offs could exist between chemi-
cal defense and concealment from predation provides an impetus for quan-
titative genetics experiments examining the potential for such trade-offs in
specific systems.

Two important groups of general hypotheses about larval defenses which
I attempt to address in this study are: 1) hypotheses about the effectiveness
of a suite of defenses against specific predatory guilds or against single spe-
cies, and 2) hypotheses about effectiveness of specific defenses against a
suite of predatory guilds or against multiple species. A related question
which I address involves generalizations about the importance of biotic in-
teractions (particularly natural enemies: Brower 1958, Bernays Sc Graham
1988) and plant chemistry (Dyer 1995) in influencing herbivores’ diet
breadths. Recent studies have demonstrated that certain specialist herbi-
vores are better protected than certain generalists against various inverte-
brate predators (Bernays 1988, Bernays Sc Cornelius 1989, Dyer Sc Floyd
1993, Dyer 1995), which suggests that natural enemies could be important
in the maintenance of narrow diet breadth or could be a selective force in
the evolution of dietary specialization. In this study, I further test this natu-
ral enemy hypothesis, and I also explore the possibility that plant chemis-
try mediates the evolution of differences in defensive capacity between spe-
cialists and generalists. Chemistry might provide such a mechanism if spe-
cialized herbivores tend to evolve the ability to sequester plant defensive
chemicals and use them as defenses against their natural enemies.

In order to generalize about the efficacies of various defenses against dif-
ferent predators and about the importance of biotic interactions and plant
chemistry in the evolution of dietary specialization, I offered specialist and
generalist lepidopteran larvae (caterpillars) with a wide variety of potential
defensive qualities to three predators representing different predatory
guilds: an assassin bug, Apiomerus pictipes (Reduviidae); a paper wasp, Polistes
instabilis (Vespidae); and the giant tropical ant, Pamponera clavata (Form-
icidae). Specifically, I asked the following questions: 1) Are different preda-
tor guilds deterred by different types of defenses? 2) What are the most ef-

J. Res. Lepid.

fective defensive mechanisms of lepidopteran larvae against a suite of preda-
tors? 3) Are specialist herbivores better protected than generalists against
a suite of predators? 4) Are noxious prey chemicals effective defenses against
a suite of predators?

The prey used in my experiments were larvae in 13 different families of
Lepidoptera that were native to a variety of micro-habitats in Costa Rica,
These caterpillars exhibited a wide variety of antipredator mechanisms
which could be compared. Apiomerus pictipesis a common sit-and-wait preda-
tor that ranges from Colorado (USA) to Columbia (Johnson 1983). It is
solitary, visually oriented, and quickly kills prey by inserting its mouthparts
and sucking, leaving behind a dry carcass. Polistes instabilis is a common for-
aging predator found from Costa Rica to Southern Brazil and Argentina
(Richards 1978). It is a solitary (i.e., it does not recruit), visually-oriented
predator, and it kills prey by biting rather than stinging. Each wasp exten-
sively chews the prey before returning to the nest to distribute ingested flu-
ids and solid caterpillar remains to the other adults and larvae (West-
Eberhard 1983). Paraponera clavata is a foraging predator common in low-
land rainforests and found from Nicaragua to the Amazon (Janzen & Carroll
1983). It is a chemically-oriented predator that forages independently or
in groups and that kills prey by using its powerful sting, using its mouth-
parts and cooperating with nestmates (pers. obs.). All three predators com-
monly prey on caterpillars (pers. obs., West-Eberhard 1983, Johnson 1983)
and are sympatric with all the caterpillars used in my study.

Materials and Methods

I conducted all experiments and most collecting in June and July, 1993 at the
following sites in Costa Rica: Palo Verde National Park, Lomas Barbudal National
Park, and private land near Lomas Barbudal. These sites are located in the
Guanacaste province of Costa Rica and are characterized by dry forest (sensu
Holdridge et al. 1971) and marsh (at Palo Verde).

Collecting

I collected most caterpillars at Palo Verde and Lomas Barbudal. I also bought
several species of caterpillars that are known to occur in Guanacaste from Finca
Mariposa, a commercial butterfly farm in La Guacima.

I either identified caterpillars to the lowest taxon possible using Stehr (1993), or
if sufficient numbers of caterpillars were available, I reared them to the adult stage
for identification. I deposited voucher specimens of most caterpillars and adults at
both the Instituto Nacional de Biodiversidad (INBio) in Costa Rea and the Uni-
versity of Colorado Entomology Museum, Boulder (Table 1).

For host plant data, I identified plant families on which I found caterpillars and
held the caterpillars in captivity for several days to verify that they actually were
using their presumed host plants as food resources. If possible, I collected enough
plant material for identification to lower taxa by park naturalists or I dried and
pressed them for identification by other tropical botanists. Voucher specimens for
some host plants are at the University of Colorado Herbarium, Boulder (Table 1).

34:48-68, 1995(1997)

Table 1a. Generalist caterpillars offered to P. clavata, P. versicolor, and A.
pictipes and the host plants upon which the caterpillars were found.

Caterpillar®

Arctiidae
(5, 5, 3)
CU:LS93GAT

Arctiidae

(5,5)

CU:PV93AWB

Eois sp.

(Geometridae)
(3, 3)

CU:TBG92
Pero sp.

(Geometridae)
(5, 3,5)
CU:LS93ATB

Geometridae
(5, 3, 5)
CU:BTB92

Gonodonta sp.
(Noctuidae)
(5, 3, 3)
IN:GON92

Pantographa limata
(Pyralidae)

(5, 3, 5)
CU:LS93SIM

Predators’’ Host Plants*"

A(Y), B(Y), W(Y) (Annonaceae)*

Co5^M5sp. (Costaceae)

Siparuna pauciflora (Monimiaceae)
Welfia georgii (Palmae)

Adiantums^. (Polypodiaceae)
Myriocarpa longipes (Urticaceae)*

A(N) , B (Y) Protium panamense (Burseraceae)

(Compositae)

Hemandiasp. (Hernandiaceae)
Nectandra hypoleuca (Lauraceae)
Colubrina spinosa (Rhamnaceae)
(Rubiaceae)

A(N),B(N) (Araceae)*

(Leguminosae)

Piper urostachyum (Piperaceae)

Sabicea sp. (Rubiaceae)

A(N), B(N), W(N) (Annonaceae)*

Diffenbachia sp. (Araceae)

Costus sp. (Costaceae)

Erythrina sp. (Leguminosae)
Pentadethra macroloba (Leguminosae)
Hampea appendiculata (Malvaceae)

A(N), B(N), W(N) Richeria dressleri (Euphorbiaceae)
Ardisia sp. (Myrsinaceae)
Passifloraspp. (Passifloraceae)*
Colubrina spinosa (Rhamnaceae)
Citrus spp. (Rutaceae)

(Solanaceae)

(Violaceae)

A(N),B(N),W(N) (Brassicaceae)

TASTY (Compositae)

Wissadula excelsior (Malvaceae)
Calathea sp. (Marantaceae)
Pithecellobium sp . (Mimosaceae)

Eicus sp. (Moraceae)

Solanum sp. (Solanaceae)

Myriocarpa longipes (Urticaceae)*

Co5^M5 sp. (Costaceae)

Manihot esculenta (Euphorbiaceae)

H amelia patens (Rubiaceae)

Paullinia pterocarpa (Sapindaceae)
Cestrum sp. (Solanaceae)*
(Solanaceae)

Goethalsia meiantha (Tiliaceae)*
Luehea seemannii (Tiliaceae)*
Myriocarpa longipes (Uritcaceae)*

A(N),B(N),W(N)

TASTY

/. E£s. Lepid.

Antheraea polyphemus AiN) ,W (Y)
(Saturniidae)

(3, 3)

IN:SAT92

Automeris rubrescens A ( N ) ,W (Y)

(Saturniidae)

(3, 3)

CU:PV93IOT

Automeris zugana A(N) ,W(Y)

(Saturniidae)

(5, 3)

CU:93IOM

Citheronia lobesis A ( N ) ,B (Y) ,W (Y)

(Saturniidae)

{5, 3, 3)

CU:PV93HHD

Godmania aesculifolia (Bignoniaceae)*
Solanumsp. (Solanaceae)*

Luehea sp. (Tiliaceae)*

[Plus 18 additional families reported in Tietz
1972]

Cordia alliodora (Boraginaceae)

Rourea glabra (Connaraceae)

Cas5wsp. (Leguminosae)*

Inga&p. (Leguminosae)

[Plus 5 additional families reported in Janzen
1984]

Cydista heterophylla (Bignoniaceae)*

Cassia sp. (Leguminosae)*

Lonchocarpus sp. (Leguminosae)*
(Sapindaceae)

Solanum hazenii (Solanacae)

[Plus 2 additional families reported in Janzen
1984]

(Anacardiaceae)

Cydista heterophylla (Bignoniaceae)*

Godmania aesculifolia (Bignoniaceae)*
Cochlospermum vitifolium (Cochlospermaceae)*
[Plus 4 additional families reported in Janzen
1984]

Erinnyis ello
(Sphingidae)
(4, 3)

CU:PV93ELL

A(N),W(Y) (Bignoniaceae)

Manihot esculenta (Euphorbiaceae)

Sapium sp. (Euphorbiaceae)

Cissus microcarpa (Vitaceae)

[Plus 2 additional families reported in Tietz,
1972 and 1 additional family reported in
Janzen, 1984]

Table 1b. Specialist caterpillars offered to P. ciavata, P. versicolor, and A.
pictipes and the host plants upon which the caterpillars were found.

Caterpillar®

Predators'*

Host Plante‘S

Euchaetes sp.
(Arctiidae)

(3,3)

CU:PV93BOA

B(Y),W(Y)

Asclepias curassavica (Asclepiadaceae)*

Arctiidae

(3)

CU:PV93MHA

W(N)

Cydista heterophylla (Bignoniaceae)*

Hesperiidae

(3)

IN:PV93HES

W(Y)

Solanumsp. (Solanaceae)*

Limacodidae

(3,3)

B(Y),W(Y)

Quercus oleoides (Fagaceae)

34:48-68, 1995(1997)

Lymantriidae B(Y),W(N) Cassias^. (Leguminosae)*

(3, 3)

CU:PV93040

Lymantriidae B(Y),W(Y) (Bignoniaceae)*

(3, 3)

IN:LS92LYM

Megalopygidae B (Y) , W (Y) Ceiba pentandra ( Bombacaceae )

(3, 3)

CU:PV93MGT

Azeta versicolor A(N)3(N),W(N) Siparuna paucijiora {Monimi2Lce3.G)

(Noctuidae)

(3, 3, 3)

CU:PV93010

Diphthera f estiva W(Y) 5o/anwmsp. (Solanaceae)

(Noctuidae)

(3)

IN:PV93NOC2

Caligo memnon memnon A(Y),B(Y),W(Y) Heliconia imbricata (Heliconiaceae)
(Brassolinae) NASTY

(5, 3, 5)

CU:LS93CAL

Agraulis vanillae W (Y) Passiflora sp. (Passifloraceae) *

(Nymphalinae)

(3)

CU:LS93AGV

Aeria eurimedia agna A(N),B(N),W(Y) (Apocynaceae)

(Ithomiinae)

(4, 3, 3)

CU:LS93AEA

Morpho peleides limpida A(Y) 3 (Y) ,W(Y) Lonchocarpus oliganthus (Leguminosae) *

(Morphinae) NASTY

(5, 3, 4)

Adelpha fessonia A(N)3(N),W(N) Randia armata {Ruhid.ce3.e)*

(Nymphalinae) TASTY

(5, 3, 4)

CU:LS93ADF

Marpesia petreus A(N)3(Y),W(Y) (Anacardiaceae)

(Nymphalinae)

(5, 3, 3)

Papilio cresophantes A(Y)3(Y),W(Y) Citrus limon (Rut3.ce2Le)*

(Papilionidae) NASTY

(5, 3, 3)

CU:LS93PAC

Papilio anchisiades idaeus A(Y)3(Y),W(Y) Citrus limon (Rutaceae)*

(Papilionidae) NASTY

(5, 3, 5)

CU:LS93PAP

J. Res. Lepid.

Anteos clorinde
(Pieridae)

(5, 3, 3)
CU:LS93ANT

A(N),B(N),W(N) Cassia fruticosa (Leguminosae)
NEUTRAL

Pyralidae
(3, 3)

CU:PV93FNT

B(Y),W(Y)

Bombacopsus quinatum (Bombacaceae)

Saturniidae

(3, 3)

CU:PV93013

A(Y),W(Y)

Ceiba pentandra (Bombacaceae)

Manduca sexta
(Sphingidae)
(5, 3, 3)

A(Y),B(Y),W(Y) Solarium (Solanaceae)

CU:LS93THW

Sphingidae

(3)

CU:PV93003

W(Y)

Piper (Piperaceae)

‘'Those species that I could not identify past the family level are identified by the family. Sample
sizes (number of caterpillars offered to ants, bugs, and wasps, respectively) are indicated in
parentheses underneath each species. Voucher specimen codes are included under those
species for which I had appropriate replicates to keep a voucher. CU = University of Colo-
rado Entomology Museum, Boulder; IN = Instituto Nacional de Biodiversidad, Costa Rica.
‘’Not all caterpillar species were available for all predators; this column indicates to which
predators each species was offered. A = ants, B = bugs, W = wasps. The predators’ average
responses (rejection) are indicated in parentheses after the letter indicating the predator. Y
= Rejected more than half of the time, N = Not rejected more than half of the time. If extract
data were available, the level of the “chemistry” category is also included in this column.
‘^Caterpillars were reared on host plants on which they were found. Although there are other
reported hosts for some species, none of them are known to feed on plants in more than 2
families. Asterisks (*) indicate those species for which voucher specimens are available at the
University of Colorado Herbarium, Boulder.

Experiments with wasps

I offered caterpillars to wasps throughout the day and at 5 different sites. Three
of the sites were areas where wasps were frequently found foraging along the side
of a dirt road within and just outside of Lomas Barbudal. The other two sites were
two different trees in Palo Verde which contained many wasp nests. Caterpillars
offered to wasps foraging along the road were placed on the ground amidst vegeta-
tion where the wasps were foraging. Caterpillars offered to wasps in the trees were
placed in small, clear plastic cups which were suspended with string from branches
of the trees. If the caterpillars crawled out of the cups before being encountered
by a wasp, they were placed back into the cup. Depending on availability, I offered
3-5 individuals of each caterpillar species (32 species) to the wasps (see Table 1),
and each site received only 1 individual of each species. Each caterpillar was inde-
pendently offered at a different spot along the road or in the tree, and the order of
presentation was haphazard (often depending on when caterpillars were found).
No site received more than 4 caterpillars in a single day.

34:48-68, 1995(1997)

I observed all caterpillar=wasp interactions until either most of an entire caterpil-
lar was carried away by wasps or at least 3 wasps had encountered and rejected it.
Rejections consisted of a wasp approaching the caterpillar and either touching it
or coming within about 20 cm without attacking; 20 cm is a distance which is well
within the field of vision of wasps (Spradbery 1973).

Experiments with bugs

Fourteen assassin bugs were collected at the 3 experimental sites and were kept
in 17 cm X 13 cm X 7 cm plastic boxes containing paper towels and twigs. When
not being used for experiments, twice a week the bugs were fed a drab, glabrous
noctuid caterpillar (voucher: PV93NOT at the University of Colorado Museum,
Boulder) which was abundant and which was palatable to various wasps, ants and
mantids (pers. obs.).

Three to five replicates of 24 species of caterpillars were offered to the bugs (Table
1 ) , and no bug received more than 6 total caterpillars or more than 1 replicate per
species. Caterpillars were placed in the plastic boxes containing bugs and were left
with the bugs for 24 hours. The bugs would either attack the caterpillar within an
hour or they would ignore it, which constituted a “rejection.”

Experiments with ants

Data were used from caterpillars offered to P. clavata in a larger study (Dyer 1995) .
In that study, caterpillars were offered to 5 ant colonies, and the numbers of ants
(within a colony) rejecting individual caterpillars were classified into the catego-
ries “no rejections” (0 ants rejecting the caterpillar), “some rejections” (fewer than
7 ants rejecting the caterpillar), and “completely rejected” (8 or more ants reject-
ing the caterpillar). Because the wasps and bugs either rejected or accepted prey
as opposed to having inconsistent responses within a colony (hence, “some rejec-
tions”), I reclassified the ant rejection category to make it comparable to data for
the bugs and wasps. Caterpillars receiving fewer than 5 rejections were considered
to be “not rejected,” while caterpillars receiving 5 or more rejections were consid-
ered to be “rejected.” This was an arbitrary categorization, but it effectively split
the “some rejections” category in half and made the P, clavata responses compa-
rable to data for the bugs and wasps.

Statistical analyses

I scored each caterpillar species for the following categorical variables: a) cater-
pillar diet breadth — generalist or specialist; b) caterpillar coloration — brightly
colored, visually cryptic, or other; c) caterpillar morphology — spines, hairs, or
glabrous; d) caterpillar size — small, medium, or large; and e) caterpillar chemis-
try — palatable extract or deterrent extract.

For the diet breadth variable, I used a taxonomic definition of specialization.
Caterpillars known to feed on fewer than 2 families of plants (according to Tietz
1972, Janzen 1984, DeVries 1987, Marquis 1991, and personal communication with
various naturalists), or caterpillars of unknown diet breadth that were found feed-
ing on only 1 plant species, were classified as “specialists” (22 species; 9 of which
had unknown diet breadths) . Since most herbivores at La Selva are monophagous

J. Res. Lepid.

or oligophagous (Marquis and Braker 1994), I assumed that it was unlikely that
unknown caterpillars would be erroneously classified as specialists. Caterpillars
found feeding on plants in greater than 3 families were classified as “generalists”
(12 species; most fed on plants in greater than 6 families).

The coloration and morphology variables were based on visual inspection of the
caterpillars. “Spiny” caterpillars had sclerotized spines at least 2 mm long. Cater-
pillars with hairs or with hairs and spines were rated as “hairy” only if more than
50% of their cuticle was covered with secondary setae that were at least 5 mm long.
“Glabrous” caterpillars had no hairs or spines.

The size statistic was based on the weight (in mg) of a caterpillar just before it
was offered to a predator. Levels of size categories were: “small” (weight < 200 mg),
“medium” (200 mg < weight < 1000 mg), and “large” (weight > 1000 mg). The size
categories were pooled in the preceding manner based on examination of a fre-
quency histogram of all the weights.

For the chemistry variable I used results from a bioassay done with crude cater-
pillar extracts offered to P. clavata (Dyer 1995). Data for this variable were only
available for 8 caterpillar species (see Table 1). The levels of this variable were:
“nasty” (caterpillars with deterrent extracts) , “tasty” (caterpillars with extracts which
attracted ants), and “neutral” (caterpillars with neutral extracts). This variable was
included to examine the defensive efficacy (against all three predators) of chemi-
cals found in caterpillars without the confounding effects of morphological and
behavioral features.

I used logit analyses to study the relative importance of these caterpillar charac-
teristics as determinants of predator rejections (see Christensen 1990 for a thor-
ough discussion of logit models) . All of the caterpillar characteristics which I ex-
amined may act as important anti-predatory traits (reviewed by Edmunds 1974,
DeVries 1987, Evans and Schmidt 1990). For all logit models I used the maximum
likelihood method for parameter estimation of linear models and Chi-square sta-
tistics for hypothesis testing (see SAS 1990). All of the models were nonhierarchical
because I either obtained significant highest-order associations in the saturated
models, or because I had specific hypotheses that I wanted to test. Since the mod-
els were nonhierarchical, I used the Newton-Raphson algorithm for parameter es-
timation and model testing (SAS 1990). I assigned values of 1 X 10'^° to cells that
contained “sampling zeroes” (sensu Bishop et al. 1975), while cells that contained
“structural zeroes” (sensu Bishop et al. 1975) were automatically deleted (see SAS
1990).

To avoid running a large model containing many cells with zeros or small values,
it was necessary to use more than one model. I chose variables for models that ad-
dressed specific questions which I wanted to ask with my experiments; in addition,
examination of frequency tables for all combinations of variables helped form de-
cisions for appropriate models (see Tabachnick and Fidell 1989). Variables that
were not significantly associated with rejection in 2-dimensional frequency tables
(using a conservative criterion of P < 0.001 because of the large number of tests)
were not included in the models.

I ran two logit models which included data from all the predators. Model 1 ad-
dressed these questions: 1) Are caterpillars’ levels of rejections dependent on the

34:48-68, 1995(1997)

Table 2. Summary of two-dimensional tables with predictors versus rejections.

Predator

Predictor

ALL

Chemistry

84.5

0.000

(n=287)

Diet Breadth

29.6

0.000

Predator

34.9

0.000

Size

31.6

0.000

Morphology

25.3

0.000

Coloration

10.0

0.007

ANTS

Chemistry

36.2

0.000

(n-103)

Diet Breadth

17.8

0.000

Size

1.9

0.386

Morphology

11.3

0.003

Coloration

0.97

0.617

BUGS

Chemistry

24.0

0.000

(n^76)

Diet Breadth

4.5

0.035

Size

28.7

0.000

Morphology

29.5

0.000

Coloration

2.5

0.297

WASPS

Chemistry

25.4

0.000

(n=108)

Diet Breadth

4.5

0.033

Size

24.6

0.000

Morphology

0.24

0.885

Coloration

16.7

0.000

type of predator?, 2) Are the presence of unpalatable chemicals in caterpillars likely
to make predators reject them more frequently?, 3) Which predictor of rejection
(chemistry or predator) is more reliable?, and 4) Are there interactions between
rejections, type of predator, and extract palatability? Model 2 addressed these ques-
tions: 1) Are specialists rejected more frequently than generalists against a variety
of predators?, 2) Is diet breadth a better predictor of rejection than type of preda-
tor?, and 3) Are there interactions between rejections, diet breadth, and type of
predator?

For the wasps and bugs I also ran a logit model for data specific to each preda-
tor. Each model asked questions about associations between caterpillar character-
istics and rejections by the predator. The bug model included morphology and size
as predictors. The wasp model included coloration and size as predictors. Models
examining predictors of ant rejections are reported elsewhere (Dyer 1995).

Results

I used results from 108 individuals of 32 caterpillar species offered to the
wasps and results from 76 individuals of 24 caterpillar species offered to
the bugs. For the ant data, I only used data for caterpillar species that were
also offered to either the wasps or bugs; this subset of the data included

J. Res. Lepid.

Table 3. Summary of log-linear models.

ModeP

Likelihood ratio
probability'"

Models using all predators

1. Chemistry (5.38 ***)

0.589

2. Diet Breadth (-5.67 ***)

Predator (5.56 ***)

Model for bugs

0.905

3. Size (3.44 ***)

Morphology (3.0 *)

Model for wasps

0.594

4. Size by Coloration (-2.85 ***)
Coloration (2.56 *)

0.290

'‘The variables shown are significant predictors of rejections from the most parsimonious
model that fit the data. Predictor variables were ranked by standardized parameter estimates,
which are given in parentheses along with asterisks to indicate significance of the estimate
(* denotes p < 0.05, ** denotes p < 0.01, *** denotes p < 0.005).

'The likelihood ratio probability is a goodness-of-fit test for the overall model, and p-values
above 0.05 indicate a good fit (SAS 1990). P-values reported here are for the most parsimo-
nious models.

103 individuals of 23 caterpillar species, values from the 2-dimensional
tables of defenses and rejections are summarized in Table 2.

For the combined predators, Model 1 revealed a signibcant = 36.0,
DF = 2, P < 0.0001) chemistry effect on rejections. Chemistry was a more
reliable predictor than type of predator, which was not significant (x^ = 0.41,
DF = 2, P = 0.814; Table 3). The predators rarely rejected caterpillars with
neutral and tasty extracts, while 98% of caterpillars with nasty extracts were
rejected (Fig. 1). There were no significant interactions between chemistry
and predators — all predators were deterred by the caterpillars with observed
chemical defenses.

Model 2 for the combined predators revealed a significant association
between predator and rejection (x^= 34.68, DF = 2, P < 0.0001) and an as-
sociation between diet breadth and rejection (x^ = 28.86, DF = 1 , P < 0.0001 ) .
Predator was a more reliable predictor than diet breadth (Table 3) and there
were no interactions between the two predictors. Ants were the most likely
to eat caterpillars, and the wasps were the most cautious (Fig. 2). The asso-
ciation between diet breadth and rejection reflects the fact that specialists
were rejected more frequently than generalists (Fig. 3).

The logit model for the bugs revealed a significant association between
size and rejection (x^ = 11.95, DF = 2, P = 0.0025) and a significant associa-
tion between morphology and rejection (x^ = 8.78, DF = 2, P = 0.012). Size
was a more reliable predictor than morphology (Table 3) and there were
no interactions between the two predictors. As the mean size of caterpillars
increased, the levels of rejection also increased (Fig. 4), Caterpillars with

34:48-68, 1995(1997)

hairs were rejected more frequently than those with other morphologies
— particularly caterpillars with spines which were never rejected (Fig. 5).

The logit model for the wasps revealed a significant interaction between
size, coloration, and rejection (x^ = 17.43, DF = 2, P = 0.0002) and a signifi-
cant association between coloration and rejection (x^ = 8.05, DF = 2, P =
0.0179). The interaction was a more reliable predictor than coloration
(Table 3) . Brightly colored caterpillars of all sizes were better protected than
caterpillars with other colorations (Fig. 6) ; however, if the caterpillars were
large, their coloration was not important (100% of the large caterpillars
were rejected by the wasps — Fig. 4).

Discussion

To some extent, the predators evaded characterization by generalizations
such as, “hairs are a good defense against invertebrate predators.” Preda-
tors varied in their propensities to reject, and each predator was influenced
by a different assemblage of caterpillar defenses. However, there were re-
sults which can be generalized for a variety of invertebrate predators (based
on the wide behavioral and taxonomic differences between the three preda-
tors) and results that can be generalized for specific predatory guilds rep-
resented in this study.

The best generalizations about caterpillar defenses against invertebrate
predators come from examining the results of the models that included all
predators. Chemistry and diet breadth were both important predictors of
rejections when considering the suite of predators and when including the
variation in predators’ inclination to reject prey. Specialists and caterpil-
lars with deterrent extracts were rejected more frequently than other cat-
erpillars by the predators, and since these predators represent very differ-
ent guilds, it may be reasonable to conclude that these qualities would pro-
tect caterpillars against many different types of invertebrate predators. Ex-
amples of the guilds that were covered by these predators include: solitary
predators (P. instabilis diXid A. pictipes), recruiting predators (P. clavata), sit-
and-wait predators (A. pictipes), flying predators (P. instabilis), visually ori-
ented predators (P. instabilis ?Lnd A. pictipes), chemically oriented predators
(P. clavata) , sucking predators (A. pictipes) , chewing predators (P. instabilis) ,
stinging predators (P. clavata), nocturnal predators (P. clavata), and diur-
nal predators (P. instabilis and A. pictipes). One caveat to broad interpreta-
tion of these results is that these predators are not necessarily representa-
tive of their foraging guilds; P. clavata, for example, is much more likely to
indiscriminately accept prey than other members of the tribe Ectatommini
(Dyer and Eolgarait, unpub. data). Thus, the results from this study do not
indicate that prey protected against these three predators should be equally
protected against any representatives of their respective foraging guilds,
rather they illustrate the effectiveness of narrow diet breadth and defen-
sive chemistry against very different types of predators.

The importance of diet breadth and chemistry as predictors of rejection
for this group of predators are also consistent with another generalization:

/ Res. Lepid.

EXTRACT PALATABILITY

Fig. 1 . The association between palatability of caterpillars’ chemical extracts and
percentage of rejections by all the predators. The y-axis represents the
percentage of individual caterpillars (n = 96 individuals; 8 species) with
specific palatabilities that were rejected by all 3 predators (ants, bugs, and
wasps). The numbers above each bar indicate the sample size; the num-
ber of caterpillars rejected is in the numerator, and the total number of
caterpillars offered (with that particular palatability) is in the denominator.

both predation and plant chemistry could affect herbivores’ diet breadth.
A scenario by which this could happen is as follows: 1) An herbivore over-
comes a specific plant defense and in the process loses access to other plants
because of trade-offs in physiological abilities to utilize plants with differ-
ent chemical compounds (Ehrlich & Raven 1964). 2) As the herbivore be-
comes more specialized as a result of step 1, it also sequesters secondary
compounds either casually (because it is eating fewer plants; Jones et al.
1989) or because of specific physiological adaptations (Bowers 1990). 3)
Specialization is further maintained by predators because specialists are
better chemically protected than more generalized herbivores (Dyer 1995).
Steps 2 and 3 are consistent with results from these experiments because
herbivores with specialized diets were better protected against a group of
predators, chemistry was an important component of their defense, and
there is evidence that some of the species used in my experiments seques-
ter noxious compounds from their host plants (Dyer 1995).

The results unique to specific predators reveal prey preferences that could
be common responses for their respective guilds. Size, for example, was

34:48-68, 1995(1997)

I —

Fig. 2. Percentages of all caterpillars rejected by each predator. The y-axis rep-
resents the percentage of individual caterpillars (n = 287 individuals; 34
species) rejected by each of the 3 predators (ants, bugs, and wasps).
The numbers above each bar indicate the sample size; the number of
caterpillars rejected is in the numerator, and the total number of caterpil-
lars offered (to that particular predator) is in the denominator.

important for the two solitary predators (the bugs and wasps) but not for
the ants which could recruit other ants and easily subdue larger prey items.
It is generally assumed that prey size is an important limitation for inverte-
brate predators (Cohen et al. 1993, Reavey 1993), but this assumption may
vary with the degree of predators’ social cooperation. The differences be-
tween the solitary (wasps and bugs) and recruiting (ants) foragers were
actually quite dramatic; the bugs and wasps barely touched large prey items
(many of which were generalists which probably were not otherwise de-
fended very well) , while the ants attacked them as voraciously as caterpil-
lars of any other size. The size categories were not ambiguous, in that all of
the predators would be in the same category as the “small” prey, since their
mass varies from 50-200 mg, while the “large” caterpillars had masses over

9g-

Hairs were an important deterrent for the bugs which have mouthparts
specialized for sucking. Hairs are probably a significant deterrent for most
hemipteran predators because they prevent insertion of a bug’s proboscis
(pers. obs., also see Bowers 1993) . Alternatively, hairs may function by warn-

J. Res. Lepid.

Q
LU
I —

—3

114/167

SPECIALISTS GENERALISTS

DIET BREADTH

Fig. 3. The association between diet breadth and percentage of rejections by all
the predators. The y-axis represents the percentage of all generalist or
specialist caterpillars (n = 287 individuals; 34 species) that were rejected
by the 3 predators (ants, bugs, and wasps). Although not all caterpillar
species were offered to all predators, the predators are treated as a group.
The numbers above each bar indicate the sample size; the number of
caterpillars rejected is in the numerator, and the total number of caterpil-
lars offered (with that particular diet breadth) is in the denominator.

ing the caterpillar of a predator’s advance before it actually has a chance to
catch the caterpillar (Tautz & Markl 1978). Indeed, many hairy caterpillars
(particularly arctiids) are fast, and an “early warning system” such as hairs
extending far from the body may make it difficult for sit-and-wait predators
to successfully attack them. It is not as clear, however, why the hairs were
effective against the ants but not against the wasps. One explanation could
be that wasps, which can fly and are more maneuverable, are able to pluck
hairs (without the prey escaping) from caterpillars more effectively than
ants (Bowers 1993). I have observed wasps and ants plucking hairs from
prey with varying degrees of success.

Coloration was important for wasps, but it was not important for ants which
are often chemically oriented. Wasps were deterred by brightly colored prey
which supports general theories about aposematism (see Cott 1940). Bugs,
on the other hand, which are also visually oriented (Johnson 1983), were
not deterred by brightly colored prey. A correlation between palatability

34:48-68, 1995(1997)

ANTS BUGS WASPS

Fig. 4. The association between size and percentage of rejections by the 3 dif-
ferent predators. The y-axis represents the percentage of all caterpillars
(n = 287 individuals; 34 species) of each particular size that were re-
jected by each of the 3 different predators (ants, bugs, and wasps). The
numbers above each bar indicate the sample size; the number of cater-
pillars rejected is in the numerator, and the total number of caterpillars
offered (of that particular size) is in the denominator.

and coloration is widely assumed to exist in the animal kingdom (e.g., Cott
1940, Edmunds 1974, Harborne 1989), and coloration has even been used
as an indicator of palatability (Sillen-Tullberg 1988). It is therefore note-
worthy that for two of the three invertebrate predators used in my experi-
ments, bright coloration is not correlated with unpalatability.

As with most studies of community patterns, there were several major limi-
tations to this study which prevent me from concluding with grand gener-
alizations. With regard to questions about the relative effectiveness of vari-
ous caterpillar defenses, I had to ignore many important defenses such as
symbioses with ants (e.g., DeVries 1991), aggregation (Bowers 1993), and
other behavioral defenses (Edmunds 1974, DeVries 1987, 1994, Evans 8c
Schmidt 1990). It is therefore impossible to conclude that any defense ex-
amined in this study is the “most important” Second, the relatively small
taxonomic sample size (number of individual species representing each
family) of this study makes it impossible to determine if a characteristic typi-

J. Res. Lepid.

m GLABROUS ■ SPINES S HAIRS

Fig. 5. The association between morphology and percentage of rejections by
the 3 different predators. The y-axis represents the percentage of all cat-
erpillars (n = 287 individuals; 34 species) of each particular morphology
that were rejected by each of the 3 different predators (ants, bugs, and
wasps). The numbers above each bar indicate the sample size; the num-
ber of caterpillars rejected is in the numerator, and the total number of
caterpillars offered (with that particular morphology) is in the denomina-
tor.

cally associated with a specific taxon is an effective defense or if some cor-
related trait of that taxon is responsible. For example, comparisons of hairy
and glabrous caterpillars could just be comparisons of traits correlated with
hairy and glabrous families (e.g., Arctiidae vs. Pyralidae), because not all
the families used in my experiments included all of the possible morpholo-
gies. I addressed this problem to some extent in another study with larger
sample sizes (Dyer 1995), and found that defenses were effective despite
taxonomic affinity.

With regard to questions about the evolution of specialization, multi-spe-
cies comparisons, such as this study, are not particularly useful in terms of
providing evidence for natural selection on particular characteristics, since
protection from natural enemies could be a consequence rather than a cause
of specialization. Actual tests of predation as a selective pressure on diet

34:48-68, 1995(1997)

CRYPTIC ■ BRIGHTS OTHER

ANTS BUGS WASPS

Fig. 6. The association between coloration and percentage of rejections by the
3 different predators. The y-axis represents the percentage of all cater-
pillars (n = 287 individuals; 34 species) of each particular morphology
that were rejected by each of the 3 different predators (ants, bugs, and
wasps). The numbers above each bar indicate the sample size; the num-
ber of caterpillars rejected is in the numerator, and the total number of
caterpillars offered (with that particular coloration) is in the denominator.

breadth would require studying intraspecific variation (see Futuyma and
Moreno 1988).

Despite these limitations, a few reasonable generalizations can be made
which address my original questions. 1 ) Predators from different guilds are
deterred by different defenses, but there are some antipredator mechanisms
that may function against many different guilds. 2) Narrow diet breadth
and the utilization of noxious chemicals are significantly associated with
rejection by a suite of predators. 3) Generalist predators are probably im-
portant in the maintenance of narrow diet breadth in caterpillars.

Acknowledgments. This research was supported by a University of Colorado Fellow-
ship, a Sigma Xi Grant in Aid of Research, a National Science Foundation Disserta-
tion Improvement Grant, and a University of Colorado Van Riper Museum grant.

J. Res. Lepid.

This manuscript was greatly improved due to the comments of Phil DeVries and
two anonymous reviewers. Deane Bowers provided much needed advice and finan-
cial support during this project as well as many helpful comments on earlier ver-
sions of this manuscript. Christine Squassoni was very helpful in the field and labo-
ratory, provided many useful suggestions about experimental designs and proce-
dures, and provided helpful comments on this manuscript. I thank the Park Ser-
vice at Lomas Barbudal and Palo Verde, the Organization for Tropical Studies, and
the staff at La Selva for logistical support. Y. Chacon, P.A. Opler, G. Phillips, J.
Corrales, the staff at Finca Mariposa, D.H. Janzen, R.J. Marquis, M.D. Bowers, and
N. Greig all helped to some degree with identification of larvae and adults. O.
Vargas, R.J. Marquis, J. Denslow, N. Greig, and individuals at INBio all helped to
some degree with identification of plants.

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Journal of Research on the Lepidoptera

34:69-82, 1995(1997)

Cooperation vs. exploitation; interactions between Lycaenid
(Lepidoptera: Lycaenidae) larvae and ants

F, Osborn and K. Jaffe

Depto. Biologia de Organismos, Universidad de Simon Bolivar, Aptdo. 89000, CARACAS 1080,
Venezuela. Email: kjaffe@usb.ve

Abstract. The larval stages of many lycaenid species are myrmecophilic,
i.e. they are associated with ants. We revised the literature and catego-
rized these associations as neutral (nonexistent, commensalistic) , coop-
erative (mutualistic, mutualistic inquiline), and parasitic (food competi-
tor, cleptoparasitic, predaceous symphile, or synechthran). The relation-
ships were also noted as being facultative or obligate. Within several of
the lycaenid taxa there has been a change in the diet from phytophagy
to aphytophagy associated with a change from cooperative to exploitative
behavior towards ants. A relatively low number of species, however, seem
to have followed the route from cooperative (mutualists) to exploitative
behavior (cleptoparasites, predaceous symphiles, synechthrans) even
though the latter may give higher returns for less investment. Even neu-
tral behavior (no relation with ants, commensals) is more probable than
exploitative behavior. We suggest that this pattern reflects both the con-
straints produced by the species specific nature of exploitative interac-
tions and the stability of cooperative interactions in evolutionary terms.

We suggest that a “reverse evolution” from obligatory to facultative rela-
tionships is evolutionarily unlikely, a phenomenon which may be ex-
plained by negentropy criteria or the irreversible nature of evolution.

Introduction

Many species of lycaenids are myrmecophilic, i.e., they are associated with
ants. Through these associations with ants, lycaenid larvae have developed
a number of morphological and behavioral adaptations. Many species of
larvae have evolved what have been termed myrmeocophilous organs, one
of the most important of these being the nectary organs which are found
on the seventh abdominal segment and secrete a substance containing sug-
ars and amino acids when solicited by the ants (Malicky 1970, Maschwitz et
ah 1975, Pierce 1983, Cushman et al. 1994).

Apart from their morphological adaptations lycaenid larvae are unusual
with respect to their diet. They may feed on lichens, homoptera, or ant
brood rather than on angiosperms, which is the normal food of lepidopter-
ous larvae. Many of the interactions involving lycaenid larvae and ants have
been described (Kitching 1987; Fiedler & Maschvdtz 1988, 1989a, Elmes et
al. 1991) and a exhaustive revision of these was undertaken by Fiedler
(1991b) . Lycaenid-ant interactions have been classified as mutualistic/ para-

Paper submitted 25 March 1995; revised manuscript accepted 12 June 1996,

/. Res. Lepid.

side, facultadve/obligate and phytophagous/aphytophagous. Although
some authors (Henning 1983) have given finer classifications, the full range
of possible associations have not been taken into account. For example,
Maculinea spp. and Liphrya brassolis are both classed as “parasites” even
though they have completely different relations with their ant hosts at a
behavioral level which implies different evolutionary pathways towards each
of these two types of relation; Maculinea spp. are attended, for example M.
alcon, M. rebeli (Cottrell 1984, Elmes et al. 1991), or ignored, for example
M. arion, M. teleius (Cottrell 1984), by the ants whilst Liphrya brassolis is at-
tacked (Johnson & Valentine 1986).

Several authors have studied the relative importance of cooperation vs.
exploitation using different models and have shown that in theory, “coop^
eration rather than “exploitation” dominate in the Darwinian struggle for
survival” (Newark & May 1992, Newark et al. 1996, Sigmund 1992). Em-
pirical evidence suggests that in the Lycaenidae this dominance of coop-
eration over exploitation may be true (Pierce 1987, Fiedler 1996).

Using data in the literature, most of it summarized by Fiedler (1991a, b),
on the types of interactions between lycaenid larvae and ants, the myrme-
cophilous organs on the lycaenid larvae, the degree of relationship (facul-
tative or obligate) and the diet of the larvae, we tentatively propose a more
detailed classification of “types of interaction.” In each case we noted the
presence or absence of the nectary organs, larval diet (spermatophytes, al-
gae, lichens; homoptera; ants; homoptera honeydew; ant regurgitations),
and type of interactions with ants. Using this information we classified eight
types of interactions the larvae may have with the ants. We then use this
classification to describe the diet changes that have occurred both between
and within subfamilies (from phytophagy to aphytophagy) and discuss these
diet changes in the context of the relative importance of cooperative/ex-
ploitative behavior of the larvae towards their ant partners.

A truly phylogenetic system of the Lycaenidae is still not available, thus the
diet changes we describe cannot yet be confirmed since without a sound phy-
logeny the directionality of such changes is difficult to assess. Nevertheless, it
is widely considered that phytophagy is a primitive trait in lycaenids (and
butterfly larvae as a whole) (Cottrell 1984, Fiedler 1991b) , thus we feel justified
in our assessments of possible evolutionary change from phytophagy to
aphytophagy in the Lycaenidae. The higher classification of the Lycaenidae
we adopt is the same as that used by Liedler (1991b), based on Eliot (1973),
with modifications by Scott and Wright (1990) . The discussion about whether
or not the Riodinidae (or Riodininae) form a monophyletic group together
with the Lycaenidae is still very much alive (Robbins 1988, Dejong et al. 1996,
Weller et al. 1996), but since the myrmecophilous organs of the Riodinidae
are clearly analogous but not homologous with those of the Lycaenidae
(DeVries 1990) we do not further discuss the Riodinidae here.

Types of interaction between lycaenid larvae and ants

The range of types of relationships that the ants may share with lycaenids

34:69-82, 1995(1997)

were classified as follows. It must be emphasized that these are “types of
behavioral interactions,” not “types of larvae.” Thus a larva that is neutral
at one stage of its life cycle may be parasitic at another, as in Maculinea spp.
(Cottrell 1984) . Parasitic larvae such as Maculinea rebeldi or M. alcon may be
generally regurgitation feeders (cleptoparasites) but during times of food
shortage may also prey on eggs and ant brood (Elmes et al. 1991).

Relationship not recorded

Larvae which have unknown relationships with ants. (Relationships re-
corded with a question mark by Fiedler [1991a, b] .)

Neutral relationships

The ants neither gain nor lose from the interaction with the lycaenid lar-
vae. The larvae, however, may neither gain or lose (No relationship) or may
gain (Commensal) from the relationship. It is very difficult to assess with
the data available which larvae are in “No relationship” with the ants, and
which are “Commensals.” The discussion about whether the larvae enter
into “enemy free space” or not must depend on studies of particular lycaenid
larvae and their relation with ants. Whether or not the ants protect the lar-
vae in any way from other predators depends on factors such as time of
occupancy of ants at the site, whether or not they have antagonistic rela-
tions with the larvae, and whether they influence in the rates of predation
or parasitism of the larvae. Since the subject of whether and which of these
larvae benefit from the presence of ants is in many cases ambiguous we have
lumped “No relationship” and “Commensal” into the same category of
Neutral relationship. Nevertheless, it is useful to dehne the two sub-catego-
ries, as they may represent the transition from a completely myrmecoxenous
state to the beginnings of an association with ants.

No relationship. The larvae do not interact with ants mutualistically, para-
sitically, or commensally. Thus neither the ants nor the larvae gain from
the relation. The larvae may avoid encounters with ants using specihc de-
fensive tactics such as Eumaeus atala (Bowers & Larin 1989) or they may be
rarely found by ants. The point is that they do not enter into ant-inhabited
“enemy free space” (Atsatt 1981). In the Curetinae, ants sometimes encoun-
ter larvae and then lick up plant sap at feeding damage, or feed at extra
floral nectaries (DeVries et al. 1986, Fiedler et al. 1995). The evidence is
ambiguous, however, as to whether or not the larvae benefit from the rela-
tion.

Commensalistic. In these associations, unlike the “No relationship” asso-
ciations, the larvae benefit from the relation, whilst the ants remain unaf-
fected. Thus they gain a twofold advantage (avoidance of ant attacks and
entering into “enemy free space”). Commensalistic relations have been
described in the Liptenini where the larvae are strictly associated with ant
columns on tree trunks where they feed on lichens or algae. The data are,
however, scanty and the proportion of Liptenini in these types of relation-
ships is unknown (Downey 1962, Atsatt 1981, Callaghan 1992). The larvae

J. Res. Lepid.

Table 1. Number of species and relative proportions of neutral, cooperative, and
exploitative interactions found between lycaenid larvae and ants: a) within the
subfamilies of the Lycaenidae, b) within the Lycaeninae.

Poritinae

MOetinae

Curetinae

Lycaeninae

Liptenini

Miletini Liphryini Curetini

i Total

Grand Total

Not recorded

0 0

122

—

Phytophagous

Neutral

0 7

64.4

215

283

30.8

Cooperative

0 0

574

62.5

Aphytophagous

Exploitative

9 0

35.6

6.7

Total

9 7

104

100

935

1039

100

Lycaeninae

Aphnaeini Lycaenini Theclini Eumaeini

Polyommatini

Total

Not recorded

104

122

Phytophagous

Neutral

101

215

26.5

Cooperative

Facultative

111

284

470

Obligate

104

Total Cooperative 67

113

318

574

70.5

Aphytophagous

Exploitative

TOTAL

124

318

376

936

100

The last column (%) refers to the relative proportion of lycaenid larvae in

a given type of

interaction with ants with respect to the total number of larvae. Larvae with a relation “not
recorded” are NOT taken into account.

supposedly gain from the relation in that the presence of ants reduces at-
tacks from predators and parasites (Atsatt 1981) whilst the ants remain
unaffected since the larvae do not compete in any way with food or other
resources. Nonetheless, Callaghan (1992) described larval behavior in 12
species from the tribe Liptenini where the larvae seem to have strictly de-
fensive relationships with ants, thus suggesting that the ants may not be
protective elements in this case and that the relationship between them and
the larvae is rather antagonistic. Nevertheless, detailed studies are required
in order to establish exactly what is the relationship between the ants and
certain Liptenini larvae. There are also certain species in the Lycaeninae
that can be classed as being commensalistic because they are or appear to
be associated with ants, but apparently do not possess a nectary organ and
thus presumably do not provide the ants with a substantial food resource,
for example Aloeides dentatis (Henning 1983).

Mutualistic (Cooperative)

This follows the standard definition of mutualism in the literature whereby
both the ants and the lycaenid larvae benefit from the association. The lar-
vae secrete a sugary nectar which the ants imbibe (Fiedler & Maschwitz 1988,
1989a, Cushman et al. 1994, Fiedler & Saam 1995). The ants in return pro-
tect the larvae from predators and parasites (Pierce & Mead 1981, Pierce et

34:69-82, 1995(1997)

al. 1987, Baylis 8c Pierce 1991) Under this definition a larva is mutualistic if
it has a functional nectary organ, if the diet is phytophagous and if it is as-
sociated with ants. Mutualists may be facultative or obligate, where the term
obligate is defined as complete dependency on a specific genus of ants
(Fiedler 1991b, 1994). Mutualists as defined here are only found in the
Lycaeninae (Table 1).

Mutualistic inquiline. Here we define a new type of interaction which is a
subdivision of the mutualists. In this case the larvae are attended by ants as
for the mutualists, but furthermore they shelter either in pavilions con-
structed by the ants or in the ant nests themselves. The larvae, however,
remain phytophagous, leaving the shelters to feed on their hostplant. Ex-
amples of species which exhibit “inquiline behavior” are Anthene emolus
(Fiedler & Maschwitz 1989a) and Paralucia aurifera (Cushman et al. 1994).
It must be emphasized again that it is the interaction that is important not
the species. Thus “inquiline behavior” may be a rare occurrence in a spe-
cies or a life history trait. The importance of this category is that it suggests
a possible intermediate stage between free-living mutualists and parasites
which live in the ant colony and feed on the ant brood.

Parasitic (Exploitative)

In these cases the lycaenid larvae benefit from the association whilst the
ants are disadvantaged. We divide the parasitic larvae in four subgroups;
food competitors, cleptoparasites (after Hoelldobler 8c Wilson 1990), pre-
daceous symphiles and synechthrans (after Wasmann 1894).

Food competitors. Here we define a type of interaction in which the lar-
vae feed on Homoptera (and Homoptera secretions), which have a
trophobiotic relationship with ants such as many species from the Miletinae
(Kitching 1987, Maschwitz et al. 1985, 1988). This definition differs from
that of Maschwitz and Fiedler (1988) who defined homopterophagous
lycaenid larvae as “indirect parasites.” We suggest, however, that “food com-
petitors” is a more precise definition. The food competitors may be further
divided into “stealthy competitors,” which are not tolerated by the ants and
feed inside shelters or cover themselves with bits of their prey to protect
themselves from ant attack, for example, Spalgis spp., and “symphilic
cleptoparasites,” which are ignored or even sometimes attended by the ants,
for example, Miletus spp. (Cottrell 1984, Fiedler 1991b).

Cleptoparasites. The larvae are food robbers (Euliphyra spp. [Dejean 8c
Beugnon 1996]) or feed on oral regurgitations from ants. Oral regurgita-
tion feeders may be either free-living {Spindasis takanonis) or may inhabit
the nests of the ants {Niphanda fusca) (Cottrell 1984). Fiedler (1991b)
defined ant regurgitation feeders as “parasites,” nevertheless Hoelldobler
and Wilson (1990) define “food robbers” which rob the ants of a food re-
source and the regurgitation feeders which receive nutrients that would
normally be destined for the ant brood (oral regurgitations) as cleptopara-
sitism (cleptobiosis in their terms) . Cleptoparasitic behavior has been re-
ported from both the Lycaeninae and Miletinae.

J. Res. Lepid.

Predaceous symphile. The larvae spend all or part of the larval phase in-
side the nests of their host ant, feeding on ant brood. By means of putative
pheromone secretions the larvae are accepted by the ants as ant brood whilst
they remain in the ant nest Qackson 1937, Cottrell 1984, Thomas et al.
1989). This definition applies to lycaenids such as Maculinea arion, M. teleius,
and Lepidochrysops spp., described simply as “parasites” in the literature, for
example (Cottrell 1984, Elmes et al. 1991).

Synechthran (following Wasmann 1894). These species of lycaenid also
feed on ant larvae, but their relation with the ants has a completely differ-
ent behavioral base than that of the predaceous symphiles. The larvae are
not welcome guests in the ant nests; rather they are treated as intruders
and attacked by the adult ants. Liphyra brassolis (Johnson & Valentine 1986)
is apparently the only known case which falls in this category in the Lycae-
nidae.

Changes in the diet within subfamilies

Changes in the diet within a subfamily have taken place in the Lycaeninae
from angiosperms to ant brood, Homoptera and regurgitations from ants,
and in the Miletinae from Homoptera, to honeydew, ant regurgitations, or
ant brood.

Changes in the diet in the Lycaeninae

Within the Aphaenini, Theclini, and Polyommatini there has been a
change in the diet from phytophagy to aphytophagy, the aphytophagous
larvae feeding on Homoptera (food competitors) or oral regurgitations from
the ants (cleptoparasites), but sometimes on ant larvae or pupae (preda-
ceous symphiles) . The phytophagous species in the Lycaeninae are either
commensals (e.g., Aloeides dentatis; all examples taken from Fiedler [1991b]
unless otherwise stated), mutualists, mutualistic inquilines, or have no rela-
tion with ants. Their behavior towards the ants is thus neutral or coopera-
tive. The aphytophagous species, however, all exploit their ant hosts. Food
competitors and/or cleptoparasites may be found in the Aphnaeini,
(Spindasis nyassae, S. takanonis, Axiocerses harpax and A. pseudo-zeritis, oral re-
gurgitations), in the Theclini {Shirozua jonasi, oral regurgitations) and the
Polyommatini {Niphanda fusca, oral regurgitations, Triclema lamias, Hom-
optera and three Maculinea spp.). These species have nectary organs and
sometimes also tentacle organs (except S. jonasi, which has neither). There
are predaceous symphiles in the tribes Theclini: Acrodipsas cuprea, A.
myrmecophila, A. illidgei; Polyommatini: two Maculinea spp. and nine
Lepidochrysops spp.; and Aphnaeini: Cigaritis acamas (Sanetra & Fiedler
1996) . As far as is known, all species possess a nectary organ, except Cigaritis
acamas which also has eversible tentacles. The Maculinea spp. are generally
specific to one ant species, at least within the same geographical region
(Thomas et al. 1989) . Lepidochrysops spp. are almost certainly species specific
(Cottrell 1984), although there is little information as regards the remain-

34:69-82, 1995(1997)

ing genera, what evidence there is points to host-ant specificity (Cottrell
1984).

Changes in the diet within subfamilies in the Miletinae
In the Miletinae there have been changes in the diet of the larvae from
Homoptera to other food sources(all examples taken from Fiedler [1991b]
unless otherwise stated) . Although the scarcity of data on this tribe does
not permit conclusions to be drawn we can state that in all cases studied
the behavior of the larvae towards the ants is exploitative. In the Miletini
there are several species reported to feed on Homoptera honeydew, these
include Miletus chinensis. Tar aka hamada, Logania malayica, L marmorata (also
Homoptera) (Fiedler 1993), A/fohwws miro/or (also Homoptera) (Maschwitz
et al. 1985, Fiedler & Maschwitz 1989b) and Lachnocnema bibulus (also ant
regurgitations). Thestor spp. (Miletini) are suspected of predating on ant
brood. In the Liphyrini Euliphyra mirifica and E. leucyania feed on oral re-
gurgitations from ants and Eiphrya brassolis (Liphyrini) feeds on ant brood.
These species do not possess nectary or tentacle organs. Of these, Lachnoc-
nema is not specific as regards the ant host, but Thestor, Miletus, Euliphyra,
and Liphyra are species specific.

Discussion

Facultative and obligate relations in the Lycaeninae
Regarding the subfamily Lycaeninae, Fiedler (1991b) discusses the pos-
sible evolutionary development from facultative mutualisms to obligate re-
lations of various types (including mutualists, inquilines, cleptoparasites,
predaceous symphiles) or alternatively an evolutionary decrease in the in-
teractions with ants (secondary myrmecoxeny) . He states that there ‘hs yet
no evidence that a reverse evolution from obligatory towards facultative
myrmecophily has ever occurred within the Lycaenidae, although such
would be possible from theory.” We propose that the theory of negentropy
provides a possible explanation for the lack of evidence for this “reverse
evolution.” This proposal assumes that the higher the order or complexity
of an organism, including in the concept of complexity higher specializa-
tions that may involve loss or simplifications of certain structures, the lower
will be the probability state of the system and the longer the evolutionary
time to produce the given state. Thus the further down a certain evolution-
ary pathway an organism finds itself the fewer available choices it will have
to return back along that pathway (Zotin & Konoplev 1978, Jaffe 1984, Jaffe
& Hebling-Beraldo 1993, Jaffe & Fonck 1994). We argue that obligate
myrmecophiles are more “complex” in that they have more finely tuned
adaptations in their associations with ants than facultative myrmecophiles.
Thus in this case negentropy is expressed as specificity of communication
with ants. (For a discussion on lycaenid/ant communication see Fiedler et
al. [1996] .) For example, the predaceous symphiles are often associated with
one or a few ant species, which implies the development of brood phero-
mone mimics, that are specific to a single (or a few closely related) ant spe-

/. Res. Lepid,

cies (Thomas et al. 1989), probably from facultative relations where the
larvae are attractive to many species of ant. A reversal of this trend would
imply a loss of specificity and thus of complexity, which would revert and
thus probably reduce the adaptive gains made in the first place. This
negentropic assumption does not exclude the possibility of posterior losses
as has taken place in the secondarily myrmecoxenous species, but predicts
that these reversions should be rare and should have specific biological
explanations, as the evolutionary process is strongly irreversible (Jaffe 1996) .

Cooperation vs. exploitation in lycaenid/ant relations

From Table 1, we may conclude that the majority of the lycaenid butterflies
maintain neutral (no relationship, commensalistic) or cooperative (mutm
alistic) interactions with ants, rather than exploitative (cleptoparasite, pre^
daceous symphile, synechthran) ones (Pierce 1987, Fiedler 1996). This fact
seems remarkable considering that exploitative behavior may give higher
nutrient returns for less investment to the lycaenid larvae. In subfamilies
without a nectary organ, i.e. where cooperative behavior has not appeared
(Table la), 64.4% of species show neutral behavior (no relation or com-
mensal), representing the subfamilies Poritiinae (60 species) and Guretinae
(7 species) and only 35.6% of the species show exploitative behavior
(cleptoparasites or synechthrans) representing the Miletinae (37 species).
In the Lycaeninae with 818 species (Table lb, excluding species for which
no information is recorded), cooperative behavior dominates, with 70.5%
of the larvae being mutualists as opposed to 3% being cleptoparasites or
predaceous symphiles. In this subfamily, 26.5% of the species have no rela-
tion with ants are or commensals, showing that even neutral behavior is more
likely than exploitative behavior. Taking the Lycaenidae as a whole (Table
la final column), 62.5% show cooperative behavior, 6.7% exploitative be-
havior and 30.8% neutral behavior towards the ants. Although these per-
centages may vary as more Lycaenid species are investigated, we suggest that
the relative proportions between exploitative larvae and cooperative/ neu-
tral larvae should remain roughly the same.

Thus, where cooperative (mutualistic) behavior is possible in the Lycae-
nidae this is the most probable evolutionary outcome, and where it is not
likely, neutral behavior is more probable than exploitative behavior. The
preponderance for mutualistic interactions over exploitative relations in
Lycaenidae lead us to suppose that cooperation must have either a higher
probability to evolve or to be maintained during evolution or both. Thus,
we postulate that cooperation is an evolutionarily more probable strategy
compared to exploitative behaviors. We propose different, but not neces-
sarily contradictory, explanations for this pattern:

1 ) A model of cooperation between species as a stable strategy was devel-
oped by Axelrod and Hamilton (1981) using the Prisoners Dilemma game.
They showed that if the probability that two individuals will continue to
interact is great enough then cooperation may be evolutionarily stable. Since
then several authors have modeled cooperation vs. exploitation using dif-

34:69-=82, 1995(1997)

ferent versions of the Prisoners Dilemma and have shown that in theory,
“cooperation rather than exploitation can dominate in the Darwinian
struggle for survival” (Nowark & May 1992, Nowark et ak 1996, Sigmund
1992) . Empirical evidence suggests that the Lycaenidae larvae benefit from
the association (Pierce et ak 1987, Robbins 1991, Fiedler Sc Hoelldobler
1992, Wagner 1993) and there is evidence showing that both partners
benefit (Fiedler Sc Maschwitz 1988, 1989a, Cushman et ak 1994, Fiedler Sc
Saam 1995). Cooperation in lycaenid/ant interactions is not necessarily a
fixed strategy (Bronstein 1994, Noe Sc Hammerstein 1994, 1995) and a coa-
lition may end or change when it becomes unproductive for one or both
partners (Enquist Sc Leimar 1993). For example, ants abandoned Polyom-
matus coridon larvae when the secretions from the nectary gland were
artificially eliminated (Fiedler &: Maschwitz 1989c). Leimar and Axen (1993)
showed that the amount of nectar secreted by larvae of P. icarus varied ac-
cording to the level of ant attendance and the larva’s need for protection.
A model of mutualism, commensalism and parasitism as evolutionarily stable
strategies in lycaenid/ant relations was developed by Pierce and Young
(1986). This model assumes that the ants enhance both the population
growth rate and the equilibrium density of the larvae by increasing the re-
alized fecundity of individual butterflies and by increasing juvenile survival,
whereas the larvae enhance the equilibrium density of the ants by increas-
ing ant food supply. Under these assumptions (albeit largely unverified)
Pierce and Young (1986) were able to demonstrate that all three types of
relation were evolutionarily stable strategies. Nonetheless, although all three
strategies are evolutionarily stable, not all have the same odds of appearing
during evolution and of avoiding extinctions in evolutionary history. Co-
operative strategies possess economic advantages which decrease their prob-
abilities of extinction and thus increase their odds of being fixed in the
genetic repertoire of more species. That is, cooperation is a highly prob-
able strategy in addition of being evolutionarily stable.

2) There are three possible strategies for exploitative behavior which the
larvae could take; a “synechthran” approach where the larvae fend off ant
attack whilst predating on ant brood, a “stealthy” approach, whereby the
larvae avoid ant attack, and a “symphilic” approach whereby the larvae de-
ceive the ants by mimicking ant brood. Thus, ants either ignore the larvae
or attend them as they predate on Homoptera or ant brood. Examples of
the first approach could be Liphrya brassolis which has an armor shaped cara-
pace in order to withstand ant attack. This type of defense does not, how-
ever, seem to have developed in lycaenid taxa other than the Liphyrini.
Examples of the second “stealthy” approach may be found in the genera
Taraka, Spalgis, and Feniseca (Miletini) where the larvae occupy silken tents
or burrows, or cover themselves with remains of their prey to avoid ant at-
tack (Cottrell 1984, Kitching 1987). The third “symphilic” approach involves
the development of a chemical mimicry system with the larvae mimicking
their homopteran prey, adult ants or ant brood. The possibility that lycaenid
larvae are chemical mimics has been studied for Aloeides dentatis, a non-

J. Res. Lepid.

mutualistic inquiline and Lepidochrysops ignota, a predaceous symphile
(Henning 1983). In both species, larval epidermal glands produced a se-
cretion that appeared to mimic the brood pheromones of the host ants,
although Henning (1983) did not identify the chemical compounds in-
volved. It is also supposed that Maculinea spp. mimic the brood pheromones
of their Myrmica ant hosts (Thomas et al. 1989), although chemical analy-
ses have not been undertaken as yet. In the Miletini many lycaenid larvae
such as Miletus spp., Lachnocnema bibulus are attended by ants even though
they do not give any reward (Cottrell 1984). All of these strategies; the
“synechthran” approach, the “stealthy” approach and the “symphilic” ap-
proach carry with them certain disadvantages. The carapace used by Liphyra
brassolis may not be 100% effective against all ant species, with the larvae
possibly incurring high mortality rates as a result. This restricts the larvae
to only associating with Oecophylla spp. The stealthy larvae may still be at-
tacked by ants in spite of their protective burrows. The symphilic larvae are
constrained by having to penetrate the complex chemical communication
systems of ants, which are highly species specific. In this sense it is notable
that the larvae mimic the brood of the ants rather than the adult ants. In
the genus Myrmica (usually hosts for larvae of Maculinea spp.) the brood
odor is not specihc to one species and Myrmica brood are transferable be-
tween the nests of different species (Brian 1975, Howard et al. 1990), al-
though Thomas et al. (1989) point out that these ants are far more discrimi-
natory under conditions of stress.

3) As far as the “symphilic” or “mimicry” approach to exploitative behav-
ior is concerned, lycaenid larvae mimics are normally specific to one spe-
cies of host ant (Cottrell 1984; Thomas et al. 1989), which is probably due
to a specificity in the chemical signals the ants use to recognize nest com-
panions and brood (Hoelldobler & Carlin 1987). Although this species
specificity of the lycaenid larvae towards their ant hosts may have led to a
diversification of some genera (e.g., Maculinea, Lepidochrysops) , this diversi-
fication is far lower than that of cooperative taxa, a hnding that contradicts
the hypotheses of Pierce (1984) who argued that species specificity should
amplify the species diversity of the Lycaenids (see also discussion in Fiedler
1991b). Nonetheless, being associated with only one species of ant carries
with it certain ecological disadvantages for the lycaenid larvae such as con-
straints on their distribution caused by a patchy distribution of their host
ant species (Jordano et al. 1992), problems of host encounter in areas with
a highly diverse ant fauna, and nutritional constraints (Fiedler 1991b). For
the predaceous symphiles exploitative behavior also carries with it a high
risk. Their host ants are generally tolerant of intruders in times of plenty,
but when food reserves are low they become increasingly intolerant and
will even eat their own brood (Thomas et al. 1989). The lycaenid larvae
must therefore be under extreme pressure to mimic their hosts as closely
as possible and it is not surprising that so few species have developed this
type of relation.

4) We may speculate that parasites normally have much shorter life cycles

34:69-82, 1995(1997)

than their hosts, as for example viral or bacterial parasites on insect or
mammal hosts. Thus, cooperative mechanisms are more likely to act in in-
teractions between two species with equivalently long life cycles. The life-
spans of ant workers and butterflies have roughly the same order of magni-
tude (they are measured in months). Even ant colonies do not live much
longer, as in most species, the mean life span of queens and colonies is a
few years. Thus exploitation of one by the other is evolutionarily unlikely.

In conclusion, a relatively high proportion of species seem to employ
cooperative or mutualistic behavior in their associations with ants rather
than exploitative or selfish behavior. We suggest that this pattern reflects
the extraordinary stability of cooperative interactions in evolutionary terms,
at least as regards lycaenid/ ant interactions.

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Journal of Research on the Lepidoptera

34:83-98, 1995(1997)

A revision of Mesogona Boisduval (Lepidoptera: Noctuidae)
for North America with descriptions of two new species

Lars Crabo^ and Paul C. Hammond^

^Thomas Burke Memorial Washington State Museum, Seattle, Washington 98195, USA
^Department of Entomology, Oregon State University, Corvallis, Oregon 97331, USA

Abstract. The North American species of Mesogona Boisduval are revised.
Pseudoglaea Grote, 1876a is treated as a synonym of Mesogona. Three spe-
cies of Mesogona occur in North America, two of which are described as
new. All are found in western North America: M. olivata (Harvey, 1874)
occurs from British Columbia south to California and Texas, while M.
subcuprean. sp. and M. rubran. sp. are restricted to Washington, Oregon,
and California. The adults and genitalia of these species are described
and illustrated. A key for identification of the adults is presented. The
larva of M. rubra is illustrated.

Introduction

Members of Mesogona Boisduval, 1840 are stout-bodied medium-sized
moths. They occur in a variety of habitats ranging from wet forest to semi-
arid steppe. The adults are active in the Fall at about the time leaves of de-
ciduous trees and shrubs turn color. Their eggs are laid in the Fall and hatch
in the Spring. The known larval foodplants include a diverse assortment of
woody plants.

There are five species in this genus, two in Eurasia and three in North
America, Until now, only one of the North American species, olivata Harvey,
1874, was described. It was placed in the monotypic genus Pseudoglaea Grote,
1876b. The two other North American species were recognized recently
from material collected in Washington and Oregon. The relationship of
the Nearctic species to Mesogona, previously thought to be restricted to Eu-
rope, became evident because one of the undescribed species resembles
M. acetosellae (Denis & Schiffermuller 1775), the genotype of Mesogona.
Closer comparison of the Palaearctic M. acetosellae to the Nearctic species
shows that they are structurally similar and thus congeneric. This revision
is limited to the North American Mesogona species because the Palaearctic
species are well known (Fibiger 1993).

Mesogona Boisduval

Mesogona Boisduval, 1840:144.

Type species: Noctua acetosellae [Denis & Schiffermuller], 1775, by sub-
sequent designation by Blanchard, 1840:512.

Paper submitted 18 October 1995; revised manuscript accepted 29 April 1996.

/. Res. Lepid.

Pseudoglaea Grote, 1876b: 18, new synonomy

Type species: Choephora blanda Grote, 1876a, by subsequent desig-
nation by Grote, 1895:95.

Description. Adult: Eyes naked, lashed. Palpi upturned with porrect third
segment, the first and second segments bearing long loose scales, the third
segment closely scaled. Frons smooth. Antennae ciliate. Thorax untufted,
covered with hairlike scales. Prothoracic tibia unarmed, slightly longer than
first tarsal segment; meso- and metathoracic tibiae with several loose rows
of stout setae (“spines”) in addition to the tibial spurs. Tarsal segments with
stout setae laterally. Male abdomen with basal coremata in all known spe-
cies. Male genitalia (Figs. 11--14): Uncus narrow, curved. Tegumen broad,
with penicillus lobes. Juxta flat, widest ventrally. Valve long and narrow,
slightly constricted mesially; cucullus rounded, with a weak corona; saccu-
lus with a costal process (sensu Forbes 1954), ys-l Xas wide as valve, ex-
tending to base of harpe; harpe nearly cylindrical, 1.5-2 Xas long as valve
width, parallel to valve at base, curved posterodorsad distally; digitus ab-
sent. Aedeagus with dorsal and ventral extensions onto base of vesica; vesica
1-2.75 X as long as aedeagus, coiled or T-shaped and bent, surface minutely
granulose and armed with two to three helds of cornuti, portion of vesica
bearing cornuti either flat, slightly raised, or a small diverticulum; the
cornuti are fragile and entire cornuti or fragments are often left in the fe-
male corpus bursae following copulation. Female genitalia (Figs. 15-18):
Bursa copulatrix uni- or bisaccate; corpus bursae curved toward right ante-
riorly, with 1-3 signa, posterior corpus bursae (M. acetosellae) or appendix
bursae heavily sclerotized; appendix bursae (if present) broadly joined to
corpus bursae posteriorly, extending to the right and anteriorly; ductus
seminalis joined to posterior corpus bursae (M. acetosellae) or to apex of
appendix bursae. Ductus bursae ys-l Xas long as bursa, joined to it
posterodorsally; ostium bursae weakly sclerotized. Anterior apophyses y2-
ys as long as posterior apophyses. Ovipositor lobes triangular, covered with
long and short hairlike setae.

Discussion: McDunnough (1927, 1928) recognized the close relationship
of M. olivata and M. acetosellae, but retained Pseudoglaea because of differ-
ences in the lengths of the distal spines of the first tarsal segments of the
first legs (“tarsal claws”) of these species. The link between the Old and
New World species is more evident now since M. acetosellae (Fig. 9) is simi-
lar to the recently discovered M. subcuprea n. sp. (Fig. 6), and both of these
species lack the long “tarsal claws” of M. olivata.

The most closely related genus is Eucirroedia Grote, 1875 from the east-
ern United States and southern Canada. This monotypic genus (type spe-
cies pampina Guenee, 1852) differs from Mesogona by the following charac-
ter states: 1 ) the vestiture of thorax has a median crest, absent in Mesogona;
2) the mid and hind tibiae bear only two weak spines while those of Mesogona
have multiple stronger spines; 3) the forewing is falcate and scalloped while
that of Mesogona has a slightly convex crenulate outer margin; 4) the harpe
of the male valve is expanded and flattened subapically and pointed dis-

34:83^=98, 1995(1997)

tally while that of Mesogona is uniform in width; 5) the juxta has a membra-
nous dorsomedian cleft, absent in Mesogona; 6) the bursa copulatrix is long
and narrow while that of Mesogona is ovoid or bisaccate.

The species of Mesogonahme often been placed in Noctuinae, as defined
by Hampson, due to the presence of tibial spines (Hampson 1903,
McDunnough 1928, Fibiger 1993) . They are more closely related to a group
of genera referred to as the “winter moths” (Xylenini, in part) , including
Eucirroedia, Metaxaglaea Franclemont, and Epiglaea Grote. Mesogona olivata
is correctly placed in Xylenini by Franclemont and Todd (1983) . In this list
Xylenini is placed in the Cucullinae (as defined by Hampson). Hampson’s
subfamily concepts are now recognized to be unnatural. Recent reevalua-
tion of the subfamilies in the trifid noctuids, outlined in Poole (1994), in-
dicates that Mesogona is a member of the subfamily Noctuinae which has
been expanded to include a large number of species previously included
in other subfamilies.

The distribution of the species of Mesogona is disjunct. The Palaearctic
species occur predominantly in Europe with the range of M. acetosellae ex-
tending east to the Altai Region of Siberia (Fibiger 1993), while the Nearc-
tic species are restricted to western North America. In Europe, larval
foodplant records include Quercus species for M. acetosellae and Salix spe-
cies for M. oxalina (Hiibner, [1803]) (Fibiger 1993).

Key to adults of North American species of Mesogona
1 .a. Hindwing gray or with gray suffusion; vesica of aedeagus with two dis-
tal bands of short thin cornuti (Fig. lib); appendix bursae overlap-
ping corpus bursae ventrally (Fig. 15); widely distributed in western
North America ............................................................................. olivata

Lb. Hindwing uniform copper-colored or reddish, without gray scales;
vesica with stout cornuti; appendix bursae not overlapping corpus
bursae ventrally; restricted to the west coast states ............................ 2

2. a. Thorax and forewings yellow-brown, with orbicular and reniform spots
strongly outlined; vesica shaped like a lopsided T with median and
subapical cornuti (Fig. 12b); appendix bursae not overlapping cor-
pus bursae (Fig. 16) .................................................................

2.b. Thorax and forewings brownish red to pink, with faint or absent forew-
ing spots; vesica coiled with one stout basal cornutus and two subapi-
cal bands of long cornuti (Fig. 13b); appendix bursae overlapping
corpus bursae dorsally (Fig. 17) ................................................... rubra

Mesogona olivata (Harvey), new combination
(Figs. 1-5, 11, 15; Map 1)

Glaea olivata Harvey, 1874:120, TL — California. Grote 1880:155, Smith
1893:221, Dyar 1903:181.

Choephora hlanda Grote, 1876a:86, TL — Washington Territory and Van-
couver Island, [British Columbia] .

J. Res. Lepid.

Pseudoglaea blanda (Grote) Grote, 1876b:18, Smith 1893:210, Dyar 1903:178,
Anderson 1904:29, McDunnough 1927:65, Jones 1951:52, Franclemont
&:Todd 1983:145.

Pseudoglaea taedata Grote, 1876b:18, TL —= Texas. Smith 1893:210, Dyar
1903:178, McDunnough 1927:65, Jones 1951:52, Franclemont & Todd
1983:145.

Cerastis olivata (Harvey) Grote, 1878:181.

Pseudoglaea decepta Grote, 1881:271, TL — Colorado. Smith 1893:210, Dyar
1903:178, Jones 1951:52, Franclemont & Todd 1983:145.

Metalepsis blanda (Grote) Dyar, 1903:132.

Metalepsis taedata (Grote) Dyar, 1903:132.

Metalepsis decepta (Grote) Dyar, 1903:132.

Mythimna blanda (Grote) Hampson, 1903:608, pi. 76, fig. 19; Barnes &:
McDunnough 1917:47.

Mythimna taedata (Grote) Hampson, 1903:608; Barnes & McDunnough
1917:47.

Mythimna decepta (Grote) Hampson, 1903:608; Barnes & McDunnough
1917:47.

Spectraglaea olivata (Harvey) Hampson, 1906:439, pi, 106, fig. 14.

Mesogona olivata (Harvey) Barnes 8c McDunnough, 1916:161.

Mythimna olivata (Harvey) Barnes & McDunnough, 1917:47, Blackmore
1927:19.

Pseudoglaea olivata (Harvey) McDunnough, 1927:65, McDunnough 1938:67,
Jones 1951:52, Franclemont & Todd 1983:145.

Description. Adults (Figs. 1-5): Males and females identical in habitus.
Distal spines of first tarsal segment of prothoracic leg twice as long as proxi-
mal spines. Ground color of head, dorsal antennae, thorax, and forewings
variable, ranging from dull tan to reddish brown, gray-brown, or cream;
median area of forewing and postmedian space at costa darker; palpi with
mixture of ground color and dark scales; abdomen fuscous. Forewing length:
15-20 mm. Forewing 2Xas long as wide; margin crenulate; lines double,
smooth, pale filled; basal line sinuous, evident only near costa; antemedian
line oblique, undulating, bent basad at costa, outer line dark; median shade
absent; postmedian line smooth, laterally convex, inner portion dark, stron-
gest in interspaces; subterminal line sinuous, indistinct, a series of dark spots
between veins; terminal line thin and dark; orbicular and reniform spots
large, pale with darker filling; claviform spot absent. Hindwing variable,
fuscous gray to reddish, always suffused with gray scales, with darker termi-
nal area and faint discal spot, fringe lighter. Male genitalia (Fig. 1 1): Valves
as in generic description; costal lobe of sacculus triangular. Vesica 2.75 X as
long as aedeagus, shaped like a lopsided T beyond basal twist with short
extension ventrad and to the right and longer distal portion curved dorsad
and to the left, two long fields of fine cornuti on distal ^3 , the proximal end
of the field of larger cornuti is raised from adjacent vesica surface. Female
Genitalia (Fig. 15): Corpus bursae approximately 2 X as long as wide, ante-

34:83-98, 1995(1997)

Map 1 . Map of part of western North America showing distribution of examined
material of M. olivata.

rior curved dorsad and to the right, with single long dorsal and ventral
signa; appendix bursae cone-shaped, curving anteriorly to overlap ventral
corpus bursae. Anterior Vs of ventral ductus bursae with a sclerotized band.

Type Specimens! Choephora blanda Grote was described from two syntypes.
One specimen was located, a male in the BM(NH) labeled: Vancouver I,
Grote Coll 82-54 / 4425 Vancouver Island / Choephora blanda Type. Grote
/ Pseudoglaea blanda Grote / Syntype / Noctuidae Brit. Mus. slide No. 4925
male. It lacks antennae as is mentioned in the description. This specimen
is here designated lectotype. The holotypes of Glaea olivata Harvey,
Pseudoglaea taedata Grote, and Pseudoglaea decepta Grote are also in the
BM(NH). Photographs of these type specimens and their genitalia have
been examined.

Diagnosis: This species is variable in color and size. The range of color is
depicted in the illustrated specimens. Individuals from semi-desert locales

J. Res. Lepid.

tend to be pale while those from more mesic forest are darker. Most speci-
mens are brownish (Figs. 1-3), but reddish morphs (Figs. 4, 5) also occur
and can be common. M. olivata is most easily separated from both other
species by the presence of gray scales on its hindwings, but can also be de-
termined without dissection by the presence of long distal spines on the
first segment of the prothoracic tarsi. These are nearly equal in length to
the proximal spines in the other species. Both this species and M. rubra n.
sp. differ from M. subcuprea n. sp. in having the distal cornuti of the vesica
in two bands. These are thin in M. olivata and stout in M. rubra. Also, the
latter species has a coiled vesica while that of M. olivata is somewhat T-
shaped. The female genitalia of M. olivata differ from the other species in
that the anterior portion of the appendix bursae overlaps the ventral cor-
pus bursae.

Early stages: The larva has been described by Crumb (1956). It is a gen-
eral feeder on deciduous shrubs and trees. Crumb lists poplar, oak, hazel,
Amelanchier Medic., alder, antelope brush, Symphoricarpos Duhamel, and
Berberish. as foodplants. It has also been reared from Quercus garry ana Dougl.
and Ceanothus velutinus Dough in Oregon Q.C. Miller, pers. comm.) and
Quercus agrifoliaNee. in California (J. Powell, pers. comm.).

Distribution and flight period: This common species occurs from south-
ern coastal and interior British Columbia south through California, Colo-
rado, and Texas (Map 1). It most likely also occurs in northern Mexico.
The distribution records suggest that it is most common in the western
portion of its range. It occurs most often in dry open forest but also lives in
shrub steppe and mesic forest habitats. M. olivata is sympatric with both other
species. Adults have been collected from late August to November, with the
earliest flight in the northern part of its range.

Mesogona subcuprea Crabo & Hammond, new species
(Figs. 6,12,16; Map 2)

Description. Adults (Fig. 6) : Males and females identical in habitus. Spines
of first tarsal segment of prothoracic leg nearly equal. Head, palpi, dorsal
antennae, thorax, and ground color of wings light yellow brown; proximal
antennae and terminal space of forewing slightly lighter; abdomen reddish.
Forewing length: 19-21 mm. Forewing broader than in M. olivata, outer
margin prominently crenulate; lines and spots similar to M. olivata', orbicu-
lar and reniform prominent with filling darker than ground color. Hindwing
light copper-colored, slightly glossy, with faint median shade and discal dot.
Male genitalia (Fig. 12): Valves as in generic description; costal lobe broad,
nearly obsolete. Vesica 2 X as long as aedeagus, shaped like a lopsided T
beyond basal twist with short extension dorsad and toward right and longer
distal portion curved ventrad, cornuti divided into a patch of equal length
spines on a median diverticulum and a large subapical patch with multiple
minute and several massive rod-like spines. Female genitalia (Fig. 16): Cor-
pus bursae 2.5 X as long as wide, anterior Vs bent 90° to the right, with I

34:83-98, 1995(1997)

long dorsal and 1 short ventral signa; appendix bursae dorsoventrally flat-
tened and heavily sclerotized with irregular ridges, extending first posteri-
orly and to the right and then anteriorly to project to right of median cor-
pus bursae without overlap; ductus seminalis joins right anterior appendix
bursae. Anterior Vs of ventral ductus bursae with broad sclerotized band.

Type specimens: Holotype, S : WASHINGTON: Kittitas Co.: Reecer Cr.
atjohnson Cyn., 900 m, 47.16°N 120.62°W, 4.IX.1989, Lars Crabo. Paratypes,
32d, 229: WASHINGTON: Same data as type locality: I7.IX.1988 (2d),
4.IX.1989 (4d,59), 1.IX.1990 (1 9 ), 4.IX.1994, Troubridge & Crabo (8d,
119); Klickitat Co.: Lyle, 4 mi [6.4 km] N., 1500’ [457 m], 12.VIIL1960,
D.F. Hardwick (Gd, 2 9); Toppenish, 29 mi [46.7 km] S., 1800' [549 m],
23.VIIL1960, D.F. Hardwick (lOd, 39); Yakima Co.: Tieton River valley.
Oak Creek at Tieton River, Elev. 525 m, 46.72°N 120.81°W, 7.IX.1990, L.G.
Crabo, riparian with Garry Oak (Id); Kusshi Canyon, 17. IX. 1949, E.C.
Johnston (Id).

We restrict the type series to specimens from Washington state. The ho-
lotype is in the Canadian National Collection (CNC). Paratypes are in, or
will be deposited in, the CNC, the United States National Museum (USNM),
University of California (Berkeley) , University of California (Davis) , Oregon
State University (Corvallis), and the personal collections of Lars Crabo
(Bellingham, Washington) and Jim Troubridge (Langley, British Colum-
bia).

Diagnosis: This species is less variable than M. olivata or M. rubra. It can
be identified by the combination of yellow-brown ground color and light
copper-colored hindwings. It is the only North American species with a
median patch of spines on the male vesica and no overlap of the appendix
bursae and corpus bursae of the female genitalia.

M. superficially resembles M. acetosellae (Fig. 9) which occurs in

Eurasia. The male genitalia of M. acetosellae (Eig. 14a) differ from those of
the North American species by having a more massive valve with a large
rounded costal lobe of the sacculus. Its vesica (Eig. 14b) is most like that of
M. subcuprea. Both species have a median patch of cornuti on a diverticu-
lum, while the other species have two distal patches and no diverticula. Fur-
thermore, both M. acetosellae And M. subcuprealiAve at least one massive spine
in the subapical group. The female genitalia of M. acetosellae (Fig. 18) dif-
fer from all of the North American species by having a unisaccate bursae
copulatrix.

Early stages: The larva of M. subcuprea has been reared on Quercus agrifolia
at Big Creek, Monterey County, California (J. Powell, unpub. data) and Q.
dumosa Nutt, from the San Gabriel Mountains, Los Angeles County, Cali-
fornia (label data, L. Crabo collection), but has not been described. It is
closely associated with oak at many localities, but must also feed on other
genera since oaks are absent from the type locality.

Distribution and flight period: M. subcuprea is known from the east slope
of the Cascade Mountains and the eastern Columbia Gorge in Washing-
ton, from the Willamette Valley and the Klamath Mountains in Oregon,

J. Res. Lepid.

Figs. 1-10. Adults and larvae of Mesogona. 1) M. olivata 6, British Columbia,
Okanogan Falls, near Vaseaux Lk. 2) M. olivata $ , California, Mono Co.,
Benton Insp. Sta. 3) M. olivata, male, Washington, Grant Co., 1 .5 mi [2.4
km] N. of Wanapum Dam on Hwy. 243, 225 m. 4) M. olivata 6 , Oregon,
Douglas Co., Umpqua River valley. Thorn Prairie, 1040 m. 5) M. olivata
6, Washington, Skagit Co., Anacortes, S. slope of Sugarloaf, 900' [274
m]. 6) M. subcuprea 6, paratype, Washington, Kittitas Co., Reecer Creek
at Johnson Canyon, 900 m. 7) M. rubra 6 , paratype, Washington,
Skamania Co., Big Lava Bed, 3000* [914 mj. 8) M. rubra S , California,
B.T.I. Exp. For., Grass Valley. 9) M. acetosellae S, Digne, Gallia mer.
10) Last instar larva of M. rubra, Oregon, Josephine Co., Cave Junction.

34:83-98, 1995(1997)

Figs. 11-14. Male genitalia of Mesogona species. Vesica of aedeagus has been
everted (bar = 1 mm for genital capsule; 2 mm for aedeagus). 11) Male
genitalia of M. olivata, Oregon, Douglas Co., Umpqua River valley. Thorn
Prairie, 1040 m (a = valves; b = vesica). 12) Male genitalia of M.
subcuprea, Washington, Kittitas Co., Reecer Creek at Johnson Canyon,
900 m (a = valves; b = vesica). 13) Male genitalia of M. rubra, paratype,
Washington, Cowlitz Co., N. shore Lewis River between Yale Lake and
Swift Creek Reservoir, 580' [177 m] (a = valves; b = vesica). 14) Male
genitalia of M. acetosellae, Digne, Gallia mer. (a = valves; b = vesica).

J. Res. Lepid.

34:83=98, 1995(1997)

and from the Klamath Mountains, the Sierra Nevada, and Coast Ranges
south to Los Angeles in California (Map 2) . Adults have been collected from
mid August until early October. It emerges approximately one week ear-
lier than M. olivata at the type locality. Adults come to light, but are more
attracted to sugar bait at some localities.

Comments: Grote’s original description of Choephora blanda, including
“forewings... yellowish fawn...” and “hindwings silky reddish... with a trace
of median line” could pertain to either M. subcuprea or some specimens of
M. olivata. This hindwing description is especially suggestive of M. subcuprea
although some specimens of M. olivata have reddish hindwings with gray
scales. The Vancouver Island syntype of blanda, designated lectotype above,
is a typical M. olivata with fuscous hindwings and two subapical bands of
cornuti on the vesica. The other syntype from Washington Territory could
not be located in collections containing Grote type specimens (J.D.
Lafontaine, pers. comm.) and is presumed lost. It is likely that the lost
syntype was also a M. olivata despite the suggestive description since the
Vancouver Island specimen and M. subcuprea are dissimilar and would prob-
ably have been recognized as different species by Grote.

M. subcuprea is moderately common in collections, especially in material
from California, but has been confused with the more common M. olivata.

The name subcuprea refers to the copper color of the hindwings of this
attractive species.

Mesogona rubra Hammond Sc Crabo, new species
(Figs. 7, 8, 10, 13, 17; Map 3)

Description. Adults (Fig3. 7, 8): Males and females identical in habitus.
Spines of first tarsal segment of prothoracic leg nearly equal. Ground color
of head, palpi, dorsal antennae, thorax, abdomen, and forewings uniform
brownish red, appearing nearly immaculate. Forewing length: 18-21 mm.
Forewing 2 X as wide as long; margin undulating; lines double, inconspicu-
ous, evident mostly as the pale filling; basal line and median shade obso-
lete; antemedian line oblique, undulating, bent slightly basad at costa; post-
median line forming a laterally convex arc, its inner line absent or evident
as small dark dots in interspaces opposite cell; subterminal line sinuous, a
series of faint dark dots between veins; terminal line dark, barely evident;
orbicular and reniform spots faint, pale, similar in shape to those of M.

Figs. 15-18. Female genitalia of Mesogona species (bar = 2 mm). 15) Female
genitalia of M. olivata, Washington, Kittitas Co., Reecer Creek at Johnson
Canyon, 900 m. 16) Female genitalia of M. subcuprea, paratype, Wash-
ington, Kittitas Co., Reecer Creek at Johnson Canyon, 900 m. 17) Fe-
male genitalia of M. rubra, California, Diablo, 3 mi [4.8 km] NE, 2100'
[640 m]. 18) Female genitalia of M. acetosellae, PODOLE POLUDN., str,
KOP u Bedrykowce, Koroszow.

/. Res. Lepid.

Map 3. Map of Pacific Coast states showing distribution of examined material of
M. rubra.

olivata but filled with ground color. Hindwing immaculate, uniform red with
a slight sheen, terminal area and fringe lighter in some specimens.

Male genitalia (Fig. 13): Valves as in generic description; costal lobe small
and rounded. Vesica 2.5 X as long as aedeagus, coiled 360°, first ventrad and
toward right and then leftward to project to left of distal aedaeagus, with a
small flattened basal cornutus, distal Vs with two large fields of cornuti con-
taining both minute hairs and long spines, the latter as two longitudinal
bands one with longer spines than the other, the proximal portion of the

34:83-98, 1995(1997)

band of shorter spines elevated from surrounding vesica like the end of an
anvil. Female genitalia (Fig. 17)i Corpus bursae rounded, slightly wider than
long with blunt extension posteriorly to the right, with 1 long dorsal and 2
long ventral signa; appendix bursae bulbous, slightly rugose, extending
anteriorly and dorsally to overlap right side of dorsal corpus bursae; ductus
seminalis joined to left anterior appendix bursae. Anterior Vs of ventral
ductus bursae with a thin sclerotized band.

