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Gregory R. Ballmer
Gordon F. Pratt
THE JOURNAL
OF RESEARCH
ON THE LEPIDOPTERA
A survey of
the last instar larvae
of the Lycaenidae
of California
VOLUME 27
NUMBER 1
SPRING 1988
Journal of Research on the Lepidoptera
27(1): 1-81, 1988
A survey of the last instar larvae of the Lycaenidae
(Lepidoptera) of California
Gregory R. Ballmer
and
Gordon F. Pratt
Department of Entomology, University of California, Riverside, Ca. 92521
Abstract. The biology and last instar larval morphology of 69 species of
California lycaenids are surveyed. Diagnostic descriptions of the
subfamilies, genera, and species are provided together with a species
key based primarily on larval morphology. Also included is a list of
confirmed larval hosts including many host species not previously
reported. Major aspects of the larval morphology of 29 exotic lycaenid
species representing six subfamilies and 17 tribes are also reviewed for
comparison.
27(1): 1-81, 1988
3
Introduction
This is the first in a series of surveys of the mature larvae of the
butterfly fauna of California. Included are 69 species of Lycaenidae
which are listed in Appendix 1. Future works will survey the larvae of
the Hesperiidae, Nymphalidae, Papilionidae, and Pieridae of California.
The Lycaenidae is perhaps the largest family of butterflies with
nearly 40% of known butterfly species (Vane- Wright, 1978). The most
recent comprehensive review of the group (based largely on adult
characters) divides it into eight subfamilies excluding the Riodinidae
(Eliot, 1973). Miller and Brown (1981) and Stehr (1987) also accord
separate family status to the Riodinidae in spite of its placement as a
subfamily of Lycaenidae by other authors (Ehrlich, 1958; Vane- Wright,
1978; Ackery, 1984; Scott, 1986). The classification of Eliot (1973) is
retained here except that the Riodininae is considered subordinate to
the Lycaenidae. Thus, the California lycaenids are divided into four
subfamilies: Lycaeninae, Polyommatinae, Riodininae, and Theclinae.
This arrangement is provisional and based largely on convenience,
since most workers are familiar with these groups. Certainly these
‘subfamilies’ are not phylogenetically equivalent; a more precise hier-
archic classification based on adult, larval, and biological characteristics
awaits a broader faunistic survey beyond the scope of this work.
Although the life histories of nearly all species treated in this key
have been published, most are inadequate for larval identification. The
majority of larval descriptions have relied heavily on coloration with
little attention to structural features. But color is perhaps the least
reliable tool for identification since larvae of many species have similar
coloration, some (especially polyphagous) species have multiple color
morphs, and ground color may change in response to different food
substrates (Orsak and Whitman, 1987). Also, larvae preserved in fluids
often lose pigmentation; only melanic pigments associated with sclero-
tized structures such as the head, legs, and setae are resistant to fading.
Overall the most reliable characters for identifying larvae are struc-
tural. The presence, absence or condition of various organs and specia-
lized setae are often diagnostic for higher taxa; the size, structure, and
distribution of specific types of setae and other cuticular structures are
often diagnostic for genera and species. Some species cannot be identi-
fied reliably by morphological traits but larval host, habitat, and
locality data may provide additional clues to their identity.
Due to the paucity of detailed descriptions of larvae of the taxa treated
in this work, a diagnostic description of each genus and a table of
comparative characters for all species are included (Table 1). Some of
the characters employed in these descriptions are new or poorly described
in the literature and much of the terminology has not been standardized;
therefore, a discussion of larval morphology and character terminology
(with a glossary) is included. This discussion is not exhaustive but deals
mostly with those characters found to be useful for identifying the
species treated here.
4
J.Res.Lepid.
Table 1 . Comparative characters of California 1
Species I.1 H.G. E.T. L.C. M.S.
lycaenid larvae
D.S. H.W. J
B.L.
N
Riodininae
A. mormo
5
-
-
.
5
-
2.28
15
10
" palmerii
5
-
-
-
5
-
1.78
12
2
C. nemesis
5
-
-
+
4
_
1.45
16
9
" wrighti
5
-
-
4-
4
-
1.60
14
6
Lycaeninae
L. arota
4
-
-
4-
2
-
1.61
17
10
" cupreus
4
-
-
4-
2
-
1.26
17
8
" editha
4
-
-
4-
3
4-
1.67
18
10
" gorgon
4
-
-
4-
2-3
_
1.74
18
10
" helloides
4
-
-
+
2
-
1.44
15
10
" hermes
4
.
.
4-
2
.
1.31
15
5
" heteronea
4
-
.
4-
2-3
4-
1.63
18
10
" mariposa
4
-
-
4-
2
-
1.53
15
3
" nivalis
4
-
-
4-
2
-
1.42
20
10
" phlaeas
4-5
-
_
4-
2
-
1.46
19
10
" rubidus
4
-
-
4-
3
4-
1.69
18
10
" xanthoides
4
-
-
4-
3-5
4-
1.87
24
10
Thedinae
A halesus
4
4-
-
-
2
.
2.34
22
10
C. (C.) comstocki
4
4-
-
-
2
4-
1.17
15
9
" " dumetorum
4
+
-
-
2
4-
1.42
19
10
" " lemberti
4
+
-
-
2
4-
1.39
17
2
" " perplexa
4
+
-
.
2
4-
1.33
15
10
* (I. ) augustus
4
+
-
-
2
4-
1.24
14
10
" " eryphon
4
+
-
-
2
4-
1.75
17
9
" "
4
+
-
-
2
4-
1.16
16
10
" " mossii
4
+
-
-
2
4-
1.40
18
10
". (M) johnsoni
5
+
-
.
2
+
1.59
17
3
" "
5-7
+
-
-
2
-
1.62
16
10
" " nelsoni
5-7
+
-
-
2
-
1.55
15
10
" " j/va
5-7
+
-
-
2
-
1.72
16
10
" " spinetomm
5
+
-
-
2
4-
1.85
21
10
" " thomei
5-7
+
-
-
2
-
1.51
15
3
H. grunus
4
-
-
4-
5
-
1.78
17
10
H. titus
4
+
.
-
6
+
2.02
24
4
M. leda
4
+
-
-
2
4-
1.03
12
10
S. auretorum
4
+
-
-
3
4-
1.56
18
10
" fre/ira
4
+
-
-
2
4-
1.19
14
10
" califomica
4
+
-
-
2
+
1.50
17
2
" fuliginosum
4
+
-
-
3
+
1.58
21
10
" saepium
4
+
-
-
2
4-
1.44
17
10
" sylvinus
4
+
-
-
3
+
1.57
17
10
" tetra
4
+
-
-
3
+
1.78
19
10
S. avalona
4
+
-
-
2
4-
1.15
13
6
" columella
4
+
-
-
2
4-
1.11
14
10
" melinus
4
+
-
-
2
4-
1.22
16
10
Polyommatinae
A.franklinii
4
-
-
-
2
-
.86
13
3
B. exilis
4
4-
+
-
2
_
.66
11
10
C. argiolus
4
4-
4-
-
2
4-
.89
12
10
27(1): 1-81, 1988 5
Table 1. con’t
Species
I.1
H.G.
E.T.
L.C.
M.S.
D.S.
H.W.
B.L.
N
E. battoides
4
+
+
_
2
+
.63
9
10
" enoptes
4
+
+
-
2
+
.67
10
10
" mojave
4
+
+
-
2
+
.68
9
10
" rita
4
+
+
-
2
+
.60
10
10
E. amyntula
4
+
+
-
2
+
1.03
14
10
" comyntas
4
+
+
-
2
+
.83
12
4
G. lygdamus
4
+
+
-
2
+
1.11
19
10
" piasus
4
+
+
-
2
+
1.17
18
10
H. ceraunus
4
+
+
-
2
+
.73
12
10
" isola
4
+
+
-
2
-
.71
11
10
I. acmon
4
+
+
-
2
+
.77
11
10
" icarioides
4
+
+
-
2
+
1.03
17
10
" lupini
4
+
+
-
2
+
1.00
15
10
" neurona
4
+
+
-
2
+
.89
12
10
" shasta
4
+
+
.
2
+
.85
12
10
L. marina
4
+
+
„
2
+
.75
11
10
L. idas
4
+
+
.
2
+
.90
14
2
” melissa
4
+
+
-
2
+
1.00
16
10
P. sonorensis
4
+
+
-
2
+
1.10
17
10
P. speciosa
4
+
-
-
2
+
.65
10
10
P. saepiolus
4
+
+
-
2
+
.95
12
6
P. emigdionis
5-7
+
+
-
2
+
1.27
16
10
1 1 = larval instars; H.G. - honey gland; E.T. = eversible tubercles; L.C. = lateroseries of crochets;
M.S. = mandibular setae; D.S. = dendritic setae; H.W. = mean head width (mm); B.L. = mean
body length (mm); N = number of specimens measured for H.W. and B.L.; + = present, - - absent.
In order to understand better the distribution of morphological
characters among higher taxa, larvae of several exotic species were also
examined. Table 2 compares the following exotic species with respect to
eight major morphological characters: Liphyrinae: Liphyra brassolis
Westwood from Australia; Lycaeninae: Heliophorus epicles (Godart)
from Malaysia; Miletinae: Feniseca tarquinius (Fabricius) from Illinois,
Spalgis epeus (Westwood) from Thailand; Polyommatinae: Anthene
seltuttus affinis (Waterhouse and Turner) and Candalides xanthospilos
(Hubner) from Australia, Eueres argiades (Palla) from J apan, Erysichton
lineata (Murray), Danis hymetus (C. and R. Felder), Lampides boeticus
(Linnaeus), Syntarucus plinius (Fabricius), a Xylomelum- feeding
member of the Theclinesthes miskini (T.P. Lucas) — onycha (Hewitson)
complex, and Zizina labradus (Godart) from Australia, and Zizula hylax
(Fabricius) from Thailand; Riodininae: Melanis pixe (Boisduval) from
Texas and Zemeros flegyas (Cramer) from Thailand; Theclinae: Arhopala
centaurus (Fabricius) and Cheritra freja (Fabricius) from Thailand,
Deudorix epijarbas (Moore) from Australia^ rora quaderna (Hewitson)
from Arizona, Eumaeus atala (Poey) from Florida, Flos areste (Hewitson)
6
J.Res.Lepid.
Table 2. Comparative larval features for some exotic lycaenidae
Species1
H.G. E.T.
L.
S.L.
L.C.
M.S.
S.S.
Chalazae
Riodininae
Hamearini
Z.flegyas
Riodinini
-
-
+
+
+
2
+
simple
M.pixe
Liphyrinae
-
-
+
+
+
24
+
simple
L. bras solis
-
-
-
+
+
2
-
stellate
Lycaeninae
H. epicles
Miletinae
-
-
+
+
+
4
+
simple
F. tarquinius
-
-
+
+
.
2
-
simple
S. epeus
Theclinae
Arhopalini
+
+
+
1
simple
A. centaurus
+
+
+
+
+
5
+
buttressed
F. areste
+
+
+
+
+
6
+
buttressed
S. quercetorum
Cheritrini
+
+
+
+
+
4
+
buttressed
C.freja
Deudorigini
-
-
+
+
+
2
?
buttressed
D. epijarbas
Eumaeini
+
-
+
+
3
+
buttressed
E. quadema
-
-
+
+
-
2
+
stellate
E. atala
-
-
+
+
.
2
+
buttressed
Hypolycaenini
H. erylus
Luciini
+
-
+
+
+
2
+
buttressed
Hypochrysops sp.
+
+
+
+
+
4
+
simple
P. kamerungae
Ogyrini
_
■
+
+
+
6
+
buttressed
O.genoveva
Theclini
+
+
+
+
+
5
+
buttressed
H. crysalus
Zesiini
-
-
+
+
+
6
+
buttressed
P. chlorinda
+
+
+
+
+
5
+
buttressed
Polyommatinae
Candalidini
C. xanthospilos
Lycaenesthini
+
+
+
+
■
2
+
buttressed
A.seltuttus
Polyommatini
+
+
+
+
■
2
+
simple
E. argiades
+
+
+
+
-
2
+
stellate
E. lineata
+
+
+
+
-
2
+
stellate
D. hymetus
+
-
+
+
-
2
+
stellate
L. boeticus
+
+
+
+
-
2
+
stellate
S.plinius
+
+
+
+
_
2
+
stellate
T. miskini-onycha
+
+
+
+
-
2
+
stellate
Z. labradus
+
+
+
+
.
2
+
stellate
Z. hylax
+
+
+
+
-
2
+
simple
1 Complete citations in text; H.G. =
honey gland; E.T. =
eversible tubercles; L.
= lenticles; S.L.
spatulate lobe on prolegs; L.C.
= lateroseries of crochets;
M.S.
— mandibular setae; S.S. = sens
setae; 4- = present,
- = absent
9 s
? •
presence
or absence not determined.
27(1): 1-81, 1988
7
from Malaysia, Hypaurotis crysalus (Edwards) from Arizona, Hypo-
chrysops apelles (Fabricius) from Australia, Hypolycaena erylus (Godart)
from Thailand, Ogyris genoveva Hewitson, Philiris kamerungae Water-
house, and Pseudalmenus chlorinda (Blanchard) from Australia, and
Surendra quercetorum (Moore) from Thailand.
The keys, diagnostic tables, and descriptions are based on observa-
tions by the authors of live and/or preserved larvae and photographs.
Descriptions of genera and higher taxa are based on the California
fauna except where noted. Distributional data has been compiled from
several sources including publications and private collectors. Larval
color descriptions are based on living material and/or color photographs.
Host plant information presented in the diagnostic section is based on
published records and field observations by the authors; only relatively
recent literature citations are given. Appendix 2 lists only those hosts
confirmed by the authors’ field collections; some reconfirm old records
but many are new. The authors of botanical names are abbreviated
according to the format of Munz and Keck (1959).
The majority of larvae examined were collected in the field by the
authors or reared from ova from field collected adults. Larval morpho-
logy was determined primarily using a binocular dissecting microscope.
The fine structures of some organs were further examined using a
compound microscope with fiber-optic illumination and/or a scanning
electron microscope (SEM). The orientation of line drawings and SEM
photographs used for illustration are standardized (unless otherwise
indicated) with cephalad to the left (lateral views) or top (dorsal views).
The key and descriptions apply to last instar larvae. Although the
number of larval instars in some Lepidoptera may depend upon environ-
mental variables and diet, most California lycaenids appear to have a
constant number of instars. Most species have four instars but the
riodinines, one polyommatine, and six theclines typically have five or
more instars. In the Lycaenidae the last instar often differs morphologi-
cally from previous instars. Since the instar in which various organs
and specialized setae first appear varies for different taxa, the best
general means of determining the instar is larval size. Although body
length increases within an instar, sclerotized body regions such as the
head remain constant in size between molts.
The head width for last instar larvae of each species is included in
Table 1 along with the body length (from anterior margin of the
prothorax, excluding the neck, to the caudal margin of abdominal
segment 10) to aid in determining larval instar and to indicate compara-
tive size among species. All measurements are in millimeters and are
based on preserved larvae. For most species these values are derived
from a single collection or pooled collections of ten individuals repre-
senting a single population. Different preservation methods, as well as
larval condition at the time of preservation, can result in different body
length and width dimensions. The problem of morphological variability
8
J.Res.Lepid.
among subspecies is discussed for a few species where it may aid in
identification. A broader discussion of all subspecies is beyond the scope
of this work.
The most effective use of the key requires a binocular dissecting
microscope since major emphasis is placed on small morphological
features. Whenever possible one should attempt to identify larvae while
they are alive because some structures are best seen while the larval
surface is dry and because coloration may be useful.
There are several effective techniques for larval preservation; the
most important considerations are to distend the larva and fix the
tissues. Commonly larvae are killed in KAAD (kerosene, ethanol, acetic
acid, and dioxane) or other fluids which both distend and fix them.
Another satisfactory method is to inject the larva through the anal
opening with a fixative such as Kahle’s fluid until it is sufficiently
distended and/or fix it in hot water (70-85 degrees C.) for about five
minutes. Larvae are usually stored in 70-80% ethanol; they should not
be put into alcohol until they have been properly fixed. Inflated, freeze-
dried, and critical point-dried larvae are also satisfactory for identifica-
tion but may be difficult to store and handle. Dry or shriveled larvae,
larval exuviae, and carcasses remaining after parasite emergence may
be identifiable after softening in 5% potassium hydroxide solution. For a
broader discussion of methods for larval preservation see Peterson
(1948) or Stehr (1987).
27(1): 1-81, 1988
9
LARVAL MORPHOLOGY
The immature stages of many species of Lycaenidae are commonly
associated with ants. This association is reflected in the specialized
morphology of the larvae among which a variety of myrmecophilous
adaptations have evolved (see Hinton, 1951; Henning, 1983b; Cottrell,
1984; Kitching and Luke, 1985).
The known mature larvae of most lycaenids (including nearly all
California species) share a few morphological features which (in com-
bination) distinguish them from all other Lepidoptera. The presence of
cuticular lenticles and a fleshy terminal lobe on the prolegs are almost
unique to this family where they occur in most known species. Other
distinctive features such as an onisciform body shape, retractable head,
eversible tubercles, and honey gland are somewhat less widely distri-
buted in the Lycaenidae. The full extent of the distribution of these and
other diagnostic features remains speculative since the larvae of most
species remain undescribed. Generalizations are further hindered by
the morphological diversity encompassed by those species which have
been investigated.
A clearer understanding of the relationships among higher taxa in
the Lycaenidae can probably by gained by broad faunal surveys of their
immature stages as suggested by Henning (1983a), Cottrell (1984), and
De Vries et al (1986), yet few such works exist. Malicky (1969a, 1969b,
1970) surveyed the larval morphology (especially ant-associated organs)
of Central European lycaenids and produced a key based on their
morphology, coloration, and host plants. Scott (1986) produced a last
instar larval key to the families, subfamilies, and some tribes of North
American butterflies; his key is more detailed than those of Peterson
(1948) and Stehr (1987), which pertain to all North American Lepi-
doptera, but fails to consider the full range of diversity in North
American lycaenids.
The shape of lycaenid larvae is often termed onisciform (shaped like a
sowbug or woodlouse, Oniscus ). The prothorax (Tl) is often the longest
segment since in most species the head is retractable into it. The body is
usually broadest and highest at the mesothorax (T2), metathorax (T3),
or first abdominal segment (Al) and gradually tapered posteriorly.
Abdominal segments ten (A 10), nine (A9), and (usually to a much lesser
extent) eight (A8) are fused to varying degrees in different groups. In
cross-section the body is typically convex dorsally and flattened ven-
traliy. The dorsum may be evenly rounded but in some species paired
dorsal prominences (flanking the middorsal line) create a trapezoidal
outline in cross-section and a saw-toothed lateral profile. Middorsal
prominences occur in some exotic species such as Cheritra freja of
southeast Asia. The junction of the lateral and ventral body regions
typically forms a fleshy lateral fold (often fringed with long setae) which
conceals the legs when at rest. When inflated (as commonly occurs
during preservation) the body may assume a more cylindric shape
10
J.Res.Lepid.
typical of other Lepidoptera. An onisciform body is typical of the
subfamilies Curetinae, Liphyrinae, Lycaeninae, Polyommatinae and
Theclinae; but the larvae of some Miletinae and many Riodininae are
much less onisciform while those of the exotic Lipteninae and Poritiinae
are not at all (Cottrell, 1984).
The lycaenid larval head is commonly much narrower than the
thoracic segments and attached to a neck-like extension of the prothorax
into which it can be withdrawn. The size of the head relative to body
width, the neck length, and degree to which the head can be withdrawn
are variable among (and to a lesser degree within) the lycaenid
subfamilies. Among the California fauna the head is smallest and neck
generally longest in the Polyommatinae while the head is largest and
neck shortest in the Riodininae. A broad nonretractable head is typical
of the known larvae of Riodininae and of some other subfamilies such as
the Miletinae (including Feniseca tarquinius of eastern North America)
and of the Florida thecline Eumaeus atala.
Other morophological features common to mature lycaenid larvae
include the presence of prolegs on A3-A6 and A10 (anal prolegs) and
numerous secondary setae. Also, with few exceptions the crochets on the
prolegs are arranged in a mesoseries divided by a fleshy lobe (fig. 72);
some species also have a lateroseries of crochets (fig. 72a). Most known
lycaenids, including all California species, possess a well defined
prothoracic shield (fig. 1). A well developed (sclerotized) suranal shield
is present in many exotic species but not in California lycaenids; a
poorly developed suranal shield is present in the riodinines and in H.
grunus.
Many lycaenids in the Curetinae, Liphyrinae, Polyommatinae, and
Theclinae possess a pair of eversible tubercles on the eighth abdominal
segment (fig. 1, 20) and/or a honey gland (Newcomer’s organ) usually
located on the seventh abdominal segment (figs. 1, 19). In the Southeast
Asian thecline, H. erylus , the honey gland is located on the eighth
abdominal segment. Analogous structures are present in some exotic
riodinines such as Anatole rossi Clench which has paired eversible
tubercles on the metathorax and paired eversible honey glands on the
eighth abdominal segment (Ross, 1964). Honey glands produce a fluid
which ants imbibe while the eversible tubercles may either attract or
excite ants by releasing semiochemicals which mimic the ants’ own
pheromone(s) (Henning, 1983a and b). Cuticular lenticles, which
probably occur in most lycaenids (see lenticle discussion below),
apparently also have a chemical communication function (Malicky,
1970; Henning, 1983a and b).
A few characters permit an easy distinction between the larvae of
riodinines and other lycaenids in California. Among the riodinine
larvae the longest setae are clustered in tufts or verrucae (figs. 30, 31,
69, 70) and are frequently longer than the head width; the prothoracic
shield is transverse (the length along the dorsal midline is about half as
27(1): 1-81, 1988
11
great as the width) and adorned with several conspicuously long setae
which extend anteriorly over the head (fig. 5). Perhaps the best
distinguishing feature of the riodinine larvae is the anteroventral
displacement of the A1 spiracles to a location just anterior and slightly
ventral to the lateral verrucae (figs. 30, 69, 70) where they may be
concealed by a cuticular fold; spiracles on the other abdominal segments
are located about midway between the dorsal and lateral verrucae. This
condition probably applies to most New World riodinines (Don Harvey,
1987 and in litt.), but not to the Old World taxa. Among the larvae of
other California lycaenids all setae are shorter than the head width and
randomly scattered over the body; the prothoracic shield is often
approximately diamond-shaped (figs. 33-54, 61, 64, 67, 71) and is
usually as long or longer than wide; no setae on the shield extend
forward as far as the anterior margin of the prothorax (fig. 1).
Some exotic riodinines possess additional features which apparently
do not occur in other Lepidoptera families. For example, the neotropical
species A. rossi (Clench), Audre epulus signatus (Stichel), and A.
susanae (Orfila) have a pair of vibratory papillae on the prothorax and
paired honey glands on the eighth abdominal segment (Ross, 1964;
Bruch, 1926; and Bourquin, 1953, respectively). Larvae of A. rossi also
possess a pair of eversible tubercles on the metathorax (Ross, 1964). The
aforementioned species are notably myrmecophilic, unlike the California
riodinines.
Chaetotaxy is the primary tool for identifying most Lepidoptera
larvae yet detailed systematic descriptions of lycaenid larval chaetotaxy
are uncommon. Clark and Dickson (1956b) proposed the use of first
instar setal patterns as a tool for elucidating phyletic relationships in
the Lycaenidae and later (1971) described the early stages of the South
African fauna. Other workers (Lawrence and Downey, 1966; Downey
and Allyn, 1979 and 1984; and Wright, 1983) provided detailed descrip-
tions of four North American species. In each of the latter works a
modified version of Hinton’s (1946) setal nomenclature was employed to
describe (primarily) the first instar chaetotaxy. These authors also
recognized a variety of structural forms of setae, some of which were
found only in later instars.
Hinton (1946) recognized two functional types of setae in the order
Lepidoptera: microscopic or proprioreceptor setae located along inter-
segmental folds and where different body parts make contact, and long
or tactile setae which are more widely distributed and may be modified
for specialized functions. The microscopic setae have received little
attention by investigators due to their small size while the structure
and distribution of tactile setae are widely employed in identifying
larvae of many families of Lepidoptera.
Tactile setae may be further categorized as primary, subprimary, and
secondary. Primary setae, found in specific body locations, are believed
to represent the archetypal lepidopteran setal pattern and, with few
12
J.Res.Lepid.
exceptions, are discernible at least in the first instar. Subprimary setae,
always few in number, also occur in fixed locations which are charac-
teristic of some families. In most groups where they occur subprimary
setae appear in later instars but in highly specialized families such as
the Lycaenidae they occur in the first instar. Secondary setae, which are
numerous in some families, are variable in number and position,
generally most abundant in the last instar, and only rarely occur in first
instars.
For most (perhaps all) lycaenids the distribution pattern of primary
and subprimary setae is obscured by numerous secondary setae in
instars following the first. Among the fauna included in this survey
there are several structurally distinct types of secondary setae whose
size, number, and distribution are often taxonomically useful.
The basic setal structure consists of a hollow shaft (usually with
lateral processes or spiculations) arising from a basal ring which
surmounts a short sclerotized prominence or chalaza. In the Lycaenidae
variations in setal structure range from long, slender, and finely
tapered to short, stout, and capitate while their lateral processes range
from long slender filaments to short, stout, pointed dentations and
minute granulations. In some taxa setal structure is nearly uniform
regardless of setal size or location but more often the longest setae are
most erect and most tapered while the shortest setae are most curved
and/or clavate. The longest setae also tend to be in locations where true
primary setae are expected to occur (i.e. dorsal, subdorsal, lateral, etc.).
Often there is a gradient in setal structure with the most clavate,
capitate, or recurved setae occurring dor sally on A7-A10. Setae with the
longest lateral processes (relative to setal length) occur most frequently
in close proximity to the honey gland and spiracles.
In the known larvae of New World riodinines most setae are gradually
tapered, filamentous, or short and multibranched while strongly bent
setae are absent or rare. Also, the longest setae (often much longer than
the head width) arise from the prothoracic shield and/or dorsal and
Interval verrucae. But in the Old World Z. flegyas setal structure is often
more complex and those setae on the prothoracic shield barely reach the
posterior cranial margin. In Calephelis the longest setae may be longer
than the body width while the shortest setae, which occur densely over
the dorsal and lateral regions, are too small to be individually discerned
without magnification. In Apodemia all body setae are gradually
tapered or filamentous and those arising from verrucae are primarily
stiff and spinelike.
For the purpose of describing the larvae treated in this survey seven
common categories of setae are defined: 1) prominent (figs. 2a, 4), 2)
dendritic (figs. 2p, 15-18), 3) sensory (figs. 6, 9-14), 4) neck (fig. 27), 5) j
mushroom (figs. 2q, 22), 6) plumose (figs. 31, 32), and 7) echinoid (fig.
31). Prominent and sensory setae occur in all lycaenid subfamilies
represented in California but dendritic and neck satae are absent in the
27(1): 1-81, 1988
13
Riodininae, mushroom setae occur only in the Lycaeninae, and plumose
and echinoid setae occur only in the Riodininae. The first four setal
categories are widespread in the Lycaenidae; the last three may occur
only in relatively small groups (subfamily, tribe, etc.) and it is likely
that other structurally equivalent setal categories could be defined for
other small phyletic groups. However, the majority of larval setae do not
fit the categories enumerated above; they comprise an unnamed assem-
blage of perhaps less specialized setae with diverse structures (figs. 2b-
2o).
Prominent setae of Lycaenidae occur in specific locations which
coincide with sites where true primary setae are found in most other
Lepidoptera. However, the number of prominent setae in a given
location is variable and often exceeds the basic number of primary setae;
they may also be absent. They are always cylindric, tapered, erect, and
straight to gently curved (figs. 2a, 4); in many species they differ from
surrounding setae only in their much greater length. This setal cate-
gory corresponds to the major setae of Lawrence and Downey (1966),
Downey and Allyn (1979), and Wright (1983). Prominent setae on the
prothorax usually occur abundantly along the anterior and lateral
margins where they can serve as the anterior most tactile receptors
when the head is withdrawn. Additional prominent setae may occur on
the prothoracic shield or near its anterior and posterolateral margins.
On all other segments prominent setae occur singly or in groups
dorsally (flanking the middorsal line), subdorsally (about half way
between the spiracles and middorsal line), and laterally (below the
spiracles, along the lateral fold) (fig. 1). Aside from the prothorax,
prominent setae tend to be most abundant on the mesothorax and
progressively less abundant posteriorly, although there is usually little
difference in their numbers on A1-A6. In species having a honey gland
there are no dorsal prominent setae on A7. Subdorsal prominent setae
are least encountered but usually occur on T2 and are more likely to
occur on A6 and A7 than on A1-A5. Lateral prominent setae usually
occur on all segments but are reduced or lacking in some species.
Prominent setae are almost always present along the anterior and
lateral margins of the prothorax and posterolaterally on A 10. In the
California riodinines most prominent setae occur on verrucae and the
prothoracic shield. In Apodemia they are primarily stiff and spinelike
but in Calephelis they are mostly long, slender, and plumose. These
latter setae are morphologically distinctive enough to be accorded a
separate setal category (plumose setae) discussed below.
Dendritic setae can be distinguished from other secondary setae by
structure and location. They may appear tree-like due to the presence of
filamentous lateral processes arising from the apical half which are
usually longer than the setal width at their point of origin. These setae
are weakly tapered to clavate, erect, straight (rarely slightly curved),
and less pigmented than other setae. They are usually restricted to a few
14
J. Res. Lepid.
locations (where lenticles may also be concentrated) such as the margin
of the honey gland and spiracles, but are more widespread in some taxa
and absent in others. Commonly at least a pair of dendritic setae occurs
at each lateral angle of the honey gland where they may be obscured if
the gland opening is retracted. The dendritic appearance of these setae
varies due to the relative length of their lateral processes. Although the
processes are usually much longer than the setal width (figs. 15-17),
they can be shorter and may not be easily seen (especially those at the
lateral angles of the honey gland) (fig. 18). In such cases dendritic setae
can usually be distinguished by their greater prominence (surrounding
nondendritic setae are often shorter and clavate-capitate or recurved).
Lawrence and Downey (1966) used the terms spiculate and dendritic for
this type of setae on the larva of E. corny ntas and likened their
appearance to Christmas trees. Other terms for dendritic setae which
appear in the literature include spiculate and hydroid setae (Downey
and Allyn, 1979) and branched hairs (Kitching, 1983).
The function of dendritic setae is not well established but there are
indications that they may be involved in chemical and/or tactile
communication with ants. Our observations indicate a direct relation-
ship between the abundance of these setae and the degree of ant-larval
association (research in progress). Thus, while larvae of all twelve
California species of Lycaeninae lack the more notable ant association
organs (honey gland and eversible tubercles), the four which hav^e
dendritic setae are the only ones which we have found associated with
ants.
Although all setae may have a sensory function, the term sensory
setae has been applied to a unique pair of setae on the lycaenid
prothoracic shield (Downey and Allyn, 1979 and 1984). They have also
been termed XD2 (op. cit. and Wright, 1983) and major setae ‘type a’
(Lawrence and Downey, 1966). But setae of homologous structure found
in other Papilionoidea and at least some Arctiidae, Geometridae,
Saturniidae, and Sphingidae occur anterior and/or anterodorsal to the
T1 spiracles and often subdorsally on other segments; their location
coincides with Hinton’s SD1 setae. These are the only primary body setae
which can be recognized in all lycaenid instars. They occur in all
California lycaenids and in nearly all exotic species examined; none
were found in the liphyrine L. brassolis nor in the miletines F.
tarquinius and S. epeus. The function of the sensory setae is unknown
and they are curiously insensitive to tactile stimulation.
The structure of the sensory setae varies among different taxa. They
may be filiform, flagelliform (fig. 14), clavate (fig. 12), spatulate (fig. 13),
or even branched (fig. 6). In most species the sensory setae have
inconspicuous lateral spicules but in others the spicules are longer,
causing the sensory setae to appear brush-like (fig. 9) . In the Lycaeninae
the lateral spicules are confined to the apex (best seen with SEM) or
absent (fig. 14). Sensory setae are more slender, at least basally, often !
27(1): 1-81, 1988
15
longer, and more flexible than other setae on the prothoracic shield;
they arise from uniquely low, button-like chalazae. In live larvae their
flexibility and fine basal attachment result in a vibratory or slow
twitching motion in weak air currents which may cause them to appear
independently motile.
Mushroom setae are known to occur only in members of the Lycaeninae
(Wright, 1983). Malicky (1969) referred to this type of seta as Baum-
chenhaare (tree setae) in his treatment of the European Lycaena
species. Under low magnification they appear as short, rounded struc-
tures resembling mushrooms but with higher magnification they can be
seen to have numerous short, stout distal and lateral processes (figs. 2q,
22). They are usually nonpigmented and much shorter than other setae
among which they are scattered over the dorsal and lateral body
surfaces. To the unaided eye they may appear as minute white speckles
in contrast to the darker larval ground color. Structural variations in
the mushroom setae, especially their lateral processes, may be useful
taxonomic characters (figs. 2q and 22) but since these cannot be seen
without SEM they are of little use in larval identification.
The lycaenid larval neck can be densely covered with minute setae
and/or spinules which give it a granular appearance. These neck setae
typically are much shorter than other secondary setae, stout (often
tooth-like), may be rounded or acute apically, and may have a few short,
stout apical or subapical spicules (fig. 27). Spinules are shorter than
neck setae and range from pointed to rounded and may be erect or
recumbent (figs. 27, 28). Both neck setae and spinules are apparent on
the posterior half of the neck where their distribution usually ends
abruptly at the junction with the nonretr actable remainder of the
prothorax. In some lycaenines and theclines neck setae are also sparsely
scattered dor sally and laterally over the remainder of the prothorax.
Under the dissecting microscope it may be difficult to discern whether
they are setae (with supporting chalazae) or merely spinules. In the
Theclinae and Lycaeninae both neck setae and spinules are abundant
on the posterior half of the neck. In the Polyommatinae there is a narrow
dorsal band of spinules at the posterior end of the neck and a more
extensive ventral patch of spinules and neck setae anterior to the
prothoracic legs. Neck setae and spinules were not observed in the
Liphyrinae, Miletinae, and Riodininae. Although differences in the
structures of the neck setae and spinules may provide characters for
taxonomic studies they are difficult to observe without SEM and are not
discussed further here.
Plumose setae occur only in the riodinines and are most apparent on
the prothoracic shield and verrucae. They are slender filaments densely
clothed with short, fine lateral processes which may confer a velvety
appearance (figs. 31, 32). These setae are mostly uniform in width
throughout most of their length but may be apically spatulate (fig. 32) or
abruptly tapered. In A. mormo a single plumose seta may arise from
16
J. Res. Lepid.
each dorsal verruca and 4 or more from each lateral verruca along with
numerous shorter spine-like setae (fig. 69), but in Calephelis the
verrucae are comprised entirely of plumose setae (fig. 70). In the latter
genus these setae vary greatly in length but many are longer than the
body width and confer to a larva the appearance of a down feather; much
shorter plumose setae occur primarily near the verrucae, on the
prothoracic shield, and near the intersegmental lines. Also in Calephelis
a few plumose setae on dorsal verrucae are relatively short, stout, and
somewhat spatulate.
Echinoid setae were found only in Calephelis larvae. Due to their
small size and density they may cause the body surface to appear
pollinose or mealy. They are rather short and stout and adorned with
relatively large lateral processes which taper from broad bases but are
apically flared (fig. 31). Other authors have referred to these setae as
many-pointed branching stars in C. wrighti (Comstock, 1928), stellate
nodules in C. nemesis (Comstock and Dammers, 1932), silvery stars in
C. borealis (Grote and Robinson) (Dos Passos, 1936), and sprocket-
shaped processes in C. muticum McAlpine (McAlpine, 1938).
Other secondary setae span a wide range of structural forms including
erect, recumbent, straight, recurved, tapered, and clavate-capitate (figs.
2b-2o). In some species these setae are uniform in structure but
commonly there is a structural gradient with the most extreme forms
occurring in specific areas. The most recurved and clavate or capitate
setae usually occur dorsally on abdominal segments 7-10 (especially
near the honey gland). The nearly continuous range of structural
variation in these setae among different species and even on individual
larvae diminishes the value of defining specific structural types for
them.
All setae on the larval body normally arise from sclerotized tubercles
or chalazae. Ventral chalazae are typically cylindric but dorsal and
lateral chalazae may be sculptured in the Polyommatinae and Theclinae.
The chalazae of most species of Polyommatinae appear stellate or
crown-like due to conspicuous lateral or distolateral points (figs. 2e, 2f,
2j, 2k, 13, 16). In the Theclinae the chalazae are most often conical with
longitudinal ridges resembling buttresses which are most prominent
basally where they fuse with the cuticle (figs. 2n, 2o, 4); similar chalazae
occur in the Curetinae (DeVries, et al, 1986). In the liphyrine L.
brassolis the dorsum is covered with highly modified flattened, shingled
chalazae bearing much smaller setae; toward the ventral margin of the
dorsal carapace these chalazae become more erect and cylindric and
appear somewhat stellate. In all Lycaeninae, Miletinae, and Riodininae
examined the chalazae are smoothly contoured and cylindric or globular
(figs. 2i, 15). Although the structural distinctions between the chalazae
of the Polyommatinae and Theclinae are true for most species examined,
there are some exceptions. In the polyommatines P. emigdionis from
California and C. xanthospilos from Australia, the chalazae appear
27(1): 1-81, 1988
17
buttressed rather than stellate. In the former species SEM photographs
show that the lateral ridges are not distobasally fused with the cuticular
surface (fig. 10) but in the latter they are very similar to typical thecline
chalazae. Larvae of the Arizona thecline, Erora quaderna , have stellate
chalazae much like those of the Polyommatinae. Also, some members of
both subfamilies have apparently nonsculptured chalazae as in the
Lycaeninae.
In many lycaenids the degree of sculpturing of the chalazae varies for
different types of setae and in different body regions. The most sculp-
tured and strongly pigmented chalazae often occur on or near the
prothoracic shield. Chalazae associated with dendritic setae are usually
less sculptured than those of other setae and may appear nonsculptured.
In the Polyommatinae the chalazae of prominent setae tend to be larger
but less stellate than those of shorter adjacent setae. The chalazae
associated with neck setae and the sensory setae on the prothoracic
shield are not sculptured. The latter are visibly low and button-like
(figs. 6, 9, 11, 12, 14) in the Lycaeninae, Theclinae, and Riodininae
whereas in the Polyommatinae they are often slightly sunken below the
cuticular surface (figs. 10, 13). Their visible dorsal surface is glassy
smooth and flat or convex, much like a lenticle with a small central pore
from which the sensory seta emerges.
Lenticles are small lens-like cuticular structures often resembling
chalazae without setae; they may be present in all lycaenids and thus
constitute an important diagnostic character. Other terms for lenticles
include perforated cupolas (Malicky, 1970), Allyn’s organs (Downey and
Allyn, 1979), and pore cupolas (Kitching and Luke, 1985). Malicky
(1970) found them in 60 species of (primarily European) lycaenids. In
this study they were found in all lycaenid species examined except L.
brassolis. The larval dorsum in that aberrant species is densely covered
with overlapping chalazae; but there are regularly placed pores which
may lead to recessed glands and/or lenticles, as suggested by observa-
tions of the inner surface of larval exuviae (Bethune-Baker, 1925).
Lenticles have a low, round, convex or flat central region surrounded
by a narrow collar; the latter may be smoothly rounded and cylindric,
buttressed, or stellate much like the chalazae of nearby setae (figs. 15,
16, 17, 19). Although the lenticles of first instar larvae are few in
number and occur in fixed locations, those of later instars are more
numerous, variable in number, and more randomly distributed. In
mature larvae lenticles tend to be sparsely scattered over dorsal and
lateral body regions but more numerous near the honey gland and
spiracles (especially on A8).
The function(s) of the lenticles remains somewhat speculative but at
least some seem to be related to myrmecophily. Malicky (1970) suggested
that lenticles have a chemical communication function since they are
most abundant in areas commonly attended by ants and at least some
have a porous surface and are associated with epidermal glands.
18
J.Res.Lepid.
Lenticles of similar structure occur in hesperiid larvae (which are not
myrmecophilous) where they have been shown to be formed by cells
which can also produce setae (Franzl et al, 1984). Henning (1983b)
found in some South African lycaenids that the body surface containing
lenticles also contains a chemical which mimics an associated ant’s
brood pheromone.
A specialized type of lenticle, found in A. halesus, has a mushroom
shape. Mushroom lenticles differ from other lenticles primarily in
being stalked or elevated above the body surface and narrowest at the
base; also, the collar surrounding the central lens is relatively broad and
divided into radial segments by narrow ridges (figs. 6, 26). Under low
magnification these lenticles appear similar in size and form to the
mushroom setae of Lycaena larvae and, likewise, confer a minutely
white speckled appearance to the larva. They are randomly distributed
over the dorsal and lateral body regions but replaced by more ordinary
appearing lenticles on the prothoracic shield and areas adjacent to the
spiracles and honey gland.
The presence of a pair of eversible tubercles (tentacular organs)
dorsolaterally on abdominal segment eight is a common trait in the
Lycaenidae; they have been reported in the subfamilies Curetinae,
Liphyrinae, Polyommatinae, and Theclinae (Cottrell, 1984). The
appearance of these organs differs little among the California species.
Ordinarily they are retracted into the body but when everted they can
be seen (in most species) to have an apical cluster of relatively long,
straight, prominently spiculate setae (fig. 20). The spicules on these
setae are slender and rather evenly distributed whereas those of
dendritic setae, which they otherwise resemble, are concentrated in the
apical half of the seta. In some exotic taxa variations in the structure of
the tubercles and in the number, size, and color of their terminal setae
are of taxonomic value (Clark and Dickson, 1956a); the eversible
tubercles of some exotic species such as Ogyris genoveua and Candalides
xanthospilos lack setae.
Among the California fauna eversible tubercles are absent in the
Lycaeninae, Riodininae, and Theclinae but well developed in all Poly-
ommatinae except A. franklinii and P. speciosa ; they may be nonfunc-
tional in some populations of E. amyntula. Generally these organs are
everted briefly at irregular intervals or in response to tactile stimuli but
in some, such asP. emigdionis , they pulsate regularly and are frequently
everted as the larva crawls. The function of the tubercles may vary
among different taxa; Clark and Dickson (1956a) felt that ‘whip’ type
eversible tubercles in Aphnaeini mechanically remove bothersome
ants; some other authors (Henning, 1983a and b; De Vries, 1984;
Kitching and Luke, 1985) suggest that the tubercles in other taxa
release a chemical which mimics an ant alarm pheromone. In preserved
larvae they are seldom everted, but their locations usually can be
discerned by the presence of a small circle of setae surrounding a bare
27(1): 1-81, 1988
19
wrinkled depression (which may surmount a low prominence) slightly
posterolateral to the eighth abdominal spiracles.
The honey gland (Newcomer’s organ or dorsal nectary organ) is a
feature present in many species of Polyommatinae and Theclinae but
apparently absent from other lycaenid subfamilies. It usually appears
as a narrow transverse middorsal furrow on A7; in the exotic H. erylus
the honey gland occurs on the eighth abdominal segment, closely
flanked by the spiracles. In response to stimulation by ants the gland
can partly evert (fig. 19) and discharge a drop of fluid which ants imbibe.
For some species this fluid has been shown to contain both sugars and
amino acids (Maschwitz et al, 1975; Pierce, 1983). In discussing the
relationship between ants and lycaenid larvae, Malicky (1969, 1970)
suggested that the honey gland provides a bribe to forestall aggression
by ants. Pierce and Mead (1981) noted that ant-tended larvae of G.
lygdamus were significantly less parasitized than untended larvae and
suggested that the honey gland attracts ants which defend the larva
much as they do other nectar sources. Henning (1983b) minimized the
importance of the honey gland and suggested that chemicals produced
by other organs (chiefly lenticles) were responsible for maintaining a
benign (even protective) attitude by ants toward lycaenid larvae. A
honey gland is not necessary for attracting ants since mymecophily
occurs in some taxa, such as Lycaena , which have no honey gland.
Among California lycaenids the honey gland is absent in the Lycaeninae
and Riodininae, but present in all Polyommatinae except A. franklinii
and in all Theclinae except H. grunus. The external gland opening is
usually surrounded by numerous lenticles and often by dendritic setae
as well. In some exotic species the base of the gland and/or the
surrounding cuticle is strongly sclerotized, but in the local fauna little
or no sclerotization is apparent. Although the surrounding setae may
provide useful diagnostic characters, the appearance of the honey gland
itself is of little use in discriminating the local fauna.
The arrangement of the crochets and presence of a spatulate lobe on
the prolegs are good characters for distinguishing most lycaenids.
Typically in this family there is a well developed bi- or triordinal
mesoseries of crochets which may be weakened or divided medially by a
fleshy lobe (fig. 72). Many taxa also have a less well developed lateral
series of crochets; in L. brassolis it is so well developed that an
essentially complete ring of crochets is apparent. In the California
fauna a lateroseries of crochets occurs in the Riodininae (except A.
mormo ), in the Lycaeninae (sometimes greatly reduced or absent on
some prolegs) and in H. grunus (fig. 72a). The fleshy lobe which often
divides the mesoseries of crochets is strongly spatulate in most species
but absent in both A. mormo and P. emigdionis.
Cephalic pigmentation is a useful diagnostic character for several
taxa. In most California Polyommatinae the cranium is uniformly dark
brown or blackish but in the Lycaeninae and Theclinae it is often
20
J.Res.Lepid.
yellowish. The cranium is usually dark in A. mormo but pale or
nonpigmented in the other three riodinine species. A narrow band of
dark pigment is usually associated with ocelli (stemmata) 1-5 (fig. 45);
ocelli are numbered as in fig. 68. One should not confuse this cuticular
infuscation with the subcuticular ocellar pigment which is often visible
in preserved specimens. A few local and many exotic polyommatine
species have light brown head color while some lycaenine and thecline
species have extensive dark cranial infuscation.
Cephalic setation is a relatively conservative trait in most lycaenid
groups. Cranial setae are usually few in number and very small (fig. 3)
except for those near the oral margin. More numerous short setae
(similar to neck setae) may occur, especially in the Lycaeninae and
Theclinae, on the frons (fig. 29) and ventral to the ocelli. The cranial
secondary setae of the Riodininae are more numerous and may be as
long as some prominent setae on the body (fig. 5).
The number of mandibular setae may be variable but is useful in
distinguishing some taxonomic groups. This character was examined
for only a limited number of specimens of each species. Although it has
been reported that riodinines have more than two mandibular setae
while other lycaenids have only two (Scott, 1986; Downey, 1987;
Harvey, 1987), there are numerous exceptions. In this study two
mandibular setae were found in all polyommatines, most theclines (fig.
55a), and in the Southeast Asian riodinine, Z. flegyas. But some
theclines have as many as six mandibular setae, while two to five were
found in the Lycaeninae (fig. 55b); 24 mandibular setae were found in
the riodinine M. pixe. Spalgis epeus has one madibular seta.
There are notable differences in the prothoracic shield among the
California taxa surveyed here. In the riodinines the shield is transverse,
rather strongly sclerotized, and adorned with numerous long setae
which overhang the head. In the other lycaenids the shield is about as
long as it is wide (sometimes longer), variably sclerotized, somewhat
recessed below the level of the surrounding cuticle (figs. 1,21), and has
setae which seldom extend much beyond its anterior margin. In the
Lycaeninae, Riodininae, and Theclinae the surface of the prothoracic
shield usually appears smooth (figs. 9, 11, 12, 14, 21). In polyommatines
the surface of the shield appears (in SEMs) honeycombed with a complex
ultrastructure of anastomozing ridges similar to, but more highly
developed than, those elsewhere on the body (figs. 10, 13); this surface
also occurs on the prothoracic shield of some theclinae such as A. hales us
(fig. 6).
Body coloration is highly variable in the Lycaenidae since most
species are cryptically colored to match their substrate. The predo-
minant ground color is green but pink, white, yellow, and brown are also
common. Except in the riodininae a disruptive pattern of contrasting
lines is often present. The latter may appear complex but usually can be
reduced to a few standard components.
27(1): 1-81, 1988
21
The following color pattern components are recognized: 1) middorsal
line, 2) subdorsal lines, 3) lateral lines, 4) lateral chevrons, 5)
transverse bar. The first three are longitudinal and generally extend
posteriorly from T2 or T3. The lateral chevrons extend posterolaterally
from the subdorsal area of one segment to near the lateral line on the
second segment behind it. These may appear as a series of parallel
diagonal lines or chevrons when the larva is viewed from above. In some
taxa there is a transverse bar of dark pigment on the first abdominal
segment. The transverse bar is always darker than the ground color and
is commonly reddish or brown; it varies in extent from a pair of
unconnected and relatively small dark dorsal spots, as in C. (C.)
dumetorum, C. (I.) augustus, and C. (M.) spinetorum (figs. 74-4b, -4d,
-5c), to a broad band extending across the dorsum toward the lateral
line, as in E. rita (fig. 74-8d). Variations in this basic pattern of lines
result from two factors: 1) each line may be highlighted dor sally and/or
ventrally by lines of contrastingly darker or lighter pigment and 2) the
degree to which each line and its bordering pigment are developed may
vary independently on each segment. Some or all lines may be absent or
reduced to discontinuous spots on some or all segments. Monophagous
and oligophagous species are usually mono- or oligomorphic while
polyphagous species are often polymorphic. These color pattern com-
ponents (except the transverse bar on Al) are illustrated in figure 1.
22
J. Res. Lepid.
KEY TO LAST INSTAR LYCAENXDAE OF CALIFORNIA
1 . The most conspicuous body setae clustered on verrucae; some setae on
pro thoracic shield extending anteriorly over head; A1 spiracles dis-
placed ventrally to a location slightly anteroventral to the lateral
verrucae (figs. 30, 69, 70); head not retractable. (Riodininae) 2.
1' Body setae not arranged in verrucae; no setae on pro thoracic shield
extending beyond anterior margin of prothorax; spiracles on first
abdominal segment in line with those on other abdominal segments
(fig. 1); head retractable ... 5.
2(1) Segments A1-A7 each with 2 pairs of dorsal verrucae consisting of
numerous fine, flexible plumose setae many of which are longer than
twice the head width (fig. 70); subdorsal verrucae absent; dorsal and
lateral body regions densely covered with echinoid setae (fig. 31)... 3 .
2' Segments A1-A7 each with 1 pair of dorsal verrucae consisting mostly
or entirely of short spine-like setae (fig. 30); subdorsal verrucae
present on segments T2-A7 (fig. 69); echinoid setae absent ... 4.
3(2) Some plumose setae on verrucae apically spatulate (figs. 32, 70); hosts
Baccharis glutinosa and Encelia calif ornica . . . Calephelis nemesis.
S' Verrucae lacking apically spatulate setae; host Bebbia juncea. . .
Calephelis wrighti.
4(2') Dorsal verrucae with darkly pigmented setae; dorsal verrucae on
segments A1-A7 (in most populations) also with one nonpigmented
plumose setae ca 3X as long as other verrucal setae (fig. 69); hosts
Eriogonum, Krameria, and Oxytheca . . . Apodemia mormo
4' All setae on dorsal verrucae on segments A1-A7 nonpigmented,
spinelike; host Prosopis. . . Apodemia palmerii.
5(1 ') Eversible tubercles present on A8 (fig. 1) and/or chalazae stellate (figs.
2e, 2f, 2j, 2k, 16, 71) and/or head uniformly black or brown (may be
darker around ocelli), ca 1/4 as wide as body; prothoracic shield not
pigmented (apparently nonsclerotized) . . . (Polyommatinae) 45.
5' Eversible tubercles absent; chalazae not stellate; head often yellowish
or bicolored, ca 1/3 as wide as body; prothoracic shield sclerotized,
often pigmented and acutely produced anteromedially ... 6.
6(5') Honey gland absent; mushroom setae present on dorsal and lateral
body regions (figs. 21-24); prothoracic shield lacking setae lateral to
sensory setae (figs. 21, 33-44); chalazae nonsculptured (figs. 2i, 2p) . . .
(Lycaeninae) 7.
6' Honey gland present on A7 and/or chalazae buttressed (figs. 2n, 2o, 4);
mushroom setae absent; prothoracic shield with some setae lateral to
sensory setae (figs. 45-54, 61, 64, 67) (Theclinae) 18.
7(3) Prominent setae absent on T2-A9; nearly all setae ca as long as
spiracle width, recumbent, truncate; sensory setae often apically
truncate-spatulate, at least 3X as long as dorsal setae on T2-A9 (fig.
27(1): 1-81, 1988
23
21); host Rhamnus crocea in San Diego Co. and northern Baja
California. . . Lycaenahermes,
7' At least some dorsal setae much longer than spiracle width and/or
erect; sensory setae not apically truncate-spatulate, seldom longer
than all dorsal setae on T2-A9; host not Rhamnus crocea ; more widely
distributed. . . 8.
8(70 Nonprominent dorsal and lateral setae on T2-A6 nonpigmented,
mostly ca 2X as long as spiracle width, recumbent, finely tapered (fig.
2i), often appearing whitish; host Eriogonum ... 9.
8' Nonprominent dorsal and lateral setae on T2-A6 more erect and/or
brownish, shorter, and not finely tapered; host not Eriogonum ... 10.
9(8) Distinct dorsal prominent setae present on T2-A8; some setae near
spiracles on A8 erect, straight and weakly dendritic (fig. 2p) . . .
Lycaena heteronea.
9' Dorsal prominent setae absent or indistinct on T2-A8; setae near
spiracles on A8 rarely erect and straight, never dendritic . . .
Lycaena gorgon.
10(8') Dendritic and short clavate-capitate setae present subdorsally and
near spiracles on A7, A8 (fig. 15) 1 1 .
10' No dendritic or clavate-capitate setae subdorsally or near spiracles on
A7, A8; on various hosts ... 13.
11(10) Head light brown but may be darker anteriorly (fig. 44); legs light
brown to nonpigmented; widely distributed mostly below 2000m . . .
Lycaena xanthoides.
11' Head dark brown, evenly pigmented throughout (figs. 42, 43); legs
dark brown; usually found above 2000m in central and northern
California... 12.
12(11') Dorsal and lateral prominent setae on T2-A8 ca 4X as long a spiracles;
no more than 25 secondary setae on prothoracic shield, most of which
are mushroom setae and the remainder are apically rounded (fig. 42)
Lycaena editha.
12' Dorsal and lateral prominent setae on segments T2-A8 ca 3X as long
as spiracles; at least 30 secondary setae on prothoracic shield of which
less than half are mushroom setae and the remainder are mostly
tapered (fig. 43) . . . Lycaena rubidus.
13(10') Head and legs predominantly dark brown (figs. 33, 34); all body setae
erect or nearly so, never strongly bent parallel to body surface; alpine
species usually found above 3000m; hosts Oxyria and R umex ... 14.
13' Legs yellowish or nonpigmented; head yellowish at least near vertex
(may be dark anteroventrally); some body setae may be strongly bent
nearly parallel to body surface; seldom found above 3000m; on various
hosts... 15.
14(13) Dorsal and lateral setae on T2-A8 shorter than sensory setae on
prothoracic shield, ca as long as spiracle width . . . Lycaena cupreus.
24
J. Res. Lepid.
14'
15(13')
15'
16(150
16'
17(160
17'
18(60
18'
19(180
19'
At least some dorsal and lateral setae on T2-A8 longer than sensory
setae on pro thoracic shield, ca 2X as long as spiracle width. . .
Lycaena phlaeas.
Longest dorsal prominent setae on T1 posterolateral to prothoracic
shield at most subequal to length of sensory setae; nonprominent setae
in same area bent caudad (fig. 23); all or most nonmushroom setae
tapered, apically pointed, erect to suberect (fig. 23), rarely bent
parallel to body surface; host R ibes . . . Lycaena arota.
Longest dorsal prominent setae on T1 posterior to prothoracic shield
longer than sensory setae; nonprominent setae in same area bent
cephalad (fig. 24); nonprominent lateral setae on T1-A9 often truncate
and recurved or bent parallel to body surface; host not R ibes ... 16.
Most nonprominent setae near posterolateral margins of prothoracic
shield bent nearly parallel to body surface (figs. 2c, 24); cephalic
infuscation extending across frons and posteriorly well beyond ocelli
(fig. 36); host Polygonum douglassii and perhaps other Polygonum and
Rumex species... Lycaena nivalis.
Most nonprominent setae near posterolateral margins of prothoracic
shield more erect, seldom bent at less than 45 degrees to body surface;
cephalic infuscation less extensive (may be confined to ocelli 1-5);
hosts Polygonum, Rumex, and Vaccinium. . . 17.
Most nonprominent dorsal setae on T2-A8 tapered, acutely pointed,
often bent nearly parallel to body surface; cephalic infuscation limited
to an arc enclosing ocelli 1-5 (fig. 40); widely distributed; hosts
Polygonum and Rumex. . . Lycaena helloides.
Most nonprominent dorsal setae on T2-A8 weakly tapered, truncate,
less bent (ca 45 degrees to body plane) (fig. 2d); cephalic infuscation
often more extensive (fig. 41); found in mountains of central and
northern California; host Vaccinium . . . Lycaena mariposa.
Honey gland absent; dorsal prominent setae present on A7; prolegs
with a lateroseries of crochets (fig. 72a); prothoracic shield smoothly
convex anteriorly and broadly rounded posterior to sensory setae (fig.
46); hosts Chrysolepis chrysophylla, Lithocarpus densiflora, and
Quercus chrysolepis . . . Habrodais grunus.
Honey gland present; dorsal prominent setae absent on A7; prolegs
lacking a lateroseries of crochets; prothoracic shield more-or-less ‘t’-
shaped, often acutely produced anteromedially and abruptly narrowed
posterior to sensory setae (figs. 45, 47-54, 61, 64, 67) ... 19.
All dorsal setae erect, straight, and tapered (figs. 7, 8) or recumbent
and ca as long as their chalazal width (fig. 2n); sensory setae often with
conspicuous lateral spicules (fig. 9) . . . 20.
Some dorsal setae (at least on A8) clavate-capitate (fig. 2g), strongly
bent, or recumbent (inclined caudad) and longer than their chalazal
width; sensory setae lacking conspicuous lateral spicules (as in fig.
10)... 38.
27(1): 1-81, 1988
25
20(19) Mushroom lenticles (figs. 8, 26) widely distributed on dorsal and
lateral body regions; dorsal prominent setae absent; prothoracic shield
white, outlined with black (fig. 45); sensory setae 2-many branched
(fig. 6); host Phoradendron . . . Atlides halesus.
20' Mushroom lenticles absent; dorsal prominent setae present or absent;
prothoracic shield not white, outlined with black; sensory setae not
branched; host not Phoradendron . . . 21.
21(20') Sensory setae longer than all dorsal setae on T3 (fig. 73), often
broadest in apical fourth (fig. 12); nonprominent dorsal setae shorter
than or subequal to spiracle width; host A rceuthobium . . . 22.
21' Sensory setae not longer than all dorsal setae on T3, not distinctly
broadest in apical fourth; some dorsal setae on T3 at least 2X as long as
spiracle width; not on Arceuthohium . . . 23.
22(21) Sensory setae ca 10X as long as other setae on prothoracic shield and
posterolateral to it on T1 (fig. 73); all or most dorsal setae on T2-A6
reclinate, ca half as long as spiracle width, subequal to chalazal
width . . . Callophrys (M.) spinetorum.
22' Sensory setae ca 5X as long as other setae on prothoracic shield and
subequal to longest setae posterolaterally adjacent to it; all dorsal
setae on T2-A6 erect, mostly 1/2- IX as long as spiracle width and
longer than their chalazal width . . . Callophrys (M.) johnsoni.
23(21) Dorsal prominences posterolateral to prothoracic shield well-developed,
with longest setae randomly distributed over them (fig. 8); host
Cupressaceae . . . 24.
23' Dorsal prominences posterolateral to prothoracic shield poorly deve-
loped, with longest setae arranged more-or-less in a transverse line
(fig. 7); host not Cupressaceae ... 27.
24(23) Hosts, Juniperus California and J. osteosperma from west end of San
Bernardino Mts. west and north in inner Coast Ranges to central
California, through Tehachapi Mts. to Walker Pass, and Mojave
Desert and Great Basin mountain ranges; also associated with J.
occidentalis from San Bernardino Mts. northward. . .
Callophrys (M.) siva.
24' Using other hosts and/or found else where. . . 25.
25(24') Restricted to vicinity of Otay Mt. in San Diego Co.; host Cupressus
forbesi . . . Callophrys (M.) thornei.
25 ' More widely distributed ; host not C. forbesi ... 26.
26(25') Host J. California from Mexican border north to San Bernardino and
Little San Bernardino Mts. . . . Callophrys (M.) loki.
26' Hosts Cupressus sargentii and Libocedrus decurrens (also J. California
rarely in inner coast ranges of central California) . . .
Callophrys (M.) nelsoni.
27(23') Cranial infuscation limited to a narrow crescent connecting ocelli 1-5
and not extending to ocellus 6 (figs. 59, 61-64), or much more extensive
across front and encompassing all ocelli (fig. 60) ... 28.
26
J.Res.Lepid.
27'
28(27')
28'
29(28)
29'
30(29')
30'
31(30')
31'
32(28')
32'
33(32')
33'
34(33')
34'
35(27)
35'
Cranial infuscation limited to ocellar area, connecting ocelli 1-5 and
extending posteriorly to anterior margin of ocellus 6 but not broadly
encompassing all ocelli (figs. 65-67) ... 35.
Cephalic infuscation limited to a narrow band connecting ocelli I-V;
host not Eriogonum or Lotus. . . 29.
Cephalic infuscation variable; host Eriogonum or Lotus . . . 32.
Frons slightly darker than remainder of head (fig. 64); host Sedum . . .
Callophrys (I.) mossii.
Fronto-clypeal area not darker than remainder of head; host not
Sedum. . . 30.
Head width greater than 1.5mm; longest dorsal setae on T2 ca 1.5X as
long as sensory setae; host Pinus . . . Callophrys (I.) eryphon.
Head width less than 1.5mm; longest dorsal setae on T2 usually less
than 1.3X as long as sensory setae; host not Pinus ... 31.
Ocellar infuscation forming a broad band connecting ocelli 1-5, ex-
tending anteriorly along margin of antennal insertion and posteriorly
half the distance from ocellus 5 to ocellus 6 (fig. 63); host Cowania
mexicana in mts. of central and eastern Mojave Desert. . .
Callophrys (I.) fotis.
Ocellar infuscation less extensive, not extending anteriorly along
margin of antennal insertion and posteriorly half the distance from
ocellus 5 to ocellus 6 (fig. 62); absent from mts. of central and eastern
Mojave Desert; host not C. mexicana . . . Callophrys (I.) augustus.
Usually below 1500m throughout cismontane California and the
desert slopes bordering the western Mojave and Colorado deserts . . .
Callophrys (C.) perplexa.
Found along central coast, in mountains of Mojave Desert, or above
2000m in Sierra Nevada northward. . . 33.
Host Eriogonum latifolium (and occasionally Lotus scoparius) along
coast from Monterey northward to Point Reyes . . .
Callophrys (C.) dumetorum.
Not found along coast of central California; host other Eriogonum
spp. ... 34.
Found mostly above 2000m in the Sierra Nevada, Siskiyou, and
Warner Mts. . . . Callophrys ( C.) lemherti.
Found in Mojave Desert mountains . . . Callophrys ( C.) comstocki.
Body with a saw-toothed dorsal profile due to paired dorsal promiences
on T3-A6 each with 1 (2 on T2) prominent seta ca 10X as long as
spiracles and 3-1 OX as long as surrounding setae (fig. 56c); hosts
Acacia and (primarily) Prosopis . . . Ministry mon leda.
Body lacking a saw-toothed dorsal profile; dorsal prominences on T2-
A6 indistinct or absent; dorsal prominent setae on segments T2-A6
absent or poorly differentiated, no more than 5X as long as spiracles;
on various hosts .. . 36.
27(1): 1-81, 1988
27
36(35')
36'
37(36')
37'
38(19')
38'
39(38')
39'
40(39)
40'
41(39')
41'
42(41')
42'
Chalazae milky, lighter than ground color (best seen in live larvae);
host Malvaceae primarily in desert and mountains from San Bernar-
dino Co. southwards . . . Strymon columella.
Chalazae not milky or noticeably lighter than ground color; many
hosts but especially Fabaceae, Malvaceae, and Polygonaceae ... 37.
Restricted to Catalina Island; hosts Lotus and Eriogonum . . .
Strymon avalona.
Widely distributed on many hosts . . . Strymon melinus.
Dendritic setae and clavate-capitate, apically truncate setae (fig. 2g)
present dorsally on A7 and A8, and laterally on Tl, T3, Al, and A7
(between spiracles and margin of honey gland); host Prunus vir-
giniana . . . Harkenclenus titus.
At least some dorsal setae on A7 and A8 recumbent, bent, or inclined
caudad (fig. 2o), not clavate-capitate; distribution of dendritic setae
variable ... 39.
Head dark brown (except along adfrontal sutures); dendritic setae on
Tl lateral to prothoracic shield; all dorsal setae on T2-A6 erect,
straight... 40.
Head at least partly yellowish or light brown; no dendritic setae on Tl
lateral to prothoracic shield; dorsal setae on T2-A6 variable ... 41 .
Legs dark brown; nondendritic dorsal setae on A7, A8 dark brown,
acutely tapered, reclinate (nearly parallel to body surface) (fig. 2n);
hosts Ceanothus, Quercus, and probably other woody perennials. . .
Satyrium californica.
Legs nonpigmented; nondendritic dorsal setae on A7, A8 nonpig-
mented, suberect, often inclined caudad at ca 45 degrees to the body
plane, more-or-less clavate and apically rounded (fig. 2o); host
Lupinus. . . Satyrium fuliginosum.
Prominent setae absent on A1-A6; dorsal setae mostly recumbent,
broadest near middle, strongly dentate, (fig. 17); dendritic setae
present subdorsally on segments T2, T3 (fig. 17); sensory setae at least
3X as long as other setae on and posterolaterally adjacent to prothor-
acic shield; host Purshia . . . Satyrium behrii.
Segments A1-A6 with some prominent setae and/or some dorsal setae
erect, cylindric; dendritic setae absent subdorsally on segments T2,
T3; sensory setae no more than 2X as long as other setae on and
posterolaterally adjacent to prothoracic shield; host not Purshia . . .
42.
Prominent dorsal setae on segments T2-A6 erect, straight, orange-
brown, forming a pair of dorsal bands each comprised of at least 18
setae per segment (fig. 56a); sensory setae tapered; 2-4 dendritic setae
present near sensory setae on prothoracic shield; host Cercocarpus . . .
Satyrium tetra.
Prominent dorsal setae on T2-A6 less numerous (or absent), variable
in pigmentation; sensory setae clavate to tapered; dendritic setae
absent on prothoracic shield; host not Cercocarpus ... 43 .
28
J.Res.Lepid,
43(42') Cephalic infuscation limited to a narrow band enclosing ocelli 1-5 (fig.
50); dorsal prominent setae on segments T2-A6 erect, ca 3-4X as long
as spiracles, host Salix . . . Satyrium sylvinus.
43' Cephalic infuscation extending across front and posterolaterally at
least to ocellus 6 (figs. 49, 51); dorsal prominent setae on T2-A6 absent
or 1-2X as long as spiracles; host not Salix ... 44.
44(43 ') Dorsal prominent setae inconspicuous, erect, ca 2X as long as spiracles
and other dorsal setae (fig. 56b); all dorsal setae cylindric, orange-
brown; sensory setae less than 2X as long as longest setae postero-
laterally adjacent to prothoracic shield; host Quercus. . .
Satyrium auretorum.
44' Dorsal prominent setae absent or obscure; dorsal and lateral setae of
two types: 1) erect, tapered, cylindric, and pale brownish and 2)
recumbent, flattened (as in fig. 17), and nonpigmented; sensory setae
ca 2X as long as longest setae posterolateral to prothoracic shield; host
Ceanothus . . . Satyrium saepium.
45(5) Prominent setae only at anterior margin of prothorax; dorsal and
lateral setae on T2-A6 recurved, clavate-capitate, ca as long as
spiracles; sensory setae tapered; hosts Chenopodiaceae (especially
Atriplex and Chenopodium ) and Sesuvium verrucosum (Aizoaceae) . . .
Brephidium exilis.
45' At least a few prominent lateral setae on one or more of segments T2-
A6 and/or sensory setae not tapered; dorsal and lateral setae on
segments T2-A6 variable in structure; host not Chenopodiaceae (except
Atriplex canescens) or S. verrucosum . . . 46.
46(45') Sensory setae spatulate or apically broadened (fig. 13) . . . 47.
46' Sensory setae finely tapered (fig. 10) . . . 53.
47(46) Chalazae strongly stellate, lateral points often much longer than
basal width of seta (fig. 2e); on various hosts ... 48.
47' Chalazae less stellate, lateral points (if present) seldom longer than
basal width of seta (fig. 2j); hosts Eriogonum and Oxy theca ... 49.
48(47) Dorsal setae on T3-A6 erect to suberect or broadly recurved and finely
tapered (fig. 2b), not sharply bent near base; dendritic setae only near
honey gland; hosts Fabaceae and Plumbago . . . Leptotes marina.
48' Dorsal setae on T3-A6 recumbent, strongly bent near base (fig. 2f);
dendritic setae near A1 spiracles and honey gland, and on prothoracic
shield; many hosts but not Plumbago . . . Celastrina argiolus.
49(47') Eversible tubercles absent; chalazae not stellate; host Eriogonum
reniforme or Oxytheca . . . Philotiella speciosa.
49 ' Eversible tubercles present; chalazae stellate; host various Eriogonum
species... 50.
50(49') Head dark blackish brown; legs much darker than body venter, nearly
as dark as the head; dorsal prominences on T2-A6 well-defined, each
with at least a pair of prominent setae directed posteromedially (fig.
27(1): 1-81, 1988
29
50'
51(50')
51'
52(51')
52'
53(46')
53'
54(53')
54'
55(54')
55'
56(55')
56'
57(56')
57'
57c); found in Great Basin mountain ranges, the east slope of the
Sierra Nevada, and foothills bordering the Mojave Desert; hosts
Eriogonum davidsonii, E. deflexum, E. microthecum, E. plumatella, E.
roseum,E. wrightii , and perhaps#, heermannii . . . Euphilotes rita.
Head medium to dark brown; legs nonpigmented to light brown, not
nearly as dark as head; dorsal prominences on T2-A6 moderately to
weakly developed, with prominent setae directed more posteriorly (fig.
57b) or absent (fig. 57a); host various Eriogonum species. . . 51.
At least two dorsal prominent setae on each of segments T2-A6 (fig.
57b); legs lightly pigmented; host Eriogonum pusillum or E. reniforme
in the Mojave Desert from late winter to early summer. . .
Euphilotes mojave.
Less than two dorsal prominent setae on each of segments T2-A6 and
often none (fig. 57a); legs nonpigmented; host not E. pusillum or E.
reniforme; widely distributed . . . 52.
Host many species of Eriogonum but not E. davidsonii , E. elongatum,
E. latifolium,E . nudum , or#, wrightii . . . Euphilotes hattoides.
Host many species of Eriogonum but not#, fasciculatum, #. heermannii,
#. microthecum , or #. ovalifolium (in California) . . .
Euphilotes enoptes.
Honey gland and eversible tubercles absent; montane in northern and
central California; host Primulaceae, especially Dodecatheon . . .
Agriades franklinii.
Honey gland present; eversible tubercles present or at least a circle of
setae marks the location where they should be; host not Primulaceae
23.. 54.
Prolegs lacking a spatulate lobe; segments T2-A6 often with a pair of
dorsal prominent setae ca 10X as long as spiracle width; chalazae
apparently buttressed (fig. 10); host Atriplex canescens. . .
Plehulina emigdionis.
Prolegs with a spatulate lobe; setation variable; chalaze not apparently
buttressed; host not A . canescens ... 55.
Dorsal setae on T2-A6 mostly erect, tapered to clavate or capitate,
shorter than spiracle width; setae around honey gland capitate (fig.
25); host Dudleya. . . Philotes sonorensis.
Dorsal setae on T2-A6 variable but not clavate or capitate; setae
around honey gland variable; host not Dudleya ... 56.
Lateral margins of spatulate lobes on prolegs pigmented, apparently
sclerotized (fig. 72b) .. . 57.
Spatulate lobes on prolegs nonpigmented, not apparently sclerotized . . .
58.
Nondendritic dorsal setae on A7, A8 erect and straight, or weakly bent
(as in Fig. 2j); usually in Astragalus seed pods . . . Everes amyntula.
Nondendritic dorsal setae on A7, A8 mostly moderately to strongly bent
30
58(56')
58'
59(58)
59'
60(58')
60'
61(60')
61'
62(61)
62'
63(62')
63'
J.Res.Lepid.
(as in figs. 2c, 2k); host various herbaceous Fabaceae including Astra-
galus, Lotus , and Vicia . . . Everes comyntas.
Dendritic setae present laterally on A6-A8 and/or most subdorsal setae
on T2-A6 erect, clavate-capitate (fig. 2j); legs nonpigmented, not darker
than body venter; host herbaceous Fabaceae, especially Astragalus,
Lotus , and Lupinus ... 59.
Dendritic setae not present laterally on A6-A8; most subdorsal setae on
T2-A6 not erect and clavate-capitate; leg pigmentation variable, may be
darker than body venter; hosts Eriogonum and various Fabaceae ... 60 .
Prothoracic shield lacking prominent setae, the sensory setae at least 2X
as long as other setae on the shield; most dorsal setae on A7, A8 (between
spiracles) moderately to strongly bent (fig. 2k), not capitate; host
Lupinus. . . Glaucopsychepiasus.
Prominent setae usually present on prothoracic shield, the sensory setae
often shorter than some other setae on the shield; dorsal setae on A7, A8
erect, mostly clavate-capitate (similar to fig. 2h) or bent apically (fig. 2j);
not only on Lupinus . . . Lycaeides idas and L. melissa.
Dendritic setae present near spiracles on A2; legs not darker than body
venter; host Astragalus, Lotus , or Lupinus . . . Glaucopsyche lygdamus.
Dendritic setae not present on A2; leg color variable; host Eriogonum or
various Fabaceae .. . 61.
All dorsal setae on T2-A6 erect, straight; leg color variable; host not
Eriogonum. . . 62.
Some dorsal setae on T2-A6 recurved or recumbent; legs not pigmented;
host Eriogonum or F abaceae . . . 64.
Legs nonpigmented, not darker than body venter; dorsal prominent
setae in T2, T3 no more than 3X as long as other dorsal setae; widely
distributed; host Lupinus . . . Icaricia icarioides.
Legs brown, much darker than body venter; dorsal prominent setae on
T2, T3 at least 4X as long as other dorsal setae; above 2000m in central
and northern California; host not only Lupinus . . . 63.
Dendritic setae present in area between honey gland and spiracles on A7 ;
chalazae on prothoracic shield much paler than legs; chalazae anterior
to prothoracic shield with lateral points less than 1/4 as long as basal
width of chalaza; host Trifolium. . . Plebejus saepiolus.
Dendritic setae on A7 only at lateral margins of honey gland; chalazae
on prothoracic shield as dark as legs; chalazae anterior to prothoracic
shield with lateral points greater than 1/2 as long as basal width of
chalaza; host prostrate Astragalus and Lupinus . . . Icaricia shasta.
64(61') Dorsal setae on A8 and A9 and nondendritic setae near abdominal
spiracles mostly bent parallel to body, broadest near middle, flattened
in body plane , and pointed (as in fig. 21). . . 65.
27(1): 1-81, 1988
31
64'
65(64)
65'
Dorsal setae on A8, A9 erect to strongly bent, cylindric, not broadest
near middle, and mostly blunt (fig. 2m); nondendritic setae near
abdominal spiracles gradually tapered to a blunt tip (as in fig.
2c) . . . Icaricia acmon, I. lupini, and/, neurona.
Longest dorsal prominent seta on T2 as long or slightly longer than
longest seta on prothoracic shield; segments T3-A6 usually with at
least 4 dorsal prominent setae (fig. 58b); no dendritic setae near
spiracles on Al; hosts F abaceae . . . Hemiargus isola.
Longest dorsal prominent setae on T2 ca 2/S-3/4 as long as longest
setae on prothoracic shield; segments T3-A6 usually with only 2 dorsal
prominent setae (fig. 58a); dendritic setae present or absent near
spiracles on Al; hosts Eriogonum and Fabaceae. . .
Hemiargus ceraunus.
32
J.Res.Lepid.
DIAGNOSTIC SECTION
The larvae of the California lycaenids share several morphological
traits which distinguish them from other families of Lepidoptera. These
features have been discussed in some detail in the morphology section
above. The presence of lenticles, a spatulate lobe on the prolegs, and an
onisciform body shape typically separate the California lycaenids from
other Lepidoptera larvae. These characters are not without exceptions:
lenticles also occur in hesperiids, a spatulate lobe is present on some
prolegs of some geometrid moth larvae while absent in two California
lycaenids, and the Riodininae are only weakly onisciform.
Riodininae
Most riodinine larvae possess an unusual arrangement of spiracles.
Those on the first abdominal segment are displaced anteroventrally far
below the latitude of the other abdominal spiracles; they are located
slightly anteroventral to the lateral verrucae (figs. 30, 69, 70) and are
often hidden by an intersegmental fold.
Other characters which distinguish these larvae from other lycaenid
larvae in California are a transverse prothoracic shield with some setae
long enough to overhang the head and the most prominent body setae
clustered in verrucae. The distribution of verrucae differs in the two
genera represented.
The number of larval instars may be variable in all four California
riodinines but the minimum (and usual) number for all populations
examined is five. Both Dos Passes (1936) and McAlpine (1938) found 8-9
instars in Calephelis borealis and C. muticum, respectively. It is not
certain whether true diapause occurs in any of the four California
riodinines but partially grown larvae of A, mormo, A. palmerii , and C.
nemesis were found on their host plants during winter. Larvae of C.
nemesis were observed to feed briefly (ca one hour) in early afternoon
each day dming winter when the temperature exceeded 12.8 degrees C.
before returning to habitual resting sites on dead foliage. Comstock
(1936) also reported intermittent winter feeding behavior in larvae of A.
palmerii. Some populations of A. mormo probably overwinter as ova.
Apodemia
There are two species of Apodemia in California. Their larvae have
paired dorsal, subdorsal, and lateral verrucae on segments T2-A8 (fig.
69); the prothorax has paired subdorsal and lateral verrucae and a
transverse shield which covers most of the dorsum. The verrucae and
prothoracic shield primarily contain stiff straight (prominent) setae;
these are dark brown in A. mormo but much lighter (often non-
pigmented) in A. palmerii. Several plumose setae arise from each
lateral verruca and the prothoracic shield, but are absent from the
27(1): 1-81, 1988
33
subdorsal verrucae. In most populations of A. mormo a single plumose
seta arises centrally from some or all dorsal verrucae (fig. 69) but these
setae are absent from the dorsal verrucae of A. palmerii. Additional
setae structurally similar to prominent setae on the verrucae, but
somewhat smaller and paler, are scattered over the dorsal and lateral
body surfaces of both species. Other distinguishing characters include a
pale green or pinkish ground color and yellowish head for A. palmerii
(fig. 74- lb) vs. a predominantly brown or violet-brown ground color
(with yellow dorsal and dorsolateral verrucae) and usually brownish
head for A. mormo (fig. 74-la).
The most widely distributed species, A. mormo , feeds primarily on
perennial species of Eriogonum , but ova have been collected (and larvae
reared) on Oxy theca perfoliata (both Polygonaceae). Krameria (Kra-
meriaceae) is a host of A. mormo in Texas (Kendall, 1976); it is also
acceptable to locally collected A. mormo. Various populations of A.
mormo occur throughout California. Apodemia palmerii occurs in the
southern deserts and feeds on Prosopis (Fabaceae).
Calephelis
There are two species of Calephelis in California; C. nemesis occurs
primarily in riparian situations in southern California, while C. wrighti
occurs in xeric habitats in southern and eastern California. The larval
hosts of the former species are Baccharis glutinosa and Encelia Califor-
nia (Emmel and Emrriel. 1973), while larvae of the latter feed on Bebbia
juncea , all in the Asteraceae. The larval ground color of both species is
cream or buff (fig. 74-lc). They have one pair of dorsal verrucae on T2,
T3, and A8, but two pairs on A1-A7; subdorsal verrucae are absent, but a
single pair of lateral verrucae occurs on T1-A8 (fig. 76). The verrucae
consist mostly of very long, slender, nonpigmented or pale brown
plumose setae which give the larva a feather-like appearance; similar
setae occur on the prothoracic shield. There are no straight, stiff, spine-
like setae as in Apodemia but echinoid setae (fig. 31) cover most of the
lateral body areas. The head is yellowish with a small amount of
brownish mottling. In C. nemesis many plumose setae (ca 1/4- 1/2 as long
as the longest ones) on each verruca are broadly spatulate apically (figs.
32, 70), while in C. wrighti no spatulate setae occur. In both species a few
short, clavate, plumose setae are present on the dorsal verrucae and
often near the spiracles. These are mostly nonpigmented, but those on
the dorsal verrucae on segments A1 and A7 are often black (especially in
C. wrighti).
Lycaeninae
The Lycaeninae is one of the smallest yet widely distributed lycaenid
subfamilies and may be better represented in California (twelve species)
than in any other region of equivalent size. Most local species are
34
J.Res.Lepid.
univoltine, diapause as ova, and use host plants in the Polygonaceae.
For many years these were placed in the genus Lycaena , but Miller and
Brown (1979) divide them among six genera. On the basis of similar
biology and/or morphology of immature stages, it is convenient to
discuss them as five groups which do not coincide with the generic
arrangement of Miller and Brown (1979). Pending further comparative
studies, we retain the single genus Lycaena for the group.
Lycaena
The larvae of all members of this genus lack a honey gland and
eversible tubercles. The length of the prothoracic shield is about twice
as great as its width. It is acutely pointed at the anterior, posterior, and
lateral extremities and is generally diamond-or ‘t’-shaped (figs. 33-44).
Although the width is greatest along the line of the sensory setae (in the
anterior third), there is typically a second lateral expansion in the
posterior third. Prominent setae are absent from the prothoracic shield;
other secondary setae on it are much shorter than the sensory setae and
never occur lateral to them. The sensory setae usually appear finely
tapered and devoid of lateral spicules; but minute lateral spicules
(visible with SEM) may occur near the apex in some species such as L.
mariposa. Mushroom setae, which are unique to this group, are scattered
over the dorsal and lateral body regions. Although mushroom setae may
occur as early as the second instar, they are more likely to appear first in
the third or fourth instar. Dendritic setae are present only in Lycaena
editha, L. heteronea, L. rubidus , and L. xanthoides. The chalazae of all
setae are nonsculptured or very weakly buttressed. Typically, a latero-
series of crochets (in addition to the mesoseries) occurs on the prolegs
but it may be greatly reduced or absent on some prolegs. Lycaena editha ,
L. rubidus , andL. xanthoides typically have three mandibular setae (L.
xanthoides rarely has up to five) while the other Lycaena species
typically have two mandibular setae. But in some populations of L.
gorgon and L. heteronea there may be three mandibular setae.
Two species which differ similarly in larval morphology and biology
from the others are L. cupreus and L. phlaeas. In California both occur
mostly above 3000m, are univoltine, and typically have four larval
instars; L. phlaeas sometimes has five instars. Both species probably
diapause as larvae. When reared from ova in the lab at 25-27 degrees C.
some larvae of L. cupreus often complete development but most enter
diapause in the third instar. Under the same conditions L. phlaeas is
continously brooded. In nature mature larvae of both species were found
in mid- July and younger larvae in August. In California the larvae ofL.
cupreus are associated with Rumex , especially R. paucifolius , and
larvae of L. phlaeas are associated with Oxyria digyna. Elsewhere L.
cupreus has also been reported to utilize O. digyna , while L. phlaeas
often uses Rumex (Ferris, 1974). The ground color of both species is
green in California, but L. phlaeas larvae from Asia and Europe may be
27(1): 1-81, 1988
35
pink or green and some California (and Old World) specimens have a
pinkish dorsal and/or lateral line (fig. 74-3b). In California they are
distinguishable from other Lycaena species by their darkly pigmented
head and legs, pale prothoracic shield (figs. 33, 34), and lack of
prominent dorsal setae. All dorsal and lateral setae on T2-A8, aside
from mushroom setae, are uniformly short, erect, tapered, and brownish.
In California L. cupreus larvae these setae are shorter than the sensory
setae and about as long as the spiracle width, while inL. phlaeas at least
some are longer than the sensory setae and about twice as long as the
spiracle width. Larvae of L. cupreus snowi (Edwards) from Colorado
have setae about as long as those of L. phlaeas. Larvae of L. phlaeas
examined from Corsica, Japan, and the eastern U.S. are similar to
California specimens in setation but have much lighter crania and legs.
Another group with similar biologies and larval morphology consists
of L. editha, L. rubidus , and L. xanthoides. Larvae of all three feed on
Rumex and are myrmecophilous; old records ofL. editha larvae feeding
on Horkelia and Potentilla (Rosaceae) (reiterated by Johnson and
Balogh, 1977 and Pyle, 1981) are not supported by our observations. The
ground color of these larvae varies from green to maroon or rust-red and
often there is a maroon middorsal line (fig. 74-3c). All dorsal and lateral
setae are erect to suberect and, except for mushroom setae, brownish.
The most distinctive features of these larvae are well developed dendritic
setae near the spiracles, especially on Tl, A7, and A8 and short clavate-
capitate setae dorsal to the spiracles, especially on A7 and A8 (fig. 15). In
each species the cranium and prothoracic shield are dark; L. editha and
L. rubidus also have dark legs. The head and prothoracic shield of L.
xanthoides are lighter than in the other species (figs. 42-44); dendritic
setae usually occur only near the spiracles on Tl, A7, and A8, but in
some populations these setae also occur on A1-A4. In L. editha and L.
rubidus larvae dendritic setae are present near the spiracles on Tl, A7,
A8, at least some of segments A1-A6, and frequently laterally on T2 and
T3. The larvae of L. editha and L. rubidus differ in prothoracic shield
setation as described in the key. Lycaena editha and L. rubidus are
sometimes sympatric and occur mostly above 2000m in central and
northern California but L. xanthoides is more widely distributed,
mostly below 2000m, and does not occur with the others. Adults of L.
editha found below 2000m in the vicinity of Mount Shasta are similar
(especially in size) to L. xanthoides and have been considered inter-
mediate between those species (Scott, 1980); but larvae from this
population (at Dunsmuir and Mount Shasta City) are more similar to
Sierra Nevadan L. editha and key to that species.
Two other closely allied species are L. gorgon and L. heteronea. They
are distinguished from the other Lycaena species by the use of Eriogonum
as a larval host and by the presence of numerous nonpigmented,
recumbent setae at least twice as long as the spiracles. These setae have
a finely granular surface giving them a whitish appearance. The larval
36
J.Res.Lepid.
color is pale turquoise to green for L. gorgon (fig. 74-2a) and dull blue-
green to green for L. heteronea (fig. 74-2c); both species are faintly
mottled but devoid of strongly contrasting markings; some L. heteronea
larvae have a pale yellow or white lateral line. The cranium is
nonpigmented in L. gorgon (fig. 37) and nonpigmented to somewhat
brownish in L. heteronea (fig. 38). Lycaena heteronea occurs mostly
above 1500m from Mount Pinos in Ventura Country northward through
the Sierra Nevada, Cascade, and Warner ranges and at sea level along
the coast near Point Reyes. Lycaena gorgon is primarily cismontane in
distribution below 1500m but also occurs up to about 2000m along the
east slope of the Sierra Nevada from southern Mono County southward
and in the Warner Mountains.
Usually these species are easily distinguished according to their
setation. Lycaena heteronea has distinct dorsal prominent setae (usually
somewhat melanic) on T2-A8 and erect dendritic setae near the spiracles
on A7 and A8 (also occasionally on T1 and rarely on the prothoracic
shield); nonprominent dorsal setae are mostly 1.5-2X as long as the
spiracle width. Lycaena gorgon lacks both prominent dorsal setae and
dendritic setae; the dorsal setae are mostly 2-4X as long as the spiracle
width. But in Lycaena gorgon larvae from the Warner Mountains of
Modoc County the setae are more erect (recumbent to suberect) than in
larvae from elsewhere and some dorsal setae may be nearly as erect as
the dorsal prominent setae ofL. heteronea. Occasionally, they also have
1-2 erect (but not dendritic) setae near the A7 and A8 spiracles. Larvae
of L. heteronea from near Mount Lassen resemble those of L. gorgon
from the Warner Mountains in setal erectness; their dorsal prominent
setae are less erect and less distinct than in other populations of L.
heteronea and are nonpigmented. In these larvae the dendritic setae
near the spiracles on A7 and A8 are poorly developed; there may be only
3-4 of them and with lateral spicules very short and confined to the apex.
In both of these populations the mandibles frequently have three setae
but other populations of both species typically have two mandibular
setae.
Lycaena hermes larvae differ greatly from those of the other Lycaena
species in host, range, and morphology. They feed on Rhamnus crocea
(Rhamnaceae) and are restricted to San Diego County and northern
Baja California, Mexico. They are bright, light green without strongly
contrasting markings (fig. 74-2b), though a pair of pale yellowish dorsal
lines may be present. These larvae may appear glabrous since all dorsal
and lateral setae on T2-A8 are sparsely scattered, subequal in length to
the spiracles, and recumbent; the setae are lightly pigmented and
weakly tapered to blunt- tipped. The sensory setae on the prothoracic
shield are at least twice as long as all other dorsal and lateral setae on
T2-A8 and subequal to the longest setae at the anterior margin of the
prothorax; they may be filiform or tapered, as in other Lycaena species,
or apically truncate-spatulate (fig. 21). The cranium is nonpigmented
and the prothoracic shield is rather small and narrow (fig. 39).
27(1): 1-81, 1988
37
The larvae of the four remaining Lycaena species are similar in
setation but different in biology. Lycaena helloides and L. nivalis feed on
Polygonum and (at least in the lab) Rumex (Polygonaceae); some
populations of L. helloides also feed on Potentilla (Rosaceae) (Shapiro,
1974). Lycaena arota feeds on Ribes (Saxifragaceae) and L. mariposa
feeds on Vaccinium (Ericaceae) (Pratt and Ballmer, 1986). They have
well-defined dorsal prominent setae on T2-A8 while most other dorsal
and lateral setae (not mushroom setae) on T2-A8 are erect to recumbent
and more-or-less tapered but apically truncate (figs. 2c, 2d). InL. arota
dorsal setae on T2 posterolateral to the shield are curved caudad (fig. 23)
while in the other species these setae are curved cephalad (fig. 24). The
ground color is green for all four species. Some populations of L. arota
(especially in southern California) have white or yellowish paired
dorsal and lateral lines (fig. 74-ld). Larvae of L. nivalis sometimes have
a maroon middorsal line and an indistinct yellowish lateral line (fig. 74-
3a); some larvae ofL. helloides andL. mariposa also have an indistinct
yellowish lateral line (fig. 74-2d). Cephalic infuscation in L. nivalis
extends across the front of the head and posteriorly well beyond the
ocelli (fig. 36), while in L. arota (fig. 35) and L. helloides (fig. 40) it is
limited to a narrow cresent connecting ocelli 1-5. In L. mariposa (fig. 41)
cephalic infuscation varies from nearly as extensive as in L. nivalis to
virtually absent. Although L. helloides is multivoltine (diapause stage
not known) and widely distributed (mostly below 2000m) the others are
univoltine and mostly confined to higher elevations in central and
northern California.
