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THE JOURNAL MAR 2 8 1989
OF RESEARCH

ON THE LEPIDOPTE^:^vafJD *

THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA

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Journal of Research on the Lepidoptera

26(1-4):1-12, 1988

Male Mate-Locating Behavior in the Desert Hackberry
Butterfly, Asterocampa leilia (Nymphalidae)

Ronald L. Rutowski
and

George W. Gilchrist*

Department of Zoology, Arizona State University, Tempe, AZ 85287-1501

Abstract. The mating system of the desert hackberry butterfly, Astero¬
campa leilia , is described with special reference to the site tenacious
mate-locating behavior of the males. Males occupy perches on or next to
the larval foodplant, desert hackberry ( Celtis pallida ). Other males are
not tolerated within several meters of a male’s perch site and are
chased away when they fly nearby. Males occupy perch sites in the
morning. Some hackberry trees are more likely to be used as perch sites
than others and males at these sites experience the highest rate of
contacts with females and other males. Females passing a perch site
are chased, courted, and, if receptive, mated. The data indicate that
males defend perch sites as a means of maximizing potential contacts
with newly-emerged, virgin females leaving the plant adjacent to their
perch site.

Introduction

Male butterflies show a wide interspecific diversity in the extent to
which they are site tenacious in their mate-locating behavior (Scott,
1974, 1975, 1982). At one extreme males are not tied to any given site
but fly widely through the environment searching for females. This
strategy has classically been referred to as patrolling. At the other
extreme males are very site tenacious and an individual may defend a
space on a hilltop or other place for several days usually during some
restricted daily activity period (Powell, 1968; Baker, 1972; Douwes,
1975; Suzuki, 1976; Davies, 1978; Bitzer & Shaw, 1979, 1983; Cal¬
laghan, 1982; Lederhouse, 1982; Alcock, 1983, 1985; Wickman &
Wiklund, 1983; Knapton, 1985; Wickman, 1985; Alcock & O’Neill,
1986). In territorial species males interact with other males in ways
that are very different from their interactions with females. Aerial
combat occurs in the form of ascending flights and wing contact.
Between these extremes of patrolling and highly territorial species
there are species in which males, although showing some site tenacity,

*Current address: Department of Zoology, University of Washington, Seattle, Washington
98195.

2 J.Res.Lepid.

may only briefly occupy a site and be less aggressive toward conspecifics.
Detailed studies of these behaviors are lacking.

Males of the desert hackberry butterfly (. Asterocampa leilia Edwards)
occupy perch sites near the larval foodplant, desert hackberry ( Celtis
pallida Torrey), that they appear to defend against conspecific males
(Austin, 1977). Our preliminary observations of this species in central
Arizona suggested that individual males do not spend much time on
specific sites. Here we describe in detail the perching behavior of the
males of this species and document the extent to which males are site
tenacious and their perch site preferences. The discussion focuses on the
ecological circumstances favoring this sort of mate-locating behavior,
especially in comparison with mate-locating techniques in other species.

METHODS

Asterocampa leilia males and females were observed and collected at two flat
or gently sloping sites near water courses in the upper Sonoran desert habitat
typical of central Arizona. Our primary site was near the Salt River about 40 km
northeast of Tempe, Arizona, and the other site, used primarily for observations
of courtship with hand-reared females, was along Sycamore Creek about 70 km
from Tempe. At both sites the large vegetation included palo verde ( Cercidium
spp.), mesquite (. Prosopsis spp.), saguaro cactus ( Carnegia giganteus (Engel-
mann) Britton and Rose) and desert hackberry ( Celtis pallida ).

At the Salt River site we identified a triangular, 1700 m2 area that extended
on the north to the beginning of a small mountain range, on the south to a line of
paloverde trees, on the west to a low ridge extending out from the mountains,
and on the east to a small dry wash. This area contained fifteen discrete clumps
of hackberry that varied in size. Males were captured and carefully marked by
writing numbers on the dorsal and ventral hindwings with a felt-tipped
marking pen (Sanford SharpieR).

On 17 mornings from 6 May to 11 June in 1985 we walked through the study
area at 30 min intervals and noted the location and identity, if marked, of each
male seen perched. We also observed the activities of males at individual perch
sites, especially those that were most often occupied, and recorded the interac¬
tions between the perch site occupant and intruding individuals of both sexes.
Wherever possible these observations were made on marked males that occupied
perch sites.

Courtship, Copulation, and Spermatophore Counts

Successful courtships and the ensuing copulations were elicited by releasing
hand-reared virgin females near perched males in the field. The females were
reared from eggs collected by placing field-caught females in cages with a sprig
of the larval foodplant.

After copulation, mated pairs were killed and stored by freezing. Later the
females were thawed, weighed, and dissected under insect Ringer’s solution.
The bursa copulatrix of each female with its contents was examined and
weighed to the nearest 0.01 mg. The male of each pair was also thawed and
weighed.

26(l-4):l-288, 1988

3

To estimate the frequency with which females mate, we collected and froze a
sample of females from the population. They were later thawed and dissected
under insect Ringer’s solution, and the contents of the bursa copulatrix of each
was examined. The wing wear of these females was assessed as an indicator of
age. Each female was placed in one of three wing wear categories: fresh — little
or no scale loss or tattering, worn — substantial scale loss or tattering on one or
two wings, and very worn — substantial scale loss or tattering on all wings.

Statistical Summary and Tests

Parametric summary statistics are given as the mean ± the standard
deviation. The results of all statistical tests were evaluated at the 0.05 level of
significance.

RESULTS

Spatial Organization and Daily Pattern of Male Activity

Male perch sites were found in only 20 locations within the Salt River
study area. A perch site was an area of approximately 1-2 m2 in all cases
except one on or immediately adjacent to a hackberry tree, confirming
Austin’s (1977) result. Some sites were occupied more frequently than
others. In 97 censuses made over 17 days the average occupation
frequency among the 20 perch sites was 21.9 ± 20.6%; however, the two
most frequently occupied sites were occupied in 83.5% and 61.9% of
the censuses, respectively. Only one of the hackberry clumps in the
study area never had a male perched next to it.

The behavior of the males followed a daily pattern. Males occupied
perch sites when they first became active in the morning. Fig. 1 shows
the number of males seen perching as a function of the time before and
after the observed time of peak activity. Peak activity was defined as the
time at which the number of sites occupied reached the maximum
number observed on a given day. The average time of peak activity was
900 MST (range: 800-1000). Late in the morning the males moved into
the shade of hackberry trees and became inactive.

Site Tenacity of Males

One-hundred and two males were captured, marked, and released. Of
these 34% were resighted at some point after release (Fig. 2). After
being marked and released, a male typically left the area and was not
seen again on that day. The probability that a marked male would be
resighted was highest the day after marking (18%). The longest time
between release and the last resighting of a male was 10 days.

A male did not occupy a given perch site for long. Fig. 3 shows the
distribution of site occupation durations observed on single days during
the study. Most males were on a site for only 30 min or less on any given
day. However, one male was observed on the most frequently occupied

4

J.Res.Lepid.

Fig. 1 . The daily pattern of perching activity for A. leilia males averaged over 17
days. The number above each indicates the sample size.

□

LlJ

Fig. 2. The probability of resighting previously marked males as a function of the
days since they were marked. The number above each bar is the number of
males in the population marked on day zero that were available on a given
day for recapture.

Fig. 3. For all sites that were seen occupied by a marked male this figure shows the
duration of site occupation as the number of consecutive census periods on
the same day that the male was seen on a given site (n = 67).

26(l-4):l-288, 1988

5

site on the study area for 5 census periods one day and then two more on
the next. It is possible our censusing activities scared males prema¬
turely from their perch sites; some males did take to wing when we
approached but they returned to their perch after a brief flight. In any
event we took some pains not to disturb males during the censuses.

Site Defense

Males perched on sand, rocks, and low vegetation (especially the
hackberry tree) (Fig. 4). From these positions a male flew out and chased
passing conspecifics, heterospecific butterflies, other insects (flies,
wasps, etc.), birds, and even thrown stones. Conspecific males flying
near a male’s perch were typically approached and chased on the wing
for several meters. In this species, no male-male interactions led to
ascending flights. Occasionally a male perched near the resident with¬
out being detected. Such intruders were not detected until they flew, at
which point they were approached and chased from the area. In 73% of
37 male-male interactions involving at least one marked male, the male
that was originally perched in the area returned alone and reoccupied
the site. This is significantly more frequently than expected from chance
(X2 = 7.81, 1 df, p < 0.05).

On occasion a male spontaneously flew up from his perch and
patrolled an area by flying back and forth in front of the hackberry tree
for a few seconds before perching again. When the resident alit after
such a patrol flight or after an interaction he typically perched on or
within a meter of his original perch.

The behavior of the males varied with the site and with the time of the
morning. The more attractive sites were more likely to be occupied for
more than one period by a single male (Table 1; X2 = 11.5, 1 dr, p < 0.05).
Males that occupied the two most popular sites tended to stay on them
throughout the activity period in spite of frequent intrusions by other
males. Eighteen perched intruders were observed at these sites in 528
min of observations during the hour surrounding the time of peak

Table 1. Howr site identity affects the number of consecutive census
periods that the site will be occupied by the same male. Sites
IN and 15 were the most frequently occupied sites.

No. of periods

Sites

All other

occupied

IN and 15

sites

1

39%

82%

2-3

39%

18%

4-5

22%

0%

Total observations

18

49

Fig. 4. Males of A. leilia perched on their sites. A male perched on staghorn cholla
(above) and on the ground (below).

26(l-4):l-288, 1988

7

activity, while no perched intruders were observed in 107 min of
observations on other sites at the same time.

Males do spontaneously abandon sites. Some sites were less likely to
be abandoned for no apparent reason than less attractive sites. For
example, during a total of 803 min of observation at sites 15 and IN only
one abandonment was observed. In contrast 6 abandonments were
observed in 111 minutes of observations at three other sites (IS, 3S, and
3N) during the same time period. There were significantly fewer
abandonments at site 15 and IN than expected from the time spent
observing there x2 = 35.5, 1 df, p < 0.05).

Competition for sites is intense as indicated by the fact that when we
sequentially removed 10 males during one hour from the most frequently
occupied site, the site was reoccupied within a few minutes by a new
male after each removal. Furthermore, the intensity of the competition
changed over the morning. This was evident in two ways. First, if a male
observed on a site in one census was not there when the site was
censused 30 min later, at the time of peak activity there was a greater
than 50% chance that the site would be occupied by a new male (Fig. 5).
Late in the morning sites were rarely reoccupied if for some reason the
male left. Second, the frequency of perched intruders waned as the
morning progressed (Fig. 6; Spearman rank correlation coefficient =
-0.87, p < 0.05).

Courtship and Copulation

The rate of appearance of wild females varied among sites. Females
appeared at a rate of 0.0177 per min (790 min of observation) and 0.0123
per min at (163 min of observation) at the first and second most
frequently occupied sites, respectively. In contrast, during a total of 203
min of observation no females were seen at several other sites when a
male was present.

A total of 6 successful and 8 unsuccessful courtships were observed
during this study. All successful courtships involved hand-reared virgin
females released near males. On three occasions during observations of
males on sites the male chased a female and did not return; the pair flew
off so quickly we were unable to determine the outcome of the interaction.

When a female flew by a perched male he immediately took wing and
followed the female. In successful courtships the female immediately
perched in vegetation near the male’s perch site. The male then landed
behind the female, moved up beside her, and began attempting to insert
his abdomen between the female’s hind wings. The female then either
remained still and permitted the male to couple or moved away from the
male for some time before becoming still and permitting copulation. In
unsuccessful courtship the female did not perch when the male appro¬
ached and in 5 cases engaged in ascending flights with the male in
pursuit. The male abandoned the female and returned to his perch after
an ascending flight interaction.

8

J . Res. Lepid.

-1 0 1 2 3 4 5 6

30 MIN PERIODS SINCE PEAK ACTIVITY

Fig. 5. The likelihood that a previously occupied site will be occupied by a new
male in the census period after the site was seen to be occupied is plotted
against the time of the post-occupation census. The sample size for each
time period is shown above each bar.

Fig. 6. The rate of appearance of undetected perched intruders is shown as a
function of the time since peak activity. The number of minutes of
observation from which the data point was calculated is shown above each
point.

Copulation averaged 49 ± 16.8 min (n = 6, range: 21 - 65 min).
During copulation a male formed a spermatophore and deposited some
loose white secretions within the female’s bursa copulatrix. The mass of
material averaged 2.49 ± 0.78 mg (n = 7, range: 1.47 - 3.73 mg) which
correspond to 2.96% ± 0.59% (n = 7, range: 1.9 — 3.71%) of the male’s
estimated precopulatory body weight. The quantity of material passed
was significantly positively correlated with the estimate of a male’s
precopulatory body weight (r = 0.797, t = 2.96, 5 df, p < 0.05).

Twenty-six females were collected in the field and the contents of the
bursa copulatrix of each was examined. In this sample 73% were fresh,
19% worn, and 7% very worn. No female carried more than one
spermatophore and three (all fresh) had bursae that were empty.

26(l-4):l-288, 1988

9

DISCUSSION

Characteristics of Male Perching Behavior

The data reveal several features of the perching behavior of A. leilia.
First, as Austin (1977) suggested, males perch most often near C.
pallida trees. Second, a perched male does not tolerate other males in
their perching area but are not likely to defend any given perch site for
long. Third, males occupy and defend perch sites for only a restricted
part of the day. Males became inactive toward the middle of the day and
roosted well within C. pallida trees on or near their site. Such temporal
restrictions on site-tenacious strategies of mate-location are common in
butterflies (Callaghan, 1982; Alcock, 1983; Wickman, 1985) and are
probably best explained by heat stress due to high midday temperatures
(Rawlins, 1980; Kingsolver and Watt, 1983) which favors abandonment
of perches in the late morning or by variation during the day in the
availability of receptive females. Fourth, some sites are strongly pre¬
ferred over others as indicated by the frequency with which they were
occupied, the rate at which undetected intruders perched on them, and
the rate with which they were abandoned by males. Similar preferences
are found in other perching species (e.g. Bitzer & Shaw, 1979; Leder-
house, 1982; Alcock, 1983). Finally, the sites that were preferred by
males were also those visited most frequently by females although it is
not clear that these females were receptive. This has been shown for
three other territorial species of butterflies (Davies, 1978; Lederhouse,
1982; Wickman, 1985).

The Function of Male Perch Site Placement and Defense

We interpret site occupation and defense as a mate-locating tactic in
this butterfly. The sites contained no nectar or water resources that
might be of interest to males or females and so defense of resources for
personal use or to gain access to females seems improbable. Why then
are the sites on or next to the larval foodplant? There are at least two
possible hypotheses. A male may perch near the larval foodplant to gain
access to females that come to oviposit. Such behavior has been observed
in bees, dungflies, odonates, and many other insects (Thornhill &
Alcock, 1983). However, there are apparently no benefits from trying to
intercept mated females in that A. leilia females mate only once.

We conclude that males perch near the larval foodplant to maximize
their chances of contacting virgin females as they emerge on their first
flight. However, this assumes that the larvae pupate on the larval
foodplant. This is likely. We have found cast pupal skins on C. pallida ,
and the pupae bear a striking resemblance to the leaves of the host
plant, suggesting that the larvae routinely pupate on the larval food-
plant.

Why do males perch at some trees and not others? Females may prefer

10

J. Res. Lepid.

certain trees as oviposition sites and therefore these trees are more
likely to produce virgin females than others. We do not at this point
know if females are more likely to oviposit on the trees preferred by
males; we only know that females are more likely to appear there. It
may also be that some trees provide better vantage points for looking for
newly-emerged butterflies. We are currently testing this hypothesis by
setting up large visual barriers and seeing if they affect male perch site
selection.

Variation in Site Tenacity and Defense

Asterocampa leilia males fall somewhere in the middle of the spectrum
of site tenacity and defense shown by male butterflies. Site tenacity and
defense are typically closely tied. Some butterflies show essentially no
site tenacity, such as the alfalfa butterfly ( Colias eury theme Boisduval).
On the other hand, males of some species perch on and defend the same
site during the activity period for several days (Davies, 1978; Suzuki,
1978; Lederhouse, 1982; Alcock, 1983, 1985; Knapton, 1985; Alcock &
O’Neill, 1986). In A. leilia even the most attractive sites were occupied
and defended for only a few consecutive 30 min census periods and
rarely for more than one day. What ecological factors have favored this
sort of behavior?

Site occupation and defense in butterflies is associated with mate
location. Hence, the form of this behavior will depend on a complex
interaction between the spatial and temporal distribution of receptive
females and the density of competitors (Rutowski, 1984; Courtney &
Parker, 1985; Odendaal et al., 1985; Alcock & O’Neill, 1986). Currently
our understanding of this interaction awaits further detailed studies of
mate-location in species that perch.

Spermatophore Size and Mating System Structure

Male butterflies expend energy in reproduction in two ways. The
first is in mate location. The second is in making spermatophores which
contain not only sperm but also accessory gland secretions that may be
used by the female as nutrients for egg production (Rutowski, 1984). In
species that engage in site defense we expect that the amount of effort
put into site defense will be a major determinant of reproductive
success. Although males that patrol also expend energy in mate location
the cost of this can be ameliorrated by their ability to feed while
searching for females. Vigilance during site defense precludes feeding
and defended sites rarely contain nectar resources.

We predict, therefore, that in species in which males defend perch
sites, the males will expend more on mate location and less on sper¬
matophore production than in species in which the males patrol in
search of mates. As expected, males of A. leilia (this study) and males of
Pararge aegeria Linnaeus (Svard, 1985), another site defending species

26(l-4):l-288, 1988

11

(Davies, 1978; Wickman & Wiklund, 1983), produce spermatophores
that are small in relation to their body weight (2-3%) compared with
other species that have been examined. Rutowski et al. (1983) surveyed
10 species of butterflies in which males patrol in search of females and
found that typically 6 to 7 percent of the male body weight was donated
in each spermatophore. Further studies are needed to test the predic¬
tion that mating system structure is linked with the investment males
make in nutrient contributions to their mates.

Acknowledgements. We thank Barbara Terkanian and Rob Gardner for assi¬
stance in the field, Dr. John Alcock, Dr. Tim Friedlander, and an anonymous
reviewer for helpful criticism of an earlier draft of the manuscript and the
National Science Foundation for financial support through NSF Grant No. 85-
00317 to R. L. Rutowski.

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Journal of Research on the Lepidoptera

26(l-4):13-26, 1988

The Biology of Seven Troidine Swallowtail Butterflies
(Papilionidae) in Colima, Mexico

Paul Spade

5059 Thorne Drive, La Mesa, CA 92041
Hamilton Tyler1

8450 W. Dry Creek Rd., Healdsburg, CA 95448
John W. Brown

Department of Entomological Sciences, University of California, Berkeley, CA 94720

Abstract. Observations on the early stages and ecology of the follow¬
ing aristolochia-feeding swallowtail butterflies are presented: Battus
philenor philenor (Linnaeus), B. polydamas lucayus (Rothschild and
Jordan), B. eracon (Godman and Salvin), B. laodamas procas (Godman
and Salvin), Parides photinus (Doubleday), P. montezuma (Westwood),
andP. erithalion trichopus (Rothschild and Jordan). Observations were
made during 1979-1982 in the state of Colima, Mexico, by Paul Spade.
Resumen. Se presenta information sobre los estadios inmaduros y la
ecologla de los siguientes papilionidos que comen aristolocareas: Bat¬
tus philenor philenor (Linnaeus), B. polydamas lucayus (Rothschild and
Jordan), B. eracon (Godman and Salvin), B. laodamas procas (Godman
and Salvin), Parides photinus (Doubleday), P. montezuma (Westwood),
y P. erithalion trichopus (Rothschild and Jordan). Los datos fueron
obtenidos durante 1979-1982 en el estado de Colima, Mexico.

Introduction

Four species of Battus Scopoli are known from the state of Colima,
Mexico: B. philenor philenor (Linnaeus), B. polydamas lucayus (Roth¬
schild and Jordan), B. eracon (Godman and Salvin), and B. laodamas
procas (Godman and Salvin). In the same region three species of Parides
Hiibner have been documented: P. photinus (Doubleday), P. montezuma
(Westwood), and P. erithalion trichopus (Rothschild and Jordan). Both
genera are confined to the New World, predominantly South America.
Battus is the smaller, containing 14 species; Parides encompasses
between about 32 and 45 species, 9 of which occur in Mexico.

Battus and Parides are members of the tribe Troidini (Papilionidae),
commonly referred to as aristolochia swallowtails (Rothschild and

deceased; forward correspondence and reprint requests to John Brown.

14

J. Res. Lepid.

Jordan 1906; Munroe 1960; Ehrlich and Raven 1965) because their
larvae feed exclusively on members of the genus Aristolochia Linnaeus
(Aristolochiaceae). This plant genus, composed primarily of tropical
vines, can be divided into two natural groups in the Americas: pentan-
drous and hexandrous species (5 and 6 stamens respectively) (Pfeifer
1966, 1970). Table 1 outlines the major characteristics and distributions
of Aristolochia species utilized by Battus and Parides in Colima.

Aristolochia species contain a number of secondary plant compounds
including aristolochic acids, benzylisoquinoline alkaloids, and sesquiter-
penoids, which in general deter both insect and vertebrate phytophagy
(Fraenkel 1959). Some members of the Troidini are known to sequester
these substances from their foodplants, rendering the butterflies dis¬
tasteful to vertebrate predators. Consequently their colors are aposema-
tically adapted (Brown, Damman and Feeny 1981). Hence adults are
models in Batesian mimetic complexes involving butterflies of the same
family in the genus Papilio Linnaeus (Brower 1958) and Eury tides
Hiibner. Various Battus and Parides species form Mullerian mimicry
complexes as well (Young 1971a, 1971b, 1972; Brown, Damman and
Feeny 1981).

The purpose of this paper is to present brief descriptions and illustra¬
tions of the early stages, summarize data on host utilization and
geographic distribution, and identify the major mimetic complexes
among the seven troidine swallowtails that occur in Colima, Mexico.

Methods and Study Area

The field work, including photography, was carried out by Paul Spade
from November 1979 to November 1982. Specimens of both butterfly
and Aristolochia species were sent by Spade to Hamilton Tyler who
arranged for their determination. Voucher specimens of the butterflies
were deposited in the San Diego Natural History Museum; no larvae
were preserved. With the assistance of Michael Parsons, a preliminary
draft of the present manuscript was completed. After the untimely death
of Tyler, John Brown was responsible for the final organization and
presentation of the paper.

Field work was conducted at 23 sites in Colima and 3 sites in the
neighboring states of Michoacan and Jalisco (figure 1). Only the Colima
localities mentioned in the text are shown in figure 1. All records of
oviposition and hostplant selection are based on the discovery of eggs
and larvae, or the observation of ovipositing females, in the field. Once
located, the early stages were brought into the lab (Spade residence in
Colima) and reared on the field-associated host at ambient temperature.
All photographs were taken in the lab. No measurements of the early
stages were noted as they were nearly identical, and most have been
presented previously elsewhere (see literature citations in Observa¬
tions).

26(l-4):l-288, 1988

15

Fig. 1. Map of Colima, Mexico.

Colima is located along the western coast of mainland Mexico. Although
one of the smallest states in the Republic, it is topographically extremely
diverse; elevation ranges from sea level to nearly 3900 m at the peak of
the Fuego Volcano. The Pacific shoreline, approximately 160 km long, is
bordered by a narrow coastal plain. Low coastal ranges rise abruptly
from the plain and quickly attain an elevation of 700—1300 m. Colima,
the capital city, is located in the east-central part of the state on a
dissected plateau which varies in altitude from 300-600 m. The interior
mountain ranges rise to a nearly uniform elevation of about 2000 m.

According to Schaldach (1963), eight distinct vegetational zones occur
within Colima, six of which are relevant to this study. These can be
summarized as follows:

(1) Arid thorn scrub. A low, semi-open scrub community composed of
spiny deciduous shrubs and legumes occurring along the coastal plain
and the basal slopes of the coastal mountains.

(2) Arid thorn forest. A relatively tall (10—12 m), homogeneous
forest of flowering trees, mostly legumes, found on hillsides and in many
of the interior valleys. These deciduous forests are often dense with a
thick undergrowth of thorny vines.

(3) Riparian gallery forest. Heavily shaded forests of tall, mainly
evergreen trees, bordering all the permanant watercourses of the lower
tropical area.

(4) Tropical deciduous forest. Tall, climax deciduous forest com¬
posed of many species of tropical trees; generally found above arid thorn

16 J.Res.Lepid.

forest at higher, moister habitats, but may occur in lower elevations in
response to local precipitation.

(5) Oak woodland. Nearly pure stands of low oaks found on the
higher ridges of the mountains. Scattered pines occur amid the oaks, but
nowhere do they form significant forests.

(6) Arid pine-oak forest. Open, dry forests of tall pines interspersed
with medium-sized oaks, occurring primarily around the volcanoes from
about 1800—2400 m. This is the highest habitat in which troidines occur
in Colima.

Observations

Battus philenor philenor (Linnaeus)

Early stages. The early stages of B. philenor in Colima closely
resemble those from elsewhere. First described by Edwards (1881), they
are summarized in numerous accounts (e.g., Klots 1951; Emmel 1975;
Opler and Krizek 1984; etc.); they are not illustrated here. The spherical
egg is russet in color. First through third instar larvae are gregarious;
later instars feed singly. The ground color of fourth and fifth instar
larvae is usually dark purple with contrasting orange-red tubercles.
However, a second color form also occurs in which the ground color is the
same orange-red as that of the tubercles; the outer two-thirds of the
anterior tubercles and the tips of both the longer thoracic and first
abdominal tubercles, are tipped with black. This second color morph also
occurs in southeastern Arizona populations of B. philenor (A. Shapiro,
personal communication).

The pupae are dimorphic, as noted by West and Hazel (1979). One
clutch of B. philenor yielded 15 brown, 3 green, and one mixed (brown
and green) pupae. Thoracic projections of the pupae of B. philenor from
Colima are less prominent than those from elsewhere.

Ouiposition and foodplants. Eggs are laid in small batches of 4— 6 on
the undersides of the leaves of the foodplant, forming a well spaced group
(as opposed to a close cluster). At Madrid, Colima, the preferred
foodplant was Aristolochia acontophylla, on which larvae of Parides
montezuma also were found. Where A. acontophylla was unavailable,
the butterfly rarely utilized A. tentaculata. At higher altitudes, larvae
occurred on A. pringlei. Many other species of Aristolochia are used by
B. philenor in other parts of its wide geographic range; these are
summarized by Scriber and Feeny (1976). Rausher (1980, 1981) provides
data on host plant selection and temporal changes in oviposition
preference.

Habitat and range. In the Colima region, B . philenor occupies relative¬
ly dry habitats receiving an annual precipitation of less than 1200 mm
and characterized by a long dry season .B. philenor occurs from sea level
to about 2400 m, reaching its upper limit in the arid pine-oak forests. In

26(l-4):l-288, 1988

17

western Michoacan, B. philenor was present on the coast at Placita and
Aquila, but just inland, in the northern region near Zapotan, no adults
were observed during a six-month period from June to November 1982.
The species was common in regions of arid thorn scrub and arid pine-oak
forest, but it was conspicuously absent from the humid forest above
Colima (city).

Battus polydamas lucayus (Rothschild and Jordan) (figures 2, 8, and 14)

Early stages. The life history of this species has been recorded by Moss
(1919), Comstock and Vazquez (1960), Young (1971a), and Brown,
Damman, and Feeny (1981). Eggs (fig. 2) are pale brown bearing 10-12
orange-brown vertical ribs of a glue-like colleterial substance that
adheres them to the leaf of the foodplant.

The first instar larva is pale straw yellow; the short, hirsute tubercles
bear long black setae. The second instar is brown-black, sparsely
marked with tan-brown; the tubercles are light brown except those of
abdominal segments 3 and 7, which are orange. The third instar is
tan-gray with contrasting transverse, oblique, brown lines on each
segment. The tubercles have a light brown base and darken towards the
apex; the lateral tubercles are twice the length of the dorsal ones. The
fourth instar larva is gray with thin, black transverse lines. All of the
tubercles are cream-white with black tips, except the lateral tubercles,
which are black with a cream base. The fifth instar is dimorphic. The pale
form (fig. 8) is yellow-brown and resembles the fourth instar, while the
other form is dark brown with black transverse lines, and black-tipped
pink tubercles. The base of the lateral prothoracic tubercles is pale pink
in the dark form, creamy yellow in the light form.

The pupae are dimorphic: one form (fig. 14) is green with pale yellow
saddle and antennae, the other form brown. The pupa has a prominent,
conical, dorsal thoracic projection, and the abdomen bears two pro¬
minent dorsolateral ridges.

Oviposition and foodplants. Females typically lay eggs in small
clusters of 2— 9 in a band around the stem of the host. In Colima, larvae
were found on Aristolochia acontophylla,A. foetida,A. odoratissima,A.
tentaculata , and A. conversiae.

Habitat and range. B. polydamas is the most common troidine in
Colima. Throughout its range in the State it occupies primarily open
habitats such as agricultural land, gardens, and arid thorn scrub; but it
also occurs in riparian gallery forest. It is common from sea level to about
500 m, ranging as high as 900 m. An exceptionally high elevation record
for this species in Colima is 1360 m at Cofradia de Suchitan.

Battus eracon (Godman and Salvin) (figures 3, 9 and 15)

Early stages. There are no details of the life history of this species in
the literature. Eggs (fig. 3) are pink with irregular rib-like globules of

Figs. 2-7. Eggs of Battus polydamas (2); B. eracon (3); B. laodamas procas
(4); Parides photinus (5); P. montezuma (6); and P. erithalion
trichopus (7).

Figs. 8-13. Last instar larvae of Battus polydamas (8); B. eracon (9); B.

laodamas procas (10); Parides photinus (11); P. montezuma (12);
and P. erithalion trichopus (13).

Figs. 14-19. Pupae of Battus polydamas (14); B. eracon (15); B. laodamas
procas (16); Parides photinus (17); P. montezuma (18); and P.
erithalion trichopus (19).

26(l-4):l-288, 1988

19

orange glue-like colleterial secretion which binds each egg within the
batch and gives them a warty appearance.

Unfortunately, field collected eggs were eaten, presumably by ants, so
first and second instar larvae were not obtained. Subsequent field
collected larvae were all in later instars. Third instar larvae are
chocolate brown with only the tips of the tubercles orange; the lateral
prothoracic tubercles are concolorous with the body and much longer
than the rest. Of the four species of Battus in Colima, B. eracon has the
shortest tubercles. The fourth instar is indistinguishable in pattern
from the third, but its ground color is maroon-brown. Fifth instar larvae
are dimorphic: the dark form (fig. 9) is the most common and is almost
entirely purple-black; only the tubercles are orange-tipped. The pale
form is brown with oblique brown-black markings similar to those of
Battus polydamas , with the lateral prothoracic tubercles entirely black.
In both forms the head and thoracic prolegs are black. The first three
instars probably feed gregariously, later instars feed singly.

The pupae are dimorphic: one form is green and yellow, the other (fig.
15) brown with greatly reduced yellow markings. Pupae average 37 mm
long and 20 mm wide, with a dorsal thoracic projection of about 5 mm,
more club-shaped than that of B. laodamas procas.

Oviposition and foodplants. A single female was observed ovipositing.
Eggs were irregularly piled in batches of 20-30, near the ground, on the
sides of stems supporting the foodplant, Aristolochia tentaculata.

Habitat and range. B. eracon is the most restricted mainland Battus ,
limited to the coastal regions of Colima, Jalisco, Michoacan, and Guer¬
rero, Mexico (Hoffman 1976; Diaz Frances and de la Maza 1978). It is
common between sea level and 200 m, where its range includes the
coastal hills at Caleras. In Colima, B. eracon is locally abundant at Cerro
de la Vieja (near Coquimatlan) and Tepames; both localities are about
500 m in elevation. It has not been observed above 610 m. In Michoacan,
B. eracon occurs at Placita, Aquila, San Telmo, and Zapotan. Its
principal habitats are riparian gallery forest, arid thorn forest, and
tropical deciduous forest; adults are avidly attracted to nectar sources
within these habitats. It is scarce in urban and arid thorn scrub areas.

Battus laodamas procas (Godman and Salvin) (figures 4, 10, 16)

Early stages. Comstock and Vazquez (1960) described the final instar
and pupa of this species. Eggs (fig. 4) are pale yellow with six irregular
vertical, rib-like strands of glue-like colleterial substance of the same
color.

First instar larvae are light brown with black-tipped tubercles bearing
long setae; the head is shiny black. Second, third, and fourth instar
larvae are entirely black. Two distinct forms and intermediates are
exhibited by fifth instar larvae: one form is entirely black, the other (fig.
10) is light brown with oblique black markings similar to those of B.

20 J. Res. Lepid.

polydamas. In the brown form the tubercles are black-tipped. All instars
feed gregariously.

The pupae are dimorphic with a green and yellow form, and a
gray-brown form (fig. 16) that has greatly reduced yellow markings. The
abdominal tubercles are reduced, forming two dorsolateral ridges.

Ouiposition and foodplants. Eggs are laid in parallel rows in batches of
20—30 on the upper surface of leaves of Aristolochia tentaculata.
Oviposition is generally between 3— 4 m above the ground and always
above 2 m.

Habitat and range. B . laodamas procas has the same distribution as B .
eracon along the Pacific coast, but ranges further inland and into higher
altitudes. It is most common from sea level to about 500 m. However, in
the mountains it has been recorded as high as 1280 m.

Parides photinus (Doubleday) (figures 5, 11, and 17)

Early stages. The life history ofP. photinus has been described in detail
by Ross (1964). The following descriptions are based on observations in
Colima. The egg (fig. 5) is dark pink with about 15 irregular, warty,
vertical strands of yellow-orange colleterial secretion. Each egg rests on
a short pedestal of the same substance, to one side of the base.

First instar larvae are gray with the dorsal tubercles of abdominal
segments 4 and 7 pale yellow, and the thoracic and anal two pair of dorsal
tubercles pale orange. The remaining tubercles are burgundy. Second
instar larvae are darker; dorsal tubercles four and seven are creamy
white. The thoracic tubercles and those at the anal end of the abdomen
are pink. The third instar is dimorphic. The lighter form is similar to the
preceding instar, but the dorsal tubercles on abdominal segments 4 and 7
are ivory-white. An oblique white line joins tubercle four with the lateral
tubercle on abdominal segment 3 which is also ivory-white. The thoracic
tubercles and the two pair of dorsal tubercles on the last abdominal
segment are creamy orange, tipped with orange-red. The dark form is
similar but the lateral and dorsal tubercles on abdominal segments 4 and
7 are ivory-white with a white stripe joining those on segment 4. All of
the remaining tubercles and the ground color are dark red. First through
third instar larva bear black setae at the tips of the tubercles. The fourth
instar is also dimorphic, the two forms similar to those of the third instar
The fifth instar is dimorphic: one form (fig. 11) is black with white and
red tubercles which have an orange cast; the other is purple-black with
dark red tubercles.

The pupa (fig. 17) is pale yellow dorsally and pale green ventrally. It
has two short, triangular, dorsolateral processes on the thorax, and the
abdomen bears a row of three pairs of triangular-shaped dorsolateral
projections.

In Colima, the duration of the early stages is fairly rapid: egg, 6 days;
first and second instar, 3 days; third instar, 3 days; fourth instar, 2 days;

26(l-4):l-288, 1988

21

fifth instar 3 days, prepupa to eclosure, 18-20 days. In Battus and
Parides species studied by Brown, Damman, and Feeny (1981), larvae
developed in 20—30 days. They suggest that in addition to temperature,
developmental rates dependent greatly upon the species of Aristolochia
utilized.

Ouiposition and foodplants. In low to mid-elevations Aristolochia
tentaculata is the larval host. At slightly higher elevation A. conversiae
is utilized. In the vicinity of the Nevada de Colima Volcano, Jalisco, just
north of the Fuego Volcano, the host is A. pringlei. Eggs are laid singly
on the leaves.

Habitat and range. In Colima, P. photinus was observed in lower arid
thorn forests at about 400 m, in oak woodlands at about 1680 m, and in
arid pine-oak forests at 2400 m. It seems to favor forest margins and
areas of disturbed or secondary vegetation. P. photinus exhibits the
greatest altitudinal range of any of the Colima troidines except for B .
philenor.

Parides montezuma (Westwood) (figures 6, 12, and 18)

Early stages. There are no details of the life history of this species in
the literature. The egg (fig. 6) is pale gray-brown with about 12
irregular, warty, vertical rib-like strands of yellow colleterial substance.
The egg rests on a pedestal of colleterial substance which is shorter than
that of P. photinus.

First instar larvae have a pinkish yellow ground color with yellow-
orange tubercles that bear long gray setae. The head is tan-brown. The
second instar larva is dark gray with the tubercles light orange except
for the dorsal ones of the mesothorax and abdominal segments 4 and 7,
which are white. The third instar is white with the tubercles red-orange
except for the lateral ones on the meso- and metathorax, and on
abdominal segments 3 and 7, which are white. There are oblique
gray-brown stripes laterally on each segment. The fourth instar is
similar in pattern, but the white tubercles are highlighted by a more
extensive pattern of darker brown stripes. The remaining tubercles are
red. The head and thoracic prolegs are black. In the fifth instar (fig. 12)
the ground color is gray-black, with numerous oblique black stripes, the
white tubercles strongly contrasting. The remaining tubercles are
burgundy. Both the ground color and tubercle color exhibit a narrow but
conspicuous range of variability in the fifth instar.

The pupa (fig. 18) is yellow with a lavender-gray lateral abdominal
band that continues onto the wings and the lateral tubercles of the
mesothorax. The same color forms a midline dorsally along the thorax,
continuing onto the frontal tubercles and those of the abdomen as well.
The dorsal saddle of the abdomen has a pink midline and two fine
dorsolateral lines of gray. Ventrally the wings and legs are greenish
yellow.

22

J. Res. Lepid.

Oviposition and foodplants. Eggs are always laid singly on the
undersides of leaves of the foodplant or on the tips of new vegetative
shoots. Ovipositing females are easily followed as they exhibit a slow,
fluttering flight close to the ground. Eggs are never deposited higher
than 1.5 m above the ground. At Estapilla, in arid thorn scrub, a female
was observed ovipositing on an Aristolochia acontophylla plant which
was only 15 cm tall. The same species of foodplant is utilized at Zapotan
and Madrid. In the latter locality P. montezuma also was observed
ovipositing on A. foetida, although no larvae were ever encountered on
this species. At Madrid an undescribed species of Aristolochia was
available, but never utilized by P. montezuma. Eggs of P. montezuma
were found once on A. cardiantha. At higher elevations A. conversiae is
utilized.

Habitat and range. P. montezuma is generally common throughout its
range from sea level to about 1360 at Cofradia de Suchitan, occupying a
variety of habitats including arid thorn and tropical deciduous forests.
Unlike P. photinus, P. montezuma does not occur in the arid pine-oak
forest.

Parides erithalion trichopus (Rothschild and Jordan) (figures 7, 13,
and 19)

Early stages. An outline description of the early stages of this species
was presented by de la Maza (1980). The egg (fig. 7) is pale gray with
about 12 irregular, warty, vertical rib-like strands of orange glue-like
colleterial secretion. Eggs are laid singly on the underside of leaves of
the foodplant, and rest on a short pedestal of colleterial substance to one
side of the base. The glue-like strands are more granular in appearance
than on the eggs of either P. montezuma or P. photinus , and the pedestal
is shorter than in those two species.

The first instar is orange-yellow except for the mesothoracic tubercles
and the dorsal tubercles of abdominal segments 4, 7, and 9, which are
translucent white. The prothoracic shield is gray-brown and the head is
shiny black. The second instar is yellow-gray with the lighter tubercles
brighter white. The lateral tubercles of the prothorax, metathorax, and
abdominal segments 7 and 9 are also white. The remaining tubercles
are translucent orange-yellow. In the third instar the ground color is
reddish brown, the darker tubercles pinkish red. The lateral tubercles of
abdominal segments 7 and 8 are white. The fourth instar is similar, but
the ground color and darker tubercles are burgundy, and the white
tubercles on abdominal segments 3 (lateral) and 4 (dorsal) are joined by
an oblique white stripe. The tips of the dorsal tubercles on abdominal
segment 5 are white. In instars one through four, the tubercles bear
setae. These are longest on the first instar and decrease in length at each
successive molt. Fifth instar larvae are dimorphic: one form (fig. 13) is
gray -black, the other dark brown. Both are marked with oblique

26(l-4):l-288, 1988

23

charcoal-black lines. The white of the lighter tubercles is quite exten¬
sive and gives the larva a clearly defined anterior, median, and posterior
pattern of lines and spots.

The pupa (fig. 19) is greenish gray; the dorsal region, the saddle of the
abdomen, and the cremaster are yellow. The dorsolateral tubercles of
the abdomen are broad and triangular in profile.

Ouiposition and foodplants. Females exhibit a slow, low, fluttering
flight while searching for foodplants. Eggs are laid singly on the
underside of leaves of the host, or occasionally near the tips of new
shoots. At Tamala, La Salada, Madrid, and Caleras, P. erithalion
utilizes an undescribed, deciduous species of Aristolochia. Other hosts
documented from the lowlands include A. tentaculata and A. mutabilis.
Aristolochia conversiae is assumed to be the host at mid elevations (e.g.,
San Antonio and Tonila) where it is the only potential foodplant
available.

Habitat and range. P. erithalion trichopus is limited to western
Mexico. In Colima it has been observed from sea level to 1520 m, but this
may not be its upper limit. The species was abundant in arid thorn
forest, less common in arid thorn scrub. With the exception of large
urban areas, P. erithalion was fairly common in all habitats.

Mimetic Associations of Adults

In Jalisco, Battus philenor and Parides alopius (Godman and Salvin),
both assumed to be unpalatable, form a Mullerian mimetic complex. In
Colima, B. philenor is the model for the palatable Papilio polyxenes
asterius Stoll, a Batesian mimic over much of their extensive concurrent
geographical ranges. B. polydamas, B. laodamas, and B. eracon are
assumed to be unpalatable, and comprise a Mullerian mimetic complex
in Colima, in which B. polydamas is the most common species. Also
associated with these species are the Batesian mimics Papilio uictorinus
morelius Rothschild and Jordan, which is common, and females of
Papilio astyalus Godart, a population similar to P. astyalus occidentalis
Brown and Faulkner, which are uncommon. Females of Papilio an-
drogeus epidaurus Godman and Salvin also may be Batesian mimics of
the Battus complex.

The three Parides species in Colima form their own Mullerian
mimetic complex, again assuming unpalatability. Papilio pharnaces
(Westwood), P. anchisiades Esper, Eurytides thymbreus aconophus
(Gray), and E. belesis (Bates) are Batesian mimics of the Parides group,
the former representing an extraordinary convergence to Parides in
both coloration and pattern. The nymphalid butterfly Biblis hyperia
(Cramer) is also a Batesian mimic of Parides in Colima.

Conclusions

Table 2 summarizes the field data of foodplant utilization by Troidini

24

J. Res. Lepid.

Table 1. Characteristics and distribution of Aristolochia species in Colima,
Mexico.

hexandrous/ growth elevations general

species_ pentandrous habit size habitat in Colima_ distribution

A. tentaculata
Schmidt

hexandrous

glabrous

small

lianas

large;
to 7 m

widespread

common to 800 m;
sparse 800-900 m;
pockets to 1036 m

Colima,
Michoacan,
Guerrero, D.F.

A. glossa
Pfeifer

hexandrous

glabrous

lianas

very

large

riparian

0-250 m

Colima and
Michoacan

A. odoratissima
Linnaeus

hexandrous

glabrous

lianas

large

riparian

0-250 m

Mexico to
Panama

A. n. sp

pentandrous

dwarf

herb

small;

2. 0-2. 5 m

rocky

hillsides

0-250 m

unknown

(Colima)

A. acontophylla
Pfeifer

pentandrous

sprawling

herbaceous

perennial

small;
less than
1.0 m

open sunny
areas in
thorn forest

below 300 m

Colima and
Michoacan

A. cardiantha
Pfeifer

pentandrous

procumbent

perennial

herb

small;
2.0 m +-

thorn

forest

ca 1000 m

Colima,
Guerrero ,
Mexico D. F.

A. foetida

H. B. K.

pentandrous

procumbent

perennial

herb

small;
less than
1.0 m

thorn

forest

200-1128 m

Jalisco, Colima,
Michoacan, and
Guerrero

A. conversiae
Pfeifer

pentandrous

procumbent

perennial

herb

small

pine-oak,

tropical

deciduous

960-1740 m

Colima,

Mexico D.F.

A. pringlei

Rose

pentandrous

twining

perennial

herb

small

moist meadows
in pine-oak
forest

2285 m

Nayarit,

Jalisco, Morelos,
and Michoacan

A. mutabilis
Pfeifer

pentandrous

procumbent

perennial

herb

small

thorn

forest

0-250 m

Colima,
Michoacan,
and Guerrero

Table 2. Summary of foodplant utilization by Colima Troidini.

philenor

polydamas

eracon

laodamas

photinus

montezuma

erithalion

A. tentaculata

+

+

+.

++

++

+

A. qlossa

?

7

A. odoratissima

+

7

7

A. n. sp.

+

A. acontophylla

++

+

++

A. cardiantha

+

A. foetida

+

7

+

A. conversiae

+

++

++

7

A. pringlei

+ +

+

A. mutabilis

+

+ + = preferred foodplant (4 or more observations)

+ = documented foodplant (1-3 observations)

? = possible foodplant (most likely available host; no observations)

in Colima, Mexico. Aristolochia tentaculata is an important foodplant to
nearly all the species studied. K. Brown (personal communication)
suggests that the relative toxicity of different species of Aristolochia
varies, and that the first choice of foodplant would be the least toxic
species available. Where troidines occur in sympatry, there may be
competition for such a foodplant. In the absence of specific biochemical
data on plant secondary compounds, A. tentaculata is suspected to
represent such a species in Colima.

26(l-4):l-288, 1988

25

From this study Battus eracon and B. laodamas procas appear to be
monophagous, although this may not be the case. It is possible that both
species utilize all three of the large hexandrous Aristolochia species
available. B . poly damas is the most polyphagous troidine; it successfully
utilizes five species of Aristolochia in Colima.

There is no unequivocal evidence from the data or observations that
Colima troidines are partitioning hosts to avoid interspecific competi¬
tion, but this is implied by regional shifts in foodplant choice. Where, for
example, several troidines are sympatric, host utilization seems to differ
from allopatric situations. Partitioning by Battus and Parides between
hexandrous and pentandrous species of Aristolochia in Colima is also
suggested. Pentandrous species are small, usually only 1— 2 m in height;
hexandrous species are typically much larger (table 1). Parides species
lay their eggs singly (Brown, Damman, and Feeny 1981); females
usually search at ground level. Parides thus are able to exploit small,
low-growing pentandrous species. In contrast, Battus species typically
lay eggs in batches. Gregarious larvae soon devour small foodplants and
have to search for additional host material. Survival is possible on
pentandrous species only where close growing stands occur. In general,
Battus species would seem to have greater survival and face less
intraspecific larval competition on large hexandrous species, where
ample foodplant is available for large numbers of actively feeding larvae.
This assumption is supported by the fact that B. eracon and£. laodamas
were observed to feed exclusively on the large hexandrous A. tentacula-
ta. Evidence to the contrary includes the preference for pentandrous
species exhibited by B. philenor, which is moderately polyphagous, and
the inability to determine preference of B. poly damas.

Acknowledgements. We wish to thank the following for reading the manu¬
script at various stages and offering valuable suggestions: John R. Arnold,
Carlos R. Beutelspacher, Keith Brown, William Burger and the botany staff at
the Field Museum, David K. Faulkner, Javier de la Maza, Michael Parsons,
Jerry A. Powell, Tommaso Racheli, and Arthur Shapiro. We also thank Kerry
Berringer of the Field Museum of Natural History, Chicago, for identifying
specimens of Aristolochia from Colima, and Jose F. Ortega Ortiz for information
on Aristolochia species in eastern Mexico.

Joyce Hayashi provided the map of Colima and helpful suggestions.

The senior author offers special thanks to Biologo-Investigador Sergio Her¬
nandez Tobias of Colima, for sharing information on collecting localities.

Literature Cited

BROWER, J. V. Z. 1958. Experimental studies of mimicry in some North American
butterflies. II. Battus philenor and Papilio troilus, P. polyxenes, and P.
glaucus. Evolution 12: 123—136.

BROWN, K. S„ A. J. DAMMAN, & P. FEENY. 1981. Troidine swallowtails (Lepidoptera:
Papilionidae) in southeastern Brazil: natural history and foodplant rela¬
tionships. J. Res. Lepid. 19(4): 199-226. |“1980”1.

26 J. Res. Lepid.

COMSTOCK, J. A. & L. VAZQUEZ G. 1960. Estudios de los ciclos biologicos en
lepidopteros Mexicanos. An. Inst. Biol. Mex. 31: 349-360.

DE LA MAZA E., R. 1980. Las poblaciones Centro- Americanas de Parides erithalion
(Boisduval) (Papilionidae: Troidini). Rev. Soc. Mex. Lepid. 5(2): 51-73.
DIAZ FRANCES, A. & J. DE LA MAZA. 1978. Guia ilustrada de las mariposas
Mexicanas. I. Papilionidae. Soc. Mex. Lepid. Publ. Esp. 3.

EDWARDS, W. H. 1881. Description of the preparatory stages of Papilio philenor ,
Linn. Can. Entomol. 13: 9-14.

EHRLICH, P. R. & P. H. raven. 1965. Butterflies and plants: a study in coevolution.
Evolution 18: 586-608.

EMMEL, J. 1975. The family Papilionidae, in, Howe, W. (ed.), The butterflies of
North America. Doubleday and Co., Inc., Garden City, New York. 633 pp.
FRAENKEL, G. S. 1959. The raison doetre of secondary plant substances. Science
129: 1466-1470.

HOFFMAN, C. C. 1976. Catologo sistematico y zoogeografico de los lepidopteros
Mexicanos. Soc. Mex. Lepid. Publ. Esp. 1.

KLOTS, A. B. 1951. A field guide to the butterflies of North America east of the
Great Plains. Houghton Mifflin Co., Boston, Mass. 349 pp.

MOSS, A. M. 1919. The papilios of Para. Nov. Zool. 26: 296—319.

MUNROE, E. 1960. The generic classification of the Papilionidae. Can. Entomol.
Suppl. 17: 1-51.

OPLER, P. A. & G. O. krizek. 1984. Butterflies east of the Great Plains. John
Hopkins Press Ltd., London. 294 pp.

PFEIFER, H. w. 1966. Revision of the North and Central American hexandrous
species of Aristolochia (Aristolochiaceae). Ann. Missouri Bot. Garden 53:
115-196.

_ 1970. A taxonomic revision of the pentandrous species of Aristolochia.

Univ. Conn. Pub. Series. 134 pp.

RAUSHER, M. D. 1980. Host abundance, juvenile survival, and oviposition prefer¬
ence in Battus philenor. Evolution 34: 342—355.

_ 1981. Host plant selection by Battus philenor butterflies: the role of

predation, nutrition, and plant chemistry. Ecological Monographs 51(1):
1-20.

ROTHSCHILD, W. & K. JORDAN. 1906. A revision of the American papilios. Nov. Zool.
13: 411-752.

SCHALDACH, W. J. 1963. The avifauna of Colima and adjacent Jalisco, Mexico.

Proc. Western Found. Vert. Zool. 1(1): 1-100.

SCRIBER, J. M. & P. FEENY. 1976. New foodplant and oviposition records for Battus
philenor (Papilionidae). J. Lepid. Soc. 30: 70—71.

WEST, D. A. & W. N. HAZEL. 1979. Natural pupation sites of swallowtail butterflies
(Lepidoptera: Papilionidae): Papilio polyxenes Fabr., P. glaucus L., and
Battus philenor (L.). Ecological Entomol. 4: 387—392.

YOUNG, A. M. 1971a. Mimetic associations in natural populations of tropical
butterflies. I. Life history and structure of a tropical dry forest breeding
population of Battus polydamas polydamas. Rev. Biol. Trop. 19: 210-240.
_ 1971b. Mimetic associations of tropical papilionid butterflies (Lepidop¬
tera: Papilionidae). J. New York Entomol. Soc. 79: 210—224.

_ 1972. Mimetic associations in populations of tropical butterflies. II.

Mimetic interactions of Battus polydamas and£. helus. Biotropia 4: 17—27.

Journal ofResearch on the Lepidoptera

26(l-4):27-31, 1988

The Mating Behavior of Papilio glaucus (Papilionidae)

Robert A. Krebs*

Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg,
VA 24061

Abstract. Male and female Papilio glaucus were released in a large
flight cage containing vegetation simulating a forest clearing. Obser¬
vations were made to study courtship behavior and the mating system
of P. glaucus. Mechanisms of female choice through solicitation of
males and rejection behavior during courtship are presented.

Introduction

The tiger swallowtail, Papilio glaucus L., has monomorphic non-
mimetic males as well as two female forms: one male-like and the other
a Batesian mimic of Battus philenor L. It has therefore been of interest
in studies of assortative mating (Burns, 1966; Platt, Harrison and
Williams, 1984) and sexual selection in mimetic species (Brower, 1963;
Silberglied, 1984; Krebs, 1986). Despite this general interest, little
information is available on its mating behavior. This note describes
courtship and mate avoidance behaviors in P. glaucus and suggests
mechanisms for female choice.

Materials and Methods

Adults were reared from eggs of field-caught females (Virginia) and females
sent by Mark Scriber, University of Wisconsin (Wisconsin and Illinois), for
experiments in 1984 and 1985. Although geographically variable, all butterflies
used were P. glaucus glaucus. Larvae were reared on fresh black cherry leaves
{Prunus serotina ) in the laboratory under a long photoperiod to prevent
diapause which allowed rearing of three generations.

The experimental population for 1985 was produced by randomly crossing
field-caught, Wisconsin, and Illinois butterflies early in the experiment and
rearing and crossing these offspring throughout the summer. Observations
therefore began with pure strain individuals presented at random and continued
with offspring of geographically mixed parentage. Observations made in the
Spring of 1984 included only reared Blacksburg individuals.

Observations were made in a flight cage (5 x 8 x 5 m) in Blacksburg, Virginia.
The cage resembled a forest clearing with small trees, including Prunus
serotina , inside. Vines lined the sides and larger trees surrounded the cage.

Because an outdoor cage was used, weather conditions could not be controlled.
All observations were made between noon and 4:30 PM on days ranging in
temperature from 22 to 33°C. Temperatures outside this range, high winds and
cloudy conditions decreased flight too much to allow for efficient testing.

A single virgin female, 1 to 3 days old, was released to one male, 2 to 4 days old.

Present address: Dept, of Zoology, Arizona State University, Tempe Az 85287

28

J. Res. Lepid.

Observations were made until either several unsuccessful courtships or a
mating occurred. The average observation time was one hour per pair.

Results

General

A total of 196 courtships using 70 P. glaucus pairs (17 in 1984 and 53
in 1985) was observed. Of these, 34 (17%) led to matings. Because
results were similar when either Virginia butterflies were presented to
each other (1984) or the geographically mixed populations (1985) were
used, structure of successful courtships versus unsuccessful ones for all
presentations are grouped when discussed below.

Successful courtships

Typical courtships leading to copulation involved an exchange of
behaviors in flight between males and females. These flights were
initiated when responsive males encountered females, usually in air (27
of 34) or on vegetation (6 of 34). Within the cage, males usually initiated
courtship. However, in nine courtships leading to copulation, females
flew toward the male, soliciting courtship. One such flight was directed
to a male on vegetation.

Following initial interactions, females flew up and away from males
which pursued 5-15 cm below and behind. Most of these courtship flights
therefore occurred along the cage top at 5 m. In two observations
butterflies released in the field ascended into and over tree tops and out
of sight.

Courtship flights were highly variable in length (x=16.5s±3.9s,
n = 18, range 0-59 s). However, only 7 of 34 matings occurred after the
first courtship. Total courtship flight time preceding copulation averaged
58 ± 12 s (n = 18). Successful males averaged 2.6 courtships (n = 34)
before they were accepted by females.

Pursuit flights continued until the females landed with wings either
open (7 of 31) or closed (24 of 31). The male hovered above the female for
a second or two before attempting to land beside her (23 of 34), but
sometimes (11 of 34) immediately landed by the female. When a female
landed with open wings, the male always hovered. Wing closing by a
female was quickly followed by the male alighting beside the female.

Three courtships leading to mating lacked usual courtship flights. In
those, the males hovered over the females perched on vegetation, landed
and were accepted. However, courtship flights had occurred previously
in all three.

A male, once beside the female, extended his abdomen to contact the
female’s genitalia. After acceptance, he relaxed, dropped below the
female and remained stationary in copula for 45 min to an hour. Most
matings occurred along the cage top, with only a few as low as three

26(l-4):l-288, 1988

29

meters. No post copulatory flights were observed unless the pair was
disturbed. In flight, females flew with males hanging below.

Unsuccessful courtships

Unsuccessful courtships were ended by either males or females. Of 162
unsuccessful courtships, designated as those interactions between
males and females which lasted at least a second, 33 encounters in air
never led to a pursuit flight. These interactions ended when males failed
to respond to females (26 males, not included in the above total, never
courted females presented to them).

Of the other 129 courtship flights, 104 broke up while males were
pursuing females. The remaining 25 ended after males interacted with
perched females. Because these males were responding positively to
females, failure to mate was probably due to behaviors on the part of
females to evade or reject courting males.

Unsuccessful courtships lasted longer than those that ended in
copulation (x = 103 ± 15 s, n = 18, p<0.02). Number of courtships,
however, did not differ, 2.72 ± .21 for unsuccessful males and 2.56 ± .26
for successful ones (p>0.3).

Discussion

Several aspects of P. glaucus mating behavior are very different from
that of other species. Most notable is the high incidence of courtship
solicitation by females. Solicitation flights were generally made directly
to males in flight, although perching males were also solicited. Females
flew across either from the side or above males within 15cm, turned, and
flew up and away. If males failed to pursue, solicitation was often
repeated.

Solicitation flights have also been observed in Pieris protodice Bois-
duval and LeConte (Rutowski, 1980), Heliconius erato L. (Crane, 1955),
Danaus gilippus Cramer (Brower, Brower and Cranston, 1965) and
Aphantopus hyperanthus L. (Wicklund, 1982). In P. glaucus , solicita¬
tions were observed for 53% of males that mated; 9 of 34 matings had
been immediately preceded by solicitation. Thirty-seven percent of all
first interactions between males and females were initiated by females.
Also, of 26 males that were not responsive to females, 85% received
solicitations.

A second unusual feature of P. glaucus courtship is lack of antennal
contact between males and females. Brower et al. (1965) described
males of D. gilippus brushing antennae of females with specialized scent
scales. Scent is important in courtship to many other butterfly species
(Thornhill and Alcock, 1983). However, as P. glaucus males court from
below and behind females, some wing contact occurs from below but
none near the female’s head. Opportunity to pass scent did occur when
females landed but only occasional wing contact was observed before the

30

J. Res. Lepid.

male attempted to land and copulate. These behaviors suggest that
visual cues are far more important than olfactory in mate choice in this
species.

With only 17% of all courtships with virgin females leading to
matings, females are able to reject and avoid males. Longer courtship
times of unsuccessful males suggested that females avoided landing
until pursuing males were evaded.

Two observed avoidance postures were closing and depressing the
wings when a male flew near, and depressing the abdomen to the
substrate to avoid genital contact when a male landed. Rejection
behaviors during pursuit included “quick landing,” a sudden stop with
wings closed and depressed, and “dropping,” a relaxed free fall into
brush. Females also employed slow descending flights, which forced
males to abandon courtship when females hovered less than 30 cm
above the ground. Two less obvious behaviors were flying through thick
brush, also observed in D. gilippus (Brower et al. 1965), and simply not
flying, a behavior I observed when females were released with a high
male density. Virgin females have been observed to reject males by like
means in other species (Rutowski, 1982, 1984).

Comparisons in this study suggested no differences between male
responses to the two female forms, mimetic and male-like. While the
experimental design did not provide for controlled comparisons of
details within courtship flights, overall mating success and male
courtship frequency were not different.

One question in this study is its application to mating behavior in
nature. Only two courtship flights, described earlier, were observed
outside of the cage, although releases, albeit unsuccessful, were at¬
tempted. However, vegetation within the cage, and its size, provided
more natural conditions than in most other cage studies.

Brower (1963) says that P. glaucus males fly around courting any
females located. No other description of a mating system exists for this
species, and no territorial behavior is known. Multiple male releases in
the cage elicited no male-male interaction. Attempts by several males to
court the same female simultaneously were observed. None were
successful. Only in mudpuddling aggregations are high densities of
males found.

Rutowski (1984) suggests that prolonged searching polygyny is the
most likely mating system to be found in butterflies. Strong rapid flight,
lack of male-male competition and dispersed abundant food and ovipo-
sition sites suggests the existence of this system in P. glaucus.

Acknowledgments. Thanks to Dave West for discussion and advice throughout
my work, Ron Rutowski for reviewing an earlier draft of this manuscript and to
Therese Markow for making me take the old data off the shelf. Two anonymous
reviewers suggested several ways to improve clarity. This research was funded
by small grants from Sigma Xi and the Virginia Academy of Science.

26(1-4): 1-288, 1988

31

Literature Cited

BROWER, L. P., 1963. The evolution of sex-limited mimicry in butterflies. Inter.
Congr. Zool. 16(4):173-179.

BROWER, L. P., J. V. Z. BROWER & F. P. CRANSTON, 1965. Courtship behavior of the
queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 50:1-39.

BURNS, J. M., 1966. Preferential mating versus mimicry: disruptive selection
and sex-limited dimorphism in Papilio glaucus. Science 153:551-553.

CRANE J., 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato
hydara Hewitson, with special reference to the social use of color. Zoologica
40:167-196.

KREBS, R. A., 1986. The effect of female mate preference on the evolution of
Batesian mimicry. M. S. thesis, Virginia Polytechnic Institute and State
University, Blacksburg, Virginia.

PLATT, A. P„ J. HARRISON, & T. F. WILLIAMS, 1984. Absence of differential mate
selection in the North American tiger swallowtail Papilio glaucus, pp. 245-
250. In R. I. Vane-Wright and P. R. Ackery (eds.), The Biology of Butterflies.
Symp. R. Entomol. Soc. Lond. No. 11.

RUTOWSKI, R. L., 1980. Courtship solicitation by females of the checkered white
butterfly, Pieris protodice. Behav. Ecol. Sociobiol. 7:113-117.

RUTOWSKI, R. L., 1982. Mate choice and lepidopteran mating behavior. FI.
Entomol. 65:72-82.

RUTOWSKI, R. L., 1984. Sexual selection and the evolution of butterfly mating
behavior. J. Res. Lepid. 23:125-142.

SILBERGLIED, R. E., 1984. Visual communication and sexual selection among
butterflies, pp. 207-223. In R. I. Vane-Wright and P. R. Ackery (eds.), The
Biology of Butterflies. Symp. R. Entomol. Soc. Lond. No. 11.

THORNHILL, R. & J. ALCOCK, 1983. The evolution of insect mating systems.
Cambridge, Massachusets: Harvard University Press.

WICKLUND, C., 1982. Behavioural shift from courtship solicitation to mate
avoidance in female ringlet butterflies ( Aphantopus hyperanthus ) after
copulation. Anim. Behav. 30:790-793.

Journal of Research on the Lepidoptera

26(l-4):32-38, 1988

A New Heritable Color Aberration in the Tiger
Swallowtail Butterfly, Papilio glaucus (Papilionidae:
Lepidoptera)

J. Mark Scriber1
and

Mark Evans

Department of Entomology, University of Wisconsin, Madison, WI 53706

Abstract. A new wing color aberration in the Eastern tiger swallow¬
tail, Papilio glaucus L., has been discovered and is called “dark cell”.

This dark scale suffusion into the dorsal forewings has a genetic basis
and appears restricted to females. We describe the new aberration and
our attempts to rear offspring of hand-pairings for elucidation of the
genetic basis of the trait.

Introduction

A number of wing color aberrations in Papilio glaucus L. have
recently been described by Clark and Clark (1951), Clarke and Clarke
(1983) and Scriber et al. (1987). Here we report a new wing color
aberration in Papilio glaucus L. which is apparently genetically based
and sex-limited in expression. Following the first appearance of this
aberration (which we call “dark cells”) in our 1982 laboratory cultures,
we hand-paired (sibs) in an attempt to elucidate the genetic basis of the
character.

Methods

Oviposition by adult females was induced by placing each wild-
captured or hand-paired female into its own clear plastic box (approxi¬
mately 10 cm deep x 15 cm x 30 cm) with a moist paper towel and
selected larval foodplant leaves. Leaf turgor was maintained in these
plants by use of floral aquapics® (water-filled plastic vials with a rubber
cap, through which leaf petioles or small branches can be inserted; see
Scriber, 1977). Heat and light were provided by an incandescent bulb
placed at a distance of approximately 0.3-0. 5 meter from the plastic
boxes.

Larvae were reared through to pupation on black cherry, Prunus
serotina Ehrh., or another foodplant (leaves were changed three times
per week) under controlled environment conditions (16:8 photo-/scoto-
phase with corresponding temperatures of 23.5°/19.5°C, respectively).

1Current address: Dept. Entomology, Michigan State University, East Lansing, MI48824

26(l-4):l-288, 1988

33

Pupae were weighed 2 days after pupation (the weight subsequently
serving as an identification number for the individual) and then placed
in cylindrical screen cages (15 cm diameter x 12 cm height) under larval
rearing conditions to permit development and eclosion as adults. Non-
diapausing individuals normally emerged within 2-3 weeks after pupa¬
tion. Other pupae were allowed at least 6 weeks before being refrigerated
in darkness (at 40-45°F for 3 months or more) to break diapause. Hand-
pairings were generally attempted 12-48 hrs after adult female eclosion
and 2-3 days after male eclosion.

Results

We first observed our new color aberration in females of a 1982 lab-
reared brood (#56). We term this aberration “dark cell” because of the
abnormal suffusion of dark scales into the normally yellow cells of the
forewings (Fig. 1). Brood #56 was the result of a yellow morph mother (a
virgin P. g. glaucus, 3rd generation lab-reared, originally from stock
collected in 1981 from Schuylkill County, Pennsylvania by William
Houtz) which was mated to a male whose mother was also from this PA
stock but whost father was a subspecies hybrid (from a P.g. canadensis
female from Clarke Co., Wisconsin mated to a P. g. glaucus male from
Pennsylvania).

Crosses #290 and #296 (Table I) are both crosses from a “dark cell”
daughter and one of her male siblings (all from brood # 56). It can be
seen that this dark cells trait occurs only in the females. The four
siblings shown here (Fig. 1) represent some of the variation in the “dark
cell” trait.

The origin of this character, and the genetic basis of its expression are
difficult to decipher because of the complexity of the parental lineages,
and the fact that no livestock currently exists from this lineage (brood
#56, #290, or #296). Only two additional related females were hand-
paired: one with a P. g. glaucus from Richland Co., Wisconsin (brood
#299; see Table 1), and the other produced no eggs.

While checking the results of our 1982-1986 hand-pairings, lab
rearings, and field captures, we discovered a total of 5 additional “dark
cell” phenotypes in our research collection (> 25,000 butterflies). Each
of these individuals resulted from a subspecies pairing involving a P. g.
canadensis male parent, and each also emerged in 1984 from hand-
pairings done in 1982. The first was one of 13 female progeny (19 male
progeny) from a dark male P. g. glaucus from Georgia meted to a P. g.
canadensis from northern Wisconsin (pairing #39). A second was one of
4 female progeny (9 male progeny) of a lab-reared yellow P. g. glaucus
female (from Pennsylvania parents) mated to a P. g. canadensis from
Bayfield County, Wisconsin (pairing #116). The third and fourth were
female siblings out of a total of 7 females (4 male sibs) from a yellow lab-
reared P. g. glaucus female (from Pennsylvania parents) mated to a P. g.

34 J. Res. Lepid.

Table 1. Adult phenotypes1 from laboratory crosses. Madison, Wiscon¬
sin (1982)

Number of Female

Mother

number

Pairing

Larvae/

Number

Offspring

Background

total

of Male

“Normal”

“Dark

(female x male)

eggs

Offspring

yellow

cell”

Female #56

(#71 x #H15)

74/132

8

3

9

Female #290*

(#56 x #56)

172/391

15

8

7

Female #296

(#56 x #56)

201/271

18

1

22

Female #299

(#56 x P. g .)

190/208

13

6

0

xThe “dark cell” aberration was first detected in offspring of female #56.
Two fertile sibling-sibling pairings (290 and #296) both resulted in
some “dark cell” phenotypes, differing from #299. One additional
daughter from #56 was mated (female #537) but produced no eggs. No
other pairings were made from the female #56 lineage.

*One deformed pharate female was not able to be classified.

canadensis from Juneau County, Wisconsin (pairing #144). The final
“dark cell” phenotype was a female from a yellow lab-reared P. g.
australis female from Florida, mated to a P. g. canadensis from Bayfield
County, Wisconsin (pairing #115). Only one living pupa from these
lineages currently exists.

Discussion

Sir Cyril Clarke and colleagues have been investigating the genetic
basis of abnormal wing coloration in Papilio glaucus for decades (see
Clarke and Clarke, 1983 for a review). They point out that color mosaics
and gynandromorph P. glaucus can be striking, because of the contrast
between the yellow background (of males and yellow morph females)
against the black/brown melanic background of the dark morph females

Fig. 1 . Four female siblings (of 22 "dark cell’’ produced in brood #296) exhibiting
variations of our "dark cell’’ aberration: Left, dorsal; Right, ventral. This
aberrant was originally detected in the parental brood (#56; see text
and Table 1)

A) pupal weight 0.8720 g; eclosed 29 Sept. 1982

B) pupal weight 1.1946 g; eclosed 29 Sept. 1982

C) pupal weight 1,6900 g; eclosed 23 Sept. 1982

D) pupal weight 1.2956 g; eclosed 20 Sept. 1982

26(l-4):l-288, 1988

35

36

J.Res.Lepid.

These dark/yellow gynandromophs and color mosaics are quite rare,
and considerable attention has been given to existing specimens. For
example, the Herman Strecker collection (currently on loan from the
Chicago Field Museum to the Allyn Museum in Florida) contains a
number of such mosaics. This valuable collection, assembled during the
second half of the 19th century, contains a number of P. g. glaucus
mosaics previously discussed in the literature (Strecker, 1878; Ehrmann,
1894; Howard, 1899; Walsten, 1977; Ehle, 1981; Shapiro, 1981b Clarke
and Clarke, 1983). The Milwaukee Public Museum (Milwaukee, Wis¬
consin) also contains (in the Neidhoefer collection) two partial color
mosaics reared by E. Dluhy in Chicago, Illinois. Color mosaics also exist
from Richmond County, NY (5 July 1971; A.M. Shapiro; currently in the
University of California Davis Collection) and from Washington County,
PA (9 May 1927; George F. Patterson Collection at Pennsylvania State
University).

Scriber and Evans (in press) describe an additional two dozen
color mosaics from Papilio glaucus (see also Scriber et al, 1987);
however, in the investigation of this entire group of color aberrations
and in investigations of Papilio glaucus from (many) institutional and
personal collections (see Scriber and Evans, 1986b), we have never
encountered material similar to the “dark cell” aberration P. glaucus
females described here (Fig. 1).

We are aware of the superficial resemblance of “dark cell” to the
melanic aberration “fletcheri” in males of P. g. canadensis , and we have
reviewed the literature and figured this form from Wisconsin previously
(Scriber and Lintereur, 1983). The “fletcheri” aberration has been noted
several times from northern Wisconsin (Ebner, 1960; Scriber and
Lintereur, 1983; and W. Gould, J. Trick, D. Robacker, D. Matusik pers.
comm.) and it is possible (though we consider it remote) that there has
been introgression from our handpairing with P. g. canadensis in the
lineage leading to the male parent of brood #56. It should be noted
however that the “fletcheri” aberration is generally believed to be
restricted to the male, and apparently toP. g. canadensis. Furthermore,
“fletcheri” exhibits significant suffusion of dark scales into both the
hind wings and fore wings, with an orange smear encroaching ventrally
as well as dorsally across the hindwings (see color figure in Scriber and
Lintereur, 1983). Our “dark cell” aberration is essentially restricted to
the forwings, and only their dorsal surface (Fig. 1).

We do not necessarily mean to imply that the “dark cell” trait
(restricted to females) can not be genetically related to the “fletcheri”
aberration (apparently restricted to males). In fact, we know that sex-
linked (female Y-chromosome) control of the dark morph color poly¬
morphism in P. glaucus females (Clarke and Sheppard, 1962) can be
transmitted by males in what we consider to have been either a
crossover event or a non-disjunction (Scriber and Evans, 1986). The
role ofP. g. canadensis introgression in these events is unclear, but such

26(l-4):l-288, 1988

37

gene flow between the two subspecies can be disruptive in a number of
ways to other morphological/color traits (Scriber, 1982; Luebke, 1986;
Rockey et al, 1987; Scriber, 1987). Unfortunately we will be unable to
conduct any further studies of this now extinct “dark cell” lineage.

Acknowledgements. This research was supported by the National Science
Foundation (DEB #79 21749, BSR #8306060), by the Graduate School and the
College of Agriculture and Life Sciences (Hatch Project 5134) at the University
of Wisconsin, Madison. In these studies and our discussions we are especially
greateful to: Thomas Allen, William Bergman, Susan Borkin, Don Caine, Susan
Heg, Sue Helgeson, William Houtz, Greg Lintereur, Dave Matusik, Jeanne
Pomraning, Howard Romack, Donald Strasburg, Bill Warfield, and Allen
Young. We are also especially thankful to Jacqueline Miller and Lee Miller for
their assistance at the Allyn Museum in examining the Herman Strecker
Collection mosaics and gynandromorphs. An anonymous reviewer made several
helpful suggestions to an earlier manuscript.

Literature Cited

CLARK, A. H. & L. F. CLARK. ( 1951) The butterflies of Virginia. Smithsonian Institute
Miscellaneous Collection, 116(7), 124-144.

CLARKE, C. & F. M. M. CLARKE ( 1983) Abnormalities of wing pattern in the eastern
tiger swallowtail butterfly Papilio glaucus. Systematic Entomology, 8, 25-
28.

CLARKE, C. A. & P. M. SHEPPARD (1962) The genetics of the mimetic butterfly,
Papilio glaucus. Ecology 43:159-161.

EBNER, J. A. (1960) A striking melanic male of Papilio glaucus. J. Lepid. Soc.
14:157-158.