Type specimens: Holotype, d : OREGON: Linn-Lane Co. [Lane County]:
H. J. Andrews For., 11 mi [17.7 km] NE. Blue River, September 3, 1986 /
J.C. Miller LEPSTUDY, HJA Admin, site, 1500' [457 m] elev., ex. UV light
trap / 1. Paratypes, 26d, 5$: OREGON: Lane Co.: Elorence, 10.IX.1960,
Blk. Lt. Trap, K. Goeden (2$ ), LIX.1995,J. Troubridge (lOd); 0.2 mi [0.3
km] E. of S. Fk. McKenzie R. on Rd. to Cougar Reservoir, 44.15°N 122.25°W,
350 m, 14. IX .1991, L.G. Crabo, powerline cut/manzanita (Id); Lincoln
Co.: Newport, 15. IX. 1961, Blk. Lt. Trap, K. Goeden (Id); Linn Co.: Santiam
Pass, Hwy. 20, 16.IX.1993 / 3-1-A (Id), 29.IX.1993 / 3-1-B (1 d), 9.IX.1993
/3-LB (ld),22.IX.1993/3-LB (Id), lO.IX. 1993 / 3-1-B (Id), 15.IX.1993
/ 3-1-B (Id); Linn-Lane Co. [Lane Co.]: same as type locality, September
2, 1986 (Id), September 11, 1986 (Id), September 1, 1987 (Id); Linn-
Lane Co.: H.J. Andrews, [larva collected] 8. IV. 1986, reared (1 $ ), [larva col-
lected] 8.IV.1986, ex. Arctostaphylos Columbiana, 86-49 (1 $ ), [larva collected]
8.IV.1986, ex. Arctostaphylos Columbiana, 86-50 (1 9 ); WASHINGTON: Cowlitz
Co.: N. shore Lewis R. between Yale L. and Swift Creek Res., 46.05°N
122.25°W, 580' [177 m], 30.VIIL1994, A. & L. Crabo, small lava bed/man-
zanita (2d); S. Cascades, Dry Cr. 300 m E. of FR81, 1 mi [1.6 km] N. of
Merrill L., 46.1 1°N 122.32°W, 1620' [494 m.], 30.VIIL1994, leg L.G. Crabo,
pumice with lodgepole pine (2d); Skamania Co.: E. side of Big Lava Bed
on FR66, 2 mi [3.2 km] S. of South Prairie, 45.89° N, 121.72° W, 3000' [914
m], 29.VIIL1994, A. & L. Crabo, Lava flow, Lodgepole pine (Id).

We restrict the type series to specimens from Lane County, Oregon and
north in Oregon and Washington. The holotype will be deposited in the
CNC. Paratypes are in, or will be deposited in, Oregon State University
(Corvallis), USNM, University of California (Davis), and the personal col-
lections of Lars Crabo (Bellingham, Washington) and Jim Troubridge (Lan-
gley, British Columbia) .

Diagnosis: Most individuals of this species are easily recognizable by the
combination of red forewings and immaculate red hindwings. Populations
of M. rubra from Lane County, Oregon northward are uniformly of the deep
red to brownish red color morphs. The populations in California and south-
western Oregon are quite variable, with pink morphs (Fig. 8) common along
with the red morphs. These vary from pale whitish pink to a darker pinkish
gray. Some of the light-colored individuals resemble M. subcuprea, but lack
the well-defined orbicular and reniform spots on the forewing of this spe-
cies. Some red M. olivata morphs are also similar to M. rubra, but have gray
hindwings and more distinct forewing markings. M. rubra is the only North

J. Res. Lepid.

American species with a coiled male vesica and dorsal overlap of the ap-
pendix bursa with the corpus bursae in the female.

Early Stages: The larva of M. rubra (Fig. 10) is reddish brown in ground
color with a finely mottled pattern, and has a pale lateral stripe. This col-
oration blends with the reddish bark of Arctostaphylos. By contrast, the larva
of M. olivata reared from Ceanothus velutinus is pale whitish gray in ground
color with fine black lines and dots, and has a broad white lateral stripe
(J.C. Miller, pers. comm.). Larvae of M. rubra have been beaten from and
reared to adults exclusively on Arctostaphylos Columbiana Piper in Lane
County, Oregon and an Arctostaphylos species, possibly A. cinerea Howell, in
Josephine County, Oregon (J.C. Miller, pers. comm.). The larvae have been
collected during April and May. It probably utilizes A. nevadensis Gray in
Washington sites where A. columbiana does not occur. However, it is prob-
ably host restricted to certain species of Arctostaphylos, since it has never been
collected along the east slope of the Oregon Cascades in habitat with A.
patula Greene.

Distribution and flight period: This species occurs in the Cascade Moun-
tains north to Skamania County, Washington, in the Klamath Mountains,
on the Pacific coast from central Oregon to central California, and in the
Sierra Nevada (Map 3). It is sympatric with both other species at many lo-
calities, including with M. olivata at the type locality. M. rubra occurs in dry
forests with Arctostaphylos species, including lava flows in the Washington
and Oregon Cascades and forested dunes on the Oregon coast. It flies from
late August to mid October.

The red color of this species resembles the bark of the foodplant. This
feature is shared by some of the other Noctuid moths which feed on Arcto-
staphylos and madrone {Arbutus menziesii Pursh. — both Ericaceae) which
both have reddish brown bark. These include Orthosia mys (Dyar) , O. pulchella
(Harvey), and O. transparens (Grote). This is likely a protective adaptation,
although it is not known that the moths rest on the plants during the day.

Comments: This species is moderately common in California collections
but has been confused with M. olivata. It was first recognized as distinct from
M. olivata during a Lepidoptera survey of the H.J. Andrews Experimental
Eorest (USDA) performed by Jeffrey C. Miller of Oregon State University.

The specific epithet refers to the prominent red color of this species.

Acknowledgements: J. Donald Lafontaine searched North American collections for
the type specimen of Choephora blanda, arranged for photographs of all Mesogona
types in the British Museum (Natural History) to be sent to the senior author, made
specimens in his care available for study, recorded locality records for Mesogona
specimens from the United States National Museum, and provided encouragement.
This study would not have been possible without his help. Jim Troubridge photo-
graphed the adults and genitalia. Jeffrey C. Miller provided the photograph of the
larva of M. rubra. Eric Metzler (Columbus, Ohio), Jerry Powell (University of Cali-
fornia, Berkeley), Steve Hayden (University of California, Davis), Ron Robertson
(Santa Rosa, California) , Jon Shepard (Nelson, British Columbia), and Jim

34:83-98, 1995(1997)

Troubridge provided access to specimens in their care. Jim Troubridge and
Jonathan and Elizabeth Pelham reviewed the manuscript and made helpful sug-
gestions. Two anonymous reviewers provided additional helpful comments. The
H.J. Andrews Experimental Forest is a Long-Term Ecological Research Site funded
by the National Science Foundation. The initial discovery of M. rubra and its larval
biology was conducted at this site as part of a comprehensive Lepidoptera
biodiversity study by Jeffrey C. Miller. We also thank John D. Lattin and the Sys-
tematic Entomology Laboratory at Oregon State University for support of this work,
including partial funding through NSF grants BSR-85-14325, BSR-85-16590, BSR-
87-17434, and BSR-90-11663.

Literature Cited

Anderson, E.M. 1904, Catalogue of British Columbia Lepidoptera. British Columbia
Provincial Museum. King’s Printer. Victoria, British Columbia.

Barnes, W. & J. McDunnough. 1916. Synonymic notes on North American Hetero-
cera. Contributions to the Natural History of the Lepidoptera of North America,
3(3):155-208.

. 1917. Check list of the Lepidoptera of boreal America. Herald Press, Decatur,

Illinois.

Blackmore, E.H. 1927. Check-List of the Macrolepidoptera of British Columbia.

Provincial Museum of Natural History. King’s Printer. Victoria, British Columbia.
Blanchard, E. 1840. Histoire naturelle des insectes; orthopteres, neuropteres,
hymenopteres, lepidopteres et dipteres. Volume 3 512, In Castenau. Histoire
naturelle des animaux, annelides, crustaces, arachnides, myriapodes et insectes.
Dumenil, Paris.

Boisduval, J.B.A.D. de. 1840. Genera et index methodicus Europaeorum Lepi-
dopterorum. Roret, Paris.

Crumb, S.E. 1956. The Larvae of the Phalaenidae. United States Department of
Agriculture. Technical Bulletin No. 1135.

Denis, D.N.C.M. Be I. Schifeermuller. 1775 [1776]. Ankiindung eines Systematisches
Werkes von den Schmetterlinge der Wiener Gegend. Bernard!, Wien.

Dyar, H.G. 1903. A list of North American Lepidoptera and key to the literature of
this Order of insects. Bulletin of the United States National Museum, No. 52.
Fibiger, M. 1993. Noctuidae Europaeae. Vol. 2. Noctuinae 11. Entomological Press,
Sor0, Denmark.

Forbes, W.T.M, 1954. Lepidoptera of New York and neighboring states. Noctuidae,
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Franclemont, J.G. Be E.L. Todd. 1983, Noctuidae. In Hodges, R.W., et ah, eds. Check
list of the Lepidoptera of America North of Mexico. E.W. Classey Ltd. and The
Wedge Entomological Research Foundation, London.

Grote, A.R. 1875. On Scopelosoma and allied genera. Canadian Entomologist
7:205-207.

. 1876a, On Noctuidae from the Pacific Coast of North America. Bulletin of

the Buffalo Society of Natural Sciences 3:77-87.

. 1876b. On Choephora and allied genera. Canadian Entomologist 8:17-18.

J. Res. Lepid.

. 1878. Descriptions of Noctuidae, chiefly from California. Bulletin of the

United States Geological and Geographical Survey of the Territories 4:169-187.

. 1880. Descriptions of Noctuidae. Canadian Entomologist 12:152-157.

. 1881. North American moths, with a preliminary catalogue of the species of

Hadena and Folia. Bulletin of the United States Geological and Geographical
Survey of the Territories 6:257-277.

. 1895. List of North American Eupterotidae, Ptilodontidae, Thyatiridae,

Apatelidae and Agrotidae. Abhandlungen des Naturwissenschaftlichen Vereins
zu Bremen 14:43128.

Guenee, a. 1852. In Boisduval, J.B.A.D. de & A. Guenee. Histoire naturelle des
insectes. Species general des lepidopteres. Tome Cinquieme. Noctuelites. Tome
1. Roret, Paris.

Hampson, G.E. 1903. Catalogue of the Lepidoptera Phalaenae in the British Museum,
Volume 4. Taylor and Francis, London.

. 1906. Catalogue of the Lepidoptera Phalaenae in the British Museum. VoL

6. Taylor and Francis, London.

Harvey, L.F. 1874. Observations on North American moths. Bulletin of the Buffalo
Society of Natural Sciences 2:118-121.

Hubner, J. [1803]. Sammlung Europaischer Schmetterlinge. Volume 4. Eulen.
Augsburg. J. Hubner.

Jones, J.R.J. 1951. An annotated checklist of the Macrolepidoptera of British
Columbia. Entomological Society of British Columbia, Occ. Paper No. 1.

McDunnough, J.H. 1927. Notes on certain Agrotid genera and species (Lepid.).
Canadian Entomologist 59:64-66.

. 1928 [1929]. A generic revision of North American Agrotid moths. Canada

Dept, of Mines, Bulletin 55.

. 1938. Check list of the Lepidoptera of Canada and the United States of

America. Part 1. Macrolepidoptera. Memoirs of the Southern California
Academy of Sciences. Vol. 1.

Poole, R.W. 1994. Noctuoidea, Noctuidae (part). In Dominick, R.B., etak, eds. The
moths of America North of Mexico, fasc. 26.1. The Wedge Entomological
Research Foundation, Washington, D.C.

Smith, J.B. 1893. A catalogue bibliographical and synonymical of the species of Moths
of the Lepidopterous Superfamily Noctuidae found in boreal America. Bulletin
of the United States National Museum 44.

Journal of Research on the Lepidoptera

34:99-118, 1995(1997)

The endangered quino checkerspot butterfly, Euphydryas
editha quino (Lepidoptera; Nymphalidae)

Rudi Mattoni/ Gordon F. Pratt, ^ Travis R. Longcore,^ John F. Emmel,^ and
Jeremiah N. George^’ ^

^Urban Wildlands Group, UCLA Department of Geography, Box 951524, Los Angeles,
California 90095-1524

^Department of Entomology, University of California, Riverside, California 92521
^Hemet, California

Abstract. With the listing of the quino checkerspot butterfly, Euphydryas
editha quino, as a federally endangered species, research into its ecology
and conservation is necessary to allow for recovery planning and man-
agement. We review systematics, distribution, natural history, and con-
servation prospects, with reference to pertinent literature about other E.
editha subspecies. Additional information is presented from museum
specimens and ongoing research on the species.

Keywords. Quino checkerspot butterfly, Euphydryas, endangered species,
conservation

Introduction

The quino checkerspot butterfly, Euphydryas editha quino (Behr) 1863
(QCB or quino), was listed as an endangered species on January 16, 1997
(62 Federal Register 2313). The basis for the listing was habitat loss, degra-
dation, and fragmentation, recognizing additional negative effects from fire
management practice. All factors are the results of intensive human eco-
nomic development of ever diminishing resources. Recent loss of the dis-
tribution area of was estimated as 50-75%, with “seven or eight popu-
lations” known in the United States with “all but three populations” con-
sisting of fewer than five individuals (Nelson 1997). Surveys over the past
year indicate that although QCB may not seem in as dire circumstance as
the listing package indicated, with at least two robust metapopulations found
in two counties and numbering thousands of individuals, we believe the
species was correctly assessed as near extinction. QCB appears headed to-
ward becoming the “passenger pigeon” butterfly — a once common wide-
spread species crashing to extinction over a few decades. This would be
especially remarkable because an average female QCB lays over 500 eggs in
a season compared with two eggs for the passenger pigeon. We summarize
herein all pertinent data regarding QCB, discuss our reasoning for project-
ing its imminent disappearance in the absence of substantial effort, and em-
phasize the rather unique event this disappearance will be among the set
of all U.S. endangered butterfly species.

Paper submitted 31 October 1997; revised manuscript accepted 1 December 1997.

100

J. Res. Lepid.

ern California and Baja California, showing distribution of nearby sub-
species of Euphydryas editha. Legend: O quino pre-1 990, • quino post-
1990, ▲ insularis, ■ augustina, ♦ new subspecies, T editha.

Systematics

The QCB is one of over 20 recognized subspecies of Euphydryas editha
(Miller &: Brown 1981). Euphydryas editha quino is the most southwesterly
distributed taxon and is parapatric with three other subspecies (Fig. 1 ) : editha
(Boisduval) 1852, augustina (W.G. Wright) 1905, and a new subspecies on
the desert slopes of the Transverse Range to the southern Sierra Nevada. A
fourth subspecies, insularis (Emmel & Emmel) 1974, occurs in southern
California on Santa Rosa Island.

In adult appearance the QCB is distinguishable from all other subspecies
by size and relative cover of red, yellow, black, and white scaling forming
both upper- and underside maculation (Fig. 2). In nominotypical editha,
black scaling predominates on the uppersides of the wings, covering ap-
proximately 50% of the wing surface, with cream spots covering about 25-
30% and orange/red scaling covering about 20-25% of the wing surface.
E. e. quino is similar to nominotypical editha in size, but differs in that the
orange/red scaling is increased and cream spots are slightly larger. E. e.
augustina is markedly smaller than quino and is similar in maculation to quino
except that there is greater development of orange/ red scaling in augustina.
The desert slope Transverse Range segregate is intermediate in size between

34:99-118, 1995(1997)

101

quino ?ind augustina, and tends to have greater development of both orange/
red and cream scaling than either of these taxa. E. e. insularis is similar to
nominotypical editha in size but differs from that subspecies by greater de-
velopment of black scaling and greater reduction of the orange/ red scal-
ing relative to the cream scaling.

There are additional defining larval characteristics, but these have not
been systematically described for all subspecies (D. Murphy 8c G. Pratt,
unpub. data) . Foodplant utilization by QCB in the wild is restricted to Plan-
tago erecta E. Morris, possibly P. ovata Forsskal [=P. Eastw.], and

Castilleja exserta (A.A. Heller) Chaung 8c Heckard [=Orthocarpus pur-
purascens Benth.]. Among E. editha subspecies, this foodplant utilization
pattern is shared with nominotypical editha and insularis. In a study that did
not include insularis, Baughman et al. (1990) presented genetic evidence
that quino is more closely related to editha than other subspecies.

A contrasting view of E. editha W2is given by Scott (1986), who recognized
only three subspecies: editha, nubigena, and beani, and stated that “Dozens
of localized races have been named, but they all fit into these three ssp.” In
our opinion Scott’s view under-represents variation (see also Baughman 8c
Murphy, in press) .

There have been two recent nomenclatorial changes with the taxon. The
first was assignment of editha to the genus Occidryas (Higgins 1978). How-
ever, the erection of Occidryas, although accepted by a few uncritical au-
thors (e.g.. Miller 8c Brown 1981), was unsubstantiated by morphological
or genetic evidence. All objective authorities synonomized it to Euphydryas.
The other matter was recognition of quino as the correct available name
for the taxon which earlier had been referred to as wrighti (Emmel et al., in
press, a). Although Gunder (1928) associated the name quino With the
Euphydryas chalcedona complex, a critical examination of Behr’s description
as well as the geographic parameters of collecting in the 1860s places quino
with the E. editha species complex. A neotype for quino has been designated
and the type locality fixed as San Diego, San Diego County, California.

The following summarizes the nomenclatorial treatment of quino and the
three other named subspecies in southern California (format based on
Miller 8c Brown 1981).

EUPHYDRYAS Scudder
editha (Boisduval) MET IT AE A.

a. e. editha (Boisduval) MET IT AE A. Ann. Soc. Ent. France, (2) 10:304
(1852). Type locality restricted to Twin Peaks, San Francisco, California,
and lectotype designated, in U.S. National Museum, by Emmel et al. (in
press, b).

= bayensisSteiYxiitzky. Canadian Ent., 69:204-205 (1937). Type locality

102

J. Res. Lepid.

Hillsborough, San Mateo Co., California. Syntypes in California Acad-
emy of Sciences, San Francisco.

b. e. augustina (W.G. Wright) MELITAEA. Butts. W. Coast: 154 (1905).
Type locality San Bernardino Mtns., San Bernardino Co., California.
Holotype in California Academy of Sciences, San Francisco.

c. e. inmlarisT. Emmel &:J. Emmel.J. Res. Lepid., 13:131-136 1974(1975).
Type locality Santa Rosa Island, Santa Barbara Co., California. Holotype
in Los Angeles County Museum.

d. e. quino (Behr) MELITAEA. Proc. California Acad. Nat. Sci., 3:90
(1863). Type locality restricted to San Diego, San Diego Co., California,
and neotype designated, in California Academy of Sciences, San Eran-
cisco, by Emmel et al. (in press, a).

= augusta (W.H. Edwards) MELITAEA. Canadian Ent., 22:21-23
(1890). Type locality vie. San Bernardino, San Bernardino Co., Cali-
fornia. Lectotype in Carnegie Museum, designated by E.M. Brown,
Trans. American Ent. Soc., 92:371 (1966).

= wrighti (Gunder). Pan-Pac. Ent., 6:5 (1929). Type locality San Di-
ego, San Diego Co., California. Holotype in American Museum of
Natural History, New York.

The name augusta has been applied to the E. editha populations in the
San Bernardino Mountains since Comstock’s publication of The Butterflies
of California in 1927. However, examination of the lectotype specimen as
well as consideration of the type locality (vicinity of San Bernardino, spe-
cifically Little Mountain northwest of the city; see Coolidge 1911, for a de-
scription of a day collecting on Little Mountain with W.G. Wright, during
which he was told that this was the type locality for Melitaea augusta) clearly
places the low elevation, phenotypically large augustaWiXh quino. The name
augustina is based on an aberrant specimen from the San Bernardino Moun-
tains; because Wright considered it a new variety (his term for subspecies),
the name can be used in a subspecific sense for the small phenotype, higher
elevation San Bernardino populations of E. editha.

Populations of E. editha on the desert slope of the Transverse Ranges (San
Bernardino and Los Angeles counties) that use Castilleja plagiotoma Gray as
a larval host represent an undescribed subspecies; this taxon is being de-
scribed by Baughman and Murphy (in press).

In spite of the importance of E. editha to population biology theory, there
has been no recent revision of the overall species group. However, the pat-
terns of variation and approximate phylogenetic relationships of the taxa
surrounding E. editha quino are fairly well defined. Because of the sensitiv-
ity of E. editha senso lato to a suite of anthropocentric environmental im-

34:99-118, 1995(1997)

103

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104

J. Res. Lepid.

Table 1 . Localities for Euphydryas editha quino and most recent date of
collection or observation. A list of museum specimens is available from the

authors upon request.

Mexico

Estrado de Baja California

N of Ensenada 1935

Las Animas Canon 1935

Mosquito Springs 1936

Rodriguez Dam, Tijuana 1977

S of Salsipuedes 1979

N of Sordo Mudo 1979

Table Mt. (near Rosarita Beach) 1979

Turn off to Ojos Negros 1981

Valle de La Trinidad, Aquaito Spring 1994
N of El Testerazo 1996

S of El Condor 1996

California

San Diego County

San Francisquita Pass 1914

Warner’s Dam 1916

South San Diego 1917

Santa Fe Ranch 1930

Lake Hodges 1932

Rancho Santa Fe 1933

AltaVista 1934

Adobe Falls, San Diego 1948

Division Street, San Diego 1948

Vista 1951

Dehesa 1957

San Miguel Mt. 1957

El Cajon 1958

La Presa, San Diego 1958

Miramar 1960

Mission Gorge 1960

Tecate Mt. 1961

Fletcher Hills near El Cajon 1963

Sweetwater Dam/Reservoir 1969

Encanto 1969

Kearney Mesa 1969

Paradise Mesa, National City 1969

Spring Valley 1969

SE of El Cajon 1970

Proctor Valley 1971

OtayLake 1973

Mt. Palomar 1975

San Diego 1976

Chula Vista 1978

Little Cedar Canyon 1979

Mesa E of Otay Reservoir 1979

Otay Mesa 1980

Dictionary Hill 1981

Brown Field 1997

Otay Mt., ridge S of O’Neal Canyon 1997
South Otay Mt., Marron Valley 1997

Jacumba 1997

North slopes of Tecate Peak 1997

Riverside County

Sage 1951

Lake Elsinore 1983

Gavilan Hills 1985

Murrieta Hot Springs 1997

Aguanga 1997

Oak Mountain 1997

Temecula 1997

Lake Skinner 1997

Orange County

Hills E of Orange Co. (Irvine) Park 1917
Anaheim 1930

Laguna Lakes 1931

Hills N of Orange Co. (Irvine) Park 1934
Dana Point 1936

liwine Park 1937

Hidden Ranch 1967

Los Angeles County

Tapia Camp, Santa Monica Mts. 1947

Pt. Dume 1954

pacts now entrained, it would be well to document geographic variation
patterns and correlated natural history characteristics into a formal revi-
sion as quickly as possible.

Distribution

The few known persistent populations of the QCB are large in area, dis-
tributed as complex metapopulations. In attempting to reconstruct historic
QCB distribution, this hypothesis implies that specimens collected prior to
1940 most likely represent samples of extensive, and not small refugial,
populations. Maps of presumed historic vegetation communities (e.g.,
Kiichler 1977) and documented specimen localities indicate that the QCB
may have had an almost continuous distribution across cismontane south-

34:99-118, 1995(1997)

105

ern California from the westernmost Santa Monica Mountains, where dense
but local concentrations of Plantago erecta still persist, across the Los Ange-
les plain and margins of the Transverse Ranges into the desert in upper
Anza-Borrego and thence south into Baja California to about the northern
San Pedro Martir (Fig. 1; Table 1) . It was abundant on coastal bluffs in Point
Dume in western Los Angeles County, Orange County (John Johnson, in
litt. 1989 and see Orsak 1977), and the northern Baja California coast
(Brown et al. 1992). All the coastal bluff populations have probably been
destroyed with the possible exception of refugial colonies in the inacces-
sible coastal region between Ensenada and Cabo Colonet. During the past
20 years most of the coastal Baja terraces have been converted to high den-
sity agriculture.

By reasonable extrapolation, the first European missionaries to southern
California made large negative impacts that are now immeasurable. In ad-
dition to direct land conversion, they caused many destructive secondary
effects including introduction of grazing animals and many preadapted in-
vasive Mediterranean plant and invertebrate species, introduction of destruc-
tive agricultural practices, general resource depletion, and modification of
native American lifestyles. With open grass- and forb lands in the general
scrub communities taking the brunt of habitat destruction, the QCB from
that moment forward likely suffered more than any butterfly species of
southern California. The importance of harvested Plantago erecta as a major
grain resource of Native Americans provides some insight as to the quanti-
ties of this plant that were available, but are now more restricted. From the
initial missionary invasion in the 1770s, the tide of acculturated humanity
has unceasingly brought on natural habitat degradation by outright destruc-
tion, fragmentation, soil ecosystem disturbance, and explosions of nonna-
tive species. Nevertheless, as recently as the early 1900s, two flora of Los
Angeles reported that P. erecta was “Very common on dry plains and in the
foothills throughout our range [Los Angeles and Orange counties]”
(Abrams 1903) and “On dry hillsides throughout the south; the common
species” (Davidson &: Moxley 1923).

Any reconstruction of the former distribution of QCB is complicated by
relying on museum specimens, which provide only presence data, and then
only for localities frequented by collectors. Our recent discovery of popu-
lations across the southern slope of Otay Mountain and north of Tecate
Peak indicates that previous collection localities were far from exhaustive.
Casual collections rather than systematic surveys are the norm for our knowl-
edge of historic butterfly distributions. The geographic extent of collection
records, taken with the historic abundance of foodplant, leads to the pre-
sumption that quino wdiS once commonly, if patchily, distributed from Point
Dume to Ensenada and inland up to 60 miles (100 km).

Recently, Parmesan (1996) surveyed Euphydryas editha popula^tions across
the entire species range, sans the Rocky Mountain populations, to test the
hypothesis that global warming should cause “net extinctions to increase
in the south and at low elevations and to decrease in the north and at high

106

J. Res. Lepid.

elevations.” After censusing 151 previously recorded populations, she con-
cluded that there indeed was a correlation, acknowledging that the rela-
tionships expected were complex, particularly with regard to habitat destruc-
tion and its effect on recolonization. Given the complex population struc-
ture of E. editha, and our observation that human impacts were almost al-
ways involved in local extirpations in southern California (even for those
areas that may seem to still have “suitable habitat”) , the role of global warm-
ing as the proximate cause of extinction of E. e. quino populations must be
carefully evaluated. We suspect that warming is perhaps an exacerbating
factor, but that increased extinction rates in southern California are pri-
marily caused by more direct anthropogenic forces.

Natural History

The studies of Paul Ehrlich and his many students and colleagues have
produced a large body of information about Euphydryas editha as a species,
mostly concerning the bay checkerspot, Euphydryas editha editha {^bayensis']
(BCB) . Most of this work is applicable to the QCB (e.g., Ehrlich 1965, Labine
1965, Ehrlich et al. 1975, 1980, Ehrlich & Murphy 1987, Ehrlich & Wheye
1984, 1986, 1988, Launer & Murphy 1994, Murphy et al. 1983, Murphy Sc
Weiss 1988, Singer 1971, 1983, Singer & Thomas 1992, Baughman et al.
1990, Dobkin et al. 1987, White 1986, Weiss et al. 1987, 1988).

Life cycle

The QCB is univoltine with adults usually flying from late February into
April (but see anomalies in phenology below). Females usually mate only
once, and are “plugged” by males, which inhibits multiple copulations
(Labine 1964). Shortly thereafter gravid females begin laying egg masses
of 120-180 eggs (Ehrlich et al. [1975] record a minimum of 39 eggs per
mass for quino in the field), which hatch in 7-10 days. Murphy et al. (1983)
experimentally demonstrated in BCB that nectar feeding is essential to
maximize egg mass production beyond the initial two masses, and in all cases
subsequent egg number per mass decreased. Total egg production ranged
from about 400-800 per female. The emergent prediapause larvae undergo
two or three obligate moults, depending perhaps on the quality of the
foodplants, and then enter an obligate diapause as either third or fourth
instar larvae (G. Pratt, unpub. data). The prediapause larvae are gregari-
ous, usually spinning a communal web, whereas postdiapause larvae are
solitary.

Surviving larvae break diapause after winter rains of the next season are
sufficient to germinate and establish foodplant. These postdiapause larvae
go through three to perhaps seven or more additional instars and then
pupate, usually among low plants near the ground or under rocks if such
occur (G. Pratt, unpub. data, White 1986). Pupae mature and eclose in about
ten days. Once larvae enter diapause their survival rates likely increase given
that postdiapause larvae can repeat diapause at least once, and perhaps
several times (D. Murphy Sc G. Pratt, unpub. data). There is also variation

34:99==118, 1995(1997)

107

in larval coloration that may be geographic. White (1986) discusses several
less studied aspects of the life history of E. editha subspecies.

Because of their dependence on annual foodplants that senesce and dry
rapidly following the last rain of a season, prediapause larvae are the stage
most susceptible to mortality. If neonate larvae cannot find foodplant within
10 cm of the egg masses, they will starve (Singer 1972, Singer & Ehrlich,
1979). Singer found approximately 99% mortality in the prediapause co-
hort leaving little room for other factors, at least in the seasons of the years
studied. Singer and Ehrlich concluded that the major population regula-
tors were density independent, highly variable weather conditions. Predia-
pause larvae (BCB) survived under three different conditions: 1) if eggs
were laid when P. erecta would remain green for five more weeks, 2) if eggs
were laid on P. erecta in soil tilled by pocket gophers ( Thomomys bottae) , which
plants have deeper root systems and are generally more robust (see Hobbs
& Mooney 1985), or 3) if larvae were able to locate the larger secondary
foodplant Castilleja exserta (Singer 1972, Ehrlich et al. 1975).

Foodplants and nectar sources

Under field conditions the QCB essentially is restricted to the two larval
foodplants, Plantago erecta ducid Castilleja exserta, throughout its range. Where
present, Plantago ovata may be used although these plants are not usually
abundant in QCB territory. P. ovata may be a long-naturalized exotic spe-
cies from the Mediterranean region (Dempster in Hickman 1993). One larva
was observed on Keckiella antirrhinoides (Benth.) Straw (G. Ballmer, unpub.
data), a plant not common in QCB range. In the laboratory females ovi-
posit and larvae feed on other Plantago, Keckiella, and Penstemon, including
plant species found at QCB localities that are not used in nature. Although
the patterns of Euphydryas editha oviposition choice and larval foodplant
specificity have been elucidated in geographical context by Singer (1971,
1982, 1983), the physiological significance remains unknown. Experimen-
tal trials have not been conducted on quino to determine host preference.

Nectar sources are almost entirely small annuals that flower in synchrony
with appearance of adult QCB. These include Lasthenia spp., Cryptantha spp.,
Cilia spp., Linanthus dianthiflora, Salvia columbariae, and annual Lotus spp.
Most perennial plants are not in flower during the average QCB flight pe-
riod. However, we observed QCB nectaring at Eriodictyon spp. late in the
season.

Phenology and microclimate

Murphy and Weiss (Murphy & Weiss 1988, Weiss & Murphy 1988; see also
Weiss et al. 1993) provided a detailed study of fine scale distribution of the
BCB in terms of relative densities of both larvae and adults to slope and
exposure (microtopography) and the resultant microclimates produced by
insolation effect. They showed that the distribution of larvae, which were
highly dumped, changed between years depending on weather patterns,
and also moved in response to climatic factors. Position of larvae across the

108

J. Res. Lepid.

27-Dec

27- Nov

28- Oct

28- Sep

29- Aug

30- Jul

30- Jun
31 -May

1 -May
1 -Apr
2-Mar

31 - Jan
1 -Jan

M I I I I 11 I I I I I I I I I II I H-H-l-t U n i l { \ I

CD C3^ lO -i-

T- t- CM CO

O) Gi Oi G)

Fig. 3. Extreme collection dates of Euphydryas editha quino from museum speci-
mens. Lines connect dates assumed to be within the same flight sea-
son. Note the fall emergence of adults in 1910, 1948, 1957, and 1976.
All of these years had significantly greater than normal rainfall in Sep-
tember and October; 1957 and 1976 were El Nino years.

microclimatic strata affected their phenology and the timing of adult emer-
gence. They also determined during the four-year study that population den-
sity centers shifted, with resultant variability in rates of postdiapause larval
development to pupation and eclosion. The complex pattern of adult emer-
gence, oviposition, and foodplant status (senescence) is described in terms
of “phasing” to weather patterns in any season (Dobkin et al. 1987). These
results illustrate that persistence of complex metapopulations depends on
maintaining large and variable habitats with a broad range of microenvi-
ronments that may not be obvious at a glance.

Adults usually fly from February through April, but substantial variation
has been recorded. Known adult flight dates are shown in Fig. 3, tabulated
from museum specimens. Late fall adult emergence in 1910, 1948, 1957,
and 1976 is correlated with significantly greater than normal rainfall dur-
ing September and October (measured in San Diego) of those years, which
may or may not be associated with an El Nino/Southern Oscillation event
(1957 and 1976 were El Nino years). These extreme emergence dates sug-
gest that larval phenology is plastic; larvae are able to break diapause virtu-
ally anytime in response to rain sufficient to establish foodplant. However,
early adult emergence dates also require sufficiently warm weather as to
not slow larval development. Dobkin et al. (1987) suggested that El Nino
years were in fact detrimental to editha, because larval development and
subsequent adult emergence were delayed by the cool, damp thermal re-
gime more than foodplant vigor was prolonged — the butterfly and the
foodplant were “out of phase.” For El Nino, this condition may have been
unique to the Jasper Ridge colony studied, because the serpentine soil is

34:99-118, 1995(1997)

109

extremely porous and excess rainfall drains quickly. Drought, too, was shown
to be detrimental to editha populations (Ehrlich et ah 1980, Ehrlich &
Murphy 1987) . In sum, weather conditions may cause the time of adult flight
to vary anywhere from October to June.

Predators^ parasitoids^ and disease

Quantitative data on predation are available for the BCB, where mortal-
ity from parasitism in mature larvae was about 5% and in pupae about 50%
(Weiss et al. 1988, White 1986). The only QCB data are for 200 larvae col-
lected at Lake Skinner, of which three were parasitized by tachinid flies (K.
Osborne, pers. comm.). No other field data concerning predation or dis-
ease are available, although ground dwelling larvae must be vulnerable to
a number of spiders, ants, and carabid beetles. Nothing is known about QCB
diseases.

Mating behavior and hilltopping

Mating behavior is an important factor in population dynamics. At loca-
tions with high population densities of the QCB, mate locating usually in-
volves actively flying males seeking perched females. Females rest on the
ground or low plants near where they eclosed, with wings spread, awaiting
males. At locations where there is topographic relief combined with dis-
persed nectar and foodplant resources, females frequently move to high
points, ridges and hilltops, where they encounter perching males (see
Ehrlich & Wheye 1984, 1986, 1988). Here, males await females and usually
defend small territories.

The latter phenomenon, hilltopping, has been described and documented
for butterflies by Shields (1967) and is defined as “a phenomenon in which
males and virgin or multiple-mating females instinctively seek a topographic
summit to mate.” According to this theory, high ground, ridges, hilltops,
or even rock formations serve as visual beacons for sexual encounters. Lar-
val foodplant or adult nectar sources may or may not be present, but males
usually defend perches and/ or patrol territory. At sites where both nectar
and foodplant resources are also associated with “hilltops,” butterfly occur-
rence is adventitious and is not necessarily hilltopping unless mating can
be shown to be the purpose of butterfly presence. Nor is it the case where
hilltop presence is the result of “random” movement across high ground.
Unequivocally discriminating mate location from resource occurrence (and
resource seeking) on “hilltops” requires statistical analysis. Shields provided
quantitative data for one species, Papilio zelicaon, whereas a summary table
of species he presents as hilltopping (including quino) is not supported by
documented evidence. Regardless, however, there is a clear tendency among
many volant insects to congregate at high ground regardless of sex or re-
sources (see refs, in Shields 1967).

While Ehrlich and Wheye (1984, 1986, 1988) presented evidence support-
ing hilltopping in E. editha, Singer and Thomas (1992) disagree. They ar-
gue that hilltopping, defined as a behavioral preference for a resource, can-

110

J. Fks. Lepid.

not be distinguished using measures of resource use (e.g., sex ratio on hill-
tops). Rather, to show hilltopping, one must observe a tendency in indi-
vidual males or virgin females to move toward hilltops, or a trend for mat-
ing location to be closer to hilltops than emergence location, neither of
which has been shown for any subspecies (Singer & Thomas 1992).

Singer and Thomas’ argument does not suggest that butterflies are not
found on hilltops; it only questions the explanation for their presence.
However, determination of the ecological and evolutionary role of the dis-
tribution of E. editha, especially quino, on hilltops is of important conserva-
tion value. If indeed quino congregate on hilltops to mate, the conserva-
tion value of those hilltops will be great.

Our observations across southern San Diego County during spring 1997
(Pratt et al. 1997) provided evidence of QCB using hilltops, although in-
sufficient data were collected to prove hilltopping as prescribed by Singer
and Thomas (1992) . Our survey team found virtually all QCB as “hilltoppers”
in the sense of appearing to be concentrated on ridges and peaks. Across
the slopes of Otay Mountain and Tecate Peak, individuals (mostly oviposit-
ing females) were found infrequently on lower slopes in comparison with
ridges. By contrast, QCB populations across extensive flat grasslands, as in
the vicinity of Murrieta, are found where there is little or no relief that pro-
vides hilltops (G. Ballmer, pers. comm.). There are also large expanses of
Plantago erecta and Castilleja exserta with abundant nectar from sites where
the species has been extirpated (Gavilan, March AFB, etc.), sites both with
and without relief. Dense, shrub-covered areas, including high relief sites,
do not have QCB populations. Thus the determination of whether a specific
upland, ridge, rock outcrop, or hill serves for hilltopping behavior remains
subject to interpretation and depends on the areography of the quino ag-
gregates in question, their place in the vegetation matrix, and population
density.

Population cycles and structure

Long-term studies initiated by Paul Ehrlich on the BCB in 1959 provided
quantitative data showing large fluctuations in population density from year
to year. As his work progressed it became apparent that the fluctuations
were caused primarily by weather patterns, principally rainfall quantity and
timing. After the major drought years, populations crashed, then variably
recovered with return of favorable rains (Ehrlich et al. 1980). In the past
two years, however, his major study population at Stanford’s Jasper Ridge
seems to have been extirpated. Although there are only anecdotal records
on the QCB, cyclic fluctuations have been recorded.

The late John Johnson (in litt. 1989) observed quinoior over 60 years in
Orange County and noted significant changes in densities over time. The
QCB was collected in abundance at Irvine County Park between 1917 and
1922 and then apparently almost disappeared until 1928. In 1933 and 1934
the species was again common, but vanished thereafter and was never seen
again. A nearby colony about 0.5 miles (0.8 km) southwest of Hidden Ranch

34:99-118, 1995(1997)

111

in Black Star Canyon, Santa Ana Mountains, was known from the 1920s to
1930s. After two decades without records James Mori found the butterfly
abundant in March 1967. A severe fire in November 1967 burned the area
and the butterfly has not been seen since. Two large reservoirs were con-
structed near Irvine Park and the whole area has been subjected to ever-
increasing trampling over the 30 years since Mori found the last QCB in
this part of the Santa Ana Mountains.

Harrison (Harrison et ak 1988, Harrison 1989) has proposed a metapopu-
lation model for the BCB, a description which probably also fits the QCB.
A metapopulation is a set of populations that are usually demographically
independent (as Ehrlich found among the three populations of BCB at Jas-
per Ridge, 1965), but that are “interdependent over ecological time”
(Harrison 1988). The evidence from edithais that local populations vary
independently and occasionally suffer extinctions, but are recolonized from
other populations. At Morgan Hill, there is a “reservoir” population that is
large, stable, and much less likely to suffer extinction, even during a bad
year. Surrounding smaller patches are periodically recolonized from the
reservoir population. Because of the sedentary nature of E. editha, these small
patches of once-occupied habitat may remain unoccupied for long periods
before being recolonized (Harrison 1989).

Current data are insufficient to describe conclusively the population struc-
ture of quino, but observed patterns and anecdotal evidence suggest that it
is similar to that of BCB. The distribution observed during 1997 surveys on
Otay Mountain was patchy, with the butterfly exploiting temporally limited
resources in some localities (post-fire chaparral, see below). Localities are
separated by several to tens of kilometers, and can be assumed to be demo-
graphically isolated. The existence of a reservoir population has yet to be
shown. QCB could have a true metapopulation structure (small patches,
low dispersal) or a core-satellite structure typified by a reservoir population
and smaller outlying habitats.

In the Gavilan Hills, Riverside County, anecdotal accounts of abun-
dance and distribution seem to be consistent with a core-satellite popula-
tion structure. At one location, on private land near Harford Spring Park,
quino W3.S abundant and always present, according to accounts from collec-
tors reaching back to the 1930s. QCB were also found on outlying patches
as far as 5 miles (8 km) distant (G, Pratt, unpub. data), but never in the
numbers or consistency as adjacent to Harford Spring Park. In 1984 the
landowner disked the presumptive reservoir population, completely destroy-
ing its habitat value. The butterfly subsequently disappeared in the surround-
ing region.

Plant community associations

The QCB is not associated with a single plant community, as are many
butterflies, but instead with open spaces within several communities. Fur-
thermore, QCB resource and climatic requirements are met, over the long
term, by dynamic relationships that we can only generally recognize and at

112

J. Res. Lepid.

present describe rather imprecisely. The butterfly is found within several
plant community types from scrub on coastal bluffs, through coastal sage
scrub, chaparral, oak woodland, to desert pinyon-juniper woodland. In all
these communities, however, it is only found in openings within the domi-
nant plant community where there is sufficient local cover of the larval
foodplants, which usually co-occur with the annual forbs that provide most
nectar for adults. Sufficient foodplant density has yet to be determined; at
Lake Skinner, QCB have occupied areas with foodplant densities as low as
one plant per square meter (K. Osborne, pers. comm.). The butterfly does
not occur in extensive open grasslands, nor does it occur in dense (without
small clearings) coastal sage scrub, chaparral, or oak woodland. Plant com-
munity structure, and not dominant species composition, is the critical fac-
tor for QCB populations. The optimum habitat for oviposition and larval
development consists of patchy shrub or small tree landscapes with open-
ings of several meters between large plants. Landscapes with alternating
open swales and dense shrub patches also provide habitat.

Among known colonies, there is usually some topographic relief such as
raised mounds, low to high hills, slopes, and ridges. The species was com-
mon on Otay Mesa before urbanization; the natural landscape was one of
vernal pool depressions alternating with a relief of mima mounds. Prior to
widespread habitat destruction, the species was apparently abundant on
coastal bluffs, which were characterized by sparse low vegetation.

Plant community identity as normally construed (i.e., dominant cover) is
less helpful in defining habitat than is consideration of larval foodplant

abundance and distribution, nectar source availability, and microtopogra-
phy. In addition, cryptobiotic crusts and episodic disturbances such fire and
light grazing contribute both to creating and maintaining suitable habitat.

Cryptobiotic crusts. In surveys for stands of Plantago erecta on Otay Moun-
tain, we observed that the species was correlated with the presence of un-
disturbed cryptobiotic crusts (also called cryptogamic or microbiotic crusts,
St. Clair &: Johansen 1993). Cryptobiotic crusts are formed in soils in arid
environments by blue-green algae, lichens, mosses, and other lower plant
species, as well as fungi and bacteria (Belnap 1993). Research has shown
that cryptobiotic crusts increase the ability of the soil to hold moisture and
decrease its susceptibility to erosion through the adhesive qualities of mu-
cilaginous polysaccharides exuded by certain blue-green aglae and fungi
(Belnap & Gardner 1993). They also improve the availability of essential
minerals (N, P, K, Ca, Mg, Fe) for higher plants and provide conditions
that promote mycorrhizal associations (Harper Sc Pendleton 1993). Crusts
are easily disturbed by trampling, especially by cattle. At Otay Mountain,
we observed that P. erecta and other native annual species (e.g., Lasthenia
sp., Castillejasp., Lepidiumsp.) were more often found in areas that had crusts
intact, as identified by their characteristic patina and the presence of small
mosses. In general, the proportion of native to exotic plant species was ob-
served to be larger in areas with intact crusts. We speculate that crusts serve
the role of “gatekeeper,” allowing the germination of native species and

34:99-118, 1995(1997)

113

perhaps inhibiting exotic species. However, crust areas have more “bare”
ground (actually occupied by lichens, small mosses, algae, etc.) than non-
crust areas, a characteristic preferred by the QCB. Cryptobiotic crusts are
also usually darker (and thereby warmer) than surrounding soils (Harper
& Pendleton 1993), making them attractive locations for QCB thermoregu-
lation. The combination of native annuals (foodplant and nectar sources)
and open ground may be encouraged by different edaphic factors (e.g.,
high clay content) in other areas. The BCB is found in grasslands defined
by serpentine soils, which, much like crusts, support sparse native vegeta-
tion.

Grazing. In areas of heavy grazing, the annual plant cover at Otay Moun-
tain was largely dominated by Erodium spp. (mostly E. botrys). In grazed ar-
eas, Plantago erecta was absent, all available space being preempted by the
prostrate storksbills. P. erecta tended to occur in areas that would be less
accessible to cattle, such as steep or rocky areas. Our observations about
cryptobiotic crusts suggest a pathway of replacement wherein trampling by
cattle disrupts the crusts, allowing establishment of the exotic Erodium, which
in turn excludes P. erecta. Cattle also disperse Erodium seeds, thus further
facilitating the invasion. Such animal-mediated disturbance has been im-
plicated elsewhere in the spread of alien plants (Schiffman 1997a), and the
quantity of seed dispersed by cattle has been shown to be enormous (Malo
8c Suarez 1995). However, light grazing may serve to maintain QCB habitat
by promoting forb-dominated, intermediate successional grassland stages,
as discussed for the southern habitat patches of the BCB by Murphy and
Weiss (1988). But too much grazing has been implicated in local extirpa-
tions (Murphy 8c Weiss 1988). Light grazing by native ungulates was his-
torically present throughout the QCB range, and emulation of it may in-
deed be necessary to maintain stable habitat areas. Also, regular disturbance
by fossorial rodents may have contributed to maintaining areas dominated
by annuals (Schiffman 1997b, Longcore, in prep.). Such disturbance by
pocket gophers has already been shown to contribute to foodplant quality
and BCB larval survival (Hobbs 8c Mooney 1985, Ehrlich 8c Murphy 1987).

Fire. Areas on the western side of Otay Mountain occupied by QCB in
1997 were in early post-burn succession. Adult QCB, Plantago erecta, and
ample nectar sources were found throughout recently burned areas. QCB
distribution was limited by the edge of the burn, which was marked by dense,
mature chaparral. Although in some areas P. erecta distribution is stable, it
can also be found tracking disturbance, with a distribution variable in both
space and time. Like other “fire-followers,” P. erecta grows well following dis-
turbance (usually fire, but also other one-time events), sets large amounts
of seed, and then thins out as the canopy is closed by the regenerating shrub
layer. The regionally dynamic metapopulation structure of the QCB is
adapted to such geographic and temporal variation in foodplant distribu-
tion.

The variable and synergistically interacting factors that contribute to ap-
propriate quino habitat make defining essential areas for species survival

114

J. Res. Lepid.

difficult. What is one year closed canopy chaparral may the next year be
covered with foodplant and flowering annuals, posing a special challenge
to conservation efforts. Protecting sufficient habitat may mean protecting
large enough areas to allow for a natural fire regime to maintain a shifting
mosaic of habitat patches.

Conservation Planning

With exception of the QCB and the BCB, all Nearctic butterflies listed
under the Endangered Species Act have restricted distributions and/or
highly specific habitat requirements. The threatened Earner blue butterfly
(Lycaeides melissa samuelis) has a 1,000-mile wide geographic distribution,
but is restricted to small dynamic successional habitat patches that support
its one foodplant. The highest extinction probability is for species found
only at single small sites. One limited catastrophe could destroy them: e.g.,
Lange’s metalmark {Apodemia mormo langei) and Palos Verdes blue butterfly
( Glaucopsyche lygdamus palosverdesensis) .

By contrast, the QCB had a large range (ca. 200 X 60 miles [320 X 100
km], now reduced by over half), occurring over a continuum of climatic
regimes from wet coastal to high desert; it is still found in several plant com-
munities although it has only two hostplants, and likely maintains substan-
tial genetic variation both hidden and expressed by local ecotypes. The key
to its conservation will be management of the surviving populations under
the assumption that they conform to a classic metapopulation structure.
The fundamental feature of this scenario is the vulnerability of any
metapopulation following the permanent loss of any of its demes (subpopu-
lations) or fragmentation that would destroy dispersal patterns that con-
nect them.

To ensure the conservation of the QCB, there must be some critical num-
ber of interconnected demes to provide a population structure with suffi-
cient habitat variation that a viable effective population size is always main-
tained in some part of the metapopulation unit (Murphy 8c Weiss 1988).
Available data do not permit even one metapopulation to be circumscribed
even though at present there are three fairly large (each ca. 40-150 square
miles [100-390 sq. km] ) areas of distribution that may support at least one
metapopulation: Otay Mountain, Temecula-Oak Mountain-Anza, and north
central Baja. Although small refuge colonies may yet be found in parts of
the historic range, as in Orange County and northern San Diego County,
these colonies will be at high risk unless appropriate management plans
are implemented to assure their survival, which may include providing cor-
ridors.

Murphy and Noon (1992) , using the northern spotted owl as an example,
provided a useful exercise in applying rigorous hypothesis tests to reserve
planning. Their approach, which was to identify the minimum number of
populations necessary to ensure species persistence, was a pioneering at-
tempt to offset the usual socioeconomic constraints in conservation plan-
ning. Their first task was to determine if the data supported rejection of

34:99-118, 1995(1997)

115

the null hypothesis that the finite rate of population change (X) was > 1.0.
The null hypothesis was rejected, leading to the recognition that their tar-
get species was in fact on the path to extinction (data concerning the QCB
concur). Murphy and Noon then proceeded to test nine more hypotheses
and concluded with a conservation map and strategy that were logically con-
sistent.

Unfortunately, field data currently available are insufficient to provide a
testable set of null hypotheses from which to design a reserve and manage-
ment program for the QCB. The only operable current reserve design ap-
proach will be to maintain large contiguous parcels of land that will con-
tain most, if not all, of the remaining metapopulations. The extent to which
quino can tolerate limited development on these parcels currently cannot
be assessed without further research on the autecology of the species. Ten-
tative conservation requirements must include care to not overgraze, atten-
tion to the fire regime, and security of core cryptobiotic crust areas to pre-
clude trampling. Whether sufficient land to preserve the species can be set
aside, either through public ownership or voluntary conservation agree-
ments with private landowners, remains to be seen.

Acknowledgments. Collection data were graciously provided by Robert Allen, Greg
Ballmer, John Brown, Tom Dimock, Bruce O’Hara, Ken Osborne, Camille
Parmesan, and John Pasko. Chris Nagano facilitated field research and found the
first quino at Marron Valley. Field surveys in southern San Diego County were sup-
ported by the U.S. Bureau of Land Management and the U.S. Fish 8c Wildlife Ser-
vice. Manuscript preparation was supported by a National Science Foundation
graduate fellowship to Travis Longcore. Editorial comment from Catherine Rich
significantly improved the manuscript; all remaining errors are the responsibility
of the authors. We especially thank David Liittschwager and Susan Middleton for
providing the striking portrait of quino on the cover of this volume. Their photo-
graphs remind us why we care.

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Journal of Research on the Lepidoptera

34:119-141, 1995(1997)

Immature stages of high arctic Gynaephora species
(Lymantriidae) and notes on their biology at Alexandra Fiord,
Ellesmere Island, Canada

Wm. Dean Morewood^ and Petra Lange^

^Department of Biology, University of Victoria, Victoria, B.C., V8W 3N5, Canada
-Department of Biological Sciences, Simon Fraser University, Burnaby, B.C., V5A 1S6, Canada

Abstract. Two species of Gynaephora are found in North America and their
geographic ranges overlap broadly in the Canadian Arctic. Despite nu-
merous studies that have addressed aspects of the biology, ecology, and
physiology of these species, confusion regarding identification of their
immature stages, originating with the original description of the first of
the two species discovered, persists even in recent literature. In this pa-
per, for the first time, all immature stages of both species are described
and most are illustrated, with emphasis on the differences between the
two species that allow for their identification.

Eggs and pupae of the two species are very similar morphologically but
usually may be distinguished by association with cocoons and also by size
at Alexandra Fiord, Ellesmere Island. In first instar larvae, the cuticle is
black in G. groenlandica but pale in G. rossii’, older larvae are readily iden-
tified by distinct differences in the color patterns of the larval hairtufts
and by the form of the hairs, being spinulose in G. groenlandica and pre-
dominantly plumose in G. rossii. Cocoons usually may be distinguished
by color but this feature is variable while the structure of the cocoons,
double-layered in G. groenlandica and single-layered in G. rossii, is defini-
tive.

Field studies conducted at Alexandra Fiord revealed some gaps and
inaccuracies in previously published life history information. Egg masses
laid on cocoons were found to suffer extensive predation by birds, a source
of mortality that was previously overlooked. There appear to be six larval
instars in G. rossii but seven in G. groenlandica rather than six as previ-
ously reported. Seasonal activity patterns of larvae were found to differ,
with G. groenlandica active only in the early part of the growing season
and G. remaining active in late summer. Foodplant preferences also
differed, partly as a result of the different food sources available at dif-
ferent times during the spring and summer. Finally, larval hairs of these
species have been found to have urticating properties, causing skin irri-
tation after extensive handling of larvae or cocoons.

Keywords. Gynaephora groenlandica, Gynaephora rossii, eggs, larvae, pupae,
cocoons, morphology, seasonal activity, foodplants

Paper submitted 23 January 1997; revised manuscript accepted 11 August 1997.

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J. Res. Lepid.

Introduction

The genus Gynaephora Hiibner ( Lyman triidae) is represented in North
America by two species, G. groenlandica (Wocke [in Homeyer] 1874) and G.
rossii (Curtis 1835). The geographic distribution of G. groenlandica \s almost
entirely limited to Greenland and islands of the Canadian arctic archipelago;
that of G. rossii includes most of the North American Arctic (excluding
Greenland) and Siberia, with isolated populations occurring in alpine ar-
eas of Japan, New England, and the southern Rocky Mountains (Ferguson
1978, M0lgaard & Morewood 1996). Gynaephora groenlandicahdiS the distinc-
tion of ranging to the most northerly point of land in Canada (Ward Hunt
Island, 83°N; Downes 1964) as well as northernmost Greenland (Wolff 1964)
and is considered to be a high arctic endemic species (Munroe 1956, Downes
1964) whereas G. rossiih^s a typical arctic/ alpine distribution.

Early accounts of arctic Gynaephora species are numerous, mostly consist-
ing of descriptions and natural history observations (Curtis 1835, Homeyer
1874, Grote 1876, Packard 1877, Scudder et al. 1879, Skinner & Mengel
1892, Dyar 1896, 1897, Nielsen 1907, 1910, Johansen 1910, Gibson 1920,
Forbes 1948, Bruggemann 1958). Later authors emphasized the apparent
adaptations of these insects and others to the extreme conditions of the
arctic environment (Downes 1962, 1964, 1965, Oliver et al. 1964, Oliver
1968). More recent studies have investigated the biology, ecology, and physi-
ology of arctic Gynaephora species in order to elucidate and understand the
various ways in which they are adapted to arctic conditions (Ryan 1977, Ryan
& Hergert 1977, Schaefer & Castrovillo 1979, Kevan et al. 1982, Kukal 1984,
Kukal & Kevan 1987, Kukal, Heinrich &: Duman 1988, Kukal, Serianni &:
Duman 1988, Kukal & Dawson 1989, Kukal et al. 1989, Kevan &: Kukal 1993,
Kukal 1995, Lyon & Gartar 1996).

Despite the attention that arctic Gynaephora species have received, there
remains confusion regarding identification of the immature stages. For
example, Kevan et al. (1982) ostensibly studied G. rossiihnt published pho-
tographs of a larva, cocoons, and even an adult that are clearly G. groenlandica.
Furthermore, Ryan and Hergert (1977) considered the two species to be
“identical in their food choices and development, and almost identical mor-
phologically”; however, there are considerable differences, both morpho-
logically and ecologically. The purpose of this paper is to describe and il-
lustrate the immature stages of G. groenlandica and G. rossii, with emphasis
on differences between the species, and to update information on their natu-
ral history as observed at Alexandra Fiord, Ellesmere Island, Canada.

Methods and Materials

Fieldwork was conducted at Alexandra Fiord (78° 53' N, 75° 55' W) on the east
coast of Ellesmere Island from 6.VI.1994 to 15.VIII.1994, from 29.V.1995 to
17.VIII.1995, and from 25.V.1996 to 13.VIII.1996. The study site consists of a small
(about 8 km^) lowland valley bounded by glaciers to the south, upland polar desert
and fellfield to the east and west, and the fiord itself to the north. This site has
been subject to a considerable amount of ecological research (cf. Svoboda & Freed-

34:119-141, 1995(1997)

121

man 1994) and is described as a “polar oasis,” noted for its relatively lush vegeta-
tion compared to the surrounding polar desert (Freedman et al. 1994), Popula-
tions of both species of Gynaephora occur at Alexandra Fiord, although G.
groenlandica is far more abundant there than is G. rossii

Larvae, cocoons, adults, and eggs of both species of Gynaephora were observed
and photographed in the field and were collected for rearing and for more de-
tailed examination. Dimensions of eggs and maximum widths of larval head cap-
sules viewed from the front were measured to the nearest 0.05 mm using a stere-
omicroscope equipped with an ocular micrometer, at a magnification of 20 X . Early
larval instars were determined by rearing larvae from eggs and measuring head
capsules shed at each moult. Head capsule width (HCW) for the final instar was
determined by measuring head capsules from larvae that had been killed by para-
sitoids after spinning cocoons, indicating that they were in their final stadium. Mean
HCW for each of the intermediate instars was estimated by extrapolating from the
mean HCW of the early and final instars according to the Brooks-Dyar Rule (Dyar
1890, Daly 1985) and these estimates corresponded well with peaks in the distribu-
tion of measured HCW for G. groenlandica. The distribution of HCW overlapped
for these intermediate instars and therefore sample statistics were calculated by
dividing the HCW distribution at the low points between peaks. Due to this over-
lap in HCW between intermediate instars and the very limited number of actual
HCW measurements for the intermediate instars of G. rossii, the given HCW for
these instars should be considered approximations only. Descriptions of the later
instars were obtained by measuring the head capsules of larvae examined in detail
and assigning these larvae to the appropriate instar. These descriptions were supple-
mented with field observations of larval phenotypes, especially larvae that were spin-
ning cocoons, indicating that they were in their final stadium. Descriptions of lar-
vae follow the terminology used by Ferguson (1978),

Photographs of larval hairs and portions of cocoons were taken through a stere-
omicroscope at a magnification of 30 X . Maximum lengths and widths of cocoons
viewed from above were measured to the nearest millimeter using a plastic ruler;
sexes were subsequently determined from the morphology of caudal segments of
the pupal exuviae (cf. Fig. 1 ) . Maximum lengths and widths of pupae in ventral view
were measured to the nearest half millimeter using a plastic ruler; very few pupae
were measured because most were left to develop within their cocoons for other
studies. Descriptions of pupae follow the terminology of Mosher (1916) and were
formulated to be comparable to those published by PatoCka (1991) . Measurements
are given as mean ± standard deviation, followed by the sample size in brackets, and
are rounded off to the level of precision of the original measurements; statistical
tests were conducted as described by Zar (1984) before rounding off the data.

Foodplant preferences were determined by recording the plant species and part
of the plant eaten by all Gynaephora larvae that were observed actively feeding on
the tundra in 1995 and 1996; these observations were limited to free-ranging larvae.

Results

Descriptions of Immature Stages

Eggs. Eggs laid in masses covered by hairs rubbed from the abdomen of

122

J. Res. Lepid.

the female, typically on the cocoon from which the female emerged but
also frequently on vegetation or the ground (Plate 1). Eggs of both species
smooth, creamy white, and roughly spherical but somewhat flattened.

G. groenlandica: 1.60 ± 0.05 mm in diameter by 1.35 ± 0.05 mm in height
(n = 10).

G. rossii: 1.40 ± 0.05 mm in diameter by 1.10 + 0.05 mm in height (n =

10).

Eggs of G. significantly smaller than those of G. groenlandica =
15.345, P < 0.0005 for diameter; t^^^^g = 15.545, P < 0.0005 for height), this
difference visible even to the unaided eye.

Larvae. Larvae of both species large and very hairy, superficially resem-
bling larval Arctiidae (Plate 2). The following general description, outlin-
ing the basic arrangement of verrucae and hairs, applicable to all instars of
both species; modifications and species-specific differences described in the
subsequent sections. Differences only noted in the specific descriptions;
larvae of a given instar correspond to the description given for the previ-
ous instar except as described otherwise.

Head capsule black and bearing many hairs. Addorsal, subdorsal,
supraspiracular, subspiracular, and subventral verrucae present on mesotho-
rax, metathorax, and abdominal segments 1 through 8. Addorsal verrucae
fused with subdorsal verrucae on prothorax and abdominal segment 9. On
prothorax, supraspiracular verrucae greatly reduced, sometimes lacking
hairs, and subspiracular verrucae enlarged and oriented anteriorly. Except
as just noted, all verrucae bearing from one to many hairs. Hairs arising
from addorsal and subdorsal verrucae generally thicker than those arising
from supraspiracular, subspiracular, and subventral verrucae. Hairs arising
from supraspiracular and subspiracular verrucae, and from dorsal verru-
cae on abdominal segment 9, up to two or three times as long as the long-
est hairs arising from addorsal, subdorsal and subventral verrucae. Cuticle,
including verrucae, entirely black except where noted below. Dorsal glands
on abdominal segments 6 and 7 whitish and well developed in all instars
except the first.

G. groenlandica: First instar HCW = 0.70 ± 0.05 mm (n = 140) . Correspond-
ing to the general description above. Cuticle between verrucae black. Hairs
arising from addorsal and subdorsal verrucae black, hairs arising from
subspiracular and subventral verrucae brown, and hairs arising from
supraspiracular verrucae mixed. All hairs spinulose.

Second instar HCW = 0.95 ± 0.05 mm (n = 85). All verrucae bearing a
mixture of black and brownish yellow hairs. Hairs arising from supra-spi-
racular, subspiracular, and subventral verrucae predominantly yellow. Hairs
arising from addorsal and subdorsal verrucae predominantly yellow on
mesothorax and metathorax, black on abdominal segments 1, 2, and 8, and
yellow on abdominal segments 3 through 5.

Third instar HCW = 1.30 + 0.05 mm (n = 30). Hairs more dense than in
previous instars, beginning to obscure underlying verrucae from which they

34:119-141, 1995(1997)

123

Plate 1 . Gynaephora groenlandica (A-D). Female ovipositing on the cocoon from
which she emerged; male still present to the right (A). Female (arrow)
ovipositing on the ground near the cocoon from which she emerged (B).
Egg mass partially depredated by foraging birds; note small tears in the
cocoon where eggs were removed (C). Egg mass (arrow) on a lichen-
covered rock (D).

124

J. Res. Lepid.

arise. Predominance of black and yellow hairs in separate tufts, as noted in
the second instar, more pronounced.

Fourth instar HCW = 1.85 ±0.10 mm (n = 46). All hairs brown except the
following. Black hairs arising from mesal portions of addorsal and subdor-
sal verrucae on abdominal segments 1, 2, and 8 forming tufts much denser
and somewhat longer than surrounding dorsal hairs. Yellow hairs arising
from addorsal verrucae on abdominal segments 3 and 4 forming tufts denser
but not longer than surrounding dorsal hairs.

Fifth instar HCW = 2.35 ± 0.15 mm (n = 230); sixth instar HCW = 3.10 ±
0.20 mm (n = 353). Hairs longer and denser than fourth instar, most nota-
bly hairs arising from supraspiracular, subspiracular, and subventral verru-
cae, and dorsal verrucae on abdominal segment 9. Some hairs arising from
dorsal verrucae on prothorax and from subdorsal verrucae on mesothorax
and metathorax as long as hairs arising from supraspiracular verrucae.
Lengths of black and yellow dorsal tufts somewhat variable, sometimes nearly
even and sometimes with black tufts distinctly longer than yellow. Black tuft
on abdominal segment 8 longer and more slender than those on abdomi-
nal segments 1 and 2, resembling more the rudimentary hair pencil that it
represents (Plate 2A).

Seventh instar HCW = 3.95 ± 0.20 mm (n = 235). Color pattern of dorsal
hairtufts on abdominal segments 1 through 5 somewhat variable. Typically,
on abdominal segments 1 through 4, hairs arising from addorsal verrucae
black and those arising from subdorsal verrucae black mesally and yellow
laterally; occasionally this pattern developed to a lesser extent also on ab-
dominal segment 5. This produces an overall appearance of four, or occa-
sionally hve, central black tufts fringed laterally with yellow (Plate 2B).
Rarely, the pattern of two black tufts on abdominal segments 1 and 2, fol-
lowed by two yellow tufts on abdominal segments 3 and 4, retained in this
hnal instar.

With the exception of the distinctive black and yellow tufts, the larval hairs
of G. groenlandica show considerable variation in overall color, depending
on how recently an individual has moulted. Freshly moulted larvae appear
silvery brown overall but the brown hairs quickly darken and then very gradu-
ally fade to golden yellow (Plate 2C) during the course of the stadium.

G. rossii: First instar HCW = 0.60 ± 0.05 mm (n = 44). Corresponding to
the general description above. Cuticle between verrucae pale. Hairs uni-
formly grey in color. All hairs spinulose.

Second instar HCW = 0.85 ± 0.05 mm (n = 41). Some hairs arising from
addorsal verrucae on abdominal segments 1, 2, and 8 plumose. One or two
hairs arising from supraspiracular verrucae on each abdominal segment
plumose. All other hairs spinulose. Cuticle generally somewhat paler be-
tween verrucae.

Third instar HCW = 1.25 + 0.05 mm (n = 26). Hairs more dense than in
previous instars, beginning to obscure underlying verrucae from which they
arise. Some hairs arising from addorsal verrucae on mesothorax and met-
athorax, as well as abdominal segments 1, 2, and 8, plumose. Some hairs

34:119-141, 1995(1997)

125

arising from subdorsal verrucae and most hairs arising from supraspiracular
and subspiracular verrucae on all segments except prothorax plumose.
Other hairs spinulose, either black or yellow, those arising from thoracic
verrucae and from addorsal verrucae on abdominal segments 3 through 5
predominantly yellow.

Fourth instar HCW approximately 1.75 mm. Grey plumose hairs denser
and more prominent, otherwise very similar to third instar.

Fifth instar HCW approximately 2.50 mm. Some to most hairs arising from
all verrucae plumose. Hairs arising from addorsal and subdorsal verrucae
quite uniform in length, giving a “clipped” appearance in lateral view. Hairs
arising from supraspiracular and subspiracular verrucae up to twice as long
as those arising from dorsal verrucae. Longer plumose hairs grey, shorter
spinulose hairs black or yellow, as in the third and fourth instars.

Sixth instar HCW = 3.55 ± 0.20 mm (n = 202). All hairs black except as
noted in the following. Thoracic verrucae bearing a mixture of black and
yellow hairs not forming distinct tufts. Addorsal and subdorsal verrucae on
abdominal segments 1 through 8 bearing dense tufts of relatively short hairs,
those arising from addorsal verrucae and the mesal portion of subdorsal
verrucae black, and those arising from the lateral portion of the subdorsal
verrucae yellow. This produces the appearance of a black tuft fringed later-
ally with yellow on each abdominal segment, the pattern becoming less dis-
tinct caudally. Variable numbers of longer grey plumose hairs arising from
all verrucae, usually obscuring the pattern of black and yellow tufts to some
extent, sometimes completely, and giving the impression of lint accumu-
lated among the larval hairs (Plate 2D). In rare individuals, grey plumose
hairs replaced by black spinulose hairs which do not obscure the pattern of
black and yellow tufts (Plate 2E). Rearing of such larvae produced either
adults of G. rossii or adults of the tachinid parasitoid Chetogena gelida
(Coquillett) , which is extremely host-specific to larvae of G. rossii, at least at
this site (Morewood, unpub. data).

In general, larvae of G. rossii smaller than larvae of G. groenlandica and
with much shorter hairs of more uniform length. Pattern of black and yel-
low dorsal hairtufts quite different in the two species and not obscured by
other hairs in G. groenlandica but usually obscured at least partially by grey
plumose hairs in G. rossii. Long spinulose (Plate 3A) or plumose (Plate 3B)
larval hairs characteristic of G. groenlandica and G. rossii, respectively, pro-
ducing a contrast in overall appearance and also quite distinct when viewed
under magnification. These hairs also readily distinguished after they have
been incorporated into cocoons (Plate 3C&:D).

Cocoons. Cocoons spun on the surface of the tundra and anchored to
the substrate, not concealed in any way but rather located in exposed sites
with maximum insolation, on substrates of vegetation, litter, bare soil, or
rock. Cocoons of G. groenlandica much larger than those of G. rossii (Plate
3E), mainly due to the difference in structure (see below).

G. groenlandica: Cocoons constructed in two distinct layers with a consid-

126

J. Res. Lepid.

Plate 2. Gynaephora groenlandica (A-C) and Gynaephora rossii (D-E). Fifth instar
larva (head to the right) with the characteristic black and yellow dorsal hairtufts
and rudimentary dorsal posterior hair pencil (A). Seventh instar larva (head
to the left) with the four black dorsal hairtufts typical of the final instar (B).
Larvae showing the range of color of larval hairs with the most recently
moulted larva on the left (C). Typical larva, showing grey tufting produced
by the plumose larval hairs (D). Larva lacking grey plumose hairs (E).

34:119-141, 1995(1997)

127

Plate 3. Gynaephora groenlandica (A, C, E, F) and Gynaephora rossii (B, D, E).
Spinulose larval hairs (A). Plumose larval hairs (B). Portions of the outer
(right) and inner (left) layers of the pupal cocoon (C). A portion of the
pupal cocoon (D). Complete cocoons of G. groenlandica (left) and G.
rossii (right) (E). Larval hibernacula; the opening in the occupied hiber-
naculum was the result of removing an overlying rock (F).

128

J. Res. Lepid.

1 cm

Fig. 1 . Pupae of Gynaephora groenlandica (left) and Gynaephora rossii (right)
in ventral view. Abbreviations: a = antenna, ab = abdominal segment,
or = cremaster, cs = cremastral setae, cx = coxa of the prothoracic leg,
11 = prothoracic leg, I2 = mesothoracic leg, I3 = metathoracic leg, lb =
labrum. Ip = labial palp, mx = maxilla.

erable air space between the layers. Outer layer ovoid, dimensions 32 ± 3
mm in length by 19 ± 2 mm in width (n = 279) , comprised of a thin layer of
silk with some larval hairs, cream colored to deep yellow or grey, depend-
ing on the number and relative proportions of black and yellow larval hairs
incorporated and the extent of weathering. Inner layer oblong-ovoid, di-
mensions 28 ± 3 mm in length by 13 ± 1 mm in width (n = 279), comprised
mainly of larval hairs tied together with silk and correspondingly deeper in
color than the outer layer. Cocoons of females, with outer layer dimensions
of 34 ± 3 mm by 20 + 2 mm and inner layer dimensions of 30 ± 2 mm by 14
± 1 mm (n = 124), significantly larger “ 6.463 for outer length, 3.576
for outer width, 12.970 for inner length, 9.770 for inner width; P < 0.0005
in all cases) than those of males, with outer layer dimensions of 31 ±3 mm
by 19 + 2 mm and inner layer dimensions of 26 ± 2 mm by 13 ± 1 mm (n =
155).

G. rossii: Cocoons constructed in a single layer roughly equivalent to the
inner cocoon of G. groenlandica, oblong-ovoid, dimensions 26 + 2 mm in
length by 13 + 1 mm in width (n = 56), comprised of a single layer of silk
with many larval hairs incorporated, dark grey to light grey, depending on
the extent of weathering. Cocoons of females, with dimensions of 27 ± 2
mm by 13 ± 1 mm (n = 17), significantly larger (t^^^^^ = 2.852, 0.0025 < P <
0.005 for length; = 2.143, 0.01 < P < 0.025 for width) than those of males,
with dimensions of 25 ± 2 mm by 12 ± 1 mm (n = 39).

34:119-141, 1995(1997)

129

Pupae* Pupae of both species (Fig. 1) reddish-brown, darkening to black
as the pharate adult matures but often retaining some areas of reddish-
brown cuticle, most notably along caudal margins of abdominal segments.
Very hairy; hairs arising from scars of larval verrucae, brown to golden yel-
low and always simple, not plumose or spinulose. Dorsal hairs long, dense,
and erect; ventral hairs much shorter, sparser, and recumbent. Labrum trap-
ezoidal with rounded corners, caudal margin varying from straight to
strongly concave. Maxillae short, slightly longer than labial palps; coxae of
pro thoracic legs distinctly visible caudad of maxillae. Pro thoracic legs (ex-
cluding coxae) border on each other for about as long as length of maxil-
lae. Antennae short, extending only about halfway to caudal margin of wings.
Wingtips separated by ends of metathoracic legs. Ventral surface of abdomi-
nal segments 9 and 10 tapering steeply towards cremaster. Cremaster short
and conical, somewhat flattened dorsoventrally, apex rounded and bear-
ing a group of short, hooked setae.

G. groenlandicai Pupae with dorsal hairs up to 4 mm in length and ventral
hairs up to 2 mm in length. Hairs usually absent from ventral surface of
abdominal segment 9 and always absent from ventral surface of abdominal
segment 10. Maxillae usually curving mesad and often meeting beyond ends
of labial palps. Ventral surface of cremaster with fine longitudinal grooves
in females, less apparent in males. Female pupae, with dimensions of 24.0
± 2.0 mm in length by 9.5 + 0.5 mm in width (n = 3), significantly larger
(t(^^4 - 3.255, 0.01 < P < 0.025 for length; t^^^^ = 7.071, 0.001 < P < 0.0025 for
width) than male pupae, with dimensions of 19.5 + 1.0 mm in length by 7.5
± 0.5 mm in width (n = 3) .

G. rossiii Pupae with dorsal hairs up to 3 mm in length and ventral hairs
up to 1.5 mm in length. Hairs always present on ventral surface of abdomi-
nal segment 9 and usually present on ventral surface of abdominal segment
10. Maxillae roughly straight or slightly curved mesad but never meeting
beyond ends of labial palps. Ventral surface of cremaster smooth. Female
pupae, with dimensions of 19.0 ±1.0 mm in length by 8.0 ±0.5 mm in width
(n = 2), larger than male pupae, with dimensions of 17.0 ±1,0 mm in length
by 7.0 ± 0.5 mm in width (n = 3), the difference statistically significant for
length (t(^^3 = 2.402, 0.025 < P < 0.05) but not for width (t^^^3 = 2.049, 0.05 <
P < 0.10), probably due, at least in part, to the small sample size.

Pupae of G. groenlandica generally larger than those of G. rossii, the differ-
ence being more pronounced for females = 3.349, 0.01 < P < 0.025 for
length; t^j^3 = 5.563, 0.005 < P < 0.01 for width) than for males (t^^^^ = 3.545,
0.01 < P < 0.025 for length; = 2.121, 0.05 < P < 0.10 for width). Consid-
erable variation was seen among individuals in exact shapes and relative
dimensions of morphological features, even in the small number of pupae
examined in detail. Therefore, differences between species, as described
above, were limited to those most consistent and clearly visible; nonethe-
less, these differences should be regarded with caution.

Differences between G. groenlandica and G. rossii in the immature stages
are outlined in Table 1. Voucher specimens, including eggs, most larval

130

/. Res. Lepid.

Table 1 . Morphological differences between high arctic Gynaephora species in the
immature stages. For measurements, the full range found in this study is given.

Stage

Morphological feature

G. groenlandica

G. rossii

Eggs

Diameter (mm)

1.55-1.70

1.35-1.45

Height (mm)

1.30-1.40

1.05-1.15

Larvae

Head capsule width (mm)

First instar

0.60-0.80

0.55-0.65

Second instar

0.90-1.05

0.75-0.90

Third instar

1.15-1.40

1.20-1.35

Fourth instar

1.50-2.00

ca. 1.75

Fifth instar

2.00-2.70

ca. 2.50

Sixth instar

2.70-3.60

2.90-4.15

Seventh instar

3.35-4.45

N/A*

Cuticle between verrucae

First instar

black

pale

Second instar

black

paler than verrucae

All subsequent instars

black

Form of larval hairs

First instar

all spinulose

Second instar

all spinulose

some plumose

All subsequent instars

all spinulose

many plumose

Color of larval hairs

First instar

black and brown

uniformly grey

All subsequent instars

varying shades of brown

longer plumose

with distinct dorsal tufts

hairs grey, shorter hairs

of black and yellow

black and yellow

Cocoons

Color

cream to deep yellow,
occasionally grey

grey

Outer layer

length (mm)

25-40

21-30

width (mm)

14-26

11-16

Inner layer

length (mm)

19-35

N/A^

width (mm)

10-17

N/A^

Pupae

Length (mm)

19.0-26.0

16.0-19.5

Width (mm)

7.5-9.5

7.0-8.0

^N/A = not applicable; these stages or structures do not occur in G. rossii

instars, pupae, cocoons, and adults of both species, have been submitted to
the Canadian National Collection of Insects, Ottawa, Ontario.

Natural History

Both species spin cocoons and pupate, adults emerge, mate, and lay eggs,
and eggs hatch all within a single summer season lasting little more than
two months; however, larval development is spread over a number of years
with larvae overwintering in each stadium (Kukal 8c Kevan 1987; Morewood
& Ring, submitted).

34:119=141, 1995(1997)

131

Fully grown larvae begin spinning cocoons very soon after becoming ac-
tive in the spring. An exceptionally heavy snow accumulation in east-cen-
tral Ellesmere Island during the winter of 1994-95, combined with a rela-
tive lack of wind, left the Alexandra Fiord lowland covered by a near-com-
plete blanket of snow at the end of May. Judging by the extremely limited
extent of snow-free ground and the subsequent rate of snowmelt, larvae
found active upon our arrival 28.V.1995 could not have been active for more
than a few days prior; however, some of these larvae had begun spinning
cocoons as early as 29. V. 1995. Cocoons may be completed within one day
or may require two or three days for completion. Similarly, pupation may
occur within one day of the cocoon being completed or may be delayed for
two or three days. Pupal development, from pupation to adult emergence,
of G. groenlandica required 15 + 5 days (n = 53) in the field in 1995; only
two G. rossii could be monitored for the complete period of pupal develop-
ment and these required 10 and 16 days. The variation in time required
for these developmental stages is due, at least in part, to variations in weather
conditions, with cool and/or cloudy weather retarding activity and devel-
opment.

Adults of both sexes have fully developed wings and males are strong fli-
ers; however, females fly very little and when they do, scarcely get off the
ground. Normally a female remains on her cocoon until she attracts a male
and, once mated, will often lay a mass of eggs there (Plate lA). Additional
eggs are laid nearby on vegetation or on the ground, with no apparent dis-
crimination among potential oviposition sites, and some females leave their
cocoons even before laying their initial egg masses (Plate IB). Of nine ini-
tial egg masses laid in the field in 1996, four were laid on cocoons whereas
five were not. Eggs laid on cocoons are very conspicuous and suffer heavy
predation by birds (Plate 1C), primarily snow buntings {Plectrophenax nivalis
Linnaeus) , by far the most abundant breeding birds at Alexandra Fiord
(Freedman 1994) . Of 39 egg masses found on cocoons during the summer
of 1994, 26 showed signs of predation and a further 11 were completely
removed before they could be protected with netting; only two egg masses
were protected before apparently suffering any predation. In contrast, egg
masses laid on the ground are quite cryptic (Plate ID) and none of these
egg masses were found to suffer any predation.

Embryonic development, as measured from the day that an initial egg
mass was laid to the day that the first larvae eclosed, for G. groenlandica was
28 ± 5 days (n = 10) in the field in 1995; for G. rossii only one female was
observed to lay an egg mass in the field and this required 31 days to begin
hatching. Upon hatching, neonates usually eat a portion, often most but
rarely all, of the chorion from which they emerged.

With the exception of neonates, larvae of G. groenlandica are active for
only a relatively short portion of the growing season, after which they spin
hibernacula and become dormant until the following spring. Regular sur-
veys, combined with incidental observations, indicated that the bulk of the
larval population was active only until the third week of June in 1994, the

132

J. Res. Lepid.

fourth week of June in 1995, and the second week of July in 1996 due to a
very late and prolonged snowmelt. Very few G. groenlandica larvae were found
active on the tundra after 1.V11.94, 15.V1L95, and 19.V1L96, and none were
found after 1.V1IL94, 31. VII. 95, and 4. VIII. 96. In contrast, larvae of G. rossii
remain active late in the growing season, with active larvae observed regu-
larly on the tundra until and including 15.VIII.94, 17.VIII.95, and 13.VIII.96,
our last days of fieldwork each year. In all three years, with the exception of
fully grown larvae that were spinning cocoons in June, more G. ro55w larvae
were found active in August than in June and July combined.

Gynaephora larvae were observed feeding on 1 1 different species of plants,
representing seven different plant families (Table 2). For G. groenlandica,
Salix arctica represented 87% of the feeding observations, most of these
being buds and expanding leaves, with Dry as integrifolia representing 7%,
Saxifraga oppositifolia representing 3%, and the remainder represented by
single or very few observations. The few feeding observations for G rossii
were almost evenly split between S. arctica and D. integrifolia, with a single
observation of a larva feeding on developing fruits of Cassiope tetragona on
the tundra (Table 2).

Hibernacula of Gynaephora larvae are spun with silk, much like pupal co-
coons except that no larval hairs are incorporated and the structure con-
sists of a single layer in both species. Larvae that are confined within enclo-
sures on the tundra generally spin hibernacula in clumps of vegetation or
in litter and incorporate litter into the structure, making it extremely cryp-
tic. Such hibernacula are rarely found on the open tundra, probably due
to their cryptic nature; however, hibernacula are commonly found beneath
or between loosely piled rocks (Plate 3F) .

Larvae and cocoons of Gynaephora generally may be handled with impu-
nity; however, the larval hairs can cause skin irritation. Extensive work dis-
secting parasitoid-killed larvae or tearing open cocoons, which contain lar-
val hairs, resulted in small (1-2 mm diameter) itchy blisters, particularly
on the sensitive skin between the fingers, and these blisters persisted for
many days.

Discussion

Identification of Immature Stages

Confusion concerning identification of immature stages of North Ameri-
can Gynaephora species dates back to the original description of G. rossii.
Curtis (1835) described an adult male in some detail and provided a color
illustration that leaves no doubt that the species was G. rossii. In contrast,
his descriptions of immature stages were rather cursory; however, the “two
tufts of black hair on the back [of the caterpillar], followed by two of or-
ange” are unmistakably those of G. groenlandica. His description of the co-
coons is unfortunately too generalized to assign to either species. The origi-
nal description of G. groenlandica, on the other hand, includes a mention
of “the characteristic Zlasyc/^fra-caterpillar hairtufts on the back and the end
segmenf (emphasis added) typical of the larvae of this species (Homeyer

34:119-141, 1995(1997)

133

Table 2. Plants on which Gynaephora larvae were observed feeding at Alexandra
Fiord, Ellesmere Island, during the spring and summer of 1995 and 1996.

Plant species

Number of observations

Part eaten

G. gromlandica

G. rossii

Salix arctica Pallas (Salicaceae)

Buds (unopened)

Expanding leaves

166

Developing catkins

Mature leaves

Senescent leaves

Dry as integrifolia M. Vahl (Rosaceae)

Leaves

Flower petals

Saxifraga oppositifolia Linnaeus (Saxifragaceae)

Flowers

Leaves

Oxyria digyna (Linnaeus) Hill (Polygonaceae)

Leaves 1 0

Arctagrostis latifolia (R. Brown) Grisebach (Gramineae)

Leaves

Festuca brachyphylla Schultes (Gramineae)

New shoots

Luzula confusa Lindeberg (Juncaceae)

Leaves

Flower head

Luzula arctica Blytt (Juncaceae)

Leaves

Flower stalk

Potentilla hyparctica Make (Rosaceae)

Flower

Vaccinium uliginosum Linnaeus (Ericaceae)

Leaves

Cassiope tetragona (Linnaeus) D. Don (Ericaceae)

Developing fruits^

Total number of observations

366

'Developing fruits were also accepted as food by G. rossii larvae held

in the laboratory;

foliage and mature fruits were not.

1874). Packard (1877) described all stages of what he thought was G. rossii,

based on specimens collected in northern Greenland.

These descriptions

are fairly accurate for G. groenlandica and Packard himself noted that the

adults differed from the description of G.

rossii given by Curtis (1835) in

that their hind wings had no “broad, blackish margin.

which is perhaps

the most obvious difference between adults of G. groenlandica and G. rossii

(cf. Plate 1 in Ferguson 1978). The brief descriptions published by Scudder
et al. (1879) as representing G. rossii are inadequate for identification of
the species; however, they did note that the original “description of the larva
does not well accord with the present specimen.” It may be that neither
Packard (1877) nor Scudder et al. (1879) knew of G. groenlandica, consid-
ering that the description of this species was published in 1874 in Germany
and therefore may not have been available to them.

134

J. Res. Lepid.

As early as 1875, G. rossii\i2id been found above treeline on Mount Wash-
ington, New Hampshire, and recognized as the same species as had been
described from the Arctic (Grote 1876). Later, Dyar (1896) described lar-
vae from the same locality and noted that they differed from the descrip-
tions published by both Curtis (1835) and Packard (1877). The following
year, he received larvae from Greenland that agreed with Curtis’ descrip-
tion, obtained an adult G. groenlandica from one of them, and concluded
that “Curtis must have mixed the species” (Dyar 1897).

Despite Dyar’s conclusion and his fairly detailed descriptions of the lar-
vae of G. rossii (Dyar 1896) and G. groenlandica (Dyar 1897), misidentifica-
tions and confusing information may be found in much more recent pub-
lished literature, as noted in the introduction to this paper. In addition,
Ryan (1977) and Ryan and Hergert (1977) presented a photograph of a
number of specimens from Truelove Lowland, Devon Island, that included
both species of Gynaephora, but the adults were not shown associated with
their cocoons. Both “light and dark color cocoons” were illustrated and Ryan
and Hergert (1977) stated that “both forms [were] found with each spe-
cies”; however, they made no mention of the structure of the cocoons and
submitted only a single specimen (a G. groenlandica female with the cocoon
from which it emerged) to the Canadian National Collection of Insects. As
described above, cocoons of both species may be light or dark in color,
depending on the extent to which larval hairs of different colors are incor-
porated into the cocoon and the extent to which the cocoons are weath-
ered, but the structure of the cocoon is species-specihc. Descriptions of lar-
vae provided by Ferguson (1978) are accurate, even though they were based
on extremely limited material; however, they may give the impression that
the differences between the two species are rather subtle when in fact these
differences produce a distinctive appearance for each species that is dis-
cernible even from a distance.

Pupae of G. groenlandica and G. rossii have not been described previously,
but both species may be identihed to genus using the key to genera pro-
vided by PatoCka (1991) . They also fit the generic description of Gynaephora
pupae except that their antennae are apparently much shorter than those
of the European species Gynaephora selenitica (Esper), as described and il-
lustrated by PatoCka (1991) . The diagrams presented here (Eig. 1) are com-
posites that attempt to illustrate “typical” pupae for both sexes of both North
American species; however, a considerable amount of individual variation
was seen, even among the small number of pupae examined. The only dif-
ferences between species that were obvious and consistent were overall size
and the length of hairs (which may be related to overall size), the presence
or absence of hairs on the ventral surface of abdominal segments 9 and 10,
and possibly the form (curved or relatively straight) of the maxillae.

It should be noted that the size differences between the two species may
not be consistent across their entire range. In fact, the adults illustrated by
Ferguson (1978) clearly show that G. ro55Mmaybe larger than G. groenlandica
from different localities. The fact that G. ro55ziwere found to be consistently

34:119-141, 1995(1997)

135

smaller than G. groenlandica in the current study may reflect the fact that
this population of G. rossii is in the extreme northern portion of the spe-
cies’ range whereas Alexandra Fiord is more central in the distribution of
G. groenlandica.