Theclinae
The most distinctive features of the California thecline larvae are the
head width (ca half as great as the body), lack of eversible tubercles, and
presence of buttressed chalazae. The pro thoracic shield is sclerotized,
frequently brownish, and lacks secondary setae longer than the sensory
setae; the head color ranges from yellowish to dark brown. The single
representative of the tribe Theclini, H. grunus, lacks a honey gland, has
a broadly diamond-shaped prothoracic shield (broadly rounded posterior
to the sensory setae), five mandibular setae, and a biordinal lateroseries
of crochets. All other species belong to the tribe Eumaeini. They have a
honey gland, a more-or-less T-shaped prothoracic shield (abruptly
constricted posterior to the sensory setae), usually two mandibular
setae, and lack crochet lateroseries on the prolegs. In general appear-
ance, thecline larvae (especially H. grunus) are most similar to those of
the Lycaeninae; they may be distinguished from the latter by the
presence of buttressed chalazae, setae on the prothoracic shield lateral
to the sensory setae, usual presence of a honey gland, and absence of
mushroom setae.
Among the Eumaeini two groups may be distinguished according to
setation. In one group consisting of Atlides , Callophrys, Ministrymon,
and Strymon all setae are cylindric, tapered, and straight or slightly
38
J.Res.Lepid.
curved with dendritic setae absent or inconspicuous (often poorly
developed or obscure) and confined to the margins of the honey gland.
Also in this group, the sensory setae are filiform to slightly clavate
(branched in Atlides) and often have conspicuous lateral spicules; the
mandibles have two setae. The other group, consisting of Harkenclenus
and Satyrium, has a broad range of setal forms including erect,
recurved, tapered, and clavate; dendritic setae occur prominently around
the honey gland and often on other segments, while other dorsal setae
on A7 and A8 are often recumbent or somewhat capitate. Also in this
group, the sensory setae are filiform, tapered, or spatulate and have
inconspicuous lateral spicules; the mandibles have 2-6 setae. Coin-
cidentally, diapause occurs as pupae in the former group and as ova in
the latter. Most California thecline species are univoltine and restricted
to a few closely related larval hosts. A few species are bi- or trivoltine,
while only S. melinus is continuously brooded and known to utilize a
wide range of larval hosts.
Atlides
The single California species, A. halesus , occurs throughout the state
but is more abundant in the south. It is often encountered in lowland
riparian habitats where the larval mistletoe host may be abundant.
Features which distinguish larvae of this species from other California
theclines include a velvety texture due to an even distribution of short
reddish-brown setae (no prominent setae), uniform green color (fig. 74-
4a) (rarely obscurely mottled), presence of mushroom chalazae (fig. 26),
branched sensory setae (fig. 6), a white prothoracic shield outlined in
black (fig. 45), and use of Phoradendron (Viscaceae) as a larval host.
There are 3-4 annual broods in the south, but probably 2-3 in the north;
the pupae overwinter.
Callophrys
The genus Callophrys was redefined by Clench (1961) to contain six
subgenera three of which ( Callophrys , Incisalia, and Mitoura) occur in
California; some authors including Miller and Brown (1981) give these
taxa full generic status. The systematics of this group is in need of
review and it seems unlikely that any study lacking comprehensive
biological, morphological, and/or biochemical data can resolve existing
controversies. The morphology of mature larvae is useful in distingui-
shing some subgenera, but of little use in distinguishing most species.
Yet the taxonomic limits of both species and subgenera are often
definable by biological differences such as host preference, habitat
selection, and number of instars.
In California, we provisionally recognize four species each in the
subgenera Callophrys and Incisalia and six in Mitoura ; Scott (1986)
recognizes only three species each of Callophrys and Mitoura in
27(1): 1-81. 1988
39
California, but his evidence is not compelling. Adding to the confusion
is a recent nomenclatural change; the name C. dumetorum , previously
applied to most lowland Callophrys populations in California, properly
refers only to those along the central coast previously known as C.
viridis (W. H. Edwards), which is now relegated to a junior synonym (J.
F. Emmel, in lift.). The name Callophrys perplexa , formerly considered
the southern California subspecies of C. dumetorum (in its former
usage), now becomes the senior synonym and must be applied to the
remaining lowland cismontane populations of this species.
There is little difference in larval morphology among the California
species of Callophrys and Incisalia (and also Strymon ). All are covered
with erect, straight, tapered setae of varying lengths which are non-
pigmented or pale brown and apically darkened. Prominent setae on T2-
A6 are absent or obscure. The sensory setae are filiform to slightly
broader in the apical half and have conspicuous lateral spicules (fig. 7);
rarely one or both may be bifurcate. Dorsal prominences on T2-A6 are
weakly to moderately developed, while those on T1 anterior and
posterior to the prothoracic shield are weakly developed. The most
prominent dorsal setae on T1 posterolateral to the shield are arranged
in one or two transverse rows (fig. 7). Relatively inconspicuous dendritic
setae occur near the honey gland. The head is yellowish-brown with
darker pigment confined to an arc connecting ocelli 1-5 but not extending
to ocellus 6 (fig. 59) or the entire anteroventral half of the head may be
dark brown, broadly enclosing all ocelli (fig. 60).
Mitoura larvae are easily distinguishable from those of the other
subgenera. Those species which utilize Cupressaceae as a larval host
have well developed dorsal prominences on T1 anterior and (especially)
posterior to the prothoracic shield often causing it to appear sunken; the
most prominent setae on the prominences posterior to the shield are
randomly distributed (fig. 8); dendritic setae are absent. In those species
which utilize Viscaceae as a larval host the dorsal prominences on T1
posterior to the shield are poorly developed and have prominent setae
arranged in a transverse row (as in the other subgenera) or absent (fig.
73); but distinct dorsal prominences are present at least on T2 and A6;
they also have dendritic setae near the honey gland.
No consistent morphological distinctions were found to separate
larvae of Callophrys and Incisalia. Larvae of species in these subgenera
are best distinguished according to distribution and host plant. In
California Callophrys larvae utilize Eriogonum (Polygonaceae) and
Lotus (Fabaceae), while Incisalia larvae feed on various hosts in other
plant families. There are four larval instars in all species of these
subgenera in California.
Cephalic pigmentation is useful in distinguishing some Callophrys
populations. Larvae of C. dumetorum (formerly C. viridis) from near
San Francisco and C. perplexa (formerly C. dumetorum) from southern
California have dark cephalic pigment narrowly confined to ocelli 1-5
40
J. Res. Lepid.
(fig. 59). In larvae of most other California Callophrys populations the
head is dorsally yellowish-brown and ventrally dark brown with dark
pigment broadly enclosing all ocelli (fig. 60), but larvae of C. dumetorum
from near Monterey are variable in cephalic pigmentation. The dorsal
profile can also be used to discriminate some taxa. The dorsal pro-
minences in C. comstocki, C. dumetorum , and C. lemberti often (but not
invariably) create a saw-toothed profile (fig. 74-4b), while those in C.
perplexa are more rounded (fig. 74-4c). All the Callophrys species use
Eriogonum as a larval host; C. perplexa also commonly uses Lotus
crassifolius and L. scoparius; C. dumetorum is reported to use L.
scoparius in the San Francisco area (Gorelick, 1971). Callophrys com-
stocki occurs in several Mojave Desert mountain ranges; C. perplexa
occurs throughout cismontane California up to ca 1500m; C. lemberti
occurs generally above 2000m in the Cascade, Sierra Nevada, Siskiyou,
and Warner Mountains; C. dumetorum is strictly coastal and associated
with Eriogonum latifolium from northern Monterey County to Sonoma
County (G. Gorelick, personal communication). Although C. comstocki
is at least partially bivoltine, the other species are univoltine. Larval
ground color is usually green, pale pink, or yellow; dorsal and lateral
lines and dorsolateral chevrons may be present or absent.
Morphological differences among Incisalia larvae are too small to aid
greatly in identification. The larval morphology of I. eryphon is most
divergent, as indicated in the species key. But the larvae of all the
species are best identified according to host and locality. As with some
members of Callophrys , the Incisalia larval head is yellowish-brown
with dark pigment confined to a narrow arc linking ocelli 1-5. Larvae of
I. augustus , the most widespread species, can be found in most areas
except the deserts on several hosts, especially A denostoma, Heteromeles,
Prunus (all Rosaceae), Ceanothus, Rhamnus (both Rhamnaceae), and
Cuscuta (Convolvulaceae); Powell (1968) also confirms that it uses
Arbutus menziesii Pursh. (Ericaceae) and Chlorogalum pomeridianum
(D. C.) Kunth. (Liliaceae). Larvae of I. eryphon occur on Pinus (Pinaceae)
usually above 2000m from the San Bernardino Mountains northward
through the Cascade, Sierra Nevada and Siskiyou Mountains. Larvae of
I. fotis feed on Cowania mexicana var. stansburiana (Rosaceae) in
mountains of the Mojave Desert. Larvae of I. mossii feed only on
Crassulaceae; primarily they utilize Sedum but some populations also
use Dudley a (J. F. Emmel, personal communication); this species occurs
in isolated cismontane colonies from the San Bernardino Mountains
northward. Incisalia augustus is partially bivoltine, especially in the
south, but the other species of Incisalia are univoltine.
Coloration can be useful in identifying live larvae of some Incisalia
species. Larvae of I. augustus (fig. 74-4d) and/, fotis are polymorphic and
often resemble members of the nominate subgenus. Their ground color
is usually green and they frequently have white or red and white
dorsolateral chevrons and lateral lines; and they often have at least a
27(1): 1-81, 1988
41
trace of a reddish lateral bar on A1 (fig. 74-4d). Larvae of I. mossii are
red or greenish-yellow and may have whitish lateral chevrons. Larvae
of I. eryphon are monomorphic with a green ground color and paired
yellowish white dorsal and lateral lines (fig. 74-5a).
The Mitoura species can be divided into two groups based on biology
and larval morphology (as described above). Those which feed on cedar,
cypress, and juniper (Cupressaceae) have 5-7 larval instars, whereas
those which feed on dwarf pine mistletoe (Viscaceae) have five larval
instars. The Cupressaceae-feeders have well developed dorsal pro-
minences on T1 anterior and (especially) posterior to the prothoracic
shield but segments T2-A6 are dorsally rounded, lacking prominences.
These larvae are dark green with white dorsal and lateral lines which
are weakened or broken intersegmentally, as illustrated for C. (M.)
nelsoni (fig. 74-5b); they are densely covered with erect straight setae ca
twice as long as the spiracle diameter; dendritic setae are absent. The
Viscaceae-feeders are more angulate with paired dorsal prominences at
least on T2 and A6 and weak lateral prominences along the lateral fold
(best developed on A7 and A8); but dorsal prominences on T1 are poorly
developed or absent (fig. 73). All dorsal setae (except those surmounding
dorsal prominences) are shorter than the sensory setae and at most
subequal in length to the spiracle width. Prominent setae subequal in
length to the sensory setae are present on T1 posterolateral to the
prothoracic shield in C. (M.) johnsoni but not in C. (M.) spinetorum. The
ground color of these larvae is yellow to olive-brown, while the dorsal
prominences are usually brighter yellow, bordered laterally by white
and dark brown, and often reddish apically (fig. 74-5c). A transverse bar
may be apparent as a darkening of the dorsal prominences on Al. These
larvae appear to glisten due to a shinier body surface and shorter,
sparser setae than in the Cupressaceae-feeders; their setae are often
reclinate toward the apices of the dorsal prominences (fig. 4); a few
inconspicuous dendritic setae occur at the margin of the honey gland.
The California Mitoura species which feed on Cupressaceae comprise
a portion of a complex of several often narrowly allopatric sibling
species and/or subspecies occurring throughout most of North America.
These may be poorly distinguishable where their ranges converge and
are perhaps best considered ecotypic components of a superspecies. Yet
in spite of a few areas of possible intergradation (Shields, 1985; Scott,
1986), the California taxa are relatively uniform throughout their
ranges which may be parallel and narrowly separate (by altitude and
habitat) over long distances. Thus, M. nelsoni occurs in association with
Libocedrus decurrens in montane habitats from San Diego County to
Oregon, while M. loki and M. siva often occur at different elevations and
in association with other hosts in the same mountains. The larvae of
these species are best distinguished according to host and locality.
Although larvae of most (probably all) of the Cupressaceae feeders
can be reared on many plants in that family, most populations are
42
J.Res.Lepid.
associated with one host species in nature (Johnson, 1978). Four
subspecies of M. siva occur in California; M. s. siva is associated with
Juniperus osteosperma in the mountains of the eastern Mojave Desert;
C. ( M .) s. juniperaria (J. A. Comstock) is associated with J. californica
from the lower northwest slopes of the San Bernardino Mountains
westward along the northern edge of the San Gabriel Mountains (where
it is also associated with J. osteosperma) to the eastern edge of the
Tehachapi Mountains; C. (M.) s. mansfieldi (Tilden) is associated with
J. californica in the inner coast ranges from Ventura County to San
Benito County; C. (M.) s. chalcosiva Clench is associated with J.
occidentalis in the Inyo and White Mountains. The status of brown M.
siva (or M. nelsoni) populations associated with J. occidentalis in the
San Bernardino Mountains, Sierra Nevada, and Modoc County is
uncertain; these may be conspecific with C. (M.) barryi Johnson described
from eastern Oregon. There are two subspecies of Mitoura nelsoni ; the
nominate one ranges from the mountains of San Diego County northward
in association with Libocedrus decurrens; C. ( M .) n. muiri (Hy. Edwards)
is associated with Cupressus sargentii and (rarely) J. californica (J.
Lane, personal communication) in coastal mountains from San Luis
Obispo County to Mendocino County. Mitoura loki occurs with J.
californica from the eastern San Bernardino Mountains southward to
Baja Californica. Mitoura thornei is known only from Otay Mountain in
San Diego County in association with Cupressus forbesii. Mitoura
nelsoni is univoltine, but at least some populations of the other species
are partially hi- or trivoltine.
Habrodais
One species of Habrodais , H. grunus , occurs in montane habitats
throughout California except in the deserts. The larval hosts are
Quercus chrysolepis and, according to Pyle (1981), Q. vaccinifolia Kell.,
Chrysolepis chrysophylla (Dough) A. DC., and Lithocarpus densiflora
(H. & A.) Rehd., all in the Fagaceae. Habrodais grunus is the only
California member of the tribe Theclini and differs considerably from
the other theclines. The most obvious differences are the lack of a honey
gland, presence of a lateroseries of crochets (fig. 72a), and an evenly
convex posterior margin of the prothoracic shield (fig. 46). It also lacks
dendritic setae and the sensory setae are finely tapered with minute
lateral spicules confined to the apex. The ground color is pale blue-green
(including the prothoracic shield) and a pair of pale yellow subdorsal
lines may be present (fig. 74-3d). A pair of prominent dorsal setae occurs
on segments T2-A8, while other dorsal setae are bent parallel to and
flattened in the body plane. This species is univoltine with egg diapause.
Harkenclenus
The single member of this genus, H. titus , ranges from coast to coast but
in California it is confined to the northeastern corner southward in the
27(1): 1-81, 1988
43
eastern Sierra Nevada to near Lake Tahoe. Larvae feed on Prunus
uirginiana and are distinctively marked reddish dorsally on T2, T3, and
A6 and dorsally and laterally on A7-A10; the remainder of the body is
green (fig. 74-5d). The head is dark brown in a band across the frons and
posteriorly to the ocelli, but lighter dorsally (fig. 47). Aside from some
members of Satyrium , this is the only California thecline with dendritic
setae present beyond the margin of the honey gland on A7. Dendritic
setae occur laterally near the spiracles or in that latitude on Tl-Al, A7,
and AS, with the greatest numbers (ca 20) on T3 and Al. All setae are
orange-brown, erect, and straight. Prominent setae on T2-A6 are
indistinct, but the longest setae (dorsally and along the lateral fold) are
ca 2.5X as long as the spiracle diameter. This species is univoltine with
egg diapause.
Minis trymoii
One member of this genus, M. leda , inhabits the southern California
deserts. It is multivoltine and larvae feed primarily on Prosopis but we
have one record on Acacia greggii (both Fabaceae). They are distinc-
tively marked green and white (fig. 74-6a) and segments T2-A6 have
dorsal prominences which confer a saw-toothed dorsal profile (fig. 56c).
Each dorsal prominence is surmounted by 1-4 prominent setae ca 3-4X
as long as the spiracle diameter; most other dorsal setae are 1-2X as long
as the spiracle diameter.
Satyrium
Seven members of this genus occur in California; all are univoltine with
egg diapause. Although morphologically more diverse than other Cali-
fornia thecline genera, the Satyrium species are united by the presence
of at least 20 dendritic setae on A7, especially along the margin of the
honey gland; four species also have dendritic setae on other segments.
Among other California theclines only if. titus shares such an abundance
of dendritic setae. Distinct dorsal prominences may be present, but more
often there are weakly developed dorsal ridges extending from T2-A6, so
that in cross-section the body appears trapezoidal with the dorsal area
flat or somewhat concave and the lateral areas sloping outward to the
lateral folds. Prominent setae of variable number and degree of dis-
tinctness occur along the dorsal ridges and the lateral fold. Other setae
may be recumbent to erect and nonpigmented to dark brown. Cephalic
pigmentation varies from yellow to dark brown.
Satyrium auretorum occurs throughout cismontane California in
chaparral and scrub oak woodland. Larvae feed on various oaks,
especially the scrub oaks, Quercus Cornelius -mulleri, Q. dumosa, and Q.
wislizenii. They are green with a yellow lateral line of variable intensity
and often ventrally bordered with pink (best developed on T2, T3, A7
and A8) (fig. 74-6b). The head is dark brown antero ventrally (across the
frons and extending posterolaterally beyond the ocelli), but yellowish
44
J. Res. Lepid.
brown apically (fig. 51). The sensory setae are tapered, ca 5X as long as
other setae on the shield, and 1.5X as long as the longest dorsal setae
posterolaterally adjacent to the shield. Prominent dorsal and lateral
setae, ca 2X as long as the spiracle width, occur in groups of 2-10 on T2-
A6 (fig. 56b). Nonprominent dorsal and lateral setae on T2-A6 are
weakly spindle-shaped, suberect, straight to moderately bent, and ca 1-
1.5X as long as the spiracles; on A7 they are shorter and more strongly
bent (often parallel to the body). These setae are reddish brown,
minutely dentate, and tapered. Dendritic setae are confined to A7.
Satyrium behrii ranges from the Little San Bernardino Mountains
west to the Mount Pinos area, northward along the east slope of the
Sierra Nevada to Oregon, and in the Panamint and White Mountains.
The larvae feed on Purshia glandulosa and P. tridentata. The ground
color is dark green with white or yellow middorsal and lateral lines and
dorsolateral chevrons on T2-A6 (fig. 74-6c). The head is mostly yello-
wish brown, but dark pigment extends across the frons and postero-
laterally beyond the ocelli (fig. 52). A pair of prominent dorsal setae ca 3-
4X as long as the spiracle width occurs on T2; lateral prominent setae
are present on A7 and A8 but usually absent elsewhere. Most other
dorsal and lateral setae are 1-2X as long as the spiracle width, bent
parallel to and flattened in the plane of the body surface, and acutely
tapered. Dendritic setae occur in groups of 4-6 subdorsally on T2 and T3
as well as around the honey gland.
Satyrium californica occurs throughout the state except in the eastern
deserts. Larvae have been found on Ceanothus and Quercus, but
additional hosts are likely since the butterfly sometimes occurs in the
absence of those plant genera. The ground color is chocolate brown
dorsally and white ventrally (fig. 74-6d); the head (fig. 53), legs,
chalazae, and most setae are dark brown. Conspicuous dendritic setae
ca 1-3X as long as the spiracle width are present on T1-A7. While dorsal
and lateral prominent setae on T2-A6 are ca 5-7X as long as the spiracle
width, other dorsal and lateral setae on those segments are sparse and
1/4- IX as long as the spiracle width. Since larvae are usually found in
early morning they may be nocturnal, as with S. edwardsii (Grote and
Robinson) in the eastern United States (Webster & Nielson, 1984).
The larva of S. fuliginosum superficially resembles that of the
polyommatine I. icarioides in habits and appearance more than the
larvae of its congeners. The ground color is light green with whitish
lateral chevrons (fig. 74-7a), the head is dark brown (fig. 48), and the
body dorsum is evenly convex. Dorsal prominent setae (ca 2-5X as long
as the spiracles) are present on T2 but absent or obscure posteriorly;
lateral prominent setae are more abundant and conspicuous on T1-T3
and A6-A10 but may be absent on the intervening segments. Dendritic
setae ca 2X as long as the spiracles are present posterolateral to the
prothoracic shield, near the spiracles (or in the same latitude) on T3-A6,
and around the honey gland. The remaining dorsal and lateral setae are
27(1): 1-81, 1988
45
numerous, ca .5-IX as long as the spiracles, erect, tapered, and light
brown. The larvae feed on Lupinus and are strongly myrmecophilous;
they probably feed nocturnally, but during the day they can be found at
the base of host plants and under nearby rocks. This species occurs along
the eastern slopes of the Sierra Nevada northward through the Cascade,
Siskiyou, and Warner Mountains.
The larva of S. saepium differs from its congeners (except S. auretorum
and S. sylvinus ) in having dendritic setae confined to the lateral
margins of the honey gland. These setae are ca as long as the spiracles,
clavate, and have very short lateral spicules. The following combination
of characters distinguish this species from all others: the sensory setae
are slightly spatulate in the apical fourth, ca 4-5X as long as the
spiracles and nearly all other dorsal and dorsolateral setae on T2-A6;
prominent dorsal setae are absent; the dorsal and lateral setae are of
two forms intermixed, one of which is brownish, suberect, and cylindric
and the other is nonpigmented, coarsely dentate, spindle-shaped, basally
bent nearly parallel to the body surface, and compressed in that plane
(as in fig. 18). Dendritic setae occur only on A7. The ground color is dull,
dark green; a yellow lateral line extends from T2 to A 10 and is most
prominent on A8. Cephalic infuscation is limited to a diffuse band across
the ocellar region (fig. 49). Larvae feed on Ceanothus and occur through-
out the state except in the eastern deserts.
Satyrium sylvinus is also widespread in California, but absent from
the Colorado and Mojave Deserts. Larvae feed on Salix (Salicaceae).
They are light green with distinct white subdorsal and lateral lines and
somewhat less distinct lateral chevrons (fig. 74-7b). The head is yello-
wish with dark pigment confined to a narrow band connecting ocelli 1-5
(fig. 50). The sensory setae are finely tapered, ca 3-5X as long as the
spiracles and other setae on th shield, and subequal to the longest dorsal
prominent setae posterolateral to the shield and on T2. Dorsal prominent
setae on T3-A6 are ca 2X as long as the spiracles and other dorsal and
lateral setae on those segments. All setae are tapered, suberect, and
nonpigmented. Nonprominent dorsal and lateral setae on T2-A6 are
erect to strongly bent; those on A7 are often bent parallel to the body.
Dendritic setae are confined to A7. The body is somewhat angulate in
cross-section due to a rather flat or slightly concave dorsum (less
noticeable in distended specimens).
Satyrium tetra has about the same distribution as S. saepium , being
found in chaparral habitats containing its host, Cercocarpus (Rosaceae).
Larvae resemble those of S. sylvinus in coloration, but have less well
developed dorsal and lateral lines and more prominent lateral chevrons
(fig. 74-7c). A slightly broader band of dark pigment surrounds the ocelli
(fig. 54). They are also distinctive in being covered with erect and
relatively short orange-brown setae. The sensory setae are slightly
spatulate, as in S. saepium , and subequal to the longest dorsal setae
posterolaterally adjacent to the shield and on T2. Dorsal prominent
46
J. Res. Lepid.
setae occur in groups of ca 20 on T2-A6 and are ca 2X as long as other
dorsal and lateral setae on those segments (fig. 56a). Clavate dendritic
setae occur on the prothoracic shield near the sensory setae and on A7.
All other dorsal and lateral setae are tapered. The body is rather flat
dorsally between the rows of dorsal prominent setae resulting in an
angulate cross-section even more pronounced than in S. sylvinus.
Strymon
Larvae of the three Strymon species in California are very similar to
each other and to larvae of C. ( Callophrys) and C. (Incisalia). They are
covered with nonpigmented (except apically darkened), erect, straight,
tapered setae; prominent setae on T2-A6 are absent or obscure (not
much longer than surrounding setae) and dendritic setae are incon-
spicuous and confined to the margin of the honey gland. They differ from
larvae of Callophrys (Callophrys) and C. (Incisalia) by their more
filiform and less prominently spiculate sensory setae, smaller head, and
by slight differences in cephalic pigmentation. Dark infuscation on the
head is confined to an arc connecting ocelli 1-5 and extending posteriorly
to ocellus 6 (figs. 65-67), whereas in C. (Callophrys) and C. (Incisalia)
the cephalic infuscation is usually either more extensive (broadly
encompassing all ocelli) or limited to ocelli 1-5. The ratio of the head
width to the distance between insertions of the sensory setae on the
prothoracic shield is usually at least 2.2 in both S. avalona and S.
melinus , about 2.0 in S. columella , and usually less than 2.0 in C.
(Callophrys) and C. (Incisalia).
Strymon melinus occurs throughout the state and probably has the
widest host range of any North American lycaenid. In California its
larvae are most often found on Eriogonum (Polygonaceae) and various
members of the Fabaceae and Malvaceae. Strymon avalona occurs only
on Santa Catalina Island and utilizes Eriogonum (Polygonaceae) and
Lotus (Fabaceae) (Gorelick, 1987). Strymon columella , largely sub-
tropical in distribution, occurs in southern California and utilizes
Hibiscus, Sphaeralcea , and other Malvaceae as larval hosts. The larvae
of S. columella can be distinguished from those of S. melinus by the
milky color of their chalazae; this character is best seen in live larvae.
Other differences include a narrower host range and smaller distri-
bution. The limited distribution of S. avalona may be the best clue to
distinguish it from S. melinus ; also, its head is slightly browner and the
ocellar infuscation is a little darker and more extensive (fig. 65), but
otherwise they are virtually indistinguishable.
Polyommatinae
All California members of this subfamily belong to the tribe Poiyom-
matini. The head is about 1/4 as broad as the body and almost always
black (but lighter brown in two local species and nonpigmented in many
27(1): 1-81, 1988
47
exotic species); most species have a honey gland and eversible tubercles;
the prothoracic shield is nonsclerotized, nonpigmented (although there
may be dark chalazae), and often has a smoothly convex anterior
margin; the bases of the sensory setae may appear sunken below the
cuticular surface due to the height and density of cuticular ridges. The
chalazae frequently appear stellate due to distolateral points. Prominent
setae (at least subequal to the length of sensory setae) commonly occur
on the prothoracic shield. The majority of species in this subfamily are
univoltine and active in spring or summer, but some species and/or
subspecies appear in late summer or fall; a few species are facultatively
bi- or trivoltine while others are multi voltine.
Agriades
One species of this genus, A. franklinii, occurs in California usually
above 3000m in the Cascade and Sierra Nevada Mts. but as low as
2000m in the Siskiyou Mts.; it is associated with wet meadows and
boggy stream and lake margins. This species is partially bi voltine with
diapause in the second instar. Eggs and larvae were found on Dode-
catheon alpinum (Gray) Greene (Primulaceae) near Sonora Pass.
The larvae mine the leaves until the last instar. This is the only
polyommatine in California which lacks both a honey gland and
eversible tubercles. Other distinguishing features include the dark
brown color of setae (even ventrally), spiracles, and legs. Distinct dorsal
and lateral prominent setae are present, and all setae are erect and
straight to slightly curved; chalazae are distinctly stellate. The ground
color is deep, bright green and there is a red middorsal line (fig. 74-7d).
Brephidium
One member of the genus Brephidium, B. exilis , is widespread in
California. It is multi voltine and occurs commonly in relatively xeric
and saline habitats where its major hosts, Atriplex, Chenopodium , and
Salsola (Chenopodiaceae) thrive; Sesuvium verrucosum (Aizoaceae) is
also used in some areas (Johnson, 1981). Larvae are various shades of
green and usually without distinct markings (fig. 74-8a); often the body
appears finely granular or pollinose. Prominent setae (2-3X as long as
the spiracles) occur only along the anterior margin of Tl, laterally on
T2, and along the posterior margin of A9-10. All other dorsal and lateral
setae are clavate-capitate, often bent parallel to the body surface, and ca
as long as the spiracle width. All setae are nonpigmented and chalazae
are weakly stellate. The sensory setae are finely tapered and at least
twice as long as all other dorsal setae on T2-A8. Dendritic setae are
absent.
Celastrina
One species, C. argiolus , occurs throughout California in many
habitats but not in the desert lowlands. In California the larvae utilize
48
J.Res.Lepid.
primarily Ceanothus (Rhamnaceae) and many hosts in the Rosaceae;
they are rarely found on Lotus (Fabaceae) (Gorelick, 1987). In Arizona
they also utilize Amorpha (Fabaceae) (Noel McFarland, personal com-
munication) and Eriogonum (Polygonaceae). Larvae prefer to feed on
flowers, buds, and immature fruit. Adults fly in spring and early
summer throughout most of California and are partially bivoltine (at
least in southern California) with pupal diapause. Most setae are bent
parallel to the body surface and arise from strongly stellate chalazae
(fig. 2f). The chalazae are crowded and their lateral points are often so
long (the span between opposite points may be nearly as great as the
setal length) that they may interdigitate; only L. marina , among other
California species, approaches this condition. The sensory setae on the
prothoracic shield are slightly expanded and flattened in the apical
third (ca 2X as broad as the basal width) and are ca 3-4X as long as other
setae on the shield; their length is subequal to a pair of prominent dorsal
setae on T2 and ca 2-3X as long as all other dorsal setae on T2-A6. A few
dendritic setae occur at the lateral margins of the honey gland and
lateral to the sensory setae on the prothoracic shield. Larvae are
polymorphic in coloration. The ground color is often whitish, pale pink
or pale green; distinct lateral lines and chevrons are lacking but a
conspicuously dark green, pink, or brownish transverse bar usually
occurs on Al. A color morph common for larvae found on Adenostoma
fasciculatum is illustrated in fig. 74-8b.
Euphilotes
The genus Euphilotes is extremely complex with four species and
numerous subspecies in California. One member, E. mojave , is often
considered a subspecies of E. enoptes (Pratt and Ballmer, 1987). All
Euphilotes larvae feed on buds, blossoms, and seeds of Eriogonum.
Several hosts may be used by one species, but most local populations use
a single host and only rarely do sympatric species (and subspecies) share
a host. All members of this genus diapause as pupae and are typically
univoltine; some populations of E. enoptes are facultatively bi- or
trivoltine (Pratt and Ballmer, 1987). Published reports of five larval
instars in E. enoptes bayensis (Langston) and E. e. smithi (Mattoni) by
Langston and Comstock (1966) and Arnold (1983), respectively, are
probably erroneous since we have found only four instars in hundreds of
rearings representing all four Euphilotes species. Larval ground color
may be white, pink, yellow, or brownish; color pattern ranges from non-
patterned to strongly marked with white, pink, yellow, and/or brown
middorsal and lateral lines and lateral chevrons (fig. 74-8c). Distingui-
shing morphological features of this genus include apically spatulate
sensory setae (fig. 13), a few dendritic setae at the lateral margins of the
honey gland and usually near the Al spiracles, moderately to weakly
stellate chalazae, and a variable number of prominent setae on T2-A6
(sometimes absent).
27(1): 1-81, 1988
49
Larvae oiE. rita are perhaps the most distinctive of the genus. Paired
dorsal prominences on T3-A6 are steeply peaked, creating a saw-
toothed lateral profile, and each usually has at least one prominent seta,
which may be directed posteromedially (fig. 57c). More prominent setae
occur laterally on T2-A10 and dorsolaterally on T2 (and occasionally on
other segments). The ground color is white or pink; markings may be
absent but usually there are reddish lateral chevrons on T2-A6 and a
transverse bar on A1 (fig. 74-8d). The head is dark brown and the legs
are a little lighter. Most nonprominent setae are strongly curved and
may be bent parallel to the body. The eversible tubercles arise from
distinct but low prominences and are everted frequently as the larva
crawls; this may be related to the fact that this is the most strongly
myrmecophilic member of the genus. This species occurs along the
desert slopes forming the southern and western borders of the Mojave
Desert, the east slope of the Sierra Nevada and in some of the desert
mountains. Host plants include Eriogonum davidsonii, E. deflexion, E.
heermannii, E . kearneyi,E. microthecum, E . plumatella, E . roseum , and
E. wrightii. Flight activity ranges from May to September for various
populations.
Larvae of E. mojave are similar to those of E. rita in distribution of
prominent setae. But segments T3-A6 are more rounded dor sally (fig.
57b); the eversible tubercles do not arise from dorsolateral prominences;
and the legs, although brown, are much lighter than the head. Popula-
tions ofE. mojave occur scattered through the Mojave Desert and desert
slopes bordering it. Larvae can be found in spring on the annuals E.
pusillum and E. reniforme.
Larvae of E. battoides andE. enoptes are best distinguished according
to host plant and locality. In both species dorsal prominences are not
apparent and dorsal prominent setae are usually absent posterior to T2
(fig. 57a); the legs are nonpigmented; the number of prominent setae in
specific locations differs for different populations; and nonprominent
setae are generally short and bent parallel to the body surface. Euphi -
lotes battoides utilizes E. fasciculatum (everywhere), E. parvifolium
(along the south coast), E. heermannii and E. microthecum (in the
eastern Mojave Desert), and E. umbellatum and various cespitose
Eriogonum species in the Cascade, Sierra Nevada, Siskiyou, and White
Mountains. Euphilotes enoptes utilizes E. nudum everywhere north and
west of the San Bernardino Mts., E. elongatum, E. davidsonii , and E.
wrightii everywhere south and east of the San Gabriel Mts., E. latifo -
Hum and E. parvifolium along the central coast, and E. elatum and E.
umbellatum in the Cascade, Sierra Nevada, and Siskiyou Mountains.
Various populations of both species fly in spring, summer, or fall.
Everes
Two species of Everes occur in California; E. amyntula is widespread
from sea level to over 3000m throughout the state (except low elevations
50
J. Res. Lepid.
of the deserts), while E. corny ntas occurs in mesic habitats generally
below 1000m from the southern San Joaquin Valley northward. One
distinctive feature easily separates this genus from all others in
California; the spatulate lobes of the prolegs have sharply defined
(somewhat scerotized and pigmented) lateral margins, especially basally
(fig. 72b). This trait also occurs in E. argiades from Japan. The ground
color is green, grey, or pinkish grey; a cream lateral line bordered with
pink may be present. Both species are at least facultatively multivoltine
with diapause in the last instar.
In California these species usually can be distinguished by setation
differences, but some populations in the north and along the Sierra
Nevada are intermediate. In larvae of E. amyntula dendritic setae are
few in number and confined to the margin of the honey gland; other
dorsal and lateral setae are erect and straight to slightly curved (rarely
bent parallel to the body surface). In E. corny ntas we found a few (ca 4)
dendritic setae around the honey gland, and others near the A1
spiracles and occasionally laterally on T3; but Lawrence and Downey
(1966) illustrate (in Illinois larvae) ten dendritic setae near the honey
gland and report that others may occur near the A2 spiracles. Many
dorsal and lateral setae in E. comyntas are curved or bent (often parallel
to the body surface), especially on A7 and A8. In some (especially
southern) populations of E. amyntula the eversible tubercles appear to
be noneversible, although their location is marked by the usual wrinkled
depression encircled by setae. This may be an adaptation to their habit
of feeding only inside Astragalus seed pods where it is less likely that
they would encounter ants. The larvae of E. comyntas and northern
California populations of E. amyntula which commonly feed externally
on various herbaceous Fabaceae have fully functional eversible
tubercles.
Glaucopsyche
There are two species of Glaucopsyche in California; their larvae are
often similar to those of Lycaeides and some Icaricia species. They are
densely covered with short, tapered, nonpigmented setae; the sensory
setae are finely tapered; chalazae are moderately stellate; and the
dorsal setae on A7 and A8 (between the spiracles) are tapered and
suberect to strongly recurved. Larvae of G. piasus have dendritic setae
laterally on T3-A3, A6, and A7; there are 6-8 prominent dorsal setae
anteriorly on T2 which are 2-3X as long as the spiracles and slightly
longer than the sensory setae; all other dorsal and lateral setae on Tl-
A6 are erect, tapered, and .5-.75X as long as the spiracles. Larvae of G.
lygdamus have dendritic setae around the A1 spiracles (also occasion-
ally around the A2 spiracles and laterally on T3) and (less conspicuously)
at the lateral margins of the honey gland. They also have 1-3 pairs of
dorsal prominent setae on T2-A6 which are .5- IX as long as the sensory
setae and 3-5X as long as the spiracles and other dorsal and lateral
27(1): 1-81, 1988
51
setae. In cismontane southern California populations of G. lygdamus,
nonprominent dorsal and lateral setae on T2-A6 are suberect to strongly
curved (often bent parallel to the body); but in populations from
northern California and east of the Sierra Nevada from the central
Mojave Desert northward these setae are more erect (never bent
parallel to the body). Larval coloration is polymorphic for G. lygdamus ,
ranging from concolorous green to pink and yellow with strong chevron
markins and a well defined dorsal line (fig. 74-9a). Larvae of G. piasus
are less polymorphic with a dull green or gray ground color and lateral
chevrons (fig. 74-9b). Larvae of G. piasus feed only on lupine, while
larvae of G. lygdamus utilize Astragalus , Lotus, Lupinus and Vicia.
Both species are univoltine, have pupal diapause, and fly in spring or
early summer.
Hemiargus
The two California species of Hemiargus are multivoltine and gener-
ally restricted to the southern and eastern regions. Their larvae greatly
resemble those of the Icaricia acmon species group with which they
share the following characters: finely tapered sensory setae, at least
four prominent dorsal setae on T2 and at least two each on T3-A6, few
dendritic setae at the lateral margins of the honey gland and occasionally
near the A1 spiracles, and the majority of dorsal and lateral setae
suberect to bent parallel to the body surface. However, the most strongly
bent setae (near the abdominal spiracles and dorsally on A7 and A8) are
somewhat spindle-shaped, flattened, and acutely pointed (fig. 21),
whereas in the /. acmon species group, setae in the same areas are
cylindric and apically blunt or truncate (fig. 2m).
Slight differences in setation distinguish these species. Larvae of H.
ceraunus sometimes have a few dendritic setae near the A1 spiracles;
also, the longest dorsal setae on T2 are no more than .75X as long as the
longest setae on the prothoracic shield; and segments T3-A6 have one
(or no) pair of dorsal prominent setae (fig. 58a). Larvae of H. isola lack
dendritic setae near the A1 spiracles; the longest dorsal setae on T2 are
at least as long as the longest setae on the prothoracic shield; and there
are usually at least two pairs of prominent setae on T3-A6 (fig. 58b).
Larvae of both species feed on members of the Fabaceae but larvae of if.
ceraunus also utilize Eriogonum (Polygonaceae). The ground color may
be green, red, brownish, or yellow; markings may be absent but often
there is a reddish middorsal line and red or yellow lateral lines and
dorsolateral chevrons (fig. 74-9c).
Icaricia
There are five species of Icaricia in California. They all have a few
dendritic setae near the A1 spiracles and lateral margins of the honey
gland, moderately stellate chalazae, and flagelliform sensory setae. The
number and relative size of prominent setae on the prothoracic shield,
52
J.Res.Lepid.
dorsally on T2-A6, laterally on all segments, and subdorsally on T2-A7
differs in each species. In all but I. icarioides the prominent dorsal setae
are somewhat curved or inclined posteriorly. Other dorsal and lateral
setae are tapered to blunt, erect to recurved and may be bent parallel to
the body surface. Nondendritic dorsal setae on A7 and A8 are usually
somewhat clavate and may be bent parallel to the body surface. Icaricia
acmon is multivoltine and I. neurona is partially bi- or trivoltine; the
other Icaricia species are univoltine. Diapause occurs in the egg for I.
shasta , but in the second instar for the other species.
Three species (I. acmon , I. lupini, and I. neurona ) constitute the
I. acmon species group. Morphological differences in larvae of these
species are slight; they are best distinguished according to host plant,
locality, and season. The following character discussion is based on
populations in the San Bernardino Mountains and does not necessarily
apply to populations elsewhere. The length and abundance of prominent
setae are generally greatest in I. neurona and least in I. lupini , thus
affording some utility in species identification. Another distinguishing
feature concerns the subdorsal prominent setae on A7 (ca midway
between the honey gland and spiracles); in I. neurona these are well
developed, while in I. lupini they are usually absent, and in I. acmon
they are usually present but weakly differentiated from surrounding
setae. Dorsal setae on A7 and A8 are semi-erect to strongly bent, (often
parallel to the body) weakly tapered to clavate, and mostly apically
blunt. Larval ground color is dark green in I. lupini , gray-green to dull
pinkish gray in I. neurona , and highly variable (including green, cream,
and maroon) in I. acmon. All three species may have a white or yellow
lateral line, which in I. acmon may be bordered with red. The latter
species may also have a contrastingly colored middorsal line and lateral
chevrons.
Icaricia acmon , the most common and widespread species, occurs from
early spring to fall in all habitats except the open desert; larvae feed on
several species of Eriogonum and Lotus. Icaricia lupini is also wide-
spread and utilizes Eriogonum as a larval host but is restricted to
montane and foothill habitats; in southern California its host is Erio-
gonum fasciculatum , but elsewhere E. umbellatum is the major host.
Icaricia neurona is restricted to montane habitats (usually above
2000m) from the southern Sierra Nevada to the San Bernardino
Mountains; hosts are various cespitose Eriogonum species, especially E.
kennedyi and E. wrightii.
Larvae of I. shasta are similar in appearance to larvae of the previous
three species, but differ primarily in the greater development of
prominent setae. They differ from all other Icaricia in having dark legs
and dark chalazae on the prothoracic shield. The ground color is brown
to maroon and there are yellowish dorsolateral chevrons and lateral
lines (fig. 74-9d); the dorsolateral chevrons may be so enlarged that the
ground color is reduced to a narrow line middor sally. They feed on
27(1): 1-81, 1988
53
several cespitose Astragalus and Lupinus species mostly above 3000m
in the Cascade, Sierra Nevada, Warner, and White Mountains.
Icaricia icarioides is a widespread species with various populations
occupying habitats from sea level to over 3000m. Larvae feed on
perennial lupines and range in ground color from green to pinkish grey.
Larval setation differs substantially from that of the other Icaricia
species; nearly all dorsal and lateral setae on T2-A6 are erect, acutely
tapered, straight, and no longer than the spiracle width. Sensory setae
are ca 4X as long as other setae on the prothoracic shield and slightly
longer than the prominent dorsal setae on T2. Dorsal prominent setae
on T3-A6 are no more than ca 1.5X as long as the spiracle width and may
be weakly differentiated from other dorsal setae on those segments.
Leptotes
Only one species, L. marina , occurs in California. It is multivoltine
and abundant throughout southern California, but less common in the
Central Valley and uncommon or absent in the central and northern
regions of the state. Larval hosts are primarily various Fabaceae and
Plumbago (Plumbaginaceae), a common ornamental which probably
accounts for the success of this species in urban areas; Adenostoma
fasciculatum (Rosaceae) is also rarely used. The ground color is variable,
ranging from pink to green and brownish violet. In general appearance
the larvae appear most similar to those of C. argiolus. They have
apically spatulate sensory setae and are covered with short erect setae
arising from strongly stellate chalazae (the lateral points of adjacent
chalazae may interdigitate) (fig. 2e); dorsal prominent setae occur on T2
and occasionally on other segments. Larvae of L. marina are uniquely
distinguishable (among California lycaenids) by the presence of
numerous broadly recurved and finely tapered setae (fig. 2b) dorsally
and along the lateral fold, intermixed with shorter erect or only slightly
bent setae. Similar recurved setae occur in L. cassius (Downey
and Allyn, 1979) from Florida and in Syntarucus plinius from the
Australasian region.
Lycaeides
Two species of Lycaeides occur in California. Lycaeides melissa ranges
throughout the state (except the Colorado and Mojave deserts) from
near sea level to over 3000m, while L. idas occurs primarily above
2000m in the Cascade, central and northern Sierra Nevada, Siskiyou,
and Warner Mountains. Larval ground color is green (fig. 74- 10a);
although some larvae have whitish middorsal and lateral lines and
dorsolateral chevrons, others are unmarked. Host plants are herbaceous
Fabaceae including Astragalus , Lotus , and Lupinus. Lowland popula-
tions of L. melissa are multivoltine while montane populations above
2000m of both species are probably univoltine; diapause occurs as ova.
54
J. Res. Lepid.
Physical distinctions between larvae of these species in California are
subtle and they are best distinguished by locality. The sensory setae are
tapered while dorsal and lateral prominent setae on T2-A6 are ca 3-4X
as long as the spiracle width. Other dorsal and dorsolateral setae on T2-
A6 are erect, ca as long as the spiracle width, tapered, and arise from
moderately stellate chalazae. Typically they have 10-40 dendritic setae
per segment near or in the latitude of the spiracles on T3-A2 and A6-A8;
fewer dendritic setae may also be present laterally on T1 and T2. Other
dorsal setae on A7 and A8 are erect and clavate-capitate, not strongly
curved or bent.
Lycaeides larvae from near Mono Lake and the Warner Mountains
differ from other populations examined. Their dendritic setae are
difficult to observe since they are much smaller and number 0-8 per
segment. Most nonprominent dorsal and subdorsal abdominal setae are
truncate and sometimes peg-like; some, especially near the honey
gland, may be sharply bent near the apex, resembling a railroad spike.
Philotes
There is one species of Philotes , P. sonorensis , which occurs through-
out cismontane southern California (mostly below 2000m) and north-
ward approximately to the latitude of San Francisco. This insect is
univoltine with pupal diapause and flies in late winter and spring;
larvae feed on Dudley a (Crassulaceae). Ground color ranges from pale
green to pink and there are no contrasting markings (fig. 74- 10b).
Prominent dorsal setae occur only on T2; other dorsal and lateral setae
on T2-A6 are erect, straight, weakly tapered to truncate, and ca .5-.7X
as long as the spiracle width. Dendritic setae at the margins of the honey
gland are clavate, longer than the spiracle width, and appear velvety
due to numerous unusually short spicules (fig. 17). Other dorsal setae on
A7 and A8 are clavate to strongly capitate (fig. 25) and mostly less than
half as long as the spiracle width. The spiracles are brown and the proleg
spatulate lobes are small and knob-like.
Philotiella
This genus is closely related to Euphilotes and contains a single
species, P. speciosa, which is nearly confined to the southeastern desert
areas of California. The rare subspecies, P. s. hohartorum (Shields),
occurs in the western foothills of the Sierra Nevada. This species in
univoltine with pupal diapause and flies in spring; larval hosts include
Eriogonum reniforme and Oxytheca , especially O. perfoliata (Polygona-
ceae). The larval ground color is green or yellowish and there may be
reddish middorsal and lateral lines and dorsolateral chevrons (fig. 74-
10c). This species was long included in the genus Philotes along with all
members of Euphilotes. The larvae of P. speciosa are most similar to
those of E. mojave in general appearance; they differ in having virtually
27(1): 1-81, 1988
55
nonstellate chalazae, lighter head pigmentation, and more erect setae.
They also differ from larvae of all other California polyommatines
except, A. franklinii , in lacking eversible tubercles.
Plebejus
There is one California species of Plehejus, P. saepiolus, which occurs
generally above 2000m from Riverside County northward. It is univol-
tine with diapause in the second instar; larvae feed on Trifolium
(Fabaceae). The larva is green and often has a white lateral line. In
many respects the larvae of this species are similar to those of Icaricia.
They differ in having dendritic setae on A 7 extending from the honey
gland laterally to the spiracles, whereas in Icaricia the dendritic setae
on A7 are confined to the margin of the honey gland. They also differ
from all Icaricia except, I. shasta , in having dark legs.
Plebulina
The single species of Plebulina , P. emigdionis, occurs only in a few
scattered colonies in and around the western Mojave Desert. It is
partially bi- or trivoltine and larvae feed on Atriplex canescens (Cheno-
podiaceae). This is the most distinctive California member of the
Polyommatinae in terms of biology and larval morphology. It is the only
one whose larvae lack a spatulate lobe on the prolegs, have more than
four instars (5-7), and are restricted to a single host plant species. Also,
the chalazae appear buttressed rather than stellate; but, unlike the
buttressed chalazae of thecline larvae, the lateral ridges do not appear
to be basally fused with the cuticle. Dendritic setae are few in number
and confined to the vicinity of the A1 spiracles and margin of the honey
gland; they and most other dorsal and lateral setae on T2-A6 are erect,
tapered, and ca half as long as the spiracle width. A pair of prominent
dorsal setae on T2 are ca 6X as long as the spiracle width. As with many
other polyommatine larvae, most dorsal setae on A7 and A8 are clavate
and strongly bent or recumbent. Lenticles are dark brown in contrast to
most chalazae which are nonpigmented. The sensory setae are filiform.
Larvae are grayish pink or green and lack contrasting markings (fig. 74-
lOd). They probably feed nocturnally; during the day they can be found
at the base of the host plant in the company of ants.
Acknowledgements. Several people have contributed toward the completion of
this work. Andrew C. Sanders of the Herbarium of the University of California
at Riverside identified the majority of larval hosts in Appendix 2. Some plant
identifications were also provided by Oscar F. Clarke, formerly of the UCR
Herbarium. John F. Emmel generously provided much information on butterfly
distributions and hosts and contributed some larval specimens. Glen A.
Gorelick also contributed some specimens and offered helpful observations
about Callophrys; John Lane and Kurt Johnson offered distributional data on
Mitoura species. Donald J. Harvey provided some helpful information about
56
J.Res.Lepid.
riodinine larval morphology. Trevor Lambkin of Brisbane, Australia provided
several exotic species from that region. We also wish to thank Robert K. Robbins
of the USNM for the loan of a Melanis pixe specimen. John D. Pinto, Vahram
Sevacherian and the California department of fish and game contributed
financial support for some SEM work. The finished manuscript benefited from
the review and comments of Lauren D. Anderson, John F. Emmel, and David M.
Wright.
27(1): 1-81, 1988
57
Literature Cited
ACKER Y, P. R., 1984. Systematic and faunistic studies on butterflies. Symp. Roy.
Ent. Soc. Lond. 11:9-21.
BETHUNE-BAKER, G. T., 1925. A revision of the Liphyrinae together with a
description of the structure of the puparium of Liphyra brassolis and of the
pupae of Aslauga uininga and A. lamborni (Lepidoptera). Trans. Ent. Soc.
Lond. 1924: 199-238, 25 pis.
BOURQUIN, F., 1953. Notas sobre la metamorphosis de Hamearis susanae Orfila,
1953 con oruga mirmecofila. Rev. Soc. Ento. Argen. 16:83-87.
BRUCH, C., 1926. Orugas mirmecofilas de Hamearis epulus signatus Stich. Rev.
Soc. Ento. Argen. 1:1-9.
CLARK, G. C. & C. G. C. DICKSON, 1956a. The honey gland and tubercles of the
Lycaenidae. Lep. News 10:37-43.
— — — , 1956b. Proposed classification of South African Lycaenidae from the
early stages. J. Ent. Soc. Sth. Africa 10:195-215
1971. Life Histories of the South African Lycaenid Butterflies . Purnell,
Capetown. 272pp.
CLENCH, H. K. 1961. Tribe Theclini in Ehrlich, P. R. and A. H. Ehrlich, How to
Know the Butterflies. W. C. Brown Co., Dubuque, pp. 209-210.
COMSTOCK, J. A., 1928. Studies in Pacific Coast Lepidoptera. Bull. So. Cal. Acad.
Sci. 27:80-89.
— - — -, 1930. Metamorphoses of five California diurnals (Lepidoptera). Bull. So.
Cal. Acad. Sci. 31:33-45.
COMSTOCK, J. A. & C. M. DAMMERS, 1932. Early stages of Melitaea leanira wrightii
and Calephelis nemesis. Bull. So. Cal. Acad. Sci. 31:9-15.
COTTRELL, c. B., 1984. Aphytophagy in butterflies: its relationship to myrme-
cophily. Zool. J. Lin. Soc. 80:1-57.
DEVRIES, P. J., 1984. Of crazy ants and Curetinae: are Curetis butterflies
tended by ants? Zool. J. Lin. Soc. 80:59-66.
DEVRIES, P. J., D. J. HARVEY & I. J. KITCHING, 1986. The ant associated organs on the
larva of the lycaenid butterfly Curetis regula Evans. J. Nat. Hist 20:621-633.
DOS PASSOS, C. F., 1936. The life history of Calephelis borealis (Lepidoptera). Can.
Ent. 68:166-170.
DOWNEY, J. C., 1987. Family Lycaenidae in Stehr, F. W., Immature Insects.
Kendall/Hunt Co., Dubuque, pp.443-446.
DOWNEY, J. c. & A. C. ALLYN, 1979. Morphology and biology of the immature stages
of Leptotes cassius theonus (Lucas) (Lepid.: Lycaenidae). Bull. Allyn Mus.
55:1-27.
~ — - — , 1984. Chaetotaxy of the first instar larva of Hemiargus ceraunus
antibubastus (Hbn.) (Lycaenidae). Bull. Allyn Mus. 90:1-4.
EHRLICH, P.R., 1958. The comparative morphology, phylogeny and higher classi-
fication of the butterflies. (Lepidoptera: Papilionoidea). Kans. Univ. Sci.
Bull. 39:305-370.
ELIOT, J. N., 1973. The higher classification of the Lycaenidae (Lepidoptera): a
tentative arrangement. Bull. Brit. Mus. (Nat. Hist.) Ent. 28:371-505.
EMMEL, T. c. & J. F. emmel, 1973. The butterflies of southern California. Natural
History Museum of Los Angeles County, Science Series 26:1-148.
FERRIS, C. D., 1974. Distribution of arctic-alpine Lycaena phlaeas L. (Lycaenidae)
in North America with designation of a new subspecies. Bull. Allyn Mus.
18:1-13.
58
J.Res.Lepid.
FRANZL, s., M. LOCKE, & P. huie, 1984. Lenticles: innervated secretory structures
that are expressed at every other moult. Tissue and Cell 16:251-268.
GORELICK, G. A., 1971. A biosystematic study of two species of Callophrys
( Callophrys) in California (Lycaenidae). J. Lep. Soc. 25 (supl. 2):1-41.
— — , 1987. Santa Catalina Island’s endemic Lepidoptera III. The
Avalon hairstreak, Strymon avalona (Lycaenidae): an ecological
study. Atala 14(1):1-12.
HARVEY, D. J., 1987. Family Riodinidae in Stehr, F. W., Immature Insects.
Kendall/Hunt Co., Dubuque, pp.446-447.
HENNING, S. F., 1983a. Biological groups within the Lycaenidae (Lepidoptera). J.
Ent. Soc. Sth. Afr. 46:65-85.
— , 1983b. Chemical communication between lycaenid larvae (Lepidoptera:
Lycaenidae) and ants (Hymenoptera: Formicidae). J. Ent. Soc. Sth. Afr.
46:341-366.
HINTON, H. E., 1946. On the homology and nomenclature of setae of lepidopterous
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy.
Ent. Soc. Lond. 97:1-37.
, 1951 . Myrmecophilous Lycaenidae and other Lepidoptera — a summary.
Trans. S. Lond. Ent. Nat. Hist. Soc. 1949-50:111-175.
JOHNSON, J. M., 1981. Unusual food plant. Utahensis 1:10.
JOHNSON, K., 1978. Specificity, geographic distributions, and foodplant diversity
in four Callophrys (Mitoura) (Lycaenidae). J. Lep. Soc. 32:3-19.
JOHNSON, K. & G. BALOGH, 1977. Studies in the Lycaeninae (Lycaenidae) 2.
taxonomy and evolution of the Nearctic Lycaena rubidus Complex, with
description of a new species. Bull, Allyn Mus. 43:1-62.
KENDALL, R. o., 1976. Larval foodplants and life history notes for some metal-
marks (Lepidoptera: Riodinidae) from Mexico and Texas. Bull Allyn Mus.
32:1-12.
KITCHING, R. L., 1983. Myrmecophilous organs of the larvae and pupae of the
lycaenid butterfly, Jalmenus evagoras (Donovan). J. Nat. Hist. 17:471-81.
— , and B. Luke, 1985. The myrmecophilous organs of the larvae of some
British Lycaenidae (Lepidoptera): a comparative study. J. Nat. Hist. 19: 259-
76.
LANGSTON, R. L. & J. A. COMSTOCK, 1966. Life history of Philotes enoptes bayensis
(Lepidoptera: Lycaenidae). Pan Pac. Ent. 42:102-108.
LAWRENCE, D. A. & J. C. DOWNEY, 1966. Morphology of the immature stages of
Everes comyntas Godart (Lycaenidae). J. Res. Lepid. 5:61-96.
MALICKY, H., 1969a. Versuch einer analyse der Okologischen Beziehungen
zwischen Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera). Tijd.
Ent. 112:213-98.
— , 1969b. Uebersicht ueber Praimaginal stadien, Bionomie and Okologie
der mitteleuropaischen Lycaenidae (Lepidoptera). Mitt. Ent. Gesell. Basel
2/3:25-91.
— , 1970. New aspects on the association between lycaenid larvae (Lycae-
nidae) and ants (Formicidae, Hymenoptera). J. Lep. Soc. 24:191-202.
MASCHWITZ, u., M. WUST & K. SCHURIAN, 1975. Blaulingsraupen als zuckerlie-
feranten fur Ameisen. Oecologia (Berlin) 18:17-21.
MCALPINE, w. S„ 1938. Life history of Calephelis muticum (McAlpine): Lepidop-
tera. Bull. Brooklyn Ent. Soc. 33:111-121.
MILLER, L. D. & F. M. BROWN, 1979. Studies in the Lycaeninae (Lycaenidae) 4. The
higher classification of the American coppers. Bull. Allyn Mus. 51:1-30.
27(1): 1-81, 1988
59
— , 1981. A catalogue/checklist of the butterflies of America north of
Mexico. Lepid. Soc. Memoir No. 2.
MUNZ, P. A. & D. D. KECK, 1959. A California Flora. University of California Press,
Berkeley and Los Angeles. 1681pp.
ORSAK, O. & D. W. WHITMAN, 1987. Chromatic polymorphism in Callophrys mossii
bayensis larvae (Lycaenidae): spectral characterization, short-term color
shifts, and natural morph frequencies. J. Res. Lepid. 25:188-201.
PETERSON, A., 1948. Larvae of Insects Part I Lepidoptera and Hymenoptera.
Edwards Bros., Inc. Ann Arbor, Michigan. 315pp.
PIERCE, N. E. & P. S. MEAD, 1981. Parasitoids as selective agents in the symbiosis
between lycaenid butterfly larvae and ants. Science 211:1185-1187.
PIERCE, N. E., 1983. The ecology and evolution of symbioses between lycaenid
butterflies and ants. Ph. D. dissertation, Harvard University, Cambridge,
Mass.
POWELL, J. A., 1968. Foodplants of Callophrys (Incisalia) iroides. J. Lep. Soc.
22:225-226.
PRATT, G. F. & G. R. BALLMER, 1986. Clarification of the larval host plant of
Epidemia mariposa (Lycaenidae) in northern California. J. Lep. Soc. 40:127.
— , 1987, The phene tics and comparative biology of Euphilotes enoptes
(Boisduval) (Lycaenidae) from the San Bernardino Mountains. J. Res. Lepid.
25:121-135.
PYLE, R. M., 1981. The Audubon Society Field Guide to North American Butter-
flies. Chanticleer Press, A. A. Knopf, New York. 916pp.
ROSS, G. N., 1964. Life history studies on Mexican butterflies II. Early stages of
Anatole rossi a new myrmecophilous metalmark. J. Res. Lepid. 3:81-94.
SCOTT, J. A., 1980. Geographic variation in Lycaena xanthoides. J. Res. Lep.
18:50-59.
— — — -, 1986. The butterflies of North America a natural history and field guide.
Stanford University Press, Stanford California. 583pp.
SHAPIRO, A. M., 1974. A salt-marsh population of Lycaena helloides (Lepidoptera:
Lycaenidae) feeding on Potentilla (Rosaceae). Ent. News 85: 40-44.
SHIELDS, O. A., 1984. Comments on recent papers regarding western Cupres-
saceae — feeding Callophrys (Mitoura). Utahensis 4:51-56.
STEHR, F. W., 1987. Immature Insects. Kendall/Hunt Co., Dubuque. 754pp.
V ANE- WRIGHT, R.I., 1978. Ecological and behavioral origins of diversity in
butterflies. Symp. Roy. Ent. Soc. Lond. 9:56-70.
WEBSTER, R. P. & M. C. NIELSEN, 1984. Myrmecophily in the Edward’s hairstreak
butterfly Satyrium edwardsii (Lycaenidae). J. Lepid. Soc. 38:124-133.
WRIGHT, D. M., 1983. Life history and morphology of the immature stages of the
bog copper butterfly Lycaena epixanthe (Bsd. & Le C.). J. Res. Lepid. 22:47-
100.
60
J.Res.Lepid.
GLOSSARY
Al, A2, A3,. . . A10: Abdominal segments 1, 2, 3,. . .10.
Adfrontal sutures: A pair of sutures extending dorsally from the
anterior mandibular articulations and converging at the stem of the
epicranial (or coronal) suture. In many Lepidoptera (but not Lycae-
nidae) two pairs of roughly parallel sutures extend from the coronal
suture to the mandibular articulations, the more mesal frontal
sutures and lateral to them the adfrontal sutures.
Allopatric: Occurring in different areas; usually pertaining to species or
subspecies whose ranges do not overlap.
Anal: Pertaining to the last adbominal segment.
Anal prolegs: The terminal pair of prolegs on abdominal segment 10.
Biordinal: Two sizes, as in two lengths of crochets arising from a single
line.
Bivoltine: Having two generations per year.
Buttressed chalazae: Chalazae with vertical lateral ridges which fuse
distally with the cuticle, appearing buttressed (fig. 2n).
Ca: About or approximately.
Capitate: Abruptly enlarged distally, especially pertaining to setae.
Caudad: In the direction of the anal or tail end; posterior.
Cephalad: In the direction of the head; anterior.
Cephalic: Pertaining to the head.
Chaetotaxy: The arrangement or distribution of setae.
Chalaza(ae): A sclerotized basal papilla from which a seta arises.
Clavate: Club-shaped; more-or-less cylindric and gradually enlarged
distally, especially pertaining to setae.
Coxa(ae): The basal segment of a true leg.
Crochet(s): A hooklike sclerotized structure at the distal end of a proleg,
usually with many others arranged in rows.
Cylindric: Circular in cross-section, especially pertaining to setae.
Dendritic seta(ae): A specialized seta (usually) with relatively long, fine
lateral spicules and often occurring in close proximity to the honey
gland and spiracles (figs. 15-18).
Distal: Toward the end of an appendage farthest from its attachment.
Dorsal: Toward or pertaining to the dorsum.
Dorsal line: A longitudinal line along the dorsum, often apparent as a
contrasting color with respect to the ground color.
Dorsum: The top of a larva when resting on a substrate; the side opposite
its legs.
Echinoid seta(ae): Short globular setae with stout, apically flared
processes found in Calephelis larvae (fig. 31).
Epicranial suture: A ‘Y’-shaped suture separating left and right halves
of the cranium which is forked anteriorly.
Eversible tubercle(s): A fleshy tubercle in many lycaenid larvae, located
slightly posterolateral to the spiracle on abdominal segment 8, which
is normally retracted and not visible.
27(1): 1-81, 1988
61
Filiform: Filamentous or thread-like; slender, long, and little or not at
all tapered, especially pertaining to setae.
Flagelliform: Whip-like; slender and finely tapered, especially per-
taining to setae.
Frons: That portion of the head between the anterior arms of the
epicranial suture (adfrontal or frontal sutures) and immediately
above the clypeus.
Frontal sutures: The anterior arms of the epicranial suture which
terminate at the anterior mandibular articulations and form the
lateral margins of the frons. In lycaenid larvae these are also known
as adfrontal sutures.
Ground color: The primary color of a larva on which a pattern of
contrasting color(s) may be superimposed.
Honey gland: A partly eversible transverse middorsal gland on abdo-
minal segment 7.
Instar: The stage between larval molts; the first instar emerges from an
ovum; the last instar immediately precedes the pupal stage.
Lateral: Pertaining to the sides.
Lateral fold: A fleshy cuticular fold below the spiracles extending
posteriorly from Tl.
Lateral line: A longitudinal line of contrasting color coinciding with the
lateral fold.
Lateral spicule: A small filamentous or spine-like lateral process of a
seta.
Lenticle: A small cuticular lens-like structure, surmounting or set into
a short chalaza-like collar, found in hesperiids and lycaenids.
Mesoseries: A band of crochets along the mesal side of a proleg.
Mesothorax: The second segment of the thorax, bearing the second pair
true legs.
Metathorax: The third segment of the thorax, bearing the third pair of
ture legs.
Middorsal: Located along the dorsal midline.
Multi voltine: Having several (continous) generations per year.
Mushroom lenticle: A stalked lenticle, narrowest at the base, found in
Atlides halesus (fig. 26).
Mushroom seta(e): A short, stout, multibranched seta, resembling a
mushroom when viewed under low magnification, found in the
Lycaeninae (figs. 2q, 22).
Neck setae: Short, stout, tooth-like setae on the neck area of lycaenid
larvae (fig. 27).
01, 02, 03,. . .06: Ocelli 1-6; numbered as in fig. 68.
Ocellus(i): One of six simple eyes or stemmata located on each side of the
head of Lepidoptera larvae (fig. 68).
Onisciform: Somewhat spindle-shaped but ventrally flattened, as in the
sow bug ( Oniscus ).
Papilla(ae): A small cuticular projection or elevation.
62 J. Res. Lepid.
Pheromone: A chemical used by an organism to communicate with
another member of its species.
Planta: The distal end of the proleg to which the crochets are attached.
Plumose seta(e): A type of seta (usually very long and filamentous)
found in Apodemia and Calephelis larvae which has numerous short,
fine lateral processes (figs. 31, 32).
Polymorphic: Having several forms or color patterns.
Posterior: Caudad.
Primary seta(e): Setae representing the archetypal setation of the
Lepidoptera, occurring in fixed numbers and locations in many
lepidopterous families but only in first instars of Lycaenidae.
Proleg: A fleshy appendage with distal crochets occurring in pairs
ventrally on the abdominal segments.
Prominence: A fleshy elevation of the body surface.
Prominent seta(e): Setae occurring in locations typical for primary setae
and which are distinguishable from surrounding secondary seta by
their greater length and/or erectness; in some cases these may be
primary setae but often they exceed the basic number of primary
setae in a given location.
Pupa: The resting stage intermediate between the mature larva and
adult in holometabolous insects.
Reclinate: Lying against or at a low angle to the body surface, especially
pertaining to setae which are not bent near the base.
Recumbent: Lying down or reclining against the cuticle, especially
pertaining to setae strongly bent near the base.
Recurved: Broadly bent back toward the base, especially pertaining to
setae.
Retractile: Retractable; able to be withdrawn, as with the head of many
lycaenid larvae.
Sclerite: A hardened (sclerotized) part of the body wall.
Secondary seta(e): Those setae occurring in addition to the basic
complement of primary setae in larvae of lycaenids and many other
lepidopterous families.
Semiochemical: Chemicals produced by one organism that incite re-
sponse in other organisms.
Sensory seta(e): A specialized pair of setae located anterodorsal to the
prothoracic spiracles; in lycaenid larvae they are on the prothoracic
shield, near its lateral margins.
Seta(e): A sclerotized hair or bristle surrounded basally by a small
cuticular ring and often arising from a chalazae.
Spatulate: Enlarged and compressed or flattened distally, as in a
spatula, especially pertaining to setae.
Spatulate lobe: A fleshy lobe (usually distally flared and flattened)
arising near the center of the mesoseries of crochets on most lycaenid
larvae.
Spicule: A spine-like projection.
27(1): 1-81, 1988
63
Spinule: A short sclerotized cuticular projection.
Spiracle: A sclerotized, cuticular pore associated with internal tracheae,
a pair of which are located laterally on the prothorax and abdominal
segments 1-8.
Stellate chalaza: A chalaza with distal or lateral pointed projections.
Stemma(ta): One of a group of lateral ocelli found in lepidopterous
larvae; ocellus.
Subdorsal: Located slightly lateral to the dorsal midline, intermediate
between the dorsal and lateral regions.
Subprimary seta(e): Those setae (additional to primary setae) occurring
in fixed locations typical of some families.
Suture: A seam where two sclerites join.
Sympatric: Occurring in the same area.
Tl, T2, T3: Referring to the prothorax, mesothorax, and metathorax,
respectively.
Tapered: Becoming gradually narrower distally, especially pertaining
to setae.
Triordinal: Pertaining to crochets of three lengths arising from a single
row.
Taxon(a): A taxonomic unit such as species, genus, family, etc.
Uniordinal: Pertaining to crochets of a single length arising from a
single line.
Ventral: Pertaining to the lower side of a larva when resting on the
substrate; the side from which the legs and prolegs arise.
Ventral prothoracic gland: An eversible gland arising midventrally
anterior to the prothoracic legs of some Lepidoptera larvae.
Verruca(e): A distinctly bounded (often sclerotized, pigmented, or
raised) area from which several setae arise.
64
J.Res.Lepid.
Appendix 1. List of the Lycaenidae of California
Riodininiae:
Apodemia mormo (C. and R. Felder, 1859)
" palmerii (W. H. Edwards, 1870)
Calephelis nemesis (W. H. Edwards, 1871)
wrighti Holland, 1930
Lycaeninae:
Lycaena arota (Boisduval, 1852)
" cupreus (W. H, Edwards, 1870)
" editha (Mead, 1878)
" gorgon (Boisduval, 1852)
" helloides (Boisduval, 1852)
" hermes (W. H. Edwards, 1870)
" heteronea (Boisduval, 1852)
" mariposa (Reakirt, 1866)
" nivalis (Boisduval, 1869)
" phlaeas (Linnaeus, 1761)
" rubidus (Behr, 1866)
" xanthoides (Boisduval, 1852)
Theclinae:
Atlides halesus (Cramer, 1777)
Callophrys (Callophrys) comstocki Henne, 1940
dumetorum (Boisduval, 1852)
" " lemberti Tilden, 1963
" " perplexa Barnes and Benjamin,
(Incisalia) augustus (W. Kirby, 1837)
eryphon (Boisduval, 1852)
" " fotis (Strecker, 1878)
" " mossii (Hy. Edwards, 1881)
(Mitoura) johnsoni (Skinner, 1904)
" " loki (Skinner, 1907)
" " nelsoni (Boisduval, 1869)
" " siva (W. H. Edwards, 1874)
spinetorum (Hewitson, 1867)
thornei (Brown, 1983)
Habrodais grunus (Boisduval, 1852)
Harkenclenus titus (Fabricius, 1793)
Ministrymon leda (W. H. Edwards, 1882)
Satyrium auretorum (Boisduval, 1852)
Satyrium behrii (W. H. Edwards, 1870)
" calif ornica (W. H. Edwards, 1862)
" fuliginosum (W. H. Edwards, 1861)
" saepium (Boisduval, 1852)
" sylvinus (Boisduval, 1852)
" tetra (W. H. Edwards, 1870)
Strymon avalona (W. G. Wright, 1905)
" columella (Fabricius, 1793)
" melinus Hiibner, 1818
1923
27(1): 1-81, 1988
65
Poly ommatinae :
Agriades franklinii (Curtis, 1835)
Brephidium exilis (Boisduval, 1852)
Celastrina argiolus (Linnaeus, 1758)
Everes amyntula (Boisduval, 1852)
" comyntas (Godart, 1824)
Euphilotes battoides (Behr, 1867)
" enoptes (Boisduval, 1852)
mojave (Watson and W. P. Comstock, 1920)
" rita (Barnes and McDunnough, 1916)
Glaucopsyche lygdamus (Doubleday, 1841)
" piasus (Boisduval, 1852)
Hemiargus ceraunus (Fabricius, 1793)
" isola (Reakirt, 1866)
Icarica acmon (Westwood and Hewitson, 1852)
" icarioides (Boisduval, 1852)
" lupini (Boisduval, 1869)
" neurona (Skinner, 1902)
" shasta (W. H. Edwards, 1862)
Leptotes marina (Reakirt, 1860)
Lycaeides idas (Linnaeus, 1761)
melissa (W. H. Edwards, 1873)
Philotes sonorensis (C. and R. Felder, 1865)
Philotiella speciosa (Hy. Edwards, 1867)
Plebejus saepiolus (Boisduval, 1852)
Plebulina emigdionis (F. Grinnell, 1905)
66
J.Res.Lepid.
Appendix 2. New and reconfirmed larval host plants.
Host species
AIZOACEAE
Butterfly species
Sesuvium verrucosum Raf.
B. exilis
ASTERACEAE
Baccharis glutinosa Pers.
Bebbia juncea (Benth.) Greene
Helianthus annuus L. ssp. lenticularis
( Dough ) Ckll.
C. nemesis, C. argiolus
C. wrighti
S. melinus
CHEN OPODIACE AE
Atriplex canescens (Pursh) Nutt.
" semibaccata R. Br.
Chenopodium sp.
Salsola iberica Sennen & Pau.
Suaeda moquinii (Torr.) Greene
B. exilis, P. emigdionis
B. exilis
B. exilis
B. exilis
B. exilis
CONVOLVULACEAE
Cuscuta sp.
C. (I.) augustus
CRASSULACEAE
Dudleya abramsii Rose
" cymosa (Lem.) Britt. & Rose
" lanceolata (Nutt.) Britt. & Rose
" saxosa (Jones) Britt. & Rose
Sedum spathulifolium Hook.
P. sonorensis
P. sonorensis
P. sonorensis, S. melinus
P. sonorensis, S. melinus
C. (I.) mossii
CUPRESSACEAE
Juniperus calif ornica Carr.
C.(M.) loki, C. (M.) siva
ERICACEAE
Vaccinium arbuscula (Gray) Merriam
myrtillus L.
L. mariposa
L. mariposa (Oregon record)
FABACEAE
Amorpha calif ornica Nutt.
fruticosa L. var. occidentalis
(Abrams) Kearn. & Peeb.
L. marina, S. melinus
L. marina
Astragalus canadensis L. var. brevidens
(Gand.) Barneby
L. melissa
" douglasii (T. & G.) Gray
lentiginosus Dougl.
palmeri Gray
whitneyi Gray
Calliandra eriophylla Benth.
Dalea searalsiae (Gray) Barneby
E. amyntula,H. ceraunus,L.
melissa
E. amyntula, G. lygdamus,
L. melissa
E. amyntula, H. ceraunus
E. amyntula
L. marina
H. isola
27(1): 1-81, 1988
67
Hoffmannseggia microphylla Torr.
Lotus argophyllus (Gray) Greene
" crassifolius (Benth.) Greene
" humistratus Greene
" nevadensis Greene
" oblongifolius (Benth.) Green
" purshianus (Benth.) Clem. &
Clem.
" procumbens (Greene) Greene
" rigidus (Benth.) Greene
" scoparius (Nutt, in T. & C.) Ottley
Lupinus andersonii Wats.
" argenteus Pursh var. tenellus
(Dougl. ex D. Don) D. Dunn
" breweri Gray
" caudatus Kell.
" excubitus Jones
" latifolius Agardh
" magni ficus Jones
Marina parry i (T. & G.) Barneby
Medicago sativa L.
Prosopis glandulosa Torr.
" pubescens Benth.
Trifolium monanthum Gray
monoense Greene
sp-
Vicia benghalensis L.
FAGACEAE
Quercus chrysolepis Liebm.
" Cornelius -mulleri Nixon &
Steele
" douglasii H. & A.
" wislizenii A. DC.
LAMIACEAE
Rosmarinus officionalis L.
Mentha piperita L.
MALVACEAE
Gossypium hirsutum L.
Hibiscus denudatus Benth.
rosa-sinensis L.
Sphaeralcea emoryi Torr. in Gray
PLUMBAGINACEAE
Plumbago auriculata Lam.
H. ceraunus
S. avalona, C.perplexa
C.perplexa, S. melinus
G. lygdamus (Oregon record)
I. acmon
L. idas
I. acmon, L. melissa
G. lygdamus, L. marina
G. lygdamus
C.perplexa, G. lgydamus,L.
marina, S melinus
G. piasus, S. fuliginosum
I. icarioides
1. shasta
G. lygdamus, S. fuliginosum
G. piasus, I. icarioides,
G. lygdamus, G. piasus,!. acmon,
S. melinus
H. isola
H. isola
L. marina
H. ceraunus, L. marina, M.
leda
A.palmerii
P. saepiolus
P. saepiolus
P. saepiolus
G. lygdamus
H. grunus, S. auretorum
S. auretorum
S. auretorum
S. auretorum, S. californica
S. melinus
S. melinus
S. melinus
S. columella, S. melinus
S. melinus
S. melinus
L. marina
68
J.Res.Lepid.
POLYGONACEAE
Eriogonum caespitosum Nutt.
cinereum Benth.
davidsonii Greene
" deserticola S . Wat s .
" datum Dougl. ex Benth.
" elongatum Benth.
" fasciculatum Benth.
heermannii Dur. & Hilg.
heradeoides Nutt.
incanum Torr. & Gray.
inflatum Torr. & Frem.
" insigne Wats.
kennedyi Porter ex Wats.
" latifolium Sm.
lobbii T. & G.
marifolium T. & G.
microthecum Nutt.
nidularium Cov.
nudum Dougl. ex Benth.
oualifolium Nutt.
panamintense Morton
parvifolium Sm. in Rees
plumatella Dur. & Hilg.
pusillum T. & G.
reniforme Torr. & Frem.
roseum Dur. & Hilg.
thurberi Torr.
umbellatum Torr.
wrightii Torr. ex Benth.
I. lupini
E. battoides
E. enoptes, E. rita, H.
ceraunus,
I. acmon, S. melinus
A. mormo
E. enoptes, S. melinus
C. perplexa, E. enoptes, H.
ceraunus, I. acmon, L. gorgon,
S. melinus
A. mormo, C. comstocki,E .
battoides, I. acmon, I. lupini,
L. heteronea
A. mormo, C. comstocki,E.
battoides
C. lemberti
E. battoides, C. lemberti
A. mormo, S. melinus
A. mormo
C. comstocki,E. battoides,
E. enoptes, I. demon, I.
neurona
A. mormo, C. dumetorum,E .
enoptes
E. battoides, I. lupini
C. lemberti, E. battoides
C. comstocki,E. battoides,
E. rita, S. melinus
C. comstocki
A. mormo, C. lemberti, E.
enoptes, I. acmon, L. gorgon, S.
melinus
E. battoides, I. acmon, I. lupini
E. enoptes
E. battoides, E. enoptes,!.
acmon, S. melinus
E. enoptes, E. rita, I. acmon, H.
ceraunus
E. mojaue
E. mojave,H. ceraunus,!.
acmon, P. speciosa
E. enoptes, E. rita, S. melinus
I. acmon
A. mormo, C. comstocki, C.
lemberti, E. battoides, E.
enoptes,!. lupini,!. neurona, L.
heteronea
C. argiolus (Arizona record), E.
enoptes, H. ceraunus, I. acmon,
S. melinus
27(1): 1-81, 1988
69
Oxyria digyna (L.) Hill
Oxytheca perfoliata T. & G.
Polygonum amphibeum L.
lapathifolium L.
R umex angiocarpus Murbeck
" californicus Rech.
" crispus L.
" paucifolius Nutt, ex Wats.
" salicifolius Weinm.
" triangulivalvis (Danser) Rech.
L.phlaeas
A. morrno
L. hellodies
S. melinus
L. editha
L. cupreus
L. xanthoides
L. cupreus, L. editha
L. editha, L. xanthoides
L. cupreus, L. rubidus
PRIMULACEAE
Dodecatheon alpinum (Gray) Greene A.franklinii
RHAMNACEAE
Ceanothus cordulatus Kell.
cuneatus (Hook.) Nutt.
" crassifolius Torr.
greggii Gray var. perplexans
(Trel.) Jeps.
leucodermis Greene
" oliganthus Nutt, in T. & G.
palmeri Trel.
Rhamnus crocea Nutt, in T. & G.
" ilicifolia Kell.
S. californica, S. saepium
S. saepium
S. saepium
C. (I.) augustus, S. saepium
C. (I.) augustus, C. argiolus, S.
saepium
S. saepium
C. (I.) augustus, C. argiolus
C. (I.) augustus, L. hermes
C . (I) augustus
ROSACEAE
Adenostomafasciculatum H. & A.
Cercocarpus betuloides Nutt, ex T. & G.
Cowania mexicana D. Don var.
stansburiana (Torr.) Jeps.
Heteromeles arbutifolia M. Roem.
Malus sylvestris (L.) P. Mill.
Prunus ilicifolia (Nutt.) Walp.
Purshiaglandulosa Curran
" tridentata (Pursh) DC.
Rubus ursinus Cham. & Schlecht.
SALICACEAE
Salix sp.
" hindsiana Benth.
" lasiolepis Benth.
C. argiolus, C. (I.) augustus, L.
marina
S. tetra
C. (I.) fotis
C. argiolus, C. (I.) augustus
S. melinus
C. argiolus, C. (I.) augustus
S. behrii
S. behrii
S. melinus
S. melinus
S. sylvinus, S. melinus
S. sylvinus
SAXIFRAGACEAE
Ribes quercetorum Greene
" roezlii Regel.
" velutinum Greene
L. arota
L. arota
L. arota
70
J.Res.Lepid.
VISCACEAE
Arceuthobium campylopodum Engelm.
in Gray
Phoradendron tomentosum (Englm. ex
Gray)
C.(M.) spinetorum
A. halesus
gland, Meg, lc= lateral
chevron, 11= lateral line, ml=
middorsal line, n=neck, p=
proleg, pds=prominent dorsal
setae, pls=prominent lateral
setae, ps=prothoracic shield,
pss=prominent subdorsal
setae, sl=subdorsal line, sp=
spiracle, ss= sensory setae.
Fig. 2. Some variations in lycaenid setal structure: a) erect, slightly
curved prominent seta (L nivalis), b) broadly recurved and finely
tapered seta (L marina), c) seta sharply bent near base and apically
truncate (L nivalis), d) semierect, apically truncate seta (L mariposa), e)
erect, straight seta with stellate chalaza (L. marina), f) recurved, apically
pointed seta with stellate chalaza (C. argiolus), g) clavate-capitate seta
{H. titus), h) clavate-capitate seta (L xanthoides), i) slender recumbent
seta (L gorgon), j) erect, apically bent seta with stellate chalaza (L
melissa), k) recurved, apically blunt seta with stellate chalaza (G. piasus),
I) spindle-shaped, apically pointed seta (H. ceraunus), m) clavate,
truncate seta (/. acmon), n) reclinate seta with buttressed chalaza (S.
California), o) slightly inclined, blunt seta with buttressed chalaza (S.
fuliginosum), p) weakly dendritic seta with nonsculptured chalaza (L
heteronea), q) mushroom seta with tapered, unbranched processes (L
arota). Bar scale ^-0.1 mm for a-k and p and 0.01mm for l o and q.
72
J. Res. Lepid.
Figs. 3-8. Some head and body setae. Fig. 3. Philotiella speciosa head. Fig. 4.
Prominent seta and other setae on mesothoracic dorsal prominence of
Callophrys (M.) johnsoni. Fig. 5. Apodemia mormo head; note long
setae on cranium and anterior margin of prothorax. Fig. 6. Branched
sensory seta on Atlides halesus. Fig. 7. Lateral view of prothoracic
dorsum of Callophrys (C.) perplexa. Note relatively low dorsal pro-
minence posterolateral to the prothoracic shield with transverse row of
prominent setae. Fig. 8. Lateral view of prothoracic dorsum of C.
(Mitoura) loki. Note relatively high dorsal prominence posterolateral to
the prothoracic shield lacking transverse row of prominent setae. Scale
bar=0.1mm for figs. 3-6 and 1.0mm for figs. 7 and 8.
27(1): 1-81, 1988
73
Figs. 9-14. Sensory setae on prothoracic shield. Fig. 9. Callophrys (M.) loki . Fig.
10. Plebulina emigdionis. Fig. 11. Apodemia mormo. Fig. 12. Cal-
lophrys (M.) spinetorum. Fig. 13. Euphilotes mojave . Fig. 14. Lycaena
xanthoides. Scale bar - 0.1 mm.
74
J.Res.Lepid.
Figs. 15-20. Dendritic setae, eversible tubercle, and honey gland. Fig. 15.
Dendritic setae around A7 spiracle of Lycaena xanthoides; note
lenticles, clavate-capitate setae, and mushroom setae. Fig. 16.
Dendritic seta near A1 spiracle on Celastrina argiolus ; note stellate
chalaza and lenticles. Fig. 17. Dorsolateral dendritic setae on
mesothorax of Satyrium behrii; note strongly dentate setae bent
parallel to body surface. Fig. 1 8. Dendritic setae near honey gland of
Philotes sonorensis (silk fiber across center). Fig. 19. Everted honey
gland of Euphilotes battoides. Fig. 20. Partially everted eversible
tubercle of Celastrina argiolus ; note strongly spiculate setae at apex.
Scale bar=0.01 mm for figs. 1 6 and 1 8 and 0.1 mm for ail other figs.
27(1): 1-81, 1988
75
Figs. 21 -26. Fig. 20. Lycaena hermes prothoracic shield. Fig. 22. Mushroom seta
of Lycaena xanthoides. Fig. 23. Lateral view of setae posterolateral to
the prothoracic shield of Lycaena arota\ note most setae directed
caudad. Fig. 24. Lateral view of setae posterolateral to the prothoracic
shield of Lycaena nivalis ; note setae directed cephalad. Fig. 25.
Capitate setae near honey gland of Philotes sonorensis. Fig. 26.
Mushroom lenticle of Atlides halesus. Scale bar=0.1mm for figs. 21 ,
23, and 24 and 0.01mm for figs. 22, 25, and 26.
76
J.Res.Lepid.
Figs. 27-32. Various setae and spinules. Fig. 27. Neck setae and spinules of
Callophrys (M.) johnsoni. Fig. 28. Spinules on neck of Plehulina
emigdionis. Fig. 29. Setae on the frons of Callophrys (I.) mossii. Fig.
30. Lateral view of abdominal segments one and two of Apodemia
mormo ; note A1 spiracle anteroventral to lateral verruca. Fig. 31.
Dorsal verruca of Calephelis nemesis ; note echinoid (lower right)
and plumose setae. Fig. 32. Spatulate tip of plumose seta of
Calephelis nemesis. Scale bar=0.01 mm for figs. 27, 28, and 32 and
0.1mm for figs. 29-31.
27(1): 1-81, 1988
77
Figs. 33-44.
Figs. 45-54.
Fig. 55.
Fig. 56-58.
Cranial pigmentation (left) and prothoracic shields (right) for 12
Lycaena species. Fig. 33. L phlaeas. Fig. 34. L cuprous. Fig. 35. L
arota. Fig. 36. L nivalis. Fig. 37. L. gorgon. Fig. 38. L heteronea. Fig.
39. L hermes. Fig. 40. L hel Icicles. Fig. 41 . L mariposa. Fig. 42. L
editha. Fig. 43. L. rubidus. Fig. 44. L xanthoides. Scale bar= 1 mm
for prothoracic shield; crania not drawn to scale.
Cranial pigmentation (left) and prothoracic shields (right) of Atlides,
Habrodais, Harkenclenus, and Satyrium species. Fig. 45. A. halesus.
Fig. 46. H. g run us. Fig. 47 ? H. titus. Fig. 48. S. fuligi nosum. Fig. 49. S.
saepium. Fig. 50. S. sy Ivin us. Fig. 51 . S. auretorum. Fig. 52. S. behrii.
Fig. 53. S. californica. Fig. 54. S. tetra. Scale bar= 1 mm for prothoracic
shields; crania not drawn to scale.
Mandibular setation of Satyrium saepium and Lycaena xanthoides,
aboral surface of left mandibles. Fig. 55a. S. saepium. Fig. 55b. L.
xanthoides. Scale bar-- 0.1 mm.
Dorsal profile and setae of abdominal segments 1 -3 of Satyrium tetra ,
S. auretorum, Ministrymon led a, Euphilotes enoptes, E. mojave, E.
rita, Hemiargus ceraunus, and H. isola. Fig. 56a. S. tetra. Fig. 56b. S.
auretorum. Fig. 56c. M. leda. Fig. 57a. E. enoptes. Fig. 57b. E.
mojave. Fig. 57c. E. rita. Fig. 58a. H. ceraunus. Fig. 58b. H. isola.
Scale bar-1 mm.
73
Figs. 59-67.
Fig. 68.
Figs. 69, 70.
Fig. 71.
Fig. 72.
Fig. 73.
Cranial pigmentation and prothoracic shields of some Callophrys and
Strymon species. Fig. 59. C. perplexa cranium. Fig. 60. C. comstocki
cranium. Fig. 61. C. (Mitoura) spinetorum cranium (left) and pro-
thoracic shield (right). Fig. 62. C. (Incisalia) augustus cranium. Fig.
63. C. (I.) fotis cranium. Fig. 64. C. (I.) mossii cranium (left) and
prothoracic shield (right). Fig. 65. S. avalona cranium. Fig. 66. S.
columella cranium. Fig. 67. S. melinus cranium (left) and prothoracic
shield (right). Scale bar - 1 mm for prothoracic shields; crania not
drawn to scale.
Lycaena xanthoides ocelli; ocelli are numbered counterclockwise
from the top. Scale bar^lmm.
Lateral view of abdominal segments 1 and 2 of Apodemia mormo and
Calephelis nemesis showing placement of spiracles and verrucae.
Fig. 69. A. mormo. Fig. 70. C. nemesis. Scale bar - 1 mm.
Prothoracic shield of Icaricia lupins. Scale bar=1mm.
Planta of prolegs of Habrodais grunus and Everes amyntula. Fig.
72a. H. grunus; note short series of lateral crochets. Fig. 72b. E.
amyntula ; note pigmented basal margins of spatulate lobe. Scale
bar= 0.5mm.
Distended last instar Callophrys (Mitoura) spinetorum larva. Note
prominent setae confined to dorsal and lateral prominences, other
setae very short and sparse. Scale bar= 1 mm.