EDWARDS, w. H. (1868) Notes on a remarkable variety ofP. turnus and descrip¬
tions of two species of diurnal Lepidoptera. Trans. Amer. Entomol. Soc., 2,
207.

EDWARDS, w. h.(1884) Butterflies of North America. Vol. II. Philadelphia
Houghton Mifflin Co.

EHLE, C. (1981) Letter to the editor. News of the Lepidopterist’s Society (No. 5,
Sept. /Oct.), p. 63.

EHRMAN, G. A. (1893) A few remarkable variations in Lepidoptera. Canadian
Entomologist, 26, 292-293.

HOWARD, L. O. (1899) An abnormal tiger swallowtail. Insect Life, 7, 44-47.
LUEBKE, H.J. (1986) Hybridization in th e Papilio glaucus group: A morphometric
study using multivariate techniques. M. S. Thesis, Univ. Wisconsin,
Madison. 102 pp.

NIJHOUT, H. F. (1981) The color patterns of butterflies and moths. Scientific
American. November: p. 140-151.

ROCKEY, S. J., J. H. HAINZE, & J. M SCRIBER. (1987) A latitudinal and obligatory
diapaus response in three subspecies of the Eastern Tiger Swallowtail,
Papilio glaucus (Lepidoptera: Papilionidae): Amer. Midi. Naturalist (in
press).

SCRIBER. J. M. (1982) Foodplants and speciation in the Papilio glaucus group,
pp. 307-314 In Proc. 5th Intern. Symp. Insect-Plant Relationships. Pudoc,
Wageningen.

38

J.Res.Lepid.

SCRIBER, J. M. (1987) Tale of tiger: Beringial Biogeography, bionomial classifica¬
tion, and breakfast choices in the Papilio glaucus complex of butterflies. IN:
Chemical Mediation of Coevolution (K. C. Spencer, ed.) (submitted Dec.
'85). (in press, AIBS).

SCRIBER, J. M. & M. H. EVANS. (1986) An exceptional case of paternal transmission
of the dark form female in the tiger swallowtail butterfly, Papilio glaucus
(Lepidoptera: Papilionidae) Journal of Research on the Lepidoptera (25: in
press).

SCRIBER, J. M. & G. L. LINTEREUR (1983) A melanic aberration of Papilio glaucus
canadensis from northern Wisconsin. Journal of Research on the Lepi¬
doptera 21: 199-201.

SCRIBER, J. M„ M. H. EVANS & D. RITLAND (1987) Hybridization as a causal
mechanism of mixed color broods and unusual color morphs of female
offspring in the eastern tiger swallowtail butterflies, Papilio glaucus. In (M.
Huettel, ed.) Evolutionary Genetics of Invertebrate Behavior. Univ. of
Florida (in press).

SHAPIOR, A. M. (1976) Seasonal polyphenism. Evolut. Biol. 9:259-333.

SHAPIRO, A. M. (1981b) Letter to the editor; Ripples. News of the Lepidopterists’
Society (Jan.-Feb., No. 1, p. 5).

STRECKER, H. (1878) Butterflies and moths of North America. Owen Press,
Reading, PA. 280 pp.

WALSTEN, D. M. (1977) Letter to the editor; Ripples. News of the Lepidopterist’s
Society (Sept.-Oct., no. 5, p. 6).

VANE-WRIGHT, R. I. & P. R. ACKERY. 1984. The biology of butterflies. Academic
Press, London.

Journal of Research on the Lepidoptera

26(l-4):39-57, 1988

Bilateral gynandromorphs, sexual and/or color mosaics
in the tiger swallowtail butterfly, Papilio glaucus
(Lepidoptera: Papilionidae)

J. Mark Scriber1
and

Mark H. Evans

Department of Entomology, University of Wisconsin, Madison, WI 53706
1Current address and reprint requests to:

Department of Entomology, Michigan State University, East Lansing, MI 48824

Introduction

Mosaic specimens of Lepidoptera may be of a sexual nature (gynan¬
dromorphs or intersexes) or homeotic (involving an inappropriate
location for a particular feature or pattern; see reviews by Sibatani,
1980, 1983a, b). Intersexuality typically arises from development errors
late in development, and involves an individual which possesses a
mosaic of traits, some of which are female, and some of which are male
(see McCafferty and Bloodgood, 1986). Gynandromorphs generally
develop a sex abnormality much earlier in development; when this
occurs at the formation of one of the first two blastomeres, it is possible
for individuals to become bilaterally differentiated with one half male
and the other half female (Clarke and Ford, 1980; Ayala and Kiger,
1984). The relationship between intersexual mosaics and gynandro¬
morphs is not entirely clear, partly because of the infrequent occurrence
of both (Ford, 1955). The combined use of laboratory crosses of species or
subspecies, and the recent development of a technique to monitor the
heteropyknotic ‘Smith’ (S) body in the nucleus of somatic cells (derived
from the Y-chromosome of female Lepidoptera) should contribute to our
understanding of the development of these abnormalities (see Cross and
Gill, 1979; Clarke and Ford, 1980, 1983; Bull, 1983).

From the Papilionidae (see Table 1) gynandromorphs have been
reported from Papilio polyxenes asterias Stoll (Edwards, 1984; Blau,
1978; and Wm. Bergman, pers. comm.); Ornithoptera victoriae Gray,
and O. priamus L., (Schmid, 1973), O. croesus Wallace (Parrott and
Schmid 1984), O. poseidon (D’Abrera 1976); Parnassius autocrator
(Sbordoni and Forestiero, 1984); Papilio androgeus (Sbordoni and
Forestiero, 1984) and Papilio glaucus L. (Skinner, 1919; Cockayne,
1935; and Clarke and Clarke, 1983). In addition to these published
records of gynandromorphs and a large number of additional records
reviewed by Cockayne (1935), several hundred sexually mosaic and
bilateral gynandromorphic specimens exist in the Lepidoptera collec¬
tion of James R. Neidhoefer (housed at the Milwaukee Public Museum).

40

J. Res. Lepid.

Table 1. Gynandromorphs (Lepidoptera) reported in the literature.

Family

Genus species

References

Saturniidae

A utomeris io F abricius

Eacles imperialis Drury

Callosamia promethea (Drury)

Cassino and Reiff, 1917, Hessel, 1964;

Muller, 1966; Manley, 1977

Pyralidae

Hedylepta accepta (Butter)

Riotte, 1978

Geometridae

Phaeoura mexicanaria (Grote)
Antepione thisoaria (Guenee)

Blanchard, 1969; Durden, 1984

Lymantriidae

Lymantria dispar (L.)

Muller, 1976

Nymphalidae

Limenitis weidemeyerii latisfacia L.

L. arthemis-lorquini (Boisduval)

L. arthemis-astyanax (Fabricius)
Speyeria atlantis dodgei (Gunder)

Grey, 1959; Perkins and Perkins, 1972;

Platt, 1983; H. Romack pers. comm., 1984

Pieridae

Conepteryx rhamni L.

G. cleopatra. L.

Pieris brassicae L.

P. protodice Bdv. & LeC

P. rapae L.

Colias Christina Edwards

C. eurytheme Bdv.

C. philodice Godart

Pontia daplidice (L.)

T atochila steradice Stgr .

Emmel, 1964; Hovanitz, 1965; Nekrutenko, 1965;
Shapiro, 1970; Gardner, 1972; Sbordoni and
Forestiero, 1984; Shapiro, 1978, 1981, and 1985.

Hesperiidae

Polistes mystic (Scudder)

P. origines (Fabr.)

Erynnis horatius Scudder & Burgess
Hesperia Columbia (Scudder)

Nielsen, 1977; Israel andCilek,

1982; Scott, 1986

Lycaenidae

Strymon bazochii Godart

Lycaena gorgon (Boisduval)

Agriades rustica rustica (Edwards)
Mitoura gryneus (Hubner)

Celastrina ebenina Clench

Cyaniris semiargus (Rott.)

C. argiolus (L.)

Opler, 1966; Riotte, 1978; Rahn, 1982;

Sbordoni and Forestiero, 1984; Shuey, 1984;

Shuey and Peacock, 1985

Papilionidae

Papilio polyxenes asterias Stoll

P. glaucus L.

P. androgeus Cramer

Ornithoptera victoriae

0. croesus Wallace

0. poseidon Doubleday

O.priamus L.

Parnassius autocrator Avinoff

Skinner, 1919; Schmid, 1973; Cockayne, 1935;
Blau, 1978; W. Bergman, pers. comm. 1983; Clarke
and Clarke, 1983; Sbordni and Forestiero, 1984;
Parrott and Schmid, 1984; D’Abrera, 1976

Table 2. Progeny ( 1982 reared) of pairing #9 ( P . g. glaucus yellow morph female x P. g. canadensis male; see text
for further details).

Eclosion

Bilateral

Males

Females

Dead

Pupae

year

Gynandromorph

eclosed

pharate1

eclosed

pharate1

pupae

still alive

1982

0

7

0

0

0

4

0

1983

1

2*

4

6

1

8

0

1984

0

0

0

6

0

0

0

(Total)

(1)

(9)

(4)

(12)

(1)

(12)

(0)

*One of these males had assymetrical external valuae.
pharate = adults dying inside the pupal case with wings formed.

26(l-4):l-288, 1988

41

Table 3. Progeny (1983 reared) of pairing #628 ( P . g. glaucus dark morph female x P. g. canadensis male; see text
for further details).

Eclosion

Bilateral

Males

Females

Dead

Pupae

year

Gynandromorph

eclosed

pharate1

eclosed

pharate1

pupae

still alive

1983

0

13

0

0

0

9

-

1984

1

25

0

4

0

10

-

1985

0

0

0

13*

0

3

-

(Totals)

(1)

(38)

(0)

(17)

(0)

(22)

(20)

*One of these females is a “yellow intermediate” and one is a “dark intermediate”, between the two typical color
morphs while all other females are typical yellow morphs (see also Scriber et al, 1987).

The bilateral gynandromorphs in this collection alone include 49
Pieridae, 70 Nymphalidae, 6 Hesperidae, 20 Lycaenidae, 1 Danaidae,
and 11 Papilionidae (S. Borkin and A. Young, pers. comm.).

It has been suggested that either viral diseases (Gardiner, 1972; Blau,
1978; Sevastopulo, 1973) or parasitism, and/or abnormal temperatures
could be responsible for inducing such abnormalities (Riotte, 1978). It
has also been observed that bilateral gynandromorphs and mosaics can
occur in multiples from the same brood (Cockayne, 1935; Ford, 1955;
Sevastopulo, 1973) and that a variety of laboratory hybrid crosses have
yielded intersexual or gynandromorphic individuals (Standfuss, 1900;
Whicher, 1915; Cockayne, 1935; Clarke and Sheppard, 1953, 1960;
Clarke et al., 1977; Clarke and Ford, 1980; Platt, 1983).

Sir Cyril Clarke and colleagues have been investigating the genetic
basis of abnormal wing coloration in Papilio glaucus for decades (see
Clarke and Clarke, 1983 for a review), and they point out that color
mosaics and gynandromorphs are sometimes strikingly visible in P.
glaucus because of the marked differences between the yellow back¬
ground of males and yellow morph females and the black/brown
background of dark morph females. The Herman Strecker collection
(currently on loan from the Chicago Field Museum to the Allyn Museum
in Florida) contains a number of such mosaics. This valuable collection,
which was assembled during the second half of the 19th century,
contains a number of P. glaucus mosaics previously described and/or
reported upon the literature (e.g. Strecker, 1878; Ehrmann, 1894;
Walsten, 1977; Ehle, 1981; Shapiro, 1981b). Edwards (1884) also figures
an individual with V2 black and V2 yellow which he describes as a female
(Edwards, 1868). Clarke and Clarke (1983) figure similar specimens
(essentially half yellow/half dark) from the Strecker collection, one of
which is a color mosaic female (from Indiana) and the other an apparent
gynandromorph from Pennsylvania.

Partial color (sex?) mosaics in Papilio glaucus are also rare, but have
been collected in Pennsylvania and previously described by Strecker
(1878), and figured by Walsten (1977), Ehle (1981), and Clarke and
Clarke (1983). In addition to the partial color mosaics we report here
(see also Scriber et al, 1987a), additional cases for Papilio glaucus are

42

J. Res. Lepid.

known to exist (e.g. personal collections of James Sternberg and David
Ritland). The Milwaukee Public Museum contains two partial mosaics
of Papilio glaucus reared by E. Dluhy in Chicago, Illinois. It is intri¬
guing that the few cases of female coloration patterns unlike their
mother’s which Clarke and Sheppard (1962; see also Clarke et al., 1976)
encountered in their studies were all tracable to a group of pupae
obtained from E. Dluhy (from Chicago, Illinois). It was suggested
(Scriber et al, 1987a) that similar chromosomally abnormal stock might
be involved in both the color mosaics and the abnormal segregation of
dark and/or yellow female forms of Papilio glaucus. We now feel that an
explanation for this abnormality may involve introgression between
P. g. canadensis and P. g. glaucus subspecies near the “blend zone” in
Wisconsin (Scriber 1982, 1983) and across the Great Lakes region
(Scriber et al., 1987b). In this paper we shall report aspects of our
laboratory studies with members of the North American tiger swal¬
lowtail, Papilio glaucus L., group.

Methods

In the past 6 years we have reared through to the adult stage or collected in
the field (from southern Florida to northern Canada) over 28,000 specimens of
the Papilio glaucus species complex. Field captured specimens have included all
three P. glaucus subspecies as well as P. eurymedon, P. rutulus, P. multicau-
datus, and P. alexiares garcia. Lab reared specimens have included pure stock of
all of these species and subspecies as well as geographic site crosses within
subspecies, subspecies-level and species-level crosses.

Oviposition by adult females is induced by placing each wild-captured or
hand-paired female into its own clear plastic box (approximately 10 cm deep x
15 cm x 30 cm) with a moist paper towel and selected larval foodplant leaves.
Leaf turgor was maintained in these plants by use of floral aquapics® (water-
filled plastic vials with a rubber cap, through which leaf petioles or small
branches can be inserted; see Scriber, 1977). Heat and light were provided by
incandescent bulbs placed at a distance of approximately 0.3-0. 5 meter from the
plastic boxes.

Larvae were reared to pupation on one of various foodplants, (leaves were
changed three times per week) under controlled environment conditions (16:8
photo-scoto-phase with corresponding temperatures of 23°/19°C, respectively).
Pupae were weighed 2 days after pupation (the weight subsequently serving an
identification number for the individual) and then placed in individual cylin
drical screen cages (15 cm diameter x 12 cm height) under larval rearing
conditions or similar laboratory conditions (21-24°C) to permit development
and eclosion as adults. Direct developing individuals normally emerged within
2-3 weeks after pupation. Other pupae were given at least 6 weeks before being
refrigerated in darkness (at 40-45°F for 3 months or more) to break diapause.
Hand-pairings were generally attempted 12-48 hrs after adult female eclosion
and 2-3 days after male eclosion. Since dark female color suppression and
obligate diapause are linked on the X-chromosome (R. Hagen and J.M. Scriber,
manuscript), we observe one year delayed emergence of F, females from both
yellow (Table 1) and dark (Table 2) females when maled with P.g. conadensis
males.

26(l-4):l-288, 1988

43

Table 4. Gynandromorphs, mosaics, and their siblings (1981-1986). a,m

Siblings in the Brood

Brood #

Rearing

Year

Gynand.

Mosaics

Fig. #

Males

Yel

Dk

Pupae (as of

7 Nov. 86)

Dead

Pupae

9

82

1

1

13b

13

0

0

12

628

83

1

2

38

17c

0

20d

22

631

83

1

10

6

6

0

2

3

1091

84

1

4

2

0

1

0

0

2025

84

1

9

1

2

0

0

0

2830

85

1

7

40

0

0

22

3

3622e

86

1

6

8

0

18

9

0

3935

86

1

8

6

0

11

13

0

4196e

86

1

5

26

1

10

20

0

4210

86

1

3

31

2

27

4

3

JMS

81

1

11

5

0

12"

0

0

658

83

1

14

7

0

10

1

0

688

83

14

20

22f

0

24g

34h

6

717

83

1

29

30

0

34

0

1

1064

84

1

12

9

0

11

0

1

1128

84

1

15

16

0

24‘

0

1

1348

84

1

28

22

18s

0

2

0

1351

84

1

27

52

37j,k

1

24

7

1534

84

1

26

0

0

0

0

0

1905

84

1

X

23

0

31

3

7

1914

84

1

23

42

0

41

14

2

1999

84

1

13

9

0

6

0

1

2030

84

1

X

7

0

2

0

0

3122

85

1

X

1

0

0

0

0

3604

86

1

24

11

0

15

18

0

3770

86

1

25

34

0

31

28

0

3800

86

1

17

29

0

28

19

1

3973

86

1

16

19

0

15

6

0

4230

86

1

30

8

2

8

1

0

Subtotals:

Total Reared to
pupae = 1282

10

32

512

98

320

240

70

“Pharate adults are included with “emerged adults”.
bOne male has asymmetrical valvae.

cOne female is a “yellow intermediate” and one is a “dark intermediate”.

dTwo of these pupae are unaccounted for as of 1984.

eAn additional emerged adult has no sex recorded on printout.

fOne male has very reduced claspers and one male has extra dark scaling.

H’wo females are slight intermediates and one female has extra yellow and unusually high density blue.
hApproximately 6 weeks after pupation, these were refrigerated to break diapause and not brought out of the
refrigerator the first time until 14 mos. later. Although they still appear viable, it is uncertain if they will eclose in
the future.

‘One female is a dark intermediate.

JSome of these may be “yellow intermediates” but did not get recorded as such. They are not all pinned now and
colors cannot be easily verified.

kFour females are “yellow intermediates” (i.e., more yellow than dark).

'Two females are “dark intermediates” (i.e., more dark than yellow).

mWild collected mosaics (see Table 3) are excluded from this table as they have no sibling data.

"Two females were “dark intermediates”.

44 J. Res. Lepid.

Table 5. Phenotype, geographic origin, and brood number of gynan-
dromorphs.

Gynandromorphs

Brood # b

Phenotypes &/or
geographic origins3

Perfect

Bilateral

< 50%

one sex

generating

aberrant

Georgia

1

2025

Ohio

3

631

3622

3935

Texas

1

1091

Pa Yel x Pgc (Juneau)

1

9

S. Car Dk x Pgc (Marinette)

1

628

Ill. Dk x Fi (GaDk x Pgc)

1

4196

W. Va. Dk x P. rutulus

1

2830

Ohio Dk x P. alexiares

1

4210

Totals

3

7

aWhen phenotype or geographic site crosses are listed, the female
background is listed first.

bFor information on siblings of aberrants, look up brood numbers on
table #1.

Results

Of the 28,000 reared adults we have observed 10 gynandromorphs
and 32 color mosaics (Tables 4 and 5). Of the obvious gynandromorphs
four of the five perfect or near perfect bilateral gynandromorphs (Figs.
1-5) were progeny of subspecies of species crosses. The fifth could also be
considered a subspecies cross if the Texas population is P. g. australis
(Scriber, 1986). Of the remaining five gynandromorphs (Figs. 6-10) only
one (Fig. 7) involves two different taxa. Of approximately 8,500 field
collected specimens during 1981-1986, we have never collected an
obvious gynandromorph.

Of the 32 color mosaics listed in Table 4, all but 6 are of either pure P.
glaucus glaucus or P. g. australis (Table 6). These mosaics are dark
morph females with varying amounts of yellow dorsally and/or ven-
trally on their wings or body. Of the 37 total P. glaucus mosaics, one was
a field collected P. g. australis from Highland Co., Florida (Fig. 22) and
two were field collected P. g. glaucus from Dane Co., Wisconsin (Figs. 18

26(l-4):l-288, 1988

45

Table 6. Phenotype, geographic origin, and brood number of color mosaics.

Gynandromorphs

Dk w /

Yel w/

Brood # b

Phenotypes &/or

exceptional

exceptional

generating

geographic origins3

color yellow

color black

aberrant

Georgia

3

1064

2030

3122

Illinois

2

1905

1999

Ohio

4

658

1128

3800

3973

Texas

2

1039 (wild)
3270 (wild)

Wisconsin

17c

JMS

688

729 (wild)
736 (wild)

Georgia x Ohio

1

3604

(Ga. x Wis.) x Ill.

1

1914

P. glaucus australis

1

no # (wild)

Ga. Dk x Pgc (Green Lake)

1

1348

Ga. DK x Pgc (Tompkins)

le

3770

Tx Dk x Pgc (Juneau)

1

1534

Tx Dk x Pgc (Wood)

1

1351

Bx (Fx (GaDk x Pgc) x Wis.) x Pga

1

717

Ill Dk x P. alexiares

1

4230

Totals

34f

3

aWhen phenotype or geographic site crosses are listed, the female background is
listed first.

bFor information on siblings of aberrants, look up brood numbers on table #1.
cOf these 17 mosaics, 14 were siblings in brood #688.

dCounties of specimen origin are shown in parentheses and all are Wisconsin
except Tompkins which is New York.

‘This is actually an intermediately colored specimen but the exceptional color is
yellow.

fOf the 34 dominantly dark color mosaics listed, 27 are shown in the following
illustrations. (All other aberrants in this table are illustrated).

46

J. Res. Lepid.

W\Y fr

4%

Igl > W

JNL &

i

, Wl

/

J} v 1 JkI'I

26(l-4):l-288, 1988

47

48

J. Res. Lepid.

Fig. 1. A bilateral gynandromorph (from brood #9, pupal wt. 1.0605, ex ova,
reared in 1982, eclosed on 21 May 1983) from a lab cross of a yellow morph P. g.
glaucus female (2nd generation lab-reared from wild stock collected by W. Houtz in
Schuylkill Co., Pennsylvania) mated to a wild collected male P. g. canadensis from
Juneau Co. Wisconsin. A) dorsal, B) ventral.

Fig. 2. A bilateral gynandromorph (from brood #628, pupal wt. 0.9310, ex. ova,
reared in 1983, eclosed on 30 May 1984) from a lab cross of a dark morph P. g.
glaucus female (reared from eggs obtained from a wild dark morph female collected
by R. Peigler in Pickens Co., South Carolina) mated to wild collected male P. g.
canadensis from Marinette Co., Wisconsin (collected 1-5 July 1983 by Don Caine).

A) dorsal, B) ventral.

Fig. 3. A bilateral gynandromorph (from brood #4210, pupal wt. 1 .1099, ex. ova
reared in 1986, eclosed on 26 Sept. 1986) from a lab cross of a dark morph P. g.
glaucus female (2nd generation lab reared from wild stock collected by J. Thorne
and MHE in July 1985 in Adams Co., Ohio) mated to a wild collected male P.
alexiares garcia from Nuevo Leon, Mexico (collected 2, 3 Aug. 1 986 by W. Warfield,
D. Robacker and MHE). A) dorsal, B) ventral.

Fig. 4. A nearly perfect bilateral gynandromorph, or sexual mosaic, (from brood
#1091 , pupal wt. 1 .1493, ex. ova, reared in 1 984, eclosed on 27 June 1 984) from a
wild collected dark morph P. g. glaucus female from Jasper Co., Texas (collected on
9 April 1984 by JMS and MHE). A) dorsal, B) ventral.

Fig. 5. A gynandromorph, or sexual mosaic, which appears to be more than 50%
female (from brood #4196, pupal wt. 1.2779, ex. ova, reared in 1986, eclosed on
29 Sept. 1 986) from a lab cross of a dark morph P. g. glaucus female (reared from
eggs obtained from a wild dark morph female collected by M. Berenbaum in
Champaign Co., Illinois in June 1986) mated to a subspecies hybrid male (whose
mother was the daughter of a dark morph P. g. glaucus female collected in Georgia
in Aug. 1985 by J. Maudsley and whose father was a P. g. canadensis collected in
Lincoln Co., Wisconsin on 3 June 1 986 by D. Ware, V. Viegut and MHE). A) dorsal,

B) ventral.

Fig. 6. A gynandromorph, or sexual mosaic, which appears to be more than 60%
female (from brood #3622, pupal wt. 1.2866, ex. ova, reared in 1986, eclosed on
12 Aug. 1986) from a pure P. g. glaucus lineage (in which the mother was the
daughter of a dark mQrph female collected in May 1986 in Hocking Co., Ohio by S.
Stribling and the father was the son of a dark morph female collected in July 1 985 in
Adams Co., Ohio by J. Thorne and MHE). A) dorsal, B) ventral.

Fig. 7. A gynandromorph, or sexual mosaic, which appears to be more than 60%
male (from brood #2830, pupal wt. 1.1341, ex. ova, reared in 1985, eclosed on 1
June 1 986) from a dark morph P. g. glaucus female (reared from eggs obtained from
a dark morph female wild collected in Clay Co., W. Virginia on 12 July 1984 by W.
Warfield and MHE) mated to a wild collected P. rutulus male (reared from eggs
obtained from a wild female collected by R. Dowell in Sacramento Co., CA). A)
dorsal, B) ventral.

Fig. 8. A gynandromorph, or sexual mosaic, which appears to be more than 75%
female but with male claspers (from brood #3935, pupal wt. 1 .5321 , ex. ova, reared
in 1986, eclosed 14 Sept. 1986) from a wild collected dark morph P. g. glaucus
female (collected in Scioto Co., Ohio on 4 July 1986 by J. Thorne and MHE). A)
dorsal, B) ventral.

Fig. 9. A gynandromorph which appears to be more than 80% male (from brood
#2025, pupal wt. 1 .1541 , ex. ova, reared in 1 984, eclosed on 1 2 May 1 985) from a
yellow morph P. g. glaucus female wild collected in Clarke Co., Georgia (collected
on 29 Aug. 1984 by J. Maudsley). A) dorsal, B) ventral.

Fig. 1 0. A gynandromorph which appears to be more than 90% male (from brood
#631, pupal wt. 0.9980, ex. ova, reared in 1983, eclosed on 25 Oct. 1984, was
paired to 2054 but produced no progeny) from a yellow morph P. g. glaucus female
wild collected in Adams Co., Ohio (collected on 9 July 1983 by W. Warfield and

26(l-4):l-288, 1988

49

MHE). A) dorsal, B) ventral.

Fig. 11. A female color mosaic (ex. ova, reared in 1 981 by D. Ritland and J. M. S.,
eclosed Aug. 1981) from a normal appearing dark morph P. g. glaucus female
(collected in Dane Co., Wisconsin in June 1981 by P. Kingsley and D. Ritland). A)
dorsal, B) ventral.

Fig. 12. A female color mosaic (from brood#1064, pupal wt. 1.2526, ex. ova,
reared in 1 984, eclosed on 9 July 1 984) from a wild dark morph P. g. glaucus female
(collected in Oglethorpe Co., Georgia on 1 4 Apr. 1 984 by J. Maudsley). A) dorsal, B)
ventral.

Fig. 13. A female color mosaic (from brood #1999, pupal wt. 1.2960, ex. ova,
reared in 1984, eclosed on 10 May 1985) from a wild dark morph P. g. glaucus
female (collected in Rock Island Co., Illinois on 25 Aug. 1984 by W. Warfield).

A) dorsal, B) ventral.

Fig. 1 4. A female color mosaic (from brood #658, pupal wt. 0.9439, ex. ova, reared
in 1983, eclosed 2 June 1984) from a wild dark morph P. g. glaucus female
(collected in Adams Co., Ohio on 9 July 1983 by W. Warfield and MHE). A) dorsal,

B) ventral.

Fig. 15. A female color mosaic (from brood #1128, pupal wt. 1.5200, ex. ova,
reared in 1 984, eclosed 1 2 July 1 984) from lab paired parents which were propgeny
of two wild dark morph P. g. glaucus females (both collected in Adams Co. , Ohio and
9 July 1983 by W. Warfield and MHE). A) dorsal, B) ventral.

Fig. 16. A female color mosaic (from brood #3973, pupal wt. 1.1323, ex. ova,
reared in 1986, eclosed 18 Sept., 1986) from a wild dark morph P. g. glaucus
female (collected in Adams Co., Ohio on 5 July 1986 by J. Thorne and MHE). A)
dorsal, B) ventral.

Fig. 17. A female color mosaic (from brood #3800, pupal wt. 1.4466, ex. ova,
reared in 1986, eclosed 7 Oct. 1986) from a wild dark morph P. g. glaucus female
(collected in Adams Co., Ohio on 6 July 1986 by J. Thorne, and MHE). A) dorsal, B)
ventral.

Fig. 18. A wild collected P. g. glaucus female color mosaic (assigned #729 and
set up for oviposition but died laying only one infertile egg: collected in Dane Co.,
Wisconsin on 7 Aug. 1983 by MHE). A) dorsal, B) ventral.

Fig. 19. A wild collected P. g. glaucus female color mosaic (assigned #736 and
set up for oviposition but died laying no eggs: collected in Dane Co., Wisconsin on 7
Aug. 1983 by W. Warfield). A) dorsal, B) ventral.

Fig. 20. Fourteen female siblings with various color mosaic patterns were
generated from one brood (#688, ex. ova, reared in 1 983) from a normal appearing
wild dark morph P. g. glaucus female (collected in Dane Co., Wisconsin on 1 Aug.
1983 by W. Warfield). Nine of those female siblings are shown here. A) dorsal, B)
ventral.

eclosed 25 Oct.

" 29 May

" 30 May

" 25 Oct.

" 25 Oct.

1 June
" 25 Oct.

" 25 Oct.

" 31 May

Pupal wt. 1.2253

" wt. 0.9834

" wt. 1.1072

" wt.

" wt.

" wt.

" wt.

" wt.

1 .2296
1.1742
1 .3444
1 .2375
1 .2555

wt. 0.9885

1984

A)

B)

C)

D)

E)

F)

G)

H)

I)

Fig. 21 . A wild collected P. g. glaucus female color mosaic (assigned #1039 and
set up for oviposition: she laid 5 eggs which produced 2 larvae. One reached the
adult stage as a normal appearing male which was not mated.) wild collected (in
Jasper Co., Texas in April 1984 by JMS and MHE). A) dorsal, B) ventral.

Fig. 22. A wild collected P. g. australis female color mosaic (collected in Highlands
Co., Florida in Apr. 1981 by B. Giebink, JMS and MHE). A) dorsal, B) ventral.
Fig. 23. A female color mosaic (from brood #1914, pupal wt. 0.9309, ex. ova,

50

J. Res. Lepid.

26(l-4):l-288, 1988

51

reared in 1 984, eclosed 28 Oct. 1 984) from a dark morph. P. g. glaucus female (the
daughter of a lab pairing of a dark morph P. g. glaucus female reared from stock
collected in Oglethorpe Co., Georgia in April 1 984 by J. Maudsley, mated to a wild P.
g. glaucus male collected in Richland Co., Wisconsin on 7 June 1984 by S. Sippl,
and JMS) lab mated to a wild P. g. glaucus male (collected in Rock Island Co.,
Illinois on 11 Aug. 1984 by W. Warfield). A) dorsal, B) ventral.

Fig. 24. A female color mosaic (from brood #3604, pupal wt. 1.4616, ex. ova,
reared in 1986, eclosed 4 Aug. 1986) from a dark morph P. g. glaucus female (the
daughter of a wild dark morph P. g. glaucus female, #31 45 collected in Habersham
Co., Georgia in Aug. 1985 by J. Maudsley) lab mated to a P. g. glaucus male (the
son of a wild dark morph P. g. glaucus female, #2868, collected in Adams Co., Ohio
on 9 July 1985 by J. Thorne and MHE).

Fig. 25. A female color mosaic (from brood #3770, pupal wt. 1 .7029, ex. ova,
reared in 1 986, eclosed 25 Aug. 1 986) from a dark morph P. g. glaucus female (the
daughter of a wild dark morph P. g. glaucus female, #31 04, collected in Athens Co.,
Georgia in Aug. 1985 by A) lab mated to a wild P. g. canadensis male (collected in
Tompkins Co., N.Y. by R. Lederhouse on 17 June 1986). A) dorsal, B) ventral.
Fig. 26. A female color mosaic (from brood #1534, pupal wt. 13389, ex. ova,
reared in 1984, eclosed 3 Aug. 1985) from a dark morph P. g. glaucus female (the
daughter of a wild dark morph P. g. glaucus female, #1038, collected in Jasper Co.,
Texas on 9 April 1 984 by JMS and MHE) lab mated to a wild P. g. canadensis male
(collected in Juneau Co., Wisconsin on 1 8 June 1 984 by W. Warfield). A) dorsal, B)
ventral.

Fig. 27. A female color mosaic (from brood #1351, pupal wt. 1.2429, ex. ova,
reared in 1 984, eclosed 1 4 May 1 985) from a dark morph P. g. glaucus female (the
daughter of a wild dark morph P. g. glaucus female, #1025, collected in Newton
Co., Texas in April 1984 by JMS and MHE) lab mated to a wild P. g. canadensis
male (collected in Wood Co., Wisconsin on 12 June 1984 by C. Plazk, Y. Allen, K.
Hale, and W. Warfield). A) dorsal, B) ventral.

Fig. 28. A femal color mosaic (from brood if 1348, pupal wt. 1.1453, ex. ova,
reared in 1984, eclosed 1 July 1986) from a dark morph P. g. glaucus female (the
daughter of a wild dark morph P. g. glaucus female with yellow discal cells, #1231 ,
collected in Oglethorpe Co., Georgia in May 1984 by J. Maudsley) lab mated to a
wild P. g. canadensis male (collected in Green Lake Co., Wisconsin on 12 June
1984 by J. Thorne, S. Sippl, and MHE). A) dorsal, B) ventral.

Fig. 29. A female color mosaic (from brood #71 7, pupal wt. 1 .3006, ex. ova, reared
in 1983, eclosed 26 Sept. 1983) from a ‘‘peppered ’’-colored yellow female (from
brood #558; a dark morph female GA Pgg x Pgc, backcrossed to a Pgg from Wise.)
mated to a wild P. g. australis male (collected in Highlands Co., Florida on 3 Aug.
1983 by JMS). A) dorsal, B) ventral.

Fig. 30. A female color mosaic (from brood #4230, pupal wt. 1 .0948, ex. ova,
reared in 1986, eclosed 4 Oct. 1986) from a dark morph P. g. glaucus female (the
daughter of a dark morph P. g. glaucus female, #3540, wild collected in Champaign
Co., Illinois in June 1986 by M. Berenbaum) lab mated to a wild P. alexiares male
(collected in Nuevo Leon, Mexico 2-3 Aug. 1986 by W. Warfield, D. Robacker, and
MHE). A) dorsal, B) ventral.

52

J. Res. Lepid.

and 19). Fourteen other P. g. glaucus mosaics are all siblings from one
wild collected female from Dane Co., Wisconsin (e.g., Fig. 20). Two field
captured females from Texas were mosaics (Table 6; Fig. 21).

Of the six lab-reared mosaics that were not of “pure” subspecies
lineage four are progeny of subspecies crosses between P. g. glaucus
females and P. g. canadensis males (Table 6). Three of these are yellow
females with the exceptional and asymmetrical color being black (Figs.
26-28) while the fourth is an intermediately colored female with the
exceptional color being yellow (Fig. 25). The fifth non-pure P. g. glaucus
mosaic is the product of a lab pairing involving all three P. glaucus
subspecies as ancestors. The mosaic is 50% P. g. australis , 12.5% P. g.
canadensis , and 37.5% P. g. glaucus (Fig. 29). A sixth mosaic arose from
a P. g. glaucus paired with a male P. alexiares garcia (Fig. 30).

Discussion

The reported frequency of color mosaics or gynandromorphs is quite
low. We have reared more than 28,000 individuals of the Papilio glaucus
complex (1981-1986) and have observed only 37 color mosaics and only 5
perfect or nearly perfect bilateral gynandromorphs. While from previous
literature records it would seem that the “blend zone” (i.e. the zone of
potential/probable hybridization) across the Great Lakes and Appala¬
chian Mountain region has accounted for most previously reported
aberrant types of P. glaucus (Edwards, 1868, 1884; Skinner and Aaron,
1888; Ehrman, 1893; Howard, 1899; Clark and Clark, 1951; Ehle, 1981;
Scriber et al, 1985; H. Romack, pers. comm.), we have in this 5-year
period observed mosaics in stock from Georgia, Texas, Illinois and Ohio
as well as Wisconsin. In addition to our Dane County field-captured
mosaics (near the zone of suspected hybridization), color mosaics exist
from Richmond County, NY (5 July 1971; A.M. Shapiro; currently in the
Univeristy of California-Davis Collection) and Washington County, PA
(9 May 1927; George F. Patterson Collection at Pennsylvania State
University), both also near the proposed blend zone (see Ritland and
Scriber, 1985; Scriber and Hainze, 1987).

Since in our studies, 4 of 5 perfect or nearly perfect bilateral gynan¬
dromorphs are progeny of subspecies or species crosses in the P. glaucus
species complex, and since 32 of 37 mosaics are progency of “pure” (i.e.
single subspecies) lines, it could be argued that factors inducing the
expression of gynandromorphic traits are likely to be different than
factors inducing the expression of mosaic traits. In fact, for the occurence
of mosaics the observed distribution between inter-taxa versus intra-
taxa crosses does not differ from the expected (n = 5105, Chi square, p =
0.25), whereas for the near perfect bilateral gynandromorphs signifi¬
cantly more resulted from the inter-taxa crosses (n = 11, 112, Fisher
Exact test, p < 0.038).

The mechanism(s) by which hybridization may catalyze these events
leading to color/sexual abnormalities is uncertain at this time. Clarke

26(l-4):l-288, 1988

53

and Clarke (1983; see also Scott, 1986) suggest that in P. glaucus
maleness is dependent upon the presence of two X chromosomes and
femaleness is dependent upon a single X chromosome (the Y chromo¬
some assumed to be relatively inert, except of course that it carries the
locus for dark morph color; Clarke and Sheppard, 1962; Clarke et al.,

1976) . If this is true (cf. Tazima, 1964; where femaleness in silk moths
may be related to the Y chromosome), then the P. glaucus sexual
mosaics in the Strecker collection may represent the results of either
non-disjunction or double fertilization (Clarke and Clarke, 1983). Three
specimens are figured by Clarke and Clarke: 1) The female half
black/half yellow may be the result of non-disjunction, i.e. XY/XO with
the gene for melanism on the Y and the XO half being yellow; 2) the
gynandromorph which is approximately half black (female) and half
yellow (male) may be the result of a double fertilization (i.e. XX/XY);
and 3) a sexual mosaic specimen (a partial mosaic described by Howard,
1899) could be the result of XX/XY (non-disjunction) or XY/XO (double
fertilization). While Clarke and Clarke were not aware of the potential
hybridization ofP. g. glaucus withP. g. canadensis , we now suggest with
our laboratory results that hybrid individuals in the “blend zone” may
indeed be naturally predisposed to non-disjunction or chromosome loss
as appears to be the case with other Papilio hybrids (see Clarke et al.,

1977) . In fact, we also have evidence suggesting the loss of a segment of
the Y-chromosome controlling the melanic (dark) background color¬
ation in brood number 674 reared in 1983 (with transfer of this
chromosome fragment independently, via a crossover or via a non¬
disjunction) with the melanic locus to at least one of the male sibs!). This
aberrant male, when mated to a virgin P. g. canadensis from northern
Wisconsin and to a yellow morph P. g. glaucus female from a yellow
morph lineage from Ohio, produced daughter progeny which were both
typical dark morph and typical yellow morph (Scriber et al., 1987).
Never before has any male P. glaucus been known to transmit the trait
for female melanism (see Scriber and Evans, 1986).

The ability to distinguish between color mosaics and sexual mosaics is
not easy, even with the distinctive pattern of the melanic form female. It
is even more difficult to distinguish between yellow morph gynandro-
morphs (where the male and female regions differ from each other both
phenotypically and in genetic constitution) and intersexes (where male
and female regions appear phenotypically different, but have identical
chromosomal constitution: see Whiting et al., 1934; Doutt and Smith,
1950; Clarke and Ford, 1980). We hope that additional studies will
clarify the genetic basis of gynandromorph production (see reviews by
Cockayne, 1935; Drescher and Rothenbuhler, 1963), and of the differ¬
ential suppression (modification) of female phenotypes dark/yellow
polymorphism in Papilio glaucus due to hybridization. Some morpho¬
logical characteristics differ between the male and female halves of the
F j specimens figured (Fig. 1 and Fig. 2) as well as the obvious differences
in melanism in Fig. 3. The usefulness of developmental physiology

54

J. Res. Lepid.

studies in this regard would be significant, especially since the color
patterns on the upper and lower surfaces of Lepidoptera wings develop
independently (see Nijhout, 1981 for a discussion), and can be modified
by environmental temperature (Ritland, 1983).

Acknowledgements. This research was supported by the National Science
Foundation (DEB #7921749, BSR #8306060, and BSR #8503464), the USDA
(#85CRCR-1-1598), and by the Graduate School and the College of Agriculture
and Life Sciences (Hatch Project 5134) at the University of Wisconsin, Madison.
In these studies and our discussions we are especially grateful to: Thomas Allen,
Yvonne Allen, Matt Ayres, Penny Barker, May Berenbaum, William Bergman,
Susan Borkin, Don Caine, Sir Cyril Clarke, Michael Collins, Robert Dowell,
Robert Hagen, Kathy Hale, Susan Heg, Sue Helgeson, William Houtz, Judy
Johnson, K.C. Kim, Robert Lederhouse, Heidi Luebke, Dave Matusik, Jim
Maudsley, Jacqueline Miller, Lee Miller, Ric Peigler, Jeanne Pomraning,
Christina Plzak, Carrie Richards, David Ritland, Dave Robacker, Sarah Rockey,
Howard Romack, Jane Schrimpf, Brian and Bradley Scriber, Art Shapiro, John
Shuey, Jane Siebenhorn, Shawn Sippl, Sam Skinner, Jim Sternberg, Donald
Strasburg, Sam Stribling, Jeff Thorne, David Ware, Vicki Ware (Viegut), Bill
Warfield, Gilbert Waldbauer, and Allen Young. We are also thankful for
Jacqueline and Lee Miller for their assistance at the Allyn Museum in
examining the Herman Strecker Collection mosaics and gynandromorphs.

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Journal of Research on the Lepidoptera

26(l-4):58-63, 1988

Opposition on Peripheral Hosts by Dispersing Pieris
napi (L.) (Pieridae)

Steven P. Courtney

Department of Biology, University of Oregon, Eugene, Oregon 97403

Abstract. Females of Pieris napi dispersing through an unfavourable
habitat were able to detect Cardamine flexuosa L. individuals hidden
from sight. Such peripheral hostplants receive more P. napi eggs than
conspecifics at the population centre; this is partly due to females laying
several eggs at a single visit, apparently a tactic adopted in response to
low hostplant density. Inverse density-dependent host selection may
have important effects on hostplant communities, such as increased
herbivore pressure on peripheral host species.

Introduction

Several recent investigations have reported that individuals within a
butterfly population may adopt different reproductive tactics in re¬
sponse to environmental variation (Wiklund, 1981; Tabashnik et al.,
1981). For instance, females experiencing low availability of foodplants
may respond by ovipositing more readily upon suboptimal hosts (Singer,
1982, 1984) or by increasing the number of eggs deposited on a host
(Parker and Courtney, 1984). Further observations are badly needed to
document the factors which influence such changes in behaviour.
However, it is normally hard to make extensive observations on females
in marginal habitats, where the insects are, by definition, scarce. I report
here observations on Pieris napi (L.) at Delamere Forest, Cheshire,
England, where individual females were regularly watched during July
and August 1982, as they entered apparently unfavourable habitats
during dispersal from population centers. Several unusual behaviours
were seen, and are described here because of their importance to
population and community processes.

Observations and Results

Two population centres for C. flexuosa were found in wet meadows to
the north-east of Delamere. Nasturtium officinale R. B and Cardamine
pratensis L. (both Cruciferae) were the predominant hostplants, with
some C. flexuosa. Both populations bordered a large coniferous planta¬
tion which contained felled areas where Bracken ( Pteridium aquilinum
L.) and Bramble (Rubus fructicosus L. agg.) grew densely. 600 m of trees
separated the two marshy areas where, in July-August, the second
brood of P. napi was in good numbers, making short flights among the

26(l-4):l-288, 1988

59

Table 1. Individuals of P. napi observed in two areas of Delamere Forest on
28.7. and 4.8.1982. Comparison of the main population (marsh) with
dispersing individuals (heath) yields a significant difference in
sex-ratio (X2 = 5.906 p< 0.025).

Marsh Heath

P. napi Males 32 17

Females 8 15

vegetation. A few butterflies were also seen in the felled areas of the
plantation, flying rapidly and directly across the apparently unsuitable
habitat. The sex ratios of individuals in the marsh and heathland areas
were significantly different: proportionately more females were seen
away from the main population areas (Table 1). This difference probably
arises more from differential conspicuousness of the two sexes in the
main population areas (Gilbert and Singer, 1975) than from differential
dispersal of the sexes (e.g., Shapiro, 1970). No differences were found in
spermatophore counts of five females from either habitat; all females
had mated once.

On 28th July, two females were observed in the felled area in
characteristic “oviposition search” style of flight, which is adopted in the
vicinity of hostplants (Chew, 1975; Wiklund and Ahrberg, 1978). Close
inspection showed that these females had located a small group of C.
flexuosa plants, growing in an old, moist ditch underneath the Bracken.
These hostplants were located 900 m away from each of the two marshy
areas. The hostplants were largely hidden from sight below the Bracken,
and subsequent observation on other P. napi individuals confirmed that
females did not visually detect the C. flexuosa patch. Butterflies were, on
several occasions, seen to switch, from rapid, direct flight to slow,
host-location flight, when in the vicinity of completely hidden C. flex¬
uosa. Olfactory stimulation of searching behaviour of Pierinae has long
been suspected; antennal chemoreceptors are known to respond to
glucosinolates, a major family of phytochemicals in Cruciferae (Den
Otter et al., 1980). Field observations of such behaviour have been
lacking until now. No females were observed to search in any other felled
or wooded area, and an extensive search revealed only two more C.
flexuosa plants in the wood; one plant bore four P. napi eggs.

On adopting the ‘oviposition search’ mode of flight, the female would
investigate the ground layer of vegetation, approaching and occasionally
landing upon small herbs. Females were sometimes seen deep under¬
neath the Bracken, flying among the fronds in their search for hosts. On
locating a host, a female would flutter around it, settle on a leaf and,
bending the abdomen underneath, deposit an egg. On two occasions (of
32 observed ovipositions) a second egg was immediately deposited. On
five other occasions the female flew up and returned to the same plant to
lay a second egg. Multiple oviposition by Pieris rapae L is well known,
but P. napi has rarely been seen to lay more than one egg at a host (F.S.
Chew, pers. comm.). In this sub-population of dispersing individuals,

60

J.Res.Lepid.

Table 2. Average height (cm.) and leaflet number (with standard error) of C.

f/exuosa individuals growing in four categories of shading by
Bracken, with the average eggload of P. napi on such plants
(Number of eggs/Number of leaflets). Data from the marsh popula¬
tion are given for comparison. All data collected 28.7. 1982.

Shading

n Height

Leaflets

Eggload

Heath

<25%

7 9.3

5.29(1.82)

1.16

25-50%

15 9.0

3.87(1.10)

1.12

51-75%

14 15.0

4.86(0.90)

0.32

>75%

26 18.1

11.69(2.61)

0.27

Marsh

35 14.1

9.20(1.31)

0.12

multiple oviposition seemed frequent, though it was never seen in the
main marshy populations.

Numerous eggs were found on the 62 C. flexuosa individuals in the
ditch. Table 2 presents these data for four groupings of plants according
to the degree of overtopping by Bracken. Seven individuals had less than
25% of the sky above obscured by Bracken (estimated by eye); these
plants received a very heavy load of P. napi eggs, with over one egg per
leaflet (far more than could be supported through larval development).
The majority of C. flexuosa were more shaded and 21 (41.9%) were
almost completely obscured by Bracken. Relatively few eggs were found
on heavily shaded plants, although these were the largest. Chew (1977)
reports that shaded individuals of Nearctic Cardamine similarly escape
P. napi oviposition. These and similar results in Anthocharis cardamines
L. have been interpreted as consequences of females restricting their
activity to areas of direct sunshine (Courtney, 1982). Note that the
eggloads, of even the heavily shaded C. flexuosa plants in the ditch, are
much heavier than those seen in the marshes where C. flexuosa was
more plentiful. Figure 1 illustrates the combined effects of shading
levels, and of host size, which also influences eggload. Similar results
obtain in Nearctic P. napi and Cardamine populations (F. S. Chew,
pers. comm.). Since a single P. napi larva would consume all the above
ground biomass of a C. flexuosa plant, probably few hosts in the ditch
escape serious grazing.

Examination of the number of eggs on the leaflets of those plants with
at least one egg suggests a tendency towards clumped distributions
(Table 3). The trend occurs in all three shading classes, but is significant
only in the most heavily shaded class. The results contrast with data
from the marsh sub-population where the distribution fortuitously
agrees well with the random expectation. These results are to be
expected if multiple oviposition occurs in the ditch, but not in the marsh.

Discussion

The very large eggloads, received by some C. flexuosa plants in the
ditch, are far in excess of the number which can be supported through
larval development. Courtney and Courtney (1982) described similar

26(l-4):l-288, 1988

61

10

A

A

20

Eggs/

Plant

10 .

20 20 50

Leaflets / Plant

Fig. 1.

The relationship between the number of leaflets on C. f/exuosa
individuals and the number of P. napi eggs deposited, in four
categories of shading (determined by eye). The value of the cor¬

relation coefficient, r, is shown

A. (<25% of Sky obscured)

B. (25-50%)

C. (51-75%)

D. (>75%)

All data summed

r = .810
r = .955
r = .779
r = .698
r = .555

p < 0.005

p < 0.001
p < 0.001
p < 0.001
p < 0.001

inverse density dependent egg distributions in P. napi and A. car-
damines and ascribed them to changes in receptivity of females. It is
suggested that females accept suboptimal oviposition sites (such as those
already bearing an egg or larva) when hosts are rare. The present
results, which link increased eggloads to observed changes in behaviour,
support the idea that females modify their behaviour in response to
resource availability.

Such behaviour may have important consequences for population and
community processes: Shapiro (1975) and Courtney and Courtney
(1982) show how both intra- and inter-specific competition are greatly
increased by clumped egg distributions. The present study suggests that
such competition may be more important in peripheral habitats and in
the offspring of dispersing individuals, than in main population
centers. Similarly, Wiklund and Ahrberg (1978) have discussed the

62

J. Res. Lepid.

Table 3. Distributions of P. nap/ eggs over leaflets of those C. flexuosa plants
with eggs (elimination of other plants from the analysis follows the
conservative procedure of assuming that such individuals were
unsuitable/unavailable for oviposition). Only three classes of
shaded plant are given forthe heathland site, thetwo lighter classes
(<25% and 25-50%) being summed (mean eggloads are very
similar as in Table 2). Values are given for X2 comparisons with
expected values from a random (Poisson) distribution.

Eggs per leaflet

Shading

0

1

2

3

4

5

9

X2

d.f.

P

Heath

<50%

35

28

17

8

3

2

—

3.13

3

n.s.

51-75%

40

10

2

1

—

1

—

1.94

2

n.s.

>75%

231

22

9

5

2

2

1

36.90

2

<0.001

Marsh

69

17

2

—

—

—

—

0.03

2

>0.975

effect of inversely density related oviposition in a community of host-
plant species which differ in their dispersion. A. cardamines attack was
most severe on low density or highly dispersed hosts. In the present
study, P. napi in the marsh were mainly supported by the hosts C.
pratensis and N. officinale. C. flexuosa individuals in the marsh suffered
fewer attacks than conspecifics growing alone in the ditch. It seems that
the C. flexuosa population in the wood suffers from being in the vicinity of
large marshland populations of other Cruciferae. It is a general predic¬
tion from the studies of Wiklund and Ahrberg, and of Courtney and
Courtney, that peripheral species surrounding major hostplant localities
may suffer disproportionately from herbivore attack.

Literature Cited

CHEW, F. S. 1975 Coevolution of Pierid butterflies and their Cruciferous food-
plants. I. The relative quality of available resources. Oecologia 20: 117—
127.

CHEW, F. S. 1977 II. The distribution of eggs on potential foodplants. Evolution
31: 568-579.

COURTNEY, S. P. 1982 IV. Crucifer Apparency and Anthocharis cardamines
oviposition. Oecologia 52: 258-265.

COURTNEY, S. P. & COURTNEY S. 1982 The ‘edge-effect’ in butterfly oviposition:
causality in Anthocharis cardamines and related species. Ecological
Entomology 7: 131-137.

DEN OTTER, C. J., BEHAN, M. & MAES, F. W. 1980 Single cell responses in female
Pieris brassicae (Lepidoptera: Pieridae) to plant volatiles and conspecific
egg odours. J. Insect Physiology 26: 465-472.

GILBERT, L. E. & SINGER, R. C. 1975 Butterfly ecology. Ann. Rev. Ecol. Syst. 6:
365-397.

PARKER, G. A. & COURTNEY, S. P. 1984 Models of Clutch Size in Insect Oviposition.
Theor. Popul. Biol. 26: 27—48.

SHAPIRO, A. M. 1970 The role of sexual behavior in density related dispersal of
Pierid butterflies. Am. Nat. 104: 367—373.

26(l-4):l-288, 1988

63

SHAPIRO, A. M. 1975 Ecological and behavioral aspects of coexistence in six
crucifer-feeding Pierid butterflies in the central Sierra Nevada. Am. Midi.
Nat. 93: 424-433.

SINGER, M. C. 1982 Quantification of host specificity by manipulation of oviposi-
tion behaviors in the butterfly Euphydryas editha. Oecologia 52: 224-229.

SINGER, M. C. 1984 Butterfly-hostplant relationships: host quality, adult choice
and larval success. Symp. R. Ent. Soc. 11: 81—88.

TABASHNIK, B. E., WHEELOCK, H„ RAINBOTT, J. D., WATT, W. B. 1981 Individual
variation in oviposition preference in the butterfly Colias eurytheme. Oeco¬
logia 50: 225-230.

WIKLUND, C. 1981 Generalist vs. specialist behaviour in Papili machaon (Lepi-
doptera) and functional aspects on the hierarchy of oviposition preferences.
Oikos 36: 163-170.

WIKLUND, C. & AHRBERG, C. 1978 Hostplants, nectar source plants and habitat
selection of males and females of Anthocharis cardamines (Lepidoptera).
Oikos 31: 169-183.

Journal of Research on the Lepidoptera

26(l-4):64-72, 1988

Enzyme electrophoresis and interspecific hybridization
in Pieridae (Lepidoptera)-The case for enzyme
electrophoresis

Hansjuerg Geiger

Zoologisches Institut, Universitaet Bern, Baltzerstrasse 3, CH 3012 Bern, Switzerland

Abstract. In a comparison of results from laboratory intertaxa hybri¬
dizations and enzyme electrophoresis in Pieridae, Lor ko vie (1986)
recognizes differences in the estimates of genetic relationships of the
taxa investigated. Lor ko vie concludes in his paper that these dif¬
ferences are due to the electrophoretic approach. It is the purpose of this
publication i) to analyze this opinion, ii) to discuss possible limitations
and pitfalls of the hybridization approach, and iii) to show that an
adequate interpretation of the data may well lead to a generally
accepted idea on the genetic relationships in Pieridae.

Introduction

In a recent paper, Lorkovic (1986) compared results from his impres¬
sive work on artificial interspecific hybridization in Pieridae with
results of an analysis of the genetic relationships in this family by
means of enzyme electrophoresis. Lorkovic (l.c.) concludes that the
observed discrepancies between the results of the two approaches are
due to the electrophoretic analysis which gives inadequate estimates of
divergence at low taxonomic levels, and limitations of the scope of the
the biochemical method. Lorkovic, further limits the significance of
enzyme electrophoresis to the study of populations and denies the
possibility to delimit taxa with this method.

However, in his discussion Lorkovic does not analyse the real extent of
the alleged discrepancies, the limitations and pitfalls of his method, or
the problem of control data. There are also a number of misunder¬
standings of the electrophoretic approach and the interpretation of the
biochemical data.

In this publication I analyze the Lorkovic paper and demonstrate the
power of the biochemical-genetic approach.

Discrepancies between and results

Lorkovic (l.c.) compares the results of his crosses with the degree of
enzyme dissimilarity (EDf) in his Table 1. (Note: The values given in
Lorkovic’s Table 1 are actually I-values, not EDf-values. A more
appropriate statistic would be Nei’s D (Nei, 1972) for the degree of
genetic differentiation). We analyse here the statistical differences

26(l-4):l-288, 1988

65

between the results of the two approaches. For this investigation we use
the correct value for the comparison between Pontia daplidice and P.
protodice (I-value =.59, not .55; neither values have ever been pub¬
lished). We also disregard the fact, that Lorkovic (l.c.) has used his data
of crosses between Euchloe crameri and the taxon graeca for the
comparison of crameri and simplonia (electrophoretic data for graeca
are not available, but there is unpublished evidence that graeca might
be another species; this may be the reason for the observed differences
between our results). Furthermore, P. daplidice in South Europe
actually consists of two species (Geiger and Scholl, 1982a) and we use
here the value for the comparison of species 2, the eastern european
species, with protodice and Pieris rapae (these values were not available
to Lorkovic, but the differences are small). If we calculate now the
correlation coefficient for a linear regression between the two sets of
data we find r = .88 (lOdF) which corresponds to a P < 1%. This is a very
good fit and it seems unjustified to emphasize the differences. Of course
this does not mean that there is absolute correlation for any individual
comparison and the reasons for any observed deviations remain to be
discussed. As Lorkovic (l.c.) already pointed out, such differences occur
mainly at the lowest taxonomic ranks.

Advantages and disadvantages of enzyme electrophoretic methods

Enzyme electrophoresis is a method that allows one to compare
populations and taxa using a set of genetic markers (loci). The zymo¬
grams obtained by this method make it possible to collect directly data
on the genetic composition at individual loci. This means that different
alleles at a locus can relatively easily be recognized. It is very important
that the genetic interpretation of the zymograms is confirmed, if
possible by analyzing the progeny of parents with various electrophoretic
phenotypes, as with some enzymes additional bands may appear that
have no direct genetic background (e.g. conformeric forms). For the
Pieridae, an extensive analysis has been carried out on Pieris brassicae
(Geiger, 1982). The pattern found corresponds perfectly with a simple
Mendelian distribution.

If we are working with population samples of one or several taxa, we
obtain two kinds of information: i) which alleles can be found at a locus
in a population or in a taxon (qualitative information) and ii) in what
frequencies (quantitative information).

The qualitative information can be used to investigate the distri¬
bution of alleles among populations. If we find e.g. a situation in which
geographically separate populations of two taxa have different alleles at
one or several loci, but share a common polymorphism at these loci in a
zone of sympatry, it seems reasonable to conclude that the two taxa are
in reproductive contact or have been so only a very short time ago. If we
do not find such a common polymorphism in sympatry this is a strong

66

J. Res. Lepid.

argument to assume interruption of gene-flow, and the existence of two
species (e.g. Geiger and Scholl, 1982b; Geiger and Shapiro, 1986;
Shapiro and Geiger, 1986).

As the genetic variants are easily distinguishable (they are, for all
practical purposes, not detectably modified epistatically) the analytical
power of such an investigation can hardly be reached with ’’classical”
methods. The qualitative information obtained by means of enzyme
electrophoresis also allows a cladistic approach (Ward, 1985). This has
not yet been done for the Pieridae, but it is planned for the future.

Quantitative information: It is one of the advantages of enzyme
electrophoretic methods that the degree of genetic correspondence
between populations or taxa can be quantified. There are a number of
different coefficients of genetic identity or distance that have been
proposed during the last 20 years. In most modern investigations the
statistic I for genetic identity or D for distance as developed by Nei
(1972) and modified by Hillis (1984), are used. The D value (D = — lnl),
used here, is an estimate of how many gene substitutions have been
accumulated per locus since interruption of gene-flow between two
populations or taxa. Of course, these values are strongly influenced by i)
the choice of loci and ii) the number of loci investigated. Therefore, it is
only possible to directly compare values of two different investigations
in those rare cases in which an identical set of loci has been scored. The
argument raised by Lorkovic (l.c.) that the fact that the values obtained
in different systematic groups are different is a serious obstacle for the
use of enzyme electrophoresis in taxonomy, is therefore only in part
valid, as most investigators use different sets of loci. Thus, Lorkovic is
perfectly correct when he states that the work of Racheli (1984) on
Parnassius apollo cannot be directly compared with our analysis in the
Pieridae, but this is only a problem if we want to relate the results of
different studies. In all cases in which identical sets have been analyzed,
as in our Pierid studies, the results are comparable

It has already been demonstrated (Geiger, 1981) that the levels of
genetic identity found in different subfamilies of the Pieridae are in fact
comparable. This is now confirmed by a much larger sample (over 100
taxa currently, all compared at the same 22 loci). However, how well do
these levels correlate with the systematic rank of the taxa? Out of this
large survey I have selected 42 taxa whose systematic rank is currently
not seriously questioned and have related the D-values with the
systematic rank. The result is summarized in Table 1. The outcome is an
excellent agreement between the systematic rank generally used for the
taxa and the D- value. Futhermore, most levels are nearly free of
overlap; only between the levels of populations and subspecies as well as
genera and subfamilies is this not true (see also Geiger and Scholl,
1984). Therefore, it seems justified to use quantified enzyme electro¬
phoretic data to discuss the systematic rank of taxa under review
(Kitching, 1985)

26(l-4):l-288, 1988

67

This result found in the Pieridae is supported by similar investi¬
gations in other organisms (e.g. Avise, 1976). Once again, the important
thing is not the absolute I- or D- value, but the correlation with the
taxonomic rank.

I agree with Lorkovic (l.c.) that it is not possible to “prove” that a
taxon is differentiated to the species level by using the I- or D-value
alone. As I have already pointed out, a qualitative analysis of the
genetic data may be conclusive in cases of sympatry. In allopatric taxa
the degree of genetic differentiation may provide important arguments
in the discussion of the systematic position of taxa with unclear rank.
Again, the strongest clues in such situations may come from a qualita¬
tive analysis. There is little else one can do in such situations as the
biological species concept can only be applied with some restrictions.
This is exactly what we have always done when arguing at the species
level. In most cases for which a substantial level of genetic differentia¬
tion has been found, this level is due to an unshared polymorphism or
fixation of different alleles at one or more loci, rather then mere
differences in allelic frequencies (Geiger, 1981; Geiger and Scholl, 1982
a and b; Geiger and Scholl, 1985; Geiger and Shapiro, 1986; Shapiro and
Geiger, 1986). A similar analytical power could only be reached by a
cladistic analysis of characters from ’’classical” or electrophoretic data.
Such a cladistic analysis of classically used characters would also be the
only real test for the genetic relationships evaluated by means of
enzyme electrophoresis.

Another argument against the use of enzyme electrophoresis used by
Lorkovic (l.c.) is that speciation probably does not take place due to
changes at the loci covered by the electrophoretic approach. This is
certainly true, but it should be clear now that we have good evidence
that after the speciation event the taxa slowly accumulate changes at
these loci. The argument is not the speciation occurs because of these
alterations, but that due to the interruption of gene-flow after the
speciation event we can very often find changes at the loci investigated.
Therefore, it is also not important that not all the variation at the
enzyme loci can be detected by routine investigations. Nevertheless, the
amount of undetected variation mentioned by Lorkovic (l.c.) is only true
for some extremely polymorphic loci not usually used in the Pieridae
(Lewontin, 1986). For all other loci most of the variation is usually
detectable.

A possible severe limitation for the electrophoretic approach may be
that in rapidly evolving groups of taxa time was too short to result in
distinct differences at loci covered with this method. It has to be
expected that such cases will occur also in the Pieridae. However, it has
to be pointed out again that there is no such case in the control data as
yet. This is a clear sign that speciation events are generally reflected by
accumulation of genetic differences at the set of loci used in the Pieridae.

The scepticism towards using these biochemical-genetic data to

68

J.Res.Lepid.

evaluate systematics in the Pieridae also may have a historical reason.
The first case in which I applied this approach in this family was the
much debated European Pieris raap j-group taxon bryoniaei Geiger, 1978).
In this investigation it was not possible to detect any genetical dif¬
ferences between alpine bYyoniae and lowland napi. This first analysis
covered relatively few loci, but the results have since then been
confirmed by a much greater number of loci (among them also the highly
polymorphic esterases, Geiger, unpublished data), individuals, and
taxa (Geiger, 1981; Geiger and Scholl, 1985). It is a remarkable result of
this extensive work that genetic differences are very often greater
among geographically close populations of napi as well as of bryoniae
than between the two taxa. There was no other choice than to interpret
these results as a support for those authors who argued for conspecifity
of the two taxa. It lies in the nature of a disputed case that this
conclusion was contradictory to the published opinions of others. But
are the enzyme data really that much in opposition to the facts
presented by such authors? To answer this question it is necessary to
discuss the situation we encounter in the field and then the laboratory
results. Eitschberger (1984), one of the most convinced proponents of the
species rank for bryoniae , reports a significant number of hybrids found
in the field. Similar observations have been made by others (e.g., Varga,
1967). This fact clearly demonstrates that gene-flow between napi and
bryoniae is not interrupted under natural conditions even in Central
Europe. Lorkovic (l.c.) points out that the two taxa show a reduced
“hybrid fertility” in his laboratory crosses. This is certainly supported
by data presented by Lorkovic (l.c.). However, his data also clearly show
that there is some “hybrid fertility” even in the F2 crosses! The degree of
this “hybrid fertility” is remarkably high, especially in the black-
crosses (Rl, see Table 2, Lorkovic, l.c.) which clearly means that the
laboratory results confirm the observation of gene-flow in nature
(morphologically intermediate individuals). The enzyme data strongly
support this view indicating that there is no sign of an interrupted
reproductive contact. Clearly, a certain degree of reduced fertility can
be observed, but it seems safe to state that the data from different
approaches are not as contradictory as they have been presented; the
opposite is true. To solve the nomenclatural problem I propose to take
advantage of the rules in the new edition of the “International Code of
Zoological Nomenclature” (1985). We now have the possibility to take
into consideration a somewhat reduced degree of fertility, and rank such
a taxon as a semispecies. This is also exactly what Lorkovic (1962) has
done in earlier papers.

Interspecific hybridization and phylogenetic relationships

In his publication Lorkovic (l.c.) uses his data from laboratory inter¬
specific hybrid crosses to test the enzyme electrophoretic data. The basic
philosophy behind the use of these hybridization results to evaluate

26(l-4):l-288, 1988

69

phylogenetic relationships is the speculation that after interruption of
gene-flow the taxa gradually accumulate characters that directly affect
the degree of genetic incompatibility. However, to use his method as a
test for the enzyme electrophoretic data Lorkovic should first demon¬
strate that the results from the interspecific crosses in the Pieridae are
in fact strongly correlated with the phylogenetic relationships. This has
not been done and is no easy task, the reason of course being that we are
dealing with a historical process and there is no method available to
reveal unequivocally the real course of evolution. There are some
methods (like cladistic analysis) that have a high potential to do so, but
all methods have their pitfalls. All we can do is to try to apply as many
methods as possible and find the most parsimonious family tree. Again,
it has to be pointed out that the high correlation between Lorkovic’s
data and the enzyme electrophoretic analysis is highly encouraging and
should be the basis for future investigations. A third approach with a
potentially high power of resolution would be a cladistic analysis, but
such an analysis is not available for the Pieridae.

One case for which our approaches give different values of evolutionary
distance has already been discussed ( Pieris napi/bryoniae). There are
two more such cases: Euchloe crameril simplonia , and Pieris rapae /
mannii. To discuss these we first have to analyse possible problems and
limitations of the interspecific hybridization approach.

I) The interspecific hybridization approach as presented by Lorkovic
(l.c.) works uniquely with postcopulative isolating mechanisms. All
precopulative factors that prevent gene-flow between taxa such as
olfactory, behavioral, ecological, and partly morphological incompati¬
bilities are excluded by this approach since the usual method of mating
is hand-pairing. The importance of such factors should not be under¬
estimated. Strictly speaking, by this method, it is only possible to
compare taxa that have only developed postmating isolating mechan¬
isms, yet much effort should be devoted to evaluating both pre- and
postmating barriers. Such premating isolating factors seem to be the
reason for the discrepancies in at least one of the above mentioned cases:
Euchloe crameri and simplonia (again, Lorkovic used graeca instead of
simplonia , a fact that itself may account for the differences). Lorkovic
(l.c., p.345) himself mentions that there is a well-expressed premating
barrier between these two taxa. Taxa that are separated by such
mechanisms do not need to develop additional strong postcopulative
mechanisms (Mayr, 1963). The approach used by Lorkovic (l.c.) will in
such cases underestimate the degree of genetic differentiation. On the
other hand enzyme electrophoresis measures the accumulated dif¬
ferences since interruption of gene-flow regardless of the true nature of
isolating mechanisms. Therefore, it seems unjustified to solely blame
enzyme electrophoresis for the observed differences in the results of the
two approaches in the sense of not revealing the true degree of
genealogical relationships. Moreover, as most of the populations of the

70

J. Res. Lepid.

two taxa are allopatric, relatively weak isolating mechanisms seem to
be sufficient to maintain genetic identity (this is also an important
problem for the hybridization approach in clear-cut allopatric taxa,
especially in taxa from different continents, islands, or mountain
ranges. In such situations theoretical problems in applying the bio¬
logical species concept also arise).

A similar problem may be the basis for the differences in the results of
the comparison of Pieris rapae/mannii. These two taxa are sympatric in
large parts of their recent distribution area. To avoid gene-flow and
maintain identity as distinct species the two taxa have obviously
developed strong postmating isolating factors. This does not mean that
speciation occured because of the same factors. The degree of hybrid
sterility will in such situations tend to overestimate the phylogenetic
distance. Furthermore, it should be noted that strong hybrid sterility
may be caused by mutations at one or a few loci and need not reflect
profound genomic differences. Complete sterility among strains within
a species may also occur due to transposable elements (hybrid dysgenesis
in Drosophila, Kidwell et al., 1977; Engels, 1983) or as a consequence of
an infection by a microorganism ( Tribolium , Wade and Stevens, 1985).

II) I have already mentioned several times the fact that enzyme
electrophoresis is primarily a method to estimate the time passed since
interruption of gene-flow (Berlocher, 1984; O’Brien et al., 1985). There
is good evidence that this is also true for the Pieridae, one indication
being the non-overlap of the levels of genetic differentiation (Fig. 1). To
make the hybridization data comparable one would have to demon¬
strate that the factors used by Lorkovic (l.c.) such as “size”, “number of
offspring”, “development” and “in viability” are also correlated with the
phylogenetic age of the taxa. Furthermore, the proposed quantification
of these factors needs also to be tested for this correlation.

IH) The interspecific hybridization approach as used by Lorkovic (l.c.)
works with individual, essentially randomly-selected animals, not
populations. The numbers of comparions are in many cases very low.
Therefore, it would be highly important to know more about the
reproductive success of randomly chosen individual butterflies. Our
own observations among European and North- American taxa show that
the degree of fertility, even among individuals of one population, may
vary enormously and may be different among taxa. In other words, we
first have to know more about the variance of fertility among individuals
of local populations before we can quantify such rates among taxa. In
some critical cases there should even be a detailed analysis comparing
the fertility among individuals of geographically distant and close
populations and especially within a zone of contact. Unfortunately, the
amount of labor required to do such an analysis may often be prohibi¬
tively great. Nevertheless, there are investigations that use this
approach (e.g., Oliver, 1978).

26(l-4):l-288, 1988

71

+

+

i—

.00

+

H - 1 - 1-

.05

- 1 - Species of different subfamilies

- 1 - Species of different genera

Species of different species-groups
Species of the same species-group
Subspecies
Local populations

-I - 1 - 1 - 1 - 1 - 1 - 1 -

i.o 1-5 D- values

Fig. 1 . Levels of genetic differences among 42 taxa whose systematic rank is
currently not debated (22 loci)

Conclusions

It was the purpose of this publication to continue the discussion on
how best to estimate the degree of the phylogenetic distance between
taxa. It has been concluded that it can not be inferred from the observed
differences between the results of the interspecific hybridization and
enzyme electrophoretic approaches that the latter method gives in¬
adequate estimates. In fact such differences only occur in some much-
debated cases for which there are good reasons to assume that the first
method may over- or underestimate the phylogenetic age of the taxa
discussed. It has been demonstrated in a set of Pierid taxa whose
systematic rank is generally not questioned that enzyme electrophore¬
tic data are highly correlated with the systematic rank of the taxa. The
generally good agreement between the results of the two approaches is
regarded as highly encouraging for future analysis.

Acknowledgements. I thank Profs. Z. Lorkovic (Zagreb) and A.M. Shapiro (UC
Davis) for very stimulating criticisms on earlier drafts of this paper and Prof. H.
Descimon (Marseille), A. Porter (Davis, CA), and M. Lortscher (Bern) for
discussions. S. Whitebread (Magden, Switzerland) read the english version of
the manuscript. If I do not agree with Prof. Lorkovic on several subjects, this is
entirely due to our differing views and does not alter my deep respect for this
eminent scientist.

Literature Cited

AVISE, J.C., 1976. Genetic differentiation during speciation. In: AYALA, F.J.(Ed),
Molecular evolution. Sinauer, Massachusetts, p. 106-122.

BERLOCHER, S.H., 1984. Insect molecular systematics. Ann. Rev. Entomol.
29:403-433.

EITSCHBERGER, U., 1984. Systematische Untersuchungen am Pieris napi-
bryoniae- Komplex. 1/1 and II. Herbipoliana, Marktleuthen.

ENGELS, R.E., 1983. The P family of transposable elements in Drosophila. Ann.
Rev. Genet., 17:315-344.

GEIGER, H.J., 1978. Die systematische Stellung von Pieris napi bryoniae :

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Biochemisch-genetische Untersuchungsbefunde. Ent. Zeitschrift, 88:229-
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- , 1981. Enzymelectrophoretic studies on the genetic relationships of

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- , 1982. Biochemisch-genetische Untersuchungen zur Systematik und

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GEIGER, H.J. & A. SCHOLL, 1982a. Pontia daplidice (Lepidoptera, Pieridae) in
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- , 1982b. Enzyme electrophoretic approach to the systematics and

evolution of the butterfly Euchloe ausonia. Experientia, 38:927-929.

- , 1984. Enzymelektrophoretisch erfassbare genetische Aehnlichkeiten

in der Familie Pieridae (Lepidoptera). Verh.Dtsch.Zool.Ges., 77:265.

- , 1985. Systematics and evolution of holarctic Pierinae (Lepidoptera).

An enzyme electrophoretic approach. Experientia, 41:24-29.