Despite the confusion that is apparent in the literature, most of the im-
mature stages of arctic Gynaephora species can be identified to species quite
readily and with little more than a cursory examination. The occasional lack
of grey plumose hairs in G. rossii larvae may cause some confusion and may
be responsible for a report of “morphs intermediate between the two . . .
species” (Kukal 1994), although the supposed intermediate morphs were
not described in that report. The species may be reliably separated by dif-
ferences in the patterns of black and yellow hairtufts and the much longer
overall hairs of G. groenlandica. Furthermore, there is strong evidence that
they are reproductively isolated at the level of mate recognition and there-
fore do not produce hybrids (Morewood, submitted). We hope that the
descriptions and illustrations provided here will help to prevent future
misidentifications.

Natural History

Gynaephora species are among the most conspicuous insects on the high
arctic tundra and observations on their natural history have been recorded
ever since the early arctic expeditions of European explorers. The first com-
prehensive study of G. groenlandica conducted by Kukal (1984) and later
published by Kukal and Kevan (1987). That study provided a significant
advance in knowledge of the natural history of this species; however, it did
contain some gaps and inaccuracies due, in part, to the fact that it was con-
ducted during a single summer season (see also Morewood & Ring, sub-
mitted).

Kukal and Kevan (1987) identified mortality factors and estimated mor-
tality rates for most of the life stages of G. groenlandica; however, the only
mortality factor they identified for eggs was “inviability.” With respect to
eggs, their study included only “six females observed in nature [which] re-
mained on their cocoons and deposited all of their eggs there” and they
concluded that the “eggs hatched within several days of their deposition”
without presenting any relevant data (Kukal & Kevan 1987). They appar-
ently found no other eggs masses in the field and this may be due to the
facts that eggs are often laid after the female has left her cocoon, such eggs
are extremely cryptic, and egg masses laid on cocoons are extremely vul-
nerable to predation by birds. Egg masses on cocoons are likely to be re-
moved before they are found and, considering the rate of predation re-
corded in 1994, it seems likely that very few eggs laid on cocoons would
escape predation long enough to hatch.

It has been known for some time that larvae of G. groenlandica limit their
activity to the early part of the growing season (Kukal & Kevan 1987). In
contrast, the fact that larvae of G. rossii are active late in the growing season
has not been reported previously from the Arctic, although Schaeffer and

136

J. Res. Lepid.

Castrovillo (1979) reported larvae of G. rossii to be active and feeding in
September on both Mt. Katahdin, Maine, and Mt. Daisetsu, Japan. This
contrast in seasonal activity may have significant consequences for the re-
spective life cycles of the two species and there are indications that it is con-
sistent across the Canadian Arctic. We collected Gynaephora larvae in the
vicinity of the Muskox River on north-central Banks Island in early August
of 1993 and this collection consisted of approximately two dozen larvae of
G. rossii but only a single larva of G. groenlandica. In addition, researchers
working on the Fosheim Peninsula of west-central Ellesmere Island in 1996
observed larvae of G. groenlandica in abundance in late June and early July
but larvae of G. rossii only in early August (A. Lewkowicz, Department of
Geography, University of Ottawa, pers. comm.).

Larvae of Gynaephora are clearly opportunistic feeders, accepting a wide
variety of plant species as food, but do show definite preferences in their
choice of foodplants. Curtis (1835) originally reported that larvae of G
groenlandica (reported as G rossii) fed mostly on Saxifraga tricuspidataRottholl
and S. oppositifolia, but the preference of this species for Salix has since been
noted repeatedly (Wolff 1964, Kukal & Kevan 1987, Kukal & Dawson 1989) .
The relatively few feeding observations for G rossii in this study probably
underestimate the variety of plants that these larvae actually eat, even at
Alexandra Fiord. This widely distributed species has been reported to feed
on many different plants, ranging from sedges to broaddeaf trees (Schaefer
& Castrovillo 1979 and references cited therein) and it has been suggested
that some isolated alpine populations show preferences for ericaceous
plants, which predominate in alpine habitats (Schaefer & Castrovillo 1979).

One of the hypotheses proposed to explain why larvae of G groenlandica
cease feeding and become dormant so early in the growing season is that
they restrict their feeding activity to the early portion of the season when
the available food has the greatest nutritional value and become dormant
when the benefits of continued feeding on foodplants of declining quality
are outweighed by the metabolic costs of remaining active (Kukal & Dawson
1989). This hypothesis is supported by observations that larvae of G
groenlandica feed primarily on buds, expanding leaves, and developing cat-
kins of S. arctica (Kukal & Dawson 1989; this study), a food source that rap-
idly declines in nutritional value as the leaves and catkins mature (Kukal &
Dawson 1989, Dawson & Bliss 1993, Klein & Bay 1994) . This may be consid-
ered an adaptation of this species for making the most efficient use of avail-
able food sources, given the constraints of the high arctic environment to
which it is endemic. In contrast, larvae of G rossii remain active late in the
growing season and appear to be less particular about seeking out food
sources of maximal nutritional value; however, the fact that G rossii larvae
consumed developing fruits, but not foliage or mature fruits, of G tetragona
suggests a similar selection of optimal food sources available later in the
summer.

The distinct double-layered structure of the cocoons of G groenlandica
may again represent an adaptation to its high arctic environment, allowing

34:119-141, 1995(1997)

137

the crucial life stages of pupation and reproduction to be completed within
the very short growing season. These cocoons are thought to act as
“microgreenhouses” and temperatures within them have been shown to be
higher than both ambient temperatures and surrounding substrate tempera-
tures (Kevan et al. 1982, Kukal 1984). Furthermore, it has been recently
reported that cocoons of G. groenlandica significantly enhance the rate of
pupal development but that those of G. rossii do not (Lyon & Cartar 1996) .
The similar pupal development times in both species found at Alexandra
Fiord might be accounted for by the difference in size of pupae of the two
species at this site. Without a development-enhancing cocoon like that of
G. groenlandica, a decrease in size of G. rossii in the northern portion of its
range may be necessary for this species to complete pupal development
quickly enough to reproduce and still leave time for the resulting eggs to
hatch before winter closes in.

In a recently published study of hibernacula and winter mortality, Kukal
(1995) apparently contradicts her previous assertion (Kukal, Serianni &:
Duman 1988, Kukal & Dawson 1989, Kukal 1990, 1991, 1993, Banks et al.
1994) that larvae of G. groenlandica move down close to the permafrost when
they become dormant in early summer. It is noteworthy that within at least
some of the cages used for that study, there were deep crevices in the tun-
dra but the larvae chose to remain on the surface and construct their hi-
bernacula in the vegetation and litter. The significance of this is that, al-
though it may be argued that G. groenlandica larvae undergo “voluntary hy-
pothermia” by virtue of the fact that they no longer thermoregulate by bask-
ing (cf. Kukal, Heinrich & Duman 1988), temperatures within such hiber-
nacula track ambient temperatures fairly closely (Kukal 1995) . Ground-level
temperatures, both ambient and within hibernacula, often exceed 20°C and
even approach 30°C during sunny weather (Morewood, unpub. data). Me-
tabolism of poikilothermic organisms in general is directly related to tem-
perature and this has been shown experimentally for larvae of G. groenlandica
(Kukal & Dawson 1989). The hypothesis that larvae of G. groenlandica mo\G
close to the permafrost where “the larval body temperatures range between
0-5°C” (Kukal 1990) and thus reduce maintenance metabolism and con-
serve energy reserves during their summer dormancy (Kukal 1990, 1991,
1993, Banks et al. 1994) must be re-evaluated in the light of more recent
discoveries regarding the location of, and temperature conditions in, lar-
val hibernacula.

Finally, the urticating nature of the larval hairs of Gynaephora has not been
reported in previously published literature but has been experienced by
other fieldworkers and may be much more severe than the small itchy blis-
ters recorded in this study. Reactions experienced by other researchers
working with Gynaephora in the field include large blisters covering most of
the hands and extensive swelling and itching of the hands (B. Lyon, De-
partment of Biological Sciences, University of Calgary, pers. comm.). It is
not known whether there is any chemical basis for these urticating proper-
ties and it may be that the irritation is a simply mechanical effect of the

138

/. Res. Lepid.

barbed hairs, as has been reported for the similar, although not closely-re-
lated, larvae of Lophocampa argentata (Packard) (Arctiidae) (Silver 1958). As
noted above, the severity of reported reactions to Gynaephora vdivies widely
among different individuals and therefore researchers who plan extensive
work involving exposure to the larval hairs would be well-advised to exer-
cise caution.

Acknoiuledgements. Thanks to J.D. Lafontaine for facilitating access to specimens in
the Canadian National Collection of Insects in Ottawa. The research of which this
represents a small part has been supported financially by a Postgraduate Scholar-
ship from the Natural Sciences and Engineering Research Council of Canada
(NSERC), an Eco-Research Doctoral Eellowship funded by Canada’s Green Plan,
and the Northern Studies Training Program of Canada’s Department of Indian
Affairs and Northern Development. Additional financial support was provided by
NSERC through an Operating Grant to R.A. Ring at the University of Victoria. Ex-
cellent logistic support was provided by the Polar Continental Shelf Project of Natu-
ral Resources Canada, through grants to R.A. Ring and to G.H.R. Henry at the
University of British Columbia. Special thanks to G.H.R. Henry for the opportu-
nity to join his field camp and to the Royal Canadian Mounted Police for allowing
us the use of their buildings at Alexandra Eiord.

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Journal of Research on the Lepidoptera

34:142-146, 1995(1997)

Notes on Boloria pales yangi^ ssp. nov., a remarkable
disjunction in butterfly biogeography (Lepidoptera:
Nymphalidae)

Yu-Feng Hsu^ and Shen-Horn Yen^

'Division of Environmental Biology, Department of Environmental Science, Policy and
Management, University of California, Berkeley, CA; Current address: Department of Biology,
National Changhua University of Education, Changhua 50058, Taiwan, R.O.C.

^Laboratory of Natural Resource Conservation, Department of Biology and Institute of Life
Science, National Sun Yat-Sen University, Kaohsiung, Taiwan, R.O.C.

Abstract. A population of Boloria pales (Denis & Schiffermuller) was found
at an alpine area in Taiwan, far away from the nearest population in west-
ern China and farther south than any previous record of the genus Boloria.

The taxon is considered a new subspecies and is described herein.

Keywords. Boloria pales, disjunction, biogeography, Taiwan, China

Introduction

The genus Boloria (sensu Scott 1986, D’Abrera 1992) is composed of ap-
proximately 30 species of small nymphalids favoring either damp and wet
habitats or rocky slopes (Shepard 1975), distributed in boreal and arctic
parts of the Holarctic region. The recognized southernmost limit of this
genus was 30° N for B. pales (Denis & Schiffermuller) in the Palaearctic
(BMNH specimens) and 35° N for B. chariclea (Esper) in the Nearctic (Scott
1986).

Two male specimens of this genus were collected by Prof. C.T. Yang from
an alpine area in the central part of Taiwan at approximately 24° N. These
specimens represent the brst record of Boloria outside the Holarctic, and
are herein classibed as a subspecies of B. pales (Denis 8c Schiffermuller).
This discovery is significant for three reasons: 1) This is unquestionably the
southernmost record for the genus and the first record of Boloria, a typical
Holarctic genus, in the Oriental region. 2) The closest B. pales colony to
Taiwan is found in the Sichuan province of western China, about 2000 km
distant (Fig. 1). This large disjunction suggests that the Taiwan population
is a relict left from a glacial period of the Pleistocene. 3) It is striking that
such a unique species has been found in a well-collected island.

The specimens from Taiwan most closely resemble B. pales palina
Fruhstorfer in western China, but has distinctive characters on wing pat-
terns and male genitalia. We describe these specimens as a new subspecies
here.

Paper submitted 7 June 1995; revised manuscript accepted 25 January 1996.

34:142-146, 1995(1997)

143

Fig. 1 . Known geographical distribution of Boloria pales in East Asia and neigh-
boring areas; squares denote confirmed sites from specimens examined
from BMNH, NCU, and NTU; circles represent province records based
on BMNH specimens.

Boloria pales subspecies yangiHsu &Yen, ssp. nov. (Figs. 2, 3, 12-15)

Male. Forewing length 18.0 mm (n = 2). Head: hairy, covered with buff
orange hairs on vertex and frons. Eyes semi-oval, naked. Labial palpus hairy,
porrect, pointed, yellow proximally, buff orange distally. Thorax: black,
covered with pale buff orange hairs. Legs hairy, buff orange. Forewing:
termen slightly concave. 12 veins, R1 independent, other R veins all extend-
ing from Rs. M2 slightly bent toward Ml; base of M3 strongly curved poste-
riorly. Upperside color pale buff orange with pale black markings. Six black,
round postdiscal spots with posterior three indented. Submarginal spot
black, prominent, fused with postdiscal spot series anteriorly. Marginal
markings obscure. Discal spots prominent, wider than postdiscal spots. Black
markings in discoidal cell composed of a distal bar, a medial bar, and two
proximal dots. Basal area covered by extensive pale black scaling. Under-
side pale buff with pale black markings. Cinnamon-rufous bar edged by
creamy yellow patches present near apex. Fringe uniformly orange.
Hindwing: nine veins, all separate. M3 bent near base. Humeral vein a short
bar, perpendicular to Sc + Rl. Upperside pale buff orange with pale black
markings. Six round postdiscal spots arranged into an arch. Submarginal
spot series prominent, triangular. Marginal markings more prominent than
forewing. Discal spots narrow. Basal area with extensive pale black scaling
extending distally covering distal end of discoidal cell and discal band in

144

J. Res. Lepid.

Figs. 2-1 1 . Boloria specimens. 2-9: Boloria pales from various geographical re-
gions. 2, 3) Taiwan; 4, 5) Sichuan, western China; 6, 7) Kazakhstan;
8, 9) Southwestern France. 10, 11) Boloria napaea from Switzerland.

34:142-146, 1995(1997)

145

Figs. 12=18. Male genitalia of 8o/or/a specimens. 12-15: Boloria pa/es yang/ Hsu
& Yen. 12) Tegumen + valvae, dorsal view; 13) Phallus, lateral view; 14)
Juxta, posterior view; 15) Left valva. 16-18: Left valva of Boloria pales
from various geographical regions. 16) Sichuan, western China; 17)
Kazakhstan; 18) southwestern France. “Amp” denotes ampulla.

Cu cells. Underside coloration variegated. Basal area cinnamon-rufous sur-
rounded by three silvery white lunules. Single prominent silvery white dot
present at base of cell Cu2; another small silvery white dot in discoidal cell.
Discal area forming a tawny band unevenly delimited distad by short black
lines. Yellow scalings present in anterior part of discal band. Postdiscal area
pale cinnamon-rufous with a series of amber-colored round spots. Silvery
white lunules present distally in cell Sc + Rl, Rs, and Cu2. Extensive yellow
scaling present in cell M3 and posterior edge of cell M2. Marginal spots
silvery white edged proximally by amber-colored scalings. Fringe uniformly
orange. Abdomen: Black covered with pale buff orange hairs, ventrally with
extensive pale yellow scaling toward distal end. Genitalia (Figs. 12-15): Scler-
ites of 9th + 10th segments ring-shaped with a medial triangular membra-
nous area dorsad. Uncus narrow, elongate, bifurcate distally. Saccus broad,
short. Valva broad, somewhat rectangular in shape; ampulla elongate with
minute teeth dorsad, forming downcurved arm with a wart-like dorsal pro-
cess at basal 74; harpe simple, setose, with distal end nearly straight; cucullus
forming a prominent, densely setaceous triangular tooth dorso-distally.
Phallus stout, short, with phallobase about as long as aedeagus. Distal end
of vesica forming two vertical semicircular lobes; both lobes spinulose ex-
ternally but asymmetrically. Bulbus ejaculatorius subterminal. Juxta form-
ing thin, flat lobe with deep dorsal cleft mesad.

Female. Unknown.

Diagnosis. B. pales yangi Hsu & Yen is similar to B. pales palina Fruhstofer
(Figs. 4, 5) of western China, but differs from it by the following charac-
ters: 1) discal spots broader than postdiscal spots on forewing upperside.

146

J. Res. Lepid.

2) fringe uniformly colored instead of checkered, 3) basal area of hindwing
underside without yellow scaling.

Biology. Host plant and early stages in Taiwan unknown. Larvae of popu-
lations in Europe utilize Tio/aspp. (Violaceae) (Higgins & Riley 1983). Ac-
cording to Wang and Huang (1993), 15 species are known to occur in
Taiwan with six species found in the vicinity of the type locality of B. pales
yangi.

Type data. Holotype: d, 24°15'N, 121°14'E. TAIWAN: [Taichung Hsien] ,
Lishan, 10.V.1964. Coll. C.T. Yang; paratype: Id, same data as holotype.
Both holotype and paratype deposited in the Insect Museum, National
Chung-Hsin University, Taichung, Taiwan, R.O.C. (NCU).

Comparative material was from the collections of the Natural History
Museum, London, U.K. (BMNH), Insect Museum, National Taiwan Uni-
versity, Taipei, Taiwan, R.O.C. (NTU), and NCU.

Discussion

The population of B. pales in Taiwan appears more closely related to those
in western China than to those in central Asia and Europe. Two possible
synapomorphies shared by the specimens from Taiwan and western China
are: 1) Reduced yellow scaling at basal area on hindwing underside (Eigs. 3,
5); 2) ampulla of valva narrow with the dorsal process wart-like, present at
basal y4, abruptly narrowed down toward the base (Eigs. 15, 16). The speci-
mens of B. palesirom central Asia (Eigs. 6, 7) and Europe (Figs. 8, 9), as well
as B. napaea (Hoffmannsegg) (Figs. 10, 11), the sister species of B. pales, all
have extensive basal yellow scaling on hindwing underside (Figs. 7, 9, 11)
and a robust ampulla on which a large dorsal process is present at basal ^/2
and attenuates toward the base (Figs. 17, 18; for B. napaea Higgins 1975).

Acknowledgements. We thank Chung-Tu Yang and Cheng-Tse Yang (NCU) for per-
mission to examine the material deposited in NCU. We are grateful to Philip R.
Ackery of Natural History Museum, London, Robert Alexis of Belgium, and Iv
Bereznoi of Russia for assistance on material from other regions. We also thank
Raina Takumi, University of California, Berkeley for reading the manuscript.

Literature Cited

D’Abrera, B. 1992. Butterflies of the Holarctic Region. Part ll. Satyridae (concl.) &
Nymphalidae (Partim). Hill House, Victoria.

Higgins, L.G. 1975. The Classification of European Butterflies. Collins, London.
Higgins, L.G. & N. D. Riley. 1983. A Field Guild to the Butterflies of Britain and
Europe. Collins, London.

Scott, J. 1986. The Butterflies of North America. Stanford University Press, Stanford.
Shepard, J.H. 1975. The genus Boloria. Pp. 243-252 in Howe, W.H., ed. The
Butterflies of North America. Doubleday 8c Company, New York.

Wang, J.C. & T.C. Huang. 1993. Violaceae. Pp. 807-834 in Huang, T.C., ed. Flora of
Taiwan, 2nd Edition. Editorial Committee of the Flora of Taiwan, Second
Edition. Taipei.

Journal of Research on the Lepidoptera

34:147-153, 1995(1997)

Yania gen. nov. and Yania sinica sp. nov. from Sichuan, China
(Lepidoptera: Hesperiidae)

Hao Huang

Qingdao Education College, Qingdao 266071, China

Abstract. Yania gen. nov. and Yania sinica sp. nov. (Hesperiidae) are de-
scribed from Sichuan, China. Yania can be placed in the Ancistroides group
of the Hesperiidae and can be distinguished from all the known genera
of this group by the following combination of characters: 1) dub of an-
tennae very gradually marked, 2) both wings with vein 5 nearer to vein 4
than to vein 6, 3) without secondary sexual characters, 4) forewing with
vein 2 nearer to wing-base than to vein 3, 5) male genitalia with uncus
deeply bifid, uncus longer than tegumen, and 6) the clasp a very simple
structure.

The new species described here was recognized when I sorted the but-
terflies I collected from Qingchenshan, Sichuan during the summer of
1991. Most specimens from the Qingchenshan Mountains were somewhat
damaged when captured or when spread. Consequently, the unique ho-
lotype of this new species lost its labial palpi and the antennae are bro-
ken at the tip (one broken below the apiculus). However, its wing vena-
tion, genital structure, and other features indicate that it belongs to a
new genus.

Yania Huang, new genus
Type species Yania sinica Huang

Male

Antennae. Half as long as costa; club very gradually marked, not con-
stricted before apiculus.

Body. Thin, weak; abdomen slightly longer than dorsum of hindwing.
Forewing. No prominent hyaline spots. Dorsum quite longer than termin.
Vein 2 arising before the origin of vein 1 1 and nearer to wing-base than to
the origin of vein 3. Vein 5 slightly closer to vein 4 than to vein 6 at its ori-
gin. Vein 11 originates midway between veins 10 and 12.

Hindwing. Costa slightly longer than dorsum. Discocellular cell slightly
shorter than half the length of hindwing. Vein 7 arising beyond the origin
of vein 2. Vein 5 well defined, not oblique and very slightly closer to vein 4
than to vein 6 at origin.

Secondary sexual characters. Absent.

Male genitalia. Uncus much longer than tegumen and deeply bifid.
Gnathos slightly shorter than uncus. Saccus significantly longer than

Paper submitted 22 February 1996; revised manuscript accepted 25 September 1996.

148

J. Res. Lepid.

tegumen and sharply pointed at tip. Clasp very simple in structure, without
a style from valva or harpe.

Etymology. The name Yania is a feminine noun based upon the given
name of my younger sister, Yan Huang.

Yania sinica Huang, new species (Figs. 1-6)

Male. Eyes smooth and blackish brown when dried. Frons nearly twice as
wide as eye, densely clad with black hairs mixed with some yellow.

Labial palpus. Unknown (both palpi missing from the holotype).

Antennae. 9.5 mm long (about half the length of forewing); club weakly
and gradually marked, segments becoming broader on apical Vs of anten-
nae with the thickest portion only twice as thick as shaft; club densely clad
with blackish scales as well as shaft on upperside, but with pale yellow scales
on underside in contrast with shaft; number of nudum segments in the
apiculus unknown as the apiculus is broken at the tip, the remaining nu-
dum segments all in bent-over portion of club, and club not constricted
before apiculus (Fig. 3).

Thorax. Clad with darker brown scales and scattered long yellow hairs.

Abdomen. Thin and weak, slightly longer than dorsum of hindwing,
densely clad with dark brown scales above, but with longer yellow and brown
scales beneath mixed with scattered yellow hairs.

Legs (Fig. 4). Densely covered with dark brown scales above, yellowish
scales beneath; fore and mid femora not apparently clad with hairs, hind
femora densely clad with long yellowish hairs beneath; tibial epiphysis red-
dish, wicker-leaf-shaped and somewhat distorted, nearly Vs times as wide as
fore tibiae, originating from the basal Vs of fore tibiae and surrounded with
long yellow and black scales; all tibiae without spines or hair-brushes, only
sparsely clad with few long yellow hairs; mid tibiae with terminal pair of spurs
which are densely clad with brown scales and blunt at tip, the inner one
(just on the inside of tibiae) slightly longer than the outer: hind tibiae with
two pairs of spurs, the upper pair somewhat shorter than the lower; all tarsi
clad with three rows of reddish spines below, which are as long as the scales
on tarsi, without any hairs; claws as in Astictopterus jama.

Wing markings. Ciliae of both wings on both sides dark brown, con-
colorous with ground color of wings. Upperside: Both wings unmarked,
uniform dark brown in color, without secondary sexual characters. Veins
not marked in color. Underside: Both wings ground color dark brown as
on upperside, with a yellowish cast. Some veins thinly clad with yellow scales.
Costal and apical areas of forewing and basal half of hindwing sparsely clad
with scattered yellow scales. Posterior marginal areas of both wings some-
what paler than other areas in color, otherwise as upperside.

Wing shape and wing venation (Fig. 5). Forewing. Length: 19.5 mm. Dor-
sum quite longer than termin. Vein 2 much closer to wing base than to vein
3 at its origin. Vein 5 slightly closer to wing base than to vein 3 at its origin.
Vein 5 slightly closer to vein 4 than to vein 6 at its origin. Vein 11 beyond

34:147-153, 1995(1997)

149

Figs. 1-2. Yania sinica c?holotype. 1) Left: upperside. 2) Right: underside. Scale
1 cm.

vein 2 at origin and about midway between veins 10 and 12. Hindwing. Costa
slightly longer than either termin or dorsum. Vein 5 well marked, not ob-
lique, very slightly closer to vein 4 than to vein 6 at its origin. Vein 7 midway
between veins 2 and 3 at origin.

Male genitalia (Fig. 6). Uncus nearly twice as long as tegumen, deeply
bifid in dorsal view, its two arms running parallel with each other. Gnathos
significantly longer than tegumen, nearly as long as the uncus arms. Saccus
also long and sharply pointed at tip in both dorsal and lateral views. Clasp
nearly rectangular in shape, with distal margin nearly plain and flat, only
bearing a small tooth in the middle, posterior angle well produced, with a
sharply pointed process. Juxta as Ancistroides nigrita. Aedeagus nearly as long
as clasp, without cornuti, its suprazonal portion nearly as long as subzonal
portion.

Female. Unknown.

Type data. Holotype: d (Figs. 1, 2). Qingchengshan, Sichuan, China,
1500m. 12.VII. 1991. Leg. H. Huang. Deposited in the Biological Labora-
tory of Qingdao Education College, Qingdao, Shandong Province, China.

Diagnosis and Discussion

Yania clearly belongs in the subfamily Hesperiinae by exhibiting a hind
tibia without erectile hair tufts, an abdomen without specialized scales, and
a forewing lacking a costal fold. According to W.H. Evans (1949:2-4),
Hesperiinae is composed of eight generic groups: Heteropterus, Astictopterus,
Isoteinon, Ancistroides, Plastigia, Hesperia, Taractrocera, and Gegenes. Yania can
be distinguished immediately from the Hesperia, Taractrocera, and Gnegnes
groups by an antennal club that is not constricted before the apiculus and
by a hindwing vein 5 that is well marked. Yania is differentiated from the
Heteropterus group in having the hindwing cell less than half the wing length,
antennae not short and legs normal (fore tibia with prominent epiphysis,
mid tibia not spined, and hind tibia with prominent upper spurs). Yania is
distinct from Plastingia with hindwing vein 5 closer to vein 4 than 6 and its

150

/. Res. Lepid.

wings not obviously produced. Yania differs from the Isoateinon group with
the hindwing median vein not co-linear with vein 4 and vein 2 before the
origin of vein 7. It would be difficult to infer to which of the remaining
groups, Astictopterus or Ancistroides, Yania is most closely related since Evans
only used the state of the second palpus segment — erect or porrect as

Fig. 3. Antennae of Yania sinica.

Fig. 4. Legs of Yania sinica (left to right): fore leg, mid leg, hind leg.

Fig. 5. Wing venation of Yania sinica.

Fig. 6. Male genitalia of Yania sinica consisting of lateral view of genital cap-
sule with left valva and aedeagus removed; dorsal view of gential cap-
sule with juxta and aedeagus removed; ventral view of gnathos and un-
cus; dorsal view of aedeagus; lateral view of aedeagus; and juxta in pos-
terior view.

34:147-153, 1995(1997)

151

the key for separation (palps of the unique holotype of Yania sinica are
missing). However, Eliot (in Corbet & Pendlebury 1992:363) questioned
the taxonomic value of palpi and rearranged the Hesperinae accordingly.
The Astictopterus group was suppressed, the Ampittia and Hesperia subgroups
placed into the Halpe group, and the genera Astictopterus and Arnetta placed
into the Astictopterus and Plastingia groups respectively. The latter rationale
is provided by wing venation and male genitalic character states. The Halpe
group differs from the Ancistroides group of Eliot (including Astictopterus)
in that forewing vein 2 arises opposite or beyond the origin of vein 1 1 and
the uncus of the male genitalia is broader in dorsal aspect. It follows then
that Yaniah^ placed in the Ancistroides growp of Eliot which comprises eight
Asian genera: lambrix, Idmon, Koruthaialos, Psolos, Astictopterus, Ancistroides,
Notocrypta, and Udaspes.

The phylogenic relationships between Yania and these eight genera are
of interest. According to Eliot’s key, mainly based on wing venation and
wing markings, Yania can be distinguished from Notocrypta and Udaspes in
the first dichotomy by a forewing without large hyaline spots; from lambrix,
Idmon, Koruthaialos, and Psolos in the second dichotomy by both wings hav-
ing vein 5 downcurved at its origin and closer to vein 4 than vein 6; from
Astictopterus in the third dichotomy by forewing vein 1 1 about midway be-
tween veins 10 and 12; leaving Ancistroides the closest allied genus to Yania.
However, Yania shares other characters which appear as important as veins
5 and 11 for generic classification. These include body aspect and the
hindwing dorsum shorter than the costa, which place Yania closest to
Astictopterus and differing from the other seven genera. With the forewing
vein 2 closer to the wing base than to vein 3 at its origin, Yania resembles
Notocrypta and Udaspes. With regard to male genitalia the most important
character for determination of generic classification appears to be the de-
gree to which the clasp is specialized. Secondary characters are relative
length of the uncus to the tegumen, length and shape of the gnathos, and
least important the shape of the uncus. I propose this hierarchy from expe-
rience with treatment of the well defined subgeneric groups. Thus, within
the Halpe group, all genera can clearly be placed into two subgroups by
specialization of the clasp: the Halpe subgroup has the cuiller of the clasp
longer and more complex (usually with heavy and branching teeth) than
in the Ampittia subgroup.

Although the shape of the uncus is variable within related genera, the
character is usually stable within a single genus. The relative length of the
uncus to the tegumen is of greater supergeneric value than uncus shape
alone. For example, the two closely related genera, lambrix 3.nd Idmon, both
have a long uncus, but in one the uncus tapers to a long point, in the other
the uncus is deeply bifid. Accordingly, Yania resembles lambrix and Idmon
in male genitalia.

In the following key for separating genera of the Ancistroides group, I
employ male genitalia as the main character. However, since the discovery
of Yania, it is not possible to decide which vein — 2, 5, or 11 ^ — is more

152

/. Res. Lepid.

important for inferring phylogeny. For this reason, the subgroups which
Eliot (1992:37) divided the group have been disregarded.

Key to the genera of the Ancistroides group

1 (6) Male genitalia with uncus substantially longer than tegumen.

(Male clasp very simple in structure. Forewing vein 11 midway
between veins 10 and 12.) ............................. (lambrix subgroup)

2 (5) Forewing vein 5 midway between veins 4 and 6 at origin. Forew-

ing vein 2 closer to vein 3 than wing base at origin. Male forew-
ing usually without brand. Body robust.

3 (4) Male genitalia with uncus tapered to a long point. Underside

hindwing with silveiy-white spots, ...................................... lambrix

4 Male genitalia with uncus deeply bifid. ............................... Idmon

5 Both wings vein 5 slightly closer to vein 4 than 6. Forewing vein 2
closer to wing base than to vein 3 at origin. Male without brand.
Body thin. .............................................................................. Yania

6 Male genitalia with uncus slightly longer than tegumen.

7 (12) Forewing vein 11 midway between veins 10 and 12. Male clasp

with cuiller not forked with harpe, or bearing a style from valva
or from cuiller. ........................................ {Ancistroides subgroup)

8 (11) Forewing vein 2 closer to wing base than to vein 3.

9 (10) Hindwing cell half wing length. Antennae longer than half length

of forewing costa. Upperside hindwing unmarked.

................................................................................... Notocrypta

Hindwing cell shorter than half the wing length. Antennae
shorter than half length of forewing costa. Upperside hindwing
with large white discal area. .............................................. Udaspes

Forewing vein 2 closer to vein 3 than to wing base at origin.

Forewing vein 11 bowed toward or briefly touching or an-
astomosing with vein 12 and remote from vein 10. Male clasp with
cuiller forked with harpe and without style from valva.

(Astictopterus group)

(14) Both wings vein 5 slightly downcurved at origin, closer to vein 4
than 6. Male without secondary sex characters. Body weak.

Astictopterus

Both wings vein 5 midway between veins 10 and 12. Males with
secondary sex characters. Body robust.

(16) Forewing origin vein 4 midway between veins 3 and 5.

Forewing origin vein 4 closer to 5 than 3. ........................... Pso/os

Suggested Phytogeny

Yania appears to represent a mixture of all genera of the Ancistroides group,
which I interpret to make it ancestral for the group. Yania shows more ap-
parent primative characters than any other genus in the group: structure

34:147-153, 1995(1997)

153

Udaspes

Notocrypta

Ancistroides

Koruthaialos

Psolos

Astictopterus

Yania

Idmon

lambrix

Fig. 7. Suggested phylogeny of Ancistroides group.

of the male genitalia is simple, veins 1 1 and 12 of the forewing are not joined,
the antennal club is formed gradually, and secondary sex characters are
absent. I lastly present the following hypothetical phylogenetic tree based
on my intuitive evaluation of the selected diagnostic characters (Fig. 7).

Acknowledgments. I wish especially to thank two anonymous reviewers for suggestions
that materially helped my presentation including substantive editing in English.

Literature Cited

Corbet, A.S. & H.M. Pendlebury. 1992. The butterflies of the Malay Peninsula. 3rd
edition revised by J.N. Eliot. Malay Nature Society, Kuala Lumpur.

Evans, W.H. 1949. A Catalogue of the Hesperiidae from Europe, Asia, and Australia
in the British Museum (N.H.). British Museum (N.H.), London.

Journal of Research on the Lepidoptera

34:154-160, 1995(1997)

A commentary on the recent book. Butterflies of Costa Rica
and their natural history: voL 2

“Books are not made to be believed, but to be subjected to inquiry”

— Umberto Eco

Although it was ordered and paid for in 1996, my copy of the new Costa
Rican riodinid field guide (DeVries, PJ., 1997. Butterflies of Costa Rica and
their natural history: voL 2, Riodinidae. Princeton University Press, 288
pages, 25 plates; ISBN: 0-691-02890-7) arrived but a few weeks ago — - a lag
of over nine months. I’ve heard rumors that the delay was due to unfore-
seen technical difficulties stemming from the vagaries of the publishing busi-
ness. But in America tardiness seems to be the corporate standard. As Fats
Waller once philosophically opined, “you P^ys your money, and you takes
your chances.” But what of that. The book arrived and could now be pe-
rused in my spare moments.

A quick glance through the riodinid book revealed a similar layout to the
previous volume. A further riffle showed what appeared to be a substantial
consignment of information on riodinid butterflies, some technical pho-
tos, tables with numbers, and to my satisfaction, the color plates appeared
to be useful for identifying riodinid specimens in the collection.

Carrying the volume to the study I pulled out a few specimens and com-
pared them to the plates. I was pleased to find that the plates were adequate
for this task. After identifying a number of specimens and writing the names
carefully on individual labels I decided to read a little about one particu-
larly odd looking species, Syrmatia nyx, on Plate 9. Great Scott!, I exclaimed,
as my whiskey glass crashed to the floor and the dog yelped from its slum-
bers by the fire. Imagine my shock at finding that not only the facing plate
for number 9, but indeed all of the facing plates were devoid of page num-
bers for the species accounts. I resolved to get to the bottom of this. Pour-
ing myself another stiff whiskey to stem the tide of annoyance, and another
for the dog to soothe its agitation, I mounted my inquiry by lunging to the
index. There I found the answer I was looking for — page 165, and so turn-
ing to the proper page I read the information about the odd little Syrmatia
nyx.

This archaic method of finding out information from the index was in-
convenient, even if it did work for the odd little riodinid. This, however,
did not solve the problem for all of the other species treated in the book.
Something had to be done. Much to the exasperation of the dog’s liver I
set to work, and the epistle presented here was born after a good many
whiskeys and words of opprobrium shouted into the night. That is to say,
users of the new riodinid volume may find the following table useful; it pro-
vides the page numbers in the text for the species illustrated on the plates.
The user can now annotate the page numbers directly on the facing plates

34:154-160, 1995(1997)

155

in the book, as I have done on my copy. Furthermore, I have noted in bold
face some errors or inconsistencies between the plates, the index, and the
species accounts.

One hopes that my missive here will be useful to current book owners,
and that the publisher will eventually correct these omissions in future edi-
tions of the book.

““ Reginald B. Swinethrottle

Department of Biology
University of Oregon
Eugene, Oregon 97403

Table 1 . Additions and corrections for facing plates. Text pages are provided for
all species illustrated in the color identification plates. Errors or
inconsistencies are noted in bold face.

Plate 1

18)

Euselasia inconspicua, p.l22

Corrachia leucoplaga, p.ll3

19)

Euselasia amphidecta, p. 124

Hades noctula, p.ll4

20)

Euselasia amphidecta, p. 124

Hades noctula, p.ll4

21)

Euselasia amphidecta, p.24

Methone cecilia chrysomela, p.ll5

22)

Euselasia gyda, p.l20

Methone cecilia chrysomela, p.ll5

23)

Euselasia gyda, p.l20

Euselasia bettina, p.ll7

24)

Euselasia gyda, p.l20

Euselasia bettina, p.ll7

25)

Euselasia leucon, p.l22

Euselasia aurantia, p.ll7

26)

Euselasia leucon, p.l22

Euselasia aurantia, p.ll7

27)

Euselasia argentea, p.l23

10)

Euselasia chrysippe, p.ll8

28)

Euselasia argentea, p.l23

11)

Euselasia chrysippe, p.ll8

12)

Euselasia chrysippe, p.ll8

Plate 3

13)

Euselasia matuta, p.ll8

Euselasia midas, p.l24

14)

Euselasia matuta, p.ll8

Euselasia midas, p.l24

15)

Euselasia leucophryna, p.ll7

Euselasia midas, p.l24

16)

Euselasia corduena, p.ll9

Euselasia rhodogyne, p.l25

17)

Euselasia corduena, p.ll9

Euselasia rhodogyne, p.l25

18)

Euselasia corduena, p.ll9

Euselasia rhodogyne, p.l25

Euselasia subargentea, p.l26

Plate 2

Euselasia subargentea, p.l26

Euselasia labdacus, p.l22

Euselasia regipennis, p.ll9

Euselasia labdacus, p.l22

10)

Euselasia regipennis, p.ll9

Euselasia labdacus, p.l22

11)

Euselasia regipennis, p.ll9

Euselasia mystica, p.l21

12)

Euselasia regipennis, p.ll9

Euselasia mystica, p.l21

13)

Euselasia aurantiaca, p.l25

Euselasia mystica, p.l21

14)

Euselasia aurantiaca, p.l25

Euselasia hieronymi, p.l21

15)

Peropthalma lasus, p.l27

Euselasia hieronymi, p.l21

16)

Peropthalma lasus, p.l27

Euselasia hieronymi, p.l21

17)

Peropthalma tullius, p.l27

10)

Euselasia eubule D, p.l24

18)

Peropthalma tullius, p.l27

11)

Euselasia eubule, p.l24

19)

Euselasia angulata, p.l26

12)

Euselasia eubule, p,124

20)

Euselasia onorata, p.l24

13)

Euselasia eucrates D, p.l23

14)

Euselasia eucrates, p.l23

Plate 4

15)

Euselasia eucrates, p.l23

Leucochimona vestalis, p. 128

16)

Euselasia inconspicua, p.l22

Leucochimona vestalis, p. 128

17)

Euselasia inconspicua, p.l22

Leucochimona lepida, p. 129

156

J. Res. Lepid.

4) Leucochimona lepida, p. 129

5) Leucochimona lagora, p. 129

6) Leucochimona lagora, p. 129

7) Mesosemia hesperina, p. 131

8) Mesosemia hesperina, p. 131

9) Mesosemia hesperina, p. 131

10) Mesosemia esperanza, p. 130

11) Mesosemia esperanza, p. 130

12) Mesosemia esperanza, p. 130

13) Mesosemia coelestis, p. 131

14) Mesosemia coelestis, p. 131

15) Mesosemia albipuncta, p. 131

16) Mesosemia albipuncta, p. 131

17) Mesosemia zonalis, p. 132

18) Mesosemia zonalis, p. 132

19) Mesosemia carissima, p. 132

20) Mesosemia carissima, p. 132

21) Mesosemia carissima, p. 132

22) Mesosemia asa, p. 135

23) Mesosemia asa, p. 135

Plate 5

1) Mesosemia grandis, p. 133

2) Mesosemia grandis, p. 133

3) Mesosemia gaudiolum, p. 133

4) Mesosemia gaudiolum, p. 133

5) Mesosemia hypermegala, p. 134

6) Mesosemia hypermegala, p. 134

7) Mesosemia ceropia, p. 133

8) Mesosemia ceropia, p. 133

9) Mesosemia lamachus, p. 134

10) Mesosemia lamachus, p. 134

11) Mesosemia telegone, p. 134

12) Mesosemia telegone, p. 134

13) Napaea eucharila, p. 142

14) Napaea eucharila, p. 142

15) Napaea eucharila, p. 142

16) Napaea theages, p. 143

17) Napaea theages, p. 143

18) Napaea umbra, p. 144

Plate 6

1) Eurybia cyclopia, p. 138

2) Eurybia caerulescens fulgens, p. 138

3) Eurybia lycisca, p. 140

4) Eurybia unxia, p. 138

5) Eurybia unxia, p. 138

6) Voltinia theata, p. 144

7) Voltinia theata, p. 144

8) Voltinia theata, p. 144

9) Eurybia patrona, p. 139

10) Eurybia elvina, p. 139

11) Voltinia radiata, p. 144

12) Necyria ingaretha, p. 149

13) Hermathena candidata, p. 146

14) Hermathena oweni, p. 146

15) Cyrenia martia, p. 150

Plate 7

1) Lyropteryx lyra cleadas, p. 148

2) Lyropteryx lyra cleadas, p. 148

3) Lyropteryx lyra cleadas, p. 148

4) Chorinea octauius, p. 156

5) Ithomeis eulema, p. 156

6) Necyria beltiana, p. 149

7) Necyria beltiana, p. 149

8) Necyria beltiana, p. 149

9) Necyria beltiana, p. 149

10) Monethe rudolphus, p. 159

11) Monethe rudolphus, p. 159

12) Cremna thasus, p. 145

13) Cremna thasus, p. 145

14) Notheme erota, p. 162

15) Ancyluris inca, p. 151

16) Ancyluris inca, p. 151 (“p. 15” in

index)

Plate 8

1) Ancyluris jurgensenii, p. 152

2) Ancyluris jurgensenii, p. 152

3) Rhetus dysonii, p. 153

4) Rhetus dysonii, p. 153

5) Rhetus dysonii, p. 153

6) Rhetus arcius, p. 153

7) Rhetus arcius, p. 153

8) Rhetus periander, p. 154

9) Rhetus periander, p. 154

10) Rhetus periander, p. 154

11) Brachyglenis dodona, p. 158

(should be dodone)

12) Brachyglenis dodona, p. 158

(should be dodone)

13) Brachyglenis dinora, p. 158

14) Brachyglenis dinora, p. 158

15) Lepricornis strigosa, p. 164

16) Lepricornis strigosa, p. 164

Plate 9

1) Cariomothis poeciloptera, p. 165

2) Cariomothis poeciloptera, p. 165

3) Cariomothis poeciloptera, p. 165

4) Cariomothis poeciloptera, p. 165

5) Syrmatia nyx, p. 165

6) Syrmatia aethiops, p. 165

7) Chamaelimnas villagomes, p. 166

8) Chamaelimnas villagomes, p. 166

9) Exoplisia cadmeis, p. 181

10) Exoplisia cadmeis, p. 181

11) Exoplisia hypochalbe, p. 181

12) Pterographium elegans, p. 192

13) Pterographium elegans, p. 192

14) Isapis agyrtus, p. 161

15) Isapis agyrtus, p. 161

16) Melanis pixie, p. 160

17) Melanis electron, p. 161

34:154-160, 1995(1997)

157

18) Melanis cephise, p. 160

19) Xenandra desora, p. 187

20) Xenandra helius, p. 187

21) Xenandra caeruleata, p. 186

22) Xenandra caeruleata, p. 186

Plate 10

1) Metacharis victrix, p. 163

2) Metacharis victrix, p. 163

3) Metacharis victrix, p. 163

4) Metacharis victrix, p. 163

5) Caria rhacotis, p. 167-168

6) Caria rhacotis, p. 167-168

7) Caria rhacotis, p. 167-168

8) Caria rhacotis, p. 167-168

9) Esthemopsis clonia, p. 187

10) Esthemopsis clonia, p. 187

11) Esthemopsis colaxes, p. 188

12) Esthemopsis colaxes, p. 188

13) Caria lampeto, p. 168

14) Caria lampeto, p. 168

15) Caria lampeto, p. 168

16) Caria lampeto, p. 168

17) Caria domitianus, p. 168

18) Caria domitianus, p. 168

19) Caria domitianus, p. 168

20) Baeotis nesaea, p. 166

21) Baeotis nesaea, p. 166

22) Baeotis zonata, p. 167

23) Baeotis macularia, p. 167

(sulphuria macularia)

24) Baeotis macularia, p. 168

(sulphuria macularia)

25) Argyrogrammana holosticta, p. 204

26) Argyrogrammana holosticta, p. 204

27) Parcella amarynthina, p. 171

28) Parcella amarynthina, p. 171

Plate 11

1) Charis auius, p. 172

2) Charis auius m, p. 172

3) Charis auius, p. 172

4) Charis gynaea, p. 173

5) Charis gynaea, p. 173

6) Charis gynaea, p. 173

7) Charis gynaea, p. 173

8) Charis hermodora, p. 173

9) Charis hermodora, p. 173

10) Charis hermodora, p. 173

11) Charis hermodora, p. 173

12) Adelotypa eudocia, p. 236

13) Adelotypa eudocia, p. 236

14) Adelotypa glauca, p. 236

15) Adelotypa glauca, p. 236

16) Parnes nycteis, p. 227

17) Adelotypa densemaculata, p. 235

18) Adelotypa densemaculata, p. 235

19) Roberella lencates, p. 212

20) Calospila trotschi, p. 234

Plate 12

1) Calephelis sixaola, p. 174

2) Calephelis sixaola, p. 174

3) Calephelis sixaola, p. 174

4) Calephelis fulmen, p. 176

5) Calephelis fulmen, p. 176

6) Calephelis fulmen, p. 176

7) Calephelis schausi, p. 176

8) Calephelis schausi, p. 176

9) Calephelis schausi, p. 176

10) Calephelis browni, p. 177

11) Calephelis browni, p. 177

12) Calephelis browni, p. 177

13) Calephelis costaricicola, p. 177

14) Calephelis costaricicola, p. 177

15) Calephelis costaricicola, p. 177

16) Calephelis sodalis, p. 177

17) Calephelis sodalis, p. 177

18) Calephelis sodalis, p. 177

19) Calephelis argyrodines, p. 178

20) Calephelis argyrodines, p. 178

21) Calephelis argyrodines, p. 178

22) Calephelis laverna parva, p. 178

23) Calephelis laverna parva, p. 178

24) Calephelis laverna parva, p. 178

25) Calephelis exiguus, p. 178

26) Calephelis exiguus, p. 178

27) Calephelis inca, p. 178

28) Calephelis inca, p. 178

Plate 13

1) Argyrogrammana venilia crocea, p. 204

2) Argyrogrammana venilia crocea, p. 204

3) Argyrogrammana venilia crocea, p. 204

4) Argyrogrammana venilia crocea, p. 204

5) Argyrogrammana leptographia, p. 204

6) Argyrogrammana leptographia, p. 204

7) Argyrogrammana leptographia, p. 204

8) Argyrogrammana leptographia, p. 204

9) Argyrogrammana barine, p. 205

10) Argyrogrammana barine, p. 205

11) Mesene silaris, p. 183

12) Mesene silaris, p. 183

13) Mesene phareus, p. 182

14) Mesene phareus, p. 182

15) Mesene phareus, p. 182

16) Mesene phareus, p. 182

17) Mesene mygdon, p. 182

18) Mesene mygdon, p. 182

19) Mesene mygdon, p. 182

20) Mesene mygdon, p. 182

21) Mesene margaretta, p. 183

22) Mesene croceella, p. 183

23) Mesenopsis melanochlora, p. 185

158

/. Res. Lepid.

24) Mesenopsis bryaxis, p. 186

25) Chimastrum argenteum, p. 188

26) Symmachia rubina, p. 189

27) Symmachia rubina, p. 189

28) Symmachia rubina, p. 189

29) Symmachia threissa, p. 189

30) Symmachia threissa, p. 189

31) Symmachia tricolor, p. 191

32) Symmachia tricolor, p. 191

Plate 14

1) Symmachia accusatrix, p. 190

2) Symmachia accusatrix, p. 190

3) Symmachia accusatrix, p. 190

4) Symmachia leena, p. 190

5) Symmachia leena, p. 190

6) Symmachia leena, p. 190

7) Symmachia probetor, p. 190

8) Symmachia probetor, p. 190

9) Symmachia xypete, p. 191

10) Symmachia xypete, p. 191

11) Symmachia xypete, p. 191

12) Phaenochitonia ignipicta, p. 194

13) Phaenochitonia ignipicta, p. 194

14) Phaenochitonia ignipicta, p. 194

15) Phaenochitonia ignicauda, p. 194

16) Phaenochitonia ignicauda, p. 194

17) Stichelia sagaris tyriotes, p. 193

18) Stichelia sagaris tyriotes, p. 193

19) Stichelia sagaris tyriotes, p. 193

20) Stichelia sagaris tyriotes, p. 193

21) Stichelia phoenicura, p. 193

22) Stichelia phoenicura, p. 193

23) Anteros allectus, p. 195

24) Anteros allectus, p. 195

25) Anteros allectus, p. 195

26) Anteros chrysoprastus, p. 195

27) Anteros chrysoprastus, p. 195

28) Anteros renaldus, p. 197

29) Anteros renaldus, p. 197

30) Anteros carausius, p. 197

31) Anteros formosus micon, p. 196

Plate 15

1) Anteros kupris, p. 196

2) Anteros kupris, p. 196

3) Sarota subtessellata, p. 202

4) Sarota subtessellata, p. 202

5) Sarota subtessellata, p. 202

6) Sarota turrialbensis, p. 203

7) Sarota chrysus, p. 201

8) Sarota chrysus, p. 201

9) Sarota myrtea, p. 199

10) Sarota gamelia, p. 199

11) Sarota spicata, p. 200

12) Sarota estrada, p. 200

13) Sarota estrada, p. 200

14) Sarota psaros, p. 201

15) Sarota gyas, p. 199

16) Sarota gamelia, p. 199

17) Sarota acantus, p. 200

18) Sarota acantus, p. 200

19) Chalodeta lypera, p. 170

20) Chalodeta lypera, p. 170

21) Chalodeta lypera, p. 170

22) Chalodeta chaonitis, p. 170

23) Chalodeta chaonitis, p. 170

24) Chalodeta candiope, p. 171

25) Chalodeta candiope, p. 171

26) Chalodeta candiope, p. 171

27) Charis iris, p. 172

28) Charis iris, p. 172

29) Charis iris, p. 172

Plate 16

1) Emesis ocypore, p. 208

2) Emesis ocypore, p. 208

3) Emesis ocypore, p. 208

4) Emesis lupina, p. 209

5) Emesis lupina, p. 209

6) Emesis lupina, p. 209

7) Lasaia agesilas, p. 179

8) Lasaia agesilas, p. 179

9) Lasaia agesilas, p. 179

10) Lasaia sula, p. 180

11) Lasaia sula, p. 180

12) Lasaia sula, p. 180

13) Lasaia sessilis, p. 179

14) Lasaia sessilis, p. 179

15) Lasaia pseudomeris, p. 180

16) Lasaia pseudomeris, p. 180

17) Lasaia oileus, p. 180

18) Lasaia oileus, p. 180

19) Lasaia oileus, p. 180

20) Calydna hiria, p. 206 (should be stemula)

2 1 ) Calydna hiria, p. 206 (should be stemula)

22) Calydna hiria, p. 206 (should be stemula)

23) Calydna venusta, p. 205

24) Apodemia multiplaga, p. 213

25) Apodemia multiplaga, p. 213

Plate 17

1) Emesis lacrines, p. 208

2) Emesis lacrines, p. 208

3) Emesis lacrines, p. 208

4) Emesis lucinda, p. 210

5) Emesis lucinda, p. 210

6) Emesis lucinda, p. 210

7) Emesis mandana, p. 209

8) Emesis mandana, p. 209

9) Emesis mandana, p. 209

10) Emesis fatimella, p. 210

11) Emesis fatimella, p. 210

12) Emesis cypria, p. 208

34:154-160, 1995(1997)

159

13) Emesis tenedia, p. 206

14) Emesis tenedia, p. 206

15) Emesis tenedia, p. 206

16) Emesis tenedia, p. 206

17) Emesis tegula, p. 209

18) Emesis tegula, p. 209

19) Emesis tegula, p. 209

Plate 18

1) Thisbe irenea, Panama, p. 215

2) Thisbe irenea, Panama, p. 215

3) Thisbe lycorias, p. 216

4) Uraneis ucubis, p. 217

5) Uraneis ucubis, p. 217

6) Juditha dorilas, p. 219

7) Lemonias agave, p. 217

8) Lemonias agave, p. 217

9) Lemonias agave, p. 217

10) Juditha molpe, p. 218

11) Juditha molpe, p. 218

12) Juditha dorilas, p. 219

13) Juditha dorilas, p. 219

14) Catocyclotis aemulius, p. 219

15) Catocyclotis aemulius, p. 219

16) Synargis mycone, p. 222

17) Synargis mycone, p. 222

18) Synargis mycone, p. 222

19) Synargis ochra sicyon, p. 223

20) Synargis ochra sicyon, p. 223

Plate 19

1) Synargis phylleus, p. 220

2) Synargis phylleus, p. 220

3) Synargis phylleus, p. 220

4) Synargis phylleus, p. 220

5) Synargis palaeste, p. 223

6) Synargis palaeste, p. 223

7) Synargis nymphidioides, p. 224

8) Synargis nymphidioides, p. 224

9) Synargis nycteis, p. 225

10) Synargis nycteis, p. 225

Plate 20

1) Rodinia calpharnia, p. 234

2) Audre domina, p. 226

3) Audre albina, p. 226

4) Audre albina, p. 226

5) Menander menander, p. 228

6) Menander menander, p. 228

7) Menander menander, p. 228

8) Menander pretus, p. 230

9) Menander pretus, p. 230

10) Menander pretus, p. 230

11) Pandemos godmanii, p. 231

12) Pandemos godmanii, p. 231

13) Periplacis glaucoma, p. 227

14) Periplacis glaucoma, p. 227

15) Menander laobotas, p. 230

16) Menander laobotas, p. 230

Plate 21

1) Synargis velabrum, p. 223

2) Synargis gela, p. 225

3) Pachythone gigas, p. 211

4) Pachythone gigas [ignifer], p. 211

5) Pachythone gigas, p. 211

6) Calospila asteria, p. 232

7) Calospila asteria, p. 232

8) Calospila asteria, p. 232

9) Calospila lucianus, p. 231

10) Calospila lucianus, p. 231

11) Calospila lucianus, p. 231

12) Calospila cilissa, p. 232

13) Calospila cilissa, p. 232

14) Calospila cilissa, p. 232

15) Calociasma lilina, p. 237

16) Calociasma icterica, p. 236

17) Calospila martia, p. 232

18) Calospila martia, p. 232

19) Calospila sudias, p. 233

20) Calospila sudias, p. 233

21) Calospila sudias, p. 233

22) Calospila argenissa, p. 233

23) Calospila argenissa, p. 233

24) Calospila argenissa, p. 233

25) Calospila argenissa, p. 233

26) Calospila zeurippa, p. 233

27) Calospila zeurippa, p. 233

28) Calospila parthaon, p. 233

29) Calospila parthaon, p. 233

30) Calospila parthaon, p. 233

Plate 22

1) Setabis lagus, p. 237

2) Setabis lagus, p. 237

3) Setabis lagus, p. 237

4) Setabis alcmaeon, p. 238

5) Setabis alcmaeon, p. 238

6) Setabis alcmaeon, p. 238

7) Setabis cleomedes, p. 238

8) Setabis cleomedes, p. 238

9) Setabis cleomedes, p. 238

10) Pixus corculum, p. 212

11) Pixus corculum, p. 212

12) Pixus corculum, p. 212

13) Nymphidium mantus, p. 248

14) Nymphidium lenocinium, p. 250

15) Nymphidium lenocinium, p. 250

16) Nymphidium olinda, p.249

17) Pseudonymphidia clearista, p. 239

18) Nymphidium onaeum, p. 251

19) Nymphidium onaeum, p. 251

20) Nymphidium azanoides, p. 250

21) Nymphidium azanoides, p. 250

160

J. Res. Lepid.

22) Nymphidium ascolia, p. 251

23) Nymphidium haematostictum, p. 250

Plate 23

1) Theope virgilius, p. 240

2) Theope virgilius, p. 240

3) Theope virgilius, p. 240

4) Theope eupolis, p. 241

5) Theope eupolis, p. 241

6) Theope eupolis, p. 241

7) Theope publius, p. 243

8) Theope publius, p. 243

9) Theope publius, p. 243

10) Theope eleutho, p. 242

11) Theope eleutho, p. 242

12) Theope eleutho, p. 242

13) Theope basilea, p. 242

14) Theope basilea, p. 242

15) Theope basilea, p. 242

16) Theope cratylus, p. 242

17) Theope cratylus, p. 242

18) Theope cratylus, p. 242

Plate 24

1) Theope matuta, p. 244

2) Theope matuta, p. 244

3) Theope matuta, p. 244

4) Theope speciosa, p. 241

5) Theope speciosa, p. 241

6) Theope speciosa, p. 241

7) Theope phaeo folia, p. 247

8) Theope phaeo folia, p. 247

9) Theope phaeo folia, p. 247

10) Theope pedias, p. 247

11) Theope pedias, p. 247

12) Theope pedias, p. 247

13) Theope herta, p, 246

14) Theope herta, p. 246

15) Theope herta, p. 246

16) Theope barea, p. 246

17) Theope barea, p. 246

18) Theope barea, p. 246

19) Theope acosma, p. 244

20) Theope acosma, p. 244

21) Theope decorata, p. 245

(should be thestias decorata)

22) Theope decorata, p. 245

(should be thestias decorata)

23) Theope eudocia, p. 244

24) Theope eudocia, p. 244

Plate 25

1) Metacharis onorata, p. 163

(should be umbrata)

2) Metacharis onorata, p. 163

(should be umbrata)

3) Theope guillaumei cecropia, p. 245

4) Theope lycaenina, p. 247

5) Theope lycaenina, p. 247

6) Mesosemia harveyi, p. 135

7) Brachyglenis esthema, p. 157

8) Brachyglenis esthema, p. 157

9) Brachyglenis nr dodona, p. 158

(should be nr dodone)

10) Brachyglenis nr dodona, p. 158

(should be nr dodone)

Immature stages of high arctic Gynaephora ( Lyman triidae) and notes

on their biology at Alexandra Fiord, Ellesmere Island, Canada 119

Wm, Dean Morewood and Petra Lange

Notes on Boloria pales yangi, ssp. nov., a remarkable disjunction in butterfly

biogeography (Lepidoptera: Nymphalidae) 142

Yu~Feng Hsu and Shen-Horn Yen

Yania gen. nov. and Yania sinica sp. nov. from Sichuan, China (Lepidoptera:

Hesperiidae) 147

Hao Huang

A commentary on the recent book. Butterflies of Costa Rica and their natural
history: vol. 2 154

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The Journal of Research

ON THE LEPIDOPTERA

Volume 34 Number 1—4 1 995( 1 997)

IN THIS ISSUE

Date of Publication: December 15, 1997

Evolution of locomotion in slug caterpillars (Lepidoptera: Zygaenoidea:

Limacodid group) 1

Marc E. Epstein

Territoriality by the dawn’s early light: the Neotropical owl butterfly Caligo

idomenaeus (Nymphalidae: Brassolinae) 14

Andre V. L. Ereitas, Woodruff W. Benson, Onildo J. Marini-Eilho, and
Roberta M. de Carvalho

A review of the genus Pttnara Doubleday, 1847 (Riodinidae) in southeast

Brazil, with a description of two new subspecies ............................................. 21

Curtis J. Callaghan

Lepidoptera of different grassland types across the Morava floodplain 39

Miroslav Kulfan, Peter Degma, and Henrik Kalivoda

Effectiveness of caterpillar defenses against three species of invertebrate

predators 48

Lee A. Dyer

Cooperation vs. exploitation: interactions between Lycaenid (Lepidoptera:

Lycaenidae) larvae and ants .........69

E. Osborn and K. Jaffe

A revision of Mesogona Boisduval (Lepidoptera: Noctuidae) for North

America with descriptions of two new species 83

Lars Crabo and Paul C. Hammond

The endangered quino checkerspot butterfly, Euphydryas editha quino

(Lepidoptera: Nymphalidae) ............................................................................ 99

Rudi Mattoni, Gordon E. Pratt, Travis R. Longcore, John E. Emmel, and
Jeremiah N. George

(contents continued inside cover)

Cover: Quino checkerspot butterfly, Euphydryas editha quino

Journal of
Research on the
Lepidoptera

Volume 35

1 996 (2000)

The Journal of Research

ON THE LEPIDOPTERA

ISSN 0022 4324

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Journal of Research on the Lepidoptera

35:1-8, 1996 (2000)

Differences in lifetime reproductive output and mating
frequency of two female morphs of the sulfur butterfly,

Colias erate (Lepidoptera: Pieridae)

Yasuyiiki Nakanishi, Mamoru Watanabe & Takahiko Ito

Dept, of Biology, Fac. of Education, Mie University, 1515 Kamihama, Tsu, Mie 5 14-8507, Japan.
E-mail: bxa@nifty.ne.jp

Abstract. Both female morphs of the sulfur butterfly Colias erate were cap-
tured in the field and dissected to investigate whether differences of re-
productive output are affected by mating frequency between them. Life-
time mating frequency of the yellow morph was significantly lower than
that of alba. In every female the immature egg load decreased with age.
During their life span the monandrous yellow morph laid about 550 eggs,
with the monandrous alba producting about 400 eggs. Polyandrous alba
laid more eggs than either the monandrous or polyandrous yellow mor-
phs. Thus, multiple mating is important for alba to increase its repro-
ductive output, supporting the field observation that alba effectively at-
tracts males.

Introduction

Some butterfly species show wing color polymorphism, such as Papilio
polytes (Watanabe 1979, Uesugi 1992) and Lycaena phlaeas (Brakefield &
Shreeve 1992). Komai and Ae (1953) reported that thejapanese sulfur but-
terfly, Colias erate, exhibits a sex-limited wing color dimorphism in females
with yellow (ancestral) and white (alba) morphs. Both morphs occur sym-
patrically although the yellow morph is never more abundant than alba (e.g.,
Watanabe & Nakanishi 1996), unlike most American species. Gilchrist
and Rutowski (1986) explained the adaptive significance of alba from the
viewpoint of reproductive success.

Emmel (1972) hypothesized that female-limited dimorphism is balanced
by differential mate selection by males. Ley and Watt (1989) studied female
limited dimorphism and concluded that the dimorphism is balanced by
differential predation on the morphs. The persistence of the two morphs
in time, however, suggests that they are equally fit. Other studies on C. erate
(e.g., Watanabe et al. 1997) demonstrated a higher frequency of mate at-
traction by the alba morph which would give them a fitness advantage. The
latter observation predicts that the yellow morph must have a compensa-
tory advantage beyond mating that equalizes their lifetime fitness. Mating
behavior of female morphs has been reported for some Colias species in
America (e.g., Gilchrist & Rutowski 1986, Graham et al. 1980). Our study
focuses on the lifetime reproductive success of the two morphs.

Recent studies have shown that female butterflies may engage in polyan-

Paper submitted 10 May 1996; revised manuscript accepted 6June 1998.

J. Res. Lepid.

dry to obtain sperm and/or nutrients that are ejaculated from males (e.g.,
Boggs & Gilbert 1979) . Wdien a mated female accepts subsequent males the
last male’s sperm has precedence (e.g., Watanabe 1988). In C. erate
Watanabe & Nakanishi (1996) pointed out that females are polyandrous.
However, there is no report on the lifetime reproductive success of the two
morphs of this species from the viewpoint of female polyandry. In this pa-
per, we examine how frequently females in the field mated during their
life span, and then estimated the fecundity for each morph.

Materials and Methods

The data were obtained mainly from the summer generations of C. erate in

Shiroiima of Nagano Prefecture, which is located in a cool temperate zone of Ja-
pan. The details of the study area have been described elsewhere (e.g., Watanabe
& Nakanishi 1996). The habitat consisted of rice fields and five ski slopes where
nectar sources and larval food plants were abundant.

We collected females engaged in various activities including feeding, roosting,
flying, copulating and ovipositing on calm sunny days from late July through mid-
August of 1989 to 1994 (n=34 days). WTeii females were captured, their abdomens
were amputated and immersed in 50% ethyl alcohol. Forewing length of each was
also recorded. The age of each female was estimated mainly by wing wear condi-
tion, and rated among 1 to 5 age classes (Watanabe & Nakanishi 1996).

Thirty seven C. larvae (mainly 3rd to 5th instar) were collected on a ski slope
during late June 1993. They were reared on clover at 25 in the laboratory (16L/
8D). All pupated and eclosed; 22 out of 37 were female. Immediately after emer-
gence, their abdomens were amputated and immersed in 50% ethyl alcohol. All
abdomens were dissected and examined for male spermatophores in the bursa
copulatrix. Eggs in the ovaries were also counted and classified into three groups
(mature, submature and immature), as has been done with the other pierid but-
terflies (Ando & Watanabe 1992, 1993, Watanabe & Ando 1993, 1994). Most statis-
tical comparisons were done with a Mann-Wliitney U-test, except for the Kendall
test on mating frequency in relation to age.

Results

Females immediately after emergence

No diseased individuals or parasitic wasps were noted in the 37 field cap-
tured larvae. Among the 22 females of the 37 emergences, the size was not
significantly different for the two morphs (Table 1). Females of both mor-
phs carried more than 750 immature eggs and 40 submature eggs. No ma-
ture eggs were found in either morph. Therefore, if no more immature eggs
were added during her life span, as in Pieris rapae (Watanabe & Ando 1993) ,
the fecundity of C. females would be about 800 for either morphs. Since
adult size and fecundity are largely dependent upon the quality and quan-
tity of food during the larval stage, we assume that each larva had consumed
similar quantities of food.

Of 474 albas captured in the field, 5 were virgin, while we found no vir-
gin yellow morphs. Such virgin albas were considered freshly eclosed. Table

35:1-8, 1996 (2000)

Table 1 . Fecundity and size for two morphs of female C. erate at emergence

reared in the laboratory, comparing those of field-captured alba having no
spermatophore (±SD).

Yellow

Alba

Field-captured
virgin alba

Number of females

Forewing length (mm)

29.8±L52

30.0±1.34

29.4±1.33

Number of immature eggs

794.2+93.57

766.8±41.87

688.8±1 88.82

Number of submature eggs

40.3±6.11

37.8±8.66

99.2±78.32

Number of mature eggs

O.OiO.OO

0.0±0.00

3.8±5.02

Total number of eggs

821.0±209.22

795.6±1 78.55

791.8±253.11

Table 2. Forewing length for two morphs of female C. erate captured in the field

(mm ± SD).

Age class

Alba

Yellow

Mann-Whitney

U-test

Monandrous I

29.5±0.97(139)

29.411.16(43)

U=96.5, n.s.

29.7±1 .59(79)

29.111.31(25)

U=86.0, n.s.

III

29.4±1 .94(48)

29.011.69(19)

U=16.0, n.s.

30.0±1.35(9)

29.112.86(4)

U=7.0, n.s.

30.2 (1)

28.810.77(3)

U^ll.O, n.s.

Polyandrous I

29.611.81(19)

30.311.44(5)

U=9.0, n.s.

29.511.38(31)

29.710.96(12)

U=43.5, n.s.

III

30.511.31(55)

29.911.49(30)

U=142.0, n.s.

29.611.51(48)

30.310.48(14)

U=18.0, n.s.

30.211.12(17)

29.610.10(5)

U=4.0, n.s.

( ):Samplesize

J. Res. Lepid.

Table 3. Frequency distributions of the number of spermatophores in the bursa
copulatrix of females in two morphs of C. erate captured in the field.

Morph Number of

Spermatophores

Age class

, I

III

143

Alba 2

Total

170

110

105

Mean

1.13

1.30

1.60

2.05^

2.2T

Yellow 2

Total

Mean

1.10

1.32

1.63

1.80^

Lyo"

Mann-Wliitney U-test, a:

U=818.0, P:

b: U=126.0, P=

-0.03

1 gives their fecundity. The number of immature and submature eggs per
female was not significantly different from reared females, although wild
females carried a few mature eggs.

Table 2 gives forewing length of field captured specimens. The difference
between morphs was not significant in each age class and body size was not
correlated with their age class. Watanabe and Nakanishi (1996) showed that
the population structure of this species was similar for each year in the same
study area.

Spermatophores in the bursa copulatrix of field-captured females

We dissected 474 alba and 166 yellow morphs in this study. No seasonal
effect on mating frequency was found, as in P. mpae (Watanabe & Ando
1993). Table 3 shows that the youngest alba (age class I) had a single sper-
matophore in the bursa copulatrix, while 5 were virgin, and 14 had been
polyandrous. The average number of matings was 1.1 for age class 1. The
number of matings for alba increased with age (Kendall Test, x=1.000,
P<0.01 ). The average number of matings in alba exceeded 2. Although the
number of matings for the yellow morph increased with age (Kendall Test,
1=0.800, P<0.05), their mating frequency was significantly lower than for
alba by age class 4 (U=818.0, P=0.05) and age class 5 (U=126.0, P=0.03).
Therefore, the yellow morph mated less than alba over their life (less than
2).

Figure 1 shows the change in the number of immature eggs with mating
frequency. Every female carried a decreased load of immature eggs within

35:1-8, 1996 (2000)

lOOO

lOO

IV'toxiatoxirous Polyaisdrotis

I ninivv I niHBfv

Age C-lass

Fig. 1 Changes in the number of immature eggs of respective age (I, II, III, IV,
and V) in relation to mating frequency in wild females of C. erate. Circles
and triangles indicate the data for yellow morph and alba, respectively.
Each bar represents SE. a and b are the results from Mann-Whitney U-
test for P=0.04 (U=10.0) and P=0.03 (U=58.0) respectively. Parenthe-
ses show one sample.

its age class. We never observed fused eggs in the ovaries, suggesting that
eggs were not consumed for somatic maintenance. If no immature eggs were
added during adult stage, the decreasing number of immature eggs assumed
due to oviposition.

For monandrous females, there was a significant difference in number
of immature eggs carried by age class 4 among alba and yellow morphs
(U=10.0, P^0.04) . Because virgin females revealed about 750 immature eggs,
about 550 eggs were laid by yellow morph and about 400 eggs by alba dur-
ing their life span. Thus when monandrous, a yellow morph female lays more
eggs than an alba morph.

In polyandrous females, alba lays more eggs than the yellow morph
(U=58.0, P-0.03, in the age class 5), with alba laying about 650 eggs and
the yellow morph laying about 550 eggs, a similar number to the mon-
androus yellow morph.

Discussion

Colias species are widely used for studies of butterfly biology (e.g,, fecun-
dity in Stern & Smith 1960; thermoregulation in Watt 1968, 1973), includ-
ing reports showing that pteridine or nitrogen pigments on the wings are
important for their reproductive success. Watt et al. (1989) showed that
population structure of Colias species co-existing with another pierid but-
terfly, Pieris napi, influenced the frequency distribution of wing color mor-

J. Res. tepid.

phs ill the Colias. Boggs and Watt (1981) also described the effect of mat-
ing frequency on population structure.

We have examined the biology of C. in Japan (population density in

Watanabe & Nakanishi 1996; courtship behavior in Watanabe et al. 1997),
in which the yellow morphs never outnumbered albas in any age class. Only
for C. scudderi, the alba outnumber yellow morph in North America (Gra-
ham et al. 1980), but no relationship between the fecundity and the mat-
ing frequency has been demonstrated.

Over a period of six years, we examined in excess of 700 females of C.
eratein our study area. Here the number of matings increased with age class,
though about half the females were captured young. Braby (1996) pointed
out that the mean number of spemiatophores correlated significantly with
age class (based on wing wear) in field females of bush brown butterflies,
Mycalesis spp.

Positive correlations between mating frequency and population density
have been noted in some butterfly species (e.g., Pliske 1973). Such studies
suggest that at high density competition among males for females becomes
more intense and the number of matings increases. Although Watanabe
and Ando (1993) showed for P. rapae that the number of active males search-
ing for mates differed between years, male density did not affect the num-
ber of matings by females. Thus females must exhibit mate choice (Rutowski
1978). For C. cra^calba, the increasing tendency in the number of matings
with age class was similar for P. rapae, while the yellow morphs were apt not
to re-mate at older ages (age class 3, 4, and 5) than albas.

There were a relatively few mature eggs and a small number of submature
eggs in the ovaries of young virgin alba, in which fecundity was estimated at
about 800. Although we have no data on the fecundity of virgin yellow
morphs from the field, laboratory populations indicated that females of both
morphs have similar fecundities. The immature eggs loaded in virgin fe-
males of C. eratewdiS the highest among other pierid butterflies inhabiting
the study area, P. rapae (Watanabe & Ando 1993) and P. melete (Ando &
Watanabe 1993).

Watanabe and Ando (1994) pointed out that monandrous females of P.
rapae laid fewer eggs than polyandrous females. Multiple spermatophores
have been shown to increase female reproductive output in some butterfly
species (e.g., Watanabe 1988, Wikhmd et al. 1993). However, in the yellow
morph of C. females, mating frequency did not correlate with the num-

ber of eggs laid during their life span. In other words, the yellow morph
does not need the extra spermatophores for oviposition, suggesting that
one mating provides a female with enough sperm to fertilize all of the eggs
(e.g., Suzuki 1978), and that spermatophores are not available energy for
egg production (e.g., Svard &: Wikhmd 1988). In fact, we observed small
but intact shaped single spermatophores in older yellow morphs. Since the
yellow morph might be mimetic to males to avoid further matings (unpub-
lished data) , they may have evolved an increased reproductive output with-
out multiple matings.

35:1-8, 1996 (2000)

Wataiiabe et al. (1997) showed that males persistently courted albas and
sometimes harassed copulating pairs involving an alba rather than an yel-
low morph in the field. In the present study, alba tended to re-mate and
polyandrous alba laid significantly more eggs than the monandrous females,
suggesting an increase of fitness as in the case of many butterfly species (e.g.,
Oberhauser 1989, Wiklund et al. 1993). Therefore, it is likely that alba ef-
fectively attract males in morphology to increase their reproductive out-
put.

Acknowledgments. We thank Dr W. Wehling (Michigan State University) for critical
reading of the manuscript. The manuscript was improved by suggestions from Dr
C.L. Boggs (Stanford University) and anonymous referees. S. Ando, K. Sato and
M. Taguchi assisted in field sun^ey. Financial support to M. Watanabe was provided
by a grant from the Inamori Foundation and by Grant-in-Aid from the Ministry of
Education, Science and Culture of Japan (No. 05640710).

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— — . 1993. Mating frequency and egg load in the white butterfly, Pieris melete
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Braby, M.F. 1996. Mating frequency in bush-brown butterflies (Nymphalidae:
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Brakefield, P.M. & T.G. Shreeve. 1992. Diversity within populations, pp. 178-196
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Emmel, T.C. 1972. Mate selection and balanced polymorphism in the tropical
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Gilchrist, G.W. 8c R.L. Rlitowski. 1986. Adaptive and incidental consequences of
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Graham, S.M., W.B. Watt & L.F. Gall .1980. Metabolic resource allocation vs. mating
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Ley, C. & W.B. Watt. 1989. Testing the ‘mimicry’ explanation for the Colias ‘alba’
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Oberhauser, K.S. 1989. Effects of spermatophores on male and female monarch
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J. Res. Lepid.

Pliske, T.E. 1973. Factors determining mating frequencies in some New World
butterflies and skippers. Ann.Ent.Soc.Am. 66:164-169.

Rutowski, R.L. 1978. The form and function of ascending flights in butterflies.

Behav.Ecol.Sociobiol. 3: 1 63-1 72.

Stern, V.M. & R.F. Smith. 1960. Factors affecting egg production and oviposition
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Suzuki, Y. 1978. Adult longevity and reproductive potential of the small cabbage
white, Pieris rapae crMaYora Boisduval (Lepidoptera: Pieridae). Appl. Ent. Zool.
13:312-313.

Sv.\RD, L, & C. WiKLUND. 1988. Fecundity, egg weight and longevity in relation to
multiple matings in females of the monarch butterfly. Behav. Ecol. SociobioL
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Uesugi, K. 1992. Temporal change in records of the mimetic Papilio poly tes

with establishments of its model Pachliopta aristolochiae in the Ryukyu Islands.
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Watanabe, M, 1979. Population size and resident ratios of the swallowtail butterfly,
Papilio polytes L., at a secondary bush community in Dharan, Nepal. Kontyu,
Tokyo 43:291-297.

■ . 1988. Midtiple matings increase the fecundity of the yellow swallowtail

butterfly, Papilio xuthus L., in summer generation. Journal of Insect Behavior
1:17-29.

Watanabe, M. & S. Ando. 1993. Influence of mating frequency on lifetime fecundity
in wild females of the small white Pieris rapae (Lepidoptera: Pieridae). Jpn. J.
Ent. 61:691-696.

-- — . 1994. Egg load in wild females of the small white Pieris rapae crucivora
(Lepidoptera, Pieridae) in relation to mating frequency. Jpn. J. Ent. 62:293-
297.

Watanabe, M. & Y. Nakanishi. 1996. Population structure and dispersals of the sulfur
butterfly Colias erate (Lepidoptera: Pieridae) in an isolated plain located in a
cool temperate zone of Japan. Jpn. J. Ent. 64:17-19.

Watanabe, M., Y. Nakanishi & M. Bonno. 1997. Prolonged copulation and
spermatophore size ejaculated in the sulfur butterfly, Colias erate
(Lepidoptera: Pieridae) under selective harassments of mate pairs by
conspecific lone males. J. Ethol. 15:45-54.

Watt, W.B. 1968. Adaptive significance of pigment polymorphisms in Colias
butterflies. 1. Variation of melanin pigment in relation to thermoregulation.
Evolution 22:437-458.

. 1973. Adaptive significance of pigment polymorphisms in Colias butterflies.

III. Progress in the study of the “alba” variant. Evolution 27:537-548.

Watt , W.B., C. Kremen & P. Carter 1989. Testing the ‘mimicry’ explanation for the
Colias ‘alba’ polymorphism: patterns of co-occurrence of Colias and Pierine
butterflies. Functional Ecology 3:193-199.

WiKLUND, C., A. Kaitala, V. Lindfors & j. Abenius 1993. Polyandry and its effect on
female reproduction in the green-veined white butterfly {Pieris napih.). Behav.
Ecol. SociobioL 33:25-33.

Journal of Research on the Lepidoptera

35:9-21, 1996 (2000)

Oviposition, host plant choice and survival of a grass feeding
butterfly, the Woodland Brown {Lopinga achine)
(Nymphalidae: Satyrinae)

Karl-Olof Bergman

University of Linkoping, Dept of Biology, S-581 83 Linkoping, Sweden, E-mail: karbe@ifm.liu.se

Abstract. Oviposition, host plant choice and sundval on different plants of
a grass-feeding butterfly, Lopinga achine, were studied in the field and in
the laboratory. Grass-feeding butterflies are generally thought to be non-
specific in their host plant choice. This seems not to be true for L. achine.
Females were selective in their host plant choice and preferred to oviposit
near Carex montana, although they do not attach their eggs to any plant.

Carex montana was also generally preferred by the larvae in laboratory
experiments among the plants available in the field. However, the larvae
preferred three species that they seldom encounter in the field {Agrostis
capillaris, Phleurn pratense and Poa pratensis) before C. montana when they
were offered these four species. Most of the larvae found in the field
(>80%) , were found on C. montana. The lan ae survived significantly better
on C. montana than on six other species in rearing experiments. The results
indicate that host plant choice occurs in two steps in L. achine. 1) the
females choose a patch to drop the egg to the ground, usually in the vicinity
of a C. montana plant 2) the newly hatched larva moves to the host plant.

The apparent dependence of the Swedish mainland L. ac/imc population
on a single host plant has important conservation implications.

Key WoRDSJ Lopinga achine, Satyrinae, host plant choice, performance, larvae,
conservation, Carex montana.

Introduction

Lopinga Scopoli (Nymphalidae: Satyrinae) is one of the threatened

Swedish butterflies that may disappear from the Swedish mainland without
conservation measures. The species is classified as endangered in three
European countries and as vulnerable in four (Heath 1981). It is one of the
few Swedish species on the Bern Convention list (Council of Europe 1993)
of endangered flora and fauna in Europe. The species is local throughout its
distribution area from the south of Fennoscandia through central Europe to
North and Central Asia and Japan (Kudrna 1986). In Sweden it lives in two
areas, in the province of Ostergotland, where I am studying it, and on the
island of Gotland in the Baltic (Henriksen & Kreutzer 1982).

Little is known about the host plant of L. achine. The female drops the eggs
to the ground and does not attach them to plants. Consequently it is difficult
to ascertain its host plants (Karlsson & Wiklund 1985). At least 15 species or
genera within Poaceae and Cyperaceae are suggested as host plants in the
literature (Nordstrom 1955, Henriksen 8c Kreutzer 1982; Karlsson & Wiklund

Paper submitted 27 May 1997; revised manuscript accepted 28 January 1999.

J. Res. Lepid.

1985; Ackery 1988; Lepidopterologeii-Aibeitsgruppe 1988;Jutzeler 1990; ).
Only three of the plants seem to be confirmed by larval findings in the field.
One larva was found on C«rcx<2/^<2Scop.andoneon C. montanah. (Cyperaceae)
(Ebert & Rennwald 1991) and lar\ae (numbers not stated) were also found
on Brachypodium sylvaticum (Huds.) PB (Poaceae) (Lepidopterologem
Arbeitsgruppe 1988).

Knowledge about host plants is still poor in many butterfly species,
especially in grass-feeding ones (Thomas 1984). Grass feeding butterflies are
generally thought to be unspecific in their choice of oviposition site (Wiklund
1984), but the studies of this are few in number. Our present knowledge of
host-plant choice and oviposition in butterflies is based primarily on studies
of Pieridae, Heliconiidae, Papilio^p^. and Euphydryas (e.g. Thompson
& Pellmyr 1991, Renwick & Chew 1994). Many butterfly species have been
shown to be more specific in their choice of habitat in the young stages than
first had been suspected. Therefore it is necessary to know the exact needs
of the immature stages to make conservation successful (Thomas 1984,
Thomas 1991, New et al. 1995).

The aim of this investigation is to study the host plant choice of Lopinga
achine and to determine its degree of specificity.

Materials and methods
Study animal and study site

L. achine f\y in one generation in June-July and hibernates in the larval stage. The
typical habitat in Ostergotland, where I study it, is partly open oak woodland ( Quercus
roburh. ) (Fagaceae) with hazel (Corylus avellanaE) (Corylaceae). This habitat is a
successional stage lasting 30-50 years before the canopy closes if not grazed. The
habitat on Gotland is different, being partly open coniferous forest with a well-
developed scrub layer of Frangula alnus Mill. (Rhamnaceae), Sorbus aucuparia L.
(Rosaceae), A intermedia (Ehrh.) Pers. and Juniperus communis E (Cupressaceae).
According to inventories up to 1997, L. achinelives in 49 populations in Ostergotland
in a small area (21 x 10km) and most of the populations have contact with each other
according to mark-recapture work. The matrix is usually open fields or spruce
plantations. Most populations are small, some hundreds of adults. Four populations
may comprise two to three thousand adults.

Oviposition

Ovipositing females were followed in the field in areas of high adult density. I used
binoculars to be able to observe the females from a distance in order not to disturb
them. Immediately after oviposition the exact place was marked and all Poaceae,
Juncaceae and Cyperaceae species within 15 cm were recorded. Plant names follow
Mossberg (1992). All females I saw ovipositing did so sitting on the vegetation. The
oviposition place is henceforth referred to as an “oviposition point”. The observa-
tions were mainly (48 out of 84 egglayings) made in the largest population and the
rest in nine other populations.

Plant species at randomly selected points were checked in the same manner as the
oviposition points. These “random points” were placed at approximately the same
distance from the edge (one meter zones) between forest and open areas as the
oviposition points to avoid vegetation differences due to influence from the forest.

35:9-21, 1996 (2000)

For example, a oviposition point 1.7 meters from the edge of the glade in the forest
has a corresponding random point between 1-2 meter from the edge.

Larval host-plant choice in the laboratory

The larval host-plant choice was tested in the laboratory using different grasses and
sedges (Table 1.) during the season when larvae normally feed. The laboratory
temperature varied between 22° and 25°C and there was limited daylight (50-80 lux)
from a small window above the petri dishes. Plant leaves were cut in 25 mm long
pieces and placed in a circle with the cut ends towards the centre of a petri dish (9
cm diameter). Each plant species was represented by one piece, except the thin
Deschampsia flexuosa (L.) Trin. (Poaceae) with several pieces in each place. Moist
filter paper covered the bottom in the dish. The leaves rested against a roll of paper
at the edge of the dish to prevent them from laying flat on the bottom. During all the
trials, only two larvae made no choice.

Eggs were collected from 20 females caught in the wild and kept together in a cage.
All of them laid eggs and the collected eggs was a mixture from these 20 females. A
newly hatched larva arising from each of these eggs was placed in the middle of each
petri dish with a fine brush. After 72 hours, the plant species were ranked according
to larval preferences: plants with the largest area eaten of was ranked as number one,
that with the second largest area eaten as number two and so on to the last one.

Host plant choice in the field

In the glades where I had found the largest numbers of flying adults, I systemati-
cally searched through every plant in the families Poaceae,Juncaceae and Cyperaceae
for larvae in a zone six meters out in the open glade and six meters under the tree
and bush cover. I noted the species upon which they were found. This was done in
four populations in the autumn (20. IX. 90-3. X. 90) and spring (22.V.91-6.VI.91).

Rearing experiments on different plant species

In 1989-90 I reared larvae on five putative hostplants: Calamagrostis arundinacea,
Carex montana, Deschampsia cespitosa, Melica nutans and Poa nemoralis. The larvae
originated from the eggs from the captured females mentioned earlier. The larvae
were reared outdoors in 18x 18x 18 cm cages with net sides. Each cage contained
10 larvae and the plants stood in water. The plants were changed every third day
for the first two weeks, and then weekly or when deteriorated. The cages were
moved into the laboratory in November and kept at 4°C until March. In February,
the larvae were offered pieces of the plants that had green shoots in the field.
Survival and weight (0.1 mg) were followed up to and including adult eclosion.
Pupae were not weighed in order not to disturb them so the last weighing before
weighing the adults was of mature larvae. The newly hatched larvae were too small
to be weighed individually so 57 were weighed together and the average was used as
a starting weight. Adults were weighed one day after eclosion.

In the second experiment (1991-92) , the larvae were reared in round plastic cages
10 cm high and 11 cm in diameter, with a net lid. The larvae came from 20 females
caught in the wild and kept individually and the offspring were mixed as evenly as
possible. Five larvae were reared in each cage. The plants roots were submerged in
water through a hole in the bottom of each cage. The plants were changed whenever
they showed signs of deterioration. Seven plants (Poaceae and Cyperaceae) were
tested: Agrostis capillarish,, Calamagrostis arundinacea, Carex montana, Dactylis glomerata,
Deschampsia cespitosa. Milium effusumC., Phleum pratenseh. Survival was followed up to

J. Res. Lepid.

Carex montana
Deschampsia flexuosa
Agrostis capillaris
Deschampsia cespitosa
Anthoxanthum odoratum
Carex pallescens
Poa pratensis

no grass
others

0 10 20

randomized points
oviposition points

frequency

Fig. 1. Grass and sedge species within 15 cm from oviposition points of Lopinga
achine and within 15 cm from randomly selected points (n=84 in both
cases). Plant species with less than five occurrences among the randomly
selected points are pooled as “others.” These species are also grouped as
“others” at the oviposition points.

and including adult eclosion. The entire experiment including hibernation was
conducted outdoors.

Statistics

All statistics were calculated using Statview 4.01 for Macintosh (Haycock et al.
1992).

Results

Oviposition

The plant frequency in the randomly chosen points was significantly
different from the frequency in the oviposition points (x^=35,7; p<0,0001;
df=8) (Fig. 1). Females preferred to oviposit near Carex montana. No female
oviposited at points lacking grasses or sedges although 23 of the 84 randomly
selected points lacked grasses and sedges. Therefore I excluded these 23
points and tested whether the frequency of C. montana differed between
oviposition points and the 61 randomly selected points with grasses and
sedges. The difference is significant (x^=7.0, p<0.01, df=l), 57 out of 84
females (68%) oviposited within 15 cm from C. montana, but it occurred at
only 28 of the 61 randomly selected points (46%) with grasses and sedges.