Fig. 74. Larval photographs of 40 lycaenid species. Species are listed from left to
right(a-d) by row (1-10) from the top. Row 1 : a) A mormo , b) A. palmerii, c)
C. nemesis, d) L. arota ; row 2: a) L gorgon, b) L hermes, c) L heteronea,
d) L mariposa ; row 3: a) L. nivalis, b) L phlaeas, c) L. xanthoides, 6)H.
g run us] row 4: a) A. halesus, b) C. (C.) dumetorum, c) C. (C.) perplexa, d)
C. (I.) augustus] row 5: a) C. (I). eryphon, b) C. (M.) nelsoni, c) C. (M.)
spinetorum, d) H. titus] row 6: a) M. leda, b) S. auretorum, c) S. behrii, d)
S. californica] row 7: a) S. fuligi nosum, b) S. syivinus, c) S. tetra, d) >4.
franklinii ; row 8: a) B. exilis, b) C. argiolus, c) E. battoides, d) E. rita\ row 9:
a) G. lygdamus, b) G. piasus, c) /so/a, d) i. shasta ; row 1 0: a) L melissa,
b) P. sonorensis, c) P. speciosa, d) P. emigdionis. All subjects greater
than life size. Orientation is standard (cephalad at left) except Id, 2d, 5a,
7b, 8c, and 9d with cephalad at right.
-I®>
S 7605
i nt JOURNAL
OF RESEARCH
ON THE LEPIDOPTERA
( n pn
I oec r 5 m
VARIES
Volume 27
Number 2
Summer 1988 (1989)
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
The Lepidoptera Research Foundation, Inc.
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Beverly Hills, California 90210
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William Hovanitz
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ISSN 0022 4324
Published By:
Founder:
Editorial Staff:
Associate Editors:
Journal of Research on the Lepidoptera
27(2): 83-95, 1988(89)
A Study of Protesilaus microdamas (Burmeister) and
the Little-known P. dospassosi (Rutimeyer) and
P. huanucana (Varea de Luque) (Papilionidae)
by
Kurt Johnson
Department of Entomology, American Museum of Natural History, Central Park West
at 79th Street, New York, New York 10024
David Matusik
Department of Entomology, Field Museum of Natural History, Roosevelt Road, Chicago,
Illinois, 60605
and
Rick Rozycki
5830 South McVicker Avenue, Chicago, Illinois, 60638
Abstract. Certain wing and genitalic characters of P. microdamas
are distinctive from other Protesilaus sens. lat.. Based on these genita-
lic distinctions, P. dospassosi is associated with Protesilaus , and P.
microdamas in particular, for the first time. P. huanucana is accorded
species status based on wing and genitalic examination of all Protesi-
laus taxa. Specimens with wings similar to P. huanucana , but sharing
the distinctive genitalic traits of P. microdamas and P. dospassosi, are
discussed relative to their possibly representing a further terminal
taxon of the genus. All of the above taxa have received little or no
previous taxonomic examination.
Introduction
Recently, we published taxonomic studies of several groups of Papi-
lionidae (Johnson, Rozycki and Matusik, 1985, 1986a, 1986b; Johnson
and Rozycki, 1986). These studies resulted because we were able to
assemble samples of several papilionids previously known from only
their types or which were apparently undescribed. This research led to
cooperation with Dr. Keith S. Brown (Universidade Estadual de Cam-
pinas, Sao Paulo, Brazil) who is preparing a synonymic list of Neotro-
pical Papilionidae, since there was mutual interest in the examination
of types and the location of specimens of certain little-known taxa. The
i present paper summarizes taxonomic results concerning taxa of the
genus Protesilaus ( sensu Hancock, 1983). Some results of this study
84
J.Res.Lepid.
were published in our paper concerning P. illuminatus Niepelt (John-
son, Rozycki and Matusik, 1986b), a taxon previously known from one
extant syntype male and accompanying female but of which we were
able to assemble recently collected specimens. The results of the pre-
sent paper concern a cluster of Protesilaus taxa which have hitherto
been either little-known or of uncertain status.
Study of male genitalia of Protesilaus indicated P. microdamas (Bur-
meister) differed from all other Protesilaus taxa in lacking the ventral
process of the valval harpe. This process is prominent in other taxa of
the group. Examination of the unique type of Papilio dospassosi
Rutimeyer (a taxon inadvertantly misplaced in Heraclides by Hancock,
1983, who had not examined the type) indicated that P. dospassosi
belongs in Protesilaus and that its holotype also lacks a ventral valval
process. Further study resulted in location of another assemblage of
specimens in Protesilaus lacking this process. We suspected these
represented an undescribed taxon. Since these specimens and P. dos-
passosi were characterized by extreme reduction of red on the hind-
wing upper surface, and since these and P. microdamas lacked the
ventral valval process, a previously unrecognized species group within
Protesilaus was suggested.
Subsequently, Keith Brown discovered that little-known Papilio
huanucana Varea de Luque (1975) matched the salient appearance of
specimens located by us which lacked the ventral valval process. Oddly,
however, dissection of representatives of the types series of P. huanu-
cana disclosed a much larger valval process than in other Protesilaus.
Hence, the present study inadvertently discovered several taxonomic
characters suggesting species status for P. huanucana , but it is doubt-
ful that this species is closely related to P. microdamas. The purpose of
the following presentation, therefore, will be to review the taxonomic
characters of P. microdamas and enumerate the several new statuses
and synonymies which result from our study of it, P. dospassosi and P.
huanucana. Also, we will discuss the specimens resembling P. huanu-
cana which lack the ventral valval process in hope that this review will
promote eventual discovery of whether authentic natural populations
exist which exhibit the wing markings characterizing P. huanucana
but lack the ventral valval process characteristic of P. microdamas and
the holotype of P. dospassosi.
Eventual resolution of the precise cladistic relationships in Protesilaus
will require a full consideration of character polarity in its own and
outgroup taxa. Such study cannot be accomplished, however, without
definition of the relevant terminal taxa. To this end, the following
treatment is provided.
Taxonomic Analysis
Both Munroe (1960) and Hancock (1983) recognized apparent
27(2): 83-95, 1988(89)
85
monophyly in a "lysithous- related group” within the genus Eurytides.
Hancock (1983) accorded this group generic status as Protesilaus.
According to these authors the group includes the following taxa,
which as noted below are tailed or untailed and mimic various other
neotropical butterfly taxa: Short-tailed or Untailed — pausanius
(Hewitson) [heliconine mimic]; protodamas (Godart) [banded, or helico-
nine mimic depending on form]; microdamas (Burmeister); phaon
(Boisduval) [banded]; chibcha (Fassl); euryleon (Hewitson); hipparchus
(Staudinger); harmodius (Doubleday); trapeza (Rothschild and Jordan);
xynias (Hewitson); ariarathes (Esper); ilus (Fabricius); branchus
(Doubleday); belesis (Bates) [troidine papilionid mimics]. Long-Tailed
— thymbraeus (Boisduval), lysithous (Hubner), kumbachi (Vogeler),
asius (Fabricius).
Morphological Structures: Protesilaus, lysithous group, taxa exhibit a
single-layered valval harpe (dark, keel-like structure centrad in Figs.
2—6) with a laterally extending spike associated just ventrad and a
variously rhomboid structure cephalo-ventrad exhibiting a ventrally
extending process. Contrastingly, the sister "marcellus group” taxa
(. sensu Munroe, 1961) display a harpe of two parallel layers (as with a
keel beneath a keel) without an emphatic associated spike and without
a ventrally extending process. The keel-like structure in Figs. 2-6
consists ventrad of two closely paralleled high ridges (drawn in thick
solid black) separated by a deep fissure (shown in white or very light
gray). The ventrad ridge is variously dentate. The keel can terminate
caudad with a variously expressed “head”, characteristically single-
edged and serrate, double-edged and serrate, or non-serrate in particu-
lar species clusters. The laterally pointing spike can be characteristical-
ly pointed, furcate, or conical, the ventral process of the rhomboid
structure variously emphatic. Characters of the keel of the valval
harpe are most useful with those of the laterally pointing spike being
less reliable though sometimes distinctive in some taxa.
Phenetic resemblance in the genitalia of short-tailed or non-tailed
members of the lysithous group of Protesilaus generally support the
clustering by D’Abrera (1981) based on characters of the wing. Four
general groups, disparate from the long-tailed members of the group,
are suggested, as shown in the accompanying figures listed below,
named in accordance with cluster names proposed by Keith Brown
(pers. comm.) and cited with the appropriate D5Abrera (1981) page
numbers: the "phaon cluster” (pp. 62-63) [Fig. 2], the “ harmodius
cluster” (pp. 64—65) [Fig. 3], the “ ariarathes cluster” (pp. 66—67) [top])
[Fig. 4], and the ''belesis cluster”: (p. 67 [bottom]) [Fig. 5]. We (Johnson,
Rozycki and Matusik, 1986b) have reviewed the major genitalic charac-
ters generally defining these clusters.
Major exception to the general morphological similarity in the groups
listed above occurs in specimens having no ventral process on the
valval harpe. Such include all specimens examined by us or Brown of P.
86
J.Res.Lepid.
Fig. 1. Upper surfaces (above) and undersurfaces (below) of A. P. micro da -
mas , male (Sapucay, Paraguay, AMNH); B. P. huanucana, male
(Tingo Maria, Peru, BMNH); C. P. dospassosi , holotype male, (Rio
Putumayo, Colombia, AMNH).
27(2): 83-95, 1988(89)
87
microdamas (figured alone by D’Abrera, 1981, p. 63), the dospassosi
type, and some specimens otherwise like P. huanucana [Fig. 6—7]. We
thus propose the following species cluster as defined by the following
key:
GENXTALIC KEY TO MICRODAMAS SPECIES CLUSTER
1. Ventral surface of valval harpe without ventr ad-protruding pro-
cess; and (less reliably) medial process fingerlike structure
pointing caudad P. microdamas,
dospassosi and some specimens otherwise like huanucana (see
treatment later in paper) [Fig. 6]
2. Ventral surface of valval harpe with ventrad-protruding process;
and (less reliably) medial process wedgelike, pointing laterally ....
......... all remaining taxa of Protesilaus ( sensu Hancock, 1983)
[Figs. 2-5]
Major complication to the identification of Protesilaus taxa by wing
characters occurs from eastern Ecuador southward through Bolivia
because of reduction of upper surface hindwing red in most lysithous
group taxa of that region. This restriction of red basically to the anal
area of the hindwing probably results from a common mimic/model
relation involving all the taxa (see Sheppard, Turner, Brown, Benson
and Singer, 1984). The following key separates these regionally sym-
patric lysithous group taxa from P. microdamas and P. dospassosi
along with P. huanucana and P. huanucana-\ike specimens lacking the
ventral valval process. Because these species are usually primarily
distinguished by the pattern of red spotting on the upper wing surface,
the following key for populations with reduced red relies on characters
of the tail and marginal wing spotting. Thus, it will not successfully
identify every specimen. However, it will be useful in identifying most.
KEY TO SUPERFICIALLY SIMILAR TAXA
1. Upper surface with both wings banded.
P. microdamas [Fig. 1A]
1A. Upper surface not banded but with a white to yellow mimetic
patch caudo-medial on forewing and red spots or orbs in anal
and/or anal-medial areas of hindwing 2
2. Upper surface margin of hindwing with yellow dots or slashes in
cells, usually from anal margin to cell M2 and/or M3 .......... 3
2 A. Upper surface margin of hind wing without yellow dots or slashes
in cells 4
3. Yellow marginal markings are slashes extending costad to cell
M2 and with tail at terminus of cell Cui thinly pointed
P. xynias [Fig. 7A]
3 A. Yellow marginal markings are small dots extending costad only
88
J.Res.Lepid.
to cell M3 and with tail at terminus of cell Cux short and stubby
. . P. trapeza [Fig. 7B]
4. Margin of hind wing with either short stubby tail or thinly
pointed tail at terminus of vein Cux ......................... 5
4A. Margin of hind wing without noticable tail and with vein termini
all about equally crennated ................................ 6
phaon CLUSTER
Fig. 2. Genitalia of the “phaon cluster" (number of dissections, parenth-
eses) A. pausanias, Jepelacio, Peru (3); B . protodamas, Gavea, Brazil
(3); C. phaon, Colombia (3); D. eury/eon eury/eon, Costa Rica (3); E.
euryleon haenshi (Rothschild and Jordan), Balzabamba, Ecuador (3);
F. euryleon pithonius (Rothschild and Jordan), Cauca Valley, Col-
ombia (3); G. illuminatus, Rio Putumayo Valley, Colombia (2).
27(2): 83-95, 1988(89)
89
aria rat hes CLUSTER
Fig. 3. Genitalia of the “ariarathes cluster” (number of dissections, parenth-
eses, other localities, brackets). A. ariarathes ariarathes , French
Guiana (3); B. ariarathes gayi f. cyamon (Grey), Middle Rio Ucayali,
Peru (3), Alto Jurua, Brazil (1) [additional studied: gayi gayi, Janjui,
Peru (1), Buena Vista, Bolivia (1); gayi metagenes (Rothschild and
Jordan), Mt. Duida, Venezuela (1)]; C. ariarathes menes (Rothschild
and Jordan), Tukeit, Guyana (3). D. ariarathes, Janjui, Peru.
harmodius CLUSTER
Fig. 4. Genitalia of the ” harmodius cluster” (number of dissections, pa-
rentheses): E. harmodius harmodius, Bolivia (3); F. harmodius
xenaides (Hewitson), Rio Pastaza, Ecuador (3); G. trapeza. Rio Napo,
Ecuador (3); H. xynias, Rio Santiago, Peru (3).
90
J.Res.Lepid.
belesis CLUSTER
Fig. 5. Genitalia of the “belesis cluster": (number of dissections parenth-
eses): A. belesis, Soyolapan, Mexico (3); B. branchus, San Jeronimo
(Chiapas), Mexico (3); C. i/lus, Sosumuco, Colombia (3).
microdamas CLUSTER
Fig. 6. Genitalia of the “microdamas cluster" (number of dissections, pa-
rentheses): A. microdamas, Sapucay, Paraguay (5, including Santis-
sima Trinidad, Paraguay); B. holotype, dospassosi ; C. specimen of
uncertain status, wings markings like huanucana but lacking ventral
valval process like taxa of microdamas cluster, Rio Santiago, Peru (4)
including specimens listed in Fig. 7; D. P. huanucana, from Ehrmann
series (CMNH), Sarajacu, Ecuador.
27(2): 83-95, 1988(89)
91
5. Red markings in anal areas of cells CU x and CU2 . 7
5 A. Red marking in anal areas reduced or obsolescent so as pattern
in CUx and CU2 not discernable . . 8
6. Ventral surface of valval harpe with large ventrad protruding
process P. huanucana [Fig. IB]
6A. Ventral surface of valval harpe without ventral process
specimens of uncertain identity referenced in text.
7. Hindwing with upper surface red spots in anal area cells C\Ji
and CU2: two median (CUi, CU2), one postmedian (CUi); dorsal
surface of keel of valval harpe moderately wide cephalad .......
P. dospassosi [Fig. 1C]
7A. Hindwing with upper surface red spots in anal area cells CUi,
CU2 and 2A: two median (CU2, 2A), one postmedian (CU2);
dorsal surface of keel of valval harpe extremely wide cephalad . . .
8
8. Ground color blackish, margin of hindwing with either short
stubby tail ( a . gayi ) or thinly pointed tail (a. ariarathes) at
terminus of vein CUi .................. P. ariarathes [Fig. 7C]
8 A. Ground color brown to lighter brown without noticable tail and
with vein termini of hindwing all about equally crenated 6
Annotated Taxonomic List (Including New Synonymies and Statuses)
P. microdamas species cluster:
Protesilaus microdamas (Burmeister), Figs. 1A, 6A
Papilio microdamas Burmeister 1878. Description Physique de la
Republique Argentine. Lepidopteres 5: 63.
Adult. Fig. 1A. Length of forewing: X of 5 males (AMNH), 40.2 mm,, range
39.0 to 42.0 mm. Male Genitalia: Fig. 6A. Location of type: unknown. Type
Locality: Corrientes, Argentina. Distribution: From sparse representation in
collections, ascertained as at least Paraguay, Matto Grosso State, Brazil and
most probably some adjacent areas.
Remarks. A banded species, P. microdamas is not confusable with any
congener. However, its taxonomic affinity has been unclear and it is extremely
rare in collections. A series of 25 males and females at the AMNH (of which
only 5 are males) from Santissima-Trinidad , Cordillera Province, Paraguay,
collected by B. Podtiaguin from May to August in an unnoted year, is the largest
series in North American institutions. Other North American museums and
British Museum (see Acknowledgments) together have fewer than 15 speci-
mens. The rarity of females of Protesilaus taxa suggests the large Podtiaguin
sample probably represents an unusually fortuitous collecting locality for the
species.
Protesilaus dospassosi (Rutimeyer), new combination, Figs. 1C, 6B
Papilio dospassosi Rutimeyer 1969. J. Lepid. Soc. 23: 255—257.
Adult. Fig. 1C. Length of forewing: holotype, male, 37.5 mm. Male Genitalia:
Fig. 6B. Location of type: AMNH. Type locality: Rio Putumayo, Colombia.
Distribution: Known only from type locality.
92
J. Res. Lepid.
Remarks. As noted previously, Hancock (1983) inadvertantly misplaced this
taxon in the genus Heraclides (tribe Papilionini) since he had not examined the
type. Keith Brown (pers. comm.) suggested the need to differentiate P. dospas-
sosi from P. morrisi Ehrmann (Ehrmann, 1921; Holland, 1927) a taxon some-
what similar in original description. We have examined the type of P. morrisi
at the CMNH, a male from “Loja, SE Ecuador, 30 November 1914, Rev. Hyde
Collection”. The type and a group of associated specimens identified by
Ehrmann all have postmedian red spots costad to cells 2 A and CU2, and by
genitalic dissection are clearly allied to P. harmodius , of which morrisi should
probably be considered a subspecies. Keith Brown informs us that he has seen a
possible specimen of P. dospassosi in the collection of the Los Angeles County
Museum.
The apparent species status and insular distribution of P. dospassosi should
be considered in light of its local sympatry with several other extremely insular
butterfly taxa equally rare in collections surveyed by us. These include: P.
illuminatus (Niepelt) (Johnson, Rozycki and Matusik, 1986b), known from only
nine specimens; the nymphalid butterfly Anaeomorpha splendida Rothschild
(only four specimens located by Johnson and H. Descimon [Universitie de
Provence, Marseilles, France] at AMNH, BMNH or Museum National d’His-
toire Naturelle (Paris) and of which species a subspecies columhiana Niepelt
(1928) was named, and is known only from, two specimens taken near the type
locality of P. dospassosi; and an apparently undescribed subspecies of Prepona
werneri Hering and Hopp known from a single specimen obtained by Johnson
and Matusik from the same collectors capturing P. illuminatus in 1981.
According to a museum survey by Johnson and Descimon, P. werneri is itself
known from fewer than 10 specimens, though the exact number is uncertain
since most are owned by private collectors.
It remains to be clarified whether further specimens of P. dospassosi will
corroborate or falsify the notion that lack of the ventral valval process, as in the
holotype, indicates sister species relationship with P. microdamas.
Other Taxa:
Protesilaus huanucana (Varea de Luque), new combination, revised
status Figs. IB, 6D
Graphium trapeza huanucana Varea de Luque, 1975. Shilap. Rvta.
Lep. 3(9): 28-32.
Adult. Fig. IB. Length of Fore wing: X of 5 specimens in Ehrmann series
(CMNH), 39.8 mm., range 38.0 - 42.0 mm. Male Genitalia: Fig. 6D. Location of
type: British Museum (Natural History). Type Locality: Tingo Maria, Peru.
Distribution: noted from dissected specimens by Keith Brown (pers. comm.) as
including southwest Colombia, eastern Peru, Acre, Rondonia and Amazonas
states, Brazil and northern Bolivia.
Remarks. Hitherto, Varea de Luque’s publication of the name huanucana
has received no further report in the literature. The taxon is distinctive, a fact
which formerly led us and Brown (at AMNH, 1972) to note it as undescribed or
not identifiable. Also, Varea de Luque suggested that huanucana might be
“quiza bona species .” It is distinguishable from congeners by the extreme
reduction of red spotting on the upper surface of the hindwing (limited to anal
27(2): 83-95, 1988(89)
93
area only) and by its cream-yellow mimetic patch on the forewing upper
surface. Amongst other taxa with regionally reduced upper surface red, it is
distinguishable by other characters (see Key). These characters, along with
those of the genitalia have led us and Brown to consider it as a valid species
pending biological studies. The existence of this taxon was also recognized by
Ehrmann who designated a type (CMNH) for a manuscript name which was
never validly published (Holland, 1927). Ehrmann’s study series was from
“Sarayoi’u, E. Ecuador” or “Sarajacu, Oriente, Ecuador” undated and from E.
T. Owen in the Buckley Collection. Our dissection of these specimens shows
that all examined have a ventral valval process and are thus P. huanucana.
Process-less Specimens Otherwise like P. huanucana :
Adult. Fig. 7D. Length of Forewing: X of 3 males, AMNH, 41.8 mm., range
41.0 to 42.5 mm. Male Genitalia: Fig. 6C. Distribution: from dissections by the
authors: AMNH — Rio Santiago, Peru; Rio Purus, Brazil; Costa Rica, Bolivia;
David Matusik Collection - Costa Rica, Bolivia.
Remarks. Among specimens generally resembling P. huanucana a number of
specimens have been found which lack the ventral valval process and are thus
like P. microdamas and P. dospassosi (Fig. 6). A high frequency of these was
found when we first began sorting from collections P. huanucana-\ike speci-
mens which we suspected represented an undescribed entity. When Brown
discovered the apparent external similarity between our series and the P.
huanucana types he further discovered these latter had a large ventral valval
process. Subsequently, by looking at a larger range of specimens, we also found
such examples. A number of lepidopterists have been consulted concerning this
and there is concensus that the smoothly edged cephalo-ventral surface of the
valval harpe on which a ventrad pointing process has not developed must be
considered as a possibly strong character within Protesilaus. Also, we and John
Rawlins (CMNH) agree there are some differences in the wing markings of the
specimens without the ventral valval process which suggest they are often
separable from P. huanucana . The former appear more brown (as opposed to
black or blackish) than P. huanucana and evidence a more evenly crennated
hind wing margin and a more yellow to ochre fore wing mimetic patch.
It is important to ascertain whether specimens lacking the ventral valval
process represent simple and insignificant variation aside from the consistency
of the valval process character in P. microdamas , are individuals representing
reversion to a primitive process-less configuration characteristic of plesiomor-
phy in Protesilaus , or whether authentic natural populations occur with the P.
microdamas- like valval harpe being taxonomically significant. If the latter is
true, such populations would constitute another important terminal taxon in
the cladistic structure of the genus.
Summary and Conclusions
Study of wing and male genital characters in the genus Protesilaus
indicates P. microdamas differs significantly from congeners and that
at least one other taxon, P. dospassosi (hitherto not placed in Protesi-
laus), shares with P. microdamas a valval harpe with a smoothly edged
ventral surface. Other Protesilaus specimens have been found with a
Fig. 7. Examples of wing patterns from "Key to Superficially Similar Taxa"
and specimens resembling P. huanucana but lacking ventral process
on valval harpe. A. P. xynias, male (left, upper surface; right, under
surface) Rio Santiago, Peru, AMNH; B. P. trapeza, male (left, upper
surface; right, under surface) Rio Napo, Ecuador, AMNH; C. P.
ariarathes, male (upper surface; extent of under surface marking
similar) Janjui, Peru, AMNH; D. example of specimens somewhat
resembling P. huanucana but lacking ventral process on valval
harpe. These may represent an undescribed taxon, male (left, upper
surface; right, under surface) Costa Rica, Bolivia, AMNH.
similar, processless, ventral harpe surface. Still others, similar in wing
facies to these latter have a large ventral process and comprise the
taxon P. huanucana . This taxon, distinctive in a number of characters
from other Protesilaus , is suggested as a species level taxon whose
biology should be studied in detail. The status of specimens like P.
huanucana lacking the ventral valval process is unresolved. If authen-
tic natural populations are found which evidence this latter character,
such populations must be suspected as representing a further terminal
taxon for the genus.
27(2): 83-95, 1988(89)
95
Acknowledgments. Numerous specialists and museum curators aided in dis-
cussing data, reviewing manuscripts or searching various collections. Below,
we list these persons, noting after each the collection surveyed: Keith S. Brown
(collection Universidade Estadual de Campinas, Sao Paulo, Brazil; Museu
Nacional, Rio de Janeiro, Brazil; Collection Museo de Historia Natural “Javier
Prado”, Lima, Peru); Tommaso Racheli (Collection Institute de Zoologia Agrico-
la, Maracay, Venezuela; Collection Tommasso Racheli); Ernesto W. Schmidt-
Mumm (Collection Ernesto W. Schrriidt-Mumm); Olaf H. H. Mielke (Collection,
Dep. de Zoologia, Universidade Federal do Parana, Curitiba, Brazil); Lee D.
Miller (Allyn Museum of Entomology of the University of Florida, Sarasota,
Florida, U.S.A.); H. T. Hannemann (Zoologishes Museum der Humbolt Univer-
sitat, Berlin, Germany); Rienk de Jong (Rijkmuseum van Natuurlijke Historie,
Leiden, Netherlands); John E. Rawlins (Carnegie Museum of Natural History,
Pittsburgh, Pennsylvania, U.S.A.), Robert K. Robbins (National Museum of
Natural History, Washington, DC, USA); Richard Vane-Wright (British
Museum, Natural History, London, United Kingdom).
Keith S. Brown, David L. Hancock (National Museum, Republic of Zimbabwe,
Bulawayo, Zimbabwe) and Frederick H. Rindge (AMNH) kindly reviewed the
manuscript. The following persons discussed the project with us and reviewed
drafts or other materials: John E. Rawlins, Ernesto W. Schmidt-Mumm and
Tommasso Racheli.
Literature Cited
D’ABRERA, B. 1981. Butterflies of the Neotropical Region. Part 1. Papilionidae
and Pieridae. Landsdowne Editions. East Melbourne, 172 pp.
EHRMANN, G. A. 1921. Some new Papilios and Ornithoptera. Lepidoptera 5:
17-19.
HANCOCK, D. 1983. Classification of the Papilionidae (Lepidoptera): a phylogene-
tic approach. Smithersia 2: 1-48.
HOLLAND, W. J. 1927. The Lepidoptera named by George A. Ehrmann. Ann.
Carn. Mus. 17: 299-364.
JOHNSON, K. & R. ROZYCKI. 1986. A new species of the anchisiades Group of
Heraclides from Venezuela (Lepidoptera: Papilionidae). J. New York Ent.
Soc. 94(3): 383-393.
JOHNSON, K., R. ROZYKI, & D. MATUSIK. 1985. Species status and the hitherto
unrecognized male of Papilio diaphora Staudinger (1891), (Lepidoptera:
Papilionidae). J. New York Ent. Soc. 93: 99-109.
JOHNSON, K„ R. ROZYCKI & D. MATUSIK. 1986a. The female of Papilio xanthopleura
Godman & Salvin (Papilionidae). J. Lepid. Soc. 40: 65—66.
JOHNSON, K„ R. ROZYCKI & D. MATUSIK. 1986b. Rediscovery and species status of
the Neotropical swallowtail butterfly Papilio illuminatus Niepelt (Lepidop-
tera: Papilionidae). J. New York Ent. Soc. 94: 516-525.
MUNROE, E. 1961. The classification of the Papilionidae (Lepidoptera). Can. Ent.
Suppl. 17, 51 pp.
NIEPELT, W. 1928. Neue Tagfalter aus Columbien. Int. Entom. zeitschr. 21: 390.
SHEPPARD, P. M., J. R. G. TURNER, K. S. BROWN, W. W. BENSON & M. C. SINGER. 1985.
Genetics and the evolution of Muellerian mimicry in Heliconius butterflies.
Phil. Trans. R. Lond., B 308: 433-610.
Journal of Research on the Lepidoptera
27(2): 96-103, 1988(89)
Hand-pairing of Papilio glaucus glaucus and Papilio
pilumnus (Papilionidae) and hybrid survival on various
food plants
J. Mark Scriber1
and
Robert C. Lederhouse1
Department of Entomology, University of Wisconsin, Madison 53706
Abstract. Hand-pairing of a female Papilio glaucus with a male P.
pilumnus resulted in the hatching of 69 larvae. Hybrid larvae survived
on species of Lauraceae, and also on species of Rutaceae and Magno-
liaceae. Hybrid larvae did not initiate feeding on black cherry. Roth
larvae and the adult males that were produced were intermediate
between the two species in a variety of morphological traits.
Introduction
The three-tailed swallowtail, Papilio pilumnus Boisduval occurs from
southern Arizona and Texas southward to Guatemala (Beutelspacher
and Howe, 1984). Howe (1975) observed oviposition on a species of Litsea
(Lauraceae) in Chiapas, Mexico, however relatively little is known
about its larval stages or field biology (but see Scott, 1986). Because of
its superficial resemblance to adult tiger swallowtails (Fig. 1). P.
pilumnus generally had been considered to be a member of the Papilio
glaucus L. species group until Brower (1959) placed it with the Papilio
troilus L. group. Three factors support the placement of P. pilumnus
with the P. troilus group. The male genitalia are more like those of P.
troilus (Brower, 1959). As originally described by Schaus (1884), P.
pilumnus larvae more closely resemble P. troilus and P. palamedes than
any of the P. glaucus group species. In addition, pupal color and
morphology (Schaus, 1884) are more like P. troilus and P. palamedes .
Tyler (1975), however, suggested that P. pilumnus is transitional
between the P. glaucus and P. troilus species groups.
In our ongoing studies of the physiological and biochemical mechan-
isms of differential foodplant use in North American Papilio glaucus
and Papilio troilus species and subspecies, we have hand-paired various
taxa in more than 4000 crosses (see Scriber, 1987a, b,c). Among the most
interesting were our pairings of virgin Papilio glaucus females with
field captured P. pilumnus males. In this paper, we describe the hybrid
offspring of these pairings and their abilities to use potential foodplants.
Current address: Department of Entomology, Michigan State University, East Lansing,
MI 48824
27(2): 96-103, 1988(89)
97
Methods:
To obtain virgin females, we first collected adult females of Papilio glaucus, P.
troilus , and P. alexiares from the field. Females were allowed to oviposit
individually on acceptable foodplant leaves (black cherry and sassafras) kept
fresh by water-filled aquapics®. Females were housed in clear plastic boxes
(12cm x 20cm x 30cm) heated by a lOOw incandescent lightbulb placed
approximately 0.5m from the boxes. Eggs were removed on leaves at 2 day
intervals after oviposition and neonate larvae were subsequently reared on
excised leaves of various species of plants (in 4cm x 15cm petri dishes with
screened ventilation) to pupation. Larvae were reared in controlled environ-
ment growth chambers (at 16:8 photo/scotophase with a corresponding tempera-
ture regime of 23.5/19.5 degrees C). Foodplant leaves were kept turgid by use of
water-filled aquapics (Scriber, 1977), and changed as needed. A mixture of
healthy-looking mature (fully-expanded) leaves and younger leaves were used
for neonate studies. After weighing, pupae were individually placed in 14 cm
screen cages until adult emergence.
Hand-pairings of virgin female butterflies to field-collected males were
conducted as in Clarke & Sheppard (1956) with the pair hanging in a screen
cylinder (approximately 12cm tall by 14cm diameter) covered by the top and
bottom of a petri dish. Females that had been in copulation at least 30 minutes
were set up in the oviposition boxes as described above. Newly eclosed larvae
were individually transferred with a camel hair brush and distributed among
the various foodplants. After females died, they were dissected and examined for
spermatophores.
Results:
Males and females of Papilio pilumnus were collected by M. Evans, D.
Robacker, and W. Warfield in the states of Nuevo Leon and Tamaulipas
in northeastern Mexico and brought to the laboratory. One P. pilumnus
female produced 2 eggs, but no larvae. A second P. pilumnus female laid
5 eggs. The single larva did not survive on red bay. A third P. pilumnus
female laid 4 eggs; the single hatchling developed on sassafras. Although
a total of five hand-pairings lasted 30 minutes or longer, in three, no
spermatophore was passed (one with P. troilus , #4245, one with P.
alexiares , #3301, and one with P. glaucus, #4235). Only one pairing
with a P. glaucus (#4231) produced fertile eggs. Of a total of 108 eggs, 69
larvae eclosed, 7 died while crawling out of their eggs, 7 additional eggs
appeared to be fertile but produced no larvae, and 25 eggs appeared to be
infertile.
No neonate hybrid larvae survived on black cherry, paper birch,
quaking aspen, or sycamore (Rosaceae, Betulaceae, Salicaceae, and
Platanaceae, respectively; see Table 1). There was no indication that
the larvae initiated feeding on these hosts. However, species from the
Rutaceae (hop tree), Magnoliaceae (sweet bay and tulip tree), and
Lauraceae (spicebush, red bay, and sassafras) were accepted by the
neonate larvae and nearly 50% survived to the second instar (Table 1).
Larvae surviving to the second instar on plants other than sassafras and
98
J.Res.Lepid.
Table 1. Neonate larval survival of Fx hybrid larvae of the cross
( Papilio glaucus female) x (P. pilumnus male).
Plant
Species
Plant
Family
Surviving(n)
to second
instar
Total
set
up
%
Survival
Prunus serotina Ehrh.
(Black Cherry)
(Rosaceae)
0
27
0
Betula papyrifera Marsh.
(Paper Brich)
(Betulaceae)
0
6
0
Populus tremuloides
Michx. (Quaking Aspen)
(Salicaceae)
0
3
0
Platanus occidentalis L.
(Sycamore)
(Platanaceae)
0
3
0
Ptelea trifoliata L.
(Hop Tree)
(Rutaceae)
1
2
50
Magnolia virginiana L.
(Sweetbay)
(Magnoliaceae)
1
4
25
Liriodendron tulipifera
L. (Tulip Tree)
(Magnaliaceae)
4
8
50
Lindera benzoin (L.)
Blume (Spicebush)
(Lauraceae)
1
4
25
Persea borbonia (L.)
Spreng. (Red Bay)
(Lauraceae)
2
6
33
Sassafras albidum
(Nutt.) Nees (Sassafras)
(Lauraceae)
5
6
83
tulip tree were switched to one of these species for rearing. Of the 14
larvae surviving the feeding trials, 5 successfully pupated (Table 2). All
resulting adults were male. No attempt to determine the fertility of
these males was made.
The adult male hybrids are intermediate in wing pattern and shape
between the P. glaucus glaucus males and P. pilumnus males on both
the dorsal (Fig. la, b, c) and ventral (Fig. 2a, b, c) sides. It is also clear
that the final larval instar reflects a composite of traits from each
species. Unlike Papilio glaucus which is a solid green color (Fig. 3a), the
hybrid larva (Fig. 3b) has a yellow line running along the side of the
body with a brown ventral color and a series of blue spots on the
abdominal segments just below this line as desribed for P. pilumnus
(Fig. 3c) (Schaus, 1884; Brower, 1959), P. palamedes (Fig. 3d), and P.
27(2): 96-103, 1988(89)
99
Table 2. A summary of data for five P. glaucus x P. pilumnus larvae
which pupated.
Larval Adult
Hatch Pupation Larval Pupal Emergence Pupal
Data Larval
(Aug 1986) Food
Date
(Sept.)
Duration
(days)
wt.
(gm.)
Date (1986)
(All Males)
Duration
(days)
18
Redbay-
Tulip Tree
20
33
no. wt.
1 Oct.
11
18
Tulip Tree
10
23
0.3229
Dead as a
pharate
adult by
late Oct
— -
19
Spicebush-
Sassafras
26
38
0.4692
7 Oct
Deformed
Adult
11
19
Sassafras
12
24
0.5089
23 Sept.
Deformed
Adult
11
19
Sassafras
12
24
0.5174
23 Sept.
11
Deformed
Adult
troilus (Fig. 3e). Unlike P. palamedes , P. troilus , and P. pilumnus , the
hybrid larva has a false thoracic eyespot without a solid black center
(Fig. 3b, 3c, 3d, 3e) which closely resembles the false eyespot of P.
glaucus (Fig. 3a). The brown larval stage (before pupation) of the hybrid
larva (Fig. 4) lacks the transverse yellow at the base of the thorax which
characterizes P. glaucus (Fig. 3a). the hybrid pupae were small (pre-
sumably due to nutritional factors) and were more troilus-hke than
glaucus-kke in general shape (Fig. 5). As described for P. pilumnus
pupae (Brower, 1959), these hybrid pupae were pinkish in color and
laterally ridged, which is unlike those of all P. glaucus group species.
Discussion:
The survival and developmental compatibility in these P. glaucus/P.
pilumnus hybrid genomes were surprising, especially when compared
with other interspecific pairings we have conducted between various P.
glaucus and P. troilus species group members. Viability of the glaucus /
pilumnus hybrids was comparable with that of interspecific hybrids
100
J.Res.Lepid.
within the glaucus group but greater than that of previous glaucus
group/ troilus group hybrids. For example, the average viability (larvae/
eggs) of the two glaucus! pilumnus pairings here was 35.2% (70.4, 0),
compared to pairings of P. glaucus females with males of P. multi-
caudatus (n=10; x=44.0%) P. eurymedon (n=!8; x=30.2%), P. rutulus
(n=8; x=31.1%), andP. alexiares (n== 15; x=56.1%). Although attempted
numerous times only one successful pairing of a P. glaucus female with
a P. troilus male has ever been obtained (i.e. copulation for more than 30
minutes), and none of the eggs produced hatched. We have achieved 3
successful hand-pairings ofP. palamedes males with female P. glaucus.
One such pairing produced viable offspring. It should be pointed out,
however, that a number of factors other than genetic compatibility are
likely to be involved in determining egg viability, and considerable
caution in the interpretation of fertility data is advised (eg. see Leder-
house and Scriber, 1987).
The ability of these hybrid larvae to eat, survive, and grow on the
Lauraceae was not surprising since this is the only family the related P.
troilus and P. palamedes may actually utilize (Scriber, 1986), and
Papilio glaucus can utilize lauraceous species to a certain extent (with
the exception of red bay) (Scriber, et al. 1975; Scriber, 1973, 1984, 1986,
1987c). The ability of hybrid larvae to use Rutaceae and Magnoliaceae
may reflect the contribution of P. g. glaucus to their genome or latent
ancestral capabilities. Although it generally has been presumed that
the Lauraceae were ancestral foodplants with a key role in the evolution
of Papilio (Forbes 1932, 1958; Munroe, 1948, 1960), the Rutaceae
(Munroe and Ehrlich, 1960; Scriber, 1972; Hancock, 1983;) and the
Magnoliaceae (Dethier, 1941, Ehrlich and Raven, 1965) could be equally
important. Feeny, et. al. (1983) summarized the potential chemical
common denominators among these plant families.
Perhaps the most interesting aspect of these hybrid glaucus! pilumnus
foodplant bioassays were the plants that larvae did not successfully use
(Table 1). In particular, black cherry ( Prunus serotina ) and paper birch
{Betula papyri fera) are two of the plant species that all members of the
Papilio glaucus species group readily consume (including P. rutulus , P.
eurymedon , P. glaucus glaucus , P. g. canadensis and P. alexiares ;
Scriber, 1987b, 1987c). The hybrid larvae did not initiate feeding on
either black cherry or paper birch apparently not recognizing them as
potential hosts. Unlike the hybrid transfer of detoxication abilities
between Salicaceae feeders (P. rutulus , P. eurymedon and P. g. cana-
densis) and Magnoliaceae feeders (P. g. glaucus , P. g. australis , P.
alexiares ; Scriber, 1987a), it remains unclear if the P. glaucus abilities
to use Betulaceae and Rosaceae were transferred to the hybrid larvae
( glaucus x pilumnus) in this study. Sample sizes are low, and additional
crosses seem warranted because there is an interesting genetic story
regarding detoxication (and perhaps phylogenetic history) to be worked
out here.
Fig. 1 . Dorsal view of adult males of: a) P. g. glaucus b) hybrid P. g. glaucus
(female) x P. pilumnus (male) c) P. pilumnus
Fig. 2. Ventral view of specimens in Fig. 1
Fig. 3. Final (5th) instar of: a) P. g. glaucus, brown and green morphs, b) hybrid P.
g. glaucus x P. pilumnus c) P. pilumnus d) P. palamedes e) P. troilus
Fig. 4. Hybrid larva (P. g. glaucus x P. pilumnus) that has turned brown prior to
pupation
Fig. 5. Ventral and lateral view of the hybrid pupae (bottom) contrasted to the same
views of P. glaucus pupae (top)
102
J.Res.Lepid.
Acknowledgements . This research was supported in part by MSU (Project
#3188), the National Science Foundation (BSR 8306060 and BSR 8503464) the
USD A (85 CRCR-1-1598) and the Graduate School and College of Agricultural
and Life Sciences (Hatch 5134) of the University of Wisconsin. Travel to Mexico
was paid for by a Romnes faculty fellowship to Mark Scriber and the Ibero-
American Studies Summer Field Research Grant Program to Mark Evans, both
of the University of Wisconsin. In addition to Mark Evans, a number of
collaborators have contributed field collected specimens over the years 1981-
1986 from which our lineages have been maintained. In particular, we would
like to thank the following people for their assistance in field collections for this
study: William Bergman, Robert Dowell, Rick Lindroth, Jim Maudsley, Ric
Peigler, David Robacker, John Shuey, Frank Slansky, Jr., John Thompson and
Bill Warfield.
Literature Cited
BEUTELSPACHER, C.R. & W.H. HOWE, 1984. Mariposas de Mexico. Ediciones Cien-
tificas. La Prensa Medica Mexicana. Copilco-Universidad Delegacion
Coyoacan, Mexico, D.F. 128 pp.
BROWER, L.P., 1959. Speciation in butterflies of the Papilio glaucus group I.
Morphological relationships and hybridization. Evolution 13:40-63.
CLARKE, C.A. & P.M. SHEPPARD, 1956. Handpairing of butterflies. Lepidopt. News.
10:47-53.
dethier, V.G., 1941. Chemical factors determining the choice of foodplants by
Papilio larvae.
Amer. Nat. 75:61-73.
EHRLICH, P R. & P.H. RAVEN, 1965 Butterflies and plants: a study in co-evolution.
Evolution 18:586-608.
FEENY, P., L. ROSENBERRY, & M. CARTER, 1983. Chemical aspects of opposition
behavior in butterflies. IN: S. Ahmad, ed., Herbivorous insect: host-seeking
behavior and mechanisms. Academic Press, NY. pp.27-76.
FORBES, W.T.M., 1932. How old are the Lepidoptera? Amer. Natur. 66:452-460.
FORBES, W.T.M., 1958. Caterpillars as botanists. Proc. 10th Int. Cong. Ent. 1:313-
317.
HANCOCK, D.L., 1983. Classification of the Papilionidae (Lepidoptera): a phylo-
genetic approach. Smithersia 2:1-48.
HOWE, W.H., 1975. The butterflies of North America. Doubleday & Co. Garden
City, NY. 633 pp.
LEDERHOUSE, R.C. & J.M. SCRIBER, 1987. Ecological significance of a post mating
decline in egg viability in the tiger swallowtail. J. Lepid. Soc. 41:83-93.
MUNROE, E.G., 1948. The geographical distribution of butterflies in the West
Indies. Ph.D.thesis, Cornell University. 555 p.
MUNROE, E.G., 1960. The classification of the Papilionidae (Lepidoptera). Can.
Entomol. Suppl. 17:1-51.
MUNROE, E.G. & PR. EHRLICH, 1960. Harmonization of concepts of higher classifica-
tion of the Papilionidae. J. Lepid. Soc. 14:169-175.
SCHAUS, w. Jr., 1884. Early stages of Mexican Lepidoptera. Papilio 4:100-103.
SCOTT, J.A., 1986. The butterflies of North America: A natural history and field
guide. Stanford University Press. Stanford, CA 583 pp.
27(2): 96-103, 1988(89)
103
SCRIBER, J.M., 1972. Confirmation of a disputed foodplant of Papilio glaucus
(Papilionidae). J. Lepid. Soc. 26:235-236.
SCRIBER, J.M., 1973. Latitudinal gradients in larval feeding specialization of the
world Papilionidae (Lepidoptera). Psyche 80:355-373.
SCRIBER, J.M., 1977. Limiting effects of low leaf-water content on the nitrogen
utilization, energy budget, and larval growth of Hyalophora cecropia (Lepi-
doptera: Saturniidae). Oecologia 28:269-287.
SCRIBER, J.M., 1984. Larval foodplant utilization by the world Papilionidae
(Lep.): latitudinal gradients reappraised. Tokurana (Acta Rhopalocero-
logica) 6/7:1-50.
SCRIBER, J.M., 1986. Origins of regional feeding abilities in the tiger swallowtail
butterfly: ecological monophagy and the Papilio glaucus australis subspecies
in Florida. Oecologia 71:94-103.
SCRIBER, J.M., 1987a. Allelochemicals and Alimentary Ecology: heterosis in a
hybrid zone? pp.43-71 In (L. Brattsten and S. Ahmad, ed.) Molecular
Mechanisms in Insect Plant Associations. Plenum Press, NY.
SCRIBER, J.M., 1987b. Tale of the tiger: Biogeography, bionomial classification,
and breakfast choices in the Papilio glaucus complex of butterflies. IN:
Chemical Mediation of Coevolution (K.C. Spencer, ed.) (in press).
SCRIBER, J.M., 1987c. Population genetics and foodplant use in the North
American tree-feeding Papilionidae. pp. 221-230 In Proceedings of the 6th
Inter. Symp. on Insect. Plant Relationships. W. Junk, Dordrecht., Nether-
lands.
SCRIBER, J.M., R.c. LEDERHOUSE, & L. CONTARDO, 1975. Spicebush, Lindera benzoin
(L.), a little known foodplant of Papilio glaucus (Papilionidae) J. Lepid. Soc.
29:10-14.
TYLER, H., 1975. The swallowtail butterflies of North America. Naturegraph.
Healdsburg, CA 192 pp.
Journal of Research on the Lepidoptera
27(2): 104-108, 1988(89)
New Host Records and Morphological Notes on Four
Tortricines (Tortricidae)
Sherri Sandberg
and
Steven Passoa
Department of Entomology, University of Illinois at Urbana-Champaign
Hypericum perforatum (Guttiferae), St. John’s- wort or Klamath
weed, a plant of European origin, has been introduced into many regions
of the world, including rangelands of the United States, Canada, South
Africa, and Australia (Harris and Peschkin, 1974; Giese, 1980). St.
John’s-wort is considered a rangeland weed because it produces the
phototoxic compound hypericin, a blister-inducing agent for livestock in
the presence of sunlight (Blum, 1941). Because there are few published
records of Lepidoptera feeding on Hypericum in North America (King-
solver et al., 1984), we now report on four native North American
tortricids reared from two species of Hypericum.
Larvae of Platynota flavedana Clemens, Choristoneura parallela
(Robinson), Sparganothis sulfureana (Clemens), and Xenotemna pal -
lorana (Robinson) were found in leaf ties on H. perforatum at several
Illinois localities; the latter two species were also found at one site in
Michigan. In addition, the last three species were collected in Illinois
from H. sphaerocarpum , a native North American species that does not
contain hypericin. For each tortricid species, as is applicable, H.
perforatum andiif. sphaerocarpum represent new host records (MacKay,
1962; Chapman and Lienk, 1971), although S. sulfureana has been
reared from an undetermined species of Hypericum (Godfrey et al., in
press). All four species are polyphagous feeders (see Table 1) with the
majority of previous host records on agricultural crops (Chapman and
Lienk, 1971). Identification of field-collected larvae was based on
individuals reared to adult. For each species of tortricid, the hosts and
collection data are provided, accompanied by morphological notes on the
immature stages to supplement the existing keys in Chapman and
Lienk (1971), MacKay (1962), and Mosher (1916).
Platynota flavedana is a pest on strawberry (Wilde and Semel, 1966).
Larvae of P. flavedana were collected on H. perforatum from the end of
June through August 1985 along roadsides of several Illinois localities:
near Monticello (Piatt Co.), Mount Vernon (Jefferson Co.), Carbondale
(Jackson Co.), and Marion (Williamson Co.). P. flavedana was common
on Hypericum in 1985 but was not found in 1986.
Sparganothis sulfureana is recorded from a wide variety of plants,
27(2): 104-108, 1988(89)
105
Table 1. Host records of 4 species of Tortricidae larvae collected and
reared on Hypericum perforatum.
SPECIES
HOST RECORDS
Choristoneura parallela
Compositae, Ericaceae, Guttiferae*,
Leguminosae, Myricaceae, Rosaceae,
Rubiaceae, Rutaceae
Platynota flavedana
Aceraceae, Begoniaceae, Compositae,
Ericaceae, Guttiferae*, Leguminosae,
Malvaceae, Rosaceae
Sparganothis sulfureana
Ericaceae, Gramineae, Guttiferae*,
Leguminosae, Pinaceae, Ranunculaceae,
Rosaceae, Salicaceae, Umbelliferae,
Verbenaceae
Xenotemna pallorana
Caryophyllaceae, Compositae, Guttiferae*,
Leguminosae, Pinaceae, Rosaceae,
Verbenaceae
* Represents a new host record
References: Beckwith, 1938; Chapman and Leinke, 1971; Deitz et al.,
1976; Freeman, 1958; MacKay, 1962; Martin, 1958; Newcomer and
Carlsen 1952; Wilde and Semel, 1966.
including many cultivated species. Considered a pest on cranberry, S.
sulfureana is commonly known as false yellowhead or sulfur leafroller
(Beckwith, 1938; Chapman and Lienk 1971). Larvae of S. sulfureana
were commonly collected on H. perforatum from late June through
August 1985 and 1986, at the same Illinois locations previously men-
tioned for P. flavedana. In addition, larvae of S. sulfureana were also
collected from H. perforatum at the University of Michigan Biological
Station (near Pellston, Michigan) in July 1985 and from H. sphaero -
carpum near Monticello and near Forrest, Illinois (Livingston Co.) in
July 1985 and 1986.
Xenotemna pallorana is a minor pest on young pines (Martin, 1958),
young apple and other fruit trees (Newcomer and Carlson, 1952). While
common, larvae were found only in July of 1985 and 1986 feeding on H.
perforatum at the same Illinois localities previously mentioned and
at the University of Michigan Biological Station. Larvae were also
collected from H. sphaerocarpum near Monticello, Illinois in July 1986.
Choristoneura parallela also has a wide host range. Commonly known
as the spotted fire worm, C. parallela is considered a pest on cranberries.
The larvae were collected on both species of Hypericum near Monticello,
106
J. Res. Lepid.
Illinois during July, 1986. They were found frequently on H. sphaero-
carpum but only once on H. perforatum at this site. Larvae were also
collected in July at a site east of Urbana, Illinois (Champaign Co.) on H.
sphaeroearpum.
Although the caterpillars of X. pallor ana and C. parallela are rela-
tively easy to recognize compared to other Hypericum- feeding tortricids
(see MacKay 1962 for descriptions), larvae of P. flauedana and S.
sulfureana can easily be confused in the field. Chapman and Lienk
(1971) illustrated the larva of S. sulfureana in color and distinguished it
from P. flavedana by the presence of a thin black line along the lateral
margins of the prothoracic shield. However, some P. flavedana may also
have this black line. A more consistent field character involves the
thoracic and abdominal pinacula of the dorsal setae. Most dorsal
pinacula in Platynota spp. are elongated lengthwise whereas in S.
sulfureana all the pinacula are round (Chapman and Lienk 1971;
MacKay, 1962). In addition, Chapman and Lienk (1971) correctly noted
that S. sulfureana may be separated from P. flavedana by the dark
dorsum contrasting with the paler ventral region. In P. flavedana the
dorsal area is concolorous with the rest of the body. MacKay (1962)
distinguished P. flavedana from other Platynota spp. by its clear
brownish-yellow head, prothoracic shield, and prothoracic pinacula.
However, this distinctive coloration is only found on the last two instars.
Younger larva have a black prothoracic shield and head (Wilde and
Semel, 1966) and thus, cannot be identified using the above characters.
Mosher (1916) separated the pupa ofP. flavedana from S. sulfureana
by the presence of flattened cremaster setae and the absence of a row of
spines on the second abdominal segment in the female. Some more
obvious morphological differences between these species (that may
prove to be a useful tool in the systematics of tortricines in general)
involves variation in the shape of the vertex and the presence of
maxillary palpi. Platynota flavedana has maxillary palpi and a round
vertex which lacks a ridge (Fig. 1). In contrast, the pupa of S. sulfureana
Fig. 1-4. Ventral view of four tortricine pupae (30x). 1. P. flavedana 2. S.
sulfureana ; 3. C. parallela ; 4. X. pallorana .
27(2): 104-108, 1988(89)
107
lacks maxillary palpi and has a ridge which extends cephalad from the
frons to the epicranial suture (Fig. 2). A similar ridge is found in C.
parallela (and other Choristoneura spp.) but it runs between the
antennal scapes (Fig. 3). X. pallor ana was not included in Mosher’s
(1918) key but the characteristic vertex (Fig. 4) readily distinguishes
this species from other Hypericum- feeding tortricids in Illinois.
It is of interest that four native generalist tortricid species have been
found commonly feeding on an introduced plant notorious for con-
taining a phototoxin. Although this occurrence seems to run counter to
current ideas on insect-plant interactions, i.e., specialists are adapted to
feed on plants with defensive chemicals whereas generalists are
deterred by them (Janzen, 1979), the larval leaf-tying habits of all these
species may shade them from the phototoxic effects of hypericin and
thus preadapt them for feeding on phototoxic plants (Sandberg and
Berenbaum in prep.).
Acknowledgements . We thank R. Brown, J. Powell, and G. Godfrey for identifi-
cation of reared adults, P. Adams for plant identification, and M. Berenbaum, G.
Godfrey, J. Neal, J. Nitao, and J. Sternburg for valuable comments on the
manuscript. This work was supported by National Science Foundation Grant
BSR 835-1407 to M. Berenbaum.
Literature Cited
BECKWITH, C. S., 1938. Sparganothis sulfureana Clem., a cranberry pest in New
Jersey. J. Econ. Entomol. 31(2):253-256.
BLUM, H. F. 1941. Photodynamic action and diseases caused by light. Am. Chem.
Soc. Mon. Ser. 85. Reinhold, Pub. Co. New York, 309 pp.
CHAPMAN, P. J. & s. E. LIENK, 1971. Tortricid fauna of apple in New York. New
York State Agr. Exp. Sta. Spec. Publ. 122 pp.
DEITZ, L. L., J. W. Van DUYN, J. R. BRADLEY, JR., R. L. RABB, W. M. BROOKS, R. E. STINNER,
1976. A guide to the identification and biology of soybean arthropods in
North Carolina. North Carolina Agr. Exper. Sta. Tech. Bull. No. 238. 264 pp.
FREEMAN, T. N., 1958. The Archipinae of North America (Lepidoptera: Tortri-
cidae). Can. Entomol. Suppl. 7. 89 pp.
GIESE, A. C., 1980. Hyper ici sin. Photochem. Photobiol. Rev. 5:229-255.
GODFREY, G. L., E. D. CASHATT, & M. O. GLENN, 1987. Microlepidoptera from the
Sandy Creek and Illinois River region: an annotated checklist of the
suborders Dacnonypha, Monotrysia, and Ditrysia (in part) (Insecta). Illinois
Nat. Hist. Surv. Publ. 7. (in press).
HARRIS, P. & D. P. peschkin, 1974. Biological control of St. John’s- wort. Can.
Agric. 19(1):13-15.
JANZEN, D. H. 1979, in Rosenthal, G. A. and D. H. Janzen (eds.). Herbivores. Their
interaction with secondary plant metabolites. Academic Press, New York.
718 pp.
KINGSOLVER, J. M., s. w. T. batra, J. A. UTMAR, 1984. A selected bibliography of
insect-vascular plant associational studies. U. S. Dept. Agric. Biblio. Lit.
Agric. No. 27. 229 pp.
108
J.Res.Lepid.
MACKAY, M. R., 1962. Larvae of the North American Tortricinae (Lepidoptera:
Tortricidae). Can. Entomol. Suppl. 28:1-182.
MARTIN, J. L., 1958. Observations on the biology of certain tortricids in young
coniferous plantations in southern Ontario. Can. Entomol. 90:44-53.
MOSHER, E. 1916. A classification of Lepidoptera based on characters of the pupa.
Bull. Illinois State Lab. Nat. Hist. 12(2):1-166.
NEWCOMER E.J. & CARLSEN, F.W. 1952. The leaf-roller moth Pandemis pyrusana J.
Econ. Entomol. 45(6): 1079-1081.
WILDE, G. & M. SEMEL, 1966. The life history of Platynota flavedana, a leaf roller of
strawberry. J. Econ. Entomol. 59(5): 1037-1041.
>
Journal ofResearch on the Lepidoptera
27(2): 109-114, 1988(89)
Notes on the biology of three Riodinine species:
Nymphidium lisimon attenuatum , Phaenochitonia
sagaris satnius , and Metacharis ptolomaeus (Lycaenidae:
Riodininae)
Curtis J. Callaghan
Rua Yeddo Fiuza 595, Petropolis, Rio de Janeiro, Brazil
Abstract. Observations are presented on the immature biology of three
riodinine species from southeast Brazil: Nymphidium lisimon attenua-
tum, Phaenochitonia sagaris satnius, and Metacharis ptolomaeus. N.
lisimon attenuatum was found to be myrmecophilous while P. sagaris
satnius larvae inhabit rolled leaves. Based on observations of oviposi-
tion behaviour, I suggest that M. ptolomaeus larvae are solitary and
non myrmecophilous.
The purpose of this paper is to present data on the biology of three
riodinine species from southeast Brazil; Nymphidium lisimon attenua-
tum Stichel, Phaenochitonia sagaris satnius , (Dalman) and Metacharis
ptolomaeus (Fabricius). Although these species are not uncommon
where found, nothing about their immature biologies has been pub-
lished to date.
Observations on the first two species were made at Fazenda Uniao, a
forest reserve belonging to the Brazilian National Railways at km 140
of the BR101 highway, Rio de Janeiro State. The vegetation is typical of
Atlantic tropical lowland forest found in the foothills of the Serra do Mar
at about 100 m above sea level. The reserve consists of patches of
secondary alternating with areas of primary forest, (fig. 1. M. ptolo-
maeus was recorded from a patch of woods near Barra de Sao Joao, an
area of transition between the restinga vegetation and the Atlantic
forest, described elsewhere. (Callaghan, 1985).
Observations on larval behaviour were made in the field and in the
laboratory.
In the following sections, each species is considered separately, with a
description of the immature stages followed by a discussion on imma-
ture biology.
Nymphidium lisimon attenuatum
Immature stages
Third (?) instar larve (fig. 3): Head light brown. Head, thorax and
abdomen covered with short setae. First thoracic segment (Tl) with
110
J. Res. Lepid.
Fig. 1 . Study area at Fazenda Uniao.
Fig. 2.
Food plant of N. lisimon attenuatum, Inga sp.
dark brown dorsal shield and numerous long setae extending cephalad;
one lateral spiracle at base of the shield, and two vibratory papillae
dorsad beneath edge of the shield. Meso- and meta-thoracic segments
(T2, T3) light brown-green mottled. Abdominal segments also light
mottled brown-green with a light green irregular band dorsad; spiracles
on A1 and A3-A7 ventrally positioned, that on A2 laterally and A8
dorsad and cephalad of the Newcomber’s organs. Segments A9 and A10
covered by a dorsal shield with numerous setae around the edge. Head
capsul 1 mm; length 10 mm. N = 4.
27(2): 109-114, 1988(89)
111
Fig. 3. Third(?) instar larvae of N. lisimon attenuatum with ants.
Discussion
Stichel’s subspecies attenuatum ranges along the coast in southeast
Brazil from Santa Catharina north to southern Bahia. Inland it inter-
grades with subspecies epiplatea Butler. N. 1. attenuatum is found
locally in disturbed forest habitats where the males perch in the
afternoon along roads and woods edges. They rest under leaves with the
body raised 45 degrees from the leaf surface.
The foodplant at Fazenda Uniao was Inga sp. (fig. 2), the same genus
associated with my other observations of Nymphidium biology. (Cal-
laghan, in prep.) The plant has broad pointed leaves with nectaries at
the base and grows commonly in open clearings to a height of 2 meters.
The Nymphidium larvae feed on the newer growth and at nectaries,
instead of older, tougher leaves. The larvae are solitary, feeding on
separate leaves, a characteristic of other myrmecophilous species.
(Callaghan, 1985) In the laboratory the larvae fed at night, remaining
motionless on the foodplant during the day.
In the field the larvae were always associated with tiny ants identified
as Wasmannia aropunctata (Roger, 1863). These gather in large
numbers on and around the larva, effectively hiding it from view, thus
apparently affording it some protection against predation. Unlike other
ants observed with larvae (i.e. Campanotus sp., (Callaghan, 1977),
Wasmannia aropunctata appear very sluggish, not taking an obvious
defensive attitude towards intruders. The consequence of this lack of
aggressive ant protection was suggested by all 4 collected larvae being
found parasitized by ichneumonoid wasps. (Hymenoptera: Tricho-
grammatidae).
Close observation of larval behaviour with ants indicated that the
Newcombers’ organs were eversible, protruding outward during the
secretion of honeydew. This physiology is similar to that observed in
1 Menander felsina larvae. (Callaghan, 1977) No eversible tubercles were
observed, such as those found on Audre larvae, (pers. obs), nor were the
vibratory papillae observed functioning.
112
J.Res.Lepid.
Fig. 4. Food plant of P. sagaris satnius fam. Melastomataceae.
Phaenochitonia sagaris satnius
Immature stages
Egg: Color white; shaped like a fat tire, diameter 0.8 mm, height 0.4
mm; micropyle 0.2 mm in diameter with many small perforations;
covering egg surface is a network of small ridges forming hexagonal
patterns with a small protrusion at each intersection. Duration: 12
days. N = 5.
First instar larva: Color uniform light green, except for head which is
slightly darker; larve pubescent with two rows of long black dorsal setae
on segments T2 to A7; four long, black setae on first thoracic (Tl)
segment pointing cephalad, long green lateral setae with one black one
found on all thoracic and abdominal segments; spiracles observed
lateraldorsad on segments A2 through A8. Segments A9, A 10 partially
covered by a small tail plate. Length 1.7 mm; head capsul 0.3 mm;
duration: 9 days.
Second Instar (Fig. 6): Head light brown, face with many short setae.
Thorax and abdomen light green with numerous small white dots; Tl
with eight long black setae pointing cephalad; segments T2 to A9 with 1
black and 5 long white lateral setae on each side per segment and 2
black dorsal setae; A 10 with 4 long black setae around the edge of the
tailplate. Length 2.5 mm, head capsul 0.4 mm; duration 8 days. N = 4.
Third Instar: Color and morphology as in second instar, except
spiracles outlined in lighter green. Length 3.5 mm; head capsul 0.6 mm;
duration: larvae died after 5 days due to unknown causes. N = 4.
Discussion
P. sagaris satnius is the central Brazilian subspecies ranging from the
coast of Sao Paulo north to Bahia then across the Planalto to Mato
Grosso. Northward it intergrades with subspecies iasis (Godman) and to
the south with subspecies phrygiana Stichel.
27(2): 109-114,1988(89)
113
Fig. 5. Ovam of P. sagaris satnius
inside rolled leaf, part of which
has been cut away.
Fig. 6. P. sagaris satnius larvae
feeding. Note frass chain.
At Fazenda Uniao a lone female was observed ovipositing about 1500
hours on a shrub identified as belonging to the family Melastomataceae.
(fig. 4) She alighted on the rolled leaf tube of an unidentified Heterocera
larva, walked to the open end and placed a small cluster of five eggs
inside the opening, (fig. 5)
Upon hatching, the larvae moved into a folded foodplant leaf provided
for them in the laboratory and proceeded to eat the inner side of the leaf,
at the same time attaching the upper and lower leaf halves together
with silk. During the second and third instars when the larvae were
placed on fresh foodplant, they would fold the leaf over by weaving
silken threads across the leaf surface, each slightly tauter than the one
placed previously until the increased tension slowly drew the halves of
the leaf together. These were then secured by numerous filaments
between the upper and lower halves, forming chambers inside the folded
leaf. The larvae always remained inside, even when feeding, undoubt-
edly being thus protected from predation.
The larvae always remained together while feeding, lining up side
by side in twos or threes. Frass excreted by the larvae stuck together,
forming a long chain behind each individual, (fig. 6) Starting two to
three days before molting the larvae would cease feeding until a day
after molting. At no time was there any evidence of myrmecophilous
organs or any behaviour patterns which would suggest their association
with ants.
Metacharis ptolomaeus
Immature stages
Egg: Shiny bronze color, 0.5 mm in diameter, 0.2 mm high. Extend-
ing from micropile is a network of raised lines forming hexagonal
114
J.Res.Lepid.
figures with a protrusion rising at each intersection. Duration: 11 days.
N = 1
First instar larva: Newly hatched larva light green, nearly white;
pubscent with four long setae extending cephalad from the edge of the
prothorax and six equally long setae extending to the rear from the
anal plate. Dorsad two rows of setae, a pair to a segment, from T2 to A7,
and numerous additional setae extending laterally from the lower edge
of each segment. Length 1 mm; head capsule 0.13 mm.
Discussion
M. ptolomaeus inhabits the coast and adjacent mountains in south-
east Brazil. It is particularly common in coastal woods and “restinga”
vegetation.
A single female was observed ovipositing in a small woods near Barra
de Sao Joao, Rio de Janeiro State. She laid a single egg at the base of a
petiole of a leaf on a small tree identified as Lacistema sp. (Flacour-
tiacea). As no ant species normally associated with myrmecophilous
riodinine species were found on the foodplant, and only a single egg was
laid, this would suggest that the larvae of Metacharis ptolomaeus are
solitary but not myrmecophilous. Unfortunately, the larva died before
fresh foodplant could be obtained.
Acknowledgements. I wish to thank Dr. Carlos Monteiro of the Universidade
Federal (Fundao) for obtaining identification of the foodplants and Dr. Woodruff
Benson of the Universidade Estadual de Campinas for determining the ant
species. Finally, my thanks to the Brazilian National Railways for permission to
collect at the Fazenda Uniao.
Literature Cited
CALLAGHAN, C. J., 1977. Studies on restinga butterflies I. Life cycle and immature
biology of Menander felsina, (Riodinidae), a myrmecophilous metalmark. J.
Lep. Soc. 31(3):173-182.
, 1982. Notes on the immature biology of two myrmecophilous Lycaen-
ida e.Juditha molpe (Riodininae) and Panthiades hitias (Lycaeninae). J. Res.
Lep. 20(l):366-42.
— , 1985(86) Notes on the biology of Stalachtis susanna (Lycaenidae:
Riodininae) with a discussion of riodinine larval strategies. J. Res. Lep.
24(3):258-263.
Journal of Research on the Lepidoptera
27(2): 115-119, 1988(89)
Portable apparatus for photographing genitalic
dissections
Tim L. McCabe
Biological Survey, New York State Museum, State Education Department, Albany,
New York 12230
Introduction
Entomologist’s and Lepidopterist’s are well aware of the value of
genitalic dissections for identifications and comparative morphology.
Dissections suitable for photography are time-consuming to produce.
Frequently, dozens of slides have to be prepared of a single species to
fully understand the range of variation. Every slide ever prepared
becomes worth seeing when one is dealing with a problematical species.
In addition, species are frequently known from a single specimen which
must be borrowed and returned or examined while visiting a museum.
Loan institutions are frequently equipped to provide photographs of
needed dissections, but this can be a burden on already understaffed
collections. The need for quality photographs of dissections is greater
than ever.
In recent years, cameras have seen a revolution in sophistication. Of
particular note to technical photographers is OTF (off-the-film) light
metering and automated flash exposures. This eliminates the chore of
calculating flash distances and taking multiple exposures at various
F-stops in an attempt to get a properly exposed picture. Lenses made for
macrophotography have also improved and dropped in price. Virtually
any semitranslucent slide-mounted subject (mouthparts, wings, fleas,
etc.) can be photographed by the illustrated set-up (Figs. 1 & 2).
Component parts total less than $1,000. A commercially available
apparatus would cost over $11,000 (for the Wild M420 Makroskop with
the MPS 45/51 Automat, Polaroid CB 101 back and necessary lenses).
Component parts
The apparatus described here (Figures 1 & 2) is comprised of (from left
to right) 1) an Olympus Varimagni Finder; 2) the OM2n by Olympus; 3)
flash-cable coupler with cable attached; 4) self winder; 5) bidirectional
monorail from Spiratone; 6) Olympus Telescopic Auto Tube 65-116;
7) objective lens mount (PM-MTob); 8) Zuiko 38 mm Macro F 3.5;
9) salvaged microscope base; 10) opal glass; 11) Olympus T32 flash with
blue filter (Electronic Flash Color Filter Set T32 — - equivalent to Kodak
Wratten 44). The rails are mounted on a board that can be clamped onto
the edge of a table. A useful accessory not depicted is an AC adapter that
116
J.Res.Lepid.
Figs. 1 & 2. Two views of the photomicrographic set-up.
plugs into the flash eliminating the need for A A batteries. Olympus has
removable focusing screens, and a microscope focusing screen (clear
field type 1-12) is necessary as the macro lenses require so much light as
to make a diffuse focusing screen appear black. The Varimagni Finder
can be independently adjusted to accomodate vision defects such as far
27(2): 115-119, 1988(89)
117
Figs. 3-6. Photographs of the same male genitalia slide of Discestra farnhami
(Lepidoptera: Noctuidae): 3) Tech Pan, shot at ASA 100; 4) Panato-
mic X, shot at ASA 64; 5) Ilford Pan F, shot at ASA 100; 6) T-Max
100, shot at ASA 320.
or near-sightedness. The T32 flash is not mounted on the board, but
merely supported on a box. The blue filter is placed over the flash to
mask the amber color of the Canada balsam commonly used to make the
specimen mount. Hardwick (1950; Preparation of slide mounts of
lepidopterous genitalia. Can. Entomol. 82:231-235) describes a suitable
Lepidoptera genitalia mounting technique. The blue filter makes only a
minimal improvement in the resulting picture and the flash can be used
without the color filter with only a slight loss in contrast. Of course, the
camera can be used without the self-winder. The flash is rested 3-4
inches behind the frosted glass but can be moved closer if full magnifi-
cation is used on a very thick slide mount. The subject should be at least
a quarter inch in front of the opal glass to prevent features of the glass
from appearing on the negative. From center to center, the monorail is
mounted 7.5 inches from the slide stage to accomodate the entire
spectrum for focusing with both the 20 and 38 mm macro lenses. A
118
J. Res. Lepid.
38 mm Zuiko MC macro lens is illustrated and I recommend the Zuiko
20 mm MC macro lens for greater magnification. Optimum resolution
for the 38 mm lens lies in the 2-6x range; that of the 20 mm lens is
4-12x. This allows full-frame pictures of subjects ranging in width from
2 to 20 mm (40 mm possible by use without extensions. The Zuiko macro
lens illustrated is a manual lens. It is now available in automatic (which
I recommend). Note, however, that the manual lens uses an adapter (the
PM-MTob) which is a universal microscope mount allowing use of
compound microscope lenses with this set-up.
Microlepidopterists will frequently have need of even greater magni-
fication than the 20 mm macro offers. Compound microscope lenses that
lack an iris have poor depth-of-field capabilities and specialty lenses are
required for high magnification. However, an iris attachment can be
added to an ordinary microscope lens to greatly enhance depth of focus.
Tips for the best shots
The Olympus Varimagni Finder has a switch allowing one to view at
1.2x or 2.5x. The greater magnification gives better critical focus. If
prints appear out of focus, remember that the Varimagni Finder has a
focusing ring that must be set for each person without eye-glasses using
the same eye each time. I made a white mark to align the focusing ring
for my own use after establishing my own critical focusing point.
Stop the macro lens all the way down (fl6) for the best depth of field.
No loss in resolution was noted at this setting. A shutter release cable
(not illustrated) will help prevent vibration during exposure.
Each lens has its own peculiar effect on the camera’s ability to
autoexpose. I find the best negatives and prints are produced by
adjusting the film speed (in the case of Panatomic X, Ilford Pan F, and
Tech Pan) one F-stop faster (+1 on the Olympus ASA ring) with the
Zuiko 38 mm macro (Figs. 3-5). T-Max 100 was pushed to ASA 320 to
obtain the least contrasty print (Fig. 6). If prints appear grainy, it is
undoubtedly because of the film. Clean dissections (dust-free surfaces,
preparations with minimal debris in mounting medium) are a must,
especially for the Lepidoptera genitalia illustrated.
Films
Kodak will soon be replacing its Panatomic X with T-Max 100 (a
faster fine-grain film with better tonal range). T-Max 100 sensitivity to
the blue filter is 1% stops more sensitive than Pan X. This means the T-
Max 100, which has an ASA of 100, will have to be pushed to ASA 320 to
obtain the desired low-contrast negative.
Fine-grained films tested include Tech Pan, Ilford Pan F, T-Max 100,
and Panatomic X. Tech Pan’s ASA is variable according to development
(Vetter, J. P. 1984. In Richard A. Morton, Ed., Photography for the
Scientist, second edit, Academic Press, 393-456), but example given was
27(2): 115-119, 1988(89)
119
taken at ASA 50 to optimize low contrast. Ilford, like T-Max, was
affected by the blue filter and will have to be pushed to produce a low
contrast negative. Agfa Pan was not tested. Tech Pan had the finest
grain (320 lines per mm versus 280 lines per mm in T-Max 100), the
lowest contrast, and was the most versatile. The next grade of films,
Plus X and others, was much too grainy to be used for this type of
photomicrography
Development
The examples (Figs. 3-6) were developed by the following process
(only the developing time varies) 1) Kodak D-76 straight (68°F) for
5 minutes for Panatomic X and Ilford Pan F (8 minutes for Tech Pan and
9 minutes for T-Max 100) (5 seconds agitation every 30 seconds); 2) Stop
Bath for 30 seconds (continuous agitation); 3) Kodak F-6 Fixer for
5 minutes (continuous agitation); 4) wash, 1 minute (two changes of tap
water); 5) Perma Wash, 1 minute (continuous agitation); 6) wash,
1 minute (two changes of tap water); 7) Photo-flo, 30 seconds; 8) dry.
Acknowledgements. I thank Christopher Supkis for the pictures of the ap-
paratus and for technical assistance. Mention of brand name is for reference
only and does not constitute endorsement of products. Contribution No. 535 of
the New York State Science Service.
Literature Cited
HARDWICK, D. F., 1950. Preparation of slide mounts of lepidopterous genitalia.
Can. Entomol. 82:231-235.
VETTER, J. P., 1984. In Richard A. Morton, Ed., Photography for the Scientist,
second edit, Academic Press, 393-456.
Journal of Research on the Lepidoptera
27(2): 120-128, 1988(89)
Census of the Butterflies of the National Audubon
Society’s Appleton-Whittell Research Ranch, Elgin,
Arizona
Richard A. Bailowitz
2774 West Calle Norado, Tucson, Arizona 85721, U.S.A.
Abstract. The surprisingly rich butterfly fauna of the Audubon
Society’s Appleton-Whittell Research Ranch is censused and an
annotated checklist presented. One hundred and three species are
included.
Introduction
The purpose of this study was to inventory and census the butterfly
fauna of the Audubon Society’s Appleton-Whittell Research Ranch. The
Ranch lies approximately 15 km southeast of Elgin in Santa Cruz
County, Arizona. The 3170 ha of the Research Ranch were set aside in
1968 by the Appleton family and later acquired by the National
Audubon Society. The Ranch serves as a sanctuary for indigenous
plants and animals and as a site for non-destructive ecological research.
In the 16 years since its founding a great deal of research has been
carried out on plants, vertebrates, and the abiotic environment; but
little has focused on the invertebrates.
The Research Ranch is comprised of patented (i.e. private), federal
(U.S. Forest Service), and State of Arizona parcels. Most of the land is
rolling grassland and oak savannah through which small fingers of
riparian and pinyon-juniper woodlands extend. The sections of nearly
pure grassland in the northern half of the Ranch produced fewer species,
and they yielded only a few specialties (species with restricted ranges)
not found elsewhere on the property (e.g. Hesperia uncas lasus ). The
deeper canyons toward the center of the Ranch— Post Canyon, Turkey
Creek, O’Donnell Creek, etc. — possessed the greatest lepidopteran
diversity. This diversity was coupled with a modicum of specialties (e.g.
Amblyscirtes texanae , Adopaeoides prittwitzi). The extreme southern
edge of the Ranch, especially Lyle Canyon and vicinity, had both high
diversity and numbers of specialties (e.g. Yvretta cams, Cyllopsis
henshawi). Proximity to the Huachuca Mtns. is responsible, at least in
part, for this.
Methods
A total of 48 visits were made to the Ranch covering 224 hours.
27(2): 120-128, 1988(89)
121
Censusing began in August 1982 and ended January 1984. Most areas
were explored on foot during the censuses and some were singled out for
special attention. These were: Lyle and Post Canyons, Turkey and
O’Donnell Creeks, the higher ridges between these drainages, Finley
and Telles Tanks, and the Headquarters area.
Records were organized into ten-day periods. At least one visit was
made during each ten-day period for the duration of the study, except for
the months of December, January, and February when suitable days
(clear, calm, temperatures over 15°C) were uncommon. Numbers of
individuals during all visits were tallied for four-hour periods to obtain
relative abundances. These were categorized as abundant- A (> 100
per-hour period), common-C (between 13 and 99 per 4-hour period),
uncommon-U (between 3 and 12 per 4-hour period), rare-R (1 or 2 per
4-hour period), and single specimen-S (1 per 10-day period. An attempt
to tie specific habitats to species occurrence was abandoned because
most species have different needs at different times and, depending
upon nectaring, water utilization, egg laying, mate location, etc., are
found at a number of locations during the lifespan of any given brood or
even on any given day. However, certain general distribution patterns
were noted.
A reference collection is housed at the Research Ranch Headquarters.
An attempt was made to secure a pair of each species (either male and
female or a dorsal and ventral view), although this was not always
possible. At least one specimen of each “resident” species is represented
in the collection. A “resident” species is defined as one which is known or
strongly suspected to breed on the Ranch (92 species, 89.3% of total).
Included in this category was Hylephila phyleus , which probably does
not survive winters on the Ranch itself but winters at nearby areas of
lower elevation. Nine species (8.7% of total) were designated as “influx”
species. These are species of regular occurrence seasonally, but they are
unable to survive winters at the Ranch or even in most areas of
southeastern Arizona. Eight “influx” species are present in the collec-
tion: Polygonus leo, Urbanus dorantes, Kricogonia lyside, Eurema
proterpia, E. boisduvaliana, Phoebis agarithe, P. sennae, and Euptoieta
hegesia. One species, Anteos clorinde, was seen but could not be netted.
The final category, which might be considered a subset of “influx”, is the
“vagrant”. “Vagrant” species do not regularly occur in the area or even
in the state but rarely find their way here. They most often appear
during and after the chubasco (strong mid and late summer rainy
season) season. Two species ( 1.9% )- -M arpesia petreus and Papilio
astyalus— are best classified as “vagrant”.
The following annotated checklist mostly follows Miller and Brown
(1981), both in sequence and systematics. However, in several cases, for
example in generic designations, the older usages of Howe (1975) are
preferred.
122
J.Res.Lepid.
Discussion
The rich diversity of Lepidoptera at the Ranch, 103 species, is affected
by a number of factors. The weather is of primary import, especially
winter temperatures and precipitation, prevailing winds, humidity
levels, and, especially, summer rainfall. For some of these variables the
data were remarkably constant between the two winters of the study
(similar winter rainfall totals and low winter temperatures for the two
years: 1981-82 vs. 1982-83). The greatest variation occurred during the
critical period of the summer chubascos. The onset of these rains was
later in 1982 (mid- July) than in 1983 (late June). The total rainfall for
the chubasco period— July, August, and September— in 1982 was 20.3
cm, approximately half that for the 1983 chubasco of 39.4 cm. Under a
regime of wet weather in Arizona and northern Sonora, Mexico, the
vegetation can be conductive to sizeable influxes of primarily tropical
species. These conditions existed during 1983 and likely contributed to
the rich Lepidoptera activity during that season.
The author’s familiarity with local land contours, drainages, plant
associations, nectar sources, watering holes, etc. also increased the day-
to-day success in the field, accounting for some of the disparity between
the two seasons. For example, on August 18th, 29 species were found in
1982 and 49 in 1983.
The late rainy season in August and September usually produced the
greatest concentrations of individuals and the highest numbers of
species (Fig. 1) for the year. During this study, the highest species count
for a single trip was made on 2 August 1983 when 51 species were
recorded. In fact, the four trips made between 27 July and 3 September
in that year produced the four highest counts, all yielding 49 or more
species per day. A secondary high was recorded in May 1983 following
Temporal Distribution
Average Number
Total Number
Species
species
Month
Per Visit
Per Month
January
3.00
3
February
1.33
4
March
14.50
21
April
22.33
35
May
34.25
55
June
31.75
47
July
37.50
59
August
37.83
67
September
37.83
67
October
33.42
57
November
26.50
41
December
3.66
9
Total
—
103
Figure 1.
27(2): 120-128, 1988(89)
123
an unusual extended wet period. Under more normal conditions, the
May numbers of individuals and species probably would have been more
in line with those of April and June.
Lows for numbers of individuals and species were found in December,
January, and February when freezing night temperatures were the
rule. The lowest temperature of the study (-10.6°C) was recorded on 24
December 1982. Insect activity is low during these months and even
lower in the canyon bottoms due to cold air drainage. Those species
favoring slopes and ridges have a better chance of maintaining adult
populations over the winter (e.g. Euptoieta claudia).
124
J.Res.Lepid.
A number of other species have been documented from the northern
Huachuca Mtns., the Mustang Mtns., Babacomari Cienega, Canelo
Cienega, and other locales in the general vicinity of the Research Ranch
but failed to turn up during the study. These species include Systasea
zampa (W.H. Edwards), Erynnis meridianus Bell, Celotes nessus (W.H.
Edwards), Oarisma edwardsii (Barnes), Stinga morrisoni (W.H.
Edwards), Atrytonopsis deua (W.H. Edwards), A. python (W.H.
Edwards), Amblyscirtes cassus W.H. Edwards, A prenda Evans,
Agathymus evansi (H. A. Freeman), Megathymus ursus Poling,
Incisalia augustus annetteae dos Passos, Phaeostrymon alcestis oslari
(Dyar), Euphilotes rita (Barnes & McDunnough), Apodemia palmerii
(W.H. Edwards), Thessalia fulvia (W.H. Edwards), Polygonia satyrus
(W.H. Edwards), Anaea aidea (Guerin-Mene ville) , and Gyrocheilus
patrobas tritonia (W.H. Edwards). Any records of these or other
especies from the Research Ranch property should be reported to the
author.
Acknowledgements. The author would like to thank the following for
guidance, encouragement, plant determinations, suggestions, map-making,
general tidbits, etc.: Jane and Carl Bock, Vern and Nancy Hawthorne, Joe and
Helen Taylor, Jack Kaiser, Doug Danforth, Arnold Moorhouse, Char Ernstein,
Renee Vitali, and the Audubon Society.
Literature Cited
BAHRE, CONRAD J., 1977. Land Use History of the Research Ranch, Elgin,
Arizona. J. Ariz. Acad. Sci. Vol. 12, Supplement 2.
BAILOWITZ, RICHARD A., 1983. The Correct Placement of Everes herrii. J. Lep.
Soc. 36(4):3Q8“309.
DAVENPORT, KEN, 1983. Geographic Distribution and Checklist of the Butterflies
of Kern County, California. J. Lepid. Soc. 37(l):46-69.
HOWE, WILLIAM H., 1975. The Butterflies of North America. Doubleday and Co.,
Inc. Garden City, N.Y. 633 pp.
McGuire, william m., 1982. New Oviposition and Larval Hostplant Records for
North American Hesperia. Bull. All. Mus. #72.
MILLER, LEE D. & F. martin BROWN, 1981. A Catalogue/Checklist of the Butterflies
of America North of Mexico. Lep. Soc. Mem. No. 2. 280 pp.
27(2): 120-128, 1988(89)
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SPECIES STATU MONTHS COLLECTED HABITAT REMARKS
27(2): 120-128, 1988(89)
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LIBYTHEIDAE
Libytheana bachmanii larvata P SSRRURR Widespread Breeds near Ranch, wanders in
128
J. Res. Lepid.
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Abundance A = Abundant (100 + seen/day at least twice/month)
C = Common (12-99 seen/day at least twice/month)
U = Uncommon (3-12 seen/day at least twice/month)
Journal of Research on the Lepidoptera
27(2): 129-134, 1988(89)
Notes on a little known ecologically displaced blue,
Agriades pyrenaicus ergane Higgins (Lycaenidae)
LG. Pljushtch
Institute of Zoology, Science Academy of the Ukranian SSR, Kiev 30, Lenina Street 15,
252601, USSR
Abstract. The distribution, ecological relationships, and early stage
features of the recently described blue, Agriades pyrenaicus ergane , are
described. This species is widely disjunct, occurring in the high
mountains of northern Spain, Yugoslavia (?), Caucasus, and in a
specialized habitat in lowland Ukraine.
Distribution
The recent discovery and description of Agriades pyrenaicus ergane in
the southeast USSR was surprising for two reasons: this is a well
studied and known region and the species represents a wide disjunction
in both distance and ecological conditions from the alpine regions of the
mountains of northern Spain and Caucasus, while in the Ukr. SSR their
habitats are found at an elevation of 200M. The subspecies was
originally described from two adult specimens labelled “Voronezh”
(Higgins, 1981). These were collected by O. V. Zuravlev near the village
of Divnogoije (fig. 1) in June 1980 according to information given in
Korshunov (1984), and a series of topotypes given to the Zoological
Museum, Biological Institute, Siberian department of the Academy of
Science. Later this subspecies was found in the Ukraine by Nekrutenko
and Pljushtch (1983) near the village of Efremovka (fig. 1). The latter is
its only known locality in the Ukraine.
Adult Behavior
In 1984 and 1985, 1 made extensive observations of the ecology of the
species near Efremovka. The adult insects flew in the latter part of May.
The males emerged first on May 9. The last males were observed May
27. The earliest females emerged on May 17 and were last observed on
June 1. The mass flight of both sexes was seen between May 19 and 24.
Although common where they are found, the butterflies are extremely
localized and specialized. They are concentrated on the steep southern
exposure chalk slopes forming the banks of the river Volchja here, as
shown in the habitat view in fig. 2. The butterflies do not fly far from
their preferred sites and have never been observed further than 50M
from their biotopes. Adult preference are weakly eroded depressions,
without shrubs or trees, and with a gradient of 30 to 50. This preference
130
J.Res.Lepid.
30°
36°
42°
Fig. 1 . Map of South East USSR showing cites and two localities of A. pyrenaicus
ergane.
appears similar to that described by Thomas (1983) for Ly sandra
bellargus in the south of England. Thomas determined that butterfly
siting was due to higher temperatures in the depressions, which
provided a microclimate more favorable to early stages as well. Fig. 3
shows concentrations of the butterfly hostplant, Androsace koso-
poljanskii Ovcz. (Primulaceae), which grows in high density in the
depressions. Although the plant is occasionally a dominant in its
specialized habitat, it is restricted to central Russian chalk hills of the
tertiary and is regarded as an endangered plant species, see Zaviruha,
Andrienko, and Protopopova (1983: p.35-36). By the same reasoning,
Agriades pyrenaicus ergane should be regarded as an endangered
butterfly. It is both a relict and highly localized.
The development of all stages of the butterfly are intimately associated
with its hostplant. The key feature in the emergence of the adults is the
flowering time of Androsace. Adult butterflies usually nectar on the
hostplant and spend some time perching upon it. The females may
oviposit on the inner surface of the sepals, among the flower buds, or the
inner part of the calyx at the base of the sepals. They also oviposit on the
thin stems or in the leaf axils. Before oviposition, the female thoroughly
probes with her abdomen, but only single eggs are deposited. Eggs are
deposited in a very brief time period (15-60 secs.). Usually the number of
females is so large to suitable oviposition substrate that several eggs
can be found on one site. Adults also nectar on Salvia nutans L., Linum
flavum L., and other plants which are in flower at the proper time.
27(2): 129-134, 1988(89)
131
Fig. 2. A chalk uncovering of steep slopes of the south exposition of river Volchja near village
Efremovka, Volchanski district, Harkov region, Ukr. SSR— the biotop of the Agriades
pyrenaicus ergane.
Fig. 3. The flowering of Androsace koso-poljanskii Ovcs. on the chalk slopes.
Fig. 4. The copulating pair of Agriades pyrenaicus ergane on its host plant.
Fig. 5. A male Agriades pyrenaicus ergane which copulating with fresh but killed female
which was being sucked out by a spider. Another male was evidently flirting with this
strange pair.
Fig. 6. The female of A. pyrenaicus ergane on the flowers of Salvia nutans L.
Fig. 7. The caterpillars of A. pyrenaicus ergane on Androsace koso-poljanskii.
Fig. 8. As 6, different view.
132
J.Res.Lepid.
Strong winds and cloud cover do not adversely affect adult flight. They
tend to fly within 6 to 15cm of the ground, below serious wind effects.
They often perch on stems, chalk stone, and paths. When on the ground
they generally rest sideways to the chalk where they are cryptically
concealed on the light surfaces.
Both sexes emerge in the daytime, mostly between 1000 and 1410. On
a sunny morning adults begin flying at 0730, with males out in mass by
0745. The females follow reaching peak density at 0800. The earliest
copulating pair was observed at 0910. The mass of copulation was
observed between 1000 and 1500. With the high population densities at
the site, as soon as a female emerged, she was seen surrounded by two or
three males. Copulation usually started before females would spread
their wings.
Prior to their first flight in the morning, adults would open their
wings, at a obtuse angle, and turn towards the sun. This apparent
thermoregulatory movement lasted several minutes. They would then
usually start nectaring at once.
An unusual event was witnessed and recorded (fig. 5) on May 19,
1984. The female of a copulating pair was killed and being fed upon by a
spider Xysticus cristatus ( CL) (det: V. E. Gurjanova). The male continued
pumping spermatophore while a second male was attempting to inter-
fere, both oblivious to the situation and danger to themselves. Another
spider, Thanatus sp., was involved in predation of the blues.
Flight time continued until 1800-1900. The butterflies tended to rest
at night at the tops of various low plants, especially showing attraction
to the flower stalks of Salvia nutans and other flowers with pink or violet
colors (fig. 6). The usually grouped in clusters of 3-4, but clusters of 7-8
were seen. At sunrise, the butterflies placed themselves so their folded
wings, underside exposed, were perpendicular to the sun.
Early Stages
In the field the egg stage lasts 10-15 days, with a 50% emergence. The
remainder collapsed, indicating infertility. The neonate larvae fed
exclusively on flower and bud tissue They enter diapause at the end of
this instar, while still very small. They move beneath lumps of chalk on
or under the soil surface. At this point they effectively disappear from
observation.
The following spring diapause breaks with the sprouting of new
vegetative growth of the hostplant in early April following snowmelt.