GEIGER, H.J. & A M. SHAPIRO, 1986. Electrophoretic evidence for speciation within
the nominal species Anthocharis sara Lucas (Pieridae) J.Res.Lepid., 25(1):
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HILLIS, D M., 1984. Misuse and modification of Nei’s genetic distance. Syst. Zool.
33:238-240.

KIDWELL, G.M., J.F. KIDWELL, & J.A. SVED, 1977. Hybrid dysgenesis in Drosophila
melanogaster : A syndrome of aberrant traits including mutation, sterility
and male recombination. Genetics, 86:813-833.

KITCHING, I.J., 1985. Allozyme variation in the milkweed butterflies (Lepi¬
doptera: Danainae). Zool. J. Linn. Soc., 86:367-386.

LEWONTIN, R.C., 1986. How important is genetics for an understanding of
evolution? In: American Society of Zoologists (Ed.), Science as a way of
knowing-III. Genetics. Amer.Zool., 26:811-820.

LORKOVIC, Z., 1962. The genetics and reproductive isolating mechanisms of the
Pieris napi-bryoniae group. J.Lep.Soc., 16:5-19 and 105-127.

- , 1986. Enzyme electrophoresis and interspecific hybridization in

Pieridae (Lepidoptera). J.Res.Lepid., 24(4):334-358.

mayr, E., 1963. Animal species and evolution. Belknap Press, Cambridge.

NEI, M., 1972. Genetic distance between populations. Am.Nat. 106:283-292.

O’BRIEN, S.J., W.G. NASH, D.E. WILDT, M.E. BUSH & R.E. BENVENISTE, 1985. A molecular
solution to the riddle of the giant panda’s phylogeny. Nature, 317:140-144.

OLIVER, C.G., 1978. Experimental hybridization between the nymphalid butter¬
flies Phyciodes tharos and P. campestris montana. Evolution, 32(3), 594-601.

RACHELI, T., R. CIANCHI & I. BULLINI, 1983. Differentiamento e variabillita genetica
di alcune sottospecie di Parnassius apollo (Lepid. Papilionidae). Atti XIII
Congr. Naz. It. Ent., Torino, Italy.

SHAPIRO, A M. & H.J. GEIGER, 1986. Electrophoretic confirmation of the species
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VARGA, z., 1967. Populationsstudien ueber Pieris bryoniae O. im Karpathen-
becken.I.Acta Biol.Debrecina 5:139-151.

WADE, M.J. & L. STEVENS, 1985. Microorganism mediated reproductive isolation in
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Journal ofResearch on the Lepidoptera

26(l-4):73-81, 1988

Systematics of Ascia (Gaiiyra)

(Pieridae) Populations in the Sonoran Desert

Richard A. Bailowitz

2774 W. Calle Morado Tucson, AZ 85745

Abstract. A breeding population of Ascia ( Ganyra ) (Lepidoptera:
Pieridae) in the josephina complex is confirmed for Arizona. The larval
hostplant is found to be Atamisquea emarginata Miers., a plant
confined to the Sonoran Desert.

This population, while closely related to Ascia josephina josepha, is
distinct in maculation, androconial pattern, genitalia, and larval host.

Its taxonomic status is discussed. Full species status is given to the
insect as Ascia howarthi (Dixey).

Introduction

During the course of a study for the National Park Service on
arthropods of the Quitobaquito Management Area in Organ Pipe Cactus
National Monument (OPCNM), Pima County, Arizona, the author
discovered an apparent population of a pierid butterfly in the Asia
josephina (Godart) complex. The discovery is noteworthy in that records
of this species complex are scarce in Arizona, in that the insect appears to
be breeding in the area, and in the distinct phenotypes of the population.

The genus Ascia is divided into two subgenera, each represented by a
single Nearctic species or species complex (Howe, 1975). An additional
taxon, A. sevata (C. & R. Felder), is recorded in Mexico. The subgenus
Ascia includes the species monuste (Linnaeus) and its subspecies. The
subgenus Ganyra Billberg includes the species (or species complex)
josephina (Godart) with its Central and North American subspecies
josepha (Salvin & Godman), howarthi (Dixey), and kuschei (Schaus) and
it also includes the species sevata.

Only josephina josepha from Ganyra has been cited within the United
States (Pyle, 1981, et al.). However, the recent studies at Quitobaquito,
OPCNM confirm the presence of an Arizona population of the josephina
complex and suggest assignment outside the subspecies josepha.

This paper examines the dimensions of the Arizona Ganyra population
and its relationships with other populations in the josephina complex. It
links the Sonoran Desert Ganyra population to a specific larval foodplant
and alters the taxonomy of the species complex.

A series of both sexes of adults of Sonoran Desert Ganyra was collected
from several wild populations. Those were further augmented by
specimens from the San Diego Natural History Museum, the California
Insect Survey collection, and the private collection of Kilian Roever. A

74

J. Res. Lepid.

small series of A.josephina from southern, eastern, and western Mexico,
as well as Texas, was also obtained. The Texas A & I collection and the
private collections of J. Brock, D. Mullins, and P. Hubbell were also used.
Steve Prchal of the Arizona-Sonora Desert Museum photographed all
stages of the life history. Measurements were done using a Lassico
Ocular Filar. An ISI-DS-130 scanning electron microscope at the Uni¬
versity of Arizona campus was used for the micrographs.

Taxonomy

Most of the taxa in the A.josephina complex were originally described
in the genus Pieris Schrank. They were later placed in the genus Ascia
Scopoli and still later in either the genus or subgenus Ganyra , depending
on the author. Godart described nominate josephina in 1819 from the
Antilles, probably Haiti and Cuba (Salvin & Godman, 1868). It is
characterized by the large size, falcate fore wing, and a large black
forewing discal cell-spot. Two other closely related Antillean subspecies
have been described: paramaryllis Comstock from Jamaica, and krugii
(Dewitz) from Puerto Rico. Both of these are somewhat smaller, with the
characteristic black forewing cell-spot narrow (Comstock, 1943).

The subspecies josepha (Salvin & Godman) is the widespread Mexican
and Central American form. It differs from nominate josephina primari¬
ly in the shape of the wings. The fore wings are not as falcate as those of
josephina and the hindwings are more squared and less produced at the
anal angle. It occurs from southern Texas (straying northward to
Kansas) through eastern Mexico into Central America and north along
the west coast of Mexico into Sinaloa.

Dixey (1915) described subspecies howarthi from Baja California Sur,
Mexico. He described it as having more noticeable marginal spotting on
the forewing and more pronounced reticulations on the ventral hind¬
wing than typical josephina, as well as being comparatively small (males
58 mm wingspread, females 52 — 54 mm).

Schaus (1920) described kuschei from Mazatlan, Sinaloa, Mexico. It is
characterized by a greater extent of the submarginal maculation on the
forewings in both males and females. He cited both sexes as having
wingspreads of 56 mm which is considerably smaller than typical
josephina.

Felder & Felder (1861) described the taxon sevata from Venezuela.
Rindge (1948) cited the Baja California population of Ganyra as A. sevata
kuschei, a new combination at that time. The original description of
sevata mentioned a solid white dorsal surface except for the apical
marginal border, wider in females than in males. Fruhstorfer (1908)
subsequently named a more northerly subspecies, A. sevata tihurtia,
from Guatemala. It differs from true sevata in the narrowing or absence
of A. sevata’ s 5 mm forewing border. The ventral hindwings of A. sevata
and A. tihurtia are also suffused with shades of violet and pink
respectively. No mention of forewing cell-spots is made for either taxon.

26(l-4):l-288, 1988

75

There are obvious close superficial relationships among the various
members of the subgenus Ganyra. A closer look will now be given to the
exact relationships between those taxa (north of Guatemala) and where
the OPCNM population lies in reference to them.

Several key points suggest that A. sevata sevata and A. s. tiburtia are
more distantly related to the josephina complex. The lack of the black
forewing cell-spot does not in itself suggest removal from the group since
Cuban, Jamaican, and Puerto Rican populations of josephina also have
this spotting weak to non-existent (Comstock, 1943). However, the
presence of a narrow to wide continuous forewing border and pink to
purple ventral hindwing coloration do set these taxa apart. More
importantly, there are significant differences in the form of the male
androconia. These average only 0.24 mm in length in sevata , approx¬
imately half that of the josephina complex members (Dixey, 1915).
Hoffman (1976) cited tiburtia from southern Mexico, a range overlap¬
ping that of josepha. This sympatry suggests distinction at the specific
level. Sevata is here considered outside the josephina complex.

The relationship between A. howarthi and A. kuschei needs clarifica¬
tion. Although kuschei was described from Mazatlan, it appears to have
been collected far north of there. There are many recent records of a
Ganyra from the Mazatlan area, all of which represent josepha , not
kuschei. The maculation, androconia, and size are all consistent with
josepha. The location “Mazatlan” was probably used in a broad sense by
Kusche. Phenotypes matching the kuschei description occur from ex¬
treme northern Sinaloa northward, where he easily may have collected.
Many specimens from Sonora and Arizona, and a few from Baja
California closely match the kuschei description, especially for broods
during the summer rains. The type locality for kuschei should probably
be amended to San Miguel, near Los Mochis, Sinaloa, Mexico, the
southernmost locality for which that taxon is known. The howarthi
phenotype, at least ventrally, is restricted to Baja California. But other
phenotypes with unmarked ventral hind wings also occur there. These
unmarked phenotypes are also widespread in central and northern
Sonora and represent the majority of the specimens taken at OPCNM.
The three phenotypes — heavily marked kuschei , ventrally marked
howarthi , and the unmarked population — all blend with each other.
Also, the OPCNM population was reared on a Sonoran Desert shrub,
Atamisquea emarginata Miers., in the family Capparidaceae. When
mapped together, the distribution of the three phenotypes duplicates
that of the probable hostplant, A. emarginata (Fig. 1). Kuschei and
howarthi , plus the unmarked phenotype are therefore considered
synonymous. Similarities in size, androconial pattern, and the distribu¬
tion of the larval foodplant all suggest this. Since howarthi has priority,
the name kuschei is suppressed as a junior synonym.

The third clarification necessary is the relationship between josepha
and howarthi. On the basis of size, howarthi and josepha represent two

76

J.Res.Lepid.

Fig. 1. Distribution of Atamisquea emarginata, probable larval host of Ascia
howarthi.

very different populations. Both sexes of howarthi average in excess of
10 mm smaller in fore wing length than those of josepha. There is no size
overlap in the males and only a single small female josepha overlaps the
size range of female howarthi. The size mentioned for howarthi by Dixey
(1915) seems inflated. He cited the male wingspread as 58 mm but the
male figured is only partially spread. His figure has a 55 mm wingspread
with a 26 mm forewing length. In contrast, a well-spread specimen with
a 26 mm forewing length has only a 45 mm spread.

The wing shape is also different in the two taxa. In howarthi , the
forewings are somewhat shortened and the hindwings are very rounded.
In josepha , and apparently even more so in nominate josephina, the

26(l-4):l-288, 1988

77

forewings are produced or even falcate while the hindwings have the
anal angles extended (Figs. 2—5).

In their maculation, the males of howarthi are generally more heavily
patterned than those of josepha, especially with forewing marginal and
submarginal spotting. Females are very similar in the two taxa, but in
josepha dark morphs are often produced where the ground color is
heavily overlaid with cinnamon brown. All specimens of this morph seen
were mid-summer captures and probably parallel the long-day form of
Ascia monuste (Pease, 1962). The absence of dark morphs in howarthi is
interesting in light of the paucity of records away from breeding colonies
(Fig. 6). Both sexes of howarthi have an additional diagnostic mark on
the ventral hindwing. Aside from the dark scaling distal to the cross-
vains at the base of cells M3 and Cul, there is a darker spot on and around
the cross- vein at the base of cell M2. While males of josepha will rarely
have that cross- vein darkened (even dorsally) it is not scaled away from
the vein as in howarthi.

While the androconial scales of howarthi and josepha are similar in
size and configuration (Dixey, 1915), their placement on the wings is
vastly different. All androconia of howarthi are confined to fore wing cells
M3, Cul, Cu2, and 2A (Fig. 4). On josepha , the androconia in these cells
are far more extensive. In addition, scent scales are present in cells M2,
Ml, and the discal cell. Furthermore,yosep/ia invariably has androconia
on the hindwing as well, in cells RS, M2, M3, the discal cell, and
occasionally in Cul.

The general configurations of the male genitalia of both taxa are
similar. Due to the larger size of josepha , the entire genital capsule is
larger in that taxon than in howarthi. However, the length of the saccus

Ascia howarthi

Ascia howarthi

Fig. 2. Venation Fig. 3. Ventral maculation Fig. 4. Androconia pattern

78

J. Res. Lepid.

Fig. 5. Phenotypic range of Ascia howarthi (top two rows, males above,
females below) and Ascia josephina josepha (bottom row).

is proportionately greater in howarthi than in josepha. Likewise, the
aedeagal elbow (Fig. 7) of josepha is proportionately larger than that
of howarthi. These genitalic differences are present but best used
cautiously.

For the genus Ascia , members of the families Cruciferae, Cappar-
idaceae, and Batidaceae have been reported as hostplants (Howe, 1975).
More specifically, Capparis fro ndosa Jacq. was cited as a hostplant for A.
josephina josepha (Jordan, 1981). Although this caper is confined to the
eastern slope of Mexico, mostly Tamaulipas and Veracruz (Standley,
1961), other members of the genus have wider distributions in Mexico
and probably serve as foodplants in other parts of the range of josepha.
Members of the genus Capparis , whose distributions include Sinaloa,
areC. flexuosah., C. verrucosa Jacq., andC. indica (L.) (Standley, 1961).
Any of these might serve as larval hosts for josepha since the northern
distributional limits of the plant and insect appear to match one another.
None of the members of the genus Capparis has been reported as far
north as Sonora. Records for josepha extend north to Mazatlan, Elota,
and Guamuchil, approximately 100 km southeast of Los Mochis in
northern Sinaloa.

Records of howarthi extend south into northern Sinaloa, near Los
Mochis. This brings the two taxa within 100 km of each other. In fact, a
somewhat questionable record of a dark morph female josepha taken
near Alamos, Sonora would bring the two populations into overlap. The
lack of an apparent cline or anything resembling intergrades suggests a
high integrity and differentiation between the two populations.

26(l-4):l-288, 1988

79

Fig. 6. Distribution of Ascia howarthi.

This differentiation between the Sonoran Desert Atamisq wea-feeding
howarthi and the Capparzs-feeding josepha of the remainder of Mexico
persists even when howarthi is compared to josephina as a whole. It
therefore appears that the differences are at the species level, not at the
subspecies level. Therefore, I propose to elevate the taxon howarthi to
species status. It is most closely related to A. josephina , less so to A.
seuata.

The genus north of Guatemala and exclusive of the Antilles would be
composed of four species as follows:

Genus: Ascia Scopoli

Subgenus Ascia Scopoli

80

J . Res. Lepid.

Fig. 7. Aedeagus of Ascia howarthi showing "elbow” to left of center.

1. monuste (Linnaeus)

a) monuste (Linnaeus)

b) phileta (Fabricius)

c) cleomes (Boisduval & Le Conte)
Subgenus Ganyra Billberg

2. josephina (Godart)

a) josepha (Salvin & Godman)

3. howarthi (Dixey)

4. seuata (C. & R. Felder)
a) tiburtia (Fruhstorfer)

Literature Cited

BOWERS, J. E. 1980. Flora of Organ Pipe Cactus National Monument. J. Ariz.-
Nev. Acad. Sci. 15: Issue 1, 11, p. 1-47.

COMSTOCK, w. P. 1943. The genus Ascia in the Antilles. Amer. Mus. Novit.
#1229: 1-7.

DIXEY, F. A. 1915. New Species and Subspecies of Pierinae. Trans. Entomol. Soc.
Lond. Part 1: 9—17.

FELDER, C. & R. FELDER. 1861. Lepidoptera nova Columbiae. Wien. Entomol.
Monatschr. V: 72—87.

26(l-4):l-288, 1988

81

FRUHSTORFER, H. 1908. Neue Sudamerikanische Pieriden. Societas Entomol.
XXII: 139-140.

HASTINGS, J. p., TURNER, R. M„ & D. K. warren. 1972. An atlas of some plant
distributions in the Sonoran Desert. Instit. Atmos. Phys., Univ. Ariz.
Tucson, AZ. Tech. Rep. No. 21.

HOFFMAN, C. C. 1976. Catalogo Sistematico y Zoogeografico de los Lepidopteros
Mexicanos. Soc. Mex. Lepid. A. C.

HOWE, W. H. 1975. The Butterflies of North America. Doubleday. New York.
633 pp.

JORDAN, C. T. 1981. Population biology and host-plant ecology of caper-feeding
pierid butterflies in northeastern Mexico. Dissertation. U. Tex., Austin.
PEASE, R. W. 1962. Factors causing seasonal forms in Ascia monuste
(Lepidoptera). Science. 137: 987—988.

PYLE, R. M. 1981. The Audubon Society Field Guide to North American Butter¬
flies. New York. Alfred A. Knopf. 916 pp.

RINDGE, F. H. 1948. Lepidoptera: Rhopalocera. No. 8 of Contributions toward a
knowledge of the insect fauna of lower California. Proc. Cal. Acad. Sci.
24: 289-312.

SALVIN, O. & F. D. GODMAN. 1868. On some new species of diurnal Lepidoptera
from South America. Ann. Mag. Nat. His. Vol. II: 141-152.

SCHAUS, W. 1920. New species of Lepidoptera in the United States National
Museum. Proc. U. S. Nat. Mus. 57: 107 — 111.

STANDLEY,P. C. (Issued) 1961. Trees and Shrubs of Mexico. Cont. U. S. Nat’l. Mus.
Vol. 23. Smith. Inst., Washington, D. C.

Journal of Research on the Lepidoptera

26(l-4):82-88, 1988

On Pieris (Artogeia) marginalis macdunnoughii
Remington (Pieridae)

S. R. Bowden

Lydeard, Merryfield Way, Storrington, W. Sussex, RH20 4NS, U.K

Abstract. Some previous work on Pieris (Artogeia) ssp. macdunnoughii
and ssp. marginalis is summarised. Experiments in which Colorado
macdunnoughii were hybridized with European P. napi L. are reported,
with particular attention to inheritance of yellow pigmentation and
viability of Fx hybrids: no females are obtained from napi 9 x
macdunnoughii cf- Macdunnoughii carries two distinct systems of
yellow coloration, both the bisexual recessive sulphurea and the
female-limited dominant flava. Whether this unusual condition has
any ecological or evolutionary significance is uncertain. It is concluded
that neither subspecies marginalis nor macdunnoughii is conspecific
with the typical napi of Europe, but Eitschberger’s opinion that
macdunnoughii is a subspecies of a species marginalis Scudder may
well be correct. However, the relative position of the subspecific taxa
venosa Scudder, microstriata Comstock and mogollon Burdick requires
experimental investigation.

Introduction

This Colorado subspecies was described as of Pieris napi L. by Barnes
& McDunnough in 1916 under the name pseudonapi. The type-locality
was “Silverton, 10,000 ft., where it is single-brooded”. The description
as very close to European napi “although in the females the black dots of
the primaries are practically obsolete” is very meagre for a subspecies;
the females varied in the amount of black markings. The completeness
of the picture is improved if we can add the authors’ description of
pallidissima, the second generation at Provo, Utah, which they seem to
have regarded as consubspecific with pseudonapi.

Remington (1954) pointed out that the name pseudonapi had been
used by Verity (1911) for a race of Pieris melete Menetries, and re-named
the Colorado insect P. napi macdunnoughii. He added that the series of
Pieris napi at Yale University from Colorado, Utah, Nevada and
Wyoming [i.e. the populations considered as subspecies macdunnoughii
and ssp. pallidissima by Eitschberger 1983] indicate that all represent a
single race. He says, “My series from the Teton Mountains of Wyoming
shows very dark and very pale individuals taken flying together” — one
supposes that the reference is to melanic markings, not to any yellow
pigmentation. I conclude that Remington, like Barnes & McDunnough
before him, regards macdunnoughii and pallidissima as showing no
differentiation worthy of separate subspecific naming.

26(l-4):l-288, 1988

83

Eitschberger (1983) treats these populations as separate subspecies of
a species Pieris marginalis Scudder, which is said to include also
mogollon Burdick (New Mexico) and six newly named subspecies as well
as ssp. marginalis (Washington Terr.) and perhaps hulda Edwards
(Alaska).

Warren (1963), working from androconial scales, had attributed ssp.
marginalis to napi and ssp. macdunnoughii to a species oleracea Harris,
but the present writer is inclined to follow Eitschberger in respect of
these two subspecies, if only on account of their special pigmentation.

The Ssp. marginalis

In 1970 I reported experiments in which a N. W. Oregon (Saddle Mt.
State Park) stock of “Pieris napi marginalis ” was crossed with the bright
yellow British napi form sulphurea Schoyen (Head’s “citronea”) and
with certain other European forms. The Oregon insects were yellow in
ground-color: very faintly citron-shaded in the males but deeper in the
females. The markings of the latter sex differed from those of napi by the
often unequal development of the melanic markings: though the fore¬
wing hind-marginal streak was generally present, the second and even
more the first discal spots were much reduced and often absent.

The breeding results seemed to indicate that the population was
homozygous for an intermediately recessive gene of the subtalba-
sulphurea series (Bowden 1963), indistinguishable from that producing
“Thompson’s pale yellow” rarely in the British Isles. The conclusion
then was that “pale yellow” heterozygotes should be sought in other
American subspecies of the napi group : it was hardly expected that
other homozygous populations would be found, but macdunnoughii
might perhaps be polymorphic in respect of sulphurea?

However, the flava (ochreous) color whch was found in the napi-
hybrid females was then attributed to the European parents and it was
not realized that the marginalis stock might also carry some flava. But
Shapiro (1985, in litt .) had noticed that some marginalis populations
had high frequencies of a buff female. Looking at the 1970 marginalis
specimens now, it seems beyond dispute that the females possess an
ochreous tinge as well as the lemon, and that the Fx hybrid females’
color is unlikely to have come entirely from the European side.

The Ssp. macdunnoughii

In 1971 I reared only two females of this subspecies from Gunnison
Co., Colorado (W.B. Watt). They were caged with two (later four)
English (Herts.) males, but these paid no attention, though brothers
were pairing with other butterflies in nearby cages. After two days three
(later five) Scottish males were supplied, but again showed no concern.
A male from French Pyrenees showed minimum interest. It appeared
that at least these two individual females had no pheromone attraction
for European males.

84

J.Res.Lepid.

Not until 1985 were we able to raise macdunnoughii in adequate
numbers (brood 1985-r), by pairing offspring of females taken in Grand
Co., Colorado, in 1984 (Shapiro). We found that the fresh females were
definitely a greenish yellow (very near the color of the forewings of
Gonepteryx rhamni L. $), the males very palely yellow, but quite
distinct from the white of napi. Apart from slight fore wing apical
blackening, our males were typically without marking, and the females
carried only the forewing discal spot that normally marks napi males;
even this was sometimes absent and the hind-marginal female streak
hardly ever showed at all.

Thus comparing the markings of these two subspecies with those of
typical napi , the females departed from type in opposite directions: the
expression of marginalis forewing markings was biassed posteriorly,
those of macdunnoughii even diminished in that direction.

Experimental

Macdunnoughii males of brood 1985-r paired in August 1985 with
British napi f. sulphurea females, but the mere 15 eggs produced only
nine pupae, of which six (presumably female) proved unable to develop.
One male hybrid emerged normally in September, a very pale lemon
yellow, with faint black forewing apices and spots, and very slight
veining below. Two more pupae colored as males and split their thoraces
but failed to eclose: these were dissected out and appeared similar to the
first male. This rather disappointing result, as far as it goes, does
indicate that the macdunnoughii lemon tint, like that of marginalis ,
belongs to the sulphurea series of alleles.

Three other pairings of macdunnoughii males (one from 1985-r, two
from the subsequent inbred 1985-qO were made with single napi females
of mainly European stock (which however carried some genes derived
from ssp. oleracea and were of funebris form — see Lorkovic 1971,
Bowden 1983). The resulting broods (1985-&, 1986-g1, gu) were large, but
nevertheless only male hybrids emerged, even after diapause; their
upperside ground-color was uniformly pure white, without yellow
toning, as expected from heterozygous sulphurea. There was indeed
one female ing11, but this was rejected. ( Marginalis and macdunnoughii
hybrids with napi usually carry radiating black markings on the distal
ends of the veins. Such are uncommon on pure ssp. napi , and napi waifs
can thus be recognised, though without certainty.) The melanic pig¬
mentation of the hybrids may be described and discussed later.

The apparently more difficult reciprocal hybridization was obtained
in August 1986, using a funebris heterozygote male of 1985-/i. The
macdunnoughii female came from brood 1985-g. It was expected that in
this case female hybrids would precede the males, if the apparent sexual
imbalance in the earlier hybrids was due to a disturbance of diapause
control. In this brood, 1986-rc, losses of larvae in early stages were
appreciable, probably 15-20%. Twenty males emerged 14-21.ix.86,

26(l-4):l-288, 1988

85

followed by a mixture of 19 cf 4- 25 9 by 23.X.86, giving a final ratio of
8 cf : 5 $• A few other pupae died.

Surprisingly, all the F: hybrid females were of a light yellow color,
rather with a flava tinge, and none were very near white. Thus this color
was inherited from the macdunnoughii mother in the dominant mode,
and was expressed only in the females.

It was thought unlikely that a straight F2 hybrid brood would be
productive. Males of the first F1, 1985-&, were back-crossed in each
direction:

JJfunebris hz., 1985-/i x cfcf 1985-& —> 1986-c, 33 cf + 24 $,
9 macdunnoughii , 1985-r x cf 1985-& — > 1986-y, 5 cf + 3 J,
the sexes here being not significantly far from equality (x2 = 1.4,
p = 0.23 and x2 = 0.5, p = .90 respectively). Brood 1986-c gave 57 adults
the same year. All the males were white, as were some of the females,
others varying from slightly tinted up to near a full flava color in two
cases. The pigmentation here was definitely not of the sulphurea series,
and as the brood had two mothers, a ratio 2 flava: 22 no-flava would be
meaningless. Fifteen individuals of 1986-c were funebris (though of
varying expression), almost exactly the anticipated number.

The other back-cross 1986-j resembled pure macdunnoughii in respect
of wing ground-color: tinted citron even in the males, very definitely so
in two of them. The one white male was small (2 x 19 mm), but the other
seven insects were large. The striking markings of several females,
possibly influenced by the funebris gene, may be discussed in a later
paper.

Pigmentation Systems

More than thirty years ago (1954) I was able to write of general
agreement that there were at least two distinct forms of napi with extra
yellow on the wings: ochreous flava Kane confined to the female and
with the color not extending to the fore wing disc underside, and the
primrose sulphurea Schoyen with the more extensive bisexual colora¬
tion. Thus all the Pieris butterflies with yellow wing-uppersides have
since been interpreted in terms of these two modes:

female only — brownish — dominant or semi-dominant genetically
bisexual — citron yellow — recessive genetically.

So the tests are:

if a problem male (white or yellow) crossed with a sulphurea
female napi produces any yellow sons, it carries sulphurea;
if a problem female (yellow or ochreous) crossed with a sulphurea
male napi produces any yellow sons, it carries sulphurea; if only
white sons, it carries flava;

if a problem female crossed with wild-type napi produces any yellow
daughters, it carries flava,
provided that numbers are adequate.

86

J. Res. Lepid.

In both flava and sulphurea the saturation of the color can vary from
very faint to intense, sometimes by the action of different alleles (cf.
Bowden 1961) and sometimes environmentally. Some sulphurea alleles
give only a very faint coloration to male butterflies: in all doubtful cases
the forewing underside disc should be examined (Bowden 1961). Also,
the sulphurea color is particularly liable to fade, especially in sunlight.
At paler levels visual discrimination between flava and sulphurea
becomes more uncertain.

It was known that both kinds of color could occur together in the same
individual — Bowden (1962) described the transference of sulphurea
genes to P. (napi) bryoniae Ochsenheimer. Unfortunately I never
considered that any natural population might constantly use a com¬
bination of the two systems of pigmentation. It was thus that I came
partly to misinterpret the Oregon marginalis coloration.

When the pigments do co-exist in uncertain proportions in the
females of a species they are difficult to distinguish visually, even in a
fresh specimen. Photographic rendering a usually imperfect. If the flava
and sulphurea systems are acting together in a population with yellow
males, it is possible to identify them by hybridizing with napi carrying
neither flava nor sulphurea. The result in the F x should be white males
plus ochre-tinted females. This is the result obtained with both mar¬
ginalis and macdunnoughii , which therefore carry flava as well
as sulphurea.

Is the matter therefore to be considered closed? Perhaps not quite.
Though sulphurea and flava pigments are generally separable by eye,
chemical composition within each class may not be constant, and
genetic controls in these Pierids may involve more than two loci.

Moreover, some biological aspects of the situation remain puzzling.
Have these fainter colors any ecological effect? Do they offer clues to the
phylogeny of taxa carrying them? Bowden (1977) figured a female
macdunnoughii extremely reflective of U.V. light, indeed much more so
than marginalis , but this phenomenon needs wider investigation in
N.W. American Artogeia.

Specific Status

How distinct from Pieris napi L. is ssp. macdunnoughii ? And how
close to ssp. marginalis ? The pterin pigmentation systems of these two
subspecies are so similar as to suggest a close relationship. How do they
react to hybridization with the European species?

Male napi have seemed rather unwilling to pair with macdunnoughii
females, but one cage pairing occurred within a quarter of an hour. The
reciprocal pairing is easy. In either case fertility is very good.

But the three large broods 9 napi x cf macdunnoughii produced
no F ! daughters (the one female obtained showed no characters negating
a pure napi origin and was probably a waif). The one brood 1986-n from

26(l-4):l-288, 1988

87

9 macdunnoughii x c? napi yielded both sexes, but females were
the less numerous (40%) and generally appreciably smaller;
their fertility was not tested.

It is appropriate to compare the results for the marginalis x European
hybrids reported in 1970:

napi 9 x marginalis cf — > 1968-6, 7 cf + 1 9 + 15 undeveloped pupae,
marginalis 9 x napi cf — > 1968-a, 9 cf + 8 9 (incl. 1 cf after diapause),
Irish napi 9 x marg. cf — » 1966-/, 38 cf -I- 1 9 + 15 undeveloped pupae,
marginalis 9 x Irish cf — > 1968-y, 32 cf + 39 9-
They agree very well with those quoted above for macdunnoughii : on
the whole it can be said for both ssp. marginalis and ssp. macdunnoughii ,
the cross 9 napi x cf m. produces no viable female offspring — it is only
too likely that the single females in 1968-6 and 19 66-/ were waifs.

Such a sexual disturbance is more serious than a diapause disorgani¬
zation, and is normally sufficient ground for a specific separation (cf.
Lorkovic 1978). One could, however, expect marginalis and mac¬
dunnoughii to be mutually fully fertile and so probably conspecific. This
should be confirmed experimentally, if possible.

Subspecific Relationships

It will be necessary also to study sexual relations between these
subspecies and adjacent ones, especially microstriata Comstock and
uenosa Scudder. Shapiro (in litt. 1979) stated that microstriata seemed
to intergrade into marginalis in N. W. California. Geographic bound¬
aries between these taxa, as well as the other adjacent “subspecies”
mentioned by Eitschberger (1983), may be very uncertain (even inde¬
finite), and it can be unsafe for us to generalize from results obtained on
constituent local demes. Differences may be in part environmental (e.g.
in altitude) rather than genetic. It would be desirable to re-consider the
criteria for separating subspecies in this area.

Our (1981) experiments with ssp. microstriata were incomplete, as
acknolwedged at the time, and there was a mistake in the report (p. 3,
para. 5). This should read:

“One good female emerged on 20.V.79, and from this one individual all

hybrid broods were derived. About 1-2 hr. after caging with a British

sulphurea heterozygote napi male she paired with him ...”
However the argument is little affected and the conclusions are un¬
changed. The female hybrids included pale yellows with the fore wing
underside disc pale yellow; a sulphurea allele must have been present in
the white microstriata female used. This could be taken as suggesting at
least introgression from marginalis or macdunnoughii into spp. micro¬
striata , which could conflict with Eitschberger’s (1983) specific separa¬
tion of venosa and marginalis.

Acknowledgements. I have C.L. Remington, W.B. Watt, F.W. Chew and
particularly C.W. Nelson and A.M. Shapiro to thank for sending living material
of the two subspecies with which this paper is concerned.

88

J. Res. Lepid.

Literature Cited

BARNES, W. & J. H. MCDONNOUGH, 1916. Notes on N. American diurnal Lepidop-
tera. Contrib. Nat. Hist. Lepid. N. Amer. 3:57-59.

BOWDEN, S. R., 1954. Pieris napi L. f. hibernica Schmidt, eine kunstliche Aberra¬
tion? Mitt. ent. Ges. Basel 4:9-15, 17-22.

_ , 1962. Ubertragung von Pieris napi- Genen auf Pieris bryoniae durch

wiederholte Ruckkreuzung. Z. Arbeitsgemeinschaft osterr. Entomologen
14:12-18.

_ , 1963. Polymorphism in Pieris : forms subtalba Schima and sulphurea

Schoyen. Entomologist 96:77-82.

_ , 1970. Polymorphism in Pieris: f. sulphurea in Pieris napi marginalis.

Entomologist 103:241-249.

_ , 1977. Pieris — the ultra-violet image. Proc. Brit. ent. nat. Hist. Soc.

9:16-22.

_ , 1983. A palaeomorph of Artogeia ? — f. funebris Lorkovic. Proc. Brit,

ent. nat. Hist. Soc. 16:76-80.

EITSCHBERGER, U., 1983. Systematische Untersuchungen am Pieris napi-bryoniae
Komplex. Marktleuthen.

LORKOVIC, Z., 1971. Pieris napi (L.) morfa funebris, osebujna nova rekombinacija
krizanja. Acta entom. Jugoslav. 7:1-9.

_ , 1978. Types of hybrid sterility in diurnal Lepidoptera: speciation and

taxonomy. Acta entom. Jugoslav. 14:13-26.

REMINGTON, C. L., 1954. A new name for the Colorado race of Pieris napi. Lepid.
News 8:75.

WARREN, B. C. S., 1963. The androconial scales in the genus Pieris, 2: the Nearctic
species of the napi-growp. Entomol. Tidskr. 84:1-4.

Journal of Research on the Lepidoptera

26(l-4):89-97, 1988

The Mating System of Three Territorial Butterflies in
Costa Rica

John Alcock

Department of Zoology, Arizona State University, Tempe, AZ 85287

Abstract. Territorial behavior was observed in three Costa Rican
butterfly species whose males perch in sunspots and open areas along a
forest stream. Males of Calaenorrhinus approximatus occupied sunspot
clearings for 1-2 hr in the early morning, with the same individual
defending a site for up to 19 days. Males of Astraptes galesus cassius
perched on the broad leaves of a piper in a tree fall clearing by the
stream, with one individual returning for 17 days as the territory
holder. Males of Mesosemia asa asa perched in streamside vegetation
with a maximum territorial tenure of 25 days. Territorial males of all
three species regularly patrolled the area about their perching sites
and responded to intruders with circling chases and ascending pursuit
flights. One mating occurred in the territory of a C. approximatus male,
and three were recorded for M. asa in or near a perch territory. The
mating systems of these species appear convergent with other butter¬
flies whose males defend landmark territories.

Introduction

Although there is a growing literature on butterfly mating systems
(reviews by Ehrlich, 1984; Rutowski, 1982, 1984), little is known about
the mating tactics of tropical butterflies. Here I describe the territorial
and mate-locating behavior of two skippers ( Celaenorrhinus approxi¬
matus Williams & Bell and Astraptes galesus cassius Evans) and a
riodinid butterfly ( Mesosemia asa asa Hewitson), demonstrating that
these butterflies have convergently evolved a number of similarities in
their mating systems. I then compare their behavior with that of a
number of temperate-zone species with similar tactics.

Materials and Methods

Calaenorrhinus approximatus Williams & Bell

On the basis of two specimens, one taken outside the study period, the
skippers I studied were assigned to Celaenorrhinus approximatus , although
another very similar species, C. eligius , also occurs in Costa Rica (John Burrns,
pers. comm.). This skipper was observed at two locations separated by about 2
km along the Rio Guacimal, a 2-3 m wide and 0.5 m deep stream that flows
through lower montane wet forest at 1300 m in Monteverde, Costa Rica. The
study took place from 26 April to 14 July 1986 with the observer capturing and

90

J. Res. Lepid.

marking six males with Liquid Paper Typing Correction Fluid while relying on
natural wing tears to identify five other individuals. I observed sites occupied by
males beginning about 0800-0900. 1 recorded the identity of the male or males at
the location over a period of several hours, while also on some days noting the
number of male-male aggressive interactions, their duration, the number of
patrol flights made by the male perching in the area, and the time spent in each
flight.

Astraptes galesus cassius Evans

The study of this skipper took place in one of the two locations where C.
approximatus was observed. Nine males were captured and marked with Liquid
Paper during the period from 7 June to 30 July 1986. On all but 5 days I checked
the perching area at intervals during the morning to record the identity of the
male or males present. On some days the butterflies were observed continuously
for 15 min to 1 hr in order to record the frequency of social interactions, and the
duration of flights taken by the male perching at the site.

Mesosemia asa asa Hewitson

I watched this riodinid butterfly at two stream sites separated by about 150 m.
I captured and marked (with Liquid Paper) eight males; five other males were
identified through their distinctive patterns of wing damage. The sites were
monitored from 7 June to 30 July 1986 to record which males perched there. Some
days one or more males was selected for continuous observation for 15 min to 1
hr, during which time social interactions, patrol flights, and their duration were
noted.

The data collected provide a picture of the daily activity pattern and the
nature of territoriality of males of the three species. In addition, for two of the
species, C. approximatus and M. asa, incidental observations of male-female
interactions permit a description of mating behavior.

Means are presented ± 1 S.D.

Results

Daily Activity Pattern of Celaenorrhinus approximatus

On sunny days males flew to and perched upon leaves within sunspots
2— 4 m in diameter that were located within 10 m of the Rio Guacimal.
At one sunspot selected for special study between 1-14 May 1986, the
first male appeared between 0801-0906 (X=0834, N=10 days). At a
second sunspot 2 km upstream the first male arrived between 0836-0910
(N=4 days). On overcast mornings, arrival times could be delayed until
1100.

Once have perched, usually on a broad-leaved plant less than 0.5 m
from the ground, the male engaged in frequent patrol flights in which he
darted about the sunspot. At both sites, resident males not only
patrolled the sunspot at which they had arrived, but also occasionally
visited and perched in one to three othersunspots up to 30 m distant.
These patrol flights lasted from 3-32 sec (X=10.4±5.7 sec; N=60 flights

26(l-4):l-288, 1988

91

by 5 males). The frequency of these flights was 1.01/min based on a total
of 80 min of observations of three males on four days.

Typically males ceased flying about sunspots at some time between
0930-1030 on sunny days with the male alighting upside down under a
leaf after a flight about the sunspot. Flight activity averaged 76.3 ± 18.5
min (N=7 days).

Site Tenacity and Territoriality

I identified six individuals at the same sunspots on 4 to 10 dates with a
mean interval between first and last sighting of 12 ±3.5 days. Some of
the returning skippers stayed only briefly on any given day but others
occupied the site for most or all of the morning activity period. These
individuals, referred to hereafter as resident males, dashed after any
conspecifics that flew near them. The two males would then chase one
another in tight circles about the sunspot. Circle chases always preceded
ascending flights, in which the two combatants flew up and away from
the sunspot in near vertical flight into the forest canopy. Some interac¬
tions consisted of a series of circle and ascending flights, with the two
rivals descending separately after an ascent and returning to the
sunspot to repeat the cycle anew.

The frequency of aerial fights was 9.5/hr (N=10.9 hrs of observation).
The mean duration of these contests was 46.4±42.0 sec (N=34 fights
recorded on 11 days). Intraspecific chases lasted much longer on average
than either patrol flights (X = 10.4 sec, see above), or flights triggered by
other passing butterflies, tachinid flies and damselflies (X=4.7±2.1 sec,
N=24 chases recorded on 4 days).

In 66 of 85 fights, only one male returned to perch in the sunspot after
a circle flight or ascending flight. In 63 of 66 cases, the sole returning
male was the resident male who had occupied the site first that day.
When two males did land in the same sunspot after an aerial chase, it
was generally only a matter of a minute or two before one male patrolled
the sunspot eliciting a new chase and (usually) the departure of the
newcomer.

Ousters of territory-defending males were observed three times. On 1
May an unmarked male with fresh undamaged wings displaced yellow
after a long series of aerial chases. Twice males that had been residents
on preceding days arrived late at their sunspot to find the site occupied
by another male; in both cases turnovers occurred, with the previous
day’s resident quickly ejecting the newcomer.

Thus the general pattern was for one male of claim a perch site for all
of the morning activity period over several days. Other males, including
previous residents, might visit the site over a number of days but they
did not remain long.

Male-Female Interactions

I saw one copulation in about 20 hr of observation. This mating was
discovered after its inception when I noticed that the resident male was

92

J. Res. Lepid.

no longer perhcing in his sunspot. In his place was an unmarked
individual. The resident was under a broad leaf within his territory
copulating with an apparently fresh female with unworn wings. If the
mating occurred at about the time when the resident disappeared from
view, the duration of copulation would be about 1 hr.

Daily Activity Pattern of Astraptes galesus cassius

Males of A. g. cassius came in the morning to a stand of broad-leaved
Piper auritum (Piperaceae) growing at the edge of the Rio Guacimal. On
six sunny mornings between 7 June and 13 July, the first male to
perch on an exposed piper leaf appeared before 0830 (0808 was the
earliest record). On cloudy days arrivals were delayed, but one or more
males perched on the piper for at least brief periods on 30 of 36 days
during the study, even if the sun did not shine. Activity ceased in steady
rain but otherwise males remained at the site until some time between
1100-1200 (the latest record was 1157). Thus males were active 2-3 hr
per day.

The leaves used as perches were 2-3 m above the ground with the
primary perch site situated in the middle of a large (roughly 15 m
diameter) clearing created by tree falls. The piper received direct
sunlight only for a short period during the latter part of the morning.
Perching males often flew out from and returned in a few seconds to the
same leaf. Longer (10-20 m) patrol flights along the stream course lasted
from 4 to 25 sec with a mean of 10.2 ± 4.2 sec (N = 79 flights of 5 males on
7 days). The frequency of patrol flights was 39.3/hr (4.5 hr of observations
of 4 different males on 8 dates).

Site Tenacity and Territoriality

Marked individuals often returned to the piper patch over many days
and stayed for several hours each morning (Table 1). The resident male
chased intruders in straight horizontal pursuits and ascending flights
that were not as structured as the circle/ascending flights of C. approxi¬
mate. Visitors usually left the area quickly when pursued. Male-male
interactions occurred an average of 16.6 per hr (N = 5.5 hr observation
of 4 residents). Chases of conspecifics lasted an average of 25.3 ±12.6 sec
(N = 39).

A change in territory ownership occurred on 7 occasions (Table 1)
during 52 days. The new males all had less damaged wings than the old
residents that they replaced after a number of chases. Previous residents
continued to return to the site on subsequent days in 4 cases (Table 1),
often coming earlier than their replacements. For example, on 24 June,
green occupied the central piper plant from 0919-0952, giving way only
when yellow finally arrived {yellow having claimed the territory on 16
June). Although past residents were always quickly displaced by the
new owner, at least one male {yellow ) succeeded in reclaiming the site

Table 1 . Identified males of Astraptes galesus cassius at one site from 7
June to 28 July 1986. R=resident territory owner, v=non-
territorial visitor, R*= males loses territory to rival.

Male June

7 8

9

10

11 12

13 14

15 16

17

18

19

20

21 22 23

pink

R— -R*

V

V

V

green

R-

. R*

V

V

V

V

yellow o/o

R -

June 24 25 26 27 28

29 30

July

1 2 3

4

5

6

7

8 9 10 11

pink

V

green

R* v

V

V

V

V

V

V

yellow o/o

R— R*

V

V

R —

blue

R — -

-R

July

12 13

14

15

16 17

18 19

20 21

22

23

24

25

26 27 28

pink

green

yellow o/o

R* v

blue

wing mark

R . -

. R*

yellow ii

R — -

R*

wing notch

1

l-

for 12 more days (Table 1) when the male that had defeated him failed to
reappear on 30 June.

Daily Activity Period of Mesosemia asa asa

Males arrived at their streamside perches between 0930-1030 (N=12
days) and they departed on sunny afternoons between 1400-1500 (N=3
days) with one record of a male present until 1516. The butterflies
perched on exposed leaves 0.5 to 1.5 m above the ground, and they faced
out into the open space created by the stream channel. Early in the
morning males often selected sites that were in full sun, but they were
active even when all perches were shaded.

During 6 hr of watching five males, I recorded 14.3 perch shifting
flights of 1-3 sec per hr. Longer patrol flights of 3 m or more lasting on
average 9.1 ±4.6 sec (N=14) occurred at the rate of just 3.8 per hr during
this same period.

Territorial Behavior

Along a 150 m stretch of stream there were only four perching sites,

94

J. Res. Lepid.

one downstream at a bend in the Rio Guacimal, and three 10-15 m apart
at an upstream bend near the piper stand occupied by males of A.
g. cassius. The relatively open forest canopy at these places permitted
the sun to penetrate to a greater degree than elsewhere.

Marked males often returned to the same perch location day after day
(Table 2), chasing away intruders in slow circling flights, or occasionally
slow ascending flights, a series of which might last as long as 4 min.
Intraspecific aerial interactions occurred at the rate of just 5/hr based on
8 hr of observation. The mean duration of these interactions was 24.7 ±
25.1 sec (N=48 chases). Interspecific chases of certain other butterflies
lasted on average 5.9 ± 4.4 sec (N = 51 chases).

Turnovers in territory ownership took place six times during 43 days
at three different perch sites (Table 2). The most dramatic of these
occurred at a perch site that had been occupied for 25 days by a male that
had lost the tip of one forewing. On 12 July an unusually large, fresh
male with no wing damage displaced the resident from a perch site next
to clipped wing’s territory. The next day the large male moved into
clipped wing’s territory and engaged him in at least 16 aerial chases.
Clipped wing left the site but continued to return at intervals on this
and the next day, each time being chased off by the newcomer.

Male-Female Interactions

Three matings of M. asa were observed at 1138 on 8 June, 1205 on 24
June, and 1248 on 21 July. All occurred when a female flew into a
territory and was pursued by the resident male. The female then landed

Table 2. Territorial males of Mesosemia asa at four territories
observed from 18 June to 30 July. Note shifts between
territories 1 and 2 by male B.

June 18 19 20 21 22 23 24 25 26 27 28 29 30 July 1 2 3 4 5 6 7 8
Territory

T1 Male A

T2

T3 MaleC

T4

MaleB

MaleD

July 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30

T1 Male A - B . E F - G . -H -

T2 MaleB - 1 - B -

T3 Male C - 1 -

T4 Male D

26(1-4): 1-288, 1988

95

on the underside of a leaf with the male alighting beside her. Copulation
followed quickly with minimal courtship; “Tf” (Burns, 1970) was
exactly 52 min in one case, while copulation lasted between 40-75 min in
the two other cases.

Discussion

Table 3 provides a summary of male behavior in the three butterflies.
There are both differences and similarities in the mating system of
these unrelated species. The butterflies differ in the timing and dura¬
tion of their daily mate-locating activity, but each species is active for
only a fraction of each day. All three species perch in relatively open
areas, with Celaenorrhinus especially dependent on sunspots. These
areas are not only relatively free from visual obstruction so that males
may detect passing females, but may also offer thermoregulatory
advantages in cool rainforest habitats. Males of the two skippers patrol
their territories often, whereas the riodinid M. asa flies from perches
infrequently. Territorial males of all three species engage intruders in
ascending flights (although speed of flight varies greatly among species).
Residents are able to control their perching areas up to two weeks or
more, but they encounter receptive females at very low frequency.

Callaghan (1982(83)) has documented similar behavior in a number
of other tropical riodinids whose males wait on perches along the very
edge of forests or in treefall areas; Scott (1974, 1975, 1982) has observed
some North American skippers in which male mate-locating activity is
concentrated in gulches and ravine bottoms. In these cases, males
appear to take advantage of natural features that channel females past
certain points. Similarly, hilltopping species (Shields, 1967; Scott,
1970), seem to be making use of conspicuous topographic features that
may guide the movements of females. Landmark-based mating systems

Table 3. A comparison of the mating system of two territorial stream-
side skippers, Celaenorrhinus approximatus and Astraptes
galesus , and the riodinid butterfly, Mesosemia asa.

C. approximatus

A. galesus

M . asa

Activity period

< 2 hr
0830-1030

ca. 3 hr
0830-1130

ca. 4 hr
1000-1400

Patrol flights - frequency: mean duration

60/hr: 10 sec

39/hr: 10 sec

3.8/hr: 9 sec

Fights-frequency: mean

duration

9.5/hr: 46 sec

16.6/hr: 25 sec

5.5/hr: 25 sec

Observed matings

1

0

3

Maximum territorial tenure

19 days

21 days

25 days

96

J. Res. Lepid.

apparently evolve when males cannot profitably search for mates at
larval foodplants or adult nectar sources (Thornhill and Alcock, 1983).

Territoriality is widespread in the landmark and sunspot mating
system group (e.g. Shields, 1976; Davies, 1978; Bitzer & Shaw, 1979(80);
Callaghan, 1982(83); Lederhouse, 1982; Wickman & Wicklund, 1983;
Alcock, 1983, 1985; Alcock & O’Neill, 1986). Males typically compete for
landmark territories with elaborate circling and vertical ascending
flights (see also MacNeill, 1964). These interactions appear to involve
demonstrations of speed and aerial agility to rivals, as well as endur¬
ance in those cases in which repeated vertical pursuit flights occur.

Con vergent evolution is also evident in the behavior of females of this
group, which fly toward perched males, leading them on a slower, more
direct pursuit flight than that seen in male-male fights. Females that
are receptive alight quickly, and males land beside them to offer only
the briefest of courtships before copulation occurs (e.g. Alcock, 1985;
Alcock & Gwynne, in press; Wickman & Wiklund, 1983).

Acknowledgements. John Bums graciously agreed to identify the skippers, a
major task given the confused state of Celaenorrhinus systematics, and he also
reviewed the manuscript. William Haber assisted in species’ identification. Ron
Rutowski provided a useful critique of the manuscript, as did an anonymous
reviewer.

Literature Cited

ALCOCK, J., 1983. Hilltopping territoriality by males of the great purple hair-
streak, Atlides halesus (Lepidoptera: Lycaenidae): Convergent evolution
with a pompilid wasp. Behav. Ecol. Sociobiol. 13: 57-62.

- , 1985. Hilltopping in the nymphalid butterfly Chlosyne calif ornica

(Lepidoptera). Amer. Midi. Nat. 113: 69-75.

ALCOCK, J. & D.T. GWYNNE, In press. The mating system of Vanessa kershawi:
Males defend landmark territories as mate encounter sites. J. Res. Lepid.

ALCOCK, J. & K. O’NEILL, 1986. Density-dependent mating tactics in the Grey
hairstreak, Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. 209: 105-
113.

BITZER, R.J. & K.C. SHAW, 1979(80). Territorial behavior of the red admiral,
Vanessa atalanta (L.) (Lepidoptera: Nymphalidae). J. Res. Lepid. 18: 36-49.

BROWER, L.P., J.V.Z. BROWER, & F.P. CRANSTON, 1965. Courtship behavior of the
queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 50: 1-39.

BURNS, J.M., 1970. Duration of copulation in Poanes hobomok (Lepidoptera:
Hesperiidae) and some broader speculations. Psyche 77: 127-130.

CALLAGHAN, C.J., 1982(83). A study of isolating mechanisms among neotropical
butterflies of the subfamily Riodininae. J. Res. Lepid. 21: 159-176.

DAVIES, N.B., 1978. Territorial defence in the speckled wood butterfly ( Pararge
aegeria ): The resident always wins. Anim. Behav. 26: 138-147.

EHRLICH, P.R., 1984. The structure and dynamics of butterfly populations. Symp.
R. Ent. Soc. Lond. 11:25-40.

26(l-4):l-288, 1988

97

LEDERHOUSE. R.C., 1982. Territorial defense and lek behavior of the black
swallowtail butterfly, Papilio polyxenes. Behav. Ecol. Sociobiol. 10: 109-118.
MACNEILL, C.D., 1964. The skippers of the genus Hesperia in western North
America with special reference to California (Lepidoptera: Hesperiidae).
Univ. Calif. Publ. Entomol. 35:1-230.

RUTOWSKI, R.L., 1982. Mate choice and lepidopteran mating behavior. Fla. Ent.
65: 72-82.

- , 1984. Sexual selection and the evolution of butterfly mating behavior.

J. Res. Lepid. 23: 125-142.

SCOTT, J.A., 1970. Hilltopping as a mating mechanism to aid the survival of low
density species. J. Res. Lepid. 7: 191-204.

- , 1974. Mate-locating behavior of butterflies. Amer. Midi. Nat. 91: 103-

117.

- , 1975. Mate-locating behavior of North American butterflies. J. Res.

Lepid. 14: 1-10.

- , 1982(83). Mate-locating behavior of western North American butter¬
flies. II. New observations and morphological adaptations. J. Res. Lepid. 21:
177-187.

SHIELDS, O., 1967. Hilltopping. J. Res. Lepid. 6: 69-178.

THORNHILL, R. & J. ALCOCK, 1983. The evolution of insect mating systems.

Harvard University Press, Cambridge, Mass. 547 pp.

WICKMAN, P.-O. & C. WIKLUND, 1983. Territorial defence and its seasonal decline in
the speckled wood butterfly (. Pararge aegeria). Anim. Behav. 31: 1206-1216.

Journal ofResearch on the Lepidoptera

26(l-4):98-108, 1988

Stratification of fruit-feeding nymphalid butterflies in a
Costa Rican rainforest

P.J. De Vries

Dept, of Zoology, University of Texas, Austin, Texas, 78712
and

Smithsonian Tropical Research Institute, Box 2072, Balboa, Panama, Central America

Abstract. 1) Paired traps showed that fruit-feeding nymphalid butter¬
flies in the subfamilies Nymphalinae, Charaxinae, Morphinae and
Satyrinae are stratified between the canopy and the understory by
species composition, and abundance, size, and color pattern. 2) Short
wing lengths and uniform underside patterns are found in the canopy,
whereas long wing lengths and underside patterns bearing eyespots
are found in the understory. 3) Wing length and color pattern cannot be
separated from taxonomic affinity, and hence, these butterflies stratify
by subfamily: Charaxinae and Nymphalinae in the canopy, Morphinae
and Satyrinae in the understory. 4) A general model is presented to
explain the apparent breakdown of stratification along forest edges and
how light levels act as barriers to maintain insect stratification.

Introduction

The tropical rainforest has been described in terms of component
layers or strata (Richards, 1966), and stratification (i.e. vertical distri¬
bution) of rainforest organisms has been documented for mammals and
birds (Allee, 1926; Dunn et al., 1968; Orians, 1969; Pearson, 1971), and
for insects (Bates, 1944, 1947; Corbet, 1961a & b; Davis, 1944; Elton,
1973, 1975; Erwin, 1983; Erwin & Scott, 1980; Garnham et al., 1946;
Galino et al., 1951; Haddow, 1945; Haddow & Corbet, 1961a; Jackson,
1961; Papageorgies, 1975; Pittendrigh, 1950a & b; Rees, 1984; Snow,
1955; Sutton, 1979, 1984; Sutton & Hudson, 1980; Wolda, 1979).

Studies on the stratification of insects other than mosquitos range
from general overviews of all insect taxa taken in a sample (Elton, 1973;
Sutton, 1984; Sutton & Hudson, 1980) to comments on a few taxa within
a sample (Corbet, 1961b; Erwin, 1984; Erwin & Scott, 1980; Jackson,
1961; Papageorgis, 1975; Rees, 1984; Wolda, 1979). Of these studies,
only two have attempted to quantify the stratification of butterflies.
Jackson (1961) documented the presence of rare Lycaenidae and
Nymphalidae in a Ugandan forest canopy, but his study ignored all
species in the understory. In Peruvian lowland rainforest, Papageorgis
(1975) found that some warningly colored butterflies and diurnal moths
tend to fly at different levels in the forest according to mimetic pattern.

26(1-4): 1-288, 1988

99

Excluding those species where males visit wet sand or plant material
for non-nutritional resources (see Boppre, 1984; Collenette & Talbot,
1926; Norris, 1936), any tropical forest community of butterflies can be
divided into two adult feeding guilds: those species that obtain the bulk
of their nutritional requirements from flower nectar (all Papilionidae,
Pieridae, Lycaenidae, Riodinidae and some Nymphalidae), and those
species that feed upon the juices of rotting fruits, fermenting sap, or
animal waste (several subfamilies of the Nymphalidae (sensu Ehrlich,
1958)). In the neotropics, only the members of the nymphalid sub¬
families Satyrinae, Morphinae, Charaxinae, and some members of the
Nymphalinae feed exclusively on rotting fruits or other non-floral liquid
as adults. These latter subfamilies, hereafter referred to as “fruit-
feeding nymphalids,” may account for over 50% of the nymphalid
species diversity in some Central American habitats and can, in
general, only be collected by baiting them with rotting fruits (DeVries,
1987).

By virtue of their feeding habits, fruit-feeding nymphalids may be
used to study stratification because individuals can be selectively
sampled with traps. In this paper I present quantitative evidence for
several patterns of stratification among these butterflies from a Costa
Rican rainforest, and discuss how forest structure may effect stratifi¬
cation.

Methods

The study was conducted from 20 October 1979 through 2 January 1980 at Finca
La Selva, Heredia Province, Costa Rica, within the area known locally as the
‘Washington Plots.’ Five trapping sites (see trap design in DeVries, 1987) within
closed canopy forest, each of which had an emergent canopy tree with a small
lightgap at its base, were chosen on the basis of their receiving at least one hour
of direct sunlight each day. The entire study was done during the rainy season, a
time when butterfly abundance is low.

One trap of each pair was placed in the canopy, the other in a small lightgap
immediately below it. Canopy traps were positioned by fastening a pulley to a
tree limb growing over a lightgap and using a rope to raise and lower the trap
from ground level. Traps were checked twice each day, and rotting banana bait
replaced regularly. All butterflies caught in the traps were killed, measured
(winglength), and determined to species and sex and the trap position of capture
was noted. Winglength data (measured from base of forewing to the forewing
apex) were supplemented for species with small sample sizes using Costa Rican
specimens from the Museo Nacional or British Museum (Nat. Hist.) collections
(Table 3); these data were log transformed for analysis. The nomenclature used
here follows DeVries (1987), and for analyses the subfamily (Table 2) Brassolinae
was collapsed into the Morphinae of Ehrlich (1958).

Results: Patterns of Stratification

The wet season depression of butterfly abundance is reflected by the low
numbers of individuals trapped: in 10 weeks the traps collected a total of

100

J.Res.Lepid.

182 butterflies in 46 species (Table 1). As one might expect, some rare
species (based on museum abundance) were common in the canopy, and
in all categories (by subfamily and trap position) significantly more
(DF = 4; G = 49.1; p < .001, DF = 1; G = 42.3; p < .001 respectively)
males were caught than females (Table 1). The trap samples also
contained previously undescribed taxa (see Singer et al. 1983; DeVries
1987). Without addressing the problems of trap effect, heterogeneity of
trap catch, or the possible effects of sampling without replacement,
these overall patterns were noted:

1. Most species tended to be trapped only in the canopy or the
understory, but a few species were found in both (Table 1).

2. Species richness was about the same in canopy and understory,
with 24 taxa trapped only in the canopy, 15 only in the understory, and 7
taxa in both (Table 1).

3. Canopy taps collected significantly more individuals than did
understory traps (Table 1).

4. Stratification occurred at the subfamily level, with members of the
Charaxinae and Nymphalinae in the canopy, and members of the
Morphinae and Satyrinae in the understory (Table 2).

5. Species trapped in the canopy had smaller mean winglengths than
those species trapped in the under story (Table 3).

6. Mean winglengths differ between subfamilies, implying that
winglength and position of capture cannot be separated from phylo¬
genetic affinity (Table 3).

7. Canopy and understory butterflies differ in possession of eyespot
patterns (Table 4), and these differences are linked to taxonomic
affiliation: Morphinae and Satyrinae have eyespots while other groups
generally do not.

Discussion

This study showed that certain genera and species of fruit-feeding
nymphalid butterflies were trapped consistently in the canopy, others in
the understory, while a small fraction of the species were found in both
canopy and understory. Overall, the data here indicate differences
between canopy and under story butterflies in abundance, species com¬
position, wing length, and color pattern (Tables 1-4). However, the
stratification of butterfly species by wing length and color pattern
cannot be separated from taxonomic relatedness. This is to say that
position of capture, size, color pattern, and subfamily are correlated
to some degree, and that similar patterns may be found in other
arthropods.

The winglength data presented here are consonant with size data
from other arthropod studies (Wolda 1979; Erwin & Scott 1980; Rees
1982; Erwin 1983), suggesting that smaller relative size may be a
general characteristic of canopy insects. This trend, however, is reversed

26(l-4):l-288, 1988

101

for Costa Rican Papilionidae, where larger winged butterflies occur in
the canopy (DeVries, unpublished data).

Stratification by color pattern in fruit-feeding nymphalids is not
likely to be explained by the mimetic resemblance hypothesis of
Papageorgis (1975) per se. In her system, predators maintain the
stratification of butterflies by selecting for similar mimetic patterns
within distinct strata. However, virtually all of the species in the
present study are cryptically colored, palatable to predators, and non-
mimetic (Chai 1986; DeVries 1987). Since predators are clearly import¬
ant in selecting the appearance of cryptic insects (Kettle well 1955,
1956; Chai 1986), the stratification of eyespots (or lack of them) found in
this study may also be due to stratification patterns of the butterflies’
predators. It is quite reasonable to assume that the species composition
of vertebrate predators (i.e., lizards and birds) differs between the
canopy and understory, and that these predators exert differing selec¬
tion pressures on butterflies. Perhaps studies on a single subfamily that
contains species found in both canopy and in the understory (e.g.,
Nymphalinae or Satyrinae) may prove fruitful for probing the effects of
how predator communities in the canopy and understory act on eyespot
pattern (and body size) of these cryptic butterflies.

Although the data here show that fruit-feeding nymphalid species are
stratified between canopy and understory when feeding, they do not
necessarily indicate where these butterflies spend their time when not
feeding. For instance, the males of some species trapped only in the
understory (Archaeoprepona Camilla, Morpho cypris ) spend much of
their time patrolling in the canopy (presumably searching for females),
and conversely, females of some rarer canopy species ( Cissia pseudo -
confusa, Megeuptychia antonoe, Catonephele orites) are known to ovi¬
posit on hostplants occurring near ground level in gaps and along forest
edges (DeVries 1986; 1987). Clearly, the location of mate seeking areas
or larval hostplants can be entirely different from where non-ovipositing
adults are found. Furthermore, these data here do not indicate whether
or not further stratification would be revealed if traps had been placed at
intermediate levels between the canopy and under story. The data do,
however, raise the question of why these butterflies show pronounced
stratification: if Newton was correct about apples, rotting fruits fall to
the ground and canopy butterflies should feed on them there. This
suggests that fruit-feeding nymphalids may eventually be shown to
have feeding specializations with respect to rainforest fruit species.

Most tropical collectors are aware that canopy flying nymphalids can
be trapped close to ground level along a forest edge. Such knowledge
implies an intuitive appreciation that stratification breaks down in
some situations. A testable, general model is offered here to explain how
different light levels maintain the stratification observed in fruit¬
feeding nymphalids, and why stratification is less pronounced in certain
habitats. The model assumes that for diurnal insects such as butterflies,

Table 1. Summary of taxa trapped during the study. See DeVries (1987) for
nomenclatural details.

Species Canopy Understory

CHARAXINAE

Prepona

omphale 6 1

Agrias

amydon 1 0

Archaeoprepona

demophon 2 1

Camilla 1 4

meander 0 1

Zaretis

itys 1 0

Memphis

morvus 5 0

cleomestra 3 0

laura 1 0

aureola 1 0

xenocles 3 0

NYMPHALINAE
Hamadryas

laodamia 21 0

arinome 16 2

amphinome 3 0

Catonephele

numilia 4 0

orites 11 0

Nessaea

aglaura 1 4

Myscelia

leucocyana 5 3

cyaniris 1 0

Eunica

monima 1 0

Callicore

lyca 1 0

patelina 2 0

Historis

odius 3 0

acheronta 3 0

Smyrna

blomfildia 3 0

Colobura

dirce 8 0

Tigridia

acesta 2 2

Total

7

1

3

5

1

1

5

3

1

1

3

21

18

3

4
11

5

8

1

1

1

2

3

3

3
8

4

26(l-4):l-288, 1988

103

Table 1. (cont’d)

Species

Canopy

Under story

Total

MORPHINAE

Morpho

peleides

0

1

1

amathonte

0

1

1

cypris

0

1

1

Antirrhea

miltiades

0

1

1

Caligo

eurilochus

0

4

4

atreus

0

9

9

illioneus

0

2

2

Catoblepia

orgetorix

0

1

1

Opsiphanes

tamarindi

0

1

1

invirae

2

0

2

cassinae

2

0

2

SATYRINAE

Cithaerias

menander

0

2

2

Dulcedo

polita

0

1

1

Cissia

pseudoconfusa

3

0

3

joycae

1

0

1

hesione

0

3

3

Megeuptychia

antonoe

11

0

11

Taygetis

Andromeda

0

5

5

xenana

1

3

4

Total

129

53

182

male-female

male-female

Subfamily

canopy

understory

Total

Charaxinae

19:5

5:2

31

Nymphalinae

59:26

7:4

96

Morphinae

2:2

15:6

25

Satyrinae

13:3

14:0

30

Total

129

53

182

104

J.Res.Lepid.

Table 2. Abundance of individuals by subfamily and position of traps. Expected
values are in parentheses. Significantly more butterflies were trapped
in the canopy than the understory [DF = 3; G = 54.67; p. < .0001].

Subfamily

Canopy

Under story

Total

Charaxinae

24 (21.97)

7 (9.03)

31

Nymphalinae

85 (68.04)

11 (27.96)

96

Morphinae

4 (17.72)

21 (7.28)

25

Satyrinae

16 (21.26)

14 (8.74)

30

Total

129

53

182

Table 3. Mean Winglengths based on Costa Rican Specimens

Species

N =

Winglength

Subfamily

Position

omphale

7

48.4

charax

both

demophon

9

55.5

charax

both

Camilla

6

59.5

charax

both

meander

8

53.7

charax

under

itys

7

35.1

charax

canopy

morvus

8

32.4

charax

canopy

cleomestra

12

32.8

charax

canopy

aureola

7

35.4

charax

canopy

xenocles

7

29.5

charax

canopy

laodamia

8

35.0

nymph

canopy

arinome

8

37.0

nymph

both

amphinome

9

37.7

nymph

canopy

numilia

7

36.2

nymph

canopy

orites

8

34.1

nymph

canopy

aglaura

8

35.9

nymph

both

leucocyana

9

29.3

nymph

both

cyaniris

10

34.1

nymph

canopy

monima

10

21.8

nymph

canopy

lyca

10

25.7

nymph

canopy

patelina

8

28.7

nymph

canopy

odius

8

56.0

nymph

canopy

acheronta

5

44.0

nymph

canopy

blomfildia

8

41.2

nymph

canopy

dirce

7

32.5

nymph

canopy

acesta

10

25.7

nymph

both

peleides

12

71.3

morph

under

amathonte

10

78.1

morph

under

cypris

8

70.0

morph

under

miltiades

9

47.3

morph

under

eurilochus

9

81.6

morph

under

atreus

8

77.7

morph

under

26(l-4):l-288, 1988

105

Table 3. (cont’d)

Species

N =

Winglength

Subfamily

Position

illioneus

6

69.9

morph

under

orgetorix

10

51.8

morph

under

tamarindi

13

48.9

morph

under

invirae

8

43.0

morph

canopy

cassinae

11

42.0

morph

canopy

menander

10

30.3

satyr

under

polita

6

34.4

satyr

under

pseudoconfusa

8

20.9

satyr

canopy

hesione

7

20.8

satyr

under

antonoe

7

31.9

satyr

canopy

andromeda

8

37.0

satyr

under

xenana

9

36.4

satyr

under

One Factor ANOVA on Winglength across trap position and subfamily.

Source

S.S.

DF

Mean sq.

F-test

between traps

1.343

2

0.672

6.716

within traps

4.000

40

0.100

p < .005

Total

5.344

42

between subfams

2.935

3

0.978

15.84

within subfams

2.409

39

0.062

p < .0001

Total

5.344

42

Table 4. Stratification of species by presence or absence of eyespot pattern.

Expected values are in parentheses. Eyespot patterns are found
with a significantly greater frequency in the understory than in the
canopy (DF = 1; G = 25.23; p = .0001). Note that this cannot be
separated from taxonomic affinity: Morphinae and Satyrinae have
eyespots.

Eyespots

Canopy

Understory

Both

Total

Present

5 (9.39)

13 (5.87)

0 (2.74)

18

Absent

19 (14.61)

2 (9.13)

7 (4.26)

28

Total

24

15

7

46

light is more important than the related factors of temperature and
humidity for explaining patterns of stratification. The model therefore
predicts that: 1) butterfly taxa usually fly in certain light levels within
any habitat, and that 2) drastic changes in light intensity act as
barriers between habitats.

106

J. Res. Lepid.

The flower-feeding butterflies Anartia fatima (Nymphalidae),
Phoebis philea (Pieridae), Battus polydamas (Papilionidae) that usually
fly in bright sun, at ground level, provide an illustration of the effects of
a light barrier. I commonly see these butterflies fly across a pasture
towards the forest edge, ascend at the forest edge, fly across the canopy,
and descend once again to ground level when the next pasture is
encountered. They do not move through the shade of the forest, but
rather they treat the canopy as an elevated pasture, despite the 40
meter difference in height be ween canopy and ground levels. In this
example, a butterfly changes vertical distance from the ground without
experiencing an appreciable change in light intensity. I strongly suspect
that butterfly species that inhabit the vegetational interface between
sunny and shaded areas (i.e., canopy /edge) treat the forest canopy and
forest edge without regard to vertical position, since light levels within
the interface should remain roughly the same regardless of height.

Stratification then, in both canopy and pasture species is probably
maintained by their preference for certain light levels. From field
observations I further reason that there are three major distribution
zones for butterflies within a closed canopy forest: open areas above and
around the canopy (high light levels), the combination of within canopy
and forest edge (medium light levels), and the shade of the forest
interior (low light levels). Canopy species can be trapped at ground level
at the forest edge because they normally inhabit the light level interface
between bright sunlight and deep shade, and like vining plants, treat
the forest edge as the canopy come to the ground. Light is considered to
be an important factor in maintaining stratification in some forest and
marine communities (Bainbridge et al. 1966; Allee et al. 1969), yet the
effect the forest edge or disturbed forests have on stratification has not
been addressed in tropical forest insects. If differences in light levels are
important for maintaining stratification in rainforest butterflies, we
might predict that in habitats without pronounced differences in light
levels (i.e., disturbed forest, in deciduous forest during the dry season, or
along the forest edge), stratification will not be as distinct as in closed
canopy forest. The study of fruit-feeding nymphalids across various
forest successional stages with the methods described herein may
provide the necessary tools for understanding the role of light levels and
forest structure in the maintainence of stratification of rainforest
butterflies.

Acknowledgements. This manuscript was improved over the years by comments
and discussion provided by T. M. Aide, R. Brown, N. Greig, D. Grimaldi, D. J.
Harvey, R. Lande, E. Leigh, M. Singer, and J. Zimmerman. For field-related
discussion and help thanks go to I. Chacon, J. Gamboa, D. Perry, D. H. Janzen,
F. G. Stiles, and the 1979-80 denizens of Finca La Selva. A special thanks is due
to D. Feener and D. Ng for statistical consultation, and to N. Greig for critical
comments. Portions of this study were supported by a Fulbright Hayes fellow¬
ship, a Smithsonian predoctoral fellowship, the Museo Nacional de Costa Rica,

26(1-4): 1-288, 1988

107

and the University of Texas at Austin. This paper is dedicated to the stratified

diversity of Clifford Brown, Horace Silver, and Art Blakey.

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ALLEE, W. C., A. E. EMERSON, O. PARK, T. PARK, & K. P. SCHMIDT, 1969. The Principles
of Animal Ecology. W. B. Saunders, Philadelphia.

BAINBRIDGE, R„ G. C. EVANS, & O. RACKHAM (EDS ), 1966. Light as an Ecological
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BATES, M., 1944. Observations on the distribution of mosquitos in a tropical
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BATES, M., 1947. The stratification of mosquitos in cages. Ecology 28:80-81

BOPPRE, M., 1984. Chemically mediated interactions between butterflies. Sym.
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CHAI, P., 1986. Field observations and feeding experiments on the responses of
rufous-tailed jacamars (Galbula ruficauda) to free-flying butterflies in a
tropical rainforest. Biol. J. Linn. Soc. 29:161-189.

COLLENETTE, C. L. & G. TALBOT, 1928. Observations on the bionomics of the
Lepidoptera of Matto Grosso, Brasil. Trans. R. Ent. Soc. Lond. 76:391-416.

CORBET, P. S., 1961a. Entomological studies from a high tower in Mpanga forest,
Uganda. IV. Mosquito breeding at different levels in and above the forst.
Trans. Roy. Ent. Soc. London 113:275-283.

CORBET, P. S., 1961b. Entomological studies from a high tower in Mpanga forest,
Uganda. XII. Observations on Ephemeroptera, Odonata and some other
orders. Trans. Roy. Ent. Soc. London 113:356-361.

DAVIS, D. E., 1944. A comparison of the mosquitos captured with an avian bait at
different vegetational levels. Rev. Ent. Rio de Janeiro 15:209-215.

De VRIES, P. J., 1986. Hostplant records and natural history notes on Costa Rican
butterflies (Papilionidae, Pieridae and Nymphalidae). J. Res. Lep.
4:290-333.

De VRIES, P. J., 1987. The Butterflies of Costa Rica and their Natural History.
Princeton University Press, Princeton.

DUNN, F. L., B. L. LIM, & L. F. YAP, 1968. Endoparasite patterns in mammals of the
Malayan rainforest. Ecology 49:1179-1184.

EHRLICH, P. R., 1958. The comparative morphology, phylogeny and higher
classification of the butterflies (Lepidoptera: Papilionoidea). Kansas Univ.
Sci. Bull. 39:305-370.