Larval host plant choice

The newly hatched larvae clearly preferred some plants to others in all
seven experiments (Table 1 ) . Carex montanav^-as, preferred in four of five trials

Table 1. Ranking of host plants by choice of newly hatched Lopinga achine larvae in seven laboratory experiments. Mean rank among
the plants are given in parenthesis and the differences between the plant species in each experiment were tested by the Friedman test

(tied P"vaiues). Empty places mean species not included in the experiment.

35:9-2T 1996 (2000)

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J. Res. Lepid.

where it occurred. However, A. capillaris, Phleum pratense and Poa pratensis
were preferred to C. montana in one trial. Other preferred species were M.
nutans and P. annua. Deschampsia flexuosa and D. cespitosa were generally
disliked. The larvae refused to eat i). completely.

Field observations of 97 lan^ae in the autumn confirmed the results of the
larval host plant choice experiments. Carex montana was used by 82 larvae
(85%) , and nine were found on the low preferred host, D. cespitosa. One larva
were found on each of P. pratensis , Festuca rubra L. (Poaceae), Luzula pilosa
(L.) Willd (Juncaceae) and an unidentified Poaceae. Two were found on
non-host material. The difference between C. montana frequency in oviposi-
tions (57 of 84) and larval occurrence (82 of 97) is significant (%^=7.0,
p=0.008, df=l).

The result was almost the same in the spring. C. montana was used by 71
larvae of 86 (83%) , and 10 used D. cespitosa. The other larvae were found on
non-host material or when pupating.

Rearing experiments

The larvae grew slowly until hibernation started in October in the third
instar (Fig. 2a) . They grew fast in spring (March-June) to the fourth (most of
the males) or fifth instar (most of the females) and pupation. The larvae were
left undisturbed and eating for a week before they were weighed in the spring
which is the reason for the apparent weight gain during hibernation. Adult
weights among females reared on different plant species differ significantly
(p=0.007, F=4.68; df^= 4, df =21, single-factor ANOVA). However, it is only
the groups reared on C. montana and D. cespitosa that differ in a post hoc test
(p=0.017, Scheffers F ). Male groups do not differ significantly from each
other (p=0.62, F=0.665, dfj=4, df,=34, single-factor ANOVA) (Fig. 2a).

There are significant differences in survival in the first experiment be-
tween larval groups feeding on C. montana and those feeding on D. cespitosa,
M. nutans and P. nemoralis {'Xp, p<0.02 three pairwise comparisons, df=l in
each comparison) but not between C. montana mid C. arundinacea (x^=2.92,
p=0.09, df=l ) (Fig. 2b) . Larval mortality was highest during the first 50 days
(July-August) and after hibernation. In the second rearing experiment, the
survival on C. montana\^2& significantly higher than on all the other species
(X^, p<0.02 six paiiivise comparisons, df=l in each comparison) (Fig. 3) . The
survival on D. cespitosamid C. arundinacea 'weiS low compared to the results in
the first rearing experiment. There is also a tendency for increased mortality
during the first days of this trial and again after hibernation but it is not as
clear as in the first trial.

Mean time to eclosion for males varied between 334 days (P. nemoralis) and
338 days (M. nutans) (Fig. 4). Mean time to eclosion for females varied
between 341 days on D. cespitosa and 348 days on C. montana. There are
significant differences between times to eclosion between females (p=0.006,
F=4.93, dfj=4, df2=21, single-factor ANOVA) but not for males (p=0.154,
F=1.79, df =4, df2=34, single-factor ANOVA).

Survival (%) § Weight (mg)

35:9-21, 1996 (2000)

2a. Mean weight increase of Lopinga achine larvae and mean of adult
weights at eclosion after development on five different grass species.
Measures of spread are omitted for clarity. The largest SEs are ±1 5,5% for
larvae and ±10,9% for adults. See Figure 2b number of larvae.

Time (months)

Figure 2b. Survival of Lopinga ac/i/ne larvae to adult butterflies on five different plant
species. The experiment was done 1 989-1 990 and started 8.VII outdoors
but hibernation took place in the laboratory November-March. n = number

of larvae.

J. Res. Lepid.

Time (months)

Figure 3. Survival of Lopinga achine larvae to adult butterflies on seven plant
species. The experiment was done outdoors and started 4. VIII. 1991.
Plant species in decreasing order of butterfly survival: Carex montana
(n=50), Phleum pratense (n=20), Agrostis capillaris (n^20), Dactylis
glomerata (n=30), Calamagrostis arundinacea (n-30), Deschampsia
cespitosa (n=30), Milium effusum (n=20). n ^ number of larvae reared.

Carex Calamagrostis Melica Deschampsia Poa

montana arundinacea nutans cespitosa nemoralis

Figure 4. Mean development time to eclosion (±S.E.) of Lopinga achine males and
females on five grass species, n = number of butterfly specimens.

35:9-21, 1996 (2000)

Discussion

It is clear that L. females prefer to oviposit near C. montana (Fig. 1),

even though they do not attach their eggs to that plant. This selective
oviposition behaviour contrasts to the suggestions of Wiklund (1984) that
the satyrines that do not attach their eggs do not bother much about were
they drop them. Polyphagy or superabundant host plants are suggested as
reasons for the behaviour (Wiklund 1984, Thompson & Pellmyr 1991).
Neither seems to be true for L. achine populations in Ostergotland when
looking at the selective host plant choice and the difference in survival on
different plants. It is important to note that C. montana is not superabundant
in large areas of the L. achine sites even though the species was the most
common grass species in the areas were the females oviposited. This indicates
that the females first make a habitat choice. After the habitat choice, the host
plant choice seem to occur in two steps in L. achine. 1 ) the females choose a
patch to drop the egg to the ground 2) the newly hatched larvae moves to the
host plant (only tested under laboratory conditions).

Lopinga achine females show the characteristic fluttering flight before
landing and oviposition (Porter 1992). This indicates that the female do not
drop the egg without regard to the environment. During this flight the
females may use shapes (Vaidya 1969, Stanton 1982), colour (Saxena &
Goyal 1978) and odour (Petersen 1954, Feeny et al. 1989) to locate host
plants. Wing fluttering increased in Papilio polyxenes in the presence of host
plant odours (Feeny etal. 1989). Many species also use contact stimuli (Chew
8c Robbins ( 1 984) and references therein ) before they oviposit. This does not
seem to be the case in L. achine females as they sometimes oviposited in a
tussock of C. montanav^h^n sitting on other plants growing together with C.
montana, for example Lathyrus linifolius (Reichard) Bassler (Fabaceae). The
search for ovipositing places may also involve microclimatic conditions
(Thomas et al. 1986, Petersen 1954) and levels of shade (Greatorex-Davies
et al. 1993).

In many species it is the female who selects host plant by her oviposition.
The newly hatched larvae cannot exercise host-plant preference in many
species, as they lack sufficient powers of movement to leave the plant on
which the eggs were laid (Singer 1971, Saxena 8c Goyal 1978, Ohsaki 1979,
Singer et al. 1 994) . However, the larva of L. achinemw^i make the final choice
itself since the female drops the egg to the ground, although near a host
plant. The larva is quite able to choose (Table Izx). It is also able to starve
longer than the newly hatched larva of Papilio machaon whose female glues
the egg to the host plant (Karlsson 8c Wiklund 1985). They stated that the
ability of the L. achine larvae to endure starvation may be regarded as an
adaptation to the females way of oviposition.

However, the plant species they can choose among are determined by the
egg laying females (Fig. 1). That is probably one reason why larvae mostly
occur on C. montana, even though they preferred A. capillaris, P. pratensemid
P. pratensis to C. montana in the experiments where all four species were

J. Res. Lepid.

offered (Table 1 ) . The larvae seldom encounter the three grasses in the field
except for A. capillaris (Fig. 1 ) , but this grass was often represented byjust one
or two leaves in the vicinity of the egg. Another reason for the significantly
higher lan^al occurrence on C. montana (83-85%) in the field compared to
ovipositions (68%), may be due to lower mortality of larvae on C. montana
compared to other species. The larva may also be able to move longer than
15 cm, the distance arbitrarily chosen when checking plant species at
oviposition points. About 10% of the larvae found in the field occurred on
D. cespitosa but the larvae rated this species low in the choice experiments
(Table 1). In the second rearing experiment it also caused high mortality
(Fig. 3). However, it relatively often occurred at the oviposition points (Fig.
1 ) . Its tussocks are large so it may be difficult to leave it if the egg hatches in
the tussock. The development to adult may succeed on it (Fig. 2b, 4).

The developmental time does not seem to be important for host-plant
choice in L. achine. The females on the preferred species, C. montana, had the
longest time to adulthood (Fig. 4) . Development time can be important if the
time available for larval growth is limited (Nylin 1988).

Summarising, L. acAmc larvae survived and succeeded best on C. montana,
and the plant species was also preferred by egg-la)4ng females and newly
hatched larvae in choice experiment among the plants available in the field
(Table 1 , Fig. 1 , 2a, b, 3) . The good correspondence between oviposition and
performance in L. acAmc indicate that the generalist behaviour suggested for
satyrines (Bink 1985) does not seem to be true for L. achine. The correlation
between oviposition preference and performance in phytophagous insects
varies much. Many studies have reported a good correspondence (e.g. Papaj
& Rausher 1987, Nylin & Janz 1993) but many have also reported low
correlations (e.g. Courtney 1981, Rausher 1979, Larsson & Strong 1992).

Carex montana is probably also the host plant for L. achine populations on
the island of Gotland. Carex montana is very common in the woods that are
habitat for L. achineXh^YG. Lopinga achinel^rv3.e from Gotland survived better
on C. montanathmi on D. glomeratav^hy E^rlssoii 8c Wiklund (1985) suggested
C. montana^s the major host plant. However, C. montana c^nnothe the single
host plant for L. achine since the populations in Finland occur in areas
without it (Hulten 8c Fries 1986). L. achine 3\so completed the life cycle on
many of the other plant species in my experiments, even though the success
rate was lower. Different populations of butterflies may evolve different host
plant preferences as in the satyrid Satyrodes eurydice]ohm\s^on (Shapiro 1974)
and the nym^\\?didEuphydryaseditha^o\sd\\\2d (Singer etal. 1994) , especially
when living in different habitats.

The dependence of at least the Swedish mainland L. ar/imcpopulations, on
a single host plant has important conservation implications. In the future,
the relations between the butterfly and the host plant may be studied to
understand the reasons for the geographic distribution and to determine if
there is a need for habitat management to ensure long-term survival.

35:9-21, 1996 (2000)

Acknowledgements. I thankjan Landiii, Niklasjanz and Soren Nylin for comments and
Ulrika Hjelni for doing one of the lar\’al choice experiments. This work was
supported by grants from the World Wide Fund for Nature (WWF) and the Swedish
Environmental Protection Agency.

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Journal of Research on the Lepidoptera

35:22-28, 1996 (2000)

The effect of environmental conditions on mating activity of
the Buckeye butterfly, Precis coenia

Alice K. McDonald and H. Frederik Nijhout

Department of Zoology, Evolution, Ecology and Organismal Biology Group, Duke University,
Durham, NC 27708-0325. E-mail: hfn@acpub.duke.edu

Abstract. The readiness of males of Precis coenia to court females depends
on time of day, temperature, and light level. Courtship activity has a tem-
perature optimum and, at that optimum, increasing light level dramati-
cally enhances courtship activity. High light level appears to be critical
for courtship activity, and high temperature cannot substitute for high
light level. The requirement for high light intensities may be related to
the behavior of males that chase females from preferred territorial
perches on bright patches of exposed substrate.

Key Words: light level, temperature, courtship. Precis coenia

Introduction

Some years ago, when we first started to raise Buckeye butterflies {Precis
coenia Hubner) for our experiments on the development of wing patterns
(Nijhout, 1991), we encountered considerable difficulties in getting this
species to mate in cages in the laboratory. Our early experimental work was
done with animals produced from eggs of gravid females caught in the field.
None of the standard measures to enhance mating in the laboratory such
as confining the adults with various species of host plants, enlarging the
mating cage, and modifying temperature and humidity, appeared to pro-
vide the necessary conditions to induce courtship and mating.

Our observations of mating activity in the field confirmed the reports by
Scott (1973, 1975a,b) that courtship and mating are most frequently ob-
served during the warmest part of the day. At these times male buckeyes
perch preferentially on bright patches of sand and from those perches chase
passing females. We attempted to mimic these conditions by placing our
mating cages in front of a south-facing window at mid-day, and found that
this immediately induced courtship behavior in males. We found that P.
coenia v^owXd mate readily and successfully even in cages as small as 10 x 10
X 10 cm as long as they were placed in direct sunlight. It was not clear, how-
ever, whether the increased light level or the increased temperature in the
mating cage was the primary stimulus for courtship and mating. Below we
present an analysis of the independent effects of temperature and light level
on mating activity.

Materials and Methods

Larvae of Precis coenia were reared in the laboratory on artificial diet and long
Paper submitted 25 May 1997; revised manuscript accepted 14 April 1998.

35:22-28, 1996 (2000)

day (16L: 8D with the lights-on signal at 6 am EST) conditions at 27 °C. Our labo-
ratory colony was derived from animals collected in the Sandhills of North Caro-
lina (Sandhills Wildlife Management Area, Richmond County). Freshly emerged
adult butterflies were sexed by using characters of the prothoracic legs, and labeled
on the ventral hind wing with a permanent fine tip laboratory marker. Males and
females were separated and animals of each sex were grouped in separate Plexiglas
cages measuring 45 x 50 x 60 cm.

Observations were made during the months of June and July 1996. For each ob-
servation session, 10 males and 5 females aged between 4 to 8 days after emergence
were randomly selected from the holding cages and transferred to an observation
cage (Plexiglas, 25 x 30 x 25 cm, except for some observations in the greenhouse
when a wire mesh cage was used, as noted below). The data presented below are
based on 6 to 10 obsen^ations sessions under each set of environmental conditions.
Observation sessions were 45 minutes long. Preceding each observation session the
mating cage was placed into the test environment for a 30 minute equilibration
period.

Observations were made at three times of day (10 am, 12 pm, 3 pm EST) , at three
different temperatures, and at three different light levels. Target temperature val-
ues were 25 °C, 33 °C, and 40 °C. Due to uncontrollable drafts and imprecise tem-
perature regulation in the greenhouse actual temperature values around these tar-
gets ranged from 23.0-28.8 °C, 30.0-38.0 °C, and 38.0-42.6 °C during the observa-
tion period. Target light level values were 2.3, 53, and 280 lux, with actual values
ranging from 2. 1-2.6, 49-56, and 100-430 lux around those targets due to variation
in solar irradiance and physical setup of the mating cages. Precision Instruments
incubators were used to provide the 33 and 40 °C temperatures under low and in-
termediate light intensities. Light level of 2.3 lux was provided by a 15 watt incan-
descent bulb, while values of 53 lux were provided by a 500 watt halogen bulb. Tem-
perature was measured with a Yellow Springs Instmment Co., Inc. Tele-thermom-
eter Model 46TUC. Light levels were measured with a Weston Illumination Meter,
Model 756. To enable observation while maintaining the necessary temperatures,
the door frame of the incubator was covered with a clear plastic sheet secured with
magnets. A clear plastic tray containing one inch of a 10% CuSO^ solution was placed
between the cage and halogen lamp (about 6 inches from the light) as a heat ab-
sorbing filter. To achieve low and medium light intensities at 25 °C, we used the
same lights in a temperature controlled room. The highest light level we used, 280
lux, is representative of the level of sunlight. It was impossible to achieve this level
with artificial lights, so measures at high light intensities were made in a climate-
controlled greenhouse. In the center of the greenhouse temperatures fluctuated
between 32-34 °C. A wire mesh cage (45 x 45 x 45 cm) was used for observations
centered around 33 °C. By placing the wire mesh cage near the cooling cells of the
greenhouse, a temperature of 25 °C could be accurately maintained. To maintain
40 °C, a Plexiglas cage was placed in a sunny location; sliding vents were used to
manually adjust the temperature inside the cage.

Assay of Mating Behaviors. Courtship and mating behavior in Precis in captivity
consist of four distinctive behaviors: nudging, chasing, head dipping, and abdo-
men curling. These differ slightly from the general nymphalid courtship behavior

J. Res. Lepid.

Figure 1. Mean frequency of individual courtship behaviors at each of different

times of day, at the intermediate temperature of 33 °C and the high light

level of 280 lux.

patterns described by Scott ( 1975b) . Nudging is defined as the brushing of the legs
of a hovering male butterfly against the wings of a perched butterfly. Often, a nudg-
ing male will land on or next to the perched butterfly. Males do not appear to dis-
tinguish between males or females at this stage in the courtship. Chasing is the
pursuit, on foot, of another butterfly. Head dipping is defined as the dipping of
the head under the abdomen or wing area surrounding the abdomen of the ap-
proached butterfly. This seems to serve to raise the female’s abdomen into a posi-
tion that the male can easily couple with. Abdomen curling consists of lateral curl-
ing of the male’s abdomen towards another butterfly in an attempt to copulate.
The frequencies of each of these behaviors during each 45 min observation period
was scored in order to obtain a quantitative estimate of the effects of the three en-
vironmental variables on mating activity.

Results

Mating activity consistently reached a maximum at intennediate tempera-
tures and high light intensities. Frequencies of individual mating behaviors
under these conditions are shown in Figure 1. The proportion of different
acts was not significantly different at 10 am and 12 noon (Chi-square,
P>0.95), while the proportions of the different acts at 3 pm differed signifi-
cantly from those at the two earlier times (Chi-square, P<0.0003). This dif-
ference appears to be accounted for entirely by a decrease in the frequency
of nudging. At 3 pm the nudging events were significantly less frequent per
unit time than they were earlier in the day (t-test, P<0.0033), whereas the
frequencies of the intermediate and later courtship events did not differ
significantly from their frequencies earlier in the day. This observation sug-
gests that in the afternoon courtship becomes more efficient in the sense

35:22-28, 1996 (2000)

10 am 12 noon 3 pm

Figure 2. Mean frequency of total courtship activity at three different times of day
and three different light intesities (all data shown is at 25 °C).

that a larger proportion of individuals that begin courtship are able to take
it to completion.

High light intensity dramatically increased mating activity at all times of
day (Figure 2). At 10 am and 12 noon a stepwise increase in light intensity
signiflcandy increased mating acti\dty (P<0.003 for all pairwise comparisons) .
At 3 pm mating activity at low and medium light intensities were not sig-
nificantly different from each other but mating activity increased signifi-
cantly at the highest light intensity (P<0.0004).

The Combined Response to Temperature and Light Level. The overall
mating activity was assessed as the sum of all observed courtship events.
These are graphed in Figure 3 as a function of both temperature and light
level at three different times of day. Contours were calculated using
SigmaPlot (Jandel Co.). In view of the fact that the numerical tally of mat-
ing activity is dominated by early courtship events (nudging, see Figure 1),
the overall mating activity was also estimated based on the intermediate and
late courtship events alone (Figure 4). Both measures of mating activity
revealed a qualitatively similar pattern. At 10 am, mating activity increased
when temperature and light level increased. The effect of increasing tem-
perature was more pronounced at high light intensities than at low light
intensities, while the effect of light level was most pronounced at interme-
diate to high temperatures (Figures 3a, 4a). At 12 pm and 3 pm, by con-
trast, there was a distinct optimal temperature for mating activity (32-34
°C; Figures 3b,c, 4b, c). At 12 pm, increasing light level had a stimulatory
effect on mating activity at optimal temperatures and below, whereas at tem-
peratures above the optimum, light level had little effect on mating activity
(Figure 4b). At 3 pm, light level affected mating behavior only within the

J. Res. Lepid.

10 am 12pm ^ 3pm

T ^ 1 — I ^ 1 r

26 28 30 32 34 36 38 40 26 28 30 32 34 36 38 40 26 28 30 32 34 36 38 40

Temperature ("C)

Figure 3. Mating activity at different times of day as a function of both tempera-
ture and light level. Mating activity (numbers on contours) was scored
as the sum of the frequencies of all 4 of the courtship behaviors (Fig. 1)
during a 45 min observation period.

10 am 12 pm ^ 3 pm

Temperature (°C)

Figure 4. Mating activity at different times of day as a function of both tempera-
ture and light level. Mating activity (numbers on contours) was scored
as the sum of the frequencies of the three late stages in courtship (chas-
ing, head dipping, abdomen curling) during a 45 min observation period.

optimal temperature range, whereas light level had little or no effect on
mating activity at temperatures below or above the optimum (Figure 4c).

Analysis of significance of pairwise comparisons of the data presented in
Figures 3 and 4 revealed the following. In Figure 3, pairs of points that dif-
fer by more than 55 events (approximately 3 contour intervals) are signifi-
cantly different from each other (t-test, P<0.05), both within and between
panels. In Figure 4, pairs of points that differ by more than 26 events (slightly
greater than 1 contour interval) are significantly different from each other
(t-test, P<0.05), both within and between panels.

Discussion

The mechanisms that regulate mating behavior result in dramatically dif-
ferent responses to temperature and light level as time of day progresses.

35:22-28, 1996 (2000)

Maximum mating activity always coincided with temperatures of 32-34 °C
and high light level (280 lux), regardless of time of day. The relative fre-
quency of nudging, the earliest event in courtship behavior, peaked at 12
pm, while the relative frequency of abdomen curling, the final event pre-
ceding copulation, was greatest at 3 pm. Therefore, although total court-
ship activity at noon was greater than at 3 pm, the final stages of courtship,
and presumably mating success, were relatively more frequent at 3 pm.

Rutowski (1991 ) has outlined three hypotheses to explain why males court
preferentially at certain times of day. First, limited thermoregulatory capacity
may restrict mating activity to periods when the environmental tempera-
tures are neither too high nor too low. Second, mating may be timed to
coincide with female emergence times. Third, mating may be timed so as
to minimize interference between species. Our results illuminate the first
of these hypotheses, but show that mating activity is not constrained strictly
by temperature. Although male Precis coenia clearly have an optimal mating
temperature of 32-34 °C, exposure to this temperature alone did not re-
sult in maximum levels of mating activity. Mating activity at 12 pm and 3
pm was consistently higher than at 10 am. Time of day, therefore, affects
mating behavior independently of temperature. Light level also has an in-
dependent effect on mating behavior. Within the optimal temperature
range, high light level increases mating activity at all three times of day,
whereas at non-optimal temperatures the effect of light level depended on
the time of day.

Scott (1975b) has noted that in nature, mating activity of Precis coenia oc-
curs mostly in late morning and early afternoon. Our results suggest that,
given the right combination of temperature and light level, mating behav-
ior can occur at most times of day, although the interaction of light level,
temperature, and time of day ensure that the bulk of mating activity is most
likely to occur in the early afternoon.

If light level is low, little mating activity occurs, even at optimal and higher
temperatures. It is not clear at present why light level should have such a
great effect on mating activity. It is possible that high light level acts indi-
rectly, by elevating the male’s body temperature. This would imply that the
optimal body temperature is substantially higher than the 32-34 °C opti-
mal environmental temperature we measured. Optimal body temperatures
for flight in insects range from 35 to 42 °C (Heinrich, 1993), so it is con-
ceivable that mating activity could also require such high body tempera-
tures. However, if the effect of light level was mediated through an eleva-
tion of body temperature, one would expect high light intensities to be more
effective at inducing mating activity at temperatures below the environmen-
tal optimum than at temperature above the optimum. The apparent tem-
perature optima in Figures 3 and 4 would then be expected to be a func-
tion of light level, with a lower temperature optimum at high light intensi-
ties and a higher temperature optimum at low light levels. Instead, the tem-
perature optima are unaffected by light level (except for a single instance
at low temperatures: Figure 4b) , suggesting that these two environmental

J. Res. Lepid.

variables do not interact, and that light level seems to be important for rea-
sons other than radiational heating.

One explanation for the evolution of high light level as a cue for mating
may be found in the fact that mating activity of the Precis coenia population
we studied occurs preferentially in open habitats on exposed patches of
sand. Such bright areas in the landscape serve as perching territories for
males (Scott, 1975b, and personal observations), and from these territories
males chase passing insects, including females and other males. Males are
chased away, and females, if receptive, land nearby and courtship begins
(Scott, 1975b). It is possible that the selection of perching territories is
guided primarily by brightness of the substrate. If the acquisition of such a
territory is important for mating success, then it seems reasonable to sup-
pose that a response to high light levels as a stimulus for courtship may have
evolved in association with the behavior by which males select especially
bright perching territories from which to chase passing females.

Acknowledgments. We are grateful to Laura Grunert and Armin Moczek for their
helpful comments on the manuscript, and to the North Carolina Wildlife Commis-
sion for permission to collect in the Sandhills Wildlife Management Area. This work
was supported in part by grants from the Howard Hughes Foundation and the
National Science Foundation

Literature Cited

Heinrich, B. 1993. The Hot-Blooded Insects. Harvard Univ. Press, Cambridge, MA.
Nijhout, H. F. 1991. The Development and Evolution of Butterfly Wing Patterns.

Smithsonian Institution Press, Washington, DC.

Rhtowski, R. L. 1991. The evolution of male mate-locating behavior in butterflies.
Amer. Nat. 138: 1121- 1139.

Scott, J. A. 1973. Mating of butterflies. J. Res. Lepid. 1: 99-127.

. 1975a. Movements of Precis coenia, a ‘pseudoterritoriar submigrant. J. Anim.

Ecol. 44: 843-850.

. 1975Z>. Variability of courtship of the buckeye butterfly, Precis coenia

(Nymphalidae). J. Res. Lepid. 14: 142-147.

Journal of Etsearch on the Lepidoptera

35:29-41, 1996(2000)

Nymphalid butterfly communities in an amazonian forest
fragment

Frederico Araujo Ramos

Departameiito de Biociencias, Centro Universitario de Brasilia, SEPN 707/907, 70.790-07,
Brasilia, Brazil, E-mail: fmmos@tba.com.br

Abstract. Species diversity and abundance of fruit-feeding nymphalid
butterflies were studied in an Amazon rain forest fragment. Butterflies
were caught in baited traps in twelve areas, selected to sample a gradient
of increasing disturbance. Measurements of six parameters of vegetation
structure were also taken to estimate the disturbance. A total of 90 but-
terfly species were trapped. The greatest alpha diversities were found at
the edge of the forest and in areas of intermediate disturbance. Canoni-
cal Correlation Analysis (CCA) showed that the composition of the spe-
cies assemblages of nymphalids was related to vegetation structure vari-
ables, especially girth at breast height and number of tree morpho-spe-
cies. The butterfly fauna appeared more similar in forested areas than in
the disturbed ones. Some species were suggested as habitat indicators and
the value of this guild of fruit-feeding butterflies in conservation programs
is discussed.

Keywords: Butterflies, nymphalids, diversity, community structure, distur-
bance, rain forest fragment, direct gradient analysis, Brazil.

Introduction

One of the main objectives of community ecology is the synthesis of the
roles of physical and biological factors that determine species abundance
and distribution within and among natural communities. After MacArthur
and MacArthur (1961) found a relationship between bird diversity and high
vegetation diversity, ecologists verified that habitat complexity is an impor-
tant factor for the structuring of local communities. Habitats that are struc-
turally more complex and heterogeneous offer more niches, and therefore
support a greater number of species (spatial heterogeneity, Pianka 1966).
In addition this idea, Connell (1978) suggested that high diversity in tropi-
cal forests is maintained by disturbances, such as tree falls. Considering such
dynamics, the forest can be seen as a mosaic of gaps in different succes-
sional stages, with different local communities, and a high regional diver-
sity.

Although biogeographic and historical conditions are extremely impor-
tant factors in structuring communities (Slansky 1972, Ricklefs 1987, Leps
and Spitzer 1990, Brown 1982, Brown 1991, Thomas 1991, Gaston 1996),
local factors also affect local butterfly diversity (Emmel and Leek 1969,
Montesinos 1985, DeVries 1994, Kitahara and Fujii 1994, Sparks and Par-

Paper submitted 7 July 1997; revised manuscript accepted 25 February 1999.

J. Res. Lepid.

ish 1995). Many studies have shown that tropical butterfly communities also
respond to physical factors of the habitat, such as topography, stratification,
gaps, edges, urbanization and habitat disturbances (Riiszczyk 1986, DeVries
1988, Raguso and Llorente-Boiisquets 1990, Brown 1991, Hill et al. 1992,
Pinheiro and Ortiz, 1992, Hill et al. 1995, Spitzer et al. 1997). As such,
multivariate analysis has proven to be an important tool when investigating
the relationships between species assemblages and environmental variables
(Leps and Sptizer 1990, Kremen 1992, Ramos 1992, Vaisanen 1992, Spitzer
et al. 1993, Spitzer et al. 1997, Blair and Launer 1997).

The objectives of this study are (1) to measure neotropical nymphalid
butterfly diversity along a gradient of disturbance, (2) to explore the spe-
cies-environment relationships through a direct gradient analysis and (3)
identify the most important butterfly species and vegetation variables, which
could be used in conservation monitoring programs.

Methods

Study site. This study was conducted in a forest fragment at the boundary of the
eastern Amazon (5°0TS, 47°32'W; 260 m), a region where the natural landscape
has been greatly modified by human activity. The study site was about 50 km north
of the transition to Cerrado. The fragment has about 1,000 ha of primary forest
with several levels of disturbance, surrounded by secondary forest in several
succesional stages, eucalyptus monocultures and cattle pasture. I selected 12 sample
units (SUs) throughout a disturbance gradient: forest understory (FUl, FU2, FU3),
forest roads (FRl, FR2, FR3), edge (EDG), highly disturbed forest understory
(DFU), highly disturbed forest road (DFR), 4-year-old secondary forest (SF4), 2-
year-old secondary forests (SF2) ?ix\d Eucalyptus pellita monocwliuYe (EUC).

Data collection. I made lepidoptera collections between June 1990 andjuly 1991.
For each of 12 SUs, three fruit-baited traps were set in line, suspended 1.0-1. 7 meters
above the ground, and 25 meters apart from one another. For each collection, the
traps were visited for 14 consecutive days. The banana and sugar cane bait was kept
moist for the duration of the trapping period. The disturbance level of each SU
was estimated using vegetation parameters obtained through the point-centered
method (Miiller-Dumbois and Ellemberg 1974), with 21 quartered points estab-
lished per SU, only for trees up to 20 cm of circumference at breast high. This
method was chosen for its simplicity and common use in phytosociological surveys.
The following vegetation variables were used: average girth at breast flight (GBH);
estimated average tree height (THG); number of tree morpho-species (NMS) estimated
by rind and leaf characteristics, with the help of a local guide; tree density Wixhin 100
m- (DEN); average horizontal cover (HOC), estimated at each sample point by an
observation made on a 50 cm square carton held 10 m from the observer in each
quarter. Cover was estimated to be within one of four categories (0-25%, 25-50%,
50-75% and 75-100% vegetation cover); average vertical cover (VEC), estimated by
the four previously mentioned vegetation cover categories, apllying a 10 cm square
frame held at a distance of 60 cm from the observer at an angle of approximately
20° in relation to zenith.

35:29-41, 1996(2000)

Table 1 . Alpha diversity of fruit-feeding butterfly species in twelve sample units
of an Amazonian forest fragment.

Forest Forest Edge Disturbed Secondarv’ Eucalyp

understor)' road forest forest monoc

FUl

FU2

FU3

FRl

FR2

FR3

EDG

DFU

DFR

SF4

SF2

EUC

Total

Number of individuals

(N) 63

114

111

106

334

267

571

490

744

604

3544

Nymphalinae richness

Satyrinae richness

Brassolinae richness

Charaxinae richness

Total species richness

(S) 18

Species diversity (H')

2.27

2.76

2.60

2.79

3.08

3.06

3.38

3.06

3.19

3.00

2.28

2.09

3.26

Data analysis. The butterfly alpha diversity of each SU was quantified by the spe-
cies richness (S) and Shannon-Wiener index (H’). To evaluate environmental ef-
fects on the butterfly community I ran a Canonical Correspondence Analysis (CCA) ,
using the program CANOCO (TerBraak 1988). The vegetation parameter estimates
of habitat disturbance were used as environmental variables in the CCA. The vari-
able tree height was removed from the analysis due to its high value of inflation,
and high colinearity with the other variables. The significance of species-environ-
ment relationships was tested using a Monte Carlo test.

Results

A total sample effort of 2,016 trap days (=3 traps x 12 SUs x 56 days)
resulted in 3,544 individuals collected, representing 90 species of five
subfamilies of Nymphalidae. The five most abundant species were
Paryphthimoides phronius, Yphthimoides spl, Yphthimoides disaffecta,
Hermeuptychia hermesRucl Cissia penelope, all belonging to the subfamily
Satyrinae, representing 45.3% of the total number of individuals col-
lected. A complete list of species abundances in each SU can be seen
in Appendix 1. The total butterfly diversity in the rain forest frag-
ment sampled was H" = 3,258.

An analysis of alpha diversity showed that edges and areas of intermedi-
ate disturbance presented higher species richness and diversity (Table 1).
Although more disturbed areas, such as eucalyptus monoculture and 2-year-
old secondary forest had higher species richnesses than forest, they had
lower species diversities, due to the high dominance of the Satyrinae spe-
cies. This pattern was not found when other groups were considered sepa-
rately: Satyrinae and Brassolinae had higher species richnesses in the dis-
turbed areas, with a peak of the edge; Charaxinae and Nymphalinae had
higher species richnesses in the road, edge and disturbed forest areas.

The CCA ordination diagram shows the relationships between butterfly
species, sample units and environmental variables (Figure 1). By compar-
ing the arrow lengths, one may evaluate the significance of the constrain-
ing vegetation variables. The arrow points roughly in the direction of the
maximum variation in the value of the corresponding variable. The spe-

J. Res. Lepid.

Figure 1. CCA ordination of the fruit-feeding butterfly communities in an Amazo-
nian forest fragment, with respect to five vegetation variables (arrows).
The species are abreviated according to the first three letters of their
generic and the first three letters of their specific names (see Appendix
1 for full names). The vegetation variables are: average girth at breast
height (GBH), estimated average tree height (THG), number of tree
morpho-species (NMS), tree density within 100 m2(DEN), estimated
average horizontal cover (HOC) and estimated average vertical cover
(VEC). The sample units are forest understory (FLU, FU2, FU3), forest
roads (FBI , FR2, FR3), edge (EDG), highly disturbed forest understory
(DFU), highly disturbed forest road (DFR), 4-year-old secondary forest
(SF4), 2-year-old secondary forests (SF2) and Eucalyptus pellita monoc-
ulture (EUC).

cies far from the origin are rare and less important to analysis. The Monte
Carlo permutation test showed that those species are significantly related
to supplied vegetation variables (99 permutations, P < 0.01).

In the first CCA axis, girth at breast height presented the highest abso-
lute value, followed by the number of tree morpho-species, both contribut-
ing to the species data fit. This axis clearly shows a gradient of disturbance
— preserved areas with a greater richness of large bole trees on the posi-
tive side, and disturbed or early succesional ones on the negative side. The

35:29-41, 1996(2000)

ordination also shows how species respond to vegetation variables: with in-
vader species typical of open areas (the small Satyriiiae Hermeuptychia
hermes, Yphthimoides spl, Y. disaffecta, Cissia penelope, Erichtodes numeria^nd
Pharneuptychia pharnaces, and the Nymphalinae Hamadryas feronia, H.
februa and Biblis hyperia) showing negative scores. With positive scores,
near the origin, are the heliophyllous species of the disturbed forests, gaps,
edges and canopy (Eryphanis polyxena, Hamadryas iphthime, H. velutina, Mem-
phis morvus, Narope cyllabarus, Nica flavilla, Pareuptychia ocirrhoe, Temenis laothoe
and Taygetes laches). The species on the right side of the diagram are typical
of the forest understory ( Colobura dirce, Morpho achiles, Nessaea obrinus, Taygetis
celia, T. echo and T. virgilia) . The second axis was primarily related to tree
density and horizontal cover, but did not form a clear gradient. The aiialy-
sis gave a large weight to some Satyrinae species, such as Cissia penelope and
Paryphthimoides phronius, which had large populations in high tree density
eucalyptus monoculture.

Discussion

The forest edge and intermediate disturbance forest presented higher
values of butterfly species richness and diversity. These environments, where
intense regeneration occurs, have high productivity and maintain high
population levels. On the other hand, the disturbance rate is high, thus
reducing the effect of competitive exclusion. A number of ecologically based
hypotheses have been proposed to explain patterns of species richness and
diversity, but not all of them are mutually exclusive (Meffe and Carroll 1997).
The productivity-disturbance hypothesis (Huston 1994) combines elements
of several other hypotheses, proposing that the high productivity and the
disturbance rate conditions of forest edges and gaps result in high species
richness. The results of this study tend to agree with this hypothesis. Addi-
tionally, the mixture of forest imderstory umbrophyllous species with open
area heliophyllous ones raises the local diversity. The fact that edges, gaps,
physiognomic transitions, and disturbed and secondary forests have high
diversity has been documented many times elsewhere (Leps and Spitzer
1990, Raguso and Llorente-Bousquets 1990, Hill et al. 1992, Pinheiro and
Ortiz 1992, 'Vainsanen 1992, Spitzer et ah 1993, Spitzer et al. 1997), For
sunloving species, drastic changes in light intensity can act as a habitat bar-
rier, while edges, gaps and canopies may be treated as a continuum of sunny,
open area. Other species, however, remain restricted to shady environments
(DeVries 1988).

The number of tree morpho-species was not a good predictor of the num-
ber of butterfly species. Because the group is herbivorous, butterfly - host
plant relationships have been explored (Gilbert and Smiley 1978, Erhardt
and Thomas 1991), although not always being meaningful (Sharp et al. 1974,
Courtney and Chew 1987, Singer and Ehrlich 1991). Besides that, the sam-
pling considered only trees, and did not consider bushes, herbs and lianas
that are host plants of several butterflies (DeVries 1987). Although not re-
lated with butterfly alpha diversity, the number of tree morpho-species was

j. Res. Lepid.

important in the fonnation of an environmental gradient, and is related to
habitat disturbance.

The ordination diagram shows that the forest assemblages are homoge-
neous, but among disturbed areas there are great variations in species com-
position and abundance. A larger constancy of forest communities has been
verified for neotropical satyrins (Brown 1991), and butterflies of South-east-
ern Asia (Leps and Spitzer 1990), but the opposite pattern was found for
Notodontidae and Arctiinae in the same forest fragment (Dubois 1993).

Although other butterfly groups such as Ithominae and Heliconinae have
been suggested as more efficient indicators (Brown 1991, Beccaloni and
Gaston 1995), the use of the fruit-feeding Nymphalidae has its utility in
conservation programs (Daily and Ehrlich 1995). The results of the CCA
ordination show that butterfly communities have a significant relationship
with vegetation variables, and suggest the use of this assemblage as an ap-
propriate indicator of habitat heterogeneity over this spatial scale. CCA can
be used to match a species assemblage to environmental factors for which
it is a good indicator, and select a subset of species as indicators for more
intensive monitoring (Kremen 1992). Since rare species have little weight
in the analysis (TerBraak 1988), common species, and not rare ones, should
be selected from this guild to be used as indicators. Thus, Hermeuptychia
hermes, Yphthimoides spl, Y. disaffecta, Cissia penelope, Erichthodes numeria,
Pharneuptychia pharnaces, Hamadryas feronia, H. februa and Biblis hyperia may
serve as indicators of disturbed environments, and Colobura dirce, Morpho
achiles, Nessaea obrinus, Taygetis celia, T. cc/^oand T. indicators of more

preser\'ed environments.

The use of higher taxa for biodiversity measurements (Williams and
Gaston 1994) can be an important management tool for situations where
taxonomic identification at the species level is difficult. For the same data
set, counting only subfamily abundance, Ramos (1992) obtained similar
ordination patterns as when counting species abundance. Another advan-
tage of this fruit-feeding guild is that it can easily be sampled with traps,
simultaneously in several points. Using appropriate criteria and guidelines,
as suggested by Sparrow et al. (1994), this nymphalid fauna may be an in-
formative species subset for monitoring programs.

The collection of vegetation variables was designed to be as simple as
possible. Of course, other local habitat variables that are important for adult
butterflies which could have been measured were not quantified. Among
the physical and structural variables are the size of the area, topography,
temperature, humidity, light, gaps, roosts and dormitories, and ground
pattern. Important biological factors for adults include food and ovoposition
site availability, predators and mimics.

Local diversity is detennined not only by local factors, but also by regional
and historical factors (Ricklefs 1987). Aside from the limits of the local
habitat structure, the local butterfly assemblage depends on the regional
species pool and historical processes such as climatic changes, isolation,
extinction and speciation. The rapid fragmentation of the Amazon rain

35:29-41, 1996(2000)

forest may be contributing to butterfly extinctions, especially larger species
with scarce resources - Morphiiiae, Brassolinae and Charaxiiiae (Brown
1991), Alternatively, the vegetation structure of disturbed forest is suitable
for sun-lovers, secondary and opportunistic species that may spread through-
out the region. Some of these butterflies are common in open biomes such
as the Cerrado. For example, Hamadryas februa, H. feronia, Erichthodes numeria
and Hermeuptychia hermes are as abundant in cerrado strictu sensu of central
Brazil (Pinheiro and Ortiz 1992) as in the disturbed areas of the fragment
studied.

Acknowledgments. The author would like to thank Dr. Vitor O. Becker for training in
lepidoptera collection, Dr. Keith S. Brown Jr. for identifying the butterflies, Drs. A,
Raw, A. F. B. Araujo, C. E. G. Pinheiro, J. S. Marinho-Filho, G. S. Dubois, J. V. Ortiz
and J. D. Hay for helpful suggestions and exciting discussions on community ecoh
ogy. Companhia Vale do Rio Doce and Dr. J. Dubois for providing field fadlities.
An anonymous reviewer for help with the language and bibliography. This work
was financially supported by the Brazilian governmental agencies CAPES and CNPq.

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J. Res. Lepid.

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Journal of Research on the Lepidoptera

35:42-60, 1996 (2000)

A Survey of the Butterfly Fauna of Jatun Sacha, Ecuador
(Lepidoptera: Hesperioidea and Papilionoidea)

Debra L. Murray

Department of Entomology, Louisiana State University, Baton Rouge, LA 70803,

E-mai 1 : dmurray @un ixl. sncc. Isu. edu

Abstract. The first extensive butterfly survey of the upper Rfo Napo ba-
sin in eastern Ecuador was conducted from 1990 to 1993. A total of 811
species was recorded at Jatun Sacha Biological Reserve. Based on species
richness comparisons with a similar site in southern Peru and extrapola-
tions from ithomiine diversity, Jatun Sacha is estimated to have approxi-
mately 1300 species of butterflies. Species richness is compared with two
other Amazonian sites (Pakitza, Peru, 1300 species and Cacaulandia, Bra-
zil, 843 species). Species and generic compositions are more similar be-
tween Pakitza and Jatun Sacha than Cacaulandia. This similarity may be
due to environmental factors. A greater percentage of Nymphalidae and
a lower percentage of Hesperiidae and Lycaenidae occur at the two some-
what disturbed sites (Jatun Sacha and Cacaulandia) than the less disturbed
site (Pakitza). Of the 228 species common to all three sites, more nympha-
lid butterfly species were found than expected based on observed spe-
cies in each family.

Key Words: Butterfly diversity, community similarities, estimations of spe-
cies richness

Introduction

The Amazon basin covers an area approximately 6 million square kilo-
meters and houses the world’s greatest diversity of plant and animal life
(Erwin 1988, Dinerstein et al. 1995). Insects are the most diverse taxon in
the neotropics, yet they have been poorly studied in this vast area (National
Academy 1992, Lamas 1989 and ref. therein, Raven 1988, Reid & Miller
1989). Even for taxonomically well known insect groups, such as the but-
terflies, there exist large gaps in our understanding of tropical species rich-
ness and factors influencing diversity (DeVries 1994, Ackery 1986). One
major hindrance is the lack of basic information available on natural his-
tory and species distributions for most Amazonian butterflies (Ackery 1986,
DeVries 1994, DeVries et al. 1997). Inventories from specific localities can
be useful in investigating changes in species compositions across landscapes,
but most of the current faunal information on Amazonian butterfly com-
munities are from Peru (Lamas 1985, 1989, Robbins et al. 1996) and areas
in Brazil (Brown 1984, 1991, Emmel & Austin 1990, Mielke 1994). There
are few published surveys of butterfly faunas in eastern Ecuador and Co-

Paper submitted 25 August 1997

35:42-60, 1996 (2000)

liimbia (Lamas 1981). Therefore our understanding of the patterns of but-
terfly diversity in these areas is very incomplete.

Biologically significant areas, such as along the eastern base of the Andes,
offer the opportunity to research factors influencing diversity and are of
particular importance to study. The eastern flank is postulated to be an area
very diverse in plant, bird, and butterfly life (Dinerstein et al. 1995, Robbins
& Opler 1996, Gentry 1988a). Gentry (1988b) found that areas of high rain-
fall and weakly defined wet and dry seasons correlated with areas of high
plant diversity. In Ecuador the only protected area in this zone is Jatun Sacha
Biological Station, located in the upper Napo basin. A flora survey at Jatun
Sacha found over 200 species of trees in one hectare plots on the reserve
(Neill & Palacios 1989) . Surveys of the fauna on the reserve have found high
species richness as well, including an extensive bird survey, which has re-
corded over 500 species (B. Bochan, pers. comm.). This diversity at Jatun
Sacha suggests the area might be equally rich in butterflies.

Here I report a survey of the Jatun Sacha butterfly fauna, which can serve
as a baseline for studies of diversity patterns at Jatun Sacha. It can also be
used for comparative studies with other localities in the region (DeVries
1994, 1996). In this paper I compare and contrast the taxonomic composi-
tions at Jatun Sacha with two other sites in the Amazon basin.

Study Sites

Jatun Sacha Biological Station is located 30 km east of the base of the
Andes (01° 04'S; 77° 36’W) and lies between the confluence of the Napo
and Arahuno rivers, its natural boundaries. Elevation varies from 400m to
450m. The uplands, typified by steep, low hills and narrow ridges with small
streams in the valleys, comprise the majority of the land. There is also a
small tract (100 hectares) in the Rio Napo floodplain with alluvial soils and
seasonal flooding. The Holdridge system would classify the lowland forests
of this area as Tropical Wet Forest (Canadas 1983). Rainfall data, recorded
since 1986, averages 3700mm annually, with no definite dry season. How-
ever, April through July are generally the wettest months and December
through February the driest months. Major floods of streams and rivers
occurs throughout the year but are more common during the wetter
months. Soil fertility is relatively rich for tropical wet forests, especially in
phosphorous and calcium, when compared to other lowland forest sites
(Clinebell et al. 1995). Storms are infrequent in the area but often cause
multiple treefalls, leaving the forest in various stages of succession (D. Neill
& W. Palacios, unpublished).

The land-use patterns in the vicinity of Jatun Sacha have undergone rapid
changes in the last decade. Before the early 1980’s the area was sparsely
populated by native Quichuans and accessed only by rivers. A road built in
1986 bisected the reserve at its northern end along the Rio Napo and greatly
increased access to the area. The influx of small scale farmers and portable
sawmills resulted in deforestation in areas accessible by the road. Currently,
tracts of land owned by farmers adjacent to the road typically have 40 to 70

/ Res. Lepid.

percent of the land cleared. Tracts in the interior are more pristine, from
50 to 100 percent primary forest. Jatun Sacha continues to expand its re-
serve and purchases lands in a piecemeal fashion as funds and land become
available. Thus the reserve is a patchwork of habitats. Its central core is
mostly primary forest (70%), and its edges are a mosaic of primary forest,
secondary forest, scrub, and pasture land (D. Neill & W. Palacios, unpub-
lished).

A brief description is presented below of the two comparative sites, Pakitza
and Cacaulandia. More complete descriptions are available from Erwin
(1991) for Pakitza and Emmel and Austin (1990) for Cacaulandia. Pakitza
is a biological station in the Reserved Zone of Parque Nacional Manu. It is
located in Madre de Dios drainage basin in Peru along the foothills of the
eastern Andes in a similar geographical zone as Jatun Sacha. The butterfly
survey for Pakitza was comprehensive and yielded 1300 species (Robbins et
al. 1996). The survey from Cacaulandia was conducted on a private ranch
in Rondonia, Brazil. Located in the rolling hills and flat plains of the Ama-
zon basin, it has both intact forest and disturbed areas. A total of 843 spe-
cies of butterflies was recorded by Emmel and Austin (1990) , although con-
tinued surveys have increased this total number to approximately 1500 spe-
cies (Austin & Emmel 1996, cited as “unpublished data”). The area of
Cacaulandia is ecologically less similar to Jatun Sacha than Pakitza, but faces
similar pressures from development.

Materials and Methods

The survey at Jatun Sacha was conducted from August 1990 to October 1993.
Hours in the field devoted to collection varied by month but covered all the months
of the year, with the exceptions of December 1990 and 1991, and October, 1992,
and data was not gathered to quantify collection effort. Collection was concentrated
in a 3 km area surrounding the station facilities. As the reserve accumulated more
land, a few specimens were taken in a 10 km area around the station. Specimens
were captured with hand held nets, butterfly traps baited with rotting fruits (DeVries
1987, 1988), artificial bait (Lamas et al. 1994), and by rearing field collected lar-
vae. Extensive use of butterfly traps at Jatun Sacha was conducted during an eco-
logical study that examined spatial and temporal diversity of the fruit feeding but-
terfly community (DeVries et al. 1997). Material from that study is included here.
The study took place from August, 1992 to October, 1993 and during that time,
baited traps were placed in both the canopy and understory for seven days a month,
a total of 105 trap days. Additional sources for species included donated specimens
or field records offered from various visiting scientists.

Identifications were conducted by comparison of my material to specimens in
the following institutions and museums: Allyn Museum of Entomology of Florida
Museum of Natural History, American Museum, Museum of Comparative Zoology,
and National Museum of Natural History, Various specialists identified particular
taxonomic groups: D. Harvey (Riodinidae) , L. Miller (Satyrinae), S. Nicolay
(Hesperiidae), and R. Robbins (Lycaenidae). Due to time constraints in the prepa-
ration and identification, some specimen determinations are tentative and are des-

35:42-60, 1996 (2000)

Table 1. Taxonomic Compositions of Butterfiy Families at Jatun Sacha,
Cacaulandia, and Pakitza. Number of species are listed in parenthesis
following the percentage of species within each family.

Family

Jatun Sacha

Cacaulandia

Pakitza

Hesperiidae

25% (198)

27% (231)

34% (442)

Papilionidae

3% (26)

2% (18)

2% (26)

Pieridae

3% (27)

4% (29)

2% (26)

Nymphalidae

38% (307)

33% (275)

28% (364)

Riodinidae

24% (194)

24% (203)

20% (260)

Lycaenidae

7% (59)

10% (87)

14% (182)

igiiated with question marks. A synoptic collection has been deposited in the Museo
de Ecuatoriana Nadonal in Quito, Ecuador.

For comparative work among the three sites, the percent of species occurring in
each family was tabulated, and a test for homogeneity across the families was calcu-
lated using a 2x2 contingency table. To compare similarity in species assemblages
between the three sites, coefficient of community indices (Pielou 1974) were cal-
culated in pairwise comparisons between Jatun Sacha and Pakitza, Jatun Sacha and
Cacaulandia, and Pakitza and Cacaulandia. Only those identified to species (spe-
cies similarities) or genus (generic similarities) were used in calculations. Lycaenidae
was not used in due to poor taxonomic resolution at the genus level and lack of
identifications in the Cacaulandia survey (59 of the 87 species were unidentified).
Using these adjusted species numbers, percentages were again calculated for fam-
ily compositions, which were used in contrasting the expected and observed spe-
cies common to all three sites.

Results

A total of 811 species were recorded at the reserve by the end of 1993
(Appendix 1). The taxonomic composition of the butterfly fauna is as
follows: Hesperiidae, 198 spp. (25%), Papilionidae, 26 spp. (3%),
Pieridae, 27 spp. (3%), Nymphalidae, 307 spp. (38%), Riodinidae, 194
spp. (24%) , and Lycaenidae, 59 spp. (7%) , Within Nymphalidae, 56 spe-
cies of Ithomiinae are those reported by Beccaloni (1995), who con-
ducted a thorough study of this group. Temporal variations in richness
and abundance were generally noted for the butterfly families, although
quantitative data was collected only for the fruitTeeding nymphalids. The
fruit-feeders were more common during the wetter months (DeVries et al.
1997), and many specimens collected during this period were fresh, indi-
cating a recent emergence. During this same time period, other families
were observed to be much less abundant, although certain species could
be common {Eurybia dardus, Urbanus simplicius, ''Theda'' tephraeus gr).
Hesperiidae, Riodinidae, and to some extent, Lycaenidae, were more abun-
dant as the rainfall decreased in August and September. Differences were
noted in the abundance of families and individual species from year to year,

/. Res. Lepid.

Table 2. Coefficient of Community Indices for Jatun Sacha, Pakitza, and

Cacaulandia.

Species similarities Generic similarities

Jatun Sacha-Pakitza

Jatun Sacha-Cacaulandia

Pakitza-Cacaulandia

especially among Riodinidae and Lycaenidae. Some species abundance
patterns were irregular. For example, I did not see Stalachtis euterpe until
January, 1993, when it was common for several months along the ridges in
the primary forest. Other examples include Metacharis regalis and Emesis

temesa.

Family compositions varied significantly among the three sites (p>0.05).
Jatun Sacha and Cacaulandia shared a greater similarity in family composi-
tions than any other pairwise comparisons (Table 1). The combination of
Riodinidae and Lycaenidae percentages is nearly identical in all three
sites (31% to 34%). However, the percentages of Lycaenidae are con-
siderably lower at Jatun Sacha, and to a lesser extent, Cacaulandia, than
at Pakitza. In contrast, Jatun Sacha shared a greater number of species
and genera with Pakitza rather than Cacaulandia. Coefficient of com-
munity values ranked Jatun Sacha and Pakitza with greatest similarity
and Pakitza and Cacaulandia with the least similarity (Table 2). Inter-
estingly, only 228 species were common to all three sites. Of those 228
species, Nymphalidae accounted for 53% (121 species) of the total num-
ber. Listed in order of abundance, the numbers of species for the other
butterfly families were: Hesperiidae (56), Riodinidae (32), Papilionidae
(12), and Pieridae (7). The number of observed overlapping nympha-
lid species was greater than expected when calculated using the family
percentages (minus unidentified species and Lycaenidae). For example,
the adjusted family compositions for Nymphalidae range from 33%
(Pakitza) to 43% (Jatun Sacha). Using the higher percentage, 98 spe-
cies of the total 228 were expected to be nymphalids, although 121 were
actually found to be overlapping. In contrast, the number of overlap-
ping hesperiid species was lower than expected.

Discussion

The survey conducted at Jatun Sacha was aimed at developing a baseline
understanding of the butterfly community of the area. A large portion of
the fauna undoubtably remains unsampled. This conclusion is supported
by the fact that unrecorded species were collected up to the end of the sur-
vey time. In addition, preliminary identifications for certain groups have
probably underestimated the number of butte idly species actually collected.
Because field collection was not standardized, estimations of the total spe-
cies richness at Jatun Sacha can not be generated through rigorous statisti-

35:42-60, 1996 (2000)

cal programs (DeVries et al. 1997). Nonetheless, some estimation can be
made from comparisons of inventories at similar localities, such as at Pakitza.
Pakitza and Jatun Sacha are both located along the eastern edge of the low-
land rainforest and share similar elevation, temperature, and annual rain-
fall, although Jatun Sacha is more aseasonal than Pakitza. Given these simi-
lar environmental factors, it is estimated that 1200 to 1300 species poten-
tially occur at Jatun Sacha. This estimate is supported by applying the model
proposed by Beccaloni and Gaston (1995), in which total ithomiine rich-
ness from an area is used to predict overall species richness. Beccaloni and
Gaston found that ithomiines were, on average, 4.5% of the total species
for an area. Given 58 species of ithomiines at Jatun Sacha, approximately
1300 species of butterflies are predicted to occur there. This suggests that
a third of the fauna has yet to be recorded, illustrating the importance of
further survey work.

Comparing faunal lists from different study sites is confounded by differ-
ences in sampling methods and climatic and ecological factors (DeVries
1994) . Misidentifications of species and nomenclature changes can also yield
misleading results. All of these factors could have influenced compari-
sons of the species assemblages between the Jatun Sacha, Pakitza, and
Cacaulandia, however differences in sampling methodologies was probably
most influential. Much of the early sampling in Cacaulandia was conducted
by participants in tour groups who may have selectively collected colorful
butterflies over some of the more drab species. Since the initial list of but-
terflies was published from Cacaulandia (Emmel & Austin 1990), the au-
thors have continued their sampling effort and have documented many
more species (Austin & Emmel 1996). Patterns of diversity reported here
may change when compared with the forthcoming update to the survey.
Sampling at Jatun Sacha used bait traps more extensively than Pakitza or
Cacaulandia, At Jatun Sacha, 189 species were trapped at rotting fruit (23%
of the butterfly fauna). At Pakitza, 130 species were trapped (10% of the
butterfly fauna) (Robbins et al. 1996) . The survey at Pakitza was conducted
on a larger scale than the other two, with intense collecting and a greater
number of experts available to identify species, although field crews varied
with each sampling period. These differences in sampling have influenced
the species recorded, and consequently, the compositions of the various
groups.

Environmental variables, most notably climatic factors, are most often cor-
related with species richness and diversity (Wright et al. 1993). In this study
the hypothesis is supported by the results of the generic and species simi-
larities. Jatun Sacha and Pakitza had the highest coefficient of community.
Pakitza and Cacaulandia, although geographically closest among the three
sites were actually the most dissimilar. This underscores the importance of
local conditions on determining species compositions.

Disturbance is another factor influencing species compositions between
the three sites. Forest areas with mild disturbances, such as those that exist
in Cacaulandia and Jatun Sacha, can experience increases in butterfly di-

J. E£S. Lepid.

versity in certain groups, such as Nyniphalidae (Brown 1982; but see also
DeVries et aL 1997). Butterfly species common to open, disturbed areas are
rare or absent at Pakitza (Robbins et al. 1996), but are quite common at
Jatun Sacha along the road bisecting the reserve. The low species richness
of Hesperiidae and Lycaenidae recorded at Jatun Sacha and Cacaulandia
could also reflect disturbance, especially atjatuii Sacha. A lepidopterist who
has been collecting in the Upper Napo area since 1978 has noted a great
decrease in the species and abundance of the Hesperiidae over the last
decade as developmental pressures increased (S. Nicolay, pers. comm.).

From the comparisons of the overlapping species, nymphalid species were
most common and found at greater numbers than expected. This suggests
broader distributions of iiymphalids than other butterfly families. This may
be due to the wide dispersing capabilities of many nymphalids, which have
been correlated with greater distributions (Hanski et al. 1993). It could also
reflect broader hostplant ranges for nymphalids or more specialized, and
hence, localized host use by other butterfly families. With our limited knowl-
edge of host use even in well studied areas such as Costa Rica (DeVries 1987,
1996; DeVries et ak 1994), examining these broader biogeographica! pat-
terns must await further investigations (but see Ackery 1988).

Human influence outside of Jatun Sacha most likely has impacted the but-
terfly fauna. Species inventories conducted while the area contains a high
percentage of pristine forest could be compared with future inventories in
a potentially much more disturbed landscape. Because degradation of the
upper Napo basin will continue, there is a critical need for more research.
For too many species, little is known beyond their site records. A great deal
remains to be discovered to complete our understanding of the butterfly
fauna, not only in documentation of the species diversity, but also their
ecology, evolution, and population dynamics.

Acknowledgements. I thank the following for contributing specimens to the list: Dave
Arenholz, George Beccaloni, Marion Murray, Andrew Neild, Stan Nicolay, Carla
Penz, Karina Soria, Alejandro Suarez, Gabriel Tapuy, and the students of the Save
the Rainforest, The survey list would not have been possible without the time and
effort to identify specimens by Stan Nicolay (Hesperiidae), George Beccaloni
(Ithomiinae), Lee Miller (Satyrinae), Don Harvey, Dave Arenholz, and Phil DeVries
(Riodioidae), and Bob Robbins (Lycaenidae). This manuscript was improved by
changes suggested by George Beccaloni (Natural History Museum), Chris Carlton
(Louisiana State University), Phil DeVries (University of Oregon), Sam Messier
(University of Colorado), David Neill (Missouri Botanical Garden), Dorothy Prowell
(Louisiana State University), and Bob Robbins (National Museum of Natural His-
tory). This work was possible through the support of Fundacion Jatun Sacha and
the United States Peace Corps,

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APPENDIX 1

The following is a list of the butterflies collected at Jatun Sacha Biological
Station. A question mark (?) following a name indicates questionable iden-
tification of the species. Species designated as “unknown” could not be iden-
tified to genus or species. The list follows the higher taxonomic classifica-
tion of Evans (1951, 1952, 1953, 1955) for Hesperiidae, Tyler et al (1994)
for Papilionidae, Klots (1933) for Pieridae, Harvey (1991) for Nymphalidae,
Forster (1964) for Satyrinae, and Harvey (1987) for Riodinidae.

Hesperiidae 198

Pyrrhopyginae; 4
Elbella theseus Bell, 1933
Passova passova Evans, 1951
Pyrrhopyge proculus cintra Evans, 1951
Pyrrhopyge aziza lexos Evans, 1951

J. Res. Lepid.

Pyrginae: 107

Achylodes thraso thraso (Hiibner, 1807)
Achylodes busims heros (Ehrmann,
1909)

Aguiia coelus (Cramer, 1782)

Agima clina Evans, 1952
Agima arimce (Hewitson,1867)

Anastriis obscurus iiarva Evans, 1953
Anastrns sempiternus simplicior
(Moschler, 1876)

Anisochoria pedaliodina Buder, 1870
Antigonus nearchus (Latreille, 1824)
Antigonus miidlatiis Hopffer, 1874
Antigonus erosus (Hiibner, 1812)
Astraptes fidgerator azul Reakirt, 1866
Astraptes alardus alardus (Stoll, 1790)
Astraptes tains (Cramer, 1777)

Astraptes fulgor (Hayuard, 1938)
Astraptes alector hopfferi (Plotz, 1882)
Astraptes cretatus cretatiis (Haward,
1939)

Astraptes anaphns anaphiis (Cramer,
1777)

Aiitochton neis (Plotz, 1882)

Autochton longipennis (Geyer, 1832)
Bolla mancoi (Lindsey, 1925)

Bolla ciipreiceps (Mabille, 1889)
Bnngalotis erythiis Cramer, 1775
Cabriiis procas purda Evans, 1952
Calliades zeutiis (Moschler, 1879)
Camptopleiira auxo (Moschler, 1878)
Carrhenes fiiscescens Mabille, 1891
Celaenorrhinns jao (Mabille, 1889)
Celaenorrhinns shema shema
(Hewitson, 1877)

Celaenorrhinns syllins (Eelder & Felder,
1862)

Charidia Incaria pocus Evans, 1953
Chrysoplectriim perniciosns perniciosus
(Herrich-Schaffer, 1869)

Cycloglypha caeruleonigra Mabille,

1904

Cyclosemia pedro Williams & Bell, 1940
Cyclosemia lathaea Hewitson, 1878
Dyscophellns euribates ein abates
(Cramer, 1782)

Dyscophellns sp.

Dyscophellns ramnsis Stoll, 1781
Ebrietas evanidns (Mabille, 1897)
Ebrietas infanda (Butler, 1876)

Enthens priassns telemns Mabille, 1898
Epargyrens socns dicta Evans, 1952
Eracon panlinns (Cramer, 1782)
Gorgythion begga planta (Moschler,

1867)

Haemactis sangninalis (Westwood,

1852)

Helias phalaenoides phalaenoides
(Hnbner, 1812)

Heliopetes alana (Reakirt, 1868)
Hyalothyrns nelens neleus (Linnaeus,
1852) ^

Mictris crispns crispns (Herrich-
Schaffer, 1869)

Milanion hemes pemba Evans, 1953
Morvina morvns Plotz, 1884
Mylon cajns (Plotz, 1884)

Mylon illineatns illineatns (Mabille &

Bonllet, 1917)

Mylon menippns (Fabricins, 1776)
Narcosins mnra (Williams, 1927)
Narcosins colossus (Herrich-Schaffer,
1869)

Nisoniades castolns (Hewitson, 1878)
Nisoniades bessns hecales (Hayward,
1940)

Oldens fridericns fridericus (Geyer,
1832)

Oldens calavins calavins (Godman &
Salvin, 1895)

Oldens matria Evans, 1953
Paches trifasciatns Lindsey, 1925
Pellicia dimidiata dimidiata (Herrich-

Schaffer, 1870)

Phanns vitrens (Cramer, 1782)

Phareas coeleste Westwood, 1852
Phocides metrodorns metrodorns Bell,
1932

Plumbago plumbago (Plotz, 1884)
Polyctor polyctor polyctor (Prittwitz,

1868)

Polytlirix endoxns (Cramer, 1782)

Polythrix cecidns (Herrich-Schaffer,

1869)

Porphyrogenes passalns passalns
(Herrich-Schaffer, 1869)
Potamanaxas hirta hirta (Weeks, 1901)
Potamanaxas flavofasciata flavofasciata
(Hewitson, 1870)

Pyrdalns corbnlo (Stoll, 1781)

Pyrgns oilens Linnaens, 1767
Pythonides assacla Mabille, 1883
Pythonides herrenins Geyer, 1838
Pythonides joviannsjovianns (Stoll,
1782)

Qnadrns deyrollei porta Evans, 1952
Qnadriis cerialis (Cramer, 1782)
Sostrata festiva (Erichson, 1848)
Sostrata pnsilla pnsilla (Godman &
Salvin, 1895)

Spathilepia clonins (Cramer, 1775)
Spioniades artemidas (Cramer, 1782)
Staphylns balsa (Bell, 1937)

Staphylns lizeri (Hayward, 1938)

35:42-60, 1996 (2000)

Tarsocteniis praecia plutia (Hewitson,
1857)

Tarsoctenus papias Hewitson, 1857
Tarsocteniis corytiis corba Evans, 1952
Telemiades epicalus sila Evans, 1953
Telemiades centrides Hewitson, 1870
Telemiades amphion misitheus
(Mabille, 1888)

Telemiades penidas (Hewitson, 1867)

Typhedanns undulatus (Hewitson,

1867)

Typhedanns orion (Cramer, 1779)
Urbanus teleus (Hubner, 1821)

Urbanus simplicius (Stoll, 1791)
Urbanus proniis Evans, 1952
Urbanus virescens (Mabille, 1877)
Urbanus viterboana viterboana
(Ehrmann, 1907)

Urbanus pronta Evans, 1952
Urbanus esta Evans, 1952
Urbanus doryssus doryssus (Swainson,
1831)

Urbanus dorantes dorantes (Stoll, 1791)
Urbanus albimargo takuta Evans, 1952
Urbanus procne (Plotz, 1881)
Xenophanes tryxus (Cramer, 1782)

Hesperiinae; 87

Anatrytone sarah (Burnes, 1994)
Anthoptus epictetus (Fabricius, 1793)
Arita arita (Schaus, 1902)

Aroma aroma Hewitson, 1867
Artines aepitus (Geyer, 1832)
Callimormus radiola radiola (Mabille,
1897)

Carystina lysiteles Mabille, 1891
Carystoides sicania orbius (Godman,
1901)

Carystoides lila Evans, 1955
Chloeria psittacina Felder, 1867
Cobalopsis potaro (William & Bell,

1931)

Cobalopsis nero (Herrich-Schaffer,
1869)

Cobalus virbius virbius (Cramer, 1777)
Conga chydea (Butler, 1870)

Corticea corticea corticea (Pldtz, 1883)
Cymaenes tripunctata alumna (Butler,
1877)

Cymaenes cavalla Evans, 1955
Cynea megalops (Godman, 1900)

Damas clavus (Herrich-Schaffer, 1869)
Decinea percosius (Godman, 1900)
Decinea sp.

Decinea derisor (Mabille, 1891)

Ebusus ebusus (Cramer, 1782)

Eutocus quichua Lindsey, 1925

Eutychide subcordata subcordata
(Herrich-Schaffer, 1869)

Eutychide complana (Herrich-Schaffer,
1869)

Flaccilla aecas Stoll, 1781
Hylephila phylaeus phylaeus (Drury,
1773)

Justinia phaetusa phaetusa (Hewitson,
1866)

Lento lento Mabille, 1878
Lycas boisduvalii Ehrmann, 1909
Metron nr. chrysogastra
Mnasilus allubita Butler, 1877
Moeris vopiscus vopiscus (Herrich-
Schaffer, 1869)

Moeris striga Geyer, 1832

Molo mango mango (Guenee, 1865)

Molo petra Evans, 1955

Morys geisa geisa (Moschler, 1878)

Mucia sp.

Nastra insignis (Plotz, 1882)

Niconiades nikko Hayward, 1948
Nyctelius nyctelius (Latreille, 1824)
Orses cynisca (Swainson, 1821)

Oxynthes corusca (Herrich-Schaffer,
1869)

Panoquina fusina fusina (Hewitson,

1868)

Panoquina evadnes (Stoll, 1781)

Papias proximus (Bell, 1934)

Papias integra Mabille, 1891
Paracarystus menestries rona (Hewitson,
1866)

Parphorus decora (Herrich-Schaffer,

1869)

Parphorus storax storax (Mabille, 1891)
Pellicula crista Evans, 1955
Pellicula criska jon Nicolay, 1980
Pellicula bryanti (Weeks, 1906)
Perichares philetes dolores (Reakirt,
1868)

Phanes almoda (Hewitson, 1866)
Pompeius pompeius (Latreille, 1824)
Quinta cannae (Herrich-Schaffer, 1869)
Racta sp.

Saliana salius (Cramer, 1776)

Saliana esperi Evans, 1955
Saliana antoninus Latrielle, 1824
Saliana triangularis (Kaye, 1913)
Saturnus tiberius suffuscus (Hayward,
1940)

Sodalia sodalis Butler, 1877
Talides sergestus Cramer, 1775)

Talides sinois sinois Hubner, 1819
Telles arcalaus (Cramer, 1782)

Thargella caura caura (Plotz, 1882)
Thespias dalman Latreille, 1824

J. Res. Lepid.

Thoon sp.

Thoon poiika Evans, 1955
Thoon taxes (Godman, 1900)

Thoon modiiis (Mabille, 1889)
Thracides phidon (Cramer, 1779)
Thracides smaragdiihis (Herrich-
Schaffer, 1869)

Vehilius stictomenes stictomenes Butler,
1877

Vehilius illudens Mabille, 1891
Vehilius vetula (Mabille, 1878)

Vehilius inca (Scudder, 1872)

Venas caeruleans (Mabille, 1828)

Vettius phyllus phyllus (Cramer, 1777)
Vettius richardi (Weeks, 1906)

Vettius artona (Hewitson, 1868)

Vettius marcus marcus (Fabricius, 1787)
Xeniades orchamus orchanius (Cramer,
1777)

Zenisjebus melaleuca (Plotz, 1882)
Papilionidaes 26

Battus crassus crassus (Cramer, 1777)
Battus polydamas polydamas (Linnaeus,
1758)

Battus belus varus (Kollar, 1850)
Eurytides dolicaon ?

Heraclides torquatus torquatus
(Cramer, 1777)

Heraclides thoas cinyras (Menetries,
1857)

Heraclides isidorus flavescens
(Oberthur, 1880)

Heraclides hyppason hyppason
(Cramer, 1776)

Heraclides chiansiades chiansiades
(Westwood, 1872)

Heraclides astyalus phanias (Rothschild
& Jordan, 1906)

Heraclides androgens androgens

(Cramer, 1776)

Mimoides ariarathes gayi (Lucas, 1852)
Mimoides xynias (Hewitson, 1867)
Mimoides pausanias pausanias
(Hewitson, 1852)

Parides anchises drucei (Butler, 1874)
Parides aeneas bolivar (Hewitson, 1850)
Parides vertumnus bogotanus (Felder &
Felder, 1864)

Parides neophilus olivencius (Bates,
1861)

Parides erithalion guillerminae
(Pischedda & Racheli, 1986)

Parides sesostris sesostris (Cramer,

1780)

Parides lysander brissonius (Hiibner,
1819)

Parides chabrias chabrias (Hewitson,
1852)

Protesilaus telesilaus telesilaus (Felder
& Felder, 1864)

Protographium agesilaus autosilaus
(Bates, 1861)

Protographium thyastes thyastinus
(Oberthiir, 1880)

Pterourus zagreus neyi (Niepelt, 1909)

Pieridae 27

Aphrissa statira (Cramer, 1777)
Archonias bellona (Cramer, 1776)
Charonias eurytele (Hewitson, 1853)
Cunizza hirlanda (Stoll, 1791)
Dismorphia theucharila (Doubleday,
1848)

Dismorphia amphiona Cramer, 1780
Enantia melite (Linnaeus, 1763)

Enantia lina (Herbst, 1792)

Eurema daira (Godart, 1819)

Eurema sp.

Eurema albula (Cramer, 1776)

Eurema xanthochlora (Kollar, 1850)
Itaballia pisonis (Hewitson, 1861)
Itaballia demophile (Linnaeus, 1763)
Leptophobia aripa (Boisduval, 1836)
Leucidia brephos (Hiibner, 1809)
Moschoneura pinthaeus (Linnaeus,
1758)

Patia oresis (Boisduval, 1836)

Perrhybis pyrrha (Cramer, 1782)
Perrhybis lorena (Hewitson, 1852)
Phoebis rurina (Felder 8c Felder, 1861)
Phoebis philea (Linnaeus, 1763)
Phoebis argante (Fabricius, 1775)
Phoebis trite (Linnaeus, 1758)
Pieriballia mandella (Felder & Felder,
1861)

Pyrisitia venusta (Boisduval, 1836)
Pyrisitia nise (Cramer, 1776)

Nymphalidae 307

Heliconiinae 22

Actinote sp.

Actinote pellenea Hiibner, 1821
Agraulis vanillae (Linnaeus, 1763)
Dionejimo (Cramer, 1780)

Dryadula phaetusa (Linnaeus, 1758)
Dryas iulia (Fabricius, 1775)

Eueides tales (Cramer, 1776)

Eueides aliphera (Godart, 1819)

Eueides isabella isabella (Cramer, 1782)
Eueides lampeto acacetes Hewitson,
1869

Eueides lybia (Fabricius, 1775)

35:42-60, 1996 (2000)

Eiieides vibilia (Godart, 1819)
Heliconius erato ladvitta Butler, 1877
Heliconius hecale quitalena Hewitson,
1853

Heliconius elevatus elevatus Noldner,
1901

Heliconius wallacei Reakirt, 1866
Heliconius sara (Fabricius, 1793)
Heliconius melponiene aglaope Felder
& Felder, 1862

Heliconius numata euphone Felder 8c
Felder, 1862

Laparus doris (Linnaeus, 1771)

Neruda aoede bartletti Druce, 1876
Philaethria dido (Linnaeus, 1763)

Nymphalinae 20

Anartia amathea (Linnaeus, 1758)
Anartia jatrophae (Linnaeus, 1763)
Anthanassa drusilla (Felder 8c Felder,
1861)

Castilia perilla (Hewitson, 1852)
Castilia angusta (Hewitson, 1868)
Castilia ofella (Hewitson, 1864)

Fresia clara clara Bates, 1864
Fresia eunice eunice (Hubner, 1807)
Fresia nauplius (Linnaeus, 1758)

Fresia sp.

Fresia perna Hewitson, 1852
Fresia pelonia pelonia Hewitson, 1852
Hypanartia lethe (Fabricius, 1793)
Junonia evarete (Cramer, 1870)
Metamorpha elissa Hiibner, 1819
Phyciodes sp.

Phyciodes aveyrana (Bates, 1864)
Siproeta stelenes Linnaeus, 1758
Tegosa claudina (Eschscholtz, 1821)
Telenassa burchelli (Moulton, 1909)

Limenitidinae 78
Adelpha boeotia (Felder 8c Felder,
1867)

Adelpha delinita (Fruhstorfer, 1913)
Adelpha iphiclus (Linnaeus, 1758)
Adelpha erotia (Hewitson, 1847)
Adelpha cytherea (Linnaeus, 1758)
Adelpha celerio (Bates, 1864)

Adelpha boreas (Butler, 1866)

Adelpha sp. 3
Adelpha sp. 2

Adelpha lerna (Hewitson, 1847)
Adelpha melanthe (Bates, 1864)
Adelpha sp. 1

Asterope degandii (Hewitson, 1850)
Baeotus japetus (Staudinger, 1885)
Baeotus deucalion (Felder 8c Felder,
1860)

Baeotus amazonicus (Riley, 1919)

Batesia hypochlora (Felder 8c Felder,
1862)

Biblis hyperia (Cramer, 1780)

Callicore cynosura (Doubleday, 1847)
Callicore lyca (Doubleday, 1847)
Callicore hystaspes (Fabricius, 1782)
Callicore hesperis (Guerin, 1844)
Callicore eunomia (Hewitson, 1853)
Callicore cyllene (Doubleday, 1847)
Catacore kolyma (Hewitson, 1852)
Catonephele acontius acontius
(Linnaeus, 1758)

Catonephele numilia numilia (Cramer,

1776)

Colobura dirce (Linnaeus, 1758)
Diaethria clymena (Cramer, 1776)
Dynamine geta (Godman 8c Salvin,

1878)

Dynamine racidula (Hewitson, 1852)
Dynamine zenobia (Bates, 1865)
Dynamine glance (Bates, 1865)
Dynamine gisella (Hewitson, 1852)
Dynamine athemon (Linnaeus, 1758)
Dvnamine artemisia (Fabricius, 1793)
Dynamine anubis (Hewitson, 1859)
Fctima iona (Doubleday, 1848)

Fctima lirides (Staudinger, 1885)

Funica eurota eurota (Cramer, 1776)
Funica sophonisba agele Seitz, 1915
Funica norica occia Fruhstorfer, 1909
Funica alpais alpais (Godart, 1824)
Funica mygdonia mygdonia (Codart,
1824)

Funica marsolia fasula Fruhstorfer, 1909
Funica amelia erroneata Oberthur,

1916

Funica clytia (Hewitson, 1852)
Haematera pyramus (Fabricius, 1782)
Hamadryas laodamia laodamia
(Cramer, 1777)

Hamadryas arinome arinome (Lucas,
1853) ^

Hamadry as amphinome amphinome
(Linnaeus, 1767)

Hamadryas feronia feronia (Linnaeus,
1758) ^

Hamadryas chloe chloe (Stoll, 1791)
Historis acheronta (Fabricius, 1775)
Historis odius (Fabricius, 1775)

Marpesia furcula (Fabricius, 1793)
Marpesia iole (Drury, 1782)

Marpesia chiron (Fabricius, 1775)
Marpesia berania (Hewitson, 1852)
Marpesia crethon (Fabricius, 1776)
Marpesia petreus (Cramer, 1776)
Marpesia themistocles (Fabricius, 1793)

J. Res. Lepid.

Nessaea obriiia lesoiidieri Le Moult,

1933

Nessaea hewitsonii hewitsonii (Felder 8c
Felder, 1859)

Nica llavilla (Godart, 1824)

Panacea prola (Doubleday, 1848)
Panacea procilla (Hewitson, 1854)
Panacea regina (Bates, 1864)
Pauiogrannna pyracmon (Godart, 1824)
Peria lamis (Cramer, 1780)

Pyrrhogyra neaerea (Linnaeus, 1758)
PvrrhogvTa otolais (Bates, 1864)
Pyrrhogyra crameri (Aurivillius, 1882)
Smyrna blomfildia (Fabricius, 1782)
Temenis pulchra (Hewitson, 1861)
Temenis laothoe (Cramer, 1777)
Tigridia acesta (Linnaeus, 1758)

Vila azeca (Doubleday, 1848)

Charaxinae 26

Agrias claudina (Godart, 1824)

Agrias hewitsonius Bates, 1860
Agrias amydon Hewitson, 1854
Archaeoprepona licomedes (Cramer,
1777)

Archaeoprepona demophoon (Hiibner,
1814)

Archaeoprepona demophon (Linnaeus,
1758)

Archaeoprepona amphimachus (Fabri-
cius, 1775)

Coenophlebia archidona Felder &
Felder, 1862

Consul fabius aequatorialis (Butler,
1875)

Fountainea ryphea ryphea (Cramer,
1776)

Fountainea eurypyle (Felder & Felder,
1862)

Hypna clytemnestra (Cramer, 1777)
Memphis morvus (Fabricius, 1775)
Memphis florita (Druce, 1877)

Memphis sp.

Memphis philumena philumena
(Doubleday, 1849)

Memphis arachne (Cramer, 1776)
Memphis xenocles (Westwood, 1850)
Memphis offa (Druce, 1877)

Memphis oenomais (Boisduval, 1870)
Memphis polycarmes (Fabricius, 1775)
Prepona pheridamas (Cramer, 1777)
Prepona laertes (Hubner, 1814)
Prepona pylene Hewitson, 1854
Siderone marthesia (Cramer, 1777)
Zaretis itys (Cramer, 1777)

Apaturinae 8

Doxocopa cherubina (Felder & Felder,
1867)

Doxocopa Clothilda (Felder 8c Felder,
1867)

Doxocopa cyane (Latreille, 1813)
Doxocopa felderi (Godman 8c Salvin,
1884)

Doxocopa laure (Drury, 1773)
Doxocopa pavon (Latreille, 1809)
Doxocopa sp.

Doxocopa agathina (Cramer, 1777)

Morphinae 8

Aiitirrhea avernus (Hopffer, 1874)
Antirrhea sp.

Morpho achilles (Linnaeus, 1758)
Morhpo adonis (Cramer, 1776)

Morpho deidamia (Hiibner, 1819)
Morpho hecuba (Linnaeus, 1771)
Morpho menelaus (Linnaeus, 1758)
Morpho rhetenor (Cramer, 1776)

Brassolinae 15

Brassolis sophorae (Linnaeus, 1758)
Caligo illioneus (Cramer, 1776)

Caligo idomeneus (Linnaeus, 1758)
Caligo eurilochus (Cramer, 1776)

Caligo placidianus (Staudinger, 1887)
Caligo euphorbus (Felder 8c Felder,
1862)

Catoblepia xanthicles (Godman 8c
Salvin, 1881)

Catoblepia berecynthia (Cramer, 1777)
Catoblepia xanthus (Linnaeus, 1758)
Eryphanis polyxena (Meerburgh, 1780)
Opoptera aorsa (Godart, 1824)
Opsiphanes quiteria (Cramer, 1782)
Opsiphanes invirae (Hubner, 1808)
Opsiphanes cassiae (Linnaeus, 1758)
Selenophanes cassiope (Cramer, 1776)

Satyrinae 68

Amphidecta calliomma (Felder &
Felder, 1862)

Amphidecta pignerator (Butler, 1867)
Bia actorion (Linnaeus, 1763)

Caeruleuptychia coelica (Hewitson,

1869)

Caeruleuptychia nr. pencillata
Caeruleuptychia sp. 2
Caeruleuptychia aegrota (Butler 1867)
Caeruleuptychia pilata (Butler, 1867)
Caeruleuptychia sp. 1
Cepheuptychia cephus (Fabricius, 1775)
Chloreuptychia herseis (Godart, 1824)
Chloreuptychia chloris (Cramer, 1782)

35:42-60, 1996 (2000)

Chloreuptychia toliimnia (Cranier,

1777)

Chloreuptychia arnaca (Fabricius, 1776)
Chloreutychia agatha (Butler, 1867)
Cissia proba (We)Tner, 1911)

Cissia terrestris (Butler, 1867)

Cissia penelope (Fabricius, 1775)

Cissia sp. 2

Cissia myncea (Cramer, 1782)

Cissia sp, 1

Cithaerias aurora (Felder & Felder,
1862)

Erichthodes erichtho (Butler, 1867)
Euptychia sp. 3
Euptychia sp. 4
Euptychia sp. 1

Euptychia picea (Butler, 1867)

Euptychia sp. 2

Haetera piera (Linnaeus, 1758)
Hermeuptychia hermes (Fabricius,

1775)

Magneuptychia analis (Godman, 1905)
Magneuptychia tricolor (Hewitson,

1850)

Magneuptychia inodesta (Butler, 1867)
Magneuptychia alcinoe (Felder &
Felder, 1867)

Magneuptychia ocypete (Fabricius,

1776)

Magneuptychia ayaya (Butler, 1867)
Magneuptychia nr. helle 1
Magneuptychia nr. helle 2
Magneuptychia nr. inani
Magneuptychia libye (Linnaeus, 1767)
Magneuptychia sp.

Manataria hyrnethia (Fruhstorfer, 1912)
Megeuptychia antonoe (Cramer, 1776)
Pareuptychia hesionides (Forster, 1964)
Pareuptychia ocirrhoe (Fabricius, 1776)
Pareuptychia sp.

Pierella lena (Linnaeus, 1767)

Pierella lamia (Sulzer, 1776)

Pierella hortona (Hewitson, 1854)
Pierella astyoche (Erichson, 1848)
Posttaygetis penelea (Cramer, 1777)
Pseudodebis sp.

Pseudodebis valentina (Cramer, 1780)
Pseudodebis marpessa (Hewitson, 1862)
Splendeuptychia nr. itonis
Splendeuptychia itonis (Hewitson,

1862)

Splendeuptychia sp. 1
Taygetis celia (Cramer, 1780)

Taygetis armillata (Butler, 1868)
Taygetis sosis (Hopffer, 1874)

Taygetis cleopatra (Felder & Felder,
1867)

Taygetis virgilia (Cramer, 1776)

Taygetis rufomarginata (Staudinger,
1888)

Taygetis thamyra (Cramer, 1779)
Taygetis laches (Fabricius, 1793)
Taygetis mermeria (Cramer, 1776)
Yphthimoides erigone (Butler, 1867)
Yphthimoides renata (Cramer, 1782)

Danainae 4

Danaus plexippus (Linnaeus, 1758)
Lycorea ilione (Cramer, 1776)

Lycorea pasinuntia brunnea Riley, 1919
Lycorea cleobaea atergatis Doubleday,
'l847

Ithomiinae 58

“Hvpoleria” orolina orolina (Hewitson,
1861)

“Hypoleria” seba oculata Haensch, 1903
“Pseudoscada” florula aureola
(Hewitson, 1855)

Aeria eurimedea negricola (Felder &
Felder, 1865)

Callithomia lenea zelie Guerin, 1844
Callithomia alexirrhoe butes Godman &
Salvin, 1898

Ceratinia tutia poecila (Bates, 1862)
Ceratiscada hymen (Haensch, 1905)
Dircenna loreta loreta Haensch, 1903
Forbestra equicola equicoloides
(Godman 8c Salvin, 1898)

Forbestra olivencia juntana (Haensch,
1903)

Godyris zavaleta matronalis (Weymer,
1883)

Godyris dircenna dircenna (Felder 8c
Felder, 1862)

Heterosais nephele nephele (Bates,
1862)

Hyalyris coeno norellana (Haensch,
1903)

Hypoleria lavinia chrysodonia (Bates,
1862)

Hypoleria sarepta aureliana (Bates,
1862)

Hyposcada anchiala ecuadorina Bryk,
1953

Hyposcada illinissa ida Haensch, 1903
Hyposcada kena kena (Hewitson, 1872)
Hypothyris moebiusi unicolora
(Tessmann, 1928)

Hypothyris mamercus mamercus
(Hewitson, 1869)

Hypothyris euclea intermedia (Butler,
1873)

Hypothyris anastasia honesta (Weymer,

/. Res. Lepid.

1883)

Hypothyris moebiusi moebiiisi (Haensch,
1903)^

Hypothyris semifliiva satura (Haeiisch, 1903)
Hypothyris anastasia bicolor (Haensch, 1903)
Hypothyris fliionia berna (Haensch, 1903)
Ithomia salapia salapia Hewitson, 1853
Ithomia salapia travella Haensch, 1903
Ithomia amarilla amarilla Haensch, 1903
Ithomia agnosia agonsia Hewitson, 1855
Mechanitis mazaens mazaeiis Hewitson, 1860
Mechanitis mazaens fallax Butler, 1873
Mechanitis mazaens visenda Butler, 1877
Mechanitis messenoides messenoides Felder
& Felder, 1865

Mechanitis polymnia dorissides Staudinger,
1844

Mechanitis lysimnia elisa (Guerin, 1844)
Melinaea mnasias abtigua Brown, 1977
Melinaea menophilns cocana Haensch, 1903
Melinaea marsans monthone Hewitson, 1860
Melinaea maeliis maenois Hewitson, 1869
Methona cnrvifascia curvifascia Weymer, 1883
Methona confnsa psamathe Godman &

Salvin, 1898

Napeogenes achaea achaea (Hewitson, 1869)
Napeogenes aethra aethra (Hewitson, 1869)
Napeogenes inachia avila Haensch, 1903
Napeogenes Stella (Hewitson, 1855)
Napeogenes sylphis cancayaensis Fox & Real,
1971

Napeogenes pharo pharo (Felder & Felder,
1862)

Oleria giinilla lota (Hewitson, 1872)

Oleria tigilla tigilla (Weymer, 1899)

Oleria sexmacnlata sexmaculata (Haensch,
1903)

Oleria lerda lerda (Haensch, 1909)

Oleria agarista agarista (Felder & Felder,
1862)

Oleria assimilis assimilis (Haensch, 1903)
Psendoscada timna timna (Hewitson, 1855)
Pteronymia vestilla sparsa Haensch, 1903
Scada reckia ethica (Hewitson, 1861)

Thyridia psidii ino Felder & Felder, 1862
Tithorea harmonia hermias Godman &

Salvin, 1898

Riodinidae 194

Adelotypa amasis (Hewitson, 1870)

Adelotypa alector Butler, 1867
Adelotypa senta (Hewitson, 1853)

Adelotypa sp. 1
Adelotypa sp. 2
Adelotypa sp. 3
Adelotypa sp. 4
Alesa amesis (Cramer, 1777)

Alesa sp.