Feeding is restricted to young leaves and terminal buds. By the third
week in April mature larvae can be found among second and third
instars, with larva densities very high. One Androsace rosette had 25
larvae (a surface area of about 2 dm2. A square meter quadrat carried
more than 100 larvae. The average density was 8.3 per rosette, or about
6 larvae per dm2. The larvae, shown in situ in figs. 7 and 8, are
cryptically colored and difficult to see among the foliage and blossoms of
27(2): 129-134, 1988(89)
133
Androsace. In the earlier instars they are darker, and usually confine
themselves to the top “cone” of the bud where they gnaw through to feed
on the internal bud contents. At this time only part of the posterior
portion protrudes and is very difficult to detect. The last instar is lighter,
and these caterpillars live in a more open situation (figs. 7,8), usually in
groups of three or four. When disturbed they drop to the ground
immediately where they remain tightly rolled up for several minutes.
The larvae are quite sedentary until time to pupate. They then vigor-
ously move about the hostplant and ground. The first pupation was
observed on April 25 in 1985 and was complete on May 1. Pupation from
prepupa to eclosion is about 15 days.
Egg: Echinoid, 0.5-0.6mm, grey, with clear micropyle dorsally. Sculp-
turing of two types, with large, smoother cells dorsally, and smaller
more prominently ridged cells laterally. Larva escapes by cutting a hole
in the lateral part of the egg.
Larva: (figs. 7, 8). Fourth instar ll-13mm, typical omnisciform. Head
completely retractile, small, black. Densely covered with secondary
setae. Background color bright green with prominent stripes. Dorsally
stripes black with lilac shading with dumb-bell shaped sectors in the
middle of each segment, and framed with white. Subdorsal stripes short,
thick dark gray extending anterio-dorsal to posterio-ventral on each
segment. Subspiracular stripe bicolor; the upper part lilac, the lower
white. The pigmentation of the stripes appears epidermal, whereas the
background green appears hypodermal. Spiracles round, lined inside in
black. Setae on the dorsal margin of spiracles longer, 5-6 times as long
as the spiracle diameter.
Pupa: (fig. 9). 9- 10mm, strongly sclerotized. Venation of the forewing
showing on the integument, this wing cover section slightly raised
above the remaining surface. Anterio- ventral part wrinkled. The head
protruding with respect to other body parts., antennae, proboscis and
legs finely differentiated. Postgenae dark, prominent. Labrum large,
heart shaped. Proboscis reaches middle of forewing covers. Pro and
mesothoracic legs short, no tarsi visible. Mesothorax strongly protu-
berant. Eight to tenth abdominal segments ventrally flexed. Cremaster
not expressed.
Conservation status Although Agriades pyrenaicus ergane is a very
abundant butterfly where it occurs, the habitat type is uncommon. It is
known from only the two localities in the region of tertiary chalk hills,
where its larval hostplant is recognized as a species of concern by its
listing the the Red Book of Plants of the USSR. A regular program of
monitoring these populations should be instituted formally, and an
investigation of ecological requirements started. It is likely similar
habitat factors to those regulating populations of Ly sandra bellargus, as
found by Thomas (1983), such as grazing management to keep sward
height reduced, may operate here. The large disjunction of this sub-
species from its vicariant alpine european and Caucasian conspecifics is
noteworthy in the argument to study and preserve these unique insects.
134
J.Res.Lepid.
Literature Cited
HIGGINS L. G. New polyommatine butterflies (Lepidoptera: Lycaenidae). —
Entomologist’s Gazette, Vol. 32, 1981, p. 230-232.
KORSHUNOV J. P. On the type-locality of Agriades pyrenaicus ergane Higgins,
1980 (Lepidoptera, Lycaenidae). — Vestnik zoologii, 1984, No 3, p. 10. (in
Russian)
NEKRUTENKO J. P., PLJUSHTCH I. G. Agriades pyrenaicus (Boisduval) in Ukrainian
SSR. — Vestnik zoologii, 1983, No 6, p. 15. (in Russian).
ZAVERUCHA B. v., ANDRIENKO T. L., PROTOPOPOVA V. V. Protected plants of the
Ukrainia. Kiev, “Naukova dumka”, 1983, 175 pp. (in Russian).
THOMAS, N. A., 1983. The ecology and conservation of Ly sandra bellargus
(Lepidoptera, Lycaenidae) in Britain. Jr. Applied Entom. 20:59-83.
Journal of Research on the Lepidoptera
27(2): 135-143, 1988(89)
Records of Hypaurotis crysalus (Edwards) (Lycaenidae) from
Western Mexico
The distribution of Hypaurotis crysalus (Edwards) in the western United
States can be predicted reliably by the range of its larval host, Quercus gambelii
Nuttall (Fagaceae). Both are widely distributed in the Rocky Mountains from
southern Wyoming, Colorado, Utah, and eastern Nevada, south through Arizona
and New Mexico. Although the host extends considerably further southward
and eastward into Texas and the Mexican states of Sonora, Chihuahua,
Coahuila, and northernmost Durango, H. crysalus has been reported only
once from Mexico (de la Maza and de la Maza, 1975, Rev. Soc. Mex. Lepid.
1(2):64), and this record was from Nuevo Leon.
I have examined two specimens of H. crysalus from western Mexico: 1 6 ,
Durango, 10 mi W El Salto, 8800’, VII-18-64 (J. Powell, Essig Entomological
Museum, University of California, Berkeley); and 1 6, Durango, Cruz de
Piedra, Sierra Madre Mts., XX-4-78 (R. Breedlove, San Diego Natural History
Museum). These localities are nearly 900 km south of the international border
(Arizona-Sonora). In addition, Richard Holland (personal communication) has
collected//, crysalus twice in Sonora: 44.9 mi S Huachinera, VII-2-79, 7300’; and
14.8 mi S Huachinera, VII-4-79, 6900’; and Javier de la Maza (personal
communication) reports a single specimen from the Sierra San Pedro Martir
of northern Baja California.
None of the specimens from Mexico was collected in association with Q.
gambelii, Holland mentioned that all oaks at the sites of his captures were
“encinals” or live oaks; Powell indicated that his speciment was most likely
associated with Quercus sideroxyla (Humb. and Bonpl.) [= Q. omissa (A.D.C.)]
(JAP#433; det, J. Tucker); de la Maza’s (1975) record from Nuevo Leon is
beyond the known eastern range of Q. gambelii ; and Q. gambelii does not occur
in Baja California. No species of oak is common to all these regions. The data
suggest that the southern limit of H. crysalus is not defined by the occurrence of
Q. gambelii , and that other species of oak must serve as larval foodplants in
Mexico.
Comstock (1927, Butterflies of California, pg. 156, published by the author)
mentioned the occurrence of H. crysalus in California on the basis of three
specimens, subsequently believed by Emmel and Emmel (1973, The butterflies
of southern California, Nat. Hist. Mus. Los Angeles Co., Sci. Ser. 26:94) to be
mislabelled, owing to the absence of Q. gambelii in California. However, the
record of H. crysalus from Baja California suggests that the California records
may indeed be valid.
I thank Richard Holland and Javier de La Maza E. for Mexican records of//.
crysalus ; John Tucker (University of California, Davis) for information on
Quercus; Robert Robbins (United States National Museum) and Jerry Powell
(University of California, Berkeley) for comments on the brief manuscript; and
Thomas Duncan (University of California, Berkeley) for allowing me access to
the Jepson-University of California Herbarium at Berkeley.
John W. Brown , Department of Entomological Sciences, University of California,
Berkeley, CA 94720.
136
J.Res.Lepid.
A Chromosome Study of Brahmaeajaponica Butler (Lepidoptera,
Brahmaeidae).
Euroasiatic species of Brahmaeidae present polymorphic populations with an
uneven geographic distribution. The disputed taxonomy of this group is also due
to the occurrence of populations that show morphological characters inter-
mediate between related species (see fig. 1).
The study of chromosomes could help to explain the affinity among different
species. In this regard only Acanthobrahmaea europaea Hartig (n = 32, 2n = 64)
has been recently investigated (Trentini and Marini, 1985: Atti XIV Congr. naz.
ital. Ent.: 299-303).
Fig. 1. Geographic distribution of Brahmaeidae. The data were obtained
from papers of Staudinger and Rebel (1901: Friedlander & Sohn
Ed., Berlin), Seitz (1911: A. Kernen Verlag, Stuttgart), Mell (1928:
A. Kernen Verlag, Stuttgart; 1929: Dtsch. Ent. Z., 5: 337-494;
1 937: Dtsch. Ent. Z., 1-1 9), Rougeot (1 971 : Masson & Cie Ed., Paris), Chu
and Wang (1977: Acta Entomol. Sin., 20: 83-85), Nassig (1980: Nachr.
ent. Ver. Apollo, N.F., 1: 77-91), Freina (1982: Entomofauna, 3(9): 129-
139), Freina and Witt (1982: Nota lepid., 5 (2-3): 81-85). Euroasiatic
species: 1 , Brahmaea certhia F.;2, Brahmeae christophi Stgr; 3, Brahmaea
ledereri Rghfr; 4, Brahmaea porphyria Chu & Wang; 5, Cailiprogonos
miraculosa Mell; 6, Acanthobrahmaea europaea Flertig. Indo-australian
species: 7, Brahmaea hearseyi White; 8, Brahmaea wallichi Gray; 9,
Brahmaea japonica Butlr. Ethiopian species: 10, Dactyloceras lucina
Drury; 11, Dactyloceras ocelligera Butlr; 12, Dactyloceras catenigera
Karsch; 13, Dactyloceras bramarbas Karsch; 14, Dactyloceras barnsi J.
& T.; 15, Dactyloceras ostentator Hering; 16, Dactyloceras Widenmanni
Karsch; 17, Dactyloceras maculata Conte.
* Freina (1 982: Ibid.) reported a new record of a population of Brahmaea
ledereri from Hakkari (Turkish Kurdistan region), that shows intermediate
features between B. ledereri and B. christophi ; for this reason the author
considers B. christophi conspecific with B. ledereri.
27(2): 135-143, 1988(89)
137
The present research reports the early results obtained on the chromosome set
of Brahmaea japonica, both males and females.
Brahmaea japonica was reared in 1986-1987 in laboratory on Ligustrum sp.
and Syringa vulgaris from ova received from Japan. Karyological observations
were carried out on eight pupae (4 males and 4 females) at one month before
adult emergence, employing the air-dried technique (Trentini and Marini,
1986: Genetica, 68: 157-160); the detailed procedure is as follows: after 0.05%
colchicine pretreatment for 2 h, testis and ovarioles were dissected out and kept
under 1% sodium citrate for 20 min, fixed in 3:1 alcohol-acetic acid, dissociated
in 60% acetic acid on a warmed slide, postfixed in Carnoy fluid, and stained with
2% Giemsa (pH 7) for 15-20 min at room temperature.
Figs. 2-5. Spermatogenesis (2, 3) and oogenesis (4, 5) of Brahmaea japonica. 2,
pachytene; 3, C-metaphase; 4, achiasmatic bivalents; 5, oogonial
C-metaphase.
138
J. Res. Lepid.
Males. In pupal testes of Brahmaea japonica very few mitoses are present
probably because the spermatogonial increase occurs in the last two instar
larvae. The found C-metaphases show 94 chromosomes; they are rod- and dot-
shaped and range from about 0.6 pm to about 2 pm (fig. 3). At the prophase of the
first meiotic division 47 bivalents are visible (fig. 2).
Females. Ovarioles still show oogonial mitoses and the start of meiosis. Fifty
mitotic C-metaphases of three specimens were scored for chromosome number:
five metaphase plates present 2n = 93, forty-three 2n = 94, and two 2n = 95. The
chromosomes are rod- and dot-shaped and their length ranges from about 0.7 pm
to about 1.5 pm (fig. 5). Some prophases of the first meiotic division with 47
bivalents were observed; they consist of parallely aligned homologues showing
their achiasmatic nature (fig. 4), as already reported in other Lepidoptera
(Suomalainen, 1965: Chromosoma (Berlin), 16: 166-184; White, 1973: Cam-
bridge Univ. Press).
With regard to the sex chromosome mechanism, the same chromosome number
found in both males and females exlcudes an XO system and indicates an XY
system, even though the sex chromosomes are undetectable in our preparations.
The only karyologically studied Brahmaeidae species, to our knowledge, are
Acanthobrahmaea europaea (n = 32; 2n = 64) and Brahmaea japonica (n = 47;
2n = 94). The two species are very different in size and wing features, moreover
they occur at the extremities of the euroasiatic region. Given the chromosome
number 2n — 94 of B. japonica, it could be supposed that B. japonica presents a
quasi-polyploidy (3n - 1) in relation to A. europaea. But two facts are contrary to
this hypothesis: 1, the chromosomes of A. europaea are clearly larger than those
of B . japonica; and 2, the genome of the two species is about the same size. We
think that the variable chromosome numbers in Brahmaeidae are probably due
to chromosomal rearrangements (fusion and dissociations), as already reported
in other non-parthenogenetic Lepidoptera (Robinson, 1971: Pergamon Press,
Oxford; White, 1973: Ibid.).
At a future time it would be valuable to examine the DNA content of both
species and the chromosome complements of some other Brahmaeidae.
This research was supported by a grant from M.P.I. 40% 1985.
Massimo Trentini and Mario Marini, Department of Biology, University of
Bologna, via S. Giacomo 9 1-40126, Italy
Re visional notes on the Genus Satarupa Moore (Lepidoptera:
Hesperiidae). I. New Synonyms of Satarupa monbeigi Oberthur.
Satarupa monbeigi Oberthur, 1921:76, pi. Y, Y bis.
= Satarupa omeia Okano, 1982:91-94, PL 1, figs. 1, 2 male; fig. 1, male genitalia
(Syn. nov.)
= Satarupa lii Okano and Okano, 1984:124-126, PI. 9, figs. 1, 2 male; figs. A,
male genitalia (Syn. nov.)
In 1982, Okano described Satarupa omeia from Omeishan, Sichuan, Peoples
Republic of China, as a new species. Two years later he described another ‘"new”
species, Satarupa lii (Okano and Okano, 1984), from exactly the same locality.
27(2): 135-143, 1988(89)
139
We consider both of Okano’s two species conspecific with Satarupa monbeigi
Oberthur, 1921, for the following reasons.
Four described taxa of Satarupa, namely S. valentini Oberthur, 1921, S. zulla
ouvrardi Oberthur, 1921, S. nymphalis khamensis Alpheraky, 1897 (=ober-
thueri Evans, 1932, = intermedia Evans, 1932) and S. monbeigi Oberthur, 1921,
have hitherto been known from West China (Evans, 1949). Okano should have
compared his two species with these four known species, but he neglected to do
so. Even if he had no opportunity to examine these species himself or through
authoritative persons, he should, at least have keyed his specimens using
Evans (1949). Instead, he compared his “new” species with S. formosibia
Strand, 1927, from Taiwan. In the descriptions he mentioned that S. omeia
“most closely resemblefd] S. formosibia Strand in almost similar appearance”,
and that S. lii was “very near to Satarupa formosibia Strand from Formosa”.
However, he did not mention that S. formosibia was the closest species that he
compared with those two species among the genus Satarupa. Our revisional
work (unpublished) suggests that S. formosibia is abnormal within this genus in
wing markings and male genitalia. Moreover, he did not refer to his own first
paper (S. omeia) in his second paper (S. lii).
We examined two males from the same locality (Omeishan, Sichuan), and
determined that those were S. monbeigi and so were Okano’s two species (based
on his figures). In figure 2 of both descriptions, the inner dot in space 7 on the
ventral side of the hindwing is vestigial, but still present. Within Satarupa the
presence of this dot separate the group of species which includes S. monbeigi
from the group which includes S. formosibia. Male genitalia of S. omeia and S.
lii appear slightly different in Okano’s figures, especially on the tip of the harpe
and the curve of the style. However, these difference appear to be either
individual variations or artificial (subjective) modifications of the figures.
Figures should be drawn carefully, with sufficient understanding of the
structures, rather than rough sketching (Kawazoe, 1973).
We thank J. N. Eliot for examination of specimens in the British Museum
(Natural History), S. Miller and A. Kawazoe for review of the manuscript.
Literature Cited
EVANS, w. H., 1949. A Catalogue of the Hesperiidae of Europe, Asia and Australia
in the British Museum (Natural History). British Museum (Natural History),
London.
KAWAZOE, A., 1973. A revisional note on some Philippine Lycaenidae and
Hesperiidae reported by Murayama and Okamura. Tyo to Ga 24:91-98.
OBERTHUR, C., 1921. Figuration Photographique de quelques Lepidopteres.
Etud. Lep. comp. 18:67-77.
OKANO, M., 1982. New or Little Known Butterflies from China (II). Artes
Liberales No. 31:91-94.
OKANO, M. & T. OKANO., 1984. A new species of Satarupa from West China
(Lepidoptera: Hesperiidae). Tokurana (Acta Rhopalocero.) Nos. 6/7:124-126.
Hideyuki Chiba, Department of Entomology , University of Hawaii, 3050 Made
Way, Honolulu, Hawaii 96822.
Hiroshi Tsukiyama, 4-18 N arashinodai#2 -303, Funabashi-shi, Chiba-Pref.
274 JAPAN
140
J.Res.Lepid.
Notes on Panacea procilla lysimache (Nymphalidae ) from Costa
Rica.
The nymphalid butterfly genus Panacea is generally thought to range from
the highlands of Chiriqui Province of western Panama southward into South
American. (DeVries, P.J. 1987. The Butterflies of Costa Rica and their Natural
History. Princeton University Press, Princeton.) Godman & Salvin (1893)
described a single specimen of the genus collected from “Chiriqui” as P.
lysimache remarking that it was the only specimen known from Central
America. In his treatment of Panacea Fruhstorfer (1912-1914. Panacea, in: A.
Seitz (ed.). The Macrolepidoptera of the World. Vol. 5, Stuttgart (Alfred Kernan.)
downgraded lysimache to a subspecies of P. procilla , a species that ranges from
Panama to the Amazon Basin, and also noted that the holotype of procilla
remained the only specimen of Panacea known from Central America. Until
recently I was aware of only four Central American specimens of P. procilla
lysimache , all from the highlands of Chiriqui in Panama (1200-2000m), and have
suggested that the butterfly was likely to be found eventually in Costa Rica from
localities in the Cordillera de Talamanca near Panama (DeVries 1987). Here I
report the first authentic Costa Rican collection of P. procilla lysimache from a
forest type very different where it has previously been collected.
On 3 August 1987 at 13:15 hours I collected a fresh male P. procilla lysimache
(Godman & Salvin, 1893) [forewing length = 45.5mm; proboscis length 24mm]
that was feeding at a sap flow on a medium-sized, mature Persea americana
(Lauraceae) tree growing on the laboratory side of the bridge at Finca La Selva,
Heredia Province, Costa Rica. The butterfly was perched head downward about
2m above the ground with the wings open and appressed to the tree trunk and
feeding alongside an individual male Myscelia cyaniris cyaniris (Doubleday,
1848).
The P. procilla lysimache individual was originally noticed at 12:00 hours
making sorties around, and perching head downward on the trunk of a
introduced Asian tree ( Averrhoa caramholla : Oxalidaceae), and it may have
been feeding on the rotting fruit that littered the ground under the tree. Both
trees where the butterfly was observed grew in a open area heavily trafficked
by humans located about 30m from the edge of a secondary forest and within
50 m of the Rio Sarapiqui.
While on the wing flutter-glide flight behavior and reddish underside made the
P. procilla individual appear much like a large Hamadry as amphinome mexicana
(Lucas, 1853). The following nymphalid species were noted to either be feeding
on the fallen fruits of Averrhoa , the sap flow of the P. americana tree, or flying
in the near vicinity at the time of capture: M. cyaniris, Archaeoprepona Camilla
(Godman & Salvin, 1884), A. demophoon gulina Fruhstorfer, 1904, Prepona
omphale octavia Fruhstorfer, 1904, Marpesia merops (Boisduval, 1836), Eueides
lybia olympia (Fabricius, 1793), Cissia hermes (Fabricius, 1775) and C. lobe
(Butler, 1870). Although I spent a additional 16 days at La Selva, no other P.
procilla individuals were seen.
There are two considerations I wish to raise regarding P. procilla in Central
America. One is that Finca La Selva (55-100m elevation) is covered mostly by
lowland Atlantic rainforest that is very different from the cloudforests of
27(2): 135-143, 1988(89)
141
Chiriqui where previous Central American records of P. procilla originate. The
fresh condition of the specimen suggests that rather than immigrating from the
mountains of the Cordillera Central or Talamanca, it eclosed either at La Selva
or in the immediate vicinity: a broad range of habitats for a rare butterfly
species. The second point of consideration is simply to wonder how a large,
garrishly colored butterfly species, that is collected commonly near human
habitations in South America, has escaped detection in Costa Rica (and
Panama) for so many years.
Acknowledgements: I thank N. Greig for asking me “is that a Hamadryas ?”
and running to get my net.
P.J. DeVries , Smithsonan Tropical Research Institute, Box 2072, Balboa,
Panama.
An Additional Natural Hostplant of Pieris Virginiensis (W.H. Edwards)
(Pieridae) in Ohio
For many years, the West Virginia white, Pieris virginiensis (W.H. Edwards),
was known to utilize only tooth wort, Dentaria diphylla Michx., as a natural
hostplant (Klots, 1935). Although other species of Dentaria were long suspected
to serve as natural hosts (Klots, 1951), only cut-leaved toothwort, Dentaria
laciniata, was subsequently reported (Shapiro, 1974; Chew, 1980; Cappucino
and Kareiva, 1985). Scott (1986) included Pennsylvania bitter cress, Cardamine
pennsyvanica Muhl., and Brassica as hosts without reference. Recently, smooth
rock cress, Arabis laevigata (Muhl.) Poir., was found to serve as an additional
host in central Ohio (Shuey and Peacock, in press). P. virginiensis will also feed
upon a number of mustards in the lab that are not utilized in nature (Shapiro,
1971; Chew, 1980).
On 25 April 1988, a female P. virginiensis was observecd ovipositing on
narrow-leaved toothwort, Dentaria multifida (Muhl.), on a rich forested stream
terrace in Delaware County, Ohio. At this site, D. laciniata is abundant and
serves as the primary host of P. virginiensis. Arabis laevigata is also fed upon
with some frequency in this area but is uncommon in occurrence. Dentaria
diphylla is absent. The single ovum deposited in D. multifida was collected and
reared to pupation on the leaves of this newly discovered host. One additional
ovum was later found on D. multifida and also reared to pupation.
Throughout its restricted range, D. multifida is generally considered un-
common, occurring in Indiana, Ohio, West Virginia, Kentucky, Tennessee,
Georgia, Alabama, and North Carolina (Montgomery, 1955; Duncan and Foote,
1975). In Ohio, D. multifida is rare and considered threatened. Post-1960
records exist for Delaware, Athens, Washington, and Morgan Counties
(McCance and Burns, 1985). The single historical Delaware County site is
located several kilometers north of the site found in 1988 (Long, 1956; Allison
W. Cusick, pers. comm.). Within Ohio, the known ranges of D. multifida and P.
virginiensis overlap only in Delaware County.
Shapiro (1971) and Chew (1980) noted thatP. virginiensis females will readily
oviposit on many species of mustards but few mustards are typically available in
the forested habitats of the butterfly. Hence, the utilization of D. multifida in
142
J.Res.Lepid.
Delaware County, Ohio is probably due to its presence in an area where P.
virginiensis is established in association with another, more common Dentaria.
In the eastern United States, the range of D. multi fida lies nearly completely
within the range of P. virginiensis and may serve as a host outside of Ohio.
With the addition of D. multifield as a host, P. virginiensis has now been found
and reared (at least to pupation) on four species of mustards in Ohio. Dentaria
diphylla and D. laciniata appear to serve as the primary hosts, while Arabis
laevigata and D. multifida are known to be utilized locally. An examination of
other species of mustards found growing in habitats where P. virginiensis occurs
will probably reveal additional natural hostplants.
Acknowledgements. Thanks are extended to Reed A. Watkins for his assi-
stance in the field.
Literature Cited
CAPPACCINO, N. & P. kareiva, 1985. Coping with a capricious environment: a
population study of a rare pierid butterfly. Ecology 66: 152-161,
CHEW, F.S., 1980. Foodplant preferences of Pieris caterpillars (Lepidoptera).
Oecologia 46: 347 -“353.
DUNCAN, W.H. & L.E. FOOTE, 1975. Wildflowers of the southeastern United States.
University of Georgia, Athens. 296p.
KLOTS, A.B., 1935. On the life history of Pieris virginiensis Edwards (Lep.,
Pieridae). J. New York Ent. Soc. 53: 139-142.
— — , 1951. A field guide to the butterflies of eastern North America.
Houghton Mifflin Co., Boston. 349p.
LONG, R.W., JR., 1956. Dentaria multifida in central Ohio. Rhodora 58: 160-161.
McCANCE, R.M., JR. & J.F. BURNS, eds. 1984. Ohio endangered and threatened
vascular plants. Ohio Department of Natural Resources, Division of Natural
Areas and Preserves. 63 5p.
MONTGOMERY, F.H., 1955. Preliminary studies in the genus Dentaria in eastern
North America. Rhodora 57: 161-173.
SCOTT, J.A., 1986. The butterflies of North America. Stanford University Press,
California. 583p.
SHAPIRO, A.M., 1971. Occurrence of a latent polyphenism in Pieris virginiensis
(Lepidoptera: Pieridae). Ent. News 82: 13-16.
— , 1974. Butterflies and skippers of New York State. Cornell University
Agriculture experiment station Circular 4: 1-60.
SHUEY J.A.&J.W. PEACOCK., In Press. A significant new hostplant record for Pieris
virginiensis (Pieridae). J. Res. Lepid.
John V. Calhoun, 369 Tradewind Ct., Westerville, Ohio 43081 and
David C. I finer, 2161 Heatherfield Ave., Worthington, Ohio 43235
27(2): 135-143, 1988(89)
143
Water pumping in Lamproptera ineges (Papilionidae)
Lamproptera meges (Zinken), called the “Green dragontail”, is a bizarre
papilionid resembling a dragonfly in flight. It inhabits the forests and is usually
found near running water. It’s wingspan is about 4-5.5 cm, and it has 4 cm long
tails. The individual shown was photographed on February 28, 1986 in Malaysia,
in the middle of the Malayan peninsula west of Cameron Highlands at about 600
m. The behavior shown is extremely unusual because it illustrates water
expulsion from the butterfly’s anus (Fig. 1). This was a rare synchronization
between the 1/250 second shutterspeed and the approximately 1/500 sec pro-
jectile-style water squirt. Butterfly was imbibing water from the mud and
periodically, about each 4-6 seconds, expelling water. One of the reasons for
puddling behavior is the acquisition of sodium ions, as documented in Papilio
glaucus by Arms, Feeny and Lederhouse (1974. Science 185:373-74).
George O. Krizek, 2111 Bancroft Place, N.W., Washington, D.C. 20008, and Paul
A. Opler, 5100 Greenview Ct., Ft. Collins, CO 80525.
Fig. 1 . Lam propter ameges expelling water anally. T aken in native habitat, Central
Malasia.
Journal of Research on the Lepidoptera
27(2): 144, 1988(89)
Book Reviews
TAGF ALTER UND IHRE LEBENSRAUME. [W. Geiger (Ed.)]. 1987.
Schweizerischer Bund fur Naturschutz, Basel. 516 pp., col. ill.; ISBN not stated.
Price Sfr. 110, — hardback.
The German title (English translation: Butterflies and their habitats) of this
book is somewhat misleading: natural history and conservation of Swiss
butterflies would have been much better choice. The book has neither a senior
author nor editor; this is from a bibliographical and taxonomic points of view
inexcusable. 16 authors and editors are listed on p. VI as members of a working
group responsible for this book. I propose to attribute the book to W. Geiger (who
was obviously the most important editor and author) to avoid future biblio-
graphical confusion. The names of contributors are listed in reversed order,
starting with the surname followed by the first name. This too can lead to
confusion. Authors of individual chapters are not stated. The book includes the
following parts: Biology of butterflies; Butterfly habitats; Decline of butterflies
in Switzerland and its causes; Conservation of indigenous butterflies; Systematic
part; Distribution and ecology of Swiss butterflies and Glossary. The book is
lavishly illustrated in colour throughout the text (adults, eggs, larvae, pupae,
habitats etc.) and in addition the adults of all species are illustrated on 25
beautiful colour plates. The species monographs are informative and feature the
following topics: description of adults, eggs, larvae and pupae; ecology of adults,
eggs, larvae and pupae; Ecology; Distribution (with maps of all species);
Phenology and Conservation. To facilitate the identification of some taxonomi-
cally difficult species, line drawings of genitalia and some other relevant
morphological characters accompany the description, but usually do not reach
the very high standard of colour illustrations. The book is extremely well
produced and will surely take an important place among the contemporary
standard works on the butterflies of Central Europe. It is being offered at a very
reasonable price. It is difficult to criticise it except for the confused authorship. I
found only a few other minor points of criticism: (1) illustrations of genitalia are
somewhat crude, (2) the names of plants are not printed in italics and (3) the
short chapter on zoological nomenclature and the subsequent treatment of some
names show, that the editors should have asked the advice of a competent
taxonomist familiar with English language and the use of the International
Code on Zoological Nomenclature (examples of confusion: unavailable names
cannot take priority over available names; synonyms do not become unavailable;
names proposed for aberrations are always unavailable). It is a great pity that
Hesperiidae have been excluded (their exclusion can be justified from a
taxonomic point of view, but not in this type of book). I recommend this book to
any student of European butterflies and congratulate the team of its authors.
Otakar Kudrna , Rhenusallee 32, D-5300 Bonn 3 (Germany)
27(2): 144-150, 1988(89)
145
MARIPOSAS MEXICANAS. Guia para su colecta y determinacion. Roberto de
la Maza Ramirez. 1987. Fondo de Cultura Economica, S. A. de 0. B., Av. de la
Universidad, 975; 03100 Mexico, D. F. 302 pp. 67 color plates. Price $60.00.
Hardbound. In Spanish. (Can be ordered directly from Javier de la Maza E.,
Boehil 340, Col. H. de Padiema, C. P. 14200, Tlalpan, B. F., Mexico.)
This is the first book to provide a comprehensive illustrated introduction to
the Mexican Rhopalocera fauna. The writer is a member of one of the most
famous iepidopterist families in Mexico, and liberally shares his extensive
knowledge of that fauna with us in this magnificently illustrated treatment.
With 67 color plates illustrating over 600 species of all families including
skippers, it will be an invaluable reference for identifying and collecting a very
substantial part of the Mexican butterfly and skipper fauna. In most cases, only
the upper side is shown, but these are usually sufficiently diagnostic to allow
ready identification of most Mexican species.
A total of 651 species are covered in the text. This is not the entire Mexican
butterfly fauna, by any means, nor is the coverage of each species exhaustive,
but the volume in total does accomplish its purpose of illustrating and describing
many of the most representative species in the country, including taxa described
by the very active Mexican lepidopterists from 1950 to date, as well as the well-
known endemics such as Papilio alexiares (which resembles the Eastern Tiger-
Swallowtail, Papilio glaucus ), Pterourus esperanza, Polygonia haroldi , and
many other colorful species in the fascinating Mexican fauna.
The Mexican butterfly fauna is one of the richest and most diverse in the
world, and well deserves the further intensive study which this work will help to
stimulate. For example, 685 species of skippers alone are known between
Guatemala and the United States border, which constitute some 40 to 45% of the
diurnal butterfly fauna of Mexico. In the family Riodinidae, some 180 species
are found throughout Mexico, 75 species of Fieri dae, 56 species of Papilionidae,
6 species of Danaidae, 35 species of Ithomiidae, 70 species of Satyridae, 15
species of Brassolidae, 4 species oi Morph :i dae, 6 species of Acraeidae, 21 species
of Heliconiidae, 200 species of true Nymphalidae, 12 species of Apaturidae, 42
species of Charaxidae, 2 species of Libytheidae, and some 230 species of
Lycaenidae. There are also at least 10 species known in the Giant Skipper
family Megathymidae. Thus the author had a formidable task in selecting the
most representative butterflies to both illustrate and briefly describe. The text
on each species includes its Latin name, figure reference, habitat description,
preferred habitat, and the states (with specific localities following each state
name) in which each species occurs. The dates of annual flight period are also
indicated for virtually all species.
In addition to the valuable summary of the Mexican butterfly fauna in
individual species accounts, de la Maza has an excellent introductory section on
the classification of butterflies, fossil butterflies, morphology, life history,
genetic phenomena, mimicry and cryptic coloration. An excellent section on
collecting, including use of live traps (a very important collecting method in the
tropics), the captive culture of butterflies, preparation of specimens, determina-
tion and conservation are discussed. The section on spreading specimens is
illustrated by beautiful full-color photographs of Morpko specimens being
spread on a mounting board. One of the most unique features of this book may be
the section discussing the butterfly in ancient Mexico, where the representa-
146
J.Res.Lepid.
tions of butterflies have been found in archaeological structures and artifacts
throughout Mexico. An additionally fascinating section is on the lepidoptero-
logy history and some of the chief lepidopterists who have been active in the
country. There is an excellent section on habitats and biogeography of the
Mexican butterflies illustrated by beautiful color maps and color pictures. In
fact, overall the spectacular illustrations and the text makes this one of the best
books on the natural history of tropical butterflies that has yet been published.
The text is in Spanish, with Latin names for all species. A lepidopterist with a
reasonable working knowledge of Spanish will have no trouble in reading the
text, which is written in a very informative, smoothly flowing manner. The
discussions of abberations, rare species, Monarch migration, and other topics
are not overly technical and can be read with ease. The text on the species
accounts is sufficiently simple and telegraphic in style that most lepidopterists
will be able to translate the habitat, locality, and date information with the aid of
a dictionary and no great difficulty, even without a previous knowledge of
Spanish.
Overall, this is an outstanding book which should be in the library of every
lepidopterist interested in the Mexican fauna and tropical butterflies in general.
The inclusion of temperate-zone genera that have reached southward into
Mexico in the temperate mountain regions, such as Polygonia, will appeal to
even North American lepidopterists who are specializing only in the U. S. fauna.
Thomas C. Emmel, Department of Zoology, University of Florida, Gainesville,
FLA 32611, USA
THE BUTTERFLIES OF INDIANA. 1987 Ernest M. Shull, Indiana University
Press, 272 pp. $25.00 U.S.
This volume is the first recent faunal treatment for a entire Great Lakes
State. As such, it should be an important library addition for those interested in
the region. The book is excellently produced, and is perhaps the most handsome
volume on butterflies currently available.
The book is organized into two sections which are subdivided into four parts.
Part one includes the Introduction and very brief accounts of butterfly biology,
collecting, and classification. These are followed by a short discussion of
butterfly conservation and the Endangered Species Act. Parts two, three, and
four are the species treatments and checklist, which make up the majority of
this volume. Each species treatment includes a description of the butterfly, and
discussions on status, distribution, habitat, and life cycle. Distribution maps are
provided for each species, and all of the species known to occur in Indiana are
figured on the 49 plates.
Shull’s writing is generally a welcome relief from most terse scientific styles,
but some of the text within the species treatments seems irrelevant, especially
the recounted observations from Mexico. Most readers will enjoy the species
discussions more than the introductory material which because of its brevity, is
very telegraphic and choppy. The botanical nomenclature used is erratic, and
some plants jump between genera (e.g., shrubby cinquefoil between Potentilla
and Dasiphora ).
27(2): 144-150, 1988(89)
147
The forward, by William Eberly, implies that the volume contributes to four
areas of butterfly biology; it serves as an identification guide; it contains
comprehensive life histories; it contains extensive listings of foodplants; and, it
presents much detailed information of the mating habits of butterflies. The
following paragraphs assess the attainment of these contributions. My critiques
are not intended to be comprehensive, but rather to alert the potential readers of
the volume to strengths and weaknesses in Shull’s work.
Identification — The plates are of exceptionally high quality and when used
with the species descriptions in the text, readily identify most of the species
found in Indiana. The treatment of various groups on the plates is very uneven:
seven plates figure 53 species of skippers while 15 plates illustrate six species of
swallowtails. Some of the specimens figured are too worn to be useful for
identification purposes (e.g. Hemiargus isola and Incisalia henrici ). Others are
misidentified; Erynnis Juvenalis appears also as E. horatius and E. lucilius;
Incisalia n. niphon as J. niphon clarki ; anxiAtlides helesus estesi as A., h. helesus.
The treatments of certain subspecies are very problematical, and likely
to confuse readers unfamiliar with evolutionary theory. Cercyonis pegala is
separated into three distinct subspecies ( alope , nephele , and olympus ), each with
its own text, distribution map, and plate figures. Since Shull demonstrates that
these subspecies occur sympatrieally, and has even collected them in the same
locality on the same day, a more parsimonious solution would have been to
accept these “subspecies” as phenotypes of a single species, and simply call it C.
pegala. A similar problem exists with Speyeria a . aphrodite and S. aphrodite
alcestis.
Life histories — The information presented in the text seems to be a brief
recounting of descriptions found throughout the general literature. Although
some of the information presented is unattributed, it is never clear if this
information is independent conformation of these life histories, or perpetuation
of standard (and often incorrect) knowledge.
Foodplants — Again, the information here seems to be recounted out of the
general literature (despite the general lack of citations). Most disturbing is the
tendency to misrepresent the hosts of the rarer species. For example, Erynnis
persius is listed as feeding on “willow (Salix) and various species of Populus”,
despite the fact that it is restricted to lupine ( Lupinus perenis ) in the lower Great
Lakes area. Lycaeides melissa samuelis is not considered to be endangered
because of the wide “variety of foodplants” that it uses, but in reality, this
subspecies feeds exclusively on lupine throughout its range. Most blatantly,
Satyrium edwardsii is reported to use scrub oak ( Quercus ilicifolia ), a tree that
does not occur in Indiana.
Mating habits — This section would be a unique contribution, but unfortun-
ately, very little of the information presented has anything to do with reproduc-
tive behavior. The bulk of these discussions are observations made after pairs
have coupled, and includes information such as the time of day, air temperature,
where the pair was found, where they were resting, and which sex flies in copula.
A few brief observations on possible courtship interactions are presented.
One of the more important contributions that treatments of state faunas
generally provide, is the resulting set of detailed distribution maps. This volume
seems to provide accurate maps for the more common and easily identified
species. However, unverified literature records are included without comment
on the maps, affectively incorporating all the errors of the past into this volume.
148
J.Res.Lepid.
For example, no self respecting Lethe creola would ever be lost in a fen in
extreme northeastern Indiana (based on a literature sight record). I also suspect
that the distributions of many of the skippers reflect erroneous literature
records. Some of the more interesting distribution patterns are really a
reflection of inadequate collecting: the counties average less than 34 recorded
species each.
Readers wanting a general feel for the butterfly fauna of Indiana and adjacent
states will certainly find this volume to be valuable. Those interested in more
specific information on ecology or hostplants will be less pleased with the text,
and will find it difficult to separate the interesting phenomena from the
recitations (inaccuracies and all) from the general literature.
John A. Shuey, 731 Kerr St., Columbus, Ohio 43215
D’ABRERA, B. 1986. Sphingidae Mundi. Hawk moths of the world. E. W.
Classey Ltd., 226 pp. 97.50 pounds Sterling.
This is a fantastic pictorial field guide to the taxonomy of the world’s
Sphingidae. It removes 98% of the taxonomic obstructions to working on or with
the ecology, behavior, morphology, etc. of this extremely important group of
moths — major pollinators as adults, major food for insectivores as adults and
larvae. It photographically figures the adults of about 1,050 species and covers
virtually the world fauna. And it should lie at the base of an explosion of
biological information about sphinx moths, quite analogous to the production of
a field guide to the birds or wild flowers of a region.
There are three roots to the production of Sphingidae Mundi — D’Abrera’s
fanaticism towards giving us picture books of major lepidoptera groups, Alan
Hayes’ quarter of a century of organizing our taxonomic knowledge about
sphingids, and the British Museum’s fanatic attention to the scientific com-
munity’s need for centralized and curated vouchers for our understanding of
animal biology. We were blessed with a mover, a curator, and a responsible
repository for the raw material with which they worked. That we should be so
forsighted as to cause the juxtaposition of these three items for the other species-
rich groups of organisms on this globe, and do it fast enough that guides can
appear now. And it will be these guides that will provide the taxonomic
foundation on which the next half century of conservation and restoration
actions will build, actions that will determine what humankind has to work
with forever in the tropics.
All taxonomic workers should take note of several traits of this book, many
good, a few not so good. First, it is extremely thorough for our knowledge to date,
yet obviously does not cover all the sphingids that will finally come to light.
Second, the photographs are crystal clear and replace endless pages of unread-
able keys, allowing both the novice and the professional to be inspired and
construct on this taxonomic base. Third, reams of detailed information about
distributions, larval stages, etc. are not necessary to fill the function of guiding
the reader to a name, a name that is itself the call number to the book on that
species’ biology. Fourth, with this table-top summary of a family in hand,
immediately a host of ecological, evolutionary, behavioral questions pop to mind
27(2): 144-150, 1988(89)
149
— questions that can now be attacked by those with very little previous
knowledge of sphingids or even Lepidoptera; the guide is an obvious magnet for
those in other areas of biology. For example, it leaps out from these pages that
African Meganoton, Coelonia and Macropoliana are either congeneric with
Neotropical Manduca , or we are confronted with the most incredible case of
convergence and parallel evolution known in biology; the same applies to the
Neotropical Xylophones and the Old World Hippotion.
On the debit side of the ledger, I hope that future generators of such guides
will view this as a working model, one that can be fine-tuned by collaboration
between the users and writers. For example, the inclusion of references to the
primary literature on geography and larval (and even adult) biology would at
best have added 5—10 pages to the total in the book — such references could be
cast in very small type. It is quite possible that Alan Hayes intended to do this
before his untimely death. Second, I suspect that there are sphingid workers
around the world who could have (would have) quite readily briefly commented
on the biology of the species of their area, comments that would have added in
the very large amount of current biological information on sphingids that does
not appear in the book. For example, it is wondered if Xylophones godmani and
Xylophones rhodina are two sexes of the same species (p. 172) when in fact both
sexes of both species have been frequently collected in the highlands of Costa
Rica in recent years by W. A. Haber, I. Chacon and myself. Third, the reader
should have been cautioned that sphingid taxonomy and taxonomic knowledge
is still in a state of flux; one can get a very positive identification from this book,
but such field determinations should still be checked by someone up on the most
recent discoveries in sphingid taxonomy. Fourth, D’Abrera was forced to pick up
the production cost out of pocket, in hopes of recuperation through sales; the
consequence is a book with a price so great that it will only be purchased by
professional lepidopterists and libraries. However, books such as these are of
enormous general biological value, and should be widely circulated and
available to realize their potential as stimulators of field studies. It is impera-
tive that the general funding community in field science come to recognize that
the subsidy of field guidebooks such as this one is as critical as is purchase of field
equipment, laboratories, and airplane tickets.
Some readers may object to my calling it a “field guide”, owing to its large size
and weight. The fact is that the “field” for a worker on sphingids (or any insect
group) generally involves both nets and mud, and tables and roofs. This book
will have no problems in the library of a biological field station, in a box in the
back of a truck, or on a lab bench. The real barrier is cost; expensive books tend
to be guarded so carefully that they are not left around for the uninitiated to
browse and contemplate.
In sum, any tropical biological station or field research area should have a
copy of this book right along with their floras and field guides to vertebrates. We
are now at the point of taxonomic understanding for sphingids whereby their
biology should be taken fully into account. Lets do it.
Daniel H. Janzen, Department of Biology, University of Pennsylvania, Philadel-
phia, PA19104
150
J.Res.Lepid.
S ATURNIID AE Ecological and behavioral observations of select Attacini .
Robert D. Weast. 1989. 53 pp. Published by the author. Available from:
S ATURNIID AE, 5324 NW 78 St. Ct., Johnston, Iowa 50131, Price: $16.90
As stated in the author’s preface, this book was written by an amateur in an
informal style for the non-professional lepidopterist, who makes up the vast
majority of the lepidopterists’ Society. The book documents attempts to colonize
the Des Moines area with Callosamia promethea, Automeris io, and Samia
cynthia ; other sections deal with life history data on Rothschildia and Eupac-
kardia; experiments and general remarks on mating biology and population
structure; and a description of a promethea X cynthia hybrid brood. At the end of
several sections the author calls for more research to test various speculations
and theories. Amateurs will enjoy the wealth of life history data and will
empathize with Weast’s enthusiasm and hard work in attempting to better
understand Saturniid population biology. I enjoyed the author’s humorous
accounts of his interactions with the unsuspecting public entitled “Close
Encounters of the Other Kind”.
Understandably, the professional will find some shortcomings in this work.
The book should contain an admonition not to release non-native species into
the wild; in Weast’s defense it should be mentioned that promethea and io occur
naturally in eastern Iowa and cynthia is an introduced exotic not prone to reach
pest status on its single introduced host. The mark-recapture data and the
Lincoln index calculations are ambiguous. More information about population
structure and male mating flight might have been derived by releasing marked
individuals and setting out traps according to a standardized grid. Weast could
have benefited from a core of student helpers and some professional guidance. I
drew two basic conclusions from his population experiments: if an established,
stable population shows a 1% survival from ova to adults (2/200 ova survive),
then it is not surprizing that one has to release a large number of individuals to
establish a colony; secondly, the tentative survival of the promethea colony
illustrates the ability of Saturniids to survive as low density, dispersed
populations.
I believe this small book points out a quandary in the pursuit of Lepidoptera
studies by both the amateur and the professional biologist. These two groups
need to enter into joint studies. The professional would benefit greatly from the
energy and enthusiasm of amateurs and the wealth of life history and other data
they collect; the amateur in turn could benefit from the experience of framing
important questions, planning appropriate experiments and observations, and
evaluating acquired data.
Michael M. Collins, 11901 Miwok Path Nevada City CA 95959 USA.
(
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Title Page: All papers must have the title, author’s name, author’s address, and any titular
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citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.
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Review: All papers will be read by the editor(s) & submitted for formal review to two
referees.
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 27 Number 2 Summer 1988 (1989)
IN THIS ISSUE
Date of Publication: July 15, 1989
A Study of Protesilaus microdamas (Burmeister) and the 83
Little-known P. dospassosi (Riitimeyer) and
P. huanucana (Varea de Luque) Papilionidae)
Kurt Johnson, David Matusik & Rick Rozycki
Hand-pairing of Papilio glaucus glaucus and Papilio 96
pilumnus (Papilionidae) and hybrid survival on various
food plants
J. Mark Scriber & Robert C. Lederhouse
N ew Host Records and Morphological N otes on F our 1 04
Tortricines (Tortricidae)
Sherri Sandberg & Steven Passoa
Notes on the biology of three Riodinine species: Nymphidium 109
lisimon attenuatum, Phaenochitonia sagaris satnius, and
Metacharis ptolomaeus (Lycaenidae: Riodininae)
Curtis J. Callaghan
Portable apparatus for photographing genitalic dissections 115
Tim L. McCabe
Census of the Butterflies of the N ational Audubon Society’s 120
Appleton- Whittell Research Ranch, Elgin, Arizona
Richard A. Bailowitz
Notes on a little known ecologically displaced blue, Agriades 129
pyrenaicus ergane Higgins (Lycaenidae)
I.G. Pljushtch
Notes 135
Book Reviews 144
COVER ILLUSTRATION: Pachytene chromosomes of Brahmaea japonica see
Trentini and Marini, pages 136-138.
76 of
c oulIRNAL OF RESEARCH
THE LEPIDOPTERA
Volume 27
Number 3-4
Winter 1988 (1989)
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
ISSN 0022 4324
Published By:
/
The Lepidoptera Research Foundation, Inc.
9620 Heather Road
Beverly Hills, California 90210
(213) 274 1052
William Hovanitz
Rudolf H. T. Mattoni, Editor
Scott E. Miller, Assistant Editor, Newsletter Editor
Emilio Balletto, Italy
Henri Descimon, France
Thomas Emmel, U.S.A.
Lawrence Gall, U.S.A.
Hansjuerg Geiger, Switzerland
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Journal of Research on the Lepidoptera
27(3-4):151-159, 1988(89)
Development of the Wing Margin in
(Lepidoptera: Nymphalidae)
C.E. Dohrmann
and
H.F. Nijhout
Department of Zoology, Duke University, Durham, North Carolina 27706
Abstract. The shape of the wings of Lepidoptera is determined in the
larval imaginal disk by the position of a peripheral “bordering lacuna”.
The portion of the imaginal disk proximal to this lacuna (the wing
epithelium) will form the wing proper, while cells distal to this lacuna
(the peripheral epithelium) undergo programmed cell death during the
pupal stage. In Precis coenia, cell death in the peripheral epithelium
begins on the ventral side by six hours after pupation and gradually
spreads throughout the epithelium over the next 72 hours. After this
period of cell death the adult wing has achieved its final form and size.
The most peripheral of the scale-forming cells on the adult wing
become enlarged between 48 and 72 hours after pupation. These scale
cells will produce the fringe of long marginal scales. Transplant
experiments show that determination of these marginal scales must
have occurred prior to pupation, and thus well prior to the period of cell
death in the peripheral epithelium. We found that in P. coenia the
marginal scales do not form a discrete size group but rather are the
extremes of a gradient in scale size that extends in from the wing
margin for at least 3 scale-cell rows. We postulate that some special
property of the wing margin, presumably originating from the border-
ing lacuna but decaying with distance, is responsible for inducing the
formation of the unusually large scales that form the marginal fringe.
Introduction
The wings of butterflies and moths develop during the larval stage as
internal imaginal disks. The wing disks undergo a substantial amount
of morphological differentiation during late larval life, so that by the
middle of the last larval instar they are usually readily identifiable as
miniature wings complete with a primitive venation pattern (Nijhout,
1985). The wing veins develop initially as a system of lacunae between
the dorsal and ventral epidermal layers of the wing disk. These lacunae
radiate out from the base of the wing disk in a branching pattern that
presages the future wing venation. In addition, a peripheral bordering
lacuna develops, that runs roughly parallel to the margin of the disk.
Suffer! (1929) showed that this bordering lacuna marks the position of
the future margin of the adult wing. During the pupal stage all cells
152
J. Res. Lepid.
peripheral to the bordering lacuna disappear and only the portion of the
imaginal disk within the periphery of the bordering lacuna will become
the wing of the adult. As a consequence, the size and shape of the adult
wing is determined by the position and path of the bordering lacuna
within the wing imaginal disk. The fine features of the wing shape, such
as tails in papilionids and saturniids, as well as the gross morphology of
the wing are determined by the path of the bordering lacuna (Suffert,
1929), and any change in wing shape, whether developmental or
evolutionary, must have its basis in an alteration of the shape of the
bordering lacuna.
Thus we can think of the adult wing shape as being produced by a
cookie-cutter-like mechanism that outlines the precise form of the adult
wing within the much larger wing imaginal disk. The margin of the
adult wing is not the margin of the imaginal disk, and this has several
implications when we think about structure and function at the wing
margin. It makes us wonder about the morphology of the adult wing
edge; about whether and how the dorsal and ventral wing surfaces
become reattached at their periphery after death of the tissue distal to
the bordering lacuna. It draws our attention to the fact that the margin
of the adult wing is further “specialized” in that it bears a distinctive
fringe of marginal scales. In most species these marginal scales are
much larger, and very different in shape and color from the scales that
deck the rest of the wing surface. Thus scale morphogenesis must be
under a different type of control at the wing margin than elsewhere on
the wing. Moreover, the wing margin appears to have an additional
functional specialization. It is clear that the margin is involved in the
determination of several elements of the wing’s color pattern (Nijhout
and Grunert, 1988), and the bordering lacuna provides an obvious
structural feature that could be the source of the requisite inductive
signal(s).
The present paper reports on a morphological study of cell death at the
wing margin of the Buckeye butterfly, Precis coenia. We document the
spatial and temporal pattern of cell death, the structure of the new wing
margin, and the fact that the specialized fringe of marginal scales
appears to be induced by a special property of the bordering lacuna at
sometime prior to pupation.
Materials and Methods
Larvae of Precis coenia were reared at a constant temperature of 27
degrees Celsius on an artificial diet as described by Nijhout (1980a).
Microscopy was done on material embedded in JB-4 Resin (Poly-
sciences), sectioned at a thickness of 1 um, and stained with Lee’s
methylene blue-basic fuchsin (Polysciences). Cell death was determined
by uptake of trypan blue according to the method of Humason (1979).
27(3-4):151-159, 1988(89)
153
Fig. 1 . A. Wholemount of imaginal disk of forewing of Precis coenia at day 5
of the last larval instar, and about 2 days prior to pupation. B.
Crossection of distal portion of wing imaginal disk at day 5 of the last
larval instar. BL, bordering lacuna; W V, lacuna of wing vein; PE,
peripheral epithelium; WE, wing epithelium. Scale bar is 0.5 mm.
Specimens for scanning electron microscopy (SEM) were air-dried,
coated with gold-palladium, and examined and photographed with a
JEOL T20 electron microscope.
Results
At the time of pupation the wing has two surfaces each of which is an
epithelial cell layer of columnar, tightly packed cells. These surfaces
meet and are continuous at the periphery so that the wing’s structure
resembles a flat bag. The shape of the adult wing is marked by a
peripheral lacuna (Fig. 1) which runs roughly parallel to the margin of
the disk (Nijhout, 1985). The results presented below will show that this
bordering lacuna divides the wing surface into a “distal epithelium”
which is located distal to the lacuna and which will die during develop-
ment, and a “ wing epithelium ” which will form the adult wing (Fig. 1).
154
J. Res. Lepid.
Fig. 2. A. Crossection of pupal wing at 48 hours after pupation. B. Crossec-
tion of pupal wing at 72 hours after pupation. Former position of
bordering lacuna is shown by arrow. PC, pupal cuticle; SC, scales of
the adult wing; VPE, ventral peripheral epithelium; DPE, dorsal
peripheral epithelium; WE, wing epithelium.
Six hours after pupation the dorsal and ventral epithelial cell layers
of the pupal wing had separated and surrounded a space filled with
hemolymph and free-floating hemocytes. Although the bordering
lacuna was no longer discernable, due to the separation of the two cell
layers, its former position was still marked by the fact that the cells
formerly distal to the lacuna, the distal epithelium, were slightly
smaller and thinner than those of the wing epithelium. The boundary
between these two cell populations formed a distinct “line” in the wing
epidermis. Further studies showed that this line now defined the
position of the future wing edge. Trypan blue staining revealed that at
this time cells in the ventral part of the distal epithelium had already
started to die. Cell death began along the former position of the lacuna
and spread distally into the ventral distal epithelium. Cell thickness
and cell density in the ventral distal epithelium decreased gradually
over the next 48 hours.
At 48 hours after pupation cell death had progressed to the dorsal part
of the distal epithelium. At this time the hemolymph space within the
pupal wing was gradually diminishing in volume and the dorsal and
ventral epithelia in the proximal region of the wing had already become
fused Fig. 2A). At 72 hours after pupation the dorsal and ventral
epithelia of the wing-proper were once more tightly apposed, and scales
27(3-4):151-159, 1988(89)
155
Fig. 3. Scanning electron micrographs of wing margin of adult wing. A.
Edge-on view showing separation of dorsal and ventral cuticles. B.
View of edge showing parallel rows of sockets of scale cells. C. View of
wing margin, with a portion of scales removed, showing gradual
increase in scale length with proximity to wing margin.
156
J . Res. Lepid.
were in the process of formation (Nijhout, 1980b). In contrast, the dorsal
and ventral parts of the distal epithelium did not become fused but
remained as a thin bag-like rim around the wing (Fig. 2B). Further-
more, no enlarged scale forming cells were detectable among the
surviving cell population in this peripheral epidermis, and no scales
were formed in this part of the wing. The scale-forming cells at the very
margin of the wing were slightly larger, and had slightly larger nuclei
than those of the generalized wing epidermis. This specialization is in
accord with the presumptive relation between ploidy level and scale size
(Henke and Pohley, 1952).
Since the peripheral tissue of the imaginal disk undergoes cell death,
no closure exists between the dorsal and ventral epidermis at the
periphery of the developing adult wing. We investigated the morpho-
logy of the adult wing margin by scanning electron microscopy, and
found that the dorsal and ventral cuticles never fuse but become simply
appressed. The absence of fusion between the dorsal and ventral cuticles
is shown by the presence of a split which separates both surfaces and
which runs at or very near the wing margin Fig. 3A).
As Fig. 3B shows, the scale forming cells are arranged in straight
parallel rows which take their course roughly parallel to the wing
margin. We found that contrary to initial expectations there was no
uniquely differentiated fringe of marginal scales. Instead the size of the
scales, and the degree of indentation of their apical margin, increased
gradually with proximity to the wing margin (Fig. 3C). Scales which
were close to the margin were much larger than scales further inside the
wing. Figure 4 shows that the size of the scales decreases exponentially
with distance from the margin.
In order to determine whether cells of the peripheral epithelium were
already programmed to undergo cell death at the time of pupation, we
excised strips of epithelium extending across the bordering lacuna and
thus consisting of both distal and wing epithelium from pupae two hours
after pupation. These strips were rotated 180° and grafted back in their
original site. Thus in these grafts cells of the peripheral epithelium were
now placed within the wing epithelium and vice versa. The results of
such a graft are shown in Fig. 5. We found that the cells of the grafted
tissue always developed according to their original fate. Cells of the
peripheral epithelium underwent normal cell death, occasionally
leaving a hole in the wing epithelium at the site they were grafted. Most
frequently, however, the epithelium contracted around the wound site,
as in Fig. 5, and no hole was evident in the area where the peripheral
epithelium died. Cells of the wing epithelium survived and when placed
so that they could establish continuity with the rest of the wing
epithelium they were retained in the adult wing. These grafts retained
their original polarity as indicated by the orientation of the scales (Fig.
5). Furthermore, large marginal scales were formed at the edge of the
grafted wing epithelium, extending into the space left in the wing by
death of the transplanted peripheral epithelial cells.
27(3-4):151-159, 1988(89)
157
300
0)
a
o
cn
100
200
Distance from Margin (microns)
Fig. 4. Graph of scale length as a function of proximity to wing margin.
Position of each scale was measured from the the aperture of its
socket to the margin.
Discussion
The development of the adult wing of Precis coenia is a process with
striking morphological changes during pupal stage. As illustrated by
Figs.l and 2 the wing starts as a flat bag-like shape. At the time of
pupation it goes through a stage at which the dorsal and ventral cell
layers become separated by a voluminous hemolymph-filled space. This
separation coincides with the peak of mitotic activity and the later
stages of color pattern determination in the wing epidermis (Nijhout,
1980b; Nijhout and Grunert, 1988). After the end of the mitotic period,
about 36-48 hr after pupation (Nijhout, 1980b), the dorsal and ventral
wing epithelia become closely appressed again but the distal epithelia
never fuse. Instead, they form a fluid filled bag-like rim around the
entire wing disk (Fig. 2B). Cell death in this peripheral epithelium
begins by 6 hours after pupation and continues until at 96 h after
pupation the bag-like rim has completely vanished. Thus it is not until
the 4th day after pupation that the shape of the adult wing is evident.
Grafting experiments (Fig. 5) show that at two hours after pupation
the cells of the distal epithelium are already programmed to die in the
course of the pupal stage. The polarity of the scale forming cells has also
been determined by this time, as was previously shown by Nijhout
(1980a).
Determination of the elongated marginal scales has also occurred by
two hours after pupation, since these scales developed normally on
transplanted wing margins. Thus determination of the characteristics
of the wing margin, including its position, fate of the peripheral
158
J. Res. Lepid.
Fig. 5. Result of a grafting experiment in which a strip of pupal epidermis
was excised, rotated 180 degrees, and reimplanted so that the former
wing margin was now located well within the field of the wing proper.
Lines at right and bottom show the limits of the graft. Arrow indicates
the position of long marginal scales at the edge of the graft. In this
specimen the wing epidermis has contracted and closed off the 'hole'
left by the degenerated peripheral epidermis.
epithelium, and character of the marginal scales must have taken place
prior to pupation. The fact that cell death in the peripheral epithelium
begins at the former location of the bordering lacuna, as well as the
existence of the gradient of scale size in the wing epithelium, suggest
that the bordering lacuna not only serves as a demarcation between
wing and peripheral epithelium but may also have an active role in
inducing these two modes of cell differentiation.
The exponential increase in size of the margin al scales with proximity
to the margin strongly suggests that a gradient of some sort is a
determining factor. The transition from normal scales to very long
marginal scales occurs over the course of 3 scale rows, or a distance of
about 80 um on the adult wing. But the adult wing of P. coenia is
expanded by a factor of about 2 from the size of the wing in the pupal
stage; thus the actual distance over which marginal cell determination
gradient extends on the pupal wing is approximately 40 um. These
findings add to the body of evidence that the bordering lacuna in the
wings disks of Lepidoptera plays specific and important roles in the
development of the wing. Not only does it control the ultimate size and
shape of the wing by outlining the areas that will undergo programmed
cell death, but it also appears to possess special properties that are 1
involved in the induction of certain elements of the color pattern
(Nijhout and Grunert, 1988) and in the induction of marginal scale
differentiation.
27(3-4):151~159, 1988(89)
159
Acknowledgements. This work was supported by Grant DCB-8517210 from
the National Science Foundation.
Literature Cited
HENKE, K., & H.-J. POHLEY, 1952. Differentielle Zellteilung und Polyploedie bei der
Schuppenbildung der Mehlmotte Ephestia kuhniella. Z. Naturforsch. AbtB
7, 65-79.
NIJHOUT, H.F., 1980a. Pattern formation on lepidopteran wings: Determination
of an eyespot. Dev. Biol. 80: 267-274.
NIJHOUT, H.F., 1980b. Ontogeny of the color pattern on the wings of Precis coenia
(Lepidoptera: Nymphalidae). Dev. Biol. 80: 275-288.
NIJHOUT, H.F., 1985. The developmental physiology of color patterns in Lepi-
doptera. Adv. Insect Physiol. 18: 181-247.
NIJHOUT, H.F. & L.W. GRUNERT, 1988. Colour pattern regulation after surgery on
the wing disks of Precis coenia (Lepidoptera: Nymphalidae). Development
102: 377-385.
SUFFERT, F., 1929. Die Ausbildung des imaginalen Flugelschnittes in der
Schmetterlingspuppe. Z. Morphol. Oekol. Tiere, 14: 338-359.
Journal of Research on the Lepidoptera
27(3-4):160-172, 1988(89)
The Morpho-Species Concept of Euphyes dion with the
Description of a New Species (Hesperiidae).
John A. Shuey
Environmental Technology & Assessment, Battelle Columbus Division, 505 King Ave.
Columbus, Ohio 43201
Abstract. Euphyes alahamae (Lindsey, 1923) and Euphyes macguirei
(Freeman, 1975) fall within the normal range of variation of Euphyes
dion (Edwards, 1879), and are reduced to synonyms of E. dion. The type
series of E. macguirei probably resulted from unique rearing condi-
tions; no specimens fitting the original description of this taxon have
ever been collected in the wild. Euphyes alahamae is at best a weakly
defined subspecies in which one of the many phenotypes of E. dion
if fairly stable. A unique Euphyes population from Bay St. Louis,
Mississippi, differs subtly, but consistently, from all 12. dion populations
and is described as Euphyes hayensis new species. Preliminary bio-
logical evidence suggests that the new species differs from E. dion in
habitat requirements and hostplant choice.
Introduction
The genus Euphyes is distributed through most of the western
hemisphere and contains approximately 20 species. The genus is
composed of four well defined species groups (Shuey, 1986) which differ
primarily in the configuration of the female genitalia. The dion group
contains six species level taxa (E. dion [Edwards], E. dukesi [Lindsey],
E. macguirei Freeman, E. pilatka [Edwards], E. herryi [Bell], and E.
conspicua [Edwards]), all of which are confined to wetland habitats. An
additional taxon, E. alahamae (Lindsey), is usually considered a
subspecies of E. dion , but often has been placed as a distinct species (e.g.,
Clark and Clark, 1951; Forbes, 1960; Miller and Brown, 1981). Because
of their restricted habitat requirements, these species are localized and
among the least collected skippers in eastern North America.
The dion group (sensu Shuey, 1986) contains two problematic names
which refer to taxa of uncertain status, E. alahamae as mentioned
previously, and E. macguirei , which has remained an enigma to most
lepidopterists since its recent description. The resolution of these taxa’s
status has been hampered by their relative rarity and the difficulty of
amassing sufficiently long series to investigate interspecific variation.
My purpose is to briefly examine the status of the two problematic taxa,
and to describe a new species from southern Mississippi.
27(3-4):160-172, 1988(89)
101
Fig. 1. Wing measurements recorded; A — ■ wing length; B — extent of
orange pattern along vein V2; C — stigma length; and, D — stigma
width.
Materials and Methods
I examined material of all the dion group species. Particulary
relevant material included several hundred specimens of the dion/
alabamae complex from throughout eastern and central North America;
the holotype male of E. alabamae ; the holotype male and allotype female
of E. macguirei ; and a series of 32 males and nine females of an
undescribed taxon from southern Mississippi.
For wing pattern analysis, 20 males and 20 females were randomly
selected from series of the dion! alabamae complex from Ohio + Indiana
and Mississippi and the undescribed taxon (only nine females of this
taxon were available). Characters measured (Fig. 1) with an ocular grid
included forewing costa length (to the nearest l/2mm), extent of the
orange pattern along vein V2 (to the nearest 1 4mm), stigma length (to
the nearest l/4mm), and stigma width (to the nearest 1/Smm).
PROBLEMATIC NAMES
Euphyes macguirei Freeman, 1975. This taxon was described from a
short series (four males and one female) of reared specimens from
Benbrook Reservoir, Tarrant Co., Texas. Although Miller and Brown
162
J. Res. Lepid.
Fig. 2. Euphyes macguirei — like
adults reared from Logan Co.,
Ohio ova. Both specimens
were reared on poor quality
cuttings of Carex lacustris.
Similarities to E. macguirei
include small size (forewing
length 15 mm and 14.5 mm
respectively), and reduced
orange pattern elements.
(1981) accepted this taxon as a valid species, most lepidopterists have
been skeptical about its status, presumably because of the unique
circumstances surrounding all of the known specimens and their
similarity to E. dion. This is probably the only butterfly in North
America which has never been captured; all specimens known to me
have been reared. Freeman (1975) listed five characteristics which
separated. E. macguirei from E. dion . Unfortunately, none of these
characteristics can withstand close scrutiny.
1. “smaller size” — Indeed, the type series is composed of specimens
that are noticeably smaller than typical E. dion. However, all of the
types were reared from a locality that also supports E. dion. Since reared
specimens are often smaller than individuals that develop under
natural conditions, it seems likely that the type series might have
resulted from stressed larvae. I have produced similar-sized and pat-
terned specimens (Fig. 2) from Ohio ova by providing larvae with very
poor quality (low moisture content) cuttings of the sedge Carex lacustris
Willd.
2. “more elongated fulvous streak throughout the cell in the 6 S on
the secondaries” — The holotype male does not have a noticeably
expanded streak on the hindwing. Furthermore, the expression of this
pattern element is variable in the E. dion/ dlabamae complex, and the
variation easily encompasses the pattern observed on the E. macguirei
holotype.
3. “the absence of fulvous markings between the stigma and the base
of the wings” — Again, this is a variably expressed pattern element in E.
dion , and occasional specimens do not have any fulvous color between
the stigma and the wing base (Fig. 7).
4. “the yellowish veins on the lower surface of the secondaries, which
are absent or else poorly defined in [E.] dion ” — The veins of fresh E.
dion are always yellow and contrast strongly with the ground color.
Freeman most likely compared reared E. macguirei with flown material
of E. dion , and mistook the natural loss of scales from the veins of E. dion
as a real pattern element.
5. “it differs” . . . “in the genitalia” — Freeman’s figure of the
genitalia does differ significantly from any known Euphyes , and if it
were accurate, might deserve generic status! However, the holotype
27(3-4) : 160-172, 1988(89)
163
Figs. 3-6. Euphyes macguirei genitalia;
Fig. 3. male genitalia, lateral view; Fig. 4. aedeagus, lateral view; Fig. 5.
female genitalia, lateral view; and Fig. 6. female genitalia, ventral
view. Scale line = 2 mm.
Fig. 7. Phenotypic variation of wing pattern in Ohio and Indiana E. dion.
These specimens were selected to show the range of variability. First
row, left to right; OH., Logan Co., 16-VII-1983 (JS); OH., Williams Co.,
18-VIM954 (OSU); OH., Williams Co., 28-VM959 (OSU); IN., Steuben
Co., 12-VIL1983 (JS); OH., Portage Co., 1 1-VII-1982 (JS). Second row,
left to right; OH., Erie Co., VII-1896 (OSU); OH., Williams Co., 6-VII-
1962 (OSU); OH., Logan Co., 16-VII-1983 (JS); OH., Williams Co., 16-
VIL1955 (OSU); OH., Williams Co., 29-VL1954 (OSU). JS - J. A. Shuey
collection; OSU = Ohio State University Collection.
male possesses genitalia (Figs. 3 & 4) which do not differ in any obvious
characteristics from the normal variation found in E. dion (Figs. 22-33),
except that they are smaller. Likewise, the allotype female genitalia
(Figs. 5 & 6) are very similar to variation within E. dion (Figs. 46-57).
Because every character which Freeman used to differentiate the
taxon E. macguirei from E. dion is questionable, and because I can find
no other characters which will separate these two taxa, I suggest that#.
macguirei be considered a synonym of E. dion (Edwards). It is best to
consider the type series of this taxon the result of unique rearing
conditions.
Euphyes alahamae (Lindsey, 1923). Described from a single male as a
race of E. dion , the status of this taxon also suffered from a lack of
material. As late as 1931, Lindsey, Bell and Williams (1931) had collec-
164
J. Res. Lepid.
lively examined only one additional specimen, a female. Clark and
Clark (1951) elevated the taxon to specific status, commenting that it
exists alongside normal dion in the Dahl Swamp of Virginia and that
the orange pattern elements reliably separated these two species. They
also noted that E. alabamae flies in late July, between the broods of E.
dion. Klots (1951) considered alabamae to be a subspecies of E. dion, but
noted that “a local colony of A. d. alabamae has been recorded from Dahl
Swamp, Accomac[k] Co., Virginia; a most unusual record”, thus,
suggesting that these names may represent distinct species. Forbes
(1960) accepted E. alabamae as a distinct species, reiterating the evi-
dence presented by the Clarks. MacNeill (1975) was more cautious,
relegating alabamae to a subspecies of E. dion, but noted that °a large
geographical region of apparent overlap of these two subspecies
suggests the need for much more information concerning their relation-
ships.” Miller and Brown (1981) re-elevated E. alabamae to specific
status without comment. Most recently, Opler and Krizek (1984)
considered the taxon to fall with in the normal variation of E. dion,
citing personal communication with John Burns. John Burns (pers.
comm.) has elaborated, stating that his position was not based on
detailed investigation, but rather an inability to differentiate between
these two taxa.
Indeed, when long series of southern “E. alabamae ” are compared
with series of northern E. dion, it is evident that wing pattern variation
is rampant, and that there is no single character that will separate these
supposed taxa. In my examination of long series of the dion! alabamae
complex from throughout eastern North America for wing pattern
variability, three trends became obvious. First, northern populations
are highly variable, and range from individuals that are bright orange
(classic dion), to individuals that have greatly reduced orange pattern
elements (classic “alabamae”) (Fig. 7). These populations occur through-
out glaciated North America, and extend south along the Atlantic
Seaboard at least to Virginia. Second, populations from the Gulf Coast
States are less variable, and have reduced orange pattern elements
(Figs. 8 & 9). These populations match the concept of E. dion alabamae,
as originally intended by Lindsey (1932). Finally, I describe as a new
species one population from extreme southern Mississippi which has
non- variable expanded orange pattern elements, and a narrow stigma
(Figs. 8 & 9).
Furthermore, male and female genitalic comparisons between
northern and southern populations failed to reveal any character that
might be useful in separating these taxa. (In fact, all the species of the
dion group are close [Shuey, 1986], and even easily recognized species
such as E. dukesi are difficult to consistently separate from#. dion using
genitalic characters alone [first noted by Lindsey, 1923].)
The original description of alabamae was, by necessity, typological.
Because Lindsey was describing a single specimen, which obviously
27(3-4):160-172, 1988(89)
165
differed from his concept of dion from northern states, no intrapopu-
lational variability was considered. Once the concept of two distinct
taxa separable by wing pattern became widespread (expanded orange =
E. dion ; reduced orange = E. alabamae ), authors such as Clark and
Clark (1951) and Forbes (1960) mistook variable populations as evi-
dence for the sympatric occurrence of two distinct species. The material
I have examined from the Dismal and Dahl Swamps of Virginia, do not
substantiate any pattern of discrete broods between the two phenotypes.
Because the alabamae phenotype can be found throughout North
America and the only apparent difference between northern and
southern populations is the reduction of phenotypic variability in the
south, alabamae (Lindsey, 1923), should be relegated to a synonym ofE.
dion (Edwards).
A population from Bay St. Louis, Hancock County, Mississippi, differs
from all known populations of Euphyes , and is here described as new.
Euphyes bayensis Shuey new species
Description. Male genitalia and female genitalia variable, placing
the species within the dion group of Euphyes , but not different in detail
from E. dion . Male stigma narrower (x= 0.52 ±0.04 mm) than in E. dion
(x=0.79±0.09 mm). Facies distinctive in several respects (Figs. 8-9); in
both sexes, the orange and melanic colors are washed-out (paler)
relative to E. dion (this difference is less noticeable ventrally); in males
the forewing orange pattern elements are expanded and completely
encircle the stigma (Fig. 8); female pattern variable, but always with
conspicuous orange pattern elements on dorsal surfaces of both wings.
Size; male forewing costa = 16.63 ±0.48 mm; female forewing costa =
18.5 ±0.53 mm.
Etymology. In the tradition of Euphyes pilatka , with which it flies,
the name refers to the type locality.
Type Deposition. The entire type series was collected at Bay St.
Louis, Hancock County, Mississippi by R. Kergosien. The holotype (19-
IX-1970) and allotype (12-IX-1970) are deposited in the Carnegie
Museum of Natural History. Paratypes are deposited as follows; three
males (18-IX-1970, 10-IX-1970, 12-IX-1970), Carnegie Museum of
Natural History; two males (9-IX-1970, 4 IX 1970) and one female (8-
XX-1970), National Museum of Natural History; two males (21-IX-1970,
12-IX-1970) and one female (8-IX-1970), American Museum of Natural
History; one male (17-IX-1970) and one female (18-IX-1970), Mississippi
Entomology Museum at Mississippi State University; one male (10-IX-
1970) and one female (18-IX-1970), Mississippi Natural Science
Museum, Jackson; two males (both 19-IX-1970), The Florida State
Museum at the University of Florida; one male (27-IX-1970), The Ohio
State University; six males (two 19-IX-1970, two 10-IX-1970, 17-IX-
1970, 19-IX-1970) and one female (18-IX-1970), J.A. Shuey collection;
166
J. Res. Lepid.
and, 10 males (two 29-VIII-1970, 12-IX-1970, 17-IX-1970, three 18-IX-
1970, two 19-IX-1970, 27-IX-1970) and three females (3-IX-1970, 21-IX-
1970, 21-IX-1970), B. Mather collection. Three additional males (10-IX-
1970, 17-IX-1970, 25-V-1971) in poor condition have been returned to B.
Mather.
Discussion
My decision to describe E. hayensis as a new species is based on
morphological and limited biological evidence and as such, is open to
alternate interpretations. Although the male and female genitalia fall
within the range of variation of E. dion (Figs. 10-57), wing pattern and
stigma configuration differ consistently between E. hayensis and all E.
dion populations (Fig. 8-9). Wing pattern differences include:
1. Color. The melanic ground color and the orange pattern elements
are paler in both sexes of E. hayensis than in E. dion (Figs. 8 & 9). These
differences are most noticeable above, and are less apparent ventrally.
This color difference is real, and is not due to wear associated with flown
specimens or fading of older specimens (all of the specimens figured
were captured between 1970 and 1973).
2. Male pattern. Males of E. hayensis have consistently expanded
orange pattern elements compared to males of E. dion from Mississippi.
Euphyes dion males from the variable northern populations (Fig. 7)
commonly approach the extent of orange pattern found, but southern
populations of E. dion are less variable and are consistently dark. A
graphic plot of one pattern element, the extent of orange along forewing
vein V2 (Fig. 58) reveals the trend towards the expansion of this element
in E. hayensis.
3. Female pattern. Females of E. hayensis have consistently greater
orange pattern elements than both northern and southern populations
of E. dion (Figs. 8 & 9). The graphic plot of one pattern element, the
extent of orange along forewing vein V2 (Fig. 59), reveals that there is
no overlap of variation between E. hayensis and Mississippi E. dion.
The most compelling morphological difference is the male stigma
which is consistently narrower in E. hayensis than in populations of E.
dion (Figs. 8 & 9). This relationship is demonstrated graphically in
Figure 60. There is minimal overlap in stigma width between E.
hayensis and E. dion.
Preliminary biological evidence for the specific differentiation of
these taxa includes:
1. Habitat. The type series of E. hayensis was captured in a brackish
marsh where it flies with Euphyes pilatka. Euphyes dion has never
before been reported as a breeding resident in a brackish habitat
(although the type locality of E. alabamae, Mobile Bay, is primarily a
brackish complex) and normally occurs in fresh water wetlands; E. dion
usually flies with E. dukesi in Mississippi (C. Bryson, pers. comm).
Similar habitat differences separate other closely related pairs of
27(3-4):160-172, 1988(89)
167
wetland butterflies and may be indicative of the speciation pattern of
species that are restricted to these habitats (e.g., E. dion andE. dukesi ;
Lyceana epixanthe (Boisduval and LeConte) and L. dorcas Kirby;
Satyroides eurydice Johannson and S. appalachia (Chermock); and
Poanes viator viator (Edwards) and P. viator zizaniae Shapiro (see
Shapiro, 1970; Shapiro and Garde, 1970; Shuey, 1985).
2. Hostplant. The only known habitat is brackish, and dominated by
sawgrass. Charles Bryson (pers. comm.) could not And Carex hyalino-
lepis Steud., the hostplant of E. dion in Mississippi, in the marsh. Thus it
seems probable that E. hayensis does not use this Carex as the host.
3. Sympatry. Two specimens referable to E. dion (based on pattern)
are known from the type locality (Fig. 61) and this species is generally
distributed throughout Mississippi. The stigmas of these specimens are
intermediate between E. bayensis and E. dion (Fig. 60). This evidence
can be interpreted in two ways. I prefer to consider this as evidence of
the sympatric distribution of closely related species. Supporting this
position are; 1, the pattern and color of these two specimens which
clearly places them as E. dion ; and 2, the absence of intermediates
between E. bayensis and E. dion from Bay St. Louis. However, the
presence of these two males could also be interpreted as indicating that
one species is represented at Bay St. Louis, and that intermediate
phenotypes have simply not yet been collected.
Obviously, the biological evidence presented here needs to be con-
firmed, and additional populations of E. bayensis need to be located.
Acknowledgements. The entire type series of E. hayensis was brought to my
attention by Bryant Mather, to whom I am deeply indebted. Other material was
obtained from J.M. Burns (National Museum of Natural History); F.H. Rindge
(American Museum of Natural History); M.D. Bowers (Museum of Comparative
Zoology); J.E. Rawlins (Carnegie Museum of Natural History); J. Liebherr
(Cornell University); C. Triplehorn (The Ohio State University); J.W. Peacock
(Marion Ohio); and J.V. Calhoun (Westerville, Ohio). Charles Bryson (Stone-
ville, Mississippi) kindly permitted me access to copies of letters written by him
to B. Mather. Roy Kendall (San Antonio, Texas) provided information about E.
macguirei. Gordan R. Stairs and Richard L. Miller, The Ohio State University,
provided access to laboratory space and photographic equipment respectively.
John W. Peacock (Marion, Ohio), David C. Iftner (Worthington, Ohio), and
Bryant Mather (Clinton, Mississippi) commented on earlier drafts. Finally,
John M. Burns briefly examined part of the type series and, unknowingly,
encouraged a lepidopterist who was otherwise discouraged about describing
such a subtly differentiated taxon.
Literature Cited
CLARK, A. H. & L. F. CLARK, 1951. The butterflies of Virginia. Smithsonian Misc.
Coll. 116:1-239.
FORBES, W. T. M., 1960. Lepidoptera of New York and neighboring states. Part IV.
Cornell Univ. Agr. Exp. St a., Mem. 371:1-188.
168
J. Res. Lepid.
Figs, 8 and 9. Phenotypic variation of wing pattern in Ruphyes bayensis n. sp.
and Mississippi E. dion.
Fig. 8. Dorsal wing pattern. First Column, E. bayensis n. sp., top to bottom (all
Bay St. Louis, Mississippi.); Holotype male, 19-IX-1970; male, 17-IX-
1970; male, 17-IX-1970; Allotype female, 12-IX-1970; female, 21-IX-
1970. Second column, E. bayensis n. sp., top to bottom; male, 18-IX-
1970; male, 19-IX-1970; male, 10-IX-1970; female, 3-IX-1970; female,
10-X-1 970. Third column, £ dion top to bottom (all Mississippi); male,
Lowndes Co., 26-VI-1973; male, Lee Co., 1-IX-1973; male, Lowndes
Co., 8-IX-1973; female. Clay Co., 9-IX-1972; female, Lowndes Co., 10-
IX-1972. Fourth column, E. dion , top to bottom; male, Clay Co., 13-IX-
1972; male, Lowndes Co., 26-VI-1973; male, Lee Co., 8-IX-1973;
female, Lowndes Co., 26-VI-1973; female, Lowndes Co., 9-IX-1973.
FREEMAN, H. A., 1975. A new species of Euphyes Scudder from Texas. J. Lepid.
Soc. 29:227-229.
KLOTS, A. B., 1951. A field guide to the butterflies of North America, east of the
Great Plains. Houghton Mifflin Co., Boston. 349p.
LINDSEY, A. w., 1923. New North American Hesperiidae (Lepid.). Ent. News
34:209-210.
LINDSEY, A. W„ E. L. BELL & R. C. WILLIAMS, JR., 1931. The Hesperioidea of North
America. Denison Univ. Bull., J. Sci. Lab. 26:1-142.
MACNEILL, C. D., 1975. Family Hesperiidae. in W. H. Howe, ed., The butterflies of
North America, pp. 423-578. Doubleday Co., Garden City, New York. 633p.
MILLER, L. D. & F. M. BROWN, 1981. A catalogue/checklist of the butterflies of
America north of Mexico. Lepid. Soc. Mem. 2:1-280.
27(3-4):160-172, 1988(89)
169
Fig. 9. Ventral wing pattern. Legend as in Figure 8.
Fig. 61 . Euphyes dion from
Bay St. Louis,
Mississippi. Left to
right; 12-IX-197Q;
2-IX-1970.
OPLER, P. A. & G. O. KRIZEK, 1984. Butterflies east of the Great Plains. Johns
Hopkins IJniv. Press, Baltimore. 294p.
SHAPIRO, A. M., 1970. Notes on the biology of Poanes viator (Hesperiidae) with the
description of a new subspecies. J. Res. Lepid. 9:109-123.
SHAPIRO, A. M. & R. T. GARDE, 1970. Habitat selection and competition among
sibling species of satryid butterflies. Evolution 24:48-54.
SHUEY, J. A., 1985. Habitat associations of wetland butterflies near the glacial
maxima in Ohio, Indiana, and Michigan. J. Res. Lepid. 24:176-186.
SHUEY, J. A., 1986. The ecology and evolution of wetland butterfly communities
with emphasis on the genues Euphyes. Ph. D. dissertation, The Ohio State
LJniv., Columbus, 145p.
170
J. Res . Lepid.
Figs. 10-21. all E. bayensis n. sp., Bay St Louis, Mississippi; 10-11, 17-IX-
1970; 12-13, 1 BIX -1970; 14-15, 19-IX-1970; 16-17, 17-IX-1970;
18-19, 12-1X 1970; and 20-21, 21-IX-1970.
Figs. 22-33. all E. dion , Mississippi; 22-23, Lowndes Co., 26-VI-1973; 24-25,
Lowndes Co., 8-IX-1973; 26-27, Lowndes Co., 3-IX-1973; 28-29,
Lee Co., 1 -IX-1 973; 30-31 , Lee Co., 1 -IX-1973; and 32-33, Clay Co.,
13-IX-1973.
27(3-4):160~172, 1988(89)
171
Figs. 34-57. The range of variation of f e m a I e g e n i ta I i s of £ uphyes ba yens is n .
sp. and Mississippi E. dion (even numbers, lateral view; odd
numbers, ventral view). Scale line = 2 mm.
Figs. 34-45. All E. bayensis n. sp., Bay St. Louis, Mississippi; 34-35, 8-IX-1970;
36-37, 10-IX-1970; 38-39, 18-IX-1970; 40-41, 3-IX-1970; 42-43,
10-X-1970; and 44-45, 12-IX-1970.
Figs. 46-57. all E. dion, Mississippi; 46-47, Lowndes Co., 164X-1973; 48-49,
Lowndes Co., 17-IX-1973; 50-51, Lowndes Co., 16-IX-1973; 52-
53, Lowndes Co., 9-IX-1973; 54-55, Lowndes Co., 8-IX-1970; and
56-57, Clay Co., 9-IX-1972.
172
J. Res. Lepid.
Extent of Orange Pattern (mm)
20
1 18
16
14
E. bayensis n. sp.
Mississippi E. dion
Bay St. Louis E. dion
2 3 4 5
Extent of Orange Pattern (mm)
Stigma Length (mm)
Figs. 58-60. Comparisons of pattern and stigma variation among E. bayensis
n. sp., and Mississippi and Ohio + Indiana E. dion.
Fig. 58. Male forewing length versus extent of orange pattern along forewing
V2. Note the two specimens of E. dion from Bay St. Louis, which fall
outside of the range of variation of E. bayensis, but within the range
of variation of Mississippi E. dion .
Fig. 59. Female forewing length versus extent of orange pattern along
forewing V2.
Fig. 60. Stigma length versus width. Note that the two specimens of E. dion
from Bay St. Louis fall between the ranges of variation for E. bayensis
and Mississippi E. dion.
Journal of Research on the Lepidoptera
27(3-4):173-185, 1988(89)
The Euphilotes battoides complex: recognition of a
species and description of a new subspecies.
(Lycaenidae)
Rudolf H.T. Mattoni
9620 Heather Road, Beverly Hills, CA 90210, USA
Abstract. Euphilotes hernardino is recognized as a species separate
from E. battoides , being cited here as a new combination. A discussion
of the background for this action is given, in addition to that for
describing a new subspecies, E. bernandino g art hi, from the Isla de
Cedros, Baja California.
Introduction
The paradox of Charles Darwin lay in the title of his immortal work.
The crux of Darwin’s thesis was that evolution proceeds from the
natural selection of individual variants. Epling and Catlin (1950) were
among the first to point out that the focus on “origin of species” was
largely a result of Darwin’s being forced to frame his arguments in
taxonomic terminology because of the lack of any knowledge of genetics
in his day. They went on to conclude that “Darwin should have
emphasized his refutation of the fixity of species,” because subsequent
workers have unfortunately come to “regard the species not only as a
taxonomic category but also as an evolutionary unit.” Epling and Catlin
conclude that the study of evolutionary processes can only be accom-
plished by testing individuals because there exist only two vehicles for
adaptive change: individuals and breeding populations.
Yet taxonomy does have a vital role in permitting communication, as
lucidly pointed out by Murphy and Ehrlich (1984). Biologists simply
cannot work without “species,” regardless of the merits of the method by
which they are defined or other philosophic value they may have, real or
imagined. Indeed, the Linnean nomenclatural system works with
superb parsimony in providing an index of relationship whether based
on phenetic, cladistic, or Gestalt methodology, I believe we all concur on
the identity of the vast majority of “species,” or clusters of similar
appearance, by intuitive recognition of form (Gestalt), at least within
the limited geographical areas with which we are familiar. Species
identity has come to be “legitimized” by such elegant techniques as
chromosome analysis, allozyme quantification, comparative bio-
chemistry, statistical analysis of morphological characters, reproduc-
tive compatibility and so forth. A great deal of this work may represent
what is a sort of fallacy of misplaced interpretation. Yes, these data do
help circumscribe the “species;” and yes, these data are of indisputable
174
J. Res. Lepid.
evolutionary significance; and yes, the cladistic concept is a valid
approach to infer phylogeny; but no, this doesn’t mean the species is a
unit of evolution. Ehrlich and colleagues (Ehrlich and Raven, 1959 and
Ehrlich and Holm 1962) discussed this subject matter in depth.
A closely related problem of taxonomy has arisen in regard to the
utility of a taxonomic nomenclatorial approach to geographic variation
within species. Since the landmark work of Wilson and Brown (1953),
“subspecies,” as the unit of geographic variation, has come to be viewed
as arbitrary. The reasoned artificiality of subspecies is lack of con-
cordance among multiple characters when these characters are quanti-
fied over the geographic range of variants. Gillam (1956) performed
neat analyses of several well known polytypic butterfly species to verify
the point. More recently, Hammond (1986) brought the arguments full
circle in showing failure of concordance between both “species” and
“subspecies” in Speyeria, he implies that neither category is more or less
arbitrary than the other.
Thus, although taxonomic categories do not explain patterns of
variation, categorization is useful for describing patterns of variation.
With all organisms, application of names, at all levels, is a matter of
responsibility, and will remain inherently controversial. Naming sub-
species is no less valid than for any other category when reponsibly
applied. As a matter of even greater issue today, in the United States,
is that subspecies have assumed federal legal status under the
Endangered Species Act. The Act provides protection to subspecies of
threatened and endangered invertebrates, while uniquely variant
populations of vertebrates can be listed. Conversely, a recent decision to
not list the butterfly Speyeria callippe callippe (Boisduval) was based on
the taxonomic assertion by Arnold (1985) that the variant populations
proposed for listing did not constitute a valid subspecies. A more recent
similar controversy did the result in the listing of Euphydryas editha
bayensis Sternitzky.
Although arguments continue on both species and subspecies “pro-
blems,” many represent a sort of continuing reinvention of the wheel.
On balance there may be no more or less evolutionary information
contained in any taxonomic category. Subspecies particularly serve to
highlight patterns of variation which may be of special biological
interest.
In the following I will perform some taxonomy, first to formally
elevate a subspecies to the species level, because it makes rational
sense. Second, I am naming a new subspecies of this species because it
represents a large disjunction in appearance and is insular. In the
process, the pattern of variation of these butterflies will be reviewed.
The species of Euphilotes
The genus Euphilotes (Mattoni, 1977) was named to circumscribe a
group of five species. Later authors familiar with the group variously
27(3-4): 173-185, 1988(89)
175
recognized three (Shields, 1975, 1977), four (Miller and Brown 1981), or
five species (Tilden and smith 1986). My present interpretation of the
genus follows.