ELTON, C. S., 1973. The structure of invertebrate populations inside neotropical
rainforest. J. Animal Ecol. 42:55-104.

ERWIN, T. L., 1983. Beetles and other insects of tropical forest canopies at Manaus,
Brazil, sampled by insecticidal fogging. In Tropical Rain Forest: Ecology and
Management , S. L. Sutton, T. C. Whitmore, & A. C. Chadwick (eds.). British
Ecol. Soc. Spec. Publ. no. 2.

ERWIN, T. L. & J. C. SCOTT, 1980. Seasonal and size patterns, trophic structure, and
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GALINO, P. S., J. CARPENTER, & H. TRAPIDO, 1951. Ecological observations on forest

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Medicine 31:98-137.

GARNHAM, P. C„ J. O. HARPER & R. B. HIGHTON, 1946. The mosquitos of the Kaimosi
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HADDOW, A. J., 1945. The mosquitos of Bwamba County, Uganda. II. Biting
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HADDOW, A. J. & P. S. CORBET, 1961a. Entomological studies from a high tower in
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Ent Soc. London 113:284-300.

JACKSON, T. H. E., 1961. Entomological observations from a high tower in Mpanga
forest, Uganda. IX. Observations of Lepidoptera (Rhopalocera). Trans. Roy.
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KETTLEWELL, H. B. D., 1956. Further selection experiments on industrial melan¬
ism in Lepidoptera. Heredity 10:287-301.

NORRIS, M. J., 1936. The feeding habits of the adult Lepdidoptera, Heteroneura.
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50:783-801.

PAPAGEORGIS, C., 1975. Mimicry in tropical butterflies. Amer. Scientist 63:522-
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PITTENDRIGH, C. S., 1950a. The ecoclimatic divergence of Anopheles bellator and
A. homunculus. Ecology 4:43-63.

PITTENDRIGH, C. S., 1950b. The ecotopic specialization of Anopheles homunculus
and its relation to competition with A. bellator. Ecology 4:64-78.

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SINGER, M. C„ P. J. DeVRlES, & P. R. EHRLICH, 1983. The Cissia confusa species group
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48:512-521.

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SUTTON, S. L., 1984. The spatial distribution of flying insects in tropical rain
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SUTTON, S. L. & P. J. HUDSON, The vertical distribution of small flying insects in the
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Journal of Research on the Lepidoptera

26(1-4):109-115, 1988

Euphydryas anicia and E. chalcedona in Idaho
(Lepidoptera: Nymphalidae)1

Clifford D. Ferris

Bioengineering Program, University of Wyoming, Laramie, Wyoming 82071,
P.O. Box 3351 University Station, Laramie, Wyoming 82071-3351

Abstract. The nymphalid species Euphydryas anicia and E. chalcedo¬
na are sympatric in several areas in Idaho and easily separated from
one another by features of the male genitalia and differences in
forewing maculation. While anicia and chalcedona are closely related,
distinct differences can be identified, thus substantiating their separa¬
tion as two species.

Introduction

The taxa Euphydryas anicia (Doubleday) and E. chalcedona (Double¬
day) have been considered historically to represent separate species,
and they have been so treated in the two most recent lists of the North
American butterflies (Miller & Brown, 1981; Hodges et al., 1983), as
well as in two recent papers (Brussard et al., 1985; Spomer & Reiser,
1985). Based upon his analysis of the male genitalia of these insects,
Scott (1978 [80]) considered these two taxa conspecific. My field studies
in Idaho indicate that E. anicia and chalcedona occur sympatrically at
several localities, and that they may be separated on the basis of
characters in the male genitalia and dorsal wing maculation.

Following the findings of Brussard et al. (1985), the generic name
Euphydryas is used in this paper rather than Occidryas Higgins.

Study Areas

Fig. 1 is a map showing the distributions of Euphydryas anicia and E.
chalcedona in Idaho. The records are based on my own collecting and
data provided by Stanford (1985). Both species occur widely in Idaho,
but to date they have been found to be truly sympatric in two localities
only. The first site is in Boundary Co. and lies east of Hwy. 95 and
northwest of Moyie Springs in the Kaniksu National Forest. Both E.
anicia and E. chalcedona were collected on July 6—7, 1985 flying
together in a small open meadow near a railroad right-of-way.

The second site is in the Bear Valley region of Valley Co. in the

1 Published with the approval of the Director, Wyoming Agricultural Experiment Station as Journal
Article No. JA 1390.

110

J.Res.Lepid.

Fig. 1 Idaho map showing distribu¬
tion by county of E. anicia
(triangles) and E. cha/cedona
(solid circles).

vicinity of the Deer Creek crossing of the Boise National Forest road
that connects Warm Lake with Hwy. 12. Specimens of the two species
were taken together along the road and on an adjoining cleared forest
slope on July 24, 1984. Elsewhere in Valley Co., chalcedona occurs in
the Payette National Forest just north of McCall, and anicia is found in
the general vicinity of Warm Lake and Stolle Meadow.

Study Material

During this study, 283 Idaho specimens (106 anicia , 175 chalcedona , 2
equivocal) were examined.

Wing Maculation

Euphydryas chalcedona throughout its range in Idaho is relatively
constant in facies. Adults are dor sally predominately black with red-
and-white maculation; ventrally the ground color is brick red. The
subspecific epithet usually applied is wallacensis Gunder (= huelle-
manni dos Passos). Two typical pairs are shown in Fig. 2.

Euphydryas anicia in Idaho is variable. In the broad sense, it can be
divided into two color groups: 1. dorsal ground color generally black; 2.
dorsal ground color red/orange. Both forms manifest pale white or
cream-colored spots dor sally. The dark form displays red markings
similar to those of chalcedona. The ventral ground color of Idaho anicia
varies from brick red to red-orange. The VHW pale intercellular
maculation varies in extent according to local colony. Because of the
variability of anicia in Idaho, no subspecific epithets are applied in this
paper.

26(l-4):l-288, 1988

111

Fig. 2. E. cha/cedona wa/lacensis from Idaho. Dorsal surfaces left; ventral
right. A. Male. Payette Nat. For., 1 mi. N. McCall, Valley Co., 1.vii.83.
B. Female. Same locality, 13.vii.84. C. Male. Canyon Creek, Idaho Co.,
31.V.85. D. Female. Kaniksu Nat. For. NW Moyie Springs, Boundary
Co., 7.vii.85. All C. D. Ferris Coll.

Specimens of anicia collected at the Boundary Co. site are brightly
colored and belong to the red/orange category. Two color forms of anicia
occur in Valley Co.: 1. a red form similar to the Boundary Co. phenoty¬
pe, but not so brightly colored; 2. specimens that generally fall into the
black category, but variable.

Some of the dark anicia specimens appear in maculation superficially
similar to chalcedona. These two species can be separated, however, on
the basis of the form of the marginal spot-band on the DFW, as
illustrated in Fig. 3. In both sexes of anicia , the DFW marginal spots
form a complete border along the outer edge of the wing. In chalcedona
wallacensis , the spot-row normally terminates about vein Cu! in the
males, and although it frequently extends to vein 2A in the females,
the spots are much reduced in size below vein Cu^ This spot-row in
anicia is clearly double at the apex of the FW, while in chalcedona
immediately basad of the marginal red spot-row, there is a blackish
band distad of a row of small whitish spots. The apical double spot-row
in anicia may present concolorous spots, or the inner row may contain
whitish spots. Ventrally, especially on the HW, chalcedona generally

112

J.Res.Lepid.

Fig. 3. DFW marginal spot-band maculation in Idaho E. anicia and chalcedo-
na.

displays much more extensive brick-red coloration than is found in
anicia , but this character is not relaible. Fig. 4 depicts specimens of E.
anicia from the Boundary Co. (A-B) and Valley Co. (C-D) populations.
Fig. 5 illustrates a pair of the dark phenotype from Stolle Meadow in
Valley Co.

Genitalic Studies

Fig. 6 depicts the diagnostic portions of the male genitalia of E. anicia
and E. chalcedona from Idaho. In this figure, drawings A and E
represent typical chalcedona , while B-D and F represent typical anicia
(G will be addressed subsequently). These illustrations show the distal
portion of the right valve and its processes with the abdomen rotated
90° from normal life position. This is the view of the genitalia seen
using a binocular dissection microscope after the abdominal hairs have
been removed (by use of a stiff brush). It is not necessary to dissect the
genitalia from the abdomen. To achieve the views shown in Fig. 6, it
may be necessary to angle the specimen relative to the microscope
objective.

As the illustrations show, E. chalcedona manifests one long and
curved process, and a short pointed process (which may be slightly
recurved). By contrast, anicia presents two long curved processes
(frequently of nearly equal length) which various authors have likened
to knitting needles. In my studies of males of anicia collected from
Chihuahua, Mexico to the southern Yukon Territory, the valvular
processes are remarkably consistent in form, and do not intergrade into
the form found in chalcedona. The two species, however, may hybridize
as discussed below.

Intermediate Specimens

Of the Idaho material, only one pair of intermediate specimens has
been found. They are illustrated in Fig. 7, with male genitalia in Fig. 6
(G). In dorsal maculation, both specimens are closer to chalcedona than
anicia , although the form of the male genitalia is closer to anicia. This
pair may represent a hybrid between the two species, but more likely it

Fig. 4. E. anicia from Idaho. Dorsal surfaces left; ventral right. A. Male,
red/orange form. Kaniksu Nat. For. NW Moyie Springs, Boundary
Co., 6.vii.85. B. Female. Same data. C. Male, red/orange form. Deer
Creek, Boise Nat. For., Valley Co., 24.vii.84. D. Female. Same data.
All C. D. Ferris Coll.

Fig. 5. E. anicia dark form from Stolle Meadow, Boise Nat. For., Valley Co.,
Idaho, 13-17.vii.83. A. Male, dorsal. B. Male, ventral. C. Female,
dorsal. D. Female, ventral. All C. D. Ferris Coll.

114

J.Res.Lepid.

E ^

B

F

\

C

G

^ J\

D ^

A = chalcedona, IDAHO CO., IDAHO

B = anicia, BOUNDARY CO., IDAHO

C = anicia, BOUNDARY CO., IDAHO

D = anicia. VALLEY CO., IDAHO

E = chalcedona, VALLEY CO., IDAHO

F = anicia, VALLEY CO., IDAHO

G = intermediate, VALLEY CO., IDAHO

Fig. 6 Male genitalia
(valvular processes)
of Idaho Euphydryas.
A, E. E. cha/cedona.
B-D, F. E. anicia.
G. Equivocal specimen.

Fig. 7. Equivocal Euphydryas pair from Deer Creek, Boise Nat. For., Valley
Co., Idaho, 24.vii.84. A. Male, dorsal. B. Male, ventral. C. Female,
dorsal. D. Female, ventral. All C. D. Ferris Coll.

26(l-4):l-288, 1988

115

is an extreme variant of the anicia phenotype. Occasional dark indi¬
viduals occur in many anicia populations, perhaps as a consequence of
thermal shock during the prepupal stage.

Various authors have suggested that paradoxa McDunnough (usually
referred to chalcedona) represents a stable hybrid between chalcedona
and anicia. It is not the intent of this paper, however, to review species
that occur outside of Idaho.

Conclusion

On the basis of my studies in Idaho and the data presented above, I
conclude that the taxa anicia and chalcedona represent closely related
but separate species. They can be distinguished by differences in the
male genitalia and FW maculation. Their flight periods overlap, with
chalcedona on the wing from mid-May into late July depending upon
geographic location and elevation. E. anicia generally appears in early
July and survives into August at suitable elevation.

Acknowledgements Prof. Nelson S. Curtis, Moscow, Idaho, kindly directed
the author to some of the collection sites visited. Dr. Richard Holland, Albu¬
querque, New Mexico, provided specimens of E. anicia from northern Mexico.
This paper was critically reviewed by Drs. Robert E. Pfadt and C. C. Bur-
khardt, Dept, of Entomology, University of Wyoming, Dr. F. M. Brown,
Colorado Springs, Colorado, and Dr. Lawrence F. Gall, Peabody Museum of
Natural History, Yale University.

Literature Cited

BRUSSARD, F.F., P. R. EHRLICH, D. J. MURPHY, B. A. WILCOX & J. WRIGHT, 1985. Genetic
distances and taxonomy of checkerspot butterflies (Nymphalidae: Nympha-
linae). J. Kans. Ent. Soc. 58(3): 403-412.

HODGES, R.W., et al., 1983. Check List of the Lepidoptera of America North of
Mexico. London: E. W. Classey, Ltd., 284 pp.

SCOTT, J.A., 1978[80]. A survey of valvae of Euphydryas chalcedona, E. c. colon,
and E. c. anicia. J. Res. Lepid. 14(4): 245-252.

SPOMER, S. M. & J. M. REISER, 1985. Observations of the life history of Occidryas
anicia bernadetta (Nymphalidae) at the type locality. J. Lepid. Soc. 39(1):
55-57.

STANFORD, R. E., 1985. Rocky Mountain butterfly distribution maps. 4th ed.
Privately published.

Journal of Research on the Lepidoptera

26(1-4):116-124, 1988

The Mating System of Vanessa kershawi: Males Defend
Landmark Territories as Mate Encounter Sites

John Alcock and Darryl Gwynne

Department of Zoology, Arizona State University, Tempe, AZ, 85287 and Depart¬
ment of Zoology, University of Western Australia, Nedlands WA, 6009

Abstract. Males of Vanessa kershawi (McCoy) occupy perch terri¬
tories in the late afternoon in sunspots on hilltops and other locations
in southwestern Australia. Perching males respond to intruders with
chases and ascending flights, with some individuals defending the
same perch site for several consecutive afternoons. Females visit
territories strictly to mate and not to use any resources in these areas.

In the absence of hilltops males wait at sites where local topography
and vegetation create passageways that might channel dispersing
females to them. Landmark-based mating systems of this sort appear
to evolve in species whose females are highly dispersed because of the
distribution of the food and oviposition resources that they exploit.

Introduction

Although the painted lady butterflies — genus Vanessa — are
common and widespread, relatively little is known of their reproductive
behavior. Accounts of V. atalanta , V. cardui , V. caryae , and V. indica
indicate that males of these species go to certain conspicuous land¬
marks, particularly hilltops, but also forest-meadow edges, where they
perch in the late afternoon (Alcock, 1984; Bitzer & Shaw, 1979(80);
Niimura, in Suzuki, 1976; Scott, 1975, 1986; Shepard, 1966; Shields,
1967). Although males of some species are territorial, defense of the
perch site does not occur in several of these species when population
densities are high (Alcock, 1984). It has been assumed that perched
males on hilltops and other landmarks are awaiting the arrival of
receptive females, but no observations of copulations at these sites have
been recorded in the literature.

This report describes the reproductive behavior of V. kershawi (Mc¬
Coy), an abundant species throughout Australia (Common & Water-
house, 1972). Some authors consider V. kershawi to be a subspecies only
of V. cardui (Zimmerman, 1958), and there is no question that the two
are very similar. We show that this butterfly uses landmarks as mate-
encounter sites but that in different locations males establish territories
at very different kinds of landmarks. We discuss the significance of this
finding for an understanding of landmark-based mating systems, as
well as documenting that landmark territoriality is associated with
mate-acquisition in this species.

26(l-4):l-288, 1988

117

Methods

The butterflies were observed on 6— 7 October at Tutanning Reserve,
approximately 25 km east of Pingelly and 175 km southeast of Perth, W.
A., on 18-19, 23-24 October and 19-20 November at Watheroo
National Park, W. A., in the south-central portion of the park about 250
km north of Perth, and from 29 October to 13 January at King’s Park, an
area of natural bushland in Perth, W. A. The descriptive data were
largely collected by selecting a site that contained a male perched on the
ground in the late afternoon and then recording the behavior of the male
or males that resided in that location for periods ranging from 10 min to
2.5 hr. In some cases resident males were captured in an insect net and
marked with Liquid Paper typewriter correction fluid through the folds
of the net before being released. The sites at which males were observed
were checked again on subsequent days to determine if some perch sites
were used repeatedly.

Means are presented ± 1 S.D.

Results

Perch Site Selection

In all three study sites, males arrived at their perching sites on the
ground in sun spots or sunny strips in the mid- to late-afternoon.
Although on some days males arrived as early as 1500 hr, the density of
perching males was greatest between 1700—1800 hr in all three places.
All males were gone from their perching areas by dusk. But the
vegetation and topography of the areas in which males chose perch sites
varied considerably among the three locations. At Tutanning, males
selected sun spots on moderately forested hilltops, particularly on the
edge of rocky escarpments, but also on the flat “plateaus” of the hilltops,
sometimes dozens of meters from the steep hillsides. The sunspots were
scattered among a forest of Eucalyptus wandoo and Casaurina huegliana.

In King’s Park there are no well-defined hilltops but instead gently
rolling terrain with gradual ascents and descents. At this site males
were found throughout the area perched on concrete pathways (Fig. 1)
and cleared firebreaks, with no obvious concentrations on the higher
elevations. The paths and firebreaks cut through a forest of Banksia ,
Casaurina and Eucalyptus marginata in places and elsewhere through
more open stretches of tall shrubs, mainly blackboys ( Xanthorrhea
preissii).

In Watheroo National Park the terrain was almost completely flat
and the forest an open one composed of pricklybark ( Eucalyptus tod-
tiana ), scattered Banksias , and zamia palms ( Macrozamia reidlei).
Males perched in sunny avenues in the woodland, particularly in one
area where low pricklybark foliage formed a green barrier; the des¬
cending sun illuminated a long strip of ground parallel to the barrier.

118

J.Res.Lepid.

Territorial Behavior

Perching males flew up at objects moving near them including stones
and chunks of wood thrown over them, as well as flying dragonflies,
other butterfly species, and even birds. In addition perched males
sometimes spontaneously flew 2-10 m away from their perch before
returning to land usually at or near the previous perching area. These
spontaneous “patrol flights” occurred at the rate of 8.7/hr based on
360 min of observation of focal males in the three locations.

If at any time, a conspecific male entered the perching area, a chase
invariably resulted. Whereas chases of non-conspecifics lasted only a
few seconds, chases of fellow males regularly lasted about 30 sec (X =
28.9 ± 17.0 sec, N = 36, range 7—75). Chases began with horizontal
dashes, roughly 5— 15 m in length and oriented back and forth over the
territory. Chases lasting more than 10—15 sec usually terminated with
rapid ascending flights that took the participants 5 or more meters high,
far away from the territory and out of view of the observer in woodland
habitats.

The frequency of male-male interactions varied from 18.4/hr (based
on 5 hr of observation) at Tutanning to 5.7/hr (11.3 hr of observation) at
King’s Park, and 4.7/hr (4.25 hr of observation) at Watheroo. Generally
only one male returned from a chase to perch at the site which had been
occupied before the interaction. Thus males used chases to monopolize
perching sites and to disperse their rivals. At King’s Park and at
Watheroo perching males were separated by at least 25 m on most days.
At the highest density site (Tutanning Reserve) a maximum of 11
males occupied an area of 1452 m2 with a minimum distance of 6.3 m
between nearest neighbors (X = 9.1 m, N = 9). Note that it was at this
site that interactions among males were most frequent.

Some males were able to control a territory for substantial periods of
time in an afternoon. At King’s Park we secured 15 records of identified
males (either marked with Liquid Paper or with distinctive wing
damage) that held their site for a minimum of 30 min in an afternoon
(up to a maximum of 140 min). Six marked males showed considerable
site tenacity by returning to their perch site on the same day as capture,
despite the trauma of netting. Residents generally succeeded in repel¬
ling intruders, winning 46 of 53 interactions sampled on seven days and
involving five different residents at King’s Park. On four occasions both
resident and intruder returned to perch in the same general area and
only in three cases did the intruder replace the resident.

Some males were also able to reclaim the same location on several
days. In the 10 day period from 9—18 November, all resident males (10)
at one territory in King’s Park were identified by marks or wing
damage, and the area checked daily. Table 1 shows that during this time
three territory owners returned to the site on more than one day. In
addition, a different male defended the same location for the four days

26(l-4):l-288, 1988

119

Fig. 1. A male Vanessa kersha wi per¬
ched in a sunspot on a con¬
crete sidewalk slab in King's
Park, Perth.

Fig. 2. A frequently occupied per¬
ching area in King's Park,
Perth. Males defended con¬
crete slabs in the left hand
lane of the track in the mid¬
dle of the figure. Note that
the surrounding vegetation
creates a tunnel of sorts over
the perching area.

Fig. 3. A copulating pair of Vanessa
kershawi that mated after the
female flew to the male's landmark
territory on a hilltop in Tutanning
Reserve, W.A.

120

J.Res.Lepid.

between 26—29 November. Thus at least under some conditions males
exhibit site tenacity.

But the competition for perch ownership also led to turnovers. On
seven occasions when observing an identified male at King’s Park or
Tutanning, a new male replaced the past owner after a chase or series
of chases. Thus some perches attracted more than one male owner in a
single day (see Table 1), and the same spot could be occupied over a
series of days by many different males. At Watheroo one site that was
held on 19, 23-24 October was also claimed a month later on 19
November. A focal territory at King’s Park was held by six different
residents during 9—18 November and at least four other males visited
the site. The recorded number of visitors is surely a gross underestimate
of the actual total because most intruders were promptly chased away
before they could be captured, marked or identified. The focal territory
at King’s Park was occupied by a resident male on 20 of 22 afternoons
when the site was checked from 29 October to 13 January. Three other
sites on a transect of King’s park paths 420 m long were also occupied on
a majority of these days (Site B = 15/22, Site C = 13/22, and Site
D = 14/22).

Male -Female Interactions

Encounters between males and females were rare. At Tutanning
Reserve two matings were recorded, both at times of peak male density:

Table 1. Resident and visitor males of V. kershawi at one perching site

in King’s Park, Perth from 9—18 November 1985. R =
resident male — present for most of observation period; v =
visitor male — present only for brief period(s) before being
chased away by the resident male; R* = new male takes site
from previous resident site during an afternoon.

November

Male

Date

9 10 11 12 13 14 15 16 171 18

A

B

C

D

E

F

G

H

I

J

R R R

v

v

v

R

v

v R R

R

R*

R*

R

R*

R

xno observations made on this day

26(l-4):l-288, 1988

121

1729 hr on 6 October and at 1733 hr on 7 October. These took place at
two different territories on the edge of a hilltop when a female flew into a
territory, and was pursued by the resident away from the perch site. The
pair flew much more slowly and more erratically than male-male
interactors, and with no ascending component to the chase. In both cases
the two butterflies landed on the foliage of a sapling wandoo eucalyptus,
approximately 24 and 5 m from the point of first encounter, respectively.
The male landed by the female, facing in the same direction. Copulation
followed quickly with no preceding wing-fluttering by the female in one
instance, although in the other the female flew off the perch several
times before alighting for good. Once the female had ceased moving, the
male simply probed with his abdomen twisting it to the side to couple
with the female, after which he turned to face directly away from his
partner (Fig. 2). In both cases copulation was still in progress lhr
after its initiation.

One mating was recorded at King’s Park at 1707 on 16 November. It
followed the same pattern, although in this case the female led the male
on a long pursuit as she flew away from and then returned to pass over
the perch territory several times before finally landing on a casaurina
cone some 3 m above the ground and about 10 m from the perch site.
Copulation was still in progress 30 min after it began.

Two other probable male-female interactions were seen at King’s
Park. One occurred at 1648 on 11 November when a male visitor with a
distinctive wing mark that had been perching near the territory holder
suddenly flew up after another visitor and departed. The pair flew off in
the distinctive, relatively slow horizontal flight that characterized
male-female pursuits but the butterflies were lost to view in the
woodland. The male did not return within a hour. The second inter¬
action took place at 1715 on 18 November when an individual, probably
a female, flew into a male’s territory and led him on a brief chase before
landing on the ground. The male perched immediately behind the visitor
and flew up when it left. The pair went into the woodland in a slow
horizontal pursuit flight and were lost in the scrub. The male did not
return within a minute as was normally the case in male-male chases.

Discussion

The term “territoriality” covers a variety of behavioral phenomena,
but in its most widely accepted sense it refers simply to the defense of
space by an individual (Brown, 1975). In this sense males of V. kershawi
are territorial, with individuals defending perches that may be visited
in the late afternoon by receptive females. The distinctive features of
male-male chases, which are very different from male-female encoun¬
ters and pursuits of heterospecifics, leave little doubt that male-male
interactions determine ownership of a perch site. Similar behavior has
been labelled “territorial” in two reviews of insect territoriality (Baker,

122 J.Res.Lepid.

1983; Fitzpatrick & Wellington, 1983; but see Scott, 1986, for argu¬
ments on the absence of territoriality in butterflies).

Some perch sites are consistently occupied by many different males
over a period of several months; other similar locations never attract a
territory owner. What properties make a perching area worth defend¬
ing? There are no flowering plants or oviposition resources in the
territories of V. kershawi ; females visit perched males solely to acquire a
mate and males defend their perching areas solely to maintain a site
from which to scan for incoming females. We suggest that the mating
system of V. kershawi is based on landmarks with local topography
dictating where females are most likely to be travelling, and this in turn
determines where males compete for waiting sites.

As is true for a host of butterflies and other insects, prominent hilltops
may serve as orientation guides or attraction points for females (Scott,
1970; Shields 1967; Thornhill & Alcock, 1983), and when hilltops are
available (as at Tutanning Reserve) males of V. kershawi wait at the
highest points. But in some regions, hilltops are absent and then males
use alternative topographic features as productive waiting sites. In
King’s Park, many territories are on walking tracks, sidewalks and
firebreaks at points where the trailside vegetation creates a passageway
likely to channel or funnel passing females toward a perched male.
Similarly at Watheroo males appear to wait in open sunny areas in the
woodland through which traveling females might be guided by the foli¬
age of plants beside the clearings. Thus, whatever their environment,
males of V. kershawi seem to take advantage of natural orientation
marks, clearings through vegetation, and foliage barriers to station
themselves at points most likely to be visited by dispersing females.
Males of other Lepidoptera that use landmarks as mate-encounter sites
may also be using topographic channels and funnels that concentrate
travelling females (Callaghan, 1982(83)).

Flexibility in the use of landmarks also occurs in other butterflies
(Scott, 1982(83)), including V. atalanta (Alcock, 1984; Bitzer & Shaw,
1979(80)), whose males station themselves on peaktops in hilly or
mountainous terrain, but wait in clearings and forest-meadow edges in
flat, forested areas.

The use of landmark and topographic guides appears widespread in
the genus Vanessa (Alcock, 1984; Bitzer & Shaw, 1979(80); Shields,
1967). The behavior of V. kershawi and V. atalanta , for example, is close
to identical in terms of sites selected by waiting males, the nature of
male-male interactions, the duration and frequency of aerial chases, the
consistency with which some territories are defended from day to day,
and the restriction of territoriality and mating to the late afternoon
(Bitzer & Shaw, 1979(80); Dimock, 1984(85)). Why should males of
these butterflies be so prone to wait at resource-less areas rather than
searching actively for females at foraging or oviposition sites? The
general rule among insects is that when either food or egg-laying

26(l-4):l-288, 1988

123

resources are concentrated, thereby concentrating females spatially,
males focus their search at these locations (Thornhill & Alcock, 1983).
In many butterflies, including Vanessa species whose larvae and adults
feed on a wide range of hosts (Common & Waterhouse, 1972), females
are often not clumped and therefore are not easy to locate (Rutowski,
1984). Under these circumstances males may be forced to wait in
portions of their environment where travelling females may occasional¬
ly appear (Rutowski, 1984). Vanessa butterflies are well-known for
their tendency to travel long distances (Johnson, 1969; Smithers, 1969).

One would predict, however, that if females of a Vanessa species
happened to become aggregated at a restricted food- or hostplant, their
males would respond by searching for mates at these productive
locations. When females of V. cardui occur in large numbers on
flowering Encelia farinosa in central Arizona, some males do search for
mates at the foodplant (Alcock, 1984). A combination of hilltopping and
searching at flowers has been reported for a few other butterflies as well
as by Scott (1982(83), 1986). Likewise, 10—20 males and females of V.
kershawi were found at a local patch of a flowering Verticordia on 19— 20
November at Watheroo National Park. The plant had not been in
bloom a month earlier when late afternoon perch defenders were
common in other areas, but by 19—20 November only a single perch
defender was located. Instead, throughout these days males frequently
engaged in what appeared to be brief (< 5 sec) horizontal sexual chases
at the foraging site. Apparently when females of Vanessa are spatially
clustered at flowers, males travel to these locations to search for
receptive partners. But if no such clusters exist, males wait at travel
points for diffusely distributed females to come to them.

Acknowledgements

This study was done while J. A. was on sabbatical leave from Arizona State
University while visiting the Department of Zoology, University of Western
Australia. UWA kindly provided a fellowship and other forms of support. We
received fine assistance in watching the butterfly from Sue and Nick Alcock and
from Sarah Brooks. Ron Rutwoski and two anonymous reviewers helpfully
criticized the manuscript.

Literature Cited

ALCOCK, J., 1984. Convergent evolution in perching and patrolling site
preferences of some hilltopping insects of the Sonoran desert. South¬
western Nat. 29: 475-480.

BAKER, R. R. 1983. Insect territoriality. Ann. Rev. Ent. 28: 65-90.

BITZER, R. J. & K. C. SHAW, 1979 (80). Territorial behavior of the red admiral,
Vanessa atalanta (L.) (Lepidoptera: Nymphalidae). J. Res. Lepid. 18:36—49.
brown, J. L., 1975. The evolution of behavior. W. W. Norton, New York. 761 pp.

124

J.Res.Lepid.

CALLAGHAN, C. J., 1982(83). A study of isolating mechanisms among neotropical
butterflies of the subfamily Riodininae. J. Res. Lepid. 21: 159—176.

COMMON, I. F. B. & D. F. WATERHOUSE, 1972. Butterflies of Australia. Angus &
Robertson, Sydney. 498 pp.

DIMOCK, T. E., 1984. Culture maintenance of Vanessa atalanta rubrica (Nym-
phalidae). J. Res. Lepid. 23: 236-240.

FITZPATRICK, G. W. & W. G. WELLINGTON, 1983. Insect territoriality. Can. J. Zool. 61:
471-486.

JOHNSON, C. G., 1969. Migration and dispersal of insects by flight. Methuen & Co.,
London. 763 pp.

RUTOWSKI, R. L., 1984. Sexual selection and the evolution of butterfly mating
behavior. J. Res. Lepid. 23: 125—142.

SCOTT, J. A., 1970. Hilltopping as a mating mechanism to aid the survival of low
density species. J. Res. Lepid 7: 191—204.

- , 1975. Mate-locating behavior of North American butterflies. J. Res.

Lepid. 14: 1 — 10.

- , 1982(83). Mate-locating behavior of western North American butter¬
flies. II. New observations and morphological adaptations. J. Res. Lepid. 21:
177-187.

- , 1986. The Butterflies of North America. Stanford University Press,

Stanford, CA. 583 pp.

SHEPARD, J., 1966. A study of the hilltopping behavior of Pieris occidentalis
Reakirt. Pan-Pac. Ent. 42: 287-294.

SHIELDS, O., 1967. Hilltopping. J. Res. Lepid. 6: 69-178.

SMITHERS, C. N., 1969. A note on migrations of Vanessa kershawi (McCoy)
(Lepidoptera, Nymphalidae) in Australia. Aust. Zool. 15: 185—187.

SUZUKI, Y., 1976. So-called territorial behavior of the small copper, Lycaena
phlaeas daimio Seitz (Lepidoptera, Lycaenidae). Kontyu 44: 193-204.

THORNHILL, R. & J. ALCOCK, 1983. The evolution of insect mating systems.
Harvard University Press, Cambridge, Mass. 547 pp.

ZIMMERMAN, E. C. 1958. Insects of Hawaii. Vol. 7. University of Hawaii Press,
Honolulu, HW. 542 pp.

Journal of Research on the Lepidoptera

26(1-4):125-140, 1988

Apodemia palmerii (Lycaenidae: Riodininae):
Misapplication of Names, Two New Subspecies and a
New Allied Species

George T. Austin

Nevade State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas,
Nevada 89107

Abstract. The subspecific names of Apodemia palmerii, A. p. palmerii
and A. p. marginalis, have been variously used for the phenotypes of
the species. Examination of the types and series of topo types indicates
that they are synonymous. New subspecific names are proposed for the
darker phenotype distributed east of the Colorado River drainage and
another in central Mexico. Southern Baja California, Mexico popula¬
tions, often referred to as Apodemia “palmerii” are described here as a
new species.

Introduction

William Henry Edwards (1870) described Lemonias palmerii from
Utah. Subsequently, Skinner (1920) named the taxon Lemonias
palmerii marginalis from a California population of the species. The
name “ marginalis” has been variously treated as a form synonymous
with Edwards’ concept (dos Passos 1964, Howe 1975, Miller and Brown
1981, Austin 1985b) or as a recognizable subspecies distinct from
nominate A. palmerii (Comstock 1927, Holland 1931, Emmel and
Emmel 1973, Tilden 1975, Austin and Austin 1980). Populations from
southern Arizona and western New Mexico generally have been
referred to as nominate A. palmerii (Comstock 1927, Holland 1974,
Howe 1975, Ferris 1976). Study of material from throughout the range
of the species and examination of the type specimens of the two
presumptive subspecies indicate errors in the application of A. p.
marginalis and that new names are needed for the A. palmerii popula¬
tions east of the Colorado River drainage and those in Mexico. Yet
another phenotype, previously referred to as A. palmerii (Rindge 1948,
Holland 1972), occurs in southern Baja California, Mexico, but it is, in
fact, an undescribed species.

Throughout this paper, butterfly size (given as mean and range) is the
length of the right primary from the base to the apex in millimeters.
Measurements are for 15 specimens unless otherwise indicated. Speci¬
mens indicated by “M” and “F” are male and female, respectively.

126

J.Res.Lepid.

Names and Populations

The description of the male oiApodemia palmerii was based on Utah
material taken by Edward Palmer (Edwards 1870). Later, Edwards
(1884b) stated that his description was based on a single male. Brown
(1967, 1968) presented ample evidence that the specimen was most
likely collected in Utah, probably in the vicinity of St. George, Washing¬
ton County during June 1870. The holotype male (Brown 1968) at the
Carnegie Museum of Natural History is a typical spring brood specimen
with entirely orange margins. Skinner’s (1920) description of A. p.
marginalis was based on two males and a female from Acme (once called
Morrison, ca. 4 miles south of Tecopa), Inyo County, California. He
distinguished it from supposed nominate A. palmerii by its orange wing
margins (hence the name “ marginalis”) and by its pallidity. No mention
was made of either the geographical or seasonal source(s) of the
material to which he compared his types. A male [holojtype and a female
[allojtype are among the type material transferred to the Carnegie
Museum of Natural History from the Academy of Natural Sciences of
Philadelphia (see also Gillham and Ehrlich 1954). These were collected
by Morgan Hebard on 8 August 1919 and represent the typical pale
late summer phenotype.

I first became aware of a possible nomenclatural problem during my
studies of southern Nevada butterflies (Austin and Austin 1980). My
series from this area was of a seasonally variable, dark to pallid, orange-
margined butterfly which was obviously distinct from the darker
southern Arizona insect. It seemed unlikely on biogeographic and
ecological bases that nearly all Colorado River basin populations were
of one phenotype while southern Utah examples should be a disjunct
population of the same sort as found in southern Arizona. I, therefore,
collected representative series from eastern California near the type
locality of A. p. marginalis , from southern Utah in the St. George region
(the designated type locality of A. palmerii ) and from southern Arizona
(Pima, Cochise and Pinal counties). I also took a small number from
western Arizona (Maricopa and Mohave counties). These along with
material borrowed from various museums and collectors indicate that
two phenotypes are indeed involved in this region but that the names
available have been inappropriately applied.

The distribution of the species Apodemia palmerii includes extreme
northeastern Baja California, Mexico; the southeastern, desert portions
of California; southern Nevada; extreme southwestern Utah; south and
eastward through southern Arizona and southwestern New Mexico to
western Texas (Fig. 1). It occurs in northwestern and northcentral
Mexico (where its distribution is incompletely known, Hoffmann 1976),
but it occurs south into the central portions of the country (to Hidalgo
and Michoacan) and in the western states of Sonora and Sinaloa. The
southern Baja California populations also traditionally have been
treated as this species.

26(l-4):l-288, 1988

127

Fig. 1. Distribution of Apodemia palmerii sspp. and Apodemia murphyi (open
symbols indicate specimens not examined).

A pale phenotype with prominant orange margins from the Colorado
River drainage and nearby areas of California, Nevada, Utah and
western Arizona generally has been referred to as Apodemia palmerii
marginalis. The darker populations from east of the Colorado River
drainage, on the other hand, have been considered nominate A . palmerii.
No other names have been applied to other variations within the
species. While Edwards’ description must serve as that for the species, it
is not detailed enough to distinguish that phenotype from others in
question. Skinner’s description unquestionably refers to the pale pheno¬
type. The application of the name “marginalis” was, in my opinion,

128

J.Res.Lepid.

probably based on comparisons with Arizona material since Arizona
specimens are more widely represented in collections than those from
southern Utah. My experience is that A. palmerii is uncommon in the
St. George region, even where good stands of its foodplants, honey
mesquite ( Prosopis glandulosa Torr.) and screwbean mesquite (P.
pubescens Benth.) are available. The species, however, is abundant in
similar situations in southern Arizona.

Apodemia palmerii palmerii (W. H. Edwards, 1870) (Fig. 2)
Lemonias palmerii W. H. Edwards (1870, p. 195), Kirby (1871, p. 650), W. H.
Edwards (1872, p. 38), W. H. Edwards (1874, p. 38), W. H. Edwards (1877,
p. 38), Kirby (1877, p. 760), Strecker (1878, p. 104), Brooklyn Entomological
Society (1881, p. 3), W. H. Edwards (1884a, p. 294), W. H. Edwards (1884b,
p. 301), Maynard (1891, p. 126), Skinner (1898, p. 43), Smith (1903, p. 6), Snow
(1907, p. 156)

Chrysobia palmerii Scudder (1876, p. 103)

Lemonias palmeri Holland (1898, p. 231), Skinner (1904, p. 16), Wright (1905,
p. 202), Haskin (1914, p. 306)

Poly stigma palmerii Dyar (1902, p. 34)

Apodemia palmerii Mengel (1905, p. 120), Stichel (1911, p. 288), Seitz (1924,
p. 700), Barnes and Benjamin ( 1926, p. 16), Stichel (1930, p. 590), McDunnough
(1938, p. 23), dos Passos (1964, p. 51), Brown (1967, p. 129), Scott (1979,
p. 191), Fisher in Ferris and Brown (1981, p. 198-199), Miller and Brown
(1981, p. 132), Pyle (1981, p. 530), Miller and Brown in Hodges (1983, p. 57)
Apodemia palmerii palmerii Stichel (1911, p. 288), Brown (1968, p. 121),
Callaghan and Tidwell (1971, p. 198), Austin (1985a, p. 128), Austin (1985b,
p. 107).

Apodemia palmeri Barnes and McDunnough (1917, p. 13), Holland (1931,
p. 213), Ehrlich and Ehrlich (1961, p. 245), Tietz (1972, p. 504), Powell in
Howe (1975, p. 270), Gillette (1983, p. 15).

Lemonias palmerii marginalis Skinner (1920, p. 175)

Apodemia palmerii from “marginalis” Barnes and Benjamin (1926, p. 16),
McDunnough (1938, p. 23), Martin and Truxal (1955, p. 20), dos Passos (1964,
p. 51), Hoffmann (1976, p. 68), Fisher in Ferris and Brown (1981, p. 198-199),
Miller and Brown (1981, p. 132), Miller and Brown in Hodges (1983, p. 57)
Apodemia palmerii marginalis Comstock ( 1927, p. 151), Comstock and Dammers
(1932, p. 37), Davenport and Dethier (1937, p. 170), Emmel and Emmel (1973,
p. 49), Tilden (1975, p. 30)

Apodemia marginalis Holland (1931, p. 213)

Apodemia palmeri marginalis Gillham and Ehrlich (1954, p. 102), Emmel
(1972, p. 3), Austin and Austin (1980, p. 23)

Apodemia palmeri form “marginalis” Tietz (1972, p. 504), Powell in Howe (1975,
p. 270)

Apodemia palmeri palmeri Austin and Austin (1980, p. 23)

Specimens of Apodemia palmerii from southwestern Utah, southern Nevada,
southeastern California, extreme western Arizona and extreme northern Baja
California Norte, Mexico (essentially the Colorado River and Death Valley

26(l-4):l-288, 1988

129

2a 2b

Fig. 2a. Apodemia palmerii subspecies (dorsal surface). Top left — A. p. arizona
holotype male, A Z: Cochise Co.; Az. 90, 14.6 mi. N Az. 82, 4 Sept. 1980,
leg. G. T. Austin. Top right — A. p. arizona allotype female, same data.
Second left — A. p. palmerii topotype male of form “marginalis”, CA: Inyo
Co.; 0.3 mi. N Tecopa, 8 Sept. 1 981 , leg. G. T. Austin. Second right — A. p.
palmerii topotype female of form "marginalis”, same data. Third left — A.
p. palmerii male of form "marginalis”, NV: Clark Co.; Whitney Mesa at
Sunset Rd., 6 Sept. 1977, leg. G. T. Austin, Third right — A. p. palmerii
female of form "marginalis”, NV: Clark Co.,; Las Vegas, Paradise Valley,
6 Sept. 1977, leg. G.T. Austin. Fourth left — A. p. palmerii male, NV: Clark
Co.; Las Vegas, Paradise Valley, 24 May 1978, leg. G. T. Austin. Fourth
right — A. p. palmerii female, same data. Bottom left — A. p. australis
holotype male, MEXICO, Durango 1 mi. S. Nombre de Dios, 1760 m, 30
Aug. 1973, leg. L. D. and J. Y. Miller, Bottom right — A. p. australis
allotype female, same data.

Fig. 2b. Apodemia palmerii subspecies (ventral surface). Same specimens as in
Fig. 2a.

drainage basins, Fig. 1) are virtually identical in their general pallidity, in the
presence of continuous orange margins on the dorsal surface of both wings, and
in seasonal variability (Fig. 2). This geographical extent encompasses the type
localities of both A. p. palmerii and A. p. marginalis. The latter name thus is
clearly synonymous with the nominate and refers to the pale late summer/
autumn form. Examination of the types confirmed this. The butterfly is often
locally abundant and flies in three to four broods in southern Nevada (Austin
and Austin 1980) and two to three broods in southern California (Emmel and
Emmel 1973).

Nominate Apodemia palmerii is seasonally variable. Early season (April-
June) specimens are generally large (male x = 11.2, range = 10.6- 11.8,; female
x = 12.5, range = 11.7-13.1, May sample) and relatively dark with a restriction
of the fulvous on the wing bases, especially on males. Later (July-October)
specimens are smaller (male x = 10.1, range 9.0—10.9; female x = 11.2, range

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J.Res.Lepid.

10.2-12.2, September sample) and paler with considerable fulvous on the
wings. The type specimen of A. palmerii was illustrated by Brown (1968).
Holland (1931) figured a paratype of form “marginalis” but his indicated “type”
of A. palmerii is a pseudotype (Brown 1968, see below). Emmel and Emmel
(1973) illustrated a male of the spring phenotype. Holland (1931), Emmel and
Emmel (1973) and Howe (1975) illustrated form “marginalis”. Comstock and
Dammers (1932) described and illustrated the life history of a California
population. The larval host plants ar eProsopis glandulosa ( = P.juliflora) andP.
pubescens (Fabaceae) (Comstock and Dammers 1932, Austin and Austin 1980).

This then leaves populations exhibiting somewhat darker phenotypes from
outside this area without names. To rectify this situation, I describe here two
new subspecies.

Apodemia palmerii arizona new subspecies Austin (Fig. 2)
Lemonias palmerii Kirby (1871, p. 650), Kirby (1877, p. 760), W. H. Edwards
(1877, p. 38), W. H. Edwards (1883, p. 9), W. H. Edwards (1884a, p. 294), W. H.
Edwards (1884b, p. 301), Maynard (1891, p. 126), Skinner (1898, p. 43), Smith
(1903, p. 6), Snow (1904, p. 337), Snow (1907, p. 156)

Chrysobia palmerii Scudder (1876, p. 103)

Apodemia palmeri Edwards (1882, p. 28), Godman and Salvin (1886, p. 468),
Barnes and McDunnough (1917, p. 13), Holland (1931, p. 213), Rindge (1948,
p. 300), Ehrlich and Ehrlich (1961, p. 245), Powell in Howe (1975, p. 270),
Austin (1978, p. 210)

Lemonias palmeri Godman and Salvin (1886, p. 468), Holland 1898, p. 231),
Skinner (1904, p. 16), Haskin (1914, p. 306), Stone (1921, p. 114).

Poly stigma palmerii Dyar (1902, p. 34)

Apodemia palmerii Mengel (1905, p. 120), Stichel (1911, p. 288), Seitz (1924,
p. 700), Barnes and Benjamin (1926, p. 16), Stichel (1930, p. 590), McDunnough
(1938, p. 23), Bauer (1954, p. 100), Martin and Truxal (1955, p. 20), dos Passos
(1964, p. 51), Brown (1965, p. 112), Lewis (1973, p. 112), Tilden (1974, p. 24),
Hoffmann (1976, p. 68), Fisher in Ferris and Brown (1981, p. 198— 199), Miller
and Brown (1981, p. 132), Pyle (1981, p. 530), Miller and Brown in Hodges
(1983, p. 57), Austin (1985b, p. 107)

Apodemia palmerii palmerii Comstock (1927, p. 151), Brown (1968, p. 123),
Holland (1974, p. 44), Ferris (1976, p. 46)

Apodemia palmeri palmeri Austin and Austin (1980, p. 23)

MALE. Dorsal ground color dark brown, sometimes nearly blackish-brown.
Basal one-third of both wings usually fulvous, often with considerable black
overscaling. White markings and their associated black outlines much as on
Apodemia palmerii palmerii. Marginal area usually of ground color, especially
apically, with small areas of fulvous in each cell (not broadly fulvous), these
fulvous areas usually broader posteriorly on both wings and usually somewhat
over scaled with ground color. Ventral ground color largely fulvous with
markings of dorsum repeated but larger. Distinct submarginal black points on
both wings.

Male genitalia virtually identical to those of Apodemia palmerii palmerii.
FEMALE. Somewhat larger in size than male with a more rounded (less
pointed) apex to primaries. Color and pattern similar to male.

TYPES (data as on labels, clarified in brackets). Holotype male — A[RI1Z
[ONA]: Cochise Co. [unty]; A[ri]z[ona State Route] 90, 10.8 mi. [les] N. [orth of]

26(1-4): 1-288, 1988

131

A[ri]z[ona State Route] 82, 7 September] 1980, leg. G. T. Austin. Allotype
female — same data as holotype. Paratypes (32M, 26F) — same data as holotype
(26M, 21F); some data as holotype except 14.6 mi. N (6M, 4F); same data as
holotype except 8.4 mi. N (IF).

DEPOSmON OF TYPE MATERIAL. The holotype, allotype, 11M and 6F
paratypes will be deposited in the type collection of the Nevada State Museum.
A pair of paratypes will be deposited in each of the following institutions: Allyn
Museum of Entomology, American Museum of Natural History, National
Museum of Natural History, Carnegie Museum of Natural History, Natural
History Museum, San Diego, and Los Angeles County Museum. The remainder
are to be retained by the author.

TYPE LOCALITY. ARIZONA: Cochise County, Arizona State Route 90, 10.8
miles north of Arizona State Route 82. The types were collected on the west side
and within 100 feet of the road. Most were perched on mesquites ( Prosopis
glandulosa ) (Fabaceae) which undoubtedly serves as the larval host plant.

DISTRIBUTION AND PHENOLOGY. Apodemia palmerii arizona occurs from
Arizona south and eastward through southwestern New Mexico to western
Texas, south into at least Chihuahua, Sonora and Sinaloa, Mexico (Fig. 1). The
subspecies has at least two (but probably more) broods in southern Arizona
(Austin 1978) and two broods In southwestern New Mexico (Ferris 1976). In
southern Arizona, Brown (1965) reported it as a late rainy season species
although Austin (1978) found it to occasionally have a large spring brood, at
least in years with spring rainfall.

ETYMOLOGY. This subspecies is named for its type locality, the state of
Arizona.

DIAGNOSIS AND DISCUSSION. The new taxon, Apodemia palmerii arizona,
is at once distinguished from nominate A . palmerii by the largely dark margins
of the dorsum of both the primaries and secondaries. This same area of A.
palmerii is broadly fulvous with considerably less or no ground coloration. A
very few A . p. arizona approach A.p. palmerii in this respect just as occasional A .
p. palmerii have the margins somewhat darkened. The dorsal ground color of A.
p. arizona is darker and with less fulvous flush basally than A. p. palmerii giving
the impression of an overall darker butterfly. Some late season females of A.p.
palmerii are very pale with the ground color approaching a pale tan (a condition
I have not seen among A. p. arizona). The ventral color is paler than the average
early season A. p. palmerii but somewhat darker than late season material. The
submarginal black points are larger and more distinctly indicated than on
nominate A. palmerii. An aberrant female from Patagonia, Arizona (5 Sept.
1951) has the postmedian and basal white markings absent on all wings; the
submarginal markings are normal.

There is no appreciable seasonal variation in Apodemia palmerii arizona and
late season specimens have the size (male x = 11.5, range = 10.6-12.0; female x
= 12.4, range = 11.6-13.2, September sample) and a comparable dark ground
color of early season A. p. palmerii. The figures in Howe (1975) adequately
illustrate A . p. arizona. Edwards (1884b) and Holland (1931, as the type of A.p.
palmerii) also illustrate this taxon. Edwards (1884b) described and illustrated
the egg and young larva from southern Arizona.

Apodemia palmerii arizona, at least, applies to southern Arizona, southern
New Mexico and adjacent northwestern Mexico populations (Fig. 1). The few
specimens I have seen from the extreme eastern portion of its distribution in

132

J. Res. Lepid.

Texas (but not near El Paso) and east central Chinhuahua (but not north¬
western) consistently have broader white markings and are somewhat larger
(female x = 13.2, range = 12.4— 14.3, N = 7) but otherwise closely fit the concept.

Material from central Mexico is yet darker dorsally and browner ventrally. It
is recognized as follows:

Apodemia palmerii australis new subspecies Austin (Fig. 2)
Apodemia palmeri Godman and Salvin (1887, p. 709), Holland (1931, p. 213),

Ehrlich and Ehrlich (1961, p. 245), Powell in Howe (1975, p. 270)

Lemonias palmeri Holland (1898, p. 231)

Lemonias palmerii Skinner (1898, p. 43)

Apodemia palmerii Seitz (1924, p. 700), Fisher in Ferris and Brown (1981,

p. 198-199), Pyle (1981, p. 530)

MALE. Dorsal ground color blackish with slight fulvous basal overscaling
on secondaries and occasionally on primaries, white markings as on other
Apodemia palmerii subspecies but often smaller in size. Outer margins with
fulvous indistinct, usually restricted to small area at anal angle of primaries and
posterior half of secondaries. Ventrum dull brownish-orange ground color, apex
of primaries and entire secondaries dark tan, markings of dorsum repeated,
black submarginal points minute.

Genitalia of typical Apodemia palmerii type with broadly rounded uncus,
hooked upper process of valve, distinctly rounded vinculum and relatively
short saccus.

FEMALE. Wings more rounded than male, color and pattern similar on both
surfaces, ventral black submarginal points larger.

TYPES, (data as on labels, clarified in brackets). Holotype male — MEXICO:
Durango; 1 mi[le] Sfouth] Nombre de Dios, 1760 m[eters elevation], desert
scrub, 30 viii [August] 1973, leg. L. D. & J. Y. Miller, Sta. No. 1973-53.
Allotype female — same data as holotype. Paratypes (all MEXICO: Durango;
43M, 20F) — same data as holotype (37M, 7F, AME); 1.5 mi. SW Durango,
1920 m, ground scrub, 28 Aug. 1973, (5M, 5F, AME); Durango, 6200', 13 Aug.
1947 (1M, 5F, AMNH); Durango, 1 Aug. 1964 (IF, CIS); Yerbanis, Cuencame
Dist., 19 Aug. 1947 (IF, AMNH); Nombre de Dios, 5900', 13 Aug. 1947 (IF,
AMNH).

DEPOSITION OF TYPE MATERIAL. The holotype, allotype, 41M and 11F
paratypes are deposited at the Allyn Museum of Entomology; 1M and 7F
paratypes are at the American Museum of Natural History; IF paratype is at
the California Insect Survey; and one pair of paratypes are retained by the
author.

TYPE LOCALITY. MEXICO: Durango; 1 mile south of Nombre de Dios, 1760
meters. The vegetation here is desert scrub with trees to 25 feet in height, some
mesquites ( Prosopis ), various other low trees with a grass undergrowth on a
valley floor ( fide L. D. Miller).

DISTRIBUTION AND PHENOLOGY. This subspecies occurs mostly in the
central mountains of Mexico from Durango to Hidalgo and Michoacan usually
above 4500' in elevation (Fig. 1). There are at least two broods with records in
April (once), May (once) and from mid July through mid September, most
records are for August.

ETYMOLOGY. This phenotype is the most southerly distributed of the
species, thus the name “australis” (= southern).

26(1-4): 1-288, 1988

133

DIAGNOSIS AND DISCUSSION. This is a very dark Apodemia palmerii
subspecies, appearing nearly black when fresh. Individuals are considerably
darker above than those from other known populations of the species and lack a
conspicuous basal flush of fulvous. The white spots average small and sometimes
appear smudged. Ventrally this butterfly is brown rather than distinctly fulvous
as are both of the more northern subspecies. The basic pattern, wing shape and the
structure of the male genitalia leave no doubt that this subspecies falls within the
range of variation expected of A. palmerii. It approximates the size of other A.
palmerii (male x = 11. 5, range = 10.6— 12.5; female x = 12.6, range = 11.3-13.2).

An even more distinctive series of populations occurs in the southern portions
of Baja California, Mexico. This heretofore has been called Apodemia palmerii
but closer examination reveals that this is a very different insect with a unique
combination of characters. This new species is here described as:

Apodemia murphyi new species Austin (Fig. 3)

Apodemia palmeri Rindge (1948, p. 300), Powell in Howe (1975, p. 270)
Apodemia palmerii Holland (1972, p. 156), Fisher in Ferris and Brown (1981,

p. 198-199), Pyle (1981, p. 530)

MALE. Dorsum with blackish-brown ground color. Late summer and fall (July-
mid November) specimens usually with well-defined fulvous basal area; winter
and spring (late November -April) specimens heavily overscaled with ground
color on this area, often black without fulvous. White markings as on Apodemia
palmerii palmerii and A . p. arizona but considerably reduced in size, especially on
primaries. Marginal area mostly of ground color except for, usually, posterior one
or two cells on each wing, these with small areas of fulvous.

Ventral ground color brownish-orange, markings of primaries repeat dorsal
pattern but tending slightly larger. Ventral secondaries with white markings
considerably larger than dorsally, those in postmedian area form a continuous
broad band. Wings narrow, apex of primaries very long and pointed, often
approaching subfalcate.

Genitalia similar to Apodemia palmerii but with several subtle differences as
discussed below.

FEMALE. Similar to male except dorsal white markings larger and more
fulvous marginally, especially posteriorly on secondaries. Shape of primaries
more rounded than on males but with distinctive tendency towards a subfalcate
tip. Seasonal variation similar to male.

TYPES. Holotype male — MEXICO: Baja California Sur; Arroyo San Bartolo,
28 Aug. 1982, leg. fJ. W.l Brown and |D. K.l Faulkner (SDNHM). Allotype
female — same data as holotype (SDNHM). Paratypes (all MEXICO: Baja
California Sur; 103M, 54F) — same data as holotype (2M, IF, SDNHM); A. San
Bartolo, 3 Nov. 1961 (2M, CM), 12 Nov. 1961 (2M, IF, CM); San Bartolo, 3 Oct.
1981 (2M, IF, SDNHM); 2 mi. S of Buena Vista, 30 Nov. 1979 (3M, IF, SDNHM);
Buena Vista at Monument Rd., 4 Jan. 1980 (IF, JB); 3 mi. S Rio Buenavista,
25 Oct. 1961 (2M, CM); 5 km S Rio Buenavista, 25 Oct., 1961 (2M, CM); 4.2 mi.
W. Miraflores, 30 Sept. 1981 (1M, SDNHM); Miraflores, 25 Oct. 1961 (1M, CM),
beach, Todos Santos, 26 July 1981 (6M, SDNHM); estuary at Todos Santos, 26
July 1981 (9M, 2F, CM), 31 July 1981 (2M, CM), 20 March 1974 (4M, IF, CM);
El Pescadero, 20 March 1974 (IF, CM); 1/4 mi W Todos Santos, 20 March 1974
(2M, SDNHM), 19-20 March 1974 (3M, GF); 14 mi N Todos Santos, 4 Oct. 1981
(1M, IF, SDNHM); Santiago, 6 Nov. 1946 (1M, 3F, SDNHM); 19 mi. SE El Cien,

134

J.Res.Lepid.

3a 3b

Fig. 3a. Apodemia murphyi and Apodemia hepburni (dorsal surface). Upper left
— A. murphyi holotype male, MEXICO: Baja California Sur; Arroyo San
Bartolo, 28 Aug. 1 982, leg. Brown and Faulkner. Upper right — A. murphyi
allotype female, same data. Second left — A. murphyi male dark
phenotype, MEXICO: Baja California Sur; 12.2 mi. SE San Perdito near
Rancho Saucito, 8 Oct. 1981, leg. F. Andrews and D. Faulkner. Second
right — A. murphyi female dark phenotype, MEXICO: Baja California Sur;
7 mi. SE Guerrero Negro, 8 Apr. 1976, leg. Doyen and Rude. Third left —
A. murphyi aberrant female, MEXICO: Baja California Sur, 1 0 mi. N Bahia
Asuncion, 25-27 April 1984, leg. Bloomfield. Third right — A. murphyi
normal female, same data. Bottom left — A. hepburni male, MEXICO:
Baja California Sur; 2 mi. SW Cadauno, 26 Aug. 1982, leg. Faulkner and
Brown. Bottom right — A. hepburni female, same data.

Fig. 3b. Apodemia murphyi and Apodemia hepburni (ventral surface). Same
specimens as in Fig. 4a.

27 Sept. 1981 (1M, IF, SDNHM); Cabo Pulmo, 4 Nov. 1946 (1M, SDNHM);
1.5 mi. SW San Jose del Cabo, 30 Sept. 1981 (3M, SDNHM); San Jose del Cabo,
17 Feb. 1940 (4M, SDNHM), 1 July 1968 (1M, CM), 22 Nov. 1961 (IF, CM); 3 mi.
N San Jose del Cabo, 22-23 Nov. 1961 (2M, 5F, CM); Cabo San Lucas, Hotel
Finisterra, 8-10 Oct. 1979 (1M, IF, SDNHM); Cabo San Lucas, 22 March 1939
(IF, AMNH), no date (1M, AMNH), 2 Apr. 1949 (1M, SDNHM), 23 Nov. 1961
(1M, CM); 20 mi. N Cabo San Lucas, 29 Sept. 1970 (IF, CM); 15 mi. S La Paz, 1
Nov. 1946 (5M, 2F, SDNHM); La Paz airport, 10 Oct. 1979 (IF, SDNHM); 7 mi.
SW La Paz, 4 Aug. 1966 (IF, SDNHM); La Paz, 17-22 Sept. 1967 (1M,
SDNHM), 9 July 1968 (2F, CM); 13 Sept. 1959 (1M, CM); La Paz, Guaycura
Hotel grounds, 6-8 Nov. 1961 (5M, IF, CM); SE shore La Paz Harbor, 5 Nov.
1961 (1M, CM), 10 Nov. 1961 (5M, CM); E shore, La Paz Bay, 8 Nov. 1961 (6M,
8F, CM); 3 mi. S Santiago, 25 Oct., 1961 (IF, CM); Las Barracas, ca. 30 km
E Santiago, 7/12 Apr. 1982 (1M, CIS); Puerto Chileno, 22 Nov. 1961 (1M, CM);
Boca de la Sierra, 17-24 Nov. 1961 (1M, 2F, CM); Ro. Palmarito, 27 Oct. -5 Nov.

26(l-4):l-288, 1988

135

1961 (2M, 2F, CM); Rancho El Salto, 28 Oct. 1961 (1M, CM); Bahia de Palmes, 20
Nov. 1961 (2F, CM); Isla Espiritu Santo, 19-23 Feb. 1936 (IF, SDNHM), 17
April 1958 (1M, LACM), 14 July 1985 (1M, SDNHM), 30 Dec., 1938 (3M, 3F,
AMNH); San Jose I., Gulf of California, 5 March 1975 (IF, UCD); Isla Partida,
17 April 1958 (1M, LACM); Bahia Agua Verde, 20 April 1958 (2M, LACM); 31
km N Todos Santos, 29 Nov. 1980 (IF, SDNHM); 7 km S Candauno [sic], 26 Aug.
1982 (2F, SDNHM); Punta Conejo, ca. 32 km SW El Cien, 9 Jan. 1977 (1M, G. T.
Austin); Todos Santos Rd., ca. 42 km N Cabo San Lucas, 14 Jan. 1977 (1M, G. T.
Austin); Muertos Bay, 24 March 1939 (1M, AMNH), 29 Dec., 1938 (1M, AMNH).

DEPOSITION OF TYPE MATERIAL. The holotype, allotype, 36M and 17F
paratypes are deposited at the Natural History Museum, San Diego; 51M and
3 IF paratypes are in the Carnegie Museum of Natural History; 6M and 4F
paratypes are in the American Museum of Natural History; 4M paratypes are in
the Los Angeles County Museum; 1M paratype is in the collection of the
California Insect Survey; IF paratype is at the Bohart Museum, University of
California, Davis; IF paratype is in the private collection of J. Brock; 3M
paratypes are in the private collection of G. S. Forbes; and 2M paratypes are in
the author’s private collection.

TYPE LOCALITY. MEXICO: Baja California Sur; Arroyo San Bartolo. San
Bartolo is on Mexico Highway 1 between La Paz and San Jose del Cabo. All
specimens examined from south of 25°N latitude are designated paratypes.

DISTRIBUTION AND PHENOLOGY. Apodemia murphyi occurs throughout
much of Baja California Sur and to extreme southern Baja California Norte,
Mexico (Fig. 1). Its northern limit appears to be the Bahia de las Animas and
Bahia de Los Angeles area on the east coast (Rindge 1948, Holland 1972). No
records of an Apodemia palmerii- like butterfly exist north of this point (Rindge
1948, Powell 1958, Patterson and Powell 1959, Holland 1972) for nearly 450 km
virtually to the United States border (one specimen of A. p. palmerii from
Mexicah, Baja California Norte, AME) although Hoffmann (1976) indicated
that A. p. “marginalis” is found in Baja California.

The insect is apparently continuously brooded and has been collected in every
month. The majority of specimens (127 of 250 examined with dates) are from
October, November and December. This may reflect collecting patterns rather
than phenological patterns of the butterfly. Fresh specimens occur throughout
the year.

ETYMOLOGY. I name this insect after Dennis D. Murphy to whom I owe
numerous debts.

DIAGNOSIS AND DISCUSSION. The taxon Apodemia murphyi is most
distinctive. It is about the size of Apodemia palmerii but the sexes are nearly the
same (male x = 11.7, range = 10.4-13.2; female x = 11.6, range = 10.8-12.7,
October sample). The basic pattern above is similar to A. palmerii arizona but
the white spots are smaller in size (but are distinct and not smudged as on some
specimens of A. p. australis). The ventral pattern is very different from any
A. palmerii , especially the secondaries with the broad (up to ca. 25% of wing
surface) and continuous white postmedian band. On A. palmerii this band is
disjunct and more of a sinuous series of spots. The shape of the primaries is
different from any A. palmerii being more drawn out and pointed towards the
apex, especially on the male, and subfalcate. The fulvous basal area of the wings
is sharply set off from the dark distal area, particularly on summer and fall
specimens. These areas grade into one another on A. palmerii.

136

J.Res.Lepid.

An interesting aberrant female was seen (MEXICO: Baja California Sur; 10
mi. N Bahia Asuncion, 25—27 April 1984, leg. Bloomfield, SDNHM; Fig. 3).
Most of the postmedian white spots of the dorsum are absent. Those present
along with the basal white spots are small. These are similar on the ventral
primaries and the typical postmedian of the secondaries is absent anteriorly and
narrowed posteriorly. The black submarginal macules and the black lines
associated with the postmedian are nearly obsolete. This is not as extreme as the
aberrant A. p. arizona noted above. Aberrations are rare among the Riodininae
with none described for the North American fauna (Kendall and McGuire 1984).

The male genitalia, while similar to those of Apodemia palmerii, differ in
several respects (note above that genitalia of A. palmerii are constant over the
entire range of the species). Overall, the structures are slightly more massive
although the aedeagus is proportionally shorter, terminating just beyond the
lower arm of the valve (extends considerably further posteriorly on A. palmerii).
The uncus is relatively narrow (dorsal aspect) and rounded (posterior aspect).
The uncus of A. palmerii is broader with more rounded lobes and flatter. The
falces are stouter on A. murphyi , the dorsal arm averages slightly longer and the
ventral arm is considerably shorter than on A. palmerii (dorsal arm: ventral
arm = 0.7 for A. murphyi , 0.5 for A. palmerii). The valves of A. murphyi are
broader and stouter with a pointed (but not hooked) upper process. The saccus is
proportionally longer on A. murphyi and the vinculum is straighter and not
broadly rounded ventrally as on A. palmerii.

Apodemia murphyi resembles Apodemia hepburni Godman and Salvin in
wing shape and the continuous postmedian band on the ventral secondaries
(Fig. 3). The latter is otherwise distinctive with a less complete spot pattern
(lacking the submarginal series among others), no clear basal fulvous area on
the dorsal primaries and a considerably narrower postmedian band on the
ventral secondaries. The figures show these characters clearly. Apodemia
hepburni is sympatric and synchronic with A. murphyi at many localities from
Mulege southward. Apodemia hepburni records extend from 26 August through
6 April; A. murphyi from 26 July through 8 April at these localities. Label data
(also fide J. W. Brown) indicates that A. murphyi nectars commonly on Bebbia
juncae (Asteraceae) and also on Melochia tomentosa (Sterculiaceae) and Bac-
charis sp. (Asteraceae). The species is associated with Prosopis glandulosa
(Fabaceae) throughout its range; this probably is a larval host plant.

Acknowledgements. I thank several curators and others for loans of specimens
from and/or space to work at their respective museums: L. D. and J. Y. Miller,
Allyn Museum of Entomology, Sarasota, Florida (AME); F. H. Rindge, American
Museum of Natural History, New York (AMNH); D. K. Faulkner, Natural
History Museum, San Diego, California (SDNHM); J. P. Donahue, Los Angeles
County Museum, Los Angeles, California (LACM); R. K. Robbins, National
Museum of Natural History, Washington, D. C. (USNM); J. E. Rawlins and C.
W. Young, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania
(CM); J. A. Powell, California Insect Survey, Berkeley, California (CIS);
R. Schuster, Bohart Museum, University of California, Davis (UCD); T. W.
Davies, California Academy of Sciences (CAS); and G. Haijes, Nevada State
Museum, Carson City, Nevada. I gratefully acknowledge the loan of specimens
from the private collections of J. Brock, Tucson, Arizona (JB); G. S. Forbes, Las
Cruces, New Mexico (FG); E. V. Gage, San Antonio, Texas; R. W. Holland,

26(l-4):l-288, 1988

137

Albuquerque, New Mexico; C. S. Lawson, Las Vegas, Nevada; S. McKown,
Boulder City, Nevada; D. Mullins, Tucson, Arizona; P. J. and S. Savage, St.
George, Utah; M. Smith, Las Vegas, Nevada; and S. Spomer, Lincoln, Nebraska.
L.D. Miller called my attention to the Apodemia palmerii australis phenotype
and provided habitat information. R. O. Kendall commented on Texas records.
J. W. Brown was helpful in providing information about Apodemia murphyi and
on Baja California in general and D. D. Murphy read manuscript drafts and
offered helpful suggestions. The manuscript was also improved from suggestions
from R. H. T. Mattoni and an anonymous reviewer.

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140 J.Res.Lepid.

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SPECIMENS EXAMINED

Apodemia palmerii palmerii

UNITED STATES: Arizona (37M, 23F; 27 March-7 September): Coconino,
Mohave, Yuma counties; CALIFORNIA (321M, 206F; 15 March-12 November):
Imperial, Inyo, Riverside, San Bernardino, San Diego counties; NEVADA
(211M, 126F; 17 April-13 October): Clark, Lincoln, Nye counties; UTAH (43M,
29F; 20 May- 13 September): Washington County. MEXICO: BAJA
CALIFORNIA NORTE (IF; August).

Apodemia palmerii arizona

UNITED STATES: ARIZONA (type series plus 407M, 335F; 17 April-24
October): Cochise, Coconino, Gila, Graham, Maricopa, Pima, Pinal, Santa Cruz,
Yavapai counties; NEW MEXICO (19M, 15F; 12 May-11 September): Dona
Ana, Hidalgo, Lincoln, Luna, Otero, Socorro counties; TEXAS (8M, 25F; 13
April-9 September): Brewster, El Paso, Presidio, Terrell counties. MEXICO:
CHIHUAHUA (2M, 6F; 14 July-31 August); SINALOA (1M, IF; 28 June);
SONORA (35M, 31F; 12 March, 3 August-25 October).

Apodemia palmerii australis

MEXICO: AGUASCALIENTES (1M; 27 August); DURANGO (type series plus
16M, IF; 30 July-20 August); HIDALGO (4M, 9F; 30 April, May, 19 July-8
August); JALISCO (1M, 26 July); MICHOACAN (15M, 3F; 9 August);
QUERETARO (2F; 20 July); SAN LUIS POTOSI (5M, 5F; 20 July-23 August);
TAMAULIPAS (4M; 20 September); ZACATAS (1M, IF; 30 August).
Apodemia murphyi

MEXICO: BAJA CALIFORNIA NORTE (1M, 3F; 30 March, 19 September-6
October); BAJA CALIFORNIA SUR (type series plus 80M, 33F; 7 March-28
June, 11 August-December including 5M at AME and 1M, IF at SDNHM
labeled Baja California Norte).

OTHER RECORDS
Apodemia palmerii

UNITED STATES: NEW MEXICO; Catron, Grant counties (Ferris 1976);
TEXAS: (31 March-7 October; Tilden 1974, Freeman 1981): Culberson, Jeff
Davis, Pecos counties (Tilden 1974, fide R. O. Kendall).

Apodemia murphyi

MEXICO: BAJA CALIFORNIA NORTE (8 May; Rindge 1948).

Journal of Research on the Lepidoptera

26(1-4):141-160, 1988

Correlations of Ultrastructure and Pigmentation
Suggest How Genes Control Development of Wing Scales
of Heliconius Butterflies

Lawrence E. Gilbert
and

Hugh S. Forrest
and

Thomas D. Schultz +
and

Donald J. Harvey*

Department of Zoology, The University of Texas at Austin, Austin, Texas 78712

Introduction

In the last two decades there have been extensive genetic studies of
the mimetic wing patterns of Heliconius butterflies (Turner, 1981;
Sheppard et al., 1985), but these analyses have not probed the precise
nature of color patterns at the ultrastructural level. Thus, while many
ecological and evolutionary phenomena have been elucidated, the
nature of the genes involved and the mode of their action in the course of
wing pattern development remains obscure.

One another front there is renewed interest in general models of
pattern formation in butterfly wings (Nijhout, 1986), but studies of
pattern development have relied upon comparisons among related
species or upon experimental manipulations of wing development
within species. Remarkably, there has been a lack of genetical analysis
applied to the problem of butterfly wing pattern development on the one
hand, and little attention given to the chemical and ultrastructural
basis of “color pattern” on the other, although excellent studies have
been carried out on chemistry (e.g., Umebachi and Yoshida, 1970;
Descimon, 1975) and ultrastructure (e.g., Ghiradella, 1974; 1985)
separately.

In an attempt to refine genetic hypotheses which explain the variation
of wing pattern observed in crosses involving different races and species
of Heliconius (Sheppard et al. 1985; Gilbert, in prep.), it appeared

+ Current Address: Division of Entomology, Peabody Museum of Natural History, Yale
University, New Haven, CT 06511

*Current Address: Department of Entomology NHB Stop 127, Smithsonian Institution,
Washington, DC 20560

142

J.Res.Lepid.

appropriate to investigate the chemical and morphological basis for
color and pattern. This paper summarizes our initial investigations of
wing scale structure and chemistry on four species of Heliconius which
have been subjects of genetic studies in the senior author’s laboratory.

The findings presented for Heliconius below provide the first clear
evidence from lepidopteran wings that genetic control of pigmentation
patterns simultaneously involves patterns of differentiation in scale
ultrastructure, a result anticipated in general terms by Descimon
(1965). Thus, beyond elucidating the connections between genes and
wing patterns in butterflies, the results suggest that Heliconius wings
may provide a useful system for addressing general questions about the
genetics of pattern development.

Material Examined

All butterflies for ultrastructural and chemical studies were reared in
greenhouses at Patterson Laboratory, The University of Texas at
Austin. Heliconius species examined included Heliconius cydno galan-
thus (stock origin, La Selva, Costa Rica), Heliconius pachinus (stock
origin, Osa Peninsula, Costa Rica), Heliconius melpomene rosina (stock
origin, Osa Peninsula, Costa Rica), and Heliconius ismenius clarescens
(stock origin, Osa Peninsula, Costa Rica). Hybrid “bar-shadow” regions
(explained below) were from FI hybrids of the H. cydno and H. pachinus
stocks above. In addition, fore wing red/brown scales were examined in
H. cydno — H . melpomene crosses. Illustrations of these species may be
found in DeVries (1987) and of their hybrids in Gilbert (1984).

Because a wide variety of methods are used in this study, necessary
details of techniques will be provided below.

SCALE MORPHOLOGY

Scales were examined by standard methods of scanning electron
microscopy. Dry wing fragments with uniform scale color were coated
with 25 A of gold-palladium in a Hummer V sputter coater and examined
at 600 and 10,000X using an ISI Super IIIA. Scale cross sections were
created by cutting wing fragment with a razor blade and searching that
area for appropriately cut scales.

Descriptive terminology used below follows the system developed by
Downey and Allyn (1975). It should also be stressed that while we are
confident in distinguishing the following major scale types, interpreta¬
tion of many morphological details is tentative.