Alesa telephae (Boisdnval, 1836)
Amarynthis meneria (Cramer, 1776)
Ancylnris aulestes (Cramer, 1777)
Ancylnris meliboeus (Fabricius, 1777)
Anteros acheus (Stoll, 1781)

Anteros allectus Westwood, 1851
Arg\Togrammana sp. 3
Argyrogrammana sp. 1
Argyrogrammana sp. 2
Argyrogrammana trochilia Westw^ood,
1851

Calospila trinitatis (Lathy, 1932)
Calospila parthaon (Dalman, 1823)
Calospila sp.

Calospila maeonides ?

Calospila rhodope (Hewitson, 1853)
Calospila emylius (Cramer, 1775)
Calydna punctata Felder & Felder, 1861
Caria trochihis Erichson, 1818
Caria sponsa Staudinger, 1888
Caria mantinea (Felder & Felder, 1861)
Caria nr. mantinea

Chalodeta theodora (Felder & Felder,
1862)

Chalodeta chaonitis (Hewitson, 1866)
Chalodeta lypera (Bates, 1868)
Chamaelimnas briola Bates, 1868
Charis nr. anins
Charis cleonus (Stoll, 1782)

Charis anins (Cramer, 1776)

Charis sp.

Cremna actoris (Cramer, 1776)
Crocozona caecias (Hewitson, 1866)
Cyrenia martia Westwood, 1851
Emesis ocypore (Geyer, 1837)

Emesis nr. lucinda 1
Emesis nr.lucinda 2
Emesis sp.

Emesis temesa (Hewitson, 1877)

Emesis fatima (Cramer, 1780)

Emesis lucinda (Cramer, 1775)
Eshtemopsis celina Bates, 1868
Ennogyra satyrus Westwood, 1851
Eurybia silaceana Stichel, 1924
Eurybia latifasciata Hewitson, 1869
Eurybia lamia (Cramer, 1777)

Eurybia nicaeas Fabricius, 1775
Eurybia sp.

Eurybia jemima Hewitson, 1869
Eurybia dardus Fabricius, 1787
Eurybia cyclopia Stichel, 1910
Euselasia uria (Hewitson, 1855)
Euselasia urites gr.

Euselasia mirania (Bates, 1868)
Euselasia sp. 1
Euselasia sp. 4

35:42-60, 1996 (2000)

Eiiselasia sp. 2
Euselasia sp. 3

Euselasia pellonia Stichel, 1919
Euselasia oiilta (Cramer, 1777)

Euselasia opalescens (Hewitson, 1855)
Euselasia sp. 8
Euselasia lysias gr.

Euselasia melaphaea (Hubner, 1823)
Euselasia lysimachus (Staudinger, 1888)
Euselasia sp. 5
Euselasia sp. 6

Euselasia euriteus (Cramer, 1777)
Euselasia issoria Hewitson, 1869
Euselasia hygenius gr.

Euselasia hahneli Butler, 1874
Euselasia gelanor (Stoll, 1780)

Euselasia sp. 7
Euselasia fabia?

Euselasia everitus (Hewitson, 1855)
Euselasia euryone (Hewitson, 1856)
Euselasia nr. euriteus
Euselasia crotopus gr. 2
Euselasia euoras (Hewitson, 1856)
Euselasia eumenes (Hewitson, 1855)
Euselasia eumedia (Hewitson, 1855)
Euselasia eulione (Hewitson, 1856)
Euselasia crotopus gr. 1
Euselasia crinon Stizhel, 1919
Euselasia arbas (Stoll, 1782)

Euselasia anica gr.

Hyphilaria parthenis (Westwood, 1851)
Hyphilaria nicia (Hubner, 1819)
Ithomiola cascella (Hewitson, 1870)
Juditha molpe (Hubner, 1808)

Lasaia agesilas (Latreille, 1813)

Lasaia sp.

Lasaia pseudomeris Clench, 1972
Leucochimona nr. philemon
Leucochimona hyphea (Cramer, 1776)
Lyropteryx apollonia Westwood, 1851
Melanis xarifa (Hewitson, 1853)

Mesene nola Herrich-Schaffer, 1893
Mesene hya Westwood, 1851
Mesophthalma idotea (Westwood, 1851)
Mesosemia sp. 3
Mesosemia steli Hewitson, 1858
Mesosemia philocles Linnaeus, 1758
Mesosemia sp. 2

Mesosemia judicialis Butler, 1874

Mesosemia sp. 1

Mesosemia eumene (Cramer, 1776)
Mesosemia nr. judicialis
Mesosemia loruhama Hewitson, 1869
Mesosemia cippus (Hewitson, 1859)
Mesosemia nr. cyanira
Mesosemia nr. ephyne
Mesosemia sp. 5

Mesosemia sp. 4

Mesosemia melpia (Hewitson, 1869)
Mesosemia gertraudis Stichel, 1910
Mesosemia Ulrica (Cramer, 1777)
Mesosemia nr. thetys
Mesosemia nr. tenebricosa
Mesosemia magate?

Mesosemia nina (Herbst, 1793)
Metacharis lucius (Fabricius, 1793)
Metacharis nr. regalis
Metacharis regalis Butler, 1867
Methone cecilia (Cramer, 1777)
Monethe albertus Felder & Felder, 1862
Mycastor nealces (Hewitson, 1871)
Napaea melampia (Bates, 1867)
Notheme eumeus (Fabricius, 1781)
Nymphidium baoetia (Hewitson, 1853)
Nymphidium cachrus (Fabricius, 1787)
Nymphidium caricae (Linnaeus, 1758)
Nymphidium leucosia (Hubner, 1806)
Nymphidium nr. derufata
Nymphidium nr. lisimon
Nymphidium sp.

Nymphidium mantus (Cramer, 1775)
Nymphidium minuta gr.

Nymphidium omois Hewitson, 1865
Pandemos pasiphae (Cramer, 1775)
Parcella amarynthina (Felder 8c Felder,
1865)

Panics philotes Westwood, 1851
Panics nycteis Westwood, 1851
Perophthalma tullius Fabricius, 1787
Rhetus periander (Cramer, 1777)
Riodina lysippus (Linnaeus, 1798)
Sarota sp. 2

Sarota acantus (Stoll, 1782)

Sarota chrysus (Stoll, 1782)

Sarota sp. 3
Sarota sp. 1
Semomesia sp.

Setabis sp.

Setabis epitus (Cramer, 1780)

Setabis salvini?

Setabis buckleyi (Grose-Smith, 1898)
Stalachtis euterpe (Linnaeus, 1758)
Stalachtis calliope (Linnaeus, 1758)
Symmachia probetor (Stoll, 1782)
Symmachia sp.

Symmachia calligraphia (Hewitson,
1867)

Symmachia accusatrix Westwood, 1851
Symmachia asclepia Hewitson, 1870
Synargis gela (Hewitson, 1853)

Synargis sp.

Synargis abaris (Cramer, 1776)

Synargis chaonia (Hewitson, 1853)
Synargis orestesa (Cramer, 1780)

J. Res. Lepid.

Syngaris ochra (Bates, 1868)

Syrmatia aethiops Staudinger, 1888
Teratophthalma pheliiia (Felder &
Felder, 1862)

Themoiie pais (Hiibner, 1820)

Theope sp.

Theope eudocia Wesnvood, 1851
Theope lycaeniiia Bates, 1868
Theope nr. thootes
Theope virgilius (Fabriciiis, 1793)

Thisbe fenestrella Lathy, 1932
Xynias christalla Grose-Smith, 1902
unknown (8)

Lycaenidae 59

“Theda” hemon (Cramer, 1775)
“Theda” bosora Hewitson, 1870
“Theda” orobia (Hewitson, 1867)
“Theda” gigantea Hewitson, 1867
“Theda” maculata (Lathy, 1936)

“Theda” ciipentus (Stoll, 1781)

“Theda” gibberosa (Hewitson, 1867)
“Theda” tephraeus gr.

“Theda” ophia Hewitson, 1868
“Theda” tephraeus (Geyer, 1837)
“Theda” phegeus (Hewitson, 1865)
“Theda” nr. gadira
“Theda” nr. augustinula
“Theda” carteia Hewitson, 1870
“Theda” ergina or ligurina
“Theda” aruma (Hewitson, 1877)
“Theda” nr. mycon
“Theda” nr. empusa
“Theda” hesperitis (Butler and Druce,
1877)

Aiawacus dolylas (Cramer, 1776)
Arawacus aetolus (Sulzer, 1776)

Areas imperialis (Cramer, 1775)
Calycopis anapa Field, 1967
Calycopis indigo (Druce, 1907)

Calycopis isobeon complex
Calycopis cerata (Hewitson, 1877)
Calycopis xenata (Hewitson, 1877)
Calycopis pisis complex 3
Calycopis pisis complex 2
Calycopis atnius complex
Calycopis calus (Godart, 1824)

Calycopis centoripa Hewitson, 1868
Calycopis pisis complex 1
Celmia celmus (Cramer, 1775)

Chalybs janias (Cramer, 1779)
Contrafacia imma Prittwitz, 1865
Cyanophrys amyntor ?

Electrostrymon eebatana Hewitson, 1868
Eumaeus minijas (Hiibner, 1809)

Evenus gabriela (Cramer, 1775)
Hypostrymon asa Hewitson, 1873

lantheda leea Venables & Robbins,
1991

Jantheda sista Hewitson, 1867
Lamprospilus orcidia (Hewitson, 1874)
Mithras nautes (Cramer, 1779)

Ocaria ocrisia (Hewitson, 1869)

Ocaria thales (Fabricius, 1793)
Panthiades bitias (Cramer, 1777)
Panthiades aeolus (=pelion) (Fabricius,
1775)

Pseudolycaena marsyas (Linnaeus,
1758)'

Rekoa palegon (Cramer, 1780)

Siderus leucophaeus (Hiibner, 1818)
Strymon ziba (Hewitson, 1868)
Theclopsis lydus (Hiibner, 1819)
Theclopsis gargara Hewitson, 1868
Theritas mavors (Hubner, 1818)
Thestius pholeus (Cramer, 1777)
Tmolus echion (Linnaeus, 1767)

Zizula cyna (Edwards, 1881)

35:61-77, 1996 (2000)

journal of Research on the Lepidoptera

Flexural stiffness patterns of butterfly wings (Papilionoidea)

Scott J. Steppaii

Committee on Evolutionary Biology, University of Chicago, Chicago, IL 60637, USA., E-mail:
steppan@bio.fsu.edu

Abstract. A flying insect generates aerodynamic forces through the ac-
tive manipulation of the wing and the “passive” properties of deformability
and wing shape. To investigate these “passive” properties, the flexural
stiffness of dried forewings belonging to 10 butterfly species was compared
to the butterflies’ gross morphological parameters to determine allom-
etric relationships. The results show that flexural stiffness scales with wing
loading to nearly the fourth power and is highly correlated with

wing area cubed

The generalized map of flexural stiffness along the wing span for
Vanessa cardui has a reduction in stiffness near the distal tip and a large
reduction near the base. The distal regions of the wings are stiffer against
forces applied to the ventral side, while the basal region is much stiffer
against forces applied dorsally. The null hypothesis of structural isom-
etry as the explanation for flexural stiffness scaling is rejected. Instead,
selection for a consistent dynamic wing geometry (angular deflection)
in flight may be a major factor controlling general wing stiffness and
deformability. Possible relationships to aerodynamic and flight habit fac-
tors are discussed. This study proposes a new approach to addressing the
mechanics of insect flight and these preliminary results need to be tested
using fresh wings and more thorough sampling.

Key Wordsj biomechanics, butterfly wings, flight, allometry, flexural stiff-
ness, aerodynamics

Introduction

A flying insect generates aerodynamic forces primarily through the ac-
tive manipulation of wing movements and the “passive” morphological prop-
erties of deformability and wing shape. The morphological parameters of
insect flight have been the subject of various investigations (Weis-Fogh 1977,
Wootton 1981, Ellington 1984, Betts 1986, Dudley 1990, Srygley 1994),
complimenting an extensive body of work on the aerodynamics of insect
and hovering flight (e.g., Jensen 1956, Weis-Fogh 1973, Nachtigall 1974,
Ellington 1980, 1984b). However, empirical measures of aerodynamically
relevant mechanical properties of wings are absent from the literature.

Present address: Department of Biological Science, Elorida State University, Tallahassee, EL

32306-1100

Paper submitted 24 September 1997; revised manuscript accepted 8 May 1998.

J. Res. Lepid.

Various measures of wing geometry have been used as surrogates for the
biomechanical properties of wings, but these can be only crude approxi-
mations given the complex structure and construction of wings. Here, I
measure the deformability of butterfly wings to determine its interspecific
scaling relationships with various wing and body size parameters. This in-
vestigation complements qualitative analyses of structure and allometry,
theoretical predictions of wing properties, and observations of flight per-
formance and behavior.

Previous studies of insect flight have investigated the aerodynamics of
flight through theoretical calculations (Weis-Fogh 1977, Ellington 1980),
allometric patterns of wing shape and wing beat (Greenewalt 1962, Ellington
1984), wing movements and deformations during flight (Wootton 1981,
Betts 1986), flight habit and behavior (Betts & Wootton 1988, Dudley 1990,
Srygley 1994), the aerodynamic effects of angle of attack or presence of
scales (Jensen 1956, Nachtigall 1974, Martin & Carpenter 1977), and com-
mon structural features of butterfly wings (Wootton 1981). To date, no study
has measured deformability of wings. This study will demonstrate the po-
tential of biomechanical approaches to understanding insect flight.

Flexural stiffness {El) is a measure of deformability, which by controlling
wing shape under aerodynamic load modifies aerodynamic forces. The flex-
ural stiffness of a structure is a function of two properties: the elastic modulus
{E, stress per unit strain) of the material that composes it; and the second
moment of inertia (/) , a function of the cross-sectional geometry. This study
will 1) determine flexural stiffness patterns within butterfly wings, and 2)
define allometric relationships among flexural stiffness and morphologi-
cal parameters. Analysis of allometric patterns can provide insights into the
importance of developmental or structural constraints relative to presump-
tive adaptations (Strauss 1990),

Some expectations for flexural stiffness patterns can be drawn from pre-
vious studies. Betts (1986) found that in a small sample of Heteroptera,
angular deformation of the wing tip was weakly correlated with angular
momentum of the wing. A principal conclusion derived from Betts (1986)
and Wootton (1981) is that dorsal transverse flexion (producing a dorsally
concave surface) is more strongly resisted by wing structure (i.e., ventrally
stiffer) than is ventral transverse flexion. Wootton hypothesized that ven-
tral flexion may reduce drag on the upstroke of wings exhibiting minimal
wing-twisting, as in Lepidoptera. These studies would predict 1 ) that stiff-
ness will decrease in the distal region, possibly associated with a flexion line
(see Wootton, 1981 for detailed explanation), and 2) ventral stiffness (e.g.,
resistance to ventrally directed forces which would produce dorsal trans-
verse flexion) will be significantly greater than dorsal stiffness.

Two alternative hypotheses regarding interspecific scaling of flexural stiff-
ness are tested. the measured index of flexural stiffness is entirely a
mechanical consequence of structural and geometric isometry. Hp the in-
dex of flexural stiffness scales so that angular deflection under proportion-
ate loading regimes remains consistent (cf. elastic similarity; McMahon,

35:61-77, 1996 (2000)

1973), The predictions based on these hypotheses are presented in the
Discussion.

Materials m'D Methods

Species selected and morphometric measures

Three individuals for each of ten species were included among a mixed dry but-
terfly set obtained from Carolina Biologic Supply Company. The 10 species were
Battus polydamas Linnaeus 1758 (Papilionidae) Parides montezuma Westwood 1842
(Papilionidae) , Danaus lotis Cramer 1779 (Nymphalidae) , Phoebis statim Cramer 1777
(Pieridae), Eurema hecabe hlnn^eus 1758 (Pieridae), Pereute cAarops Boisduval 1836
( Pieridae ), Asda wowMste Linnaeus 1758 (Pieridae), Pyrr/wgyra wmcrm Linnaeus 1758
(Nymphalidae), the heliconiine Dione juno CiRmer 1782 (Nymphalidae), and the
pierid Catopsillia scylla Linnaeus 1764. Two living Vanessa cardui Linnaeus 1758
(Nymphalidae) were included, and their wings measured both immediately after
death and after three weeks of desiccation. Species were identified according to
Lewis (1974). For each specimen, total body mass and mass of the right fore- and
hindwing separately, were weighed with a Mettler H80 electro-balance (0.1 mg pre-
cision). Fore- and hindwings were drawn to scale using a camera lucida attached to
a Wild microscope at magnification x6. These outlines were then digitized to de-
termine wing area.

Flexural stiffness measures

The principal set of measurements consisted of force /deformation curves from
forewings under cantilever loading to produce transverse bending (Fig. 1). These
curves were generated for all 1 1 species. Cantilever loading was chosen over alter-
natives such as three- and four-point bending because, in natural flight, the base of
the wing is fixed relative to the body while the remainder of the wing is aerody-
namically loaded along its length as nearly perpendicular to the plane of the wing
as possible. The 10 dried species were compared for allometric patterns in wing
area (5), wing loading (dry body mass/wing area; rj, and flexural stiffness (El) as
a function of dry body mass (m) . Calculated wing loading will underestimate actual
wing loading because dried specimens were used. All wings were loaded both dor-
sally and ventrally. As described in this paper, loading from the dorsal direction
(dorsal loading) results in a dorsally convex surface, which is equivalent to ventral
transverse flexion in other studies.

Two Vanessa cardui adults were tested two to three days after emergence from chry-
salides, They were killed by pinching their thorax and then placed in a freezer for
five minutes, immediately after which they were weighed. After the V. cardui were
loaded in the tensiometer, they were allowed to dry for two to three weeks, then
weighed and loaded again to provide an estimate of the effects that drying had
produced upon the properties of the wings. A detailed map was made of stiffness
along the span of a single Vanessa cardui wing. Use of dried wings hinders accurate
estimation of flexural stiffness under natural conditions. For allometric studies
though, the effects of drying need only be consistent across taxa. If drying does
vary in its effects along the wing, this could bias interpretation of the wing maps

Basal attachment regions of individual forewings were glued using cyanoacrylate

J. Res. Lepid.

Figure 1. Diagrammatic representation of the method by which the wings were
loaded for the stiffness measures. The rectilinear loading bar was dis-
placed horizontally into the wing as indicated by the arrow. Measurements
were taken at specified distances perpendicular to the line between wing
base and tip. Remainder of the tensiometer apparatus not shown.

between two glass microscope slides. Spacers were placed between the glass slides
to prevent crushing of the wing. Only one to two millimeters were grasped in this
way, allowing the remainder of the wing to flex freely. Any discrepancy in the esti-
mate of the actual place of attachment will affect stiffness calculations near the base
much more than near the tip, because flexural stiffness varies with “beam” length
to the third power. For example, an underestimate of 0.4 mm at 10% of wing length
in the finely sampled Vanessa cardui (27 mm total length) would underestimate stiff-
ness by 30%, while the same error at 90% of wing length would only underestimate
stiffness by 5%. Wings were positioned with the span oriented perpendicular to the
loading bar (Fig. 1).

The other principal wing deformations of camber and torsion are very impor-
tant in wing aerodynamics, but are more difficult to measure accurately. Transverse
flexion is observed widely in lepidopteran wings (Wootton 1981) and is amenable
to experimental control. The loading bar was positioned using a millimeter scale
at predetermined distances from the secured wing base (20%, 40%, 60%, and 80%
of wing span) perpendicular to wing span. Another measurement was made at ap-

35:61-77, 1996 (2000)

proximately 0.5 mm less than 100% wing span because loading at 100% wing span
would result in the bar slipping off the wing. The wings were loaded in cantilever
bending by fixing the glass slide grips to the carriage of a tensiometer. The loading
bar, whose position could be adjusted with an accuracy estimated at ±0.4 mm, was
displaced horizontally into the wing from either the dorsal or ventral directions.
The diameter of the loading bar used in most measurements (including the de-
tailed mapping) was 1.0 mm. Some of the wings wore loaded with a 2.5 mm diam-
eter bar.

Fore wings were loaded in a tensiometer designed and assembled by M.
LaBarbera. Displacement of the wing at the loading bar was measured by an LVDT,
linear variable differential transformer (7307, Pickering, New York, USA), with a
linear range of 2.5 mm attached to the carriage of the tensiometer. Force was mea-
sured by a force transducer (FTD-6-10 10 g, Schaevitz, New Jersey, USA), accurate
to ±7x10'® N at the most sensitive setting. The LVDT was calibrated by inserting the
core rod a distance measured using an attached scale (±0.05 mm). The force trans-
ducer was calibrated by hanging known weights from the transducer when aligned
vertically. Force and displacement were recorded on a chart recorder (2200, Gould,
Ohio, USA). In regions of linear response of force to displacement, the slope was
used to estimate the force (F) and displacement (D). These variables were then
used to calculate flexural stiffness by the formula:

f:/=(f^lV(3*d) (1)

where Elis flexural stiffness in N m-, Fis force in Newtons, D is displacement at the
loading bar in meters, and L is the length of the wing segment under bending
(Wainwright et al. 1982) . This formula applies to a cantilever beam of uniform EL
The region between 60% and 80% of the wing span showed relatively constant stiff-
ness. An index of flexural stiffness, EI(W), was derived for each wing by averaging
the dorsal and ventral stiffnesses at 60% and 80% wing spans. Averaging these four
measures also reduced the expected error in Tithat were due to errors in position-
ing the loading bar.

It must be emphasized that each position’s Ells calculated assuming uniform ma-
terial properties throughout the section under load. Therefore, the maps of El do
not plot local stiffness, but rather the integral of stiffness of the wing up to that
position. Although this should not significantly affect the overall pattern, dorsal
versus ventral differences basally could obscure discrimination of differences dis-
tally. For example, ventral stiffness in the tip region may actually be greater than
that calculated for mean El, but deflections for a given load may be similar because
of greater deformation in the basal region under ventral loading.

Allometric patterns were determined by regressing morphological parameters
and the index of flexural stiffness. Species means were used rather than individual
measurements to avoid inflating the degrees of freedom in statistical tests, because
within species values are naturally correlated due to phylogenetic relatedness. Re-
duced major axes (RMA) were calculated rather than least squares regressions be-
cause RMA is more appropriate for allometric investigation (Rayner 1985).

Degree of distastefullness for each species to avian predators was provided by R.

J. Res. Lepid.

Figure 2. Log wing loading versus log dry body mass for 30 individuals repre^
seating 10 species. The RMA equation for the log-transformed data is
/^^=0.516 mm-o-3^1 (r^^O.SIO).

Sr)'gley (Chai 1986, 1988).

Results

Morphometric scaling

The slope of a regression line on a log-log plot defines the exponent in a
power function relationship of the form y=ax'L Log-transformed measures
of wing area {S) were regressed against log-transformed total mass for the
10 dry species means. Isometric scaling would produce a regression line with
slope of 2/3 The reduced major axis (RMA) slope obtained for

the 10 dry species means, 0.582, is not significantly less than 2/3 (r^=0.846) .
Wing loading shows weak positive allometry; wing loading scales with the
square root of mass (RMA=0.516, r~=0.81), almost significantly different
(P=0.06) from the null hypothesis of isometric scaling (all 30 indi-

viduals shown in Figure 2). Additionally, wing area scaled isometrically with
dry wing mass n=10), therefore, wings are not becoming proportion-

ately thicker (ignoring wing architecture like pleating). No strong conclu-
sions regarding any taxonomic pattern can be drawn, given the small sample
size, although nymphalids appear to have relatively higher wing loadings
than pierids.

Flexural stiffness maps

The effect of drying on wing stiffness was estimated by measuring two Vanessa
cardui wings immediately after killing the butterflies and then again after
two to three weeks of drying. Drying appears to significantly increase stiff-
ness, but the overall pattern of stiffness across the wing remains roughly

35:61-77, 1996 (2000)

Position

Figure 3. Effect of drying on wing stiffness averaged for the two Vanessa cardui.
El (10 ® kg m^) plotted on log scale. E/ values represent stiffness of en-
tire wing up to measurement point.

similar with peak stiffness in the middle region (Fig. 3). Differences do exist
between the patterns in the two condidons, primarily in the distal and proxi-
mal measurements (e.g., low dorsal stiffness at the tip for dry wings), and
these may be due small errors in positioning the bar. Stiffness decreases
rapidly in the distal 1.0 mm, and positioning errors are magnified in the
basal region because of the cubic relationship between length and stiffness.
Flexural stiffness {El) for 10 species was determined for five positions along
the wing both dorsally and ventrally. E/ values ranged over two orders of
magnitude from 2.3x10'^ N m^'to 1. 49x10'^ N m^ (Table 1). Because wings
varied so greatly in stiffness, values were normalized by dividing each wing’s
set of measurements by the maximum stiffness measured for that wing. The
10 wing maps so derived could then be compared as a proportion of maxi-
mum stiffness for each wing position. The normalized stiffness maps are
displayed in Figure 4a. A single factor ANOVA showed that for the pooled
data set (dorsal plus ventral), all adjacent positions (e.g. 40% with 20% and
60%) were significantly different in E/ except for the 60% and 80% pair.
The relative constancy in this region is one of the reasons that E/at 60%
and 80% were averaged to give E/( W) . An average wing is clearly stiffer under
dorsal loading along the basal 40% of wing span. More pronounced than
at 40%, the dorsally loaded wing is 55% stiffer at 20% of wing span
(P<0.001). The distal 40% is less stiff under dorsal loading than ventral load-
ing, but the difference is less pronounced and not statistically significant.

J. Res. Lepid.

5 "

”2

CJ3

l-H

1— 1

r-H

. ^

C/3

X— K

r—i

r-H

V-

■n

C/)

J>«l

c/3

i-H

■o

^ ^

X-

•D

r-H

r<H

r-H

— \

CX5

r-4

C/3

..LJ

*5.

J>-

r-H

■q

f-H

r*H

r— 1

r-H

f-H

Q-

M—

C/3

r“

cr>

^ -

l^-H

C/D

CM ’

’S

Jo-

f 3

r-H

•r—

1— 1

T-H

T—H

’?

CAl

0“

C/3

r— H

dry

H—

"i_

a (p)

• S2

E-

CM "

c/3

ecies

■g

’£

”S

Dionejuno

hS
' ^

’C

•S

g s
S q

g ‘C
•S .2

C/D

■0

f—

\—

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(—

-o

•o

35:61-77, 1996 (2000)

• Dorsal
□ Ventral

Figure 4a. Map of normalized stiffnesses for the mean values for 1 0 butterfly spe-
cies under dorsal and ventral loading. Original measurements were nor-
malized as a proportion of the maximum stiffness measured for each
wing. Standard deviation bars shown. Ail positions are significantly dif-
ferent from each other except 60% and 80% (P<0.05 ) . E/ values repre-
sent stiffness of entire wing up to measurement point.

Percent of Wing Span

Figure 4b. Mean ratios of dorsal versus ventral stiffnesses by wing position. Stan-
dard error bars shown. Only at 20% of wing span are dorsal and ventral
differences significantly different {P<0.001).

J. Res. Lepid.

The 60% position shows a significant difference only at the 90% confidence
level while the 40%, 80%, and 100% positions do not show significant dif-
ferences (P>0.2). The relative stiffnesses of dorsal versus ventral are sum-
marized in Figure 4b.

Figure 4a illustrates a possible common pattern across species but is a
rather crude map of dorsal and ventral flexural stiffness along wing span.
It also blends together slightly different stiffness patterns among species.
To complement this data set, a dried forewing of V. carduiw^s mapped with
much finer resolution, at approximately 1.4 mm intervals (Figure 5a). The
general pattern is in agreement with the averaged wing map. The wing is
dorsally stiffer (i.e., against ventral flexion) in the basal 60%, particularly
in the basal 40%. The distal 20% to 30% seems to be slightly stiffer ven-
trally. Wlien Elis plotted on a log-scale, two features stand out (Figure 5b).
First, stiffness from 40% to 85% of wing span is relatively constant compared
to the rest of the wing. Second, within the basal 25%, the wing is dorsally
much stiffer than ventrally; on average about three times stiffer. The accu-
racy of £7 estimates is lowest very near the base (e.g. <3 mm), due to small
errors in distance measures from the actual base of the wing.

Flexural stiffness and morphological parameters

The index of flexural stiffness, EI{W), was regressed against several com-
mon wing parameters, using mean values for each species. It was hypoth-
esized that by structural necessity, EI{ 14^ would be correlated with wing load-
ing, and indeed, E1(W) scales with wing loading to nearly the fourth power
(3.9) with a moderate correlation coefficient of 0.598. Longer, more heavily
loaded wings would need to be stiffer to prevent excessive deformation.
However, E1(W) is more strongly correlated with dry body mass (r-^0.814,
RMA slope=1.80) . The correlation of EI(W)with relative wing thickness (total
dry wing mass/ total wing area) drops to 0.417 (RMA s!ope=0.928). The
strongest correlation is with wing area; r-=0,91 1 (Figure 6) . £/fVF) scales with
wing area cubed (S^-/ 90% confidence interval, 2.46-3.73).

Figure 6 is slightly curvilinear. The power function provides a much bet-
ter fit to the data than a simple linear model (r^^O.785) which predicts zero
stiffness at 10 cm~. The remaining apparent curvilinearity is likely taxon spe-
cific. The two nymphalid species are approximately 40% less stiff than pre-
dicted by the regression, whereas the smaller of the papilioiiids is 64% stiffer
than predicted. These would result in deflections 60% more or less than
expected respectively.

The residuals from a polynomial regression constrained to pass through
the origin were compared for two groups: those palatable to birds and un-
palatable. The mean residuals were not significantly different between the
two groups (P>0.5), indicating that palatable butterflies do not have rela-
tively stiff wings.

Discussion

Various selective forces and phylogenetic constraints have been proposed
to account for insect wing morphology. The functional constraint of then

35:61-77, 1996 (2000)

% of Wing Span

Figure 5a. Wing stiffness map for a dry Vanessa cardui individual. Below is a
diagram of the wing, drawn to the same scale as the X-axis of the stiff-
ness map. The loading bar was oriented parallel to the Y-axis. El values
represent stiffness of entire wing up to measurement point.

moregulation may well have been significant during the early evolution of
insect wings (Kingsolver and Koehl 1985). However, thermoregulation is
probably of little importance to major scaling and structural patterns in
butterflies because only the proximal 15% of the wing surface plays a sig-
nificant role in conductive heat transfer to the body (Wasserthal 1975) and
the combination of pigmentation and behavior significantly effect ther-
moregulation in species that utilize the entire wing (Kingsolver 1985).
Strauss’ (1990) study of shape allometry in nymphalids suggests that aero-
dynamic (i.e., functional) constraints may be less important than sexual (i.e.,

J. Res. Lepid.

% of Wing Span

Figure 5b. Log-scaled wing stiffness map for a dry Vanessa cardui individual,
illustrating the relatively constant stiffness from 40% to 85% of wing span
and the large differences between dorsal and ventral stiffness basaliy.

Figure 6 . Index of flexural stiffness, Ei(W), versus wing area, S. Logdog scale.
RMA equation is lnE/W-3.1 S-9.78 (r2-0.911).

35:61-77, 1996 (2000)

display related) selection. Butterflies have unusually large wings used to
attract mates, to confuse or warn predators, for camouflage, and for other
display-related functions.

Scaling

The wing and body morphology measured in this study do not scale iso-
metrically among the butterfly species sampled. Although wing thickness
seems to scale isometrically, wing area shows a slight negative allometry with
dry body mass. As a consequence, wing loading shows positive allometry. In
addition, EI{ W) increases more rapidly than any of the other parameters,
and is most highly correlated with wing area. These results do not indicate
strong selection for an optimal wing loading that is size-independent.

The impact of allometrically induced variation in propulsion related forces
has been examined in other organisms. Because flying squirrel patagium
did not scale so as to minimize allometric variation in wing loading,
Thorington and Heaney (1980) concluded that other selective factors must
be involved, resulting in size related differences in gliding habit and ma-
neuverability. In response to isometric scaling, changes in the geometric
alignment and utilization of propulsive limbs in mammals can compensate
for size-dependent increases in mechanical stresses (Biewener 1989). These
compensations can significantly limit maneuverability and accelerative abil-
ity. Possible examples of compensation in butterflies include flight habit
and wing-stroke frequency. Indeed, Betts and Wootton (1988) found ten-
dencies in flight mode among a small sample of butterflies to be associated
with size and shape parameters of wings, including wing loading.

The results in this study can be compared to those reported elsewhere
(Greenewalt 1962, Kokshaysky 1977, Dudley 1990). Greenewalt’s analysis is
generally in accord with the wing area/body mass result, but in disagree-
ment with wing thickness. Greenewalt found that wing area increased with
the 0.60 power of wing mass, and thus wing thickness increased with the
1.34 power of wing span. The result from this study is almost significantly
different from Greenewalt’s figure (P<0.10). It should be noted at this point
that reanalyses of the original data (Magnan 1934, Sotavalta 1947) show a
slightly weaker relationship but a similar slope than he reported (r-’=0.702
versus 0.772; RMA=0.652 versus his mean regression line 0.634). The re-
analysis standardized sample sizes at one individual per species (n=20). As
Kokshaysky (1977) also noted, the number of data points graphed (35)
exceeded those listed in the regression table (33) and the number with
complete data (23).

Two hypotheses of flexural stiffness allometry were tested; structural isom-
etry and consistent dynamic wing geometry. For a beam with rectangular
cross-section, /, the second moment of area, is a product of width* thickness^.
Assuming isometry, width and thickness will be proportional to L, yielding
by substitution, la U. Area is proportional to L^, and thus, £/ should scale
with area S^. The hypothesis that El scales isometrically with wing area is
rejected because the allometric coefficient of 3.1 is significantly different
from 2.0 (P<0.02).

J. Res. Lepid.

Alternatively, aerodynamic constraints could result in angular deflection
remaining constant; i.e., £7 compensates for scaling in mass and wing area
so as to maintain a size independent dynamic wing geometry. This concept
is congruent with the elastic similarity which McMahon (1973, 1975) devel-
oped and applied to a variety of issues including tree shape and quadraped
locomotion. Deformation may be the most important structurally controlled
property of lepidopteran wings affecting aerodynamics. Greenewalt (1975)
argued that if wing thickness scales isometrically, angular deflection should
remain constant (since his results did not indicate isometry, he concluded
that angular deflection must show negative correlation with size). However,
under the assumption that deflection of the wing scales isometrically (D/
L=constant c), rearrangement of eq. 1 yields a prediction for EL

£/=F*LV3c (2)

If, instead of inputting the experimental force that was used to calculate
El, we assume that the principal forces acting on the wing are proportional
to body weight, and replace £/with EI(W), then eq. 2 predicts that £7 is
proportional to weight x wing area (L^, assuming on average, wing shape
scales isometrically). Multiplying wing loading by the area yields the total
force acting on the wing; total body weight. (In addition, the virtual mass
of the accelerated air can range from 0.3 [Diptera] to 1.3 [Odonata] times
the wing mass [Ellington, 1984]. Virtual mass has not been taken into ac-
count in this analysis.) The results are close to the prediction; £7scales with
(77z*3V (r^=0.882). The hypothesis of constant angular deflection cannot

be rejected.

Wing stiffness patterns

The reduction in distal stiffness matches the expectation of previous work-
ers. In Heteroptera, significant reduction in inertial stresses may be achieved
by lightening the fore wing distally (Betts 1986) , thereby reducing stiffness.
Betts views transverse [ventral] flexion as improving aerodynamics by “op-
timizing camber and angle of attack ..., minimizing adverse aerodynamic
forces at stroke reversal, ... creating favourable unsteady forces at stroke re-
versal” (1986, p. 298). Wootton (1981) felt that ventral flexion would pref-
erentially reduce drag on the upstroke. The hypothesis of a structural basis
for the limited dorsal flexion seen in previous studies is not strongly sup-
ported by the results of this study. The differences in the magnitude of £7
appear to be less than the difference between dorsal and ventral deflections
described by Betts and Wootton. Distal deflection will be affected by load-
ing distribution in addition to structural properties. Differences in distal
load may be due to differences in angular velocity, or related to the effects
of angle of attack stemming from camber and torsion elsewhere on the wing.
For example, Pieris supi nates its wings on the upstroke to an angle

of attack near zero, thus significantly reducing the force generated during
the upstroke (Ellington 1980).

35:61-77, 1996 (2000)

Perhaps the most striking result of the present work is the very low stiff-
ness near the wing base. The thickening of the veins and wing structure
observed near the base would be expected to increase the second moment
of area, /, and therefore flexural stiffness. Although the smaller chord width
near the base will reduce I, this reduction in width alone would seem insuf-
ficient to account for the magnitude of change documented here given that
thickness increases near the base would increase stiffness. Some functional
advantages may be suggested. Low ventral stiffness basally may permit wing
geometries that facilitate the “clap and fling” mechanism for generating
lift (see Weis-Fogh [1973] for description) . This stiffness pattern would seem
to be disadvantageous during normal flapping flight, where a stiff wing
would transmit muscle power to the surrounding air more efficiently. If
greater ventral flexibility is found to be aerodynamically disadvantageous,
then these results imply that the requirements for initial take off using clap
and fling impose the greater functional constraints and stronger selective
forces on wing design.

Alternatively, the low stiffness at the base relative to the center of wing
span may act to increase wing accelerations at stroke reversal in much the
same manner as a whip. This flexibility may also reduce inertial stress, es-
pecially at stroke reversal. Basal curvature appears greatest near stroke re-
versal in high speed photos of butterflies in flight (Dalton 1975). These
possibilities need to be tested further as well as testing whether the biome-
chanical properties of the glue and apparatus used to grasp the wing base
account for some of the reduced stiffness measured near the wing base.

No association was found between relative stiffness and palatability to avian
predators. A relationship might be expected if palatable species must be
stronger fliers to escape predators (Srygley 1994) and if stronger fliers have
stiffer wings. The findings here can be compared with those of Srygley
(1994) who found that palatability was most strongly associated with posi-
tions of centers of body and wing mass, which related to flight speed and
turning performance, but was less strongly associated with measures of wing
shape.

At present, improved understanding of the phylogenetic and ecological
contexts of butterfly flight are most needed in order to synthesize the bio-
mechanical and performance studies. There appears to be a strong phylo-
genetic component to relative wing stiffness, with the nymphalids having
relatively flexible wings and the papilionids having stiff wings. Future stud-
ies with greater taxonomic sampling should incorporate explicitly the phy-
logenetic relationships in order to avoid inflating significance levels, using,
for example, independent contrasts rather than raw species values in the
regression (Felsenstein 1985). Particularly important is the need to incor-
porate flight performance and flight habit parameters in studies such as
Betts and Wootton (1988) and Dudley (1990), along with structural bio-
mechanics and ecological correlates on comparable species.

The results of this study should be viewed as preliminary and subjected
to further testing and refinement. Fresh rather than dried wings must be

J. Res. Lepid.

measured to avoid the assumptions of proportional effects of drying, both
among species and across wings. Applying the load to the wing along a chord
of constant rotational radius may be preferable to the transverse orienta-
tion used here. Local rather than integrated stiffnesses should be measured.
The wing orientation chosen by Betts and Wootton (1988; fig, 2), which is
rotated approximately 20° posteriorly relative to this study, may be more
representative of loadings experienced during natural flight. The orienta-
tion used in this study is sometimes observ^ed at stroke reversal (Betts &
Wootton 1988). Furthermore, neither camber nor torsion were examined,
and deformations and wing movements usually involve all three. However,
this Study introduces an approach based on direct measurement of the bio-
mechanical properties of wings that has heretofore not been addressed.
Biomechanical studies are currently the missing link between studies of al-
lometry, flight performance, ecology, wing geometry, and theoretical aero-
dynamics.

Acknowledgements. Sincere thanks must go to M. LaBarbera for his generous contri-
butions of time, expertise, encouragement, and for the use of his experimental
equipment. R. Srygley generously provided the information on palatability. M.
Morgan, R, Srygley, K. Roy, R. Robbins, W. Watt, and four anonymous reviewers
provided suggestions that significantly improved the content of this manuscript.

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Chai, P. 1988. Wing coloration of free-flying Neotropical butterflies as a signal
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Journal of Research on the Lepicloptera

35:78-89, 1996 (2000)

The number of copulations of territorial males of the
butterfly Callophrys xami (Lycaenidae)

Carlos Cordero*

Instituto de Ecologi'a, Uiiiversidad Nacional Aiitonoma de Mexico, Apdo. Post. 70-275, 04510
Coyoacan, D. F., and Centro de Investigaciones Fisiologicas, Universidad Autonoma de
Tlaxcala, Apdo. Post. 262, 90070 Tlaxcala, Tlaxcala, Mexico. F-mail: ccordero@xolo.conahio.gob. mx

Rogelio Macias

Institute de Fcologia, A.C., Km. 2.5 antigua carretera a Coatepec, 91000 Xalapa, Veracruz,
Mexico.

Gabriela Jimenez

Institute de Fcologia, Universidad Nacional Autonoma de Mexico, Apdo. Post. 70-275, 04510
Coyoacan, D. F., Mexico.

Abstract. The number of copulations by different males and in different
territories was evaluated in the field in the butterfly Callophyys xami
(Lycaenidae). The total number of copulations per male and per hour was
very low (.0027 and .0029 copulations / male / h in 1989 and 1990,
respectively). There was high variance among males in the number of
copulations. Data from the few males observed copulating more than once
suggests a mating advantage for big, long lived males. Variation among
territories in the number of resident males, frequency of occupation and
number of copulations suggests variation in territory quality. Frequency of
occupation was not correlated with the territory variables measured, and
there were no differences in any territory variable between territories in
which copulations were observed and those in which no copulation was
observed. Furthermore, there were no between-years correlations in
frequency of occupation and number of copulations in the territories
studied in two different years. The location of territories maybe important
in determining territory quality.

Keywords; Callophrys xami, Lycaenidae, copulation, mating success, territo-
riality

Introduction

The fitness of male insects is difficult to determine in the field (Thornhill
and Alcock, 1983). Although the number of observed copulations has been
used frequently as a measure of male fitness (Thornhill and Alcock, 1983),
it is not possible to be confident about such a measure without knowledge of
male mating costs, female copulation frequency, spenn competition pat-

*To whom correspondence should be addressed

Manuscript accepted 14 March 1999.

35:78=89, 1996 (2000)

terns (Smith, 1984) and postcopulatory female choice criteria (Eberhard,
1996). However, some studies suggest that the number of copulations
achieved by a male is an important fitness component at least in some species.
One line of evidence supporting this suggestion is the fact that several aspects
of the male phenotype seem to be specific adaptations to increase the
number of copulations (reviews in Darwin, 1871; Thornhill and Alcock,
1983; Choe and Crespi, 1997).

In the butterfly Callophrys xami Reakirt (Lycaenidae) the number of
copulations seems to be an important fitness component for males since they
spend all their active adult lifetime defending territories that lack concentra-
tions of larval and adult food resources (Cordero and Soberon, 1990) and
that are used only as mating stations (Cordero and Soberon, 1990; Cordero,
1993). Laboratory observations suggest that copulation inhibits female
sexual receptivity for a number of days (if it is the first mating of the female,
these days correspond to the days in which oviposition rates are higher) , and
dissection of field collected females indicates a relatively low degree of
polyandry (Cordero and Jimenez, unpublished data).

In this paper we report field observations of copulations by territorial males
of the butterfly C. xami Differences in the number of copulations performed
by different males, and ocurring in different territories, are described and
some factors possibly affecting such differences are discussed.

Methods

Tlie study was conducted in the Pedregal de San Angel ecological reserve,
maintained by the Universidad Nacional Autonoma de Mexico in the south of
Mexico City. This zone is characterized by volcanic soil, rough topography, markedly
seasonal rainfall regime, and xerophytic shrubby vegetation. C xami is a multivoltine
butterfly that can be found throughout the year at relatively low numbers, reaching
its highest density from October to January (Soberon et at, 1988). The main larval
food plant in the study area is the perennial Echeveria gibbijlora (Crassulaceae), an
abundant species (Soberon et al, 1988; Larson et ai, 1994).

Study periods were chosen to coincide with population density “peaks” (Soberon
etai, 1988; personal observation); observations were made between November 1 and
December 20, 1989 and between November 10 and December 6, 1990. Most
territorial males observed were captured and individually marked on the wings with
felt-tip pens and their right forewing length was measured through the mesh of the
net with a calliper (in the laboratory, male wing length is correlated with adult body
weight at emergence: r = 0.91, p < 0.001, n = 28; Cordero, unpublished data).
Individuals were assigned to one of three wing wear categories: (1) similar to a
recently emerged adult (wings mostly green with intact margins), (3) very worn male
(wings mostly brown with worn margins), and (2) all individuals intermediate
between ( 1 ) and (3) . Longevity was defined as the number of days elapsed between
the first and the last observation of the male. Territory limits were determined as
explained in Cordero and Soberon (1990). We measured the (i) maximum length
and the (ii) “cross” length (length of the perpendicular axis crossing through the
middle point of (i)); territory area was approximated as (i) x (ii); the ratio (i) / (ii)
was used as a measure of territory “shape”.

The study period of each year was divided in two parts. During the first part we

J. Res. Lepid.

measured the frequency of occupation of each territory (= number of days the
territory was occupied by a territorial male / number of days the territory was
censused), determined the identity of each male defending the territory, and
recorded all copulations observed. In this part of the study we made observations in
25 territories in 1989 and in 19 in 1990. One observer walked along transectsjoining
several territories two times per day between 1000 and 1500 h, the daily territorial
defense period (DTDP; Cordero and Soberon, 1990), during 31 days in 1989 and
during 11 days in 1990, and observed each territorial male (if present) for at least two
minutes. The observation period was longer if, for example, the male was interacting
with a conspecific male or courting a female. The average number of days (±SD) each
territory' was censused was 26,2 ±1.8 (median = 26; range: 22--30) in 1989, and
10.2 ±0.6 (median = 10; range: 9“11) in 1990.

During the second part of each study period we estimated the probability of
copulating twice in a day (previous work indicated that the maximum number of
successful copulations per day that a male can achieve is two, since a male’s first
copulation of the day lasts 32 min on average, while the second copulation of the day
lasts several hours; Cordero, 1993). We made focal observations of territorial males
throughout the DTDP and recorded all copulations observed, during nine days in

1989 and 12 days in 1990, The number of males with focal observations was 15 in 1989
and 16 in 1990; the total number of hours of focal obsevations was 200 hours in 1989
(40 five hours periods of focal observations) and 130 hours in 1990 (26 five hours
periods of focal observations) . The number of days of focal obsen^ations per male
varied from 1 to 6 in 1989 (mean ± SD = 2.7 ± 1,7, median = 2), and from 1 to 3 in

1990 (1.6 ±0.8, median = 1). Focal observations were made in 14 territories in 1989
and in 1 1 territories in 1990. The number of days of focal observations per territory
varied from 1 to 6 in 1989 (2.9 ± 1.6, median = 2), and from 1 to 3 in 1990 (2,4 ±0.7,
median = 3). All observations were made on sunny days since C. xami is not active
under cloudy conditions. All summary statistics are given as mean ± standard
deviation and/or median and range (minimum-maximum).

Results

Throughout the study periods of 1989 and 1990, we observed territorial
males (Cordero, 1997) and sexually receptive females. All successful court-
ships observed (n = 15) began inside territories and involved territorial
males. Copulations were observed between 1100 and 1500 h in 1989 and
between 1230 and 1500 h in 1990.

Number of copulations by different males

We observed a total of 27 copulations (Table 1). Although we marked and
observed 159 territorial males (99 in 1989 and 60 in 1990), only 21 males
(three of them unmarked) were observed copulating (12 in 1989 and 9 in
1990), Three males were observed copulating more than once (two, three
and four times) . Only one male was observed copulating two times in a day
(this was the male that mated four times in 1989). We suspect that another
male mated twice in one day (this was the male that mated three times in
1989), since this male was observed arriving at the territory at 1111 h and
copulating at 1 137 h for more than 268 min (observation was interrupted at
1605 h); therefore, it is possible that this male copulated before the begin-

35:78-89, 1996 (2000)

Table 1. Distribution of copulations observed in different territories in each part
of the study periods of 1 989 and 1 990. T : territory.

1989 1990

Number of copulations: Number of copulations:

First part

Second part

First part

Second part

3-4N

Pnm

3^‘

8-9

ICh2

^ Territory in which a male copulated twice in a day.

’^Although in this territory no focal observations were made, we casually
observed one copulation during the second part of the study.

^ Territory in which no focal observations were made.

Territory in which a male probably copulated twice in a day.

ning of observations (focal observations began at 1004 and sometimes males
began territory defense before 1000 h) and that the long copulation ob-
served was the second of the day (remember that a male’s second copulation
of the day last several hours, while his first copulation of the day lasts on
average 32 min) . Therefore, only in one (possibly two) of the 40 five h periods
of focal observations in 1989 we observed two copulations; no male copulat-
ing twice in a day was observed in any of the 26 five h periods of focal
observations in 1990.

As expected from the different sampling methods employed during the
first and second part of each study period, the proportion of marked males
observed copulating in the first part of the study (10 / 144 = 6.9%; 8/92 in
1989 and 2/52 in 1990) was lower than the proportion observed copulating
during the second part (10 / 31 = 32.3%; 4/15 in 1989 and 6 /16 in 1990).
Seven of the nine copulations performed by the three males observed
copulating more than once were observed during the second part of the
study (including the two copulations performed in the same day by a male).

The number of copulations per hour calculated from the pooled focal
observations was similar in both years of study: 0.04 copulations / h
(= 8 copulations / 200 h of focal sampling) in 1989 and 0.046 copula-
tions / h (- 6 copulations / 130 h of focal sampling) in 1990. The number
of copulations per male and per hour calculated from the pooled focal
observations was almost identical in both years of study:
0.0027 copulations / male / h (= 8 copulations / 15 males / 200 h) in 1989
and 0.0029 copulations / male / h (= 6 copulations / 16 males / 130 h) in
1990.

J. Res. Lepid.

Characteristics of males

A total of 99 territorial males in 1989 and 60 in 1990 were individually
marked. No significant differences between years were found in wing length
(1989: 1.64 ± 0.1 cm, range: 1.36--1.89, n = 90; 1990: 1.65 ± 0.09 cm, range:
1.4-1.83, n = 55; t = -0.29, p = 0.77), longevity (1989: 4.8 ± 5.1 days, median
= 2, range: 1-20, n = 99; 1990: 4.9 ±5.9, median = 2, range: 1-28, n = 57; Mann-
Wliitney U = 2819, p = 0.99) and wing wear at the moment of being marked
(1989: median = 1, range: 1-3, n = 90; 1990: median = 1, range: 1-3, n = 57;
U = 2419.5, p = 0.51). There was no correlation between wing length and
longevity (1989: r^ = 0.13, p = 0.22, n = 90; 1990: r^ = 0.11, p = 0.43, n = 52).

Due to the sampling methods employed in this study, we cannot look for
a relationship between male traits and number of copulations in the data.
However, the characteristics of the three males observed copulating more
than once suggest that male size and longevity could be positively correlated
with copulation success. The male with the most copulations (four) was also
the biggest male observed in both years (wing length = 1.89 cm); this male
was also the only one observed copulating twice in a day. The longevity of this
male was 14 days, longer than that of 89.9% of the males observed in 1989.
The male that was observed copulating three times in 1989 was bigger (wing
length = 1.72 cm) than 73.3% of the males observed that year. The longevity
of this male was 11 days, longer than that of 85.9% of the males observed in

1 989. This male probably copulated twice in a day (see previous section) . The
male that was observed copulating two times in 1990 was bigger (wing
length = 1.72 cm) than 74.5% of the males observed that year. The longevity
of this male was 18 days, longer than that of 94.7% of the males observed in

1990. Therefore, the characteristics of the multiply mated males indicate
that a study of the possible (positive) effect of wing length and longevity on
male mating success would be particularly interesting in this butterfly (see
Appendix).

Number of copulations in different territories

The 17 copulations of 1989 and the 10 copulations of 1990 were observed
in seven territories each year, although only three of these were the same in
both years (Table 1). To explore the relation between territory variables
(maximum length, “cross” length, maximum length / “cross” length and
area) and the frequency of occupation of the territory, and to compare the
characteristics of territories in which copulations were observed with those
in which no copulations occurred, only the data obtained during the first
part of the study periods were analized. This decision was made considering
that during the second part of both study periods the sampling effort was very
heterogeneous (the number of days of focal observations per territory varied
from 1 to 6 in 1989 [CV = 55.2%] and from 1 to 3 in 1990 [CV = 29.2%]),
whereas during the first part of both study periods it was much more
homogeneous, and, therefore, comparable, between territories (the num-
ber of days each territory was censused varied from 22 to 30 in 1989 [CV =
6.9%], and from 9 to 11 in 1990 [CV = 5.9%]).

35:78-89, 1996 (2000)

Table 2. Comparison of characteristics of territories in which copulations were
observed with those of territories in which no copulations were observed. Values
are median (range). Statistics from Mann-Withney U test are given.

Territory variable

Year

Territories in
which copulations
were observed

Territories in

which no
copulations
were observed

Maximum

1989

5.17 (2.69-6.2)

3.26 (2.32-7.1)

24,5

length (m)

1990

3.81 (2.54-6.2)

3.88 (2.32-7.1)

.89

“Cross” length (m)

1989

3.27 (1.5-4.35)

2.5 (1.35-4.85)

.19

1990

3.02 (2.14-4.35)

2.68 (2-3.86)

.67

Maximum length /

1989

1.49 (.87-3.21)

1.16 (1.01-2.49)

.48

“Cross” length

1990

1.23 (1.03-1.84)

1.36 (1.01-2.25)

.67

Area (m-)

1989

16.93 (7.21-26.97)

7.78 (4.54-33.61)

.14

1990

10.85 (6.22-26.97)

8.48 (4.56-27.41)

.54

Frequency

1989

.69 (.52-.92)

.33 (.04-1)

21.5

.024

of occupation

1990

.85 (.64-.91)

.4 (0-1)

.36

^ This difference is not significant if we perform a sequential Bonferroni
adjustment of significance levels using as a family of tests (Chandler, 1995)
the five U tests of 1989, and using a = 0.1, as suggested by Chandler (1995):
k - 5, a/k = .02.

Average frequency of occupation of territories during the first part of the
study periods was 0.47 ± 0.29 (median = 0.5, range: 0.04-1) for 1989 and
0.6 ± 0.37 (median = 0.8, range: 0-1 ) for 1990. The frequency of occupation
of territories was not correlated with any of the territory variables measured
(Spearman correlations, all p > 0.26). Average number of copulations in
territories during the first part of the study periods was 0.36 ± 0.76 (median
= 0, range: 0-3) for 1989 and 0.24 ±0.44 (median = 0, range: 0-1) for 1990.
There were no significant differences between territories in which copula-
tions were observed and territories in which no copulations were observed in
maximum length, “cross” length, maximum length / “cross” length and area
(Table 2). The frequency of occupation of territories in which copulations
were observed was higher than that of territories in which no copulations
were observed in 1989 (Table 2), but no difference was detected in 1990.
However, even the 1989 difference is not significant if we perform a sequen-
tial Bonferroni adjustment of significance levels (see Table 2).

/. Res. Lepid.

Twelve territories were observed in both years. Considering only the data
collected during the first part of both study periods, there were no significant
between-years correlations in the frequency of occupation of these territo-
ries (r^ = 0.55, p = 0.078, n = 11) or in the number of copulations (Gamma
correlation, y = -0.09, p = 0.87, n = 11) observed in these territories.
Therefore, the “quality” of a territory in a given year was not a predictor of
that in the next. In fact, the territory that in 1989 had the maximum number
of observed copulations (five or, probably, six; Table 1) and the second
highest frequency of occupation (0.94; maximum = 1 ) , was not occupied by
a territorial male in any of the more than 10 days in which it was censused in
1990.

Discussion

Male copulation frequency

As is common in insects exhibiting lek territoriality {e.g. Alcock, 1983,
1987; Alcock and O’Neill, 1986; Table 3), the overall rate of copulations
observed in C. xami was low: 0.0027 and 0.0029 copulations / male / h in
1989 and 1990, respectively. Low copulation rates are expected since lek
mating systems are favored when receptive females are scarce and widely
dispersed (Thornhill and Alcock, 1983; Rutowski, 1991), and such condi-
tions seem to apply to the population of C. xami in the Pedregal de San
Angel (Cordero and Soberon, 1990).

This study suggests that there was relatively high variance in copulation
success between territorial males. First, although most males were not
observed copulating, some males copulated up to four times, including one
(probably two) male that was observed copulating two times in a day. Second,
one third of the copulations observed (nine out of 27) were performed only
by three males. Although we were not able to obtain estimates of male
lifetime reproductive success, these results, together with information indi-
cating that females exhibit a low level of polyandry (in a sample of 28 field
collected females, 78.6 % had only one or no spermatophore in their corpus
bursae, and the mean number of spermatophores found in non-virgin
females was 1.37 ±0.6 [Cordero and Jimenez, unpublished data]), suggests
that there is high variance in male fitness and, therefore, that the opportu-
nity for sexual selection in males is high. Sexual selection may be acting in
favor of an increase in male wing length and longevity if the mating
advantage suggested by the characteristics of the few males that mated more
than once is real. However, the relationships between male phenotypic traits
and copulation success still needs clarification.

Number of copulations and territory variables

The substantial variation observed between territories in frequency of
occupation, numbers of males and number of copulations suggests that
territories of C. xami vary in quality. However, none of the territory variables
measured affected the frequency of occupation or the number of copula-
tions (Table 2). In species with non-resource based territoriality, such as C.

35:78-89, 1996 (2000)

Table 3. Mating behavior of butterflies in which male copulation success and/or
phenotypic traits associated to male copulation success have been studied in the field ^

Species

d MF

d TSRMF

$ ME*^

$ TSRMF

Reference

Papilionidae

Papilio

polyxenes

.13 ±.49
(0-3)

ST""

1.3 ±.54
(0-3)

wvy"

Lederhouse
(1981, 1982)

Atrophaneura

alcinous

.43±L3P

(0-5)

ED'', L^
ME", WL"^"

1.0

—

Suzuki & Ma-

tsumoto (1992)

Luehdorfia

japonica

FA", WW^,

-1.0'

—

Tsubaki & Ma-

tsumoto (1998)

Nymphalidae

Coenonympha

pamphilus

.0198

.083

TB", WL""

.97 ±.05
(0-3)

—

Wickman

(1985)

Danaus

plexippus

2.98 ± 2.65
(0-11)

FA"^", PL'^",

3.50 ±1.22
(1-6)J

FA", PL''", Frey et aL

MT."", (1998)

Euphydryas

editha

Ednr

1.27 ±.46
(1-2)'

■—

Baughmann

(1991)

Heliconius

hewitsoni

—

BL", VOP,

1.0

—

Deinert et al.
(1994)

Pieridae

C philodice
eriphyle

—

1.21

(0-3)

WM'P"

Watt et al.

(1986)

Colias

eurytheme

—

(3)"

—

Watt et al.

(1986)

Pirns

napi°

PW"

2.03 ±.11
(1-5) p

—

Wiklund &

Kaitala (1995)

Lycaenidae

Jalmenus

evagoras

.97 ± 2.56
(0-7) ‘

L", ED",
WL"

1.0

—

Elgar & Pierce
(1988)

Callophrys

xami

.0027 (0-4)
.0029 (0-2)^

WL"", L""

1.37 ±.60
(0-3)

WL''",

This study
& Cordero

(1998)

/. Res. Lepid.

^ MS: male mating system according to the classification of Thornhill and Alcock (1983). LT:
lek polygyny. PM: pupal mating. SC: scramble competition polygyny. MF: mating frequency.
TSRMF: traits statistically related (’^), possibly related (^’^) or not related to MF. BL: body
length. ED: adult emergence date. FA: fluctuating asymmetry in forewing and hindwing radius
length. G: genotype. L: longevity. ME: mating experience. PL: parasitism level, PW: pupal
weight. ST: ME depends on specific territory. TB: territorial behavior (species with territorial
and non-territorial males) . \\3L: wing length (in the case of L.japonica this was measured as the
forewing and hindwing radius length). WW: wing wear.

^ Mean ± SD (range) of spermatophore number of mated females.

Mean ± SD (range) for the second brood of 1975.

Mean lifetime number of copulations ± SD (range).

^ Number of matings estimated by assessing degree of scale loss from claspers.

‘ fifty out of 51 field collected females had one spermatophore and one had two (Matsumoto
and Susuki,1995).

s Number of copulations/ male/ census. Upper figure: non-territorial males; lower figure:
territorial males.

^ Studied in a big outdoors mating cage.

' Mean lifetime number of copulations ± SE (range).

' Pliske (1973), cited in Drummond (in Smith, 1984), estimated a mean number of spermato-
phores (maximum) = 2.23 (8).

^ Relative number of matings estimated by marking male genitalia with powdered fluorescent

dye,

' Data from Ehrlich Sc Ehrlich (1978), cited in Drummond (in Smith, 1984).

Mean (range) (Drummond in Smith, 1984).

" Maximum number of spermatophores (Gwyiine in Smith, 1984).

“ Butterflies were raised in captivity and released in the field.

P Mean ± SE (range).

P Number of copulations / focal male / hour of focal observation (minimum number of
copulations per male - maximum number of copulations per male). Upper figure: 1989 study
period; lower figure: 1990 study period.

xami, it has been proposed that female “rules of movement” may be respon-
sible for territory location and quality (Bradbury, 1 985; Cordero and Soberon,
1990; Rutowski, 1991; Wickman et al, 1995). Although female movement in
C. xami\i?is not been studied, casual observations suggest that territories are
located in the confluence of natural or manmade trails, which are used by
females for their displacement through the habitat (Cordero and Soberon,
1990). If this suggestion is true, differences in territory quality may result
from the specific location of territories with respect to areas of high probabil-
ity of female transit, which may vary with time (as suggested by the lack of
between-years correlations in occupation frequency and number of copula-
tions in territories).

Male copulation success in other butterflies

Field estimates of male copulation success are scant. In Table 3 we

summarize the information on the mating behavior of butterflies in
which male copulation success and/ or phenotypic traits associated with male

35:78-89, 1996 (2000)

copulation success have been studied in the field. Unfortunately, a formal
quantitative comparison is prevented by the different methods employed to
estimate copulation success (Table 3).

The copulation success of males has been shown to be affected by a variety
of factors, such as weather conditions (Davies, 1978), adult emergence date
(Elgar and Pierce, 1988), body size (Deinert et aL, 1994; Elgar and Pierce,
1988), longevity (Elgar and Pierce, 1988), mating experience (Suzuki and
Matsumoto, 1992) , type ofbehaviour (territorial vs. non-territorial; Wickman,
1985), female mate choice (Rutowski, 1981-83) and fluctuating asymmetry
(Tsubaki and Matsumoto, 1998). A positive effect of body size on male
mating frequency has been found in two (four, if the possible cases of C.
pamphilus 3.nd C. xamiRve true) species (Table 3): Jalmenus evagoras3.nd Pieris
napi] while in Heliconius hewitsoni body length is negatively correlated with
mating success. The first species exhibits pupal mating, a mating system that
involves direct male-male competition (the same as lek polygyny, the mating
system of C. pamphilus and C. xami) , the second species exhibit scramble
competition polygyny, a mating system with indirect male-male competition,
and the third species also exhibits pupal mating. These data suggest that big
body size (or correlated traits) confer advantages in different male compe-
tition settings, although there may be situations in which small size may be
advantageous, as in H. hewitsoni. However, in Atrophaneura alcinous (Susuki
and Matsumoto, 1992) and Danaus plexippus (Frey et al, 1998), species
showing scramble competition polygyny, no relation between male size and
mating success was found. These observations are in accord with other
studies that indicate that male size and resource holding power are corre-
lated in some butterfly species (Rosenberg and Enquist, 1991) but not in
others (Alcock, 1994). In the three species in which it has been investigated,
a correlation between male longevity and number of copulations achieved
has been found {Atrophaneura alcinous and J. evagoras) or is suspected (C.
xami). These species have different mating systems, pertain to different
families, and have very different adult body sizes {Atrophaneura alcinous is
much bigger than the two lycaenids) .

Appendix

A prospective comparison of all marked males observed copulating (CM)
during the first and second parts of both study periods, with all marked males
not observed copulating (NCM) supports the suggestion that copulation
success may be correlated with wing length and longevity. CM had longer
wing length (CM: 1.71 ±0.1 cm, median = 1.72, range: 1.51--1.89, n = 14;
NCM: 1.64 ±0.1, median = 1.65, range: 1.36-1.88, n = 131; Mann-Whitney U
= 569, P = 0.0197) and lived longer (CM: 10.1 ± 5.6 days, median = 9,
range: 2-20, n = 18; NCM: 4.1 ±5, median = 2, range: 1-28, n = 138; U = 397,
P = lO"^) than NCM, but the degree of wing wear at the moment of being
marked was not different (CM: 1.4 ± 0.7, median = 1, range: 1-3, n = 15;
NCM: 1. 6 ±.7, median = 1, range: 1-3, n = 132; U = 845.5, P = 0.29). The mean
number of copulations of the CM was 1.3 ± 0.8 (median = 1, range: 1-4,

J. Res. Lepid.

11 = 18). We stress that these comparisons are based in data obtained from a
heterogeneus, non-random, and probably biased sampling of males.

Aknowledgements. We thank W. Eberhard, L. Egiiiarte, H. Drummond, J. L. Osorno,
C. Macias, R. Torres and two anonymous reviewers that provided useful comments
on previous versions of the manuscript. We also thank several friends (altruistic field
asisstants) for helping in the observation of butterflies. During the course of this
work the first author was supported by a Consejo Nacional de Ciencia y Tecnologia
(Mexico) scholarship.

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WiKLUND, C. & A. Kaitala. ( 1995) . Sexual selection for large male size in a polyandrous
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Journal of Research on the Lepidoptera

35:90-136, 1996 (2000)

Egg size in butterflies (Lepidoptera: Papilionoidea and
Hesperiidae): a summary of data

Enrique Garcia-Barros

Departnient of Biology, Universidad Autonoma de Madrid, 28049-Madrid, Spain, E-mail:
garcia. barros@sdi. uam.es

Abstract. A table summarising the estimated egg volumes, and adult wing
lengths, of 1184 species of butterflies is presented. The estimates were
primarily derived from published sources. They are expected to be rea-
sonable approximations, although some amount of measurement error
cannot be discarded. They may constitute a useful preliminary data base
for studies relating egg size to adult size.

Introduction

The allometry of egg, or neonate size, to adult body size has frequently
prompted the interest of evolutionary biologists. This relationship is of spe-
cial relevance, first, because of the many potential implications of body size
(Calder, 1984), And, second, because the ratio between body size and egg
size may have relatively direct effects on the number of eggs that can be
produced (that is, potential fecundity), depending on the availability of
resources and other anatomical, or physiological characteristics (e.g., Reiss
1989).

The allometry of egg to body size has been subject for study in several
kinds of organisms (examples in Reiss 1989), but work on arthropods, in-
cluding insect orders, is comparatively recent (Blueweiss et al. 1978, Berrigan
1991, Blackburn 1991, Reavey, 1992). The first attempts to determine the
interspecific correlation between egg size and ecological variables, or be-
tween egg size and parent body size in butteflies were those of Nakasuji
(1987) and Wiklund et al (1987), based in small sets of grass-feeding
Hesperiinae and Satyrinae, and north-European Pieridae. This not very
relevant number of comparative studies is in contrast with the potentially
important amount of descriptive material that has been published on but-
terfly life histories. While it is true that detailed data such as egg weight are
scarce, generalizations based on relatively rough estimates such as egg vol-
ume probably can make general patterns arise, and thus help subsequent
workers to determine the kind of data needed, and the taxa where such
patterns probably occur. After a first approach to the interspecific allom-
etry of egg to body size based on a selection of Holarctic butterfly species
(Garcia-Barros & Munguira 1997) it was evident that a wider sample, in-
cluding data from the tropical and subtropical areas, could be attempted
without much aditional effort. Consequently, I collected data on butterfly
egg descriptions to obtain approximate estimates of egg volume. These were
used in a comparative study that will be presented elsewhere. Because of

Paper submitted 24 November 1998; revised manuscript accepted 9 Februaiy 1999.

35:90-136, 1996 (2000)

the volume of the data base (which includes more than 1200 species), I
considered the possibility of presenting them as a separate publication rather
than as a summary of, e.g., genus, tribe, or family means. This has disad-
vantages (essentially, the amount of paper required), but also some advan-
tages, namely the accessibility of the data to other entomologists. This is
the main purpose of the present work. While the data are far from com-
plete, and probably not free of error, they will be available for further study
in a compyled printed form, and may help others to fill the gaps or even
serve as a basis for a more comprehensive, world-based data base on butter-
fly sizes. While this work might be used to compyle information on egg
morphology or related details, it is important to note that much descrip-
tive material that did not contain information on egg size is not quoted here.

Methods

The data collected included an estimate of egg size, and another of adult size,
for each butterfly species. The coverage of the sample was determined by the ac-
cessibility of the data (I do not intend it to be exhaustive). Data collection was ter-
minated when the number of species covered was judged to suffice for a prelimi-
nary comparative approach. Although some unpublished material was included,
the largest part of the estimates derives from the literature alone.

Egg volume. This was calculated using the formula for a regular ellipsoid using
the egg maximum diameter {ed) and length or heigth {el): Egg volume^ 1/
^{k) {edr) {eJ) . An empyrical approach demonstrated that egg volumes calculated
this way may constitute reasonable estimate for eggs with a rounded profile. The
fit was not so good for other egg shapes, and was specially rough for very ‘square’
profiled eggs. Because the descriptions available in the literature are frequently
based in very small sample sizes, a potentially important measurement error must
be presumed anyway. Wdien two or more descriptions (that included egg width and
length) were available, the average was calculated. Eggs described as spheroid,
spheric, or nearly spheric were considered to be of spheric shape unless this was
contradicted by existing figures, or by information form the most closely related
species. When scale figures were available, these were used to estimate the egg di-
mensions. Estimated egg volumes in mm^ are given in the table under the heading
EV.

Adult wing length. The length of the forewing, measured from the base to the
apex, was used to estimate adult size. Actually, the average between the male and
female sexes was estimated. Any available source was employed but, since collec-
tion material was available for a small part of the species, the live-sized specimens
photographed in the plates by D’Abrera (1977-1995) were used by default. Wlien
measurements from the adults used in the rearings, or from the same country or
geographic area as the egg descriptions were available, these were incorporated.
Estimated wing lengths in mm are given in the table under the heading WL.

Results

The data collected cover 1184 species. The results are presented in a single
table with five columns. The first of these is the species number (1 to 1184) .

J. Res. Lepid.

The second column corresponds to the species names (with a few excep-
tions, followed by the author and date of description). I am afraid that the
taxonomy had to be eclectic. The specific rank has been mantained for some
taxa that should probably be treated as subspecies (in case of duobt, this at
least ensures identification). The third and fourth columns give the esti-
mated egg volumes (EV) and wing lengths (WL). The last column (Sources)
gives numbered references; the sources are given after the table (see
‘Sources’).

Some further details on the taxonomic arrangement are given below (see
‘Taxonomic arrangement’). Numbers in small case (^ to ^^) refer to notes
on the taxonomy, sources for taxonomic arrangement, or other pertinent
detail, given at the end of the text. Finally, I have added one Appendix that
summarises the taxonomic (or, when phylogenetic approaches exist, da-
distic) relations between the species included. This is in parenthetical no-
tation, and the numbers correspond to those in the species list. This ar-
rangement is used in the comparative interspecific study of the data, that
will be presented separately. It may be of use to assess the effect of taxo-
nomic relationships in association with this data set, and it (or parts of it)
can be used as input for computer programs as a means to reconstruct the
tree structure adopted.

Species

Sources

Coeliades forestan (Stoll, 1782)

0.17

28.0

122

C. keithloa (Wallengren, 1857)

0.39

27.5

122

Allora dolleschallii (Felder, 1860)

0.42

22.5

219

Acleros mackenii (Trimeii, 1868)

0.31

15.0

122

Kedestes barberae (Trimen, 1873)

0.71

15.0

122

K. macomo (Trimen, 1862)

0.57

14.0

122

K. niveostriga (Trimen, 1864)

0.67

12.5

122

Cymaenes Iripunctus (Herrich-Schaffer, 1865)

0.25

13.5

Lerema comelius (Latreille, [1824])

0.95

18.0

Aeromachus inachus (Menetries, 1859)

0.11

12.5

182

Metisella malgacha (Boisdiival, 1833)

0.20

14.0

122

M. metis (Linnaeus, 1764)

0.15

13.3

122

Thoressa varia (Murray, 1864)

0.45

15.5

182

Tsitana uitenhaga Evans, 1937

0.51

15.5

122

Calpodes ethlius (Stoll, [1782])

0.65

25.0

129

Nyctelius nyctelius (Latreille, [1824])

0.82

18.0

45; 260

Panoquina nero (Fabricius, 1798)

1.87

22.0

129

P. sylvicola (Herrich-Schaffer, 1865)

0.15

18.0

41;45

Parosmodes moranlii (Trimen, 1873)

0.82

14.5

122

Borbo cinnam (Wallace, 1866)

0.06

14.0

182

B.fallax (Gaede, 1916)

0.38

16.0

122

B.fatuellus (Hopffer, 1855)

0.31

18.5

122

B. mpflr (Waterhouse, 1932)

0.31

17.0

297

B. lugens (Hopffer, 1855)

0.33

15.0

122

Gegenes niso (Linnaeus, 1764)

0.31

14.5

122

Pamara guttalla (Bremer & Grey, 1853)

0.17

18.5

158;182

P. monasi (Trimen Sc Bowker, 1889)

0.21

13.0

122

P. naso (Fabricius, 1789)

0.13

14.5

182

Pelopidas agna (Moore, [1866])

0.30

13.5

182

35:90-136, 1996 (2000)

P. jansonis (Butler, 1868)

0.35

18.0

182

P. mathias (Fabridus, 1798)

0.32

17.5

182

P. (Hubner, [1821])

1.06

22.0

122

Polytremis pellucida (Murray, 1874)

0.50

18.5

182

Isoteinon larnprospilusY eider 8c Felder, 1862

0.36

18.0

182

Amblyscirtes aenus Edwards, 1878

0.28

12.3

Lerodea eufala (Edwards, 1869)

0.30

13.0

Hidari irava (Moore, [1858])

1.56

27.8

295

Lotongus calathus (Flewitson, 1876)

3.66

24.5

295

Unkana ambasa (Moore, [1858])

3.32

31.3

295

Artitropa erinnys (Trimen, 1862)

2.51

24.0

122

Moltena fiara (Butler, 1870)

3.77

24.0

122

Zophopetes dysmephila (Trimen, 1868)

1.18

21.0

122

Ocybadistes knightorum Lambkin 8c Donaldson, 1994

0.15

10.7

280

Potanthus flavum (Murray, 1875)

0.30

15.0

182

Choranthus radians (Lucas, 1857)

0.15

14.5

Hesperia comma (Linnaeus, 1758)

0.43

15.3

53;156;233

H. nabokovi (Bell & Comstock, 1948)

1.06

19.0

209

Ochlodes ochracea (Bremer, 1861)

0.20

14.0

182

0. venatus (Bremer 8c Grey, 1862)

0.25

15.1

53;182;233

Poanes hobomok (Harris, 1862)

0.34

15.0

Polites baracoa (Lucas, 1857)

0.13

11.5

P. mystic (Edwards, 1863)

0.27

15.5

P. origenes (Fabricius, 1793)

0.34

14.5

P. sabuleti (Boisduval, 1852)

0.15

11.8

Pompeius vema (Edwards, 1862)

0.24

13.3

Wallengrenia egeremet (Scudder, 1864)

0.18

14.5

W. misera (Schaus, 1902)

0.26

14.0

Ancyloxypha (Fabricius, 1793)

0.14

11.5

129

Oarisma powesheik (Parker, 1870)

0.15

12.6

Thymelicus acteon (Rottemburg, 1775)

0.21

12.5

233

T. leoninus (Butler, 1868)

0.12

15.5

182

T. lineola (Ochsenheimer, 1808)

0.24

13.0

T. sylvaticus (Bremer, 1861)

0.13

13.5

182

T. sylvestris (Poda, 1761)

0.38

13.6

60;233

Carterocephalus palaemon (Pallas, 1771)

0.16

13.8

53;182;233

C. sylvicola (Meigen, 1728)

0.11

12.5

182

Leptalina unicolor (Bremer 8c Grey, [1852])

0.22

15.5

182

Anisynta Waterhouse 8c Lyell, 1912

0.15

14.0

108

Hesperilla crypsigramma (Meyrick 8c Lower, 1902)

0.69

13.7

121

H.furua Sands &: Kerr, 1973

0.69

14.4

121

H. malindeva Lower, 1911

0.89

18.0

121

H. Atkins, 1978

0.68

15.9

121;149

H. jexgwWato Herrich-Schaffer, 1869

0.55

14.7

121

Mesodina cyanophracta Lower, 1911

2.10

17.7

292

Neohesperilla crocea (Miskin, 1889)

0.20

14.9

251

N. senta (Miskin, 1891)

0.35

13.6

251

N. xanthomera (Meyrick & Lower, 1902)

0.28

13.3

251

N. xiphiphora (Lower, 1911)

0.40

10.5

251

Pasma tasmanica (Miskin, 1889)

0.39

19.2

186

Toxidia rietmanni (Semper, 1879)

0.27

17.2

186

Trapezites genevievae (Atkins, 1997)

1.27

18.0

293

T. Zacc/ioZrfes Waterhouse, 1903

1.15

18.6

178

T. maheta (Hewitson, 1877)

0.66

14.9

161

T. phygalioidesW2d.er\\ouse, 1903

1.60

15.9

178

T. praxedes (Plotz, 1884)

0.59

16.9

161

T. heteromacula Meyrick 8c Lower, 1902

0.52

13.4

114

100

101

102

103

104

105

106

107

108

109

110

111

112

113

114

115

116

117

118

119

120

121

122

123

124

125

126

127

128

129

130

131

132

133

134

135

136

137

138

139

140

141

142

143

J. Res. Lepid.

T. sciron Waterhouse & Lyell, 1914

0.37

15.0

231

T. waterhousei Mayo Sc Atkins, 1992

0.15

13.8

300

T. Atkins, 1997

0.81

21.0

293

Antipodia atralba (Tepper, 1882)

1.47

17.0

152

A. chaosloln (Meyrick, 1888)

0.83

16.3

152

Croitana arenaria (Edwards, 1879)

0.45

12.1

180

C. croites (Hewitson, 1874)

0.88

12.5

188

Herimosa albovenata (Waterhouse, 1940)

0.56

13.5

244

Proeidosa polysema (Lower, 1908)

0.75

14.0

Abantis paradisea (Butler, 1870)

0.34

21.5

122

Celaenorrhinus mokeezi (Wallengren, 1857)

0.28

20.0

122

Eagris nottoana (Wallengren, 1857)

0.34

14.3

122

Eretis djaelaelae (Wallengren, 1857)

0.17

14.0

122

Netrobalane Canopus (Trimen, 1864)

0.26

18.5

122

Anastrus sempitemus (Butler & Druce, 1872)

0.27

21.0

125

Erynnis afranius (Lintner, 1878)

0.25

16.0

E. tages (Linnaeus, 1758)

0.11

13.5

53;60;233

E. tristis (Boisduval, 1852)

0.30

18.9

E. zarucco Lx\c‘a&, 1857

0.15

20.0

Alenia sandasler (Trimen, 1868)

0.09

11.5

122

Carcharodus alceae (Esper, 1790)

0.20

14.0

304

Gomalia elma (Trimen, 1862)

0.25

13.5

191

Heliopetes ericetonim (Boisduval, 1852)

0.21

17.5

12;24

Hesperopsis libya (Scudder, 1878)

0.51

13.0

Pyrgus alveus (Hubner, 1803)

0.33

13.5

233

P. communis (Grote, 1872)

0.05

13.5

49;24

P. malvae (Linnaeus, 1758)

0.11

11.5

53;60;233

P. oileus (Linnaeus, 1767)

0.04

15.0

Spialia aslerodia (Trimen, 1864)

0.09

11.5

122

S. depauperata (Strand, 1911)

0.21

12.5

122

S. dromus (Plotz, 1864)

0.23

11.5

122

S. mafa (Trimen, 1870)

0.13

11.5

122

S. nanus (Trimen Sc Bowker, 1889)

0.13

11.0

122

S. sataspes (Trimen, 1864)

0.16

12.0

122

S. spio (Linnaeus, 1767)

0.21

12.5

122

Syrichtus proto (Oschsenheimer, 1808)

0.39

14.5

102

Sarangesa motozi (Wallengren, 1857)

0.20

17.0

122

S. phidyle (Walker, 1870)

0.17

16.0

122

Tagiades flesus (Fabricius, 1781)

0.25

23.5

122

Aguna albistria (Plotz, 1881)

0.30

16.0

135

Cabares potrillo (Lucas, 1857)

0.26

18.0

Codatractus aminias (Hewitson, 1867)

0.30

23.5

Poly gonus leo {GmeVin, [1790])

0.34

23.0

3;47

Typhedanus undulatus (Hewitson, 1867)

0.27

21.0

Urbanus dorantes (Stoll, [1790])

0.28

22.0

47;66;260

U. Simplicius (Stoll, [1790])

0.34

20.0

Archon apollinus (Herbst, 1798)

0.26

28.3

239

Elypermnestra hellos (Nickerl, 1846)

0.24

26.9

239

Pamassius szecheriyii Frivaldszky, 1886

0.45

33.0

239

P. Grum-Grshimailo, 1891

0.65

28.5

239

P. delphius (Eversmann, 1843)

0.47

31.0

239

P. imper ator Ohevt\\\\Y , 1883

0.68

40.1

239

P. autocr ator Avinoi^, 1913

0.67

36.0

239

P. /oxta5 Piingeler, 1901

0.64

33.5

239

P. waxmfnm Staudinger, 1891

0.71

36.0

239

P. acre Gray, 1853

0.59

27.9

239

P. 5mo Gray, 1853

0.72

24.5

239

35:90-136, 1996 (2000)

144

P. Staiidinger, 1889

0.80

25.5

239

145

P. hardwickii Gr3y, 1831

0.43

29.1

239

146

P. ariadne (Lederer, 1853)

0.97

34.5

239

147

P. glacialis Biider, 1 866

1.06

37.3

239

148

P. rnnemosyne (Linnaeus, 1758)

0.97

30.8

239;249

149

P. nordmanni (Nordmann, 1851)

1.30

33.0

239

150

P. stubbendorf a M.€n€tiies, 1849

0.66

32.5

239

151

P. actius (Eversmann, 1843)

1.01

31.0

239

152

P. apollo (Linnaeus, 1758)

1.12

39.5

239

153

P. apollonius (Eversmann, 1847)

2.54

39.5

239

154

P. epaphus OhertXiiir, 1879

0.76

27.0

239

155

P. Staudinger, 1882

1.85

39.0

239

156

P. jacquemontii'^oi&duv2L\, 1836

0.87

33.5

239

157

P. phoebus (Fabricius, 1793)

0.89

36.0

239

158

P. tianschanicus Oherthur, 1879

0.97

39.5

239

159

Sericinus montela Gray, 1853

0.20

37.0

295

160

Allancastria cerisyi (Godart, 1822)

0.17

28.0

239

161

Zerynthia polyxena (D. 8c Schiff., 1775)

0.20

28.3

162

Z. rumina (Linnaeus, 1758)

0.24

25.3

304

163

Bhutanitis linderallii Atkinson, 1873

1.05

59.0

295

164

Luehdorfia japonica Leech, 1889

0.53

31.0

295

165

L. longicaudata Lee -

0.60

35.0

295

166

L. puziloi (Erschoff, 1872)

0.41

29.5

295

167

Eury tides celadon (Lucas, 1852)

0.42

38.2

168

E. epidaus (Doubleday, 1846)

0.52

43.0

169

E. belesis (Bates, 1864)

0.52

46.0

170

Iphiclides feisthameli (Duponchel, 1832)

1.59

41.2

234;304

171

/. podalirius (Linnaeus, 1758)

1.58

39.5

60;234

172

Gmphium eurypylus (Linnaeus, 1758)

0.52

38.0

173

G. sarpedon (Linnaeus, 1758)

0.76

45.5

174

G. arisleus (Cramer, [1775])

0.29

39.6

200

175

G. antipathes (Cramer, [1775])

0.80

55.0

176

G. angolanus (Goeze, 1779)

0.38

41.1

259

177

G. antheus (Cramer, [1779])

0.50

48.0

259

178

G. leonidas (Fabricius, 1793)

0.64

47.5

259

179

G. morania (Angas, 1849)

0.50

37.2

259

180

G. policenes {Crmner , [1775])

0.50

40.0

259

181

Pharmacophagus antenor (Drury, 1773)

4.61

68.5

290

182

Battus polydamas (Linnaeus, 1758)

0.66

48.0

43;66;94

183

Troides aeacus (Felder & Felder, 1860)

6.12

81.0

295

184

T. amphrysus (Cramer, 1782)

7.64

77.0

295

185

T. andromache {Sta.udinger, 1892)

6.85

70.0

295

186

T. Helena (Linnaeus, 1758)

2.18

81.0

295

187

T. hipolitus (Cramer, [1775])

5.40

87.5

295

188

T. miranda (Butler, 1869)

6.69

91.0

295

189

T. brookiana (Wallace, 1855)

2.54

83.5

295

190

T. alexandrae (Rothschild, 1907)

20.5

100.0

87;295

191

T. priamus (Linnaeus, 1758)

9.36

91.5

295

192

Parides areas (Cramer, [1777])

0.68

39.0

94;101

193

P. childrenae (Gray, 1832)

0.71

48.5

194

P. iphidamas (Fabricius, 1793)

0.55

39.5

119

195

P. photinus (Doubleday, 1844)

6.79

47.5

196

P. alcinous (Klug, 1836)

1.55

54.0

239

197

P. horishana (Matsumura, 1910)

9.39

70.0

295

198

P. polyeuctes (Doubleday, 1842)

3.07

57.0

295

199

P. semperi (Felder & Felder, 1861)

5.93

73.5

295

200

P. varuna (White, 1868)

2.41

55.0

295

201

202

203

204

205

206

207

208

209

210

211

212

213

214

215

216

217

218

219

220

221

222

223

224

225

226

227

228

229

230

231

232

233

234

235

236

237

238

239

240

241

242

243

244

245

246

247

248

249

250

251

252

253

254

255

256

257

J. Res. Lepid.

Fachliopta aristolochiae (Fabriciiis, 1775)

1.32

48.0

295

Cressida cressida (Fabriciiis, 1775)

1.10

50.5

295

Papilio Fabriciiis, 1793

3.02

75.9

210;222;238

P. hellanichus We\\Ats,on, 1868

1.24

45.0

21;81

P. Linnaeus, 1771

0.83

48.5

P. anchisiades'Es^er, 1788 13

0.82

52.5

P. agestor {GxdLj , 1832)

1.83

56.5

295

P. epycides (Hewitson, 1862)

0.46

33.0

295

P. laglaizei Depuiset, 1877

0.38

51.0

109

P. clytia Linnaeus, 1 758

0.73

52.0

295

P. maraho (Shiraki & Sonan, 1934)

1.42

64.5

295

P. demolion Cramer, 1779

0.99

51.0

295

P. gigon Felder & Felder, 1864

0.91

70.0

295

P. cMc/icnor Guerin-Meneville, 1829

2.53

66.5

295

P. Linnaeus, 1758

0.66

50.5

259

P. a/cxanor Esper, 1799

0.39

42.0

235

P. indra Reakirt, 1 866

0.68

38.5

71;82;103

P. hospiton Genee, 1839

0.50

40.0

197;235

P. machaon Linnaeus, 1758

0.68

40.0

60;197;304

P. hippomdes Yelder 8c Felder, 1864

0.73

53.0

235

P. echerioides Trimen, 1868

0.75

42.8

259

P. ophidicephalus Oherthur, 1878

2.14

61.0

259

P. dardanus Ero\m, 1776

1.59

46.9

259

P. demodocus Esper, 1 798

0.96

46.0

259

P. demoleus (L.)

0.60

41.0

211

P. euphranor Tnm^n, 1868

0.97

54.3

259

P. pam Linnaeus, 1758

1.77

48.5

Dismorphia amphiona (Cramer, [1777])