The species E. enoptes (Bdv.), battoides , (Behr), rita (B. & McD.),
pallescens (Tilden & Downey, and spaldingi (B. & McD.) constitute
five distinct morphospecies each clearly defined by several con-
cordant characters of both male and genitalia and early stage mor-
phology. The latter three species are allopatric, although spaldingi
overlaps the other two in broad distribution. The three are clearly
sister species, but should be regarded as distinct by virtue of several
unique characters states. E. mojave should be rationally regarded as a
species differentiated from E. enoptes by small but consistent differ-
ences in wing facies, female genitalia, and hostplant preference, in
addition to sympatry (with and without synchrony) of several popul-
ations with other enoptes subspecies. Certain patterns of variation,
distribution and natural history in the E. battoides set of entities imply
a single species concept here cannot palpably describe the observed
pattern of variation.
E. battoides is distinguished from its congeners at least by male and
female genetalia, egg chorion morphology (Mattoni, unpubl.), fourth
instar larva chaetotaxy and pattern (Ballmer and Pratt, 1988) and
obligate univoltinism. As with all species of Euphilotes, populations are
delimited by the spacial distribution and flowering times of their
usually specific Eriogonum hostplants (see Shields, 1975). A number of
subspecies have been described to reflect this variation. These are
summarized as follows, with hostplant data from Shields (1975, 1977)
and Pratt (unpubl.)
subspecies
distribution
flight
time
Eriogonum hostplant
battoides (Behr)
Alpine Sierra, CA
July, Aug.
lobbii, incanum, polypodum
oregonensis (B. & McD.)
Casoades, OR
July
marifolium, umbellatum
intermedia (B. & McD.)
No. CA, So, OR
July
marifolium, incanum
glaucon (W.H. Edws)
E. CA, WA, OR,
ID, MT, NV, B.C.
May-July
umbellatum, heracloides
f/avum, sphaerocephalum
comstocki (Shields)
Tehachapi Mts. CA
Aug
umbellatum
centralis (B. & McD.)
CO, UT, NM, AZ
July-Aug
umbellatum, jamesi
corymbosum
baueri (Shields)
CA, NV
May
ovaliforlium, kennedyi
bernandinio (B & McD)
S & Cent CA, Baja CA
Apr-July
fasciculatum, cinereum
martini (Mattoni)
Mojave; CA, AZ
Apr-May
fasciculatum
allyni (Shields)
El Seg. Dunes, CA
July-Aug
parvifo/ium
garthi (new)
Cedros Island, Baja CA
Mar-June
fasciculatum
ellisi (Shields)
E. CA, NV, AZ, CO, UT
July-Sept
corymbosum, heermannii,
microthecum heermannii
The relationships among the above taxa are more complex than given
and may be more accurately dealt with as several species. The model of
one monophyletic grouping exhibiting simple geographio polytypy does
not square with the data: e.g. sympatry and synchrony of battoides and
intermedia at Gold Lake, CA, sympatry and allochrony of glaucon and
baueri at Westgard Pass, CA; sympatry and allochrony of martini and
176
J. Res . Lepid.
ellisi in several mojave desert range; sympatry and allochrony of ellisi
and an undescribed taxon in northern Arizona, and the parapatry
(sympatry?) and synchrony of glaucon and bernardino at several sites
along the east slope of the southern Sierra Mevada. As an initial step in
attempting to more clearly reflect the pattern of variation in the group, I
propose the following concept:
Euphilotes bernardino (Barnes & McDonnough 1917) new combination
The species includes the cluster of four closely related taxa commonly
recognized as subspecies of battoides: bernardino (B. & B. 1917), martini
(Mattoni, 1954), allyni (Shields, 1975), and garthi (new spp.). Syn-
apomorphic characters of the species include: 1) exclusive hostplants
Eriogonum fasciculatum, E. cinerium, and E, parvofolium 2) small
mean adult size (wingspan <11.0 mm) and 3) fourth instar larva
morphology and pattern (Pratt, unpub.). The suite of variable wing
pattern characters which discriminate the four subspecies is given in
Table 1.
Although recognition of E. bernardino as a “species” is based on weak
wing characters, size and larval hostplant, and larval characters which
may not stand up to scrutiny of the many populations not surveyed, the
sympatric criterion is consistent. Future work may well discriminate
other species in the remaining “ battoides ” group. The taxon glaucon, for
example, passes the sympatry test in some localities has a large
geographic distribution, shows extensive wing pattern variation, and
uses several foodplants. Complete information on glaucon over its range
and in relation to its nearest nieghhors is not sufficient to override the
consideration of conservatism.
Classification of the populations of E. bernardino
INTRAPOPULATION VARIATION AND WING PATTERN
TERMINOLOGY
The degree of variation in wing pattern elements in adult Euphilotes
is shown in Figure 1. Pairs of specimens were selected from five series of
both E. enoptes and E. battoides to show extremes of both upper and
underside variation in both sexes. Such extremes are frequent when
dealing with series and underline the care that should be taken in
arriving at taxonomic descision in the group of butterflies.
The description of wing characters in polyommatine blues has been
very inconsistent in the past for lack of a standardized nomenclature to
apply to the various elements of pattern which repeat through the
group. Nabokov (1943) attempted to rectify the matter by suggesting a
detailed terminology. I have in large part followed his system, which is
graphically presented as figure 2. Interspaces are designated by the
named vein anterior to the space. The lower part of figure 2 diagram
matically classifies fringe types which are found throughout the tribe
27(3-4):173-185, 1988(89)
177
Fig. 1 . Intrapopulation variation exhibited by selected pairs of Euphi/otes.
Top two rows, left to right. E. enoptes ancilla , J UNS. Montana, 9
mile canyon, 20 vi 82. S. Kohler. E. pa/lescens pa/lescens. § UNS.
Nevada, Lincoln Co. 2 mi NE Hancock Summit, 24 viii 78, 0. Shields.
E. battoides centralis, cf UPS. Colorado, Chaffee Co. O'Haver Lake,
30 vi 68, R. Mattoni.
Bottom two rows. E. Battoides intermedia, cf UNS, California,
Siskiyou Co. Castle Lake, 21 vii 77, T. Dimock. E. battoides e/lisi, 2
UNS. Arizona, Coconino Co. 9 mi. E. Winona, 20 viii 79, R. & L.
Mattoni. E. battoides baueri, 2 UPS. California, Inyo Co. White Mt.
Rd. 2. mi. N. Hwy. 168. 5 vi 76, R. 8c N. Mattoni.
TERMEN
WING MEMBRANE - ■ ' >
| T f TERMINAL CILIA
SUBTERMINAL CILIA
I TERMINAL LINE (MELANIC SCALES)
TERMEN
Fig. 2. Upper. Nomenclature for wing pattern elements in Euphi/otes (and
most Polyommatine blues). Diagrammatic representation of UNH
macules and marks across Mn (cut-away) and M2. Interspaces named
for anterior vein.
Lower. Fringe types as character states at CU^
178
J. Res. Lepid.
Table 1. Compa rative data of samples of populations of Euphilotesbattoides: garthi (type series, data intext),a//y,n/(ES = CA,
Los Angeles Co., El Segundo Dunes, Chevron Refinery, 25 VII 65, Mattoni, leg.), allyni (PV = CA. Los Angeles Co.,
Palos Verdes Peninsula, Crenshaw, 27 VII 83, Mattoni, leg.), bernardino (CA, Los Angeles Co., Santa Monica Mts.,
Mulholland Dr. & Sepulveda, various dates, Mattoni, leg.), martini [AZ, Yavapai Co. 1-1 7 at Bumblebee cutoff, 17 IV 79,
Mattoni, leg.), f. = Frequency. Boldface numbers indicate character state sets unique to that subspecies. Refer to Fig.
2 for numeration.
garthi
allyni
ES
PV
bernardino
martini
Wingspread-mm
Males Mean
10.1
10.3
10.8
10.0
10.9
Range
8.7-10.9
9.4-11.0
10.0-11.5
9.3-10.5
10.0-12.3
N
11
10
10
7
11
Females Mean
9.9
10.0
10.3
9.5
10.7
Range
9.4-10.4
9.4-10.5
9.8-10.9
8.5-10.2
9.9-12.0
N
5
10
10
8
10
Males-Upperside
FW-Marginal
1.26
1.10
1.01
0.59
0.86
Bandwidth-mm
HW-f. with aurora
1.0
0.8
0.9
0.1
0.3
f. with checkered fringe M3
1.0
0
0
0
0.7
Underside
f. with halos
0
0
0
0
0.8
FW-width PM macule mm M
1.20
0.99
1.00
0.80
0.70
f. dissociated PM macs.
0
0.6
0.9
0.7
1.0
f. without marg. mac. R4+r
0.5
0.9
0.4
0.3
0.5
f. Cu2 Suffusion
0.5
0
0.1
0
0.6
HW-f. dissociated aurora
1.0
10
1.0
0.7
0.2
f. fringe type 4
0.7
0
0.1
0
0
Females-Upperside
HW-width aurora M3 mm
0.88
1.59
1.87
1.50
0.96
f. distinct marg. macs.
0.2
1.0
1.0
1.0
0.8
Underside
f. with halos
0
0
0
0
0.5
FW-width PM mac. mm M
1.43
1.09
1.13
0.93
0.85
f. with aurora
0.2
0.9
0.8
0.5
0.6
f. dissociated PM macs.
0
0.2
0.3
0.4
1.0
f. without marg. mac. R4+5
1.0
0.6
0.1
0.5
0.4
f. Cu2 Suffusion
0.6
0.7
0.7
0
0.4
HW-f. dissociated aurora
1.0
1.0
1.0
0.7
0.2
f. fringe type 4
0.8
0.1
0.1
0
0
Foodplant
fasciculatum
parvifolium
cinereum
fasciculatum
fasciculatum
Notes
1. allyni 6 homoeotic DV transposition of cyanic scales over distal part of Cu2-UNF.
2. ma/t/'n/dimorphicforanelongated tear shaped posterio-distal pointed UNF PM macule in M2 in. 3 males and .8 females and
subsequently noted in other population of martini.
and are useful characters. The cross section of the termen illustrates
how the illusion of various fringe patterns is produced. The set of
character states used in this study are given in Table 1.
Fig. 3. Subspecies of Euphi/otes bernardino . UPS, 4 specimens above, UNS
the same 4 specimens below. About 0.9 life size.
Rows 1 and 2, cf and J E. b. martini, Arizona, Yavapai Co. Bumble-
bee turnoff of 1 —17, 17 iv 79 R. Mattoni.
Rows 3 and 4, cf and J E. b. bernardino. California, Los Angeles Co.
Mulholland Hwy. various dates, May, R. Mattoni.
Rows 5 and 6, cf and $ E. b. allyni, California, Los Angeles Co.
Chevron plant, El Segundo, 25 vii 65. R. Mattoni.
Rows 7 and 8, cf and $ E. b. garthi from type series: see data in text.
27(3=4):173-185, 1988(89)
179
180
J. Res. Lepid.
Relationship of subspecies
There are 4 subspecies ofE, bernardino (Table 1). Of these, one has not
been formally described, although recognized for some time (Rindge,
1948):
Euphilotes bernardino garthi Mattoni new subspecies
Males. Fig. 3. Table 1. Distinguishable in every specimen available
from all other subspecies by 1) UPF marginal bandwidth, 2) UPH
marginal band not dissociated, 3) UNS macules, particularly the post
median (PM) set, extremely large, PM set arranged without dissociation
between interspaces, 4) UNH fringe type 4.
Females. Fig. 3. Table 1. Distinguishable in all specimens by 1) UPH
aurora not extending distally to wing margin such that marginal
macules are not differentiated, 2) underside characters as in males.
Genitalia. Indistinguishable in either sex from any member of the E.
battoides complex.
Type material. Holotype male, Baja California Norte, Isla de Cedros,
canyons west of Punta Norte, 1 IV 1983 (Faulkner and Brown).
Paratypes some locality as holotype, dates as follows: 2 8 8 30/III, 4 88
4 9$ 1/IV, 1 6 2/1 V, 1819 1/VII, 288 3/VII, all 1983 all leg. Faulkner
and Brown. 3 88 “Mexico, Cedros Island, 15/III/39” no. leg cited
(presumably F. Rindge) colln. LACMNH. Disposition of types.
Holotype, 5 male paratypes and 4 female paratypes deposited in the
SDNHM; 1 male and 1 female paratype deposited in CAS, San Francisco;
3 male and 1 female paratypes despoited in the LACMNH; 1 male and 1
female paratype deposited in the Instituto de Biologia, National
University of Mexico, Mexico City; 1 male and 1 female deposited in the
USNM, Washington.
Distribution. E. bernardino garthi is an apparent disjunct population of
the species endemic to Cedros Island. Although the indicated larval
hostplant occurs throughhout the island, the insect was only found in
March and April at low elevations in the washes and canyons of the
north end of the island, and at higher elevations in July.
Natural History. The larval hostplant in all likelihood is Eriogonum
fasciculatum Bentham with which the adults were exclusively associ-
ated. The insect also appears univoltine, with an extended emergence
taking place as the season extends altitudinally. This pattern corre-
sponds to the development of foodplant flowering which is essentil for
adult nectaring, oviposition, and larval growth. A report of Faulkner
and Brown discusses Cedros Island and its butterfly fauna in detail.
Etymology. The subspecies nomen is a patronym honoring Dr. John
Garth for his early work on the biology of Baja California and especially
Isla de Cedros.
27(3-4):173-185, 1938(89)
181
Discussion
E. bernardino garthi is an endemic of lsla de Cedros, where it
probably evolved in isolation since the eustatic sea level rise after the
last glaciation. Indications of evolutionary history might be inferred
from study of any E. bernardino populations their foodplants on the
adjacent mainland, Natividad island which was also connected to the
mainland, and San Benito island which is oceanic. Simultaneously, in-
sight might be shed on the emigration potential of Euphilotes, which is
unknown from all the Channel Islands (Miller, 1985), although Santa
Rosa and Catalina islands have populations of proper foodplants.
Quantitative data on wing characters, determined to the be variable
over the whole array of Euphilotes species, are given for the four
subspecies of E. bernardino in Table 1. Certain character states can be
used to classify all specimens of the species almost unequivocally into an
appropriate subspecies following Table 1. The single character state
which may serve to identify each taxon is the relative amount of
melanin in the underside macules. The character is expressed by the
width of the PM macule of fore wing M3 in in table 1. I illustrate the
character in a short series of specimens of each taxon in figure 3, which
also provides information on variability in wing pattern as well as other
characters. The cline of increasing darkening exhibited by each sub-
species from the desert to coastal environments is concordant with two
additional characters of the males: 1) upperside cyanic overlay and 2)
marginal band width. It must be emphasized that these dines are sharp
step dines, with the steps corresponding to the subspecies limits. E.
bernardino bernardino populations on the desert edges of the San
Bernardino and San Jacinto mountains appear somewhat lighter than
cis-montane populations, but these are not clinal in other traits which
might confuse them with martini , Scoring individuals from bernardino
colonies at Lytle Creek (south side of the San Gabriel mountains) and
Horsethief Canyon (north side of the San Bernardino range) showed
them to be statistically identical in character states to the data given in
Table 1.
The two darkest subspecies, garthi and allyni, are associated with hot
daytime weather during their flight times, with most moisture coming
from frequent dense fogs and not rainfall. The two no doubt evolved
independently under what may be similar environmental conditions,
the intervening 700 km are occupied by populations of bernardino.
The distribution of the four subspecies is shown in figure 4. The data
are largely from Shields (1977) plus a few newer records. The occurrence
of bernardino is probably almost continuous, corresponding with the
continuous distribution of its hostplant Eriogonum fesciculatum across
most of southern California. However, following the coast ranges north
of Santa Barbara the hostplant becomes increasingly disjunct. The
182
J. Res. Lepid.
subspecies martini is completely disjunct, nowhere directly meeting
bernardino. Through the desert mountains of eastern California,
southern Nevada, and western Arizona, martini occurs as a series of
isolated colonies. From central Arizona east it is more continuous as it
occurs over the belt of Eriogonum fasciculatum which grows in a band
along the south slope of the Mogollon Plateau and then ranging into
southern Arizona and probably Sonora.
The precise distributional boundaries of bernardino are not defined
where it ranges into west central Nevada. The two populations in fig. 4
were cited by Shields (1977) from Churchill Co., who made the speci-
mens available. Although highly suggestive of bernardino , they must
remain unassigned until further collections are available. The popula-
tions were associated with Eriogonum heermanii and Austin (pers
comm.) found similar populations, also on E. heermanii , in the southern
Toiyabe mountains. These represent the first documentation of Euphi-
lotes bernardino on Eriogonum heermanii.
Across the area which would provide any contact zone between
Euphilotes bernardino bernardino and E. b. martini , populations are
found as isolates in desert mountain “islands.” There are no data, other
than anecdotal, to indicate the two “blend” in any manner as suggested
by both Langston (1969) and Shields (1977). The Beatty, Nye County
(Shields, 1977) specimens were scored and completely overlap the data
given in Table 1, including dimorphism for the same peculiar PM
macule as cited. The martini population of the Providence Mountains
was also identical by the same criteria. The term “blend” is a very
unfortunate term which has found its way into wide use in the
literature. By implication “blend” is usually taken to mean the result of
gene flow causing blending of character states. However, in all but the
most rigorously tested cases, it is not possible to discriminate between
hybridization or introgression and adaptive selection along an envi-
ronmental gradient (Endler, 1977), but see Collins (1984) for a well
documented study in the Lepidoptera.
The fine grain distribution of the bernardino and allyni interface is
well established, although precise classification of the interface popula-
tion (s) is open to interpretation. E. bernardino allyni occurs only on the
historic El Segundo sand dunes, which comprised four distinct segments
prior to the urbanized destruction of southern California. It is extinct on
two segments (Mattoni, 1989). It is solely restricted to Eriogonum
parvifolium as larval hostplant, although females will oviposit on both
E. cinereum and fasciculatum in field and choice experiments. Mattoni
(unpub.) has evidence that the latter two species are toxic to neonate
larvae from El Segundo Dunes stock. Today Euphilotes bernardino
allyni is known from only three sites: 1) the 1 Ha type locality at the
Chevron refinery preserve, 2) on about 10 Ha at the Los Angeles
International Airport (LAX) dunes property and 3) on a <0.5 Ha site at
Malaga cove. The latter, at the northwest base of the Palos Verdes
27(3-4):173-185, 1988(89)
183
Fig. 4. Distribution map of E. bernardino and its subspecies discussed in text
peninsula, is isolated by one kilometer from the south where the
buckwheat Eriogonum cinereum becomes common, growing intermixed
with E. parvifolium along the seacliff. At this point there is a shift in
butterfly ecotype to a taxon which is best referred to bernardino on the
basis of natural history, although phenetically it overlaps the wing
pattern of allyni . At the higher elevations on Palos Verdes, the host-
plant occurs as pure stands of E. cinereum , but also includes a few
colonies of E. fasciculatum which grown in the canyons of the north
slope. It is not known if the butterfly feeds on the latter plant.
Examination of the topographic survey maps, and aerial and other
photographs taken prior to significant urbanization in the 1930’s
showed that the scrub communities of both the El Segundo Dunes and
184
J. Res. Lepid.
the Palos Verdes penisula were surrounded and isolated by low forb
meadows. Further, the penisula was isolated from the major dunes site.
The latter is reflected in two butterfly distributions: the now extinct
Palos Verdes Blue, Glaucopsyche lygdamus palosverdesensis Perkins
and Emmel, which evolved independently from G.l. australis Grinnell, a
still abundent species on the dunes; and the occurrence of an Apodemia
mormo virgulti (Behr) ecotype on the dunes which is absent from Palos
Verdes. These findings indicate that the Euphilotes bernardino
bernardino populations found today at Palos Verdes are relicts from
some time since the last glaciation when a continuous belt of Eriogonum
fasciculatum must have connected to th coastal sage communities to the
north. Euphilotes bernardino allyni on the other hand must have
evolved in situ and in isolation during the formation of the El Segundo
sand dunes over the past 4-6000 years.
Conclusions
The nature, meaning and proper use of species and subspecies
concepts will no doubt remain an idle and infinite speculative endeavor.
However, for the purposes of the above description of patterns of
variation of the bernardino part of the E. battoides complex, the general
application of kind (species) and reasonably concordant geographically
distributed kind (subspecies) suffices. There is no evolutionary con-
notation inherent in either category itself, although two modal sorts of
subspecies variant classes are included: bernardino and martini with
large geographic distributions and inclusion of many probable ecotypic
clusters (genetically differentiated populations adapted to local envi-
ronmental conditions) , and allyni and garthi which are highly re-
stricted endemic populations each of which may be, or recently were,
essentially panmictic.
Acknowledgements. Michael Collins, Paul Opler and Paul Hammond all
provided thoughtful and sometimes pungent critiques of this paper, much of
which is incorporated. An earlier version was read and commented upon
by Dennis Murphy, Oakley Shields, John Emmel, Gordon Pratt and Dave
Faulkner. Data were provided by the above plus Richard Bailowitz and
George Austin. John Brown and Dave Faulkner very generously provided
the type series of garthi , all existing information about it, and urged the
patronym.
Literature Cited
ARONLD, R. A., 1985. Geographic Variation in Natural Populations of Speyeria
callippe (Boisduval) (Lepidoptera: Nymphalidae). Pan Pac. Entom. 61:1-23.
BALLMER, G. & G. PRATT, 1989. A Survey of the Last Instar Larvae of the
Lycaenidae of California. J. Res. Lepid. 27:1-82.
COLLINS, M. M., Genetics and Ecology fo a Hybrid Zone in Hyalophora (Lepi-
doptera: Saturniidae). Univ. Calif. Publ. Entom. vol. 104: 93p. Berkeley.
27(3-4):173-185, 1988(89)
185
EHRLICH, P. R. & R. W. HOLM, 1962. Patterns and Popultions. Science 158: 652-657
EHRLICH, P. R. & P. raven, 1969. The Differentiation of Populations. Science
165:12281232.
ENDLER, J. A., 1977. Geographic Variation, Speciation, and Clines. Princeton
Univ. Press. NJ.
EPLING, C. & W. GATLIN, 1950. The Relation of Taxonomic Method to the Explana-
tion of Organic Evolution. Heredity 4:313-325.
FAULKNER, D. & J. BROWN, 1989. Butterflies of Isla de Cedros, Baja California
Norte, Mexico. J. Res. Lepid. 27:
GILLHAM, N. w., 1956. Geographic Variation and the Subspecies Concept in
Butterflies. Syst. Zool. 5:110-120.
HAMMOND, P. C., 1986. A Rebuttal to the Arnold Classification of Speyeria
callippe (Nymphalidae) and the defense of the subspecies concept. J. Res.,
Lepid. 24:197-208.
LANGSTON, R. L., 1969. Philotes of North America: America: A Synonymic List
and Distribution (Lycaenidae). J. Lepid. Soc. 23:49-62.
MATTONI, R. H. T., 1989. The endangered El Segundo Blue Butterfly. Unpub-
lisher.
MILLER, S., 1985 (1968). Butterflies of the California Channel Islands. J. Res.
Lepid. 23:282-296.
MURPHY, D. D. & P. R. EHRLICH, 1984. On Butterfly Taxonomy. J. Res. Lepid.
23:19-23
NABOKOV, V., 1944. Notes on the Morphology of the Genus Lycaeides (Lycaenidae:
Lepidoptera). Psyche 51:104-138
RINDGE, F., 1948. Contributions Toward a Knowledge of the Insect Fauna of
Lower California. No. 8. Lepidoptera: Rhopalocera. Proc. Cal. Acad. Sci. 4th
series 24:298-312.
SHIELDS, O., 1975. Studies on North American Philotes IV. Taxonomic and
Biological Notes, and New Subspecies. Bull. Allyn Museum No. 28. 36 pp.
, 1977. Studies on North American Philotes (Lycaenidae) V. Taxonomic
and Biological Notes, continued. J. Res. Lepid. 16-1-67.
tilden, J. w. & A. c. smith, 1986. A Field Guide to the Western Butterflies.
Houghton Mifflin, Boston.
WILSON, E. O. & W. L. BROWN, 1953. The subspecies concept and its taxonomic
application. Syst. Zool. 2:97-111.
Note added in proof: A recent paper by O. Shields and J. Reveal (1988.
Sequential evolution of Euphilotes (Lycaenidae, Scolitantidini) on their
plant host Eriogonum (Polygonaceae; Eriogonoideae). J. Linn. Soc.
33:51-91) was received after this paper was in final proof. Shields
proposed therein to elevate bernardino to species status, an action
supported by the above, with the exception of E. battoides ellisi. This
combination is illogical because of sympatry (but allochrony) with E.
bernardino martini. Consideration of ellisi as a subspecies of bern-
ardino is insupportable because of chaetotaxy (Pratt, unpublished),
foodplant, adult pattern and size, and seasonal adaptedness. These
characters firmly place it in the battoides group.
Journal of Research on the Lepidoptera
27(3-4):186-191, 1988(89)
Genetic experiments with a calverleyi- like mutation
isolated from Papilio bairdi oregonius (Papilionidae)
David V. McCorkle
Biology Department
Western Oregon State College, Monmouth, Oregon 97361
and
Paul C. Hammond
2435 E. Applegate, Philomath, Oregon 97370
Abstract. A major aberration in the wing pattern of the black Papilio
polyxenes aster ius was discovered and named calverleyi in 1864.
Recently a similar mutation was isolated in the yellow P. bairdi
oregonius. Genetic experiments suggest that this trait is inherited as a
simple Mendelian recessive, although possible deleterious effects from
this trait may increase mortality rates among the homozygotes in
certain family lines. In order to compare the oregonius mutation with
the original calverleyi phenotype, it was necessary to combine the
oregonius gene with the black wing pattern. This was accomplished by
hybridizing the oregonius stock carrying the gene with black forms of
P. bairdi, P. polyxenes asterius, and P. joanae, and successfully
producing a phenotype nearly identical to the original calverleyi
aberration.
Introduction
The Papilio machaon complex is represented in North America by
many differentiated populations that have been traditionally regarded
as distinct taxonomic species (Howe, 1975; Tyler, 1975). However, P.
indra Reakirt is the only member of this group that is particularly
distinct in morphology, including larval color pattern, pupal morpho-
logy, and adult male genitalic structure. In addition, artificial hybrids
produced by crossing P. indra with other members of the machaon
complex are apparently not viable, and did not survive beyond the first
larval instar in one experiment (Emmel & Emmel, 1964).
All other taxa within the machaon complex are partially or completely
inter-fertile (Clarke & Sheppard, 1955), and can be hybridized and
back-crossed for various genetic experiments of the type reported in the
present paper. Mating crosses are accomplished using the hand-pairing
technique described by Clarke (1952).
Nevertheless, three or four groups of populations may be recognized
as distinct biological species based upon reproductive isolation and
27(3-4): 186- 191, 1988(89)
187
ecological segregation in zones of sympatry. These are outlined as
follows.
1. Papilio machaon Linnaeus. This Eurasian species also includes
two subspecies distributed in the arctic and alpine regions of Alaska and
Canada. However, Sperling (1987) has recently documented extensive
hybrid swarms between one of the subspecies and several members of
the P. polyxenes group in central Canada. This suggests that these
groups may be regarded as conspecific, despite the divergence in wing
color pattern and allozyme patterns observed by Sperling (1987). Larval
foodplants ar e Artemisia arctica Less and Umbelliferae (Tyler, 1975). P.
machaon populations are monomorphic for the yellow color form of the
adult except in hybrid suture zones.
2. Papilio polyxenes Fabricius. This group consists of at least seven
well-differentiated subspecies or semispecies that are allopatric through-
out much of North America, extending from Newfoundland to British
Columbia and southward to Cuba and the Andes of South America.
Larval foodplants are Umbelliferae and Rutaceae. The group is poly-
morphic with both yellow and black color forms in the adult, but the
subspecies asterius Stoll used in the present experiments is mono-
morphic for the black form.
3. Papilio hairdi Edwards. This group includes four or five subspecies
that are widely distributed in the arid regions of western North
America, and are sympatric with members of the polyxenes group
throughout their distribution. Artemisia dracunculus L. is apparently
the only larval foodplant. As with the previous group, P. bairdi
populations are polymorphic in adult color. Of those used in the present
experiments, the typical subspecies consists primarily of the black form,
while the subspecies oregonius Edwards is monomorphic for the yellow
form.
4. Papilio joanae Heitzman. This is a local endemic restricted to
central Missouri. It is weakly differentiated from P. polyxenes asterius ,
but does exhibit both reproductive isolation and ecological segregation
from sympatric populations of this latter species (Heitzman, 1973).
Larval foodplants of P. joanae are restricted to certain Umbelliferae.
The species is monomorphic for the black form.
A very colorful aberration in the wing pattern of P. polyxenes asterius
was named calverleyi by Grote (1864), and was illustrated in a color
plate. In this variant, the black submarginal borders normally found in
swallowtails of the machaon complex are completely absent on both the
fore and hind wings, so that the yellow median area of the wings extends
to and fuses with the yellow submarginal spots.
The original specimen was a male captured August, 1863 on Long
Island, Queens Co., New York. A female of similar aberration type was
subsequently captured in April, 1869 near Enterprise, Florida (Mead,
1869). Both specimens were illustrated in color by Edwards (1884), and
the female was also illustrated by Holland (1899).
188
J. Res. Lepid.
Fig. 1. Top row (left) normal oregonius male, (middle) “cal” oregonius male,
(right) “cal” oregonius female with extensive orange. Middle row (left)
normal oregonius female, (middle) “cal” oregonius female, (right) normal
H3 joanae hybrid female (Jo-Or-Bd-TC 86-1 ). Bottom row (left) normal H3
asterius hybrid male (As-Or-Bd-TC 86-1), (middle) “cal” H3 asterius
hybrid male (As-Or-Bd-TC 86-2), (right) “cal” H3/'oar?ae hybrid female (Jo-
Or-Bd-TC 86-1) with extensive orange.
On July 13, 1984, D. V. McCorkle captured a near-normal female ofP.
bairdi oregonius along the Columbia River at Celilo in Wasco Co., j
Oregon. The specimen displayed an unusually large amount of orange
in the median area of the ventral hindwing. From the progeny of this
female, a brother-sister mating (Or 84-1 F-2) was performed in an
attempt to intensify the orange coloration through inbreeding. Of
approximately 40 progeny produced from the sibling mating, 33 speci-
mens were of a normal phenotype and 7 specimens were of a calverleyi -
like phenotype (abbreviated “cal”) in which the black wing borders were
completely absent (fig. 1). As a consequence, the normal black and blue
coloration of the wing borders is replaced by yellow and orange pig-
mentation. The numbers of the “cal” and normal phenotypes are not
significantly different from the 3:1 ratio that we would expect by simple
recessive inheritance for this mutation (x2p < .28). Because both of the
sibling parents must have been heterozygote carriers of the trait to
produce this ratio, the mutation must also have been carried by one of
27(3-4):186-191, 1988(89)
189
Table 1. Experimental crosses used in the production of “cal”
phenotypes.
Phenotypes
Generation Mating No. Parentage male x female normal “cal”
F2
Or 84-1 F-2
Celilo oregonius
(sibling cross)
33
7
HI
Or-Bd 85-6
bairdi x Celilo oregonius
(Or 84-1 F-2)
59
0
H2
As-Or-Bd 86-3
Or-Bd 85-6 x asterius
20
0
H2
Jo-Or-Bd 86-1
Or-Bd 85-6 x joanae
10
0
H3
As-Or-Bd-TC 86-lAs-Or-Bd 86-3 x Celilo
oregonius (Or 86-1)
46
3
H3
As-Or-Bd-TC 86-2Celilo oregonius (Or 86-3
F-2) x As-Or-Bd 86-3
4
1
H3
Jo-Or-Bd-TC 86-lJo-Or-Bd 86-1 x Celilo
38
7
oregonius (Or 86-1)
the original wild parents at Celilo, either the male or the female.
Although several other wild butterflies from Celilo were tested for this
trait by inbreeding, the “cal” phenotype did not appear in other family
lines.
Since the “cal” specimens displayed the same extensive orange
coloration on the hindwings as their normal parents and wild grand-
parent, it was thought that this trait might be linked with the “cal”
mutation, and would thus serve to identify heterozygote carriers of
“cal”. Unfortunately, subsequent crosses decoupled the orange colora-
tion from the “cal” phenotype, proving that these traits are indepen-
dently inherited (fig. 1).
Of course, the basic color background for the above “cal” mutation is of
the yellow oregonius form, rather than the black asterius form of the
original caluerleyi specimens. For those readers unfamiliar with the
genetics of the Papilio machaon complex, the black form is a simple
Mendelian dominant over the yellow form (Clarke & Sheppard, 1955).
Thus, we decided to test the hypothesis that the oregonius “cal”
mutation is similar to or identical with the original caluerleyi aberra-
tion in asterius. This was accomplished by combining the oregonius
“cal” mutation through hybridization with the black form, and pro-
ducing a black caluerleyi- like phenotype very similar to that of the
original specimens obtained by Grote and Mead in the 19th century (fig.
190
J. Res. Lepid.
1). It should be noted that the original Mead female is of the early spring
form with a well developed yellow median band and discal bar. By
contrast, our specimens are of the summer form in which these yellow
markings are mostly absent in black females.
As shown in Table 1, we crossed our Celilo oregonius stock carrying
the “cal” trait with a black P. bairdi bairdi Edwards stock that was
originally obtained from near Flagstaff, Arizona. In the first hybrid
generation (HI), all progeny were of a normal phenotype. Because we
were also conducting an unrelated experiment with these butterflies,
we crossed the HI bairdi X oregonius hybrids with P. polyxenes asterius
from Warsaw in Benton Co., Missouri. Again, the H2 progeny were all of
a normal black phenotype. Next, the H2 ( bairdi X oregonius ) X asterius
hybrids were back-crossed to the original Celilo oregonius stock (progeny
of Or 84-1 F-2). Although some of our H3 broods produced only normal
phenotypes, two crosses did yield the “cal” mutation combined with the
black phenotype, and these closely resemble the original calverleyi
specimens. We also replicated this experiment by substituting P.joanae
from Warsaw, Missouri for the asterius parent in the H2 hybrid cross,
and again obtained black calverleyi- like specimens in the H3 back-cross
to Celilo oregonius (fig. 1).
The “cal” mutation may represent some type of deletion in the genetic
information needed to produce the black wing borders in the Papilio
machaon complex. This could result from a simple point-mutation at a
control locus. However, it could also be the result of a major deletion of a
chromosomal arm, perhaps even an entire chromosome. We have not yet
attempted any karyotype studies to check this possibility. However,
“cal” homozygotes appear to exhibit various pleiotropic and/or epistatic
effects from this mutation in addition to the black wing borders. Most
“cal” individuals show reduced vigor and poor fertility, while their
normal siblings show normal vigor and fertility. As yet, we have only
obtained two larvae from a “cal” homozygote (neither survived), and all
of our breeding experiments have been conducted with heterozygote
carriers of “cal”. This has not been easy, because the carriers do not
differ in phenotype from non-carriers. Moreover, in many family lines
which produce “cal” phenotypes, there is often a sharp deficit in the
number of “cal” homozygotes. For example, in one of our H3 hybrid
back-crosses to Celilo oregonius (As-Or-Bd-Tc 86-1), 49 progeny were
obtained, but only 3 were of the “cal” phenotype. This is a very
significant deviation from the 3:1 ratio that we expected to obtain
(X2 < .003), and suggests that the homozygotes of “cal” may suffer
exceptional mortality during development. Such deleterious effects
may be expected if the mutation is the result of a chromosomal deletion.
In conclusion, the information that we have obtained in studying the
“cal” mutation demonstrates the value of conducting inbreeding experi-
ments with butterflies. First, these studies provide some insight into the
recessive genetic variation carried by natural butterfly populations as
27(3-4):186-191, 1988(89)
191
recently noted by Dimock & Mattoni (1986). Second, specific genetic
variants provide insight into how the butterfly genome is structured
and functions. In the case of the “cal” mutation, it provides us with the
knowledge that a specific part of the genome is responsible for producing
the black wing borders in the Papilio machaon complex, and that this
segment of genetic information is independent of other parts of the
genome that encode for the remaining components of the wing pattern
including general coloration, submarginal spots, black discal bars, and
the anal spot of the hindwing.
Acknowledgements. We would like to thank J. R. Heitzman and Ken Hansen
for providing breeding stock used in these experiments. In addition, David A.
West provided helpful suggestions for improving the manuscript.
Literature Cited
CLARKE, C. A., 1952. Hand pairing of Papilio machaon in February. Ent. Rec.
64:98-100.
CLARKE, C. A. & P. M. SHEPPARD, 1955. A preliminary report on the genetics of the
machaon group of swallowtail butterflies. Evolution 9:182-201.
DIMOCK, T. E. & R. H. T. MATTONI, 1986. Hidden genetic variation in Agraulis
vanillae incarnata (Nymphalidae). J. Res. Lepid. 25:1-14.
EDWARDS, W. H., 1884. The Butterflies of North America. Vol. 2, plate 11.
Houghton, Mifflin Co. Boston.
EMMEL, J. F. & T. C. EMMEL, 1964. Genetic relationships of Papilio indra and
Papilio polyxenes. J. Res. Lepid. 3:157-158.
GROTE, A. R., 1864. Description of a new species of North American Papilio. Proc.
Ent. Soc. Philad. 2:441-442.
HEITZMAN, J. R., 1973. A new species of Papilio from the eastern United States
(Papilionidae). J. Res. Lepid. 12:1-10.
HOLLAND, w. J., 1899. The Butterfly Book. Doubleday & McClure Co., New York.
HOWE, W. H., 1975. The Butterflies of North America. Doubleday & Co., Inc.,
Garden City, New York.
MEAD, T. L., 1869. Papilio (var?) calverleyi captured in Florida. Am. Nat. 3:332.
SPERLING, F. A. H., 1987. Evolution of the Papilio machaon species group in
western Canada (Lepidoptera: Papilionidae). Quaest. Ent. 23:198-315.
Journal of Research on the Lepidoptera
27(3-4):192-196, 1988(89)
The Life History of Automeris zephyria (Saturniidae)
PaulM.Tuskes
3808 Sioux Ave, San Diego, C A 92117
and
Michael J. Smith
7320 Amsterdam Ave., Citrus Heights, CA 95621
Abstract. Au tomeris zephyria occurs in the mountain ranges of central
New Mexico, and a small portion of western Texas. There is one
generation per year, and the adult flight season extends from mid-May
to mid-July. The only confirmed natural larval host plant is Salix.
Larvae were reared to maturity on Cercocarpus, Cercis, Prunus, and
Quercus. The ground color of the mature larva is primarily yellow with
a thin light blue mid-dorsal line. Pupation occurs among debris on the
ground during August and September.
Introduction
Automeris zephyria Grote inhabits the central mountain ranges of
New Mexico, south into the portion of the Guadalupe Mountains that
extends into western Texas. Since its initial description in 1882 little
information has been published on the biology of zephyria. Ferguson
(1972), reviewed the available collecting data and illustrated the adults.
Adult Automeris zephyria are attracted to lights and have been
captured from mid-May to mid-July, with most records from early to
mid-June. In June 1985, zephyria was common at lights (34 6 6, 8 9 9 ,
VI-8-9-1985) in High Rolls (Sacramento Mts., Otero Co. N.M., elev. 1993
m). The High Rolls habitat is a piny on-juniper woodland, with scattered
oaks representing elements of southwestern Madrean Evergreen
Woodland (Brown 1982, Little 1976). Most trees are under 12 m in
height and widely separated. A low growing Salix is a common member
of the riparian habitat. Other zephyria habitats in the area were visited.
Karr Canyon Picnic Ground (elev. 2430 m), 9.2 km S. of High Rolls, is |j
coniferous woodland, while Pine Campground near Cloudcroft (elev.
2690 m) is Sierran Montane Conifer Forest habitat (Brown 1982). The
mountain systems in New Mexico and Texas inhabited by zephyria
(Sangre de Cristo, Sandia, Capitan, Sacramento, and Guadalupe) have
sufficient elevation to contain the Sierran Montane Conifer Forest
community. With the exception of Sitting Bull Falls in the Sacramento
Mts. (elev. 1472 m), most locations where zephyria is collected exceed
1750 m.
27(3-4):192-196, 1988(89)
193
Figs. 1=2. Mature sixth instar larva of A u to men's zephyria : 1, lateral view; 2.
dorsal view.
At High Rolls, both sexes were attracted to lights from about one
hour after dark until observations ended at 0030 h. In captivity mating
occurred after 2330 h (N = 4). Pairs remained together until nearly
dawn and then separated. Oviposition began the following night. Eggs
were deposited in clusters on the sides of paper bags. Due to the arti-
ficial substrate the number of eggs that would be naturally deposited
in a cluster is unknown.
As with all U.S. Automeris, the ova are white, and when fertile,
develop a black dot on the top of the egg. The only confirmed natural
larval food plant is an unidentified species of Salix (Kenneth Hansen,
pers. comm.). Since 1972, we have reared larvae on four different
occasions. In captivity larvae successfully developed to maturity on
Salix sp. (willow), Cercocarpus betuloides Nutt, (mountain mahogany),
Cercis canadensis L. (redbud), and four species of Oak, Quercus phellos
L. (willow oak), Q. nigra L. (water oak), Q. oblongifolia Torr. (Mexican
Blue Oak), and Q. alba L. (white oak). Richard Peigler has reared
zephyria on Prunus serotina Ehrh. (Donahue, 1979). Upon emergence
from the eggs, larvae feed gregariously. As with other Automeris and
Hemileuca species larval clusters may divide and reunite numerous
times during a 24 hour period. After the fourth instar larvae tended to
feed singly.
Larvae mature and pupate during late August and September (N =
108). Prior to pupation they leave the host plant and construct cocoons
on the ground among debris. Based on a review of the flight data and our
rearing experience, there is one generation per year. Suggestions that
the two and one half month flight periods may represent two gener-
ations (Collins and Weast 1961, Ferguson 1972) appear incorrect.
Larval Description
The larval descriptions are based on material reared from ova deposited by a
female collected at the Tunnel Inn, High Rolls, Sacramento Mts., Otero Co.,
N.M. Twenty-three larvae from Sunspot, Otero Co. and five from Dark Canyon,
Guadalupe Mts., Eddy Co., N.M. were also reared to maturity and examined.
194
J. Res. Lepid.
Calipers were used to measure various characters at the end of each instar.
Preserved larvae are in the collection of both authors and will be deposited in an
institutional collection upon completion of larval studies.
First instar. Head: Diameter 1 mm. Brown with sparse short gold setae.
Body: Ground color yellowish green. Length 7.5 mm, width 1.5 mm. All scoli
black. Dorsal meso- and metathoracic scoli slightly enlarged and forked. Re-
maining scoli appear as simple shafts. Prolegs, true legs, sublateral and ventral
surfaces brownish yellow.
Second instar. Head: Diameter 1.3 mm. Reddish brown with sparse short
gold setae. Body: Ground color yellowish green and reddish brown. Length 11-12
mm, width 1.6 mm. Thoracic, caudal, and dorsal abdominal scoli black with
black spines. Dorsolateral scoli reduced in size with black shaft and yellow
spines. Lateral and sublateral scoli reduced in size; shafts yellow with yellow
spines. Dorsal area yellowish green with brownish red mid-dorsal line. Brownish
red intersegmental bars in line with dorsal scoli. Lateral surface reddish brown
with two thin yellowish green lines extending length of abdomen: first passes
just ventral of lateral scoli; second passes through base of sublateral scoli.
Prolegs, true legs, and ventral surface, red.
Third instar. Head: Diameter 1. 9-2.0 mm. Dark brown with short light
brown setae. Body: Ground color yellow. Length 12-15 mm, width 3 mm. Dorsal
scoli with black shafts and yellow and black spines. Dorsolateral scoli similar to
dorsal scoli but V2 the length. Lateral and sublateral scoli reduced in size; yellow
with trace of black on some spines and shafts. Segmental area yellow with black
dot between dorsal and dorsolateral scoli. Lateral intersegmental area black
and crossed by numerous yellow lines that extend length of larva: first connects
distal edge of each dorsal scolus; second passes mid way between dorsal and
dorsolateral scoli; third passes through base of lateral scoli; fourth passes
through base of each sublateral scolus. Sublateral surface black. Ventral
surface brown. Prolegs, true legs, and spiracles, reddish brown.
Fourth instar. Head: Diameter 2. 3-2. 9 mm. Reddish brown with short white
secondary setae. Body: Ground color yellow. Length 22-25 mm, width 4 mm.
Dorsal scoli elongated with black shafts; spines yellow with black tips. Dor-
solateral, lateral, and sublateral scoli similar, but latter reduced in size. Mid-
dorsal line black. Dorsal and dorsolateral surfaces yellow with three horizontal
thin black intersegmental lines. Lateral and sublateral intersegmental areas
and posterior and anterior of segments black with two thin horizontal white
lines: first touches base of each lateral scoli; second connects sublateral scoli;
both lines disrupted by yellow segmental coloration. Lateral yellow segmental
areas with small black bar between dorsolateral and lateral scoli; similar but
longer black line occurs just posterior of small black bar. Ventral surface
brownish red. Prolegs and true legs, red. Spiracles brown.
Fifth instar. Head: Diameter 3. 7-4. 2 mm. Reddish brown with short white
secondary setae. Body: Ground color yellow. Length 28-35 mm, width 7 mm.
Dorsal and dorsolateral scoli with black shafts and yellow spines; some spines
with black tips. Lateral and sublateral scoli reduced in size, with black shafts
and yellow spines. Thin mid-dorsal line bluish gray and bordered by thin black
stripe. Segmental area yellow. Intersegmental area with numerous thin hori-
zontal stripes; progressing from the outer edge of mid-dorsal line to a point even
with dorsolateral scoli, series of lines as follows: yellow, black, greenish yellow,
black, yellow, black, greenish yellow, black. Three well developed inter-
27(3-4):192-196, 1988(89)
195
segmental lines extend below these to lateral scoli: first solid white, well
developed, extending from abdominal segment one (Al) to (A8); second solid
black intersegmental patch extends from mesothorax (T2) to A9; third white
inverted “v”, connecting base of lateral scoli from A2 to A7. Sublateral and
ventral surface black with a few white pinacula. Ventral intersegmental area
black or red. Prolegs red, with red patch posterior to upper portion of leg. True
legs, red. Spiracles light brown.
Sixth instar. Head: Diameter 4. 9-5. 9 mm. Frons black, adfrontal area and
clypeus brown. Body: Ground color yellow. Length 46-57 mm, width 9-10 mm.
Dorsal, dorsolateral, and lateral scoli shaft with black tips and yellow base;
spines predominantly yellow, some with black tips. Sublateral scoli yellow. Mid-
dorsal line bluish gray and bordered on each edge by thin black stripe.
Segmental area yellow. Lateral and dorsal lateral abdominal surfaces with
yellow, black, greenish yellow and white stripes as in fifth instar. A red patch
with white pinacula and short white secondary setae occurs both posterior and
anterior of upper proleg base. Ventral surface black with white pinacula. Mid-
ventral area reddish. Prolegs and true legs red. Spiracles light brown.
The larvae of zephyria differ markedly from those of A. cecrops pamina
(Neum.). Although their shapes are similar, mature pamina larvae are light
gray-green with a few thin white, grayish, and black lines on the dorsal and
dorsolateral surfaces. On the lateral surface, pamina has two prominent
diagonal white lines that converge as they reach the lateral scoli of the
succeeding segment. In zephyria, these two corresponding white lines are
roughly parallel (Figs. 1 & 2). The ground color of the mature larva is yellow and
there are numerous prominent yellow, light green, black, and yellowish green
lateral lines present on the dorsolateral and lateral surfaces. At present, there is
no indication that these two species occur sympatrically. Automeris io neo-
mexicana B. & Benj. also occurs in New Mexico, but their larval ground color is
green with a red and white lateral line extending the length of the abdomen.
Thus, zephyria larvae are easily separated from related species. A key to the last
instar Automeris larvae of the United States (Tuskes 1986) and color larval
illustration of zephyria and io (Donahue 1979) and cecrops pamina (Packard
1914) have been published.
Acknowledgements. We thank Peter Jump, Kenneth Hansen, Jim Coleman
and Noel McFarland for records, and Michael Collins and Richard Peigler for
suggestions on the manuscript.
Literature Cited
BROWN, D.E. (EDITOR), 1982. Biotic communities of the American Southwest
United States and Mexico. Desert Plants (4)1-4. Univ. Ariz. Pub., 342 pp.
COLLINS, M.M. & r.d. weast, 1961. Wild silk moths of the United States. Collins
Radio Co., Cedar Rapids, Iowa. 138 pp
DONAHUE, J.P., 1979. Strategies for survival, the cause of a caterpillar. Terra,
17(4)3-9.
196 J. Res. Lepid.
FERGUSON, D.C., 1972. The moths of America north of Mexico. Fasc. 20. 2B
Bombycoidea (in part). Classey, London, 155-275 pp.
LITTLE, E.L., 1976. Southwestern trees — a guide to the native species of New
Mexico and Arizona. Agriculture Handbook No. 9., U.S. Dept of Agric. 109
pp.
PACKARD, A.S., 1914. Monograph of the bombycine moths of North America, part
3. Mem. Natl. Acad. Sci., 12: pp. i-ix, 1-276, 503-16, pis. 1-113.
TUSKES, P.M., 1986. The biology and immature stages of Automeris randa, and
Automeris iris hesselorum. J. Lep. Soc. 39(3)163-170.
Journal of Research on the Lepidoptera
27(3-4):197-212, 1988(89)
Three new species of Paradirphia (Saturniidae:
Hemileucinae) from Mexico and Central America with
notes on the immature stages
Claude Lemaire
La Croix de Baux, F-84220 Gordes, France
and
Kirby L. Wolfe
Entomology Department, San Diego Natural History Museum, San Diego, CA 92112
Abstract. Observation of the early stages and subsequent study of the
genitalia revealed that, in addition to P. semirosea and P. coprea, three
new species are involved in the P. semirosea complex in Mexico and
Central America. P. semirosea and P. coprea are redescribed and
lectotypes are designated. P. boudinoti and P. valverdei are described
from northeastern and southern Mexico, respectively, and P. wini-
fredae from Costa Rica and Panama. Type specimens are figured and
male and female (when known) genitalia of the five species are
illustrated. Species distribution is discussed and mapped. The imma-
ture stages ofP. semirosea , P. boudinoti and P. valverdei are described
with reference to larval food preferences in the laboratory.
Introduction
Paradirphia , with Phricodia coprea Draudt as type species, was
originally described by Michener (1949: 146) as a subgenus of Or mi-
scodes Blanchard. Nine species were later included in the subgenus by
Michener (1952: 445) in his major work on the Saturniidae of the
Western Hemisphere. Paradirphia was then cited at full generic rank
by Beutelspacher (1978, 1984) and Lampe (1986). The new status is
entirely justified based on obvious differences in the general appearance
and distinctive characters in the genitalia.
Paradirphia ranges from Mexico to Bolivia; it is mostly, if not
exclusively, a montane genus, P. geneforti (Bouvier) ranging up to
2800 m in N Ecuador. It is represented by 10 species (including the new
ones) in Mexico and Central America and three in South America where
it is an exclusive inhabitant of the Andes. This paper was initiated by
the junior author’s observation of marked variability in the larvae of
moths in which the wing pattern is so similar that all were originally
believed to be P. semirosea (Walker). Subsequent study of the genitalia
revealed that three species were involved in the reared material;
further investigations led to evidence of at least four species in the P .
198
J. Res. Lepid.
semirosea/P. coprea complex in Mexico, two hitherto unpublished. A
third new species from the same group was found among specimens from
Costa Rica and Panama.
Paradirphia semirosea and P. coprea will be redescribed prior to
descriptions of the new taxa.
Paradirphia semirosea (Walker)
(figs. 1, 2, 3, 11, 13, 16, 17, 18, 23, 24)
Dirphia semirosea Walker, 1855: 1359
Phricodia semirosea Walker; Draudt, 1930: 781
Dirphia semirosea Walker; Bouvier, 1935: 256
Dirphia semirosea Walker; Hoffmann, 1942: 243
Ormiscodes (Paradirphia) semirosea (Walker); Michener, 1952: 445
Phricodia semirosea ab. roseana Draudt, 1930: 781 (infrasubspecific name)
The species cited by Lampe (1986: 273) asP. semirosea isP. boudinoti n. sp. (see
below).
Male. Head dark brown, labial palpi three-segmented, dark brown, usually
scattered with purplish scales. Antennae pale stramineous, quadripectinate to
the apex; apical rami shorter than basal rami, those of outer side less than one-
third as long as those of the inner side of flagellum. Thorax covered with brown
to red brown hairs intermixed with longer gray hair-like scales on the tegulae;
legs dark brown, densely scattered with carmine red; epiphyses large, covered
with long hairs, about as long as two thirds of the tibia; a single subapical spur
on metathoracic tibia. Abdomen dorsally black, broadly ringed with carmine
red, ventrally dark brown with intermixed carmine scales. Forewing above
brown, more or less suffused with purplish, especially on lower half of baso-
median area and both sides of submarginal band; veins and fringes brown; lines
cream white, angled as shown in figs. 1, 2, and 3, the antemedian three-
sectioned, the postmedian usually continuous, emphasized with white dots at
the intersection of the veins. Forewing below dark brown, more or less suffused
with purplish; postmedian line straight, shaded with brown. Hindwing above
purplish, in some specimens darkened with blackish brown especially on baso-
median area; postmedian line black with two small, usually fused, subcostal
white spots. Hind wing below about same coloration as above; postmedian line
white, usually strongly contrasting, with or without white dots. The absence of
discal spots is a characteristic wing pattern feature of the P. semirosea group.
Length of forewing 30-35 mm.
Female. Antennae shortly bipectinate to the apex. Epiphysis absent.
Ground color usually lighter than in male with an extension of the lightest areas
tending toward purplish pink or pink instead of purplish. Averaging larger than
the male; fore wing 35-40 mm.
P. semirosea is the most variable of the five species studied. Pink forms were
named by Draudt (1930:781) aberration roseana. Provisions of the Code do not
apply to this name published at an infrasubspecific rank (Art. 45a). The
markings also vary, especially the postmedian lines, above and below.
27(3-4):197-212, 1988(89)
199
Male genitalia (figs. 18A, B, 23A). Uncus down-curved apically, simple,
slightly notched at the apex. Valves very broad, trilobed, the lower portion of the
proximal lobe connected to the transtilla; a very strong spine, posteriorly
produced, arising from the inner side of the valvula. Lateral arms of the
transtilla medially fused in a strongly sclerotized sub trapezoidal ventral plate.
Juxta deeply concave, broadly fused to the anterior portion of the valves with
lateral sides posteriorly produced as strongly sclerotized processes. Aedeagus
straight; the vesica has a strong hook-like cornutus.
Female genitalia (fig. 18C). Sclerotization of the eighth sternum post-
vulvar with medial portion posteriorly prominent and laterally fusing to the
eighth sternum and to the anapophyses. Sclerotization of the eighth tergum
bilobed. Ductus bursae chitinized; pyriform bursa moderately bulky; ductus
seminalis arising from the right side very close to the ductus bursae. Post-
apophyses slightly longer than the anapophyses. Ovipositor well developed,
covered with fine setae.
Types. P. semirosea was described by Walker (1855: 1359) from one male
and one female. The male is hereby designated as the lectotype.
Lectotype: One male, locality unknown (43-58 = presented in 1853 by E.
Doubleday Esq.) (genit. preparation D. Goodger) (British Museum, N. H.)
(examined).
Distribution (fig. 24). MEXICO. VERACRUZ: 62 mi (100 km) SW of
Nautla, 1290 m; Las Minas, 1385 m; Naolinco de Victoria; Orizaba, 1243 m.
CHIAPAS: San Cristobal de Las Casas, 2160 m; Oxchuc; 11 mi (18 km) W of
Ocosingo, 1375 m; Pinola; Santa Rosa Comitan; Zapalota (= La Trinitaria); Las
Delicias. OAXACA: 5 2 mi (84km) NE of Guelatao, 1400m, El Portillo del Rayo,
1450 m. GUATEMALA. ALTA VERAPAZ: Mpio. San Cristobal Verapaz,
Hacienda Baleu, 1850 m; Coban, 1200 m; BAJA VERAPAZ: Pantic, 1600 m;
Santa Elena (La Cumbre); NE of Volcan Acatenango, 2200 m. COSTA RICA.
CARTAGO: Tapanti, 1400 m; Moravia de Chirripo, Platanillo, 1150 m; id.,
Tausito, 1200 m; Cantina de Rio Macho, 1200 m.
P. semirosea is the most widely distributed species of Paradirphia in
Central America where it is mainly recorded from moderate elevations from
1100 to 1500 m. The absence of records between Guatemala and Costa Rica is
probably due to lack of collecting rather than to a gap in distribution.
Immature stages. See under group heading, also figs. 11, 13, 16, 17.
Material examined. Large series from the above cited localities; 22 speci-
mens dissected.
Paradirphia coprea (Draudt)
(figs. 9, 10, 19, 24)
Phricodia coprea Draudt, 1930: 781
Dirphia coprea Draudt; Bouvier, 1935: 258
Dirphia coprea Draudt; Hoffmann, 1942: 243
Ormiscodes (Paradirphia) coprea (Draudt); Michener, 1949: 146
Ormiscodes (Paradirphia) coprea (Draudt); Michener, 1952: 445
Male. Antennae noticeably shorter than in P. semirosea, stramineous. P.
coprea differs fromP. semirosea mainly by the more uniform, duller brown of the
200
J. Res. Lepid.
wings above and below as a result of less contrast between the light and dark
areas, especially on the baso-median and the postmedian areas of the forewing.
The markings are as in P. semirosea , but the postmedian line is less continuous,
tending to fade between the dots on the veins. Forewing (lectotype) 38 mm.
Female. Same coloration and markings as in male. Forewing (paralecto-
type) 38 mm.
Male genitalia (fig. 19A, B). Differ from those of P. semirosea by the much
shorter lateral sides of the juxta which are not posteriorly produced in strongly
sclerotized processes, and in the extreme reduction of the cornutus. However, in
some specimens doubtfully referred to this species (see distribution) a small
hook-like cornutus is present.
Female genitalia (fig. 19C). The single specimen examined (paralectotype)
presented a weaker structure than in P. semirosea with the bursa much smaller;
the anapophyses are noticeably shorter than in the previous species.
Types. P. coprea was described by Draudt (1930: 781) according to three
pairs from Cuernavaca, Mexico, all in his own collection. As the latter was
destroyed during the Second World War, there was little hope of finding
syntypes to identify this species with certainty. A search among museums
where Draudt’s type material is occasionally found (Museum fur Naturkunde
der Humboldt-Universitat zu Berlin, British Museum (N.H.), Musee d’Histoire
naturelle de la ville de Geneve) was unsuccessful, but one male and one female
syntypes were located in the American Museum of Natural History, specimens
from the collection of the late Frank Johnson who probably purchased them
from Draudt or Niepelt. Several other types of Saturniidae from the same source
are likewise preserved. The male syntype is hereby designated as the lectotype,
the female as paralectotype. Both were examined, and although old and
somewhat faded (figs. 9, 10), still show the main characters of the wing pattern.
Lectotype: One male, Mexico, Morelos, Cuernavaca, VI. 1912, n° 263, genit.
preparation (in glycerine) C. Lemaire, n° 5217 (coll. Draudt, coll. Frank
Johnson, American Museum of Natural History). Paralectotype: one female,
same locality and collections, VII. 1909, n° 264, genit. preparation C. Lemaire,
n° 5218.
Distribution (fig. 24). MEXICO: Type locality. The distribution as reported
by Hoffmann (1942: 243): “Tierra templada de la cuenca superior del Rio Balsas,
Morelos, Sierra Vole. Transversal (hasta 2000 m). Jalisco” may refer to several
different species. Specimens from the following localities are doubtfully identi-
fied as P. coprea : STATE OF MEXICO, Malinalco. GUERRERO, vicinity of
Acuitlapan, 10 mi (16 km) NE of Taxco, 5000 ft (1524 m). OAXACA, Candelaria
Loxicha, 550 m. In these specimens, unlike the lectotype, the vesica has a small
hook-like cornutus. Although they do not otherwise differ, their identification
will remain uncertain until additional topotypical material is available for
comparison.
Immature stages. Unknown.
Material examined. Lecto- and paralectotype and seven questionable
specimens; all dissected.
Paradirphia valverdei Lemaire & Wolfe new species
(figs. 4, 12, 15, 20, 24)
Male. Antennae more rusty yellow than in both previous species. Long hair-
like scales of tegulae black, little intermixed with gray. Ground color of wings
27(3-4):197-212, 1988(89)
201
above and below much darker brown than in P. coprea, almost black, with
shades of purplish brown scarcely contrasting and confined to both sides of
submarginal band on forewing and postmedian area of hindwing. Lines pure
white and very contrasting; postmedian of forewing reduced to dots on the veins.
Fore wing length: 37-40 mm (holotype = 40 mm).
Female. The very damaged single known example (progenitor of the larvae)
was unfortunately lost. It was collected at light in Veracruz, 62 mi (100 km) SW
of Nautla, 1290 m, 1. VIII. 1984, K. Wolfe, M. Valverde.
Male genitalia (fig. 20). A much larger structure than inP. coprea. Shape of
valves differing with inner lobe (fused to the transtilla) larger, and posteriorly
oriented spine noticeably stronger. Lateral sides of juxta more prominent than
in P. coprea but not as much as in P. semirosea , and less sclerotized as in
latter; connection of juxta to anterior protion of valves differs from P. semirosea
(figs. 18, 23 A). Cornutus minute or entirely lacking from vesica.
Types. Holotype: male, Mexico, Oaxaca 52 mi (84 km) NE of Guelatao,
1400 m, 27. VII. 1987, K. Wolfe, M. Valverde, D. Mullins. Paratypes: two males,
same locality, data and collectors; five males, 53 mi (85 km) NE of Guelatao,
1475 m, 30.VII.1984, K. Wolfe, M. Valverde; one male, Oaxaca, 45 mi (72 km)
NE of Guelatao, 2000 m, 29. VII. 1984; four males, Oaxaca, 54 mi (87 km) SW of
Tuxtepec, 1260 m, 25. VII. 1984, same collectors; one male, Veracruz, 62 mi (100
km) SW Nautla, 1290 m, 3. VIII. 1984, same collectors.
The holotype and three paratypes are in the collection of the Museum national
d’Histoire naturelle, Paris, five paratypes are in the collection of the junior
author, one para type is in the San Diego Natural History Museum, one paratype
each will be deposited in the Natural History Museum of Los Angeles County,
the American Museum of Natural History, the Allyn Museum of Entomology,
Sarasota, Florida, and the Coleccion Entomologica del Instituto de Biologia de
la Universidad Nacional Autonoma de Mexico.
Distribution (fig. 24). P. valverdei is known only from the above cited
localities at moderate elevations in Oaxaca and Veracruz where it is sympatric
and synchronic with the more numerous P. semirosea.
Immature stages. See under group heading, also figs. 12, 15.
Material examined. 14 males; seven dissected.
This species is named after Marvin D. Valverde to show gratitude for his
contribution to the collecting and the rearing of the material studied.
Farudirphm boudinoti Lemaire & Wolfe new species
(figs. 7, 8, 14, 21, 24)
Male. Antennae straw yellow. Like P. coprea and P. valverdei , this species
differs from P. semirosea by the reduction of contrast between light and dark
areas on wings above and below. Coloration not as dark as in P. valverdei, and
one specimen from Mexico, Coahuila, Saltillo, with the purplish as contrasting
as in lectotype ofP. semirosea. Lines cream white as inP. semirosea , postmedian
of forewing usually reduced to vein-dots but, in some specimens, as continuous
as in P. semirosea. Forewing 33-36 mm (holotype = 35 mm).
Female. Slightly larger than male, same markings and coloration. Fore-
wing (allotype) 40 mm.
Male genitalia (fig. 21 A, B). Distinguished from three previous species by
hypertrophy of inner lobe (fused to transtilla) and much longer spine of valves.
Lateral sides of juxta not prominent as in P. coprea . Most characteristic feature
202
J. Res. Lepid.
Table 1. Larval description of Paradirphia. P. semirosea, P. boudinoti, P. valverdei, sixth (last) instar.
P. semirosea
P. boudinoti
P. valverdei
Head (0.5 mm)
orange, adfrontal sutures
black
coral pink, adfrontal
sutures black
blue green, adfrontal
sutures black
Integument:
— Dorsal and lateral areas
red-brown, speckled
whitish
yellow, densely covered
with black vermiculations
light green, broadly but
indistinctly checkered
with red
— Ventral area
duller, similar
similar
bluegreen
Dorsal band
broad, dark brown
almost inconspicuous,
white
greenish white
Subdorsal, upperand
lower sub-spiracular lines
white
greenish white
white
Spiracularband
dark brown
denser black markings
copper red
Subspiracularband
white
white
white
Thoracic legs
orange
orange
pale green
Abdominal legs
brown
black
blue green tipped with
black
Anal plate
pinksurrounded with
black
orange
blue-green
Paranal lobes
pink surrounded with
black
orange thinly surrounded
with black
blue green surrounded
with black
Spiracles
orange surrounded with
black, then white
chestnut brown circled
with black, then orange
white
orange circled with white
Scoli
rusty orange brown,
prothoracic spines, dorsal
and subdorsal spines of
abdominal segment 9
black
orange, spines orange
interspersed with black
green, prothoracic
spines, dorsal and
subdorsal spines of
abdominal segments
black
of armature is wedge-like shape of aedeagus; vesica lacking cornutus.
Female genitalia (fig. 2 1C). Differing from P. semirosea and P. coprea by
less prominent medial portion of ventral plate. Narrow pre vulvar belt present in
both dissected specimens. Bursa noticeably bulkier than in P. semirosea .
Types. Holotype: male, Mexico, Tamaulipas, Gomez Farias, Rancho del
Cielo, 1127 m, 8-1 l.V. 1985, J. Boudinot. Allotype: female, same locality, dates
and collector. Paratypes: eight males, one female, same locality, dates and
collector; 18 males, San Luis Potosi, Ciudad del Maiz, El Platanito, Torre
Forestal, 1160 m, 26.VII-1.VIII.1984, E. C. Welling (all in the Museum national
d’Histoire naturelle, Paris); two males, San Luis Potosi, 16 mi (26 km) E of
Ciudad del Malz, 1140 m, 2. VII. 1983, K. Wolfe, M. Valverde (K. Wolfe
collection); one male, San Luis Potosi, El Naranjo, 5. VIII. 1975, T. W. Taylor
(Natural History Museum of Los Angeles County); five males, five females,
Puebla, Villa Juarez; four males, Puebla, San Juan Apulco (Allyn Museum of
Entomology, Sarasota). One paratype each will be deposited in the American
Museum of Natural History and in the Coleccion Entomologica del Instituto de
Biologla de la Universidad Nacional Autonoma de Mexico.
Distribution (fig. 24). P. boudinoti is widely distributed in NE Mexico from
Coahuila to Puebla, its southernmost range meeting the northernmost of P.
semirosea. Additional records are from COAHUILA, Saltillo, 1599 m, and
HIDALGO, 70 mi (113 km) S of Tamazunchale, 1700 m. P. boudinoti probably
lives in drier areas than P. semirosea.
27(3“4):197-212, 1988(89)
203
Immature stages. See under group heading, also fig. 14.
Material examined. 32 specimens; 11 dissected.
This species is named after Jacques Boudinot of the Department of Entomo-
logy of the Museum national d’Histoire naturelle, Paris, to express gratitude for
the collecting of a great part of the type material during his mission to Mexico in
July and August 1985.
Paradirphia winifredae Lemaire & Wolfe new species
(figs. 5, 6, 22, 23, 24)
Male. Antennae straw yellow. Red scales on thorax and legs, dorsal rings on
abdomen paler pink, not carmine as in four previous species. Ground color dark
gray brown as in P. boudinoti; purplish zones on forewings confined to
postmedian area, especially between postmedian line and submarginal band
Fig. 1. Paradirphia semirosea male, Mexico, Veracruz, Las Minas, 1385 m,
ab ovo, Escondido, California, on Robinia pseudoacacia.
Fig. 2. Paradirphia semirosea female, Chiapas, 11 mi. (18 km) W. of
Ocosingo, 1375 m, 21 .VII. 1987 (K. Wolfe, M. Valverde, D. Mullins).
Fig. 3. Paradirphia semirosea male (extreme pink phase), Chiapas, 15 mi
(25 km) W of Ocosingo, 1325 m, 10.VIII.1985 (K. Wolfe, M. Valverde).
Fig. 4. Paradirphia valverdei new species, paratype male, Mexico, Oaxaca,
52 mi (84 km) NE of Guelatao, 1400 m, 27.VIII.1987 (K. Wolfe, M.
Valverde, D. Mullins).
Fig. 5. Paradirphia winifredae new species, paratype male, Costa Rica,
Cartago, Tapanti, 1540 m, 10. VII. 1988 (K. Wolfe, M. Valverde).
Fig. 6. Paradirphia winifredae new species, paratype female, Costa Rica,
Cartago, El Empalme, 2000 m, 6. IV. 1978, (K. Wolfe, M. Valverde).
Fig. 7. Paradirphia boudinoti new species, paratype male, San Luis Potosi,
16 mi (27 km) E of Cd. Maiz, 1140 m, 2.VII.1983 (K. Wolfe, M.
Valverde).
Fig. 8. Paradirphia boudinoti new species, allotype female, Mexico,
Tamaulipas, Gomez Farias, Rancho del Cielo, 1127 m, 8/11.1985 (J.
Boudinot) (Museum national d'Histoire naturelle, Paris).
Fig. 9. Paradirphia coprea lectotype male, Mexico, Morelos, Cuernavaca,
VI. 1912 (American Museum of Natural History).
Fig. 10. Paradirphia coprea paralectotype female, same locality, VII. 1909
(American Museum of Natural History).
Fig. 1 1 . First instar larvae of Paradirphia semirosea (similar to first instar of
P. valverdei and P. boudinoti ), Mexico, Chiapas, 18 mi (29 km) W of
Ocosingo, ab ovo, Escondido, California, on Robinia pseudoacacia.
Fig. 12. Fourth instar larva of Paradirphia valverdei (similar to fourth instar
of P. semirosea and P. boudinoti ), Mexico, Veracruz, 62 mi (100 km)
SW of Nautla, 1290 m, ab ovo, Escondido, California, on plum.
Fig. 13. Larva of Paradirphia semirosea sixth (last) instar, Mexico, Chiapas,
11 mi (18 km) W of Ocosingo, 1375 m, ab ovo, Gordes (France) on
Robinia pseudoacacia.
Fig. 14. Larva of Paradirphia boudinoti sixth (last) instar, Mexico, Hidalgo, 70
mi 8 of Tamazunchale, 1700 m, ab ovo, Escondido, California, on
Robinia pseudoacacia.
Fig. 15. Larva of Paradirphia valverdei sixth (last) instar, same data as fig. 12.
Fig. 16. Eggs of Paradirphia semirosea, Mexico, Chiapas, same data as fig.
13.
Fig. 17. Pupae of Paradirphia semirosea, same data as above.
II
204
J. Res. Lepid.
27(3-4):197-212, 1988(89)
205
206
J. Res. Lepid.
where they form a contrasting, well delineated band from costa to inner margin.
Lines less conspicuous than in all previous species, postmedian usually reduced
to white vein-dots, the one on the inner margin line being as small as those on
veins Culb to Ml. P. winifredae averages larger than P. semirosea. Forewing
34-39 mm (holotype — 39 mm) versus 31-35 mm in examined P. semirosea males
from Costa Rica.
Female. Larger and darker than male, with purplish brown zone on fore-
wing proximal to submarginal band less conspicuous. White vein-dots on ante-
and postmedian lines especially small, except subcostal. Forewing (allotype)
40 mm.
Male genitalia (figs. 22 A, B, 23B). Resembling those of P. semirosea in
having lateral sides of juxta well prominent. Differing in shape of valves and in
connection of inner portion of valves to juxta, as shown in fig. 23. Vesica has
strong hook-like cornutus as in P. semirosea.
Female genitalia (fig. 22C). Same structure as in P. semirosea.
Types. Holotype: male, Costa Rica, Cartago, Tapanti, 1660 m, 22. VIII. 1984,
F. Beneluz (genit. preparation in glycerine, C. Lemaire, n° 5075). Allotype:
female, Costa Rica, Puntarenas, Monteverde, Rio Guacimal, Nuboso, 1550 m,
8. IX. 1983, J.-M. Cadiou, W. Haber (genit. preparation in glycerine, C. Lemaire,
n° 5102). Paratypes: two males, Cartago, Tapanti, 1600 m, 8.VI., 15. XII. 1985, F.
Beneluz, Museum national d’Histoire naturelle, Paris; one male, Cartago,
Tapanti, 1540 m, K. Wolfe, M. Valverde; one female, Cartago, El Empalme,
2000 m, 6.VIIII.1978, same collectors (collection of junior author); two males,
Alajuela, Volcan Poas, 2350 m; one male, Cartago, 16 km S of Cartago on Pan
American Highway, 1800 m; one male, Puntarenas, Monteverde, 1300 m; two
males, San Jose, Parque Nacional Braulio Carillo, Estacion Zurqul, 1500 m
(collection of the University of Pennsylvania, Philadelphia); 15 males, Panama,
Santa Clara de Chiriqul, 1600 m, 5. VI. 1968, C. Moinier (Museum national
d’Histoire naturelle, Paris). One paratype each will be deposited in the Natural
History Museum of Los Angeles County, the American Museum of Natural
History, and the Allyn Museum of Entomology.
Distribution (fig. 24). In addition to the above cited localities, there are
records from Panama, Chiriqul, road from Gualaca to Fortuna, km 32, Hornito,
1000 m; El Hato del Volcan, Quebrada Tisingal, 1400 m; Boquete, Alto Quiel,
1700 m. Although occurring in neighboring areas, P. winifredae and P.
semirosea are probably allopatric or only occasionally sympatric. In Monteverde,
Costa Rica, where both species occur in the same area of montane rainforest, P.
winifredae has been collected at higher elevations than P. semirosea.
Immature stages. Unknown.
Material examined. 36 specimens; 18 dissected.
This species is named after Winifred Hallwachs, for her contributions to the
knowledge of the Saturniidae of Parque Nacional Santa Rosa, Guanacaste,
Costa Rica.
Immature stages (P. semirosea , P. boudinoti , P. ualverdei )
Egg (fig. 16). Diameter ca. 1.5 mm, yellow to greenish yellow,
becoming gray about five days before hatching.
Larva (Figs. 11-15). Length ca. 2.5 mm (first instar) to 65-70 mm
27(3-4):197-212, 1988(89)
207
(last instar). There are six instars (five molts) and spination and pattern
of markings are typically hemileucine. The arrangement of scoli is as in
Leucanella leucane (Geyer) (see Lemaire, 1971: 30), and is as follows:
Thoracic segments, abdominal segments 1, 2, 7 bear four pairs
(subdorsal, prespiracular, upper and lower subspiracular); abdominal
segments 3-6, three pairs (lower subspiracular absent); abdominal
segment 8 has subdorsal pair fused into a single dorsal scolus + three
pairs as in abdominal segment 7 ; abdominal segment 9 has dorsal scolus
as in segment 8, but removed to posterior end of segment + three pairs
(upper subdorsal, lower subdorsal, subspiracular); abdominal segment
10 has paranal scoli present. In the first instar, subdorsal and pre-
spiracular pairs on thoracic segment and dorsal scolus on abdominal
segments 8 and 9 are apically forked.
There are usually distinctive generic characters in the larvae of
Hemileucinae, such as the rosette-type dorsal scoli in Hemileuca and
the hypertrophied upper subdorsal scolus of abdominal segment 9 in
Periphoba (see Gardiner, 1982: 145, P. arcaei , figured as “P. hircia”).
The most characteristic features in the larvae of Paradirphia studied
are 1) the absence of obviously predominant scoli, and (2) the slightly
longer subdorsal and prespiracular scoli on the prothoracic segment and
longer lower subdorsal pair on abdominal segment 9. Structure of the
different groups of scoli is unusually indistinct.
There are distinctive specific characters in the larvae of Paradirphia
which were studied, especially in the color of the integument; sixth
instar larvae of P. semirosea, P. valverdei, and P. boudinoti are respec-
tively orange and brown, green and red, and yellow and black.
Lampe (1986: 273) described the immature stages of P. boudinoti
logically referring to them as P. semirosea , unaware of the features
which gave rise to this study.
A comparative description of the 6th instar larvae of P. semirosea , P.
valverdei , and P. boudinoti is given in table 1.
Pupa (fig. 17). Unlike most of the Hemileucinae, the larvae of the
species of Paradirphia reared do not spin cocoons. Before pupation, the
larvae leave the plant in search of a pupation site. There are no traces of
silk in the pupal chamber formed in the soil at a depth of 10 cm or more.
The pupa of P. semirosea is ca. 30 mm long, black and smooth, with
thoracic segments rounded. Cremaster is simple, prominent, bearing a
tuft of strong hooks at the anal end.
Larval hostplant preferences (in the laboratory). P. semirosea
preferred Robinia pseudoacacia (Leguminosae) over a variety of other
plants offered in California and France; P. boudinoti accepted plum
( Prunus : Rosaceae) in France, plum and R. pseudoacacia in California,
and Malus (Rosaceae) in Germany; P. valverdei preferred plum in
California. Native host plants are unknown.
208
J. Res. Lepid.
Fig. 18. Genitalia of Paradirphia semirosea . A. Male, aedeagus removed,
ventral view; B. Aedeagus, lateral view; C. Female, ventral view.
Scale line = 1 mm.
Fig. 19. Genitalia of Paradirphia coprea. A. Male (lectotype), aedeagus
removed, ventral view; B. Aedeagus, lateral view; C. Female (Para-
lectotype), ventral view. Scale line = 1 mm.
27(3-4):197-212, 1988(89)
209
Fig. 20. Male genitalia of Paradirphia va/verdei new species. A. Aedeagus
removed, ventral view; B. Aedeagus, lateral view. Scale line = 1 mm.
Fig. 21. Genitalia of Paradirphia boudinoti new species. A. Male, aedeagus
removed, ventral view; B. Aedeagus, lateral view; C. Female, ventral
view. Scale line = 1 mm.
210
J. Res. Lepid.
Fig. 22. Genitalia of Paradirphia winifredae new species. A. Male, aedeagus
removed, ventral view; B. Aedeagus, lateral view; C. Female (allo-
type), ventral view. Scale line = 1 mm.
Fig. 23. Juxta and anterior portion of the valves in genitalia of Paradirphia. A.
P. semi rosea; B. P. winifredae. Scale line = 1 mm.
27(3-4):197“212, 1988(89)
211
Fig. 24. Geographical distribution in Mexico and Guatemala of the species of
Paradirphia studied.
Acknowledgements. We thank F. H. Rindge (American Museum of Natural
History) for the loan of the syntypes of P. coprea of which the identification was
thus made possible. A. Watson and D. Goodger (British Museum (Natural
History) kindly dissected the genitalia and confirmed the identification of the
lectotype of P. semirosea. Equally we thank H. J. Hannemann (Museum fur
Naturkunde der Humboldt Universitat zu Berlin), J. Donahue (Natural History
Museum of Los Angeles County), D. H. Janzen, (Department of Biology,
University of Pennsylvania) and L. and J. Miller (Allyn Museum of Entomology
of the Florida State Museum) for the loan of specimens. F. Beneluz, J.-M.
Cadiou, W. Haber, W. Hallwachs, D. H. Janzen, R. Lampe, C. Moinier, D.
Mullins, T. Porion, L. Schwartz, M. Valverde and N. Venedictoff have either
collected or reared the studied material. Thanks also to R. S. Peigler and two
anonymous reviewers for their helpful comments and suggestions.
Literature Cited
BEUTELSPACHER, C. R., 1978. Familias Sphingidae y Saturniidae (Lepidoptera) de
Las Minas, Veracruz, Mexico. An. Inst. Biol. Univ. Nal. Auton. Mex.,49, Ser.
Zook (1): 219-230.
BEUTELSPACHER, C. R„ 1984. Una nueva especie mexicana del genero Para-
dirphia Michener (Lepidoptera: Saturniidae). An. Inst. Biol. Univ. Nal.
Auton. Mex., 54 (1983), Ser. Zook (1): 123-127.
212
J. Res. Lepid.
BOUVIER, E.L., 1935. Etude des Saturnioides normaux, famille des Hemileucides,
deuxieme partie. Ann. Sc. nat., Zool., (10)18 (2eme vol.): 217-418.
DRAUDT, M., 1929-1930. 12. Familie Saturnidae [sic] in Seitz, A., Die Gross-
Schmetterlinge der Erde, 6 (Die amerikanischen Spinner und Schwarmer):
713-827. A. Kernen, Stuttgart.
GARDINER, B. O. C., 1982. A silkmoth rearer’s handbook, 3rd edition. The Amateur
Entomologist, 12, XII + 255 pp.
HOFFMANN, C. C., 1942. Catalogo sistematico y zoogeografico de los lepidopteros
mexicanos, Tercera Parte. Sphingoidea y Saturnioidea. An. Inst. Biol.,
Mexico, 13: 213-256.
LAMPE, R. E., 1986. Die Praimaginalstadien von Paradirphia semirosea Walker,
1855 (Lep.: Saturniidae). Ent. Z., 96(19): 273-288.
LEMAIRE, C., 1971. Revision du genre Automeris Htibner et des genres voisins,
Biogeographie, Ethologie, Morphologic, Taxonomie (Lep. Attacidae) (lere
partie). Mem. Mus. natl. Hist, nat., (N.S.), serie A, Zool., 69, 232 pp.
MICHENER, C. D., 1949. New genera and subgenera of Saturniidae (Lepidoptera).
J. Kansas Ent. Soc., 22(4): 142-147.
MICHENER, C. D., 1952. The Saturniidae of the Western Hemisphere, Morphology,
Phylogeny, and Classification. Bull. Amer. Mus. Nat. Hist., 98(5): 335-502.
WALKER, F., 1855. List of the specimens of lepidopterous insects in the collection
of the British Museum, 6:1259-1507. London, by order of the Trustees.
Journal of Research on the Lepidoptera
27(3-4):213-221, 1988(89)
A List of the Butterflies and Skippers of Mount
Revelstoke and Glacier National Parks, British
Columbia, Canada (Lepidoptera)
David L. Threatful
P. 0. Box 190, Revelstoke, British Columbia, Canada VOE 2SQ1
Abstract. An annotated list of 63 species of butterflies and skippers
found in Mount Revelstoke and Glacier National Parks, British
Columbia, Canada, has been complied. Eight additional species are
considered to be likely additions to the known fauna, and one previous
record to be a mislabelled European specimen. The alpine species
present on the two highest peaks near Revelstoke are also listed.
Introduction
Mount Revelstoke and Glacier National Parks are located west of the
Rockies in the Columbia Mountains of southeastern British Columbia,
Canada. The Monashee, Selkirk, and Pucell Ranges form the portion of
the Columbia Mountains in the vicinity of the parks. Mount Revelstoke
National Park (M.R.N.P. henceforth) encompasses 25,900 hectares and
is located in the western part of the Selkirk Mountains. It is approxi-
mately bounded by latitudes 51°00'-51°15' North, longitudes 117°50'-
118°15' West. Glacier National Park (G.N.P. henceforth) encompasses
135,000 hectares and is located in the Selkik and Purcell Ranges, with
the Beaver River Valley separating the two ranges in the park. G. N. P.
is bounded by latitudes 51°00'-51°30' North, longitudes 117°10'-118°00'
West. The town of Revelstoke lies at the southwest edge of M.R.N.P. on
the Columbia River between the Monashee and Selkirk Ranges.
Other than a small amount of collecting by Mark Hobson and John G.
Woods, no previous study has been conducted on the butterflies of these
parks. The reason for the paucity of collecting is probably the inhospit-
able climate and terrain. The mountains rise precipitously from the
deep valley floors (450 m) up to a maximum of 3387 m elevation. The
only access roads are the Trans-Canada Highway (which skirts
M.R.N.P. and bisects G.N.P.) and a road to the summit of Mt. Revelstoke.
Many of the mountain trials are long and strenuous to climb. Dense
coniferous forest (Columbia Forest 456-1220 m, subalpine forest above
1220 m) covers most of the area below treeline, with the only breaks in
the forest being due to cliffs, streams, avalanches, and the activities of
man. Treeline is at about 1800-2000 m or occasionally higher, above
Annual precipitation in the parks is generally 150-200 cm, but parts
of G. N. P. receive up to 350 cm. Much of the precipitation occurs in the
1Current Address: 1501 32nd Street, #14, Vernon, British Columbia V1T 5K4
214
J. Res. Lepid.
winter, resulting in heavy snow packs which are slow to melt. Summers
are generally warm with frequent cool rainly intervals. At the lower
elevations (Columbia River Valley floor) there are only a little over 4
frost-free months (127 days), with the area near the town of Revelstoke
(elevation 456 m) being somewhat warmer and drier (100-150 cm) than
the rest of the park area. Much of the butterfly fauna resident in the
Rocky Mountains (Banff and Jasper National Parks, Alberta) is missing
from this area, probably due to the high precipitation and late spring
combined with the lack of open habitats below treeline.
Data for this report were compiled from collections made in 1980,
1981 and 1983 and general observations from 1965 to 1979. All species
reported were collected by the author under the authority of a Parks
Canada volunteer agreement. The Biosystematics Research Institute,
Ottawa, Ontario, Canada, confirmed the identification of all specimens
to species level. A representative collection has been placed in the parks
collection at Glacier Park nature center and selected specimens re-
tained by the Biosystematics Research Institute. Specific collection data
have been placed in the fauna files maintained by the park naturalists
at Glacier Park nature center. Sixty-three species are confirmed for the
parks. An additional eight are possible additions. A species list for the
alpine areas of Mt. Cartier, Selkirk Range, and Mt. Begbie, Monashee
Range (both 12-13 km south of Revelstoke) is given at the end. Mt.
Begbie was collected 7 August 1983 and Mt. Cartier on 4 August 1983.
These specimens are in the collection of John H. Shepard. Experience
with the British Columbia butterfly fauna leads me to believe that the
majority of the species regularly occurring in the parks are now
documented.
The scientific names employed generally conform to those used in
Howe (1975). The subspecies designation should be treated with caution
because the taxonomic status of many species in this area is only poorly
known.
For each species, the abundance, habitat, altitudinal range, flight
period, and park(s) in which it is found are given. Five terms are used to
describe abundance:
(1) Common: a species usually encountered every day in numbers,
(2) Uncommon: a species encountered on most days usually in small
numbers,
(3) Rare: a species of which few are encountered, and encounters are
infrequent during a year,
(4) Extremely rare: a species not seen most years with few records for
any given location,
(5) Local: a species known only from restricted localities and habitats.
May be common or rare depending upon the circumstances.
The descriptions of habitats and elevations are based on observation
within the parks. Flight seasons are normally fairly constant, although
during inclement weather emergence may be delayed by two or three
weeks. There are a few species that appear to be found in only one of the
27(3-4):213-221, 1988(89)
215
parks. Question marks indicate that certain species may occur in both
parks, although thus far having been found only in one park.
Systematic Account
Hesperiidae (Latreille 1809)
1. Thorybes pylades (Scudder 1870): Abundance variable, rare to
uncommon and local; forest edges, clearing, and roadsides up to 550 m;
mid-May through June; M.R.N.P. only.
2. Erynnis icelus (Scudder and Burgess 1870): Common: open forest
edges and roadside clearings; 456-914 m; May to early June; both parks.
3. Pyrgus centaurae loki (Evans 1953): Local and uncommon; dry
alpine tundra and occasionally moist tundra; 2042-2134 m; July; both
parks (Selkirk and Purcell Ranges). This species and Hesperia comma
are found at higher elevations than any other Hesperiidae within the
parks.
4. Pyrgus ruralis (Boisduval 1852): Rare to uncommon; dry grassy
clearings, gravel road shoulders with short grass, and dry open areas
where Dryas grows; 456-549 m; late April through June; M.R.N.P.
only(?).
5. Carterocephalus palaemon mandan (W. H. Edwards 1863):
Uncommon to common; wet grassy bogs close to forest edges; 457-
1219 m; June and July; both parks.
6. Thymelicus lineola (Ochsenheimer 1808): Rare; open grassy areas
such as fields and roadsides; one record for M.R.N.P. at 549 m in June.
Appeared at Sicamous, B. C. 72 km west of Revelstoke several years ago
and is rapidly spreading in all directions. The population within the
parks is still expanding. The nearest collection site outside the parks
was 15 km south of Revelstoke at 456 m in June 1981.
7. Hesperia comma manitoba (Scudder 1874): Rare and local; open
subalpine forests, clearings, bogs and rockslides near timberline; 1859-
2042 m; July and early August; both parks (Selkirk and Purcell
Ranges).
8. Polites themistocles (Latreille 1824): Rare; grassy openings near
forest edges and fields up to 610 m; June to early July; M.R.N.P. only.
9. P. mystic (W. H. Edwards 1863) ssp.: Rare; grassy openings near
forest edges and fields up to 549 m; June and early July; M.R.N.P. only.
10. Ochlodes sylvanoides (Bosiduval 1852) ssp.: Very common;
forest openings and edges, roadsides; up to 549 m; late July to mid-
September; M.R.N.P. only.
11. Amblyscirtes vialis (Edwards 1862): Uncommon; clearings,
forest edges, and along forest roads up to 762 m; late May to early July;
M.R.N.P. only.
Papilionidae Latreille 1809
12. Papilio zelicaon zelicaon (W. H. Edwards 1852): Common to
uncommon; forest edges, alpine meadows, mountain tops, riparian
areas; 456-2438 m; mid-May to August; both parks.
216
J. Res. Lepid.
13. P. glaucus canadensis (Rothschild and Jordan 1906): Common;
forest edges, clearings, riparian areas, and open areas generally; p up to
1219 m; late May to mid-July.
14. P. rutulus rutulus (Lucus 1852): Extremely rare, three records
for district; one record from G.N.P. (habitat, date, and elevation un-
known), one record from Revelstoke (456 m) and one record in Rogers
Pass (1300 m). The main blend zones for P. rutulus and P. glaucus are
further west in the Okanagan-Shuswap Districts and south at the north
end of Kootenay Lake.
15. P. eurymedon (Lucas 1852): Uncommon; riparian and open areas
close to open forest edges and clearings, sometimes in association with
Ceanothus velutinus ; up to 640 m; late May to mid-July; M.R.N.P. only.
Pieridae Duponchel 1832
16. Neophasia menapia tan (?) (Scudder 1861): Uncommon; near
forest edges; 518-2042 m; mid-July to mid-September; M.R.N.P. only(?).
17. Pieris occidentalis occidentalis (Reakirt 1866): Uncommon to
common; roadsides, open forest edges, clearings, dry alpine tundra; 456-
2438 m; late April to October; both parks (Selkirk and Purcell Ranges).
At low elevations the early spring specimens are darker and sometimes
smaller than summer specimens. At high elevations there is only the
summer form present (mid- July to mid- August).
18. P, napi (Linnaeus 1758) ssp.: Common to uncommon; clearings in
dense forest and forest edges along roadsides; 456-1859 m; late April to
mid- August; both parks. At low elevations the summer brood is usually
lighter than the spring brood. At high elevations only the darker form
appears to be present.
19. P. rapae (Linnaeus 1758): Uncommon to common; widespread,
but mostly near human habitations; 456-549 m; late April to early
October, M.R.N.P. only (?).