Type I scales. Yellow/white (Fig. 1 A, B, C, D; Fig. 5B; Fig. 6A)

In Heliconius , yellow and white scales appear to represent the same
morphological type. Average spacing of scute peaks (= lamellae of
Ghiradella, 1985) along the ridge is approximately the same as the
inter-ridge distance. Obverse membrane obscures the scale internal

26(l-4):l-288, 1988

143

structure. Many variable sized windows may occur in the membrane,
especially in the central region of the scale. Transverse flutes (= micro¬
ribs of Ghiradella, 1985) run over the membrane surface between, and
perpendicular to, longitudinal ridges. Such flutes are evenly spaced and
occur at a density of 8-10 per inter-scute interval.

White and yellow scales appear to lack crossribs, and scale cross
sections show trabeculae primarily below longitudinal ridges.

The spacing of ridges in Type I scales is narrower on the dorsal wing
surface, so that dorsal wing scales often have 1.5 to 2 times the number
of ridges per scale width as do ventral wing scales (compare Fig. IB vs.
1A). This ultrastructural difference may help account for the sheen and
richer colors of the dorsal versus the ventral wing surfaces.

Type II scales. Black (Fig. 2A, B, C, D; Fig. 5A, C; Fig. 6B)

The melanic scales of Heliconius possess longitudinal ridges con¬
nected by ladder-like crossribs, most of which are supported by trabe¬
culae. Crossribs appear more narrow than ridges and are arranged in
poorly aligned rows. There is usually no obverse membrane in melanic
scales and flutes are visible only on the vertical walls of longitudinal
ridges.

As in Type I scales, ridges are spaced more widely on ventral wing
scales than on dorsal scales. In both H. cydno galanthus and H.
pachinus, dorsal scales noted as “dull” proved to have more widely
spaced ridges than those noted to be “shiny” (e.g., Fig. 2C vs. 2B).

Type IT Hybrid “bar-shadow” scales. Black (Fig. 3 A, B, C)

The ‘"bar shadow” is a region of altered reflectance on the melanic
region of a hybrid Heliconius ventral hindwing. This region corresponds
to the location of yellow scales in one parent race of a cross, the other
parent of which possess a totally melanic hindwing. The bar shadow is
used to diagnose hybrid genotypes in ecological genetic studies (Mallet,
1986).

These scales are identical to Type II scales except that roughly 5% of
spaces between crossribs are covered by obverse membrane (Fig. 3). In
some cases the membrane is intact over the inter-rib space, but in most
cases these scales resemble partly dissolved tissue draped over chicken
wire. This subtle change is visible to the naked eye, but not under light
microscope. This scale type might be viewed as a small step toward a
morphological hybrid of Type I and Type II scales. However the
membrane, where present, lacks supportive flutes and the scales are
otherwise identical to Type II scales.

Type HI scales. Reds and browns (Fig. 4A, B, C, D; Fig. 5D; Fig. 60

“Red and brown” scales in Heliconius include orange, orange-brown,
brown, pink, and red scales. These share a basic morphology, Type III.
Crossribs often appear to be wider than those of Type II scales, and one

144

J.Res.Lepid.

or two strengthening flutes pass over each crossrib, connecting adjacent
ridges. In addition, obverse membrane appears to be retained over each
crossrib and immediately adjacent to longitudinal ridge. This may
account for the thicker appearance of ridges and ribs in Type III scales,
as well as the angular appearance of crossribs.

Cross sections of Type III scales do not reveal trabeculae supporting
presumptive crossribs, but they can be seen supporting longitudinal
ridges, and may simply occur less frequently than in Type II scales.
Without further detailed work, it is not possible to exclude the possibility
that strips of membrane supported by flutes function as pseudo-crossribs
rather than overlay them as suggested above. Close examination of
Type III scales suggests their closer relationship to Type I than to Type
II scales. For example, the inter-ridge space on the left extreme of the
brown scale in Fig. 4A is virtually identical to the obverse membrane of
a dorsal Type I scale (e.g., Fig. IB). These characteristics gradually
change to typical Type III features toward the center of the scale. Like
Types I and II, Type III scales typically possess more narrow spacing of
the longitudinal ridges on the dorsal wing surface.

Scale Chemistry

White Scales: No pigment

White scales of Heliconius oydno possess a highly reflective quality or
sheen quite unlike the flat white of Pieris. Under light microscope at low
power, these scales are brighter where two or more overlap. These
observations suggested a structural rather than chemical basis for the
white color.

To test this possibility, scales were immersed in a solution whose
refractive index is near that of chitin (1.55). When single scales were
observed in such a solution (xylene or Permount) against a black
background, they became essentially transparent. Scales in xylene
regain their white luster when the liquid evaporates. Comparisons with
yellow and black scales indicate that luster, but not color, disappears in
these liquids. Uric acid tests were negative on chromatographs of Type I
scale areas. White scales in Heliconius are therefore due to structural
features of the scale rather than pigments.

Yellow Scales: 3-hydroxykynurenine

A small circle of the yellow part of the wing was cut out with a cork
borer, and positioned carefully at the aperture of the Cary Recording
Spectrophotometer. Measurement of the UV spectrum revealed peaks
at 282nm and 405nm (a rather broad peak). These are essentially
identical to the peaks produced by 3-hydroxykynurenine in 0.1 N NaOH
(pH 14) (285 and 395nm).

Extraction of the pigment using water or dimethyl sulfoxide and

26(l-4):l-288, 1988

145

rerunning the spectrum of this extract in 0.1N NaOH or 0.1N HC1 gave
the following values:

X max 0.1N NaOH
280(285)

395(395)

X max 0.1N HCl
252(252)

312 shoulder(312)

3-hydroxykynurenine peaks (in parentheses)

These data indicate that the yellow pigment in Heliconius spp. is the
alkaline form of 3-hydroxykynurenine. This pigment is previously
described from Heliconius (Brown, 1967).

An interesting question concerns the maintenance of 3-hydroxyky-
nurenine in its alkaline form in the wing scales of Heliconius. Chro¬
matographic studies were carried out to elucidate this phenomenon.
Fragments of yellow wing areas of H. pachinus were ultrasonicated,
then agitated in 80% methanol. This extract was spotted on Whatman
No. 1 filter paper and subjected to one dimensional chromatography
using BAW, n-butanol/acetic acid/water (4:1:1) as solvent. A ninhydrin
test (lg of ninhydrin dissolved in 50ml acetone; chromatograph was
immersed in this solution, allowed to dry, and heated at 110°F until color
developed) revealed that an amino acid or peptide was located in the
identical spot with the alkaline form of 3-hydroxykynurenine (revealed
by UV light).

A comparison with white scales using the same procedure showed the
identical ninhydrin sensitive spot, but no yellow pigment. Two-dimen¬
sional chromatographic studies of all basic amino acids using the same
solvent, BAW, did not duplicate the spot derived from Heliconius wing
extract. It is therefore likely that a peptide or small polypeptide is
responsible for keeping 3-hydroxykynurenine in the alkaline state. The
precise location of this complex within the scale is not yet determined.

Black Scales: melanin

Chromatographic evidence verified that black scales contain melanin
but tryptophan is also present in extracts of black scales. Melanie scales
embedded in paraffin, sectioned, and examined with light microscope
revealed that pigment is found in the walls of ridges and in crossribs.

Brown and Red Scales: xanthommatins

Wings were extracted with dimethyl sulfoxide (DMSO) or with 2%
HCl in methanol. The spectrum of the extracts and of standard com¬
pounds are given in Table 1. There are two problems of interpretation
with these data. First, the spectra of xanthommatin and dihydroxan-
thommatin are notoriously difficult to reproduce because of rapid
decomposition and slight changes in state of reduction (see Linzen,
1974). Second, it was not possible to make comparisons of extracted

146

J.Res.Lepid.

Table 1. Extraction methods and absorbance values (nm) of chemical
standards and wing pigments of Heliconius (* Values from
Denys, 1982)

PIGMENTS

EXTRACTION

ABSORBANCE MAXIMA (nm) CONDITIONS

Brown from

H. ismenius

DMSO

DMSO

pH7-7.5

Acid Methanol

5N HC1

2% Digitoxin
pH6.5

2% Digitoxin
pH 10.4

440-450,365

1

DMSO acidified
to 5N HC1

450,360

2

acid methanol

450, no distinct
peak at 360

3

Red from

H. melpomene

DMSO

490,365

4

DMSO extract +
acetone/ether,
resulting ppte.
in H20 pH7.0

465,368

5

acid methanol

448, no distinct
peak at 360

6

Red from

H. pachinus

acid methanol

458,360-380

7

acid methanol,
oxidised with
NaN02

448

8

Brown from
pure sample
xanthommatin

dissolved as
indicated

440

475-480,

370-375

*450

*478

9

Red from
pure sample
dihydro¬
xanthommatin

dissolved as
indicated

505-510

475-360

500(shoulder),

390

10

Reduced with
NaBH4

490,370

11

1

2

3

4

5 | 6

pigments from Heliconius wings with standard samples of xanthom-
matins under absolutely identical or controlled conditions. However,
taken as a whole, the data indicate first that the major brown or red
pigments of Heliconius butterflies are xanthommatin and dihydroxan-
thommatin and second, that variations in color from bright red to brown
are due to variations in the state of oxidation of dihydroxanthommatin
(or the state of reduction of xanthommatin).

First note the correspondence in the spectral maximum between the
brown pigment of H. ismenius and that of xanthommatin (Table 1, row 1
and 2 versus row 9). Further note the DMSO extract of H. melpomene
red pigment (Table 1, row 5, col. 1), the spectrum of which peaks near
that of dihydroxanthommatin under reduced conditions (Table 1, row
11, col. 3). Obviously these extracts of red dihydroxanthommatin are in
various stages of oxidation to xanthommatin.

Given the in vitro instability of the reduced red form of xanthom-
matins, the observed stability of various shades of orange and red on the
wings of various races of H. melpomene and H. erato presents an

26(l-4):l-288, 1988

147

interesting mystery. However, the likelihood that 3-hydroxy kynurenine
is maintained in an alkaline state by a gene product suggests that such
associations may allow races to “select” localized pH conditions within
Type III scales by modification of a peptide or protein associated with
xanthommatin pigment, and thereby affect subtle variation in the
actual color displayed on the wing.

Evidence from H. cydon X H. melpomene crosses indicates genetic
control of separate factors maintaining the reduced form of xanthom¬
matin (red) in H. melpomene. Thus, in all FI hybrids of H. pachinus or H.
cydno with red forewing banded H. melpomene races (illustrated in
Gilbert, 1984), brown scales appear on the ventral side of the dorsal
fore wing red band, and with appropriate crosses, one can convert the
dorsal red forewing band of hybrids to brown (Gilbert, unpublished
data).

Discussion

Different hypotheses can be proposed for how genes determine final
color patterns in Heliconius. At one extreme, scale morphology and
pigmentation would be separately determined by independently re¬
gulated genes such that any combination of structure and color could
occur. This possibility is not the case in Heliconius because of melanic
scales (including bar-shadow scales), which are consistently found to
have a particular subset of ultrastructures, scales with Type I ultrastruc¬
ture which consistently lack melanin or xanthommatin, and Type III
scales are never white, yellow, or melanic.

At the other extreme, genes which determine pigment production in a
developing scale might pleiotropically determine its ultrastructure.
This would be the case if the product of a single gene directly or
indirectly regulates both morphological events and the pigment path¬
way within a scale. In Heliconius , this possibility appears to hold true at
the level of major pigmentation differences (eg. xanthommatin vs.
melanin). However, some pigment variation such as brown vs. red in
Type III scales, or white vs. yellow in Type I scales, represents minor
pigment variation within the major categories. We hypothesize that
such minor variation in Type III scales is based on variation in genes
coding for those peptides which act to stabilize pigments at particular
oxidation states in the scales.

Any useful model for scale development in Heliconius should explain
the observed correlations of structure and pigmentation (summarized in
Table 2) in genetic and chemical terms. It should also be in general
accord with current knowledge of scale development, pigment chemi¬
stry, and genetics. Fortunately, development of scale pigmentation has
been carefully studied in another nymphalid genus, the pigments
involved are relatively well-studied in other systems, and extensive
classical genetics is available for Heliconius.

148

J.Res.Lepid.

With respect to scale development, Nijhout’s (1980) observations and
experiments provide a useful model for the development of different
colored melanic scales in Precis (Nijhout, 1980, p.287).

1. Enzymes for pigment synthesis are insoluble but active within
cuticle of the scale.

2. Substrates for pigment synthesis circulate in the hemolymph and
are produced in sequence.

3. Substrates can gain access to scales at all times.

4. Scales in each presumptive color region possess only a single
enzyme and are capable of utilizing only a single substrate.

Nijhout (1980) also observed that longitudinal ridges formed before
melanin deposition, indicating that the pigment per se only stiffens the
scale, but does not direct its morphogenesis. We therefore assume that
in Heliconius , any pleiotrophic effects of genes involved in pigment
pathways on scale structural distinctiveness is not via the pigment, its
precursors, or substrates. Rather it seems most likely that the product of
a “scale selector” gene acts as a turn-on switch for other genes involved
in scale ultrastructure on the one hand, and genes for pigment pathway
enzymes on the other.

Our interpretations of the chemistry of yellow, red, and brown
variation in Heliconius benefit from the extensive genetic and bio¬
chemical work on Drosophila eye color variation which is based on the
same ommochrome pathway (Summers et al ., 1982). Xanthommatin
pigments derive from tryptophan via intermediates such as kynurenine
(Linzen, 1974), but two lines of evidence suggest that the substrate for
xanthommatin production is 3-hydroxy kynurenine. First, in Drosophila
eyes, normal xanthommatin production depends on external kynurenine
and/or 3-hydroxykynurenine supplied via the hemolymph (Summers et
al., 1982). Second, Linzen (1970) reviews evidence that a) in holometa-
bolous insects, including Lepidoptera, tryptophan accumulation in
hemolymph and other tissues is transitory and b) 3-hydroxykynurenine
is the metabolite most likely to persist at elevated levels. Thus, it is
reasonable to assume that the substrate for xanthommatin in Heliconius
Type III scales is 3-hydroxykynurenine.

Similarly, although melanins arise ultimately from oxidation of
tyrosine, the substrates for melanin production are likely to be dopa or
dopamine if Heliconius follows the usual pattern for insects (Wiggles-
worth, 1972) and for Precis wing scales (Nijhout, 1980).

Association of xanthommatin and other ommochromes with specific
proteins in silkworm blood (see Linzen, 1974 for review) make our
suggestions about mechanisms of color fine- adjustment and stability a
credible working hypothesis. On the other hand, reports of ommochrome-
binding protein in cecropia moth eyes (Ajami and Riddiford, 1971) have
not been verified in parallel studies on Drosophila (Wiley and Forrest,
1979), nor have the subtle variations in Drosophila eye color been
adequately explained.

26(l-4):l-288, 1988

149

Additional parts of the Heliconius scale puzzle are provided by genetic
evidence. Certain genes for xanthommatin scales (Type III) are domin¬
ant (Sheppard et al., 1985) or epistatic (in single dose) to those for
melanic scales (Type II) (Gilbert, in prep.) in H. melpomene. Other
dominant or epistatic genes, active in the same regions of the wing, may
replace yellow scales (Type I) with melanic or xanthommatin containing
scales (see Sheppard etal ., 1985). We suggest these observations are due
to our hypothesized scale selector genes, the interaction of which
generally produces an unambiguous scale type in the following order of
dominance or epistasis: III > II > I. Bar shadow scales on the ventral
hindwing may represent an exception to this rule if they indeed possess
intermediate features.

Genetic variation for pigmentation within scale types appears to have
no common theme. In Type I scales, white is dominant to yellow (see FI
of H. cydno X H. pachinus, (Gilbert, 1984). This is counter to what we
would expect if the heterozygote simply possesses one half the amount of
yellow pigment. We hypothesize a gene involved with transport of 3-
hydroxykynurenine into the developing Type I scale. The bar shadow
variant of Type II scales (ID probably reflects dosage of Type II selector
gene, M, (but only expresses on one wing surface!). Color variation of
xanthommatin pigments may be due to a structural gene for the binding
peptide as previously discussed.

At this stage of knowledge, many alternative models of Heliconius
scale development might be equally difficult to reject. With this caveat,
we present a model which is consistent with the observed relationships
of scale structure and color (Table 2) and which assumes as valid, the
foregoing points about scale development, pigment chemistry, and
genetics. Finally, for simplicity, we develop the model as a series of
binary choices which depend upon the state of scale selector genes in
cells which give rise to the scales. The following model should be
considered a tentative scheme rather than a well-substantiated theory.

During the course of development, cells would be fated to give rise to a
particular scale type at a particular wing location by the combination of
selector genes which are switched on or off. The threshold conditions for
such switching might allow trichogen cells of the same genotype to end
up as different scales types depending upon the strength of morphogen
signals at that location (see Nijhout, 1986).

A fundamental decision in scale development seems to be between
Type I versus Type II or III, because Type I scales do not require pigment
to stiffen, and are apparently not manufacturing complex pigments
from simple substrates. For simplicity, we consider this scale type the
null state, that scale type which develops if no other selector genes are
switched on.

Next, if Type II selector switch gene M is turned on, cells are fated to
develop Type II or Type III scales. Given that only M is on, and given
appropriate positional information, the M + signal would turn on

150

J.Res.Lepid.

Table 2. A summary of the
relationship between scale mor¬
phological types and scale pig¬
mentation observed in the four
Heliconius species and two inter¬
specific crosses of this study. Type
IF refers to the bar shadow scale
type, yellow, brown, and red refer
to 3-hydroxykynurenine, xan-
thommatin, and dihydroxan-
thommatin respectively (see text).

PIGMENTATION

none

yellow

brown

T3

a

melanic

I

X

X

II

X

II'

X

m

X

X

morphological programs and melanin pathway enzymes. However, the
scale would not melanize and stiffen until dopa or dopamine circulate in
the hemolymph.

In keeping with a binary decision model of genetic determination, we
suggest that the selector gene for Type III scales, X, can only be
expressed in M 4- cells, and that its signal initiates Type III morphology
and turns on genes for xanthommatin pathway enzymes. Genetic
evidence summarized above indicated that M++X+ cells give rise to
Type III, xanthommatin containing scales.

Thus, it appears that the xanthommatin pathway inhibits in melanin
pathway by a method similar to its inhibition by another oxidative
pathway, xanthopterine synthesis (Wigglesworth, 1972). Since in
Heliconius pupae, homozygous and heterozygous fore wing Type III
patches develop xanthommatin well ahead of the melanization of Type
II areas (Gilbert, unpublished observation), it may be that the pigment
itself inhibits the oxidation of substrates of the melanin pathway as is
the case with xanthopterine and melanin. In explaining the epistasis of
X over M in determining scale morphology, it may be less complicated to
assume that morphology is a direct result of pigment-scale interaction.
However, as one reviewer pointed out, in the absence of further
information, independent determination is a better null hypothesis. In
our model therefore, the X+ signal overrides M++ to redirect morpho¬
genesis, and acts separately on genes involved with morphology and
pigmentation as M is hypothesized to do.

This scheme of developmental genetic control is summarized by
Figure 7. This diagram also shows the final genetically controlled
decisions which occur after scale type is established which we have
discussed above. For each branch, the gene dosages necessary for each
state of a scale is indicated by plus (one gene dose) or zero (null).

26(l-4):l-288, 1988

151

Summary

Scanning electron microscopy reveals three morphological categories
of wing scales in Heliconius butterflies. Type I, white or yellow scales,
possess an obverse membrane between longitudinal ridges and lack
conspicuous crossribs. Type II, melanic scales, have ladder-like, regular
crossribs supported by trabeculae. Type III, red or brown scales, are
characterized by crossribs which feature flutes and a thicker, more
angular appearance. In hybrids, whose parents possess Type I and Type
II scales on the hind wing bar respectively, the “bar shadow” scales
which replace the yellow bar appear to be a slightly modified version of
Type II scales.

Spectroscopic analyses reveal that yellow, red, and brown pigments
are tryptophan derived 3-hydroxykynurenine, dihydroxanthommatin,
and xanthommatin, respectively. White is a structural color expressed
when yellow pigment is not present, while red and brown are different
oxidation states of xanthommatin. Chromatographic evidence suggests
the possibility that unstable forms of pigments in this pathway are
maintained by association with peptides in the scale. Thus, although
substantial color variation occurs within scale morphological types, it is
chemically trivial. These observations are supplemented by evidence
from the literature to develop an hypothesis for the relationship
between genes, scale pigmentation, and scale structure (Figure 7).

Because of the variety of scale morphology and pigment chemistry
within the Lepidoptera, it is not possible to assess the degree to which
this scheme for Heliconius wing color pattern constitutes a model for
other groups. However, it will be surprising if the Heliconius system
described here turns out to be other than a variation on a theme common
to all butterflies and moths. Indeed, a similar correlation of color and
structure has been described for zygaenid moths (Burgeff and Schneider,
1979).

More generally, Heliconius wings may contribute to some of the
unsolved problems of the genetics and development of tissue specific
ommochrome pigmentation. This is because one can work with scale
specific regulation of the pathway on the wings within species having
distinctively patterned genotypic varieties or races, rather than rely on
constitutive mutants.

Acknowledgements. We gratefully acknowledge the help of Ian Millett who
conducted preliminary work on wing pigments, Bob Riess, who carried out most
of the SEM work, and Sharon Bramblett, who helped rear butterflies and typed
the various versions of the manuscript. We thank Susan Weller, Jim Bull, and
Suzanne Dyby, all of whom read and criticized the manuscript. We also thank
two constructively critical reviewers whose suggestions improved the paper.
This was work was supported by a University of Texas University Research
Institute grant R-511 and (indirectly) by NSF BRS-8315399 to LEG.

152

J.Res.Lepid.

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BURGEFF, H. & L. SCHNEIDER, 1979. Elektronenmikroskopische Untersuchungen
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DENYS, C. J., 1982. Ommochrome pigments in the eyes of Euphausia superba.
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DESCIMON, H., 1965. Ultrastructure et pigmentation des ecailles des Lepidopteres.
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DESCIMON, H., 1975. Biology of pigmentation in Pieridae butterflies. In: Chemistry
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DEVRIES, P. J., 1987. The Butterflies of Costa Rica and their Natural History.
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DOWNEY, J. C. & A. C. ALLYN, 1975. Wing-scale morphology and nomenclature.
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GHIRADELLA, H., 1974. Development of UV reflecting butterfly scales: how to
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GHIRADELLA, H., 1985. Structure and development of iridescent Lepidopteran
scales: the Papilionidae as a showcase family. Ann. Entomol. Soc. Amer. 78:
252-267.

GILBERT, L. E., 1984. The biology of butterfly communities. In: The Biology of
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LINZEN, B., 1970. Zur Biosynthese von Ommochromen, I. Einbau 35S-markierter
Vorstufen in Ommine. Hoppe-Seyler’s Z. physiol. Chem. 351:622-628.
LINZEN, B., 1974. The tryptophan —> ommochrome pathway in insects. Adv.
Insect Physiol. 10: 112-246.

MALLET, J., 1986. Hybrid zones of Heliconius butterflies in Panama and the
stability and movement of warning colour dines. Heredity 56:191-202.
NIJHOUT, H. F., 1980. Ontogeny of the color pattern on the wings of Precis coenia
(Lepidoptera: Nymphalidae). Dev. Biol. 80:275-288.

NIJHOUT, H. F., 1986. Pattern and pattern diversity on lepidopteran wings. Biosci.
36:527-53.

SHEPPARD, P. M„ J. R. G. TURNER, K. S. BROWN, W. W. BENSON, & M. C. SINGER, 1985.
Genetics and the evolution of Mullerian mimicry in Heliconius butterflies.
Phil. Trans. Roy. Soc. Lond. 308:433-613.

SUMMERS, K. M„ A. J. HOWELLS, & N. A. PYLIOTIS, 1982. Biology of eye pigmentation
in insects. Adv. Insect Physiol. 16:119-166.

TURNER, J. R. G., 1981. Adaptation and evolution in Heliconius : A defense of
neoDarwinism. Ann. Rev. Ecol. Syst. 12:99-121.

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WIGGLESWORTH, V. B., 1972. The Principles of Insect Physiology. 7th ed. London,
Chapman and Hall. 827 pp.

WILEY, K. & H. S. FORREST, 1979. Drosophila melanogaster lacks eye-pigment
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154

J.Res.Lepid.

Fig. 1. Type I, white or yellow scales. All cover scales viewed perpendicular to
surface at 10k. A. White scale, ventral forewing, H. cydno galanthus. B.
White scale, dorsal forewing, H. cydno galanthus. C. Yellow scale, dorsal
hindwing, H. pachinus. D. Yellow scale, dorsal forewing, H. pachinus. Note
on bottom left of D, where ridge spacing increases, obverse membrane of
dorsal scale resembles that of a ventral scale.

J

Fig. 2. Type II, melanic scales. All viewed approximately perpendicular (± 1 0°). A.
Ventral forewing, H. cydno galanthus. B. Dorsal hindwing (shiny scale), H.
cydno galanthus. C. Dorsal forewing (dull area), H. cydno galanthus. D.
Dorsal forewing, H. ismenius.

155

26(l-4):l-288, 1988

156

J.Res.Lepid.

zwunmm

IIWuwih

Sssasss

Fig. 3. Type II, "shadow" scales. These ventral hindwing scales lie in zones of
altered reflectance and are diagnostic of hybrids between forms with yellow
hindwing bars X forms with all black hindwings. A. Shadow region of a H.
cydno galanthus X H. pachinus, FI hindwing. B. Non-shadow region of a
H. cydno galanthus X H. pachinus, FI hindwing. C. Shadow region of H.
cydno galanthus X H. pachinus, FI hindwing, D. Shadow region of H.
cydno galanthus X H. pachinus, backcross hindwing.

26(l-4):l-288, 1988

157

9 *

**}**>$*

•»*

* -:nm i

Fig. 4. Type III, red or brown scales. A. Ventral hindwing, H. cydno galanthus
(brown). B. Ventral hindwing, H. pachinus (basal red spot). C. Ventral
forewing, hybrid H. cydno with H. melpomene forewing band (brown). D.
Dorsal forewing, hybrid H. cydno with H. melpomene forewing band
(brown).

if h<

Fig. 5. Angled views of various scale types (all of H. pachinus) showing scutes, all
at 10k. A. Dorsal hindwing, melanic, H. pachinus. B. Ventral hindwing,
yellow, H. pachinus. C. Ventral hindwing, melanic, H. pachinus D. Ventral
hindwing, basal red scale, H. pachinus.

158

J. Res. Lepid.

26(l-4):l-288, 1988

159

Fig. 6. Cross sections of various scale types, all at 1 0k. A. Ventral forewing, white
(Type I), H. cydno. B. Ventral forewing, melanic (Type II), H. cydno. C.
Basal red spot, ventral hindwing, H. pachinus. Dorsal hindwing, melanic, H.
cydno.

160

J.Res.Lepid.

trichogen

cell

M+

II or m

xanthommatin pathway

(T)3-OH-K— ►
T xanthommatin

X+

m

i

o+,++

00

(off)
-► (on)

melanin pathway

dopa — ► melanin

X- I,

- • —

II or IT

0+

red

in

m

brown

melanic

IF

bar shadow

M-

3-OH-K

I

00

0+,++

yellow

white

undifferentiated cells mature scales

Fig. 7. Hypothetical scheme for genetic control of Heliconius wing scale develop¬
ment based on morphological, chemical, and genetic information discussed
in text. Solid circle represents time that morphological characteristics of
mature scale begin to be established. M and X are selector genes regulating
morphological decisions and pigment pathways as shown. Effect of genes
which act within major scale categories are indicated on the final branches
of the diagrams in terms of doses (indicated by Type I scales vary in
terms of a gene which affects transport of 3-hydroxykynurenine (3-OH-K)
to the developing scale, one dose (o+) gives a white scale. Type IF scales
probably represent scales heterozygous for M (o+). Type III scales vary
according to a structural gene for a pigment binding peptide. One dose (o+)
stabilizes xanthommatin in its reduced state. (See text)

Journal of Research on the Lepidoptera

26(1-4):161-172, 1988

A mutant affecting wing pattern in Parnassius apollo
(Linne) (Lepidoptera Papilionidae)

Henri Descimon

Laboratoire de Systematique evolutive, Universite de Provence, 3 place Victor Hugo,
13331 Marseille Cedex 3, France.

and

Jean-Pierre Vesco

14bis rue Montplaisir, 84600 Valreas, France.

Abstract. A mutant affecting wing pattern has been observed re¬
peatedly and over a large number of years in a population of P. apollo
from the upper Durance basin in France. It is dominant and morpho¬
logically modifies the postcellular region of forewings and the posterior
part of hind wings, inducing a mask-like design in the former and
obliterating the second eyespot in the latter. The frequency of the
mutant in the population was 1 to 2% in the late 1970’s. It has markedly
decreased since.

Introduction

Scores of aberrations have been described in Parnassius apollo.
However, no genetic work has been carried out although breeding of this
species has been practised for some time. The main difficulty is obtaining
mating in captivity. One of us has mastered the problem by hand¬
pairing. The method may permit production of a practically indefinite
number of successive generations and thus genetic experimentation.
The present paper, the first of a series with such experiments, involves a
very spectacular aberration.

Materials and methods

Ova of either field-collected or bred females are obtained by placing females
singly in a plastic-gauze cage of ca 1 liter (this device will be described with more
details in a later paper). Oviposition is induced either by filtered sunshine or by
a 60-100W incandescence bulb placed 20 to 50cm from the cage. In all cases,
overheating must be carefully avoided. Ova are deposited singly or in small
batches upon 1) a cellulose towel placed on the bottom of the cage, 2) foodplant
fragments ( Sedum sp, Sempervivum) which are not necessary to elicit laying, or
3) the cage walls. Regular feeding, once or twice a day, by a honey-water mixture
(1:10) is essential. We have observed that old, almost exhausted wild females
will recommence laying if carefully fed for several days. In all cases, it is

162

J.Res.Lepid.

preferable to allow the females to lay for only a limited period (1-2 hours per day)
and to keep them quiescent in a cool, shady place the rest of the time. Under
these conditions, females can live 3 weeks or more and lay between 100 and 200
ova.

The ova generally diapause and are best refrigerated for at least 2 months at
0-4°C. However, a small portion of ova (1 to 5% in French populations, but much
more in Spanish ones) hatch immediately and may be reared to adults by the end
of summer (this observation implies the possibility of a potential partial second
generation under natural conditions).

For breeding larvae, it is important to maintain a condition of cool or even cold
air while using a heat radiating light: this condition can be satisfied using either
sunshine or artificial light. It is possible to greatly accelerate caterpillar growth
rate by continuous lighting, pupae being obtained within 10 days. Foodplants
are various species of Sedum, according to their availability. S. album , of low
vegetation, is especially convenient for starting young caterpillars, which do not
spin silk and are consequently unable to climb over elevated plants. Some
cultivated species are refused ( e.g . S. acre) and may be toxic. The broods must be
well ventilated, covering with gauze is unadvisable. Glass or plastic pans with
appropriately high walls are convenient, since the walls are impassable barriers
to the non-climbing caterpillars. Palik’s method (1980), using cellophane walls,
is more sensitive to use and can cause trapping of young larvae at the base of the
plastic sheet. The offspring of a mutant female was lost in this way in 1980,
which delayed the completion of the present study until 1984.

Copulation is easily obtained between bred individuals. The butterflies may
pair freely even in a small cage (for instance a 50cm side cube), provided there is
sufficient sunshine. However, hand-pairing affords the most reliable control of
partner choice. We used Clarke and Sheppards’s method (1953). Although
Parnassius are markedly more difficult to pair than Papilio, success is generally
complete when conditions are good and the operator skilled. Key factors are that
males must be excited by sunshine and the females young. Although females
can be kept ready for mating in a refrigerator at 4°C for several days, the
freshest are best. Pairing lasts several hours (the couple is left still for this time
under attenuated light). One male is capable of fecundating at least three
females. In the first mating, a large, well formed sphragis is secreted; in the
second one, this appendage is rudimentary and is absent in the third.

In all cases, however, fecundation is complete. We recall here that the
sphragis is not a “laying pouch” as once stated, but a true “chastity belt”,
precluding further fecundation. The presence of fecund females with no sphragis
or with a rudimentary one in the field is a strong indication that males can
practise several successive matings. It is also possible that a female could be
fecundated at least twice, first by an old male no longer able to secrete a sphragis
and again by a young male. Such an event would be exceptional yet possible to
check by counting the spermatophores present in the bursa copulatrix.

Results

FIELD OBSERVATIONS: For obvious reasons, we will not give the
exact location where mutant individuals have been observed. It gener¬
ally lies in the upper Durance basin, in french southern Alps. The
habitat is a large set of rather smooth, sunny barren slopes, inter¬
mingled with mowed meadows, at ca. 1800m elevation. Few trees are

26(l-4):l-288, 1988

163

present, a condition quite probably due to forest destruction by man.
The substrate is essentially formed by moraines and screes, with some
thalwegs, not very accentuated, and a small stream. Sedum and
Sempervivum are quite abundant and provide food for Parnassius
apollo. The Parnassians themselves are very abundant over a large
area. We carried out a mark-release-recapture study over a precisely
defined small area of the flight locality. This experiment allowed us to
estimate the population flying upon this area to 400-500 individuals
(Napolitano, Cooke and Descimon, in preparation). The total area of the
locality is much larger and extrapolation of the data allows to estimate
the order of magnitude of the population being at least 10,000. By direct
behavioral observation, it was seen that the butterflies move freely from
one point to another over distances of at least one kilometer, as confirmed
by the capture of marked individuals at distances of this order from
their previous marking area. However, inhospitable zones circumscribe
flight areas to some extent. In such inhospitable zones, individuals are
casually seen, but are much scarcer by comparison. Other high density
flight areas exist some kilometers away from the main one, but they are
markedly smaller.

The first aberrant individuals were taken August 9, 1977, with three
taken on one day. Only later on did we realize that, since the aberration
was recurrent, it was probably due to mutation and that its frequency
was worth further investigation. Still later, we discovered in the
correspondence of the senior author a letter from Lucien Jean, who
mentioned the capture of an ’’aberrant apollo” by another collector, Mr.
Dreano, in the same locality. The letter was accompanied with a color
slide which allowed us to verify that the female aberrant collected by
Mr. Dreano belonged to the same type we found. This specimen had been
captured around 1975.

The population has been followed regularly to 1981 and less inten¬
sively since 1982. We attempted to count all individuals seen to obtain a
gross estimation of the mutant frequency (Table I). In most cases, counts
were made by direct sighting, without marking correlation, so results
must be considered approximate. At face value the frequency of the
mutant decreased from ca. 2% to ca. 0.5% in five years. However, the
1985 capture of a normal female which produced mutant offspring
indicated it had been mated by a mutant male and that the gene was
still present in the population at that time.

A substantial fraction of the mutant individuals was secured, in
particular 3 females for laying. Foolishly, we made the faulty assump¬
tion that the mutation was recessive, which would have implyed that
removing the thus supposedly homozygous individuals was not detri¬
mental. This supposition was quite unfortunate, as we will see further.

It is worth noting that, when mutants were observed, they generally
were in a group of 2-5 individuals flying in a restricted “pocket”
surrounded by areas where none was to be found. It seems that this
pocket correspond to the laying area of the mother female. Sometimes,

J.Res.Lepid.

164

Table I. Number and percentages of “Zorro” mutant vs. “normal”
phenotypes of P. apollo.

Year

Numbers of P. apollo observed

Percentage

1977

“normal” phenotype

“Zorro” mutants

1977

ca 600*

12 (8 6, 4 9)

2.0

1978

co 800*

14(12 6,2 9)

1.7

1979

1184**

7 (6 d, 1 9)

0.6

1980

136**

1 9

0.3

1981

ca 300*

0

0.0

1983

ca 200*

1 6

0.5

1985

ca 200*

^ g***

0.5

1987

co 150*

0

0.0

* Individuals counted but not marked; in this case, we applied a
correction coefficient keeping in account multiple captures and deduced
from marking-releasing experiments.

** Individuals marked before being released.

*** Individual not observed, but existence deduced from the offspring of
a “normal” female.

mutants were observed flying slightly more awkwardly than the normal
butterflies and, in some cases, the degree of wing damage indicated they
were less sturdy. Behavior was unaffected, but phenotype modification
was discernible during flight to an experienced eye from a distance.
When at rest in the absence of sunshine, Parnassius display a charac-
teristical protective behavior. They open their wings in a horizontal
plane and reveal their posterior eyespots. This display is accompanied
by a kind of stridulation obtained by brushing the ventral face of
hindwings with posterior legs (Descimon, 1965). The resultant noise
resembles bruising silk and is perceptible to human ear from at least 1
meter. This behavior seems to occur in all species of Parnassius,
including both P. mnemosyne, in which the hindwings do not have red
eyespots, and the very ornate blue and red Himalayan species (F.
Michel, pers. comm.). The mutants also display this behavior, but it’s
effect is entirely different to human observer’s eye: attention is drawn
from the hindwings, where the posterior eyespot is missing, to the
forewings with their striking mask-like design.

BREEDING. In 1983, a male was secured and handpaired with a
virgin female from the Luberon (Vaucluse, France). A 1:1 segregation
appeared in the offspring and, since the mutation had never been
observed among thousands of butterflies in the Luberon population, the
mutation must be dominant. The gene was subsequently introduced
into other stocks, including those from the Mercantour ( Alpes Maritimes,
France) and the Causse du Larzac (Aveyron, France). Further, a new
mutant strain, isolated from the original Durance locality, was recovered

26(l-4):l-288, 1988

165

by chance. As already mentioned above, a normal female was collected
amongst a group of normals. This female yielded mutants in a 1:1 ratio,
which indicated that it had been fecundated by a heterozygous mutant
male. In all mutant x normal crosses a 1:1 segregation appeared (table
II). The only heterozygote x heterozygote cross (table II No. 6) yielded a
high proportion of mutant individuals. This result is puzzling. Indeed,
the other crosses do not depart significantly from a 1:1 ratio, which
precludes considering a superiority of heterozygotes over normal
homozygotes. In No.6 cross, the excess of mutant phenotypes is not only
too high to allow considering the possibility of homozygous mutant
lethality (2:1 ratio, X2 = 9.6, p<0.01), but even to fit with the expected 3:1
ratio (X2 = 5.4, p<0.02). We, however, cannot draw conclusions from a
single cross. Actually, the genetic composition of this brood was fairly
heterogenous, which could give rise to “hybrid breakdown” phenomena
and distort the ratios. In particular, we have indications that the Causse
du Larzac population is peculiar; for instance, its ova are on the average
twice as heavy as those from Alpine populations. The concerned cross
was the only laboratory-bred of a series of identical parentage which
was used to make an experiment of founding an artificial population of
P. apollo on the Sainte Baume mountain, where it is absent. Now, this
experiment, which involved the deposition of 1,000 ova upon favorably

Table 2. Results of crosses with “Zorro” mutant of Parnassius apollo.

Number of
the cross

1

Parents

8 : “Zorro” from “Zorro“ 9 x
“normal” <5 from Mercantour
9 : wild, normal, Briangon

Offspring

“normal”

‘Zorro’

21 24

(9 d, 12 9) (9 d, 15 9)

8 : wild, “normal”, Causse du
Larzac

9 : “Zorro” from “Zorro” 9 x
wild “normal” 8 , Luberon

16 17

(7 d, 9 9) (10 d, 7 9)

8 : “Zorro” from a “normal” 9
caught in the wild
9 : “normal” from all-normal F x
from the original locality

40 39

(14 d, 16 9) (19 d, 10 9)

8 : wild, “normal”, Mercantour

4 9 ; “Zorro” from 3

5 8 : wild “normal”, Luberon
9 : “Zorro” from 3

12

8 : “Zorro” from 2
9 : “Zorro” from 2

28

166

J.Res.Lepid.

sited foodplants, has been unsuccessful. Moreover, the mortality in the
brood was high (only 30 adults from 110 ova), in spite of having been
pampered. More experiments are needed but, unfortunately, they are at
present impossible.

MUTANT GENE EXPRESSION AND VARIATION. The average
mutant phenotype is represented on figure 1 (A: male, B: female). On the
forewings, it is the region posterior to vein 7 which is modified: the discal
spot is elongated distally into a point following vein 7. Basally, this spot
shows a tendency to be connected to the median (discocellular) spot of
the cell by two black streaks following the anterior and posterior limits
of the cell. The marking gives mutant butterflies a striking aspect, as if
they bear a black mask. The postdiscal row of spots is shifted distally
and partly obliterated in 5-6 and 6-7 intervals. No obvious modification
is to be noticed in 2-3 and 3-4 intervals, but the spot of the lb-2 interval,
very characteristic of the species, is conspicuously shrunken and divided
into two parts by lc rudimentary vein. On the hind wings, the anterior
eyespot is absolutely unaffected, while the posterior one, which lies in

2 cm

Fig. 1. “Zo/ro” mutation of Parnassius apollo.

A - male mutant, wild collected, 2 VIII 78.

B - female mutant, wild collected, 14 VIII 78.

C - male, normal phenotype, same locality, 28 VII 79.
D - female, normal phenotype, same locality, 6 VIII 78.

26(l-4):l-288, 1988

167

3-4 and 4-5 intervals, is extremely modified. It is dissociated into two
parts, as if a factor following vein 4 inhibited the formation of the eye-
spot pattern. The series of anal spots in lb, lb-2 and 2-3 is diminished
but still present. Premarginal black scales lunules and marginal
hyaline band are slightly displaced basally.

It is rather obvious that such a remarkable mutation deserves a
name. Here we meet with a problem which has not been considered
seriously for Lepidoptera. In species such as Drosophila melanogaster,
which has not been plagued by a crowd of aberrational names given by
mere “curios” collectors, matters are clear. Mutational names are
indispensalbe working tools, with simple and clear terminological
rules. In all animals, the practice of giving latin names to any infrasub¬
specific form has been rejected by the International Commission of
Zoological Nomenclature. We believe that this does not preclude using
non-linnean names which would follow the rules of genetic nomenclature
in the case of variants which have in fact been studied genetically. The
controversial situation arises in the case of “classically named aberra¬
tions” or morphs which later prove to be mutants. At first sight, it seems
advisable to retain the old names. However, some cases would be quite
puzzling; for instance, for the white female of Colias croceus , should we
use the old name of “Helice” given by Hiibner or, following the American
authors, the generic name of “AZ6a”, which assume that all
white female mutants are homologous? In any event, we propose here
the mutational name of “Zorro” for the described variant of P. apollo.
Names that we would have preferred, such as mephisto or diaholicus ,
have been already given to Parnassius variants or subspecies; otherwise,
the selected name will recall that nomenclature need not be such a
serious topic, after all. . .

Field-collected, as well as bred examples of “Zorro” display variability.
We noted above this is not due to incomplete dominance. The argument
is reinforced by the fact that the probability of occurrence of homozygous
mutant individuals in natural populations is very low (practically equal
to variant frequency, that is 2 to 0.5 percent). Moreover, when variation
is observed within bred individuals, it must be due mainly to interaction
of the mutant gene with its genetic background, since rearing conditions
were kept relatively constant. Fig. 2 shows some of this variability;
individual 1 is among the least accentuated and 2 among the most.
There is also an interaction with sexual dimorphism. If venation is not
markedly modified on forewings, some striking abnormalities may be
observed on hindwings. In all cases, a rudimentary cell is present at the
outer extremity of the cell between Ml and M2 normal veins, but much
more spectacular modifications can also occur: the cell is open between
Ml and M2; Ml is often branched and in some cases a complex system of
supplementary cells is formed. These modifications are often assy me¬
trical. Figure 3 gives an idea of these atypical vein patterns.

Fig. 2. Variation in the expression of “Zorro” mutation in Parnassius apollo.

1 - male, wild, 13 VIII 77.

2 - D°, D°, 27 VII 78.

3 - D°, D°, 14 VIII 78.

4 - D°, D°, 28 VII 79.

5 - Female, D°, 2 VIII 78.

6 - D°, D°, 10 VIII 77.

7 - D°, bred (Brood Number 1, see table II).

8 - D°, D° (Brood Number 2)

Fig. 3. Abnormalities of venation in “Zorro” mutation of P. apollo.

1 . Normal female.

2. "Zorro" female (n° 6 of fig. 2), with supplementary distal vein.

3. D°, brood n° 6: supplementary distal vein plus intracellular
rudimentary vein.

4. D°, male, brood n° 6: D°, with different branching.

5. D°, female, brood 5: supplementary cell with atrophy of normal cell
closure.

6. D°, female, brood 5: supplementary distal cell.

All figures represent right hindwing in the region where median veins (Ml , M2, M3)
take rise from the cell, except n° 6, where it is left hindwing which has been
photographed.

170

J.Res.Lepid.

Discussion

From a morphogenetic point of view, “Zorro” is one of the most
striking aberrations in Parnassius. A thorough survey of the previous
described forms in available literature ( e . g. Bryk, 1935, Eisner, 1966)
did not reveal any close equivalent. It may be remarked that, in both
pairs of wings, the mutation modifies only the posterior half, while the
anterior one remains unaffected. Pattern and venation are most per-
turbated at the suture between the imaginal disk compartments which,
according to the studies of Sibatani (1981), follows the axis of symmetry
of the cell and the corresponding distal part of the wing. It is obvious
that “Zorro” could provide a choice tool for the study of the development
of wing pattern, using for example the methodology of Nijhout (1985).
Unfortunately, it is probable that practical difficulties would render
such a study rather difficult.

From the point of view of evolutionary genetics, the history of this
mutation appears clear. It arose once, at one locus of one individual of
the population. It is more difficult to understand how its frequency
reached 1 or 2 percent. If we assume the size of the population,
previously estimated to roughly 10,000, the original frequency must be
around 0.5 xl0~4. To reach thelO-2 frequency observed in 1977, the
“Zorro” allele therefore must have been multiplied by 200. The simplest
explanation would be to assume the population had been reduced to few
individuals in at least one year, the mutation having been preserved by
chance (or having appeared) during the time of the population con¬
traction. Further, its frequency would have been amplified in parallel to
the population increase. We can provide some observational support to
this hypothesis: the senior author and his brother Robert Descimon
have collected and observed butterflies very regularly in the region
until the present time and P. apollo was noted as very scarce at the end
of the 1960’s. It is further noteworthy that the “Zorro” mutation was not
observed in the locality where it was later discovered. Many P. apollo
were seen during early 1960’s.

Could selection have played a role in the variation of the frequency of
the mutation? From 1977 to 1981 “Zorro” has obviously decreased. It is
very unfortunate that we did not surmise that this aberration could be a
dominant but postulated it was a recessive, with a gene frequency of
about 0,14, providing the observed “homozygote” frequency of 2 percent.
Under these conditions, we incorrectly believed that securing and
killing some individuals was not detrimental. Actually, we introduced a
massive selection coefficient, “destroying our own subject of study”,
according to the accurate expression of Dubois (1983). Fortunately, only
a portion of the population was screened and the mutation was not
eradicated in totality. The most distressing consequence of this thought¬
less action is that we are now hindered from drawing conclusions.
Would “Zorro” frequency have decreased anyway? It has been clearly
observed that the mutants appear a trifle handicapped in flight activity.

26(l-4):l-288, 1988

171

At the larval stage, however, no disadvantage appears to exist; in one
brood, an excess of mutants has been observed. The experiments should
be repeated with a larger sample. The modification of wing pattern does
not obviously impair its deterrent effect. To human eye, it is even more
frightening! Therefore, we may not rule out the hypothesis that the
mutation was slowly increasing its frequency when we clumsily
intervened.

Although rather unusual, the dominance of “Zorro” seems to be best
interpreted in terms of physiological genetics. The observed variation in
expression does not seem related to overdominance but to the inter¬
ference with the entire genotype. It would not be relevant to hypothesize
that the mutation should have become dominant after a process of
“evolution of dominance” (Ford and Sheppard, 1966), since none of the
conditions for it appear here.

We have planned to “repair our fault” by breeding and releasing
mutant individuals (with, of course, no mixture with foreign strains)
into the locality. To do so, we would introduce yet another perturbation
into the population. The best would be to proceed but with seriously
controlled and monitored conditions. Only accidental difficulties have
delayed this operation. We also intend to undertake again experiments
of creating artificial populations like the one previously mentioned,
which has been probably unsuccessful. Such attempts (which are
debatable if not carefully designed) have proven successful and quite
instructive in some cases (Descimon, 1976, Holdren and Ehrlich, 1981).

Cases of decreasing mutation frequency in natural populations by
collecting are already known. The most striking is probably the
“honnoratii” form of Zerynthia rumina in the region of Digne
(Alpes de Haute Provence, France). The problem, which has elicited
some row in the local press (with ridiculously exaggerated considera¬
tions, especially about the prices fetched on butterfly market) led to the
promulgation of a law forbidding all insect collecting in the concerned
department. Notwithstanding the inadequacy of enforcing the law, it is
almost certain that it has improved the chance of maintaining the
mutant gene. Z. rumina “ honnoratii ” has been observed in recent years
(P. Bonnet, pers. comm.).

We believe that, for “Zorro” as for “honnoratii” , the best protection is
to breed and distribute them to collectors, who would be deterred from
painstakingly seeking for them in nature. Further, lowering the venal
value would render the “black market” less likely. One would pass from
“hunting and gathering” to “farming”. We strongly suggest “desperate
hunters” not only search for new aberrations, but breed them, obtaining
at the same time not only fine collection items, but genetic information.
Such a practice has been frequent for some time in England - it must be
generalized.

Acknowledgements. We thank Rudi Mattoni for having kindly helped us edit the
present work.

172

J. Res. Lepid.

Literature Cited

BRYK, F., 1935. Lepidoptera Parnassiidae pars ii (subfam. Parnassiinae).
Tierreich 65: i-li, 1-788, 698 figs.

CLARKE, C.A., 1952. Hand pairing of Papilio machaon in February. Entomol. Rec.
J. Var. 64: 98-100.

DESCIMON, H., 1965. Quelques comportements protecteurs des Lepidopteres.
Alexanor IV: 61-66.

- , 1976. L’acclimatation des Lepidopteres: un essai d’experimentation

en Biogeographie. Alexanor IX: 195-204.

DUBOIS, A., 1983. Renforcements de populations et pollution genetique. C. R. Soc.
Biogeogr. 59: 285-294.

FORD, E.B. & P.M. SHEPPARD, 1966. Natural selection and the evolution of domin¬
ance. Heredity 21: 139-147.

EISNER, C., 1966. Parnassidae-Typen in der Sammlung J.C. Eisner. Zool.
Verhand. 81: 1-190, PI. 1-84.

HOLDREN, C.E. & P R. EHRLICH, 1981. Long range dispersal in checkerspot butter¬
flies: transplant experiments with Euphydryas gillettii. Oecologia (Berlin)
50: 125-129.

NIJHOUT, F., 1985. The developmental Physiology of color patterns in Lepidoptera.
Adv. Insect Physiol., 18: 181-247.

PALIK, E., 1980. The protection and reintroduction in Poland of Parnassius apollo
Linnaeus (Papilionidae). Nota lepid. 2: 163-164.

SIBATANI, A., 1983. A Compilation of Data on Wing Homeosis in Lepidoptera. J.
Res. Lepid. 22: 1-46.

Journal of Research on the Lepidoptera

26(1-4):173-176, 1988

Mimicry by illusion in a sexually dimorphic, day-flying
moth, Dysschema jansonis (Lepidoptera: Arctiidae:
Pericopinae).

Annette Aiello

Smithsonian Tropical Research Institute, Box 2072 Balboa, Ancon, R. Panama
and

Keith S. Brown, Jr.

Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas,
C. P. 6109, Campinas, Sao Paulo 13.081 Brazil

Abstract. Sexual dimorphism and development in Dysschema jansonis
(Butler, 1870) are discussed, and wing patterns of females and males
are compared with those of their models, female Parides spp. (Papilion-
idae) and transparent-winged ithomiine nymphalids, respectively.

Two questions are addressed: Why does D. jansonis not display
more accurate mimicry? How does D. jansonis create the illusion of
mimicry?

Introduction

Species of the genus Dysschema Hiibner, 1818 are well-known
examples of diurnalism in moths (Klots & Klots, 1959), and sexual
dimorphism of wing pattern and color in Lepidoptera (Watson &
Whalley, 1975). They are presumed mimics of unpalatable butterflies of
various groups (Fisher, 1958; Gilbert, 1984; Watson & Whalley, 1975),
and it is possible that they themselves are unpalatable, Mullerian
mimics. The following remarks concern D. jansonis in the Republic of
Panama.

Sexual Dimorphism

In general appearance, males and females of D. jansonis are strik¬
ingly different (Figure 1). However, a careful inspection reveals that the
same wing pattern elements are present in both sexes. The dorsal
forewings of the two sexes are quite similar, except that the pattern
shows slightly more contrast in males. The hind wing pink forms a large
blotch at the posterior margin in females, but is reduced to a tiny dot or
is absent in males. The dorsal yellow, restricted to a round mark near
the anterior margin of the hind wing in females, occupies the basal three
fourths of the wing in males. In both sexes, the ventral side closely
resembles the dorsal side except that in the male the forewing markings
are more pronounced on the ventral surface than on the dorsal, and in

174

J. Res. Lepid.

Fig. 1 . Left column, from top: Dysschema jansonis males (D, YCF), (D, WCF),
Oleria rubescens (D), D. jansonis male (V, WCF); Middle column, from
top: Eurytides ilus (D), Papilio anchisiades (D); Right column, from top: D.
jansonis females (D), (V), Parides areas female (D).

Figure codes: D = dorsal, V = ventral, YCF = yellow color form, WCF =
white color form.

the female the ventral forewing bears a yellow band that is only faintly
evident on the dorsal side.

Sexual Differences During Development

Five males and six females of D. jansonis were reared from a clutch
of 13 eggs laid by a single female, 10. XI. 1982 during daylight hours,
on the underside of a leaf of Spiracantha cornifolia (Compositae) at
the Summit Observatory in the Canal area of the Republic of
Panama (Aiello Lot 82-62). All 13 eggs hatched on 22. XI. 1982. One
larva died in its first stadium, another died as a late instar, but the other
11 completed development to adult.

Females passed through eight larval stadia and required 72-80
days (x = 75.83 days, s = 3.19) to complete development from egg to
adult, whereas males had only seven larval stadia dnd completed
development in 66-69 days (x — 67.60 days; s = 1.14). The late larva that
died, did so on day 69 as a ninth instar. One can only guess at the sex of
that individual; possibly it was an exceedingly small male who had
difficulty achieving the minimum weight necessary for pupation.

Early stadia were synchronized; all 13 eggs hatched on the same
day, and molting occurred synchronously through stadium six. Larvae
in the first through third stadia aggregated, and fed by scraping the
upper leaf surface either while strung out along one edge of a leaf, or
while lined up parallel to one another to form a feeding front. Fourth

26(1-4): 1-288, 1988

175

instars began to show conspicuous size variation, and from then on were
solitary and ate whole leaf instead of scraping the surface. Molts to
seventh and eighth stadia were asynchronous; the molt to stadium
seven occurred over two days, and that to stadium eight took place over
four days. Following the molt to stadium seven, all larvae were
transferred to separate cages and given individual numbers. Six indivi¬
duals that were larger than the others were numbered 1-6, and all
proved to be females; the smaller larvae (numbered 7-11) turned out to
be males.

Adult eclosions in both sexes took place between 1500 and 1700 hours.

Mimicry in Females

Dysschema jansonis females resemble females of the aposematic
butterflies Parides spp. (Papilionidae), when in flight. Like Parides
(Figure 1), they have yellow and pink markings against a black
background. However, in Parides the yellow is on the forewings and the
pink is on the hind wings, while in D. jansonis both color elements are
located on the hind wings. This seemingly important discrepancy does
not detract from the mimetic effect when the moth is in flight; the
human observer, at least, sees the rapidly fluttering moth as a fast-
moving Parides. The illusion is made possible partly by exposure of the
yellow band, on the underside of the forewing, during the high wing-
stroke flight of this moth, and partly by the fact that when the wings are
spread they are somewhat translucent, and the underside yellow shows
through them from above.

There exist a number of other Parides look-alikes with much more
convincing mimicry. In Panama, Eury tides ilus and Papilio anchisiades
(Papilionidae) (Figure 1) have the yellow on the forewings and the pink
on the hind wings as does Parides. A probable explanation for
D. jansonis’ s “imperfect” mimicry of Parides can be found in the resting
posture of this moth. Both sexes rest, with the fore wings covering the
hind wings, on foliage or bark where they blend with their background
or resemble a bit of dead leaf. A yellow band or large white area on
the dorsal forewing might interfere with this resting-posture crypsis.
D. jansonis has overcome this constraint by employing mimetic illusion,
and as a result possesses two distinct modes of visual protection.

Another example of mimicry by illusion can be found in the nymphalid
butterfly Consul fabius , which resembles a dead leaf when at rest with
its ventral cryptic pattern showing, and a tiger-striped Heliconius when
in flight with the dorsal pattern exposed. The illusion occurs when
C. fabius flies in sunlight and the dorsal mimetic pattern shows through
the wings from below as well (DeVries, 1987).

Mimicry in Males

Dysschema jansonis males resemble various genera of transparent¬
winged ithomiine nymphalid butterflies (Figure 1). Although scaled,

176

J.Res.Lepid.

the wings of D. jansonis males appear transparent when in flight, an
illusion made possible by the fact that dorsal and ventral surfaces are
identically patterned, and enhanced by exposure of the strongly marked
ventral forewing during flight. In addition, when the wings are spread,
the paler areas are translucent, a fact which can be demonstrated by
holding a pinned specimen to the light. Males’ fluttery flight further
contributes to the ithomiine resemblance.

When at rest, with the forewings covering the hind wings, transmitted
light and wing pattern contrast are reduced, and the moths blend well
with mottled bark or dried leaves. Reduced contrast can be demonstrated
by placing a pinned specimen over a dark background.

Male Color Dimorphism

Some males have white, instead of yellow, hind wings, and while it
has been supposed (Druce, 1884; Hering, 1925) that the two forms are
mere color variations of the same species, no evidence has been
presented until now. Of the five males reared from the single clutch of
13 eggs, three were the white color morph and two were the yellow.
Presumably both are ithomiine mimics; transparent-winged ithomiines
include species with clear, and species with yellow-tinged wings.

Acknowledgements. We are grateful to Gordon B. Small who observed the
female D. jansonis during oviposition and collected her egg clutch, to Phil
DeVries who commented on the manuscript, to Diomedes Quintero and the
University of Panama for loan of the Oleria specimen, and to Carl Hansen who
photographed the adult specimens.

Literature Cited

DEVRIES, P. J., 1987. The Butterflies of Costa Rica and Their Natural History.

Princeton University Press. Princeton, New Jersey, xxii -I- 327.

DRUCE, H., 1884. Pericopidae pp. 105-112, In Biologia Centrali- Americana.

Insecta. Lepidoptera-Heterocera I. 490 pp.

FISHER, R. A., 1958. The Genetical Theory of Natural Selection. Dover. New York,
xiv -1- 291 pp.

GILBERT, L. E., 1984. The Biology of Butterfly Communities, pp. 41-54, In R. I.
Vane-Wright and P. R. Ackery (eds.), The Biology of Butterflies. Symposium
of the Royal Entomological Society of London, No. 11. Academic Press.
London, xxiv + 429 pp.

HERING, M., 1925. Arctiidae: Pericopinae pp. 425-455, In A. Seitz, editor,
The Macrolepidoptera of the World. II Division: The Macrolepidoptera of
the American Region. Volume 6. The American Bombyces and Sphinges.
1304 pp.

KLOTS, A. B. & E. B. KLOTS, 1959. Living Insects of the World. Hamish Hamilton.
London. 304 pp.

WATSON, A. & P. E. S. WHALLEY, 1975. The Dictionary of Butterflies and Moths
in Color. Exeter Books. New York, xiv -I- 296 pp.

Journal of Research on theLepidoptera

26(1-4):177-186, 1988

“Black-Light” Induction of Photoperiod-controlled
Diapause Responses of the Viceroy Butterfly, Limenitis
archippus (Nymphalidae)

A. P. Platt
and

S. J. Harrison1

Department of Biological Sciences, University of Maryland Baltimore County, 5401
Wilkens Avenue, Catonsville, Maryland 21228

Abstract. The nearctic viceroy butterfly, Limenitis archippus
(Cramer), is a typical long-day insect. The 2nd and early 3rd instars are
photosensitive. Half-grown larvae respond to short-day (autumn)
photoperiod by constructing hibernacula and entering diapause. Long-
day photoperiod induces rapid growth and direct development.
Previous experiments testing this facultative response have used
fluorescent bulbs emitting both UV and visible light (320-700 nm).
However, both diapause and direct development can be induced in
larvae exposed to “black-light” photoperiod regimes containing mainly
the near UV (violet) and UV portions of the spectrum (320-436 nm).

The stemmata and dorsolateral abdominal saddlepatch areas of the
2nd and 3rd instar larvae represent possible photoreceptors for mediat¬
ing these responses.

Introduction

Many temperate insects respond to changes in daylength through
hormonally regulated physiological mechanisms influencing direct
development or diapause (Lees, 1955, 1960; Danilevskii, 1965; Andre-
wartha & Birch, 1973, Saunders, 1976, 1977; Tauber & Tauber, 1973,
1976; Beck, 1980). Typical long-day insects exhibit diapause when
exposed to short daylength (< 11 hr light per 24 hr day) but show direct
development under longer photoperiods. Such environmentally induced
responses are termed facultative. The photosensitive stages vary
between species, and may involve eggs, larvae, pupae, or adults (Wig-
glesworth, 1970, 1972). Such responses exhibit variability within and
between broods in a single species. Temperate insect strains from
different latitudes or altitudes, exhibit similar variation. (Tauber &
Tauber, 1972; Beck, 1980; Ujiye, 1985). Diapause is related to insect
endocrine control and metabolism (Gilbert et al, 1960; Harvey, 1962; de

'Present address: EA Engineering, Science and Technology, Inc. (formerly Ecological
Analysts, Inc.) 15 Loveton Circle, Sparks, MD 21152

178

J.Res.Lepid.

Wilde, 1965; Harvey and Haskell, 1966), and is under polygenic control
in most species. Recently, a dual system involving photoperiod and
temperature for regulating circadian rhythms and diapause has been
proposed (Beck, 1977; Neumann & Kruger, 1985).

Facultative diapause in the nymphaline butterfly Limenitis ( Basil -
archia) archippus (Cramer) has been studied by Clark & Platt (1969),
Hong & Platt (1975), Frankos & Platt (1976), and Williams & Platt
(1987). This species enters larval diapause when exposed to short-day
photoregimes at room temperature. Second and early 3rd instar larvae
respond to photoperiod over a seven to ten day period. Larvae preparing
to diapause grow more slowly than their long-day siblings. Those reared
in either continuous light or in complete darkness exhibit nearly 100%
mortality (Platt, pers. obs.). Diapause initiation and termination take
place while development is arrested in the 3rd instar. Thus, the
diapause responses of Limenitis are not complicated by developmental
processes associated with metamorphosis, as they are in other species
which diapause as pupae.

Complex innate behavior precedes larval diapause and can be con¬
veniently studied in short-day larvae. This behavior is genetically
based (Clark & Platt, 1969; Hong & Platt, 1975), but is expressed only
under short-day conditions. Short-day larvae grow slowly and construct
hibernacula (overwintering chambers) from the basal portions of poplar
(. Populus spp .) and willow ( Salix spp.) leaves ( Salicaceae ) on which they
feed. These tubular structures are made by chewing the leaf in a
characteristic manner, and then covering the remaining basal leaf
surfaces with silk to form an enclosed curving tube which remains open
at its outer (distal) end. The hibernaculum is attached to the foodplant
twig with a substantial silk girdle formed around the leaf stem and
surrounding next season’s bud (Edwards, 1884; Scudder, 1889; Weed,
1926; Klots, 1951). The hibernaculum remains attached to the plant
throughout the winter months following leaf drop. Short-day larvae
crawl into these chambers prior to diapausing. Physiological changes
such as water loss and glycerol accumulation accompany diapause
(Frankos & Platt, 1976). Diapause onset can be reversed by switching
short-day larvae to long-day conditions (LD 16:8), even after hiberna¬
culum formation and entry have occurred. However, the reciprocal
transfer of 2nd and 3rd instar larvae (from long-day to short-day
photoregimes) usually results in larval death (Clark & Platt, 1969).

Few papers have addressed the wavelengths (colors) of light which
influence insect diapause. Beck (1980) states that the most effective
wavelengths are between 400-550 nm for most species. Biinning and
Joerrens (1960) found that blue light induced diapause in Pieris

2Larvae were not reared in photochambers containing the insect “black-light” bulbs,
because plants placed in these chambers wilted within 12 hrs, and their leaves dried up.
This evidently resulted from the light quality emitted by these particular bulbs.

26(l-4):l-288, 1988

179

brassicae L. (Pieridae) during early photophase, but that red light
promoted diapause later in the diel photoperiod cycle. However, Beck
(1980) criticized their conclusions, which he believed could be explained
more simply by assuming that the red light regime was equivalent to
total darkness in their experiments. The spectral limits for diapause
induction in Limenitis have not been previously determined.

In this paper we show that Limenitis larvae exhibit normal diapause
reponses when grown in photochambers containing fluorescent “black-
light” bulbs emitting a partial spectrum of violet-blue and ultraviolet
(UV) wavelengths. UV light is known to be an important spectral
component of many insects, especially Lepidoptera (Silbergleid, 1979).
Clark & Platt (1969) suggested that the dorsolateral grey- white abdo¬
minal saddlepatch of young larvae may represent a photosensitive
region. This pale saddlepatch makes the small larvae cryptically
patterned, and helps them resemble bird droppings when at rest in
curved positions on the leaves and twigs of their foodplants. The
stemmata (ocelli) of larvae also are known to possess UV receptors
(Ichikawa & Tatda, 1982).

Materials and methods

All larvae were reared from eggs in light-tight wooden photochambers (61 x
61 x 42 cm inside dimensions) at room temperature (25 ± 2°C). The inside wahs
of the chambers were painted a non-fluorescent flat white. Experimental
chambers contained single 15" G. E. fluorescent “black-light” bulbs emitting a
partial spectrum between 320-436 nm [so-called “poster lights” Fig. 1A)]. The
quahty of hght emitted by these bulbs is a deep, dim violet. The “poster light”
bulbs are dark purple in color, but the wavelengths emitted are not the same as
those of “black-lights” used commonly to attract and trap insects (Fig. IB)2.
Control chambers contained G. E. cool white fluorescent bulbs identical in size
and wattage, which emitted wavelengths throughout the visible spectrum and
some UV as weh (Fig. 1C).

The emission spectra were measured as follows: Light from each type of source
passed through an Oriel computer-driven monochromator to a photomulti¬
plier (PM) tube (Hamamatsu R-928) through an evacuated quartz window. The
lamp’s output was measured at each nm from 300-600 nm by averaging the
digitized photocurrent of the PM tube (200 measurements per nm), without
correction for the spectral response curve of the PM tube. These data were
subsequently normalized to the peak value. The different emission spectra
obtained from the three kinds of fluorescent tubes are shown for comparison in
Fig. 1.

The larvae were reared on rooted cuttings of weeping willow ( Salix babylonica
L.) in moist conditions using long-day (LD 16:8) and short-day (LD 8:16)
photoperiods. The onset of photophase occurred at 8:00 a.m. EST in all
chambers. A total of 81 larvae was reared during the experiments. *

Individual larvae were taken from stock cultures maintained in the labora¬
tory, or were collected as diapausing larvae in hibemacula. Eggs from three L.
archippus females, which had been hand-paired using Platt’s (1969) method,
were used in the induction experiments (Table 1). Eggs from each brood were

180

J.Res.Lepid.

divided evenly between the control and experimental conditions. Diapause
termination studies were done using overwintering L. archippus larvae obtained
by collecting hibernacula from the vicinity of Conowingo Dam, Cecil Co.,
Maryland (Table 2). Some wild-collected larvae had been parasitized by wasps,
as noted in the table.

Photochambers were checked daily to determine the condition of the eggs,
developing larvae, and the foodplants. Plants and larvae were misted with
water each time they were examined. The laboratory in which the photo¬
chambers were kept was light-tight, and was totally darkened before the
chambers themselves were opened. Under these conditions the “black-lights”
emitted so little visible light that it was difficult to find the individual eggs and
larvae in the experimental chambers.

Results

Results of the diapause induction experiments are shown in Table 1 .
The experimental “black-light” and control white light chambers gave
comparable overall results. All surviving larvae on long-day underwent
direct development whereas the majority of short-day larvae entered
diapause. Only four short-day larvae showed direct development under
“black-light”. None of the short-day insects reared in white light
developed directly. Mortality was minimal (11.5%) for the early larval
instars and was evenly distributed among the groups.

Chi square analyses of the Table 1 data show no significant differences
exist between the “black-light” and control regimes when comparing
the numbers of larvae which died, developed directly, or those which

2

diapaused. For short-day, a 2 x 3 contingency test yielded X x — =

2

4.43, with p > 0.10. For long-day, a 2 x 2 analysis gave S x =

0.31, with p x* 0.05. Thus, the null hypothesis that the larval responses
are the same in both the experimental and control photoregimes is
upheld in both cases.

All of the diapause termination experiments (Table 2) were carried
out using long-day photoperiods. Resumption of larval activity following
diapause is independent of photoperiod in L. archippus. Repeated
observations by Platt (unpubl.) show that diapause termination can
take place either in long-day or short-day photoperiods. Our present
experiments show that diapause termination occurs equally well in

3In a preliminary experiment to determine whether larvae would survive in the “black-
light” photochambers, four eggs of the banded purple butterfly, Limenitis arthemis
(Drury), from Vermont were placed on a willow plant in experimental (“black-light”)
conditions. One larva soon disappeared, and could not be found in the dim light. The other
three grew well, underwent normal metamorphosis, and produced perfect adults. Both L.
arthemis and L. archippus exhibit similar facultative diapause responses (Greenfield &
Platt, 1974). The availability of strains was the sole criterion for choosing which species to
use in the main experiments.

100

80

60

40

20

0

3

100

80

60

40

20

0

3<

100

80

60

40

20

0

3(

. Cc

Wc

Wc

36

A. POSTER "BLACK-LIGHT"

z>

_i

CD

<0

ro

1

»0

500

600

B. INSECT "BLACK-LIGHT

LlI

>

I

<T>
IT) ID

A-

400

500

600

C. COOL WHITE LIGHT

D

WAVELENGTH (nm)

nparative emission spectra of G. E. flourescent bulbs (No. F15T8, 15
:). The poster “black-light" (A) emits a partial spectrum with most
elengths within the UV range (between 313-400 nm), and peaking at
nm. Additional visible lines occur at 405 nm and 436 nm, producing a
, deep violet color. Both the insect “black-light" (B — shown for
iparative purposes only), and the cool white tube (C) have additional
ssion peaks at 546 nm and between 577-579 nm as well. These spectra
q measured for us by T. W. Cronin of UMBC using a microspectro-
tometer as described in the methods section.

182

J.Res.Lepid.

Table 1. Diapause induction in Maryland L. archippus larvae sub¬
jected to “black-light” (experimental) and white light
(control) photoperiods.

Photophase

Short-day (LD 8:16) Long-day (LD 16:8)

“Black-light” White light “Black-light” White light

Initial

No. of eggs

21

20

10

10

No. & %

dying

1 (4.7%)

2 (10%)

3 (30%)

1 (10%)

No. & % of

survivors

diapausing

16 (80%)

18 (100%)

0 (0%)

0 (0%)

No. & % of

survivors
showing direct
development

4 (20%)

0 (0%)

7 (100%)

9 (100%)

both the “black-light” and white light chambers under long-day condi¬
tions. Emerging larvae in both groups completed development success¬
fully. All mortality encountered in these experiments was attributed to
larval parasitism, which was equal in both test groups. A 2 x 2 chi

2

square test of these data gave 2 x — (Yates) = 0.00, with p > 0.90. All of

the larvae used in these experiments had been wild-collected. Thus,
they had been exposed to ambient (outdoor) photophase until we
collected them in December and January prior to the experiments. The
larvae exhibited no apparent physiological or metabolic difficulties
adjusting to either the artificial photoregimes or other laboratory
conditions to which we subjected them.

Discussion

Insects in general, and Lepidoptera in particular possess visual
sensitivity which peaks in the UV range between 320-400 nm (Lutz,
1933; Goldsmith, 1961; Ferris, 1972; Goldsmith & Bernard, 1974;
Platt, et al, 1984). However, Lepidoptera and other insects can perceive
colors in the visible portion of the spectrum as well (Burkhardt, 1964;
Ichikawa & Tateda, 1982). Insect perception of infrared wavelengths

26(1-4): 1-288, 1988

183

Table 2. Diapause termination in over- wintering L. archippus larvae
collected in hibernacula near Conowingo Dam (Cecil Co.)
MD. Larvae were reared under long-day (16L:8D) photo¬
period at room temperature with high moisture in the
chambers.

Black light

White light

No. of hibernacula with
live larvae

10

10

No. & % dying
(all parasitized)1

4 (40%)

3 (30%)

No. & % of survivors
maturing to adults

6 (100%)

7 (100%)

1The braconid wasp parasite was Apanteles limenitidis Riley. Parasitized larvae
emerged from their hibernacula (terminated diapause) and fed briefly. Soon
they became inactive and never molted beyond third instar. Wasp maggots
emerged from the dying larvae and spun tiny yellow silk cocoons nearby. Each
larva yielded a single wasp. Adult wasps emerged about one week later. The
parasites obviously regulate their own cyclic development to correspond with
that of their host, since the larvae had been parasitized the previous autumn
prior to entering their hibernacula.

(600 nm and above) has not been well-documented in most species.
Often the individual ommatidia of insect compound eyes are specialized
for receiving a specific spectral range. Individual stemmata (important
larval photoreceptors) also are known to respond to specific wavelengths
in the same manner (Ichikawa & Tateda, 1982). However, these
stemmata (larval ocelli) are not believed to be functional in diapause
responses (Tanaka 1950, a, b, c, 1951, a, b).

Insects can respond to low intensities of light. Chapman (1969) states
that intensities above 1.0 foot-candle (f.c.) are sufficient to induce photo¬
responses. Above this level, light intensity is not important in insect
photoreception. In both our experimental and control photochambers
the light intensities were well above these minimal levels. Earlier
studies on L. archippus larvae done in our laboratory revealed that
white light intensities as low as 0.10 f.c. were sufficient for inducing
either larval diapause or direct development, although adults from
larvae reared under such extreme conditions often possessed malformed
wings (Frankos & Platt, pers. obs; Platt, 1984).