0.28

33.0

132

D. foedora (Lucas) ^

0.38

29.0

D. spio (Godart, 1819)

0.51

33.0

260

Leptidea sinapis (Linnaeus, 1758)

0.15

20.2

53;60;233

Colotis antexnppe (Boisduval, 1836)

0.18

21.3

259

C. auxo (Lucas, 1852)

0.25

21.1

259

C. danae (Fabricius, 1775)

0.14

23.7

259

C. evagore (King, 1829)

0.08

18.3

259

C. evippe (Linnaeus, 1758)

0.11

20.5

259

C. erone (Angas, 1849)

0.17

26.0

C. ione (Godart, 1819)

0.08

25.5

C. vesta (Reiche, 1849)

0.08

21.5

C. eHs (King, 1829)

0.28

23.6

259

Eronia cleodora (Hubner, 1823)

0.28

29.2

259

Hehomia glaucippe (Linnaeus, 1758)

1.04

50.0

295

Nepheronia argia (Fabricius, 1775)

1.52

39.4

259

N. Imquettii (Boisduval, 1836)

0.67

29.4

259

Pareronia hoebera (Eschscholtz, 1821)

1.43

36.0

139

Anthocharis cardamines (Linnaeus, 1758)

0.16

21.1

233;53;60;233

A. cethura Felder & Felder, 1865

0.11

17.0

Euchloe ausonia Hubner, 1803

0.24

23.0

E. helemia (Esper, [1800])

0.12

22.9

175;281

E. craiwm Butler, 1869

0.13

23.3

175;281

E. Back, 1990

0.11

20.0

201

E. tagis (Hubner, [1804])

0.06

19.0

175;281

E. hyantis (Edwards, 1871)

0.11

21.0

Pinacopteryx eriphia (Godart, 1819)

0.09

29.3

259

Aporia crataegi (Linnaeus, 1758)

0.11

32.3

53;60;204

Appias epaphia (Cramer, [1779])

0.13

28.7

259

Belenois aurota (Fabricius, 1793)

0.12

25.8

259

35:90-136, 1996 (2000)

258

B. creona (Cramer, [1775])

0.12

27.0

259

B. gidica (Godart, 1819)

0.08

29.0

259

260

B. thysa (Hopffer, 1855)

0.30

33.4

259

261

B. zochalia (Boisduval, 1836)

0.13

28.7

259

262

Cepora nerissa (Fabricius, 1775)

0.08

29.5

295

263

Delias descombesi (Boisduval, 1836)

0.21

41.5

295

264

Dixeia charina (Boisduval, 1836)

0.08

22.2

259

265

D. pigea (Boisduval, 1836)

0.16

24.9

259

266

Ixias pyrene {LinndiQus, 1764)

0.19

33.5

295

267

Leptophobia caesia (Lucas, 1852)

0.41

27.0

268

Mylothris agathina (Cramer, [1779])

0.26

31.5

259

269

M. rueppellii (Koch, 1865)

0.28

31.5

259

270

M. ^nw??wBuder, 1869

0.24

27.8

259

271

Perrhybris lypera (Kollar, 1850)

1.00

35.3

272

Phulia nymphula (Staudinger? Blanch.?) ^

0.05

17.0

184

273

P. rosea '

0.07

15.0

184

274

Pieris brassicae (Linnaeus, 1758)

0.18

29.8

53;60;233

275

P. rapae (Linnaeus, 1758)

0.13

24.5

53;60;233

276

P. virginiensis¥.dyN2Lvds, 1870

0.10

24.4

184;206

277

P. napi (Linnaeus, 1758)

0.14

22.5

53;60;233

278

P. brionniae (Hubner, 1805)

0.31

22.5

172;208

279

P. callidice Wuhntr, 1800

0.05

23.7

184

280

P. occidentalis (Reakirt) ^

0.05

23.7

184

281

P. protodice (Boisduval & Le Conte, 1833)

0.05

23.4

184

282

Pontia daplidice (Linnaeus, 1758)

0.14

22.5

53;60

283

P. helice (Linnaeus, 1764)

0.12

23.0

259

284

P. chlorodice (Hubner, 1808)

0.13

22.0

285

Prioneris thestylis (Doubleday, 1842)

0.29

46.0

295

286

Tatochila mercedis (Eschscholtz, 1821)

0.04

27.0

184

287

T. microdice ’

0.10

21.0

184

288

T. sterodice ^

0.07

28.0

184

289

T. vanvolxemii (Capr.) ^

0.14

30.0

184

290

Leptosia alcesta (Stoll, 1870)

0.13

22.0

291

Anteos clorinde God3.Yt, 1823

0.16

44.0

292

Catopsilia florella (Fabricius, 1775)

0.21

35.0

259

293

C. pomona (Fabricius, 1775)

0.13

34.5

295

294

C. pyranthe (Linnaeus, 1758)

0.17

36.0

295

Colias alfacariensis Ribbe, 1905

0.13

24.5

53;62;181

296

C. croceus (Geoffroy, 1785)

0.13

25.5

53;207;233

297

C. electo (Linnaeus, 1763)

0.09

24.3

259

298

C. hyale (Linnaeus, 1758)

0.10

24.0

53;60;62

299

C. myrmidone (Esper, 1781)

0.10

23.0

207

300

C. palaeno (Linnaeus, 1761)

0.13

25.0

53;212

301

Eurema brigitta (Stoll, [1780])

0.06

20.4

259

302

E. jucunda '

0.03

24.0

303

E. lisa (Boisduval & Le Conte, 1829)

0.03

17.0

304

E. albula (Cramer, [1775])

0.41

18.5

305

E. desjardinsii (Boisduval, 1833)

0.10

20.0

306

E. hecabe (Linnaeus, 1758)

0.18

22.5

307

Gonepteryx cleobule (Hubner, 1825)

0.06

37.0

215

308

G. rhamni (Linnaeus, 1758)

0.21

28.1

53;60;205;233

309

Nathalis iole 'boisduval, 1836

0.02

14.3

310

Phoebis cipris (Fabricius) ^

0.08

32.5

311

Gandaca harina (Horsfield, [1829])

0.14

24.0

139

312

Abisara neophron (Hewitson, 1860)

0.06

23.8

294

313

Hamearis lucina (Linnaeus, 1758)

0.22

15.2

53;60;233

314

Euselasia hieronymi (Salvin & Godman, 1868)

0.05

15.0

136

315

316

317

318

319

320

321

322

323

324

325

326

327

328

329

330

331

332

333

334

335

336

337

338

339

340

341

342

343

344

345

346

347

348

349

350

351

352

353

354

355

356

357

358

359

360

361

362

363

364

365

366

367

368

369

370

371

J. Res. Lepid.

E. hygenius (Stoll, 1790)

0.06

13.5

279

Napaea beltimia Bates, 1869

0.09

17.0

227

N. orpheus (Hewitson, 1847)

0.08

14.3

Mesosemia acuta Hewitson, 1873

0.07

20.0

Calephelis borealis (Grote & Robinson, 1866)

0.05

13.5

C. nilus (Felder, 1861)

0.05

10.5

136

C. raivsoni McAlpine, 1939

0.06

12.5

136

C. perditalis (Barnes & McDunnough, 1918)

0.06

13.3

111

Caria ino (Godman & Salvin, 1866)

0.05

11.0

136

Staudinger, 1888

0.06

16.0

136

Metacharis ptolomaeus (Fabricius, 1783)

0.03

17.5

194;216

Panara thisbe (Fabricius, 1781)

0.03

19.0

Apodemia mormo (Felder & Felder, 1859)

0.27

15.5

24;136

A. nab' Edwards, 1874

0.22

15.5

136

A. palmeri (Edwards, 1871)

0.22

11.5

28;136

A. walkeri (Godman & Salvin, 1886) -

0.10

13.6

136

Emesis emesia (Hewitson, 1867)

0.05

13.0

136

E. mandana (Cramer, [1780])

0.15

22.5

136

E. tegula Godmmi & Salvin, 1886

0.21

19.5

136

E. tenedia Felder 8c Felder, 1861

0.11

19.0

136

E. zela Butler, 1870 ^

1.06

14.1

Audre susanae Oriila., 1935

0.18

15.5

A. colchis (Felder, 1865)

0.10

22.5

Lernonias caliginea (Clench, 1964)

1.00

17.9

L. epone (Godart, 1825)

0.16

18.5

Stichelia sagaris (Cramer, [1777])

0.13

12.0

194

Xenandra agria (Hewitson, 1847)

0.07

19.5

Cliaris calicene? (Hewitson, 1866)

0.08

13.0

Synargis phillone (Godart, [1824])

0.12

21.5

Baliochila aslanga (Trimen, 1873)

0.03

16.0

Durbania aniakos a Trimen, 1862

0.15

14.9

Durbaniopsis saga Trimen, 1883

0.17

16.8

Alaena amazoula Boisduval, 1847

0.15

13.5

A. margaritacea Eltringmn, 1929

0.17

12.8

Pentila tropicalis (Boisduval, 1847)

0.08

17.7

Lachnocnema bibulus (Fabricius, 1793)

0.03

13.5

L. durbani Trimen, 1887

0.04

12.8

Thestor basutus (Wallengren, 1857)

0.13

18.0

T. brachycerus (Trimen, 1883)

0.07

14.1

T. dicksoni Riley, 1954

0.07

21.1

T. dukeivan Son, 1951

0.08

14.5

T. holmesivan Son, 1951

0.08

16.8

T. protwnnus van Son, 1941

0.14

19.6

T. n/c)/? Pennington, 1956

0.09

17.2

T. yildizae van Son, 1941

0.07

16.0

Taraka hamada (Druce, 1875)

0.05

14.5

299

Curetis acuta Moore, 1877

0.29

22.8

299

Aloeides aranda (Wallengren, 1857)

0.21

14.8

A. clarkiTite 8c Dickson, 1968

0.21

14.5

A. damarensis (Trimen, 1891)

0.13

16.6

A. dentatis (Swierstra, 1909)

0.17

16.5

240

A. depictaTite 8c Dickson, 1968

0.21

18.3

A. gowaniTite 8c Dickson, 1968

0.26

17.7

A. pallida (Riley, 1938)

0.34

17.1

A. pierus (Cramer, [1779])

0.20

15.1

A. sp indet. ^

0.24

16.0

Aphnaeus hutchinsonii (Trimen, 1887)

0.38

18.3

35:90-136, 1996 (2000)

372

Ar^rocupha malagrida (Wallengren, 1857)

0.61

15.2

373

Axiocerses //oan^Grose-Smith, 1900

0.21

15.9

374

ChTfsoritis zeuxo (Linnaeus, 1764)

0.17

13.4

375

Crudaria leroma (Wallengren, 1857)

0.15

15.6

376

Oxychaeta dicksoni GAbriel, 1947

0.17

16.3

377

Phasis braueri Dickson, 1968

0.74

19.8

378

P. thero (Linnaeus, 1764)

0.74

18.8

379

Poecilmitis Pennington, 1962

0.23

14.4

380

P. anTim Pennington, 1953

0.17

12.8

381

P. Pennington, 1967

0.17

12.8

382

P.felthami (Trimen, 1904)

0.28

12.5

383

P. lycegenes (Trimen, 1864)

0.23

12.4

384

P. ly Sander Bennington, 1962

0.19

13.5

385

P. nigricans (Aurivillius, 1925)

0.21

13.9

386

P. palmus (Cramer, 1781)

0.19

12.8

387

P. pyroeis (Trimen, 1864)

0.28

14.1

388

P. thysbe (Linnaeus, 1764)

0.19

12.8

389

P. wrawm Pennington, 1962

0.24

13.8

390

Spindasis ella (Hewitson, 1865)

0.17

14.1

391

S. natalensis (Westwood, 1852)

0.18

17.0

392

S. to/eawowA Matsumura, 1906

0.11

14.7

299

393

Trimenia wallengrenii (Trimen, 1887)

0.89

19.5

394

T. arg^roplaga Tiickson, 1967

0.53

17.6

395

Tylopaedia sardonyx (Trimen, 1868)

0.42

22.0

85;141;153

396

Lycaena alciphron (Rottemburg, 1775)

0.11

18.6

60;302

397

L. clarki (Clark & Dickson, 1971)

0.05

14.3

398

L. (Haworth, 1802)

0.08

19.8

53;60

399

L, epixanthe (Boisduval & Le Conte, [1835])

0.14

12.8

163

400

L. helle (D. & SchifL, 1775)

0.07

12.8

401

L. helloides (Boisduval, 1852)

0.08

15.0

18;24

402

L. hippothoe (Linnaeus, 1761)

0.08

17.3

403

L. orus (Cramer, 1782)

0.06

10.5

404

L. phlaeas (Linnaeus, 1761)

0.06

14.6

405

L. tityrus (Poda, 1761)

0.10

14.3

53;60

406

L. virgaureae (Linnaeus, 1758)

0.20

16.4

407

Heliophorus epicles (Codart, [1824])

0.06

16.0

295

408

Arhopala muta (Hewitson, 1862)

0.12

16.5

295

409

A. bazalus (Hewitson, 1862)

0.09

17.3

299

410

A. ganesa Moore, 1857

0.09

16.0

299

411

A. japonica Murray, 1857

0.14

19.0

299

412

Acrodipsas illidgei (Waterhouse & Lyell, 1914)

0.14

13.9

199

413

Paralucia aurifera (Blanchard, 1848)

0.14

11.8

202

414

P. pyrodiscus (Rosenstock, 1885)

0.23

12.5

202

415

P. spinifera Ed\s’ards 8c Common, 1978

0.10

9.3

202

416

Theda betulae (Linnaeus, 1758)

0.28

19.3

53;60;233;261

417

Theda ? phydela Hewitson, 1 869

0.03

16.5

418

Cordelia comes (Leech, 1890)

0.11

15.5

295

419

Laeosopis roboris (Esper, [1793])

0.27

22.5

304

420

Ussuriana takarana (Araki & Hirayama, 1941)

0.11

23.0

295

421

U. stygiana (Butler, 1881)

0.20

22.5

299

422

Shirozua jonasi (Janson, 1877)

0.43

22.0

105;264;299

423

Artopoetes pryeri (Murray, 1873)

0.26

23.0

105;299

424

Coreana raphaelis (Oberthur, 1880)

0.14

19.5

299

425

Chrysozephyrus rarasanus (Matsumura, 1939)

0.31

19.0

295

426

C. hisamatsusanus (Nagami & Ishiga, 1937)

0.11

19.0

295;299

427

C. ataxus (Doubleday & Hewitson, 1852)

0.14

22.5

295;299

428

C. brillantinus (Staudinger) ^

0.40

20.0

261;299

100

429

430

431

432

433

434

435

436

437

438

439

440

441

442

443

444

445

446

447

448

449

450

451

452

453

454

455

456

457

458

459

460

461

462

463

464

465

466

467

468

469

470

471

472

473

474

475

476

477

478

479

480

481

482

483

484

485

/ Res. Lepid.

C. smaragdinus Bremer, 1864

0.32

18.5

299

Hahrodais grunus (Boisduval, 1852)

0.38

16.8

141;153

Neozephyrus quercus (Linnaeus, 1758)

0.18

16.3

233;261

N. japonicus Murray -

0.11

16.0

261;299

Sibataniozephyrus fujisanus (Matsumura, 1910)

0.25

16.8

250;299

S. kuafuiHsii & Lin, 1994

0.30

16.6

277

Iratsume orsedice (Butler, 1882)

0.13

18.8

105;299

Japonica adusta Riley ^

0.19

19.5

299

J. lutea (Hewitson, 1865)

0.24

20.0

261;299

J. saepestriata (Hewitson, 1865)

0.16

21.0

299

Favonius jezoensis (Matsimiura, 1915)

0.20

20.0

299

F. latifasciatus Shir ozu 8c Hayashi, 1959

0.16

20.0

299

F. orientalis (Murray, 1875)

0.18

19.8

299

F. saphirinus (Staudinger, 1887)

0.16

17.8

261;299

F. taxila (Bremer, 1861)

0.15

18.3

299

F, ultramarinus (Fixsen, 1877)

0.21

19.5

261;299

F. 'ywasai Shirozu, 1948

0.13

19.5

299

Araragi entheum (Janson, 1877)

0.11

15.3

105;299

Wagimo sign.atus (Butler, 1882)

0.18

15.5

261;299

Antigius attilia (Bremer, 1861)

0.19

17.0

261;299

A. hutleri (Fenton, 1881)

0.22

16.8

261;299

Ogyris genoveva Hewitson, 1853

0.13

18.0

Myrina dermaptera (Wallengren, 1857)

0.25

16.8

M. silenus (Trimen, 1879)

0.28

19.9

Eooxy tides tharis (Geyer, 1837)

0.32

16.0

236

lolaus situs (Westwood, 1852)

0.15

17.7

/. aemulus (Trimeii, 1895)

0.09

12.8

/. alienus (Trimen, 1898)

0.17

18.1

1. aphnaeoides (Trimen, 1837)

0.13

15.0

259

/. mimosae (Trimen, 1874)

0.15

15.8

1. sidus (Trimen, 1895)

0.-13

16.2

1. howkeri (Trimen, 1864)

0.21

18.1

Hypolycaena philippus (Fabridus, 1793)

0.05

16.5

Leptomyrina hirundo (Wallengren, 1857)

0.06

13.4

L. gorgias (Stoll, 1790)

0.19

14.5

L. tara (Linnaeus, 1764)

1.00

12.4

Capys alphaeus (Cramer, [1777])

1.34

20.0

C. dijunctus Trimen, 1885

0.31

18.5

Deudorix antalus (Hopffer, 1855)

0.17

16.6

I), dmoc/iflra Grose-Smith, 1887

0.28

18.0

D. Hewitson, 1869

0.37

19.5

D. epijarbas (Moore, 157)

0.19

21.0

Artipe eryx (Linnaeus, 1771 )

0.35

17.5

299

Rapala aurata (Bremer, 1864)

0.10

15.8

299

Callophrys rubi (Linnaeus, 1758)

0.13

13.8

60;233

C. avis (Chapman, 1909)

0.10

14.5

C. loki (Skinner)

0.13

13.2

C. nelsoni (Boisduval)

0.17

13.0

C. ferrea Butler, 1866

0.05

14.5

299

Evenus regalis {Crmner, [1775])

0.18

25.0

141;153

Satyrium pruni (Linnaeus, 1758)

0.17

16.4

233;299

S. iyonis Ota Sc Kusimoki, 1957

0.22

14.3

299

S. mera (Janson, 1873)

0.10

16.5

299

S. esculi (Hubner, [1806])

0.31

16.1

302

S. spini (D. & Schiff., 1775)

0.15

16.0

S. w-album (Enoch, 1782)

0.14

15.5

233;261;299

Eumaeus debora (Hewitson?) '

0.21

30.0

35:90-136, 1996 (2000)

101

486

E. toxea Godart, 1824 ^

0.45

22.0

487

E. minijas (Hiibiier, 1809)

0.46

22.5

75;141;153

488

Candalides cyprotus (Olliff, 1886)

0.11

14.8

179

489

C. gilberti Waterhouse, 1903

0.13

14.5

274

490

Anthene amarah (Guerin, 1847)

0.04

12.8

491

A. hutleri (Trimen, 1881)

0.10

14.3

492

A. definita (Butler, 1899)

0.03

14.3

493

A. kersteni (Cramer, [1780])

0.03

14.1

494

A. lemnos (Hewitson, 1878)

0.04

15.6

495

A. otacilia (Trimen, 1868)

0.12

12.2

496

A. sp indet. ^

0.17

12.5

497

A. talboti Stempier, 1936

0.06

12.5

498

Cupidopsis cissus (Godart, 1819)

0.08

17.6

499

C. jobates (Hopffer, 1885)

0.07

15.8

500

Pseudonacaduba sichela (Wallengren, 1857)

0.02

13.8

501

Nacaduba kurava (Moore, 1857)

0.04

12.0

288;299

502

Aciizera lucida (Trimen, 1883)

0.03

10.5

503

A. stellata (Trimen, 1883)

0.03

9.0

504

Cacyreus Pennington, 1962

0.06

11.8

505

C. lyngeus (Cramer, 1872)

0.04

14.7

506

C. marshalli 'Sutler, 1897

0.04

13.5

85;221

507

C. palemon (Cramer, 1782)

0.04

10.5

508

C. virilis (Aurivillius, 1924)

0.03

15.2

509

Harpendyreus nolohia (Trimen, 1868)

0.15

14.7

510

Lampides boeticus (Linnaeus, 1767)

0.04

16.5

85;112

511

Jamides alecto (Felder 8c Felder, 1860)

0.13

20.5

299

512

J. bochus (Stoll, 1782)

0.03

16.5

299

513

Leptotes brevidentatusE lie, 1958

0.04

13.8

514

L. cassius (Cramer, [1775])

0.03

12.0

127

515

L. pirithous (Linnaeus, 1767)

0.04

13.5

85;112

516

Tarucus bowkeri (Trimen, 1883)

0.08

13.5

517

T. sybans (Hopffer, 1885)

0.06

12.9

518

Tuxentius calice (Hopffer, 1855)

0.03

11.5

519

T. melaena (Trimen, 1887)

0.06

12.2

520

Zintha hintza (Trimen, 1864)

0.08

13.0

521

Zizeeria knysna (Trimen, 1862)

0.03

11.5

522

Z. maha Kollar, 1848

0.04

12.8

299

523

Zizina antanossa (Mabille, 1877)

0.06

13.0

524

Z. Otis (Fabricius, 1787)

0.04

10.5

299

525

Zizula hylax (Fabricius, 1775)

0.01

9.9

85;246

526

Brephidium exilis (Boisduval, 1852)

0.02

8.5

16;141;153

527

B. metophis (Wallengren, 1860)

0.02

9.0

528

Cupido lorquinii (Herrich-Schaffer, 1847)

0.04

10.2

189;302

529

C. minimus (Fuessly, 1775)

0.02

10.6

60;233

530

Everes argiades (Pallas, 1771)

0.03

13.0

60;299

531

E. comyntas (Godart, 1828)

0.06

14.5

532

E. lacturnus (Godart, [1824])

0.03

12.0

220;299

533

E. fischeri (Eversmann, 1843)

0.02

12.3

299

534

Pithecops corvusYruhstorier, [1919]

0.05

13.5

299

535

/*. Doherty, 1889

0.05

12.5

299

536

Azanus jesous (Guerin, 1847)

0.03

12.3

537

A. mirza (Plotz, 1880)

0.02

12.5

538

A. moriqua (Wallengren, 1857)

0.02

11.9

539

A. natalensis (Trimen, 1887)

0.02

13.2

540

Eiochrysops hippocrates (Fabricius, 1793)

0.03

10.8

541

E. messappus (Godart, 1819)

0.04

10.5

542

Celastrina argiolus (Linnaeus, 1758)

0.06

14.5

102

543

544

545

546

547

548

549

550

551

552

553

554

555

556

557

558

559

560

561

562

563

564

565

566

567

568

569

570

571

572

573

574

575

576

577

578

579

580

581

582

583

584

585

586

587

588

589

590

591

592

593

594

595

596

597

598

599

/. Res. Lepid.

C. sugitanii Matsiimura -

0.04

13.8

299

Actyolepis puspa (Horsfield, [1828])

0.04

15.0

299

Megisba malaya (Horsfield, 1828)

0.03

12.5

299

Udara alhocaerulea (Moore, 1879)

0.05

16.0

299

Cdaucof)syche alexis (Poda, 1761)

0.09

15.0

G. (Boisduval, [1828])

0.07

14.3

302

G. piasus (Boisdiival, 1852)

0.05

17.0

14;24;141;153

G. lygdamus (Doiibleday, 1841)

0.06

14.5

9;141;153

G. lycormas (Buder, 1868)

0.07

17.5

299

Maculinea iolas (Oscheriheimer, [1816])

0.06

18.9

189;302

M. alcon (D. & Schiff., 1775)

0.05

17.9

53;60;189

M. arion (Linnaeus, 1758)

0.05

19.9

4;53;60;189

M. nausithous {Bergstrasser, [1779])

0.06

16.7

189;302

M. rebeli (Hirschke, 1904)

0.05

16.9

53;189;302

M. teleius (Bergstrasser, [1779])

0.05

17.0

53;299

M. arionides Staudinger ^

0.10

19.5

299

Pseudophilotes abencerragiis (Pierret, 1837)

0.02

10.1

302

P. panoptes (Hubner, [1813])

0.03

10.8

302

P. bavnis (Eversmann, 1832)

0.04

14.0

263

P. barbagiae Prins 8c Porten, 1982

0.08

11.5

301

Euphilotes rita (Barnes & McDunnough, 1917) "

0.12

10.9

E. enoptes (Boisduval, 1852)

0.35

11.8

Philotiella speciosa (Edwards, 1877)

0.04

8.5

Sinia divina (Fixsen, 1887)

0.08

20.0

299

Scolitantides orion (Pallas, 1771)

0.08

12.5

299

Euchrysops barkeri (Triinen, 1893)

0.11

17.0

E. dolorosa (Triinen, 1877)

0.08

15.0

E. malathana Boisduval, 1833

0.04

17.8

E. osiris (Hopffer, 1885)

0.07

18.8

E. cnejus (Fabricius, 1798)

0.05

14.8

299

Lepidochrysops asteris (Godart, 1819)

0.06

19.7

L. bacchus Riley, 1938

0.06

14.2

77;85

L. dukei Cottrell, 1965

0.06

13.9

L. /sete Cottrell, 1965

0.06

16.6

77;85

L. methymna (Trimen, 1862)

0.09

19.4

77;85

L. oreasTite, 1964

0.09

14.5

L. patricia (Trimen, 1877)

0.06

21.5

L. puncticilia (Trimen, 1883)

0.05

14.0

77;85

L. trimeni (Bethune-Baker, 1823)

0.10

20.8

L. V ariabilis Colti'tW, 1965

0.06

17.1

77;85

Orachrysops lacrimosa (Bethune-Baker, 1923)

0.06

17.9

Oboronia omata (Mabille, 1890)

0.04

17.0

298

0. Stempffer, 1950

0.04

15.0

298

Polyommatus damon (D. & Schiff., 1775)

0.07

15.0

P. thersites (Cantener, 1834)

0.09

14.0

P. semiargus (Rottemburg, 1775)

0.05

15.0

P. albicans (Gerhard, 1851)

0.06

17.2

117;304

P. bellargus (Rottemburg, 1775)

0.05

15.8

60;233

P. coridon (Poda, 1761)

0.06

16.5

60;117;233

P. eros (Oschenheimer, 1807)

0.05

12.9

189;302

P. golgus {Uuhncr, [1813])

0.06

13.4

189;302

P. icarus (Rottemburg, 1775)

0.05

15.2

53;60;112;233;304

P. nivescens (Keferstein, 1851)

0.06

15.0

189;302

Ghilades trochilus (Freyer, 1844)

0.03

9.5

Plebeius pylaon (Waldheim, 1832)

0.05

14.7

189;302

P. argus (Linnaeus, 1758)

0.10

14.0

53;233;299

P. argyrognomon (Bergstrasser, [1779])

0.10

15.0

60;299

35:90-136, 1996 (2000)

103

600

P. (Linnaeus, 1761)

0,07

13.8

53;60

601

P. 5Mfco/arzM5 Eversmann, 1851

0.11

16.5

299

602

P. agestis (D. & Schiff., 1775)

0.04

13.0

233

603

P. artaxerxes (Fabriciiis, 1793)

0.04

12.0

233

604

P. morronensis Kihhe, 1910

0.03

11.9

189;302

605

P. nicias (Meigen, 1829)

0.11

13.2

189;302

606

P. an/CT-05 (Freyer, [1838])

0.07

14.0

263

607

P. emigdionis (Grinell, 1905)

0.12

11.4

608

Hemiargus hanno Stoll “

0.01

11.5

609

Niphanda fusca (Bremer & Grey, 1853)

0.07

20.5

299

610

Lihythea geoffroy Godart, 1 820

0.05

37.5

196

611

L. /aMara Westwood 8c Hewitson, 1851

0.13

25.3

259

612

Libytheana carinenta (Cramer, [1777])

0.06

22.8

2;24

613

Philaetria dido (Linnaeus, 1763)

1.00

50.0

64; 140

614

P. wemickei (Rober, 1905)

0.95

48.5

140

615

P. Pygmalion Fruhstorfer

0.95

48.5

140

616

Podotricha telesiphe (Hewitson, 1867)

0.39

38.0

140;273

617

Dryadula phaetusa (Linnaeus, 1758)

1.06

40.5

64; 140

618

Agraulis vanillae (Linnaeus, 1758)

0.41

37.6

64; 140

619

Dionejuno (Cramer, [1779])

0.30

38.4

64; 140

620

D. moneta (Hubner, [1821])

0.37

38.0

140

621

D. glycera Felder, 1861 ^

0.51

35.0

140

622

Dryasjulia (Fabricius, 1775)

0.66

43.5

64;66;75;140

623

Eueides vihilia Stichel, 1903

0.34

32.0

140

624

E. pavana (Menetries, 1857)

0.34

32.5

140

625

E. lineataS^iWin, 1868

0.30

34.5

140

626

E. procula Doubleday, 1847 ^

0.27

34.5

140

627

E. lampeto Bates, 1862 ’’

0.37

37.0

140

628

E. isabella (Cramer, 1781)

0.43

33.7

64;140

629

E. lybia (Fabricius, 1775)

0.26

31.5

140

630

E. tales (Cramer, [1780])

0.37

35.5

140

631

E. aliphera (Godart, 1819)

0.11

30.0

64; 140

632

Neruda godmani (Staudinger, 1882)

0.37

37.0

140

633

N. metharme (Erichson, 1848)

0.26

41.5

140

634

N. floerfc (Hiibner, 1816)

0.23

36.5

140

635

Laparus doris (Linnaeus, 1771)

0.36

40.0

64;140

636

Heliconius xanthodes (Bates, 1862)

0.47

38.5

140

637

H. li'a/Zam Reakirt, 1866

0.49

40.0

64;140

638

H. bumeyi (Hubner, 1816)

0.47

46.0

140

639

H. egeria (Cramer, [1775])

0.51

46.0

140

640

H. astraea Staudinger, 1 896

0.51

45.0

140

641

H. nattereriY elder, 1865 ^

0.37

41.0

140

642

H. numata (Cramer, [1780])

0.66

40.0

64;140

643

H. ismenius (Latreille, 1817)

0.44

43.5

140

644

H. pardalinus (Bates, 1862)

0.59

42.0

140

645

H. hecale (Fabricius, 1777)

0.70

44.0

128;140;182

646

H. ethilla Godart, 1819 ^

0.70

42.0

38;140

647

H. atthis (Doubleday & Hewitson, 1847)

0.44

40.5

140

648

H. cydno (Doubleday & Hewitson, 1847)

1.10

42.0

102;140

649

H. pachinus Sadvin, 1871

0.84

42.0

140

650

H. heurippa (Hewitson, 1854)

0.84

44.0

140

651

H. Hewitson, 1867^

1.08

42.0

140

652

H. Noldner, 1901

0.89

42.0

140

653

H. luciana

0.95

42.0

140

654

H. besekei Menetries, 1857

0.51

34.5

140

655

H. melpomene (Linnaeus, 1758)

0.76

38.0

64; 140

656

H. charitonius (Linnaeus, 1767)

0.49

40.0

128;140

104

657

658

659

660

661

662

663

664

665

666

667

668

669

670

671

672

673

674

675

676

677

678

679

680

681

682

683

684

685

686

687

688

689

690

691

692

693

694

695

696

697

698

699

700

701

702

703

704

705

706

707

708

709

710

711

712

713

J. Res. Lepid.

H. hermalhena (Hewitson, 1853)

0.33

41.0

140

H. erato (Linnaeus, 1758)

0.59

40.5

51;64;75;128;140

H. Latreille, 1817 '^

0.79

39.5

140

H. telesiphe (Doubleday, 1847)

0.54

42.0

140

H. ricini (Linnaeus, 1758)

0.31

34.4

64; 140

H. Staudinger, 1896

0.33

34.5

140

H. leucadia Bates, 1862

0.34

38.0

140

H. Sara (Fabricius, 1793)

0.28

33.0

64; 140

H. antiochus (Linnaeus, 1767)

0.37

42.0

140

H. Staudinger, 1875

0.40

37.5

140

H. congener Weyiner

0.26

38.0

140

H. eleuchia Hewitson

0.34

39.0

140

H. sapho (Drury, 1782)

0.26

37.0

140

H. hecalesia Hewitson, 1853

0.42

46.5

128;140

Argynnis paphia (Linnaeus, 1758)

0.38

33.0

53;60;233

Argyreus hyperbius (Linnaeus, 1763)

0.23

39.0

116

Brenlhis daphne (D. & SchifL, 1775)

0.55

23.0

B. hecate (D. & Schiff., 1775)

0.28

23.7

69;304

B. ino (Rottemburg, 1775)

0.27

20.0

B. mo/irfn Wyatt, 1969

0.35

26.1

241

Fabriciana adippe (Linnaeus, 1767)

0.28

27.5

53;60;233;304

F. niobe (Linnaeus, 1758)

0.30

28.0

53;304

F. auresiana (Fruhstorfer, 1908)

0.32

25.5

102

Issoria lathonia (Linnaeus, 1758)

0.12

21.0

53;60

Mesoacidalia aglaja (Linnaeus, 1758)

0.37

29.0

53;60;233

Pandoriana pandora (D. & Schiff., 1775)

0.08

34.0

304

Speyeria aphrodite (Fabricius, 1787)

0.15

33.0

S. atlantis (Edwards, 1872)

0.18

28.8

S. cybele (Fabricius, 1775)

0.27

39.8

S. idalia (Drury, 1773)

0.26

41.5

S. hydaspe (Boisduval, 1869)

0.22

27.9

S. nokomis (Edwards, 1862)

0.68

38.2

S. callippe (Boisduval, 1852)

0.22

29.3

Boloria aquilonaris (Stichel, 1908)

0.18

19.0

B. bellona (Fabricius, 1775)

0.21

20.0

B. dia (Linnaeus, 1767)

0.17

17.0

53;60

B. euphrosyne (Linnaeus, 1758)

0.19

21.3

53;233

B. selene (D. & Schiff., 1775)

0.15

20.8

53;60;233

B. eunomia (Esper, 1799)

0.14

19.5

Euploieta hegesia (Cramer, [1779])

0.19

30.5

Phalanta phalantha (Drury, 1770)

0.21

27.6

131

P. eurytis (Doubleday, 1847)

0.17

26.2

131

Acraea petraea Boisduval, 1847

0.15

25.4

A. oio/ornw Boisduval, 1847

0.23

20.5

A. nohara Boisduval, 1847

0.18

22.7

A. caldarena Hewitson, 1877

0.28

25.0

A. oncnen Hopffer, 1855

0.17

24.3

A. nata/icn Boisduval, 1847

0.23

31.5

A. zetes (Linnaeus, 1758)

0.32

34.0

A. neo^w/e (Doubleday, 1848)

0.19

27.5

70 '

A. horta (Linnaeus, 1764)

0.22

30.3

A. flganice (Hewitson, 1852)

0.19

35.8

A. (go/n Trimen, 1889

0.08

24.0

A. encedon (Linnaeus, 1758)

0.98

29.5

A. esebria Hewitson, 1861

0.12

29.3

A. eponina (Cramer, [1780])

0.12

21.9

A. ca/^fra Hopffer, 1855

0.12

23.8

35:90-136, 1996 (2000)

105

714

A. Hewitson, 1863

0.10

25.0

715

A. anacreon Tiimen, 1868

0.19

25.5

716

A. Boisduval, 1833

0.98

22.0

717

Pardopsis punctatissima (Boisduval, 1833)

0.21

16.8

718

Aglais urticae (Linnaeus, 1758)

0.20

23.8

60;233

719

Araschnia levana (Linnaeus, 1758)

0.99

18.0

720

Cynthia cardui (Linnaeus, 1758)

0.15

29.4

2;60;233;304

721

Inachis io (Linnaeus, 1758)

0.11

28.6

53;60;233

722

Nymphalis antiopa (Linnaeus, 1758)

0.24

33.8

2;53;60

723

N. milberti (Godart, [1824])

0.05

23.8

724

N. polychloros (Linnaeus, 1758)

0.23

30.5

60;233

725

Polygonia comma Harris, 1862

0.16

28.5

726

P. c-album (Linnaeus, 1758)

0.20

23.3

60;233;304

727

P.faunus (Edwards, 1862)

0.25

27.5

728

P. interrogationis (Fabricius, 1793)

0.45

29.0

729

P. progne (Cramer, [1775])

0.21

25.5

730

Vanessa atalanta (Linnaeus, 1758)

0.12

30.0

2;60;233

731

Antanartia schaeneia (Triinen, 1879)

0.47

26.2

131

732

A. hippomene (Hubner, 1806)

0.17

24.4

131

733

Amnosia Doubleday, 1849

1.05

43.5

295

734

Anartia amathea (Linnaeus, 1758)

0.15

26.0

130

735

A.fatima Godart, 1820 13

0.08

28.5

94; 130

736

A, jatrophae {Ukwwsiews, 1763)

0.10

20.0

44;66

737

A. lytrea (Godart, 1819)

0.18

30.5

738

Junonia coenia Hubner, 1822

0.07

25.5

739

J. evarete (Cramer, 1782)

0.17

27.5

740

J. oenone (Linnaeus, 1758)

0.15

25.6

131

741

]. hierta (Fabricius, 1798)

0.11

25.0

131

742

J. terea (Drury, 1773)

0.13

27.2

131

743

/. natalica Felder & Felder, 1860

0.18

27.2

131

744

Precis iphita (Cramer, [1779])

0.17

38.0

295

745

P. octavia (Cramer, [1777])

0.21

29.0

746

P. orithya (Linnaeus, 1758)

0.14

26.5

131;295

747

P. ceryne (Boisduval, 1847)

0.18

24.2

131

748

P. archesia (Cramer, [1779])

0.20

30.3

131

749

P. Trimen, 1879

0.27

29.5

131

750

Siproeta epaphus (Latreille, 1811)

0.53

44.6

93;94

751

S. slelenes (Linnaeus, 1758)

0.59

46.5

43;94

752

Catacroptera cloanthe (Stoll, [1781])

0.52

29.7

131

753

Protogoniomorpha parhassus (Druiy, 1782)

0.63

43.8

131

754

Hypolimnas misippus (Linnaeus, 1764)

0.15

45.0

131

755

H. deceptor {liiimQn, 1873)

0.15

39.5

131

756

H. anthedon (Doubleday, 1845)

0.15

44.5

131

757

Euphydryas aurinia (Rottemburg, 1775)

0.21

21.5

53;60

758

E. beckeri (Herrich-Schaffer, 1851)

0.16

23.4

304

759

E. maturna (Linnaeus, 1758)

0.14

21.0

53;60

760

E. phaeton (Drury, 1773)

0.21

28.0

761

Thessalia leanira (Felder 8c Felder, 1860)

0.34

19.7

762

Chlosyne harrisii (Scudder, 1862)

0.08

20.2

763

C. nycteis (Doubleday, 1847)

0.08

20.5

764

Melitaea britomartis (Assmann, 1847)

0.11

17.0

765

M. cinxia (Linnaeus, 1758)

0.09

19.8

53;60;233

766

M. diamina (Lang, 1789)

0.07

18.0

53;60

767

M. didyma (Esper, [1779])

0.19

22.0

53;60

768

M. phoebe (D. & SchifL, 1775)

0.08

23.2

157

769

Mellicta athalia (Rottemburg, 1775)

0.14

19.5

53;60;233

770

M. aurelia (Nickerl, 1850)

0.05

16.5

106

771

772

773

774

775

776

777

778

779

780

781

782

783

784

785

786

787

788

789

790

791

792

793

794

795

796

797

798

799

800

801

802

803

804

805

806

807

808

809

810

811

812

813

814

815

816

817

818

819

820

821

822

823

824

825

826

827

J. Res. Lepid.

M. Diiponchel, [1832]

0.18

20.9

79;304

Eresia eutropia Hewitson, 1874

0.17

25.0

100

Phyciodes campestris (Behr, 1863)

0.14

17.6

2;34

P. thaws (Drury, 1773)

0.05

17.5

Atlantea tulita (Dewitz, 1877)

0.26

31.0

260

Colobiim dirce (Linnaeus, 1764)

0.36

36.0

31;54

Historis acheronta (Fabriciiis, 1775)

1.53

47.5

260

H. odius (Fabriciiis, 1775)

0.91

57.5

303

Smyrna blomjildia (Fabriciiis, 1782)

0.32

40.0

Sea sophronia (Godart, [1824])

0.26

32.0

258

Eunica bechina (Hewitson, 1852)

0.21

30.2

229

Sallya natalensis (Boisduval, 1847)

0.09

26.2

131

S. boisduvali (Wallengren, 1857)

0.11

23.3

131

S. trimeni (Aurivillius, 1889)

0.09

23.8

131

Ariadne merione (Cramer, [ 1 777] )

0.12

32.9

295

Eurytela dryope {CAY2LmeY, [1775])

0.27

28.5

131

E. hiarbas (Drury, 1782)

0.27

25.9

131

Byblia acheloia (Wallengren, 1857)

0.13

25.0

131

B. ilithyia (Drury, 1773)

0.13

25.0

131

Hamadryas fehrua (Hubner, 1823)

0.65

36.0

Adelpha cc/mo Bates, 1864

0.35

30.5

A. iphiclus (Linnaeus, 1758)

0.28

28.0

A. syma (Godart, [1824])

0.34

22.5

Cymothoe alcimeda (Godart, [1824])

0.30

27.0

131

Limenitis archippus (Cramer, [1775])

0.34

44.0

L. arthemis (Drury, 1773)

0.69

39.0

L. Camilla (Linnaeus, 1764)

0.39

28.0

60;193

E. populi (Linnaeus, 1758)

1.03

40.0

181

L. reducia (Staudinger, 1901)

0.26

25.3

170;304

Pseudacraea lucretia (Cramer, [1775])

1.41

35.8

131

P. boisduvalii (Doubleday, 1845)

4.61

43.0

131

Pantoporia hordonia (Stoll, 1790)

0.73

24.5

295

P. peiius (Linnaeus, 1758)

0.70

30.0

Neptis praslini Boisduval, 1832

0.48

29.0

185

N. saclava (Boisduval, 1833)

0.27

23.8

131

N. /ac/a Overlaet, 1955

0.58

27.5

131

Bebearia orientis (Karsch, 1895)

0.82

32.7

259

Dophla evelina (Stoll, 1790)

11.5

54.0

295

Euthalia amanda (Hewitson, 1862)

2.71

40.5

295

Lexias dirtea (Fabriciiis, 1793)

2.71

46.5

295

Mahaldia formosana (Friihstorfer, 1899)

2.87

45.0

295

Tanaecia iapis (God^LYt, [1824])

2.59

33.0

295

Hamanumida daedalus (Fabriciiis, 1775)

1.54

31.6

131

Marpesia petreus (Cramer, 1778)

0.13

40.0

Cyrestis pantheus (Lathi, 1901)

0.38

28.5

259

Charaxes bernardus (Fabriciiis, 1793)

2.74

46.5

295

C. varanes (Cramer, 1764)

1.58

40.5

23;86;131;195

C. fulvescens (Aurivillius, 1891)

1.86

47.9

C. paphianusWmd, 1871

0.90

29.4

195

C. zoolina (Westwood, 1850)

0.50

27.8

131;195

C. Grose-Smith, 1883

1.77

37.8

195

C. candiope {God3.Yt, [1824])

1.65

44.6

23;131;195

C. jasius (Linnaeus, 1767)

2.74

42.9

304

C. epijasius Reiche, 1850

0.98

47.3

23;195

C. satumus EudeY, 1865

3.00

45.4

131;195

C. pelias (Cramer, [1776])

2.14

41.1

52;131;195

C. castor {CYmiicY, [1775])

2.14

52.5

23;131;195

35:90-136, 1996 (2000)

107

828

C. hrutus (Cramer, [1779])

3.23

42.2

23;131;195

829

C. pollux (Cramer, [1775])

1.54

39.1

131;171;195

830

C. dowsetti Henning, 1988

4.15

50.8

195

831

C. druceanus Bntler, 1869

3.61

41.4

131;171;195

832

C. numenes (Hewitson, 1865)

1.77

47.7

23;195

833

C. tiridates (Cramer, [1777])

4.19

56.1

23;195

834

C. bohemaniY eider, 1859

5.24

44.7

195

835

C. xiphares (Cramer, 1781)

3.02

51.2

131;195

836

C. wawfima Rothschild & Jordan, 1901

4.19

51.8

195

837

C. cithaeron Felder, 1859

3.59

45.5

23;86;131;195

838

C. achaemenes F elder 8c Felder, 1867

2.14

38.5

131;195

839

C. etesipe (Godart, [1824])

4.19

41.8

23;195

840

C. jahlusal^rimen, 1862

1.13

26.6

131;195

841

C. eupale (Drury, 1782)

0.52

31.2

23;195

842

C. dilutus Rothschild, 1898

0.52

28.3

23;195

843

C. anticlea (Drury, 1782)

0.29

27.5

23;195

844

C. baumanni Rogenhoier, 1851

0.28

27.6

23;195

845

C. catachorus Staudinger, 1896

0.52

36.8

195

846

C. etheodes (Cramer, [1777])

0.52

37.3

23;195

847

C. mariepsYan Someren & Jackson, 1957

3.05

39.3

195

848

C. karkloof\an Someren &Jackson, 1957

1.77

35.7

195

849

C. pondoensis Van Someren, 1967

2.14

33.2

195

850

C. nyikensisVan Someren, 1975

3.05

38.7

195

851

C. ethalion Boisduval, 1847

1.61

35.4

131;195

852

C. cedreatisHeW\Xs,on, 1874

0.52

38.0

195

853

C. chintechiVan Someren, 1975

1.77

36.8

195

854

C. chittyiRydon, 1980

1.15

34.3

195

855

C. howarthi Minig, 1976

1.15

34.3

195

856

C. fulgurata Aurivillius, 1889

0.52

33.3

195

857

C. phaeus Hewitson, 1877

1.77

34.6

195

858

C. fionae Henning, 1977

1.77

27.3

115;195

859

C. viola Butler, 1865

0.90

31.9

195

860

C. Butler, 1881

0.90

39.1

195

861

C. vansoniVan Someren, 1975

0.90

31.6

23;129;195

862

C. berkeleyiVan Someren & Jackson, 1957

1.77

34.7

195

863

C. martini Yan Someren, 1966

1.44

33.0

143;195

864

C. gallagheri Yan Son, 1962

1.44

35.2

195

865

C. guderiana (Dewitz, 1879)

1.77

32.8

195

866

Prothoe calydina (Hewitson, 1855)

2.02

50.5

295

867

Agrias amydon (Hewitson, [1854])

3.56

44.0

155

868

A. daudina (Godart, [1824])

14.9

45.5

166

869

Archaeoprepona demophoon (Linnaeus, 1758)

7.92

52.4

113

870

Noreppa chromus (Guerin, 1844)

3.22

51.5

871

Prepona omphale (Hubner, 1819)

5.04

48.4

30;32;98

872

Fountainea ryphea (Cramer, [1775])

0.52

29.5

248

873

Consul fabius (Cramer, [1776])

0.52

36.9

104

874

Euxanthe eurinome (Cramer, [1775])

4.19

45.5

23;195

875

E. wakefieldi (Ward, 1873)

1.19

45.0

131;195

876

E. Grose-Smith, 1889

4.19

52.0

195

877

Palla usheriButler, 1870

0.63

42.8

23;195

878

Apatura iris (Linnaeus, 1758)

0.70

37.5

60;233

879

Asterocampa celtis (Boisduval 8c Le Conte, [1835])

0.45

25.6

57;187

880

A. leilia (Edwards, 1874)

0.42

24.1

881

A. dyton (Boisduval 8c Le Conte, [1835]

0.20

27.6

2; 187

882

A. texana (Skinner, 1911) ®

0.37

28.3

65;187

883

A. idyja (Geyer, [1828])

0.29

31.2

174;187

884

Sephisa princeps (Fixsen, 1887)

0.99

39.0

282

108

885

886

887

888

889

890

891

892

893

894

895

896

897

898

899

900

901

902

903

904

905

906

907

908

909

910

911

912

913

914

915

916

917

918

919

920

921

922

923

924

925

926

927

928

929

930

931

932

933

934

935

936

937

938

939

940

941

j. Res. Lepid.

Morpho anaxibia (Esper, 1777)

7.00

73.0

M. catenanus (Peri7% 1811)

1.52

63.0

M. Hercules (Dalman, 1861?) ^

2.15

71.5

M. menelaus (Linnaeus, 1758)

7.78

88.0

M. peleides Kollar, 1 850

2.09

68.0

223

Antirrhaea philoctetes (Linnaeus, 1764)

4.09

47.0

214

Amathusia phidippus (Linnaeus, 1763)

1.68

55.0

295

Faunis canens (Hubner, [1819])

1.38

37.0

295

F. phaon (Erichson, 1834)

1.30

38.5

295

Taenaris artemis (Snellen, 1860)

1.65

47.5

287

T. catops ? ^

1.65

47.0

287

T. onolaus ? *

1.77

47.0

159

Thauria aims (Westwood, [1858])

5.05

58.0

295

Zeuxidia amethystus Butler, 1865

2.66

56.0

295

Z. aurelius (Cramer, [1777])

6.41

70.5

295

Z. doubledayi Westwood, 1851

3.57

59.0

295

Pierella hyalinus (Gmelin, 1788)

0.52

44.0

213

Melanitis constantia (Cramer, [1777])

1.77

42.0

265

M. leda (Linnaeus, 1763)

0.66

39.0

61;182

Gnophodes parmeno (Butler, 1880)

0.82

36.5

Kirinia roxelana (Cramer, [1777])

0.18

31.3

263

Lasiommata maera (Linnaeus, 1758)

0.56

24.9

53;60;262

L. megera (Linnaeus, 1767)

0.38

22.4

53;60;233;262

L. petropolitana (Fabricius, 1787)

0.40

21.4

262

Lethe diana Butler, 1866

0.52

27.0

182

L. dura (Marshall, 1882)

0.86

38.5

295

L. europa (Fabricius, 1775)

1.38

34.0

295;182

L. Fruhstorfer, 1914

0.24

33.3

278

L. rohria (Fabricius, 1787)

0.85

33.5

295

L. sicelis (Hewitson, 1866)

0.70

33.0

218

L. verrna (Kollar, 1884)

0.58

27.0

295

Lopinga achine (Scopoli, 1763)

0.72

26.5

262

Neorina lowii (Doubleday, [1849])

3.55

57.0

295

Pararge aegeria (Linnaeus, 1758)

0.36

21.9

53;60;233;262

P. xiphia (Fabricius, 1775)

0.92

26.1

262

P. xiphioides (Staudinger, 1871)

0.49

25.3

262

Satyrodes eurydice (Johannsen, 1763)

0.60

23.8

S. portlandia (Fabricius, 1781)

0.45

30.0

Aeropetes tulbaghia (Linnaeus, 1764)

0.76

46.0

Paralethe dendrophilus (Trimen, 1862)

0.52

38.0

Zethera pimplea (Erichson, 1834)

1.79

42.5

295

Elymnias agon das ? *

1.54

42.5

162;243

E. casiphoneGeyer, [1827]

2.22

46.0

295

E. melias (Felder, 1863)

1.77

43.0

295

E. nesaea (Linnaeus, 1764)

1.95

37.0

295

Bicyclus anynana (Butler, 1879)

0.52

21.2

259

B. safitza Hewitson, 1851

0.57

25.5

Mycalesis anaxioides (Marshall, 1883)

1.97

32.0

295

M. gothama Moore, 1857

0.55

25.5

182

M. maiaenas Eiewltson, 1864

0.96

25.5

295

M. perseus (Fabricius, 1775)

0.49

23.7

245

M. sirius (Fabricius, 1775)

0.58

24.8

245

M. terminus (Fabricius, 1775)

0.56

24.8

245

Orsotriaena medus (Fabricius, 1775)

0.52

23.0

265

Henotesia perspicua (Tri m e n m 1873)

0.47

21.5

Ragadia luzonia Felder & Felder, 1863

0.28

23.3

147

Acrophtalrnia artemisFelder Sc Felder, 1861

0.07

18.8

147

35:90-136, 1996 (2000)

109

942

Hypocista angi/jteto Waterhouse & Lyell, 1914

0.11

18.0

190

943

H. irius (Fabricius, 1775)

0.27

19.0

190

944

Tisiphone helena (Olliff, 1888)

1.41

36.0

232

945

Ypthima asterope (Moore, 1857)

0.39

24.0

176

946

Y. impuraElwes 8c Edwards, 1873

0.47

19.0

259

947

Y. loryma Hewitson, 1865

0.27

20.5

295

948

Y. praenubila Leech, 1891

0.70

30.4

295

949

Coenyra aurantiaca Aurlvillius, 1910

0.37

20.0

950

C. hebe (Trimen, 1862)

0.47

17.0

259

951

Melampias stenipteraVan Son, 1955

0.42

19.3

259

952

M. huebnerivm Son, 1955

0.47

20.0

953

Strabena tamatavae (Boisd., 1833)

0.70

19.0

183

954

Physcaeneura panda (Boisduval, 1847)

0.72

19.0

955

Cassionympha cassius (Godart, 1823)

0.55

19.0

956

Neita durbani (Trimen, 1887)

0.63

21.0

957

N. extensa (Butler, 1898)

0.73

23.5

958

Pseudonynpha hippia (Cramer, 1782)

0.69

19.5

959

P. trimenii (Butler, 1868)

0.65

21.0

960

P. magus (Fab., 1793)

0.34

19.5

961

P. magoidesM^n Son, 1955

0.47

21.0

962

P. detectaliYivaGin, 1914

0.44

19.5

963

Stygionympha vigilans (Trimen, 1887)

1.38

24.5

964

S, wichgrafivan Son, 1955

0.58

22.5

965

S. irrorala (Trimen, 1873)

0.47

17.5

966

Megisto cymela ^

0.31

20.2

967

Neonympha areolala (Abbot 8c Smith, 1779)

0.69

18.1

968

Taygetis andromeda (Cramer, [1779])

1.15

35.0

164

969

Coenonympha oedippus (Fabricius, 1787)

0.13

21.0

182

970

C. arcania (Linaeus, 1761)

0.49

18.1

53;60;233;262

971

C. glycerion (Borkhauseii, 1788)

0.29

17.0

972

C. hero (Linnaeus, 1761)

0.23

16.8

145

973

C. iphioidesSxsLwdingtY, 1870

0.42

19.1

262;304

974

C. leander {Esper , [1784])

0.29

16.7

304

975

C. tullia (Muller, 1764)

0.23

18.9

53;233

976

C. Oberthur, 1881

0.40

14.7

192

977

C. corinna (Hubner, 1804)

0.30

15.0

284;304

978

C. dorus (Esper, [1782])

0.53

16.7

262;304

979

C. elbana Staudinger, 1901

0.17

13.3

285

980

C. pamphilus (Linnaeus, 1758)

0.20

16.6

53;60;123;233;262

981

C. saadiKoWm, 1848

0.07

17.0

148

982

C. thyrsis Ereye^x, 1845

0.24

14.1

169

983

Aphantopus hyperantus (Linnaeus, 1758)

0.26

21.2

53;60;233;262

984

Cercyonis oetus (Boisduval, 1869)

0.36

21.8

275;24

985

C. pegala (Fabricius, 1793)

0.45

29.8

2;90;230;275

986

Hyponephele lupinus (Costa, [1836])

0.13

22.7

230;262

987

H. lycaon (Kuhn, 1774)

0.22

22.3

60;262;230

988

H. maroccana (Blachier, 1908)

0.08

19.0

102

989

Maniola jurtina (Linnaeus, 1758)

0.08

25.8

53;60;230;262

990

M. rzwrag Ghiliani, 1852

0.08

22.8

230

Pyronia bathseba (Fabricius, 1793)

0.27

20.4

230;262

992

P. cecilia (Vallantin, 1894)

0.11

19.6

6;230;262

993

P. tithonus (Linnaeus, 1771)

0.13

18.9

53;233;262

994

Proterebia afra (Fabricius, 1787)

1.04

22.8

160

995

Erebia eriphyle (Freyer, 1836)

0.26

17.5

996

E. euryale (Esper, [1805])

0.45

21.3

304

997

E. ligea (Linnaeus, 1758)

0.45

24.3

53;60

998

E. manto (D. 8c SchifL, 1775)

0.36

20.0

142

110

J. Res. Lepid.

999 £■. w/a Staudiiiger, 1886

0.50

24.0

198

1000 E. meolans (Priinner, 1798)

0.58

22.9

5;60;165;169;262

1001 E. palarica Chapman, 1903

0.76

26.7

262

1002 E. medusa (D. & Schiff., 1775)

0.40

21.6

53;60;146;263

1003 E. aethiopella (Hoffmannsegg, 1806)

0.30

18.5

1004 E. cassioides (Hochenwartz, 1793)

0.42

18.4

262

1005 E. (Hubner, [1824])

0.47

25.0

252

1006 E. gorge (Esper, [1805])

0.39

19.1

304

1007 E. hispania Sutler, 1868

0.42

19.5

59;262;266

1008 E. Herrich-Schaffer, [1847]

0.40

21.1

59;263

1009 E. claudina (Borkhausen, 1789)

0.30

17.0

138

1010 E. epiphron (Knoch, 1783)

0.26

17.5

233

1011 E. lefebvrei (Boisdiival, 1828)

0.59

21.2

304

1012 E. Herrich-Schaffer, [1846]

1.05

21.3

263

1013 E. melas (Herbst, 1796)

0.87

21.0

253

1014 E. neoridas (Boisduval, 1828)

0.47

22.8

170;262

1015 E. scipio (Boisduval, 1832)

0.53

22.0

1016 E. zapateri Oberthur, 1875

0.47

24.5

88;262;267

1017 E. triarius (Prunner, 1798)

0.82

24.5

304

1018 E. aethiops (Esper, 1777)

0.72

23.2

60;263

1019 E. niphonica (Janson, 1877)

0.48

22.0

218

1020 Calisto batesiMxchener, 1943

0.17

14.0

291

1021 C. row/w5a Lathy, 1899

0.30

15.0

291

1022 C. grannus Elites, 1939

0.34

16.5

291

1023 C. herophile Huhner, 1823

0.18

17.5

1024 C. hysius (Godart, 1819)

0.18

17.0

291

1025 C.pw/rM/a Lathy, 1899

1.29

23.5

291

1026 Oeneis glacialis {Moll, 1783)

1.21

28.8

1027 0. jutta (Hiibner, 1806)

0.59

28.8

1028 0. polixenes (Fabricitis, 1775)

0.51

23.1

1029 Arethusana arethusa (D. & Schiff., 1775)

0.30

24.0

53;262;263

1030 Kanetisa circe {Y2LhT\cix\s, 1775)

0.33

37.1

53;262

1031 Minois dryas (Scopoli, 1763)

0.64

32.7

60;262

1032 Berberia abdelkader {Vi^iYxet, 1837)

2.68

35.0

1033 B. lambessanus (Staudinger, 1901)

2.78

66.5

1034 Satyrus actaea {Esper , 1780)

0.51

26.1

256;257;262

1035 S. a7na5'vww5 Staudinger, 1861

0.61

26.4

263

1036 S'. Staudinger, 1892

2.55

28.8

263

1037 S. ferula (Fabricius, 1793)

1.03

28.9

53;262;283

1038 Chazara briseis (Linnaeus, 1764)

0.33

31.6

10;53;60;137;262

1039 C. prieuri (Pierret, 1837)

0.40

34.0

7;10;262

1040 Pseudochazara cingovskii Gross, 1973

0.90

26.0

134

1041 P. graeca (Staudinger, 1870)

0.63

26.3

133

1042 P. hippolyte {Esper, 1784)

0.61

25.0

262;266;304

1043 P. lydia (Staudinger, 1878)

0.71

28.6

263

1044 P. mnizechii (Herrich-Schaffer, [1851])

0.71

28.3

263

1045 Hipparchia alcyone (D. & Schiff., 1775)

0.59

30.2

8;60;217

1046 H.fagi (Scopoli, 1763)

0.71

32.2

53;304

1047 H. neomiris {God?Lrt, [1824])

0.39

24.5

269

1048 H. ellena (Oberthur, 1894)

0.66

31.5

1049 H. aristaeus (Bonelli, 1826)

0.19

28.6

269

1050 H. azorina (Strecker, 1899)

0.56

24.1

150

1051 H. pellucida (Stauder, 1924)

0.23

29.8

263

1052 H. semele (Linnaeus, 1758)

0.25

26.8

53;60;217;233

1053 H. leighehiKxidrn2i, 1976

0.42

32.9

268;304

1054 H. sbordoniKudrm., 1984

0.38

29.9

304

1055 El. hansii (Austaut, 1879)

0.34

24.0

35:90=136, 1996 (2000)

111

1056

H. statilinus (Hufnagel, 1766)

0.29

26.9

217;262

1057

H. powelli (Oberthiir, 1910)

0.27

24.0

102

1058

H.fidia (Linnaeus, 1767)

1.07

29.7

10;217;254;262

1059

H. wyssii (Christ, 1889)

0.91

30.5

226

1060

Melanargia halimede (Menetries, 1859)

0.51

31.0

151;262

1061

M. russiae (Esper, 1783)

0.44

29.0

151;262;271

1062

M. larissa (Geyer, [1828])

0.38

25.9

151;263

1063

M. hylata (Menetries, 1832)

0.67

30.0

151

1064

M. gTMwi Standfuss, 1892

0.55

26.0

151

1065

M. titea (Klug, 1832)

0.41

28.5

151;263

1066

M. galathea (Linnaeus, 1758)

0.55

25.9

60;64;151;233

1067

M. lachesis (Hubner, 1790)

0.69

28.5

151;257;262

1068

M. arge (Sulzer, 1776)

0.26

28.0

154;255

1069

M. ines (Hoffmannsegg, 1804)

0.65

25.1

173;262;304

1070

M. occitanica (Esper, 1793)

0.66

26.5

262;270

1071

M. pherusa (Boisduval, 1833)

0.70

27.0

286;304

1072 Dira clytus (Linnaeus, 1764)

0.38

29.0

1073

D. oxylus (Trimen, 1881)

0.52

34.5

1074

D. swanepolei (van Son, 1939)

0.70

35.0

1075

D.jansei (Swiestra, 1911)

0.58

31.3

1076 Dingana dingana (Trimen, 1873)

0.70

29.0

1077

D. bowkeri (Trimen, 1870)

0.35

23.5

1078

Torynesis mintha (Geyer, 1837)

0.90

26.5

1079

Tarsocera cassus (Linnaeus, 1764)

0.47

26.0

1080

Anetia ihina Geyer, [1833]

1.28

46.5

242

1081

A. briarea (Godart, [1819])

0.44

45.0

228

1082

Idea hypermnestra (Westwood, 1848)

2.05

73.7

144

1083

/. Erichson, 1834

1.94

61.0

168;295

1084 Euploea Sylvester {V2LhY\ci\\s, 1793)

0.20

39.5

296

1085

E. (Cramer, [1777])

0.92

51.5

168;295

1086 E. darchia (Macleay, 1827)

0.13

34.0

289

1087

E. rrawm Lucas, 1853

1.36

45.0

144

1088

Amauris crawshayi 'Sutler,

1.16

40.0

168

1089

A. echeria (Stoll, [1790])

1.36

39.5

61;168

1090

A. albimaculata (Butler, 1875)

0.79

34.6

1091

A. ochlea (Boisduval, 1847)

1.36

40.1

1092 Ide(ypsis juvenata {CidLin^Y, [1777])

1.22

41.0

168

1093

Parantica luzonensis (Eelder 8c Eelder, 1863)

0.57

38.0

295

1094

P. aspasia (Fabricius, 1787)

0.99

41.5

168;295

1095

P. vitrina (Felder 8c Felder, 1861)

0.76

35.0

168;295

1096

Tirumala petiverana (Doubleday, [1847])

0.52

46.8

168

1097

T. limniace {Cramer, [1775])

0.63

48.0

168

1098

T. hamata (Macleay, 1827)

0.54

41.0

168

1099

T. ishmoides Moore, 1883

0.42

44.0

168

1100

Danaus chrysippus (Linnaeus, 1758)

0.48

40.0

61;168

1101

D. gilippus {Cramer, [1775])

0.53

33.5

168

1102

D. erippus (Cramer, [1775])

0.72

48.8

31;168

1103

D. plexippus (Linnaeus, 1758)

0.50

45.8

168

1104

D. genutia (Cramer, [1779])

0.51

43.0

168;272

1105

D. melanippus (Cramer, [1777])

0.55

41.5

168

1106

D. philene (Stoll, [1782])

0.39

39.8

168

1107

Athesis clearistaiyouhleday, 1847

0.79

38.0

247

1108

Patricia dercyllidas (Hewitson, 1864)

0.53

35.0

247

1109

Tithorea harmonia (Cramer, [1777])

0.19

39.0

247

1110

T. tomcma Hewitson, 1853

0.80

41.0

247

nil

Aeria eurimedea (Cramer, [1779])

0.51

24.0

124

1112

A. olena^N eymer , 1875

0.18

23.5

247

112

J. Res. Lepid.

1113

Melinaea ethra (Godart, [1819])

0.52

45.5

247

1114

M. ludovica (Stoll, [1780])

0.83

41.8

247

1115 Athyrtis mechanitisleQldee, 1862

0.50

44.5

247

1116

Eutresis hypereia Doubleday & Hewitson, 1847

0.65

44.0

247

1117

Paititia neglecta (Miiller, 1886)

1.07

34.5

247

1118

Placidula eurynassa (Felder, 1860)

0.25

38.5

237

1119

Methona thernisto (Hiibner, [1818])

0.86

47.5

247

1120

Thyridia psidii (Linnaeus, 1758)

0.18

41.5

247

1121

Scada karschina (Herbst, 1792)

0.30

24.5

247

1122

Sais rosalia (Crcimer, [1779])

0.23

29.5

247

1123

Mechanitis lysimnia (Fabricius, 1793)

0.35

37.5

31;247

1124

Callithomia lenea (Cramer, 1782)

0.26

32.0

247

1125

Talamancana lonera (Butler & Druce, 1872)

0.36

35.0

247

1126

Velamysta pupilla (Hewitson, 1874)

0.37

32.0

247

1127

Ithomia ellara (Hewitson, 1874)

0.31

33.0

247

1128

/. r/rymo Hubner, 1816

0.15

24.5

247

1129

Miraleria cymothoe (Hewitson, 1854)

0.24

28.5

247

1130 Napeogenes harbona (Hewitson, 1869)

0.48

29.5

247

1131 Hyalins frater (Salvin, 1869)

0.47

29.5

247

1132

H. oulita (Hewitson, 1858)

0.55

31.5

247

1133

Rliodussa cantobrica (Hewitson, 1875)

0.18

26.5

247

1134 Hypothyris euclea {God^LXt, [1819])

0.23

30.8

247

1135

H. leprieuriY eisi\\?ime\, 1835“

0.23

27.0

247

1136

H. ninonia (Hubner, 1806)

0.27

28.5

247

1137

H. semifulva Salvin, 1869

0.21

29.5

247

1138 Epityches eupompe {Gey er, 1832)

0.16

28.5

247

1139

Oleria aquata (Weymer, 1875)

0.22

23.0

247

1140

O. asiraea (Cramer, [1775])

0.35

25.0

247

1141

0. zelica (Hewitson, 1856)

0.76

26.5

107

1142 Hyposcada cyrene {Latreille , 1811)

0.81

31.0

247

1143

H. virginiana (Hewitson, 1856)

0.77

31.5

247

1144

Ollantaya canilla (Hewitson, 1874)

0.82

32.5

247

1145

(?) susiana (Felder, 1862)

0.78

37.0

247

1146 Hyalenna pascua (Schaus, 1902)

0.22

28.0

247

1147

Dircenna dew (Hiibner, 1823)

0.15

36.5

247

1148

D. relala Butler 8c Druce, 1862

0.13

35.0

1149 Pterony mia carlia Sch^LUS, 1902

0.12

22.5

247

1150

P. pmnuba (Hewitson, 1870)

0.45

25.5

247

1151

P. thabena (Hewitson, 1869)

0.25

24.0

247

1152

P. notilla Butler & Druce, 1872

0.63

27.0

106

1153 Episcada clausina (Hewitson, 1876)

0.15

23.5

247

1154

E. philoclea (Hewitson, 1854)

0.14

23.5

247

1155

Prittwitzia hymenaea (Prittwitz, 1865)

0.11

23.5

247

1156

Ceratiscada canaria Brown 8c D'Almeida, 1970

0.18

23.5

84;247

1157 Dygoris dirce,nna {^ elder , 1867)

0.16

35.5

247

1158

Godyris duilia (Hewitson, 1852)

0.40

41.0

247

1159

G. hewitsonii (Haensch, 1903)

0.27

34.5

247

1160 Hypoleria adasa (Hewitson, 1854)

0.26

23.0

247

1161

H. cassotis (Bates, 1864)

0.13

25.5

118

1162 Hypomenitis dercetis (Doubleday & Hewitson, 1847)

0.28

28.0

247

1163

Greta andromica (Hewitson, 1854)

0.35

27.0

247

1164

G. cyrcilla (Hewitson, 1854)

0.34

31.5

247

1165

G. diaphanus (Drury', 1773)

0.15

26.0

276

1166

G. nero (Hewitson, 1854)

0.19

25.5

1167

Pseudoscada erruca (Hewitson, 1855)

0.21

24.0

247

1168

P. quadrifasciataUdlhoi, 1928

0.25

25.0

247

1169 Mcclungia salonina {HeWiXson, \^bb)

0.22

24.5

247

35:90-136, 1996 (2000)

113

1170

Heterosais edessa (Hewitsoii, 1854)

0.19

30.0

247

1171

(?) derama (Haenscli, 1905)

0.30

25.0

247

1172

Brassolis isthmia Bates, 1864

1.60

49.0

11;177

1173

Caligo eurilochus (Cramer, [1775])

3.88

89.5

1174

C. mnenion (Felder & Felder, 1866)

4.31

86.0

1175

C. illioneus (Cramer, [1775])

3.98

69.5

1176 Dynastor darius {^^hxicxws, 1775)

5.07

53.5

120;225

1177 D. Doubleday, 1849

11.5

52.5

224

1178 Eryphanis aesacus 1850)

5.54

67.3

167

1179

E. polyxena (Meerburgh, 1775)

5.28

59.0

126

1180 E. reevesi (Doiibleday & Westwood, 1849)

4.19

50.5

1181

Opoptera sulcius (Staudiiiger, 1887)

1.39

38.0

1182 Opsiphanes cassina Felder, 1862

4.04

39.0

66;110

1183

0. quiteria (Stoll, [1780])

1.65

54.9

167

1184

0. tamarindi (Felder, 1861)

4.00

47.9

68;110

Taxonomic arrangement. The high level taxa were arranged after De Jong
et al. (1996). Other relevant references are given in the notes 14 to 23 be-
low. A detailed arrangement of the species is given in the Appendix. This
was constructed after varied sources, phylogenetic approaches having be-
ing given priority. The families and subfamilies included in the Table are
as follows (for each taxon, the first and last species numbers are given):

Hesperiidae^'^

1-132

Coeliadinae

1-3

Hesperiinae

4-67

Trapezitinae

68-95

Pyrginae

96-132

Papilionidae^^

133-227

Parnassiinae

133-166

Papilioninae

167-227

Pieridae

228-311

Dismorphiinae

228-231

Pierinae

232-290

Coliadinae

291-311

Lycaenidae^®

312-609

Riodininae^^

312-343

Poritinae

344-349

Miletinae

350-360

Curetinae

361

Lycaeninae

362-609

Nymphalidae^^

610-1184

Libytheinae

610-612

Heliconiinae^^

613-717

Nymphalinae

718-775

Limenitinae

776-815

Charaxinae

816-877

Apaturinae

878-884

Morphinae

885-900

J. Res. Lepid.

Satyrinae-^

Danainae^^

Ithomiinae"

Brassolioae-^

901--1079

1080-1106

1107-1171

1172-1184

Notes 1-23 to the Table and the taxonomic arrangement.

^No reference was found to reliably quote the author of the species name.
-No reliable information was found on the date of description of the spe-
cies or the genus where the species was originally described.

^According to Clark 8c Dickson (1971 ), a species formerly confused with A.
taikosama (Wallengren), of which I have been unable to stablish the cor-
rect identity.

^After some authors, a subspecies of Callophrys (Mitoura) gryneus (Hiibner,
1819).

'’As E. minyas in the original reference (see Fiedler 1991).

^Clark 8c Disckson (1971) refer to Anthene sp, close to A. talboti Stempffer,
but I have been unable to secure the correct identity of the species.
’According to Emmel 8c Emmel (1989) this record might be adscribed to
Euphilotes mojave (Watson 8c Comstock) .

^As ‘forms’ of H. sapho (Drury) in D’Abrera (1984).

^Following Friedlander (1988), probably better as a subspecies of A. clyton
(Boisduval 8c Le Conte).

afra, sin.: P. phegea (e.g., Hesselbarth et al. 1995).
iiproposed for a new genus by Brown et al. (1994), I have not traced fur-
ther references.

^-Proposed for a new genus by Brown et al. (1994), formerly in Pteronymia.
^^References 94, 128, and 182, give estimates of the egg volumes based in
their own estimates or former references.

^'^=Hesperioidea aucL, arrangement following Bridges (1994)

^^Arrangement after Miller (1988), other references in Collins 8c Morris
(1985).

^'’Following De Jong et al. (1996) for the relationships among subfamilies,
and Fiedler (1991) for other details.

^^Based in the provisional consensus provided by De Vries (1997).

^^After Dejong et al. (1996) up to subfamilies, and other details after Harvey
(1991) unless otherwise stated.

^^Arrangement based in a strict consensus of the results of Brown (1981)
and Brower (1997).

-^Taxonomy simplified from Miller (1968) (see Harvey 1991).
-^Relationships between species are a consensus based on Acery 8c Vane-
Wright (1984), Kitching (1985), Vane-Wright et al. (1992), and Sourakov
&: Emmel (1996).

^‘“Relationships among species after Brown et al. (1994).

“^Brassolinae was kept independent from Morphinae.

35:90-136, 1996 (2000)

115

Sources. Numbers 1 to 304 correspond to those quoted in the data table:
l“Scudder 1873; 2-Scudder 1889; 3-Dyar 1897; 4-Gillmer 1904; 5-Chapman
1905; O^Powell 1905a; 7-Powell 1905b; S^Rebel 1910; 9-Bower 1911; 10-
Oberthiir 1914; 11-Dunn 1917; 12-Coolidge 1923a; 13-Coolidge 1923b; 14-
Coolidge 1923c; 15-Coolidge 1923d; 16-Coolidge 1924a; 17-Coolidge 1924b;
18-Coo!idge 1924c; 19-Cleare 1926; 20-Hayward 1926; 21 -Hayward 1926; 22-
Hayward 1926; 23-Van Someren & Van Someren 1926; 24-Comstock 1927;
25-Stubenrauch 1929; 26~Hayward 1931; 27-Comstock & Dammers 1932; 28-
Conistock 8c Dammers 1932; 29-Comstock Sc Dammers 1932; 30-Le Moult
1932; 31~Hoffmann 1933; 32-Lichy 1933; 33-Dos Passos 1936; 34-Dos Passos
1936; 35-Dethier 1938; 36-Djou 1938; 37-Hoffman et ak 1938; 38-Hoffmann
1938; 39-Tsang 1938; 40-Dethier 1939; 41-Dethier 1939; 42-Comstock 1940;
43-Dethier 1940; 44-Dethier 1941; 45-Dethier 1942; 46-Dethier 1942; 47-
Dethier 1942; 48-Dethier 1943; 49-Dethier 1944; 50-Bourquin 1948; 51-
Bourquin 1949; 52-Dickson 1949; 53-Sarlet 1949-1957; 54-Beebe 1952; 55-
Urbahn 1952; 56-Bourquin 1953; 57-Comstock 1953; 58-Zikan 1953; 59-De
Lesse 1954; 60-D5ring 1955; 61-Van Son 1955; 62-Jarvis 1956; 63-Clark 8c
Dickson 1957; 64-Beebe et ak 1960; 65-Comstock 1961; 66-Comstock 8c
Vazquez 1961; 67-Malo & Willis 1961; 68-Harrison 1963; 69-Niculescu 1963;
70-Van Son 1963; 71-Emmel 8c Emmel 1964; 72-Hayward 1964; 73-Ross
1964a; 74-Ross 1964b; 75-Ross 1964c; 76-Clark 8c Dickson 1965; 77-Cottrell
1965; 78-Lawi'ence 8c Downey 1966; 79-Templado 1966; 80-Clark 8c Dickson
1967; 81-Hayward 1967; 82-Emmel 8c Emmel 1968; 83-Heitzman 8c Heitzman
1969; 84-Brown 8c d’Almeida 1970; 85-Clark 8c Dickson 1971; 86-Rydon 1971;
87-Straatman 1971; 88-Bodi 1972; 89-Dujardin 1972; 90-Emmel 8c Mattoon
1972; 91-Mcalpine 1972; 92-Quick 1972; 93-Young 1972a; 94-Young 1972b;
95-Young 1972c; 96-Young 1972d; 97-Atkins 1973; 98-Muyshondt 1973; 99-
Young 1973a; 100-Young 1973b; 101-Young 1973c; 102-Young 1973d; 103-
Emmel 8c Emmel 1974; 104-Muyshondt 1974; 105-Shirozu 8c Hara 1974; 106-
Young 1974a; 107-Young 1974b; 108-Atkins 1975; 109-Straatman 1975; 110-
Young 8c Muyshondt 1975; 111-De la Maza 8c De la Maza 1976; 112-Martin
1976; 113-Muyshondt 1976; 114-Atkins 8c Miller 1977; 115-Henning 1977;
116-Lambkin 8c Lambkin 1977; 117-Schurian 1977; 118-Young 1977a; 119-
Young 1977b; 120-Aiello 8c Silberglied 1978; 121-Atkins 1978; 122-
Pennington 1978; 123-Roos 1978; 124-Young 1978; 125-Dias 1979; 126-Dias
1979; 1 27-Down ey 8c Allyn 1979; 128-Duniap-Pianka 1979; 129-Henning
1979; 130-Silberglied et ak 1979; 131-Van Son 1979; 132-Aiello 1980; 133-
Aussem 1980; 134-Aussem 8c Hesselbarth 1980; 135-Dias 1980; 1 36-Down ey
& Allyn 1980; 137-Roos 1980; 138-Roos & Amscheid 1980; 139-Yata& Eukuda
1980; 140-Brown 1981; 141-Downey 8c Allyn 1981; 142-Roos 8c Arnscheid
1981; 143-Henning 1982; 144-Kirton et ak 1982; 145-Roos et ak 1982; 146-
Arnscheid 8c Roos 1983; 147-Eukuda 1983; 148-Hesselbarth 1983; 149-
Johnson & Valentine 1983; 150-Oehmig 1983; 151-Wagener 1983; 152-Atkins
1984; 153-Downey 8c Allyn 1984; 154-Eitschberger 8c Racheli 1984; 155-
Furtado 1984; 156-Heath et ak 1984; 157-Martm 8c Templado 1984; 158-
Nakasuji 8c Kimura 1984; 159-Parsons 1984; 160-Roos et ak 1984; 161-Sands

116

J. Res. Lepid.

et al. 1984; 162-Wood 1984; 163-Wright 1984; 164-Young 1984; 165-
Arnscheid & Roos 1985; 1 66-Casagrande & Mielke 1985; 167-Cubero 1985;
168-Kitching 1985; 169-Boillat 1986; 170-Boudinot 1986; 171-Callaghan
1986; 172-Eitschberger 8c Strohle 1986; 173-Eitschberger et aL 1986; 174-
Eriedlander 1986; 175-Hiiertas 1986; 176-Roos 1986; 177-Young 1986; 178-
Atkins 1987; 179-Atkins 8c Heinrich 1987; 180-Atkins 8c Miller 1987; 181-
Benz etal. 1987; 182-Nakasuji 1987; 183-Roos 1987; 184-Shapiro 1987; 185-
Wood 1987; 186-Atkins 1988; 187-Eriedlander 1988; 188-Graham 1988; 189-
Munguira 1988; 190-Wood 1988; 191-Benjamini 1989; 192-Boillat 1989; 193-
Boudinot 1989; 194-Callaghan 1989; 195-Henning 1989; 196-Johnson 8c
Valentine 1989; 197-Marini 8c Trentini 1989; 198-Roos 8c Arnscheid 1989;
199-Sanison 1989; 200-Valentine &:Johnson 1989; 201-Back 1990; 202-Braby
1990; 203-Eitschberger 1990; 204-Eitschberger 1990; 205-Eitschberger 1990;
206-Eitschberger 1990; 207-Eitschberger 8c Strohle 1990; 208-Eitschberger
8c Strohle 1990; 209-Emmel 8c Emmel 1990; 210-Enmiel 8c Garraway 1990;
211-Goyle 1990; 212-Koppel 1990; 213-Urich & Emmel 1990; 214-Urich 8c
Emmel 1990; 215-Ziegler &:Jost 1990; 216-Callaghan 1991; 217-Garcia-Barros
8c Martin 1991; 218-Hara 1991; 219-Johnson 8c Doherty 1991; 220-Samson
1991; 221-Sarto 8c Maso 1991; 222-Turner 1991; 223-Urich 8c Emmel 1991;
224-Urich 8c Emmel 1991; 225-Urich 8c Emmel 1991; 226-Wiemers 1991;
227-Brevignon 1992; 228-Brower et al. 1992; 229-Ereitas 8c Oliveira 1992;
230-Thomson 1992; 231-Williams et al. 1992; 232-Braby 1993; 233-Dennis
1993; 234-Eitschberger 1993; 235-Eitschberger 1993; 236-Eiedler 1993a; 237-
Freitas 1993; 238-Garraway et al. 1993; 239-Hauser et al. 1993; 240-Henning
et al. 1993; 241-Leestmans 8c Carbonell 1993; 242-Llorente-Bousquets et al.
1993; 243-Merrett 1993; 244-Atkins 1994; 245-Braby 1994; 246-Braby 8c
Woodger 1994; 247-Brown 8c Freitas 1994; 248-Caldas 1994; 249-Freina 1994;
250-Hsu 8c Lin 1994; 251-Johnson et al. 1994; 252-Jutzeler 1994a; 253Jutzeler
1994b; 254-Jutzeler 1994c; 255-Jutzeler 1994d; 256-Jutzeler 8c Leestmans
1994a; 257-Jutzeler 8c Leestmans 1994b; 258-Otero 1994; 259-Pringle et al.
1994; 260-Smith et al. 1994; 261-Dan tchenko et al. 199; 262-Garcia-Barros
8c Martin 1995; 263-Hesselbarth et al. 1995; 264-Hirukawa 8c Kobayashi 1995;
265-Johnson et al. 199; 266-Jutzeler 1995a; 267-Jutzeler 1995b; 268-Jutzeler,
Grillo 8c De Bros 1995; 269-Jutzeler, Pitzalis 8c De Bros 1995; 270Jutzeler et
al. 1995a; 271-Jutzeler et al. 1995b; 272-Meyer 1995; 273-Penz 1995; 274-
Samson & Wilson 1995; 275-Sourakov 1995; 276-Sourakov 8c Emmel 1995;
277-Yen &Jean 1995a; 278-Yen &Jean 1995b; 279-Zanundo etal. 1995; 280-
Atkins 1996; 281 -Caballero 1996; 282-Dantchenko et al. 1996; 283-Jutzeler
1996; 284-Jutzeler 8c De Bros 1996; 285Jutzeler, Biermann 8c De Bros 1996;
286-Jutzeler et al. 1996; 287-Merrett 1996; 288-Meyer 1996a; 289-Meyer
1996b; 290-Parsons 1996; 291-Sourakov 1996; 292-Williams & Atkins 1996;
293-Atkins 1997; 294-Callaghan 1997; 295-Igarashi 8c Fukuda 1997; 296-
Meyer 1997a; 297-Meyer 1997b; 298-Sourakov & Emmel 1997; 299-Teshirogi
1997; 300-Williams 8c Atkins 1997; 301-Leigheb 8c Cameron-Curry 1998; 302-
M.L. Munguira, unpublished data on Spanish Lycaenidae; 303-F. Urich,
unpublished; 304-E. Garcia-Barros, unpublished.

35:90-136, 1996 (2000)

117

Acknowledgements. I wish to thank several persons provided either unpublished in-
formation (on eggs, or adult butterflies) , useful advise on the specialised literature,
or help in locating and obtaining the data: M.L. Munguira, J. Fernandez-Haeger,
A. Vives, D. Jutzeler, SJ. Johnson, PJ. Merrett, F.C. Urich, P.R. Acker)% R. de Jong,
and T. Racheli.

Appendix. Relationships among the species included in the data set (see
'Sources' for the main references), in parenthetical notation.

((((1,2), 3), (((4, (5, 6, 7), (8, 9), (10, (11, 12), 13, 14), (15, 16, (17, 18)), 19, ((20, 21, 22, 23, 24), 25, (26, 27, 28), (29, 30,
31,32),33),34,(35,36),(37,38,39),(40,41,42),(43,44),((45,(46,47),(48,49),50,(51,52,53,54),55,(56,57)),(58,
59, (60,61 ,62,63,64) ) ) , ( (65,66) ,67) ) , (68, (69,70,71 ,72,73) ,74, (75,76,77,78) ,79,80, ( (81 ,82,83,84,85) ,86, (87,
88),89),((90,91),(92,93),94,95))),((96, 97, 98,99,100), (101, (102, 103,104,105)),(106, 107, 108, 109, 110, (111, 112
,113,114),(115,116, 117,118,119,120, 121), 122), ((123, 124),125),(126,127,128,129,130,(131, 132)))))), ((((133,
134,(135,(136,137), 138, (139,140), 141,142,(143,144),145,(146,147,148,149,150),(151,152,153,154,155,156,
157,158))),(l®ae0,(161,162),m(16ia65,ie6))),(((167,16aie9),((170,171),((17^B),((174,175),(17dl77,mi79,180))))),((181,
(182,((((183,184,185, 186, 187,188),189),(190,191)),((192,193,194,195),(196, 197,198, 1993X))),(201,202)))),(((203,
204),((2(B,206),((2073)8)3)93031))), ((21233214)35, (216,(217,(2ia(2193)))))221,222223,(224,225)226,
227))))),(((22822930)31), (((((232332313523637238239)240)241242, (243244)245), ((246247),(248249,
250313233)254),(25536, ((257,25839), (26031))32,263,(2bl265)26637, (26826030), 31,(272,273),
(((3435),(3627738)), ((27928031), (2ffi283)284))285,(286287288289)290)),(291,(29233294),(295296,
297298299300),(301302203201305206),(307208)209310311))),((((312313),(314315),(((316317)318),((319,
320,321, 322), 323,324,325,326),((327,33329,330),(331,332,333, 334,335)), ((336, 337),(338,339)),(340,341),342,
^)),(((((344345346),((317A18)349)),(((35031),(352333543553563735839))360))361),(((362363361,
365366367, 368,369, 370),371372,373, 374,375,376,(377378), (379, 38038132, 383, 384385, 386,387,388389),(390,
391,392),(393,301),395),((396,397,398,399,4OO,4O1,4O2,4O3,4(>1,4O5,4O6),4O7),(((4O8,4O9,41O,411),(412,(413,414,
415))),((416,417,418, 419,(420, 421),422,423),(424,(425,426,427,428,429),430,(431,432),(433,434),435,(436,437, 438),
(439,440,441,442, 443,444,445)),(446,447,(448,449))),450),((451,452),453,(454,(455,456,457,458,459),460),(461,(462,
(463,4bl))),((465,466),((467, 468,469), 470),471,472),(((473,474),(475,476),477),478, ((479,480,481), (482, 483,484)),
(485,486,487))),((4^,489), (490,491, 492,493,494,495,496, 497),((498,499)300301, (502303), ((5043053063073(B),
509,510, (511312)), (513,514315),((516317),(518319))320,((521,522),(523,524)),525,(526327),((528,529),(530,
531332333),(534335)),(536337338339),(540341),((542343)344345346),(((M7348349350351)352,(553354,
555,556357358)),(559,560361362),(563,561),565366,567),((5683e9370371, 572), (573374375376377378379,
580381382)383,(584385)),(((586387)388,(589390391),(5923933943*))396,(597,(598399,600,601),((602,603,
601),e05,606),607),e08)),e09)))),(((610,611),612),(((((613,(614,615)),((616,(617,(618, (619,(620,621))))), (622,(((623,
624,625),(626,627,628,629,630,631)),((632,633,634),(635,(636,(637,638),(B9,610),(641, (612,643), &14, 645,616, 647,
(648,649, 650, 651),(652,653),654,655),(656,657,658,(659,660),(661,662),(663,664),665,666,(667, (668,669)),
670)))))))), (671, 672,(673, 674,675, 676),(677,678,679), 680, 681, 682, (683, 684, 685, 686, 687, 688, 689)), ((690,691),(692,
693,604),^),696,(^7,698)),((((699,7OO,7O1),(7O2,7O3,7O4)),((7O5,(7O6,7O7)),7O8)),(7O9,((71O,711),((712,713),
(714,(715,716)))))), 717),((718, 719, 720, 721, (722,723,724), (725, 726, 727,728, 729),730,(731,732)),(733, (734,735, 736,
737),(738,739,740,741,742,743),(744,745,746,747,748,749),(750,751),752,753,(754,755,756)),(((757,758),759,760),
(761,(762, 763),(761,7ffi, 766, 767,768),(7e9,770,771)),(772, (773, 774)),775)),((776,(777,778),779),((780,781,(782,783,
784)),(785,(786,787),(788,789)),790),(((791,7^,793),794,(795,796,797,798,799),(800,801)),((802,803),(804,805,
806)),(807,808,809, 810,811, 812, 813), (814,815))),((816,(817,818),((819,820),(821,(822,((((823,824,825),826,827,
828,829,830,831),(832,833,834,835,836,837)),((838,839),(810,((841,842),((813,844),(845,846,847,848,849,850,851,
852,853,854,855,856,857,858, 859,860,861,862,863,864,865)))))))))),866,((867,868),869,870,871),(872,873),((874,

118

J. Res. Lepid.

875), 876), 877), (878, ((879,880), ((881, 882), 883)), 884), (({885, 886, 887, 888, 889), 890), (891, (892, 893), (8&4,895, 896),
897, (898, 899, 900))), (901,((902,903),904),((905,(906,907,908),(909,910, 911, 912, 913, 914,915), 916,917,(918,919, 920),
(921,922),923,924),925,(926,927,928,929),((930,931),(932,933, 9^,935,936, 937), 938, 939),(940, 941), (((942, 943), 914),
((945,946, 947,948), (949, 950), (951, 952),953, 954, 955,(956, 957), (958, 959, 960, 961, 962), (963,964, 966)), (966, 967,968),
(969,(970, 971,972, 973, 974,975), (976,977,978,979,980, 981,9^)),(983,(984,985),(986,987,988),(989, 990), (991,992,
993)),(994,((995, 996997^), 999,(1000,1001), 1002,(1003,1004,1005,10061007, 1008),(1009,1010),(1011, 1012,1013, 1014,
1015,1016), 1017, (1018,1019))), (1020, 1021, 1022,1023, 102il0K),(1026,1027,1028),((1029,1030,1031),((1032,1033), (1034,
1035, 1036, 1037))),((1038, 1039), (1040,1041,1042,1043,1044),(((1045,1046,1047,1018),(1049,1050,1051,((1052, 1053),
1054))), ((1055,1056,1057),(1058,1059)))),((1060, 1061), (((1062,1063,1064,1065),(1066,1067)),1068, (1069,(1070,
lO71)))),((lO72,lO73,lO74,lO75),(lO76,lO77),lO761O79))),(((lO0O,lO81),((lOffi,lO83),((l(B4,(lO85,lO86)),lO87)),((l^^
(1089, (1090,1091))), (1092,(10®, (1094,1095))), ((10961097, 1098, 1099), ((1100, 1101), (1102, 1103),(1164,1106, 1106))))),
((1107,1108), ((1109,1110),((1111,1112),((11161114),(1115,(11161117)),((1118,1119), (1120, (1121,(1122,1123)))),((1124,
1125, 1126(((((1127,1128),1129), 1130), ((1131,1132), (1133(1134,1135, 11361137))), 1138), ((imil40, 1141), (1142,1143),
1144,1145))), (((1146(1147,1148)), (1149,1150,1151, 1152)), (((1153, 1151,1155),1156), (1157, (1158,ll»),(lie0, 1161), llffi,
(1163, 1164,1166,1166), (1167,1168),llffl, 1170, 1171))))))))),(1172,(1173,117ill75), (11761177), (1178,1179, 1180),1181,
(1182,1183,1184)))))))

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Book Reviews

THE WILD SILK MOTHS OF NORTH AMERICA, by Paul M. Teskes,
James P. Tuttle and Michael M. CoUins, 1996. Cornell University Press,
Ithaca, NY, IX + 250 pages, including 30 color plates. ISBN 0-8014-3130-
L Price: $75 US.