20. Anthocaris sara Lucus 1852 ssp.: Common; open forest edges,
roadsides, and clearings; 456-640 m; late April to early July; M.R.N.P.
only (?).
21. Colias philodice eriphyle W. H. Edwards 1876: Common; open
areas such as roadsides, clearings, forest edges, and dry areas of alpine
meadows; 456-1829 m; late April to October; both parks.
22. C. eury theme Boisduval 1852: Rare to uncommon; roadsides,
fields, clearings, open forest edges; 456-762 m; mid-July to October;
M.R.N.P. only. This species is probably a migrant to this area since it is
not seen every year.
23. C. nastes streckeri Or u m Gr i sc him ai 1 o 1985: Uncommon and
local; barren mountain ridges and dry alpine tundra; probably mid- July
to mid- August; one record for 2499 m on Dawn ML, Purcell Range,
G.N.P.
24. C. pelidne minisni Barnes 1895: Extremely rare; one record at
2438 m in G.N.P. on a ridge north of Dawn Mt. on the border of G.N.P.
(could be a wind blown stray from lower down), 17 August 1981.
27(3-4):213-221, 1988(89)
217
25. Lycaena cupreus henryae (Cadbury 1937): Rare and local; wind
swept barren ridges and rockslides; 2017-2134 m; mid-July to late
August; G.N.P. on the ridges near Dawn Mt. (Purcell Range) and on
Avalanche Crest (Selkirk Range).
26. L. helloides (Boisduval 1852): Common, local in G.N.P. ; open
areas such as roadside, forest edges, and fields; 459-909 m; mid-May to
September; both parks. At least double-brooded at Revelstoke, probably
single-brooded in most of the park areas.
27. L. mariposa Reakirt 1866 ssp.: Uncommon to common; forest
clearings, edges of bogs, riparian areas, moist clearings near trails; 488-
1463 m; late June to late August; both parks.
28. Satyrium acadica colinensis (Watson and W. P. Comstock 1920):
Common; forest edges in association with Salix ssp., clearings, and
riparian areas; 456-914 m; late June to early September; both parks.
Material from this area is of uncertain affinity, but is closest to acadica
coolinensis (J. H. Shepard, in litt.).
29. Callophrys spinetorum (Hewitson 1867): Extremely rare; one
record at 946 m along a roadside (Trans-Canada Highway) forest edge in
early July in G.N.P.
30. C. rosneri rosneri K. Johnson 1976: Rare to uncommon; damp
roadsides close to forest edges; up to 457 m; mid-May to mid-June;
M.R.N.P. only.
32. C. augustus iroides (Boisduval 1852): Common; roadsides close
to forest edges and in clearings; 456-762 m; late April to early June;
M.R.N.P. only.
32. C. eryphon eryphon (Boisduval 1852): Common; open forest
edges close to roadsides; 456-914 m; late April to early June; both parks.
33. E veres amyntula albrighti Clench 1944: Uncommon to rare;
open forest edges around clearings, sometimes attracted to damp spots
along roadsides; 456-549 m; mid-May to mid-June; M.R.N.P. only.
34. Celastrina argiolus nigrescens (Flectcher 1903): Common; open
forest edges, damp forest roads, clearings, and riparian areas; 456-945
m; mid- April to early July; both parks. This is the first species to appear
in the spring in the parks, other than those which overwinter as adults.
Eliot and Kawazoe (1983) consider nigrescens to be a hybrid population
between spp. lucia and echo.
35. Glaucopsyche lygdamus Columbia (Skinner 1917): Uncommon;
clearings, avalanche paths, forest edges, mountain meadows; 456-1829
m; late April to mid-August; both parks.
36. Lycaeides idas atrapraetextus (Field 1939): Rare; along road-
side gravel banks close to forest edges; 731-1311 m; late June to mid-
August; M.R.N.P. only(?). The species name is idas , rather than
argyrognomon (Berstrasser), as a consequence of I.C.Z.N. Opinion 269
and work by L. G. Higgins (C. D. Ferris, in litt.).
37. Plebejus saepiolus arnica (W. H. Edwards 1863): Common;
roadsides, open forest edges, bog edges, damp grassy meadows, fields;
456-1219 m; June to August; both parks.
218
J. Res. Lepid.
38. Agriades rustica megalo W. H. Edwards 1927): Rare, occasion-
ally locally common; rockslides, barren rocky ridges, open subalpine
forest edges; 1981-2164 m; mid-July to early September; both parks
(Selkirk and Purcell Ranges). A. franklinii is a low elevation, coastal
arctic species with Leguminoseae foodplants. A. rustica is a montane
species which feeds on Saxifraga (C. D. Ferris, in litt.).
Nymphalidae Swanison 1827
39. Speyeria atlantis beani (Barnes and Benjamin 1926): Common;
mountain meadows, forest edges, bog edges, clearings; 456-1676 m; late
June to early August; both parks. The commonest and most variable
Speyeria in area.
40. S. hydaspe sakuntala (Skinner 1911): Uncommon to common;
damp places along forest edges and riparian areas, mountain meadows
and subalpine forest clearings; 456-1829 m; early July to September;
both parks.
41. S. mormonia opis (W.H. Edwardes 1874): Uncommon; mountain
meadows adjacent to open subalpine forests; 1219-1981 m; July to early
September; both parks.
42. Boloria selene atrocostalis (Huard 1927): Uncommon and
local; edges of wet grassy bogs and meadows; up to 917 m; late May to
mid-August; G.N.P. only.
43. B. epithore chermocki (E. and S. Perkins 1966): Common; open
forest edges, edges of bogs, clearings, riparian areas, mountain
meadows; 456-1920 m; late May to mid-August; both parks.
44. B. astarte astarte (Doubleday and Hewitson 1847): Rare, local;
barren windswept ridges and scree slopes; 2438-2621 m; mid- July to
mid-August; Dawn Mt. summit (Purcell Range) and Avalanche Crest
(Selkik Range) in G.N.P. , Mt. Williamson (2045 m); in M.R.N.P.
45. Phyciodes tharos (Drury 1773) ssp.: Common; roadsides, fields,
clearings, forest edges; 456-914 m; late May to early August; both parks.
Subspecific status uncertain.
46. P. campestris campestris (Behr 1863): Uncommon; roadside
clearings, open forest edges; 456-945 m; late May to early August; both
parks.
47. P. mylitta mylitta (W.H. Edwards 1861): Extremely rare; dry
roadsides near clearings, forest edges, open fields; 456-457 m; mid-May
to late September; M.R.N.P. only. Also found 19 air km. south of
Revelstoke in a field in the Akolkolex River area at 456 m (May 18,
1970). Possible two broods south of Revelstoke, and at least one brood in
the park.
48. Euphydryas anicia anicia (Doubleday and Hewitson 1848):
Common; mountain meadows, rockslides, subalpine forest edges, clear-
ings, ridges; 1859-2073 m; late June to early September; both parks
(Selkirk and Purcell Ranges).
49. Polygonia satyrus (W.H. Edwards 1869): Common; open forest,
damp forest roads, forest edges, and riparian areas; 456-1219 m; in
27(3-4):213-221, 1988(89)
219
flight late March to October, adult overwinters; both parks. Subspecies
‘ neomarsayas ’, sometimes attributed to this area, is probably not a valid
subspecies but simply a form (C.D. Ferris, in litt.).
50. P. faunus rusticus (W.H. Edwards 1874): Common; open forest,
damp forest roads, forest edges, and riparian areas; 456-1829 m; in flight
late March to October, adult overwinters; both parks.
51. P. zephyrus (W.H. Edwards 1870): Uncommon; open forest edges,
mountain meadows, subalpine clearings; 456-1981 m; in flight late
March to October, adult overwinters; both parks. Found at higher
elevations and visits flowers (Compositae) more frequently than the
other two Polygonia species in this area.
52. Nymphalis vau album watsoni (Hall 1924): Uncommon at times;
forest edges along damp roadsides; 456-945 m; in flight late March to
October, adult overwinters; both parks. There are major population
fluctuations every few years.
53. N. californiea herri Field 1936: Rare; open forest edges, road-
sides, clearings; 456-549 m; in flight late March to October; M.R.N.P.
only(?). Migrates into area. Adult probably does not overwinter in the
parks.
54. N. antiopa antiopa (Linneaus 1758): Common; riparian areas,
forest edges, clearings, and damp forest roads; 456-1036; in flight late
March to October, adult overwinters; both parks.
55. N. milberti milberti (Godart 1819): Common; mountain mea-
dows, forest edges, riparian areas, clearings; 456-1981 m; in flight late
March to October, adult overwinters; both parks.
56. Vanessa cardui (Linnaeus 1758): Common during some years;
open sunny areas, clearings, meadows; 456-1524 m; May to October,
adult does not overwinter, but instead migrates into the area some
years; both parks.
57. V. annabella (Field 1971): Rare to uncommon in some years;
roadsides close to forest edges; 456-1219 m; May to October, adult
probably does not overwinter, but instead migrates in during some
years; both parks. Not seen every year.
58. Vaqnessa atalanta rubria (Furhstorfer 1909): Rare to uncommon
during some years; open forest edges, riparian areas, clearings, 456-
1067 m; May to October, adult probably does not overwinter, but instead
migrates in some years; both parks. Not seen every year.
59. Limenitis lorquini burrisoni (Maynard 1891): Common; ripa-
rian areas, damp forest roads, and forest edges; 456-1097 m; late- June
to early August; both parks. Seldom seen visiting flowers but comes to
moisture, manure, and mud.
Satyridae Boisduval 1833
60. Cercyonis pegala boopis (Behr 1864): Locally common; grassy
clearings along forest edges; one locality at 549 m along the main trail to
the summit of Mt. Revelstoke, mid- July to late August, M.R.N.P. only.
61. Oeneis chryxus chryxus (Doubleday and Hewitson 1849): Rare
220
J. Res. Lepid.
and local; bases of rockslides close to grassy clearings, edges of sub-
alpine forests and rocky alpine draws; 1524-1981 m; July to mid-
August; G.N.P. (Selkirk and Purcell Ranges) only(?).
62. Oeneis melissa beani Elwes 1893; Rare and local; rocky screes
and barren windswept short grass ridges; 2134-2438 m; late June to late
July; G.N.P. (Selkirk and Purcell Ranges) only(?).
Danaidae Duponchel 1844
63. Danaus plexippus (Linnaeus 1758): An extremely rare migrant;
one record at 457 m in August, 1973 on the south edge of M.R.N.P.; one
record at Revelstoke in July 1957.
Possible additional species
There are a few additional species which might occur in the parks,
either because they have been collected elsewhere in the Revelstoke
District or because suitable habitats have not been completely
sampled.
64. Erynnis persius fredericki H. A. Freeman 1943: This species
could enter G.N.P. through the Beaver River valley. A mid-May to early
July flight period would be expected.
65. Polites coras (Cramer 1775): Another species which could enter
G.N.P. through the Beaver River valley. A mid-June to August flight
period would be expected.
66. Parnassius phoebus smintheus Doubleday 1847: One specimen
was taken 13 km south of Revelstoke in June 1980 at 456 m, and six
specimens were taken on Mt. Cartier at 1615 m 4 August 1983. One
female taken 37 km southeast of Revelstoke on the Akolkolex Forestry
Road at elevation 677 m on 19 July 1983. Nomenclature follows that of
Ferris (1976). This species may occur in the parks where its foodplant
(, Sedum spp.) grows in open rocky areas.
67. P. multicaudatus (W. F. Kirby 1884): A single specimen taken in
the Akolkolex Valley 30 km southeast of Revelstoke in July 1970 at 640
m.
68. Euptoieta claudia (Cramer 1775): One specimen seen but not
collected on Dawn Mtn., Purcell Range, G.N.P. at 2499 m 12 August
1981.
69. Boloria euphrosyne (Linnaeus 1758): Jones (1951) listed this
species as occurring in the Revelstoke area. B. euphrosyne is not found
in the Nearctic, his record is a mislabelled European specimen (J. H. ,
Shepard, in litt.).
70. Limenitis arthemis rubrofasciata (Barnes and McDunnough
1916): One specimen collected 13 km south of Revelstoke at 456 m on
June 23, 1982, near the junction of the Old South Highway and the
Akolkolex Forestry Road. Currently in the collection of J. H. Shepard.
71. Erbia epipsodea epipsodea Butler 1868: Apparently absent
from the parks, but might occur in G.N.P. in Grizzly Creek area at 1890-
2075 m.
27(3-4):213-221, 1988(89)
221
72. Oeneis jutta chermocki Wyatt 1965: Suitable habitat is present
in the Beaver River Valley in G.N.P. at 945-1219 m. A likely flight
period would be late June to mide-July. May be biennial and missed on
the off years.
Alpine species found on Mt. Begbie and Mt Cartier
1. Pyrgus centaurae loki (Evans 1953): 2316 m on Mt. Begbie. This
species and L. cupreus may also occur on Mt. Cartier.
2. Hesperia comma manitoba (Scudder 1874): 2316 m on mt.
Cartier.
3. Pieris occidentalis occidentalis (Reakirt 1866): 2286-2408 m on
Mt. Begbie; 2469 m on Mt. Cartier.
4. Colias nastes streckeri (Grum-Grischimailo 1895): 2194 m on
Mt. Begbie; 2347 m on Mt. Cartier.
5. Lycaena cupreus henryae (Cadbury 1937): 2225 m on Mt. Begbie.
6. Agriades rustica megalo (W. H. Edwards 1927): 2225-2732 m on
Mt. Begbie; 1615-2316 m on Mt. Cartier.
7. Boloria astarte astarte (Doubleday and Hewitson 1847): 2225-
2732 m on Mt. Begbie; 2408-2610 m on Mt. Cartier.
8. Euphydryas anicia anicia (Doubleday and Hewitson 1848): 2316
m on Mt. Begbie; 1646-2286 m on Mt. Cartier.
9. Oeneis chryxus chryxus (Doubleday and Hewitson 1849): 2316
m on Mt. Begbie; 2225-2316 m on Mt. Cartier.
10. Oeneis melisa beani Elwes 1893: 2732 m on Mt. Begfbie; 2469
m on Mt. Cartier.
Acknowledgements. I would like to express my sincere appreciation to John G.
Woods, Chief Park naturalis of Mount Revelstoke and Glacier National Parks.
Mr. Woods suggested the projecct and provided encouragement throughout the
study, as well as helping me to produce the final report for Parks Canada
(Western Region). He also kindly supplied helicopter access to some of the
mountains. The staff of the Biosystematics Reseach Institute, especially Dr. D.
H. Kritsch, were most supportive. Dr. Clifford D. Ferris encouraged me to seek
journal publication of this report, and offered valuable information and cri-
ticism of previous drafts of this manuscript. Jon H. Shepard assisted in specimen
identification, as well as giving encouragement. I am particularly grateful to
Crispin S. Guppy. Without his assistance in revising the original Parks Canada
report this publication would not have been possible.
Literature Cited
ELIOT, J. N. & A. KAWAZOE, 1983. Blue Butterflies of the Lycaenopsis-group.
British Museum (Natural History), London. 296 pp.
FERRIS, C. D., 1976. A Proposed revision of Non-arctic Parnassius phoebus
Fabricius in North America (Papilionidae). J. Res. Lep. 15(l):l-22.
HOWE, WILLIAM H., ed., 1975. The Butterflies of North America. Doubleday and
Co., Inc., Garden City, N.Y.
JONES, J. R. J. LEWELLYN, 1951. An Annotated Check List of the Macrolepidoptera
of British Columbia. Ent. Soc. B.C. Occasional Paper No. 1.
Journal of Research on the Lepidoptera
27(3-4):222-232, 1988(89)
Hybridization of the Mexican tiger swallowtail, Papilio
alexiares garcia (Lepidoptera: Papilionidae) with other
P . glaucus group species and survival of pure and hybrid
larvae on potential host plants
J. Mark Scriber1
Mark H. Evans
and
Robert C. Lederhouse1
Department of Entomology, University of Wisconsin, Madison, WI 53706
Abstract. Mexican tiger swallowtails, Papilio alexiares garcia were
collected in Nuevo Leon and Tamaulipas. Males and virgin females
were hybridized with other P. glaucus group species. Crosses with P.
glaucus had normal egg viability and a 1:1 sex ratio of hybrid adults.
Fewer crosses with other species and subspecies were made, and the
results were more variable. Inheritance of the dark female morph
appeared to be the same in P. a. garcia as in P. glaucus. Pure P. a. garcia
neonate larvae survived best on Prunus serotina, the natural host, and
on other Rosaceae and Oleaceae with intermediate survival on species
of Rutaceae, Magnoliaceae, Platanaceae, and Betulaceae. Salicaceae
and Rhamnaceae species were of little value as larval hosts. In general,
hybrid survival was similar but showed influences of the P. glaucus
subspecies that was the female parent.
Introduction
Two Mexican tiger swallowtail butterfly subspecies have been de-
scribed ( Papilio alexiares alexiares Hopffer and P. a. garcia Rothschild
and Jordan), but little is known about their biology (Brower, 1958;
Scriber, 1973; Frances & Elvira, 1978; Beutelspacher & Howe, 1984).
The subspecies P. a . alexiares ranges throughout the states of Hidalgo,
Puebla, and Veracruz, northeast of Mexico City at altitudes from 500 m
to 2600 m. Both sexes are the yellow tiger-striped morph (Beutelspacher
& Howe, 1984; Tyler, 1975; Jorge Llorente Bosquets, pers. comm.). P. a.
garcia is found further to the north in Tamaulipas, Nuevo Leon, and
San Luis Potosi (Fig. 1) and is reported to have only dark morph females
(Beutelspacher & Howe, 1984; Lee D. Miller, pers. comm.). On the basis
of male genitalia, Brower (1959) suggested thatP. alexiares was more
1Current address and reprint requests to senior author: Department of Entomology,
Michigan State University, East Lansing, MI 48824
27(3-4):222-232, 1988(89)
223
closely allied to the western species (P. rutulus Lucas, P. eurymedon
Lucas, and P. multicaudatus Kirby) than to P. glaucus. L. Genetic
distance data derived from allozyme electrophoresis in our laboratory
support this contention (Hagen and Scriber, in prep.)
Recently, black cherry {Prunus serotina Ehrh.) has been observed to
be one of the natural hosts of P. a. garcia (Evans et al., 1988; Fig 2). In
this paper, we report larval acceptance and survival on various potential
foodplant species used elsewhere in North America by the Papilio
glaucus and/or troilus species groups. We also present data detailing
various interspecific hand-pairings of P. a. garcia with other Papilio
glaucus species group members. These data provide additional insights
into the genetics of the dark morph female color polymorphism in the
Papilio glaucus species group (see Clarke & Sheppard, 1959; 1962;
Scriber, 1985; Scriber et al., 1986; Scriber & Evans, 1987 for discussion).
Methods
Both male and female P. a. garcia were collected in Nuevo Leon and
Tamaulipas, Mexico in March and April, June, and August and September
1984, 1986 and 1987. Enveloped specimens were either mailed or carried on ice
to our laboratory.
Male P. a. garcia were hand-paired to virgin P. a. garcia females or virgin
females of other Papilio glaucus group species. Field-collected and laboratory-
mated females were set up in plastic boxes (10 cm x 20 cm x 27 cm) with a sprig
of black cherry, Prunus serotina , under saturated humidity. The boxes were
placed 0. 7-1.0 m from continuously lighted 100 watt incandescent bulbs.
Females were fed a mixture of 1 part honey to 4 parts water at least once daily.
Most females were allowed to oviposit until death. After they died, hand-paired
females were dissected, and the presence of spermatophores was determined.
Any female not containing a spermatophore was eliminated from analysis.
Field-collected females were not routinely dissected for this study because
virgin Papilio females are rarely collected (Burns, 1968; Makielski, 1972;
Pliske, 1972; Platt et al., 1984; Lederhouse & Scriber 1987a).
Eggs were collected and counted at 2-day intervals except on weekends.
Larvae were removed as they hatched, and the remaining eggs were monitored
for 10 days after the last larva hatched. Egg viability was the proportion of the
total eggs laid that hatched as larvae. Using fine camel-hair brushes, first instar
larvae (neonate) were gently placed on fresh leaves of various potential
hostplants for bioassays of consumption and survival. Leaf moisture was
maintained using aquapics, and fresh leaves were provided 3 times per week
throughout larval development. Larval survival equaled the percent of first
instars set up on a host that successfully molted to the second instar. Means
were calculated with each mother considered a replicate. Some progeny of field-
collected P. a. garcia females were used in subsequent matings.
Results and Discussion
The pattern of oviposition of 26 field-collected and 36 hand-paired P.
a. garcia was similar to that of the 3 P. glaucus subspecies (Table 1). In
224
J. Res. Lepid.
general somewhat more than half of the females that were set up laid
some eggs. Of those females than laid eggs, field-collected females were
more likely that hand-paired females to produce larvae from their
clutches (X2, p < 0.01 in each case). The mean viability of P. a. garcia
clutches laid by field-collected females was similar to those of compar-
able females of each P. glaucus subspecies (Lederhouse & Scriber,
1987a). There was considerable clutch to clutch variability in larval
hatching.
Spermatophores were passed during hand-pairings between P. a.
garcia males and females and other P. glaucus group species (Table 2).
Table 1. Oviposition characteristics of field-collected and laboratory reared
and hand-mated females of Papiiio alexiares garcia and P. glaucus
subspecies. A subsample of females that had laid more than lOeggs
was used to calculate mean egg viability.
Phenotype
No.
females
% laying
eggs
% layers
with larvae
n
Egg Viability (%)
mean range
P. alexiares
field
26
65.4
76.5
11
51.3
10.6-80.9
hand-paired
36
58.3
28.6
5
34.6
4.5-73.3
P. g. glaucus
field
959
54.6
70.4
63
59.3
1.6-100.0
hand-paired
191
87.4
26.3
35
52.7
8.3-100.0
P. g. canadensis
field
730
48.4
65.2
50
55.9
2.4-95.0
hand-paired
69
82.6
19.3
9
29.7
2.4-95.5
P. g. australis
field
85
70.6
73.3
31
58.7
0.7-97.1
hand-paired
0
—
—
--
—
—
Fig. 1. Typical habitat of Papiiio alexiares garcia west of Cola de Caballo,
Nuevo Leon, Mexico at an elevation of about 1000 m.
Fig. 2. Black cherry tree (Prunus se rot in a) where P. a. garcia larvae were
collected. The tree was at about 1 1 00 m elevation on Chipinque Mesa,
Nuevo Leon, Mexico.
Fig. 3. Adult P. a. garcia collected in Nuevo Leon, Mexico. A. Male dorsal and
ventral, 15 April 1984. B. Female dorsal and ventral, 23 March 1985.
Fig. 4. Representative hybrid adults from a yellow Ohio P. g . glaucus female
and a P. a. garcia male (pairing 1071). A. Male dorsal and ventral. B.
Female dorsal and ventral.
Fig. 5. Representative hybrid adults from a dark Ohio P. g. glaucus female
and the same P. a. garcia male (pairing 1100). A. Male dorsal and
ventral. B. Female dorsal and ventral.
Fig. 6. Larvae of P. a. garcia reared on black cherry. A. Neonate. B. Larva
molting into the final instar found on black cherry in the field. C. Final
(fifth) instar.
27(3-4):222-232, 1988(89)
225
|
226
J. Res. Lepid.
Table 2. Oviposition characteristics of P. giaucus species-group females
hand-paired with Papi/io alexiares garcia males and P. a. garcia
females hand-paired with P. giaucus species-group males. The
female parent is listed first. Mean and range of viabilities of hybrid
eggs are presented. All females were dissected, and only those
containing a spermatophore are considered. Mean number of eggs
and percent egg viability is presented only for females with at least
one larvae.
Phenotype
Mated
females
% laying
eggs
% layers
with larvae
Egg
Mean
Egg Viability {%)
Mean Range
P. g. giaucus
x P. alexiares
24
87.5
76.2
173.9
66.1
28.4-96.9
P. g. canadensis
x P. alexiares
7
100.0
71.4
71.0
35.7
2.1-67.9
P. g. australis
x P. alexiares
6
83.3
100.0
105.8
34.8
11.5-59.1
P. rutu/us
x P. alexiares
1
100.0
0.0
_
_
P. alexiares
x P. g. giaucus
3
100.0
33.3
15.0
60.0
—
P. alexiares
x P. g. canadensis
1
100.0
100.0
1.0
100.0
—
P. alexiares
x P. g. australis
1
100.0
100.0
21.0
9.5
—
P. alexiares
x P. rutu/us
1
100.0
100.0
113.0
33.6
_
P. alexiares
x P. eurymedon
1
100.0
0.0
-
-
-
Nearly all females laid eggs. The mean viability of eggs from female P.
g. giaucus and P. a. garcia males was equivalent to that of field-collected
pure subspecies (Table 1). Egg viability of other hybrid crosses was
lower, but not lower than that of hand-paired pure subspecies. Sex ratios
at adult emergence totaled 240 males to 211 females for P. g. giaucus
females x P. a. garcia males, 37:32 for P. g. australis females x P. a.
garcia males, and 8:10 for P. g. canadensis females x P. a. garcia males
(Table 3). None of these ratios differs significantly from an expected of
1:1 (X2, p > 0.20 in each case). These results are further evidence of high
genetic compatibility between P. giaucus and P. a. garcia.
The crosses between male P. a. garcia and female P. g. giaucus or P. g.
australis were particularly interesting since these are the only
members of the entire North American tiger swallowtail group that
have dark female polymorphism (Fig. 3; Clarke & Sheppard, 1962;
Scriber et al., 1987; Lederhouse & Scriber, 1987b). Our data (Table 3)
27(3-4):222-232, 1988(89)
227
Table 3. Hybrid pairings of the two color morphs of P. g/aucus subspecies
females with P. alexiares garcia males and the resulting offspring.
Papi/io g/aucus females were reared from mothers collected in
Ohio, Illinois, Wisconsin and Florida.
Mating
number Phenotypes
Dead
pupae
Males
Yellow
females
Dark
females
P. g. g/aucus
1071 Yellow OH x male 1
7
58
42
1
1072
Yellow OH x male 2
3
8
8
0
1100
Dark OH x male 1
14
61
1
49
4210
Dark OH x male 3
3
32
2
27
4211
Dark IL x male 4
7
47
2
52
4227
Dark OH x male 5
0
5
0
2
4228
Dark IL x male 6
0
5
0
5
4230
Dark IL x male 3
0
8
2
9
4234
Dark OH x male 7
0
3
0
7
4458
Dark OH x male 8
0
5
0
1
4471
Yellow OH x male 9
0
8
1
0
P. g.
3547
austra/is
Dark FL x male 10
0
27
0
27
4581
Yellow FL x male 1 1
1
8
3
0
4587
Yellow FL x male 12
0
1
1
0
4598
Yellow FL x male 13
0
1
1
0
p. g-
1625
canadensis
Yellow Wl x male 14
0
0
2
0
3368
Yellow Wl x male 15
0
0
1
0
4457
Yellow Wl x male 16
0
2
5
0
4470
Yellow Wl x male 17
1
6
2
0
suggest that the same genetic basis is likely to be involved for all 3 taxa:
a Y-linked gene for melanism without color suppressors in males
(Scriber, 1985; Scriber et al., 1987). Dark females (XY°) generally
produce dark daughters regardless of the color of the mother of their
mate, and yellow females (XY) generally only produce yellow daughters
(Table 3). Occasionally, a female offspring of the opposite color from her
mother will be produced (Hagen & Scriber, 1989). Possible explanations
of such results are discussed elsewhere (Clarke at al. 1976, Scriber et al.
1987). Hybrid pairings of the same P. a. garcia male with a yellow
female and a dark female produced daughters of the expected phenotype
(pairing 1071, Fig. 4, pairing 1100, Fig. 5). The same pattern held for P.
g. australis females crossed with P. a. garcia males (Table 3). All hybrid
females from P. g. canadensis females crossed with P. a. garcia males
were yellow (Table 3); P. g. canadensis females lack the dark gene
(Scriber et al. 1987).
The newly eclosed first instar larvae of P. a. garcia and P. glaucus
subspecies hybrids with P. a garcia males exhibited differential sur-
vival in no-choice tests on leaves of 27 plant species from 10 plant
families (Table 4). For pure P. a garcia, neonates survived best on black
cherry ( Prunus serotina Ehrh.), its natural host, other Prunus species,
Table 4. No-choice feeding bioassays of Mexican Papilio aiexiares garcia and its hybrids. The female parent of hybrid larvae is
listed first. The top value for survival is the mean percent; the lower value is the standard deviation.
228
J. Res. Lepid.
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J. Res. Lepid.
and white ash ( Fraxinus americana L.). Intermediate levels of survival
were shown by P. a. garcia neonates on hoptree ( Ptelea trifoliata L.,
Rutaceae), tuliptree (. Liriodendron tulipifera L., Magnoliaceae), syca-
more ( Platanus occidentalis L., Platanaceae), and paper birch ( Betula
papyrifera Marsh., Retulaceae). Although sample sizes are small in
some cases, plant species in the Salicaceae and Rhamnaceae are of
minimal usefulness as food plants for the Mexican tiger swallowtail.
First and final instar P. a. garcia larvae are shown in Figure 6.
In general, hybrid survival was similar to that of pure P. a. garcia , but
showed the influence of the particular subspecies of P . glaucus that was
the female parent (Table 4). Hybrid survival was uniformly high on
black cherry, choke cherry ( Prunus virginiana L.), and white ash.
Hybrids from P. g. glaucus and P. g. australis mothers exhibited higher
survival on hosts in the families Rutaceae, Lauraceae, and Magno-
liaceae. Hybrids from P. g. canadensis mothers had enhanced survivor-
ship on Salicaceae hosts (Table 4).
The Magnoliaceae and Salicaceae are believed to represent major
adaptive radiations in host use for North American Papilio from a
possible Lauraceae or Rutaceae root (Scriber, 1983; 1986). Larvae ofP.
g. glaucus and P. g. australis readily grow on Magnoliaceae but mostly
die on Salicaceae; P. g. canadensis , P. rutulus, and P. eurymedon larvae
exhibit the opposite abilities (Lindroth et ah, 1986, 1988; Scriber et ah,
1986). Should P. alexiares represent the ancestral stock (from south-
western Pleistocene refugia) for a P. glaucus species group radiation, we
are not surprised that P. alexiares larvae possess some capabilities to
detoxify and process foodplants from all 4 plant families. We continue to
expand our studies to assess the degree of phylogenetic affiliation of P.
alexiares garcia with other P. glaucus group taxa.
Acknowledgements. This research was supported in part by the National
Science Foundation (BSR 8306060, BSR 8718448), USDA grants #85-CRCR-l-
1598 and #87-CRCR-l-2851, and the Graduate School and College of Agricul-
tural and Life Sciences (Hatch 5134) of the University of Wisconsin. We would
particularly like to thank Dr. Robert Dowell, Dr. John Thompson, and Wayne
Wehling for their help in collecting P. eurymedon , P. multicaudatus, and P.
rutulus , and Dr. James Nitao for photographing larvae. We are also grateful for
the continuing assistance of Dr. David Robacker and William Warfield in
Mexico and to the University of Wisconsin Ibero-American Studies Program (to
MHE) and the Romnes Faculty Fellowship (to JMS) for travel support to Mexico.
We are thankful for the assistance of Dr. Lowell Nault, Dr. Raphael Rodriquez,
and the Center for International Maize and Wheat Improvement in Mexico, and
to the Colleges of Natural Science and Agriculture of Michigan State University,
MAES Project 8051.
Literature Cited
BROWER, L. P., 1985. Larval foodplant specificity in butterflies of the Papilio
glaucus group. Lep. News 12:103-114.
27(3-4):222-232, 1988(89)
231
BROWER, L. P., 1959. Speciation in butterflies of the P. glaucus group. I. Morpho-
logical relationships and hybridization. Evolution 13:40-63.
BEUTELSPACHER, C. R. & W. H. HOWE., 1984. Mariposas de Mexico. La Prensa
Medica Mexicana, S. A., Mexico 127 pp.
BURNS, J. M., 1968. Mating frequency in natural populations of skippers and
butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci.
61:852-859.
CLARKE, C. A. & P. M. SHEPPARD, 1959. The genetics of some mimetic forms of
Papilio dardanus Brown and Papilio glaucus L. J. Genetics 56:236-260.
CLARKE, C. A. & P. M. SHEPPARD, 1962. The genetics of the mimetic butterfly,
Papilio glaucus. Ecology 43:159-161
CLARKE, C. A., P. M. SHEPPARD, & U. MITTWOCH, 1976. Heterochromatin poly-
morphism and colour pattern in the tiger swallowtail butterfly Papilio
glaucus L. Nature 263:585-587.
EVANS, M. H., D. ROBACKER, W. WARFIELD, & J. M. SCRIBER, 1988. Larval foodplants of
the Mexican swallowtail butterfly, Papilio alexiares garcia. J. Lepid. Soc.
(submitted).
FRANCES, A. D. & J. M. ELVIRA, 1978. Guia ilustrada de las mariposaa Mexicanas.
Parte I. Familia Papilionidae. Special Pub. Sociedad Mexicana de Lepi-
dopterologia, A-C. 15pp.
HAGEN, R. H. & J. M. SCRIBER, 1989. Genetic analysis of sex-linked loci in the tiger
swallowtail butterfly. J. Heredity (in press).
LEDERHOUSE, R. C. & J. M. SCRIBER, 1987a. Ecological significance of a postmating
decline in egg viability in the tiger swallowtail. J. Lepid. Soc. 41:83-93.
LEDERHOUSE, R. C. & J. M. SCRIBER, 1987b. Increased relative frequency of dark
morph females in the tiger swallowtail Papilio glaucus (Lepidoptera:
Papilionidae) in S-central Florida. Amer. Midi. Nat. 118:211-213.
LINDROTH, R. L„ J. M. SCRIBER, & M. T. S. HSIA, 1986. Differential responses of tiger
swallowtail subspecies to secondary metabolites from tulip tree and quaking
aspen leaves. Oecologia 70:13-19.
LINDROTH, R. L., J. M. SCRIBER, & M. T. S. HSIA, 1988. Chemical ecology of the tiger
swallowtail: mediation of host use by phenolic glycosides. Ecology 69:814-
822.
MAKIELSKI, S. K., 1972. Polymorphism in Papilio glaucus (Papilionidae): Main-
tenance of the female ancestral form. J. Lepid. Soc. 26:109-111.
PLATT, A. P., S. 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.
Academic Press, London.
PLISKE, T. E., 1972. Sexual selection and dimorphism in female tiger swallow-
tails, Papilio glaucus L. (Lepidoptera: Papilionidae): A reappraisal. Ann.
Entomol. Soc. Am. 65:1267-1270.
SCRIBER, J. M., 1973. Latitudinal gradients in larval feeding specialization of the
world Papilionidae (Lepidoptera). Psyche 80:355-373.
SCRIBER, J. M., 1983. The evolution of feeding specialization, physiological
efficiency, and host races in selected Papilionidae and Saturniidae. In R. F.
Denno & M. S. McClure (eds.). Variable plants and herbivores in natural and
managed systems. Academic Press, New York, pp. 373-412.
SCRIBER, J. M., 1985. The ecological and genetic factors determining geographic
limits to the dark morph polymorphism in Papilio glaucus. Bull. Ecol. Soc.
Amer. 66:267.
232
J. Res. Lepid.
SCRIBER, J. M., 1986. Origins of the regional feeding abilities in the tiger
swallowtail butterfly: Ecological monophagy and the Papilio glaucus
australis subspecies in Florida. Oecologia 71:94-103.
SCRIBER, J. M. & M. H. EVANS, 1986. An exceptional case of paternal transmission of
the dark form female trait in the tiger swallowtail butterfly, Papilio glaucus
(Lepidoptera: Papilionidae). J. Res. Lepid. 25:110—120.
SCRIBER, J. M., M. H. EVANS & D. B. RITLAND, 1987. Hybridization as a causal
mechanism of mixed color broods and unusual color morphs of female
offspring in the eastern tiger swallowtail butterfly, Papilio glaucus. In M. D.
Huettel (ed.), Evolutionary genetics of invertebrate behavior. Plenum
Publishing Corp. pp. 119-134.
SCRIBER, J. M., M. T. S. HSIA, P. SUNARJO & R. LINDROTH, 1986. Allelochemicals as
determinants of insect damage across the North American continent:
biotypes and biogeography. In. G. R. Waller (ed.), Allelochemicals: role in
agriculture, forestry and ecology. Amer. Chem. Soc. Symp. Series 330, pp
439-448.
TYLER, H., 1975. The swallowtail butterflies of North America. Naturegraph,
Healdsburg, CA. 192 pp.
''N
Journal of Research on the Lepidoptera
27(3-4):233-256, 1988(89)
The Butterflies oflsla de Cedros, Baja California Norte,
Mexico
John W. Brown
and
David K. Faulkner
Entomology Department, San Diego Natural History Museum, San Diego, California
92112, U.S.A.
Abstract. Isla de Cedros is an arid Pacific island off the western coast of
Baja California Norte, Mexico. The island supports a depauperate
butterfly fauna consistent with other offshore islands which exhibit
varying degrees of faunal reduction when compared to their mainland
counterparts. The 23 butterfly species recorded from Isla de Cedros
reflect 2 broad categories of presumptive biogeographic origin:
1) species of Neotropical origin, which are distributed throughout the
peninsula; and 2) species of Nearctic origin, some of which occur
throughout the peninsula, and others confined to the Californian
province of the adjacent peninsula. The 80 year history of entomo-
logical activity on the island is outlined; the physiography of the area is
briefly discussed; and the 23 butterfly species are listed with capture
records and taxonomic comments. Additionally, an endemic species,
Mitoura cedrosensis is described and illustrated.
Introduction
The butterfly fauna oflsla de Cedros, Baja California Norte, Mexico,
has been sampled on numerous occasions over the past 80 years, most
recently by the authors in 1981 and 1983. A total of 23 butterfly species
has been recorded from the island, including 1 endemic species and 1
endemic subspecies. This number is considerably less than the number
of species that would be found in comparable habitats on the adjacent
mainland. This fact is consistent with other offshore islands which
exhibit varying degrees of faunal reduction, generally dependent upon
their size and distance from continental masses (MacArthur and Wilson,
1967; Pielou, 1979; Langston, 1980). Geologic evidence of a previous
landbridge to the peninsula of Baja California suggests the past
opportunity for the development of a more diverse fauna than is
currently evident. Pielou (1979) suggests that upon separation from the
mainland, continental islands have an over-saturated biota, and that a
period of floral and faunal reduction ensues until the number of species
on the island falls to an appropriate equilibrium level. Clear evidence of
faunal reduction has been given by Wilcox (1978) for the lizard faunas of
several Baja California islands. In the butterfly fauna of Cedros,
234 J. Res. Lepid.
however, the island appears to be in an under-saturated (non-equilib-
rium) condition.
The peninsula of Baja California can be divided into 3 major biotic
provinces: a northwestern Californian region, a central desert region,
and a southern subtropical thorn scrub region which includes the cape.
Floral characteristics of the northwestern province occur as disjuncts
southward on scattered higher peaks forming outposts of this region as
far south as the mountains of the cape. Such an outpost occurs in the
higher elevations of Isla de Cedros. Before the origin of the deserts in the
late Quaternary, these southern relicts were presumably more nearly
continuous with the northwestern region (Gould and Moran, 1981). As a
consequence of this outpost effect, serveral Californian elements reach
their southern limit on Isla de Cedros, considerably disjunct and
isolated from the southern end of their contiguous peninsular popu-
lations to the north.
Collecting History
Although seldom a primary destination, Isla de Cedros has histori-
cally provided a stop-over for boat expeditions traveling along the
Pacific coast of Baja California. The following outline briefly sum-
marizes the historical accounts of entomological activity on the island.
1905. California Academy of Sciences Expedition to the Galapagos
Islands. On Cedros 18 July 1905. F.X. Williams, entomologist.
1922. California Academy of Sciences Expedition to the Eastern
Pacific Islands. On Cedros 22 July 1922. G. Hanna and J. Slevin,
collectors.
1925. California Academy of Sciences Expediton to Revillagigedo
Islands. On Cedros 2-6 June 1925. H. H. Keifer, entomologist.
1932. Allan Hancock Pacific Expedition. On Cedros 25 February
1932. J. S. Garth, entomologist.
1934. Allan Hancock Pacific Expedition. On Cedros 10 March 1934. J.
S. Garth, entomologist.
1937. Allan Hancock Pacific Expedition. On Cedros 10 and 12 July
1937. J. S. Garth, entomologist.
1937-1939. Several boat trips to Baja California by F. Rindge family.
F. H. Rindge, entomologist.
1941. Allan Hancock Pacific Expedition. On Cedros 28 February 1941.
J. S. Garth, entomologist.
1949. Velero IV Gulf of California Cruise. On Cedros 4-5 March 1949.
J. S. Garth, entomologist.
1981. San Diego Natural History Museum Expedition to Northern
Baja California. On Cedros 20-23 March 1981. D. Faulkner and F.
Andrews, entomologists.
1983. San Diego Natural History Museum Expedition to Isla de
Cedros. On Cedros 28 March-5 April 1983. J. Brown and D. Faulkner,
entomologists.
27(3-4):233-256, 1988(89)
235
1983. Diamaresa Expedition to Pacific Islands Adjacent to Baja Cali-
fornia. On Cedros 30 June-2 July 1983, 13 July 1983. D. Faulkner, D.
Weissman, D. Lightfoot, and V. Lee, entomologists.
Although there have been a number of visits to the island in the past
80 years, few of the expeditions spent more than a brief time on the
island, making only short trips into the more accessible localities, such
as Canon de la Mina in the north. This is reflected in the few Lepidoptera
specimens available for examination as well as the low number of
species recorded until recently.
Physiography
Geology. Isla de Cedros is a rather large (348 km2), rugged, moun-
tainous island (Fig. 1) situated about midway down the western side of
the peninsula of Baja California, Mexico (Fig. 2). Oriented north to
south, the island is about 34 km in length and varies from about 4 to 15
km in width. The southeastern extremity, Punta Morro Redondo, is
separated from the mainland by a narrow and shallow strait 22 km
wide. Projecting northwest from the mainland, Punta San Eugenio
represents the southern connection of a presumed landbridge that once
united Cedros with the peninsula (Gentry, 1950). It is likely that
migrant species regularly reach Cedros by “island hopping” from Punta
San Eugenio to Isla Natividad, and from there to Cedros.
The island’s montane spine is bisected into a northern and a southern
range by a deep gorge called El Gran Canon or El Arroyo Grande. The
highest point, Cerro de Cedros, in the southern half of the island,
Fig. 1. Eastern coast of Isla de Cedros, looking south from Punta Norte.
236
J. Res. Lepid.
Fig. 2. Map of Isla de Cedros; all localities mentioned in the text are figured.
27(3-4):233~256, 1988(89)
237
reaches an elevation of 1200 m (3950’) - The uplifted sedimentary strata
reflect a history of tremendous geologic disturbances. The granodioritic
rocks present are of pre-Cretaceous and Pliocene origin (Wiggins, 1980).
A reconnaissance of the geology of Isla de Cedros (Kilmer, 1977)
indicates that the island was formed by an uplift of late Jurassic
metamorphic and igneous rock at a point where the Pacific plate was
subducted beneath the western margin of the North American plate.
The possible geological relationship of Isla de Cedros with the Cali-
fornia Channel Islands emphasizes the relationship between the small
but striking relictual floral elements common to the two areas (Moran
and Benedict, 1981).
Climate. The climate of Isla de Cedros is generally temperate owing
to its proximity to Mediterranean climatic regimes; however, long, hot,
dry spells are common. Cedros is near the southern edge of California’s
winter Pacific storm tract, and at the northern extreme of southern Baja
California’s tropical summer storm pattern. Precipition records, as a
result, indicate extreme inconsistency in both seasons; in some years
little or no rain reaches the island. Figure 3 provides climatological data
adapted from Hastings and Humphrey (1969).
In the vicinity of Isla de Cedros, generally to the north and west, there
are often low, dense mists or fog banks which are common in all seasons
but particularly in the summer months (Libby, Bannister, and Linhart,
1968; Lewis and Ebeling, 1971). The abundant moisture provided by
this condition has great influence in producing the luxuriant desert
vegetation which occurs during certain seasons on parts of the western
slopes (Nelson, 1921), and sustains the stands of Monterey pine that
occur on the west and northwest escarpments of the island’s northern
range.
Flora
Because of its accessibility by ship, and more recently be cargo plane,
the flora of Isla de Cedros has been rather extensively studied (Moran,
1972). Hale (1941) estimates that 97% of the island is covered by desert
scrub vegetation similar to that occurring throughout the Vizcaino-
Magdalena region of the adjacent peninsula. The most striking and
conspicuous plants occurring over most of the island are the elephant
tree (. Pachycormus discolor (Benth.) Cov.) and the mescal {Agave
sehastiana (Greene) Gentry) (Fig. 4). In small isolated areas the desert
scrub gives way to other types of vegetation, most notably coastal sage
scrub, chaparral, and even coniferous forest. The most remarkable
departure from the desert vegetation is the closed-cone pine forests
dominated by Pinus radiata var. cedrosensis J. T. Howell which occur in
2 major populations in the mountains (Libby, Bannister, and Linhart,
1968). Several Californian floral elements reach their southern limit on
238
J. Res. Lepid.
co
3
1§ 24-1
0
o
co
0
0
u.
O)
0
•D
C
0
w.
3
22-
20“
18-
16-
Mean Monthly Temperature
i 1 1 1 — ™r 1 r 1 1 1 1 1
J FMAMJ jasond
Mean Monthly Rainfall
Fig. 3. Climatological data adapted from Hastings and Humphrey (1969).
Above: Annual precipitation (in mm). Below: Average annual
temperature (in °C).
Cedros including California juniper ( Juniperus californica Carr.),
lemonade berry ( Rhus integrifolia (Nutt.) Rothr.), chamise ( Adenostoma
fasciculatum var. obtusidolium S. Wats.), and California sage brush
( Artemisia californica Less.). The flora of the island includes 245
vascular plants, of which 216 species are native and 29 species intro-
duced (Moran and Benedict, 1981). Of the native flora, 16 species are
endemic to Isla de Cedros and are discussed by Moran (1972).
Butterfly Fauna
The 23 butterfly species recorded from Isla de Cedros represent 6
families: Hesperiidae (2 species), Pieridae (7 species), Lycaenidae (9
27(3-4):233-256, 1988(89)
239
Fig. 4. Above: Fog-enshrouded Canon de la Mina at the north end of the
island. The tall, white-flowered, endemic Eriogonurn mo/ie is con-
spicuously abundant. Below: Characteristic vegetation near the
light-house at Punta Norte, dominated by Agave sebastiana and
Opuntia species.
240
J. Res . Lepid.
species), Riodinidae (2 species), Nymphalidae (2 species), and Danaidae
(1 species). The 23 species reflect 2 extremely broad categories of
biogeographic origin: the Neotropical and the Nearctic.
Species of Neotropical origin represented in the island’s fauna are
widespread forms that occur the entire length of the peninsula, ex-
tending more or less from South or Central America northward into
southern California. Species in this category include Erynnis funeralis,
Phoebis sennae, E urema nicippe, Strymon columella , Brephidium exilis ,
Leptotes marina , Hemiargus ceraunus, and Danaus gilippus. These
species represent approximately 35% of the total butterfly fauna.
Elements of Nearctic origin illustrate 2 patterns of mainland distri-
bution: a) species distributed throughout the peninsula, including
Pyrgus albescens, Pieris protodice, Colias eurytheme, Strymon melinus,
Celastrina ladon, Apodemia mormo, Calephelis wrighti, Vanessa
cardui , and Vanessa annabella, comprising approximately 39% of the
butterfly fauna; and b) species typically confined to the Californian
province of the adjacent peninsula, represented on Cedros by disjuncts
or isolated relict populations, including Pieris beckerii, Anthocharis
sara, Anthocharis cethura, Mitoura cedrosensis new species, Philotes
sonorensis, and Euphilotes battoides. All of these Californian elements
reach their southernmost distributional limits on Cedros. Included in
this group are the 2 endemic taxa. The Californian province elements
account for approximately 26% of the species recorded from the island.
Thus were it not for a broad zone of distributional overlap between the
widespread Neotropical and widespread Nearctic species, the island’s
fauna would most likely reflect an even more depauperate conditon than
is currently illustrated.
Approximately 50% of the species known from the adjacent mainland
(species pool) occur on Isla de Cedros. This is consistent with the finding
of Langston (1980) regarding the faunal composition of Santa Cruz
Island which is located off the western coast of California. The two
islands share 9 species of butterflies representing widespread Neo-
tropical, widespread Nearctic, and Californian province elements. The
species in common all exhibit a high degree of vagility.
Latitude seems to have little effect on the phenology of the Cali-
fornian elements. Although Isla de Cedros is 500 km (310 mi) south of
the California-Baja California border, species’ flight periods closely
resemble those of their southern California counterparts. Several of the
univoltine species do, however, exhibit extended flight periods giving
the appearance of more than a single brood, i.e., Philotes sonorensis and
Euphilotes battoides . Langston (1975) has shown that species occurring
near the Pacific coast (of California) often display this tendency,
probably in response to mild winters, periods of inclement spring
weather, and moderate summer temperatures, which in turn contribute
to the staggered development of the various larval hostplants.
27(3-4):233-256, 1988(89)
241
We examined 457 specimens representing 23 species. An additional 2
species, Danaus plexippus (L.) and a large dark papilionid, both reported
as sight records by David Weissman, are mentioned here but are not
included in the species accounts. All observations were made by the
authors during 1981 and 1983.
Unless otherwise indicated, all specimens listed in the species
accounts were collected by Faulkner and Brown, and are deposited in
the San Diego Natural History Museum. Specimens collected by J.
Garth are in the collection of the Allan Hancock Foundation at the
University of Southern California, Los Angeles. Additional depositories
are abbreviated as follows: CAS, California Academy of Sciences, San
Francisco; and LACM, Los Angeles County Museum of Natural History.
Species Accounts
HESPERIIDAE
1. Erynnis funeralis (Scudder and Burgess).
First reported from Cedros by Rindge (1948), we collected £. funeralis
on both the north and south ends of the island. It was encountered more
often at mid-to-low elevations, frequently “patrolling” canyons. Several
species of Lotus (Fabaceae) occur on the island, and one or more of these
probably serve as larval hosts. E. funeralis occurs the entire length of
the peninsula of Baja California, and there appear to be no phenotypic
differences between mainland and insular populations.
MacNeill (1975) indicates that funeralis has considerable dispersal
ability and has been shown to be a pioneer species in several insular
situations.
Specimens examined: Punta Norte, 30 March 1983 (2 males), 1 April
1983 (2 males); vicinity El Pueblo, 4 April 1983 (1 female).
2. Pyrgus albescens Plotz
MacNeill (1975) states thatP. albescens andP. communis (Grote) are
ecologically isolated as well as (genitalicly) distinct. On this basis, they
appear to represent separate species and were treated as such by Miller
and Brown (1981). That treatment is followed here.
P. albescens is a widespread inhabitant of the hot, arid lowlands of the
southwestern United States and adjacent Mexico. It was one of the more
common butterflies encountered on Cedros in both spring and summer
of 1983. It was particularly abundant in disturbed areas in the vicinity
of El Pueblo, especially in association with the weedy, introduced Malva
parviflora L. (Malvaceae). Several specimens were also collected on the
south slope of Cerro de Cedros, near the summit, in association with
Sphaeralcea fulva Greene (Malvaceae). No phenotypic differences are
apparent between peninsular and insular populations.
Specimens examined: El Pueblo, 29 March 1983 (1 male), 4 April 1983
242
J. Res. Lepid.
(10 males), 13 July 1983 (2 males); Punta Norte, 31 March 1983 (1
female), 1 April 1983 (1 male, 1 female), 3 July 1983 (2 males); vicinity
Cerro de Cedros, 3 April 1983 (4 males, 1 female), 1 July 1983 (2 males);
Gran Canon, 2 July 1983 (1 male).
PIERIDAE
3. Pontia protodice Boisduval and LeConte
As P. protodice occurs commonly throughout much of the United
States and northern Mexico, and in a variety of habitats, it was not
surprising to find this species on Isla de Cedros. Specimens were
collected on both the north and south ends of the island. Some of the
possible cruciferous hosts available include Descurainia, Sisymbrium ,
and Thelypodium. Although seasonally polyphenic, P. protodice is quite
homogeneous in phenotype throughout its range (no subspecies), in-
cluding Isla de Cedros.
Specimens examined: vicinity Punta Norte, 28 February 1941 (1
male), leg : J. Garth, 30 March 1983 (1 male), 1 April 1983 (1 male);
vicinity El Pueblo, 3 April 1983 (1 female), 4 April 1983 (3 males).
4. Pontia beckerii Edwards
A common pierid of the western United States, P. beckerii , generally
inhabits hot, shrubby, semi-arid habitats (Howe, 1975). Only in southern
California and northwestern Baja California does it occur on or near the
coast. The population on Cedros represents a slight southern disjunct
from northern Baja California. The larval host, Isomeris arborea Nutt.
(Capparidaceae), occurs commonly on the eastern side of the island
(Hale, 1941); a single larva was collected on I. arborea in a disturbed
area near El Pueblo. Specimens of P. beckerii from Cedros are indistin-
guishable from those of southern California.
Specimens examined: El Pueblo, 29 March 1983 (2 males), 3 April
1983 (1 male, 1 female), 4 April 1983 (1 female); Punta Norte, 1 April
1983 (1 female); vicinity Cerro de Cedros, 1 July 1983 (2 males).
5. Anthocharis sara Lucas
Widespread through the western United States, and extending south
into northern Baja California, A. sara reaches its southernmost distri-
bution on Isla de Cedros. Capture records from February through April
may indicate two broods, as is the case in coastal southern California.
Although some insular populations from California are subspecifi-
cally distinct (Emmel and Emmel, 1973), specimens from Cedros appear
to represent nominate A. sara . However, in about 10% of the male
specimens, the black scaling at the posterior end of the bar located near
the apical end of the DFW cell, extends basally forming a slight hook
(Fig. 5). Although these individuals have a distinct appearance, this
character is not consistent within the population sampled.
Specimens examined: vicinity Punta Norte, 25 February 1932 (2
27(3-4):233-256, 1988(89)
243
males, 1 female), 28 February 1941 (1 male, 1 female), all leg : J. Garth,
30 March 1983 (10 males, 1 female), 31 March 1983 (3 males, 2 females),
1 April 1983 (7 males, 1 female), 2 April 1983 (5 males, 1 female);
vicinity El Pueblo, 3 April 1983 (1 male).
6. Anthocharis cethura (Felder and Felder)
Restricted to the extreme southwestern United States and adjacent
northern Mexico, A. cethura reaches its southernmost limit on Isla de
Cedros. Although first collected on Cedros by John Garth in 1932, its
occurrence there was not noted until Rindge’s (1948) publication. In the
spring of 1983 A. cethura was collected on both the north and south ends
of the island. Although it was uncommon, generally observed singly in
canyons or on hilltops, previous collectors have found it to be much more
abundant. The authors collected a single larva on Sibara pectinata
(Greene) Greene (Brassicaceae) which is widely distributed on Cedros.
Other potential larval hosts available include Thely podium lasiophyl-
lum (Hook, and Arn.) and Descurainia pinnata (Walt.) (both Brassi-
caceae). Specimens from Cedros are probably best referred to nominate
A. cethura.
Specimens examined: vicinity Punta Norte, 25 February 1932 (1
male), 28 February 1941 (12 males, 3 females), all leg : J. Garth, 31
March 1983 (1 male), 1 April 1983 (1 male), 2 April 1983 (1 male);
vicinity El Pueblo, 29 March 1983 (1 male), 3 April 1983 (2 males).
7. Colias eury theme Boisduval
This widespread species was encountered only sparingly on Cedros.
Specimens were observed in spring and summer of 1983. The only
example collected, however, was a damaged adult retrieved from a
spider’s web. Several legumes on the island are available as potential
larval hostplants.
Specimen examined: vicinity Punta Norte, 3 July 1983 (1 female).
Fig. 5. Anthocharis sa-
ra, male, upper-
surface, Isla de
Cedros.
244
J. Res. Lepid.
8. Phoehis sennae rnarcellina (Cramer)
Although probably not a breeding resident, P. sennae was commonly
observed on both ends of the island in the summer of 1983. Captures
were made in the late afternoon as the adults were settling on Rhus.
Larvae ofP. sennae are known to feed on Cassia (Fabaceae), none of
which are available on Cedros. A well-known disperser-rnigrator, P.
sennae is frequently encountered far from its breeding areas, which
appears to be the case on Isla de Cedros.
Specimens examined: vicinity Funta Norte, Canon de la Mina, 3 July
1983 (2 males).
9. Eurema nicippe (Cramer)
E. nicippe is widespread throughout most of southern North America;
it occurs the length of Baja California. As with the preceding species,
nicippe does not appear to be a breeding resident on Cedros owing to the
absence of Cassia as a larval host. It is possible that other legumes are
utilized, but the flight-worn condition of specimens and their rapid
unidirectional flight together seem to indicate that specimens taken on
Cedros represent migrants from the adjacent mainland.
Specimens examined: vicinity Cerro de Cedros, 1 July 1983 (1 male);
vicinity Punta Norte, Canon de la Mina, 3 July 1983 (1 male).
LYCAENIDAE
10. Mitoura cedrosensis new species
Figures 6 and 7
Male: forewing length x = 11.4 mm (range 11.0-12.0 mm; n = 14).
Frons and vertex fuscous; eyes mesially edged with white; antennae
black, white annulate, the club black with a fulvous tip. Upperside:
both wings fuscous to mahogany brown with marginal, apical, and basal
darkening. A thin terminal white bar on hindwing between tornus and
Cu2. A short, thread-like tail at termination of Cu2 of hindwing, black
tipped with white. Only a small black tooth at Cu1? also tipped with
white. Forewing scent patch well developed although variable in color.
Underside: forewing rich mahogany brown with a fine postmedian line
composed of 5 white dashes. Faint traces of maroon purple over-scaling
apically, and faint basal darkening. Hind wing with a diffuse inconsi-
stent maroon postbasal band; occasionally bordered at outer margin by
a thin white line from M3 toward costal margin, absent to very faint in
some specimens. Terminal area aqua gray with a variable row of poorly
defined black dots. In Cux-Cu2 a Thecla spot composed of 2 longitudin-
ally arranged black dots divided by a poorly defined orange-brown
lunule. Entire hind wing surface rather melanistic in appearance, with
a faint iridescent luster.
Female: forewing length x = 11.4 mm (range 11,0 12.0 mm; n = 9).
Upperside: as in male but without scent patch, and color more consi-
stent rich reddish brown; darkening confined to marginal area. Under-
side: as in male with little or no consistent differences.
27(3-4):233-256, 1988(89)
245
Fig. 6.
Fig. 7.
Mitoura cedrosensis, female, uppersurface, Isla de Cedros.
Mitoura cedrosensis, female, undersurface, Isla de Cedros.
246
J. Res . Lepid .
Genitalia: Two specimens of each sex are illustrated in Figure 8. As
noted by Brown (1983) for related species, variation, as exemplified
between the two specimens of each sex examined, is substantial.
Comparison with illustrations in Brown (1983) gives a brief account of
the related southern California taxa. The only character which may be
of diagnostic value is the dor so- ventral shape of the male saccus. The
female genitalia and male valvae appear to be of less taxonomic value,
although quantitative differences may be evident in larger samples
which could be statistically validated. Although the saccus shape may
be of diagnostic value in differentiating the loki, thornei , and nelsoni
groups, this character does not lead to any conclusions regarding
reproductive isolation (Shapiro, 1978).
Type material: All Isla de Cedros, Baja California Norte, Mexico;
holotype, male, Punta Norte, 28°22'N, 115°12'W, 20-22 March 1981;
allotype, Punta Norte, 28°22'N, 115°12'W, 20-22 March 1982. Thirteen
male and 8 female paratypes as follows: Punta Norte, 20-22 March 1981
(6 males, 5 females), 1 April 1983 (2 males, 1 female), 31 March 1983 (3
males, 1 female), 30 March 1983 (1 male), 3 July 1983 (1 male); vicinity
Cerro de Cedros, 1 July 1983 (1 female).
Disposition of types. Holotype and allotype are deposited in the
SDNHM. Paratypes deposited in the following institutions: Los Angeles
County Museum of Natural History, Los Angeles, California; California
Academy of Sciences, San Francisco, California; and Universidad
Biologla de Mexico, Mexico City, Mexico.
Remarks. Mitoura cedrosensis is closely related to M. loki (Skinner).
It represents an insular, southernmost outpost of the California juniper-
feeding Mitoura complex and is endemic to Isla de Cedros. The nearest
known population of M. loki occurs approximately 300 km to the north
in the vicinity of Mike’s Sky Ranch in the Sierra San Pedro Martir, Baja
California Norte.
M. cedrosensis is easily distinguished from M. loki by its smaller size1
and by the fuscous purplish brown of the ventral hindwing surface
replacing the hindwing green overscaling of loki. When compared to the
newly described M. thornei Brown (1983) from southern California, M.
cedrosensis is smaller and the markings on the hindwing are slightly
darker, more fuscous, and less well-defined. The thin, white border at
the outer edge of the postbasal band present in both thornei and loki is
reduced or absent in cedrosensis. There is some question regarding the
specific status of thornei and cedrosensis , both of which might be
considered as subspecies of M. loki by some authors (Shields, 1984). M.
cedrosensis is not similar to the unusual M. nelsoni (Boisduval) known
from Isla Guadalupe, Baja California Norte, Mexico (Powell, 1958;
Brown, 1983). The presence of basal markings representing the inner
1Student’s t-test comparing 2 sample means indicates statistically significant difference
in forewing length between samples of M. loki and M. cedrosensis (P < 0.001).
27(3-4):233-256, 1988(89)
247
Fig. 8. Selected characters showing variation in male and female genitalia
in Mitoura cedrosensis. Upper Row, Left: Dorsal view of saccus;
Right: lateral view of valva. Saccus and valvae outlined by solid line
in first specimen and dotted line in second. Center Row: Cornuti
(both specimens identical). Bottom Row: Female, ductus bursa
(sclerotized) and lamella antivaginalis.
margin of the postbasal band (hindwing underside) clearly separates M.
cedrosensis from any M. nelsoni or M. siva (Edwards) populations.
Adults of M. cedrosensis were found in close association with Cali-
fornia juniper ( Juniperus calif ornica Carr., Cupressaceae), which is
undoubtedly the larval host (Brown and Faulkner, 1984). A captive
female readily oviposited on the juniper, but the eggs were not viable.
California juniper, which generally exhibits a medium tall stature,
grows almost prostrate in the canyons and slopes of the north end of the
island. This aspect is so striking that the juniper was originally thought
to be an endemic species closely related to J. californica (Gentry, 1950).
Large stands of the juniper occur in scattered areas throughout much of
the island, and it is suspected that the Mitoura has a distribution
comparable to that of its larval host. Adults were collected in both the
spring and summer probably representing 2 broods, consistent with
other low elevation southern California Mitoura populations, i.e., M.
loki and M. thornei.
11. Strymon columella istapa (Reakirt)
All specimens of S. columella taken on Cedros were collected on
hilltops or prominent knolls in the area northwest of El Pueblo on the
south end of the island. The larval hostplant in southern California,
Sida hederacea (Dougl. ex Hook.) Torr. (Malvaceae), is not known from
Cedros, but several other malvaceous plants are present. Although
i currently referred to S. columella istapa, specimens from Baja Cali-
fornia and adjacent southern California are undoubtedly subspecifically
248 J. Res. Lepid.
distinct from mainland istapa (Clench, in litt.). Insular and peninsular
specimens are indistinguishable.
Specimens examined: vicinity El Pueblo, 29 March 1983 (1 male), 3
April 1983 (1 male); vicinity Cerro de Cedros, 1 July 1983 (5 males, 1
female).
12. Strymon melinus pudica (Hy. Edwards)
S. melinus is probably the most widespread Nearctic hairstreak. It
occurs throughout much of the United States, extending into northern
Mexico, and occupies a tremendous variety of habitats from mountains
to deserts. S. melinus was first reported from Cedros by Rindge (1948),
and it was encountered commonly in both spring and summer of 1983.
This insect is a frequent hilltopper and was collected on both ends of the
island. A number of potential larval hosts are available including
Malua, Phaseolus , and Eriogonum. Larvae were abundant on the flower
heads of the endemic Eriogonum molle Greene (Polygonaceae) in the
summer of 1983. All Baja California material is best referred to
subspecies pudica.
Specimens examined: vicinity Punta Norte, 28 February 1941 (1 male
leg: J. Garth, 30 March 1983 (2 males, 1 female) 31 March 1983 (1 male),
1 April 1983 (3 males), 3 July 1983 (1 female), ex-larva, emerged 25 July
1983 (1 female), ex-larva, emerged 31 July 1983 (1 male); vicinity El
Pueblo, 29 March 1983 (10 males), 3 April 1983 (2 males, 1 female), 4
April 1983 (2 males); vicinity Cerro de Cedros, 1 July 1983 (2 males, 6
females); Gran Canon, 2 July 1983 (2 males); Punta Prieta, 5 July 1983
(2 males).
13. Brephidium exilis (Boisduval)
B. exilis occurs throughout Baja California, ranging from the coasts to
the deserts and from Tijuana to La Paz. On Cedros it was most
commonly encountered in heavily disturbed areas where weedy A tri-
plex and Chenopodium (Chenopodiaceae) formed dense clumps. Two
such habitats include the vicinity of the fishing village at Punta Norte,
and near El Pueblo at the south end of the island. Specimens from
Cedros are indistinguishable from those collected elsewhere on the
peninsula. B. exilis is also known from all the California Channel
Islands (Miller, 1984).
Specimens examined: vicinity Punta Norte, 25 February 1932 (1
male), 28 February 1941 (1 female), both leg: J. Garth; Punta Norte 20-
22 March 1981 (1 male, 1 female), 30 March 1983 (1 male, 1 female), 1
April 1983 (2 males, 1 female), 2 April 1983 (1 female); vicinity El
Pueblo, 29 March (1 male, 1 female), 4 April 1983 (3 males), 13 July
1983 (1 female); Morro Redondo, 5 April 1983 (1 male); Cerro de Cedros,
1 July 1983 (1 male); Gran Canon, 2 July 1983 (2 males, 1 female).
14. Leptotes marina (Reakirt)
L. marina was first reported from Cedros by Rindge (1948). Although
not collected by us in the spring, L. marina was quite common in the
27(3-4):233-256, 1988(89)
249
summer of 1983. This species occurs on both the north and south ends of
the island, particularly in the lowlands and in disturbed areas. L.
marina ranges the length of Baja California, extending north into
California; it has also been collected on Santa Catalina, Santa Cruz, and
Anacapa Islands (Emmel and Emmel, 1973; Langston, 1980; Miller,
1984).
Specimens examined: vicinity Punta Norte, 25 February 1932 (1
male), leg: J. Garth, 3 July 1983 (2 males, 1 female); vicinity Cerro de
Cedros, 1 July 1983 (2 males, 1 female).
15. Hemiargus ceraunus gyas (Edwards)
Although quite rare and localized in the spring, H. ceraunus was
abundant and widespread in the summer of 1983. This species was
collected almost everywhere on the island, although most commonly at
low elevations.
H. ceraunus gyas is distributed the length of Baja California and in a
variety of habitats. It is multiple brooded and several genera of
Fabaceae, including Astragalus, which is available on Cedros, are
utilized as larval hosts.
Specimens examined: vicinity Punta Norte, 25 February 1932 (1
female), leg: J. Garth, 3 July 1983 (5 males, 1 female); Punta Morro
Redondo, 5 April 1983 (5 females); vicinity Cerro de Cedros, 1 July 1983
(16 males, 6 females); Gran Canon, 2 July 1983 (2 males, 1 female);
Punta Prieta, 5 July 1983 (6 males); El Pueblo, 13 July 1983 (1 male).
16. Philotes sonorensis (Felder and Felder)
Restricted to California and adjacent Baja California, P. sonorensis
reaches its southernmost limit on Isla de Cedros, slightly disjunct from
the nearest peninsular population. First collected on Cedros by J. Garth
in 1941, P. sonorensis was quite common in the canyons of the north end
of the island in the spring of 1983. Typically one of the earliest spring
fliers in coastal areas peaking in February and March, our captures in
late March and April seem unusually late; the July record is extraor-
dinary. Several species of Dudleya (Crassulaceae) are available as
larval hosts; a single larva was observed feeding on Dudleya pachy phy-
lum (Moran and Benedict, 1981).
Specimens examined: vicinity Punta Norte, 28 February 1941 (1
male), leg: J. Garth, 20-22 March 1981 (3 males, 1 female), 30 March
1983 (4 males), 31 March 1983 (7 males), 1 April 1983 (8 males), 3 July
1983 (1 female).
17. Euphilotes hattoides garthi Mattoni
Rindge (1948) first reported E. hattoides from Cedros; he also re-
cognized that this insular population was phenotypically distinct.
Shields (1975) later referred to two males from Cedros as conforming to
his description of E. hattoides allyni. Mattoni (1988) recently described
the Cedros population as garthi , and examined its relationships within
the hernardino cluster of subspecies.
250
J. Res. Lepid.
E. battoides garthi is endemic to Isla de Cedros and probably occurs
throughout the island wherever Eriogonum fasciculatum (Benth.)
(Polygonaceae) is found. We encountered it most frequently in the
canyons and washes of the north end of the island in spring 1983. E.
battoides garthi represents the southernmost subspecies of the E.
battoides complex. The nearest battoides population occurs about
130 km northeast in the northern central desert region of the peninsula
(Brown and Faulkner, 1984).
Specimens examined: canyons west of Punta Norte, 30 March 1983 (2
males), 1 April 1983 (5 males, 4 females), 2 April 1983 (1 male), 1 July
1983 (1 male, 1 female), 3 July 1983 (2 females); Cedros Island, no
further locality data, 15 March 1939 (3 males), no leg data, LACM, 18
March 1939 (1 female), no Ige data, CAS.
18. Celastrina ladon echo (Edwards)
C. ladon echo , the westernmost subspecies of the widespread Nearctic
ladon complex, has an extensive range from British Columbia to Baja
California (Langston, 1975). It is also known from several islands off the
western coast of California (Emmel and Emmel, 1973; Langston, 1979;
Miller, 1984). The population on Cedros represents an isolated and
disjunct outpost. The echo blue was the most common lycaenid en-
countered on Cedros; in the spring of 1983 it was particularly abundant
in the canyons and washes of the north end of the island; and in July
1983, it was most common in the vicinity of Cerro de Cedros. The
abundance of freshly emerged adults in both spring and summer
indicates that the species is at least double brooded on Cedros. Larval
hostplants encompass several families and many genera including
Rhus and Lotus , both available on Cedros.
Specimens examined: vicinity Punta Norte, 25 February 1932 (2
males, 2 females), 28 February 1941 (21 males, 3 females), leg : J. Garth,
20-22 March 1981 (1 female), 30 March 1983 (6 males, 2 females), 31
March 1983 (6 males, 1 female), 1 April 1983 (2 males, 2 females), 2
April 1983 (4 males), 3 July 1983 (2 males, 2 females); vicinity Cerro de
Cedros, 1 July 1983 (3 males); Gran Canon, 2 July 1983 (1 male, 2
females).
RIODINIDAE
19. Apodemia mormo virgulti Behr
Figures 9 and 10
A dark segregate of Apodemia mormo virgulti with greatly reduced
hind wing orange occurs in central and north central Baja California.
Opler and Powell (1961) have indicated that these dark populations
may warrant subspecific recognition. Specimens from Cedros are consi-
stent in maculation and color, and represent the extreme in this
phenotype.
27(3-4):233-256, 1988(89)
251
Fig. 9. Apodemia mormo virgulti, female, uppersurface, Isla de Cedros.
Fig. 10. Apodemia mormo virgulti, female, undersurface, Isla de Cedros.
Apodemia mormo occurs in the canyons and washes of the north end of
the island, generally associated with Eriogonum fasciculatum (Benth.)
(Polygonaceae). It is known from the south end but is much less common
there. The presence of A. mormo adults from March through July
appears to illustrate the extended flight periods of coastal species
previously suggested by Langston (1975). Although A. mormo extends
all the way to the cape region of the peninsula, in the form of Apodemia
mormo maxima (Weeks), Isla de Cedros is near the southernmost
distribution of the virgulti- like phenotype.
Specimens examined: Punta Norte, 20-22 March 1981 (15 males, 5
females), 30 March 1983 (2 males), 31 March 1983 (2 females), 1 April
1983 (1 male, 1 female), 2 April 1983 (1 female), 3 July 1983 (1 male, 3
females); vicinity Cerro de Cedros, 1 July 1983 (1 male, 1 female).
20. Calephelis wrighti Holland
C. wrighti occurs throughout Baja California. First collected on
Cedros by Hanna and Slevin in 1922, C. wrighti was one of the more
common butterflies that we encountered in 1983. It was collected on
both the north and south ends of the island, commonly in association
with Behhia juncea (Benth.) Green (Asteraceae), the larval host.
Considerable confusion exists in older literature between C. wrighti
and C. nemesis (Edwards). Rindge’s (1948) records of Calephelis nemesis
australis (Edwards) from Cedros are almost certainly misdetermined
specimens of C. wrighti.
Specimens examined: Cedros Island, 22 July 1922 (1 male, 1 female),
le: Hanna and Slevin, CAS; El Pueblo, 29 March 1983 (1 male, 1 female),
El Pueblo, 3 April 1983 (2 males, 1 female), 4 April 1983 (3 males, 4
females); Punta Norte, 30 March 1983 (2 males), 31 March 1983 (2
females), 2 April 1983 (1 male); Gran Canon, 2 July 1983 (2 males);
Punta Prieta, 5 July 1983 (1 male, 1 female).
252
J . Res. Lepid.
NYMPHALIDAE
21. Vanessa cardui (Linneus)
In the spring of 1983 V. cardui was abundant throughout southern
California and northern Baja California, exhibiting one of its periodic
unidirectional migrations. The butterfly was extremely common on
Cedros, with both flight-worn and freshly emerged adults evident.
Larvae were plentiful on both the weedy Malua parviflora L. (Malva-
ceaae) and Lupinus sparsiflorus Benth. (Fabaceae). In years of excep-
tional abundance such as 1983, an extremely wide variety of larval
hostplants are exploited by V. cardui (Emmel and Emmel, 1973).
First reported from Cedros by Rindge (1948), V. cardui is commonly
encountered throughout the peninsula of Baja California.
Specimens examined: vicinity Punta Norte, 28 February 1941 (4
males, 2 females), leg: J. Garth; El Pueblo, 29 March 1983 (1 male, 2
females), 4 April 1983 (1 male), ex-larvae, ex-Lupinus , (4 females)
emerged as follows: 20 April 1983, 19 April 1983, 18 April 1983, and 16
April 1983; Punta Norte, 30 March 1983 (4 males, 6 females), 31 March
1983 (1 male, 1 female), 1 April (1 female), 2 April 1983 (1 male, 1
female); vicinity Cerro de Cedros, 3 April 1983 (1 female).
22. Vanessa annabella (Field)
Another of the widespread vanessas, annabella , was encountered in
large numbers on Cedros in the spring of 1983. It was particularly
common at low elevations and in disturbed areas. V. annabella occurs
the length of the peninsula of Baja California; insular and peninsular
specimens are indistinguishable. Vanessa uirginiensis (Drury), a notor-
ious pioneer species present in many insular situations, was absent
from Cedros; its hostplant, Gnaphalium , is known from the island.
Specimens examined: Punta Norte, 30 March 1983 (1 female), 31
March 1983 (1 male, 1 female), 1 April 1983 (1 female), 2 April 1983 (2
females); El Pueblo, 29 March 1983 (2 males, 2 females), 4 April 1983 (2
males, 1 female); vicinity Cerro de Cedros, 1 July 1983 (1 male).
DANAIDAE
23. Danaus gilippus strigosus (Bates)
A common inhabitant of the desert regions of southern California and
Arizona, the striated queen occurs the length of the peninsula of Baja
California, and in a variety of habitats. Although rather uncommon in
the spring, with 1 or 2 individuals observed each day, D. gilippus was
quite common in the summer of 1983 on both ends of the island. This
species was reported from Cedros by Rindge (1948); his record is from
the fall when the butterfly is probably more common. The only potential
larval host available on Cedros is Asclepias subulata Decne. (Asclepia-
daceae). Specimens of D. gilippus collected on the island may represent
breeding residents as well as strays from the mainland.
27(3-4):233-256, 1988(89)
253
Specimens examined: Punta Norte, 1 April 1983 (2 males) vicinity El
Pueblo, 4 April 1983 (1 male); vicinity Cerro de Cedros, 1 July 1983 (1
female).
Discussion and Summary
Is la de Cedros supports an exceedingly depauperate butterfly fauna,
primarily as a consequence of a limited mainland species pool. The
Viscaino-Magdalena region of the peninsula directly adjacent to Cedros,
lies near the southern end of the Californian Province influence, and
considerably beyond the northern extremity of the Cape Province
influence.
The biotic diversity of the Californian Province attenuates north of
Isla de Cedros, with strays rarely occurring as far south as 28°N.
Seasonal meteorological patterns do not favor immigration from this
direction. Californian elements present on Cedros presumably repre-
sent relict populations separated from their contiguously distributed
mainland populations since the Pleistocene.
Most of the Neotropical species inhabiting the Cape Region scarcely
extend northward into the Viscaino Desert. Even such well known
dispersers as Phoebis agarithe and Ascia monuste are yet to be recorded
from Cedros. Apparently the combination of the Viscaino Desert and the
Pacific Ocean together present an almost impenetrable barrier to Cape
Province species' immigration to the island. The lack of suitable larval
hostplants would also act to preclude these species from establishing in
the event they were to be introduced to the island. Elements of
Neotropical affinity present on Cedros represent widespread species
occurring from Central America northward to at least southern
California.
A speculative explanation for the conspicuous absence of butterflies
common to the Californian province, such as Icaricia acmon , E veres
amyntula, Caliophrys dumetorum , and representatives of the genus
Satyrium , can be extracted from the equilibrium theory of island
biogeography. Since most of the above species occur sympatrically in
cismontane Baja California with many of the island’s resident species, it
is possible that several of these missing butterflies were formerly
resident on the island. Their absence may be partially explained by the
reduction in floral and faunal diversity of the island which occurred as a
result of its over-saturated biota following its separation from the Baja
California peninsula. Many of the expected but absent species may have
gone extinct on Cedros, and because prevailing conditions did not favour
southward dispersal, were never reintroduced.
According to Pielou (1979), a low species diversity exists on islands
not only as a function of land area and distance from the mainland, but
also with aspects of community complexity acting to maintain this
status. The fragmentary or patchy occurrence of suitable larval host-
254
J. Res. Lepid.
plants makes it difficult for immigrant species of moderate host speci-
ficity to become established. The effects of patchy habitats in insular
situations are discussed by Powell (1981) regarding the introduction of
insect species onto Santa Cruz Island, California.
It should also be noted that man’s presence on and his introduction of
herbivores to Isla de Cedros has had little impact on the island’s native
flora and fauna. Man’s activities have been restricted to the south-
eastern end of the island and in a lesser degree to the areas in and
around the fishing village and abandoned copper mine on the northeast
end. There is no current effort at agriculture, and its inherent ecological
impact, owing to the poor soil conditions and undependable rainfall.
There is barely enough groundwater from springs to provide for the
growing needs of the village population, with none available for
irrigation. Feral grazing animals, especially goats and pigs, are re-
stricted to the southeast end of the island and have overall resulted in
only minor impact on the island’s native vegetation. This point is in
sharp contrast to the effect that uncontrolled feral animals, mainly
goats, have had on the flora of other Coastal Pacific islands, such as
Santa Catalina Island (Coblentz, 1980) and San Clemente Island
(Faulkner, personal observation) in California, and Isla Guadalupe
(Moran and Lindsey, 1950) in Baja California, Mexico.
Acknowledgements. We thank John Garth of the Allan Hancock Foundation
for sharing collecting notes and specimens he compiled during numerous visits
to Isla de Cedros. We are also indebted to David Weissman for including us in a
July 1983 collecting expedition to the island as well as the crew of the sloop
Diamaresa, supplied by the Island Packers Company, Ventura, California, for
providing transportation, meals, and lodging during the 2-week trip. Dr. Jerry
A. Powell and Scott E. Miller both contributed significant editorial suggestions.
Shirley Latislaw provided the map of Isla de Cedros and Rudolf H. T. Mattoni
provided the genitalic drawings and discussions.
Literature Cited
BROWN, J. W., 1983. A new species of Mitoura from southern California (Lepi-
doptera: Lycaenidae). J. Res. Lepid. 21(4):245-254 [“1982”].
— — & D. K. FAULKNER, 1984. Distributional records of certain Rhopalocera
in Baja California, Mexico, with the description of a new subspecies of
Papilio (Heraclides) astyalus (Godart) (Lepidoptera: Papilionidae). Bull.
Allyn Mus. 83:1-9.
COBLENTZ, B. E., 1980. Effects of feral goats on the Santa Catalina Island
ecosystem, in Power, D. M. (ed.), The California Islands: Proceedings of a
Multidisciplinary Symposium. Santa Barbara Museum of Nat. Hist.,
pp. 167-172.
EMMEL, T. C. & J. F. EMMEL, 1973. Butterflies of southern California. Nat. Hist.,
Mus. Los Angeles Co., Sci. Ser. 25:1-148.
GENTRY, H. S., 1950. Land plants collected by the Velero, III, Allan Hancock
Pacific Expeditions, 1937-1941. Allan Hancock Foundation Pub. Univ. So.
Calif. 13:5-246.
27(3-4):233-256, 1983(89)
255
GOULD, F. N. & R. MORAN, 1981. The grasses of Baja California, Mexico. San Diego
Soc. Nat. Hist. Mem. 12:1-140,
HALE, G., 1941. A survey of the vegetation of Cedros Island, Mexico. 96 pp.
Unpublished thesis, Univ. of Calif., Los Angeles.
HASTINGS, J. R. & R. R. HUMPHREY (eds.), 1969. Climatological data and statistics for
Baja California. Technical Reports on the Meteorology and Climatology of
Arid Regions, no. 18. Univ. of Ariz., Inst, of Atmospheric Physics.
HOWE, W. H., 1975. The Butterflies of North America. Doubleday and Co., Inc.
Garden City, New York.
KILMER, F. H., 1977. Reconnaissance geology of Cedros Island, Baja California,
Mexico. Bull. So. Calif. Acad. Sci. 76:91-98.
LANGSTON, R. L., 1975. Extended flight periods of coastal and dune butterflies in
California. J. Res. Lepid. 13(2):83-98. [“1974"].
— , 1975. The genus Celastrina, in Howe, W. H., The Butterflies of North
America. Doubleday and Co., Inc. Garden City, New York.
, 1980. The Rhopalocera of Santa Cruz Island, California. J. Res. Lepid.
18(1):24— 35. [“1979”].
LEWIS, L. R. & P. E. EBELING, 1971. Sea Guide , volume 2, Baja, SEA Publications,
Inc. Newport Beach, Calif.
LIBBY, W., M. BANNISTER & Y. LINHART, 1968. The pines of Cedros and Guadalupe
Island. J. Forestry 66:846-853.
MacARTHUR, R. H. & E. O. WILSON, 1967. The theory of island biogeography.
Monographs in population biology no. 1. Princeton Univ. Press.
MacNEILL, C. D., 1975. The family Hesperiidae, in Howe, W. H., The Butterflies of
North America. Doubleday and Co., Inc. Garden City, New York.
MATTONI, R. H. T„ 1988. A new subspecies of Euphilotes battoides (Behr) with
notes on relationships among the Euphilotes battoides bernardino (Barnes
and McDunnough) group of subspecies (Lepdioptera: Lycaenidae). J. Res.
Lepid. (in press).
MILLER, L. D. & F. M. BROWN, 1981. A catalogue/checklist of the butterflies of
America north of Mexico. Lepid. Soc. Mem. 2:1-280.
MILLER, S. E., 1984. Butterflies of the California Channel Islands. J. Res. Lepid.
23(4):282-296.
MORAN, R., 1972. Las plantas vasculares de la Isla de Cedros. Calafia 2(l):34-36.
& M. BENEDICT, 1981. Dudley a pachyphytum, of Isla Cedros, Mexico.
Cactus and Succulent Journal 53:132-136.
& G. LINDSEY, 1950. Guadalupe Island. Desert Plant Life 22:3-9.
NELSON, E., 1921. Lower California and its natural resources. Mem. Nat. Acad.
Sci. 16:1-194.
OPLER, P. A. & J. A. POWELL, 1961. Taxonomic and distributional studies on the
western components of the Apodemia mormo complex (Riodinidae). J. Lepid.
Soc. 15:145-171.
PIELOU, E., 1979. Biogeography. 315 pp. John Wiley and Sons, New York.
POWELL, J. A., 1958. Additions to the knowledge of the butterfly fauna of Baja
California Norte. Lep. News 12:26-32.
— =, 1981. Five insects believed to be newly established or recolonized on
Santa Cruz Island, California (Dermaptera, Lepidoptera). Bull. So. Calif.
Acad. Sci: 79(3):97-108.
RINDGE, F. H., 1948. Contributions toward a knowledge of the insect fauna of
Lower California, Lepidoptera: Rhopalocera. Proc. Calif. Acad. Sci. 24(8):
289-312.
256 J. Res. Lepid.
SHAPIRO, A.M., 1978. The assumption of adaptivity in genital morphology. 1979.
J. Res. Lepid. 17(l):68-72.
SHIELDS, O., 1975. Studies on North American Philotes (Lycaenidae). IV. Taxono-
mic and biological notes, and new subspecies. Bull. Allyn Mus. 28:1-36.
— — — , 1984. Comments on recent papers regarding western Cupressaceae
feeding Callophrys (Mitoura). Utahensis 4(4):51-56.
WIGGINS, I. A., 1980. Flora of Baja California. 1025 pp. Stanford Univ. Press.
WILCOX, B. A., 1978. Supersaturated island faunas: a species-age relationship for
lizards on post-Pleistocene landbridge islands. Science 199:996-998.
Addenda. Colias cesonia Stoll has been documented recently from
Isla de Cedros, bringing the species total to 24. The single record
probably represents a stray from the peninsula rather than an indi-
genous population.
Journal of Research on the Lepidoptera 27(3-4):257-258, 1988(89)
Opinion.
Parallelism and Phylogenetic Trees
James A. Scott
60 Estes Street, Lakewood, Colorado 80226
Nearly all of Brock’s (1988) statements in his criticism of my (Scott
1986) phylogeny of the advanced Ditrysia and Macrolepidoptera mere-
ly (but correctly) demonstrate that many character states of one Ditry-
sia group also independently occur in one or more other Ditrysia
groups. But every worker on Ditrysia knows this; I knew of most of
these independent occurrences when I wrote Scott (1986). Merely
because there is a parallelism of a trait in several taxa does not
invalidate the use of the trait as a shared derived trait for either taxon;
if parallelism is real, the structure must by definition be a shared
derived trait in each group in which it occurs. And most of the parallel-
isms that Brock cites involve primitive Ditrysia (Tineoid superfamilies
or Cossoidea-Castnioidea-Zygaenoidea); one of the major points of
Scott (1986) is that the phenetic distance between these lower Ditrysia
and the Macrolepidoptera is so great that direct phylogenetic links
between them are inconceivable. Brock fails to acknowledge the vast
morphological gap (demonstrated by Scott 1986) between macro-
lepidoptera and lower Ditrysians such as Cossoidea-Castnioidea-
Zygaenoidea and the Tineoid superfamilies; just counting the number
of morphological differences between these groups and the Macro-
lepidoptera families clearly shows that they are not direct ancestors of
any Macrolepidoptera, so parallelisms involving them are not directly
relevant to Macrolepidoptera. Brock uses independent occurrence of
many traits as justification for not proposing any phylogenetic scheme
at all (his 1971 tree-like drawing resembling a phylogenetic tree was
not derived from any list of characters using repeatable methods). But
the fact of evolution means that the ancestor of every Ditrysia group
had certain character states when it branched away from the remain-
ing Ditrysia; therefore it is our job to deduce those character states
were. Avoiding making a tree merely because of the complication of
parallelism in some traits is not progress; progress is constructing
trees and selecting the most likely tree, and listing the exact character
changes involved so that other workers can verify or change the tree;
progress is studying the characters in detail and the distribution of
characters within taxa and reassigning those taxa that were misplaced
(some reassignments may eliminate false parallelisms).
Brock’s criticism (1988) also contains some misstatements of fact:
Scott (1986) did not claim that secondary setae are absent in Noctuoi-
dea; Scott wrote (p. 35) that “Noctuoidea. . .generally lack secondary
setae” and his Table 1 shows that they are sometimes present. The
Pyraloidea-Macrolepidoptera ancestor pupa was obtect in the sense
258
J. Res. Lepid.
that only abdomen segments 5-6 moved. Sphingidae pupae occur in an
earthen cell, but do any have true dense-silk cocoons? HesseFs (1969)
figures clearly show that only Papilionoidea-Hesperioidea and some
Cossidae have an aortic enlargement (“chamber”); the aorta is not en-
larged much in other groups. The anapleural cleft IS a sulcus in
Hesperiidae (fused, no longer a cleft). Abdominal segment 2 sternal
apodemes are especially small in Rhopalocera. Maxillary palpi are 2-
segmented in Baronia (Papilionidae) as well, but still they are very
small in all Macrolepidoptera. Mandible remnants are not protuberant
in Rhopalocera; in this group the name mandible remnant (and the
erroneous name pilifer) does not represent an actual functional struc-
ture as it does in Cossoidea where the mandible remnants are definite
bumps. Thus the name mandible remnant in Rhopalocera is not useful
for morphological comparison, but is useful only for the convenience
of Lepidopterist’s descriptions; stating that mandible remnants are
larger in Rhopalocera is misleading because the correct functional
statement is that the sulci surrounding the absent “mandible rem-
nant” are farther from each other. Lepidopterists’ common practice of
naming an ordinary expanse of exoskeleton as though it is some real
functioning structure is frequently misleading; the truth is often that
the area is just another undistinguished portion of body wall, and the
functional structures that changed in the ancestor of the taxon are
actually the sulci (which strengthen the cuticle during locomotion) or
membranes (which allow movement of legs wings neck etc.)
Independently-possessed character states make the search for the
true Ditrysia tree difficult, but do not justify the abandonment of the
effort. Brock’s comments do not mean that Scott’s (1986) phylogenetic
tree is wrong and should be changed. Brock should apply his expertise,
and give us his phylogenetic tree, complete with character changes
clearly placed on the branches of the tree (not some pseudo-phylogene-
tic gradistic tree conjured up with unknown methods, divorced from
real data). And because parallelisms are common, perhaps a numerical
taxonomy phenetic classification of the Ditrysia would be useful,
merely to convince Lepidopterists that some superfamilies cannot be
direct ancestors of some other superfamilies. I agree with Brock that
too many Lepidoptera taxonomists refuse to apply their knowledge to
study of phylogeny; these Lepidoptera taxonomists only seem to care
about species/genera-level taxonomy, and once in a while they stray a
bit by proposing a new obscure family; they “worship the god of genita-
lia” as they prepare drawings of the male and female genitalia that
distinguish their species.