Our experiments demonstrate that exposure to short-day “black-
light” photophase regimes induces facultative larval diapause in L.
archippus. Photoregimes consisting of the blue-violet and UV portions

184

J.Res.Lepid.

of the spectrum will either induce and terminate diapause, or produce
direct development, just as well as those consisting of white light do.
Only four of 20 larvae in one short-day “black-light” group failed to
diapause, but even here 80% of the larvae made hibernacula and
entered diapause as did all under white light. The failure of all larvae to
enter diapause is not unexpected for such a complicated life history trait
which is under polygenic control (Hong & Platt, 1975; Williams & Platt,
1987). Failure to diapause occurs in natural populations of Limenitis
spp. in the late summer and autumn. As the ambient photoperiod
decreases, a few prediapause larvae continue to show direct develop¬
ment late in the season, when most others are diapausing (Weed, 1926;
Greenfield & Platt, 1974). The “black-light” portion of the spectrum
may not be the only portion to which these larvae respond. This portion
may not even be necessary for diapause induction, but it is sufficient for
eliciting the same responses that occur in white light. Clark & Platt
(1969) suggested that the grey- white saddle patch of 2nd and 3rd stage
Limenitis larvae may represent a photosensitive region. Future experi¬
ments are planned to investigate the possibility that the saddle patch is
photosensitive, as well as a cryptic pattern.

Acknowledgements. We thank C. F. Stroup, Martin Marietta Enviromental
Systems, and T. W. Cronin, F. E. Hanson, and P. G. Sokolove of UMBC, and P. F.
Brussard of Montana State University for helpful comments on our manuscript.
T. W. Cronin also measured the emission spectra of the three fluorescent bulbs
for us, and Fran Baldwin and G. C. Ford, Graphics Illustrators, UMBC made the
figures. T. F. Williams of UMAB provided help with the statistical analyses.

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Journal ofResearch on the Lepidoptera

26(l-4):187-200, 1988

Suppression of the Black Phenotype in Females of the
P.glaucus Group (Papilionidae)

David A. West

Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg,
VA 24061 USA

and

Sir Cyril A. Clarke

Department of Genetics, University of Liverpool, Liverpool, L69 3BX, England

Abstract. Crosses between Papilio g.glaucus and P.eurymedon pro¬
duced fertile offspring of both sexes, and backcrosses to P. g.glaucus
revealed an autosomal suppressor in P. eurymedon which prevents
expression of the black female phenotype of P. g. glaucus. Similar
suppressors are demonstrated in P. rutulus, P. multicaudatus and
P. g. canadensis.

Introduction

It is well known that in Papilio glaucus Linnaeus 1764, black females
usually produce black daughters and yellow females yellow ones,
irrespective of the provenance of the male. Occasional exceptions to this
rule have been reported over many years and various explanations
given (Clarke and Sheppard, 1959; Clarke et al., 1976). The present paper
highlights the importance of autosomal genes which suppress, to a
greater or less extent, the black female wing pattern. This can occur in
the wild or be laboratory produced, and the phenomenon occurs in
various species of the group.

In Section I we report a new example, bred by one of us (D.A.W.) in
hybrids between black glaucus and Papilio eurymedon Lucas 1852.

Section II deals with further examples of suppression of black by
P. rutulus Lucas 1852, P. glaucus canadensis Rothschild & Jordan 1906
and probably by Papilio multicaudatus Kirby 1884 (C.A.C.).

Section I: Suppression of Black in Hybrids Between P. glaucus and
P. eurymedon (D.A.W.)

Papilio eurymedon (Fig. 1&2) is closely related to two other North
American Swallowtails, Papilio glaucus (Fig. 11-13) and Papilio rutulus
(Brower, 1959). Although laboratory crosses between P. eurymedon and
P. rutulus have not been reported, Wagner (1978) described an almost
certain male hybrid of these species from Idaho. In 1956 hybrids were

188

J. Res. Lepid.

obtained between black and yellow female glaucus and male P. euryme-
don (see Clarke and Sheppard, 1957) but only male insects were
produced. In the present paper an account is given of female hybrids
which were obtained (D.A.W.), and these are very informative as
regards a gene suppressing the black of glaucus .

Materials and Methods

P. eurymedon were provided by Michael Collins from the vicinity of Nevada
City, Nevada Co., California and Monitor Pass, Mono Co., California. They were
collected as pupae from native food plants, Ceanothus integerrimus Hooker &
Arnott (Rhamnaceae) and Prunus virginiana, var. demissa (Nuttall) (Rosaceae),
in late summer 1984 and 1985 and eclosed in the laboratory in Blacksburg the
following springs. P. glaucus were reared from local stock (Montgomery Co.,
Virginia) or from Wisconsin and Illinois stock provided by J. Mark Scriber. The
latter originated from south of the principal zone of interaction of P. g. glaucus
and P. g. canadensis in Wisconsin (Scriber et al. 1986 and see Section II). The
insects were hand-paired, and the females laid eggs on Prunus serotina Ehrhart
(a local foodplant of P. glaucus ). Larvae were reared on this species, either in
tight plastic boxes or on leafy stems kept turgid in Aquapics® in ventilated
plastic canisters. Rearing was on natural mid-summer day length supple¬
mented to 15 to 16 h by artificial light.

Samples of P. glaucus from Virginia and West Virginia and of P. eurymedon
from northern California and western Oregon were used for morphological j
comparisons with the hybrids. P. eurymedon differs from P. glaucus most
conspicuously in ground color, which is white or very pale yellow in the former
but bright yellow in males and some yellow females of the latter. Some yellow j
females of P. glaucus are more orange-yellow. The black stripes and wing
margins ofP. eurymedon are broader than those ofP. glaucus. As an index of the
extent of the black pattern, the width of the under fore wing black margin from ,
its inner edge to the margin of the wing along vein Cux was divided by the length
of vein Cux from the cell to the wing margin. The resulting ratio separates P.
glaucus from P. eurymedon.

Results

Table 1 summarizes the successful crosses and Table 2 the unsuccess¬
ful ones. The Fx hybrids (Broods 85.1 and 86.1, Figs. 3, 4, 15) are nearly
intermediate between the parental species in the black pattern, in the
larval thoracic eye spots (as in Clarke and Sheppard, 1957, Fig. 1) and in
pupal proportions. As Clarke and Sheppard (1957) noted, the hybrid
pupae do not show the green/brown dimorphism of P. eurymedon , all
being brown as in P. glaucus. In fact the female pupa of brood 85.2 had
some green markings, but they were in areas in which occasional P.
glaucus pupae are green and not in the well-defined zones in which that
color is found in green P. eurymedon pupae. In both sexes the ground
color of the wings is nearly as yellow as that of P. glaucus. The
submarginal spots on the upper and under sides of the hindwing of P.

Table 1. Successful matings between Papilio glaucus and P. eurymedon.

26(l-4):l-288, 1988

189

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190 J.Res.Lepid.

Table 2. Unsuccessful matings between Papilio glaucus and P.
eurymedon.

Mating No.
and date

Source of Parents
Mother

Father

Remarks

86.3

28 May 86

yellow glaucus
(unknown)

eurymedon
(Nev. City, CA)

14 infertile eggs
(no spermatophore)

86.4

31 May 86

eurymedon
(Nev. City, CA)

glaucus

(unknown)

41 infertile eggs

4 June 86

"

glaucus

(unknown)

28 infertile eggs
(1 tiny spermatophore)

86.5

14 July 86

yellow glaucus
(unknown)

Fx ‘A’ ex 86.1

12 infertile eggs

19 July 86

//

Fx ‘L’ ex 86.1

unsuccessful pairing

19 July 86

rt

Fx ‘Q’ ex 86.1

very long pairing
(1 spermatophore?)

86.6

14 July 86

Fx ‘B’ ex 86.1

Fx ‘D’ ex 86.1

20 infertile eggs
(1 spermatophore)

86.10

17 July 86

Fj ‘N’ ex 86.1

glaucus

(unknown)

2 infertile eggs
(very long pairing, no
spermatophore)

86.13

29 July 86

Fx ‘Y’ ex 86.1

Fx ‘P’ ex 86.1

© #

39 infertile eggs
(1 spermatophore)

86.14

29 July 86

yellow glaucus
(unknown)

Fx T* ex 86.1

23 infertile eggs
(1 spermatophore)

glaucus are variable in size, shape and color but are usually rounder and
oranger than those of P. eurymedon. The hybrids are also variable in
these spots but are closer to P. glaucus in color and intermediate in size
and shape. In all respects the male hybrids in broods 85.1 and 86.1
resemble those described by Clarke and Sheppard (1957) from the
reciprocal species cross.

Brood 85-3, a backcross of a male Fx from brood 85.1 to the black
female form of P. glaucus , produced males (Fig. 5) and females of both
forms: two black ones with a somewhat intermediate phenotype (Fig. 6),
resembling occasional specimens taken in Virginia (Fig. 14), and two
yellow ones (Fig. 7). One of the yellow females was backcrossed again, to
a male P. glaucus (brood 86.9). All of the female pupae developed, and
they produced six black and one yellow adults (Fig. 9 and 10; male in

26(l-4):l-288, 1988

191

Figs. 1-14. Species and hybrids of Papilio. 1. 8
eurymedon (Nevada Co., California) Forewing base-to-
tip = 52 mm, all other figures to the same scale; 2. 9
eurymedon (Lake Co., Oregon); 3. F, 8 (brood 86.1 , 9
eurymedon x 8 glaucus)', 4. F, 9 (as in Fig. 3); 5.
Backcross 8 (brood 85.3, black 9 glaucus x 8 ex 85.1);
6. Backcross 9 (as in Fig.5); 7. Backcross 9 (as in Fig. 5);
8. 2nd backcross 8 (brood 86.9, yellow 9 ex 85.3 x 8
glaucus)', 9. 2nd backcross 9 (as in Fig. 8); 10. 2nd
backcross 9 (as in Fig. 8); 11. <5 glaucus (Montgomery
Co., Virginia); 12. 9 glaucus (reared, Montgomery/Giles
Co., Virginia); 13. 9 glaucus (Montgomery Co., Virginia);
14. 9 glaucus (Giles Co., Virginia, 1970).

192

J.Res.Lepid.

Fig. 8). Fig. 15 shows that the variation in the fore wing black margin
ratio is inherited in hybrids and backcrosses as a quantitative trait with
little evidence of dominance.

Discussion

Brood 86.1, and backcross and F2 broods 86.7, 86.9, 86.11 and 86.12,
show that the cross eurymedon 9 x glaucus 6 can be completely
compatible, to the extent that fertile males and females are produced.
All pupae of brood 86.1 eclosed 12-13 days after pupation, the normal
minimum time for imaginal development. Taken with Clarke and
Sheppard’s results (1957) and brood 85.1, in all of which only male
hybrids eclosed, brood 86.1 suggests that there is variation in the
crossability of the two species and that Haldane’s Rule need not apply
(Haldane, 1922). Thus one of the reasons that Brower (1959) judged P.
glaucus more closely related to P. rutulus than to P. eurymedon is
sometimes removed. More crosses will be needed to assess the variation
in cross-compatibility among these three species.

The ground color of yellow female P. glaucus is often just like that of
males, and the genes responsible for it should therefore be autosomal (or
X-linked), being found in both sexes. The black pigment, which Clarke
and Clarke (1983) believe is “added” in black females, is of course Y-
linked in its inheritance (Clarke and Sheppard, 1962; Clark and Clarke,
1983). The female hybrids in brood 86.1 support the view that yellow is
autosomal or X-linked, because they are nearly as yellow as P. glaucus
but are carrying a Y chromosome from P. eurymedon The blue scaling of
P. glaucus females appears also to be autosomal (or X-linked) since,
although P. glaucus males usually lack the blue, and P. eurymedon has
reduced blue scaling, the female hybrids have nearly full expression as
in P. glaucus. The male parent evidently carried the genes, but they
were expressed only in females.

The two yellow daughters in 85.3 (Fig. 7) were a surprise, since all the
daughters should be carrying the mother’s Y chromosome, and there is
ample evidence that black is Y -determined (Clarke and Sheppard, 1962).
This suggests that P. eurymedon has a suppressor of that black pheno¬
type, as discussed below in P. g. canadensis (Scriber et al., 1986) and P.
rutulus (Clarke et al., 1976). The possibility of a suppressor was tested
by a further back cross of one of the yellow daughters in brood 85.3 to a
male P. glaucus (brood 86.9). Black reappeared in six of the seven female
offspring (not different from a 1:1 ratio) and this could have been
because the yellow female parent was heterozygous for the suppressor.
The suppressor cannot be X-linked, because the mother in 86.9 passed it
to her yellow daughters but would have passed an X only to her sons. It
also cannot be Y-linked, because an Fx male passed it to some of his
daughters (brood 85.3). There are therefore good grounds for invoking
an autosomal suppressor of black in eurymedon , though another remote

26(l-4):l-288, 1988

193

P eurvmedon cftf

1 — 1 — 1 1 — 1 1 1 1 — 1

P eurvmedon 9 9 r— i

— t

i — i — i n

t

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2nd glaucus backcross 99

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.32

.34

.36 .38 .40 .42 .44 .46 .48 .50 .52 .54 .56 .58 .60 .62

under forewing black margin

cell to margin

Fig. 15. Distributions of the relative width of the forewing black marigin of P.

glaucus, P. eurymedon and hybrids between them. Relative width is
given as the ratio of black margin to the distance from cell to margin,
measured along vein Cu^ Means are marked by arrows.

possibility must be mentioned. The suppressor could have been intro¬
duced from glaucus of Wisconsin stock by the male parent of the original
cross 85.1, but arguing against this are the facts that the Wisconsin
male came from south of the range of P. g. canadensis and had a black
mother and, more importantly, that among the black glaucus broods
raised in Blacksburg from that particular stock there were none with
suppression of the black phenotype.

Section II: Suppression of Black in Hybrids Between P. g. glaucus and
P. rutulus, P. g. canadensis , and P. multicaudatus (CA.C.)

There are hybrid zones where naturally occurring crosses can take
place between some of these species (see map in Scriber et al. 1986) and
all are easy to initiate using the hand-pairing technique.

Hybrids between P. g. glaucus and P. rutulus (Table 3)

In 1955 we hand-paired a female rutulus with a male glaucus and
obtained 10 males and 8 yellow females (Clarke and Sheppard, 1955),

194

J. Res. Lepid.

Table 3. Crosses between black glaucus females and rutulus males
where suppression might occur. Some of the black insects had
yellow “brackets” on the forewings, but these are probably
normal variants.

Type of

Brood

Parent

Offspring

Cross

No.

Mother

Father

dd

Black Interm.

99 99

Yellow

99

Fx

14013

black

glaucus

rutulus

California

18

0

2+

0

15416

rutulus

Vernon, B.C.

1

0

0

2

backcross

8455

glauclrut

Fx hybrid

8

4

0

2

"

9608

"

"

13

6

0

1

"

9699

"

*

8

3

1

0

»

14173

"

"

18

3

4

0

2nd back.

14655

"

14173

25

16

2

0

9769

ex bl. glauc. 9
x ( glauclrut ) Fx<d

14

8

0

0

"

14639

"

"

16

4

0

0

"

14642

"

similar to 9769
and 14639

21

17*

1

0

3rd back.

14761

"

ex 14639

9

4

1

0

"

14763

*

"

17

21

0

0

* 6 with yellow showing through papery wings, 11 normal black.

+ Ecdysterone treated.

and in the same year found a similar situation (unpublished) when the
mother was yellow glaucus and the male rutulus. However, importantly,
the Fi cross between a black female glaucus x rutulus male, brood
14013, only yielded females when ecdysterone was injected into the
pupae. Then females intermediate in color (and resembling those in
brood 85.3, above) were produced. This suggested that here rutulus
carried a gene which partially suppressed the black pigment (Clarke et
al., 1976, Clarke and Willig, 1977, Clarke and Clarke, 1983). Later, in
our brood 15416, (previously unpublished) a black glaucus female
mated to a male rutulus from Vernon B.C., Canada, produced one male
and two yellow females, suppression of the black here being complete.
There was also evidence of suppression on back crossing some male
glaucus! rutulus F 1 hybrids to black glaucus females, since we occasion¬
ally obtained broods where the female offspring segregated for black
and yellow wing color. On the whole, however, yellow was less evident
in the backcross (Table 3).

More difficult to assess from the point of view of the suppressor are
minor degrees of yellow in black females. Sometimes the yellow shows
through and sometimes there are yellow “brackets” (see Clarke and
Clarke, 1983, Figs. 8 and 10). These might be due to partial suppression

26(l-4):l-288, 1988

195

or be normal variants of the form, particulary as there is some evidence
that the black pigment is laid down late in development (Clarke and
Willig, 1977).

Hybrids between P. g. glaucus and P. g. canadensis (Table 4—8)

The ranges of these two subspecies overlap, and canadensis also has
common ground with P. eurymedon and P. rutulus. All these three
butterflies are monomorphic, none having a black form, and it would
not be surprising if black in P. g. glaucus were suppressed in the
northernmost part of its range, where canadensis will have naturally
hybridized with it. This is what in fact happens, but we are stating the
case with hindsight, for it was Mark Scriber and his colleagues who
greatly clarified the problem of the five species mentioned in this paper
(personal communication 1983, Scriber et al., 1986).

Tables 4,5,6 and 7, taken from Scriber et al. (1986), give the details of
families where these fundamental principles obtain. Table 4 shows that
black is suppressed in Fx hybrids between P. g. glaucus and P. g.
canadensis. The suppressor is transmitted by F x males in backcrosses to
black glaucus females, since both yellow and black daughters appear
(Table 5), but not by “suppressed” yellow backcross females in crosses
with glaucus males (Table 6). These results are consistent with Scriber
el al.’s (1986) hypothesis that the suppressor is X-linked. We ourselves
have two similar examples in Table 8. The first (brood 18288) derives
from a black female glaucus mated to canadensis , and this produced
yellow females (cf. Table 4); the second (brood 18344) was bred by using
a yellow “suppressed” female 18288 mated to a Georgian male (well
south of the canadensis -glaucus interface) and demonstrates the re¬
appearance of black (cf. Table 6).

Thus the matter at first sight seems simple - the suppressor is the
result of a single autosomal or X-linked gene dominant in effect, and in
the examples given there is clear-cut segregation black/yellow. But
there are problems.

In Scriber’s F2 results (Table 7) half of the female offspring should be
black, but no black appeared. Furthermore, in the backcross to black
(see tables 3 and 8) the ratios are not strictly Mendelian, and in practice
both in the field and in the laboratory clear-cut black/yellow are often
blurred; all grades from black through intermediates to yellow are
encountered. An attempt to explain the genetics of this is made in the
general discussion; all we wish to emphasize here is that a suppressor
mechanism undoubtedly exists in P. g. canadensis.

Hybrids between P. g. glaucus and P. multicaudatus (see Table 9)

In the Fj broods using either black or yellow female glaucus and
hand-mating them to male multicaudatus we only obtained male
offspring (68 in one brood), but in backcrosses of glaucus females to
hybrid males females were obtained in 8 broods and the sex ratio was

196 J.Res.Lepid.

Table 4.

Offspring of crosses between black P. g. glaucus females and
P. g. canadensis males (from Scriber et al., 1986).

Pairing

Code

Males

Females

Yellow Black

Pupae still in diapause
at time information
was given

74

2

1

0

0

73

18

17

0

7

112

4

0

0

5

4

1

3

0

5

39

18

4

0

26

75

49

2

0

54

113

7

0

0

10

Table 5. Offspring of crosses between black P. g. glaucus females and

F i hybrid males, P. g. canadensis 9 x P. g. glaucus 8 (from

Scriber et al., 1986).

Pairing Male

Female

Code

Yellow

Black

6 8

1

3

14 7

5

4

15 40

11

20

Table 6.

Offspring of crosses between various P. g. glaucus males and
yellow females from the backcross of F 1 ( glaucus x canadensis)
8 x black glaucus 9 (from Scriber et al., 1986).

Pairing

Male

Females

Code

Yellow Black

148

2

0 2

154

1

0 7

157

2

0 7

225

4

0 1

155

3

0 1

not significantly upset. Seven of these broods were to yellow glaucus
females and therefore not informative as regards suppression, but in the
single backcross to black which produced butterflies (brood 6329, table
9) the offspring were two males and two yellow females, one of them

26(l-4):l-288, 1988

197

Table 7. Offspring of crosses between yellow Fx females (cross 73,
Table 4) and F x males (ex black glaucus 9 x canadensis $ )
(from Scriber et al., 1986).

Pairing

Code

Male

Yellow

Females

Black

272

29

28

0

288

19

12

0

279

11

6

0

Table 8.

Hybrids between P. glaucus and P. g. canadensis bred from
Scriber stock by CAC in England.

Parents Offspring

Brood

No.

Mother Father

Males

Black

Females

Yellow

Females

18288

Black canadensis

Wisconsin

3

0

3

18344

Yellow glaucus

18288 (Georgia)

8

8

0

Table 9. Hybrids between P. glaucus and P. multicaudatus (CAC)

Brood

No.

Parents

Mother Father

Males

Offspring

Black

Females

Yellow

Females

6139

black

glaucus

multicaudatus

68

0

0

6329

black

glaucus

6139

2

0

2

described as “dark yellow.” Unfortunately these insects are lost, but
there is no reason to doubt the record, particularly in the light of later
developments with rutulus, eurymedon and canadensis.

More work is obviously indicated and we have pupae of both species
(June 1987).

198

J.Res.Lepid.

General Discussion

The purpose of this paper has been to bring together the evidence
showing that in four monomorphic yellow Tiger Swallowtails there are
suppressors which can inhibit the expression of the black form of Papilio
glaucus. In P. eurymedon the suppressor is clearly autosomal, since it
can be transmitted from a father or a mother to a daughter, but the
critical evidence is lacking for the other species. If the suppressor in P. g.
canadensis were also autosomal, the expected frequency of “suppressed”
yellow daughters in the F2 broods of Scriber et al. (1986, Table 7) would
be 3/4 (rather than 1/2, as from the hypothesis of X-linkage), but the lack
of black daughters in those broods is still significant. In all other broods
the expectations are the same for both modes of inheritance. Why these
suppressors are present is not known. If P. g. canadensis and the three
western species are derivatives of P. glaucus populations with female
yellow/black dimorphism, then the suppressors may have been part of
the genetic system that cut them off from the ancestral species.
Otherwise the existence of suppressors in populations far from the
nearest overlap with P. g. glaucus (e.g., P. eurymedon in central
California) is difficult to explain. On the other hand, the history of
speciation in the glaucus group, perhaps excepting P. g. cnadensis, is
unknown.

Although it was thought possible (Clarke et al., 1976) to associate the
presence of the nuclear heteropyknotic body only with the genetic factor
for black, and therefore to be able to determine if the western “sup¬
pressed” species still carry that factor, it now appears that the heter¬
opyknotic body is not a sure marker for black (Cross & Gill, 1979).

Suppression is common in Batesian mimetic butterflies, many species
of which have mimetic patterns limited to females, although inherit¬
ance is autosomal. The mechanism of suppression of sex-limited traits is
not known, but it appears to exist even in monomorphic species, since
male hybrids do not show the mimetic patterns (Clarke and Sheppard,
1972). Crosses with non-mimetic species, however, usually produce
more or less intermediate females, with little evidence of interspecific
suppression (seeP. fuscus x P . polytes, Clarke and Sheppard, 1972, Plate
42h).

The intermediate forms and unusual Mendelian ratios mentioned
earlier would formerly have been explained by differences in expres¬
sivity of the gene or genes, but recent studies of the human X chromo¬
some using the techniques of molecular biology suggest other possibilites.
For example, an inherited form of mental deficiency appears to be
transmitted as an X-linked recessive. The lesion responsible is an X
chromosomal abnormality known as the fragile site, which can be induced
in cell cultures by various procedures or chemicals. However, as with
suppressors, problems soon arose. Occasional males with the fragile X
were mentally normal, and some carrier females were affected. Pembrey

26(l-4):l-288, 1988

199

et al. (1985) erected a two-stage hypothesis by which the fragile X occurred
as a mutant in the male X chromosome but produced no ill effect. However,
at the formation of gametes in the daughter of such a male, recombination
took place such that the fragile X mutation became more damaging, with
half of the daughter’s sons being handicapped and half her female
offspring carriers. Of particular relevance to race crosses in butterflies are
hybrid studies with the fragile X chromosome (Ledbetter et al., 1986).
There the threshold for initiation of fragility by various chemicals (for
example caffeine) is much reduced if the human X is isolated in a rodent
genetic background. It seems not unlikely that everyone has a fragile-X
site if the chromosome is suitably manipulated. Possibly the non-suppres¬
sion of black pigment in P. g. glaucus is similarly the exceptional state in
the glaucus group, with the other species carrying the potential for black
permanently suppressed. That possibility could be tested with molecular
techniques which have the potential of probing for suppressors in both
humans and butterflies.

Acknowledgements. We thank Michael Collins, David McCorkle and J. Mark
Scriber for material. One of US (C.A.C.) is indebted to the Nuffield Foundation
for generous support.

Literature Cited

BROWER, L.P., 1959. Speciation in butterflies of the Papilio glaucus group. I.
Morphological relationships and hybridization. Evolution 13:40-63.

CLARKE, SIR CYRIL & CLARKE, LADY F.M.M., 1983. Abnormalities of wing pattern in
the eastern tiger swallowtail butterfly, Papilio glaucus. Systematic
Entomology 8:25-28.

CLARKE, C.A. & SHEPPARD, P.M., 1955. The breeding in captivity of the hybrid Papilio
rutulus female X Papilio glaucus male. Lepidopterists’ News 9:46-48.

- , 1957. The breeding in captivity of the hybrid Papilio glaucus female X

Papilio eurymedon male. Lepidopterists’ News 11:201-206.

- , 1959. The genetics of some mimetic forms of Papilio dardanus, Brown,

and Papilio glaucus, Linn. Journal of Genetics 56:236-259.

- , 1962. The genetics of the mimetic butterfly Papilio glaucus. Ecology

43:159-161.

- , 1972. The genetics of the mimetic butterfly Papilio polytes L. Philoso¬
phical Transactions of the Royal Society of London B 263:431-458

CLARKE, C.A., SHEPPARD, P.M., & MITTWOCH, u., 1976. Heterochromatin polymorphism
and colour pattern in the tiger swallowtail butterfly Papilio glaucus L.
Nature 263:585-587.

CLARKE, C.A. & WILLIG, A., 1977. The use of a-ecdysone to break permanent diapause
of female hybrids between Papilio glaucus L. female and Papilio rutulus Lucas
male. Journal of Research on the Lepidoptera 16:245-248.

CROSS, W. & GILL, A., 1979. A new technique for the prospective survey of sex
chromatin using the larvae of Lepidoptera. Journal of the Lepidopterists’
Society 33:50-55.

200 J.Res.Lepid.

HALDANE, J.B.S., 1922. Sex ratio and unisexual sterility in hybrid animals. Journal
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LEDBETTER, D.H., LEDBETTER, s.A. & NUSSBAUM, R.L., 1986. Implications of fragile X
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163.

PEMBREY, M E., WINTER, R.M. & DAVIES, K.E., 1985. A premutation that generates a
defect at crossing over explains the inheritance of fragile X mental retardation.
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and London.

WAGNER, W.H Jr., 1978. A probable natural hybrid of Papilio eurymedon and P.
rutulus (Papilionidae) from Idaho. Journal of the Lepidopterists’ Society
32:226-228.

Journal ofResearch on theLepidoptera

26(1-4):201-218, 1988

New Species and new Nomenclature in the American

Acronictinae

(Lepidoptera: Noctuidae)

Douglas C. Ferguson

Systematic Entomology Laboratory, Agricultural Research Service, USDA, c/o National
Museum of Natural History, Washington, D. C. 20560.

Abstract. Described as new species are Acronicta sinescripta from
South Carolina, Louisiana, and Florida; A. kendallorum from Chi¬
huahua, Mexico; A. perblanda from the region between North Carolina
and Missouri southward to northern Florida and Louisiana; and
C ryphia cyanympha from South Carolina, northern Florida, and eastern
Texas. C. cyanympha is the only member of the tribe Bryophilini
definitely known from eastern North America. A. kendallorum was
reared from larvae on Froelichia arizonica (Amaranthaceae); A. per¬
blanda is associated with cypress swamps, although it has not been
reared; and C. cyanympha is related to a Caribbean group believed to
feed on lichens of the genus Usnea. The host of A. sinescripta is
unknown. A. perblanda is divergent and only doubtfully referred to the
genus Acronicta. Agriopodes jucundella Dyar and Bryolymnia huastea
Schaus from Puerto Rico, which appear closely related to C. cyanympha,
are transferred to the genus C ryphia Hubner and considered synonyms,
with jucundella being the older name. Agriopodes teratophora (Herrich-
Schaffer) is transferred to the genus Anterastria Sugi in the Acontiinae.

Introduction

The three species from the southeastern United States treated in this
paper have been known and recognized as undescribed for nearly 20
years, and questions concerning their identity have arisen repeatedly.
It is time that they were named. I include also a new species of Acronicta
from northern Mexico that might eventually be found to occur in the
border region of the United States. Three of the new species are in the
tribe Acronictini and placed in the genus Acronicta Ochsenheimer,
although one of them is referred to this genus somewhat doubtfully, as I
explain later. The fourth, a species of Cryphia Hubner, belongs to
the tribe Bryophilini as currently understood. This moth, Cryphia
cyanympha , n. sp., is particularly interesting because it is the only
bryophiline definitely known to be present in eastern North America,
belongs to a small group otherwise known only from the Greater
Antilles, and is almost certainly a lichen feeder. Although Cerma galva
Strecker, 1898 (=C ryphia galva, Franclemont and Todd, in Hodges, et
al., 1983), was described from Clyde, Wayne County, New York, its type

202

J.Res.Lepid.

has since been identified as a specimen of the western Cryphia olivacea
(Smith, 1891) (R. W. Poole, pers. comm.), a species not otherwise known
from the eastern United States. Strecker evidently cited a false type
locality.

My assignment of cyanympha to the genus Cryphia followed a review
of the species formerly listed under Agriopodes Hampson in North
American lists, as well as a cursory investigation of Bryolymnia
Hampson, and of species formerly referred to Bryophila Treitschke. It is
in agreement with the arrangement of Franclemont and Todd (in
Hodges et al. 1983: 136), in which species thought to be related to
Agriopodes fallax (Herrich-Schaffer), the type species of Agriopodes
(Figs. 31, 32), were left in that genus, and the remainder were combined
with species transferred from Cerma Hubner to form the American
section of the genus Cryphia (formerly Bryophila in much of the Old
World literature) (Acronictinae: Bryophilini). This has nothing to do
with the Cryphia of previous North American literature, in which the
name was misplaced in the Acontiinae as a result of a misunderstanding
of the type species. Those moths are now in the genus Hyperstrotia
Hampson. The type species of Cryphia is the European C. deceptricula
(Hubner). It is still uncertain whether C. cyanympha and its Caribbean
relatives should be regarded as congeneric with any other species
assigned to the Bryophilini, but within the present classification Cryphia
is where they fit best. A thorough revaluation of Cryphia , Bryophila,
Bryoleuca Hampson, and other related Old World genera and their type
species is needed but beyond the scope of the present paper.

It seems clear that the North American species long known as
Agriopodes teratophora (Herrich-Schaffer) (= Bryophila teratophora
Herrich-Schaffer, [1854]) is misplaced and belongs in the genus Anteras-
tria Sugi, 1982. This generic name was proposed for a closely related
Japanese counterpart of teratophora originally described as Erastria
atrata Butler, 1881 (Sugi, in Inoue et al., 1982, vol. 1:818; vol. 2:383, pi.
197, fig. 57). The American species therefore becomes Anterastria
teratophora (Herrich-Schaffer, 1854), NEW COMBINATION, and it
should be transferred to the Acontiinae, tribe Eustrotiini, following
Sugi’s arrangement. That would place it somewhere near Lithacodia
Hubner in the North American fauna. However, the male genitalia of
both species of Anterastria are disconcertingly similar to those of species
of Cryphia treated in this paper, so much so that one might even be led to
consider them congeneric on that basis. If the reassignment of Anteras¬
tria to the Acontiinae is correct, the resemblance to Cryphia must be a
result of convergence. Anterastria teratophora was illustrated most
recently by Coveil (1984: pi. 25, fig. 19).

Acronicta sinescripta Ferguson, new species

Diagnosis. This is a southeastern species that most closely resembles Acronicta
oblinita (J. E. Smith), especially the evenly gray, almost unmarked form insolita

26(l-4):l-288, 1988

203

Grote; but it differs in the configuration of dark markings associated with the
reniform spot, the regular rather than dentate postmedial line (if visible), the
grayer hindwings (white in both sexes of oblinita ), and the absence of dark
terminal dots along the outer margin of the hindwing. Also, on the underside,
the forewing is much darker than the hindwing in sinescripta, almost as pale in
oblinita , and both wrings beneath lack the discal spot present in oblinita.
Another species with which sinescripta could be confused is A. lanceolaria
Grote, which does have traces of similar dark markings associated with the
reniform. However, lanceolaria is larger, has a more produced and pointed apex
on the forewing, lacks the basal streak and other dark forewing markings of
sinescripta , often has a fairly prominent, sinuous, pale postmedial line, especially
in females and, like oblinita , an almost white hindwing. A. sinescripta may be
distinguished from A. arioch Strecker, which looks like a large, pale form of
oblinita, by essentially the same differences that separate it from the latter
species.

Further description. About same size and shape as A. oblinita. Male antenna
laminate, finely setose, whitish-scaled dorsally, tapering to an almost simple
tip; female antenna filiform, finely ciliate ventrally, weakly whitish- scaled
dorsally; palpus slender, about as long as distance across front diagonally in
both sexes, white ventrally, grayish or mixed black and white apically, black
dorsally beneath eye; front narrower than eye in male, about as wide as eye in
female, almost flat, with mixture of white and dark-gray or blackish scales; eye
bordered posteriorly by a few dark scales but without heavy black border;
vestiture of body fight gray; head and thorax lightly streaked with darker scales
and appearing slightly darker than abdomen; tegula and patagia similarly
gray, unmarked; legs normal, stoutish, gray, unmarked; hindtibia somewhat
swollen. A. sinescripta, oblinita, and others of this group lack the dark facial
band comprised of black contiguous areas of front, palpi, tegulae, and eyes that
create the black-masked appearance of most Acronicta species.

Wings elongated like those of oblinita, but forewing a little less pointed, its
shape intermediate between those of oblinita and lithospila Grote. Upperside of
fore wing uniformly bluish gray, resulting from even mixture of whitish and
gray scales; almost unmarked, although variable vestiges of pattern are
present, as follows: thin, black, basal dash on cubital stem running about 6 mm
toward middle of wing; thin, weak, black longitudinal streak running through
outer end of cell, appearing to pass through and be interrupted by reniform spot,
although reniform not really visible; this streak, usually dividing to form
vaguely defined, longitudinal cell at that point, then continues as a faint, dark
fine on M2 toward outer margin; other short, faint, black dashes may occur in
outer third of forewing on R4, M1? and Cu2. Some specimens show traces of a
postmedial fine that is evenly curved, subparallel to and quite near outer
margin; others show a wide, very diffuse, dark transverse band through
postmedial area of forewing or both wings. Hindwing whitish, variably dusted
with gray scales, especially toward apex and costa. Fringes concolorous with
wings or slightly paler. Underside of forewing gray, darkest mesially, usually
paler near costa and in outer third. Underside of hindwing about like upperside,
darkest toward costa. Length of forewing: holotype, 19 mm; other 66, 18-19
mm. Allotype, 20 mm; other 9 9, 19-21 mm.

Male genitalia (Figs. 16, 17). Valve broad, rounded, simple, with clasper
consisting mainly of one large, stout process that does not quite reach costa;

204 J.Res.Lepid.

uncus normal but delicate; juxta partly spinulate; vesica with cluster of about 14
medium-sized cornuti with fine, attenuated points.

Female genitalia (Fig. 29). These were compared with genitalia of oblinita
and lepusculina Guenee, representatives of two groups to which I thought
sinescripta might be related. It differs obviously from oblinita in having more
elongated, pear-shaped bursa copulatrix, nearly twice as long as that of oblinita ;
longer, much more swollen, sclerotized and rugose ductus bursae; and more
abruptly and deeply cleft posterior margin on eighth sternite. Ductus bursae
adjoins corpus bursae apically, and entire bursa copulatrix and eighth sternite
are of different form, the corpus bursae long and slender posteriorly, ending in
zone of heavy sclerotization where it meets the very short ductus bursae, and the
eighth sternite unnotched but with wide, shallow concavity on posterior
margin. On basis of female genitalia, sinescripta would appear more closely
related to oblinita than to lepusculina.

Types. Holotype 6, 4.2 mi. NE of Abita Springs, St. Tammany Parish,
Louisiana, 8 April 1984, V. A. Brou (Fig. 1). Allotype 9, same locality and
collector, 16 June 1984 (Fig. 2). Paratypes: 1 9, same locality and collector, 20
August 1982; 10 6 6, same locality and collector, 17, 30 April, 7 May, 11, 15, 30
June, 3, 7, 8, 20 July 1983; 1 9 , same locality and collector, 16 July 1983; 18 8 6,
same locality and collector, 3 March, 1, 3, 4, 7, 11, 21 April, 4, 5, 14, 15, 18, 23, 26
June, 1, 21 (2), 28 August 1984; 1 9 , same locality and collector, 21 June 1984; 1
9, MTF [Mississippi Test Facility, Natl. Air and Space Admin.], Hancock Co.,
Mississippi, 2 July 1973, R. Kergosien; 1 9, Gainesville, Florida, 30 March
1929; 4 6 6, Archbold Biological Station, Lake Placid, Highlands Co., Florida,
29 March 1959 (d genitalia slide 6751, J. G. Franclemont — Figs. 16, 17), 30, 31
March, 2 April 1959, J. G. Franclemont; 1 9 , some locality and collector, 3 April
1959; 1 9, The Wedge Plantation, McClellanville, South Carolina, 20 August
1968, D. C. Ferguson (Fig. 3); 3 9 9, same locality, 23 May, 23 June, 7 August
1971, R. B. Dominick and C. R. Edwards. All as far as known were collected at
light. Holotype and allotype in U.S. National Museum of Natural History;
paratypes in National Museum, Canadian National Collection, Florida State
Collection at Gainesville, British Museum (Natural History), and collections of
V. A. Brou, J. G. Franclemont, and Bryant Mather.

Distribution. Coastal South Carolina to Highlands County, Florida, and
westward through the Gulf States to Louisiana.

Early Stages. Unknown.

Remarks. I have known of this species since I collected a specimen in 1968,
and shortly afterward found a second specimen, from Gainesville, in the
collection of the U. S. National Museum. Several others were soon sent by other
collectors, but all were females. When I prepared an earlier draft of this paper in
the 1970’s and sent it to Dr. Franclemont for his comments, I learned that he had
collected four males and a female in Florida in 1959. He kindly responded by
sending the slide from which my illustration of the male genitalia (Figs. 16, 17)
was then prepared. Then in 1982, 1983, and 1984, Vernon A. Brou, at one
locality in Louisiana, collected three times as many specimens as were previously
known, and these include such fresh, bright examples that I chose two of them as
holotype and allotype.

Acronicta kendallorum Ferguson, new species

Diagnosis. A gray-dusted, medium-sized Acronicta of rather nondescript
appearance, perhaps most resembling a dark A. sperata Grote or rubricoma

26(1-4): 1-288, 1988

205

Guenee, with similar wing shape and pattern but differing from these and
apparently from all other North American species of the genus in having
pectinate male antennae and very short, hairy palpi in both sexes. Male
genitalia normal for the group and similar to those of A. impressa Walker,
although armature of valve differently oriented, juxta differently shaped, and
vesica with more and longer cornuti. Known only from a small reared lot from
southwestern Chihuahua, Mexico.

Further description. Male antenna bipectinate with branches so short and
closely set as to appear laminate, each lamellum about as long as thickness of
shaft and orange brown; shaft scaled dorsally with black and white scales;
female antenna simple; palpus of both sexes not or only slightly protruding
beyond front, clothed in hairlike scales; front slightly protuberant; eye of male
about average for genus, about as wide as front, that of female three-fourths as
wide as front; tongue moderately developed, not large; ocellus present, small;
legs heavily clothed with long scales, all spurs present; epiphysis large but not
quite reaching end of fore tibia.

Thorax and abdomen gray, resulting from mixture of dark-gray and white
scales; front and vertex gray, with thin white margin adjacent to eye; dark mask
that passes from palpi back through eye in many Acronicta species only vaguely
apparent, not contrasting; legs gray, tarsi dark with white ring marking end of
each segment.

Upperside of forewing with an almost equal mixture of white and blackish
scales and with usual pattern of antemedial and postmedial bands and orbicular
and reniform spots present but not very distinct; dark scales in these elements of
pattern seen to be symmetrically aligned in transverse rows under magnifica¬
tion; costa with about eight small black spots that indicate otherwise largely
obsolescent transverse lines or bands of forewing pattern; black and white
checkered fringe preceded by weak, discontinuous, dark terminal line. Hindwing
pale gray, nearly white, with slight dusting of darker gray-brown scales toward
outer margin, especially along veins; a weak, dark terminal line; and white,
unmarked fringes. Underside of wings almost unmarked; forewing light gray¬
ish with very diffuse, gray reniform; hind wing nearly white. Length of
forewing: holotype 6, 15 mm; other 66, 14, 15 mm; allotype 9, 16 mm.

Male genitalia. As illustrated (Figs. 20, 21).

Female genitalia. Not dissected.

Types. Holotype 6 , Creel, ca. 2134 m (7000 ft.), Chihuahua, Mexico, reared 10
April 1979 from larva on “? Froelichia arizonica Thornber” (Amaranthaceae),
Roy O. and C. A. Kendall (Fig. 4). Allotype 9, same data but emerged 30 May
1979 (Fig. 6). Paratypes: 2 6 6 , same data but emerged 7 May and 27 June 1979;
2 last-instar larvae in alcohol, same data but preserved 21 Sept. 1978. Holotype,
allotype, and paratype larvae deposited in collection of U.S. National Museum
of Natural History; paratypes returned to R. O. Kendall.

The exact type locality is about halfway up a small hill adjacent to the Motel
Parador de La Montana. Creel is about 177 km (110 mi.) southwest of the city of
Chihuahua in the region of the Sierra Tarahumara, and north of Copper Canyon
and Urique Canyon.

Distribution. Known only from the type locality.

Early stages. Type material was reared from larvae collected from a plant
tentatively identified by R. O. Kendall as Froelichia arizonica Thornber
(Amaranthaceae). Many larvae were present, some of them feeding on another,
still unidentified plant that had long, succulent stems and peltate leaves. Only a

206

J.Res.Lepid.

few were kept for rearing because an acceptable substitute food could not be
found. Twenty-five larvae were preserved (R. O. Kendall, pers. comm., 1985).
Larvae that matured and pupated in late September produced adults the next
spring, following a pupal diapause.

The larva (Fig. 15) is of a type that seems to place it in that section of the genus
that includes impressa Walker and perdita, noctivaga, and sperata of Grote, but
is conspicuously adorned with large, pale verrucae, contrasting boldly with the
very dark integument. It also has a wide but segmentally interrupted and
somewhat sinuous, lateral (subspiracular) band. The pale markings may have
been white or yellow in life. The head and thoracic legs are nearly black, and the
dark-brown body integument has a finely granulate texture. The verrucae bear
numerous long, whitish, barbed hairs, and centrally, from the dorsal verrucae
especially, arise one to four erect, blackish, barbed spines, almost as long as the
whitish hairs. Length of two larval paratypes: 33, 47 mm.

Remarks. I am pleased to name this species for its collectors, Roy O. and
Connie A. Kendall, through whose efforts not only the adult but also the larva
and food plant have been made known to us.

Acronicta perblanda Ferguson, new species

Diagnosis. This is a small, delicate, light-gray species easily recognized by
the straight black streak that runs often without interruption from the base to
near the tornus of the fore wing. The genitalia are distinctive, and there is a
question as to whether it is a true Acronicta ; but the pattern, including the
black markings of the head and legs, are typical of the lobeliae-laetifica-
interrupta group. A. perblanda would look very much like a diminutive lobeliae
were it not for the absence of the black dash associated with the reniform spot
and the exaggerated length of the basal and tornal dashes that may unite them
as one continuous streak. The species is southeastern and rare in collections.
Circumstantial evidence from collecting sites suggests an association with bald
cypress or with some plant that grows in cypress swamps.

Further description. Antenna filiform, minutely ciliate beneath, that of male
about as stout as longest tarsal segment, that of female much more slender than
longest tarsal segment, both with mixture of white and gray scales dorsally;
palpi of both sexes quite long, exceeding front by one-third their length, black
with whitish scales ventrally on proximal half, at tip, and a few on inner
surfaces; distal palpal segment small but visible; front somewhat protuberant,
clothed in white scales with a few gray ones mixed in, except that scales of lower
lateral corners, near palpus, are black; black areas of palpus, front, and tegula,
together with eye, forming a unified band that creates a masked effect in usual
manner of many Acronicta species; thorax and abdomen gray, clothed with
mixture of gray and whitish scales; tegula margined laterally with black scales;
short, black, middorsal streak just behind head, between patagia; scales on top
of head, between and behind bases of antennae, white; broad border of black
scales adjacent to eyes posteriorly; legs whitish with mixture of gray scales,
except mid- and hindtibiae, which are marked with a very distinctive, heavy,
full-length black streak on outer side; inner (dorsal) tibial spur of each pair at
least twice as long as outer one.

Fore wing somewhat elongated, about same shape as that of A. lobeliae but
much smaller; ground color of upperside pale gray, resulting from mixture of
white and light gray-brown scales, all markings black; most conspicuous

26(l-4):l-288, 1988

207

marking is long streak from base to point on outer margin just anterior to
tornus; basal and tomal (anal) black dashes characteristic of such species as
lobeliae and interrupta are longer in perblanda , being thinly joined across
median space or nearly so, thus forming what usually appears as a continuous
streak; smaller black dash in space between Cu2 and 2nd A, immediately behind
constricted or interrupted mid-zone of long streak, although this may be
obsolescent; a few black scales along inner margin posterior to lesser dash;
basal, antemedial, and what is probably postmedial line clearly outlined with
black scales toward costa; basal line short, curved, much as in A. interrupta ;
antemedial and postmedial lines, if apparent, sharply inclined outwardly from
costa, then acutely angled back toward base at radius, thus forming long
dentations at those points; antemedial continuing posteriorly to form rounded
lobe inwardly and lesser, blunted tooth outwardly before reaching large
longitudinal streak; short dashes usually present at outer margin in folds
between Mx and M2, and Cux and Cu2, and shorter ones in spaces between other
veins, giving fringes a checkered appearance; dark discal spots hardly apparent.
Black scales that form markings of forewing overlapped and offset, not arranged
in straight rows that give ridged or striated effect characteristic of other species
of Acronicta with similar markings. Hindwing dull grayish brown with whitish,
faintly checkered fringes. Underside of forewing brownish, of hindwing light
gray, with faint, very diffuse postmedial bands; hindwing with diffuse discal
spot. Length of forewing: holotype 9 , 12.0 mm; other 9 9 , 13.0 mm; allotype d,
11.0 mm; other SS, 10.5-12.0 mm.

Male genitalia. As shown (Figs. 18, 19). Everted vesica may be seen to have 10
long, very slender cornuti and numerous short ones.

Female genitalia. As shown (Fig. 30). Ductus bursae as long as corpus bursae,
nearly cylindrical, somewhat rugose, sclerotized, rigid.

Types. Holotype 9, McClellan ville, South Carolina [Wedge Plantation], 20
April 1970, R. B. Dominick and Charles R. Edwards; USNM Genitalia Slide No.
52,174 (Fig. 7). Allotype S , same locality and collectors, 25 April 1973 (Fig. 8).
Paratypes: 1 d, Hilliard [Nassau County], Florida, “4-2-46,” F. H. Chermock; 1
d, Gainesville, Florida, 13 April 1963, blacklight trap, H. A. Denmark; 1 d, Jet.
State Highway 101 and 181, Cartaret County, North Carolina, 30 April 1974, J.
Bolling Sullivan (Fig. 9); 1 d, Edgard, St. John Parish, Louisiana, 13 April 1979,
V. A. Brou; 1 9 , same locality and collector, 30 March 1982; 2 d d, 4.2 mi. NE of
Abita Springs, St. Tammany Parish, Louisiana, 24 April 1983, 22 April 1984,
V. A. Brou; 1 9, same locality and collector, 7 April 1984; 1 d, cypress-tupelo
swamp with associated southeastern flora, Otter Slough Wildlife Area, Stoddard
Co., Missouri, 28 May 1983, at blacklight, J. R. Heitzman (Fig. 10).

The holotype and allotype, which are deposited in the U.S. National Museum
of Natural History, were taken in a light trap at the edge of the Santee Delta, on
the south bank of the Santee River. Paratypes are in the Canadian National
Collection, the Florida State Collection of Arthropods at Gainesville, and in the
collections of V. A. Brou, Abita Springs, Louisiana, J. R. Heitzman, Indepen¬
dence, Missouri, and J. B. Sullivan, Beaufort, North Carolina.

Distribution. In cypress swamp areas from Cartaret County, just south of Cape
Hatteras, North Carolina, and Stoddard County, Missouri, to northern Florida
and southern Louisiana.

Early stages. Unknown. However, the occurrence of adults almost entirely in
April (one taken in Louisiana on 30 March and one in Missouri on 28 May)
indicates that the species is univoltine and probably overwinters as a pupa. The

208 J. Res. Lepid.

localities all appear to have been in the vicinity of bald cypress, Taxodium
distichum (L.) Rich.

Remarks. The female and to a lesser degree the male genitalia show such
obvious similarities to those of Acronicta ybasis Dyar (1918: 345), a Mexican
species, that the relationship must be close. Both have the wide, sclerotized,
regularly cylindrical ductus bursae. The size, wing shape, and slender build are
also similar, but the forewing pattern is different, ybasis having a distinct
reniform and orbicular, and short basal and anal dashes only.

The correct taxonomic disposition of this species is not likely to be made
without much more comprehensive revisionary study. But meanwhile, for
purposes of collection arrangement, I would simply add it to the end of the list of
North American Acronicta species, inasmuch as there seems to be nothing
closely comparable except the Mexican A. ybasis.

J. G. Franclement suggested to me that the moth listed from Florida as a
Catabena species by Kimball (1965: 104) is probably the same as A. perblanda.
The specimens were supposed to be in the Los Angeles County Museum of
Natural History but cannot now be found.

Cryphia cyanympha Ferguson, new species

Diagnosis. This is a small, green, recently discovered noctuid known from
South Carolina, northern Florida, and eastern Texas. Its predominantly blue-
green forewing might lead to confusion with Cyathissa percara (Morrison), an
unrelated species of similar size and coloring that also occurs in the Southeast;
but on closer inspection it is easily seen that the forewing pattern is different.
The pattern and male genitalia are characteristic, and the illustrations should
make identification simple. This moth belongs to a group of three or four closely
related species that occur on islands of the Caribbean, at least on Cuba,
Hispaniola and Puerto Rico, and of which only one seems to have been described
(see remarks). The closest known species in the continental United States may
be the southwestern Cryphia viridata (Harvey) (Figs. 23, 24).

Further description. Male antenna heavily ciliate, with setae about as long
as width of shaft; female antenna slender, with short setae; shaft in both sexes
with alternating transverse bands of light and dark scales dorsally. Head, scape,
and thorax with light-green to whitish scales, patagia black tipped; front
smooth, almost square; palpi of both sexes fairly long, nearly reaching top of
front, blackish with third segment white; ocelli very small, inconspicuous;
tongue moderately but not highly developed. Legs fairly short, rough-scaled,
with long spurs, irregularly blotched and banded with dark-brown and whitish
scales. Abdomen light gray brown, concolorous with upperside of hindwings,
with admixture of black scales beneath.

Upperside of forewing light bluish green with black and white markings as
follows: basal line black, prominent between costa and radius, otherwise
obsolete except for triangular projection on cubitus and a few black scales near
inner margin; antemedial and postmedial bands black, bordered with white, the
former white on inner side, the latter on outer side; inner white border one or
two scales wide, outer white border two or three scales wide; antemedial band
evenly excurved, reflexed outwardly at 2nd anal; black part of antemedial band
widening greatly at costa, less so at inner margin, with a broad rectangular

26(l-4):l-288, 1988

209

process extending outwardly as a black bar to end of cell, which is two-thirds
distance to postmedial band, this being followed posteriorly by some black scales
forming a second, less definite, rectangular configuration adjacent to the other
but less than half its length; postmedial line missing near costa, thence sinuous,
excurved at Mi and at 2nd A, incurved about M3, inclined outwardly and
somewhat expanded at inner margin where its black scales closely approach
those of antemedial band; black triangular spot at costa opposite end of discal
bar, this probably being remnant of medial band otherwise wanting in this
species; postmedial third of forewing with scattered black scales, especially in
middle area near outer margin and near costa, probably representing fragmen¬
tary submarginal band; black scales at outer margin form a weak terminal line,
preceded by border of white scales; fringes rather long, comprised of black-
tipped gray scales, with vague greenish rays opposite ends of R5, M2, and M3;
costa margined with a few black scales between basal and antemedial bands,
and with three small black patches in distal third, the first being opposite
postmedial band and probably marking point at which it meets costa. Hindwing
grayish brown, unmarked, thinly scaled, becoming slightly paler basally, and
with proximal tier of fringe scales concolorous, outer tier whitish. Underside of
forewing grayish brown, almost unmarked; basal two-thirds of costa dark, outer
third with three dark spots separated by areas of whitish scales; outer margin
with dark terminal line and moderately large, subcircular, light-greenish spot at
apex; discal spot faint. Underside of hindwing an even mosaic of white and
blackish-brown scales, except that there are fewer dark scales in space between
cubitus and third anal vein; outer margin with dark terminal line, somewhat
interrupted at vein endings and fading out toward inner angle. Length of
forewing: holotype, 8 mm; other 6 6, 7-8 mm; 9 9, 7-8 mm.

Male genitalia. As figured (Figs. 27, 28). Similar to those of Cryphia viridata
(Figs. 23, 24) but with simpler clasper, no transtilla, various differences in shape
of other components, and very large cornutus on vesica, plus a closely set,
seemingly deciduous clump of much smaller cornuti. Although moth looks more
like jucundella, it seems no closer in genital characters to that species (Figs. 25,
26) than to viridata.

Female genitalia. Of a generalized type that could be acronictine or almost
anything, but with good species characters. Although genitalia of female
holotype of C. huastea Schaus were badly mutilated by whoever dissected them,
making comparison of cyanympha with that species difficult, it appears that
cyanympha has a funnel-like ostium only about half as large, a heavily
sclerotized elbow in the ductus bursae lacking in huastea , a corpus bursae
without numerous spicules that may be seen in huastea , and two very delicate
signa, which together would equal no more than half the size of the one
invaginated, cuplike, spiculate signum of huastea. Bursa of huastea either
bilobate or distorted in such a way as to appear so. Bursa of cyanympha entire.
Female genitalia of C. viridata also have funnel-shaped ostium but of different
shape; otherwise simple, offering little basis for comparison, having simple,
membranous, slightly elongated bursa copulatrix without either spicules
or signa: ductus bursae shaped much like that of cyanympha but entirely
membranous.

Types. Holotype 6 , University of Florida Preserve, Welaka, Putnam County,
Florida, 24 March 1987, D. C. Ferguson (Fig. 12). Paratypes: 67 6 6, same
locality and collector, 24, 25, 28 March 1987; 16 6 6, same locality, site 4, live

210

J.Res.Lepid.

oak xeric hammock, 9-10 June 1986, J. B. Heppner and J. Powell; 89 8 6 , same
locality, some labelled Site 4 as above, others Site 5, slash pine-palmetto
flatwoods, 17-21 March 1986, J. B. Heppner; 1 8 , Gainesville, Alachua Co.,
Florida, 3 Oct. 1983, E. C. Knudson; 2 9 9, Withlacoochee State Forest, vie. Kirk
Hill, Hernando Co., Florida, 6 Sept. 1986, L. C. Dow; 1 9, Rock Spgs., Kelly
Park, 6 mi. N. of Apopka, Orange Co., Florida, 18 Feb. 1984, L. C. Dow; 1 6,
McClellanville [Wedge Plantation], South Carolina, 12 May 1970, R. B. Domi¬
nick and C. R. Edwards; 1 9 (Fig. 11), same locality and collectors, 26 Apr. 1970
(Fig. 11); 2 8 8, New Waverly, Texas, 14 June 1964. A. & M. E. Blanchard; 2 88,
Town Bluff (Dam B), Tyler Co., Texas, 15 Sept. 1975. Holotype and many
paratypes in U.S. National Museum; others in Florida State Collection,
Gainesville, and in the collections of E. C. Knudson, H. D. Baggett, and L. C.
Dow. Paratypes will be distributed to other collections.

Distribution. Charleston Co., South Carolina; Alachua, Putnam, Orange,
and Hernando counties, Florida; Tyler Co., Texas.

Early stages. Unknown, but the closely related C. jucundella (Dyar) from
Puerto Rico is evidently a lichen feeder. In the original description Dyar (1922:
11) wrote: “Mr. Wolcott states that the larvae fed on lichens, being of the same
color as the lichen, marked with brown, a beautiful example of protective
resemblance.” The type of jucundella (Fig. 13) is a small, bright (although
somewhat damaged) specimen that looks as though it could have been reared,
and pinned beside it in the drawer where it was originally placed by Dyar was a
sample of a lichen, but without explanatory labels. The lichen was identified for
me by Mason Hale, Department of Botany, National Museum of Natural
History, as a species of the Usnea longissima group. Many lichens, including a
similar Usnea species, occur conspicuously on the trunks and branches of trees
at the type locality of cyanympha.

Remarks. The closest known relatives of Cryphea cyanympha are small, green
species that have been collected in Cuba, the Dominican Republic and Puerto
Rico. One of these is Agriopodes jucundella Dyar (1922: 10) ( =Cryphia jucun¬
della (Dyar), new combination) (Figs. 13, 25, 26) from Puerto Rico. Bryolymnia
huastea Schaus (1940: 203) (Fig. 14), also from Puerto Rico, is almost certainly a
junior synonym of jucundella and was misplaced in Bryolymnia. The type
species of Bryolymnia, Dacira roma Druce of Mexico and Central America, is a
larger, stouter moth with a well-developed corona and an incipient cucullus in
the male genitalia, unlike species of Cryphia and other genera now placed in the
Acronictinae. Although the holotypes of jucundella and huastea are both before
me and appear conspecific, the former is a male and the latter a female, so that
their genitalia cannot be compared. Two other similar species, apparently both
undescribed, are represented by specimens in the U.S. National Museum from
the Greater Antilles. One male from Cuba appears identical to cyanympha
except that the valve lacks the clasper entirely.

Within the continental United States, Cryphia cyanympha is a distinctive,
easily recognized species, but I illustrate for comparison and to support the
generic placement the male genitalia of C. viridata and C . jucundella, as well as
those of the type species of Agriopodes, A. fallax, which may be seen to differ
conspicuously in the form of the juxta, valve, and aedeagus. A colored illustra¬
tion of fallax was given by Coveil (1984: pi. 26, fig 16).

26(l-4):l-288, 1988

211

Acknowledgements. The lepidopterists who kindly made available most of the
material on which this paper is based are as follows: H. D. Baggett, V. A. Brou,
L. C. Dow, J. G. Franclemont, J. R. Heitzman, J. B. Heppner, R. O. Kendall, E. C.
Knudson, J. D. Lafontaine, B. Mather, J. B. Sullivan, and the late R. B.
Dominick and C. P. Kimball. John G. Franclemont and the late Edward L. Todd
reviewed an earlier draft of the manuscript, Charles V. Covell, Jr. and F.
Christian Thompson reviewed the present version, and Robert W. Poole
provided answers to numerous questions. The drawings are by four illustrators:
Linda Heath Lawrence and Mary Lou Cooley of the Systematic Entomology
Laboratory, Young Sohn of the Department of Entomology, National Museum
of Natural History, and by me. I did the photography.

Literature Cited

COVELL C.V. JR. 1984. A Field Guide to the Moths of Eastern North America.

Houghton Mifflin Co., Boston. 496 pp., 64 pis., 76 text figs.

DYAR, H.G., 1918. Descriptions of new Lepidoptera from Mexico. Proc. U. S. Natl.
Mus. 54:335-372.

DYAR, H. G., 1922. New American moths and notes. Insecutor Inscitiae Menstruus
10:8-18.

HODGES, R. W., ET AL. 1983. Check fist of the Lepidoptera of America north of
Mexico. E. W. Classey Ltd. and The Wedge Entomological Research
Foundation, London, xxiv + 284 pp.

INOUE, H., S. SUGI, H. KUROKO, S. MORIUTI & A. KAWABE, 1982. Moths of Japan.

Kodansha Co. Ltd., Tokyo. Vol. 1: 966 pp.; vol. 2: 552 pp., 392 pis.
KIMBALL, C. P., 1965. Lepidoptera of Florida. Arthropods of Florida and neigh¬
boring land areas, vol. 1. Div. of Plant Industry, Florida Dept. Agric.,
Gainesville. 365 pp., 26 pis.

SCHAUS, W., 1940. Scientific survey of Porto [sic] Rico and the Virgin Islands
12(2): 175-290. New York Acad. Sci.

212

J.Res.Lepid.

Fig. 1 -6. New Species of Acronicta. 1 , A. sinescripta, holotype; 2, A. sinescripta,
allotype; 3, A. sinescripta , 9 paratype, Wedge Plantation, McClellanville,
S. C., 20 Aug. 1968, D. C. Ferguson; 4, A. kendallorum, holotype; 5, A.
kendallorum, 8 paratype, same data as for holotype but emerged 27
June 1979; 6, A. kendallorum, allotype. About natural size.

26(l-4):l-288, 1988

213

Figs. 7-14. Acronicta and Cryphia species. 7, A. perblanda, holotype; 8, A.

perblanda, allotype; 9, A. perblanda, 6 paratype, Cartaret County, N.
C., 30 Apr. 1974, J. B. Sullivan; 10, A. perblanda, 6 paratype, Otter
Slough Wildlife Area, Stoddard County, Mo., 28 May 1983, J. R.
Heitzman; 11, C. cyanympha, paratype 9, McClellanville, S. C., 26
Apr. 1970, R. B. Dominick & C. R. Edwards; 12, C. cyanympha,
holotype; 13, C. jucundella, holotype, Puerto Rico; 14. C. huastea,
holotype, Puerto Rico. About twice natural size.

214

J.Res.Lepid.

Fig. 1 5. Acronicta kendallorum, paratype larva (preserved in alcohol). Dorsal and
lateral views.

26(1-4): 1-288, 1988

215

Figs. 16-19. Acronicta species, 8 genitalia. 16, A. sinescripta ; 8 paratype,
Highlands County, Florida, J. G. Franclemont slide No. 6751; 17,
aedeagus of same specimen; 1 8, A. perblanda, 8 paratype, Hilliard,
Florida, D. C. Ferguson slide No. 1494 (Canadian Natl. Coll.); 19,
aedeagus of same specimen.

A

216

J.Res.Lepid.

Figs. 20-22. Acronicta kendallorum and Cryphia cyanympha, genitalia. 20, A.

kendallorum , holotype, main part of 6 genitalia. 21, aedeagus of
same specimen. 22, Cryphia cyanympha, 9 genitalia of paratype
shown in Fig. 11.

26(l-4):l-288, 1988

217

Figs. 23-28. Cryphia species, male genitalia. 23, C. viridata, San Diego, California,
USNM slide 52,177; 24, aedeagus of same specimen; 25, C. jucun-
della, holotype; 26, aedeagus of same specimen; 27, C. cyanympha,
paratype, McClellanville, S. C., USNM slide 52,101 ; 28, aedeagus of
same specimen.

218

J.Res.Lepid.

Figs. 29-32. Acronicta species, 9 genitalia, and Agriopodes fallax, 6 genitalia.

29, A. sinescripta, paratype, Gainesville, Florida, USNM slide 52,1 16;

30, A. perblanda, holotype; 31, A. fallax, Halifax County, Nova
Scotia, USNM slide 52,106; 32, aedeagus of same specimen.

Journal of Research on the Lepidoptera

26(l-4):219-224, 1988

The identity of Sphinx brunnus Cramer and the
taxonomic position of Acharia Huebner (Lepidoptera:
Limacodidae)

Vitor 0. Becker1

Centro de Pesquisa Agropecuaria dos Cerrados, Caixa postal 700023, 73300-Planaltina,
DF, Brasil

and

Scott E. Miller2

Bishop Museum, Box 19000-A, Honolulu, Hawaii 96817

Abstract. The identity of Sphinx brunnus Cramer is established and
Acharia Huebner is found to be a senior synonym of Sibine Herrich-
Schaeffer. Taxa previously included in Sibine are listed. A lectotype for
Acharia brunna is designated and illustrated. Sibine zellans Dyar and
S. berthans Dyar are recognized as new junior synonyms of Acharia
brunna.

Introduction

The identity of Sphinx brunnus Cramer has not been recognized since
its description in 1777. Consequently, the placement of the genus
Acharia Huebner has been uncertain since its publication. Huebner
([1819]) included Acharia in his ’’Stirp III. Glaucopen, Glaucopes” as
”Coitus[= genus] 2”, between ’’Coitus 1. Aclytia Huebner” and ’’Coitus 3.
Macrocneme Huebner”, both currently placed in the Ctenuchinae
(Arctiidae). Kirby (1892: 166) included Acharia in the Charadrinae
[= Ctenuchinae], after Teucer Kirby [ =Telioneura Felder], and de¬
signated brunnea [a misspelling] as the type species, as Huebner
originally also assigned another species, Sphinx coras Cramer, to the
genus (corns is currently considered a synonym oiPhobetron hipparchia
(Cramer), also Limacodidae). Hampson (1898-1920), however, did not
mention either Acharia or brunnus in his monographs on the Syn-
tomidae [= Ctenuchinae] and Arctiidae. Therefore, Acharia brunna has
remained unplaced until Fletcher and Nye (1982: 2) recently included it
in the Limacodidae, as suggested to them by Becker.

These identity problems resulted from the inaccuracy of Cramer’s
figure and the unavailability of the type specimens to previous authors.
Cramer’s illustration (Fig. 1) depicts a uniformly brown-colored moth
with falcate hindwings. No entomologist has ever located a specimen
that matches the figure.

Sphinx brunnus was described from an unspecified number of speci-

220

J.Res.Lepid.

mens from Surinam originally in the C. van Lennep collection. This
collection passed on to Felder, then to Rothschild, and finally to the
British Museum (Natural History) (BMNH). A specimen, presumably
belonging to the type series, has been found at the BMNH. We examined
this specimen (Fig. 2), have figured its genitalia (Fig. 4), and designate
it here as the lectotype. The specimen matches Cramer’s illustration
very well, except for the transluscent areas along the hind wing termen
(Fig. 2). There are two reasons why no subsequent specimen has been
found to match either Cramer’s illustration or the type specimen. First,
none of the plain brown specimens in collections have falcate hindwings

Fig. 1. Reproduction of figure 147c from Cramer, 1777.

Fig. 2. Lectotype male of Acharia brunna (BMNFI) (forewing length about 1 4 mm).
Fig. 3. Flolotype male of Sibine zellans , from Para [= Belem], Brazil (USNM
40684).

26(l-4):l-288, 1988

221

as shown by Cramer (Fig. 1). Secondly, the type specimen was painted to
match the figure, and the paint has partially pealed off (especially along
the hindwing margins), giving the impression that the species belongs
to the auromacula group of the genus Sibine herrich-Schaeffer, which
have the hindwings partially transluscent and with a falcate ter men.
However, none of the species belonging to the auromacula group
0 auromacula Schaus, intensa Dyar, barbara Dyar, sibinides Dyar, and
blanda Dyar) have plain brown forewings. They all have two well
defined white dots, a round one below the cell, and an elongate one
beyond the cell apex, crossing R3+R4 and R5.

In the extensive series of Sibine in the National Museum of Natural
History (USNM) and the Becker collection, we have found only two
males which have plain dark brown wings that match the lectotype of
Acharia brunna. These are the holotype of S. zellans Dyar (Fig. 3) (Para
[=Belem], Brazil, [no date], A.M. Moss) and a syntype of S. berthans
Dyar (Villa Rica [=Villarrica], Paraguay, March 1926, F. Schade). S.
berthans was described from two syntypes, a male and a female from the
same locality, both in USNM. We hereby designate the male as
lectotype. Without revision of the genus, we cannot be sure that the
female paralectotype is conspecific with the lectotype, and do not treat it
further here.

The external margin of the hindwings is straight in Sibine zellans ,
but slightly convex in S. berthans. However, the genitalia of both are
identical and match perfectly those of Acharia brunna. The only differ¬
ences mentioned in the original descriptions (Dyar, 1927: 547) were that
berthans has the “wings rather less pointed” and the “spines of the penis
[=cornuti of aedoeagus] are finer and more numerous than in zellans.”
We are convinced that they are all conspecific. We regard the differences
in wing shape as individual or local variation, and the slight differences
in comuti as insignificant. Although it is widely distributed (Surinam,
through the Amazon Basin, into Paraguay), apparently brunna does not
readily come to light and is therefore rare in collections.

Dyar (1935) reviewed the species of Acharia (as Sibine ), and gave a
key with poor quality illustrations of wing surfaces. The genus, which
includes several agriculturally important species, needs revision. With¬
out a modern revision of all the species, it is not possible to assess the
significance of the morphological variation discussed above!’

Checklist

Except for berthans and zellans , which are newly synonymized under brunna ,
the taxa are listed as they presently stand in the literature. We expect that
examination of type material of the older names, as well as newly accumulated
material in collections, will result in many taxonomic changes. Forbes (1942)
has suggested several taxonomic changes, but they should not be adopted
without examination of the relevant types.

Acharia Huebner, 118191

222

J.Res.Lepid.

Fig. 4-6. Male genitalia of lectotype oiAcharia brunna : 4: ventral view (manica not
shown); 5: lateral view, 6: aedoeagus, with vesica everted.

Sibine Herrich-Schaeffer, 1855, n. syn.

Empretia Clemens, 1860, n. syn.

Eupretia Walker, 1865, misspelling
Eupalia Walker, 1866, n. syn.

Episibine Dyar, 1898, n. syn.
affinis (Moeschler, 1883) (Sibine) n. comb,
alicians (Dyar, 1935) (Sibine) n. comb, [as subspecies of nitens]
apicalis (Dyar, 1900) (Sibine) n. comb,
auromacula (Schaus, 1896) (Sibine) n. comb,
ausa (Dyar, 1935) (Sibine) n. comb, [as subspecies of stimulea]
barbara (Dyar, 1905) (Sibine) n. comb,
blanda (Dyar, 1935) (Sibine) n. comb,
bonaerensis (Berg, 1878) (Streblota) n. comb,
brunna (Cramer, 1777) (Sphinx) n. comb.

berthans (Dyar, 1927) (Sibine) n. syn., n. comb,
zellans (Dyar, 1927) (Sibine) n. syn., n. comb,
clarans (Dyar, 1927) (Sibine) n. comb,
didactica (Dyar, 1927) (Sibine) n. comb.

differentiata (Bryk, 1953) (Sibine) n. comb, [as subspecies of joyceansl

26(l-4):l-288, 1988

223

dorans (Dyar, 1927) (Sibine) n. comb.

eucleides (Dyar, 1905) (Sibine) n. comb.

extensa (Schaus, 1896) (Sibine) n. comb.

francescans (Dyar, 1927) (Sibine) n. comb.

fusca (Stoll, 1780) (Phalaena) n. comb.

gertrudans (Dyar, 1927) (Sibine) n. comb.

geyeri (Fletcher, 1982) repl. name (Sibine) n. comb.

nesea (Geyer, [1833]) (Streblote) n. comb,
giseldans (Dyar, 1927) (Sibine) n. comb,
helenans (Dyar, 1927) (Sibine) n. comb,
horrida (Dyar, 1905) (Sibine) n. comb,
hyperoche (Dognin, 1914) (Sibine) n. comb,
intensa (Dyar, 1905) (Episibine) n. comb,
iolans (Dyar, 1927) (Sibine) n. comb,
joyceans (Dyar, 1927) (Sibine) n. comb,
laberia (Dyar, 1935) (Sibine) n. comb,
laurans (Dyar, 1927) (Sibine) n. comb,
lophostigma (Dognin, 1910) (Sibine) n. comb.

megasomoides (Walker, 1866) (Eupalia) n. comb, [currently subspecies of
rufescens]

minuscula (Bryk, 1953) (Sibine) n. comb, [as subspecies of auromacula]
modesta (Cramer, 1777) (Phalaena) n. comb,
nesea (Stoll, 1780) (Phalaena) n. comb.

vidua (Sepp, [1848]) (Phalaena) n. comb,
fumosa (Walker, 1855) (Nyssia) n. comb,
nitens (Dyar, 1905) (Sibine) n. comb,
norans (Dyar, 1927) (Sibine) n. comb,
ophelians (Dyar, 1927) (Sibine) n. comb,
pauper (Dyar, 1918) (Sibine) n. comb,
permessa (Dyar, 1918) (Sibine) n. comb,
priscillans (Dyar, 1927) (Sibine) n. comb,
quadratilla (Dyar, 1935) (Sibine) n. comb,
quellans (Dyar, 1927) (Sibine) n. combv
reletiva (Dyar, 1927) (Sibine) n. comb,
rollans (Dyar, 1927) (Sibine) n. comb,
rufescens (Walker, 1855) (Nyssia) n. comb.

determinata (Walker, 1865) (Nyssia) n. comb,
plora (Schaus, 1896) (Sibine) n. comb,
pallescens (Dognin, 1901) (Sibine) n. comb,
sabis (Dyar, 1935) (Sibine) n. comb,
sarans (Dyar, 1927) (Sibine) n. comb,
sibinides (Dyar, 1905) (Episibine) n. comb,
stimulea (Clemens, 1860) (Empretia) n. comb.

ephippiatus (Harris, 1869) (Limacodes) n. comb,
subalbicans (Dyar, 1935) (Sibine) n. comb,
tontineans (Dyar, 1927) (Sibine) n. comb,
trimacula (Sepp, [1848]) (Phalaena) n. comb,
varia (Walker, 1855) (Nyssia) n. comb,
violans (Dyar, 1927) (Sibine) n. comb
ximenans (Dyar, 1927) (Sibine) n. comb.

224

J.Res.Lepid.

Acknowledgements. We thank the curators of the collections consulted,
especially Allan Watson of the BMNH. The photographs were taken by Victor
Krantz of the Smithsonian Institution. Research for this paper was done while
both authors had Smithsonian fellowships. D.R. Davis, M.E. Epstein and N.L.
Evenhuis reviewed the manuscript.