The Saturniidae, or wild silk moths, have historically captured the attention of
lepidopterists and others often attracted by the large size and rich colors of many of
these moths, which number more than 1 200-1300 species worldwide. This beautiful
book covers about 70 species in 18 genera that occur within the limits of the
continental United States and Canada. The authors’ many years of experience with
these remarkable insects have been condensed and translated into an easily read-
able tome replete with black and white photographs, maps and drawings. Thirty fine-
quality color plates illustrate in life-size all adult moths treated, with smaller
photographs of the last instar caterpillars of all but two species. That the authors
were able to rear and photograph so many species of moths reveals just part of the
dedication, enthusiasm and labor required to produce this outstanding work.

The authors are well-known in the United States for their contributions to
satumiid research. Paul Tuskes has published numerous papers on the U.S.
Saturniidae; James Tuttle, a police detective-lieutenant, has been an officer of the
Lepidopterists’ Society, and collects, rears and photographs wild silk moths; Michael
Collins is a research associate with the Carnegie Museum of Natural Histor)', and is
especially interested in spedation and natural hybridization.

The text is divided into two main sections, both in small print, allowing ample
information to be packed in: Part One, entitled Behavior and Ecology, discusses
such topics as metamorphosis and development, parasitism, diseases, species con-
cepts and taxonomy, collecting, rearing, and silk moth impact on human culture.
Part Two, Species Accounts, contains the color plates, and presents a description of
each subfamily (three in U.S, and Canada) , genus and species. Each species receives
about one or more pages of coverage, including general comments, adult diagnosis,
variation and biology, immature stages and rearing notes. I was gratified to see that
the striking photographs of caterpillars were presented in the natural “hanging
down” position beneath the limb instead of the reverse, as is often the case. Two
appendices list host-parasitoid records and satumiid hybrids. An extensive bibliog-
raphy of cited literature is especially valuable for the student.

However, there is a tendency to overlook or disregard recent taxonomic opinions
and conclusions by other U.S. and international saturniologists. An obvious ex-
ample is the arbitrary decision to reinstate the genus Sphingicampa Walsh 1864,
removing all species except molinafrom the genus Hiibner [1819], on the

basis that molina, which is the type species for Syssphinx, differs morphologically and
in the genitalia from the others. The authors “feel that the North American species
are phylogenetically closer to each other than to Syssphinx molinad' This is followed
by the statement that “The genus Sphingicampa is obviously related to Anisota and
Dryocampa 3.nd to the tropical genus Adelocephala” Adelocephala Dnponchdi 1841 is
not a valid generic name, because it is actually ajunior objective synonym of Anisota
Hiibner [1820], which occurs mainly in North America, Such a provincial approach

138

J. Res. Lepid.

dismisses or ignores the landmark 1988 updated revision of Claude Lemaire (The
Saturniidae of America: Ceratocampinae) who worked with the much greater
number of species found throughout the new world, and the 1982 work of Fletcher
and Nye of the British Museum (The Generic Names of Moths of the World, Vok 4)
and others. Thus, this otherwise excellent book is a bit weak in its taxonomic
treatment, and will perpetuate some confusion among its readers.

The geographical area covered reflects the focus of many U.S. saturniolo-gists, an
area limited to North America north of the Mexican border. This political boundary’
separates a faunistically extremely rich territory from the rest of North America, and
hampers or discourages study of its insects by U.S. investigators. But because more
than half of North American silk moth species reside there, I would like to see a book
integrating the saturniid fauna of the entire North American continent. Also, a more
complete introductory overview of worldwide Saturniidae and recognition of inter-
national saturniid researchers would have been welcome. Nevertheless, for areas
north of the border this book represents an impressive reference work that belongs
in the library of every serious lepidopterist.

Kirby L. Wolfe, 3090 Cordrey Drive, Escondido, CA 92029-5112

GARDEN BUTTERFLIES OF NORTH AMERICA: A GALLERY OF
GARDEN BUTTERFLIES AND HOW TO ATTRACT THEM. Rick
Mikula. 1997. 143 pp. Willow Creek Press, Minocqua, WI. $29.50.

Are butterflies “disappearing?” Butterfly gardening books certainly are not. This
one has what might be called a charismatic, or at least media-friendly, author. Rick
Mikula, son of a coal miner and a sales clerk, dropped out of college, did a stint in
the Navy, work in his native Hazelton, Pennsylvainia as a machinist, and somehow
got “into” butterflies. He started the Hole-in-Hand Butterfly Farm in Hazelton in
1980, selling monarchs and other species for weddings, garden parties, and the like.
Nowadays he raisees 50 species and reportedly sells 25,000 a year shipping FedEx
in summer (because its trucks are cooler) and UPS in winter (because its brown
tnxcks are warmer) . He’s been profiled in Peoplemid the Wall Street Joumal?Lnd on the
Discovery Channel. He designed the Butterfly Emporium at Bollywood, Dolly
Parton’s resort-theme park in Pigeon Eorge, Tennessee. No academic stuffed shirt
he!

So how is the book? OK. The sections on butterfly biology’ and butterfly gardening
are pretty standard. The lists of recommended garden plants are uneven. Those on
pages 35, 43 and 46 have no scientific names for the plants, while that on page 44
does. The numerous color photos appear mostly from life, though several of them
give hints of being posed, perhaps with chilled specimens. Only one is a blatant fake.
It’s on page 1 16 and purports to be a Colias philodice. It’s a winter form of C. eurytheme,
and is rather obviously dead. The male Pieris rapae on page 112 is suspect, too.

Mikula has inexplicable Pierid problems. On page 118 are two photographs of
what are supposed to be Phoebis sennae but are in fact two rather different-looking
male Colias eurytheme. On page 124 there is a real Phoebis sennaes identified as C.
eurytheme. There are other slip-ups, too. On page 63 Mikula says “All swallowtails
perform a ritual called hilltopping,” which might be a surprise to quite a few of them.
On page 33 is a lovely photograph identified as “Eastern Tiger Swallowtails puddling
in Guadalupe Mountains National Park” — they’re P. multicaudatus. On page 28 two

35:137-140, 1996(2000)

139

photos share a caption: “Supply host plants for butterflies to lay eggs on as well as
nectar plants for feeding.” One picture is a male Hy lephila phyleus The other
is a very un-skippery clutch of eggs laid on a tendril of something. And so on.

In short: this is a fairly pretty book, neither the best nor worst of the lot of butterfly-
gardening books in print. There is no compelling reason to buy it unless one collects
butterfly gardening books or stuff peripherally related to Dolly Parton.

Arthur M. Shapiro, Center for Population Biology, U.C. Davis, Davis, CA 93616

BUTTERFLIES ON BRITISH AND IRISH OFFSHORE ISLANDS:
ECOLOGY AND BIOGEOGRAPHY* Roger Dennis and Tim Shreeve.
Gem Publishing Company, Wallingford. 131 pp, ISBN 0-906802-06-7.
£16.00.

Although a slim volume of 131 pages, this book is a tour de force on the ecolog)'
and biogeography of butterflies on British and Irish offshore islands. No more, no
less. It is actually two books in one, the first a checklist of butterflies species found
on British and Irish islands, the second a rigorous analysis of those data. The
checklist alone represents an enormous effort on the part of Dennis and Shreeve,
with records gleaned from every conceivable source, as evidenced by the extensive
list of personal communications and the 571 references in the bibliography.

Dennis and Shreeve present their analysis of the data in a series of short, dense
chapters. They kindly provide a short explication of the their statistical methods,
which include many multivariate ordination and clustering techniques, for those
who may be rusty in that realm. The review is warranted, because the authors put the
data through serious manipulation in their attempts to explain the variation in
species incidence on islands. After starting by placing species number on islands in
the obvious context of island biogeography, the authors proceed to explore all of the
issues that confound the basic relationship described by MacArthur and Wilson.
Their first analytical chapter explores the determinants of species richness. The
following two chapters explore the affinities among island butterfly faunae (they are
nested) and among incidence ofbutterfly species on islands (confirming the nested-
species subsets across islands). Following a chapter predicting butterfly species
number on islands, they present an interesting discussion of migration. Drawing
from the extensive literature assembled for the book, they document instances of
butterflies in hostile habitats that belie the conventional characterizations of
butterfly populations as “closed” or “open.” In the light ofrecords of butterflies from
supposedly closed population structures observed over open ocean, they call for
increased attention to the spatial and temporal variation in mobility and its
implications for metapopulation structure. The next chapter reapplies an ecologi-
cal explanation for butterfly incidence on islands earlier developed by Dennis. The
final two substantive chapters consider intraspecific variation on islands and histori-
cal (Holocene) influences on patterns ofbutterfly abundance on islands.

The writing is straightforward, almost terse, and packed on the page. The authors
use an inordinate number of occasionally non-in tuitive abbreviations for variables,
forcing the reader to repeatedly check back to previous pages to decipher tables.
Even more annoying is the use of numbers to designate points on graphs that in
some instances refer to islands (listed elsewhere) and other times to butterfly species
(listed in yet another location). In this age of computerized publishing and graphic

140

J. Res. Lepid.

design the layout could have been much kinder to the reader by at least using words
instead of abbreviations in the tables and highlighting species or islands discussed
in figtires.

The minor annoyance caused by the typesetting does not diminish from the
thorough treatment of the subject. The book makes an excellent case study of the
issues in island biogeography, but the reader must relate it to the wider literature
without the help of the authors. And although the cover claims that the author’s
findings have ramifications for butterfly conservation, the authors do not discuss
them. The work does illustrate the importance of amateur observation of butterflies
to scientific inquiry and even provides an appendix describing how to make effective
observations of butterflies and moths on islands.

Travis Longcore, UCLA Department of Geography, Box 951524, Los Angeles, CA 90095-
1524

Journal of Research on the Lepidoptera

35:141-142, 1996 (2000)

Notes

A Million White Butterflies (Pieridae) At Ouray National Wildlife Refuge^
Utah

Key Words: Pontia protodice, Pieris rapae, superabundance, Glycyrrhiza, censusing

On 8.VIIL1996 the authors visited Ouray National Wildlife Refuge, Uintah County
in northeastern Utah. The 5000 hectare Refuge is located about 25 km west of
Venial. Part of the refuge is accessible by an 8 km loop road through marshes and
fresh water impoundments along the Green River. Near the entrance to the loop
road we noted large numbers of white butterflies (Pieridae), both Cabbage White,
Pieris rapae Linnaeus, and Checkered 1/VOiite Pontia protodice Boisduval & LeConte,
nectaring on Rabbitbrash {Chrysothamnus nauseosus, PalL, Asteraceae). As we
continued on to the loop road the butterflies appeared extremely abundant- We
estimated the numbers of white butteiTlies within 20 m of the road using order of
magnitude categories. We classified stretches of road as having 5-50, 50-500,
500-5000, or 5000-50,000 butterflies per 100 m (column 1 in table 1) . For example,
we estimated between 50 and 500 individuals per 100 linear meters, along the first
1 km of road. Then after a short spell of low density, the number of butterflies
increased and we began finding them nectaring on Tamarisk or Salt Cedar ( Tamarix
gallica French, Tamariaceae). A sample of 15 plants with between 10 and 40
butterflies nectaring, showed an average of 9% Checkered Wliites among the more
abundant Cabbage Butterflies.

As we proceeded through the marshes, we encountered many Tamarisk bushes on
either side of the road, and as we turned to parallel the river, the roadside ditches
were clogged with stands of American Wild Licorice {Glycyrrhiza lepidota Pursh,
Fabaceae) , in a band ranging from 10-20 m wide, occupying one or both sides of the
road. At this point the butterflies were so numerous that we could only estimate
them by the thousands. Some licorice plants had over 100 individuals nectaring at
flower clusters partially hidden under foliage. In the densest area we used 4 spot
counts (1 m radius circles) which yielded a mean of 48.3 individuals per 3.14 m^, to
validate our estimate. These yielded an estimate of 30,800 for a 100 m long segment
(2000 m^), comfortably close to the midpoint of our range (27,500). We noted
whether the butterflies occurred on one or both sides of the road, and clocked
distances with the odometer to estimate the length of each segment. We ended a
segment at a point where the density seemed to change markedly. Table 1 shows the
calculation for the 8 km route. Taking the midpoint of the estimated range for each
segment (2nd column in table 1) as representative, and multiplying by the length
of each road segment and the number of sides occupied by butterflies, yielded an
estimate of over 1 ,000,000 butterflies within 20 m of the road along the 8 km route.
(There were very few further away because of lack of nectar sources) . We doubt that
the number was less than 750,000 nor more than 1.5 million.

Although one occasionally reads about “millions” of butterflies, these usually refer
to migratory movements occurring over periods of hours or days. This is by far the
largest localized aggregation we have ever seen. Ironically, we were unable to
identify the larval host plants that the Pierids might have been using. We observed

142

/. Res. Lepid.

Table 1 . Estimation of abundance of white butterflies at Ouray National Wildlife
Refuge, Utah, 8 August 1996.

Range of

Midpoint

Length

estimate

value for

of road

Number of

Estimate

per 100 ni

100 m

segment

sides

for

of roadside

segment’

(meters) -

occupied^

segment

50 - 500

275

960

2,640

5- 50

27.5

320

50 - 500

275

480

1.5

1,980

500 - 5000

2750

1600

1.25

55,000

5000- 50000

27500

2080

1.7

972,400

500 - 5000

2750

800

44,000

50 - 500

275

1280

7,040

5- 50

27.5

960

1.5

396

Total Estimate

1,083,544

This value multiplied by length

of segment (column 3) and number of sides

occupied (column 4), divided by 100, yielded die segment estimate in the 6th
column.

- Converted from mileage on odometer

^ Either one or both sides of the road were occupied by butterflies, depending
mainly on distribution of Glycyrrihiza. For example, a value of 1.7 indicates that
both sides were occupied for most of the segment.

that P. protodice occurred most commonly where P. rapaew^LS also common. Such
superabundant aggregations are probably not rare, but linear distribution along the
roadside ditches afforded an unusually favorable opportunity for estimating num-
bers.

We thank our companions Guy Tudor and Michelle LeMarchant for their
patience, and botanist Tamara Naumann for identifying Glycyrrhiza.

Michael Gochfeld and Joanna Burger, Environmental and Occupational Health Sciences
Institute, 170 Frelinghuysen Road, Piscataway, NJ 08854 gochfeld@eohsirutgers.edu

INSTRUCTIONS TO AUTHORS

Manuscript formats Two copies must be submitted, double-spaced, typed, with wide
margins. Number all pages consecutively. Italicize rather than underline scientific names
and emphasized words. Footnotes are discouraged. Do not hyphenate words at the right
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standard time. Abbreviations must follow common usage. Dates should be cited as: day-
Arabic numeral; month-Roman numeral; year- Arabic numeral (e.g. 6. IV. 1992). Numerals
must be used for ten and greater e.g. nine butterflies, 12 moths.

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The Journal of Research

ON THE LEPIDOPTERA

Volume 35

1996(2000)

IN THIS ISSUE

Date of Publication: March 15, 2000

Differences in lifetime reproductive output and mating frequency of two

female morphs of the sulfur butterfly, Colias erate (Lepidoptera: Pieridae) 1

Yasuyuki Nakanishi, Mamoru Watanabe, and Takahiko Ito

Oviposition, host plant choice and survival of a grass feeding butterfly, the

Woodland Brown {Lopinga achine) (Nymphalidae: Satyrinae) 9

Karl-Olof Bergman

The effect of environmental conditions on mating activity of the Buckeye

butterfly, Precis coenia 22

Alice K. McDonald and H. Frederik Nijhout

Nymphalid butterfly communities in an amazonian forest fragment 29

Frederico Araujo Ramos

A Survey of the Butterfly Fauna of Jatun Sacha, Ecuador (Lepidoptera:

Hesperioidea and Papilionoidea) 42

Debra L. Murray

Flexural stiffness patterns of butterfly wings (Papilionoidea) 61

Scott J. Steppan

The number of copulations of territorial males of the butterfly

Callophrys xami (Lycaenidae) 78

Carlos Cordero, Rogelio Macias, and Gabriela Jimenez

Egg size in butterflies (Lepidoptera: Papilionoidea and Hesperiidae):

a summary of data 90

Enrique Garcia-Barros

Book Reviews 137

Note

141

Cover: Butterfly. Abstract sketch by Pavel Tocik, 1997.

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journal of Research on the Lepidopieia

36:1-15, 1997 (2000)

A Study of the Riodinid Butte
Nepal (Riodinidae)

Ave. Siiba 130-25 Casa 6, Bogota, Colombia,

Curtis John Callaghan

>odona in

Abstract. I present and discuss the adult habits of five riodinid species from
the Kathmandu valley, Nepal; Dodona egeon (Westwood, 1851), Dodona
eugenes (Bates, 1867), Dodona ouida (Hewitson, 1865), Dodona dipoea
(Hewitson, 1865) ?iv\d Dodona adonira (Hewitson, 1865) , including ovipo-
sition, feeding, perching and distribution, and describe the immature
biolog)' and lar\'al habits of D. egeon, D. eugenes and D. dipoea for the first
time. The food plant for D. egeon is Myrsine capitellataV^^3.\\. (1824) and for
D. eugenes and D. dipoea is Myrsine semiserrata Wall. (1824), both family
Myrsinaceae. Principal adult food resources were bacteria and algae found
on wet earth and leaves and to a lesser extent pollen and faeces. All species
had a proboscis modified with numerous small lateral projections to assist
in absorbing nutrients. I conclude that sympatric Dodona species use
perching in different micro- habitats as a mechanism to maintain species
isolation.

Key Words: Nepal, Oriental Region, Riodinidae, immature biolog)', adult
habits.

Introduction

Although the butterfly fauna of the Oriental region has arguably been
studied more than any other tropical region, its riodinid fauna has been sadly
neglected. Aside from short mentions in species lists and general faunal
books starting with “Seitz”, the only works dealing with riodinid biology are
Sevastopoulo (1946) andjohnston&jolmston (1980). As a start in fdling this
void, this paper presents field and laboratory observations on the biology and
habits of five Dodona species from Kathmandu valley, Nepal; Dodona egeon
(Westwood, 1851), Dodona eugenes (Bates, 1867), Dodona ouida (Hewitson,
1S65) , Dodona dipoea (Hewitson, 1865) Dodona adonira (Hewitson, 1865).
I describe the immature biologies of D. egeon, D. eugenes Rud D. dipoeaior the
first time and include field observations and discussions on ovipositing,
feeding, perching and distribution.

Materials and Methods

I made field observations in the hills surrounding Kathmandu Valley in southern
Bahktapur District and Lalitpur District during December 1995 and February to
September, 1996. The study area (fig. 27) extended from Suryebinayak ridge south
to Godawari and Pulchok peak, all forming part of the Pulchok massive, a range of
hills on the southeast side of Kathmandu valley, with altitudes from 1300 m to 2762
m. Between 1400 m and 2100 m is Schhna-RJiododendron-O'ak forest, the dominant tree
species being Schitna wallachii, Quercus glauca mid Rhododendron arboreurn. Above 2000

Paper submitted 17 September 1996; revised manuscript accepted 16July 1997.

/. Res. Lepid.

in S. wallachii is replaced by Quercus lamellosa. Parts of this formation not used for
agricultural terraces have been altered into scrubland by the gathering of fuel wood
and animal fodder (fig, 23); however, on the steeper slopes and partially protected
areas like the Suryebinayak ridge and the headwaters of Nag creek (fig. 24 ), less
disturbed forest remnants are found. Between 2100 m and 2400 ni is a transition zone
between Rhododendron arboreiimmid the Quercus seynicarpifoliaiorest'which continues
to Pulchok peak (2762m) (Kliadka et a/,1984).

I discovered food plants and larvae through observing oviposition, and on one
occasion hired local people to search for eggs and larvae. Five immature D. egeonw^r^
studied, 42 D. eugenes and 3 D. dipoea. All larvae and eggs were raised in petri dishes,
each larva receiving a unique reference code for recording its development. I
examined immature stages with a binocular lupa. Some larvae and parasites were
preserved in formaldehyde, and adults in papers. Voucher specimens are in the
collection of the author.

Results

Dodona egeon (Westwood, 1851) (fig 2,3 )

D. egeon ranges from Central Nepal east to western China and Burma. In
Nepal, it has been recorded as far west as Baghmg, Baghmg District (fig. 28
). It flies between 1000 m and 2235 m, with an average locality elevation of
1400m.

Dodona egeon, immature stages

EGG: Diameter 0.7 mm, height 0.6 mm. Color reddish-brown when first
laid, changing to white before hatching. Surface smooth. Micropyle is a tiny
depression on top of egg. Duration: 7 days. n=5.

FIRST INSTAR LARVA: Length 2.5 mm. Thorax and abdomen slightly
dorsally compressed with segments T2 through A8 protruding laterally at
base; larv^a initially transparent, turning light green upon feeding. Head dark
yellow, face setose with black spot in center; headcapsule width 0.4 mm. T1
light yellow, transverse pro thoracic shield high, bifurcated dorsally with 5
long setae projecting cephalad on each side and one long lateral setae.
Segments T2 through A8 light green, each with four white dorsal tubercles
and a ‘V’ shaped forked dorsal setae from each, and one long unforked setae
and several small ones on each lateral protrusion; anal shield triangular with
6 setae around edge and 6 dorsad. Spiracles light green, lateral/ posterior on
T1 and superior to lateral protrusions on A1 to A8. Duration: 5 days. n=4.

SECOND INSTAR LARVA: (fig. 4) Length 5.0 mm. Thorax and abdomen
dorsally compressed, segments T2 through A8 with larger lateral protru-

Fig. 1 D, egeon foodplant, Myrsine capitellata
Fig. 2 D. egeon female on leaf of foodplant.

Fig. 3 D. egeon perching male
Fig. 4 D. egeon second instar larva
Fig. 5 D. egeon third instar larva
Fig. 6 D. egeon fourth instar larva
Fig. 7 D. egeon fifth instar larva
Fig. 8 D. egeon pupa

36:1-15, 1997 (2000)

/. Res. Lepid.

Fig. 9 D. eugenes male feeding

Fig. 10 D. ei/genes foodplant, Myrsine semiserrata

Fig. 1 1 D. eugenes second instar larva

Fig. 12 D. eagenes third instar larva

Fig. 13 D. eagenes fourth instar larva

Fig. 14 D. eugenes fifth instar larva

Fig. 1 5 D. eugenes pupa

36:1-15, 1997 (2000)

sioiis. Head yellow-brown, face setose with dark spot in center; headcapsnle
width 0.8 mm. T1 light yellow-brown, bifurcated ridge on prothoracic shield
lower than first iiistar with 5 long setae projecting cephalad from each side
and a cluster of long lateral setae. T2 through A8 greenish white dorsad with
row of green dorsal spots flanked by a row of smaller elongated green spots
and 4 white tubercles, each with two unitaiy setae; dorsal spots on T2, T3 and
A3-A7 larger. Each segment protrudes laterally at base with a cluster of long
setae at tip and numerous shorter setae dorsad; anal shield light brown with
6 setae around edge and 4 dorsad. Spiracles as on first instar. Duration: 6
days. n==3.

THIRD INSTAR LARVA: (fig. 5 ) Length 10.7 mm. Thorax and abdomen
light green with darker green markings, dorsally compressed with segments
T2 through A8 protruding laterally. Head light brown, setose; headcapsnle
width 1.5 mm. T1 light green with long setae on rim projecting cephalad,
short line dorsad. T2 through A8 lighter green-yellow dorsad, darker green
laterally, dark green spots on each segment fonning triangular pattern with
base cephalad and apex as a large dorsal spot on posterior segment margin.
Lateral protrusions green-white, triangular with numerous long setae; anal
shield rounded with four white spots dorsad with one setae each and more
setae around edge. Spiracles as on first instar. Duration: 6 days. n=3.

FOURTH INSTAR LARVA: (fig. 6) Length 16.0 mm. Thorax and abdo-
men less dorsally compressed. Head light green, face setose with white spots
forming circular pattern dorsad (fig. 7) ; headcapsnle width 2.0 mm. T1 light
green with short forward projecting setae and a short green line dorsad. T2
through T8 light green, dorsal pattern as in third instar but fainter with
numerous short setae; triangular lateral protrusions on segments T2 through
A8 less prominant, with long lateral setae; anal shield thick with four white
spots dorsad and long setae around edge. Spiracles as on first instar.
Duration: 3 days. n=3.

FIFTH INSTAR LARVA: (fig. 7) Length 22.5 mm to 28 mm. Head as in
fourth instar, headcapsnle width 2.7 mm. T1 light green mottled with white
spots, short line dorsad. T2 through A8 light mottled green with light brown
spiracles; otherwise as in fourth instar; anal shield mottled light green with
long setae around edge and dorsad. Two days before pupating, larva turns
uniform light green. Duration: 7 days. n=3.

PUPA: (fig. 8) Length 1 9.2 mm; width at widest point 1 2.0 mm. Color light
green with light blue and yellow markings. Pupa attached by a cremaster and
a girdle which crosses dorsum at AL T1 crest indented with cerated, yellow
edge; light blue dorsal line from T1 to A9 flanked on each side by a broken
blue line on T1-T3 and blue spots A1 to AlO; yellow spiracles on T1 and A2
through A8, wing cases white. Duration: 10 days. n= 2.

Dodona eugenes (Bates, 1867) (fig. 9)

Z). eugenes is found from Nepal east to Burma and central China. In Nepal
it ranges across the country between 1600 m to 2700 m, with an average
locality elevation of 1870 m.

/. Res. Lepid.

Dodona eugenes, immature stages

EGG: Diameter 0.7 mm, height 0.6 mm. Color cream when first laid,
changing to brown as the laiwa matures. Sides smooth. Duration: 6 days. n=4.

FIRST INSTAR IJUIVA: Length 3.0 mm. Thorax and abdomen tubular
with segments T2 through A8 protruding slightly laterally; larva initially
transparent, turning light green upon feeding. Head black with setae on
face, headcapsul width 0.4 mm. T1 light green with prothoracic shield as
bifurcated black transverse ridge, with 6 long setae from each side and a
lateral/posterior white spiracle and one long lateral setae. T2, T3 light
green, shorter setae dorsad and one long lateral setae from fleshy protrusion
at segment base. Segments A1 through A8 each with 2 pairs of forked “Y”
shaped dorsal setae, lateral protrusions with one long and 3 short black setae
on each side and numerous shorter setae dorsad; anal shield with 8 long
black setae around edge and 4 dorsad. Spiracles white, lateral on A1 to A8.
Duration: 7 days. n=7.

SECOND INSTAR LARVA: (fig. 11 ) Length 4.2 mm. Thorax and abdo^
men dorsally compressed with segments T2 through A8 protruding laterally
at base. Head light brown with setae and sometimes two spots or a faint bar
on face, headcapsule width 0.7 mm. T1 light green with prothoracic shield
as high, transverse bifurcated dorsal ridge with 6 long setae and a brown spot
caudad on each side; several long lateral setae at base cephalad of spiracle.
T1 covers neck initially, but before molting, neck is exposed. T2 through A8
gray-green dorsad, light brown/ green laterally, a row of faint gray-brown
dorsal spots, each flanked by a short line and numerous shorter setae on each
segment; lateral protrusions pronounced with long laterally projecting
setae; anal shield fleshy with black setae around edge and dorsad. Spiracles
as on first instar. Duration: 5 days. n=5.

THIRD INSTAR LARVA: (fig. 12) Length 7.0 mm. Head light green,
rounded, setose, some individuals with a brown bar across face; headcapsule
width 1.2 mm. T1 light green with dark green line dorsad flanked by a light
brown spot with 6 long setae projecting cephalad and several long lateral
setae; neck exposed on molting. Segments T2 through A8 gray- green dorsad
with numerous short setae and a dorsal spot flanked by two smaller ones, and
a short lateral line cephalad at segment division, largest spots on T2-3, A3-A6;
laterally light green, lateral basal projections rounded, with long setae; anal
shield rounded with setae around edge and dorsad. Spiracles white, lateral
on T1 and A1 to A8. Duration: 6 days. n=17.

FOURTH INSTAR LARVA: (fig. 13) Length 12.0 mm. Thorax and abdo-
men dorsally compressed with segments T2 through A8 protruding laterally
along base. Head light green, with circular pattern of white dots on face and
numerous setae, headcapsule width 1.7 mm. T1 light green, neck covered
initially by prothoracic shield, then exposed upon molting (fig. 11, 13);
dorsad long setae projecting cephalad, a short dark green line dorsad, two
brown spots fainter, or lacking. T2 through A8 light olive green dorsally,
darker green laterally, with same pattern of dark green spots as third instar;
anal shield with setae around edge and two white tubercles dorsad with one
setae each. Spiracles light brown. Duration: 5 days. n=42.

36:1-15, 1997 (2000)

FIFTH INSTAR LARVA: (fig. 14) Length 19 mm to 24/28.4 mm. Olive
green dorsad, laterally mottled wliite/green. Head as in fourth instar,
headcapsule width 2.7 mm. Prothoracic shield covers neck with short setae
on cephalad rim. Segments T2-A8 dorsal pattern same as fourth instar, but
becoming fainter as larva matures; dorsal spots on T2-3, A3-A6 black,
connected by a line of tiny black setae, dorsad with tiny white setae with black
heads. Anal shield mottled green with setae around edge and dorsad.
Spiracles black. Prepupa uniform light green, black spiracles prominent.
Duration: 12 days. n==26.

PUPA: (fig. 15) Length 13-20 mm, width at widest point 5- 6.4 mm. Color
light green with light blue and yellow markings . Pupa attached by a
cremaster and a girdle which crosses dorsum at Al. T1 with indented crest
with ragged edge tinged with yellow. Dorsal blue line from T1 to T9, flanked
on T1 by a shorter blue line, and from T2 to A9 with a broken light blue line;
spiracles outlined in yellow at T1/T2 and A2-A8; wing cases darker green
outlined dorsad by faint blue markings; AlO yellow, pointed. Duration: 10
days. n= 20.

Dodona dipoea (Hewitson, 1865) (fig. 16)

D. dipoea ranges from central Nepal to Assam and north Burma. In Nepal,
it has been recorded as far west as Pokhara valley, and between 1500 m to
2870 m with an average locality elevation of 2200 m. Farther west it is replaced
by Dodona durga (Kollar, 1844).

Dodona dipoea, immature stages

EGG: Unknown.

FIRST INSTAR LARVA: Unknown.

SECOND INSTAR LARVA: (fig. 18) Length 4.3 mm. Thorax and abdomen
tubular with segments T2 through A8 protruding slightly basad. Head light
green/ brown with face setose and two dark brown spots above sutures,
headcapsule width 0.7 mm. T1 dark gray- green dorsad, laterally lighter
green, prothoracic shield high, bifurcated, with 6 long brown setae extend-
ing over head on each side, and a black lateral/posterior spiracle. T2
through A8 brown-green dorsad, flanked by two rows of white spots, inner
row elongated on T2-A1, narrower A2-A8; laterally light brown, with long,
white setae from basal protrusions and black spiracles on A1-A8. Anal shield
flat, triangular with setae around edge and dorsad. Duration: At least 5 days.
n=L

THIRD INSTAR LARVA: (fig. 19) Length 5.5 mm. Head light yellow-
brown, pubescent, initially with dark bar across face; headcapsule width 1.2
mm. T1 light brown dorsad with transverse bifurcated ridge on prothoracic
shield separated by short, red-brown dorsal line and with 6 long setae on each
side, laterally with long setae and a black spiracle; before molting, neck is
exposed as with D. eugenes.T2 through A8 dark green dorsad with two rows of
elongated white marks flanked on each side by a white, irregular line and
covered with short setae; laterally lighter green with some white mottling.

36:1-15, 1997 (2000)

lateral projections at base small with long setae. Anal shield larger, triangular
with black setae. Spiracles as on second instar. Duration: 8 days. n=2.

FOURTH INSTAR LARVA: (fig. 20) Length 13 mm. Lar\a laterally com-
pressed, from A3 tapering to point caudad. Head as in third instar, headcapsule
width 1.7 mm. Prothoracic shield light green, low, with 9 long setae on each
side, lateral spiracle black; T2-T3 olive green with reddish brown line dorsad,
darker green laterally. T2-A8 with short, bristle- like setae dorsad, lateral
projections reduced with long, white setae; A1-A8 darker mottled green,
faint trace of reddish brown dorsal line, flanked on either side by a faint,
irregular lighter green line. Anal shield elongated, pointed with black setae.
Spiracles black. Duration: 7 days, n=2.

FIFTH INSTAR LARVA: (fig. 21 ) Length 14.5 mm to 20 mm. Head round,
yellow/green, face setose, headcapsule width 2.4 mm. T1 uniform light
green, bristle-like short brown setae on cephalad rim, black posterior/lateral
spiracle, numerous brown, bristle-like setae dorsad. T2- A8 uniform, light
mottled green, with red/ brown line from T2 to A8, widest on T2-T3; body
covered with short, bristle- like setae, lateral setae at base shorter. Anal shield
more elongated, pointed. Prepupa lighter green. Duration: 8 days. n=2.

PUPA: (fig. 22) Length 13 mm, width at widest point 6.0 mm. Color light
green with light blue and yellow markings . Pupa attached by a cremaster and
a girdle which crosses dorsum at A1 . T1 with bifurcated dorsal crest reduced.
Dorsal blue line from A1 to T9, spiracles outlined in dark green on T1-T2 and
A2-A8; wing cases darker green. AlO pointed, yellow. Duration: 10 days. n=
2.

Dodona ouida Hewitson, 1865 (fig. 25)

D. ouida ranges from Nepal east to central China . In Nepal it is found across
the countiy between 1450 m and 2900 m, with an average locality elevation
of 2000 m.

Dodona adonira (Hewitson, 1865) (fig. 26)

D, adonira is found from Nepal east to Assam, Sikkim, Burma and northern
Thailand. In Nepal, it is recorded as far west as the Pokhara valley, and
between 1451 m and 2353 m with an average habitat elevation of 1854 m.

Discussion

Food plant and species distribution

The food plant of D. egeon, Myrsine capitellataWRll. (1824) (Myrsinaceae)

(fig.l), is distributed from Central and Eastern Nepal to Burma and Indo-
China. M. capitellata, “Seti Kath” in Nepali, grows on exposed, degraded
lower slopes of the Schima-Rhododendron-o^k forest zone below 1 600 m where
it is very common, and to a lesser extent in secondary forests. It grows to a
tree, 4 to 9 m tall, with large (6-20 cm long), elliptic- lanceolate leaves
crowded near the branch tips, with small red glands on the ventral edge and
small round pinkish fruit.

The food plant of D. dipoeam\d D. eugenes, Myrsine semiserrataW^W. (1824)

36:1-15, 1997 (2000)

Fig. 16 D. dipoea male

Fig. 17 D. c//poea foodplant, Myrsine semiserrata

Fig. 18 D. dipoea second instar larva

Fig. 19 D. dipoea third instar larva

Fig. 20 D. dipoea fourth instar larva

Fig. 21 D. dipoea fifth instar larva

Fig. 22 D. dipoea pupa

/. Res. Lepid.

Fig. 23 Degraded scrub habitat, Bhamare creek, Godawari.

Fig. 24 Mature Schima-Rhododendron -oak forest, Nag creek, Godawari.
Fig. 25 D. Guides male perching.

Fig. 26 Male D. adonira feeding on stream bed.

36:1-15, 1997 (2000)

(Myrsiliaceae) (fig. 10, 17), “Kali Kath” in Nepali, ranges from Nepal
through Burma to central China and is found most commonly on wooded
slopes in the upper ScMma- Rhododendron zone from 1600 m to 2300 m.
It grows into a small tree 6 m high with lanceolate, entire denticulate leaves
with small, red dotted glands on the ventral margin.

The reported food plant of D. ouida and D. adonira , Maesa chisia Biich.
(Myrsiliaceae) (Sevastopoulo, 1946), is found from east Nepal to north
Burma. It grows into a small shrub or tree with 5 to 17 cm long lanceolate ,
glabrous, crenate leaves with small clusters of white flowers or fruit. Known
locally as “Bilauni”, it is used as an insecticide and grows on the edges of
disturbed forests and along streams below 1800 m. It is also the food plant of
two other local riodinids, Abisara fylla ( Doubleday, 1847) and Zemeros Jlegyas
(Cramer,! 780) (Sevastopoulo, 1946; Callaghan, unpublished data).

The distribution of Dodona in the study area follows that of their food
plants. The range of D. egeon is the same as M. capitellata, under 1700 m, while
D. dipoea?ind D. share the same distribution as M. semiserrata (fig. 27) ,

1700 to 2300 111. There is a slight overlap of food plant and Dodonar^\\ge.?> near
the 1700 111 contour on Bhaniare creek. As an experiment, I successfully
raised two D. eugenes larvae on M. capitellata. However, I found no evidence
in the field suggesting that the two species oviposit on each other’s food
plant.

D. ouida perch on the highest summits, above the range of M. chisia

, but return to the lower valleys for feeding. D. adonira has not been recorded
from the study area above 1900 ni, except for an old (1963) and dubious
record from Phuchok peak.

Larval Habits

The development time for D. citgcwcs averaged 51 days and Z). egeon 46 days
from oviposition to eclosure. The fifth instar had the greatest duration and
increase in body size . Early instar larvae spent all their time on the food plant,
feeding on plant tissues between the veins, consuming all leaf tissues only
from the third instar. Up to the fifth instar, the laiwae fed at any time of the
day, alternating with periods of inactivity when they rested against the center
vein of the leaf. Fifth instar larvae fed mainly at night. During molting and
pupation, the larvae moved off the food plant, resting on the sides of their
container, which suggests a similar behaviour in the field. Larvae raised in
the same container cohabited peacefully.

Besides crypsis, Dodona larvae defend themselves by raising the front half
of the body and snapping vigorously with their mandibles. Young D. dipoea
larvae raised both front and rear portions, and they may also regurgitate
black stomach contents. Despite these tactics, parasitism accounted for 32 %
of the field collected D. eugenes 19% by an unidentified dipterid and

1 3 % by an /c/irzcMmm wasp. The fly larvae appeared in the fifth instar, causing
the lar\a to enter the prepupal stage prematurely. A single fly larva emerged
through the body wall of each infected larva and quickly pupated. The wasp
larvae emerged through the body wall during the fourth instar, 5 to 7 per
larva, and quickly pupated, forming white cocoons.

]. Res. LepuL

Fig. 27 Study area, showing Dodona species distribution. D. egeon □
D. adonira ■ D. ouides • D. eugenes A D. dipoea A

36:1-15, 1997 (2000)

Fig. 28 Distribution of Dodona in Nepal. D. egeon □ D. adonira ■ D. ouides •

D. eugenes A D. dipoea A

Hilltop

Slope

i i , .i

Bill

oiream

WTjr^-r-TT

0000 1000 1100 1200 1300 1400 1500 1600
Hours

Species

D. ouida
D. dipoea
D. eugenes
D, egeon

Distribution of Observations. O = mean.

+++++0+++++

xxxxxxOxxxxx

********f«k******

Fig. 29 Spacing of male Dodona perching activity by locality and time of day.

/. Res. Lepid.

Adult Habits

Capture records for the Dodona species in the Kathmandu valley (C. Smith,
pers comm.) and my own observations suggest that most species fly through-
out the year. D. eugenes and I), dipoea are recorded from February to
December, D. cgcoTzfrom February to November, and I). oMirfa from January
to December. D. adonira flies from March to November. There are definite
peaks in the populations of these butterflies. Adults are most common in
April and May before the start of the rains. I observed a virtual population
explosion of D. egeon at Suryebineyak on April 21 ,1996. At Godavari popula-
tions peaked the middle of May, then dropped precipitously at the beginning
of the rains in June, increasing with the onset of dry weather in September.

The adults of both sexes fed mainly on algae, salts and bacteria growing on
damp soil, leaves and rocks near streams. Other food resources were pollen,
nectar and excrement. They were assisted by a modified proboscis with a
wide, flat tip supporting numerous small black lateral projections, resem-
bling a small brush when extended. When feeding, the proboscis was in
constant motion over the substrate. Evidently the greater area of the probos-
cis created by the projections facilitates the absorption of nutrients. This
modification is found in other Old World riodinid genera such as Abisara
(Callaghan, unpublished data).

WTien feeding, wings may be raised or flat (fig. 9, 16). In either case, the
lobes and tails at the anal angle of the hindwing bend outwards resembling
two eyes when viewed from behind, or, in the case of D. eugenes and D. egeon,
as eyes with antennae. The white scaling around the lobes of Z). ouides mimic
the butterfly’s eyes (fig 25) . These modifications suggest they ser\e to decoy
predators, like the “false heads” of Theclinae.

Feeding times also differed between species and sexes. D. egeon males and
females fed from 0845 to 1130, then from 1410 to 1540. D. owziZa females fed
between 1015 and 1120, then 1300 to 1530 and males froml330 to 1530. D.
cwgcTzcs females fed from 0830 to 0930, then 1200 to 1530 and males from
1215 to 1530. Female D. dipoea fed throughout the day and males fed from
noon to 1500. Three or four species may feed together. The feeding times,
particularly for the males, reflected perching times discussed below.

I observed oviposition activity in D. eugenes and D. egeon. D. eugenes ovipos-
ited at 1130 and D. egeon at 0910, 0945, 1115 and 1215. Females of both
species landed on the food plant leaf dorsal surface, walked to the edge, and
placed a single egg on the ventral surface by reaching underneath with the
abdomen. They then flew off in search of another plant.

Dodona species used perching behavior for locating mates, in which males
await females in prominent locations and at certain times. The hypothesis
that closely related, or congeneric riodinid species use different perching
times and localities as an isolating mechanism has been advanced previously
(Callaghan, 1982), and is examined here with respect to Dodona.

The results of my observations on perching behaviour for four Dodona
species are shown in figure 29. The micro-habitat type and the hours over

36:1-15, 1997 (2000)

which perching took place, and the mean of the obseiTations are shown for
each species.

Z). males (fig. 25) perched on ridges, especially on summits where up
to 6 or 7 individuals vied for preferred spots. Perching was from 0945 to 1400,
with maximum activity between 1100 and 1200. Males rested with wings
together on dorsal leaf surfaces from 1 to 3 meters high, but never on the
highest branches.

D. dipoea males perched on the hillsides along trails or on prominent
bushes from 0934 to 1300 with peak activity at 1200. They defended their
perches vigorously from other males, thus spacing themselves over the
habitat. Wlien a female appeared, the males would follow and when she
landed, hover in the air above, beating their wings to spread pheromones, as
suggested by the long scent hairs which cover both surfaces of the male
hindwings.

D. eugenes males also perched on hillsides from 0945 to 1200 with peak
activity at 0945, congregating around prominent vegetation, but withotit the
aggressiveness of Z). ouida or D. dipoea. Their resting position was the same.

D. egeon males perched along streams or gullies on the upper branches of
prominent trees from noon until 1345, defending their perching spots
vigorously against other males.

The foregoing observations suggest that sympatric Dodona species use
perching in different micro habitats as a mechanism to maintain species
isolation. Only D. eugenes and D. dipoea males perch in similar habitats, but
never together, and the peak of D. citgcncs perching activity is earlier..

Acknowledgements. I thank Mr. Colin Smith of the Annapurna Museum, Pokhara, for
his orientation to the study area, advice and distribution records for Dodona-, Drs T.C.
Majupuria and Krishna K. Shrestha of Tribhuvan University for identification of the
plant species, Mr. Mehendro S. Limbu of Godawari for his help in the field work, and
two anonomous reviewers for their helpful comments.

Literature Cited

Callaghan, C.J., 1982. A study of isolating mechanisms among neotropical butterflies
of the subfamily Riodininae.J. Res. Lepid. 21:159-176.

Johnston & Johnston, 1980. This is Hong Kong: Butterflies. Government Printer,
Hong Kong.

Kh.\dk.v, R.B. et a/. 1984. Ecology of Godawari Hills: a case study. In T.C. Majupuria,
ed. Nepal- Nature’s Paradise. Wdiite Lotus, Bangkok, pp. 408-426.
Sevastopoulo, D.G., 1946. The early stages of Indian Lepidoptera. Partiv.J. Bombay
Nat. Hist. Soc. 46: 253-269.

Smith, C., 1993. Illustrated checklist of Nepal’s butterflies. Rohit Kumar, India. 126

pp.

Journal of Research on the Lepidoptera

36:16-23, 1997 (2000)

On the correct placement of Erebia epipsodea Butler, 1868
within the genus Erebia Dalman, 1816 (Lepidoptera:

Satyridae)

Alexei G. Belik

P.O. Box 2108, Saratov 49, RU-410049, Russian Federation. E-mail: belik@san.ru

Abstract. It is demonstrated that the Nearctic species Erebia epipsodea But-
ler, 1868 is the closest relative to the Palaearctic species Erebia medusa
(Denis & Schiffermiiller) , [1775] and has no affinity with the species of
the Alberganus in which it was placed previously. This conclusion

is suggested by certain details of the male genitalic structure, but is con-
firmed by the structure of the female genitalia. Therefore E. epipsodea is
removed from the Alberganus species group and placed into the Medusa
group of species.

Since the time of the original description, the position of Erebia epipsodea
Butler, 1868 within the system of the genus Erebia Dalman, 1816 was not
stable. While describing it, Butler (1868) has clearly stated that the new
species is very similar to Erebia psodea (Hiibner, 1804): “Ate supra forma et
coloribus fere Psodeae (Hbn.)... Alae anticae subtus velut in Psodea sed magis
rufescentibus..E . In contrast to the explanation in Bird et al. (1995), the spe-
cific epithet “epipsodea” is given exactly in this connection: “epi” in Greeks
means “on”, “towards” and “psodea” is [at present] the name of a South-
east European subspecies of Erebia medusa (Denis & Schiffermiiller), [1775],
In the time of Butler the name Erebia psodea (Hiibner, 1804) was in com-
mon usage for the species called at present Erebia medusa (Denis &
Schiffermiiller), [1775].

Wien the structure of the male genitalia of E. epipsodea and E. medusawAS
studied and compared, the first species was placed far from the second one
on the basis that the male genitalia of both species look quite different
(Chapman 1898). Chapman divided the genus Erebiainto two sections and
nine groups. E. medusa placed in the section “A” group “VII”; E. epipsodea
ill section “B” group “VIH”.

However, even knowing this, at the same time Elwes again placed E.
epipsodea near E. medusa as its closest relative, basing this on the dear exter-
nal similarity of both species (Elwes 1898).

Warren refuted this point of view in his monumental work on the genus
Erebia (Warren 1936). He divided the genus into 15 specific groups, plac-
ing both discussed species in different groups, taxonomically distant from
each other. E. medusaw^'s placed into “IX. Medusa Group” while E. epipsodea
was placed into “XL Alberganus Group”.

Warren (1936) iiad noticed very characteristic features in the genitalia
of E. epipsodea: branches of juxta heavily chitinized and covered with teeth,

Paper submitted 17 May 1998; revised manuscript accepted 1 November 1998.

36:16-23, 1997 (2000)

Fig. 1. Erebia epipsodea: left valva, lateral view. USA, Montana, Missoula Co.,
Miller Creek, 12.VI.1982, S. Kohler leg.

Fig. 2. Erebia theano: left valva, lateral view. Canada, Manitoba, Churchill,
20.VII.1981, P. Klassen leg.

Fig. 3. Erebia alberganus: left valva, lateral view. Switzerland, Wallis, NE
Hohtenn/Lonza, Alp Tatz - Alp Laden, 11. VII. 1977, C. Hauser leg.

Fig. 4. Erebia medusa: left valva, lateral view. Russia, Chita region, Yablonovyy
mountain range, vie. Yablonovo, 20.VI.1995, A. Belik leg.

Fig. 5. Erebia kozhantshikovi: left valva, lateral view. Russia, Yakutia, Oymiakon
distr., vie. Ust’-Nera, 25.VI.1993, S. Sazonov leg.

and coarse teeth on the aedoeagus (Fig. 6). He noted that the presence of
these structures makes E. epipsodea a unique species within the whole ge-
nus. However he was certainly disoriented by two things. First is the gen-
eral superficial similarity of the form of the valvae in E. epipsodea male geni-
talia (Fig. 1) to those of the species of the Alberganus group. Though not
exactly resembling any species of the Alberganus group, the outline and
comparative sizes of valvae elements in E. epipsodea are especially similar to
those of some Nearctic representatives oi E. theano (Tauscher, 1806) (Fig.
2) . For the comparison, the shape of the valvae of E. alberganus is also shown
here (Fig. 3). Second is the clearly considerable difference in the form of
the valvae between the genitalia of £. epipsodea 3.nd ot' E. medusa (Fig. 4).

After the exhaustive work of Warren (1936) there were no further attempts

/. Res. Lepid.

Fig. 6. Erebia epipsodea: aedoeagus and juxta, lateral view. Canada, Manitoba,
Riding Mountains, 21. VI. 1982, P. Klassen leg.

Fig. 7. Erebia medusa: aedoeagus and juxta, lateral view. Russia, Chita region,
Yablonovyy mountain range, vie. Yablonovo, 20.VL1995, A. Belik leg.
Fig. 8. Erebia alberganus: aedoeagus and juxta, lateral view. Switzerland, Wallis,
NE Flohtenn/Lonza, Alp Tatz - Alp Laden, 11. VII. 1977, C. Hauser leg.

at critical revision of the genus Erebia. Kurentzov (1970), reviewing system-
atics and distribution of the genus Erebia both in the Eastern Palaearctic
and partly in the Nearctic region, mentioned E. epipsodea as a member of
the Alberganus Sipe.cies group. Later there were two publications byjapanese
authors. First of these publications was the paper of Murayama (1975) , which
was a brief illustrated abstract of Warren’s “Monograph of the genus Erebia'
rather than a new critical review of the genus. Published recently was the
well illustrated work of Kogure & Iwamoto (1992; 1993). In both these pa-
pers E. epipsodea was also placed into the Alberganus species group, though
the latter authors stated: “This species is placed in Group XI, Alberganus
group because of structural characteristics of the male genitalia, but its
morphological characteristics such as the size and the pattern of the wings
are similar to those of E. medusa" (Kogure & Iwamoto 1993).

The question about a close relationship between E. medusa and E. epipsodea
was raised again by Pringle (1992). It is demonstrated in that article that
male genitalia of is. medusa have the same characteristic features that War-
ren (1936) considered as unique for epipsodea. The branches of the juxta
are heavily chitinized and covered with teeth, and there are well developed
teeth on the aedoeagus (Fig. 7). For the comparison, the aedoeagus and
the juxta of F. alberganus iire also illustrated here (Fig. 8) to show the shape
of these structures in members of the Alberganus group. The author’s study
of specimens of E. medusa from various localities (from West Europe to
Transbaikal Siberia) has confirmed the data reported by Pringle (1992)
[Note: in all examined species the vesica is without cornutij. Warren seems

36:16-23, 1997 (2000)

a — length of tegumen
b — length of uncus

Fig. 9. Erebia epipsodea: tegumen and uncus, lateral view. USA, Montana,
Missoula Co., Miller Creek, 12. VI. 1982, S. Kohler leg.

Fig. 10. Erebia medusa: tegumen and uncus, lateral view. Russia, Chita region,
Yablonovyy mountain range, vie. Yablonovo, 20.VI.1995, A. Belik leg.
Fig. 1 1 . Erebia alberganus: tegumen and uncus, lateral view. Switzerland, Wallis,
NE Hohtenn/Lonza, Alp Tatz - Alp Laden, 11.VIL1977, C. Hauser leg.

to have completely overlooked these important details in the male genita-
lia of E. medusa.

Studying die morphology of various species of Erebia, the author has no-
ticed that the male genitalia of E. epipsodea and E. medusa have two other
similar features, which at the same time distinguish E. epipsodea from all
species of the Alberganus group. Sometimes these features are not clearly
developed, but on material from series it is quite notable. The first feature
is the comparative length of uncus and tegumen. In E. epipsodea and E. me-
dusa the uncus is shorter than the tegumen (Figs. 9-“10), in species of the
Alberganus growp the uncus is of equal length to the tegumen or even some-
what longer (Fig. 11). The second feature is that both in E. epipsodea and E.
medusa the uncus with gnathos is connected to the tegumen with a rather
acute angle, which varies from near 45° to 60° (Figs. 9-10). In members of
the Alberganus group the uncus with gnathos is connected to the tegumen
with a less acute angle, from 60° to 90° (Fig. 11). Numerous examples of
these facts maybe observed in the figures of Warren (1936: Figs. 334-338,
357-385); more examples of male genitalia of some North American spe-
cies of the Alberganus group are shown by Troubridge &: Philip (1983: Figs.
46-51).

However, all the mentioned features (phenetic similarity of E. epipsodea
with E. medusa and notable external difference of E. epipsodea from all spe-
cies of Alberganus group; the same features in male genitalic structures of

J. Res, Lepid.

p. a. — - papiilae anales

a. p. apophyses posteriores

!. p. — lamella postvaginalis

I. a. — lamella antevaginalis

0. b. — ostium bursae

p. !. a. — processus lamellae antevaginalis

d. b. c. — ductus bursae copulatrix

caudal end

Fig. 12. Erebia epipsodea: female genitalia, ventral view. Canada, Manitoba,
Riding Mountains, 5. Vi. 1977, P. Klassen leg.

Fig. 13. Erebia medusa: female genitalia, ventral view. Russia, Chita region,
Yabionovyy mountain range, vie. Yablonovo, 20.VI.1995, A. Belik leg.
Fig. 14. Erebia aiberganus: female genitalia, ventral view. Switzerland, Wallis,
NE Flohtenn/Lonza, Alp Tatz - Alp Laden, 1 1. VII. 1977, C. Hauser leg.
Fig. 15. Erebia medusa: sterigma (female genital plate), ventral view. Russia,
Chita region, Yabionovyy mountain range, vie. Yablonovo, 20.VL1995,
A. Belik leg.

E. epipsodea and E. medusa, which are lacking in the male genitalia of spe-
cies of the Aiberganus group) seem to be not quite enough to remove E.
epipsodea from the Aiberganus group and to place it into Medusa group.
There is still the shape of the valvae in the male genitalia of E. epipsodea,
which is not consistent with the idea of the affinity of E. epipsodea with E.
medusa.

The author believes that the form of the valvae in male genitalia within
the genus Erebia is a less stable trait, more subjected to adaptive radiation
and specialization during the evolutionary process of speciatioii. For ex-
ample, E. kozhantshikovi Sheljuzhko, 1925 undoubtedly belongs to the

36:16-23, 1997 (2000)

Alberganus group, but the form of the valvae (Fig. 5) may be veiy different
from the generalized shape of valvae in this group. At the same time the
form and comparative sizes of the uncus and tegumen, and the form and
chitinization of the juxta and aedoeagus seem to be much more conserva-
tive. So in certain cases the intrageneric arrangements of Warren, when
based primarily on the form of the valvae, are not natural.

The author’s study of the comparative morphology of female genitalia in
the genus Erebia has revealed new and indisputable proof that E. epipsodea
belongs to the Medusa group and has no relationship to the Alberganus
group.

The female genitalia of E. epipsodea (Fig. 12) are very similar to those of
E. medusa (Fig. 13). Both species have a structure in the female genitalia
the shape of which is very uncommon for the genus Erebia as a whole: a
very short flat triangular process associated with lamella antevaginalis (pro-
cessus lamellae antevaginalis), which is directed anteriorly. Therefore, the
ostium bursae opens freely to the ventral side. In females of most species of
the genus Erebia that were studied by the author, and in members of the
Alberganus gYoxvp in particular, the processus lamellae antevaginalis (of vary-
ing form, usually bifurcated at the distal end) is well developed. It is directed
caudally and therefore covers the ostium bursae from the ventral side. This
is illustrated for the case of alberganus (de Prunner, 1798) (Fig. 14); other
members of Alberganus group have female genitalia of similar shape. Fur-
thermore, in the female genitalia of E. epipsodea and E. medusa the lamella
postvaginalis has a characteristic convexity (Figs. 12-13, 15), while in spe-
cies of the Alberganus growp the lamella post\-aginalis is quite flat (Fig. 14).
[Notes the author believes that the structure of the bursa copulatrix has no
significant taxonomic value for the intrageneric systematics of the genus
Erebia. In all species examined, it has the same structure (with two signa,
identical in all species). Therefore the bursa copulatrix is not illustrated
on Figs. 10--13.]

Conclusion

Summarizing the preceding argument, it is clear that the Nearctic spe-
cies E. epipsodea is the closest relative of the Palaearctic species E. medusa,
having no affinity with members of the Alberganus group. So herein E.
epipsodea is removed from the Alberganus group and placed into the

Medusa species group of the genus Erebia.

Appendix: Material examined and the range of variations
The conclusions presented in this paper, to be meaningful, could not be
based merely on the study of single specimens. During the preparation of
the present paper, genitalia were examined of a representative series of
specimens from each discussed species and, for completeness of compari-
son, from all species of the Alberganus group:

Erebia epipsodea: 103 , 59 ; from Idaho, Wyoming, Montana and Manitoba.

/. Res. Lepid.

Erebia medusa: \1 6 , 79; from Norway, Austria, Italy, Bulgaria, Ukraiua,
Cisbaikal Siberia and Transbaikal Siberia.

Erebia alberganus: lOd, (39 ; from France and Switzerland.

Erebia maurisius (Esper, 1803): 5<?, 19; from Altai and East Sayan Mtns.
Erebia theano (Tauscher, 1806): 37d, 13 9; from Altai, East Sayan Mtns.,
Yakutia, Magadan region, Yukon, Manitoba, Montana, Wyoming, Colo-
rado.

Erebia youngi¥{o\\3.nd , 1900: 26 from Yukon;

Erebia dabanensis Erschoff, 1871: 20d, 49 ; from Polar Ural Mtns, Putorana
Plateau, East Sayan Mtns. and Magadan region.

Erebia anyuica Kurentzov, 1966: 13c^ , 2 9 ; from East Sayan Mtns. and Yakutia.
Erebia occulta Roos & Kimmich, 1983: 46,29 from Yukon.

Erebia kozhantshikovi?A\e\]\\z\\ko, 1925: b6 ,29 from Yakutia.

Erebia lafontaineiTrowhYidge. 8c Philip, 1983: 26 from Alaska.

A number of non-critical individual variations were seen in the genitalic
structures of all above-mentioned species. In the male genitalia, these indi-
vidual variations affect mainly the form of the valvae, while in the female
genitalia they affect the general shape of the sterigma and the form of the
processus lamellae antevaginalis.

Acknorvledgements. Serious work in the field of lepidopteraii systematics is impossible
without study of numerous scientific publications, the majority of which are for-
eign ones and inaccessible here in the heart of Russia because of known reasons.

The author expresses his sincere gratitude to the following persons: both Mr.
John B. O’Dell (St. Albans, England) and Mr. Willy De Prins (Antwerp, Belgium)
for their long-standing great help in providing him with many foreign literature
sources; Mr. Norbert G. Kondla (Genelle, Canada) for his veiy^ kind help with mod-
ern North American lepidopterological literature including the book “Alberta
Butterflies”; Mr. Kuniomi Matsumoto (Tokyo, Japan) for his most friendly help with
Japanese literature sources and for very' useful translations of them into English.

Special thanks of the author are addressed to Dr. Kenelm W. Philip (Eairbanks,
AK, USA) and to Dr. Clifford D. Ferris (Laramie, WY, USA) for constructive com-
ments on the present paper and for friendly correction of author’s English through-
out the text.

Literature Cited

Bird, C.D., Hilchie, G.J., Kondla, N.G., Pike, E.M., Sperling, F.A.H. 1995. Alberta
Butterflies. Edmonton, The Provincial Museum of Alberta, VIII-i-349 p.

Bulter, A.G. 1868. Catalogue of the diurnal Lepidoptera of the family Satyridae in
the collection of the British Museum. London, printed by order of the Trustees,

211 p.

Ch/\pman, T.A. 1898. A review of the genus Erebia, based on an examination of the
male appendages. Transactions of the Entomological Society of London 1898:
209-239.

Eewts, H.J. 1898. A revision of the genus Erebia. Transactions of the Entomological
Society of London, 1898:169-207.

36:16-23, 1997 (2000)

Kogure, M. 8c Ivvamoto, Y. 1992. Illustrated catalogue of the genus Erebia in color.
Yadoriga 150:2-33 [in Japanese] .

— — . 1993. Illustrated catalogue of the genus Erebia in color (II). Yadoriga, 154: 2-
38 [in Japanese].

Kurentzov, A.I. 1970. The butterflies of the far east USSR. Leningrad, “Nauka”
Publishing House, 164 p., 14 pi. [in Russian].

Murwama, S. 1975. A general view of the genus Erebia in the world. Gekkan-Mushi
54:9-14; 56:3-9; 57:3-6 [in Japanese].

Pringle, G. 1992. A note on the Satyrid butterflies, Erebia medusa (D.& S.) and Erebia
epipsodea Butler. British Journal of Entomology and Natural History 5:15-16.

Troubridge, J.T. 8c Philip, K,W. 1983. A review of the Erebia dabanensis complex
(Lepidoptera: Satyridae), with descriptions of two new species. Journal of
Research on the Lepidoptera 21 (2):107-146.

W.ARREN, B.C.S, 1936. Monograph of the genus Erebia. London, printed by order of
the Trustees, Adlard and Son Ltd., VII-i-407 p., 104 pi.

Journal of Research on the Lepidoplera

36:24-30, 1997 (2000)

Pontia occidentalis (Pieridae) Near Sea Level in California: a
Recurrent Enigma

Arthur M. Shapiro

Center For Population Biolog\% University of California, Davis, CA 95616

Abstract. Two definite and one probable Pontia occidentalis have been
taken near sea level in the Sacramento Valley of California in 27 years. This
species normally breeds above 1500m at this latitude. All were taken in
October, flying with the lowland sibling species P. protodice. The only
explanation of these captures that is at all parsimonious entails long-range
downslope dispersal, a seldom-documented event in montane non-migra-
tory butterflies.

Introduction

Pontia occidentalis (Reakirt), the Western White, and P. protodice
(Boisduval and LeConte), the Checkered White, are sibling species that
largely replace each other altitudinally in California. They are, however,
frequently synipatric in the western Great Basin, and intermittently so on
the mid-west slope of the Sierra Nevada (Shapiro 1992). P. occidentalis \s
not known to be resident anywhere in north-central California below
lOOOm, and its breeding range at the latitude of Sacramento is upslope
from 1500m. Most collections contain misidentified individuals of both
species, leading to erroneous distributional reports, but Shapiro (1977)
recorded a definite P. occidentalis wq/ay sea level in the Sacramento Valley.
This was noteworthy for at least three reasons: it was the first record of this
species in the California Central Valley, the first in Sacramento County,
and one of surprisingly few records of apparent long-range downslope
dispersal by a montane California butterfly. Low-altitude species, in
contrast, are commonly recorded high in the mountains and most
common Central Valley species have been recorded in most or all of the
Sierran counties. It is not clear that this strong asymmetry is purely a
function of either flight season or area, though both are likely to play
roles in it (Sheehan, Richerson and Shapiro, in preparation).

I have tracked the dynamics of P. protodice in both space and time in the
vicinity of Sacramento for 27 years, and the presence/absence of both
species along a permanent 10-station transect across California parallel to
Interstate Highway 80. Pontia protodice^inciwAi^.^ tremendously in abundance
and distribution in the Valley and indeed in most of its range, but in most
years the largest populations occur on dredge tailings along the American
River in northeastern Sacramento County; the capture of P. occidentalis
reported by Shapiro (1977) was made there. Since the early 1970s the
population density of P. protodice there has varied through four orders of
magnitude, and with several apparent local extinctions. Among many thou-
sands of individual Pontia examined here and elsewhere in the Sacramento
Manuscript accepted 29 April 1999.

36:24-30, 1997 (2000)

Valley, the 1976 P. occidentalis remained unique until 1995, when a second
(albeit problematic) individual was taken some 100m from the site of the
earlier capture! A third was then taken in nearby Yolo County in 1998. The
conditions of these captures are unusual enough as to require comment.

The first collection was a dark female of the “winter” phenotype “ca/ycc,”
taken 17.X.1976 at Rossmoor Bar, Rancho Cordova, Sacramento County
(19.7 111) amidst a dense flight of P. protodice. Both sexes were present, mostly
fresh, and presenting variable but normal early- autumn phenotypes easily
distinguished from P. occidentalis (fig. 1 ) .

The second specimen, a male, was taken at Rossmoor Bar 19 years later,
13.x, 1995 also in the company of numerous P. protodice, again of normal
seasonal phenotypes (fig. 2). This individual is somewhat ambiguous. It is
strikingly different from the others collected the same day, valuing in the
direction of P. occidentalis in most characters. Had it been taken in the
western Great Basin in an area of sympatry it would have been relegated to
the roughly 1% of wild specimens I cannot assign confidently to either
species, and suspect to be hybrids. These are quite variable among them-
selves, but most - including the 1995 Rossmoor Bar male - have been rather
closely duplicated among laboratory hybrids. Similar specimens also occur
in areas of sympatry in Colorado. Such ambiguous individuals are occasion-
ally taken within apparently pure ocalim/a/A populations, but this is the first
and only one I have gotten in an ostensibly pure protodice population. I
revisited the site at two-week intervals for the remainder of the season,
finding nothing unusual.

The third specimen, like the 1976 one, is a heavily-marked “cfl/ycc,” in this
case a male. It was taken among normal protodice at Willow Slough, Yolo
County (14.5 m), 10.X.1998 and is strikingly different-looking from them
(fig. 3) . Willow Slough is approximately 30 km due west of Rancho Cordova.
The site is a weedy, overgrown floodplain; the butterflies were nectaring at
Aster. I revisited Willow Slough three times from mid-October into early
November but found no more P. occidentalis. This is the first record of P.
occidentalis in Yolo County; it was not expected. P. protodice is often found at
Willow Slough in autumn, but is not persistent. For example, it was found
there in 6 of 22 Fourth ofjuly counts since 1977, and was common only twice
(1977 and 1992).

All three specimens are deposited in the Bohart Museum of Entomology,
UCD.

Discussion

These three P. occidentalis were captured within 7 calendar days (X.10-1 7) ,
but in different years. This hints at a common process giving rise to all three
records. The obvious candidate is downslope dispersal.

In all three cases the weather pattern during the preceding week was the
same, with strong high pressure and a gentle NNE (i.e., downslope) wind at
the surface and aloft, giving fair, warm conditions. This is a very common
autumnal pattern. I have reviewed my long-term records and can find no

/. lies. Lefnd.

Fig. 1 . Female P. occidentalis and several P. protodice collected with it, Rancho
Cordova, Sacramento Co., CA, 17.X.1976, upper and lower surfaces.

36:24-30, 1997 (2000)

Fig. 2. Male P. occidentalis/hybrld (?) and P. protodice collected with it, Rancho
Cordova, Sacramento Co., CA, 13.X.1995, upper and lower surfaces.

/. Res. Lepid.

Fig. 3. Male P. occidentalis and P. pro tod/ce collected with it, Willow Slough, Yolo
Co., CA, 10.x. 1998, upper and lower surfaces.

36:24-30, 1997 (2000)

pattern of dowiislope dispersal by other montane butterfly species under
these conditions; however, such dispersal is ver)^ rarely seen at all in non-
migratory montane species.

October is usually the month of maximum density and maximal areal
occupation in the Valley for P. protodice, but it is difficult to see how this could
account for the occurrence of P. occidentalis. Furthermore, October is not
usually the month of greatest abundance for P. occidentalis in its normal
montane range. Shapiro (1992) reviewed the dynamics of both species at
1500m on the Sierran west slope, where neither is a permanent resident. P.
occidentalis, whose nearest permanent population (at 1900 in) is less than 15
km away, dispersed to my Lang Crossing site in 8 of 20 years, and bred in 4.
This site is monitored biweekly from snowmelt through late October - early
November. Of 17 dates when it was recorded there, 6 were in August and 4
each in July and September - only 1 in October. There is no evidence of a
regular seasonal downslope movement, although we know P. protodice moves
upslopefrom the Nevada desert in late spring (Shapiro 1992). In some years
the densest populations of P. occidentalis ?iX 1500m on the Sierran east slope
do occur in October, where breeding occurs on Cruciferous weeds in
irrigated alfalfa. The three Valley captures, however, do not coincide with
known outbreaks of P. occidentalis on the east slope, and the dispersal
distances required are on the order of 200 km, including the crossing of the
Sierran crest. It is, however, noteworthy that both the 1976 and 1998
specimens correspond closely to the mean phenotypes flying at both 1500m
on the east slope and 2100m on the crest (Donner Pass) at that time. The
1995 specimen is too idiosyncratic for such a comparison, but would not be
“out of place” at either elevation as a putative hybrid; I have similar individu-
als taken in autumn at Sierra Valley, Sierra Co., an area of sympatry. See
Shapiro (1976) for phenotypic exemplars.

Shapiro and Geiger (1986) demonstrated electrophoretically that under
conditions of mutual abundance in sympatry, hybridization between these
two species must be a rare event since no heterozygotes were found for a
species-specific fixed allelic difference. It may occur more often when one
species is much more abundant than the other. Hybridization appears to be
more frequent in Colorado (J. Kingsolver, D. Wiernasz, personal communi-
cation).

When the species status of P. protodicem\d P. occidentalisw^s still unclear, the
occurrence of occidentalis within what should be pure protodice populations
could be ascribed to intrapopulational variation. This “explanation” is no
longer tenable, at least for the 1976 and 1998 specimens, which are unam-
biguously occidentalis using Chang’s (1963) and my own wing characters.
Neither specimen would arouse any special comment if labeled as coming
from 3000m in the High Sierra.

Shapiro (1977) observed that the similar habitat preferences and behav-
iors of the two species could account for a dispersing P. occidentalis lingering
in a prime proto dice hdhiVaX, such as Rossmoor Bar. That hypothesis remains
tenable.

/. Res. Lepid.

The idea that two and possibly three P. occidentalis would disperse in
different years from the montane Sierra to the floor of the Sacramento Valley
at exactly the same season - two to the exact same location! - and “join up” with
resident populations of the sibling species P. protodice, where they were then
accidentally discovered, strains credulity. Nonetheless, it is the only hypoth-
esis that is at all parsimonious, and it suggests that there may indeed be an
inconspicuous, low-density downslope movement by this species in autumn
that we should be looking for.

Acknowledgments. I thank Michael Plotkin for acceding to my impulsive request to be
dropped by the side of the road at Willow Slough on 10.X.1998, rather than at my
house or lab. Without that bit of serendipity I would be much less perplexed.

Literature Cited

Chang, V.C.S. 1963. Quantitative analysis of certain wing and genitalia characters of
Pieris in western North America. Journal of Research on the Lepidoptera 2: 97-
125.

StrvpiRO, A.M. 1976. The biological status of Nearctic taxa in the Pieris protodice-
occidentalis (Pieridae). Journal of the Lepidopterists’ Society' 30: 289-300.

. 1977. Apparent long-distance dispersal by Pieris occidentalis (Pieridae) .Journal

of the Lepidopterists’ Society 31: 202-203.

— — . 1992. Twenty years of fluctuating parapatry and the question of competitive
exclusion in the butterflies Pontia occidentalis and P. protodice (Lepidoptera:
Pieridae). Journal of the New York Entomological Society 100: 311-319.
Shapiro, A.M. & H.J. Geiger. 1986. Electrophoretic confirmation of the species status
of Pontia protodice and P. occidentalis (Pieridae). Journal of Research on the
Lepidoptera 25: 39-47.

Journal of Research on the Lepidoptera

36:31-44, 1997 (2000)

Effects of microclimate and oviposition timing on
prediapause larval survival of the Bay checkerspot butterfly,
Euphydryas editha hayemis (Lepidoptera: Nymphalidae)

Erica Fleishman, Alan E. Launer, Stuart B. Weiss, J. Michael ReecE, Carol L.
Boggs, Dennis D. Murphy-, and Paul R. Ehrlich

Center for Conservation Biology, Department of Biological Sciences, Stanford University,
Stanford, CA 94305, E-mail: eJJeish@leland.stanford.edu

Abstract. We tested empirically whether microclimate and relative timing
of oviposition affected prediapause larv^al survival and development rates
in the federally threatened Bay checkerspot butterfly, Euphydryas editha
bayensis (Nymphalidae). Most mortality in Bay checkerspot butterflies
occurs among prediapause larvae. Because phenology of the butterfly’s
lan^al hostplant, Plantago erecta, has been thought to drive prediapause
larval survival patterns, we also tested whether P. erecta senescence and
density over time varied among microclimatic zones. We found that
microclimate had a significant effect on P. erecta phenolog)'. Changes in
density of edible P. erecta among microclimatic zones were out of phase
temporally, but otherwise were similar. In the year of our study, neither
microclimate nor oviposition date tended to affect prediapause laiwal
survival, but both variables had significant effects on prediapause larval
development rates. Because temperature and precipitation patterns in the
butterfly’s environment vary' from year to year, whether microclimate and
oviposition date significantly affect prediapause larval survival and devel-
opment also may vary' annually. At least in some years, however, senescence
of P. erecta may not cause prediapause larval mortality. Our results support
the hypothesis that topographic heterogeneity is critical to the long-term
viability of the Bay checkerspot butterfly as well as other species that
inhabit temporally variable environments.

Keywords: Euphydryas editha bayensis, invertebrates, conservation, microcli-
mate, grasslands

Introduction

Spatial extent of suitable habitat is a fundamental consideration in conser-
vation planning for viable populations of virtually all species. Certain land-
scape attributes that must be emphasized in conservation planning for
invertebrates, however, differ from those that traditionally have received
attention in conservation efforts targeting large vertebrates (Ehrlich and
Murphy 1997). Habitat area is a primary concern for conservation of large
vertebrates. These animals often require sizable protected zones in which
population sizes can be maintained at or above a probabilistically safe

^ Address: Department of Biolog)', Tufts University, Medford, MA 02155
^ Present address: Department of Biology/314, University of Nevada, Reno, NV 89557

Paper submitted 14 April 1999; revised manuscript accepted 4 November 1999.

/. Res. Lepid.

baseline— for example, a 99% probability of remaining extant for 1000 years
(Shaffer 1981, Boyce 1992). Not only geographic extent per se but also
topographic heterogeneity of protected areas may be critical for the conser-
vation of many invertebrates and small vertebrates, including the Bay
checkerspot butterfly (Euphydryas editha bayensis) (Nymphalidae:
Nymphalinae) (Ehrlich and Murphy 1987, Weiss et al 1987, 1988, Laimer
and Murphy 1994). Spatial heterogeneity is important because invertebrate
population dynamics frequently are density-independent and highly sensi-
tive to climatic variability (Andrewartha and Birch 1954, Pollard and Yates
1993, DeVries et al 1997, Crisp et al 1998, Shaffer et al 1998).

The Bay checkerspot butterfly, which inhabits patches of native serpentine
soil-based grassland south of San Francisco, California, was listed in 1987 as
threatened under the U.S. Endangered Species Act. Serpentine-based soils
have a physical and chemical composition that limits the invasion of intro-
duced Eurasian grasses, and thus can provide refugia for native vegetation
(Ki'uckeberg 1 954, 1 984, Walker 1 954, Thomas 1961, Turitzin 1981, Huenneke
et al 1 990) . The viability of these native grasslands and of the Bay checkerspot
butterfly currently is jeopardized by suburban development (Murphy and
Ehrlich 1980, Ehrlich and Murphy 1981, 1987). Consemng sei'pentine
patches in the region is essential because the Bay checkerspot butterfly is
structured as a “mainland-island” metapopulation in which local demo-
graphic units frequently go extinct and temporarily unoccupied habitat
patches are recolonized (Ehrlich etal 1975, 1980, Murphy and Ehrlich 1980,
Ehrlich and Murphy 1981, 1987, Harrison et al 1988).

Prediapause Bay checkerspot butteidly laiwae suffer far greater mortality
than any other life stage (Singer 1972, Ehrlich et al 1975, 1980, Weiss et al
1988, Cushman et al 1994). Previous field studies estimated that survival of
prediapause laiwae rarely exceeds 10% annually (Singer 1972, Ehrlich et al
1975, 1980, Singer and Ehrlich 1979, Dobkin et al 1987, Weiss et al 1988).
Two interacting factors— microclimate and timing of oviposition during the
growing season— -are thought to affect rates of prediapause survival.
Prediapause larval suiwival is believed to be highest among offspring of early-
flying females that oviposit on cool north-facing slopes (Weiss et al 1987,
1988, Murphy et al 1990). On these slopes, the butterfly’s larval hostplants
[Plantago erecta (Plantaginaceae) and less commonly Castilleja densifloraor C.
exserta (Scrophulariaceae) ] remain edible until relatively late in the flight
season (Weiss et al 1987, 1988). Paradoxically, the females that fly earliest
tend to be those that fed and pupated on warmer south-facing slopes, where
hostplants senesce early and prediapause survival rates are thought to be
lowest (Ehrlich etal 1980, Weiss etal 1988, Murphy a/. 1990). Eggs laid well
into the flight season may be too late to produce larvae that survive on any
slope (Weiss et al 1988). For example, Cushman et al (1994) estimated that
just 1 week into the flight season, female reproductive success was less than
25% of that on the 1st day of the flight season. To date, estimates of
prediapause larval suiwival over space and time have been based on measure-
ments of hostplant senescence (Cushman etal 1994) rather than measured

36:31-44, 1997 (2000)

directly. The purpose of this study was to test empirically the influence of
microclimate and relative timing of oviposition on prediapause larval sur-
vival. In addition to quantifying hostplant senescence and density over time
in different microclimatic zones, we monitored the survival and develop-
ment rates of prediapause Bay checkerspot larvae that resulted from eggs laid
in different microclimatic zones on different dates during the flight season.

Study system

Euphydryas editha bayensis is univoltine. Adults fly for 3-5 weeks betw^een late
February and early May (Weiss et al 1988) . Females lay masses of 20-200 eggs
near the base of larval hostplants (Singer 1972, Weiss et al. 1988). Newly-
hatched larv^ae feed until they reach the 3rd or 4th instar and then enter an
obligatory diapause that lasts through the dry season (approximately May-
November) (Ehrlich 1965, Singer 1972). If hostplants senesce before larvae
reach the middle of the 3rd instar, the larvae starve prior to or die during
diapause (Singer 1972, Singer and Ehrlich 1979). Wlien the rainy season
begins, surviving larvae break diapause and feed on newly germinated
Plantago erecta until February or early March (Singer and Ehrlich 1979, Weiss
et al. 1988). Adults emerge following 10-20 days of pupation and generally
live for 1-2 weeks (Ehrlich 1965, Murphy et al. 1983, Cushman et al. 1994).

Extreme weather events can have markedly deleterious effects on Bay
checkerspot butterfly metapopulations (Singer and Ehrlich 1979, Ehrlich et
al. 1980, Murphy and Ehrlich 1980, Murphy et al. 1990). Wlien seasonal
precipitation is average or slightly above average, and the rainy season is not
prolonged, the geographic distribution of the butterfly tends to expand and
population sizes often increase. When precipitation patterns are extreme
(drought or deluge), however, or when the start of the flight season is
delayed by cool and cloudy weather, the geographic distribution of the
butterfly tends to shrink and its abundance tends to decline (Singer and
Ehrlich 1979, Ehrlich etal. 1980,Dobkin etal. 1987, Weiss etal. 1987, Murphy
et al. 1990).

Because variation in aspect and tilt affects solar exposure and retention of
soil moisture, local topography within habitat patches mediates hostplant
senescence and therefore plays a key role in enabling Bay checkerspot
butterfly metapopulations to survive extreme weather events (Ehrlich and
Murphy 1987, Weiss et al. 1987, 1988). For example, south-facing slopes
receive more solar radiation on clear days, thus are wanner and drier than
north-facing slopes. Plantago erecta on south-facing slopes often senesce 3-4
weeks prior to those on cooler north-facing slopes (Weiss et al. 1988).
Because hostplants on relatively cool slopes remain edible long into the
spring, those slopes are believed to serve as “core” habitat for the Bay
checkerspot butterfly. The availability of even a few cool slopes within a
habitat patch can prevent its butterfly population from being extirpated
during a short or mild drought. The importance of warmer slopes to the
persistence of Bay checkerspot butterfly populations should not be underes-
timated, however (Harrison et al. 1988, Weiss et al. 1988). Even very warm

/. Res. tepid.

slopes contribute to loiig-teriii viability of the Bay checkerspot butterfly by
providing diverse eaiiy-season nectar, which can increase female fecundity
and lifespan (Ehrlich and Murphy 1981, 1987, Murphy et al 1983, Boggs
1997). Proximity' of different microclimatic zones also is important because
postdiapause larvae that disperse from cooler to warmer slopes may advance
their adult emergence dates by a week or more, thus increasing their chances
of reproductive success (Weiss et al. 1987, Cushman et al. 1994). In sum,
survival and reproduction of the butteiily can occur under most macroclimatic
conditions in a patch of habitat that includes a range of slope classes (Weiss
et al. 1988).

Methods

Our experiments were conducted at Kirby Canyon, Santa Clara County, Califor-
nia, USA (37°1 1' N, 121°40' W) in spring 1993. This site includes approximately 1350
ha of serpentine soil-based grassland and is the butterfly’s largest remaining habitat
patch. The site is believed to serve as an important source of emigrants that
recolonize adjacent habitat patches from which the butterfly has been extirpated
(Harrison et al. 1988).

We selected 5 slopes as representatives of their microclimatic zones (Weiss el al.
1988, Cushman et al. 1994). Each was classified as very warm (south- and west-facing
slopes, tilt >17“), warm (south- and west-facing slopes, tilt >11“), moderate (all
aspects, tilt <11“), cool (north- and northeast-facing slopes, tilt >11“), and very cool
(north- and northeast-facing slopes, tilt >17“). Replication of microclimatic zones
was not tractable in terms of time and personnel requirements.

Plantago erecta phenology and density

To test the null hypothesis that Plantago erecta phenology does not vary among
microclimatic zones, we monitored the phenolog}' of 200 individual P. crccto through
the Bay checkerspot butterfly flight season. Prior to the flight season, when virtually
all P. erecta appeared edible (no visible senescence) and displayed only vegetative
growth, we randomly selected 40 P. erecta in each of the 5 microclimatic zones. We
monitored the phenolog)’ of each plant every 3-4 d over a period of 63 d, until all
plants had senesced. Phenology was ranked on a qualitative scale from 1 to 5 (1 =
strictly vegetative growth, 2 = partial flower, 3 = full flower, 4 = partial senescence, 5
= full senescence).

For each plant, we calculated the number of days between the start of the flight
season and each phenological stage (from partial flower through full senescence).
We conducted experimentwise comparisons of phenology (days from the start of the
flight season to each phenological stage) with a nested analysis of variance using the
General Linear Models Procedure (SAS 1990). Because microclimatic zones were
subsampled rather than replicated, we used the interaction term as the error sums
of squares; i.e., we calculated the E-value for each of the 4 analyses by dividing the
microclimatic zone mean square by the mean square for individual P. crccto within all
microclimatic zones. P-values reported for this and later analyses are for Type III
sums of squares. When there was a significant microclimatic zone effect, we com-
pared zones with Duncan ’s Multiple Range Tests. The significance level for these and
later Duncan’s Multiple Range Tests was set at alpha = 0.05.

We tested 2 hypotheses concerning tlie density of edible Plantago erecta during the
Bay checkerspot butterfly flight season. First, we tested whether the density of edible

36:31-44, 1997 (2000)

P. necta \’?ccwq\ among microclimatic zones at any given point in the flight season.
Approximately once a week through the flight season, in each microclimatic zone,
we measured the distance between 50 randomly selected, edible P. ejecta and the
nearest neighboring edible P. erecta. Plants were selected each week; we did not
monitor the same plants over time. Measurements were made on 7 d over a 45 d
period in all microclimatic zones. On Day 56, we only measured plants in the cool and
very cool zones because we were unable to find 50 edible P. erecta in the other 3
microclimatic zones. We tested the effect of microclimatic zone on P. erecta density
for each day on which measurements were made with analysis of variance using the
General Linear Models Procedure (SAS 1990). When there was a significant micro-
climatic zone effect, we used least-squared differences to compare zones. The
significance level for the latter tests was set at alpha = 0.05.

Second, we tested whether density patterns of edible Plantago erecta across time
(rather than on individual days) varied among microclimatic zones. This hypothesis
was tested with a General Linear Model /-test for detecting differences among
regression lines (Neter et al. 1990).

Larval survival and development

To test the hypothesis that prediapause larval survival and rates of prediapause
larval development did not vary' among microclimatic zones and oviposition dates,
we carried out the following protocol on each of 3 consecutive weeks during the
flight season. Weeks 1, 2, and 3 approximately corresponded to days 7, 14, and 21 of
the flight season. On the 1st day of each week, we captured at least 100 adult female
Bay checkerspot butterflies at Kirby Canyon. We fed them a sugar solution ad libidum
to encourage oviposition and then returned them to the field. In each microclimatic
zone, we placed 20 females in cylindrical cages over edible Plantago erecta (one
butterfly per cage). After several hours, we checked each caged site for presence or
absence of an egg mass. Butterflies were removed from the cages and released in the
area of capture.