Literature Cited
BROCK, J. P. 1988. Reply to Scott’s criticism. J. Res. Lepid. 26:240-247.
SCOTT, J. A. 1986. On the monophyly of the Macrolepidoptera, including a re-
assessment of their relationship to Cossoidea and Castnioidea, and a re-
assignment of Mimallonidae to Pyraloidea. J. Res. Lepid. 25:30-38.
Journal of Research on the Lepidoptera
27(3-4):259-271, 1988(89)
Notes
A Significant New Hostplant record for Pieris virginiensis (Pieridae)
Since Klot’s (1935, J. New York Entomol. Soc. 53:139-142) original description
of the larva of Pieris virginiensis , this butterfly has been presumed to be
monophagous, feeding exclusively on Dentaria. More recently, Shapiro (1971,
Entomol. News 82:13-15) noted that the biology of this butterfly seemed
completely tied to the phenology of its ephemeral hostplant. He found that in the
laboratory, P. virginiensis females will oviposit on, and larvae will develop on
several species of mustards. But that under natural conditions, Dentaria is the
only host, probably because it is the only mustard which is usually present in
this butterfly’s habitat.
Chew (1980, Oecologia 46: 347-353) reiterated this position noting that
populations with which she was familiar had only two species of mustards to
choose between, D. diphylla Michx. and D. laciniata Muhl. . Likewise, the
population studied in depth by Cappuccino and Kareiva (1985, Ecology 66:152-
161) could choose between only these two hostplants. These authors quantita-
tively assessed and reaffirmed the close relationship between P. virginiensis s
biology and the phenology of Dentaria.
In central Ohio, P. virginiensis occurs in isolated populations inhabiting
wooded creek bottoms, usually with shale banks. In Morrow County, Ohio, we
observed females ovipositing at a site where Dentaria is abundant in a creek
bottom, but another mustard, Arabis laevigata (Muhl.) Poir, occurs as widely
scattered plants on surrounding shale banks. From a distance, we noted P.
virginiensis females settling on the Arabis, but we were unable to observe actual
oviposition. However, upon examination, we found that these plants held
several Pieris ova, presumably deposited by one or more of the females we
observed. Later, by searching Dentaria we located two additional ova. Since all
of these ova could have been deposited by P. rapae, which is also common in this
area, we reared them on their original oviposition substrates. From these
rearings we obtained two P. virginiensis pupae reared on Dentaria and one
reared on Arabis.
These observations confirm the suggestion by Shapiro and Chew that P.
virginiensis is usually monophagous not because female oviposit only on
Dentaria, but because Dentaria is usually the only mustard available in their
restricted habitat for them to oviposit on. Our preliminary observations indicate
that Arabis may be more attractive to ovipositing females than Dentaria.
Several Arabis plants located by us had several ova attached (one plant had six)
while many Dentaria plants had to be searched to locate our two ova. However,
Arabis is rare relative to Dentaria at this site, and we assume that Dentaria is
the primary ovipositional substrate.
Arabis may be more attractive to ovipositing females, and because it is not as
ephemeral as Dentaria it might allow more time for completion of larval
development. However, its rarity in this habitat, and sometimes high egg load
(indicating possible defoliation with no nearby mustards on which the larvae
could relocate) limit the possibility that P. virginiensis could adopt a biological
strategy that would allow it to become less dependent on Dentaria.
260
J. Res. Lepid.
Acknowledgments. We thank Dr. A. Shapiro, University of California, Davis,
for his help in preparing this note.
John A. Shuey, 731 Kerr Street, Columbus, Ohio 43215, John W. Peacock, 185
Benzler Lust Road, Marion Ohio 43302
Description of the Hitherto Unknown Female of Acerbas suttoni Russell
(Hesperiidae)
The hitherto unknown female of Acerbas suttoni Russell is described as
follows. The conspecificity of A. latefascia and A. suttoni are discussed below.
Acerbas suttoni Russell, 1984, Ent. Rer., 44:154-156; Figs 4a, b, 5, 6.
Female (Fig. 1): Forewing 20 mm. Head, palpi, ventral thorax, costa of legs,
bases of fore wing and ventral hindwing with green reflection. Antenna black,
long, 3/5 length of costa. Abdomen dark brown; segments with faint white
hairs on posterior margin. Dorsal forewing: dark brown, detached hyaline spots
in spaces 2 and 3, small upper cell spot, no apical and lower cell spots. Dorsal
hindwing: white median band from dorsum to vein 6, obscured in space lb. Cilia
brown, becoming paler toward tornus. Ventral forewing: similar to dorsal side,
but dorsum paler. Ventral hindwing: blackish brown, median band conspicuous
and sharply defined; break in space lb; trace of band reach to costa.
Material examined: Lambarese, 100 km N. of Palopo, Sulawesi, Indonesia. 28.
VI. 1966 (Bernice P. Bishop Museum, Honolulu).
Three species of Acerbas have been described from Sulawesi, of which only A.
azona Hewitson, 1866 has been known for a longtime. De Jong (1982, Ent. Ber.,
42:88-90) described A. latefascia from one female specimen from N. E. Sulawesi.
He suggested that A. latefascia could be considered a subspecies of A. duris
Mabille, 1883, though he mentioned that the examination of the male would be
necessary to establish the exact relationship of these two taxa. Two years later,
Russell (1984, Ent. Ber., 44:154-156) described A. suttoni from one male from
Central Sulawesi. He mentioned that A. suttoni was the nearest to A. duris
dorka Evans, 1949 from Borneo in appearance. However, he did not suggest the
relationship between A. latefascia and A. suttoni. I suggested (in litt.) the
conspecificity of A. latefascia and A. suttoni to both of the authors before I found
Fig. 1 Female of Acerbas suttoni: dorsal and ventral view.
27(3-4):259-271, 1988(89)
261
Fig. 2. Distribution map of Acerbas duris complex.
the female specimen described here in the Bishop Museum. De Jong (pers.
comm.) suggested that further examination was necessary. Russell (pers.
comm.) denied my suggestion because he felt that the two taxa were clearly
differentiated and that only slight sexual dimorphism were known in this
genus. After examining the male and the female of A. suttoni (I could not
examine A. latefascia directly, but with a photograph), I retain my opinion that
the two taxa could be the same species. The female markings do not differ
markedly between A. latefascia and A. suttoni. The only significant difference is
that the hindwing median band is clearer and wider in A . latefascia than in A .
suttoni. This degree of difference, however, is not uncommon within intraspecific
variation. Biogeographically, the two taxa are allopatric, and are, no doubt,
congeneric with A. duris as both authors suggested (Fig. 2). I believe A.
latefascia and A. suttoni should be treated as subspecies of a single species, but I
withhold conclusive judgment. A. suttoni is now known from two males (the
holotype and another in Tsukiyama collection in Japan) and the single female
which are described here. A. latefascia is only known from single type female.
Discovery of male A. latefascia is desirable to confirm my suggestion.
I thank H. Tsukiyama for permitting the examination of his collection, R. de
Jong and A. Russell for useful comments. Special thanks to S. Miller for review
of the manuscript and G. Uchida for photography.
Hideyuki Chiba, Department of Entomology , University of Hawaii, 3050 Made
Way, Honolulu, HI 96822, U. S. A.
262
J. Res. Lepid.
Homosexual Pseudocopulation in Eucheira socialis (Pieridae).
Eucheira socialis Westwood is a bizarre endemic Mexican Pierid displaying a
variety of degenerate morphological and behavioral traits associated with
intense inbreeding and gregariousness (Shapiro et al., in preparation). Its
mating behavior is extremely simplified; there is essentially no courtship; males
approach and attempt to copulate with females, which either accept them or
walk away rapidly. Both sexes mate multiply, and in some populations
copulation may occur in the male-superior-dorsally position characteristic of
Orthopterans and Coleopterans as well as most other non-Lepidopterous
insects. Simultaneous multiple courtships of a single female are frequent, and
at times males will attach to the side of the abdomen of an already-copulating
female. Males have also been observed copulating with recently-dead females,
and with each other.
Fig. 1 . Living (right) and dead male Eucheira socialis in amplexus. The live male
walked about vigorously dragging his dead partner.
The illustration (fig. 1) is of a homosexual pseudocopulation between two
males from a single sibship originating in the state of Hidalgo, reared in the
laboratory. The pair was first noticed at 1600 hrs, 2. VI. 1987. At that time both
males were alive. At 0900 the following day they were still in amplexus, but one
was apparently dead. The photograph was taken at 1445, 3.VI; a few minutes
later, under the influence of the strobe lights, the surviving male began to flap
his wings violently and dislodged his burden. The dead male had been clasped
tightly onto the living male to the left of the dorsal abdominal midline. There
was no evidence of extrusion of any spermatophore material, and the survivor
lived until 9. VI, the last three days in refrigeration.
Studies of E. socialis have been made possible by a grant from the UC MEXUS
program.
Arthur M. Shapiro, Department of Zoology, University of California, Davis, CA
95616.
27(34):259-271, 1988(89)
263
Effect of Refrigeration on Hatching of Eggs of the Tasar Silk Moth
Antheraea mylitta Drury (Saturniidae)
Antheraea mylitta Drury is a semidomesticated Tasar Silk Moth with three
generations a year: July- August, September-October and November-December.
After the third broad, seed cocoons are preserved in commercial tasar insect-
aries until mid- June for egg production for first crop. During this period the
pupae usually diapause from Winter (December-F ebruary) until the following
summer (March- June). However, it has been recently observed that from May to
mid- June emergence occurs which leads to the production of fertile eggs. Their
resultant larvae cannot be reared due to lack of quality food plant leaves and
excessive outdoor temperature (39±4°C). The situation requires means to
preserve those unseasonal eggs until a favourable rearing time.
Considerable information is available on the effect of low temperature on
mulberry ( Bombyx mori) silkworm eggs (Yakoyama, 1962; Tanaka, 1964; Datta
etal, 1972; Devaiah & Thontadarya, 1975; Govindan & Narayana Swamy, 1986;
Narayana Swamy & Govindan, 1987; Tayade et al 1987) and eri (Phibsamia
ricini) silkworm eggs (Govindan et al, 1980, Chowdhury, 1982; Vishwakarma,
1982-83), but no such literature is available on the Tasar silkworm eggs.
Consequently an attempt was made to study the effect of refrigeration on
hatchability of the eggs of Tasar Silk Moth Antheraea mylitta as follows.
29,000 freshly oviposited eggs were collected at random from healthy coupled
female moths of the STV (Sukinda tri-voltine) race of Antheraea mylitta from
Mayurbhanj district of Orissa, India on 22 May 1987. These were kept under
room temperature (31 ± 2°C) as a common stock. Every day at 9 A.M., from the
first to seventh day following oviposition, 4000 eggs were taken at random from
the common stock and divided into four equal parts, each subjected to 24, 48, 72
or 96 hours of refrigeration (10 ± 1°C). Following treatment, the eggs were
allowed to incubate at room temperature until hatching. The remaining 1000
eggs served as the control. The hatching percentage of the refrigerated eggs
were noted and compared with the control. The experiment was repeated five
times during the same period under the same conditions.
Results and summarized in Table 1. The control eggs kept at room tempera-
ture (31 ± 2°C) showed 82.44 percent hatch. The refrigeration of 0 day old eggs
(fresh) for 24 and 48 hours indicated 82.34 and 82.28 percent hatch respectively,
not significantly different from the controls. The same eggs when refrigerated
for 72 & 96 hours showed reduction in hatching percentage. Cold treatment to 1
day old eggs for 24 hours also gave satisfactory hatching (82.32%), but in the
other treatments, as 1 day old eggs refrigerated for 48, 72 and 96 hours and 2, 3,
4, 5, and 6 day old eggs refrigerated for 24, 48, 72 and 96 hours, there was
reduced hatch.
The effect of refrigeration on Antheraea mylitta eggs of different ages
indicated that the eggs beyond 1 day old were more susceptible to damage at
lower temperatures. Vishwakarma (1982-83) observed that Philosamia ricini
eggs beyond third to fifth day old were more susceptible to low temperature (7 ±
2°C). Datta etal (1972) found increasing percentage of mortality in Bombyx mori
eggs under low temperature refrigeration (5 to 7°C). Govindan et al (1980)
reported that refrigeration oi Sarnia cynthia ricini Boisuduval eggs beyond 5
days old had adverse effect on hatching. Narayana Swamy and Govindan (1987)
264 J. Res. Lepid.
Table 1. Mean Hatching percentage of A. my/itta eggs refrigerated for
different time and at different ages.
Day after
oviposition
Age of
Eons -
HATCHING
PERCENTAGE
(Day)
24 hours
refrigeration
48 hours
refrigeration
72 hours
refrigeration
48 hours
refrigeration.
First
0
(Fresh
eggs)
82.34
82.28
74.48
78.14
Second
1
82.32
42.62
24.54
26.86
Third
2
46.30
50.04
72.06
32.82
Fourth
3
72.26
62.74
63.26
46.10
Fifth
4
60.36
62.22
74.10
24.42
Sixth
5
76.38
74.54
56.26
62.38
Seventh
6
72.56
62.12
44.22
60.32
reported that the hatching percentage of Bombyx mori eggs of blue stage
reduced with increase of refrigeration period from first day (83.70%) to the
seventh day (21.60%).
In general, percentage of hatching of Antheraea mylitta eggs declined with
increase of cold period with few exceptions (Table-1). A similiar trend was also
observed by Datta et al. (1972) in Bombyx mori eggs. Tayade et al (1987)
concluded that short refrigeration is better to minimise adverse effect on
hatching percentage of Bombyx mori eggs. Narayana Swany and Govindan
(1987) observed that the refrigeration of eggs of pure Mysore race of Bombyx
mori at blue stage negatively affected yield. Govindan and Narayan Swamy
(1986) reported that multivoltine silk worm eggs of Bombyx mori at eye spot
stage can be refrigerated for one day without decreasing yield.
Literature Cited
CHOWDHURY, S.N., 1982. Eri Silk Industry, Directorate of Sericulture and
weaving, Govt, of Asam, Gauhati. p. 1-177.
DATTA, R.K., s. sengupta & S.N. BISWAS, 1972. Studies on the preservation of
multivoltine silkworm eggs at low temperature. Indian J. Seric. 11 (1):
20-27.
DEVAIAH, M.C. & T.S. THONTADARYA, 1975. Effect of refrigeration on the hatching
of silkworm Bombyx mori Linnaeis eggs. Curr. Res. 4:65-66.
GOVINDAN, R. & T.K. NARAYANA SWAMY, 1986. Influence of refrigeration of eggs of
multivoltine silkworm, Bombyx mori L. at eye spot stage on rearing
performance. Sericologia 26(2): 151-155.
GOVINDAN, R., M.C. DEVAIAH, H.R. RANGASWAMY & C. THIPPESWAMY, 1980. Effect of
refrigeration of eggs of eri silkworm Sarnia cynthia ricini Boisuduval
on hatching. Indian J. Eric. 19(1): 13-15.
27(3-4):259-271, 1988(89)
265
NARAYANA SWAMY, T.K. & R. GOVINDAN, 1987. Effect of refrigeration of eggs of pure
Mysore race of silkworm Bombyx mori L. at blue stage. Entomon 12(2): 105=
107.
TAYADE, D.S., M.D. JAW ALE & P.K. UNCHEGAONKAR, 1987. Effect of refrigeration on
hatching of eggs of multivoltine Bombyx mori L. Sericologia 27(2): 297-299.
TANAKA, Y., 1964. Sericology. Published in English by the Central Silk Board,
Bombay, India. P. 1=277.
VISHWAKARMA, S.R., 1982-83. Effect of refrigeration of Silk worm, Philosamia
ricini Hutt eggs on the hatching (Lepidoptera: Saturniidae). Indian J. Seric.
21=22:36=39.
YOKOYAMA, T., 1962. Synthesized Science of Sericulture. Published in English by
the Central Silk Board, Bombay, India. P. 1=398.
A K. Dash & B.K. Nayak, State Sericultural Research Station, Orissa, Baripada-
757 001, India.
A Melanie Aberration of PhUotes sonorensis (Lycaenidae) from
California
The Sonora Blue, PhUotes sonorensis (Felder & Felder) with ts exquisite color
pattern of iridescent light blue, black and white markings, and red spots, is one
of California’s most beautiful butterflies. Locally common in the nondesert
portions of California, it is found in the mountains of Santa Barbara County,
Fig. 1.
Aberrant P. sonorensis: left, dorsal; right, ventral
Fig. 2. Habitat of P. sonorensis in Mission Canyon,
(see text to-spelling it is doubt)
266
J. Res. Lepid.
flying in F ebruary-March of each year. The author has collected this Blue in
Mission Canyon, in the Santa Ynez Mountains, and in Oso Canyon, in the San
Rafael Mountains.
The Santa Ynez Mountains, predominantly chaparral-oak woodpland, form a
2,000 ft to 4,000 ft. wall behind the cities of Carpinteria, Santa Barbara, and
Goleta Valley. Large colonies of Philotes sonorensis are to be found on its south
slope. Mission Canyon is a large watershed below La Cumbre Peak where a
number of small waterways converge to form Mission Creek, which runs down
through the Santa Barbara Botanic Gardens, behind the Santa Barbara
Museum of Natural History, through the city of Santa Barbara, and out to the
Pacific Ocean. There is a bridge in upper Mission Canyon, which crosses the
creek at the 1,400 ft. elevation, and above this bridge the foodplant, Dudley a
lanceolata (Nutt.) Britt. & Rose. Crassulaceae is abundent on the rocky hill-
sides, and supports a large colony of this Blue. (Figure 2)
Here in Mission Canyon a male melantic aberration of Philotes sonorensis,
was caught by Robert F. Denno, February 23, 1961. (Figure 1) This is a striking
aberration, with the black spots on both the forewings and the hindwings
smeared across the wings. Both wing surfaces are affected.
Richard Carl Priestaf, 5631 Cielo Avenue, Goleta, California 93117.
A Replacement Name for Lycaena editha nevadensis Austin
(Lycaenidae)
It has come to may attention that the name Lycaena editha nevadensis Austin
(J. Res. Lepid. 23:83, 1984) is an invalid junior primary homonym of Lycaena
nevadensis Oberthur (Etud. Ent. 20:pl. 4, fig. 54, 1986). To rectify this, I propose
the following replacement name for L. e. nevadensis :
Lycaena editha obscuramaculata
The description, types and type locality remain as in Austin (1984, op. cit .: 83-
88). The new name reflects the faintness of the maculation on the ventral
hindwing, characteristic of the subspecies. At the time of the original descrip-
tion, specimens were known only from northern Elko and Humboldt counties,
Nevada. Subsequently, I have examined material from southwestern Idaho
(Canyon County, CM) and Ruby Valley, Elko County, Nevada (AMNH).
I thank C. A. Bridges for pointing out the homonymy to me and F. H. Rindge
(American Museum of Natural History, AMNH) and J. E. Rawlins and C. W.
Young (Carnegie Museum of Natural History, CM) for allowing me to examine
specimens in their care.
George T. Austin, Nevada State Museum and Historical Society, 700 Twin Lakes
Drive, Las Vegas, Nevada 89107.
27(3-4):259-271, 1988(89)
267
Sex Characters of the Pupae of the Banded Moth, Cochylis hospes
Walsingham (Lepidoptera: Cochylidae)
The banded sunflower moth Cochylis hospes Walsingham, is a destructive
pest of commercial sunflower seed (Charlet and Busacca 1986, Charlet L. D. and
J. D. Busacca. 1986. Insecticidal Control of Banded Sunflower Moth, Cochylis
hospes (Lepidoptera: Cochylidae), Larvae at Different Sunflower Growth Stages
and Dates of Planting in North Dakota. J. Econ. Entomol. 79:648-650. Beregovoy,
personal communication). Increased cultivation of the sunflower and economic
loss due to banded sunflower moth damage has led to research into the biology
and control of this species. A description of sex characters useful in sexing the
pupae has not been published. Sexing the pupae is useful for behavioral or
physiological research where adults must be kept separate. The genital pri-
mordia and the length or diameter of the antennae have been used to sex pupae
of the sunflower moth Homoeosoma electellum Hulst (Rogers C. E. 1978. Sexing
pupae and adults of the sunflower moth. S. W. Entomol. 3:305-307). These
morphological characters were examined in the banded sunflower moth to
determine their usefulness in sexing pupae.
Pupae were obtained from a laboratory culture of C. hospes established at the
Metabolism and Radiation Research Laboratory, Fargo, North Dakota. A
Fig. 1. Ventral view of C. hospes
pupae showing sex char-
acters Arrows show genital
primorida. Solid lines indi-
cate posterior limit of
antennae. Ant = antenna
The genital primordia and length of
the antennae are useful in sexing the
pupae of C. hospes. The genital opening
of the male pupae is a single opening
on the 9th abdominal sternite (Fig.
1A). The genital opening of the female
is on the 8th abdominal sternite. The
female genital opening is longer than
in the male and appears divided into
two openings (Fig. IB). The deve-
loping antennae extend to the margin
separating the 2nd and 3rd abdominal
sternites in females. In males,
the antennae extend to the margin
separating the 3rd and 4th abdominal
segments (Fig. 1).
The genital primoridia are conclu-
sive morphological features to identify
the sex of C. hospes pupae. The lengths
of the developing antennae are reliable
sex characters, but experience is re-
quired to use this character. In the
authors experience, these morpho-
logical features were 100% reliable.
268
J. Res. Lepid.
dissecting microscope at 20X was used to examine 50 pupae. Conformation of
sex in relation to morphology was determined by dissection of the pupa for testes
and ovaries.
Reliable methods to sex C. hospes pupae should be useful in research on the
behavior and biology of this important pest species.
Acknowledgements. The assistance of Kelly Jones and Sharon Grugel in rearing
C. hospes in the laboratory is acknowledged.
John F. Barker, United States Department of Agriculture, Metabolism and
Radiation Research Laboratory, P.O.B. 5674, Fargo, ND 58105
Laboratory Rearing of Sandia xamixami (Lycaenidae, Eumaeini).
The population dynamics of Sandia xami in a small volcanic area near Mexico
City has been studied since 1984. The high numbers of eggs required to perform
life-table experiments lead us to attemp the rearing of S. xami in laboratory
conditions.
S. xami flies from central Mexico to the southern part of Texas and Arizona
(Scott, J.A. 1986. The butterflies of North America. Stanford University press.
Stanford, California. 583 pp.). In the Valley of Mexico S. xami can be found all
year with peaks of abundance in August-October, J anuary-March and, perhaps,
April and May (Soberon, J., C. Cordero, B. Benrey, P. Parlange, C. Garcia-Saez
and G. Berges. 1988. Ecol. Entom. 13(1): 71-76.). S. xami feeds on several
Crassulaceae species (Ziegler, J.B. and T. Escalante. 1964. Jour. Lep. Soc. 18:
85-89) but in the ecological reserve on the National University of Mexico
Campus at Mexico City, the main food plant is Echeueria gibbiflora. The larvae
eats the leaves, flowers and stem of the plant. S. xami may be regarded as a leaf-
miner on the exceptionally thick leaves of Echeueria. The life cycle was partially
described by Ziegler and Escalante (op. cit.). The territorial behavior of S. xami
has been described by Cordero (1986. Defensa territorial en la mariposa Sandia
xami. B. Sc. Thesis. Fac. de Ciencias. UNAM. 75 pp.) and Cordero and Soberon
(submitted) and their oviposition patterns by Soberon et al (op. cit.).
Early Stages
To obtain the eggs in the laboratory, a fertilized female is placed in a cage (fig.
1) built according to Munger, F. and T.T. Harris (1970. Jour. Res. Lep. 8: 169-
176.). A 100 watts tungsten lamp is placed over the insectary a providing a 8:16
LD. One or two pots with Echeueria are placed inside the insectary. The females
lay most eggs on the surface of the plant, although it is not uncommon to find
eggs on the pot. A single female can produce up to 200 eggs in a three week
period (fig. 2). Peak egg-laying takes place in the first week.
Every morning eggs are removed using a fine camel hair brush slightly
dampened with tap water. The eggs are then placed in square (1.5 cm side) cuts
of Echeueria leaves over a filter paper and inside plastic Petri dishes.
Larvae that have emerged from eggs are fed with squares of Echeueria which
are replaced as required. The humidity inside the Petri dishes is kept high by a
drop of water every three days. The Petri dishes are kept at room temperature. A
single, medium-sized leaf (10 cm long) provides food for one larvae to mature.
Larvae are handled with fine camel hair brushes during the first two instars.
27(3-4):259-271, 1988(89)
269
Later instars can be manipulated with coarse brushes or entomological forceps.
When larvae are ready to pupate, they stay still at the edge of the Petri dish
and remain in this state for three to four days.
When adults are ready to emerge (16 to 20 days from beginning of pupation
depending on temperature of the year, x = 16.9 days, S.E. = 1.2), the pupae are
placed in a closed plastic box with enough space for the spreading of the wings to
take place.
Mortality is usually low at every instar, with the exception of the first. New
larvae can be easily damaged by handling. A sumary of several laboratory life
tables is presented in fig. 3.
The adults are maintained in the insectaries on a diet of 10% sucrose on water
with a few drops of commercially available hydrolyzed vegetable proteins
(“Jugo Maggi”, trade mark, similar to soy sauce). Small cubes of plastic foam
moisted with this solution are placed at the end of 10 cm wood sticks attached
vertically to a base of clay. The adults must be placed by hand upon the foam
cubes. This can be easily performed by gently persuading them to attach
themselves to one finger and then placing the butterfly on the foam. At room
temperature in Mexico City (around 20°C) adults can survive for as long as 40
days.
Mating
Inducing butterflies to copulate in laboratory conditions is seldom an easy
task. Hand-pairing has been successful for large butterflies (Clarke, C.A. and
P.M. Sheppard. 1956. Jour. Lep. Soc. 10: 47-53.), but the Lycaenids are more
difficult to hand-pair because the genital armature is more deeply hidden than
in other families (Clarke and Sheppard, op. cit.). We tried the hand-pairing
methods, but none was successful. We have developed two techniques, described
as follows.
1) We placed pairs of laboratory-raised butterflies in portable cages made of
green net cloth and wire (fig. 4). These are then hung outside the laboratory, in
direct sunlight. Matings occur within the hour. At first we used both wild-
caught and laboratory specimens, but wild males always refused to mate.
Seventeen attempts, during cloudless weather, using laboratory reared butter-
flies yielded 15 successful matings. The two failures were apparently due to
female refusal because of unknown causes. This method works quite well, but it
relies on availability of sunlight and, perhaps, on a good ventilation of the cages
(R. Mattoni, personal communication).
2) It is also possible to obtain matings with males in the wild. Many
territories are located in conspicuous places that are usually occupied by a male
(Cordero, op. cit/, Cordero and Sberon, op. cit.). A laboratory female, one or two
days old, is placed in a small card box with the lid attached to a string. The box is
fixed to the end of a 1.5 m wood pole. The box is then placed as close as possible to
the perching male and the lid opened by pulling the string. When the female
emerges, a mating flight usually ensues, with a high probability of a successful
pairing. In our area the vegetation includes Buddleia trees, which can be 4 or 5
meters tall. If the mating takes place on a tree, recapturing the couple may be
impossible, but when the mating occurs in an accessible place, recapture after a
period of 1 hour has always yielded a fertilized female. We have released 16
females and recovered 7 fertilized females.
Although this method requires the localization of an occupied territory and is
less reliable than the first, it can be used to mantain heterozygocity.
270
J. Res. Lepid.
5 5 cm
Fig. 1 Insectary for the maintenance
and oviposition of adult Sandia
xami, built according to Monger
and Harris (1970).
Fig. 2 Mean oviposition per day for
female Sandia xami, in labora-
tory conditions with 8:16 ID
light. The eggs were removed
every morning. Mean ± stand-
ard error. N = 7.
Fig. 3 Mean survivorship curve of
seven laboratory life tables.
Bars are standard errors. In
parenthesis mean and stand-
ard error of duration of stage
in days. Eggs (6.89 ± 0.07); LI
= first instar (5.36 ± 0.06); L2
= second instar (4.71 ± 0,07);
L3 = third instar (5.40 ± 0.10);
L4 = fourth instar (6.72 ±
0.13); Pp = prepupal larvae
(3.70 ± 0.04); P - pupa (18.39
± 0.25); A - adult (31.34 ±
1.18).
2 6 cm
h~ — — I
Fig. 4 Mating portable cage.
27(3-4):259-271, 1988(89)
271
Acknowledgements. We are thankful to Dr. Jorge Llorente, of the Faculty of
Sciences of the UNAM, and Dr. Kurt Johnson, of the American Museum of
Natural History for their suggestion of the first method for inducing matings
and Dr. Mattoni for his comments on this manuscript. Paulina Parlange, Betty
Benrey, Gerardo Berges and Carlos Cordero were very helpful at different
stages of this work.
Gabriela Jimenez C. & Jorge Soberon M., Centro de Ecologia, UNAM, Apdo.
Postal 70—275, Ciudad U nicer sitaria, Mexico 04510 DE., MEXICO
THE END OF NATURE: 1989. Bill McKibben. Random House, N.Y. 230 pp.
$19.95
Armageddon 1: Nature O.
The End of Nature, a landmark book for this century on the philosophy of the
relationship of man to the environment, was not written by a scientist, but by a
reporter. To the academic clique this lack of credentials may be looked upon
with suspicion, but the clarity of thinking, mastery of fact, cool objectivity and
charm of writing are both very impressive and very moving.
I don’t believe any of our members, who are almost universally in regular
touch with nature, will fail to grasp or disagree with the central thpsis of the
book: nature has come to an end. Nature here is that idea describing the set of
interactions among wild organisms that we think of as the planetary eco-
system. In the meantime we all go blithely consuming, travelling, and making
investment decisions like life as we know it will all go on forever. In the mean-
time the almost certainly entrained global warming trend is signalling continu-
ing disintegration of the environment with a foreseeable end to the lifestyles
we have grown to accept. Deductions from the impact of human resource
depletion is nothing new, of course, but what McKibben shows is that the wild
nature in which we evolved is no more. We now live in a man-made world.
The destruction of nature is not only irreversible, but will in all likelihood be
compounded by the “fixes” technology has and will generate. This pessimistic
conclusion will probably not be accepted in the popular weltanschaung. The
laws of nature, as thermodynamics and relativity, have not been repealed,
many forests are still green, there are masses of moth species in some tropical
places and there are a few aborigines around. But this is a managed home. My
job with “restoration and management” of endangered species at the El Segundo
sand dunes focuses on the absurdity. This tad of nature only now exists at our
pleasure. The catena is gone.
You cannot fail to read this book. It is not strident or hysterical. There is no
preaching or demands for change in life style. It is reflective, disturbing and
very topical.
R. H . T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210, USA
Journal of Research on the Lepidoptera
27(3-4):272-276, 1988(89)
PORTRAITS OF SOUTH AUSTRALIAN GEOMETRID MOTHS. McFarland,
Noel. 1988. 400 pp. + inserts in pocket. Pufol. by author, P. O. Box 1404, Sierra
Vista, AZ 85636, USA. Price postpaid $80.00 USA, $85.00 outside USA.
Softcover.
In the time of the closing of the american mind, along comes McFarland’s
Portraits as a refreshing throwback to a gentler kinder era. The price is a bit
steep for a black and white, typewriter font softcover, but I recommend that
you at least strongly urge your library to buy the work as a book you cannot
live without. However, this should be done quickly, as I understand the limited
500 copies are selling well. Rationally, any success for so obscure a focus
(geometrids of South Australia) is surprising, if not miraculous. The answer
lies partly in the superb quality of the graphics. According to information the
prints were processed with a 300 mesh screen. The results are the highest
quality half tones around. The lOVs x 13% size delivers the figures as portrait
scale, and there are about 1500 illustrations. The cover weight is no greater
than this Journal, however, which makes the book awkward to handle, es-
pecially if one reads same in a lounge chair. The typewriter typeface of the text
suggests a transitory presentation, but on refection may not be inappropriate
for a work of this sort, which is more diary than scientific paper. That is its
charm and value.
The natural history writing style is strictly descriptive, but is straightfor-
ward and clear. The exquisite detail and documentation provides minimal
quantitative information, yet of its kind, every angle is covered. Further, the
subject is pursued with a passion not only of the subject matter, but for the
precision of the topic discussed. Casual asides provide an empathetic feel for
working conditions: photographic equipment deluging Mrs. Henley’s dining
room table while chowing down on Andrew’s fish and chips. The biology of the
geometrids is completely covered for 72 species, each treated in its own
“chapter” with morphology and behavior data of adult through egg, including
parasites. Data on breeding technique are thorough to a fault, including
considerations on failure. I would suggest no-one working with geometrids at
any venue could do without this work.
Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210, USA
THE MOTHS AND BUTTERFLIES OF GREAT BRITAIN AND IRELAND.
Volume 7, part 1, Hesperidae-Nymphalidae. The Butterflies. A. M. Emmet and
J. Heath, eds. 1989. Harley Books, Colchester (England). 370 pp. 24 col. pi.
£49.50.
This splendid book unquestionably is the new definitive text on british
butterflies. The work is beautifully produced including painted color illustra-
tions that are absolutely first class. The book may be regarded as an exemplar
of its type. It is divided into three sections. The first is a chapter by A. M.
Emmet on the early lepidopterists which gives the origin of the vernacular
names of all species. Since the historical subject is regional, this part would
27(3-4):272-276, 1988(89)
273
have limited interest to workers outside the U. K. It is noteworthy, however,
that the common names of british butterflies have a long heritage and are
remarkably stable, far more so than the scientific names (see below). The
second, all too brief, chapter by M. G. Morris and J. A. Thomas is titled “Re-
establishment of insect populations, with special reference to butterflies”, but
sets forth the topic of conservation as well. Because of the premier position of
these authors in butterfly conservation, this chapter is must reading for all
lepidopterists. The emphasis of these authors on the subject of re-introductions
is fraught with opinion and tinged with some hostility, yet the points are
righteous and very apt to the circumstances of diminishing biodiversity every-
where. The positions of Morris and Thomas are particularly close to me, for
both trying to re-introducing butterfly species into a highly disturbed habitat
and also attempting to restore a working ecosystem involving plants and other
animals as well. Time and expense wasted in trivial arguments on myopic
positions with bureaucrats brings their discussion home.
The major section of the book is descriptive. It assumes one is a knowledge-
able student of the field. The systematic and distributional data are given in
some detail including highlights of the life histories of all resident native
species. The richest informational material lies in the distribution data, which
is essentially presented in the form of 10 km sq UTM maps, with the time
frame of occurrence categorized into pre-1940, etc. Maps for regular migrants
present quantitative data. Every species ever recorded from Britain is cited,
including some downright bizarre observations. Obvious escapes, as are
apparently becoming more commonplace with the advent of butterfly houses,
are omitted. The “natural” migrants are fascinating for information provided
on potential vagility of different species. Historical evidence is summarized for
species which were or could have been breeding residents in kinder and gentler
times as Aporia crataegi, Lycaena virgaureae and Cyaniris semiargus. Thus
the importance of these species beyond their curiosity status for collectors is
validated. For future generations information on historic distributions of all
organisms is extremely important, particular assuming a hopeful change in
human behavior towards greater rationality in dealing with the conservation
of nature.
In spite of all of its goodness, the book is not perfect. Perhaps its most
damaging feature is the very spotty attribution of work cited, particularly in
the sections dealing with biology. The bibliography is overall disappointing,
based, no doubt, by the editors assuming the trail to more arcane works could
be followed by the conscientious. The major attention to adult appearance sub-
stantially weakens the book, particularly as early stages are so well known for
the british fauna. A key to mature larvae would have been simple and would
not have materially added to length. Chaetotaxy maps of larvae with SEM
micrographs are not unreasonable to expect from a work of this class given the
resources and talent of its some 30 editors, associates, and authors.
In contrast to the stability of common names virtually since the time of
Linneaus, the scientific name picture is disturbing. The use of Eurodryas is
especially inappropriate, standing in sharp contrast to conservatism in the use
of Argynnis and Lycaena . Although the nomenclature topic has been written
into near banality, the strongest argument is for conservatism in a world
where the primary objective of taxonomy is communication and stability and
not the needs of specialists. A last criticism is the inaccessibility of informa-
274
J. Res. Lepid.
tion. The recent guide by Thomas, Butterflies of the British Isles, employs a
clever bar graph showing the phenology of life cycles. Thomas’ presentation,
combined with tables of foodplants, nectar sources, vagility, etc. in a few pages
would impart information at a glance that otherwise requires lengthy rooting
through the text. In spite of a few warts, lepidopterists everywhere can look
forward to completion of the whole series as a landmark.
R. H. T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210, USA.
PRIMITIVE GHOST MOTHS. 1989. E. S. Nielsen and N. Kristensen. Mono-
graphs on Australian Lepidoptera: Volume 1. 206 pp. + xii. CSIRO Publica-
tions. East Melbourne. $A60. Hardbound.
Here is a work of real systematic science in the event you are looking for a
role model. Although de facto interest in Hepialids may not qualify them as a
popular group, they are of great importance to understanding the phylogeny of
Lepidoptera. This book only treats one genus, Fraus Walker, which includes
only 25 species. Of these, 17 are newly described.
The general importance of the work, however, lies in the 116 pages of back-
ground material that prefaces the remainder of the book-the taxonomic re-
vision proper. This background material includes a truly extraordinarily
detailed morphology of all stages of the moths, but emphasizing the Adult.
Included is a virtual atlas of both integument and internal anatomy illustrated
by supurb SEM photographs. One cannot praise the quality of the illustrative
material too highly, which appears in a total of 435 figures. A comprehensive
biology review of 5 pages follows the 98 page morphology section, then 7 pages
on diversity and distribution, and 4 on phylogeny. The latter includes a dis-
cussion on the overall classification of the Hepialidae and relationship of the
family to other Lepidoptera. There is a cladogram of relationships, based on
the thorough information presented herein.
The book is well produced and bound. It is obviously not a work for the
general audience, but at least every institution involved with the systematics
of Lepidoptera has the responsibility of acquiring this book. Every lepidopter-
ist with concerns in morphology and/or primitive moths will find it invalue-
able. Note well the very reasonable price for a work of such high intellectual
and physical standards.
R. H. T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210, USA.
THE BUTTERFLIES OF HISPANIOLA, 1989, A. Schwartz, University of
Florida Press, Gainesville, pp. 580.
Butterfly books that are based on an author’s firsthand field experience
occupy a special place on the bookshelf. Through this type of book one may
27(3-4):272-276, 1988(89)
275
gain insight into how the butterflies of some exotic land look and act, and what
they do for a living. In a summary of over ten years worth of field trips and
museum work by himself and a score of field assistants, Albert Schwartz
provides us with a new, hardcover book — The Butterflies of Hispaniola.
For the hard bitten types who buy butterfly literature this book may prove
useful. Between the covers of The Butterflies of Hispaniola lie numerous flower
visitation records, and a wealth of no nonsense locality, temperature and
seasonality data on 196 species of butterflies. A colleague with experience
assures me that to find a species of Hispaniolan butterfly, just use Schwartz’s
data — if the habitat hasn’t been destroyed, the butterfly will be there. The
taxonomy used in the book is a composite of the families employed by Riley
(1975) and the generic names of Miller & Brown (1981), but no classification
showing relationships above the species level is used. Nonetheless, inters-
persed into many of the species accounts are notes that titillate the “I want to
see this species in the field” juices of somebody like myself, especially since
many of the butterflies are endemic to Hispaniola and of biogeographical
interest. The discussion provides an ecological characterization of the entire
butterfly fauna, and the genus Calisto is used as a model sounding board for
the author’s ideas on the evolutionary history of the entire area treated. My
only complaint about the discussion is that I had trouble with some of the
author’s ecological terms (e.g., quasi-cloud forest, pseudoforest, p. 504; euryx-
enophiles, stenoxerophiles, eurymesophilic, stenomesophilic p. 505).
The person who wants an aesthetically pleasing reference on general but-
terfly biology, or who wants the book simply to identify butterflies of His-
paniola may find this book disappointing. In a modern treatment one expects
to see illustrations of the butterflies for convenient identification. Instead of
identification plates there is a decidedly onerous key; 24 pages in English, and
then again in 24 pages in Spanish. The key is brimming with couplets like:
“15. UP pale gray with darker gray to blackish markings. . . P. oileus; 15'. Not
so. . .16; 16. UP orange. . .17; 17'. Not so. . .23 (p. 538). Regardless of what
keys that have crossed my field of vision, I am always mystified by what “Not
so” or similar statements mean. Perhaps because in my experience just about
everything falls into these categories. The stoic who enjoys wrestling with the
mysteries of keys may grin at my admission of sophomoric ineptitude with this
key. However, nowhere in the book are we told how to separate the butterflies
into families or genera — one must wade through every single couplet hunting
a name for the specimen in hand. In short, I found the only means of
identification in this book to be undiluted chloroform in print.
Surely Hispaniolan butterflies are more than a chunks of dead matter with
vaguely described color patterns. I think the 15 black and white photographs
of habitats (which are poor) should have been replaced with illustrations of
butterflies. With a minimum of 13 specimens per plate, the keys could be
eliminated and replaced with a line in the text on how to tell similar species
apart, and the reader could easily identify the butterflies — even the 25 species
of Calisto described by Schwartz and colleagues that are not illustrated in
Riley (1975) or Brown & Heineman (1972). Perhaps the author and the
publisher might wish to consider adding a set of plates to be sold with the book.
The lack of information on early stages and general references to butterfly
natural history was disheartening. After acknowledging how few life histories
of Hispaniolan butterflies are known, Schwartz states, “But intensive research
276
J. Res. Lepid.
on caterpillars is not for visiting biologists; such work must be carried out by
resident scientists (p. 2)”. However, no attempt is made to help the reader into
the literature, or even give the known hostplants in the species accounts. For a
reference work I am puzzled why no reference to Vane- Wright & Ackery
(1984), Ackery & Vane- Wright (1984), Scott (1986) is made in the literature
cited — literature of decided importance to the butterfly fauna of Hispaniola.
To make use of this book, copies of Brown & Heineman (1972) and the out of
print Riley (1975) should be kept near at hand. This will allow the reader to
consult the plates and broader information content provided in the latter
works. To potential buyers I recommend going to the local library and
examining a copy of Butterflies of Hispaniola firsthand, then compare it to
Riley’s (1975) Butterflies of the West Indies , or Brown & Heineman’s (1972)
Butterflies of Jamaica. Dollars is not much money for a hardcover book, but I
cannot honestly recommend this book to anyone except those who are already
experts on Hispaniolan butterflies.
Literature Cited
ACKERY, P. R. & R. I. VANE-WRIGHT. 1984. Milkweed butterflies: their cladistics
and biology. London: British Museum (Nat. Hist.), Entomology.
BROWN, F. M. & B. HEINEMAN. 1972. Jamaica and its Butterflies. E. W. Classey,
London.
MILLER, L. D. & F. M. BROWN. 1981. A catalogue/checklist of the butterflies of
America north of Mexico. Lep. Soc. Mem. 2: 1-280.
SCOTT, J. A. 1986. The Butterflies of North America. A natural history field
guide. Stanford Univ. Press, Stanford.
VANE-WRIGHT, R. I. & P. R. ACKERY (EDS.) 1984. The Biology of Butterflies. Symp.
Roy. Ent. Soc. 11: 1-429.
P. J. DeVries, Dept of Zoology, University of Texas, Austin, Texas 78712
I
■
'
INSTRUCTIONS TO AUTHORS
Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, with wide margins. Number all pages consecutively and put
author’s name at top right corner of each page. Underline all words where italics are
intended. Footnotes, although discouraged, must be typed on a separate sheet. Do not
hyphenate words at the right margin. All measurements must be metric. Metric altitudes
and distances should include imperial equivalents in parenthesis. Time must be cited on a
24-hour basis, standard time. Abbreviations must follow common usage. Dates should be
cited as example: 4. IV. 1979 (day-arabic numeral; month-Roman numeral; year-arabic
numeral). Numerals must be used before measurements (5mm) or otherwise up to number
ten e.g. (nine butterflies, 12 moths).
Title Page: All papers must have the title, author’s name, author’s address, and any titular
reference and institutional approval reference, all on a separate title page. A family
citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.
Abstracts and Short Papers: All papers exceeding two typed pages must be accompanied
by an abstract of no more than 300 words. An additional summary is not required.
Name Citations and Systematic Works: The first mention of any organism should include
the full scientific name with unabbreviated author and year of description. New
descriptions should conform to the format: description of male and/or female, type data,
diagnosis, distribution, discussion. There must be conformity to the current International
Code of Zoological Nomenclature. We strongly urge deposition of types in major museums,
all type depositories must be cited.
References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbreviations must conform to the World List of
Scientific Periodicals. Do not underline periodicals. If four or less references are cited,
please cite in body of text not in Literature Cited. Journals and serials not listed in the
World List are to be abbreviated according to the Serial Publications on the British
Museum (NH), 3rd edition (1980) or given in full.
Tables: Tables should be minimized. Where used, they should be formulated to a size
which will reduce to 11 x 19 cm (or 4V2 x 7V2 inches). Each table should be prepared as a
line drawing or typed with heading and explanation on top and footnotes below. Number
with Arabic numerals. Both horizontal and vertical rules may be indicated. Complex tables
may be reproduced from typescript.
Illustrations: Color can be submitted as either a transparency or print, the quality of
which is critical. Black and white photographs should be submitted on glossy paper, and, as
with line drawings, must be mounted on stiff white cardboard. Authors must plan on
illustrations for reduction to page size. Allowance should be made for legends beneath,
unless many consecutive pages are used. Drawings should be in India ink at least twice the
final size. Include a metric scale or calculate and state the actual magnification of each
illustration as printed. Each figure should be cited and explained as such. Each illustration
should be identified as to author and title on the back, and should indicate whether the
illustration be returned, which will be at the authors expense.
Legends should be separately typed on pages entitled “Explanation of Figures”. Number
legends consecutively with separate paragraph for each page of illustrations. Do not attach
to illustration. Retain original illustrations until paper finally accepted.
Review: All papers will be read by the editor) s) & submitted for formal review to two
referees.
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 27 Number 3-4 Winter 1988 (1989)
IN THIS ISSUE
Date of Publication: 10 December, 1989
Development of the Wing Margin in Precis Coenia 151
(Lepidoptera: Nymphalidae)
C.E. Dohrmann & H.F. Nijhout
The Morpho-Species Concept of Euphyes dion with the 160
Description of a New Species (Hesperiidae)
John A. Shuey
The Euphilotes battoides complex: recognition of a species and 173
description of a new subspecies
Rudolf H.T. Mattoni
Genetic experiments with a calverleyi-like mutation isolated 186
from Papilio bairdi oregonius (Papilionidae)
David V. McCorkle & Paul C. Hammond
The Life History of Automeris zephyria (Saturniidae) 192
Paul M. Tuskes & Michael J. Smith
Three new species of Paradirphia (Saturniidae: Hemileucinae) 197
from Mexico and Central America with notes on the
immature stages
Claude Lemaire & Kirby L. Wolfe
A list of the Butterflies and Skippers of Mount Revelstoke and 213
Glacier National Parks, British Columbia, Canada
(Lepidoptera)
David L. Threatful
Hybridization of the Mexican tiger swallowtail, Papilio 222
alexiares garcia (Lepidoptera: Papilionidae) with other
P. glaucus group species and survival of pure and hybrid
larvae on potential host plants
J. Mark Scriber, Mark H. Evans & Robert
C. Lederhouse
The Butterflies of Isla de Cedros, Baja California Norte, 233
Mexico
John W. Brown & David K. Faulkner
Opinion: Parallelism and Phylogenetic Trees 257
James A. Scott
Notes 259
Book Reviews 272
Cover Illustration: Collage of scanning electron micrographs of
developing wing cross section of Precis coenia by Dohrmann and
Nijhout, pages 151-159
27 VOLUME INDEX
[1962 - 1988(89)]
Journal of Research on the Lepidoptera
compiled by:
Greg A. Kareofelas and Carol W. Witham
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
ISSN 0022 4324
Published By:
Founder:
Editorial Staff:
Associate Editors:
The Lepidoptera Research Foundation, Inc.
9620 Heather Road
Beverly Hills, California 90210
(213) 274-1052
William Hovanitz
Rudolf H. T. Mattoni, Editor
Scott E. Miller, Assistant Editor
Emilio Balletto, Italy
Henri Descimon, France
Philip DeVries, U.S.A.
Thomas Emmel, U.S.A.
Lawrence Gall, U.S.A.
Hansjuerg Geiger, Switzerland
Otakar Kudrna, Germany
Arthur Shapiro, U.S.A.
Atuhiro Sibatani, Japan
Karel Spitzer, Czechoslovakia
Manuscripts and Notices Material may be sent to the Editor at:
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STATEMENT OF OWNERSHIP AND MANAGEMENT
THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a year by THE
LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of publication and the general business
office are located at 9620 Heather Road, Beverly Hills, California 90210. The publisher is THE
LEPIDOPTERA RESEARCH FOUNDATION, INC. The Editor is R. H. T. Mattoni at the above address.
The Secretary-Treasurer is Leona Mattoni at the general business office. All matters pertaining to
membership, dues, and subscriptions should be addressed to her, including inquiry concerning mailing,
missing issues, and change of address. The owner is THE LEPIDOPTERA RESEARCH FOUNDA-
TION, INC., a non-profit organization incorporated under the laws of the State of California in 1965. The
President is R. H. T. Mattoni, the Vice President is John Emmel, the Secretary-Treasurer is Leona
Mattoni. The Board of Directors is comprised of Barbara Jean Hovanitz, Leona Mattoni, and R. H. T.
Mattoni. There are no bond holders, mortgages, or other security holders.
Printed by Art Printing Works Sdn. Bhd., 29 Jalan Riong, 59100 Kuala Lumpur, Malaysia.
27 VOLUME INDEX
[1962 - 1988(89)]
Journal of Research on the Lepidoptera
compiled by:
Greg A. Kareofelas and Carol W. Witham
27 VOLUME INDEX
[1962 - 1988(89)]
Journal of Research on the Lepidoptera
CONTENTS
About the Index p. iii
Electronic Copies Available p. iv
Publication Dates p. iv
Author Index p. I
Family/Genus Index p. 39
Geopolitical Index p. 66
Africa p. 66
Asia (north of Himalayas) p. 66
Caribbean Islands p. 66
Central America & Mexico p. 67
Europe p. 68
Indo-Australia (south of Himalayas) p. 69
North America (north of Mexico) p. 69
South America p. 74
Miscellaneous Subject Index p. 76
ii
ABOUT THE INDEX
This 27 Volume Index to the Journal of Research on the Lepidoptera is organized
into four parts: author index, family/genus index, geopolitical index, and
miscellaneous subject index. Each part is organized to provide the maximum
usefulness within the limited space considerations. A description of each index,
with its contents and possible uses, follows:
Author Index
This is a complete index to the Journal of Research on the Lepidoptera. All
articles, book reviews, notes and opinions published in volumes 1-27 are
included. The citations are alphabetical by first author with cross-
referencing for other authors.
Family/Genus Index
Whenever possible, each published article has been indexed by the taxa
discussed. The citations are alphabetical by family/genus. Note that the
nomenclature conforms to that in use at the time of article’s publication;
some cross-referencing, within families, may be required to find all articles
pertinent to a given taxa. This is not a complete index as many articles do
not contain a specific reference to species, genus or family. After locating
the articles of interest in this index, the user should also refer to the author
index for other possible publications.
Geopolitical Index
Regional studies (i.e. checklists) and newly described taxa are indexed by
geographical/geopolitical region. The citations are grouped by continent
and/or geographic region and then by country. Within the United States,
the articles are further indexed by state. Again, this is not a complete index
and should be used in conjunction with the author index.
Miscellaneous Subject Index
This index includes many articles of general interest which would not be
found in the taxa or regional indices. Articles included are bibliographies,
book reviews and opinions as well as topics of special interest such as
aberrants and gynandromorphs as well as genetics and general laboratory
techniques. All articles describing new taxa are also included. The subjects
have been grouped into rather broad headings to eliminate cross-referencing.
Only a small portion of the articles are included in this index. Again, we
suggest that you refer to the author index for additional articles of interest.
TECHNICAL INFORMATION
Data management for this index was performed on an IBM PC-XT in DBase III+
(Ashton-Tate). Final camera-ready copy was prepared in Word Perfect 5.0
(Word Perfect Corp.) and printed on a Laserjet IIP (Hewlett-Packard).
in
ELECTRONIC COPIES AVAILABLE
This index is available in electronic form for IBM compatible computers with
DBase III+, or IV. The database file contains author(s), year, title, volume,
number and pages. The file is suitable for preparation of bibliographies and
can be searched using wildcard queries, etc. Diskette copies (please specify 5.25
inch or 3.5 inch) are available for $10.00 from: The Lepidoptera Research
Foundation, Inc., The Journal of Research on the Lepidoptera, 9620 Heather
Road, Beverly Hills, California 90210.
VOLUME DATES
References are to volume, number and page. Date of publication has been
omitted to conserve space. There are two multi-number volumes [26(1-4) and 27(3-
4)] and one supplement [i7(s)]. A list of volumes with seasonal dates follows.
Year of publication is listed, in ( ), if different than seasonal date.
1(1) August, 1962
1(2) January, 1963
1(3) March, 1963
1(4) May, 1963
2(1) July, 1963
2(2) September, 1963
2(3) November, 1963
2(4) December, 1963
3(1) March, 1964
3(2) June, 1964
3(3) September, 1964
3(4) December, 1964
4(1) March, 1965
4(2) June, 1965
4(3) September, 1965
4(4) December, 1965
5(1) March, 1966
5(2) June, 1966
5(3) September, 1966
5(4) December, 1966
6(1) March, 1967
6(2) June, 1967
6(3) September, 1967
6(4) December, 1967
7(1) March, 1968
7(2) June, 1968
7(3) September, 1968
7(4) December, 1968
8(1) March, 1969
8(2) June, 1969
8(3) September, 1969
8(4) December, 1969
9(1) March, 1970
9(2) June, 1970
9(3) September, 1970
9(4) December, 1970
10(1) March, 1971
10(2) June, 1971(72)
10(3) September, 1971(73)
10(4) December, 1971(73)
11(1) March, 1972(73)
11(2) June, 1972(73)
11(3) September, 1972(73)
11(4) December, 1972(73)
12(1) March, 1973
12(2) June, 1973
12(3) September, 1973(74)
12(4) December, 1973(74)
13(1) March, 1974
13(2) June, 1974
13(3) September, 1974
13(4) December, 1974
14(1) March, 1975
14(2) May, 1975
14(3) September, 1975
14(4) December, 1975
15(1) March, 1976
15(2) June, 1976
15(3) September, 1976
15(4) December 1976
16(1) March, 1977
16(2) June, 1977
16(3) September, 1977
16(4) December, 1977
17(1) March, 1978(79)
17(2) Summer, 1978(80)
17(3) Autumn, 1978(80)
17(4) Winter, 1978(80)
1 7(S) Supplement, 1978(79)
18(1) Spring, 1979(81)
18(2) Summer, 1979(81)
18(3) Autumn, 1979(81)
18(4) Winter, 1979(81)
19(1) Spring, 1980(81)
19(2) Summer, 1980(81)
19(3) Autumn, 1980(81)
19(4) Winter, 1980(81)
20(1) Spring, 1981(82)
20(2) Summer, 1981(82)
20(3) Autumn, 1981(83)
20(4) Winter, 1981(83)
21(1) Spring, 1982(83)
21(2) Summer, 1982(83)
21(3) Autumn, 1982(83)
21(4) Winter, 1982(83)
22(1) Spring, 1983
22(2) Summer, 1983
22(3) Autumn, 1983(84)
22(4) Winter, 1983(84)
23(1) Spring, 1984
23(2) Summer, 1984
23(3) Autumn, 1984
23(4) Winter, 1984(85)
24(1) Spring, 1985
24(2) Summer, 1985
24(3) Autumn, 1985(86)
24(4) Winter, 1985(86)
25(1) Spring, 1986
25(2) Summer, 1986(87)
25(3) Autumn, 1986(87)
25(4) Winter, 1986(88)
26(1-4) Autumn, 1987(88)
27(1) Spring, 1988(89)
27(2) Summer, 1988(89)
27(3-4) Winter, 1988(89)
IV
AUTHOR INDEX
[Volumes 1-27, 1962-1988(89)]
Journal of Research on the Lepidoptera
Adams, Phillip A.
A rubber stamp method for producing specimen labels 2(3):225-228
A rapid method for producing insect labels 1 5(3): 1 69- 172
Adams, Phillip A. and J. E. Heath
An evaporative cooling mechanism in Pholus cichemon
(Sphingidae) 3(2):69-72
Ae, Shigeru
Hybrids between Papilio memnon and Papilio protenor 3(l):55-62
Agarwal, Anil Kumar
Digestive enzymes of sugarcane pink borer, Sesamia inferens Walker
(Noctuidae) 1 5(3): 1 53- 1 62
Digestive enzymes of a sugarcane borer, Chilotraea infuscatellus
Snell 1 7(3): 1 80- 187
Aiello, Annette and Keith S. Brown, Jr.
Mimicry by illusion in a sexually dimorphic, day-flying moth, Dysschema
jansonis (Lepidoptera: Arctiidae: Pericopinae) 26(1-4):173-176
Alcock, John
The mating system of three territorial butterflies in Costa
Rica 26( l-4):89-97
Alcock, John and Darryl Gwynne
The mating system of Vanessa kershawi : males defend landmark territories
as mate encounter sites 26(1-4):! 16-124
Andersen, William A. (see Simmons, Robert S. and William A. Andersen)
(see also Simmons, Robert S., William A. Andersen and Philip J. Kean)
Anthony, George Scott
Population structure of Oeneis melissa semidea (Satyridae) from the
Presidential Range, New Hampshire 7(3): 133-1 48
Araujo, Aldo M. (see Menna-Barreto, Ymara and Aldo M. Araujo)
Archer, D. M. (see Lees, E. and D. M. Archer)
Arnold, Richard A.
Conservation and management of the endangered Smith’s Blue Butterfly,
Euphilotes enoptes smithi (Lepidoptera: Lycaenidae) 22(2): 135-153
Austin, Anna T. (see Austin, George T. and Anna T. Austin)
Austin, George T.
Book Review - Miller and Brown: A Catalogue/Checklist of the Butterflies
North of Mexico 19(4):241-243
A new subspecies of Lycaena editha (Mead) (Lycaenidae) from
Nevada 23(l):83-88
Lowland riparian butterflies of the Great Basin and associated
areas 24(2):1 17-131
Apodemia palmerii (Lycaenidae: Riodininae): Misapplication of names, two
new subspecies and a new allied species 26(1-4):125-140
Book Review - Tilden and Smith: A Field Guide to Western
Butterflies 26(l-4):278-283
1
2
AUTHOR INDEX
J. Res. Lepid.
Austin, George T. (continued)
Notes: A replacement name for Lycaena editha nevadensis Austin
(Lycaenidae) 27(3):266
Austin, George T. and Anna T. Austin
Butterflies of Clark County, Nevada 1 9( 1 ): 1 -63
Austin, George T. and Douglas Mullins
A new Limenitis weidemeyerii W. H. Edwards from southeastern Arizona
(Nymphalidae) 22(4):225-228
Bacheler, Jack S. and Thomas C . Emmel
Genetic control of maculation and hindwing color in Apantesis
phalerata 13(1 ):49-56
The chromosomes of Apantesis phalerata , A. radians , and their hybrid in
Florida populations (Arctiidae) 1 3(3): 1 62- 168
Bailowitz, Richard A.
Systematics of Ascia ( Ganyra ) (Pieridae) populations in the Sonoran
Desert 26( 1 -4):73-8 1
Census of the butterflies of the National Audubon Society’s
Appleton-Whittell Research Ranch, Elgin, Arizona 27(2): 1 20- 128
Balletto, Emilio and Otakar Kudrna
An annotated catalogue of the Burnets and Foresters (Lepidoptera:
Zygaenidae) named by Roger Verity 24(3):226-249
(see also Kudrna, Otakar and Emilio Balletto)
Ballmer, Gregory R. and Gordon F. Pratt
A survey of the last instar larvae of the Lycaenidae (Lepidoptera) of
California 27(1):1-81
(see also Pratt, Gordon F. and Greg R. Ballmer)
Baragano, J. R. (see De Viedma, M. G., J. R. Baragano, A. Notario, M.
Rodero and C. Iglesias)
Barker, John F.
Notes: Sex characters of the pupae of the banded moth Cochylis hospes
Wilsingham (Lepidoptera: Cochylidae) 27(3):267-268
Barton, H. E. (see Selman, Charles L. and H. E. Barton)
Bauer, William R. (see Buckett, John S. and William R. Bauer)
Baughman, John F. (see Murphy, Dennis D. and John F. Baughman)
Beard, P. (see Urquhart, F. A., P. Beard and R. Brownlee)
Becker, Vitor O. and Scott E. Miller
The identity of Sphinx brunnus Cramer and the taxonomic position of
Acharia Huebner (Lepidoptera: Limacodidae) 26(1 -4):2 19-224
Biggs, James D. (see Shapiro, Arthur M. and James D. Biggs)
Bishop, J. A. (see Sheppard, P. M. and J. A. Bishop)
Bitzer, Royce J. and Kenneth C. Shaw
Territorial behavior of the red admiral, Vanessa atalanta (L.) (Lepidoptera:
Nymphalidae) 18(1 ):36-49
Bowden, S. R.
On Pieris ( Artogeia ) marginalis macdunnoughii Remington
(Pieridae) 26(l-4):82-88
Bowman, D, E.
Notes: A range extension and dark phenotype of Hemileuca
chinatiensis 24(1 ):85
Bradley, J. D., D. S. Fletcher and P. E. S. Whalley
Kloet and Hincks’ Check list of British Lepidoptera Insects (Lepidoptera)
Edn. 2. A reply to criticisms 13(4):265-266
J. Res. Lepid.
AUTHOR INDEX
3
Breedlove, D. E. (see Emmel, John F., Oakley Shields and D. E. Breedlove)
(see also Shields, Oakley, John F. Emmel and D. E. Breedlove)
Brock, J. P.
Opinion: Reply to Scott’s criticism 26(l-4):240-247
Brower, A. E.
Variations of Parasemia pcirthenos 1 1(3): 183- 186
Brown, F. Martin
W. H. Edwards’ life histories of North American Coenonympha. 3(2): 121-128
Comments on the genus Cercyonis Scudder, with figures of types
(Satyridae) 4(2):131-148
Euphydryas editha gunnisonensis , a new subspecies from western
Colorado 9(l):21-23
Book Review - Beattie: Rhopalocera directory 1 5(3): 173-175
(see also Miller, Lee D. and F. Martin Brown)
Brown, John W.
Observations on Phoebis sennae (Pieridae) 17(3)168-169
Notes on the life history and Baja California distribution of
Chlorostrymon simaethis sarita (Skinner) (Lepidoptera:
Lycaenidae) 20(4):207-213
A new species of Mitoura Scudder from southern California (Lepidoptera:
Lycaenidae) 21(4):245-254
Notes: Records of Hypaurotis crysalus (Ed) (Lycaenidae) from western
Mexico 27(2): 135
(see also Spade, Paul, Hamilton Tyler and John W. Brown)
Brown, John W. and David K. Faulkner
The butterflies of Isla de Cedros, Baja California Norte,
Mexico 27(3):233-256
Brown, Keith S., Antoni J. Damman and Paul Feeny
Troidine swallowtails (Lepidoptera: Papilionidae) in southeastern Brazil:
natural history and foodplant relationships 1 9(4): 1 99-226
(see also Aiello, Annette and Keith S. Brown, Jr.)
Brown, Richard M.
Larva and habitat of Callophrys fotis bayensis 8(2):49-50
A revision of the North American Comadia (Cossidae) 1 4(4): 1 89-2 1 2
A new genus and species from the southwestern United States (Noctuidae:
Acontiinae) 25(2): 1 36- 1 45
(see also Wells, James F. and Richard M. Brown)
Brownlee, R. (see Urquhart, F. A., P. Beard and R. Brownlee)
Brussard, Peter F. (see Vawter, A. Thomas and Peter F. Brussard)
Buckett, John S.
Collecting of Annaphila spila with notes on the "crimson winged" group of
the genus 2(4):303-304
Revision of the North American genus Behrensia 3(3): 1 29- 1 44
Identity of Heliosea celeris melicleptroides 4(l):79-80
The Noctuid moth Annaphila baueri with notes on its habits. . . 4(3): 185- 189
A reevaluation of Annaphila casta (Noctuidae) 4(3):199-204
Rediscovery of Annaphila casta Hy. Edw. in California
(Noctuidae) 5(l):37-38
The little known moth Euxoa sculptilis (Harvey) in Arizona, with
descriptions, illustrations, and notes on Euxoa violaris (Grote and
Robinson) (Noctuidae) 5(4):255-261
4
AUTHOR INDEX
J. Res. Lepid.
Buckett, John S. (continued)
Discovery of a larval hostplant for Annaphila lithosina with notes on the
species (Noctuidae: Amphipyrinae) 5(4):262-264
Description of a new species of Xylomiges from California. . . . 6(l):23-30
A new species of Feralia 6(1):43-51
A new species of armyworm - genus Faronta 6(4):268-274
Identity of the moth "Stretchia" behrensiana with new synonymy
(Noctuidae) 7(l):57-63
Species in the genera Polia and Euxoa 7(2):8 7-94
Identity of the moth Loxagrotis pampolycala from the southwestern US
and Mexico (Noctuidae) 8(3): 118-1 28
Revision of the Nearctic genus Philtraea Hulst with notes on biology and
description of new species (Geometridae) 9(l):29-64
Identity of the moth Oncocnemis semicollaris J. B. Smith 10(3):248-254
A new species of Nephelodes Guenee for the Great Basin. . . . 1 1(4):260-268
(see also Opler, Paul A. and John S. Buckett)
Buckett, John S. and William R. Bauer
Review of the depicta group of the genus Annaphila 3(2):95- 101
Petaluma , a new genus 3(3): 193- 196
A new species of Oncocnemis from the western United States (Noctuidae:
Cuculliinae) 5(4):197-208
A new species of Polia Ochsenheimer from California and notes on Polia
discalsis (Grote) (Noctuidae: Hadeninae) 5(4):221-228
Homonymy of the "new genus" Petaluma and proposal of the name
Petalumaria 6(1):52
Buckett, John S. and Ronald H. Leuschner
Rediscovery and redescription of the moth Lithophane vanduzeei
(Barnes) 4(4):281-286
Buckett, John S. and L. P. Lounibos
The little known species Luperina venosa 4(4):227-232
Buckett, John S. and T. A. Sears
Variation in color and maculation in Nemoria pulcherrima from the Sierra
Nevada of California. Lepidoptera: Geometridae 7(2):95-98
Calhoun, John V.
Notes: Aberrant Polyommatinae (Lycaenidae) from Ohio and
Florida 26(l-4):264-266
Callaghan, Curtis J.
Notes on the immature biology of two Myrmecophilous Lycaenidae:
Juditha molpe (Riodininae) and Panthiades bitias
(Lycaeninae) 20(l):36-42
A study of isolating mechanisms among Neotropical butterflies of the
subfamily Riodininae 2 1(3): 1 59- 176
Notes on the biology of Stalachtis susanna (Lycaenidae: Riodininae) with a
discussion of Riodinine larval strategies 24(3):258-263
Notes on the biology of three Riodinine species: Nymphidium lisimon
attenuatum , Phaenochitonia sagaris satnius , and Metacharis ptolomaeus
(Lyceanidae: Riodininae) 27(2): 1 09- 1 14
Callaghan, Curtis J. and Kenneth B. Tidwell
A checklist of Utah butterflies and skippers 1 0(3): 191 -202
Callaghan, Curtis J. and Norman B. Tidwell
Addition to: "A checklist of Utah butterflies and skippers". . 1 1 (3): 1 99-200
Castrovillo, Paul J. (see Schaefer, Paul W. and Paul J. Castrovillo)
J. Res. Lepid.
AUTHOR INDEX
5
Chacon, Patricia and Marta de Hernandez
Immature stages of Odonna passi florae Clark (Lepidoptera: Oecophoridae):
Biology and morphology 20(l):43-45
Chang, Vincent C. S.
Quantitative analysis of certain wing and genitalia characters of Pieris in
western North America 2(2):97-125
(see also Hovanitz, William and Vincent C. S. Chang)
(see also Hovanitz, William, Vincent C. S. Chang and Gerald Honch)
Cheverton, Mark R. (see Thomas, Chris D. and Mark R. Cheverton)
Chiba, Hideyuki
Notes: Description of the hitherto unknown female of Acerbas suttoni
Russell (Hesperiidae) 27(3):260-261
Chiba, Hideyuki and Hiroshi Tsukiyama
Notes: Revisional notes on the genus Satarupa Moore (Lepidoptera:
Hesperiidae) I. New synonym of Satarupa monbeigi
Oberthur 27(2): 1 38- 1 39
Chua, K. E., J. C. E. Riotte and C. Gilmour
Investigation of selected species of the genus Orgyia (Lymantriidae) using
isoelectrofocusing in thin layer polyacrylamide gel 1 5(4):2 1 5-224
Clark, Gowan C. and C. G. C. Dickson
The life history of two species of South African Eurema 4(4):252-257
The life histories of South African Colotis erone, C. tone , C. vesta and
Leptosia alcesta (Pieridae) 6( 1 ):3 1 -42
South African Eurema 8( 1 ): 1 8- 1 9
Clark, J. F. Gates
A new genus and two new species of Oecophoridae from Columbia
(Lepidoptera) 20(l):46-49
Clarke, Sir Cyril A. (see West, David A. and Sir Cyril A. Clarke)
Clarke, Sir Cyril A. and Axel Willig
The use of alpha-ecdysone to break permanent diapause of female hybrids
between Papilio glaucus L. female and Papilio rutulus male. 16(4):245-248
Clench, Harry K.
Callophrys (Lycaenidae) from the northwest 2(2): 151-1 60
A synopsis of the west Indian Lycaenidae, with remarks on their
zoogeography 2(4):247-270
A new species of Riodinidae from Mexico 3(2):73-79
A review of the genus Lasaia (Riodinidae) 1 0(2): 1 49- 1 80
Comstock, John Adams
Notes on the early stages of two California geometrids 1 (3): 1 95-200
Early stages of a southern California Geometrid moth, Drepanulatrix hulsti
hulsti (Dyar). l(4):245-248
Notes on the early stages of Drepanulatrix monicaria (Guenee)
(Geometridae) 2(3):201-203
The eggs and first instar larvae of three California moths. . . . 5(4):2 1 5-2 1 9
An additional food plant record for Papilio thoas autocles R. & J. . 5(4):220
Old Timers 14(2):90-92
Early work on the Megathymidae 14(2):98-99
Comstock, John Adams and Christopher Henne
Studies in life histories of North American Lepidoptera. California
Annaphilas 3(3): 1 7 3- 1 9 1
Studies in the life histories of North American Lepidoptera, California
Annaphila II 5(1): 15-26
6
AUTHOR INDEX
J. Res. Lepid.
Comstock, John Adams and Christopher Henne (continued)
Studies in the life histories of North American Lepidoptera. California
Annaphila III 6(4):257-262
Early stages of Lygomorpha regulus 6(4):275-280
Cooper, William J. (see Miller, Thomas A., William J. Cooper and Jerry W.
Highf ill)
Cornes, Michael A. (see Larsen, Torben B., John Riley and Michael A.
Comes)
Courtney, Steven P.
Notes: Notes on Tomares mauretanicus (Lycaenidae) in
Morocco 21(3):205-206
Oviposition by the mistletoe-feeding Pierid butterfly Mathania leucothea
(Mol.) in Chile 24(3):264-270
Oviposition on peripheral hosts by dispersing Pieris napi (L.)
(Pieridae). 26(l-4):58-63
Coutsis, John G.
Description of the female genitalia of Hipparchia fagi Scopoli, Hipparchia
semele Linnaeus (Satyridae) and their related taxa 22(3): 161 -203
Covell, Jr., Charles V., Irving L. Finkelstein and Abner A. Towers
A new species of Narraga (Geometridae, Ennominae) from Georgia, with
biological notes 23(2): 161-1 68
Crawford, C. S.
Decapitation-initiated oviposition in Crambid moths 3(l):5-8
Primary geo-orientation in sod webworm moths 9(2):65-74
Crowe, Charles R.
The climatological tool in lepidoptera research 4(l):23-36
A possible new hybrid copper 8(2):5 1 -52
Damman, Antoni J. (see Brown, Keith S., Antoni J. Damman and Paul Feeny)
Dash, A. K. and B. K. Nayak
Notes: Effect of refrigeration on hatching of eggs of the tasar silk moth
Antheraea mylitta Drury (Saturniidae) 27(3):263-265
De Benedictis, John A.
On the taxonomic position of Ellabella Busck, with descriptions
of the larva and pupa of E. bayensis (Lepidoptera:
Copromorphidae) 23(l):74-82
The pupa of Lotisma trigonana and some characteristics of the
Copromorphidae (Lepidoptera) 24(2): 1 32- 135
de Jong, Rienk
The biological species concept and the aims of taxonomy. . . . 21(4):226-237
Book Review - Bridges: Lepidoptera: Hesperiidae. Notes on Species -
Group Names 24(4):379-381
de Hernandez, Marta (see Chacon, Patricia and Marta de Hernandez)
de Maeght, James Mast (see Descimon, Henri and James Mast de Maeght)
Descimon, Henri and Jean-Pierre Vesco
A mutant affecting wing pattern in Parnassius apollo (Linne) (Lepidoptera
Papilionidae) 26( 1 -4): 161-172
Descimon, Henri and James Mast de Maeght
Semispecies relationships between Heliconius erato cyrbia Godt. and H.
himera Hew. in southwestern Ecuador 22(4):229-237
De Viedma, M. G., J. R. Baragano, A. Notario, M. Rodero and C. Iglesias
Artificial raising of Lignicolous Lepidoptera 24(4):372-374
J. Res. Lepid.
AUTHOR INDEX
7
DeVries, Philip J.
Observations on the apparent Lek behavior in Costa Rican rainforest
Perrhybris pyrrha Cramer (Pieridae) 17(3): 142- 144
Hostplant records and natural history notes on Costa Rican butterflies
(Papilionidae, Pieridae & Nymphalidae) 24(4):290-333
Stratification of fruit-feeding nymphalid butterflies in a Costa Rican
rainforest 26(l-4):98-108
Book Review - Schwartz: The Butterflies of Hispaniola 27(3):274-276
DeVries, Philip J. and Isidro Chacon Gamboa
A new species of Adelpha (Nymphalidae) from Parque Nacional Braulio
Carrillo, Costa Rica 20(2):123-126
Dickson, C. G. C. (see Clark, Gowan C. and C. G. C. Dickson)
Dimock, Thomas E.
Type locality and habitat - Cynthia annabella 10(4):265-266
Opinion: Patronyms in Rhopaloceran nomenclature 23( 1 ):94- 101
Notes: Six homoeotic Vanessa atalanta rubria (Nymphalidae) 23(2): 176
Notes: Culture maintenance of Vanessa atalanta rubria
(Nymphalidae) 23(3):236-240
Notes: A homoeotic Agraulis vanillae incarnata (Nymphalidae). . . 23(4):332
Dimock, Thomas E. and Rudolf H. T. Mattoni
Hidden genetic variation in Agraulis vanillae incarnata
(Nymphalidae) 25( 1 ): 1 - 1 4
Dohrmann, C. E. and H. F. Nijhout
Development of the wing margin in Precis coenia (Lepidoptera:
Nymphalidae) 27(3): 1 5 1 - 1 59
Donahue, Julian P.
A melanic form of Pieris rapae 6(4):266
Donahue, Julian P. and Claude LeMaire
A new species of Ormiscodes (Dirphiella) from Mexico (Saturniidae:
Hemileucinae) 1 3(2): 123-1 30
Dornfeld, Ernst J.
A field captured scale-deficient mutant of Anthocharis sara. . . . 9(l):25-28
Dos Passos, Cyril F.
The correct name for the subspecies of Limenitis weidemeyerii occurring in
Arizona (Nymphalidae) 1 2( 1 ):2 1 -24
A note on Oeneis jutta harperi , its author and date of publication
(Satyridae) 1 5(4):2 1 1-213
Some little-known U. S. publications on Lepidoptera 1 20(2): 111-1 22
Douwes, Per
An area census method for estimating butterfly population
numbers 1 5(3): 1 46- 1 52
Downey, John C. (see Lawrence, Donald A. and John C. Downey)
Ducros, Patrick (see Feltwell, John and Patrick Ducros)
Eaton, Jr., Theodore H.
Caterpillar versus dinosaur? 1 (2): 1 14-1 16
Ebner, James A. and Clifford D. Ferris
A new subspecies of Colias palaeno (Linnaeus) from Baffin Island, N.W.T.,
Canada (Pieridae) 1 6(3): 1 55- 1 6 1
Ehrlich, Paul R. (see also Murphy, Dennis D. and Paul R. Ehrlich)
Ehrlich, Paul R. and Dennis D. Murphy
Butterfly nomenclature: A critique 20( 1 ): 1 - 1 1
Nomenclature, taxonomy, and evolution 20(4):199-204
8
AUTHOR INDEX
J. Res. Lepid.
■382
152
Ehrlich, Paul R. and Dennis D. Murphy (continued)
Butterflies and biospecies 2 1 (4):2 1 9-225
Book Review: Pyle: The Audubon Society Handbook for Butterfly
Watchers 24(4):381-
Ehrlich, Paul R. and Darryl Wheye
Some observations on spatial distribution in a montane population of
Euphydryas editha 23(2): 143-
Emmel, John F. (see Emmel, Thomas C. and John F. Emmel; and Shields,
Oakley and John F. Emmel)
(see also Shields, Oakley, John F. Emmel and D. E. Breedlove)
Emmel, John F. and Thomas C. Emmel
Larval food-plant records for six western Papilios 1 (3): 191-1 93
Genetic relationships of Papilio indra and Papilio polyxenes. . . 3(3): 157- 158
Emmel, John F. and Oakley Shields
Larval foodplant records for Papilio zelicaon in the western United States
and further evidence for the conspecificity of P. zelicaon and P.
gothica 17(1 ):56-67
The biology of Plebejus ( Icaricia ) shasta in the Western United States
(Lycaenidae) 1 7(2): 129- 1 40
Emmel, John F., Oakley Shields and D. E. Breedlove
Larval foodplant records for North American Rhopalocera. Part
2 9(4):233-242
Emmel, Thomas C.
Colias philodice in Chiapas, Mexico 1(3): 194
New gynandromorph of Colias philodice from Colorado 3(l):63-64
Methods for studying the chromosomes of lepidoptera 7(1 ):23-28
Population biology of the Neotropical Satyrid butterfly, Euptychia hermes.
1. Interpopulation movement, etc 7(3): 153-1 65
Estimation of natural mutation rates for albinism in two species of
Satyrid genus Cercyonis 8(2):65-68
Dispersal in cosmopolitan butterfly species ( Pieris rapae) having open
population structure 1 1 (2):95-98
The butterfly faunas of San Andres and Providencia Islands in the
western Caribbean 14(1 ):49-56
Book Review - D’Abrera: Butterflies of South America 23(2): 171-172
(see also Bacheler, Jack S. and Thomas C. Emmel)
(see also Emmel, John F. and Thomas C. Emmel)
Emmel, Thomas C. and John F. Emmel
Composition and relative abundance in a temperate zone butterfly
fauna.... 1(2):97-
Life history of Satyrium sylvinas dryope 7(2): 123-
Two new subspecies of Euphydryas chalcedona from the Mojave desert of
Southern California 1 1 (3): 141-1 46
A new subspecies of Euphydryas editha from the Channel Islands of
California .' 1 3(2): 1 3 1 - 1 36
Emmel, Thomas C. and Charles F. Leck
Seasonal changes in organization of tropical rain forest butterfly
populations in Panama. . 8(4): 133-1 52
Enns, Wilbur R. (see Heitzman, Roger L. and Wilbur R. Enns)
Ericksen, C. H.
Further evidence of the distribution of some boreal Lepidoptera in the
Sierra Nevada 1(1 ):89-93
108
125
J. Res. Lepid.
AUTHOR INDEX
9
Evans, Mark H. (see Scriber, J. Mark and Mark H. Evans)
(see also Scriber, J. Mark, Mark H. Evans and Robert C. Lederhouse)
Faulkner, David K. (see Brown, John W. and David K. Faulkner)
Federici, Brian A. (see Santiago-Alvarez, Candido and Brian A. Federici)
Feeny, Paul (see Brown, Keith S., Antoni J. Damman and Paul Feeny)
Felt well, John
Migration of Hipparchia semele L 1 5(2):8 3-9 1
The depredations of the large white butterfly ( Pieris brassicae )
(Pieridae) 1 7(4):2 1 8-225
Feltwell, John and Patrick Ducros
Notes: Further migration of Hipparchia semele (L.) in 1976 and
1980 20(1):53
Ferguson, Douglas C.
Response to J. C. E. Riotte’s Review of the Lymantriid
fascicle 17(4):265-267
New species and new nomenclature in the American Acronictinae
(Lepidoptera: Noctuidae) 26( 1 -4):20 1-218
Ferris, Clifford D.
Two new forms of Plebejinae from Wyoming 8(3):9 1 -93
A new subspecies of Euphydryas from Wyoming (Nymphalidae). 9( 1 ): 1 7-20
Polymorphism in two species of Alaskan Boloria 1 1 (4):255-259
Notes on Arctic and Subarctic collecting 13(4):249-264
Book Review: Butterflies of the World by H. L. Lewis 13(4):278-280
A note on Oeneis Melissa (Fabricius) in the western United States
(Satyridae) 1 4(4):2 1 3-2 1 5
A proposed revision of non-Arctic Parnassius phoebus Fabricus in North
America (Papilionidae) 15(1 ): 1 -22
Butterfly collecting in Labrador and Newfoundland 1 5(2): 1 06- 1 08
A note on the subspecies of Parnassius clodius Menetries found in the
Rocky Mountains of the United States (Papilionidae) 15(2):65-74
Biochemical studies of the larval hosts of two species of Lycaena Fabricius
(Lycaenidae) 17(1 ):27-32
Notes: On Colias hecla Lefebvre re a recent paper by Oosting and Parshall
(Lepidoptera: Pieridae) 20(l):52-53
Field study of Phyciodes batesii (Reakirt) and P. tharos (Drury) from a site
in the Black Hills, South Dakota (Lepidoptera: Nymphalidae:
Melitaeinae) 20(4):235-239
Polymorphism in Satyrium calanus (Huebner) from Wyoming and Colorado
(Lepidoptera: Lycaenidae: Theclinae) 2 1(3): 188-1 94
Speyeria atlantis phenotypes in the southern Rocky Mountains
(Lepidoptera: Nymphalidae: Argynninae) 22(2): 101-1 14
A second phenotype of Satyrium calanus (Heubner) from Wyoming
(Lepidoptera: Theclinae) 23(4):297-302
Notes: Field notes on Clossiana improba harryi Ferris (Lepidoptera:
Nymphalidae) 25(l):71-72
Book Review - Kudrna: Butterflies of Europe 25(2): 155
Euphydryas anicia and E. chalcedona in Idaho (Lepidoptera:
Nymphalidae) 26( 1 -4): 1 09- 1 1 5
(see also Ebner, James A. and Clifford D. Ferris)
Ferris, Clifford D. and Michael S. Fisher
Charidryas flavula Barnes and McDunnough (Nymphalidae): A question of
identity 1 6(3): 133-1 40
10
AUTHOR INDEX
J. Res. Lepid.
Ferris, Clifford D. and D. R. Groothuis
A new subspecies of Boloria eunomia from Wyoming 9(4):243-248
Finkelstein, Irving L. (see Covell, Jr., Charles V., Irving L. Finkelstein and
Abner A. Towers)
Fisher, Michael S.
The heathii- white banding aberration in the Strymoninae
(Lycaenidae) 1 5(3): 177-181
(see also Ferris, Clifford D. and Michael S. Fisher)
Flanders, S. E.
Did the caterpillar exterminate the giant reptile? 1(1 ):85-88
Fletcher, D. S. (see Bradley, J. D., D. S. Fletcher and P. E. S. Whalley)
Forbes, Gregory S.
Description and taxonomic implications of an unusual Arizona population
of Apodemia mormo (Riodinidae) 1 8(3):20 1 -207
Forrest, Hugh S. (see Gilbert, Lawrence E., Hugh S. Forrest, Thomas D.
Schultz and Donald J. Harvey)
Fosdick, Michael K.
The Neotropical Nymphalid butterfly, Anartia amalthea 1 1 (2):65-80
Fox, Richard M. ,
Affinities and distribution of Antillean Ithomiidae 2(3): 173-1 84
Forelegs of butterflies I. Introduction: Chemoreception 5( 1 ): 1 - 1 2
Freeman, H. A.
Type localities of the Megathymidae 2(2): 137-141
The effects of pH on the distribution of the Megathymidae 3( 1 ): 1 -4
Larval habits of Agathymus mariae 3(3): 1 45- 1 47
New skipper records for Mexico 5(l):27-28
Remarks on the genus Zera Evans in Mexico with a new
record 5(3):181-184
Speciation in the Agathymus (Megathymidae) 5(4):209-214
The status of some Hesperiidae from Mexico 6(l):59-64
Polyctor polyctor in Mexico 6(3): 195- 196
Freeman, T. N.
New Canadian species of leaf-mining lepidoptera of conifers. 4(3):209-220
A new species of Epinotia Hubner from British Columbia
(Olethreutidae) 5(1):13-14
A new species of Nepticula on bur oak in Ontario (Nepticulidae). 6( 1 ): 1 9-2 1
(see also Mutuura, A. and T. N. Freeman)
Fried lander, Timothy P.
The biology and morphology of the immature stages of Asterocampa idyja
argus (Bates) (Lepidoptera: Nymphalidae) 24(3):209-225
Egg mass design relative to surface-parasitizing parasitoids, with notes on
Asterocampa clyton (Lepidoptera: Nymphalidae) 24(3):250-257
A new squash borer from Mexico (Lepidoptera: Sesiidae). . . . 24(4):277-288
Taxonomy, phylogeny and biogeography of Asterocampa Rober 1916
(Lepidoptera: Nymphalidae: Apaturinae) 25(4):2 1 5-338
Funk, R. S.
Book Review: Annotated check list of the butterflies of Illinois by R. R.
Irwin and J. C. Downey 12(4):243-244
Furuta, Kimito (see Schaefer, Paul W. and Kimito Furuta)
Gage, Edward V.
Some preliminary notes about the immature stages of Eacles oslari
(Citheronidae) 1 5(3): 1 75- 1 76
J. Res. Lepid.
AUTHOR INDEX
11
Gage, Edward V. (continued)
(see also Perkins, Jr., Edwin M. and Edward V. Gage)
Gall, Lawrence F.
Measuring the size of Lepidopteran populations 24(2):97- 1 1 6
Book Review - Vane-Wright and Ackery: The Biology of
Butterflies 25(2): 1 49- 1 55
Book Review - Scott: The Butterflies of North America. A Natural History
and Field Guide 26(l-4):270-275
Gamboa, Isidro Chacon (see DeVries, Philip J. and Isidro Chacon Gamboa)
Gardiner, Brian O. C.
Genetic and environmental variation in Pieris brassicae 2(2): 127-1 36
The rearing of Dirphiopsis eumedide (Saturniidae) 4(4):287-291
Notes on Eacles penelope (Saturniidae) 5(3): 177-1 80
Rearing Euleucophaeus rubridorsa and E. lex 6(l):53-58
Gynandromorphism in Pieris brassicae L 1 1 (3): 1 29- 1 40
The early stages of various species of the genus Dirphia
(Saturniidae) 1 3(2): 101-1 14
Melanie Papilio machaon larvae 1 5(3): 1 84
The early stages of Leucanella memusae ssp. gardinerii Lemaire
(Saturniidae) 1 5(4):20 1 -205
Garth, John S. and James Wilson Tilden
Yosemite butterflies: An ecological survey of the butterflies of the
Yosemite sector of the Sierra Nevada, California 2( 1 ): 1 -96
(see also Perkins, Jr., Edwin M. and John S. Garth)
Geiger, Hansjurg
Enzyme electrophoretic studies on the genetic relationships of Pierid
butterflies (Lepidoptera: Pieridae) I. European taxa 1 9(4): 181-1 95
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera) - The case for enzyme electrophoresis 26(l-4):64-72
(see also Kudrna, Otakar and Hansjurg Geiger)
(see also Shapiro, Arthur M. and Hansjurg Geiger)
Geiger, Hansjurg and Arthur M. Shapiro
Electrophoretic evidence for speciation within the nominal species
Anthocharis sara Lucas (Pieridae) 25(1): 15-24
Gilbert, Lawrence E., Hugh S. Forrest, Thomas D. Schultz and Donald J.
Harvey
Correlations of ultrastructure and pigmentation suggest how genes control
development of wing scales of Heliconius butterflies. . . 26( 1 -4): 141-1 60
Gilchrist, George W. (see Rutowski, Ronald L. and George W. Gilchrist)
Gilmour, C. (see Chua, K. E., J. C. E. Riotte and C. Gilmour)
Glick, Perry A. (see Kendall, Roy O. and Perry A. Glick)
Gorelick, Glenn Alan
A new subspecies of Callophrys dumetorum from Washington and
Oregon 7(2):99-104
Grey, L. P.
On the Gunder collection of Argynnids 8(2):55-64
Groothuis, D. R. (see Ferris, Clifford D. and D. R. Groothuis)
Guppy, Richard
Further observations on "hilltopping" in Papilio zelicaon 8(3): 1 05- 117
Gupta, M. L. (see Narang, R. C. and M. L. Gupta)
Gwynne, Darryl (see Alcock, John and Darryl Gwynne)
Habeck, Dale H. (see Heitzman, Roger L. and Dale H. Habeck)
12
AUTHOR INDEX
J. Res. Lepid.
Hadley, Mark
Ocellus variation and wingspan in Attacus atlas Linnaeus, Is there a
relationship? 16(3):141-145
Haeger, J. Fernandez
Notes: Notes on the biology of Brephidium exilis (Boisduval)
(Lycaenidae) 26(1 -4):254-255
Hammond, Paul C.
The colonization of violets and Speyeria butterflies on the ash-pumice
fields deposited by Cascadian volcanoes 20(3): 1 79- 191
Opinion: A rebuttal to the Arnold classification of Speyeria callippe
(Nymphalidae) and defense of the subspecies concept. . . . 24(3):197-208
(see also McCorkle, David V. and Paul C. Hammond)
Hammond, Paul C. and David V. McCorkle
The decline and extinction of Speyeria populations resulting from human
environmental disturbances (Nymphalidae: Argynninae). . 22(4):217-224
Hanson, William R.