Literature Cited

CRAMER, P., 1775-1780. De Uitlandsche Kapellen voorkomende in de drie
Waerelddeelen Asia, Africa en America. Amsterdam, S.J. Baalde. Volume 2,
152 pp., pis. 97-192.

DYAR, H.G., 1898. Notes on certain South American Cochlidiidae and allied
families. J. New York Ent. Soc. 6:231-239.

DYAR, H.G. 1927. New species of American Lepidoptera of the families Limaco-
didae and Dalceridae. J. Wash. Acad. Sci. 17:544-551.

DYAR, H.G., 1935. Limacodidae. in A. Seitz (ed.), Die Gross-Schmetterlinge der
Erde. Stuttgart, Alfred Kernen. Volume 6, pp. 1104-1 139.

FLETCHER, D.S. & I.W.B. NYE. 1982. in I.W.B. Nye (ed.), The generic names of moths
of the world, volume 4. London, British Museum (Natural History), xiv +
192 pp.

FORBES, W.T.M., 1942. The Lepidoptera of Barro Colorado Island, Panama. No. 2.
Bull. Mus. Comp. Zool. 90:265-406.

HAMPSON, G.F., 1898. Catalogue of the Lepidoptera Phalaenae. Syntomidae.

London, British Museum (Natural History). Volume 1. 559 pp.

HAMPSON, G.F., 1900. Catalogue of the Lepidoptera Phalaenae. Arctiadae [sic].

London, British Museum (Natural History). Volume 2. 589 pp.

HAMPSON, G.F., 1901. Catalogue of the Lepidoptera Phalaenae. Arctiadae [sic]
and Agaristidae. London, British Museum (Natural History). Volume 3. 690

pp.

HAMPSON, G.F., 1914. Catalogue of the Lepidoptera Phalaenae. Amatidae and
Arctiadae [sic]. Supplement Volume 1. London, British Museum (Natural
History). 858 pp.

HAMPSON, G.F., 1920. Catalogue of the Lepidoptera Phalaenae. Lithosiadae and
Phalaenoididae. Supplement Volume 2. London, British Museum (Natural
History). 619 pp.

HUEBNER, J., 1816-[1825]. Verzeichniss bekannter Schmettlinge. Augsburg. 431

pp.

KIRBY, W.F., 1892. A synoptic catalogue of Lepidoptera Heterocera. (Moths.) Vol.
I. Sphinges and bombyces. London, Gurney and Jackson, xiii + 951 pp.

Journal of Research on the Lepidoptera

26(l-4):225-235, 1988

The Life History of Hemileuca magnifica (Saturniidae)
With Notes on Hemileuca hera marcata

Stephen E. Stone1

18102 East Oxford Drive, Aurora, Colorado 80013

DeanE. Swift

Box B, Jaroso, Colorado 81138
and

Richard S. Peigler2

303 Shannon Drive, Greenville, South Carolina 29615

Abstract. Hemileuca magnifica is a diurnal saturniid inhabiting
Climax Sagebrush Associations in southern Colorado and northern
New Mexico. The larval host is exclusively Artemisia tridentata in
Costilla County, Colorado and northern Taos County, New Mexico. The
Albuquerque, Bernalillo County, New Mexico colony is found on A.
filifolia. The species has a two-year life cycle, overwintering first as ova
and the second winter as pupae. The ova are deposited in rings typical of
Hemileuca species. The young larvae feed gregariously and as they
mature, disperse widely. Observations of insect parasitism and avian
predation are reported.

Introduction

Until recently, Hemileuca magnifica (Rotger) (Saturniidae) was re¬
presented in private and museum collections by very few specimens
(Ferguson 1971) and virtually nothing was known of its life history. The
present paper provides new information on habitat, behavior, and
biology. Hemileuca magnifica was considered a subspecies of H. hera by
the original author and Ferguson (1971), but more recently the moth
was given full specific rank (Ferguson 1983).

Discussion

Habitat

Hemileuca magnifica is a cryptically colored black, white and orange
saturniid of moderate size inhabiting the Upper Sonoran Life Zone in

1 Natural Resource Specialist, U. S. Department of the Interior, National Park Service,
Denver, Colorado

2Museum Associate in Entomology, Los Angeles County Museum of Natural History

226

J.Res.Lepid.

southern Colorado and northern New Mexico. The Upper Sonoran Life
Zone, which includes the Sagebrush-Piny on- Juniper Belt, extends from
at least Colorado and Utah south to central Arizona and New Mexico.
The altitudinal range of this zone is from about 1,370 to 2,280 m and its
limits vary with and are influenced by differences in exposure, soil, and
moisture conditions (Gregg 1963, Patraw and Janish 1977).

Sagebrush, {Artemisia tridentata Nuttall: Compositae) is found at
elevations between 1,820 and 2,500 m and ranges through western
North America from South Dakota to British Columbia and south to
New Mexico, northern Arizona and Baja California and is, therefore,
not restricted to the Upper Sonoran Life Zone. The plant can cover vast
areas to the almost total exclusion of other plant species and can grow
well over two m in height in conditions of favorable soils and abundant
precipitation, but one m or less is normal (Coyle and Roberts 1975,
Elmore and Janish 1976, Gregg 1963, Kearney and Peebles 1942,
Patraw and Janish 1977, Rydberg 1906, Tidestrom 1925, Wiggins 1980,
Wooton and Standley 1915).

Artemisia tridentata is the dominant shrub of the Climax Sagebrush
Association southern San Luis Valley, Costilla County, Colorado where
most of the work reported in this study took place. Occasional associated
plant species within this study area include rabbitbrush {Chrysotham-
nus sp.: Compositae) and saltbush {Atriple x sp.: Chenopodiaceae) (Lamb
1975). The Climax Sagebrush Association at Albuquerque, New Mexico
is dominated by Artemisia filifolia Torrington.

Seasonal temperatures in the study area range from 35°C during July
and August to extremes of — 35°C and lower in December and January.
Annual precipitation averages less than 20 cm (Colorado Climate
Center, pers. comm.) with occasional summer thunderstorms. The
majority of moisture, however, is received as winter snows.

Range

Hemileuca magnifica is associated with Climax Sagebrush Associa¬
tions (Fig. 1) at elevations between 2,000 and 2,500 m. In Costilla
County, Colorado the known range of H. magnifica extends from just
north of Fort Garland south through Mesita (type-locality) and Jaroso.
It extends further south in New Mexico through Taos County to
Albuquerque, Bernalillo County and then west to just northeast of
Pintado, McKinley County (Jim Coleman, pers. comm., and Ferguson
1971). Thus, H. magnifica inhabits a limited, roughly triangular area on
the border of northern and northwestern New Mexico and south central
Colorado. However, Townsend (1893) stated that on 25. VII. 1892 he
observed a number of large brownish and blackish Hemileuca larvae
feeding on A. filifolia east of Navajo Springs, Arizona. It is reasonable to
believe the species Townsend observed were H. hera hera (Harris) or H.
magnifica. Navajo Springs, Apache County, Arizona shares the same
Great Basin influence as Pintado, McKinley County, New Mexico
(Martin and Hutchins 1980), which is the westernmost known locality

26(l-4):l-288, 1988

227

Fig. 1. Typical climax Sagebrush habitat which supports Hemileuca
megnifica.

for H. magnified. As discussed below, populations of H. magnified and H.
h. hera are not known to be sympatric nor parapatric.

Biology

The Hemileuca hera group is composed of three taxa, H. h. hera, H. h.
marcata (Neumoegen), and H. magnifica , which have long been asso¬
ciated with sagebrush communities (Holland 1903, McFarland 1974,
Packard 1914, Ferguson 1971, Furniss and Carolin 1977). Recently, H.
h. hera in California has been shown to prefer A. tridentata as host,
although when larval populations are dense, various species of Lupinus
(Mimosoideae) and Eriogonum (Polygonaceae) are also used (Tuskes
1984). The population of H. magnifica in the vicinity of Albuquerque,
New Mexico feeds only on A. filifolia (Jim Coleman, pers. comm.) while
those of Taos County, New Mexico and Costilla County, Colorado feed
exclusively on A. tridentata as does the colony of H. h. marcata in
Klamath County, Oregon (Melvin Parker, pers. comm.).

It is noteworthy that H. magnifica is not distributed uniformly
throughout all suitable range, but seems to be highly localized in
concentrated populations, as is H. h. hera in California (Tuskes, pers.
comm.). The preferred habitat seems to be areas where the host is
growing in sandy-loamy-clay soils and is fairly tall, dense and lush.
Coarse gravelly areas, where the host plant is shorter and less fully
developed, do not appear to support populations of H. magnifica. Even
where lush and stunted sagebrush communities are adjacent, the
stunted areas are devoid of H. magnifica. Patrolling (mate seeking)

228 J.Res.Lepid.

males stay within the more lush habitats and larvae are found only in
such areas.

Adults in J aroso are on the wing from early August to late September
with the main flight in late August. Males fly from approximately 1100
to 1500 hours (Mountain Daylight Time). Ovipositing females may be
on the wing until 1800 hours. Adults hatch from 0900 to 1200 hours
emerging from the pupa and climbing a short way into nearby vegeta¬
tion, usually A. tridentata , where they expand and harden their wings
which takes about an hour. As soon as males harden their wings they fly
off in search of a mate. It is to the advantage of male saturniids to cover
as much suitable habitat as quickly as possible in their search for
potential mates, which is why they are such wide ranging and active
fliers (Janzen 1984). It would appear males flying in late afternoon are
those that have been unsuccessful in locating a virgin female earlier in
the day and are still actively seeking a mate, or have mated earlier in
the day and are seeking another mate.

Females start emitting pheromone as soon as their wings are hardened
and usually do not fly until after copulation. As discussed by Tuskes
(1984) for Hemileuca in general, newly emerged females usually remain
on lower stems of the plant while emitting pheromone. Mating requires
one to three hours. After separation, the females seldom fly to deposit
the first egg ring but, instead, crawl up the plant to form a ring of cream
colored eggs on an apical shoot of the host. The rings are typical of the
genus Hemileuca. The first ring contains ca. 50 to 75 ova. During the
next few days, the female deposits several additional egg rings, but each
contains a smaller number of 30 to 35 ova. The female may deposit 100
to 200 ova during her life of five to seven days. Ova normally overwinter
the first year.

Although theoretically equal in numbers to males, females are
uncommonly encountered. It would appear that females which have
mated need not range far seeking suitable hosts for oviposition. After
ovipositing, which takes anywhere from 30 minutes to more than an
hour, the female may or may not immediately fly again in search for
another host. Our observations indicate the female may perch and rest
for several hours or until the next day depending on her burden of ova,
age, strength and condition.

Females mate once, males may mate several times as opportunity
provides. Males searching for pheromone plumes emitted by virgin
females search exactly as described by Collins and Tuskes (1979),
Collins and Weast (1961), and Janzen (1984). The males fly corsswind at
rapid speed searching for pheromone plumes emitted by females. Upon
encountering a plume they follow it upwind to the female by an ever
decreasing series of lateral oscillations. When males are from one to two
m from the female, flight slows and is more direct. When males are less
than one m from the female, they fly very slowly and frequently hover
within a few cm of the female before landing.

Our observations indicate the male crosswind flight often exceeds

26(l-4):l-288, 1988

229

Fig. 2. Adult male H. magnifica
perched for the night on
Artamisia. P.

400 m. If no plume is encountered the male drops downwind several
hundred m and begins another crosswind search pattern. Patrolling
males generally cruise widely just over the tops of the sagebrush and
seem to prefer draws, washes and saddles, perhaps using these topo¬
graphic features as natural fly ways. A single caged pheromone-emitting
female may attract hundreds of males within a few hours.

After the daily flight is over, adults, especially males, tend to perch on
the high exposed tips of plants where they spend the night (Fig. 2).
Tuskes (pers. comm.) observed similar behavior in populations of H.
eglanterina (Boisduval), H. nuttalli (Strecker), and H. h. hera in east
Sierra country in California. Approximately one hour after sunrise of
the next morning they position themselves in an easterly, southeasterly
or southern orientation with the dorsal surface fully exposed to the sun
in order to sufficiently warm to begin the day’s activities.

In Jaroso, Colorado ova hatch from mid to late June. In Albuquerque,
New Mexico ova hatch from early to mid May (Jim Coleman, pers.
comm.). Larvae bear urticating spines and are gregarious until mid to
late third instar when they disperse widely. More mature larvae
generally prefer apical, succulent growth. During the day these older
larvae tend to gravitate to this exposed growth and are easily seen (Fig.
3). Larvae were also observed in shadier portions of the host, probably
shifting between these two areas as a means of thermoregulation (see
Capinera et al. 1980). During the night larvae retreat to the interior of
the host where their color patterns provide excellent camouflage.

230

J.Res.Lepid.

Fig. 3. Last instar larva of H.
magnifica in a natural day¬
time feeding position.

Larvae reared indoors on cut host do poorly. Ferguson (1971) stated
that H. h. hero larvae have successfully been reared in screened
cylinders placed in sunlight for an hour or so each day. Hemileuca
magnifica larvae can be reared indoors but require spacious airy
conditions and the radiation of a “Gro-Light” (Tuskes, pers. comm.).
They do best if reared outdoors on living hosts, but will mature indoors
on cut food if it is fresh. The later instars cannot stand crowding and will
die if crowded. These findings are consistent with those of McFarland
(1974) and Smith (1974).

Pupation is a few cm under the soil surface without any kind of cocoon.
The pupae normally overwinter for a single year, but individuals
holding over for two or more years are known.

Adult Coloration

The black, white and orange colors of H. magnifica adults are an
excellent camouflage while they are perched deep inside sagebrush
plants, especially during the most vulnerable time of expanding and
hardening their wings. If the moths are distasteful to birds as Ferguson
(1971) suggests, then moths of this genus would be interesting inas¬
much as they are cryptic while resting yet aposematic in flight. The
black and white of wings blend well with the light and shadows within
the host and help to break up the adult outline. The orange tufts and
rings on the body are a very close match to an orange lichen ( Xanthoria
polycarpa (Ehrh.) Olir.: Teloshistaceae) found commonly on sagebrush
stem and trunk bark, especially in the Jaroso area. The evolution of

26(l-4):l-288, 1988

231

highly developed orange patches and rings on adult H. magnified may
be directly related to the lichen. While in repose within the sagebrush
plant the orange color further helps to conceal the adults and may
contribute to reduced predation pressure.

Parasitoids and Predators

A group of 50 H. magnifica larvae ranging in age from 3rd instar to
mature were collected on A. tridentata 24. VIII. 1985 at Jaroso,
Colorado. The following parasitoids were found.

Diptera. Exorista sp. (Tachinidae); medium sized flies, of which
several specimens were reared. Chetogena sp. (Tachinidae); smaller
sized flies, of which several specimens were also obtained. The species
are not presently identifiable, according to N. E. Woodley who deter¬
mined them. Arnaud (1978) cited no parasitic tachinid records for H.
magnifica.

Hymenoptera. Microdontomerus fumipennis Crawford (Torymidae),
determined by E. E. Grissell. These small wasps with prominent
ovipositors are known to attack several hosts (Krombein et al. 1979,
Peigler 1985). Numerous wasps emerged from a single host.

Cotesia electrae (Viereck) (Braconidae), determined by P. M. Marsh,
formerly classified in the genus Apanteles. ( Apanteles electrae has been
recorded as a parasitoid of a Hemileuca species in Texas [Peigler 1985],
Agapema galbina [Collins and Weast 1961], H. h. hera and other
saturniids, especially Hemileucinae [Krombein et al. 1979] in western
North America.)

When the C. electrae larvae are mature, they exit the host and spin
silken cocoons directly on the host body surface (Borror and DeLong 1970
[as Apanteles ]). A population of 10 to 20 parasitic C. electrae larvae per
parasitized H. magnifica larva host is common.

Voucher material of Chetogena and Microdontomerus were deposited
in the U. S. National Museum of Natural History, and Exorista and
Cotesia in the Los Angeles County Museum of Natural History.

Jim Coleman (pers. comm.) has reported robins ( Turdus migratorius
Ridgeway) occasionally feeding on mature H. magnifica larvae in the
Albuquerque colony.

Larva Description

A description of a mature H. magnifica larva (Fig. 4) follows based on
living material from Jaroso, Colorado. There usually are six instars
although seven is not uncommon (see Lemaire 1979: 231, footnote).
DESCRIPTION OF MATURE LARVA OF HEMILEUCA MAGNIFICA
HEAD: Lustrous black to reddish brown with abundant white secondary
setae; diameter 4. 5-5.0 mm. BODY: Length (including head) 60.0-72.0 mm:
width 9.0-1 1.0 mm. Ground color dark brown approaching black, rarely tan to
light brown. Body with four pairs of off-white lines which may be reduced to a
series of inconspicuous dashes. Subspiracular line most developed, broad, and
quite pronounced, running through subspiracular and subdorsal scoli and

232

J.Res.Lepid.

Fig. 4. Last instar larva of H.
magnifies.

variably developed, but always less than subspiracular line. Subdorsal line
between subdorsal and dorsal scoli, and significantly reduced or absent.
Dorsal line between dorsal (rosette) scoli, reduced to narrow white dashes or
dots, if present. Ventral scoli present on pro-, meso-, and metathorax and
abdominal segments one, two, seven and nine. Dorsal scoli present as two
rows of short dull yellow rosettes with black tips, on metathorax and next
seven abdominal segments; on abdominal segment eight a median rosette; on
pro- and mesothoracic segments rosettes containing larger black scoli re¬
sembling subdorsal and spiracular ones. Subdorsal and lateral scoli long,
black, and yellow tips on branches. Numerous white secondary setae over
entire body surface. True and prolegs black, feet white on inner side. Spiracles
black. Abdominal and especially thoracic segments with white dots or
markings ventrally, anterior to each proleg or true leg.

Since no descriptions of the early stages of H. hera marcata have been
published, we offer the following description based on a mature larva in
alcohol. The specimen was collected as a first instar larva on A.
tridentata at Klamath Falls, Klamath County, Oregon by Melvin
Parker and reared to maturity on A. tridentata by Dean Swift in Jaroso,
Costilla County, Colorado.

DESCRIPTION OF MATURE LARVA OF HEMILEUCA HERA MARCATA
HEAD: Lustrous black with abundant white secondary setae; diameter 4.0
mm. BODY: Length (including head) 55.0 mm; width 7. 0-9.0 mm. Ground
color purplish brown. Body with four pairs of off-white lines. Subspiracular
line most developed, broad, running through subspiracular scoli, with dark
evaginations from ventral edge. Lateral line between subspiracular and
subdorsal scoli, closer to latter, prominent and unbroken, but only half as
thick as subspiracular line. Subdorsal line between two aforementioned lines

26(1-4):1-288,1988

233

in thickness, unbroken. Dorsal line barely visible, except on thoracic seg¬
ments, running between dorsal scoli. Ventral scoli present on pro-, meso-, and
metathorax and abdominal segments one, two, seven and nine. Dorsal scoli
present as two rows of short dull yellow rosettes with black tips, on
metathorax and next seven abdominal segments; on abdominal segment
eight a median rosette; on pro- and mesothoracic segments (and three on
abdominal segment nine) rosettes containing larger black scoli resembling
subdorsal and lateral ones. Subdorsal and lateral scoli long, black, with
yellow tips on branches. Numerous white secondary setae over entire body
surface. True legs black, prolegs white with dark patches on outer sides.
Spiracles black. Ventral surface predominantly off-white, with a few small
dark median patches on each segment.

Preserved mature larvae of H. magnified and H. h. marcata have been
deposited in the Los Angeles County Museum of Natural History.

Diagnosis

Based on our above observations of mature larvae of H. magnified and
H. h. marcata and the description of H. h. hera by Tuskes (1984) plus a
color slide of H. h. hera which he kindly sent to us, we are able to make
the following brief comparisons. The larva of H. h. hera has a slender yet
conspicuous subspiracular stripe, but less developed than the lateral
one; in H. h. marcata and H. magnified this ventralmost stripe seems to be
widest and the most conspicuous. The dorsal rosette scoli of H. h. hera
appear to be dull yellow, like the ones in H. magnified and H. h. marcata.
The ventral surfaces of H. h. hera and H. magnified are predominantly
blackish, whereas H. h. marcata is mainly whitish throughout the
ventral area. As would be expected due to the size of the imagines, the
larva of H. magnifica is larger than those of the other two.

Comparison of H. h. hera,H. h. marcata and H. magnifica convinces us
that these three taxa form a compact group, or a subgroup within the
eglanterina group of Tuskes (1984, Table 1). The adults of these three
differ from their nearest relatives (viz H. nuttalli and H. eglanterina) by
lacking yellow or orange scales on the wings. The hera subgroup also
differs from the other members of the eglanterina group by being almost
exclusively feeders on sagebrush.

As mentioned above, Ferguson (1983) elevated H. magnifica to full
specific status; the same author (Ferguson 1971) earlier speculated that
both H. magnifica and H. h. marcata could be full species distinct from
H. h. hera. Since subspecies by definition should have ranges that
overlap or at least are in contact, and show blending of characters, we
consider it significant that no such zones are known for H. h. hera and H.
magnifica. These two species are allopatric as far as known. Any pair of
full species could be either allopatric or sympatric, but if two taxa are
subspecies, they should show blending in a parapatric pattern. As
discussed earlier, the status of H. h. hera and H. h. marcata is less
certain.

Presently the response of each species males to calling females of the
other taxon is unknown. Tuskes (1984) discussed a blend zone of H. h.

234

J.Res.Lepid.

hera and H. h marcata and described what appears to be intergrade
adults. He referred to H. h. marcata as a “form” but did not state
whether the H. h. hera population in California contacts the H. h.
marcata in Oregon, or if the two populations are disjunct. The “ H . h.
marcata population in Oregon, or if the two populations are disjunct.
The “H. h. marcata ” he figured from California is not as extreme in
reduction of black markings as in specimens of true H. h. marcata.
backcrosses and sibmatings could be reared. Such hybrid work may
answer such questions as the nature and degree of phenotypic variation
of hybrid individuals, ova fertility and fecundity, survival and viability
of hybrid individuals throughout development (which is directly related
to phylogenetic divergence), and isolating mechanisms (see Peigler and
Williams 1984).

Acknowledgements. The authors extend special thanks to Claude Lemaire for
kindly reviewing the manuscript of this paper and for making many beneficial
suggestions and encouraging comments. We also thank Paul M. Tuskes for
giving us his information regarding Hemileuca hera hera, H. nuttalli, H.
eglanterina and H. magnifica and for reviewing the manuscript of this paper,
offering many helpful suggestions. Michael J. Smith also reviewed the manu¬
script and offered helpful suggestions. Jimmie C. Coleman and Melvin Parker
supplied information and material. Lawrence F. Van Horn offered useful
editorial suggestions. Larry Schlichenmayer provided foodplant. N. E. Woodley,
E. E. Grissell, and P. M. Marsh of the Systematic Entomology Laboratory, U. S.
Department of Agriculture, identified parasitoids. William A. Weber identified
the lichen. The Colorado Climate Center provided meteorological data. Sincere
thanks must be expressed to Becky Debs for putting the drafts on the computer.
The senior author expresses deep thanks to his wife Karen and daughter
Shaunna.

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Journal of Research on the Lepidoptera

26(l-4):236-239, 1988

Opinion. Opinion is intended to promote communication between
lepidopterists resulting from the content of speculative papers. Comments,
viewpoints and suggestions on any issues of lepidopterology may be
included. Contributions should be as concise as possible and may include
data. Reference should be limited to work basic to the topic.

Are We Studying Our Endangered Butterflies to Death?

Dennis D. Murphy

Center for Conservation Biology, Department of Biological Sciences, Stanford University,
Stanford, California 94305

A number of studies suggest that handling has adverse effects on
butterflies. Singer and Wedlake (1981), for instance, were able to
recapture only 2% of the Graphium sarpedon that had been captured,
marked, and released on a riverside beach in Sarawak. Subsequently,
they were able to mark the swallowtails without handling.

“The procedure employed was to crawl towards the insects extremely
slowly, taking several minutes to reach them. On the first occasion
that this was tried, the marker pen was held in an outstretched hand,
but it was found to have dried out by the time it reached the feeding
butterflies. Subsequently we wormed our way up the beach on our
stomachs with both hands outstretched, so that we could delay
uncapping the pen until after its arrival among the butterflies.”
After using these methods, Singer and Wedlake found that recapture
rates soared to 21%. Morton (1984) in a temperate zone study likewise
found that marking the lycaenids Polyommatus icarus and Lysandra
coridon , without handling, also resulted in increased recapture rates.

Direct impact of handling on butterfly behavior and longevity,
however, has been difficult to document. Powell (pers. com.) has observed
that freshly emerged adults of the endangered Lange’s metalmark
butterfly ( Apodemia mormo langei ) assume their nocturnal resting
position when marked immediately after emerging from their pupae.
Some individuals retain this position for extended periods which could
subject them to higher rates of predation. Reid (1985 and pers. comm.)
found that handling of the endangered mission blue butterfly ( Plebejus
icarioides missionensis ) results in about 10% mortality. He has for that
reason discontinued the use of mark-recapture techniques in long-term
studies on San Bruno Mountain. Many field workers suggest that mark-
recapture methods probably should never be employed with small¬
winged, swift-flying species, for example the threatened Pawnee

26(l-4):l-288, 1988

237

montane skipper ( Hesperia leonardus montana) presently under study
in Colorado.

Despite this mounting evidence that many butterflies exhibit greatly
disturbed behavior or suffer physical damage from marking and recap¬
ture, mark-recapture remains one of the initial steps in nearly every
conservation effort focussing on butterflies. This is especially ill-advised
because the data obtained from such studies rarely are useful in the
protection of endangered species. For instance, mark-recapture studies
and intensive statistical analysis have revealed few differences in
population parameters between the gravely endangered bay checker-
spot butterfly ( Euphydryas editha bayensis) and the widespread chalce-
don checkerspot ( Euphydryas chalcedona chalcedona ) on Jasper Ridge
Biological Preserve at Stanford University (Murphy, et al. 1986). The
study suggested that “new approaches both to mark-recapture analysis
and the study of endangered species are needed to generate more useful
information concerning the conservation status of invertebrate popula¬
tions.” Murphy et al. further conclude that methodologies which mini¬
mize physical damage to butterflies and their habitats should be
emphasized when studying endangered or threatened species.

The limits of distributions and relative densities of butterflies within
their habitats constitute the critical information usually sought by con¬
servation biologists. This information nearly always can be ascertained
through simple observation and use of a low impact “sampling” tech¬
nique such as that of Pollard (1977), which involves repeatedly walking
transects while recording the number of butterflies observed. The
purported advantage of mark-recapture techniques over transect
observations lies in the estimate of absolute (rather than relative) popu¬
lation sizes which may be generated from field data. But this advantage
usually is moot since endangered butterflies are virtually always
restricted to small habitat patches where population sizes are small.
Little practical value is gained from establishing that populations that
are obviously small are indeed small.

Two additional problems are prevalent when mark-recapture tech¬
niques are used to derive population size estimates for conservation
studies. First, the variances associated with these estimates are often as
large as the population estimates themselves. The only way to reduce
that variance is to increase the handling, marking, and recapturing of
the endangered butterflies. Second, exact population size is relatively
unimportant when developing conservation agendas. For example, the
entire two acres of habitat owned by Chevron Oil, in Southern Cali¬
fornia, must be protected and maintained free of alien plant species to
ensure the persistence of the El Segundo blue butterfly, whether 100,
300, or 1,000 individuals are present in any given year.

If mark-recapture studies offer little guidance in conservation efforts,
why then are they allowed — and even promoted? The answer may rest
in the fact that most studies are commissioned in response to govern-

238

J. Res. Lepid.

ment regulations which require private land developers to assess the
potential impact of proposed habitat disturbance to endangered species.
The lead agency in these situations is either the United States Fish and
Wildlife Service or a state agency with a similar mandate for species
protection. Limited by tight budgets and small staffs emphasizing fish
and game resource management, these agencies often must seek advice
from outside “experts,” especially on matters pertaining to invertebrate
conservation. Many consultants take advantage of this situation to
promote more labor-intensive (read ‘more expensive’) field methodo¬
logies, such as mark-recapture studies, when simpler approaches are
adequate to achieve most conservation objectives.

A particularly egregious example is a recent study involving the
federally protected Smith’s blue butterfly ( Euphilotes enoptes smithi)
on Marina State Beach in Monterey County, California (Arnold, 1986).
This study proposed no new approach to the management of this
endangered butterfly, yet it potentially subjected a population to severe
impacts from handling. Nominally trained “students” were used to
mark more than 400 adult blue butterflies, which together subsequently
were handled another several hundred times. The stated objectives of
the study were “to 1) determine the present distribution and population
numbers of Smith’s blue, at Marina State Beach; 2) determine which
habitat areas of the dunes are important for mate location, foraging, and
larval development; 3) evaluate different census methods for their
appropriateness in future monitoring of Smith’s blue during and after
implementation of the revegetation program; and 4) provide manage¬
ment recommendations for Smith’s blue at Marina State Beach.”

As discussed above, the first and second goals largely could be met by
simple survey and transect procedures, without handling butterflies. To
use the most fragile, federally protected butterfly species as a lepidop-
teran guinea pig in pursuit of the third goal defies any sense of logic or
conscience. Certainly a non-endangered, taxonomically related species
could have been used to develop a low impact protocol for application to
an endangered species. The fourth goal underscores the lack of per¬
tinence of most mark-recapture studies to habitat management. The
key recommendations stated at the conclusion of the study were “to
control, and where feasible, eradicate alien flora that can outcompete
native vegetation such as the butterfly’s buckwheat foodplants” and to
“promote natural dune dynamics, which facilitates natural seeding
establishment by the native flora.” Not only were those recommenda¬
tions obvious from a cursory view of dunes blanketed with South
African iceplant, but just such a restoration project was already in
process. In other words, hundreds of endangered butterflies were
subjected to physical impairment and potential mortality associated
with handling for no good reason.

All this does not mean that no value whatsoever can be derived
from mark-recapture studies of endangered butterfly species. Interha-

26(l-4):l-288, 1988

239

bitat dispersal can be crucial to the overall persistence of butterflies
where adjacent populations exist or empty (but suitable) habitat
patches are available. Gall (1984), for instance, used mark-recapture to
demonstrate age-dependent dispersal in female Boloria acrocnema in
an isolated Rocky Mountain habitat patch. He speculated that late
season female dispersal may contribute to the colonization of new
habitat patches. But one must ask whether similar insights into the
population dynamics of species less physically hardy than Boloria
acrocnema warrant the potential damage from intensive handling. I
contend that they do not.

This paper is presented as a plea to government agencies, environ¬
mental consultants, and field biologists to restrict mark-recapture
studies to non-endangered species, or to endangered species proven
hardy enough to withstand human handling. In the latter circum¬
stances, mark-recapture should be employed only to answer specific
questions crucial to recovery strategies. The use of untrained field
workers in such studies should be strongly discouraged. The prepon¬
derance of evidence indicates that the handling of lycaenid butterflies,
in particular, usually results in the taking of individuals — an unlawful
act under most permits to handle endangered species and one which
should not be encouraged in the name of conservation.

It seems the ultimate irony that recent attempts at conserving our
butterflies often have amounted to studying them to death.

Literature Cited

ARNOLD, R. A., 1986. Ecological studies of the endangered Smith’s blue butterfly
at Marina State Beach in 1986. Final Report (unpublished).

EHRLICH, P. R. & D. D. MURPHY, 1987. Conservation lessons from long-term studies
of checkerspot butterflies, Jour. Cons. Biol. 1:122-131.

GALL, L. F., 1984. Population structure and recommendations for conservation of
the narrowly endemic alpine butterfly, Boloria acrocnema (Lepidoptera:
Nymphalidae). Biol. Cons. 28:111-138.

MORTON, A. C., 1984. In R. I. Vane-Wright and P. A. Ackery eds. The Biology of
Butterflies (Symposium of the Royal Entomological Society of London, No.
11). Academic Press, London.

MURPHY, D. D„ M. S. MENNINGER, P. R. EHRLICH, & B. A. WILCOX, 1986. Local
population dynamics of adult butterflies and the conservation status of two
closely related species. Biol. Cons. 37:201-223.

POLLARD, E , 1977. A method for assessing change in the abundance of butterflies.
Biol. Cons. 12:115-132.

REID, T., 1985. San Bruno Mountain area habitat conservation plan activities
report — 1983-1984. San Mateo County (unpublished).

SINGER, M. C. & P. WEDLAKE, 1981. Capture does affect probability of recapture in a
butterfly species. Ecol. Ent. 6:215-216.

A

26(l-4):240-247, 1988

Opinion.

Reply to Scott’s Criticism

J. P. Brock

Glasgow, Scotland (U.K.)

Differences of interpretation amongst authors dealing with the same set
of facts are only to be expected in the literature of phylogenetics.
However, when an author’s work receives heavy refutation on grounds
which (a) grossly misrepresent his own viewpoint, and (b) lean heavily
on wrongful statements of fact, then it is necessary and right that these
misrepresentations be exposed. The paper by Scott (1986) in this journal
is the subject of the following commentary.

The following notes on the distribution of character states of Ditry-
sian Lepidoptera show that Scott’s factual evidence runs contrary to
established knowledge.:

Larval Characters:

(1) The loss of L3 on the prothorax is not restricted to Pyraloidea and
‘Macrolepidoptera’ — it is also found in the Scardiinae (Tineidae),
Zygaenoidea, Epermeniidae and Glyphipterigidae-sercsw Brock. Scott
gives no explanatory comment on his placing of Carposinidae in
Pyraloidea — and his placing of Mimallonidae in that superfamily runs
contrary to characteristics of adult and pupa, his observations on larval
morphology having been known since the work of Fracker (1915). In
addition, some Pyraloidea ( sensu Scott) actually possess the ‘missing’ L
seta (Forbes, 1923). (2) Scott offers no reason for assuming that the
Pyraloid ‘L’ group is primitively unisetose on abdominal segment nine.
It is often bi- (sometimes tri-) setose in that superfamily. Again, the
supposed absence of two of the three ‘L’ setae on the same segment for
‘Macrolepidoptera’ is in opposition to the presence of two ‘L’ setae on
segment nine in Notodontidae and Drepanidae (Hassenfuss, 1969). The
same reduction trend is seen in many members of the Tineoid complex
(MacKay, 1972). (3) Scott is correct in stating that all ‘Macros’ have LI
and L2 separate on the abdomen. However, these setae are also
dissociated in the (extralimital) Immidae (Common, 1979), and in
monotrysian Heteroneura, Tineoidea s. str. (excluding Psychidae) and

26(l-4):l-288, 1988

241

Yponomeutoidea-serasw Brock. Other correlations of chaetotaxy could
be taken to show that the ‘Macros’ evolved from Cossoidea. The
condition of the ‘L’ setae in Mimallonidae (see above) could equally well
serve as evidence that these setae were associated in the ‘Macro’
ancestor — as indeed suggested by Forbes. These ‘wide-gap’ character
states occur separately in different families of Tineoidea sensu stricto.
(4) The presence of secondary setae cannot be taken as a shared-derived
trait of ‘Bombycoidea-Sphingoidea-Hesperioidea-Papilionoidea’. Similar
secondary vestiture is found in the (extralimital) Zygaenidae — and
it is certainly not ‘absent in Noctuoidea’ as claimed by Scott (cf.
Arctiidae, Ctenuchidae, Notodontidae, Acronictinae, etc.!). (5) Reduc¬
tion of crochets from circlet to mesoseries is found within many
ditrysian superfamilies, and the ‘butterfly triserial condition’ is found in
some Cossoidea and Pyraloidea — even in groups as remote as Gelechio-
idea (Forbes, 1923). Intermediate stages are found in Geometroidea, as
well as in Rhopalocera (amongst ‘Macrolepidoptera’).

Pupal Characters:

All characters of the ‘Macrolepidopterous’ pupa are found also in the
‘Micro’ groups Yponomeutoidea and Gelechioidea — while Scott’s
ancestor for Pyraloidea-Macrolepidoptera is clearly an incompletely
obtect pupa. ‘Spinose-protruded’ pupae also occur in some Bombycoidea
and Forbes ( 1923) records pupal protrusion for certain Pyraloidea. Some
Psychidae have ‘incomplete’ male pupae, and obtect females!

Looking in greater detail at Scott’s listing of pupal characters, it must
be stated that: (1) Maxillary palpi are not ‘lost’ in ‘Macros’ — as stated by
Mosher (1916) they are developed externally in many Noctuidae. (2) In
placing Sphingidae in his Hesperioid-Papilionoid lineage, Scott cites
‘loss of cocoon’ as a shared derived trait — yet some Sphingidae are
cocoon-builders. (3) Contrary to Scott’s claim, neither Zygaenoidea nor
Sesioidea have two rows of spines per segment. The primitive condition
is one row for Zygaenidae, the spination becoming more diffuse in
advanced forms. In Sesioidea, there is a single row per segment in
Choreutidae — double rows only in Sesiidae and Brachodidae (Heppner
and Duckworth, 1981). Similarly, the Tineoid superfamilies do not
‘generally have one row. . in this group, the Yponomeutoidea and
Gelechioidea are never spinose/protruded — excluding genera of Tortri-
coid-Sesioid-Zygaenoid relationships wrongly placed in Yponomeu¬
toidea by Common (1970) — see Brock, (1967, 1971), Heppner and
Duckworth ( loc . cit. above), Heppner (1977) and Kyrki (1984). Lyone-
tiid pupae are also non-spinose, and two spine rows have apparently
evolved independently in some members of the families Psychidae and
Gracillariidae (Mosher, 1916). (4) Mandible remnants are not ‘definite
bumps in Cossoidea-Castnioidea, weakly developed in butterflies’; the
‘Cossoid-Castnioid condition’ is widespread amongst ‘moths’ and the (non-

242

J.Res.Lepid.

unique) ‘butterfly condition’, far from being weakly developed, involves
an enlargement of these structures — so that right and left ‘mandible
remnants’ almost meet along the median line (Mosher, 1916). In the
same way, development of the clypeo-labral suture is variable in
Ditrysia, and cannot be used as a ‘shared-derived trait’. (5) Having used
the presence of fore leg femur in the pupae of ‘nearly all moths’ (cf:
‘shared-derived traits for monophyly of Hesperioidea-Papilionoidea’),
Scott goes on to list the exclusive presence of the same trait as evidence
for primitiveness of Geometroidea and Noctuoidea within his Macro-
lepidoptera. Loss of a visible fore-leg femur is also quite widespread
amongst Microlepidoptera (Mosher, 1916). (6) Scott uses the presence of
an epicranial suture/cleavage line as a further indicator of the primi¬
tive position of Geometroidea-Noctuoidea within ‘Macros’, yet fails to
state the presence of this trait in some Bombycoidea; his substitution of
temporal for ‘epicranial’ is based on an unlikely pupal-imaginal homo¬
logy (Scott, 1985), and his statement ( loc . cit ., and in the present diatribe)
that the epicranial cleavage line is absent in Lycaenidae (contrary to
Mosher) is incorrect.

Adult Characters:

(1) Vestigiation of the Cup vein in the fore wing is widespread
amongst lower ditrysians (including some Cossoidea-Zygaenoidea), and
in any case this vein is well developed in certain Bombycoidea. It cannot
therefore be a ‘shared-derived trait’ of Macrolepidoptera. (2) Sharplin’s
summary of character states associated with the wing base in Lepidop-
tera (Sharplin, 1964) clearly states that Cossoidea, Castnioidea and
Zygaenoidea either exhibit intermediate conditions between primitive
and advanced Ditrysia, or else they carry advanced (‘Macro’) character
states. Scott’s manipulation of the Sharplin data requires support from
described morphological observations — together with some indication
as to how Sharplin herself came to be misled. (3) The ‘discrimen’ (mesal
lamella) of the mesothorax shows variable development in Ditrysia, and
its strengthening in ‘Macros’ is not an exclusive (or universal) attribute
of these superfamilies (Brock, 1971). (4) The looped heart of ‘Macro¬
lepidoptera’ is also developed in many Cossidae and Limacodidae —
although not in Pyraloidea. Scott also states that ‘moths’ other than
some Cossidae ‘lack a chambered heart’, following Hessel (1969), while
the latter author states quite clearly {loc. cit.) that most ‘Macro-moths’
do have a chambered heart. Hessel’s distinction between the groups
cited by Scott was not dependent upon presence! absence of the heart
chamber — but on transverse versus horizontal orientation of the
chamber itself. (5) Scott presents no evidence whatsoever for his asser¬
tion that the Bombycoidea-Sphingoidea evolved from a tympanum¬
bearing ancestor. According to his scheme, the abdominal tympanum of
Pyraloidea was ancestral to that of Geometroidea and Noctuoidea — yet

26(l-4):l-288, 1988

243

many Pyraloidea and Geometroidea lack a tympanum, and there is no
definite evidence that this is a secondary development in either case.
There is also evidence to suggest that the Noctuoid tympanum evolved
in situ (i.e., on the thorax), rather than being a later advance on an
originally abdominal tympanum. (6) The presence/absence of ocelli,
chaetosemata and haustellum in Ditrysia follows such a widespread
pattern of correlation in the taxonomic hierarchy, that no importance
can be attached to this, above the level of family (Brock, 1971). (7) The
loss of the upper sector of the precoxal suture/sulcus cannot be brought
forward as a shared-derived trait for Bombycoidea-Sphingoidea-
Rhopalocera. As stated by Brock (1971), this trend is widespread in
other ditrysian superfamilies in which the suture is sometimes retained.
Scott’s treatment of this suture also implies two misidentifications in
Brock (loc. cit .) — firstly, in his use of quotation marks: ‘precoxal suture’
of Brock; in fact, the term paracoxal is nothing more than a doubtfully
necessary replacement term of Matsuda (1970) for the widely used
precoxal. Secondly, Scott states (here, and in Scott, 1985) that the
structure identified as ‘precoxal suture’ by me for Hesperiidae, is in fact
the secondary sternopleural suture (a term originated by me to replace a
misnamed suture of Ehrlich, 1958, still misapplied by Kristensen,
1976). If a secondary sternopleural suture does occur in Hesperiidae
other than those I examined, then (a) Scott must name and figure
examples, and (b) we require a more acceptable explanation from Scott
as to why the precoxal suture vestige figured by me is ‘misidentified’. (8)
The distal enlargement of the antenna reported by Scott for Sphingidae
(as a shared derived trait with Rhopalocera) is absent in many Sphingids
(and present in many diurnal moths — e.g.: Agaristinae, Castniidae,
Zygaenidae, Callidulidae, etc.). Its presence is clearly correlated with
diurnal activity. (9) The ‘areole’ of the fore wing radial system does not
‘occur in most moths’ — it is limited to a few families of Noctuoidea and
Geometroidea and one of Bombycoidea, amongst ‘Macros’. It is unlikely
that true homology exists between the ‘areoles’ of these groups, or
between any one of them and the (similarly sparsely distributed) ‘areole’
of lower Ditrysia (Brock, 1971). (10) The R4/5 branching basad of R1 in
the radial system of Rhopaloceran pupae is found also in adult Castni¬
idae (see Forbes, 1923). Published sources of data on pupal tracheation
of ‘moth’ groups are too fragmentary to allow broad generalisations to be
made at this stage. (11) Fusion of the anapleural cleft, listed as a shared-
derived trait for Hesperioidea-Papilionoidea by Scott (contrary to
Kristensen, 1976 — following Brock, 1971) is a feature of some diurnal
‘moths’ (and in any case, is not apparent in Hesperiids). (12) ‘Mesal
fusion of the metathoracic furcal arms’ is also listed as a shared derived
trait of Hesperioidea-Papilionoidea by Scott, but this occurs also in
Pyralidae (although not in other Pyraloidea — see Brock, 1971). Scott
also erroneously refers to fusion of furcal arms — whereas it is the
dorsal laminae of these structures which form the fusion (this same

244

J.Res.Lepid.

character is listed by Kristensen, 1976 as a ‘probable synapomorphy’ of
the same group). (13) The reduction of sternal apodemes (of abdominal
sternite two) in Rhopalocera is a widespread trait in Ditrysia —
including Limacodidae, Saturniidae, Thyrididae, and of course, many
tympanum-bearing families (Brock, 1971). (14) Scott mentions simil¬
arities between Hesperiidae and ‘Macro-moths’ in compound eye struc¬
ture. Following Yagi and Koyama (1963), this correlation lies with the
more primitive Bombycoidea alone — although these workers report a
similar correlation with the ‘non-Macro’ Cossidae — a fact not men¬
tioned by Scott. The same authors found a natural lineage between the
compound eye conditions of Pyraloidea-Geometroidea-Noctuoidea, this
apparently unconnected with the Cossoid/Hesperioid/lower Bombycoid
eye — although many convergencies were evident between diurnally
active members of quite remote families. These broader findings are
entirely ignored by Scott. In the lack of any good functional explanation
of the data on the compound eye, we do not know how (or if) the Cossid
condition could form a transition to that of the supposed Pyraloid
lineage — although its (theoretical) connections with Hesperioidea and
Bombycoidea are clear enough. (15) Scott lists the reduced condition of
the maxillary palpi in ‘Macrolepidoptera’ as diagnostic, yet three-
segmented palpi are reported for Carthaeidae of Bombycoidea by
Common (1966, cited in Brock, 1971). Reduction of palpi is found in
many ‘Microlepidoptera’, including Cossoidea, Zygaenoidea, and even
Tineoidea (Psychidae).

Given the high degree of polyphyly in the characters listed, it is not
possible to use them either as diagnostic features of suprafamilial
groups — or as monophyletic character changes along the branches of
Scott’s phyletic tree. Many provide useful suprageneric characters, but
all are known to be unstable at higher levels. Scott also uses primitive
character states as indicators of affinity (as indeed in his parallel paper
on butterfly phylogeny, Scott 1985). This ‘phenetic’ approach is univers¬
ally rejected by all cladists. In the same way, Scott’s use of ‘secondary
loss’ characters cannot have much relevance to phylogeny reconstruc¬
tion.

Scott’s attack on certain phyletic relationships suggested by me
completely overlooks the indirect ancestor concept (see Cracraft, 1974),
yet this too is regarded as a fundamental principle by phylogenetic
systematists. No direct ancestor relationship was ever proposed by me,
for any ditrysian superfamily pair — and to argue against theories of
indirect relationship on these grounds is nothing more than the knocking-
down of a ‘straw man’ hypothesis.

Scott’s table of characters actually shows ‘plus-minus’ entries (pro¬
viding ‘modified weighting’) for some characters listed in the accom¬
panying text as either ‘plus’ or ‘minus’ for the same taxa. An expansion
of this table to incorporate all other characters (and all other Ditrysian
superfamilies — with many more ‘pluses’ and ‘minuses’ corrected to

26(l-4):l-288, 1988

245

‘plus-minus’) would agree pretty well with the general conclusions of
Brock (1971) regarding the lability of the vast majority of characters
available for higher classification and phylogeny reconstruction at
superfamily level and above in Ditrysia.

Looking more deeply into the question of misrepresentation of my
own views — I find no mention whatsoever of the fact that my ‘selection
of characters’ was empirically based ( loc . cit ., p. 30). My final analysis
was stated, character-by-character, at some length — far from being the
‘intuitive’ system Scott claims it to have been. That analysis was placed
firmly in perspective with the broadest conclusions of the classical
authors working with the early stages, with actual quotations from the
contributions of Hinton (1952) and Chapman (1896) incorporated in the
text. In the same way, the broader conclusions of Yagi and Koyama
(1963) and of Hessel (1969) are either ignored or misrepresented by
Scott.

Based on the widespread manifestation of polyphyletic trends at and
above superfamily level in Ditrysia, I had argued that an explanation
could be found in the phenomenon of gradistic evolution (following the
terminology of Huxley; cf. Huxley, 1942/63) — and that the higher
groups Macrolepidoptera and Microlepidoptera were in fact, gradistic
constructs — this hypothesis being based on three main facts: (1) the
diagnostic traits of ‘Macro’ grade had virtually all evolved in several
‘Micro’ superfamilies, (2) the majority of ‘Micro’ grade characters are to
be seen distributed amongst primitive members of the ‘Macro’ super¬
families, and (3) most of these trend characters seemed correlated with
the change-over from endophagous to exophagous larval habits. This is
not the place to go into this hypothesis is detail, but Scott omits all
reference to this central theme of my earlier paper. Subsidiary to that
thesis, was an arguable hypothesis for cladistic relationships between
members of ‘Macro’ and ‘Micro’ grade, based upon those trends which
seemed the least labile. The gradistic concept of large-scale evolu¬
tionary change presents an obstacle to those cladists who are certain
that phylogeny can always be worked out on the basis of ‘synapomorphy’
and gradism has the further ‘wrong’ attribute of being highly ‘non-
parsimonious’. Whichever way we look at the data for Ditrysia, there
are virtually no ‘synapomorphies’ for units above superfamily — with
many superfamilies themselves based on polythetic distribution of
character states for included families. Basic to the question of gradistic
evolution is the related typological problem of ‘wide phenetic gaps’
between ‘Micro’ and ‘Macro’ superfamilies. For example, arguments
concerning the possibility of an indirect ancestry relationship between
Castnioidea and Rhopalocera invariably cite ‘Micro’ grade features of
Castnioidea (many of them lost in advanced Castniids!) as evidence of
‘unbridgeable gaps’ between these taxa. The strongest of these ‘gap’
character complexes lie with the larval and pupal stages — yet no one
objects to similar ‘gaps’ existing within families which display both

A

246

J.Res.Lepid.

endo- and exophagous biology (e.g., Pterophoridae, see Brock, 1971).
Secondly, the same authors seem unconcerned by the presence of
‘incomplete’ and obtect pupae within single families like Yponomeut-
idae ( sensu Common) and the Lyonetiidae of many recent authors.
Either these features are indicative of ‘unbridgeable gaps’ (equals
Special Creation?) or else they are potentially rapid adaptations (as
suggested by their taxonomic correlations).

The exceedingly widespread expression of parallelism at higher levels
in Ditrysia surely points to a major feature of the evolutionary process
itself, rather than an attempt by Brock to upset the theories of some
cladists and pheneticists. The way forward from this point must surely
be to look more deeply into the morphology of the lower Ditrysia, and to
re-examine larval and pupal morphology — looking for functional
explanations in these data — along with that presented by Yagi and
Koyama on the compound eye and by Hessel on the aorta and associated
structures. Anything less stands in grave danger of being categorised
as what H.E. Hinton once described as ‘a kind of juggling with the facts
— by persons with limited direct knowledge of the facts’. Granted, there
are alternative theories to the one I proposed — but we require better
representation both of existing knowledge and of other proposed
theories, than that now put forward by Scott. Impediments to such
progress are numerous — many families of Ditrysia are poorly repre¬
sented in collections and often early stages are little known — even
undiscovered. Functional explanations of morphological trends are
often evasive. Some authorities in a position to help with certain of
these problems are actually hostile to the continuation of work of this
kind — many lepidopterists will only support work in ‘safe areas’ which
pose no apparent threat to established classifications — ‘the stability of
Lepidoptera classifications since Herrich-Schaeffer’ having been one
‘justification’ for a personal attack on my work by an ‘authority’ at the
British Museum in London, which probably did more than anything to
impede further progress.

Literature Cited

BROCK, J. P. 1967. The systematic position of the Choreutinae (Lep: Glyphipteri-
gidae). Ent. mo. Mag. 103:245-6.

_ , 1971. A contribution towards an understanding of the morphology

and phylogeny of the Ditrysian Lepidoptera. J. nat. Hist., 5:29-102.
CHAPMAN, T. A. 1896. On the phylogeny and evolution of the Lepidoptera from the
pupal and oval standpoint. Trans. R. ent. Soc. Lond. (1896):567587.
COMMON, I. F. B. 1966. A new family of Bombycoidea (Lep.) based on Carthaea
saturnioides Walker from Western Australia. J. ent. Soc. Qd. 5:29-36.

_ , 1970. Lepidoptera. in Insects of Australia. C. S. I. R. O. Melbourne

Univ. Press.

_ , 1979. The larva and pupa of Imma acosma (Turner) and I. vaticina

Meyr. (Lep: Immidae) and the taxonomic relationships of the family. J.
Aust. ent. Soc. 1979; 18:33-38.

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247

CRACRAFT, J. 1974. Phylogenetic models and classification. Syst. Zool. 23(1):71-
90.

EHRLICH, P. R. 1958. The comparative morphology, phylogeny, and higher classi¬
fication of the butterflies (Lep: Papilionoidea). Univ. Kans. Sci. Bull.
39:305-370.

FORBES, w. T. M. 1923. Lepidoptera of New York and neighboring states. Mem.
Cornell Univ. Agric. Exp. Sta. 68: Ithica, New York.

FRACKER, S. B. 1915. The classification of Lepidopterous larvae. Contr. ent. Labs.
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Dennis D. Murphy, Department of Biological Sciences, Stanford University,
Stanford, California 94305

26(l-4):254-266, 1988

Notes

Notes on the biology of Brephidium exilis (Boisduval) (Lycaenidae)

Brephidium exilis (Boisduval) is a common butterfly throughout much of the
western United States, where its distribution stretches as far north as Oregon.
Since Edwards (1894, Can. Ent. 26: 37-38), several authors (Coolidge, 1924,
Ent. News 35: 115-121; Shapiro, 1972, J. Lepid. Soc. 27(2): 157-158 and
Johnson, 1984, J. Res. Lepid. 23: 104—106) offers data concerning aspects
related to its host plant and behavior.

In September and October of 1985 I studied some aspects of the biology of this
species on the U. C. Davis campus. Around Davis, Yolo Co., California, B. exilis
forms populations considered colonizing or fugitive (Shapiro, 1980, Atala 8(2):
46-49). In this locality I have seen this species often flying around Bassia
hyssopifolia (Pallas) Volk., Suaeda fruticosa (L.) Forsk, Atriplex argentea Nutt,
ssp, expansa (Wats) Hall & Clem, and Salsola australis R. Br. (all Chenopo-
diaceae) and Amaranthus graecizans L. (Amaranthaceae). In this paper some
aspects of the biology of this species in a colony which was flying around S.
australis and A. graecizans are described. Both plants were growing on waste
ground. S. australis grew in a semispherical fashion, reaching a height of about
1 m with some 60 individual plants spread over a distance of 100 m between the
two furthest apart. A. graecizans , on the other hand, is a creeping plant. In the
study area its distribution and abundance are similar to the latter. Both are
annual and reach maximum development during late summer and early
autumn.

In September, S. australis is in full flower. B. exilis flies preferentially around
the bushes of this species, upon which courting, mating and egglaying are
carried out. The light bluish-green eggs are laid near the flowers of S. australis.
For this purpose the female walks up and down the branches and after trying
out different flowers, lays just one egg close to one of them, flying off then to
search for another place to lay the next egg. Oviposition occurs also on A.
graecizans, although egglaying on this foodplant is less frequent.

In spite of their abundance, the caterpillars are difficult to find on the food
plant. On S. australis they feed on the flowers introducing the forepart of their
body into the axils of the bracteol which supports the flower. The B. exilis
caterpillars frequently attract ants. Conomyrma insana Buckley (Formicidae:
Dolychoderinae) is often found around them and occasionally several speci¬
mens around the same caterpillar. Due to the crypsis of the caterpillars on the
food plant, the best way to locate them is to follow the ants which attend them.
C. insana may play an important role in protecting the caterpillars from fly
parasitization. It has been observed that when a larva is approached by the fly
Aplomia theclarum Scudder (Tachinidae), attendant ants become very agi¬
tated, forcing the tachinid to fly off again. Similar bahavior has been reported
by Coolidge (1924, loc. cit.). Nevertheless a large number of caterpillars
collected in the field and reared in the laboratory contained the parasitoid
larva, which emerges in the last larval instar. The fly’s pupal stadium lasts
about 12 days.

With the first autumn rainfalls the population of B. exilis decreases rapidly.
According to Coolidge (1924, loc. cit.) hibernation takes place as pupa. Shapiro

26(l-4):l-288, 1988

255

(pers. comm.) states that no live pupa have been found near Davis after
Christmas, and that the species resumes flight in late Spring (April to June in
different years), suggesting annual reimmigration.

Acknowledgments. J. McCaskill identified the plants, P. S. Ward the ant and
P. H. Arnaud the fly. A. M. Shapiro contributed comments on this note. I am
very grateful to all concerned. My stay in Davis was supported by a grant of the
Consejeria de Educacion, Junta de Andalucia, and by grant 3126/83 of the
Spanish CAICCYT.

J. Fernandez Haeger, Departamento de Ecologia, Facultad de Ciencias 14071-
Cordoba, Spain

Courtship of a model ( Adelpha ; Nymphalidae) by its probable Batesian
mimic ( Limenitis ; Nymphalidae)

Silberglied (1977, in How Animals Communicate , T. Sebeok, ed., Indiana
Univ. Press, Bloomington, Indiana) suggested that optimal mating behaviors of
organisms involved in mimicry associations should involve communication
modes not shared by vertebrate predators. Because predation pressure causes
natural selection for convergence in flight behaviors as well as gross wing
pattern characteristics between mimics and their models, behaviors associated
with species recognition during courtship are expected to not rely heavily on
visual cues whether the species involved is a model or a mimic.

Observations on interspecific courtships allow insights concerning which
traits are important for mate recognition within species. Shapiro (1985, J. Res.
Lep. 24: 79-80) reported a case of confounded courtship where in the male of a
presumed model species ( Erynnis propertius Scud. & Burg., Hesperiidae)
pursued a female mimic ( Euclidea ardita Franc.; Noctuidae). Contrary to the
prediction above, the male appeared to be relying exclusively on visual cues. In
the following case the sexes are reversed — the pursuing male is the mimic.

Adelpha bredowii californica (Butler) and its presumed Batesian mimic
Limenitis (Basilarchia) lorquini lorquini (Boisduval) are sympatric below 2100
m. elevation throughout the Coast Ranges and Sierra Nevada in California.
Both species are dark brown with a prominent creamy white band across both
sets of wings and an orange tip on the forewing. These butterflies fly together
in the same canyons, and males often compete for territories in the sunny
patches of stream beds. The mimicry relationship has never been formally
tested, but is widely inferred from both the sympatry and the documentation of
Batesian mimicry in other species of Limenitis (e.g., Platt, Coppinger, &
Brower, 1971, Evolution 25: 692-710).

While collecting L. lorquini and A. b. californica for laboratory studies, I
encountered a male lorquini courting a female 6. californica. The sighting
occurred at 1355 hrs, 22 April 1986, in Mix Canyon, Solano Co., California
(approx. 10 km. north of Vacaville), and lasted until 1359 hrs, when the pair
was lost from sight. When initially encountered, the courtship was in progress,
with both butterflies fluttering one to two meters above the road at the edge of
a small sunlit area. After about thirty seconds, the female began flying faster,

256

J. Res. Lepid.

with the lorquini in hot pursuit (i.e., within 10 cm). The pair made a number of
wide circles in a large (30 m) sunny area above a stream adjacent to the road,
ending when the female alit on a shrub ( Quercus dumosa Nutt., Fagaceae; a
possible foodplant of b. calif ornica) . The lorquini male immediately alit beside
her and nudged her with his curled abdomen in repeated attempts to copulate.
The female avoided this by turning her abdomen away from the male or by
short flicks of the wings. These behaviors continued for approximately one
minute, then the male flew to a nearby (2 m) bush and perched while the
female basked. When the female flew again after an interval of approximately
90 seconds, the male gave chase, and the pair was lost from sight. The weather
was sunny and clear with little wind, temperature approximately 26°C. A. b.
californica males were quite common in the canyon, but only eight lorquini
males were seen over the span of three hours. Additionally, three b. californica
females and one lorquini female were collected. These densities are representa¬
tive of most spring seasons.

Because I did not see the beginning of the encounter, it is not clear whether
the female above initiated the fluttering courtship flight using wing pattern
cues from the male Limenitis , or if receptive females simply initiate the
fluttering flight whenever pursued. It does appear clear, however, that the
female Adelpha rejected the inappropriate suitor only after she had begun the
fluttering courtship display. Whether this was on the basis of visual or pher¬
omonal cues remains unknown. The male lorquini seemed completely unaware
of his faux pas throughout the encounter. As in Erynnis reported by Shapiro
(op. cit.), male Limenitis appear to rely exclusively on visual cues during
courtship.

Adam H. Porter, Dept, of Zoology, University of California, Davis, CA 95616

A Bibliography of Euphydryas

Checkerspot butterflies of the genus Euphydryas are among the most well
studied Lepidoptera, and have become key organisms for testing ecological and
evolutionary theory. Here we have compiled a bibliography of papers con¬
cerning this genus. We suspect that this bibliography will be a useful resource
both to those working directly with Euphydryas and to those with a more
general interest in butterfly ecology. The topics included cover distributional
notes, population dynamics, population genetics, host plant and parasitoid
interactions, and behavior. We have endeavored to make this bibliography as
complete as possible, but in an effort to produce a bibliography of manageable
size we have excluded most taxonomic descriptions. Those for the most part are
referenced in Gunder (1929), Miller and Brown (1981, A Catalogue/Checklist of
the Butterflies of America North of Mexico. The Lepidopterists Society Memoir
No. 2), Kudrna (1985, Butterflies of Europe; Concise Bibliography of European
Butterflies. AULA-Verlag Wiesbaden), and various works by Higgins.

We gratefully acknowledge contributions to this list by M. D. Bowers, N. E.
Stamp and R. R. White.

26(l-4):l-288, 1988

257

BESNOIST, G., 1970. Note sur l’abondance d Euphydryas cynthia au Lac d‘Allos.
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BETZ, J. T., 1956. Melitaea aurinia Rott. en Algerie. Rev. Franc Lepid. Paris
15:143-144.

BOROWSKY, R., 1977. Detection of the effects of natural selection of protein
polymorphisms in natural populations by means of a distance analysis.
Evolution 31:341-346.

BOURGOGNE, J„ 1960. Euphydryas intermedia Men. dans les Hautes Alpes.
Alexanor 1:253-254.

BOWE, J. J., 1972. Another larval foodplant for Euphydryas phaeton (Drury)
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BOWERS, M. D., 1978. Overwintering behavior in Euphydryas phaeton (Nympha¬
lidae). J. Lepid. Soc. 32:282-288.

BOWERS, M. D„ 1979. Unpalatability as a defense strategy of checkerspot butter¬
flies with special reference to Euphydryas phaeton (Nymphalidae). Ph. D.
Dissertation, Univ. of Mass.

BOWERS, M. D., 1980. Unpalatability as a defense strategy of Euphydryas
phaeton. Evolution 34:586-600.

BOWERS, M. D., 1981. Unpalatability as a defense strategy of western checkerspot
butterflies ( Euphydryas ). Evolution 35:367-375.

BOWERS, M. D., 1983. Mimicry in North American checkerspot butterflies:
Euphydryas phaeton and Chlosyne harrisii (Nymphalidae). Ecol. Entomol.
8:1-8.

BOWERS, M. D., 1983. Iridoid glycosides and larval hostplant specificity in
checkerspot butterflies ( Euphydryas : Nymphalidae). J. Chem. Ecol. 9:475-
493.

BOWERS, M. D., 1985. Host plant choice of checkerspot larvae: Euphydryas
chalcedona, E. colon, and hybrids (Lepidoptera: Nymphalidae). Psyche
92:39-48.

! BOWERS, M. D., 1986. Population differences in larval host plant use in the
checkerspot butterfly Euphydryas chalcedona. Entomol. Exp. App. 40:61-69.

BOWERS, M. D„ I. L. BROWN, & D. R. WHEYE, 1985. Bird predation as a selective agent
in a butterfly population. Evolution 39:93-103.

BOWERS, M. D, & G. M. PUTTICK, 1986. Fate of ingested iridoid glycosides in
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BRETHERTON, R. F., 1981. On rearing the Spanish Fritillary Eurodryas desfon-
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BROWN, I. L. & P. R. EHRLICH, 1980. Population biology of the checkerspot butterfly
Euphydryas chalcedona. Structure of the Jasper Ridge colony. Oecologia
47:239-251.

BROWN, K. S., 1965. Some unusual butterfly records from central California. J.
Lepid. Soc. 19:171-175.

BRUSSARD, P. F. & P. R. EHRLICH, 1970. Contrasting population biology of two
species of butterfly. Nature 227:91-92.

BRUSSARD, P. F„ P. R. EHRLICH, & M. C. SINGER, 1974. Adult movements and
population structure in Euphydryas editha. Evolution 28:408-415.

j BRUSSARD, P. F. & A. T. VAWTER, 1975. Population structure, gene flow, and natural
selection in populations of Euphydryas phaeton. Heredity 34:407-415.

I BRUSSARD, P. F„ P. R. EHRLICH, D. D. MURPHY, B. A. WILCOX, J WRIGHT, 1985. Genetic
distances and the taxonomy of checkerspot butterflies (Nymphalidae:

A

258

J.Res.Lepid.

Nymphalinae) J. Kans. Ent. Soc. 58:403-412.

CHAPMAN, T. A., 1905. Cryptic form and colouring of Melitaea larvae. Entomo¬
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CLARK, A. H., 1927. Notes on the melitaeid butterfly Euphydryas phaeton (Drury)
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COMSTOCK, J. A., 1940. Some notes on the early stages of Euphydryas gillettii
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COMSTOCK, J. A., 1958. Random notes on early stages of Lepidoptera. Bull. S. Cal.
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CULLENWARD, M. J., P. R. EHRLICH, R. R. WHITE, & C. E. HOLDREN, 1979. The ecology
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DOBKIN, D. S., I. OLIVIERI, P. R. EHRLICH, 1987. Rainfall and the interaction of
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DOUDOROFF, M., 1935. Notes on two local butterflies. Pan-Pac. Entomol. 11:144.

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EHRLICH, P. R., 1961. Intrinsic barriers to dispersal in the checkerspot butterfly,
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EHRLICH, P. R. & D. D. MURPHY, 1987. Monitoring populations on remnants of
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EHRLICH, P. R„ D. D. MURPHY, M. C. SINGER, C. B. SHERWOOD, R. R. WHITE, & I. L. BROWN,
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EHRLICH, P. R., & D. WHEYE, 1985. Some observations on the spatial distribution in
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EHRLICH, P. R., & D. WHEYE, 1986. Non-adaptive “hilltopping” behavior in male
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EHRLICH, P. R., & R. R. WHITE, 1980. Colorado checkerspot butterflies: isolation,
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population. Evolution 37:389-403.

SINGER, M. C., 1984. Butterfly-host plant relationships: host quality, adult choice,
and larval success. In: Vane-Wright and Ackery. eds. The Biology of
Butterflies. Academic Press, London.

SINGER, M. C. & P. R. EHRLICH, 1979. Population dynamics of the checkerspot
butterfly Euphydryas editha. Fortrschr. Zool. 25:53-60.

SPOMER, S., 1985. Observations on the life history of Occidryas anicia bernadetta
(Nymphalidae) at the type locality. J. Lepid. Soc. 39:55-57.

STAMP, N. E , 1979. New oviposition plant for Euphydryas phaeton. J. Lepid. Soc.
33:203-204.

STAMP, N. E., 1980. Effects of group size on an egg clustering butterfly. Ph. D.
Dissertation, Univ. of Maryland.

STAMP, N. E., 1980. Egg deposition patterns in butterflies: why do some species
cluster their eggs rather than deposit them singly? Amer. Natur. 115:367-
380.

STAMP, N. E„ 1981. Effect of group size on parasitism in a natural population of
the Baltimore checkerspot, Euphydryas phaeton. Oecologia 49:201-206.

STAMP, N. E ., 1981. Parasitism of single and multiple egg clusters of Euphydryas
phaeton (Nymphalidae). J. N. Y. Ent. Soc. 89:89-97.

26(l-4):l-288, 1988

263

STAMP, N. E., 1981. Behavior of parasitized aposematic caterpillars: advantage to
the parasitoid or host? Am. Nat. 118:715-725.

STAMP, N. E., 1982. Aggregation behavior in Baltimore checkerspot caterpillars,
Euphydryas phaeton (Nymphalidae). J. Lepid. Soc. 36:31-41.

STAMP, N. E., 1982. Selection of oviposition sites by the Baltimore checkerspot,
Euphydryas phaeton (Nymphalidae). J. Lepid. Soc. 36:290-302.

STAMP, N. E., 1982. Behavior interactions of parasitoids and Baltimore checker-
spot caterpillars (. Euphydryas phaeton ). Environ. Ent. 11:100-104.

STAMP, N. E., 1982. Searching behavior of parasitoids for webmaking caterpillars:
a test of optimal searching theory. J. Animal Ecol. 52:387-395.

STAMP, N. E., 1984. Effect of defoliation by checkerspot caterpillars ( Euphydryas
phaeton and sawfly larvae ( Macrophya nigra and Tenthredo grandis ) on
their host plants ( Chelone spp.). Oecologia 63:275-280.

STAMP, N. E., 1984. Interactions of parasitoids and checkerspot caterpillars
Euphydryas spp. (Nymphalidae). J. Res. Lepid 23:2-18.

STERNMITZ, F. S., D. R. GARDNER, F. J. ODENDAAL, & P. R. EHRLICH, 1986. Euphydryas
anicia utilization of iriodoid glycosides from Castilleja and Besseya (Scophu-
lariaceae) J. Chem. Ecol. 12:1456-1468.

TEMPLADO, J., 1975. La regulacion natural de las poblaciones de Euphydryas
aurinia Rott. Bol. Estn. Ecol. Madrid 4:77-81.

THOMAS, C. D., & M. C. SINGER, 198'/ . Variation in host preference affects movement
patterns within a butterfly population. Ecology: in press.

THOMAS, C. D., D. NG, M. C. SINGER, J. L. B. MALLET, C. PARMESAN, & H. L. BILLINGTON,
1987. Incorporation of a European weed into the diet of a North American
herbivore. Evolution: In Press.

THORNE, F., 1970. Habitat: Euphydryas editha wrighti. J. Res. Lepid. 7:167.

TILDEN, J. W., 1958. Some notes on the life history of Euphydryas editha bayensis.
J. Lepid. Soc. 12:33-35.

TOLIVER, M., 1971. A record of Euphydryas anicia (Nymphalidae) in Oklahoma.
J. Lepid. Soc. 25:246.

TURATI, E., 1910. Zwei neue italienische Melitaea aurinia Formen. Ent. Z.
Frankf. a.M. 23:233.

VAWTER, A. T. & J. WRIGHT, 1986. Genetic differentiation between subspecies of
Euphydryas phaeton (Nymphalidae: Nymphalinae). J. Res. Lepid. 25:25-29.

VARGA, Z., & G. SANTHA, 1973. Verbrietung und taxonomische Gliederung der
Euphydryas maturna L. in So-Europa (Euphydryas-S tudien 1.) Acta Biol,
debrecina 10/11 (1972/73):213-231.

VERITY, R., 1928. An essay on the geographic variation of the Rhopalocera
exemplified by Melitaea aurinia Rott. Entomologists Rec. J. Var. 40:41-45,
86-91, 97-101.

VINE-HALL, J. H., 1979. Euphydryas aurinia Rott. present in Cumbria. Entomo¬
logist’s Rec. J. Var. 91:24-25.

WEISS, S. B., R. R. WHITE, D. D. MURPHY, & P. R. EHRLICH, 1987. Growth and dispersal of
larvae of the checkerspot butterfly Euphydryas editha. Oikos, in press.

WHEELER, G., 1915. The genus Melitaea. Proc. S. Lond. ent. nat. Hist. Soc. 1914-
1915:1-16.

WHEYE, D., & P. R. EHRLICH, 1985. The use of flourescent pigments to study insect
behavior: investigating mating patterns in a butterfly population. Ecol.
Entomol. 10:231-234.

WHITE, R. R., 1973. Community relationships of the butterfly Euphydryas editha.

264

J.Res.Lepid.

Ph. D. Dissertation, Stanford Univ.

WHITE, R. R., 1974. Foodplant defoliation and larval starvation of Euphydryas
editha. Oecologia 14:307-315.

WHITE, R. R., 1979. Foodplant of alpine Euphydryas anicia (Nymphalidae). J.
Lepid. Soc. 33:170-173.

WHITE, R. R., 1980. Inter-peak dispersal in alpine checkerspot butterflies
(Nymphalidae). J. Lepid. Soc. 34:353-362.

WHITE, R. R., 1986. Pupal mortality in the bay checkerspot butterfly. J. Res.
Lepid. 25:52-62.

WHITE, R. R., 1987. The trouble with butterflies. J. Res. Lepid. in press.

WHITE, R. R., & M. P. LEVIN, 1981. Temporal variation in insect vagility: implica¬
tions for evolutionary studies. Am. Midi. Nat. 105:348-357.

WHITE, R. R., & M. C. SINGER, 1974. Geographical distribution of host plant choice in
Euphydryas editha (Nymphalidae). J. Lepid. Soc. 28:103-107.

WILCOX, B. A., & D. D. MURPHY, 1985. Conservation strategy: the effects of
fragmentation on extinction. Amer. Nat. 125:879-887.

WILLIAMS, E. H., 1981. Thermal influences on oviposition in the montane butterfly
Euphydryas gillettii. Oecologia 50:342-346.

WILLIAMS, E. H., & M. D. BOWERS, 1987. Factors affecting host plant use by the
montane butterfly Euphydryas gillettii (Nymphalidae). Am. Midi. Nat. (in
press).

WILLIAMS, E. H., C. E. HOLDREN, & P. R. EHRLICH, 1983. The life history and ecology of
Euphydryas gillettii Barnes (Nymphalidae). J. Lepid. Soc. 38:1-12.

WILLIAMS, K. S., 1981. The coevolution of the checkerspot butterfly Euphydryas
chalcedona and its larval host plants. Ph. D. Thesis, Stanford Univ.

WILLIAMS, K. S., 1983. The coevolution of Euphydryas chalcedona butterflies and
their larval host plants. III. Oviposition behavior and host plant quality.
Oecologia 56:336-340.

WILLIAMS, K. S., D. E. LINCOLN, & P. R. EHRLICH, 1983. The coevolution of Euphy¬
dryas chalcedona butterflies and their larval host plants. II. Maternal and
host plant effect on larval growth, development, and food-use efficiency.
Oecologia 56:330-335.

WILLIAMS, K. S., D. E. LINCOLN, & P. R. EHRLICH, 1983. Coevolution of Euphydryas
chalcedona butterflies and their larval host plants. I. Larval feeding
behavior and host plant chemisty. Oecologia 56:323-329.

Dennis D. Murphy and Stuart B. Weiss , Department of Biological Sciences,
Stanford University, Stanford, CA 94305.

Aberrant Polyommatinae (Lycaenidae) from Ohio and Florida

Aberrant Polyommatinae have recently been illustraded by Wright (1979, J.
Lepid Soc. 33: 266) and Neil (1983, J. Lepid. Soc. 37: 258). Presented here are
three extreme aberrations in pattern not previously recorded.