We monitored the life stage of each group of offspring in the field every' 2-3 d for
47 d, until all animals had either entered diapause or disappeared. Development
usually was synchronous within each group. We scored the life stage of each group
on a scale from 1-6(1= egg mass, 2-5 = 1st through 4th instars, 6 = diapause) . Mortality
of egg masses or 1st or 2nd instar larvae often can be observed directly. Prior to 3rd
instar, disappearance also implies mortality (D.A. Boughton, unpublished manu-
script) . Many 3rd instar larvae disperse from the hostplant where they were deposited
as eggs. These larvae are cryptic and extremely difficult to track as they move through
the habitat. Dispersing 3rd instar larvae can molt and enter diapause after feeding
briefly (D.A. Boughton, unpublished manuscript) . They also, however, may stance or
be depredated. Therefore, our hypotheses addressed survival to 3rd instar rather
than to diapause. Because we were not able to monitor individual larvae, our
measurements of survival and development corresponded to survival or develop-
ment of at least 1 individual animal from each group.

We conducted Goldstein’s A:*-tests (Goldstein 1964), controlling first for oviposi-
tion date and then for microclimatic zone, to test the hypothesis that survival to 3rd
instar did not vary among microclimatic zones and oviposition dates. When there was
a significant effect of microclimatic zone or oviposition date, we used Goldstein’s
tests to compare survival at different life stages (i.e., survival between egg and 1st
instar, 1st and 2nd instar, and 2nd and 3rd instar).

To test the hypothesis that larval development rates did not vary among microcli-

/. Res. Lepid.

Table 1. Effect of microclimatic zone on phenology of Plantago erecta. Values

are mean ± o days from the start of the Bay checkerspot butterfly flight season
to each phenological stage. Black lines indicate means that are not significantly

different (alpha - 0.05).

Microclimatic zone

Phenological stage

veiy warm

warm

moderate

cool

very cool

partial flower

13.6 ±8.7

11.0±6.7

11.3±6.0

24.7 ±5.5

28.9 ±6.0

full flower

17.9 ±8.6

15.4 ±6.3

15.3 ±5.5

28.7 ±7.0

34.3 ±6.6

partial senescence

26.0 ±6.0

23.4 ±4.2

25.2 ±3.8

38.2 ±5.8

43.8 ±3.8

full senescence

34.4 ±6.3

31.0 ±6.0

33.2 ±6.1

45.8 ± 4.0

49.3 ±4.4

made zones and oviposition dates, we calculated the number of days between
oviposition and each larval instar for each group of offspring. We conducted
experimentwise comparisons of the days to 1st and 2nd instar with a two-way analysis
of variance using the General Linear Models Procedure (SAS 1990). Small sample
sizes precluded comparison of later life stages. When there was a significant effect of
microclimatic zone or oviposition date, we carried out among-zone and among-week
comparisons with Duncan’s Multiple Range Tests.

Results

Plantago erecta phenology and density

Numbers of days in each microclimatic zone from the start of the flight
season to each Plantago erecta^henologic^X stage are presented in Table 1 . We
rejected the hypothesis that P. erecta phenolog)^ does not vaiy among
microclimatic zones. The experimentwise effect of microclimatic zone on P.
crcctophenology was statistically significant (P<0.01) for each phenological
stage (partial flower: = 62.0, full flower: = 63.5, partial senescence:

F^ = 143.6, full senescence: F^ = 90.6) . P. crcc^aphenolog)' was not distinct
in each microclimatic zone, however (Table 1). Phenology of plants in the
very warm, warm, and moderate microclimatic zones often was not signifi-
cantly different (Table 1). Phenology of plants in the cool and very cool
zones, by contrast, grouped neither with each other nor with plants in any of
the warmer zones (Table 1).

Distances in each microclimatic zone from edible P. erecta to nearest
neighboring edible individuals throughout the Bay checkerspot butterfly
flight season are presented in Table 2. In each microclimatic zone, nearest
neighbor distances across the flight season tended to decrease as new P. erecta
germinated, then to increase as P. erecta senesced. The effect of microcli-
matic zone on nearest neighbor distances of edible P. crccto was statistically
significant for each of the distinct points in time at which measurements were
made, although the percentage of the variance in nearest neighbor distance

36:31-44, 1997 (2000)

Table 2. Effect of microclimatic zone on density of apparently edible (no visible
senescence) Plantago erecta. Values are mean ± a nearest neighbor distances
in mm. Degrees of freedom are 4,245 for days 1-45 and 2,98 for day 56. Black
lines indicate means that are not significantly (alpha = 0.05) different. *** = p<

0.0001.

Day

Microclimatic zone

veiy^ warm

warm

moderate

cool

very cool

1 28.1128.4

8.3111.0

10.219.9

19.5112.0

64.3156.3

29.9***

0.328

8 17.6120.5

8.11 8.9

9.2112.6

20.0116.8

32.8126.9

15.0***

0.196

14 16.9155.6

3.41 7.3

10.1112.8

11.0110.4

23.0120.7

10.8***

0.150

21 22.2±34.4 7.2110.5 19.6126.9 16.1120.2 29.914.9 4.6*** 0.070

28 62.9184.0 41.4132.6 40.7145.5 25.2120.4 28.5133.5 4.7*** 0.07

33 55.9153.4 68.1153.4 49.7±43.8 24.8117.9 41.4141.2 6.8*** 0.100

45 129.4194.2 114.9187.2 135.2182.7 38.3136.1 43.1142.9 21.4*** 0.259

56 312.21121.0 161.51100.0 46.1*** 0.320

explained by microcliiiiadc zone often was small (Table 2). This result
indicates that the relative timing of P. erecta germination and senescence
varies among microclimatic zones. Significant differences {P < 0.05) in
nearest neighbor distances among individual microclimatic zones are shown
in Table 2. At the beginning of the flight season, edible P. crccto densities were
greatest in the warm, moderate, and cool zones and lower in the very warm
and very cool zones. From roughly the middle to the end of the flight season,
the density of edible P. erecta was greatest in the cool and very cool zones.

Density patterns of edible P. across the season as a whole (rather than

on individual days) did not vary among microclimatic zones (Pjc.„ = 0.69,
Pyo5 crit ~ 2.23, P> 0.05). In other words, density patterns among zones were
out of phase temporally, but otherwise were similar.

Larval survival and development

Differences in Plantago erecta phenology Rve thought to be a key mechanism
by which microclimate affects survival of prediapause Bay checkerspot
butterfly larvae. We assumed a priori that the slopes on which we conducted
our experiment had different microclimates (Weiss et al. 1988, Cushman et
al. 1994). This led to the hypothesis that P. erecta senescence dates on each
of the 5 experimental slopes would differ significantly. Our analysis of P.
erecta phenology, however, rejected this hypothesis. Therefore, for analyses

/. Res. LepicL

Table 3. Number of groups of larvae with at least one representative surviving at

each life stage.

Week 1

egg

1st ill star
2iid iiistar
3rd instar
Week 2

egg

1st instar
2iid instar
3rd instar
Week 3

egg

1st instar
2nd instar
3rd instar

Microclimatic zone

warm group

cool

very cool

of larval survival and development, we grouped animals that had been
deposited in the very warm, warm, and moderate microclimatic zones. We
then tested whether (a) survival to 3rd instar and (b) development rates to
1st and 2nd instar differed significantly among 3 microclimatic zones (warm
group, cool, and very cool) and among oviposition dates (weeks 1, 2, and 3) .
Sample sizes are presented in Table 3.

In most cases (8 of 9 tests), microclimatic zone did not have a statistically
significant effect on sunival to 3rd instar (Table 4) . The single exception was
that groups deposited in the middle of the flight season (week 2) had a
greater probability of surviving to 3rd instar in the cool zone than in warm
microclimatic zones. This largely was due to different probabilities of survival
to 1st instar (x* = 2.725, P< 0.01). Probabilities of survival from 1st to 2nd
instar and from 2nd to 3rd instar were not significantly different between
warm and cool zones on week 2 (lst-2nd: x*' = 0.656 ns, 2nd-3rd: x* = 1.288
ns).

Likewise, only 1 of 9 tests showed a significant effect of oviposition date on
survival to 3rd instar (Table 4). Groups deposited in warm zones on week 1
had a significantly higher probability of surviving to 3rd instar than did
groups deposited in that zone on week 3. Survival from 1st to 2nd instar was
higher in warm zones for those deposited on week 1 than on week 3 (x* ^ -
2.800, P< 0.01). Suivival to 1st instar, and from 2nd to 3rd instar, however,
was not significantly different between weeks 1 and 3 (egg-1 st: x* = -1 .896 ns,
2nd-3rd: x*' = -0.256 ns).

Both microclimatic zone and oviposition date had a significant effect on
rate of development from oviposition to 1st instar (microclimatic zone:

36:31-44, 1997 (2000)

Table 4. Goldstein’s x*--tests for survival to 3rd instar. * =

P<0.05 (x*>

■k-k

P<0.01 (x*> 2.576),

*** = P< 0.001 (x*

> 3.291).

Within week

Within microclimatic zone

week 1

r*'

warm group

warm-cool

-0.351

weeks 1-2

1.786

warm-very cool

-0.962

weeks 1-3

1.989*

cool-veiy cool

-0.648

weeks 2-3

0.111

week 2

cool

warm-cool

-2.350*

weeks 1-2

-0.393

warm-very cool

-1.498

weeks 1-3

0.711

cool-very cool

0.244

weeks 2-3

1.000

week 3

very cool

warm-cool

-0.972

weeks 1-2

0.474

warm-very cool

-1.934

weeks 1-3

0.570

cool-very cool

-0.641

weeks 2-3

0.000

= 5.30, P < 0.01, oviposition date: = 44.80, P < 0.0001) and from

oviposition to 2iid iiistar (microclimatic zone: = 4.92, P= 0.01, oviposi-
tion date: = 27.13, P< 0.0001). The interaction of zone and date was not

significant {P- 0.19) and therefore was removed from the model. Groups in
warm zones developed more quickly than those in the cool zone (Table 5).
Surprisingly, groups deposited in the very cool zone on week 1 also devel-
oped to 1st and 2nd instar more quickly than groups deposited in the cool
zone on week 1 (Table 5) . Relatively high densities of edible P. erecta (that is,
limited senescence) may have accelerated the developmental rate of groups
in the very cool zone. Flowever, it is also possible that the accuracy of
estimates of development rates in the veiy cool zone was affected by small
sample sizes (Table 3). Within each microclimatic zone, mean rates of
development were significantly different on weeks 1, 2, and 3. Groups that
were deposited later in the flight season developed significantly more quickly
(Table 5). As discussed below, the latter result was not independent of
annual weather.

Discussion

It long has been assumed that interactions among topographic heteroge-
neity, hostplant senescence, and timing of oviposition mediate survival of
prediapause Bay checkerspot butterfly larvae and, by extension, population
sizes and geographic distribution of the butteiTly (e.g., Singer 1972, Ehrlich
etal 1975, 1980, Ehrlich and Murphy 1987, Weiss etal 1987, 1988, Cushman
et al. 1994). In our experiment, microclimate had statistically significant
effects on Plantago erecta phenology and density of edible individuals. In
terms of P. erecta phenology, we found that microclimatic zones tended to
group into three classes: warm, cool, and very cool. Similarly, by the middle
of the flight season, when members of the earliest experimental cohort of
offspring began to reach 1st instar and thus to feed, nearest neighbor

/. Res. Lepid.

Table 5. Development times (mean ± o) in d from oviposition to 1st and 2nd
instar. Black lines indicate means that are not significantly (alpha = 0.05)

different.

Microclimatic zone

1st instar

warm group

cool

very cool

week 1

15.9 ±2.0

17.0 ±2.3

15.7 ±0.6

week 2

12.0 ±2.1

14.8 ±1.7

week 3

11.2±1.4

1L3±0.4

1L9±L4

2nd instar

week 1

18.1 ±1.8

19.3 ±2.1

16.8 ±1.1

week 2

13.7 ±0.8

16.6 ±1.5

week 3

1L9±2.6

14.7 ±1.2

distances of edible P. erecta often grouped among the very warm, warm, and
moderate zones.

We found that microclimate had significant effects on rate of development
to 1 St and 2nd instar of Bay checkerspot butterflies. Oviposition date also had
a significant effect on laiv^al development rates to 1st and 2nd instar,
although daily weather patterns represent a potential confounding factor.
Because differences in annual weather patterns have complex ramifications
for plant senescence and invertebrate population dynamics, whether ovipo-
sition date significantly affects larval development may vary anntially.

Surprisingly, in the year that our study was conducted, neither microcli-
mate nor oviposition date tended to affect survival to 3rd instar of the Bay
checkerspot butterfly. Again, the effects of oviposition date on prediapause
larval survival may depend upon annual fluctuations in temperature and
precipitation. Caveats about temporal variability admittedly are frustrating;
scientists and managers naturally would prefer clear-cut rather than equivo-
cal experimental results. Yet variability and uncertainty are integral aspects
of natural systems that inevitably must be addressed in developing conserva-
tion plans for species or ecosystems. Recent advances in conceptual develop-
ment and implementation of adaptive management, which seeks to apply
scientific principles to decision-making in the face of uncertainty, reflect
growing recognition of the need to study and respond to shifting ecological
conditions (McLain and Lee 1998, Slocombe 1998). Similarly, Gaston et al.
(1998) argue that inability to conclusively accept or reject an ecological
hypothesis should be viewed as an opportunity to focus on drivers and
ramifications of variation rather than a deficiency of theory or method.

The absence of an effect of microclimate or oviposition date on larval
survival in this experiment also may be in part an artifact of our study design.
There is no tractable way to monitor individual prediapause larvae over many
days if the larvae are allowed to disperse freely. Therefore, we quantified

36:31-44, 1997 (2000)

siimval at the group level rather than at the level of individual animals. If we
had been able to track individuals, and most individuals deposited in the
same egg mass starved before reaching 3rd instar or diapause, our sur\aval
estimates would be reduced dramatically. Conversely, our sundval estimates
might increase if many individuals that disappeared in fact suiwived to 3rd
instar or to diapause. It is conceivable, although nearly impossible to
quantify, that microclimatic zone and oviposition date have significant
effects on the number of individuals per group that survive to diapause. We
therefore agree with the inference of previous investigators that most
reproductive females are likely to have some reproductive success, although
the number of offspring per female that survive to diapause often decreases
at later oviposition dates (Cushman et al 1994).

Our results suggest that at least in some years, it is erroneous to assume that
apparent senescence of P. erecta implies larval mortality (Ehrlich et al. 1975,
1980, Singer and Ehrlich 1979, Ehrlich and Murphy 1987, Cushman et al.
1994). Eor example, our data contradict the estimates of Cushman et al
(1994), which were based on hostplant senescence, that eggs laid after day
15 of the flight season (assuming a 28-day period of development from egg
to diapause) or day 19 of the flight season (assuming a 24-day period of
development) have no chance of reaching laiwal diapause. In our experi-
ment, at least 1 individual from 4-31 % of the egg masses laid on day 14 of the
flight season (which developed to 4th instar in 25-28 days) suiwived to 3rd
instar (the earliest stage at which lar\ae can enter diapause. Singer 1972).
Similarly, at least 1 individual from 4-25% of the egg masses laid on day 21 of
the flight season (which developed to 4th instar in 1 6-2 1 days) sinwived to 3rcl
instar. Again, our data cannot address the absolute number of individuals
that sunaved, only the fraction of groups that had survivors. Moreover, the
data of Cushman et al. were gathered in spring 1992, which was slightly
warmer and drier than in 1993.

There are several possible explanations why we found that laiwae survived
after the majority of their hostplants had senesced. Eirst, laiwae may have
developed on P. crccto that senesced later than most other P. erecta in the same
microclimatic zone. Second, although P. erecta that have begun to senesce
generally have been considered inedible (e.g., Cushman et al. 1994),
prediapause Bay checkerspot butterfly larvae can eat P. erecta seeds that are
green and developing even if the plant’s flowers are dead (M.C. Singer,
personal communication) . Third, the mobility of 3rd instar larvae is consid-
erable (mean = 17 mm in 10 min on warm sand; N. Mehdiabadi, Harrison,
and C. Boggs, unpublished data), and these lan^ae may be able to seek out
edible P. erecta even if those plants are few and far between. Eourth, it is
probable that prediapause Bay checkerspot butterfly larvae are facultative
cannibals (E. Fleishman, personal observation) that eat their siblings if
edible hostplants are not available.

Previous work (e.g., Singer 1972, Wliite 1974, Ehrlich et al. 1975, Weiss et
al. 1988, Cushman et al. 1994) suggested that survival of prediapause Bay
checkerspot butterflies occurs at the group level. In other words, if egg

/. Res. Lepid.

masses each contained 100 eggs, then 99% mortality could imply that all
individuals in one group simaved and all individuals in 99 other groups
starved. Our experiment suggests that survival instead may be spread widely
among groups. Whether the former or latter scenario is more accurate has
important ramifications for population dynamics and viability of the threat-
ened Bay checkerspot butterfly. As distribution of survival among groups
increases, so should the effective size (N^) of the butterfly population, as well
as its ability to withstand stochastic genetic events that can reduce probabili-
ties of long-term population viability (Allendorf 1986, Frankham 1996,
Rabinowitz et al 1986).

Although hostplants senesce earlier in warm microclimatic zones than in
cooler zones, distribution of offspring in warm as well as in cool zones likely
increases the long-term viability of populations of the Bay checkerspot
butterfly. For example, larvae that survive to diapause on warm slopes may
have relatively high reproductive fitness as adults because they eclose earlier
than individuals on cooler slopes in the subsequent year, when they have a
good chance of finding mates and can lay eggs while hostplants are still young
and edible (Weiss et al 1988). Also, macroclimate in coastal California is
notoriously unpredictable. Timing of P. crccto senescence relative to the Bay
checkerspot butterfly flight season, and the magnitude of the difference in
senescence timing among microclimatic zones, varies among years.
Postdiapause larval densities in warmer microclimatic zones tend to increase
in years following a relatively cool and wet flight season (e.g., Weiss et al.
1988).

Topographic heterogeneity likely is key to the persistence of numerous
residents of native grasslands and other temporally variable environments.
The need for topographic refugia may be especially pronounced among
native annual plants, invertebrates, and other species with relatively short
generation times or habitat requirements that vary thoroughout their life
cycle.

Research on checkerspot butterflies (Euphydryas) in the western United
States has been conducted virtually uninterrupted for the past 35 years.
Biological studies of such duration are notable both for their rarity and for
their ability to provide vital information for single- or multiple-species
conservation planning (Ehrlich and Murphy 1987, Stohlgren et al. 1995,
Heikkinen 1998). Nonetheless, our study emphasizes that it is critical to
examine empirically our assumptions about long-term study systems.

Acknowledgments. Thanks to D. Palmquist for statistical advice and J. Hellmann, C.
Thomas, and an anonymous reviewer for comments on earlier versions of the
manuscript. Field assistance was provided by R. Bell, E. Bjorkstedt, M. Fagan, K. Gish,
J. Hodgson, J. Hoekstra, S. Motland, M. Nielsen, D. Pollock, K. Switky, and A. Weiss.
Thanks to Waste Management, Incorporated and Thomas Reid Associates for access
to tlie study site and logistic support. Research was funded by Peter and Helen Bing
and Waste Management, Incorporated. All work involving Bay checkerspot butter-
flies was conducted under appropriate permits from and agreements with the U.S.
Fish and Wildlife Service.

36:31-44, 1997 (2000)

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Journal of Research on the Lepidoflera

36:45-78, 1997 (2000)

The Lepidoptera of Marine Corps Air Station Miramar:
Calculating Faunal Similarity among Sampling Sites and
Estimating Total Species Richness

John W. Brown

Systematic Entomology Laboratory, U.S. Department of Agriculture, PSI, ARS, c/o
National Museum of Natural History, Washington, DC 20560-0168 U.S. A., E-mail:
jbroxun @sel. bare, usda.gov

Katherine Bash

Department of Zoology, University of Texas, Austin, Texas 78712 U.S.A

Abstract. An intensive 3-year survey of the Lepidoptera of Marine Corps
Air Station Miramar in southwestern San Diego County, California, was
conducted from October 1995 through September 1998. Sampling rneth-
odolog)' included blacklight trapping (364 nights), diurnal collecting
(148 days), and pheromone “baiting.” About 646 species of Lepidoptera
were documented from the Station, including 20 (or more) undescribed
moth species and one “sensitive” butterfly species - Hermes copper,
Lycaena hermes (Edwards) . Two species were newly recorded for the United
States - Dryadaula terpsichorella (Busck) (Tineidae) and Metapluera polos?
Busck (Gelechiidae). While the species accumulation curve reached a
convincing asymptote, it is highly unlikely that all species of Lepidoptera
present on the Station were sampled. Four methods extrapolated or
estimated the fauna to be between 706 and 922 species. Based on the family
Geometridae, faunal similarityamongasubset of 10 permanent blacklight
sites ranged from 0.29 to 0.69. We briefly discuss how Lepidoptera
inventories may provide insight into identification of areas of high conser-
vation value.

Key Words: Insecta, Lepidoptera, faunal siiu/ey, inventory, coastal sage
scrub, conservation, species richness.

Introduction

Over the last decade the maintenance of biological diversity has become an
issue of both local and global concern. The values of maintaining biodiversity
have been discussed by numerous authors and were summarized best by
Ehrlich (1990) as ethical, aesthetic, economic, and the provision of “ecosys-
tem services.” Before we can attempt to maintain biodiversity we must know
its components; i.e., it is impossible to establish goals and/ or methods for the
long-term management and protection of biological resources without
knowing what resources are present. Therefore, the process of inventory
represents the first critical step in all efforts to effectively maintain biodiversity.

Because it is virtually impossible to inventory, monitor, and manage all
aspects of a local or regional biota, specific taxa may be selected as “indicator”
or “umbrella” species (or groups) based on their ability to reflect the diversity
or health of an ecosystem and their ability (or inability) to respond to

Paper submitted 2 October 1999; revised manuscript accepted 4 November 1999.

/. lies. Lepid.

changing environmental conditions. New (1998) and others (e.g., Eyre &
Rnshton 1989, Sutton & Collins 1991, Ki emen et ah 1993, Oliver & Beattie
1994) present convincing arguments that the use of insects for documenting
biodiversity, assessing ecosystem health, monitoring environmental change,
and identifying areas of high conservation value has many advantages over
the use of vertebrates or vascular plants (but for differing points of view see
Howarth & Ramsey 1991, Scott et al. 1 993, Noss & Cooperrider 1 994) . Owing
to insect abundance and diversity, the complex interrelationships between
them and other organisms form the most prevalent and comprehensive
elements of the fabric of all terrestrial and freshwater aquatic biological
communities (Powell 1995). Insects frequently exhibit rapid and perceivable
responses to habitat modification (e.g., local or regional changes in abun-
dance, extinction and colonization of habitat patches, range expansions and
contractions) (e.g., Kempton& Taylor 1974, Taylor etal. 1978, Pollard 1979,
Razowski 1985, Murphy & Weiss 1991); non-biased, standardized techniques
are available for sampling many types of insects (e.g., Merritt & Cummings
1978); large numbers of individuals can be sampled reliably over short
periods of time (e.g., Murphy & Weiss 1988a, b); collections of insects can be
stored easily and efficiently and maintained for verification of data and
future use; and there are fewer societal and ecological constraints to collect-
ing insects (e.g., Murphy & Weiss 1988b). The use of Lepidoptera as an
exemplar taxon for estimating overall insect diversity has advantages that
include the relative ease of identification at the species level (for many
families), standardized sampling methodology (e.g., Thomas & Thomas
1994), and a high correlation with the spatial, architectural, and taxonomic
diversity of vascular plants (e.g., South wood et al. 1979, Brown & Opler 1990,
Panzer &: Schwartz 1998). In addition, Lepidoptera comprise the richest
group of phytophagous insects in California.

A suiwey of the Lepidoptera of Marine Corps Air Station (MCAS) Miramar
(= the Station), situated in southwestern San Diego County, California, was
conducted from October 1995 through September 1998 by personnel
associated with the San Diego Natural History Museum, under contract with
the U.S. Ncivy. V\4iile the primary goal of the survey was to determine the
presence/absence of one butterfly species (i.e., Euphydryas editha quino
(Wright)) listed as endangered and two others (Lycaena hermes (Edwards)
and Euphyes vestris harbisoni Brown &: McGuire) formerly recognized as
candidates for listing by the U.S. Fish and Wildlife Sendee, the fieldwork
resulted in considerable information on local Lepidoptera diversity and
phenology. This information may represent a baseline against which the
effect of future environmental changes may be assessed.

Our purposes are to present an inventory of the Lepidoptera documented
from MCAS Miramar; provide a cumulative or summaiy seasonal phenology
for each species and the Lepidoptera community in general; estimate total
species richness of the Station based on a number of different assumptions;
discuss faunal similarity and site complementarity among a subset of 10

36:45-78, 1997 (2000)

blacklight sampling sites; and identify features that maybe useful in assessing
the potential consen ation value of any area based on its Lepidoptera fauna.

Materials and Methods

Study Site

The study site, MCAS Miramar (formerly Naval Air Station Miramar), is an
approximately 10,500-hectare (23,000-acre) property owned and managed by the
U.S. Marine Corps, situated in the southwestern portion of San Diego County,
California (Fig. 1), at about 33°N, 117°W. The Station extends nearly 16 km (10
miles) east-to-west, roughly from Santee Lakes to Interstate Highway 805, and about
5 km (3 miles) north-to-south, from Miramar Road to State Route 52. Elevations on
the Station range from about 30 to 250 m above mean sea level.

Wdiile portions of the Station are highly disturbed/developed, with aircraft
runways, warehouses, family housing, and an extensive road system, the vast majority
supports native biotic communities, including coastal sage scrub, scrub oak chapar-
ral, southern mixed chaparral, chamise chaparral, southern willow scrub, sycamore
alluvial woodland, oak riparian woodland, and valley needlegrass grassland. The
Station also supports extensive acreage of vernal pool habitat, a rare and highly
depleted, ephemeral wetland community. This situation is not unusual in California
or elsewhere in the United States, where militaiy’ reservations often represent
bastions of biodiversity in otherwise highly urbanized landscapes. Sampling was
restricted to portions of the Station where native habitat occurs; we sampled about
50% of the entire site.

Several of the plant communities on the Station are highly depleted assemblages
restricted to southern California. Among these are coastal sage scrub, the object of
significant conservation efforts and the focus of the State of California’s Natural
Communities Conservation Plan. This community supports a large number of plant
and animal species listed as rare, threatened, or endangered by the resource
agencies, the most notable of which is the coastal California gnatcatcher {Polioptila
californica califomica) , a small gray songbird. Coastal sage scrub occurs in a mosaic
distribution with other native scrub and wetland communities, each of which
contributes to the overall stability and long-term viability of the natural landscape.

Collecting Methods

Three general techniques were utilized to collect Lepidoptera; 1) blacklight
trapping; 2) diurnal collecting; and 3) pheromone “baiting.” Each of the methods
is described below.

Blacklight Trapping. Through a trial-and-error approach, 10 permanent black-
light trapping sites were established during the first three months of the program;
three additional sites were added during the second year (Fig. 1). The sites were
located in all major habitat types on the Station as characterized by the base
Geographic Information System vegetation mapping, including coastal sage scrub,
chaparral, grassland, oak woodland, willow woodland, and sycamore/oak woodland.
The thirteen sites are characterized in Table 1.

Regardless of season, ambient temperature, or phase of the moon, once a month
each of the sites was sampled using two, three, or four blacklight traps, each equiped
with a 15-watt ultra-violet light. The traps were deployed in the evening and retrieved
the following morning. Blacklight trapping was conducted on 364 nights over the 3-
year period. The frequency of trapping nights per month from October 1995
through September 1998 is illustrated in Fig, 2.

Marine Corps Air Station

J. Res. Lepid.

Fig. 1. yap of Marine Corps Air Station Miramar, with locations of sampling sites.

Table 1. Blacklight sampling sites.

36:45-78, 1997 (2000)

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Fig. 2. Frequency of blacklight sampling by month ; x-axis = month of the year, y-axis
= number of sampling dates; first bar of each month = 1 995-1996; second
bar = 1996-1997; third bar = 1997-1998.

Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep

Fig. 3. Frequency of diurnal sampling by month; x-axis = month of the year, y-axis
= number of sampling dates; first bar of each month = 1 995-1996; second
bar = 1996-1997; third bar - 1997-1998.

Diurnal collecting. Diurnal collecting was concentrated from the middle of
Febrnar)' through the end of June to coincide with peak adult activity of the three
target species of butterflies (i.e., Euphydryas editha quino, Lycaena hertnes, Euphyesvestris
harbisoni) . Additional diurnal collecting was conducted sporadically throughout the
remainder of the year. During the visits, meandering transects were walked while
searching for adult Lepidoptera, primarily butterflies. Diurnal collecting was con-
ducted on 148 days. The frequency of diurnal collecting per month from October
1995 through September 1998 is illustrated in Fig 3.

Because many species of butterflies are attracted to surface moisture, artificial

36:45-78, 1997 (2000)

Fig. 4. Generalized annual pattern of adult Lepidoptera activity on the Station; x-axis
= months of the year, y-axis = number of species collected/observed.

Fig. 5. Species accumulation curve; x-axis == cumulative number of sampling nights,
y-axis = cumulative number of species collected/observed.

“pxiddles” were created during several diurnal sur\'eys in May, June, and July 1996,
by spraying water onto the surface of dirt roads. Also, because males of many butterfly
species exhibit hilltopping behavior, brief visits frequently were made to the most
prominent hill in the vicinity of the survey locality.

During the first full year of diurnal surveys (i.e., 1996), minimal time was spent
examining vegetation for the presence of larval Lepidoptera; no time was invested
in subsequent years. Wlien lar\'ae were discovered, they were taken to the laboratory
and kept in half-gallon cardboard containers along with cuttings of the larval food
plant.

Pheromone baiting. Aluminum pie plates filled with ethylene glycol (anti-freeze)
(= pan traps) and baited with a synthetic sesiid moth pheromone were deployed at

/. Res. Lepid.

several of the permanent blacklight sites from about the middle of April through the
middle of August 1996.

In October 1996 we used synthetic pheromone of Hemileuca electra (Wright)
(Saturniidae) to determine the presence/absence of this species on the Station. In
February' 1997, we used a virgin female Saturnia walterorum Hogue 8c Johnson
(Saturniidae) in an effort to attract males of this species to determine the presence
of a resident population on the Station.

Data Analyses

The first date of capture was recorded for each of the identified species; for most
unidentified species this date was not recorded because there was uncertainty
regarding the taxonomic integrity of most of the “morphospecies.” A species
accumulation cim^e (Fig. 5) was derived by plotting the cumulative number of
species documented from the Station against cumulative number of sampling
nights; only identified species were included (n ~ 600).

For each identified species, all months of capture were recorded. The total
number of species documented for each month (January through December) was
tallied and used to generate a histogram illustrating the seasonal phenolog)^ or
temporal distribution of the entire Lepidoptera community. Although flight periods
of some species var\' from year to year depending on environmental cues (e.g., timing
of rainfall, winter/spring temperatures) , such annual fluctuations probably contrib-
ute little to the overall pattern of community phenology.

Using the family Geonietridae as an exemplar taxon, we constructed a matrix of
species by blacklight sampling site (using sites 1-10 as a subsample). We used this
matrix to evaluate faunal similarity and site complementarity. Faunal similarity (FS)
was calculated by the following equation: FS = C/ (A+B)-C, where “A” is the number
of species recorded from site A, “B” is the number species recorded from site B, and
“C” is the number of species shared by sites A and B. Complementarity (D)
(dissimilarity), which is defined as the inverse of faunal similarity, was derived using
the following equation: D = 1-FS, where FS equals faunal similarity.

Nomenclature, Arrangement of Taxa, and Disposition of Material

Scientific nomenclature and the sequence of families, genera, and species follow
Hodges et al. (1983). Specimens from fieldwork conducted in 1995-1996 are
deposited in the Entomolog)^ Department of the San Diego Natural Histor) Museum;
specimens from 1997 are deposited in the National Museum of Natural History,
Smitlisonian Institution, Washington, D.C.; specimens from 1998 are deposited in
the Essig Museum of Entomology, University of California, Berkeley. The last
material was used in development of the species accumulation curv e and the overall
species inventory, but not was included in the compilations of species’ phenology or
calculations of site complementarity because it was not examined by us.

Results and Discussion

Fieldwork resulted in the collection of about 30,000 specimens represent-
ing about 646 species (Appendix A). Because numerous specimens of
microlepidoptera are not yet identified (and are unlikely to be identified in
the foreseeable future), it is likely that the actual total may exceed 700
species.

36:45-78, 1997 (2000)

Nocturnal Lepidoptera

As would be expected, the Noctuidae (161 species; 25%) and Geonietridae
(90 species; 14%) comprise the largest portions of the Lepidoptera fauna.
Other families that are well represented in the fauna of the Station include
Pyralidae (including Crambidae) (75 species; 12%) and Tortricidae (60
species; 9%).

Among the material collected in blacklight traps were 20 (or more)
nndescribed species, including at least one species of Amydria ( Acrolophiclae)
(D. Davis, pers. comm.), a species of Lampronia (Incnrvariidae) (D. Davis,
pers. comm.), two species of Gnorimoschema (Gelechiidae) (J. Powell, pers.
comm., Povolny 1998), at least nine species of Blastobasidae (D. Adamski,
pers. comm.) , three species of Tortricidae, and four species of Noctuidae (T.
Mnstelin, pers. comm.). Also collected in blacklight traps were about 40
specimens of Metapleura potosi Bnsck (Gelechiidae), a Mexican species
previously unrecorded from the United States, and a single specimen of
Dryadaula terpsichorella (Bnsck) (Tineidae), also new to the U.S. fauna.

Among the Tortricidae are a new species oi Decodes (Powell & Brown 1998) ,
a new Epinotia (R. Brown, pers. comm.), and a new Eucosma (related to E.
hazelanaYAots) . Among the Noctuidae are a Aseptis, a new Xylomoia, a new
Lacinipolia, and a new genus (and species) near Miodera. It is highly unlikely
that any of the new species is restricted to the Station. For example, the new
Decodes is known from two of the California Channel Islands, Silverwood
Audubon Sanctuaiy (in central San Diego County), and northwestern Baja
California, Mexico (Powell 8c Brown 1998). The new genus of Noctuidae is
known from San Diego, Riverside, and San Bernardino counties, from sea
level to about 1400 m (Mnstelin, unpubl.).

Diurnal Lepidoptera

Species collected only during diurnal sampling represent about 1 1 % of the
Lepidoptera fauna of the Station. Four species of yucca moths (2 Tegeticula
and 2 Prodoxus; Prodoxidae) were collected from the flowers of Yucca whipplei
(Liliaceae). Five species of deaming moths (Sesiidae) were collected in pan
traps baited with pheromone. A single specimen oiHemaris diffinis (Boisduval)
(Sphingidae) was collected and a second individual observed. The latter
species previously was known in San Diego County only from the interior
montane region.

Other diurnal Lepidoptera included one species of fairy moth (Adela sp. ) ,
one species of Plutellidae (Pliniaca bakerella Busck), and a small number of
noctuids (e.g., 5c/imi<2spp.), geometrids (e.g., Stamnodess^Y^.) and pyralids,
about half of which were not duplicated in blacklight samples.

Cnephasia longana (Haworth) (Tortricidae) was one of about 10-12 species
collected both diurnally and in blacklight traps; it was abundant on the
Station. Powell (1997) chronicled the spread of this Palaearctic moth
southward and northward from the San Francisco Bay area, reporting it as far
south as Santa Rosa Island. The presence of this species on MCAS Miramar

/. Res. Lepid.

represents a southern range extension of approximately 300 km from its
previously documented range.

Using a synthetic pheromone for the diurnal buckmoth (Hemileuca electro) ,
we attracted males of this rapid-flying species. It was abundant on the section
of the Station known as Parcel G (Rubinoff 1998), but it is likely to be
common wherever flat-top buckwheat (Eriogonum fasciculatum Benth.;
Polygoiiaceae) is common. Using a virgin female Saturnia walterorum, we
successfully attracted males of this species on the Station.

Fifty-one species of butterflies were recorded, approximately 40% of the
butterfly fauna of San Diego County (i.e., about 125 resident species).
Furthermore, it is likely that a few additional butterflies species are present
and were not detected. For example, several common urban skippers
(Hesperiidae) present in adjacent developed areas were not detected on the
Station.

During 1996, single individuals of Hermes copper (Lycaena hermes) were
obseiwed at one location on the Station on two occasions. In 1997 this species
was found at five different sites; it was common in both 1997 and 1998.
Neither quino checkerspot {Euphydyras editha quino) nor Harbison’s dun
skipper (Euphyes vestris harbisoni) was observed.

Larval Lepidoptera

Larvae of only four species of Lepidoptera were collected in 1996: Orgyia
vetustaon Lotus scop arius (Fabaceae) ; Hemileuca electraon Eriogonum fasciculatum
(Polygoiiaceae); Apantesis nevadensis on Lotus, Eriogonum, and Erodium
(Geraniaceae); and Vanessa virginiensis on Gnaphalium californicum
(Asteraceae). All but the Vanessa were successfully reared to maturity.

Temporal Distribution

Appendix A presents a cumulative summary of the temporal distribution
(for 1995-1997) of the identified Lepidoptera based on all survey methods;
for most of the unidentified species, data are not presented. Based on the
larger moths, butterflies, and identified small moths, the potential number
of species per month ranged from a high of 226 in April to a low of 87 in
December, A histogram of the overall annual pattern of adult Lepidoptera
activity on the Station is presented in Fig. 4, In general, activity increases from
January, peaking in April and May, stays relatively high through September,
drops in October, and declines dramatically in December.

Wlien we examine one family, such as the Geometridae, we find consider-
able deviation from the overall pattern described above. For example,
geometrid adult activity was lowest in September (n = 14 species), increased
through late winter and early spring (December through February), and
peaked in March (n = 48 species). The Geometridae represented approxi-
mately 37 % of the species sampled injanuary. This deviation from the overall
pattern is not surprising: many geometrids fly in the winter and thus have
been given the common name “winter moths.”

Even during the period of lowest moth activity (i.e., December) , at least 87

36:45-78, 1997 (2000)

species (about 13.5% of the Lepidoptera fauna) were collected. Although
the vast majority of species exhibited distinct seasonal patterns, a few species
were present almost year-round: Pseudochelaria scabrella (Busck), Holcocera
gigantella (Chambers), Eucsoma pulverulenta (Walsingham), Amorbia cuneana
(Walsingham) , Pero radiosaria (Hulst) , Aethaloidia packardaria (Hulst) , Pherne
subpunctata (Hulst), Apantesis proxima (Guerin-Menetries), Agrotis ypsilon
(Hufnagle) , and Spodoptera exigua (Hiibner) . This pattern is most evident in
Tortricidae, Geometridae, and Noctuidae - families that include many
polyphagous, pest species that are opportunistic in their larval food plant
selection.

Faunal Similarity and Site Complementarity

Using the family Geometridae (n = 90 species) as an exemplar taxon, we
calculated faunal similarity and its inverse, complementarity (dissimilarity),
among a subset of 10 permanent blacklight sampling sites. The number of
species documented per site varied from 21 (23% of the geometrid fauna;
site 8) to 56 (62%; site 3); one species was collected only diurnally on a
meandering transect. This substantial variability suggests that although the
native habitat on the Station has a rather homogeneous appearance, plant
communities and the features that determine them (e.g., slope, exposure,
soil type) strongly influence the fauna, resulting in localized assemblages of
species. This finding corroborates the a priori (and highly logical) assump-
tion that in order to maximize the number of species sampled, it is imperative
to maximize the number of plant communities and microhabitats sampled.
It also suggests that blacklights may attract only geometrid species that are in
the immediate vicinity, suggesting localized or patchy distribution of many
species. These data may be biased by the fact that blacklight traps are not as
effective for geometrids as they are for noctuids (J. Powell, pers. comm.).

Faunal similarity (FS), as defined in the Materials and Methods section, is
equivalent to Jacard’s coefficient of similarity. This measure, emphasizing
shared presence and disregarding shared absence, is useful in many biogeo-
graphic and conservation contexts. Tables 2 and 3 present the “unreduced”
and “reduced” faunal similarity values, respectively, for a subset of 10
blacklight trapping sites (i.e., sites 1-10). As illustrated in Table 2, the highest
combined species richness for any two sites was 73 (81% of all geometrids)
(sites 6 and 7) and the lowest was 42 (47% of the geometrids) (sites 4 and 8).
The highest number of shared species was 41 (by sites 2 and 7, and sites 3 and
7), the lowest was 13 (by sites 4 and 8). The low values for both combined
species richness and shared species can be explained by the fact that sites 4
and 8 yielded the lowest numbers of species (i.e., 34 and 21, respectively) of
the 10 sites. Faunal similarity (Table 3) was highest (0.69) between sites 2 and
7 and lowest (0.29) between sites 1 and 8 and 6 and 8. Mean faunal similarity
was lowest for site 8 (0.32), a grassland area, and highest for site 2 (0.55), a
site supporting coastal sage scrub and sparse sycamore woodland.

Complementarity (dissimilarity) (Tabled) varied from 0.31 (between sites
2 and 7) to 0.71 (between sites 1 and 8, and sites 6 and 8). While

J. Res. Lepid.

Table 2. “Unreduced” values of faunal similarity (based on Jacard’s coefficient
of similarity) for a subset of 10 blacklight sampling sites. Highest and lowest

values in bold face.

123456789 10

Site 1

Site 2 36/60 ^

Site 3 37/65 39/67 -

Site 4 24/56 30/54 28/62 ^

Site 5 30/59 33/60 34/65 25/55 -

Site 6 33/68 37/68 39/71 28/61 35/63 -

Site 7 36/60 41/59 41/65 26/58 32/61 32/73 =

Sites 15/52 18/53 18/59 13/42 16/48 17/59 21/54 -
Site 9 37/59 38/62 40/66 25/59 33/60 36/69 38/62 17/54 »

Site 10 23/66 29/64 31/68 23/54 26/60 32/66 26/63 16/48 29/65 ^

Table 3. Faunal similarity (based on Jacard’s coefficient of similarity) for a
subset of 10 blacklight sampling sites. Highest and lowest values in bold face.

Site 1

Site 2

0.60

Site 3

0.57

0.58

Site 4

0.43

0.56

0.45

Site 5

0.51

0.55

0.52

0.45

Site 6

0.49

0.54

0.55

0.46

0.56

Site 7

0.60

0.69

0.63

0.45

0.52

0.44

Sites

0.29

0.34

0.31

0.33

0.29

0.39

Site 9

0.63

0.61

0.42

0.55

0.52

0.61

0.31

Site 10

0.35

0.45

0.46

0.43

0.48

0.41

0.33

0.45

Table 4. Complementarity (dissimilarity) for a subset of 10 blacklight sampling
sites. Highest and lowest values in bold face.

Site 1

Site 2

0.40

Site 3

0.43

0.42

Site 4

0.57

0.44

0.55

Site 5

0.49

0.45

. 0.48

0.55

Site 6

0.51

0.46

0.45

0.54

0.44 ■

Site 7

0.40

031

0.37

0.55

0.48

0.56

Sites

0.71

0.66

0.69

0.67

0.71

0.61

Site 9

0.36

0.39

0.58

0.45

0.48

0.39

Site 10

0.65

0.55

0.54

0.57

0.52

0.59

0.69 -

0.67 0.55

36:45-78, 1997 (2000)

coniplementarity is a relative indicator of dissiinilarity, its value does not
precisely reflect the presence of “different” species at the sites being com-
pared. For example, site 8 had the highest complementarity values, but
supported few species not present on other sites. Its overall high
complementarity is the result of its low species number (i.e., the absence of
shared species with other sites or “mismatches” based on absence) rather
than its uniqueness (i.e., the presence of different species or species not
present on other sites).

Species Richness Estimates

The species accumulation ciin^e (Fig. 5) shows that the rate of encounter-
ing new species increased throughout the first year and began to reach an
asymptote by October 1996. During the first year each successive month
added an average of about 40 species to the inventory. In contrast, during the
entire second year only 27 species total were added. This suggests that the
first year of the survey successfully captured about 95% of the fauna. Only six
additional species were added in the third year. Because sampling was
conducted with greater frequency during the first year, these findings may be
slightly biased; i.e., it is likely that uniform sampling during each of the three
years would have produced slightly different results.

VMiile we have documented about 646 species of Lepidoptera from Station
(not all of which are identified and many of which are undescribed), and the
species accumulation cim^e has reached a convincing asymptote, this num-
ber (646) may underestimate the fauna. The number of species potentially
present can be estimated or extrapolated by at least four different methods:
(I ) comparison with plant species richness; (2) extrapolation from butterfly
species richness; (3) evaluation of taxonomic components of the docu-
mented fauna; and (4) “Chao 1,” a non-parametric statistical model.

Powell (1995) has found Lepidoptera species richness to be 1.5™ 3.0 times
plant species richness at other sites in California (e.g.. Big Creek in Monterey
County). Because the flora of MCAS Miramar includes about 615 species of
vascular plants (Wier 8c Brown, impubk), a conservative estimate of the
Lepidoptera based on Powell’s findings would be 922 species. Based on
inventories of other sites in California, this estimate is too high; it may reflect
the fact that the Station has been subject to numerous intensive botanical
surveys that may be cumulatively more thorough than those of the sites used
for Powell’s calculations. Alternatively, Powell’s values of 1.5-3. 0 may not be
applicable over a broad ecological range. The latter seems unlikely given that
Powell has investigated habitats as diverse as redwood forest and coastal sand
dunes. It is more likely that such extrapolations are not meaningful owing to
the considerable difference in size of the study areas; i.e., the Station is nearly
seven times the size of Powell’s most thoroughly suiweyed site, Big Creek.

Throughout the western United States, butterfly species typically repre-
sent about 7% of the Lepidoptera species richness at any particular site or
geographic region (Powell 1995). Because butterflies are diurnal, easily
observed, and comparatively easily identified, resident butterfly species

/. Res. Lepid.

richness can be clocuineiited fairly accurately in the western United States.
If we extrapolate from the total number of butterfly species we obseiwed (n
= 51 ) , we would expect the total Lepidoptera fauna to be about 729 species.
This value seems a little high, but certainly is within a reasonable expected
range based on other California inventories.

W9ien we examine the relative components of the general “taxonomic”
categories of Lepidoptera (i.e., Microlepidoptera, Pyraloidea, butterflies,
Macrolepidoptera), we find that we are conspicuously low in the Microlepi-
doptera category. That is, Macrolepidoptera and Microlepidoptera typically
each represent about 40% of the Lepidoptera fauna at most sites in Califor-
nia (Powell 1995). Wliile we recorded about 287 species of Macrolepi-
doptera, we recorded only about 222 identified species of Microlepidoptera.
Many species of Microlepidoptera are confined to highly localized areas that
support their larval food plant, are not efficiently captured in blacklight
traps, are exceedingly small and difficult to prepare, and are difficult to
identify. If the Microlepidoptera richness of the Station is comparable to the
sampled Macrolepidoptera richness (i.e., 287 species) , the total Lepidoptera
fauna would be about 65 species greater than we documented, or about 71 1
species. This value is fairly consistent with our estimates of the number of
undetermined Microlepidoptera in our samples and the probability that this
group has been under-sampled using our methodolog)^

Chao 1 (Chao 1984) is a n on-parametric statistical model that can be used
to estimate species richness from samples (e.g., Colwell & Coddington 1994) .
It is represented by an easily calculated mathematical equation: + a“

/2b, where is the estimated species richness, is the observed species
richness (n ~ 646), “a” is the number of species represented by a single
specimen (n ~ 62), and “b” is the number of species represented by two
specimens (n ~ 32). The equation focuses on the number of species
represented by one and two individuals because these are likely indicators of
under-sampling. Using this equation we derive an estimate of about 706
species. This number is clearly a conser\^ative estimate (or under-estimate)
because with the exception it is based entirely on species that we have
been able to identify at least to morphospecies.

If we disregard the conspicuous outlier (n = 922), based on the three other
estimates described above, the total number of species on the Station maybe
between 706 and 729, with an average estimate of 715 species.

The most logical interpretation for the “false” asymptote of the species
accumulation curve is that the pool of species that can be detected reliably
using the methodologies we employed was nearly exhausted. This suggests
that utilization of additional or different collecting techniques would add
species to the inventory, in particular, the diverse leaf-mining fauna is poorly
represented in our samples.

It is apparent that to sample thoroughly the entire Lepidoptera fauna of
any site, one must employ a variety of techniques: diurnal sampling (i.e.,
approximately 1 1 % of the fauna of MCAS Miramar was detected only by
diurnal sampling), blacklighting (which yields the vast majority of moth

36:45-78, 1997 (2000)

species), pheromone “baiting” (which may be the best way to sample
adequately Sesiidae and some Saturniidae), and larval collecting (particu-
larly for leaf-mining and other Microlepidoptera) . Wliile our survey did not
demonstrate that multiple years significantly increase the number of species
detected (i.e., an increase of only about 5% was observed during the second
year) , it is likely that multiple years of effort are necessaiy to document some
species. Our results are biased by the fact that nearly 45% of our entire
sampling was conducted during the first year. According to Powell (1995),
multiple-year surveys are less affected by fluctuations in year-to-year abun-
dance and also have a higher likelihood of documenting vagrant and
migrant species that may not be resident. The latter is not exclusive to
multiple-year surveys; for example, it is highly likely that Magusa orbifera
(Walker) and Ascalapha odorata (L.), each recorded once from the Station,
are non-residents.

One of the greatest difficulties encountered while conducting Lepidoptera
surveys that focus on the entire order is the paucity of taxonomic expertise
to provide determinations of the samples, particularly for Microlepidoptera.
Wdiile species names are not vital for compiling an inventoiy and estimating
species richness, they are extremely useful. We found that the morphospecies
concept of identification was inadequate for determining the number of
species because of the similarity of many Microlepidoptera, especially
Gelechioidea. Although we received considerable assistance from many
taxonomists, because of the large number of specimens, it was impossible to
obtain determinations of all the material. We suspect that our blacklight
samples may include 5-10% more species that, when identified, will be “new”
to the inventory.

Conservation Context

Although we cannot assume that any of the species documented on the
Station are restricted to sensitive coastal sage scrub habitat, it is highly likely
that most contribute to the functioning of the larger biotic landscape, which
includes coastal sage scrub, through pollination, herbivoiy, or as prey. There
is little doubt that Lepidoptera phenology (e.g., timing of larval availability
as a food source for birds and small mammals) and density (e.g., amount of
prey resources available for predators and the amount of herbivore pressure
on plants) play a major role in determining the success or failure of many
biotic functions of the community.

From a conservation perspective, what sort of information can be extracted
from the Lepidoptera survey of MCAS Miramar? Wliile there are numerous
ways of examining biotic complexity or ecosystem health, we focus on three
criteria that may be useful in assessing the potential conservation value of any
site using Lepidoptera: (1) presence of endemics or rare taxa; (2) presence
of “weedy” species; (3) and species richness and complementarity.

The presence of numerous “regional endemic” species (e.g., Lycaena
hermes. Decodes /ic/ixPowell & Brown, Eucosma williamsiV Crambidia dusca
Barnes & McDunnough, and numerous undescribed species discussed

/. Hi’S. Lepid.

above) indicates that there probably is considerable native habitat on the
Station that is still intact, and that the general area supports biological
resources of “regional” significance. Although none of these species is listed
as threatened or endangered, and none is restricted to the Station, there may
be few Other places in southern California where as many co-occur. Hence
the Station may support an “assemblage” of regional endemics that exceeds
that found at other coastal southern California localities. The occurrence of
assemblages of rare species is a common phenomenon wherever rare or
depleted native habitats are present. For example, vernal pools throughout
the Central Valley of California support numerous plant species listed (or as
candidates for listing) as rare, threatened, or endangered, plus one or more
species of “sensitive” fairy shrimp (Anostraca) and at least one “sensitive”
amphibian.

The abundance of several widespread, weedy species, including Spodoptera
exigua, Pseudaletia unipuncta (Haworth), Agrotis ypsilon (all Noctuidae),
Cmephasia longana (Tortricidae), and others indicates the presence of dis-
turbed or degraded habitat. Wliile occurrence of these species can be
explained in part by the adjacency of urbanization, there is little doubt that
resident populations of these polyphagous “pests” are present in degraded
habitat on the Station. It is likely that the native Lepidoptera fauna has
suffered from the introduction of invasive weeds that serve as host plants for
weedy moth species. On the other hand, numerous weedy species common
in adjacent disturbed and/or urban areas are not present on the Station, or
are present in exceedingly low density. For example, Trichoplusia ni (Hubner) ,
Spodoptera ornithogalli {G\\^.\\€^.) , Peridroma saucia (Hiibner) (all Noctuidae),
Platynota slultana (Walsingham) , Crocidosema plebejana Zeller (both
Tortricidae), and Cadra cautella (Walker) (Pyralidae) are common or abun-
dant in adjacent urbanized and disturbed areas, but were uncommon or rare
on the Station. Monitoring strategies capable of detecting changes in the
abundance of these weedy species may provide insight into the affects of
future environmental change and/ or habitat perturbation on the Station.

Wliile the preseiwation of maximum biodiversity may seem intimately
linked with the preseiwation of areas of highest species richness, this may not
always be the case. Areas of highest species richness may represent areas of
greatest range overlap of common or widespread species. The number of
common, weedy species in an urban area may exceed the number of native
species in a depauperate, unique native habitat. Hence the total number of
species alone may say little about the overall conservation value of a site or
region. Other factors adding to the complexity of using species richness as
a measure of conservation value are differences in sampling strategies
leading to the richness numbers being compared, or the absence of com-
parative numbers altogether. For the Lepidoptera of MCAS Miramar, nei-
ther average species richness derived from the three methods of extrapola-
tion (n = 715) nor the documented value (n = 646) provide useful conserva-
tion information because there are no other numbers to compare, i.e,, no
other sites in California of this size have been surveyed as thoroughly.

An alternative to focusing conseiwation efforts on areas of high biodiversity

36:45-78, 1997 (2000)

is to focus on landscape diversity and site complementarity. Maximizing
coiisen'ation of the greatest variety of habitats, plant communities, slopes,
exposures, etc. almost certainly will lead to the preseiwation of the greatest
number of species. Evaluating site complementarity, likewise, may help
identify scenarios that capture the greatest number of species. For example,
areas of gabbro-derived or serpentine soils typically support exceedingly
depauperate floras because the unusually high magnesium and iron content
of the soil inhibits the growth of most plant species (Kiuckeberg 1954, 1969) .
Consequently, these areas would receive little or no consideration in conser-
vation efforts focused on areas of high diversity. However, these soils typically
support an endemicflora (Raven & Axelrod 1978) that has exceedingly high
conservation value. Under a strategy of conserving landscape diversity and
areas with high complementarity, these unique areas would receive attention
comparable to areas of high species richness. Just as high species richness
may not always be an indication of high conservation value, high
complementarity, likewise, may lead to conservation decisions that are not
optimal for maximizing the preservation of biodiversity. For high
complementarity to be an effective criterion, it must be the result of high
cumulative species richness (the denominator of FS; see Table 2) as well as
a low number of shared species. A site that supports only a subset of a more
diverse site may have a high complementarity value with a diverse site, but
make no contribution to the cumulative species richness.

In summary, Lepidoptera inventories may be used to focus conservation
efforts towards areas of endemism (e.g., areas that support assemblages of
regional endemics) and away from areas that support an abundance of weedy
species. Species richness, per se, may be of little assistance in assessing
conservation value, but a landscape approach that evaluates site
complementarity may be highly useful in capturing the greatest species
richness.

The growth of conservation biology and concern for the biological conse-
quences of environmental change has stimulated a new and intense interest
in ecological monitoring. However, before the results of monitoring pro-
grams can be interpreted, a baseline inventory is absolutely vital. We believe
that the results of this inventory of MCAS Miramar represent a baseline
against which changes in the Lepidoptera fauna of the Station can be
measured.

Acknowledgments. Fieldwork and data compilation for this study were conducted with
the financial support of the United States Navy, Naval Air Station (NAS) Miramar,
under contract number N6871 b95-LT-C0048 to the San Diego Natural History
Museum. Data analyses were completed, in part, with funding from the National
Science Foundation’s “Research Experiences for Undergraduates” program, which
supported Katherine Bash’s participation in the Smithsonian Institution’s 1997
Research Training Program. We thank the following for field logistic support and/
or contract administration and coordination: Tamara Conkle, MCAS Miramar
(formerly NAS Miramar); Tommy Wright, formerly Southwest Division, Naval
Facilities Enginerring Command; and Phil Unitt, San Diego Natural History Mu-
seum. Field work was conducted by Norris Bloomfield, John W. Brown, John Brown,

/. Res. Lepid.

Jr., and David Faulkner, widi Bloomfield shouldering the lion’s share. Steve McElfresh,
University of California, Riverside, provided synthetic pheromone of Hemileuca
elecira to Dan Rubinoff, who conducted field work on this species; Thomas Eichlin,
California Department of Food and Agriculture, Sacramento, California, provided
sesiid pheromone; and Tomas Mustelin provided the female Saturnia walterorum.

The following provided determinations of specimens: David Adamski, USDA,
Systematic Entomology Laborator)', National Museum of Natural History, Washing-
ton, D.C. (Blastobasidae); Richard Brown, Mississippi State University (Epinotia);
Don Davis, National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. (Tineidae); Thomas Eichlin, California Department of Food and Agricul-
ture, Sacramento (Sesiidae); Douglas Ferguson, USDA, Systematic Entomology
Laborator)', Smithsonian Institution, Washington, D.C. (Geometridae) ; Peterjump,
Santa Paula, California {Acrolophus, Acrolophidae); Lauri Kaila, Finnish Museum of
Natrual History, Helsinki (Elachistidae) ; Ron Leuschner, Manhattan Beach, Califor-
nia (Noctuidae, Geometridae, and Pyralidae) ; Tomas Mustelin, San Diego, Califor-
nia (Noctuidae); Jerry Powell, Essig Museum of Entomology, University of Califor-
nia, Berkeley (microlepidoptera); and Ron Robertson, Santa Rosa, California
(Noctuidae).

We thank the following for comments on the manuscript, which enhanced its
quality and clarity: Tamara Conkle, MCAS Miramar, San Diego, California; Marc
Epstein, National Museum of Natural History', Washington, D.C; Sonja Scheffer,
Systematic Entomology Laboratory, USDA, Beltsville, Maryland; David R. Smith,
Systematic Entomology Laboratory', National Museum of Natural History, Washing-
ton, D.C.; Paul A. Opler, Colorado State University, Fort Collins, Colorado; andjerry^
A. Powell, University of California, Berkeley.

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/. Res. Lepid.

APPENDIX A. Adult flight period for Lepidoptera based on capture records and
observations; months abbreviated by capital letters; + = observed or captured, -

= not observed.

Months J FMAMJ JASOND

Nepticulidae

Stigmella (?) sp. 1

Stigmella (?) sp. 2 . . . .

Stigmella (?) sp. 3 - _ . +

Opostegidae

Opostega bist.riguella Braun + - + +

Opostega sp. +

Tischeriidae

Tischeria sp. +

Incurvariidae

Lampronia sp. + _ . .

Adela flammeiisella Chambers - - - +

Prodoxidae

Grey a sp.

Tegeticula maculata (Riley)

Tegeticula yuccasella (Riley)

Prodoxus marginatus Riley
Prodoxus cinereus Riley
Prodoxus aenescens Riley

Acrolophidae

Cephi tinea obscurostrigella (Ch.) - - +

Amydria arizonella Dietz
Amydria cunristrigella (Dietz) - + +

Amydria confusella Dietz + + +

Amydria oblique lla Dietz _ _ _

Amydria erecta {Er?iun)

Amydria n. sp. _ . .

Amydria sp (d^rk) . _ .

Acrolophus kearfotti (Dyar) . _ -

Acrolophus laticapitanus _ _ _

Acrolophus pyramellus {E. 8c McD.)-
Acrolophus variabilis (W<i\s.) _ _ .

+ - + + +

+ - - - +

+ + + + -

+ - - + +

+ + + +

+ ---

+ + + + +

+ + +

+ + + + +

+ + +

+ +
+ +

+ +

Tineidae

Tinea occidentalis Chambers + - + +

Opogona sp. 1 - - + -

Opogona sp. 2 . . _ _

Opogona omoscopa (Meyrick) - - + -

Dryad aula terpsichorella (Busck) - _ . -

ca. 2 undetermined Tineidae

+ + + + + +

+ - + + + +

+ . - . -

36:45-78, 1997 (2000)

Months

J FMAMJ J ASOND

Bucculatricidae

Pamlnicoptera sp.

Bucculatrix sp. +

Gracillariidae

Caloptilia sp. 1
Caloptilia sp. 2
Caloptilia sp. 3
Caloptilia sp. 4

ca. 2 undetermined Gracillariidae

Oecophoridae

Ethmia arctostaphylella (Wals.)
Ethmia discostrigella (Chambers) -
Inga concorella (Beutenmuller)
Pleurota albastrigiilella (Kearf.)
ca. 2 undetermined Oecophoridae

+ +

Elachistidae

Coelopoeta glutinosi Walsingham
Elachista coniophora Braun
Elachista lurida Kaila

Blastobasidae

Symmoca signatella (H.-S.)
Holcocera gigantella Chambers
Holcocna n. sp. 1
Holcocera n. sp. 2
Hypatopa interpunctella (Dietz)
Hypatopa n. sp. 1
Hypatopa n. sp. 2
Hypatopa n, sp. 3
Hypatopa n. sp. 4
Blastobasis n. sp. 1
Blastobasis n. sp. 2
Blastobasis n. sp. 3

-h

-t-

-h +

-t- +

-t-

+ +

Coleophoridae

Coleophora accordellaWlsm.
Coleophora sp. 2
Coleophora sp. 3
Coleophora sp. 4
Coleophora sp. 5
Coleophora sp. 6
ca. 5 undetermined sp.

+ + + ----

Momphidae

Mompha eloisella (Clemens)
Mompha sp.

+ + +

/. Res. Lepid.

Months

Cosmopterigidae

Anlequera acertella (Biisck)

Cosmopterix sp.

Stagmatophora iridella Biisck

Stagmatophora enchrysa (Hodg.)

Pyroderces sp.

Anoncia sp. 1

Anoncia sp. 2

Stilbosis sp.

Walshia miscecolorella

ca. 2 undetermined Cosmopterigidae

■

Scythrididae

ScytJiris sp. 1

Scythris sp, 2

Scythris sp, 3

Gelechiidae

Isophrictis sp.

Aristotelia elegantella (Cham.)

Aristotelia sp. 1 (tan)

Aristotelia sp. 2 (rust)

Exotelia californica (Busck)

Exotelia graphicella (Busck)

Leucogoniella californica (Keifer)

Telphusa sedulitella (Btisck)

Pseiidochellaria scabrella (Busck)

-t-

-f

Tfleiopsis baldiana (Bar. 8c Bus.)

H-

Gelechia sp. 1

Gnorimoschema powelli Povolny

”

Gnorimoschema sp. 1

”

Gnorimoschema saphirmella (Ch.)

Chionodes figurella (Busck)

”

Chionodes notandella (Busck)

Chionodes ochreostrigella (Cham.)

Chionodes sp, 1

Chionodes sp. 2

Chionodes sp. 3

Chionodes sp. 4

Chionodes sp. 5

Eilatima sp. 1

Eilatima sp. 2

Aroga morenella (Busck)

A nacampsis lacteusochrella ( Ch . )

Dichomeris baxa Hodges

Metopleura potosi Busck

Gelechiidae sp. 1

Gelechiidae sp. 2

Gelechiidae sp. 3

36:45-78, 1997 (2000)

Months J FMAMJ J ASOND

Gelechiidae sp. 4

ca. 22 undetermined Gelechiidae

Alucitidae

Alucita hexadactylon L. _ _ _ +

Carposinidae

Bondia comonana (Kearfott) - +

Plutellidae

Euceratia securella Walsingham
Plmiaca bakerella Busck
PhUella albidorsella (Wals.)

Plutella xylostella (L.) . .

Plutella sp. (tan)

Ypsolopha cervella (Walsingham) -

Yponomeutidae

Ze//ena sp.(?) -h -f

Sesiidae

Paranthrene robinae (Edwards)
Synanthedon poiygoni (Edwards)
Synanthedoff resplendens (Edw.)
Synanthedon exitiosa (Say)

Penstemonia hennei Engelhard

+ +

+ -h - -

+ + +

Cossidae

Prionoxystus robiniae (Peck) _ _ _ _ +

Tortricidae

Episimus argutanus (Clemens)
Bactra furfumna (Haworth)

Bactra verutana Zeller
Bactra priapeia Heinrich
Endothenia hebesana (Walker)
Endothenia nubilana (Clemens)
Rhyacionia frustrana (Comstock) -
Phaneta apacheana (Wals.) +

Phaneta misturana (Heinrich)
Phaneta pallidarcis (Eleinrich)
Phaneta subminimana (Eleinrich) -
Phaneta sp. (brown) +

Eucosma ridingsana (Robinson)
Eucosma sandiego Kearfott
Eucosma n. sp. (nr. hazelana)
Eucosyna costastrigulana Kearfott -
Eucosma williamsi Powell
Eucosyna pulveratana (W 'As.) -i-

Months

Eucosma nr. passerana (Wals.)
Eucosma sp. 1
Eucosma sp. 2

Epiblema streniiana (Walker)
Suleima lagopana (Walsingham)
Sonia filiana (Busck)

Pseudexentra habrosana (Heinrich)
Chimoptesis chrysopyla Powell
Crocidosema plebejana Zeller
Epinotia siskiyouana Heinrich
Epinotia subplicana (Wals.)
Epinotia sagittana McDnnnongh
Epinotia Columbia (Kearfott)
Epinotia bigemina Heinrich
Epinotia kasloana McDnnnongh
Epinotia signiferana Heinrich
Epinotia n. sp.

Ancylisnv. simuloides (McD.)
Ancylis mediofasciana (Clemens)
Cydia latifeneaniis (Wals.)
undet. Olethreutinae sp. 3
undet, Oledireutinae sp. 4
Acleris senescens (Zeller)

Acleris foliana (Walsingham)
Cnephasia longana (Haworth)
Decodes fragarianus (Bnsck)

Decodes asapheus Powell
Decodes helix Powell & Brown
Anopina triangulana (Kearfott)
Argyrotaenia niscana (Kearfott)
Argyrotaenia franciscana (Wlsm.)
Archips argyrospila (Walker)

Clepsis peritana (Clemens)
Sparganothis senecionana (Wlsm.)
Platynota stultana (Walsingham)
Amorbia cuneana (Walsingham)
Henricus umbrabasanus (Kearfott)
Lorita scarificata Meyrick
Cochylis carmelana Kearfott
Saphenista (?) sp. 1
Saphenista (?) sp. 2
Saphenista (?) sp. 3

Hesperiidae

Erynnis tristis (Boisdnval)

Erynnis funeralis (Send. & Burg.)
Pyrgus albescens Plotz
Heliopetes ericetorum (Boisdnval)
Hylephila phyleus (Drury)

36:45-78, 1997 (2000)

Months

J FMAMJ J ASOND

Atalopedes campestris (Boisdiival) -
Ochlodes agricola (Boisduval)
Ochlodes sylvanoides (Boisduval)
Poanes melane (Edwards)

Lerodea eufala (Edwards)

PapUionidae

Papilio zelicaon Lucas
Papilio cresphontes Cramer’

Papilio rutulus Lucas
Papilio eurymedon Lucas

+ + +

-h

Pieridae

Pontia protodice (Bois. & LeCon.) -
Pieris rapae (h.)

Anthocharis sara Lucas
Colias eiirytheme Boisduval
Colias eurydice^o\sdu\2iV
Phoebis sen nae {L.)

Nathalis iole Boisduval

Lycaenidae

Lycaena hermes (Edwards)

Satyrimn sylvinum (Boisduval)
Satyrium tet.ra (Edwards)

Satyrium saepium (Boisduval)
Callophrys dumetorum (Bois.)
Incisalia augustinus (Kirby)

Stryinon melinus Hiibner
Brepliidium exile (Boisduval)
Leptotes marina (Reakirt)

Celastrina ladon (Cramer)

Philotes sonorensis (Feld. & Feld.) -
Euphilotes bernardino (B, & McD.) -
Glaucopsyche lygdarnus (Doubl.)
Icaricia acmon (West. & Hew.)

+ +

-h

Riodinidae

Apodemia mormo (Feld. & Feld.) -

Nymphalidae

Agraulis vanillae (L.)

Nymphalis antiopa (L.)

Vanessa viginiensis (Drury)

Vanessa cardui (L.) -t-

Vanessa annabella (L.)

Vanessa atalanta (L.)

Junonia coenia (Hiibner)

Speyeria callippe (Boisduval)

+ + + -t + + + + +

-t- + + + - - - - +

+ + + -h + - + +

+ + + ___■

/. Res. Lepid.

Months

F M A M J

J A S O N D

Chlosyne gabbli (Behr)
Euphydryas chalcedona (DoiibL)
Limenitis lorquini (Boisduval)
Adelpha bredowii (Geyer)
Coenonympha tullia Westwood
Danaus plexippus (L.)

Danaus glUppus (Cramer) ’

Limacodiae

Monoleuca ocddentalis B. & McD. -
Crambidae

Scop aria palloralis Dyar - +

Eudonia rectilinea (ZeUer)

Eudonia spenceri Muiiroe
Usingenessa briinnidalis (Dyar)

Petrophila jaliscalis (Schaiis)

Microtheoris ophionalis (Walker)
Nannobofys conimortalis (Grote)
Mim.oschinia rufofascialis (Steph.) -
Hellula rogatalis (Hiilst)

Slegea powelli Mu n roe
Abegesta reluctalis (Hiilst)

Lipocosma albibasalis^. ScMciy.
Dicymolomia metalliferalis (Pack.) -
Achyra occidentalis (Packard)

Pyrausta riapaealis (Hulst)

Pyrausta nr. roseivestalis Mun.

Pyrausta pikitealis Sc
Pyrausta volupialis (Grote)

Pyrausta morenalis (Dyar)

Pyrausta coccinea Warren
Pyrausta laticlavia (Gr. & Rob.) - +

Pyrausta fodinalis (Lederer)
lldea profundalis (Fuck^rd)

Udea octosigrialis (Hulst)

Lamproserna sinaloanensis Dyar
Lineodes integra (Zeller)

Choristostigma elegantalis Warren - +

Mecyna mustelinalis (Packard)

Mimorista subcostalis Hamp.

Noniophila neartica Munroe + -i-

Spoladea recurvalis (Fabricius)

Lygropia octonalis (Zeller)

Diastictis Jracturalis (Zeller) +

Crambus occidentalis Grote
Crambus rickseckerellus Klots
Crambus cypridalis Hulst
Agriphila undata (Grote)

-f

-h

_1_

_J_

-h

36:45-78, 1997 (2000)

Months

J FMAMJ J AS

O N D

Agrtphila attenuata (Grote)
Agriphila angulata (B. & McD.)
Mkrocrambus sp. 1
Mkrocramhus sp. 2
Parapediasia teterrella Zincken
Euchromius califomkalis (Pack.)
Hemiplatytes epia (Dyar)

+ -f + + +

+
+

+ + +
+ + + + +

+ + + +

+ +

-h

Pyralidae

Pyralis electalis Hiilst
Pyralis cacamka Dyar
Herculia phoezalis Dyar
Acallis gripalis (Hulst)

Arta epicoenalis Ragonot
Jocara tra balls (Grote)

Tallula fieldi Barnes & McD.
Galleria mellonella (Linnaeus)
Achroia grisella (Fabricius)
Macrotheca angalalis B. & McD,
Macrotheca ponda (Dyar)
Rhodophaea caliginella (Hiilst)
Myelopsis alatella (Hiilst)

Ambesa rmlsinghami (Ragonot)
Nephopteryx bifasciella Hiilst
Sarata pullatella (Ragonot)
Sarata dophnerella Ragonot
Lipographis fenestrella ( Packard )
Adelphia ochripunctella (Dyar)
Elasmopalpus lignoselhis (Zeller)
Eumysia fuscatella (Hulst )

Honora dotella Dyar
Homeosom.a electellurn (Hulst)
Homeosoma uncanale Hulst
Phycitodes mucidella (Ragonot)
Laetilia coccidivora (Comstock)
Laetilia zaynacrella Dyar
Rhagea stigmella (Dyar)

Olycella subumbrella (Dyar)
Eremberga craebates (Dyar)
Ozamia fuscomaculella (Wright)
Ephestiodes gilvescentella Ragonot
Ephestiodes erythrella Ragonot
Ephestiodes griseus Nuenzig
Manhatta setonella (McD.)
Sosipatra rileyella (Ragonot)
Anagasta kuehniella (Zeller)
Cadra cautella (Walker)

Arivaca albidella (Hulst)

/. Res. Lepid.

Months

Pterophoridae

Ptnophorus rrhigoris (Wals.)

Platyptila carduidacyla (Riley)

Anstenoptila marmarodactyla (Dy.)

Stenoptilia zophodactyla (Dup.)

EmmeliJia monodactyla (L.)

ca. 3 undetermined Pterophoridae

Geometridae

Profit arne siibalbaria (Packard)

I tame qiiadrilinearia (Packard)

Itame semivolata (Dyar)

Itame extemporata B. & McD.

Itayne giienearia (Packard)

Elpiste marcescaria (Guenee)

Elpiste metanemaria (Hulst)

Semiothisa pictipennata (Hidst)

Semiothisa californiaria (Hulst)

Semiothisa iieptaria (Guenee)

Semiothisa s-signata (Packard)

Hesperumia sulphurararia Packard

Neoalcis californiaria (Packard)

Glaucina epiphysaria Dyar

Glaucina magnifica Grossbeck

Hulstina exhumata (Swett)

Hulst ina wrightiaria (Hulst)

Pterotaea lamiaria (Strecker)

Anacamptodes fragilaria (Gross.)

Gochisea sinuaria B. & McD.

-h

Phigalia plumogeraria (Hulst)

Paleacrita longiciliata Hulst

Lomographa elsinora (Plidst)

Sericosema juturnaria ( Guenee )

Eudrepanulatrix rectifascia (Hidst)

Drepanulatrix unicalcaria (Guen.) +

Drepanulatrix hulstii (Dyar)

Drepanulatrix bifilata (Hulst)

Drepanulatrix (piadraria (Grote)

Drepanulatrix foeminaria (Guen.)

Drepanulatrix carnearia (Hulst)

Drepanulatrix falcataria (Pack.)

Drepanulatrix monicaria (Guen.)

Pero radiosaria (Hulst)

Pero mcdunnoughi (Cass. & Swett)

Aethaloidia packardaria (Hulst)

Parexcelsa ultraria Pearsall

Slossoriia rubrotincta Hulst

Sicya laetula Barnes & McD.

Plataea personaria (Edwards)

36:45-78, 1997 (2000)

Months

Eusarca falcata (Packard)
Somatolopha shnplicianaB. 8c McD
Phmie subpimctata (Hulst)

Synaxis hirsutaria Barnes & McD.
Synaxis mosesiani Sala
Prochoerodes truxaliata (Giienee)
Chlorosea banksaria Sperry
Nemoria pulcherrima (B. 8c McD.)
Nemoria pistaciaria (Packard)
Nemoria damnniata (Dyar)
Nemoria leptalea Ferguson
Nemoria glaucomarginaria

(Barnes 8c McDunnough)
Dichordia illustraria (Hulst)
Synchlora aerata (Fabricius)
Cheteoscelis faseolaria (Guenee)
Chlorochlamys triangularis Prout
Lobocleta ossularia (Geyer)
Lobodeta granitaria (Packard)
Lobodeta plemyraria (Packard)
Idaea bonifata (Hulst)

Idaea eremiata (Hulst)

Cydophora dataria (Hidst)
Cydophora nanaria (Walker)
Dysstroma mancipata (Guenee)
Plydriomena albifasdata (Pack.)
Hydriomena nubilofasdata (Pack.)
Triphosa californiata (Packard)
Ardiirhoe neomexicana (Hulst)
Perizoma custodiata (Guenee)
Antidea switzeraria (Wright)
Stamnodes albiapicata Grossbeck
Stamnodes reckseckeri Pearsall
Stamnodes affdiata Pearsall
Stamnodes annellata (Hulst)
Stamnodes coenonymphata (Hulst)
Stamnodes cassinoi Swett
Epirrhoe plebeculata ( Cjuen ee )
Zenophleps lignicolorata (Packard)
Orthonama centrostrigaria (Woll.)
Venusia duodecelimeata (Packard)
Eupitheda maestoma (Hulst)
Eupitheda misturata (Hulst)
Eupitheda rotundopuncta Packard
Eupitheda zelmira Swett 8c Cass.
Eupitheda acutipennis (Hulst)
Eupitheda shirleyata Cass. 8c Sw.
Eupitheda neoadata Packard
Nasusina inferior (Hulst)

/. Res. Lepid.

Months

JFMAMJJASOND

Nasusina vaporata (Pearsall)
Ttichopteryx veritata Pearsall

Lasiocampidae

Tolype gleriwoodi Barnes
Phyllodesyna amerkana (Harris)
Gloveria medusa (Strecker)
Malacosoma constrictum (Edw.)

+ +

+ + + +

+ + + + - - 4

+ +

+ + - -

+ +

Saturniidae

Hemileuca electra (Wright)

Saturnia walterorum Wog. 8c ]o\\w. - - +

Antheraea polyphemus {Crmwer) -
Hyalophora euryahis (Boisduval) + + + +

Sphingidae

Manduca sexta (Linnaeus)
Sphinx perelegans Edwards
Smerinthus cerisyi Kirby
Pachysphinx occidentalis (Edw.)
Erinnyis ello (Linnaeus)
Heynaris diffmis (Boisduval)
Hyles lineata (Fabricius)

Notodontidae

Clostera apicaiis (Walker)
Datana perspicua Gr. & Rob.
Furcula cinerea (Walker)
Furcula scolopendrina (Bois.)

Dioptidae

Phryganidia califomica Packard

+ + + - + +

+ + + + + +

+ + - + + +

+ + - + + 4

4 4

Arctiidae

Crambidia dusca B. & McD.
Cisthene liberomacula (Dyar)
Cisthene deserta (Felder)

Cisthene dorsimacula (Dyar)
Cisthene perrosea (Dyar)

Cisthene faustinula (Boisduval)
Fycornorpha grotei (Packard)
Estigmene acre a (Drury)

Spilosoma vestalis Packard
Arachnis picta Packard
Apantesis hewletti B. & McD
Apamiesis nevadensis (Gr. & Rob.)
Apantesis proxim.a (Guer.-Mene.)
Hemihyalia edwardsii (Packard)
Ctenucha brunnea Stretch

4 4

4 4-4
4 4 4 4
4-44
4 4 4 4
4 4 4 -

4 4

36:45-78, 1997 (2000)

Months

Lyman triidae

Orgyia vetusta (Boisdnval)

Noctuidae

Tetanolita palligera (Smith)

-t

-h

Myctei'ophora geometriforrnis Hill

-t

Hem.eroplanis finitima ( Smith )

-h

-H

Hemeroplanis incusalis (Grote)

Melipotis indomita (Walker)

Melipotis jucunda Hhbner

Bulia deducta (Morrison)

-h

Synedoida ochracea (Behr)

-h

-t-

Synedoida edwardsi (Behr)

-H

-t-

-h

Synedoida fumosa (Strecker)

Synedoida tejonica (Behr)

-h

Ascalapha odorata (Linnaeus)

Zale insiida (Smith)

-h

Zale termina (Grote)

Caenurgia togataria (Walker)

Catocala ilia (Cramer)

Catocala cleopatra Strecker

Catocala verrilliana Grote

-h

Trichoplusia ni (Hiibner)

-h

Pseudeva palligera (Grote)

Autographa biloba (Stephens)

Autograplia calif ornica (Speyer)

-h

Meganola fuscula (Grote)

-h

Nola apera Druce

-h

-H

-1-

-t-

-I-

Tripudia balteata Smith

Cobubatha dividua (Grote)

Copibryophila angelica Smith

-1-

Eumicremma mirmna (Gnenee)

Tarachidia cajidefacta (Hhbner)

-h

Conochares alter (Smith)

Conochares arizonae (Edwards)

Acontia cretata (Gr, & Rob.)

-h

Acronicta othello Smith

-h

Cryphia nanoides Franclemont

-h

Cryphia viridata (Har\'ey)

Cryphia albipuncta (B, Sc McD.)

-t-

Apamea albina (Grote)

Apamea chief acta (Grote)

-1-

Oligia marina (Grote)

Oligia tusa (Grote)

Oligia violacea (Grote)

-h

Cobalos angelicus Smith

Xylomoia sp.

Benjaminiola colorada (Smith)

-t

Mammifrontia riley Benjamin

-f +

/. f^s. Lepid.

Months

JFMAMJJASOND

Archanara oblonga (Grote) + + + +

Helotropha reniformis (Grote) -

Aseptis perfurnosa {Hmvipson) -

Aseptlsu.s^.

Aseptis paviae {Siveckev) + + + + + -

Aseptis pausis (Smith)

Asepth genetrix {Gvoi^)

Aseptis susquesa (Smith)

Chyton ix dwesta (Gvole)

Properigea albiniacu la {B. Sc McD.) - - + + + +

Properigea suffusa (B. & McD.) -

Pseud obey omima Jallax (Hamp.) + + + -- -- -- + + +

Magasa orbifera (Walker)

Protoperigea posticata -

Alicranthetis triplex (Walker) + + + + + + - + + - + +

Platyperigea extirna (Walker)

Platyperigea mona (B, & McD.) -

Spodoptera exigiia (Hiibner) + + - + + + + + + + -

Spodotera frugipera (Smith)

Spodotera ormthogalli (Cmenee) -

Galgula partita Giienee

Draudtia leucorena (Smith) + - + + _ +

Draudtia funeralis (Hill) + + + + + + -- + - + +

Polenta tepperi (Morrison)

Lineostristiria ollvaUs (B. & McD.) -

Nocloa rivulosa Smith
Cosmia calami (Har\'ey)

Homoglaea calif ornica (Smith) + +

Homoglaea carbonaria (Harvey) + +

Pseudoglaea olivata (Harv'ey) + + -

Agrocliola purpurea (Grote) +

Feralia februalis Grote +

Plnomella opter Dyar _ + + +

Plei'omellodia cinerea (Smith) + +

Catabena lineolata Walker

Homoncocnemis fortis (Grote) - +

Oncocnemis nita Smith -

Oncocnemis ragani Barnes - - - + + + + + + -

Lepi polys persdpta Cmenee + + + -1--1-.

Lepipolys behrensi (Grote)

Behrensia conchiliformis Grote + + +

Cucullia serraticornis (Lintner) + + -

Cucullia excissica Dyar -

Cucullia eulepis (Grote)

Discestra chart.aria (Grote) -

Tripedia nova (Smith)

Admetovis similaris Barnes
Polia nipana (Smith)

Lac/nipolia cuneala (Grote)

36:45-78, 1997 (2000)

Months

JFMAMJJASOND

Lacinipolia n. sp.

Lacinipolia stricta (Walker) -

Lacmipolia strigicolUs (Wallen.) - -- ----f-f----

Lacmipolia quadrilineata (Grot.) - - +

Lacinipolia patalis (Grote)

Dargida procincta {Groie)

Pseudaletia unipuncta (Haw.) - + + -i--i---!- + --i--h +

Leucania farcta (Grote)

Leucania februalis (Hill) _ + +

Leucania oaxacana Schaus - +

Perigonka tertia Dyar + + + +

Stretchia inferior Smith - + +

Orthosia erythrolita (Grote) + - - - - - + +

OrtJiosia transparens (Grote) + + + + + -

Orthosia praeses {Grote) - _ + -j-

Orthosia mys (Dyar) +

Orthosia ferrigera (Smith) - + + _

Orthosia terminata (Smith) + + + +

Orthosia behrensiana (Grote) _ +

Orthosia arthrolita (Harvey) - +

Orthosia pacifica (Harvey) _ +

Orthosia hibisci (Guenee) + + +

Egira hiemalis (Grote) +

Egira crucialis (Harvey) - + +

Egria cognata (Smith) _ + +

Egira curialis (Grote) + + +

Egira rubrica (Harvey)

Anhimella contrahens (Walker) -

HommiJwdes communis (Dyar) . + +

Protorthodes rufula (Grote) +

Protorthodes alfkerii {Civote)

Protorthodes variablis {'&, Sc MED.)

Protorthodes melanopis (Hamp.) - + + .

Ulolonche disticha (Morrison)

Zoster opoda hirtipes Grote + + - --

Miodera stigmata Smith + + + +

New gen., new sp. + + +

Tricholita fistula Harvey .

Agrotis vetusta Walker .

Agrotis venerabilisWMi^er -

Agrotis ipsilon (Hufnagel) -i- + + + --}- + -t + -h + -h

Agrotis subterranea (Fabriciiis) - +

Euxoa atomaris (Smith) .

Euxoa auxiliaris (Grote) -

Euxoa septentrionalis (Walker) -

Euxoa Olivia (Morrison) -

Euxoa tocoyae (Smith)

Euxoa simulana McDimnough -

Euxoa brunneigera (Grote)

J. Res. Lepid.

Months

J F M A M J J

S O N D

Euxoa selenis (Smith) + +

Euxoa heririetta {Smith)

Euxoa perexcellens (Grote) +

Euxoa difformis (Siiiith)

PseudortJiosia variabilis Grote) + +

Hemieuxoa rudens (Harvey) - + . + + . +

Peridroma saucia (Hiibner) + + + + - + - - - - +

Anom,ogyna infimatis (Grote) - + +

Adelphagrotis indeterminata (W.) -.- + -j- +

Abagrotis kirkwoodi Buckett +

Abagrotis denticulata McD.

Par abagrotis form alls (Grote) - - + + -!- + + + + ” +

Heliothodes diminutivus (Grote) =-- + + + » + -.- =

Helicoverpa zea (Bodie)

Heliothis phloxiphagus G. &R.

Sfhinia pulchripennis (Grote) - - + +

Schinia buta Smith
Schinia oleagina Morrison
Schinia oculata Smith

^ species observed only, not collected

INSTRUCTIONS TO AUTHORS

Manuscript format; Two copies must be submitted, double-spaced, typed, with wide
margins. Number all pages consecutively. Italicize rather than underline scientific names
and emphasized words. Footnotes are discouraged. Do not hyphenate words at the right
margin. All measurements must be metric. Time must be cited on a 24-hour basis,
standard time. Abbreviations must follow common usage. Dates should be cited as; day-
Arabic numeral; month-Roman numeral; year- Arabic numeral (e.g. 6. IV. 1992). Numerals
must be used for ten and greater e.g. nine butterflies, 12 moths.

Electronic submission: The Journal is now being produced via desktop publishing,
allowing much shorter publication times. Although typewritten manuscripts may be
considered, those submitted on computer disk are highly preferred. After being notified of
your paper's acceptance, submit either a Macintosh or IBM disk (3.5 inch) version. Include
on your disk both the fully formatted copy from your word processing program and a text-
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Please note in your cover letter any special characters that are used in either the body of
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Title material: All papers must include the title, author's name, author's address, and any
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include the full scientific name with unabbreviated author and year of description. There
must be conformity to the current International Code of Zoological Nomenclature. We
strongly urge depositing of types in major museums, all type depositories must be cited.

References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbreviations should not be used; write out the entire
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please cite in body of text not in Literature Cited. For multiple citations by the same
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Illustrations: Color can be submitted as either a transparency or print, the quality of
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Include a metric scale. Each figure should be cited and explained as such. Each illustration
must be identified by author and title on the back. Indicate whether you want the
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If figure is prepared on a computer, send a copy in electronic format upon acceptance (see
Electronic Submission, above).

Legends should be separately typed on pages entitled “Explanation of Figures.” Number
legends consecutively with separate paragraph for each page of illustration.

Review: All papers will be read by the editor! s) & submitted for formal review to two
referees.

The Journal of Research

ON THE LEPIDOPTERA

Volume 36 i 1997 (2000)

IN THIS ISSUE

Date of Publication: March 15,2000

A Study of the Riodinid Butterflies of the Genus Dodona in Nepal

(Riodinidae) 1

Curtis John Callaghan

On the correct placement of Erebia epipsodea Butler, 1868 within the

genus Dalman, 1816 (Lepidoptera: Satyridae) 16

Alexei G. Belik

Pontia occidentalis (Pieridae) Near Sea Level in California: a Recurrent

Enigma 24

Arthur M. Shapiro

Effects of microclimate and oviposition timing on prediapause larval
survival of the Bay checkerspot butterfly, Euphydryas editha bayensis
(Lepidoptera: Nymphalidae) 31

Erica Eleishinan, Alan E. Launer, Stuart B. Weiss,]. Michael Reed,

Carol L. Boggs, Dennis D. Murphy, and Paul R. Ehrlich

The Lepidoptera of Marine Corps Air Station Miramar: Calculating
Eaunal Similarity among Sampling Sites and Estimating Total
Species Richness 45

John W. Broxun and Katherine Bash

Cover: Photograph of final instar larva of the Saturniid moth Rothschildia
crycmrt collected on an unidentified Rubiaceae near Selva Verde lodge, Costa
Rica. © Mike Collins, 1990.

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