On 19 April 1986, at Shade River State Forest, Meigs Co., Ohio, an aberrant
female Glaucopsyche lygdamus lygdamus (Doubleday) was collected in a rich
stream valley near Forked Run State Park. The individual was found in

26(l-4):l-288, 1988

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association with wood vetch, Vicia caroliniana Walt., which grows abundantly
along a SE-facing roadbank. This plant is the primary hostplant of the species in
Ohio. Between 1030 h and 1200 h the author and D. C. Iftner collected and
observed several G. lygdamus flying in the vicinity of wood vetch and visiting
damp soil along a dirt township road that traverses the valley.

The aberrant female (Fig. lb) is striking in that the normal basal and
postmedian black spots on the ventral surfaces of the hindwings are totally
lacking. The small discal bars are the only remnants of normal maculation. The
postmedian row of spots on the ventral surface of each forewong is dis-placed
inwardly and is found just beyond the cell. The ventral surface of the left
fore wing possesses five spots while the opposite wing exhibits only four. The
dorsal surfaces are normal. Although G. lygdamus is highly variable both
individually and geographically, this aberration illustrates a phenotype never
before observed in Ohio.

Two aberrant individuals of the subspecies Hemiargus thomasi bethunebakeri
W. P. Comstock and Hemiargus ceraunus antibubastus Hubner were captured
in extreme southern Florida. The former specimen (Fig. Id) is a worn male
taken 14 December 1982 on Key Largo, Monroe Co. Ventrally, the forewings
possess enlarged and lengthened white areas that taper basally. The hindwings
lack the black portions of the basal and costal spots. The postdiscal white band is
expanded and tapers inwardly.

The individial of H. c. antibubastus (Fig. If) is a male in good condition taken 1
September 1982 on Stock Island, Monroe Co., Florida. The specimen has a
grossly atypical pattern ventrally. The wing surfaces are almost completely
suffused with white scales, creating a submarginal band of dark spots on the
forewings. The hindwings slightly differ individually. The two black costal spots
of the right hindwing are enlarged and the outermost spot appears “smeared”.
Two dark postdiscal bands extend completely across the wing. The left hindwing

Fig. 1 . Aberrant and normal specimens of three species of Polyommatinae a.
Normal male G. lygdamus, b. Aberrant female G. lygdamus, c. Normal
male H. thomasi, d. Aberrant male H. thomasi, e. Normal male H.
ceraunus, f. Aberrant male H. ceraunus.

266

J. Res. Lepid.

lacks the innermost black costal spot but a ’’smeared” outermost spot is present.
A dark postdiscal band extends across the wing and a partial second band
extends from the inner margin to CUj.

The specimens figured are in the collection of the author.

John V. Calhoun, 369 Tradewind Ct. Westerville, Ohio 43081

Ommochromes in Libytheidae

Ommochromes are a group of pigments that occur as granules in insect eyes
and in various other organs and tissues (Robinson, 1971, Lepidoptera Genetics,
Pergamon Press, Oxford). The red meconium of teneral adults, which contains
ommochromes, is widespread in Pierinae and Nymphalinae and is also known to
occur in a few Coliadinae, Charaxes, and certain Parnassiinae (Shapiro, 1982, J.
Res. Lepid. 20: 97—102). Some Pierinae have red eggs which may be a
manifestation of ommochromes (Shapiro, op. cit.); ommochromes can also be
brown in color (Robinson, op. cit.). Since pterins and ommochromes are often
associated in insect eyes (including Lepidoptera) and tend to be affected by the
same mutation, it is thought that certain pterins are involved in the enzymatic
processes that form ommochromes (Robinson, op. cit.; Gilmour, 1961, The
Biochemistry of Insects, Academic Press, New York & London). It was recently
discovered that some Libytheidae contain pterin pigments in their wings
(Shields, 1987, J. Chem. Ecol., in press).

The meconium of two freshly emerged Libytheana bachmanii bachmanii
Kirtland from College Station, Texas, was light reddish orange to moderate
orange with brown speckles, i.e. rust tan or testaceous in color (T. Friedlander, in
litt.). When this species is ready to pupate, it spins a button of red silk on the
underside of a leaf (Edwards, 1881, Canad. Ent. 13: 226-229). When first laid,
the egg of Libythea laius Trimen is whitish but in two days turns to a pale, dull
salmon tint; similarly, Libythea celtis celtis Fuessly eggs turn brownish-pink
(Shields, 1985, Tokurana 9: 1—58). These data suggest the Libytheidae may also
contain ommochromes.

Oakley Shields, 4890 Old Highway, Mariposa, California 95338.

26(l-4):267-288, 1988

Book Reviews

THE BUTTERFLIES OF NORTH AMERICA — A NATURAL HISTORY AND
FIELD GUIDE. Scott, James A., Stanford University Press, Stanford, Cali¬
fornia. 583 pages, hardback. $49.50.

This is the new definitive treatment of North American butterflies and
skippers. It replaces the book edited by W. H. Howe that was published in 1974.
Howe’s book appeared 44 years after the previous compendium by W. H.
Holland. The reader may ask ‘Is it really necessary to have another new
butterfly book so soon after Howe’s treatment?’ or ‘How does Scott’s book
compare with previous books?’

Howe’s book was really written in the late 1960’s but took more than 5 years
in the production process so the span is a bit more than mere subtraction of years
would suggest. Scott attempted to keep his text as current as possible up to about
a year before actual publication. Twenty-one authors contributed to the Howe
book and the text was edited by F. M. Brown and H. K. Clench. In contrast, the
entire text of the new book was written by a single author — a monumental
undertaking. The two books differ in many other ways as well. The Howe book is
illustrated with 97 plates with beautiful color paintings by the editor, whereas
the new work includes 64 color plates of photographs of butterfly specimens
and shots of living individuals. I consider Howe’s plates superior, although
many species not found in the Howe book are newly illustrated by Scott. The
larger and lighter butterflies are best shown by Scott’s photographic plates, but
the smaller darker butterflies — often too many on a single plate — lack
sufficient contrast to be shown well. In addition, most of the specimens are from
Scott’s own collection, and the ventral views show the ravages on the butterflies
of having died between thumb and forefinger, i.e. missing legs and vestiture.

Scott’s book places a much greater emphasis on behavior, host plants, and
ecology than any previous North America-wide treatment. The book has 153
pages of introductory chapters, followed by 348 pages of species treatments, and
ending with 45 pages of appendices.

The introductory chapters are almost like a separate book on butterfly
evolution, structure, and biology. This reviewer is amazed at the amount of
technical material included. Many readers will find the introductory sections
like an introductory text in its content, but will find many of the technical terms
softened by the author. I found a few bothersome, e.g. ‘jaws’ instead of
‘mandibles,’ and ‘blood’ instead of ‘hemolymph,’ but most readers will not mind.
Scott also uses mechanomorphisms such as tracheae being ‘constructed like
vacuum cleaner hose.’ Most of the material is accurate and factual, but there are
some inaccuracies such as the statement on p. 42 that ‘butterflies only fly when
the sun shines’ and that on p. 26 that implies females of all or most species are
larger than their males. Anyone who has observed butterflies in eastern North
America can vouch that many species fly in numbers on cloudy days. Likewise,
males of many pierid species are larger than the females, and most lycaenids
have no significant size difference between the sexes. It is difficult to ascertain
which information is the result of Scott’s research, which is opinion, and which
has been taken from other works. In some cases literature sources are referred to
by the author’s last name and first initial, but no indication of the literature
citation is to be found later in the book, in other cases this reviewer found

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instances of results from works other than Scott’s given with no attribution. In a
similar vein some of the line illustrations that appear throughout the book,
mostly drawn by Scott, are taken directly from the works of others without
acknowledgement. Much of the introductory material will prepare the reader
for the species treatments that follow, e.g. chapters on behavior, reproduction,
migration, and geographic variation, but others are of no use later on. Examples
of these are the excellent keys to first instar and mature larvae, pupae, and
adults, but on page 158 the reader is told “To identify an adult butterfly, first
compare the butterfly you have found with the butterflies illustrated on the
color plates, and settle on its probable identity.” Why, one wonders, were the
keys included?

The species treatments will be the ‘meat’ for most lepidopterists, and I will
devote most attention to them. First it is worth comparing the taxa included
with those in the 1981 Miller and Brown Catalogue/Checklist (Lepid. Soc. Mem.
No. 2). At the suprageneric level Scott follows the Ehrlich classification —
recognizing five families of Papilionoidea (Papilionidae, Pieridae, Lycaenidae,
Libytheidae, and Nymphalidae) and one of Hesperioidea (Hesperiidae); while,
in contrast, Miller and Brown split the Nymphalidae into 5 separate families
while recognizing the others. At the subfamily level the splitting approach is
further executed by Miller and Brown as they recognize several subfamilies that
in the eyes of this reviewer are placed at least one level too high, i.e.
Argynninae, Melitaeinae, Marpesiinae, Charaxinae, Apaturinae. Scott’s sub¬
family treatment is suitably conservative, fitting well with those for other Lepi-
doptera Superfamilies. Scott’s apparent recognition of the unnatural higher
groups ‘Macrolepidoptera’ and ‘Rhopalocera’ in his only black mark in the
higher classification discussion.

At the generic level the contrast between Scott versus Miller and Brown
continues. Miller and Brown raised subgenera and some species group names to
genera, and considered North American butterflies to belong to 241 genera,
while Scott, in a more conservative treatment, considered the same species to
comprise only 203 genera, 38 fewer than the former work; 5 other genera in
Scott’s treatment are due to Hawaiian species(3) and Mexican species(2) newly
reported in North America.

At the species level Miller and Brown listed 763 species, but felt that 14 were
improperly reported; thus they attributed 749 species to the North American
fauna. In contrast, Scott includes 679 species, but 6 of these are Hawaiian not
included by Miller and Brown, 2 listed as probably not North American, and,
finally, 17 are newly reported for North America: Staphylus azteca, Arteurotia
tractipennis , Autochton cincta, Amblyscirtes unnamed (sic!), Vettius fantasos,
Synapte syraces, Piruna cingo, Callophrys Herodotus , Boloria natazhati,
Chlosyne marina, Precis genoveva, Epiphile adrasta, Oeneis alpina, Erebia
occulta, E. kozhantshikovi, Phoebis orbis, and Papilio victorinus. This makes the
comparable species number for Scott’s book 654 — 95 species fewer than Miller
and Brown! What is the reason for this disparity? The answer seems to be split¬
ting on the part of prior revisers reported by Miller and Brown and some
lumping on the part of Scott. Examples of needless inclusions as full species in
Miller and Brown are hybrids ( Colias boothii\hccla x nastes ] and Anthocharis
dammersi [ sara x lanccolata] as well as many members of closely related para-
patric and allopatric species complexes, e.g. Astcrocampa, Cocnonympha, and

26(l-4):l-288, 1988

269

Agathymus. On the other hand, Scott eliminates many of these excesses, but
probably goes too far in the other direction by lumping as conspecific good bio¬
logical species. Some of these are each other’s closest relatives, but Scott seems
to use any evidence of hybridization in nature as cause for lumping. Unfor¬
tunately, the rationale for these actions is usually not given. In addition, some
species are lumped under their closest Palaearctic relatives even though no
biological studies to justify the actions have been reported. Some of these may
make sense; examples are Pontia occidentalis under Pontia callidice, and
Euchloe ausonides under Euchloe ausonia.

For 30 years or more, the trend in animal systematics has been to deem-
phasize subspecies. Examples are the current checklists for North American
birds and mammals that exclude listing of any subspecies. Scott’s book includes
430 subspecies above the given nomenotypic taxon for each species, whereas
Miller and Brown list 745 subspecies above those necessitated by listing the
species itself. This is a strong reduction of 315 names, and most of the missing
names will leave no voids in any meaningful discussion of geographic variation.
Yet, Scott does apply the criteria for which subspecies to include in his book very
unevenly. Virtually all of the subspecies described by Scott himself are
mentioned, but others that seem more significant are omitted or lumped.

Whether intentional or not, Scott’s systematics is unorthodox. Some popula¬
tions are mentioned as ‘subspecies unnamed’ — a meaningless designation, and,
more importantly, the authors are given for none of the included scientific
names. The reader is referred to the Miller and Brown treatment for authors’
names and citations, yet many species not listed by those authors are given and
many new combinations are presented.

Looking at the species discussions themselves one finds the real heart of the
book, and there is little to fault. Under each species there are 4 paragraphs —
one for identification features of the adults, one to describe habitat, geographic
range, and larval host plants, one for a thumbnail sketch of the early stages, and
one to describe voltinism and adult behavior. A small but accurate range map is
included in the margin for most species. Scott’s researchs have been nearly
exhaustive and the pages devoted to the species treatments are absolutely
packed with valuable information. This reviewer found that the given southern
terminus of the range for many species extending into the Neotropics fell far
short of the known southern limits; apparently Scott did not look at any
collections or much literature on Neotropical butterfly ranges. Some examples
are Eurytides philolaus — Scott gives south to Honduras, actually south to
Costa Rica, and Apodemia multiplaga — Scott gives south to Puebla, Mexico,
actually south to Costa Rica.

Some of the wording in the species treatments is a bit bothersome; for example
butterflies do not ‘sip mud,’ but they may sip moisture from mud or wet sand.

The appendices are valuable. They contain lists of species from islands in the
vicinty of North America — Iceland, Greenland, Bermuda, and Hawaii; and
techniques for collecting and studying butterflies. Scott continues the ‘pinching
technique’ for the dispatch of specimens, instead of the use of poison which does
not cause the loss of legs or distortion of the body. The techniques described are
valuable and usually expedient. Especially valuable are techniques for study¬
ing life stages and marking butterflies. Many of the line drawings used to
illustrate the genitalia dissection technique paragraphs are borrowed from
other works without attribution.

270

J.Res.Lepid.

Ending up the book is a woefully inadequate bibliography (see above), an
excellent ‘hostplant catalogue’ — but numbered cross references to butterfly
names earlier in text reduce the value for quick reference, an excellent glossary
for the nontechnical reader, and a long detailed index to general subjects and
butterfly names.

It will probably be at least a decade before another North America- wide
compendium appears. For the person interested in observing butterflies as a
naturalist or for people interested in conservation, the book has minimal value.
Despite its title the book is not one you would want to put in your back pocket for
a hike and it does not cater to non-collectors. In summary, the book has a lot of
positives and some detractions, but all butterfly collectors and biologists who
study butterflies will want the book on their shelves.

Paul A . Opler, U. S. Fish and Wildlife Service, 1 025 Pennock PL, Ft. Collins, CO
80524

To judge from its self-congratulatory Preface, this most recent in the spate of
books on North American butterflies represents the comprehensive treatise —
superceding and subsuming the many regional treatments and few synthetic
volumes to have appeared in the past several decades. It is indeed such a bid on
an authoritative treatise, like Howe (1975) for the whole fauna, and Opler &
Krizek (1984) for the eastern states. And although its title might imply so, it is
neither a field guide (Pyle’s several books are) nor really a natural history
(Opler & Krizek’s is of that genre, if anything can be).

Scott’s book is among the lengthiest produced in recent years, bursting its
seams with biological and taxonomic minutiae at every turn of the page. It
therefore warrants an exhaustive review, so please bear with me. Since other
opinions are always instructive, I’ve sprinkled in excerpts from Paul Opler ’s
initial review of this book in the Audubon Naturalist News (Vol. 13(1): 6), Art
Shapiro’s in the Journal of the Lepidopterists’ Society (Vol. 40(4):356-357), and
Jeff Glassberg’s in the Mulberry Wing (Vol. 2(4): 4-6).

1. Generalities

The prose is staccato. As a consequence, the book doesn’t read as smoothly as it
could, wanting still for further grammatical editing. Opler’s review termed the
text “concise, sparse, and direct.”

The positive points of the book are at once identifiable. These include: (1)
novel keys to the newly hatched and full-grown larvae, and pupae of North
American butterflies; (2) much hostplant information, with copious attendant
details; (3) exhuastive treatment of butterfly physiology and behavior; (4) a
well-armed assault on the phyletics of Lepidoptera using largely morphological
characters; and (5) basically sound, diagnostic photographs of the Nearctic taxa.

The size of the book and these novel sections thus produce a powerful,
impressive first feel. But that’s where the buck stops, because these good
portions are permeated and surrounded by some really bad ones.

The most horrifying problem with the book is a nearly total absence of
accountability throughout: (1) no proper references in the text to the original

26(l-4):l-288, 1988

271

scientific literature; (2) frequent failure and/or incompleteness in delimiting
Scott’s observations from those made by others, leading de facto to inappropriate
attribution of much work; and (3) an insensitivity to providing such documenta¬
tion for the reader’s benefit and the scientific record, particularly for the many
new claims that are made.

Well into the book, on page 22, one excavates the first attributed piece of
information — “(J. Osborne)” — in regard to ligaments in the anal region of
larvae. Such incomplete parenthetical teasers are all the reader is ever offered
in the text of this book. Unfortunately, these are useless as reference fodder,
since the reader hasn’t the slightest clue as to whether they refer to published
papers (proper attribution would be the usual “[whomever, year]”), letters to the
author (proper attribution would be “[whomever, in litt.]”), or conversations or
telephone calls (proper attribution would be “[whomever, pers. comm.]”).

Glassberg’s review put it only slightly differently: “It is usually impossible to
know if some piece of information is 1) generally accepted as true by the
scientific community, 2) controversial and based on the publications of another
author, 3) based upon the published studies of the present author, or 4) based
upon the unpublished studies or opinions of the present author. [Points] 2 and 3
are unfortunate in that the reader accepts information as ‘true’ although it is
known to be controversial. [Point] 4, presenting information in a pseudo-
authoritative textbook form when no published, preferably refereed data exist
to support one’s statements is akin to intellectual dishonesty.” And Opler’s
review termed Scott’s referencing “unconscienably brief,” which clearly logs in
as a glowing compliment.

What useable citations are provided for the reader, then? Scott’s bibliography
numbers a scant 25, of which 22 simply are other general works. That leaves 3 to
cover the available original scientific literature on the North American lepidop-
teran fauna. Two of these prove to be papers by Scott; no references to hallmark
papers in lepidopterology, and no references to solid technical research.

What a ridiculous false economy of attribution! Opler & Krizek’s book cites
254 scholarly articles — a full order of magnitude more — for the eastern taxa
alone. Ferris & Brown (1981) cite 143 for just the Rocky Mountain region
species . . . and so forth. Never before have I read a comparably sized recent work
on any insect group that claims to he a standard in its field make so many novel
assertions that simply can’t be traced. Even if one entirely ignores these matters
of good science and ethics, a major referencing problem still remains — namely,
that the budding lepidopterist receives not a hint on where to turn in the
literature for further enlightenment.

In short, the lack of accountability renders the book essentially useless as an
educational tool. Appropriately, most serious workers have already all but
dismissed it in this context.

2. Biological & ecological considerations

Scott’s training is in butterfly behavior and ecology. Part I (Biology and
Ecology, pp. 9-118) of the book thus addresses his forte, and most of it is
essentially accurate in detail, though unreferenced.

Of special value are the extended discussions of the immature stages. The
larval keys, as noted earlier, are most welcome additions. Lacking is compar¬
able treatment of the eggs, with no mention being made of the available

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exhaustive Palearctic lepidopteran egg atlas by Doring (1955), and Hinton’s
(1981) vast scanning electron micrographic egg survey of all insects. Although
Immature Insects (Stehr, 1987) is too recent to have been cited, it is a volume
that the interested reader should consider browsing regardless.

The section “Evolution and the Fossil Record” (pp. 84-101) is chock full of
potentially revolutionary information for the systematist, if he/she can manage
to plow through it (here is where a few skillful tables would have saved much
paper), and can obtain further details as to its origin.

Despite its basic solidity, Part I is punctuated with “just so” fables (affec¬
tionately dubbed “offhand, undocumented zingers” in Shapiro’s review), many
of which prove to be absurd biological errors. An exemplar is the repeated
assertion that the V-shaped impressions of bird beaks not infrequently found on
wild-collected specimens indicate that “butterflies often escape when the bird
tries to swallow” (e.g., pp. 3, 70). Apparently, Scott doesn’t often consult studies
on moth biology published alongside those on butterflies in the Journal of the
Lepidopterists ’ Society. Therein, the morphology of bird beak-marks — which
represent voluntary releases on the part of the bird, for any of a variety of
reasons — and escape-tears are rigorously categorized, and their genesis
cleverly documented experimentally (e.g., Sargent, 1973, J. Lepid. Soc. 27:175-
192, and his 1976 underwing treatise, Legion of Night: note also the many
comparable studies on butterflies during the middle part of this century that
Sargent cites).

Lastly, consider the foodplant compilations. They are exhaustive, they are
largely accurate. But, on balance so have been the foodplant compilations in the
other recent butterfly books. Moreover, Opler & Krizek also adequately delimit
local versus regional preferences; Scott does not. Tietz (1971) gives us an
encyclopedic array of literature references with which we can trace foodplant
records as we wish; we can’t do so with Scott’s listings. Scott slaps Tietz’s cheek
several times and admonishes us to check “every reference” therein for errors,
flagging these records in his species accounts with a T superscript. But Tietz
(1971) never pretended to trap all or even most of the published errors — that
work was offered as a bibliographic tool. It would have been much more valuable
for Scott to have superscripted his own records in this book with an S, and to
have assigned other appropriate superscripts to original literature references
he considered valid, and to unpublished foodplant information contributed to
him by other lepidopterists.

3. The geography of Nearctic butterflies

Scott’s range maps often inappropriately fill in truly discontinous distri¬
butions. This is immediately clear in a number of instance with the eastern taxa
(e.g., Satyrium edwardsii in the upper midwest, Poanes massasoit in the
northeast, and Mitoura hesseli throughout its range), and a scan of the Opler &
Krizek maps suggests the problem could be endemic throughout Scott’s book.
Compare, for example, Opler & Krizek’s maps for other selected eastern
skippers (map 223 p. 229, map 241 p. 243) with Scott’s (bottom p. 440, top p. 450).

Of course, some discrepancies between these two books likely reflect advances
in regional sampling since 1984, when Opler & Krizek’s book came out. Other
discrepancies may point to old records, and/or possibly document extinctions
and range contractions over time, information of crucial importance. How would
we tell? In the texts would be where to root around for clarification. Let’s do so.

26(l-4):l-288, 1988

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The range information provided by Scott for Problema bulenta (p. 450), for
example, is so terse as to be misleading at best: “Habitat Lower Austral Zone
freshwater marshes.” Opler & Krizek offered instead (p. 242): “To date, the Rare
Skipper is known only from lower brackish reaches of the Wicomico River
(Maryland), Chickahominy River (Virginia), Cape Fear River (North Carolina),
Santee River (South Carolina), and Savannah River (Georgia).”

Consider Mitoura hesseli. The species’ sole larval host is Atlantic White
Cedar, Chamaecyparis thyoides. One can see from any recent tree treatise (such
as Elias, 1980, The Complete Trees of North America , pp. 132-133) that this
cedar has large gaps in its largely eastern coastal distribution. Opler & Krizek’s
map for Hessel’s Hairstreak shows a tracking of much, but not all, of the range of
this larval foodplant, with hopscotch-like jumps in the butterfly’s distribution
from Virginia southward (no surprise). Opler & Krizek’s map also correctly
shows no records from Connecticut, something that has vexed us residents for
years (a singleton was finally located in the east-central extreme of the state in
1986, by Chris Maier of the Connecticut Agricultural Experiment Station).
Scott doesn’t speak to the disjunctions in the cedar’s range, and fills in the entire
eastern seaboard for the hairstreak’s home territory, from northern Florida
clear up to southern Maine.

One might possibly excuse this sort of sloppy accounting if “poetic license”
were being taken with all the maps (a tack of debatable utility in and of itself in
anything other than a coffee-table book). However, Scott’s maps for western
species often sport minute zigs, zags, and holes. Thus, it wasn’t a matter of the
resolution capabilities of Stanford University Press’ printers either . . .

4. Taxonomic decisions

Like Shapiro, I heartily endorse a return to conservative generic nomencla¬
ture for North American butterflies. Opler & Krizek did so for the eastern
species, and now Scott blessedly finishes the task off for all the taxa. As Shapiro
also notes, generic nomenclature is as much art as it is hard science; it will
always remain a matter of taste.

Now, delimiting species boundaries for the great majority of our taxa has
never posed much of a problem, whatever flavour of taxonomy was practiced on
them over the years. North American Lepidoptera volumes treat this great
majority identically (as does this new book). But these easy taxa are really only
of passing alpha taxonomic interest — the volatile ones that resist initial
sorting are the creatures of intrigue. Within this troublesome residuum is
where one hunts for an author’s taxonomic mettle.

In Part III (pp. 157-504) Scott essentially adopts the following taxonomic
approach: (a) if allopatry and/or parapatry, then subspecies; (b) if (a) and/or
differences slight, and/or variation not eyeballable, then synonymize. He
summarily lumps the bulk of the tough Nearctic species and their subspecies —
and then some, even combining several old and new world taxa — with neither
justification nor deftness. As a colleague remarked to me, “it would have been
more helpful to point out to the reader where taxonomic controversy lies rather
than to try to hide it with sweeping proclamations lumping (many] taxa.”

Shapiro offers a good rebuttal of the lumping in western Hesperia , and
discussion of the problem of unilaterally lumping all Cupressaceae feeding
Mitoura save hesseli into gryneus (fortunately this is an instance where Scott
cross-references his foodplants to the various taxa, so that at least some of the

274

J.Res.Lepid.

intriguing host-specificity issues in these hairstreaks are still extractable).

Typical of the plight of other lumped taxa are the Chlosyne. Scott dumps (pp.
306-308) a bevy of names into the single pan- western species gabbii, including,
among others: gabbii, neumoegeni, sabina, acastus, vallismortis, dorotheyi,
whitneyi, and damoetus. Here Scott narrowly evades embarrassment, via a
footnote tucked into the bottom of page 307 at the last minute, in which he must
acquiesce that at least gabbii, acastus, and damoetus need to be retained at the
species level — as they always had been before him, from Emmel & Emmel
(1973) through Howe (1975) to Ferris and Brown (1981).

Even with its avowed emphasis on including biological facts about butterflies,
Scott’s book overlooks some well-known instances of sibling species awaiting
formal recognition. Consider the two virtually certain sibling Megisto in the
northeast. Here the staccato text is so brief that it’s wrong, as Scott (p. 237)
offers: “One flight, June-E July northward.” Opler & Krizek (p. 185) offer: “The
Little Wood Satyr is univoltine over most of its range . . . the two emergences in
Virginia and Pennsylvania (late and May through June and then early July to
mid- August) and some other locations that follow each other so closely strongly
indicate that two cryptic siblings must be involved.”

Shapiro’s review of the taxonomy ended by saying “Scott may be right a lot of
the time.” However, this begs the whole issue. Let me put it to you another way:
my wife is also “right a lot of the time” if I ask her to sort a box of butterflies (she’s
a pianist, with no training in lepidopterology, but owns a keen eye and a bolt of
common sense). The point, of course, is that every one of you readers is also “right
a lot of the time.” You’re all expert taxonomists to a certain degree. I would
never hold my wife to figuring out the tough taxa in that box of butterflies . . . but
you should insist that an author of a major treatise be able to work deftly with all
the taxa — not just the trivially simple ones — and show temperance, and a
capacity for something other than summary execution of names.

5. What the book offers the butterfly enthusiast of the 1980s

Where has lepidopterology been during the past decade, and where is it
heading? For one, Lepidoptera conservation has gone through its painful
growing stage and come into its own, and is gathering more steam as time moves
past. Personal and community awareness of the joys and benefits of butterflies
are soaring, thanks to the path-breaking efforts of Bob Pyle and others.
Butterfly photography is booming, as is butterfly watching and gardening. And
natural history work, notably rearing, is taking hold again. Popular interpre¬
tive books are all the recent rage.

This represents a fundamental shift in emphasis in North American lepidop¬
terology. We are beginning to experience what the British and others in the
Palearctic have in their recent past history. What does Scott’s book say to these
trends of the times? Essentially nothing.

Absent from the book are segments on butterfly gardening, and scarcely a
paragraph can be found on awareness, watching, and photography. Discussion
of these aspects of butterflies’ lives is what one might expect from a properly
executed “Natural History” (witness the subtitle of this book). Opler’s review
torpedoed Scott’s passages on Conservation and Extinctions (pp. 110-111) as “a
rationalization that says collecting of rare species is all right and condones
secrecy for local colonies of rare butterflies. This unbalanced discussion does not

26(l-4):l-288, 1988

275

mention endangered species efforts on behalf of insects, completely omits the
Xerces Society, and fails to mention the butterfly habitat conservation accom¬
plishments of the past ten years.” I had hoped that Scott’s “The Butterfly
Census” (pp. 111-112) would be a discussion of the utility of butterfly counts and
other relative censusing procedures, but instead I found it to contain statistics of
lukewarm interest, such as guesses of the number of individual butterflies
currently residing on the planet.

6. Conclusions

What do I recommend? If you already own the other books mentioned above,
then I can’t advise adding this one to your library. Only Parts I and II of Scott’s
volume have lasting value, and even so, the lack of accountable referencing
palls their use as sources for science and otherwise. I will recommend that you
think seriously about adding Opler & Krizek (1984) to your library, even if you
presently live west of their covered region. They established a true standard for
all of us. Future butterfly books for North America should follow their healthy
lead.

7. Postscript

Stanford University Press deserves a healthy reprimand for letting such a
heterogeneous volume out of the house by itself without further review. At the
barest minimum, their editors should have stripped the pompous crowing from
the Preface and elsewhere — and replaced those passages with appropriate
modesty and circumspection, as befitting any large book in any field.

Stanford University lepidopterists have certainly sung loud songs in the
1980s about the supposed poor merit of other major works on North American
butterflies (see the common names and generic nomenclature squabbles in this
journal and elsewhere). I guess “what goes around comes around” — Stanford
should wag its tail between its legs and beat a hasty retreat, now that the Scott
book wears the name of that institution on its spine. Oh, incidentally, I happen
to be an alumnus of those same Stanford butterfly laboratories, but happily cast
my allegiance elsewhere in this instance.

Lawrence F. Gall Entomology Division Peabody Museum Yale University New
Haven , CT 06511 USA

Like the migrating monarchs which return annually to their besieged
Mexican wintering grounds, lepidopterists themselves make a similar pil¬
grimage to the bookstores. They return with gutted wallets to their infarcted
bookshelves with yet another butterfly book. This obsession with acquiring books
on butterflies is rivaled only by butterfly collecting itself, so aptly compared to
masturbation in an article we have long since lost. That author suggested that
butterfly collecting certainly feels good, but you’re really not sure why. You
really don’t want anyone to see you doing it. But, nevertheless, when you’re
done, you know you’re going to do it again.

We need to crowbar a space between a multivolumed “butterflies of the world”
by a creationist and a couple of previous “butterflies of North America” one by a

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minister and another by an artist. (Actually, we can now get rid of the
latter — the one with, inexplicably, the neotropical butterflies on the cover, and
Mexican butterflies for the frontpiece.) Voila, just enough space for the best book
to date on North American butterflies. Refreshingly, this one is by a biologist,
frankly all too rare an event. Very simply, Jim Scott knows more about his
subject than anyone else alive. Scott also might euphemistically be called a free
spirit. His unique opinions, which pervade this book, will amaze and enrage just
about everybody.

The text is incredibly rich in detail on many aspects of natural history and
butterfly biology. The larval host plant lists are truly awesome, seemingly every
published record and many more are included. The introductory chapters are by
far the best of their kind anywhere. All this in Scott’s typically turgid prose
which is scattered with the Scottisms for which he has often been criticized.
Butterflies “sipping mud,” indeed.

A rigorous text on North American butterflies will, by necessity, fall
somewhere between research reports in professional journals and a simple
guide with species descriptions. This sophisticated work, in comparison to
previous treatments, leans in the former direction. The decision, therefore, to
forego citations and bibliography, we think, was a mistake. Although some
ideas and observations are attributed to their sources with an initial and last
name, many, even most, are not. This gives the impression that a given
statement is common knowledge in butterfly biology, or that the source is Scott
himself — frequently neither are the case. Worse, many illustrations bear
striking resemblence to ones in previous works, a debt which is generally not
adequately acknowledged.

By numbering the butterfly species he includes, Scott cannot help but send the
reader to Miller and Brown’s 1981 checklist for comparison. Where Miller and
Brown list 763 species, Scott finds just 679, a decrease of 12%. This is not due to
five years of rampant extinction among the butterflies, but instead, to the most
dazzling exercise in taxonomic lumping of all time. Indeed, that large drop in
species is buffered by inclusions of several newly recorded southern migrant
species, the recognition of several new sibling taxa, and the amalgamation of
several nearctic taxa with palearctic species. Thus the decrease in species-level
taxa is actually on the order of 16%, much the result of a number of truly thought
provoking “new” combinations, which will have many lepidopterists doing
homework.

The rationale presented for several of these new treatments is rich and solidly
argued; for instance, that for one of Scott’s pet groups, the Callophrys. For other
species, such as Euphydryas chalcedona and Papilio glaucus, the lumping
reflects common sense and a recognition that there is insufficient information to
support specific status for many long-recognized taxa. Yet the linking of some
putative species into widespread, exceedingly variable superspecies, such as
Mitoura gryneus and Coenonympha tullia, will surely invite some argument.

With Scott’s taxonomy and nomenclature in print (which largely supports
that of Opler and Krizek’s recent book) now we can finally bury the ludicrous
nomenclature that brought us “Gaides,” “ Hyllolycaena ,” “Artogeia” and the
rest. We strongly suggest that Scott’s work constitute the guiding nomenclature
for all students of the North American butterflies. The fractionation of genera
and blanket acceptance of poorly supported species descriptions in the checklists

26(l-4):l-288, 1988

277

of Miller and Brown and the field guides of Pyle and of Tilden and Smith can now
be laid to rest.

Scott, however, has picked an unfortunate time to become extraordinarily
cavalier about subspecies. Notwithstanding arguments over the “value” of
subspecies and their evolutionary meaning, the subspecies is recognized by the
United States Fish and Wildlife Service as the lowest invertebrate taxonomic
category which can be conferred protection under the Endangered Species Act.
The practice of wholesale lumping of subspecies has but one real world result —
our ability to conserve those entities it compromises. To suggest, for instance,
that Euphydryas editha consists of but three ecological and genetic segregates
that are phenetically separable into subspecies is preposterous. Scott wants the
range of Euphydryas editha editha to encompass all of California west of the
Sierra Crest. But Sierra foothill populations differ from coastal populations in
flight phenology (more than a month), in host choice, in oviposition behavior,
and by wing markings, which differ dramatically enough that effectively every
specimen can be assigned to its foothill or coast source at a glance. We are not
sure what Euphydryas editha rubincunda means in an evolutionary context
(whether or not it is a species in statu nascendi), but assuredly it is an entity
sufficiently genetically distinct from Euphydryas editha bayensis to warrant
subspecific recognition. Hardly a coincidence, all of the subspecies named
previously by Scott himself are included, and virtually all names in his
“favorite” genera are acknowledged (few of which are significantly better than
of many which he has dispensed).

For most of us blindly driven butterfly book collectors, the most important
feature of any work is the quality and accuracy of the figures. One of our
colleagues has suggested that in Scott’s book, “the illustrations suck.” That is,
perhaps, a bit strong. The illustrations, however, do not make for a really lovely
coffee-table book. The plates are crowded, the orientation of specimens is
inconsistent and many are damaged. But, having seen the original prepublica¬
tion prints, the reproductions ended up surprisingly good. Arguably they are
more useful for identifying specimens in hand than the figures in most other
books. (Which naturally brings up the subtitle, claiming that this is a “field
guide.” We, for two, are having a bit of difficulty figuring out into which pocket
this three pounds of “field guide” is supposed to fit!) The plate legends and figure
arrangements, however, are really cumbersome. While finding the species
writeups from the illustrations is simple, linking text to plate is a royal pain. It
is nice however, finally, to see a butterfly book include lots of photos of living
butterflies, larval stages, and eggs.

But, more difficult to follow yet are the identification keys. Instead of using
traditional couplets, Scott’s keys list traits with a reference number at the end of
each intended to guide the reader. This make it almost impossible to enter the
key at any point except at the beginning. Most enthusiasts would probably know
the difference between a papilionid and a nymphalid, but separate keys for
major groups of butterflies are not easily located within the cumbersome
general key.

Acknowledging that by the time this review appears, all real butterfly buffs
will have this book (and it probably will be remaindered, for that matter), we
nonetheless toss in our nickle’s worth that this will probably be the book on
North American butterflies for a very long time. We can’t imagine anyone

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trying to follow, or wanting to follow, Scott in such an endeavor.

We do, however, additionally suggest that before throwing away Howe’s book,
that you save the color plates which, when laminated, make particularly festive
placemats for those special occasions.

Dennis D. Murphy and John F. Baughman, Department of Biological Sciences,
Stanford University, tanford, California 94350

A FIELD GUIDE TO WESTERN BUTTERFLIES. Tilden, J. W. and A. C.
Smith, 1986. 370 pp., 48 plates (24 in color) plus text figures; Houghton Mifflin
Company, Boston. Price $12.95, soft (also available in hardback).

Timing may not be everything, but the appearance of Tilden and Smith’s
“new” book sandwiched between the publication of the two best recent books on
North American butterflies by Paul Opler and George Krizek (1984) and Jim
Scott (1986) certainly will not add to its popularity. A Field Guide to the Western
Butterflies has the standard Peterson Field Guide Series format with introduc¬
tory chapters, species accounts, plates (with the familiar arrow system pointing
to key identifying marks), and appendices including a checklist with “all”
western butterflies and their subspecies. It covers North America west of the
100th meridian (with Alaska and Hawaii) and “partial coverage of northern
Mexico.”

As a western version of Alexander B. Klots’ ( 1951 ) landmark A Field Guide of
the Butterflies of North America, East of the Great Plains, this book has been long
overdue, and should fill a glaring gap in the Peterson series. But, Tilden and
Smith’s book apparently has been in the works for well over a decade and
unfortunately it shows . . . over and over again. The last decade has seen an
explosion of information on the biology of butterflies. That explosion has been fed
by an immense increase in knowledge of the distributions and evolutionary
relationships of our butterflies. A number of state and regional monographs
(Emmel and Emmel 1973, Dornfeld 1980, Ferris and Brown 1981) and a variety
of checklists and research papers on the western butterflies make these insects a
very well known group in a well studied region. In this light, Tilden and Smith’s
work is something of a relict; the amount of quite readily available information
lacking is really shocking.

Considering all the recent attention paid to North American butterfly
taxonomy, we cannot be more disappointed that one more book is using an
unacceptable nomenclature. This situation is not only unnecessary, but is
particularly ill-advised in a guide for lay persons. There is virtually no explana¬
tion of the origins of Tilden and Smith’s taxonomic treatment except on page 1
where they stated that they followed Klots’ sequence of families. Not so. Klots
recognized 11 families and followed Nymphalidae with Libytheidae whereas
Tilden and Smith inexplicably discuss 12 families and place the libytheids
between the Pieridae and Riodinidae. The order of families used is hardly most
important but it succeeds in thoroughly obscuring relationships among butter¬
flies as we know them. The recognition of 12 families of butterflies itself is totally
unsupported. The evidence is abundant that butterflies belong at most to 5 or
perhaps 6 families (Ehrlich 1958, Ackery 1984, Scott 1984).

26(l-4):l-288, 1988

279

The nomenclature appears to follow Miller and Brown (1983). Yet as Opler and
Krizek’s and Scott’s recent books indicate, responsible workers continue to use a
much more conservative nomenclature. Tilden and Smith have never met a
taxon they didn’t like at any taxonomic level. They include the widely discounted
genera “Occidryas” , “Artogeia” , “ Hyllolycaena ”, and “Falcapica”. Their pre¬
sentation of the Euphydryas is typical. That generic name cannot be found
except with a notation stating that the name was “ replaced ” in 1978. Higgins
(1978) provided no evidence whatsoever for his generic demolition. Nonetheless,
ignored from the same year was the much more careful work of Scott (1978)
suggesting that Euphydryas anicia, E. chalcedona , and E. colon form a rassenk-
reis and hence constitute a single species.

If the Euphydryas species are confusing in this book, the presentation of
Euphydryas below the species level is positively stultifying. No fewer than 21
subspecies of E. editha and of E. “ anicia ” are listed in the “checklist/life list.” Yet
only three of each are illustrated and discussed in the text. Those three account
for but a percent or so of the overall distribution of either species. In contrast, all
nine checklist subspecies of E. “ chalcedona ” are presented in the text, but only
two are illustrated. How, for instance, E. c. kingstonensis (“salmon-colored, dark
markings small”) is to be distinguished from neighboring E. c. corralensis (“spots
light red, dark markings narrow”) is not clear. The lay person who could identify
a checkerspot to subspecies has to be using another book.

Unfair example? We don’t think so. In the western states, rimmed and divided
by young mountain ranges, confounding species groups are hardly the excep¬
tion. Euphilotes, Chlosyne, Speyeria, Phyciodes, Pontia, Euchloe, Erynnis,
Thorybes, Hesperia , and the “ machaon ” swallowtails are but a few of the
troublesome groups containing hundreds of taxa. For these and other genera
and species groups, this book will confuse rather than guide in the field.

The treatment of subspecies in general is terribly uneven. Granted, some were
described too late to be included. But where is Plebejus shasta charlestonensis
(Austin 1980)? Scott (1981a) named 18 new subspecies, but only 7 of these are
included. Most (or all) the other eleven represent very distinct phenotypes but
are omitted for some unknown reason. For some species all recognized subspe¬
cies are mentioned in the species accounts, their distributions given and they are
briefly characterized. For others, there is no mention except that there are some
similar subspecies. Still others are grouped as sets of similar phenotypes.

Arguments for and against the use of common names have been well aired but
one thing on which all agree is that for common names to be useful, they must be
standardized (Murphy and Ehrlich 1983a, Pyle 1984, Austin 1985a). This book
certainly is not a step in that direction. Tilden and Smith present 41 species of
satyrines corresponding to those in Pyle (1981) but only 23 of these share the
same common names. Considering the principally lay audience of this book, this
lack of nomenclatural concordance is hardly trivial. Tilden and Smith go on to
give each subspecies a common name as well. Suffice to say that even the bird
people did away with that attempt long ago.

Not surprisingly, these points alone add up to an immensely confusing field
guide, which fails at the most fundamental level for a field guide, it does not
facilitate the association of names with organisms in hand. For example, the
guide is useless in the field for separating any of the Euphydryas species (less the
extremely distinct E. gillettii). As Murphy and Ehrlich (1983b) pointed out,
Euphydryas editha usually may be distinguished from members of the E.
chalcedona complex by an easy diagnostic feature — the former lack white spots

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J. Res. Lepid.

on the dorsal portions of the plural region of the abdomen. That simple “in hand”
character solves the confusion nearly everywhere where species overlap in the
west.

Then again, perhaps, we should give Tilden and Smith the same break we
might give A Field Guide to the Stars or A Field Guide to the Animal Tracks.
Actually, many of our complaints with their book center on subspecies level
information, and no other contribution in this series attempts to do so much at
that level of taxonomic resolution. A Field Guide to the Western Birds, for
instance, dabbles merely with the very well differentiated subspecies. But, while
that does limit the information content, the information included is largely
correct.

A Field Guide to the Western Butterflies, by comparison is a veritable
cacophony of errors when it comes to the distributions of butterflies. Many of the
errors simply repeat old ones, which suggests that the authors have not kept up
with the literature. Taking, for example, an area with which both of us are
familiar — the state of Nevada — you’ll get an idea of the extent of the
distributional disinformation. Although Austin and Austin ( 1980) is listed in the
bibliography, a substantial proportion of their distributional records are not
included. Other information is somewhat newer but certainly was available (e.g.,
Ferris and Brown 1981, season summaries of the Lepidopterists’ Society, etc.).
Anyway, despite what Tilden and Smith say, Cyllopsis pertepida, Calephelis
wrighti, Ministrymon leda, Callophrys comstocki, Euphilotes battoides martini,
E. enoptes dammersi, and Systasea zampa are all resident in southern Nevada
and Coenonympha California, Anthanassa texana, Phyciodes phaon, Heliopetes
domicella, and Chiomara asychis all stray to southern Nevada. Basilarchia
archippus lahontani is restricted to northern Nevada; those elsewhere are a
different phenotype. The hybrid B. weidemeyerii latisfascia X B. lorquini
lorquini occurs at several locations in western Nevada. Speyeria egleis toiyabe
occurs in central Nevada and not north-central Nevada. Anthocharis cethura is
not in central Nevada but in southern and western Nevada (A. pima synony¬
mous). Polydryas arachne occurs in southeastern Nevada. Hypaurotis crysalus
is known from eastern Nevada. Callophrys affinis occurs to central Nevada.
Incisalia fotis occurs in parts of western Nevada. Lycaena nivalis near browni
occurs in northeastern Nevada. Plebejus lupini occurs east at least as far as
eastern Nevada. Euphilotes enoptes ancilla is more than Rocky Mountain in
distribution, occuring through much of northern Nevada (see also Shields
1977a). E. rita pallescens is not in southern Nevada but in central Nevada (see
also Shields 1977b). E. spaldingi is in eastern Nevada. Megathymus color-
adensis is only in southern Nevada. Euphyes “ ruricola ” (= E. vestris ) is on the
east slope of the Sierra Nevada in extreme western Nevada. Hesperia lindseyi is
in northwestern Nevada. Pseudocopaeodes eunus occurs also in central and
western Nevada. Erynnis propertius is in extreme western Nevada (all Austin
1985b). Cercyonis pegala blanca is a synonym of C. p. stephensi and the latter is
not a form of C. p. ariane — C. p. ariane does not occur in Nevada. Polygonia hylas
is a subspecies ofP. fa unus (Ferris and Brown 1981). Speyeria zerene pfoutsi is a
synonym of S. z. platina (see Grey and Moeck 1962). The Speyeria mormonia
mormonia phenotype is not found in Utah and S. m. arge is a synonym.
Charidryas palla vallismortis is not a C. palla subspecies but closer to C. acastus
(fide J. F. Emmel). Thessalia leanira cerrita is a synonym of T. 1. wrighti (the
types were both from virtually the same place) and the Mojave Desert popula-

26(l-4):l-288, 1988

281

tions presented as “ cerrita ” are unnamed. The type locality of Neophasia
menapia is not in Utah, but probably in western Nevada since Pierre Lorquin
was never in Utah (see Miller and Brown 1981, Scott 1981a). Apodemia palmerii
palmerii and A. p. marginalis are synonymous. Calephelis nemesis dammersi is
the summer form of C. n. californica. The form “ines” of Ministrymon leda also
occurs in the early spring. Satyrium dryope does not occur in Mono County,
California; what is there is a large, pale, and sometimes tailless S. sylvinus.
Plebejus icarioides ardea is a synonym (actually an aberration) of P. i. fulla.
Hesperia pahaska occurs in S. Nev. not “S. Neb.” A number of taxa are attributed
to the Nevada fauna which apparently have no basis in fact: Oeneis nevadensis,
Lycaenax. xanthoides, L. mariposa and Poanes taxiles. And, it is certainly a lapse
to separate Anthocharis and Euchloe from Pieris (s.l.) by the Coliadinae.
Anthocharis and Euchloe belong together with other whites in a single subfami¬
ly, the Pierinae, distinct from the Coliadinae (Geiger 1980).

These are problems culled on a quick glance from the standpoint of just one
state!

We have not checked all the hostplants listed but spot checking some of those
identified as mistakes by Scott (1981b) shows that many are again repeated
incorrectly. More obvious, and important, to a field guide are the mistakes in
habitat associations. Speyeria egleis, in Nevada, is a hilltopper and an edge
species, occurring along shaded roads and openings. Speyeria mormonia is
montane occurring mostly in wet meadows. Phyciodes orseis herlani does not
occur in “mountain meadows”; it is found along wood edges and in forest
openings. Chlosyne neumoegeni uncommonly occurs in desert scrub, but nearly
always along washes. Chlosyne lacinia is common in agricultural areas. Phoebis
sennae is a desert species in parts of its western range. Great Basin Anthocharis
sara are mainly found in the pinyon-juniper woodlands. Callophrys lemberti also
occurs along washes and hillsides at relatively low elevations in the Great Basin.
Polites sabuleti tecumseh occurs in upper montane and subalpine meadows and
is not restricted to the alpine. And, what is meant by “Occidryas“ (these are all
Euphydryas, Brussard et al. 1985) chalcedona being “a dominant in its range”?
The statement certainly has no biological meaning.

And the illustrations, do they really measure up to the standards of a good field
guide? Tilden and Smith might get a gentlemen’s “C” on that front. Most of the
black and white plates are clear, although some such as Plate 25 are a little fuzzy.
The outlines of many of the more pale butterflies were darkened to make them
stand out against a white background. This looks unnatural and easily could
have been cured with a darker background. Some, such as Plate 44, look a little
dark overall. The Peterson Field Guide Series, as a whole, generally has
excellent color but this one suffers. Several (such as Plates 9, 12, 14) are far too
yellow. Plate 21 is far too pale, the lycaenid plates are either too dark or are off
color. Plate 13 figure 2 looks to be a Speyeria coronis rather than S. zerene and S.
callippe nevedensis is much greener than figure 6. Plate 14 figure 4 is a pale S.
mormonia eurynome (S. m. mormonia is not in Utah). All the plates lack the
crispness of the plates in other field guides of this series.

For our money, we’ll take Bob Pyle’s field guide in the Audubon series, or the
Ehrlichs’ ( 1961 ) classic. Neither tried to do as much; both did what they did well.
And, we will continue to look forward to Opler giving us a “West of the Great
Plains” as good as the “East of the Great Plains”. It won’t fit in our pockets, but at
least we’ll wish it did.

282

J. Res. Lepid.

Literature cited

ACKER Y, P. R. 1984. Systematic and faunistic studies of butterflies, chapter 1 in
The Biology of Butterflies, Academic Press, New York.

AUSTIN, G. T. 1980. A new Plebejus ( Icaricia ) shasta (Edwards) from southern
Nevada (Lycaenidae). J. Lepid. Soc. 34: 20-24.

AUSTIN, G. T. 1985a. Commentary: proliferation of common names. News Lepid.
Soc. 1985(1): 2.

AUSTIN, G. T. 1985b. Nevada butterflies: preliminary checklist and distribution. J.
Lepid. Soc. 39: 95-118.

AUSTIN, G. T. AND A. T. AUSTIN. 1980. Butterflies of Clark County, Nevada. J. Res.
Lepid. 19: 1-63.

BRUSSARD, P. F., P. R. EHRLICH, D. D. MURPHY, B. A. WILCOX AND J. WRIGHT. 1985.
Genetic distances and the taxonomy of checker spot butterflies (Nymphali-
dae; Nymphalinae). J. Kansas Ent. Soc. 58: 403-412.

DORNFELD, E. J. 1980. The butterflies of Oregon.. Timber Press, Forest Grove, OR.
EHRLICH, P. R. 1958. The comparative morphology, phylogeny and higher clas¬
sification of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci.
Bull. 39: 305-370.

EHRLICH, P. R. AND A. H. EHRLICH. 1961. The butterflies. W. C. Brown, Dubuque,
Iowa.

EMMEL, T. C. AND J. F. EMMEL. 1973. The butterflies of southern California. Nat.

Hist. Mus., Los Angeles Co., Sci. Series 26.

FERRIS, C. D. AND F. M. BROWN. 1981. Butterflies of the Rocky Mountain states.
Univ. Oklahoma Press, Norman.

GEIGER, H. J. 1980. Enzyme electrophoretic studies on the genetic relationships of
pierid butterflies (Lepidoptera: Pieridae) I. European taxa. J. Res. Lepid. 19:
181-195.

GREY, L. P. AND A. H. MOECK. 1962. Notes on overlapping subspecies. I. An example
in Speyeria zerene. J. Lepid. Soc. 16: 81-96.

HIGGINS, L. G. 1978. A revision of the genus Euphydryas Scudder (Lepidoptera:

Nymphalidae). Entomol. Gazette 29: 109-115.

KLOTS, A. B. 1951. A field guide to the butterflies Houghton Mifflin, Boston, MA.
MILLER, L. D. AND F. M. BROWN. 1981. A catalogue/checklist of the butterflies of
America north of Mexico. Lepid. Soc. Mem., no. 2.

MILLER, L. D. AND F. M. BROWN. 1983. In Hodges, R. W. et al. (eds.), Check list of the
Lepidoptera of America north of Mexico. E. W. Classey, London.
MURPHY, D. D. AND P. R. EHRLICH. 1983a. Opinion — Crows, bobs, tits, elfs and
pixies: the phoney “common name” phenomenon. J. Res. Lepid. 22: 154-
158.

MURPHY, D. D. AND P. R. EHRLICH. 1983b. Biosystematics of the Euphydryas of the
central Great Basin with the description of a new subspecies. J. Res. Lepid.
22: 254-261.

OPLER, P. A. AND G. O. KRIZEK. 1984. Butterflies east of the Great Plains. John
Hopkins University Press, Baltimore, MD.

PYLE, R. M. 1981. The Audubon Society field guide to North American butterflies.
A. A. Knopf, New York.

PYLE, R. M. 1984. Rebuttal to Murphy and Ehrlich on common names of
butterflies. J. Res. Lepid. 23: 89-93.

SCOTT, J. A. 1978. A survey of the valvae of Euphydryas chalcedona, E. c. colon,
and E. c. anicia. J. Res. Lepid. 17: 245-252.

26(l-4):l-288, 1988

283

SCOTT, J. A. 1981a. New Papilionoidea and Hesperioidea from North America.
Papilio (new series), no. 1.

SCOTT, J. A. 1981b. Book review: Butterflies of the Rocky Mountain states, by C.

D. Ferris and F. M. Brown. J. Res. Lepid. 20: 58-64.

SCOTT, J. A. 1984. The phytogeny of butterflies (Papilionoidea and Hesperioidea).
J. Res. Lepid. 23: 241-281.

SCOTT, J. A. 1986. The butterflies of North America, a natural history and field
guide. Stanford Univ. Press, Stanford, CA.

SHIELDS, 0. 1977a. Studies on North American Philotes (Lycaenidae). V. Taxono¬
mic and biological notes, continued. J. Res. Lepid. 16: 1-67.

SHIELDS, 0. 1977b. Distribution on Shijimiaeoides rita , especially S. r. rita and S.
r. coloradensis (Lycaenidae). J. Res. Lepid. 16: 162—172.

George T. Austin, Nevada State Museum and Historical Society, 700 Twin Lakes
Drive, Las Vegas, Nevada 89107 and Dennis D. Murphy, Department of
Biological Sciences, Stanford University, Stanford, California 94305.

BUTTERFLIES OF THE HIMALAYA. Mani, M. S. 1986. 181 pp. Dr. W. Junk,
publisher, Dordrecht. Price: $79.50.

Victims of crime often speak of feeling “violated.” To buy this book is to
invite that feeling. The price, $79.50, is not a misprint. It comes out to
440/page, which is about what the most expensive technical monographs run
nowadays. Professionals — scientists, scholars, physicians and the like — have
gotten used to paying such prices. Publishers justify them on the ground that
technical monographs have small audiences and small press runs and are thus
very expensive to produce. The prices are softened a bit by our ability to write
such expenses off on our income tax returns as “professional expenses,” and the
availability of grant money to buy publications needed for research. Even as
prices skyrocketed in the past decade, the esthetics of technical books deter¬
iorated. To save money, fewer and fewer such books are actually set in type.
More and more, they are printed by photo-offset from typescript (“camera-ready
copy”); they thus look like legal briefs between covers and are ugly and jarring to
the eyes of those who truly love books. We have gotten used to that, too, in the
same sense as one of my math teachers once informed the class (referring to
homework) that “you can get used to hanging if you do it long enough.”

Butterflies of the Himalaya is set in type, and that is about the only good thing
one can say about it. Because it actually looks like a book, one is unprepared for
what lies beyond the title page. There are many words to describe this book, and
most of them are unprintable.

There is no evidence that the manuscript was ever copy-edited or the proofs
checked for typos — at least not by anyone who reads English. In a haphazard
sample of ten pages I found over 5 typos per page (many of them both egregious
and obvious) and 2 grammatical errors, usually involving omissions of particles
or the like. The impression of slovenliness is further compounded when one gets
to the illustrations.

There are a few color photos of habitats, reduced to the size of a block of four
postage stamps and virtually useless. Anyone who wants a visual impression of

284

J. Res. Lepid.

the Himalaya would do much better with a backfile of National Geographic. The
captions are confusing at best. The term “steppe” is used there and in the text for
what we would call “woodland,” as in California foothill woodland. The photo of
the “abode of high-altitude butterflies” shows a snowfield. The “high-altitude
meadow with Thermopsis in bloom, the home of Colias butterflies,” is a close-up
of Thermopsis in flower. There is a picture of “Papilio butterflies congregated in
swarms.” There is no explanation of why they are congregated in swarms.
Maybe they really are Danaid-mimicking Graphium (not Papilio; Mani himself
uses Graphium), but they look like Danaids to me and are acting like them, too.

After this the illustrations deteriorate rapidly. Many of the specimens figured
are abominably set. None has any data given. Even the sexes are not specified.
The Parnassius are shown in color, photographed against a distracting sky-blue
background full of vague shadows. The black and white illustrations defy
description. There are four photos to a page, arranged in a rectangle 2x2
starting at the bottom of the page. They are to different (unspecified) scales;
some are out of focus; and — unbelievably — some are cut off at the edges, like
the headless snapshots of Uncle Harry at the family reunion. But the best is yet
to come: the caption-identifications are very nearly random vis-a-vis the
animals illustrated. Sometimes the names are in the wrong positions; some¬
times they refer to photos on different plates; and sometimes they refer to
species not illustrated at all. Consider the Pieridae. On plate XIV, “Pontia
daplidice” is actually a Euchloe; the real daplidice appears on the same plate as
P. glauconome; on plate XV “Pieris dubernardi” is actually Pontia callidice
kalora , and “Pontia callidice ” is an Appias; on plate XVI “ Euchloe ausonia” is a
Pieris. . . and so it goes.

The taxonomy in the text is nearly random, too. On p. 34 under the heading
“Zerynthidae” (note family ending), the first sentence begins “The butterflies of
this subfamily are recognised ...” and the paragraph concludes a few lines later
with: “This was formerly considered a subfamily of Papilionidae.” I will not try
to correct all the equally absurd errors, but to give the flavor of this magisterial
work, I note that Argynnis (p. 121) includes Brenthis; Limenitis is consistently
misspelled Liminitis; Vanessa (pp. 118-120) incorporates Vanessa sens. str. as
well as Nymphalis, Inachis, and Polygonia (!). . .and so it goes.

The distribution maps show no physiographic features — just a schoolboy’s
outline map of the Indian subcontinent and Southeast Asia with a handful of
symbols; the lettering is childish and the symbols are indistinctly drawn
freehand. Rarely have so many uncircular dots appeared together in public.

The Hesperiidae are not included.

I bought this book because the publisher’s advertisement promised a con¬
cluding synthesis of biogeography. Its principal point — that the Himalaya has
a remarkably unbalanced, undiverse, immature fauna with nearly all the
endemism confined to the alpine zone - — could have been made in a five-page
paper, and should have been.

Mani did us all a great favor 25 years ago by publishing his Introduction to
High-Altitude Entomology, My copy, now worn and tired, literally guided me
into the high-altitude research I now do in the Andes, though as I got further and
further into my work I realized more and more that much in Mani’s book is
inadequately documented and even dead wrong. Still, I know he knows his
Himalaya, and I deeply regretted his failure to attend and present his scheduled
paper at the Latin American Congress of Zoology in Arequipa. I thought this

26(l-4):l-288, 1988

285

book would substitute. But this book is a disgrace to Mani and to the publisher.
It should never have appeared in book form at all; it could easily have been
published as a long faunistic paper in any of several respectable British or
Indian journals, in which form it could have reached its proper clientele at
sensible cost and under better editorial control. Once a decision was made,
however, to publish it as a book the publisher had an obligation to assure quality
at all levels, and this he failed to meet. The book was actually set and printed by
IBH Publishing in New Delhi, but it bears Junk’s imprint and Junk is
responsible for it. Junk used to be a very respectable, if always pricey, publisher
of important technical monographs. Now, apparently, it is living up to its name.
It is still pricey, but it now publishes junk.

Anyone who buys this book is an accomplice to his own victimization. As a
professional, I can write it off on my tax return — but I am almost ashamed to
admit I bought the damned thing.

Arthur M. Shapiro, Department of Zoology, UC Davis, Davis, CA 95616.

BREEDING OF BUTTERFLIES AND MOTHS

Ekkehard Friedrich, 1986. Translated from German by S. Whitebread, supple¬
mented and edited by A. M. Emmet. 176 pp., 46 figs.; Harley Books, Colchester
(Great Britain). ISBN 0 946589 20 8. Price £5.95 (hardback).

Friedrich’s “Handbuch der Schmetterlingszucht” (2nd edition 1983) has become
a “classic” for all breeders of butterflies and moths in German speaking
countries. The present first English edition is rather late on the market. The
book consists of two distinct parts. In the first part (“Basic principles”) the
methodological description of breeding of butterflies and moths is provided; it
includes topics as breeding equipment and techniques, hand pairing, cages,
storage of eggs and pupae, hibernation,, transport, conservation, artificial and
semisynthetic diets (the last topic rewritten by B. O. C. Gardiner) etc. The
second part of the book (“Rearing descriptions”) gives detailed descriptions of
breeding many European Lepidoptera, species by species, divided into three
subsections: butterflies, “larger” moths and “micro” moths (the last subsection
written by A. M. Emmet). The book closes with a useful select bibliography,
information on entomological societies, addresses of specialist booksellers,
suppliers of equipment and seedmen. The first part of the book under review is
well illustrated, but lacks information on some modern techniques and equip¬
ment (e.g incubators are now common in most laboratories and surely not out of
reach of many amateur lepidopterists: they enable the breeder to maintain the
desired temperature and humidity, the better models can maintain the desired
photoperiod and different night and day temperatures). Since the book contains
instructions on how to kill adult butterflies and moths, I would have expected to
find instructions on how to preserve early stages, particularly the caterpillars,
utilizing modern methods (e.g. freeze drying). All in all: Friedrich’s book is a
most welcome addition to lepidopterological literature in English language; it
will surely and deservedly become a best seller. Compared with the German
edition, it is much better produced (printing, quality of paper, contents) and the

286 J.Res.Lepid.

superior English hardback sells for less than a half of the price of the inferior
German paperback.

Otakar Kudrna, Rhenusallee 32, D-5300 Bonn 3, WEST GERMANY

WIESEN UND BRACHLAND ALS LEBENSRAUM FUR SCHMETTERLINGE.
(in German; English summary) Erhardt, Andreas. 1985. Birkhauser Verlag,
Basel, Boston, & Stuttgart. 154 pp., 2 color plates, 89 text-figs. sFr. 69.

The socioeconomic context of European agriculture is changing. Pastoral
methods established in the Alps over many centuries are being supplanted by
intensive fertilizing and mowing of the better pastures, and abandonment of
the poorer ones. Because so much of the diurnal Lepidopteran fauna of the
region is associated specifically with montane and subalpine meadows man¬
aged in the traditional ways, Andreas Erhardt undertook to examine the
impact of the new methods on that fauna. His results are alarming. They are
presented here in a magnificently-produced monograph which is a sophisti¬
cated ecological study, though superficially packaged as a coffee-table book.

Using adaptations of European phytosociological techniques (generally un¬
familiar to Americans, and coolly received by most plant ecologists this side of
the Atlantic), Erhardt characterizes the vegetation of traditional and modern
pastures and its correlation with Lepidopteran fauna. He finds that both
intensified fertilization and cutting cycles and abandonment are catastrophic
for many species, including some of special concern. Indeed, entire species may
be lost from the Alps if protective measures are not taken. Ironically, “protec¬
tive measures” in this case means setting aside some carefully selected pas¬
tures and managing them as they have been managed since Wilhelm Tell’s
day.

How is it possible that native butterflies and moths could owe their continued
existence to the maintenance of grazing disclimaxes ? Lepidoptera are, by and
large, not animals of climax forest. The common notions of “balance of nature”
and “pristine communities” are also exceedingly naive. Throughout the Qu¬
aternary (Pleistocene and Holocene = Recent), climate has fluctuated and with
it, plant and animal communities. Human impacts provide an additional layer
of disturbance, but that disturbance benefits some organisms as it damages
others. Thus, many meadow butterflies and moths of the Northern Hemisphere
are ill-adapted to the climax forest biomes currently dominant there. They
represent relicts of the drier steppe-tundra conditions around the continental
glaciers, and have kept going only through natural and man-induced deforesta¬
tion which produced local “mini-steppes” for them. The most stable of these
have probably been the grazing disclimaxes of Erhardt’s montane meadows,
and it should not be all that surprising to find them supporting rich and
specialized faunas. The British learned too late that grazing was prerequisite
for the persistence of their Large Blue, another beast at the edge of its climatic
tolerance. In that case, grazing acted by producing conditions favorable to the
ant species on which the Blue was dependent. A glimpse at the distributions of
the genus Maculinea quickly underscores how marginal M. arion was in
Britain. The same is true of many Alpine meadow species. Ironically, the
massive tree die-off in central Europe (which seems to be a synergistic effect of

26(l-4):l-288, 1988

287

pollution and climatic change) may yet come to their rescue.

Erhardt’s study must not be generalized glibly. In northern Europe a higher
percentage of species appears well-adapted to brush- or heathland, so abandon¬
ment of pasture could be a more favorable development there. In North
America we know almost nothing of these matters. In the West, fire suppres¬
sion has reduced the extent of successional habitat and increased that of
mature forest. It has also reduced runoff and hence adversely impacted natural
wetlands. One would predict a generally negative effect on butterflies, which
might be mitigated by logging and moderate grazing. On the other hand,
intensive grazing would be expected to have devastating effects in arid and
semiarid climates and in fragile alpine or subalpine meadows with peaty soils.
Erhardt’s study serves as a model of what can be done. Ultimately, it may help
us understand the global determinants of ecological diversity; presently, it
points the way to local investigations and prudent assessment of risk. It should
be read widely by ecologists, conservationists, and managers.

Unfortunately, the proverbial incompetence of Americans in foreign lan¬
guages will impede its full impact being realized here. There is a concise
one-page summary in English (p. 143) which is sufficiently tantalizing to get
the timid reader of German reaching for his dictionary. There is also what
amounts to a somewhat longer abstract published as an article in the English
literature (Erhardt, A. 1985. Diurnal Lepidoptera: sensitive indicators of
cultivated and abandoned grassland. J. Appl. Ecol. 22: 849-861).

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

THE MACHINERY OF NATURE. The Living World Around Us - and How It
Works. Paul R. Ehrlich. Simon and Schuster, New York, 320 pp. $18.95.

If your book budget for the year is $20.00, 1 suggest you spring for this tour de
force of exceptionally clearly written contemporary ecology. I was so taken with
the work that I read it through with complete attention in two sittings. Since
Ehrlich is probably our most prominent lepidopterist, the book is filled with
examples from his and his colleagues work with butterflies, in particular
Euphydryas editha. Ehrlich, no slouch as a popular science writer, here gives us
his most highly readible work to date. He generously ascribes a part of his
writing success to the input of many peers to whom he offers his work for
criticism prior to publication. I make this point to emphasize the value of having
ones work read and commented on by others.

Development of the book follows what would be a logical sequence in
constructing a course in general ecology, from the specific to the general, with
lots of exciting side trips. The flavor of the whole comes in the introduction,
subtitled “butterflies, ecosystems, and people,” wherein Ehrlich starts with a
checkerspot anecdote, goes on to state that nothing is more important today
than understanding how nature works, and concludes that a quasi religious
movement to alter popular awareness of nature may be our only hope in saving
civilization.

The first chapter deals with physiological ecology, the adaptive relationship of
individuals to their environment. Citing many of the exquisite attributes which

288

J.Res.Lepid.

deliver the miracle of survival he goes on to the question of why. Chapter two
starts with a butterfly story, the checker spot classic of Jasper Ridge, generalized
to an understanding of the forces regulating numbers in a population. Anyone
with the slightest real interest in Lepidoptera should know this as litany. The
text is all this while threading us through evolutionary theory and its most
recent arguments. Behavioral ecology, subtitled sex and society follows. Com¬
munity ecology is discussed in chapters entitled “Who lives where and why” and
“Who lives together and how.” Life support systems is last, with a concluding
essay as epilogue. The message is that natural ecosystems are rapidly being
destroyed and that we still know precious little about their detail of operation.
Ehrlich makes a call to action for public support and efforts of all informed
citizens to educate. We must all work together to amplify this call while there is
some bit of nature left.

Rudolf H.T. Mattoni, 9620 Heather Road, Beverly Hills, CA. 90210, USA

TAGF ALTER 1. H.-J. Weidemann. 1986. Neumann-Neudamm Verlag,
Melsungen (Germany). 288 pp., coll, ill.; ISBN 3-7888-0500-5. DM 38, —
hardback. (In German).

This small, richly illustrated pocket book is the first of two volumes intended
to present the natural history of German butterflies (the Alps excluded). The
first volume deals with families Papilionidae, Pieridae and the first part of
Lycaenidae. It consist of a general part dealing with butterfly ecology and a
systematic part featuring individual German and contrary to the authors
introduction, many non-German species. Of particular value are colour photo¬
graphs of early stages (mostly taken in captivity, some are marked in this
respect) and the fact that the author lists only those foodplants which were
accepted by larvae kept in captivity. Most interesting is the authors original
attempt to relate German butterfly species to certain phytoecological units. The
descriptions of most species and their early stages are very poor; time and time
again the author tells us just that the caterpillar looks like a typical Lycaenide
larva. This is as useful as D’Abrera’s infamous statement “cf as illustrated”.
Much information has been taken from other publications, without any reference
to the source of information. Thus Weidemann deals with the K- and r-strategy
in butterflies using a formula from a German ecology textbook and the
ecological properties of species exhibitions either K- or r-strategy are taken
from an English textbook. The species are classified by the formula without ever
obtaining the data (i.e. populations sizes) of the species concerned. Further, this
distinction is in connect as all butterflies are r selected (see Pianka, E.R. 1970.
Amer. Nat. 104:592-597). The attempt to present information on population
ecology of German butterflies is a change for the good, as such information, in
contrast to British Butterflies, is almost non-existant. For a country where
widespread extirpations have already occurred the situation is deeply troubling.
The book pays more attention to butterfly life histories than any standard
German publication. It does not reach the standard of the much larger Swiss
publication “Tagfalter und ihre Lebensraume” reviewed in this journal, nor of a
similar English pocketbook “RSNC Guide to the butterflies of the British Isles”
by J. A. Thomas.

Otakar Kudrna, Karl -Straub -Str. 21, D-8740 Salz (Bad Neustadt), Germany.

The Mating System of Vanessa kershawi : Males Defend
Landmark Territories as Mate Encounter Sites

John Alcock& Darryl Gwynne 116

Apodemia palmerii (Lycaenidae: Riodininae):

Misapplication of Names, Two New Subspecies
and a New Allied Species

George T. Austin 125

Correlations of Ultrastructure and Pigmentation Suggest
How Genes Control Development of Wing Scales
of Heliconius Butterflies

Lawrence E. Gilbert, Hugh S. Forrest,

Thomas D . Schultz & Donald J . Harvey 141

A mutant affecting wing pattern in Parnassius apollo
(Linne) (Lepidoptera Papilionidae)

Henri Descimon& Jean-Pierre Vesco 161

Mimicry by illusion in a sexually dimorphic,
day-flying moth, Dysschema jansonis
(Lepidoptera: Arctiidae: Pericopinae).

Annette Aiello & Keith S. Brown, Jr. 173

“Black-Light” Induction of Photoperiod-controlled
Diapause Responses of the Viceroy Butterfly,

Limenitis archippus (Nymphalidae)

A.P. Platt & S.J. Harrison 177

Suppression of the Black Phenotype in Females of
th eP.glaucus Group (Papilionidae)

David A. West & Sir Cyril A. Clarke 187

New Species and new Nomenclature in the American
Acronictinae (Lepidoptera: Noctuidae)

Douglas C. Ferguson 201

The identity of Sphinx brunnus Cramer and the
taxonomic position of Acharia Huebner
(Lepidoptera: Limacodidae)

Vitor O. Becker & Scott E. Miller 219

The Life History of Hemileuca magnified (Satumiidae)

With Notes on Hemileuca hera marcata

Stephen E. Stone, Dean E. Swift & Richard S. Peigler 225

Opinion: Are We Studying Our Endangered Butterflies
to Death?

Dennis D. Murphy 236

Opinion: Reply to Scott’s Criticism

J.P. Brock 240

Bibliography 248

Notes 254

Book Reviews 267

THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA

Volume26 Number 1-4 Autumn 1987 (1988)

IN THIS ISSUE

Date of Publication: November 15, 1988

Male Mate-Locating Behavior in the Desert Hackberry
Butterfly, A sterocampa leilia (Nymphalidae)

Ronald L. Rutowski & George W. Gilchrist 1

The Biology of SevenTroidine Swallowtail Butterflies
(Papilionidae) in Colima, Mexico

Paul Spade, Hamilton Tyler & J ohn W. Brown 13

The Mating Behavior of Papilio glaucus (Papilionidae)

Robert A. Krebs 27

A New Heritable Color Aberration in the Tiger Swallowtail
Butterfly, Papilio glaucus (Papilionidae: Lepidoptera)

J. Mark Scriber & Mark Evans 32

Bilateral gynandromorphs, sexual and/or color mosaics
in the tiger swallowtail butterfly, Papilio glaucus
(Lepidoptera: Papilionidae)

J. Mark Scriber & Mark H. Evans 39

Oviposition on Peripheral Hosts by Dispersing
Pieris napi (L.) (Pieridae)

Steven P. Courtney 58

Enzyme electrophoresis and interspecific hybridization
in Pieridae (Lepidoptera)— The case for enzyme
electrophoresis

Hansjuerg Geiger 64

Systematics of Ascia ( Ganyra) (Pieridae) Populations
in the Sonoran Desert

Richard A . B ailo witz 7 3

On Pieris (Artogeia) marginalis macdunnoughii
Remington (Pieridae)

S.R. Bowden 82

The Mating System of Three Territorial Butterflies
in Costa Rica

JohnAlcock 89

Stratification of fruit-feeding nymphalid butterflies
in a Costa Rican rainforest

P.J. DeVries 98

Euphydryas anicia and E. chalcedona in Idaho
(Lepidoptera: Nymphalidae)

Clifford D. Ferris 109