Estimating the density of an animal population 6(3):203-247
Hanson, William R. and William Hovanitz
Trials of several density estimators on a butterfly population. . 7(l):35-49
Harmsen, R. (see Ward, P. S., R. Harmsen and P. D. N. Hebert)
Harmsen, R., P. D. N. Hebert and P. S. Ward
On the origin of austral elements in the moth fauna of south-eastern
Ontario, including a number of species new for Canada. . 1 2(3): 1 27-1 34
Harris, T. T. (see Munger, Francis and T. T. Harris)
Harrison, S. J. (see Platt, Austin P. and S. J. Harrison)
Harvey, Donald J. (see Gilbert, Lawrence E., Hugh S. Forrest, Thomas D.
Schultz and Donald J. Harvey)
Hayes, Jane Leslie
Colias alexandra : A model for the study of natural population of
butterflies 23(2):1 13-124
Heath, J. E. (see Adams, Phillip A. and J. E. Heath)
Hebert, P. D. N. (see Harmsen, R., P. D. N. Hebert and P. S. Ward)
(see also Ward, P. S., R. Harmsen and P. D. N. Hebert)
Heitzman, John Richard
The complete life history of Staphylus hayhursti 2(2): 170-172
Early stages of Euphyes vestris 3(3): 1 5 1 - 1 54
The habits and life history of Amblyscirtes nysa (Hesperiidae) in
Missouri 3(3): 1 54- 1 56
The life history of Amblyscirtes belli in Missouri 4(l):75-78
A new species of Papilio from the eastern United States
(Papilionidae) 1 2( 1 ): 1 - 1 0
Heitzman, John Richard and Roger L. Heitzman
The life history of Amblyscirtes linda (Hesperiidae) 8(3):99-104
Hesperia metea life history studies 8(4): 187- 193
New butterfly records for the United States (Hesperiidae and
Libytheidae) 10(4):284-286
Atrytonopsis hianna biology and life history in the Ozarks. . . 13(4):239-245
Heitzman, Roger L.
Life history studies of Idaea obfusaria (Walker) 1 2(3): 145-1 50
An annotated checklist of the Missouri Geometridae 1 2(3): 1 69- 179
A new species of Hypagyrtis (Geometridae) 13(1 ):43-48
J. Res. Lepid.
AUTHOR INDEX
13
Heitzman, Roger L. (continued)
Studies of the ova and first instar larvae of Geometridae (Ennominae).
1 1 3(3): 1 49- 1 56
(see also Heitzman, John Richard and Roger L. Heitzman)
Heitzman, Roger L. and Wilbur R. Enns
Descriptions of a new species of Eupithecia and the male of E. cocoata
Pearsall (Geometridae) 16(2):75-82
Male genitalic illustrations and notes on the Larentiinae (Geometridae) of
Missouri 1 7(3): 145-1 67
Heitzman, Roger L. and Dale H. Habeck
Taxonomic and biological notes on Bellura gortynoides Walker
(Noctuiidae) 18(4):228-235
Henne, Christopher
Life history studies on the lithosina-miona-casta complex of the genus
Annaphila 6(4):249-256
(see also Comstock, John Adams and Christopher Henne)
(see also Hogue, Charles L., Frank P. Sala, Noel McFarland and
Christopher Henne)
Heppner, John B.
The distribution of Paratrytone melane and its spread into San Diego
County 10(4):287-300
Habitat: Adela bella in Florida 13(l):67-72
Habitat: Brephidium pseudofea (Lycaenidae) 1 3(2):99- 1 00
Book Review: Bradley, Tremewan and Smith: British Torricoid moths,
Cochylidae and Tortricidae: Tortricinae 15(4):208-210
A new Tortyra from Cocos Island, Costa Rica (Lepidoptera:
Choreutidae) 1 9(4): 1 96- 1 98
Revision of the Oriental and Nearctic genus Ellabella (Lepidoptera:
Copromorphidae) 23(l):50-73
Book Review - Tarmann: Generic revision of the American Zygaenidae,
with descriptions of New Genera and Species 24(4):383-384
Herlan, Peter J.
A new subspecies of Limenitis archippus 9(4):2 17-222
Highfill, Jerry W. (see Miller, Thomas A., William J. Cooper and Jerry W.
Highfill)
Hoegh-Guldberg, Ove
Pupal sound production of some Lycaenidae . 1 0(2): 127-1 47
Hogue, Charles L.
A standard method for mounting whole adult lepidoptera on slides
utilizing polystyrene plastic l(3):223-235
A new species of Basilodes from the southwestern United States
(Noctuidae) 4(4):275-280
Hogue, Charles L., Frank P. Sala, Noel McFarland and Christopher
Henne
Systematics and life history of Saturnia ( Calosaturnia ) albofasciata in
California (Saturniidae) 4(3): 173-1 84
Holland, Richard
Butterflies of middle and southern Baja California 1 1 (3): 147-1 60
Aberrant New Mexican butterflies 19(2):88-95
Parallel albinism in two Theclines (Lycaenidae) 2 1 (3): 158
Honch, Gerald (see Hovanitz, William, Vincent C. S. Chang and Gerald
Honch)
14
AUTHOR INDEX
J. Res. Lepid.
Hovanitz, William
The distribution of the species of the genus Pieris in North
America l(l):73-83
Argynnis and Speyeria 1(1 ):95-96
Geographical distribution and variation of the genus Argynnis. I.
Introduction. II. Argynnis idalia 1 (2): 1 17-123
The relation of Pieris virginiensis Edw. to Pieris napi L. species formation
in Pieris ? 1(2): 124- 134
Geographical distribution and variation of the genus Argynnis. III.
Argynnis diana 1(3):20 1-208
The origin of a sympatric species in Colias through the aid of natural
hybridization 1 (4):26 1 -274
The origin of a sympatric species in Colias through the aid of natural
hybridization 2(3):205-223
Book Review - Fox and Fox: Introduction to Comparative
Entomology 3(1):8
Book Review - C. F. dos Passos: A Synonymic List of the Nearctic
Rhopalocera 3( 1 ): 1 8
Book review - Dos Passos: Synonymic list of Nearctic
Rhopalocera 3( 1 ): 1 9-24
The origin of a sympatric species in Colias through the aid of natural
hybridization 3(l):37-44
A Colias Christina gynandromorph 4( 1 ):4 1
Colias christina-alexandra intergradation. Cover illustration 4(1):42
Alaska refreshments 4(2): 113
Parallel ecogenotypical color variation in butterflies (cover
illustration) 4(2): 114
Ecological color variation in some Argynnis of the western United
States 6(3): 197-1 98
Natural habitats - Philotes sonorensis 6(3): 199-202
Man-made habitat for Colias eurytheme 6(4):267
Present and Ice Age Life Zones and distributions 7(1 ):3 1 -34
Habitat - Argynnis callippe laurina 7(1 ):50
Habitat - Pieris beckeri 7(1 ):56
Habitat: Specific type locality, Plebejus icariodes missionensis. . . . 7(2): 1 22
Habitat: General type locality, Glaucopsyche lygdamus xerces , Plebejus
icariodes pheres 7(2): 126
Habitat - Zerene caesonia eurydice 7(4): 182, 190
Habitat - Euchloe hy antis andrewsi 8(1): 16- 17
Habitat - Argynnis nokomis 8(1):20
Habitat - Colias philodice eriphyle and Colias eurytheme 8(4): 182
Habitat - Oeneis chryxus Stanislaus 8(4): 194
Book review - Ronald W. Hodges: The Moths of North America Fasc. 21
Sphingoidae 9( 1 ): 1 0
Book review - Malcolm Barcant: Butterflies of Trinidad and
Tobago 9(1):24
Habitat - Colias vautieri 9(2): 100
Habitat - Yramea cytheris 9(2): 126
Habitat - Argynnis adiaste 9(3): 168
Book review - Brown and Heineman: Jamaica and its Butterflies. 1 0(2): 1 48
Cover Illustration: Variation in Colias nastes of Lapland 1 2(3): 1 80
J. Res. Lepid.
AUTHOR INDEX
15
Hovanitz, William (continued)
Ecological color variation in a butterfly and the problem of "protective
coloration" 1 7(S): 1 0-25
Parallel ecogenotypical color variation in butterflies 17(S):26-65
(see also Hanson, William R. and William Hovanitz)
Hovanitz, William and Vincent C. S. Chang
The effect of various food plants on survival and growth rate of
Pieris 1(1 ):2 1 -42
Three factors affecting larval choice of food plant 1(1 ):5 1-61
The effect of hybridization of host-plant strains on growth rate and
mortality of Pieris rapae 1 (2): 157-1 62
Change of food plant preference by larvae of Pieris rapae controlled by
strain selection, and the inheritance of this trait 1(2): 1 63- 1 68
Selection of ally 1 isothiocyanate by larvae of Pieris rapae and the
inheritance of this trait 1(3): 1 69- 182
Ovipositional preference tests with Pieris 2(3): 1 85-200
Comparison of the selective effect of two mustard oils and their
glucosides on Pieris larvae 2(4):28 1-288
Adult oviposition responses in Pieris rapae 3(3):1 59- 1 72
The alteration of host plant specificity in larvae of Pieris rapae by
induction 4( 1 ): 1 3-2 1
Hovanitz, William, Vincent C. S. Chang and Gerald Honch
The effectiveness of different isothiocyanates on attracting larvae of
Pieris rapae l(4):249-259
Howarth, T. G.
Notes on the biology of Lamproptera curius Walkeri Moore (Lepidoptera:
Papilionidae) 15(1 ):27-32
Huntzinger, David H. (see Tilden, James Wilson and David H. Huntzinger)
Iglesias, C. (see De Viedma, M. G., J. R. Baragano, A. Notario, M. Rodero
and C. Iglesias)
Irwin, Roderick R.
Euphyes dukesi and other Illinois Hesperiidae 8(4): 183-1 86
Further notes on Euphyes dukesi 1 0(2): 185-188
Jaros, Josef (see Spitzer, Karel and Josef Jaros)
Jimenez, Gabriela and Jorge Soberon
Notes: Laboratory rearing of Sandia xami xami (Lycaenidae:
Eumaeini) 27(3):268-271
Johnson, John W.
Two new California Catocala subspecies (Noctuidae) 20(4):245-248
The immature stages of six California Catocala (Lepidoptera:
Noctuidae) 23(4)303-327
Johnson, John W. and Erich Walter
A new species of Coloradia in California (Saturniidae,
Hemileucinae) 18(1 ):60-66
The immature stages of Catocala erichi Brower (Lepidoptera:
Noctuidae) 23(3):23 1 -235
Johnson, Kurt
The Butterflies of Nebraska 1 1 ( 1 ): 1 -64
Post Pleistocene environments and montane butterfly relicts on the
western Great Plains 1 4(4):2 1 6-232
16
AUTHOR INDEX
J. Res. Lepid.
Johnson, Kurt, David Matusik and Rick Rozycki
A study of Protesilaus microdamas (Burmeister) and the little-known P.
huanucana (Varea de Luque) (Papilionidae) 27(2):83-95
Johnson, Kurt and Eric L. Quinter
Commentary on Miller and Brown vs. Erhlich and Murphy et al .:
Pluralism in systematics and the worldwide nature of kinship
groups 21(4):255-269
Johnson, Kurt, Eric L. Quinter and David Matusik
A new species of Calisto from Hispaniola with a review of the female
genitalia of Hispaniolan congeners (Satyridae) 25(2):73-82
Johnson, Samuel A.
Notes: Lateral perching in Brephidium exilis (Boisduval) (Lycaenidae) in
Texas 23(1 ): 1 04- 1 06
Justice, John A. (see Scott, James A. and John A. Justice)
Kamm, J. A.
Antennal sensilla of some Crambinae 1 6(4):20 1 -207
Kasy, Dr. Fritz
Das naturhistorische Museum in Wein und seine
Lepidopterensammlung 13(1 ):63-65
Kean, Philip J. (see Simmons, Robert S., William A. Andersen and Philip J.
Kean)
Keiper, Ronald
Field studies of Catocala behavior 7(2): 1 13-121
Daytime vision by the moth, Exyra ridingsi 7(2):131-132
Kendall, Roy O. and Perry A. Glick
Rhopalocera collected at light in Texas 10(4):273-283
Kilduff, Thomas S.
A population study of Euptychia Hermes in northern Florida. 1 1 (4):2 1 9-228
Kingsolver, Joel G.
Butterfly thermoregulation: Organismic mechanisms and population
consequences 24( 1 ): 1 -20
Kolyer, John M.
The feeding of coloring matters to Pier is rapae larvae 4(3): 1 59- 172
Vital staining of Colias philodice and C. eurytheme 5(3): 137-1 52
Note on vital staining of Actias luna silk 7(l):29-30
Note on damaged specimens 7(2): 1 05- 111
Development of the markings on the pupal wing of Pieris rapae
(Pieridae) 8(3):69-90
Vital staining as evidence for wing circulation in the cabbage butterfly
Pieris rapae 1 1 (3): 161-173
Kolyer, John M. and Anne Marie Reimschuessel
Scanning electron microscopy on wing scales of Colias eurytheme. 8( 1 ): 1 - 1 5
Krebs, Robert A.
The mating behavior of Papilio glaucus (Papilionidae) 26( l-4):27-31
Krizek, George O. and Paul A. Opler
Observations on Problema bulenta 25(2): 1 46- 1 48
Kudrna, Otakar
On the nomenclature of Colias alfacariensis Berger 1948 (Lepidoptera:
Pieridae) 20(2):103-1 10
Book Review - Dowdeswell: The Life of the Meadow Brown. . . . 20(3):192
An annotated catalogue of the butterflies (Lepidoptera: Paplionoidea)
named by Roger Verity 21(1):1-1 06
J. Res. Lepid.
AUTHOR INDEX
17
Kudrna, Otakar (continued)
Book Review - Larsen and Larsen: Butterflies of Oman 21(3):210
Book Review - Heath: Threatened Rhopalocera (butterflies) in
Europe 22(2):1 59-160
Book Review - Dabrowski: Ginace i zagrozone gatunki motyli
(Lepidoptera) w. faune Polski 22(2):160
Book Review - Brooks and Knight: A Complete Guide to British
Butterflies 23( 1 ): 1 08
Book Review - Whalley: The Mitchell Beazley Pocket Guide to
Butterflies 23( 1 ): 1 08- 1 09
Book Review - Bjorn: The Butterflies of Northern Europe. . . 23(1 ): 1 09- 1 10
Book Review - Larsen: Butterflies of Saudi Arabia and its
Neighbors 24(l):93-94
Book Review - Kock: Wirbestimmen Schmetterlinge 24(1):94
Book Review - Landing: Factors in the Distribution of Butterfly color and
Behavior Patterns - Selected Aspects 24(4):375-376
Book Review - Sbordioni and Forestiero: The World of Butterflies and II
Mondo delle Farfalle 24(4):382-383
Book Review - Friedrich: Breeding of Butterflies and
Moths 26(l-4):285-286
Book Review - Weidemann: Tagfalter 1. H-J 26(l-4):288
(see also Balletto, Emilio and Otakar Kudrna)
Kudrna, Otakar and Emilio Balletto
An annotated catalogue of the Skippers (Lepidoptera: Hesperiidae) named
by Roger Verity 23(1 ):35-49
Kudrna, Otakar and Hansjurg Geiger
A critical review of "Systematische Untersuchungen am Pieris
napi-bryoniae- Komplex (s.l.)" Lepidoptera: Pieridae) by Ulf
Eitschberger 24(l):47-60
La Due, Lillian
Oxygen consumption and metabolic rate of Papilio zelicaon pupae in a
state of delayed eclosion 3(4):197-206
Lakshmi, G. Vijaya (see Rao, N. Nageswara and G. Vijaya Lakshmi)
Lamas, Gerardo
Occurrence of the "Elymi" aberrant phenotype in Vanessa carye (Huebner)
(Nymphalidae) 22(2):1 15-1 17
Landing, Benjamin H.
Opinion: Rebuttal to Murphy on factors to the distribution of butterfly
color and behavior patterns - selected aspects 25(l):67-70
Langston, Robert L.
Extended flight periods of coastal and dune butterflies in
California 13(2):83-98
The Rhopalocera of Santa Cruz Island, California 18(1 ):24-35
Larsen, Torben B., John Riley and Michael A. Cornes
The butterfly fauna of a secondary bush locality in Nigeria. . . 18(1 ):4-23
Lawrence, Donald A. and John C. Downey
Morphology of the immature stages of Everes comyntas Godart. 5(2):61-96
Leck, Charles F.
Butterflies of St. Croix 1 2(3): 1 6 1 - 1 62
(see also Emmel, Thomas C. and Charles F. Leck)
Lederhouse, Robert C. (see Scriber, J. Mark and Robert C. Lederhouse)
(see also Scriber, J. Mark, Mark H. Evans and Robert C. Lederhouse)
18
AUTHOR INDEX
J. Res . Lepid.
Lees, E. and D. M. Archer
Diapause in various populations of Pieris napi L. from different parts of
the British Isles 1 9(2):96- 1 00
LeMaire, Claude
A new subspecies of Hemileuca maia from central Texas (Attacidae,
Hemileucinae) 1 8(3):2 1 2-2 1 9
A new species of Automeris cecrops (Attacidae: Hemileucinae). 18(4):236-240
(see also Donahue, Julian P. and Claude LeMaire)
LeMaire, Claude and Richard S. Peigler
Samia watsoni Obertheur color plate 18(1 ):67
LeMaire, Claude and Kirby L. Wolfe
Three new species of Paradirphia (Saturniidae: Hemileucinae) from Mexico
and Central America with notes on the immature stages. . 27(3): 1 97-2 1 2
Leuschner, Ronald H.
California coastal Eupithecia with description of a new species
(Geometridae) 4(3): 191-1 97
(see also Buckett, John S. and Ronald H. Leuschner)
Lintereur, Greg (see Scriber, J. Mark and Greg Lintereur)
Lorkovic, Zdravko
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera) 24(4)334-358
Lounibos, L. P. (see Buckett, John S. and L. P. Lounibos)
Lundgren, Lennart
The role of intra- and interspecific male:male interactions in Polyommatus
icarus Rott. and some other species of blues (Lycaenidae). 16(4):249-264
Mallet, James (see Singer, Michael C. and James Mallet)
Mallorie, H. C. (see Thomas, Chris D. and H. C. Mallorie)
Marini, Mario (see Trentini, Massimo and Mario Marini)
Marshall, Larry D.
Protein and lipid composition of Colias philodice and C. eurytheme
spermatophores and their changes over time 24( 1 ):2 1 -30
Masters, John H.
Ecological and distributional notes on Erebia disa (Satyridae) in central
Canada 7(l):19-22
Records of Colias gigantea from southwest Manitoba and
Minnesota 8(3): 1 29- 1 32
Ecological and distributional notes on Erebia discoidalis (Satyridae) in the
north central states 9( 1 ): 1 1-16
Concerning Colias eurytheme alberta Bowman (Pieridae) 9(2):97-99
Concerning Colias Christina mayi 9(4):227-232
Heliconius cydno in Venezuela with descriptions for two new
subspecies 10(4):267-272
Habitat: Oeneis macounii Edwards 10(4):30 1-302
Habitat: Oeneis jutta ascerta Masters & Sorenson 1 1(2):94
Butterfly records for three northwest Wisconsin counties. . . . 1 1 (3): 175-1 82
Lack of melanism in Colias (Cover illustration) 1 1 (4):2 1 8
Concerning Heliconius cydno aberration "larseni" Niepelt. ... 1 1(4):25 1-254
A list of the butterflies of the Willow River State Park,
Wisconsin 14(1 ):57-59
Variation in Colias alexandra Christina Edwards (Pieridae) in southwest
Manitoba 1 4(3): 1 48- 157
J. Res . Lepid .
AUTHOR INDEX
19
Mather, Bryant
Euphyes dukesi 2(2): 161-1 69
The southern limits of the range of Pier is napi and P.
virginiensis 3(l):45-48
Speyeria cybele in Mississippi. (Argynninae: Argynnis) 5(4):252-253
Euphyes dukesi - additional record 5(4):253-254
Mathew, George
Variations in the wing venation of Pteroma plagiophleps Hampson
(Lepidoptera: Psychidae) 24(4):359-363
Mattoni, Rudolf H. T.
Distribution and pattern of variation in Philotes rita 4(2):8 1-101
The Scolitantidini I: Two new genera and a generic rearrangement
(Lycaenidae). 16(4):223-242
Editorial 16(4):243-244
Dr. William Hovanitz, 1915-1977 17(S):l-6
Editorial: Extinction of the British Large Blue Butterfly 1 8( 1 ): 1 ,3
Book Review - Dornfeld: The Butterflies of Oregon 18(1 ):68
The Scolitantidini II. The World’s smallest butterfly? Notes on Turancinci,
and a new genus and species from Afghanistan
(Lycaenidae) 18(4):256-264
Book Review - Miller and Brown: A Catalogue/Checklist of the Butterflies
North of Mexico 19(4):243-244
Book Review - Eliot and Kawazoe: Blue Butterflies of the Lycaenopsis -
Group 23( 1 ): 1 11-112
Notes: Three intersubfamilial matings in nature (Lycaenidae). 24(l):86-87
Book Review - Miyata: Handbook of Moth Ecology 24(1):88
Book Review - King and Leppla: Advances and Challenges in Insect
Rearing 24(1):88
Book Review - Tekulsky: The Butterfly Garden 25(2): 1 56
Book Review - Menke and Miller: Entomology of the California Channel
Islands 25(2):1 56
Book Review - Ehrlich: The Machinery of Nature 26( l-4):287-288
The Euphilotes battoides complex: recognition of a species and description
of a new subspecies 27(3): 173-1 85
Book Review - McKibben: The End of Nature 27(3):271
Book Review - McFarland: Portraits of South Australian Geometrid
Moths 27(3):272
Book Review - Emmet and Heath: The Moths and Butterflies of Great
Britain and Ireland. Vol 7, Part 1, Hesperiidae - Nymphalidae, The
Butterflies 27(3):272-274
Book Review - Nielsen and Kristensen: Primitive Ghost Moths. . 27(3):274
(see also Dimock, Thomas E. and Rudolf H. T. Mattoni)
Mattoni, Rudolf H. T. and Marvin S. B. Seiger
Techniques in the study of population structure in Philotes
sonorensis l(4):237-244
Mattoon, Sterling O. (see Scott, James A. and Sterling O. Mattoon)
Matusik, David (see Johnson, Kurt, David Matusik and Rick Rozycki)
(see also Johnson, Kurt, Eric L. Quinter and David Matusik)
McAlpine, Wilbur S.
The butterfly genus Calephelis 1 0( 1 ): 1 - 1 25
Observations on life history of Oarisma pawesheik (Parker)
1870
1 1(2):83-93
20
AUTHOR INDEX
J. Res. Lepid.
McCabe, Timothy L.
Speyeria idalia 16(1 ):68
The larva of Acronicta spinigera Guenee (Noctuidae) 17(3): 173- 179
Portable apparatus for photographing genitalic dissections. . . 27(2): 1 15-1 19
McCabe, Timothy L. and Richard L. Post
North Dakota butterfly calendar 15(2):93-99
McCaffrey, J.
Notes: Dione moneta poeyii Butler (1873) in New Mexico (Lepidoptera:
Nymphalidae) 23( 1 ): 1 06- 1 07
McCorkle, David V. (see Hammond, Paul C. and David V. McCorkle)
McCorkle, David V. and Paul C. Hammond
Genetic experiments with a calverleyi- like mutation isolated from Papilio
bairdi oregonius (Papilionidae) 27(3): 1 86- 191
McFarland, Noel
The moths (Macroheterocera) of a chaparral plant association in the Santa
Monica Mountains of southern California 4(l):43-73
A moth sheet 5(l):29-36
Overcoming difficulties with the pupae of Euproserpinus phaeton
mojave 5(4):249-252
Spring moths of a natural area northeastern Kansas 6( 1 ): 1 - 1 8
Rearing techniques for speeding up larval stages of some root or
stem-boring Lepidoptera 7(3): 1 66
Botanical names in entomological papers and habitat studies. . . 9(2):89-96
Notes on describing, measuring, preserving and photographing the eggs of
lepidoptera 10(3):203-214
Egg photographs depicting 40 species of Southern Australian
moths 10(3):215-247
Some observations on the eggs of moths and certain aspects of first instar
larval behavior 12(4):199-208
3 Stacks of the eggs of Hemistola hatching 13(1 ):2 1 -22
Live Geometrid (cover illustration) 14(1 ):60
Retention of cast head capsules by some Nolid immatures in four Old
World countries 1 7(4):209-2 1 7
(see also Hogue, Charles L., Frank P. Sala, Noel McFarland and
Christopher Henne)
McHenry, Paddy
The generic, specific and lower category names of the Nearctic butterflies.
Part 1 - the genus Pieris . 1(1 ):63-7 1
The generic, specific and lower category names of the Nearctic butterflies.
Part 2 - the genus Colias 1(3):209-221
Generic or subgeneric names closely related to Argynnis 2(3):229-239
The generic, specific and lower category names of the Nearctic butterflies.
Part 3 - Argynnids 3(4):231-268
"Corrections” to paper by P. McHenry 4( 1 ): 1 2
The generic, specific and lower category names of the Nearctic butterflies.
Part 4 - the genus Euptoieta 4(3):205-208
The generic, specific and lower category names of the Nearctic butterflies.
Part 5 - The genus Heliconius 6(l):65-68
The generic, specific and lower category names of the Nearctic butterflies.
Part 6 - the genus Dryas 6(4):263-265
The generic, specific and lower category names of the Nearctic butterflies.
Part 7 - the genus Dryadula 7(2): 112
J. Res . Lepid.
AUTHOR INDEX
21
McHenry, Paddy (continued)
The generic, specific and lower category names of the Nearctic butterflies.
Part 8 - the genus Agraulis 7(2): 1 27- 1 30
McLeod, L.
Controlled environment experiments with Precis octavia Cram. . . 7(1):1-18
Controlled environment experiments with Precis octavia Cram. . 8(2):53-54
Menna- Barreto, Ymara and Aldo M. Araujo
Evidence for host plant preferences in Heliconius erato phyllis from
southern Brazil (Nymphalidae) 24( 1 ):4 1 -46
Meyers, Linda (see Stimson, John and Linda Meyers)
Miller, Jacqueline Y.
The head capsule of selected Hesperioidea 9(4): 1 93-2 1 4
Miller, Lee D.
Systematics and zoogeography of the genus Phanus
(Hesperiidae) 4(2):1 15-130
A review of the West Indian "Choranthus" 4(4):259-274
On Mexican Satyridae with description of a new species 7(1 ):5 1-55
Nomenclature of wing veins and cells 8(2):37-48
Miller, Lee D. and F. Martin Brown
Butterfly taxonomy: A reply 20(4): 193-198
Miller, Scott E.
Supplementary notes on the distribution of Epargyreus clarus in southern
California (Hesperiidae) 15(4):206-207
Paratrytone melane in San Luis Obispo County, California
(Hesperiidae) 1 6(2): 1 3 1 - 1 32
Book Review - Gilbert: A Compendium of the Biographical Literature on
Deceased Entomologists 17(3):207
Preface to reprints on ecogenotypical color variation in butterflies by
William Hovanitz 17(S):7-9
Publications of William Hovanitz 17(S):66-76
Book Review - Hollis (ed.): Animal Identification: A Reference Guide. Vol
3 Insects 21(3):209-210
Notes: Type locality of Papilio indra pergamus (Lepidoptera:
Papilionidae) 23(2): 175
Butterflies of the California Channel Islands 23(4):282-296
(see also Becker, Vitor O. and Scott E. Miller)
(see also Orsak, Larry J. and Scott E. Miller)
Miller, Thomas A.
Notes: Weights and dimensions of Hyalophora euryalis and pupae
(Lepidoptera: Saturniidae) 24(l):83-84
Miller, Thomas A., William J. Cooper and Jerry W. Highfill
Susceptibility of eggs and first-instar larvae of Callosamia promethea and
Antheraea polyphemus to Malathion 25( 1 ):48-5 1
Mohamed, U. V. K. and Humayun Murad
Studies on the excretory system of the fully grown larva of Callograma
festiva Donov. (Noctuidae) 1 6(4):2 1 3-221
Mohanty, P. K. and B. Nayak
Karyology of three Indian Lasiocampid moths (Lepidoptera). 19(4):227-229
Chromosome studies in sixteen species of Indian Pyralid moths
(Pyralidae) 20(2):86-96
On the supernumerary chromosomes of Tarache tropica Guen.
(Lepidoptera: Noctuidae) 20(3): 1 70- 173
22
AUTHOR INDEX
J. Res. Lepid.
Mohanty, P. K. and B. Nayak (continued)
Chromosomes of seven species of Indian Sphingid moths. . . . 21(4):238-244
Karyotypes of some Indian Noctuid moths (Lepidoptera). . . . 22(4):238-248
Montesinos, Jose Luis Viejo
Diversity and species richness of butterflies and skippers in central Spain
habitats 24(4):364-371
Montgomery, Johnson C. (see Shields, Oakley and Johnson C. Montgomery)
Mori, James R. (see Shields, Oakley and James R. Mori)
Morton, Ashley C.
Book Review - Singh: Artificial Diets for Insects, Mites and
Spiders 1 7(3):207-208
Rearing butterflies on artificial diets 1 8(4):22 1 -227
Muller, Joseph
Is air pollution responsible for melanism in lepidoptera and for scarcity of
all orders of insects in New Jersey? 1 0(2): 1 89- 1 90
Aberrant species of New Jersey Lepidoptera 1 5(3): 1 44- 1 45
Mullins, Douglas (see Austin, George T. and Douglas Mullins)
Munger, Charles
An improved method for rearing the monarch butterfly 1 2(3): 1 63- 1 68
Munger, F. (see Urquhart, Francis A., N. R. Urquhart and F. Munger)
Munger, Francis and T. T. Harris
Laboratory production of the monarch butterfly, Danaus
plexippus 8(4): 169- 176
Munro, J. Alex
Biology of the Ceanothus stem-gall moth, Periploca ceanothiella (Cosens),
with consideration of its control 1 (3): 183-1 90
Murad, Humayun (see Mohamed, U. V. K. and Humayun Murad)
Murphy, Dennis D.
On the status of Euphydryas editha baroni with a range extension of E.
editha luestherae 2 1(3): 194- 198
Book Review - Arnold: Ecological studies of six endangered butterflies
(Lepidoptera, Lycaenidae): Island biogeography, patch dynamics,
habitat preserve 22(4):267-269
Book Review - Blab and Kudrna: Hilfsprogramm fur Schmetterlinge.
Okologie und Schutz von Tagfaltern und Widderchen. . . . 23(2): 1 69- 1 70
Bibliography 1982 No. 1 23(4):328-331
Book Review - Cater (ed): Love Among the Butterflies: The Travels and
Adventures of a Victorian Lady 23(4):335-336
Bibliography 1982-1983 No. 2 24(l):72-75
Book Review - Hodges et al: Checklist of the Lepidoptera North of
Mexico 24(l):95-96
Bibliography 1983-1984 No. 3 24(3):27 1 -275
Book Review - Landing: Factors in the Distribution of Butterfly Color
and Behavior Patterns - Selected Aspects 24(4):375-376
A response to Landing: On factors in the distribution of butterfly color
and behavior 25(3):2 1 3-2 1 4
Opinion: Are we studying our endangered butterflies to
death? 26(l-4):236-239
Bibliography 1984-1985 No. 4 26(l-4):248-253
(see also Ehrlich, Paul R. and Dennis D. Murphy)
J. Res. Lepid.
AUTHOR INDEX
23
Murphy, Dennis D. and John F. Baughman
Book Review - Scott: The Butterflies of North America. A Natural History
and Field Guide 26(l-4):275-278
Murphy, Dennis D. and Paul R. Ehrlich
Opinion: Crows, bobs, tits, elfs and pixies: The phoney "common name"
phenomenon 22(2): 1 54- 158
Biosystematics of the Euphydryas of the central Great Basin with the
description of a new subspecies 22(4):254-261
On butterfly taxonomy 23( 1 ): 1 9-34
Murphy, Dennis D. and Stuart B. Weiss
Notes: A bibliography of Euphydryas 26(l-4):256-264
Murty, A. S. and N. Nageswara Rao
Chromosome numbers in two species of Ergolis (Lepidptera:
Nymphalidae) 15(1 ):23-26
Note on the chromosomes of Byblia ilithyia (Drury)
(Nymphalidae) . 1 5(3): 1 29- 1 3 1
Mutuura, A. and T. N. Freeman
The North American species of the genus Zeiraphera 5(3): 1 53- 1 76
Muyshondt, Alberto (see Young, Allen M. and Alberto Muyshondt)
Nakamura, Ichiro
Japanese literature 20(2): 127-1 28
Japanese literature 20(3): 1 34- 135
Narang, R. C. and M. L. Gupta
Chromosome studies including a report of B-chromosome in a wild
silkmoth, Sonthonnaxia maenas (Doubleday) (Saturniidae:
Saturniinae) 1 8(3):208-2 1 1
Nayak, B.
Asynaptic meiosis in three species of Lepidopteran males. . . . 17(4):240-244
(see also Dash, A. K. and B. K. Nayak)
(see also Mohanty, P. K. and B. Nayak)
(see also Padhy, Kunja Bihari and B. Nayak)
Neck, Raymond W.
History of scientific study on a larval color polymorphism in the genus
Chlosyne (Nymphalidae) 14(1 ):4 1 -48
Lepidopteran foodplant records from Texas 15(2):75-82
Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) Nymphalidae
II. Additional larval food plant data 16(2):69-74
Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) (Nymphalidae)
III. Adult resources 1 6(3): 1 47- 1 54
Role of an ornamental plant species in extending the breeding range of a
tropical Skipper to subtropical southern Texas
(Hesperiidae) 20(3): 1 29- 1 33
Nekrutenko, Yuri P.
The hidden wing-pattern of some Palearctic species of Gonepteryx and its
taxonomic value 3(2):65-68
Three cases of gynandromorphism in Gonepteryx 4(2): 1 03- 1 07
Tertiary Nymphalid butterflies and some phylogenetic aspects of
systematic lepidopterology 4(3): 1 49- 158
A new subspecies of Gonepteryx amintha (Pieridae) from Yunnan,
mainland China 1 1 (4):235-240
24
AUTHOR INDEX
J. Res. Lepid.
Nentwig, Wolfgang
A tropical caterpillar that mimics faeces, leaves and a snake (Lepidoptera:
Oxytenidae: Oxytenis naemia ) 24(2): 136-141
New, T. R.
Notes: An early season migration of Catopsilia pomona (Lepidoptera:
Pieridae) in Java, Indonesia 24(l):84-85
Newcomer, E. J.
The synonymy, variability and biology of Lycaena nivalis 2(4):27 1-280
Life histories of Papilio indra and Papilio oregonius 3(l):49-54
Three western species of Polites 5(4):243-247
Nicolay, S. S.
Illustrations and descriptions of species of some Pyrrhopyginae from
Panama (Hesperiidae) 1 3(3): 181-1 90
Nicolay, S. S. and G. B. Small, Jr.
Illustrations and descriptions of some species of Pyrrhopyginae from
Costa Rica, Panama and Columbia (Hesperiidae) 19(4):230-239
Nielsen, M. C.
Gynandromorphic Polites skippers (Hesperiidae) 1 6(4):209-2 1 1
Nijhout, H. F. (see Dohrmann, C. E. and H. F. Nijhout)
Notario, A. (see De Viedma, M. G., J. R. Baragano, A. Notario, M. Rodero
and C. Iglesias)
Novak, Ivo and Karel Spitzer
The relationship between migration and diapause during phylogeny and
ontogeny of some Lepidoptera 1 0(2): 181-1 84
Oehmig, Steffen
Hipparchia azorina (Strecker, 1899) (Satyridae) biology, ecology and
distribution on the Azores Islands 20(3): 1 36-160
Oliver, Charles G.
Notes: Celastrina ladon (Lycaenidae) female ovipositing on Sambucus
canadensis , a plant unsuitable for larval development 20(1):54
Oosting, Daniel P. and David K. Parshall
Ecological notes on the butterflies of the Churchill region of Northern
Manitoba 1 7(3): 1 88-203
Opler, Paul A.
Studies on the Nearctic Euchloe. Parts I, II 5(l):39-50
Studies on the Nearctic Euchloe. Part 3. Complete synonymical treatment.
Part 4. Type data and type locality restrictions 5(3): 1 85- 1 95
A gynandromorph of Lycaena gorgon 5(4):230
Studies on Nearctic Euchloe. Part 5. Distribution 7(2):65-86
Studies on Nearctic Euchloe. Part 6. Systematics of adults. .... 8(4): 153-168
Studies on Nearctic Euchloe - Part 7. Comparative life histories, hosts and
the morphology of the immature stages 1 3( 1 ): 1 -20
Book Review - Scott: The Butterflies of North America. A Natural History
and Field Guide 26(l-4):267-270
(see also Krizek, George O. and Paul A. Opler)
Opler, Paul A. and John S. Buckett
Seasonal distribution of "Macrolepidoptera" in Santa Clara County,
California 9(2):75-88
Correction 9(3): 1 56
Orsak, Larry J.
Recent captures of Anthocharis cethura catalina Meadows 1 4(2):85-89
Type locality for Calosaturnia walterorum Johnson (Saturniidae). 1 5(4):2 1 4
J. Res. Lepid .
AUTHOR INDEX
25
Orsak, Larry J. and Scott E. Miller
Habitat: Lycaena heteronea clara (Lepidoptera: Lycaenidae). . 17(3):204-206
Orsak, Larry J. and Douglas W. Whitman
Chromatic polymorphism in Ccillophrys mossii bayensis larvae
(Lycaenidae): Spectral characterization, short-term color shifts, and
natural morph frequencies 25(3): 1 88-20 1
Paclt, Juraj
The nomenclature in an important British check list (1972). Part
1 12(4):21 1-212
The nomenclature in an important British check list (1972) Part 2:
Corrections of family-group names for Geometridae
(lepidoptera) 1 3(3): 179-1 80
The nomenclature in an important British check list (1972). Part 3. Correct
gender for generic names derived from classical without change of
termination 13(4):267-270
The nomenclature in an important British check list (1972) Part 4: Correct
gender for some other generic names 17(1 ):24-26
Padhy, Kunja Bihari
Chromosome aberrations in the holocentric chromosomes of Philosamia
ricini (Saturniidae) 25(l):63-66
Padhy, Kunja Bihari and B. Nayak
Supernumerary chromosomes in the domesticated eri-silkmoth, Philosamia
ricini (Saturniidae: Lepidoptera) 20(1): 16- 17
Palm, Cheryl Ann (see Shapiro, Arthur M., Cheryl Ann Palm and Karen L.
Wcislo)
Parnell, J. R. (see Turner, T. W. and J. R. Parnell)
Parshall, David K.
Notes: Further notes regarding Colias hecla Lefebvre (Lepidoptera:
Pieridae) at Churchill, Manitoba 20(4):250
(see also Oosting, Daniel P. and David K. Parshall)
Passoa, Steven (see Sandberg, Sherri and Steven Passoa)
Peacock, John W. (see Shuey, John A. and John W. Peacock)
Pease, Jr., Roger W.
Variation of Uthetheisa ornatrix (Arctiidae) including a new species from
St. Croix, Virgin Islands 1 0(4):26 1 -264
Peigler, Richard S.
Rectification of a recent paper on Leucanella memusae gardineri. 16(4):222
Demonstration of reproductive isolating mechanisms in Callosamia
(Saturniidae) by artificial hybridization 19(2):72-81
Book Review - Barlow: An introduction to the Moths of South East
Asia 23(1):110-111
(see also LeMaire, Claude and Richard S. Peigler)
(see also Riotte, J. C. E. and Richard S. Peigler)
(see also Stone, Stephen E., Dean E. Swift and Richard S. Peigler)
Pellmyr, Olle
Plebeian courtship revisited: Studies on the female-produced male
behavior- eliciting signals in Lycaeides idas courtship
(Lycaenidae) 21(3):147-157
Perkins, Jr., Edwin M. and Edward V. Gage
On the occurrence of Limenitis archippus X L. lorquini hybrids. 9(4):223-226
26
AUTHOR INDEX
J. Res. Lepid.
Perkins, Jr., Edwin M. and John S. Garth
Limenitis weidemeyerii angusti fascia X L. astyanax arizonensis = (?) ab.
doudoroffi (Gunder) 1934 1 1(4):229-234
Perkins, Jr., Edwin M. and T. F. Perkins
A bilateral gynandromorph of Limenitis weidemeyerii lati fascia
(Nymphalidae) 1 1(3): 1 95- 1 96
Perkins, T. F. (see Perkins, Jr., Edwin M. and T. F. Perkins)
Persson, Bert
Diel egg-laying activity of Agrotis exclamationis (Noctuidae). 10(4):255-260
Petersen, Bjorn
The male genitalia of some Colias species 1 (2): 135-1 56
A method for breeding Pieris napi and Pieris bryoniae l(4):275-279
Comparative speciation in two butterfly families: Pieridae and
Nymphalidae 5(2):1 13-126
Petterson, M. A. and R. S. Wielgus
Acceptance of artificial diet by Megathymus streckeri
(Skinner) 1 2(4): 1 97-1 98
Pettus, David (see Simpson, Robert G. and David Pettus)
Pfeiler, Jr., Edward J.
The effect of pterin pigments on wing coloration of four species of
Pieridae 7(4):183-189
Philip, Kenelm W. and Peter Roos
Notes: Notes on Erebia occulta (Lepidoptera: Satyridae) 24( 1 ):8 1-82
(see also Troubridge, James T. and Kenelm W. Philip)
Pike, E. M.
A critique of the genus Boloria (Nymphalidae) as represented in "The
Butterflies of North America", with corrections, additions and a key to
species 1 8(3): 153-1 66
Platt, Austin P.
Stubby-winged mutants of Limenitis (Nymphalidae) - Their occurrence in
relation to photoperiod and population size 23(3):217-230
Platt, Austin P. and S. J. Harrison
"Black-light" induction of photoperiod-controlled diapause responses of
the viceroy butterfly, Limenitis archippus (Nymphalidae). 26( 1-4):1 77-1 86
Pljushtch, I. G.
Notes on a little known ecologically displaced blue, Agriades pyrenaicus
ergane Higgins (Lycaenidae) 27(2): 1 29- 1 34
Poore, Dennis M.
Nantucket Pine Tip Moth, Rhyacionia frustrana , in Kern County,
California: Integrated control and biological notes (Lepidoptera:
Tortricidae, Olethreutinae) 19(2):65-67
Porter, Adam H.
Notes: Courtship of a model (Nymphalidae: Adelpha) by its probable
Batesian mimic (Nymphalidae: Limenitis) 26(l-4):255-256
Porter, James W.
An annotated list of butterflies for northwestern Ohio 4(2): 1 09- 112
Post, Richard L. (see McCabe, Timothy L. and Richard L. Post)
Powell, Jerry A.
Biology and immature stages of Australian Ethmiid moths
(Gelechioidea) 20(4):214-234
Discovery of two new species and genera of Shaggy Tortricids related to
Synnoma and Niasoma (Tortricidae: Sparganothini) 24( 1 ):6 1-71
J. Res. Lepid.
AUTHOR INDEX
27
Powell, Jerry A. (continued)
Records of prolonged diapause in Lepidoptera 25(2):83-109
Pratt, Gordon F. (see Ballmer, Gregory R. and Gordon F. Pratt)
Pratt, Gordon F. and Greg R. Ballmer
The phenetics and comparative biology of Euphilotes enoptes (Boisduval)
(Lycaenidae) from the San Bernadino Mountains 25(2): 121-135
Priestaf, Richard Carl
Notes: Abnormal chrysalis of Papilio zelicaon (Papilionidae) 21(4):270
Notes: A melanic aberration of Philotes sonorensis (Lycaenidae) from
California 27(3):265-266
Pyle, Robert Michael
Opinion: Rebuttal to Murphy and Ehrlich on common names of
butterflies 23(l):89-93
Quinter, Eric L. (see Johnson, Kurt and Eric L. Quinter)
(see also Johnson, Kurt, Eric L. Quinter and David Matusik)
Rahn, Russell A.
A dwarf form of Euptoieta claudia 11 (3): 174
Rao, N. Nageswara
A study of the meiotic chromosomes of Ixias marianne (Cramer)
(Pieridae) 1 7(3): 1 70- 1 72
(see also Murty, A. S. and N. Nageswara Rao)
Rao, N. Nageswara and G. Vijaya Lakshmi
On the meiotic chromosomes of Argina stringa Cram (Arctiidae). 17(1 ):5 1 -52
Reimschuessel, Anne Marie (see Kolyer, John M. and Anne Marie
Reimschuessel)
Reinthal, Walfried J.
Butterfly aggregations 5( 1 ):5 1 -59
Riley, John (see Larsen, Torben B., John Riley and Michael A. Cornes)
Riotte, J. C. E.
On the distribution of some Skippers in Ontario 1 1 (2):8 1 -82
New food plant for Darapsa pholus (Cramer) 12(4):209-210
New food plant for Darapsa pholus (Cramer) 13(4):247-248
Significant additions to the Lepidopterous fauna of southeastern
Ontario 1 5(2): 1 0 1 - 1 02
Gynandromorphs in Hawaiian butterflies and moths 1 7( 1 ): 1 7- 1 8
Book Review - Ferguson: The Moths of America North of
Mexico 1 7(4):260-264
Moths of North America north of Mexico, Supplemental literature:
I 19(2):68-7 1
Moths of North America north of Mexico, Supplemental literature:
II 22(2): 131-1 34
(see also Chua, K. E., J. C. E. Riotte and C. Gilmour)
Riotte, J. C. E. and Richard S. Peigler
A revision of the American Genus Anisota (Saturniidae) 1 9(3): 101-1 80
Riotte, J. C. E. and G. Uchida
Butterflies of the Hawaiian Islands according to the stand of late
1976 17(l):33-39
Ritland, David B.
The effect of temperature on expression of the dark phenotype in Papilio
glaucus (Papilionidae) 25(3): 1 79- 187
Rodero, M. (see De Viedma, M. G., J. R. Baragano, A. Notario, M. Rodero
and C. Iglesias)
28
AUTHOR INDEX
J. Res. Lepid.
Roever, K ILIAN
Bionomics of Agathymus (Megathymidae) 3(2): 1 03- 1 20
Instar determination of Agathymus larvae 3(3): 1 48- 1 50
Roos, Peter (see Philip, Kenelm W. and Peter Roos)
Rosenberg, Risa H.
Notes: Description of the larvae of Coenonympha haydeni Edwards
(Lepidoptera: Satyridae) 24(4):394-395
Ross, Gary N.
Evidence for lack of territoriality in two species of
Hamadryas 2(4):24 1-246
Life history studies on Mexican butterflies. 1 3( 1 ):9- 1 7
Life history studies on Mexican butterflies. II, Anatole rossi. . . . 3(2):8 1 -94
Life history studies on Mexican butterflies. III. Nine Rhopalocera from
Ocotal Chico, Vera Cruz 3(4):207-229
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico 1 4(2): 103-1 24
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued ) 14(3): 169- 188
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued ) 14(4):233-252
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico ( continued ) 15(1 ):4 1 -60
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued) 1 5(2): 1 09- 1 28
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued) 15(3): 185-200
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued ) 1 5(4):225-240
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (concluded ) 1 6(2):8 7- 1 30
Rozycki, Rick (see Johnson, Kurt, David Matusik and Rick Rozycki)
Ruszczyk, Alexandre
Distribution and abundance of butterflies in the urbanization zones of
Porto Alegre, Brazil 25(3): 1 57- 1 78
Rutkowski, Frank
Notes: Hide and/or seek 2 1 (3):207
Rutowski, Ronald L.
Courtship behavior of the dainty sulfur butterfly, Nathalis iole with a
description of a new, faculative male display (Pieridae). . 20(3): 161-1 69
Courtship leading to copulation in the cloudless sulphur, Phoebis sennae
(Pieridae) 22(4):249-253
Sexual selection and the evolution of butterfly mating
behavior 23(2): 1 25- 1 42
Rutowski, Ronald L. and George W. Gilchrist
Male mate-locating behavior in the desert hackberry butterfly,
Asterocampa leilia (Nymphalidae) 26( 1 -4): 1-12
Sala, Frank P.
The Annaphila astrologa complex, with descriptions of three new
species 2(4):289-301
Synaxis mosesiani Sala; a new Synaxis from southern
California 9(3): 185-191
J . Res. Lepid.
AUTHOR INDEX
29
Sala, Frank P. (continued)
(see also Hogue, Charles L., Frank P. Sala, Noel McFarland and
Christopher Henne)
Sandberg, Sherri and Steven Passoa
New host records and morphological notes on four Tortricines
(Tortricidae). 27(2):104-108
Santiago-Alvarez, Candido and Brian A. Federici
Notes on the first-instar and two parasites of the clover cutworm
Scotogramma trifollii (Noctuidae: Hadeninae) 17(4):226-230
Schaefer, Paul W. and Paul J. Castrovillo
Gynaephora rossii (Curtis) on Mt. Katahdin, Maine, and Mt. Daisetsu,
Japan, and comparisons to records for populations from the Arctic
(Lymantriidae) 18(4):241-250
Schaefer, Paul W. and Kimito Furuta
A black-backed larval mutant of Lymantria dispar (L.) (Lepidoptera:
Lymantriidae) in Japan 1 8(3): 1 67- 170
Schaefer, Paul W., William E. Wallner and Mark Ticehurst
Notes: Incidence of the black backed larval mutant of Lymantria dispar
(L) (Lepidoptera: Lymantriidae) in Ukrainian SSR 23( 1 ): 1 03- 1 04
Schultz, Thomas D. (see Gilbert, Lawrence E., Hugh S. Forrest, Thomas D.
Schultz and Donald J. Harvey)
Scott, Glenn R. (see Scott, James A. and Glenn R. Scott)
Scott, James A.
Hilltopping as a mating mechanism to aid survival of low density
species 7(4): 191 -204
A list of Antillean butterflies 9(4):249-256
Mating of butterflies 1 1 (2):99- 127
Survey of ultraviolet reflectance of Nearctic butterflies 1 2(3): 151-1 60
Adult behavior and population biology of two skippers mating in
contrasting topographic sites 1 2(4): 181-1 96
Lifespan of Butterflies 12(4):225-230
Early stages and biology of Phyciodes orseis 12(4):236-242
Mate-locating behavior of the western North American
butterflies 1 4( 1 ): 1 -40
Early stages of Phyciodes pallida , P. orseis , and P. mylitta
(Nymphalidae) 14(2):84
Variability of courtship of the buckeye butterfly. Precis coenia
(Nymphalidae) 1 4(3): 1 42- 147
Amblyscirtes "Erna" a form of Amblyscirtes aenus 15(2):92
The identity of the Rocky Mountain Lycaena dorcas-helloides
complex 17(1 ):40-50
A survey of valvae of Euphydryas chalcedona, E. c. colon , and E. c.
anica * 1 7(4):245-252
Geographic variation in Lycaena xanthoides 18(1 ):50-59
Hibernal diapause of North American Papilionoidea and
Hesperioidea 1 8(3): 171 -200
Notes: Mate locating behavior of Gnophaela latipennis vermiculata G. & R.
(Pericopidae) 20(1):51
Book Review - Pyle: The Audubon Society Field Guide to North American
Butterflies 20(l):55-58
Book Review - Ferris and Brown eds.: Butterflies of the Rocky Mountain
States 20( 1 ):58-64
30
AUTHOR INDEX
J. Res. Lepid.
Scott, James A. (continued)
An apparent Interspecific Fx Hybrid Speyeria (Nymphalidae).20(3):174-175
Mate-locating behavior of western North American butterflies. II. New
observations and morphological adaptations 2 1 (3): 177-187
A review of Polygonia progne (oreas) and P. gracilis (zephyrus)
(Nymphalidae) including a new subspecies from the southern Rocky
Mountains 23(3): 1 97-2 1 0
The phylogeny of butterflies (Papilionoidea and
Hesperioidea) 23(4):24 1 -28 1
Notes: The origin of Satyrium calanus albidus 23(4):334
On the monophyly of the Macrolepidoptera, including a reassessment of
their relationship to Cossoidea and Castnioidea, and a reassignment of
Mimallonidae to Pyraloidea 25(l):30-38
Opinion: Parallelism and phylogenetic trees 27(3):257-258
Scott, James A. and John A. Justice
Intergradation between Callophrys dumetorum oregonensis and Callophrys
dumetorum af finis in northwestern U.S. (Lycaenidae) 20(2):81-85
Scott, James A. and Sterling O. Mattoon
Early stages of Speyeria nokomis (Nymphalidae) 20(1): 12- 15
Scott, James A. and Glenn R. Scott
Ecology and distribution of the butterflies of southern central
Colorado 17(2):73-128
Scott, James A. and Ray E. Stanford
Geographic variation and ecology of Hesperia leonardus
(Hesperiidae) 20( 1 ): 1 8-35
Scriber, J. Mark and Mark H. Evans
An exceptional case of paternal transmission of the dark form female
trait in the tiger swallowtail butterfly, Papilio glaucus (Lepidoptera:
Papilionidae) 25(2): 110-1 20
A new heritable color aberration in the tiger swallowtail butterfly, Papilio
glaucus (Papilionidae: Lepidoptera) 26(l-4):32-38
Bilateral gynandromorphs, sexual and/or color mosaics in the tiger
swallowtail butterfly, Papilio glaucus (Lepidoptera:
Papilionidae) 26(l-4):39-57
Scriber, J. Mark, Mark H. Evans and Robert C. Lederhouse
Hybridization of the Mexican tiger swallowtail, Papilio alexiares garcia
(Lep: Pap) with other P. glaucus group species and survival of pure and
hybrid larvae on potential host plants 27(3):222-232
Scriber, J. Mark and Robert C. Lederhouse
Hand-pairing of Papilio glaucus glaucus and Papilio pilumnus
(Papilionidae) and hybrid survival on various food plants. . 27(2):96-103
Scriber, J. Mark and Greg Lintereur
A melanic male aberration of Papilio glaucus canadensis from northern
Wisconsin 2 1(3): 1 99-20 1
Sears, T. A. (see Buckett, John S. and T. A. Sears)
Seiger, Marvin S. B. (see Mattoni, Rudolf H. T. and Marvin S. B. Seiger)
Selman, Charles L.
The flight periods of several sibling species of moths 12(4):217-224
Selman, Charles L. and H. E. Barton
Spring moth activity in relation to locality, temperature, and air
pressure 9( 1 ): 1 -9
J. Res. Lepid.
AUTHOR INDEX
31
Sette, Oscar Elton
Variation in the silvering of Argynnis ( Speyeria ) callippe in the interior
mountain area of south central California 1(1 ):3-20
Shapiro, Adrienne R. (see Shapiro, Arthur M. and Adrienne R. Shapiro)
Shapiro, Arthur M.
Melanie tendencies in Phalaenid and Geometrid moths in eastern
Pennsylvania 3( 1 ): 1 9-24
Polymorphism in Sunirci bicolorago (Noctuidae) 4(1): 1-5
Antepione thisaria and Xanthotype : A case of mimicry 4( 1 ):6- 1 1
Origin of autumnal "false broods" in common Pierid
butterflies 6(3): 181-1 93
The biology of Poanes viator 9(2): 109- 123
Postglacial biogeography and the distribution of Poanes viator and other
marsh butterflies 9(3): 1 25- 155
An interfamilial courtship (Nymphalidae, Pieridae) 1 1 (3): 1 97-1 98
Altitudinal migration of butterflies in the central Sierra
Nevada 1 2(4):23 1 -235
Natural and laboratory occurrence of "Elymi" phenotypes in Cynthia
cardui (Nymphalidae) 13(1 ):57-62
The butterfly fauna of the Sacramento Valley, California. . . . 13(2):73-82
Altitudinal migration of central California butterflies 1 3(3): 157-161
Butterflies of the Suisun Marsh, California 1 3(3): 191 -206
The genetics of subspecific phenotype differences in Pieris occidentalis
Reakirt and of variation in P. o. nelsoni W. H. Edwards
(Pieridae) 14(2):61-83
Why do California tortoiseshells migrate? 14(2):93-97
Supplementary records of the butterflies in the Sacramento Valley and
Suisun Marsh, lowland central California 1 4(2): 1 00- 1 02
The role of watercress, Nasturtium officinale as a host of native and
introduced pierid butterflies in California 1 4(3): 158-1 68
Habitat: Pieris occidentalis Nelsoni (Pieridae) 1 5(2): 1 03- 1 05
Habitat: Pieris occidentalis (Pieridae) 15(3): 182- 183
Autumnal false broods of multivoltine butterflies at Donner Pass,
California 16(2):83-86
Evidence for two routes of post-pleistocene dispersal in Poanes viator
(Hesperiidae) 1 6(3): 1 73-1 75
Photoperiod and temperature in phenotype determination of Pacific slope
Pierini: biosystematic implications 1 6(4): 1 93-200
Weather and the liability of breeding populations of the checkered white
butterfly, Pieris protodice Boisduval and Le Conte 17(1):1-16
The assumption of adaptivity in genital morphology 17(1 ):68-72
Book Review - Cracraft and Eldredge (ed.): Phylogenetic Analysis and
Paleontology 18(3):220
Book Review - Herbivores: Their interaction with secondary metabolites.
Edited by Gerald A. Rosenthal and Daniel H. Janzen 1 9( 1 ):64
Canalization of the phenotype of Nymphalis antiopa (Lepidoptera:
Nymphalidae) from subarctic and montane climates. ...... 1 9(2):82-87
Notes: A recondite breeding site for the monarch ( Danaus plexippus
Danaidae) in the montane Sierra Nevada 20(1):50-51
Notes: An interfamilial courtship (Lycaenidae - Pieridae) 20(1):54
32
AUTHOR INDEX
J. Res. Lepid.
Shapiro, Arthur M. (continued)
The biological and systematic significance of red fecal and meconial
pigments in butterflies: A review with special reference to the
Pieridae 20(2):97-102
A new record of Vanessa virginiensis "ab. ahwashtee" from northern
California (Lepidoptera: Nymphalidae) 20(3): 176-178
Notes: An aberration of Glaucopsyche lygdamus (Lycaenidae) with a
complete Scolitantidine dorsal pattern 20(4):240
Notes: A reared gynandromorph of Tatochila (Pieridae) 20(4):240-242
Two homoeotic Pieris rapae of Mexican origin (Pieridae). . . . 20(4):242-244
An apparent "intersexual" Colias eurytheme (Pieridae) 20(4):244
Taxonomic uncertainty, the biological species concept, and the Nearctic
butterflies: a reappraisal after twenty years 2 1 (4):2 1 2-2 1 8
Notes: A recessive lethal "wingless" mutation in Tatochila
(Pieridae) 22(4):262-263
Book Review - Young: Population Biology of Tropical Insects. . . 23( 1 ): 1 12
Book Review - D’Abrera: Butterflies of South America 23(2): 172-174
Polyphenism, phyletic evolution, and the structure of the Pierid
genome 23(3): 1 77- 1 96
Notes: A complex gynandromorph of Pontia daplidice
(Pieridae) 23(4):332-333
Notes: Mating confusion between a mimic and its model: Erynnis
(Hesperiidae) and Euclidea (Noctuidae) 24(l):79-80
Notes: A melanic Colias euxanthe stuebeli from Peru (Pieridae). . . 24(1 ):87
Book Review - Hafernik: Phenetics and Ecology of Hybridization in
Buckeye Butterflies (Lepidoptera: Nymphalidae) 24(1):89-91
Book Review - Opler and Krizek: Butterlies East of the Great Plains: An
Illustrated Natural History 24( 1 ):9 1-93
The impact of Pierid feeding on seed production by a native California
crucifer 24(2): 1 9 1 - 1 94
Notes: An intersubfamilial courtship (Lycaenidae) 24(2):195
Book Review - Mani: Butterflies of the Himalaya 26(l-4):283-285
Book Review - Erhardt: Wiesen und Brachland als Lebensraum fur
Schmetterlinge 26(l-4):286-287
Notes: Homosexual pseudocopulation in Eucheira socialis
(Pieridae). 27(3):262
(see also Geiger, Hansjurg and Arthur M. Shapiro)
Shapiro, Arthur M. and James D. Biggs
A hybrid Limenitis from New York 7(3): 1 49- 1 52
Shapiro, Arthur M. and Hansjurg Geiger
Electrophoretic confirmation of the species status of Pontia protodice and
P. occidentalis (Pieridae) 25(1 ):39-47
Shapiro, Arthur M., Cheryl Ann Palm and Karen L. Wcislo
The ecology and biogeography of the butterflies of the Trinity Alps and
Mount Eddy, Northern California 1 8(2):69- 1 5 1
Shapiro, Arthur M. and Adrienne R. Shapiro
The ecological associations of the butterflies of Staten Island. 1 2(2):65- 1 26
Shapiro, Irene
Courtship and mating behavior of the fiery skipper, Hylephila phylaeus
(Hesperiidae) 1 4(3): 125-141
Shaw, Kenneth C. (see Bitzer, Royce J. and Kenneth C. Shaw)
J. Res. Lepid.
AUTHOR INDEX
33
Shaw, Nirmal K., Michael C. Singer and Deborah R. Syna
Notes: Occurrence of homosexual mating pairs in a checkerspot
butterfly 24(4):393
Sheppard, Jon H.
The genus Lycaeides in the Pacific Northwest 3(l):25-36
Sheppard, P. M. and J. A. Bishop
The study of populations of Lepidoptera by capture-recapture
methods 1 2(3): 1 35- 1 44
Shields, Oakley
Callophrys ( Mitoura ) spinetorium and C. ( M .) johnsoni : Their known range,
habits, variation, and history 4(4):233-250
The butterfly fauna of a yellow pine forest community 5(2): 1 27- 1 28
Fixation of the type locality of Lycaena phlaeas hypophlaeas and a
foodplant correction 6(1):22
Hilltopping: An ecological study of summit congregation behavior of
butterflies on a southern California hill 6(2):69- 178
A partial bibliography of the world distribution and zoogeography of
butterflies 1 3(3): 1 69- 1 78
Toward a theory of butterfly migration 1 3(4):2 1 7-238
Fossil butterflies and the evolution of Lepidoptera 1 5(3): 1 32- 1 43
Studies on North American Philotes (Lycaenidae) Part V. Taxonomic and
Biological Notes (continued ) 1 6( 1 ): 1 -67
Distribution of Shijimiaeoides rita, especially S. r. rita and S. r.
coloradensis (Lycaenidae) 1 6(3): 1 62- 172
Book Review - Pyle: The Audubon Society Field Guide to North American
Butterflies 20(1):55
International Nepal Himalaya Expedition for Lepidoptera Palaearctica
(INHELP) 1977, Report No. 1: Introduction and Lycaenidae. 20(2):65-80
Book Review - Nelson and Platnick: Systematics and Biogeography:
Cladistics and Vicariance 21(3):208-209
Notes: A revised, annotated checklist of world Libytheidae. . 22(4):264-266
Book Review - Futuyma: Coevolution 23(2): 174
Notes: Revisions to the checklist of world Libytheidae 24(1):85
Notes: Ommochromes in Libytheidae 26(l-4):266
(see also Emmel, John F. and Oakley Shields)
(see also Emmel, John F., Oakley Shields and D. E. Breedlove)
Shields, Oakley and John F. Emmel
A review of carrying pair behavior and mating times in
butterflies 12(l):25-64
Shields, Oakley, John F. Emmel and D. E. Breedlove
Butterfly larval foodplant records and a procedure for reporting
foodplants 8(l):21-36
Shields, Oakley and Johnson C. Montgomery
The distribution and bionomics of arctic-alpine Lycaena phlaeas subspecies
in North America 5(4):231-242
Appendix to distribution of Lycaena phlaeas 5(4):265-266
Shields, Oakley and James R. Mori
Another Anthocharis lanceolata X A. sara hybrid 17(1 ):53-55
Shuey, John A.
Habitat associations of wetland butterflies near the Glacial Maxima in
Ohio, Indiana, and Michigan 24(2): 176-1 86
34
AUTHOR INDEX
J. Res. Lepid.
Shuey, John A. (continued)
Opinion: Comments on Clench’s temporal sequencing of Hesperiid
communities 25(3):202-206
The morpho-species concept of Euphyes dion with the description of a new
species (Hesperiidae) 27(3): 1 60- 1 72
Notes: A significant new host plant record for Pieris virginiensis
(Pieridae) 27(3):259-260
Shuey, John A. and John W. Peacock
Notes: A bilateral gynandromorph Celastrina ebenina
(Lycaenidae) 24(2): 1 95- 1 96
SlBATANI, ATUHIRO
A compilation of data on wing homoeosis in Lepidoptera 22( 1 ): 1 -46
Compilation of data on wing homoeosis on Lepidoptera: Supplement
1 22(2): 1 18-125
Simmons, Robert S. and William A. Andersen
Eighteen new or scarce butterflies for the state of Maryland. . 9(3): 1 75- 1 84
Notes on Maryland Lepidoptera No. 7, No. 8, and No. 9. .... 17(4):253-259
Notes: Notes on Maryland Lepidoptera No. 11: Six new butterflies for the
state of Maryland 23( 1 ): 1 02- 1 03
Simmons, Robert S., William A. Andersen and Philip J. Kean
Notes: Notes on Maryland No. 10: Three new butterfly records for the
state of Maryland 20(4):249
Simpson, Robert G. and David Pettus
Records of Limenitis hybrids from Colorado 1 5(3): 1 63-1 68
Sims, S. R.
Reproductive diapause in Speyeria (Lepidoptera:
Nymphalidae) 23(3):21 1-216
Singer, Michael C. (see Shaw, Nirmal K., Michael C. Singer and Deborah R.
Syna)
Singer, Michael C. and James Mallet
Notes: Moss feeding by a Satyrine butterfly 24(4):392
Slansky, Jr., Frank
Latitudinal gradients in species diversity of the new world swallowtail
butterflies 1 1 (4):20 1 -2 1 7
Small, Jr., G. B. (see Nicolay, S. S. and G. B. Small, Jr.)
Smith, Michael J. (see Tuskes, Paul M. and Michael J. Smith)
Soberon, Jorge (see Jimenez, Gabriela and Jorge Soberon)
Spade, Paul, Hamilton Tyler and John W. Brown
The biology of seven Troidine swallowtail butterflies (Papilionidae) in
Colima, Mexico 26( 1 -4): 1 3-26
Spitzer, Karel
Seasonality of the butterfly fauna in southeastern Vietnam
(Papilionidae) 22(2):126-130
(see also Novak, Ivo and Karel Spitzer)
Spitzer, Karel and Josef Jaros
Notes on Gncithmoc erodes petri fraga Diakonoff 1967 (Lepidoptera:
Tortricidae) associated with Barringtonia trees 24(2):187-190
Stamp, Nancy E.
Interactions of parasitoids and checkerspot caterpillars Euphydryas spp.
(Nymphalidae) 23( 1 ):2- 1 8
Stanford, Ray E. (see Scott, James A. and Ray E. Stanford)
Stegner, R. W. (see Urquhart, F. A. and R. W. Stegner)
J. Res. Lepid.
AUTHOR INDEX
35
Stimson, John and Linda Meyers
Inheritance and frequency of a color polymorphism in Danaus plexippus
(Lepidoptera: Danaidae) on Oahu, Hawaii 23(2): 153-1 60
Stone, Stephen E., Dean E. Swift and Richard S. Peigler
The life history of Hemileuca magnified (Saturniidae) with notes on
Hemileuca hera marcata 26(l-4):225-235
Swift, Dean E. (see Stone, Stephen E., Dean E. Swift and Richard S. Peigler)
Syna, Deborah R. (see Shaw, Nirmal K., Michael C. Singer and Deborah R.
Syna)
Tang, A. P. S. (see Urquhart, F. A. and A. P. S. Tang)
Thomas, Chris D.
Notes: An effect of the colony edge on gatekeeper butterflies Pyronia
ti tonus L. (Satyridae) 21(3):206-207
Thomas, Chris D. and Mark R. Cheverton
Notes: On the behavior and flight patterns of the neotropical butterfly,
Anartia fatima Fab (Nymphalidae) 21(3):202-204
Thomas, Chris D. and H. C. Mallorie
Notes: Oviposition records and larval foodplants of butterflies in the
Atlas Mountains of Morocco 24(l):76-79
Thorne, Fred
The distribution of an endemic butterfly Lycaena hermes 2(2): 1 43- 1 50
Habitat: Euphydryas editha wrighti 7(3): 167- 168
Threatful, David L.
A list of the butterflies and skippers of Mount Revelstoke and Glacier
National Parks, British Columbia, Canada 27(3):2 1 3-22 1
Ticehurst, Mark (see Schaefer, Paul W., William E. Wallner and Mark
Ticehurst)
Tidwell, Kenneth B. (see Callaghan, Curtis J. and Kenneth B. Tidwell)
Tidwell, Norman B. (see Callaghan, Curtis J. and Norman B. Tidwell)
Tilden, James Wilson
General characteristics of the movements of Vanessa cardui. . . . 1(1 ):43-49
The Argynnis populations of the Sand Creek area, Klamath Co., Oregon,
Part 1 1 (2): 1 09- 1 13
An analysis of the North American species of the genus
Callophrys 1(4):28 1-300
The genus Panoquina occurring in Texas 4(l):37-40
A previously unrecognized subspecies of Philotes speciosa 6(4):28 1-284
Concerning the names and status of certain North American members of
the genus Phyciodes 8(3):94-98
Comments on the Nearctic members of the genus Precis
Huebner. 9(2):101-108
Specific entities of the subgenus Icarica Nabokov (Lycaenidae). 1 2( 1 ): 1 1 -20
A name for Glaucopsyche lygdamus behrii auct., not Edwards
1862 1 2(4):2 1 3-2 1 5
Junonia and Precis. A correction 1 2(4):2 1 6
A proposed terminology for the types of diapause occurring in the order
Lepidoptera 15(1 ):33-39
Urbanus simplicus (Stoll), a new record for California 15(1 ):40
Observations of predation on Lepidoptera in Alaska 1 5(2): 1 00
Attempted mating between male monarchs 18(1 ):2
(see also Garth, John S. and James Wilson Tilden)
36
AUTHOR INDEX
J. Res. Lepid.
Tilden, James Wilson and David H. Huntzinger
The butterflies of Crater Lake National Park, Oregon 1 6(3): 1 76- 1 92
Tindale, Norman B.
A butterfly-moth (Lepidoptera Castniidae) from the Oligocene shales of
Florissant, Colorado 24( 1 ):3 1 -40
Towers, Abner A. (see Covell, Jr., Charles V., Irving L. Finkelstein and
Abner A. Towers)
Trentini, Massimo and Mario Marini
Notes: A chromosome study of Brcihmaea jciponicci Butler (Lepidoptera:
Brahmaeidae) 27(2): 1 36- 1 38
Troubridge, James T. and Kenelm W. Philip
A review of the Erebia dabanensis complex (Lepidoptera: Satyridae), with
descriptions of two new species 2 1 (2): 1 07-1 46
Tsukiyama, Hiroshi (see Chiba, Hideyuki and Hiroshi Tsukiyama)
Turner, John R. G.
A little-recognized species of Heliconius butterfly 5(2):97- 112
Correction to "A little-recognized species of Heliconius butterfly". 5(4):267
Turner, T. W. and J. R. Parnell
The identification of two species of Junonia Hubner (Lepidoptera:
Nymphalidae) : J. evarete and J. genovena in Jamaica 24(2): 1 42- 153
Tuskes, Paul M. and Michael J. Smith
The life history of Automeris zephyria (Saturniidae) 27(3):192-1 96
Tyler, Hamilton (see Spade, Paul, Hamilton Tyler and John W. Brown)
Uchida, G. (see Riotte, J. C. E. and G. Uchida)
Urquhart, F. A.
Book Review - Harris: Butterflies of Georgia 1 1 (2): 1 28
Reduction of abdominal scales of the monarch butterfly imago as a result
of cauterizing the Abd PPM of the pupa 1 1 (4):24 1 -244
Urquhart, F. A., P. Beard and R. Brownlee
A population study of a hibernal roosting colony of the monarch butterfly
{Danaus plexippus) in northern California 4(4):22 1-226
Urquhart, F. A. and R. W. Stegner
Laboratory techniques for maintaining cultures of the monarch
butterfly 5(3):129-136
Urquhart, F. A. and A. P. S. Tang
The effect of cauterizing the PPM of the pupa of the monarch
butterfly 9(3):157-167
Urquhart, F. A. and N. R. Urquhart
Announcement 17(4):268
Urquhart, Francis A., N. R. Urquhart and F. Munger
A continuosly breeding population of Danaus plexippus in southern
California 7(4):169-181
Urquhart, N. R. (see Urquhart, F. A. and N. R. Urquhart)
(see also Urquhart, Francis A., N. R. Urquhart and F. Munger)
Vawter, A. Thomas and Peter F. Brussard
Allozyme variation in a colonizing species: The cabbage butterfly Pieris
rapae (Pieridae) 22(3):204-216
Vawter, A. Thomas and Janet Wright
Genetic differentiation between subspecies of Euphydryas phaeton
(Nymphalidae: Nymphalinae) 25(l):25-29
Vesco, Jean-Pierre (see Descimon, Henri and Jean-Pierre Vesco)
J. Res . Lepid .
AUTHOR INDEX
37
Wallner, William E. (sec Schaefer, Paul W., William E. Wallner and Mark
Ticehurst)
Walter, Erich (see Johnson, John W. and Erich Walter)
Ward, P. S. (see Harmsen, R., P. D. N. Hebert and P. S. Ward)
Ward, P. S., R. Harmsen and P. D. N. Hebert
Checklist of the Macroheterocera of south-eastern Ontario. . . . 13(1 ):23-42
Wcislo, Karen L. (see Shapiro, Arthur M., Cheryl Ann Palm and Karen L.
Wcislo)
Weiss, Dalibor
A new Parnassius eversmanni race from northeast Siberia
(USSR). 9(4):2 1 5-2 1 6
Information on availability of holotypes of the described taxons at a
public institution (Rhopalocera) 16(4):208
Weiss, Stuart B. (see Murphy, Dennis D. and Stuart B. Weiss)
Wells, James F. and Richard M. Brown
Larval migration of Hyles lineata (Fab.) 13(4):246
Wescott, Richard L.
An aberrant Oregon Swallowtail, Papilio oregonius Edwards from
Oregon 18(4):255
West, David A. and Sir Cyril A. Clarke
Suppression of the black phenotype in females of the P. glaucus group
(Papilionidae) 26( 1 -4): 1 87-200
Whalley, P. E. S. (see Bradley, J. D., D. S. Fletcher and P. E. S. Whalley)
Wheye, Darryl (see Ehrlich, Paul R. and Darryl Wheye)
White, Raymond R.
Pupal mortality in the Bay checkerspot butterfly (Lepidoptera:
Nymphalidae) 25(l):52-62
Opinion: The trouble with butterflies 25(3):207-212
Whitman, Douglas W. (see Orsak, Larry J. and Douglas W. Whitman)
Wielgus, Dale (see Wielgus, Ronald S. and Dale Wielgus)
(see also Wielgus, Ronald S., Joseph R. Wielgus and Dale Wielgus)
Wielgus, Joseph R. (see Wielgus, Ronald S., Joseph R. Wielgus and Dale
Wielgus)
Wielgus, Ronald S.
The rearing of Papilio indr a kaibabensis 8(4): 177-181
A search for Speyeria nokomis coerulescens (Holland) (Nymphalidae) in
southern Arizona 1 1 (3): 187-1 94
Artificial Diet: The key to the mass rearing of Megathymus
larvae 1 3(4):27 1 -277
(see also Petterson, M. A. and R. S. Wielgus)
Wielgus, Ronald S. and Dale Wielgus
Some techniques for the rearing of Megathymus larvae 1 1(4):245-250
Wielgus, Ronald S., Joseph R. Wielgus and Dale Wielgus
Additional notes on the distribution and foodplant preferences of
Megathymus coloradensis navajo 9(3): 169- 174
Willig, Axel (see Clarke, Sir Cyril A. and Axel Willig)
Wolfe, Kirby L. (see Lemaire, Claude and Kirby L. Wolfe)
Wright, David M.
Life history and morphology of the immature stages of the Bog Copper
butterfly Lycaena epixanthe (Bsd. & Le C.) (Lepidoptera:
Lycaenidae)
Wright, Janet (see Vawter, A. Thomas and Janet Wright)
22(1):47-100
38
AUTHOR INDEX
J. Res. Lepid.
Young, Allen M.
On the evolutionary distance between Asclepiadaceous-feeding Danaida
and Apocynaceous-feeding Ithomiids 1 8(4):25 1 -254
Notes: Natural history notes on Brassolis isthmia Bates (Lepidoptera:
Nymphalidae: Brassolinae) in northeastern Costa Rica. . . . 24(4):385-392
Young, Allen M. and Alberto Muyshondt
Notes on Caligo memmon Felder and Caligo atreus Kollar (Lepidoptera:
Nymphalidae: Brassolinae) in Costa Rica and El Salvador. 24(2): 1 54- 175
FAMILY/GENUS INDEX
[Volumes 1-27, 1962-1988(89)]
Journal of Research on the Lepidoptera
Amatidae/Lygomorpha
Early stages of Lygomorpha regulus. Comstock and Henne . . . 6(4):275-280
Arctiidae/Apantesis
Genetic control of maculation and hindwing color in Apantesis phalerata.
Bacheler and Emmel 13(1 ):49-56
The chromosomes of Apantesis phalerata , A. radians , and their hybrid in
Florida populations (Arctiidae). Bacheler and Emmel . . . 1 3(3): 1 62- 1 68
Arctiidae/Argina
On the meiotic chromosomes of Argina stringa Cram (Arctiidae). Rao and
Lakshmi 17(l):51-52
Arctiidae/Dysschema
Mimicry by illusion in a sexually dimorphic, day-flying moth, Dysschema
jansonis (Lepidoptera: Arctiidae: Pericopinae). Aiello and
Brown 26( 1-4): 1 73- 1 76
Arctiidae/Parasemia
Variations of Parasemia parthenos. Brower 1 1(3): 183-1 86
Arctiidae/Uthetheisa
Variation of Uthetheisa ornatrix (Arctiidae) including a new species from
St. Croix, Virgin Islands. Pease 10(4):261-264
Attacidae/Automeris
A new species of Automeris cecrops (Attacidae: Hemileucinae).
LeMaire 18(4):236-240
Attacidae/Hemileuca
A new subspecies of Hemileuca maia from central Texas (Attacidae,
Hemileucinae). LeMaire 1 8(3):2 1 2-2 1 9
Notes: A range extension and dark phenotype of Hemileuca chinatiensis.
Bowman 24(1):85
Brahmaeidae/Brahmaea
Notes: A chromosome study of Brahmaea japonica Butler (Lepidoptera:
Brahmaeidae). Trentini and Marini 27(2): 1 36- 138
Castniidae
A butterfly-moth (Lepidoptera Castniidae) from the Oligocene shales of
Florissant, Colorado. Tindale 24( 1 ):3 1 -40
Choreutidae/Tortyra
A new Tortyra from Cocos Island, Costa Rica (Lepidoptera: Choreutidae).
Heppner 1 9(4): 1 96- 1 98
Citheronidae/Eacles
Some preliminary notes about the immature stages of Eacles oslari
(Citheronidae). Gage 1 5(3): 1 75- 1 76
COCHYLIDAE/COCHYLIS
Notes: Sex characters of the pupae of the banded moth Cochylis hospes
Wilsingham (Lepidoptera: Cochylidae). Barker 27(3):267-268
Copromorphidae/Ellabella
Revision of the Oriental and Nearctic genus Ellabella (Lepidoptera:
Copromorphidae). Heppner 23(l):50-73
39
40
FAMILY/GENUS INDEX
J. Res. Lepid.
Copromorphidae/Ellabella (continued)
On the taxonomic position of Ellabella Busck, with descriptions of the
larva and pupa of E. bayensis (Lepidoptera: Copromorphidae). De
Benedictis 23(l):74-82
Copromorphidae/Lotisma
The pupa of Lotisma trigonana and some characteristics of the
Copromorphidae (Lepidoptera). De Benedictis 24(2): 1 32- 135
COSSIDAE/COMADIA
A revision of the North American Comadia (Cossidae).
Brown 1 4(4): 1 89-2 1 2
Crambinae
Decapitation-initiated oviposition in Crambid moths. Crawford . 3(l):5-8
Primary geo-orientation in sod webworm moths. Crawford . . . 9(2):65-74
Antennal sensilla of some Crambinae. Kamm 1 6(4):20 1 -207
D AN AID AE/ D AN AUS
A population study of a hibernal roosting colony of the monarch butterfly
( Danaus plexippus) in northern California. Urquhart et al. 4(4):22 1 -226
Laboratory techniques for maintaining cultures of the monarch butterfly.
Urquhart and Stegner 5(3): 1 29- 1 36
A continuously breeding population of Danaus plexippus in southern
California. Urquhart et al 7(4): 1 69- 181
Laboratory production of the monarch butterfly, Danaus plexippus.
Munger and Harris 8(4): 1 69- 176
The effect of cauterizing the PPM of the pupa of the monarch butterfly.
Urquhart and Tang 9(3): 157-1 67
Reduction of abdominal scales of the monarch butterfly imago as a result
of cauterizing the Abd PPM of the pupa. Urquhart 1 1 (4):24 1 -244
An improved method for rearing the monarch butterfly.
Munger . 1 2(3): 1 63- 1 68
Attempted mating between male monarchs. Tilden 18(1 ):2
Notes: A recondite breeding site for the monarch {Danaus plexippus
Danaidae) in the montane Sierra Nevada. Shapiro 20(1):50-51
Inheritance and frequency of a color polymorphism in Danaus plexippus
(Lepidoptera: Danaidae) on Oahu, Hawaii. Stimson and
Meyers 23(2): 1 53- 1 60
Gelechiidae
New Canadian species of leaf-mining lepidoptera of conifers.
Freeman 4(3):209-220
Biology and immature stages of Australian Ethmiid moths (Gelechioidea).
Powell 20(4):214-234
Geometridae
Notes on the early stages of two California geometrids.
Comstock 1 (3): 1 95-200
Melanie tendencies in Phalaenid and Geometrid moths in eastern
Pennsylvania. Shapiro 3( 1 ): 1 9-24
An annotated checklist of the Missouri Geometridae.
Heitzman 1 2(3): 1 69- 179
Studies of the ova and first instar larvae of Geometridae (Ennominae). I.
Heitzman 1 3(3): 1 49- 1 56
The nomenclature in an important British check list (1972) Part 2:
Corrections of family-group names for Geometridae (lepidoptera).
Paclt 1 3(3): 1 79- 1 80
J. Res. Lepid.
FAMILY/GENUS INDEX
41
Geometridae/Antepione
Antepione thisaria and Xanthotype : A case of mimicry. Shapiro . . 4( 1 ):6- 1 1
Geometridae/Drepanulatrix
Early stages of a southern California Geometrid moth, Drepanulatrix hulsti
hulsti (Dyar). Comstock l(4):245-248
Notes on the early stages of Drepanulatrix monicaria (Guenne)
(Geometridae). Comstock 2(3):201-203
Geometrid ae/Eupithecia
California coastal Eupithecia with description of new species
(Geometridae). Leuschner 4(3): 191-1 97
Descriptions of a new species of Eupithecia and the male of E. cocoata
Pearsall (Geometridae). Heitzman and Enns 16(2):75-82
Geometrid ae/Hemistola
3 Stacks of the eggs of Hemistola hatching. McFarland 13(1 ):2 1 -22
Geometrid ae/Hypagyrtis
A new species of Hypagyrtis (Geometridae). Heitzman 13(l):43-48
Geometrid ae/Idaea
Life history studies of Idaea obfusaria (Walker). Heitzman . 1 2(3): 1 45- 1 50
Geometrid ae/ L arentiinae
Male genitalic illustrations and notes on the Larentiinae (Geometridae) of
Missouri. Heitzman and Enns 1 7(3):145-1 67
Geometrid ae/Narraga
A new species of Narraga (Geometridae, Ennominae) from Georgia, with
biological notes. Covell et al 23(2): 161-1 68
Geometrid ae/Nemoria
Variation in color and maculation in Nemoria pulcherrima from the Sierra
Nevada of California. Lepidoptera: Geometridae. Buckett and
Sears 7(2):95-98
Geometrid ae/Oenochroma
Live Geometrid (cover illustration). McFarland 14(1 ):60
Geometrid ae/Philtraea
Revision of the Nearctic genus Philtraea Hulst with notes on biology and
description of new species (Geometridae). Buckett 9(l):29-64
Geometridae/Xanthotype
Antepione thisaria and Xanthotype : A case of mimicry. Shapiro . . 4( 1 ):6- 1 1
Hesperiidae
New skipper records for Mexico. Freeman 5(l):27-28
The status of some Hesperiidae from Mexico. Freeman 6(l):59-64
The head capsule of selected Hesperioidae. Miller 9(4): 1 93-2 1 4
New butterfly records for the United States (Hesperiidae and
Libytheidae). Heitzman and Heitzman 10(4):284-286
On the distribution of some Skippers in Ontario. Riotte 1 1 (2):8 1 -82
Adult behavior and population biology of two skippers mating in
contrasting topographic sites. Scott 1 2(4): 181-1 96
Hibernal diapause of North American Papilionoidea and Hesperioidea.
Scott 1 8(3): 171 -200
Role of an ornamental plant species in extending the breeding range of a
tropical Skipper to subtropical southern Texas (Hesperiidae).
Neck 20(3): 1 29- 133
An annotated catalogue of the Skippers (Lepidoptera: Hesperiidae) named
by Roger Verity. Kudrna and Balletto 23(l):35-49
42
FAMILY/GENUS INDEX
J. Res. Lepid.
Hesperiidae (continued)
Opinion: Comments on Clench’s temporal sequencing of Hesperiid
communities. Shuey 25(3):202-206
Hesperiid ae/Acerbas
Notes: Description of the hitherto unknown female of Acerbas suttoni
Russell (Hesperiidae). Chiba 27(3):260-261
Hesperiid ae/Amblyscirtes
The habits and life history of Amblyscirtes nysa (Hesperiidae) in Missouri.
Heitzman 3(3):154-156
The life history of Amblyscirtes belli in Missouri. Heitzman . . . 4(l):75-78
The life history of Amblyscirtes linda (Hesperiidae). Heitzman and
Heitzman 8(3):99-104
Amblyscirtes "Erna" a form of Amblyscirtes aenus. Scott 15(2):92
Hesperiidae/ Atrytonopsis
Atrytonopsis hianna biology and life history in the Ozarks. Heitzman and
Heitzman 13(4):239-245
Hesperiid ae/Choranthus
A review of the West Indian "Choranthus". Miller 4(4):259-274
Hesperiid ae/Epargyreus
Supplementary notes on the distribution of Epargyreus clarus in southern
California (Hesperiidae). Miller 15(4):206-207
H esperiidae/ E r ynnis
Notes: Mating confusion between a mimic and its model: Erynnis
(Hesperiidae) and Euclidea (Noctuidae). Shapiro 24(l):79-80
Hesperiid ae/Euphyes
Euphyes dukesi. Mather 2(2): 161-1 69
Early stages of Euphyes vestris. Heitzman 3(3): 151-1 54
Euphyes dukesi - additional record. Mather 5(4):253-254
Euphyes dukesi and other Illinois Herperiidae. Irwin 8(4): 183-1 86
Further notes on Euphyes dukesi. Irwin 1 0(2): 185-188
The morpho-species concept of Euphyes dion with the description of a new
species (Hesperiidae). Shuey 27(3): 1 60- 172
Hesperiid ae/Hesperia
Hesperia metea life history studies. Heitzman and Heitzman . 8(4): 187-1 93
Geographic variation and ecology of Hesperia leonardus (Hesperiidae).
Scott and Stanford 20( 1 ): 1 8-35
Hesperiidae/Hylephila
Courtship and mating behavior of the fiery skipper, Hylephila phylaeus
(Hesperiidae). Shapiro 1 4(3): 125- 141
Hesperiid ae/Oarisma
Observations on life history of Oarisma pawesheik (Parker) 1870.
McAlpine ll(2):83-93
Hesperiid ae/Panoquina
The genus Panoquina occurring in Texas. Tilden 4(l):37-40
Hesperiid ae/Paratrytone
The distribution of Paratrytone melane and its spread into San Diego
County. Heppner 10(4):287-300
Paratrytone melane in San Luis Obispo County, California (Hesperiidae).
Miller 1 6(2): 131-1 32
Hesperiid ae/Phanus
Systematics and zoogeography of the genus Phanus (Hesperiidae).
Miller 4(2):1 15-130
J. Res. Lepid.
FAMILY/GENUS INDEX
43
Hesperiidae/Poanes
The biology of Poanes viator. Shapiro 9(2):109-123
Postglacial biogeography and distribution of Poanes viator and other marsh
butterflies. Shapiro 9(3): 1 25- 155
Evidence for two routes of post-pleistocene dispersal in Poanes viator
(Hesperiidae). Shapiro 16(3): 1 73-1 75
Hesperiidae/Polites
Three western species of Polites. Newcomer 5(4):243-247
Gynandromorphic Polites skippers (Hesperiidae). Nielsen ... 1 6(4):209-2 1 1
Hesperiidae/Polyctor
Polyctor polyctor in Mexico. Freeman 6(3): 195- 196
Hesperiidae/Problema
Observations on Problema bulenta. Krizek and Opler 25(2):146-148
Hesperiidae/Pyrrhopyginae
Illustrations and descriptions of species of some Pyrrhopyginae from
Panama (Hesperiidae). Nicolay 1 3(3): 181-1 90
Illustrations and descriptions of some species of Pyrrhopyginae from
Costa Rica, Panama and Columbia (Hesperiidae). Nicolay and
Small 19(4):230-239
Hesperiidae/Satarupa
Notes: Revisional notes on the genus Satarupa Moore (Lepidoptera:
Hesperiidae). I. New synonym of Satarupa monbeigi Oberthur. Chiba
and Tsukiyama 27(2): 1 38- 1 39
Hesperiidae/Staphylus
The complete life history of Staphylus hayhurstii. Heitzman . . 2(2): 170- 172
Hesperiidae/Urbanus
Urbanus simplicus (Stoll), a new record for California. Tilden . . . 15(1 ):40
Hesperiidae/Zera
Remarks on the genus Zera Evans in Mexico with a new record.
Freeman 5(3):181-184
Incurvariidae/Adela
Habitat: Adela bella in Florida. Heppner 13(1 ):67-72
ITHOMIIDAE
Affinities and distribution of Antillean Ithomiidae. Fox .... 2(3): 173-1 84
Libytheidae
New butterfly records for the United States (Hesperiidae and
Libytheidae). Heitzman and Heitzman 10(4):284-286
Notes: A revised, annotated checklist of world Libytheidae.
Shields 22(4):264-266
Notes: Revisions to the checklist of world Libytheidae. Shields . . 24(1):85
Notes: Ommochromes in Libytheidae. Shields 26(l-4):266
Limacodidae/Sphinx
The identity of Sphinx brunnus Cramer and the taxonomic position of
Acharia Huebner (Lepidoptera: Limacodidae). Becker and
Miller 26(l-4):219-224
Lycaenidae
A synopsis of the west Indian Lycaenidae, with remarks on their
zoogeography. Clench 2(4):247-270
A possible new hybrid copper. Crowe 8(2):5 1 -52
Pupal sound production of some Lycaenidae.
Hoegh-Guldberg 10(2): 1 27-1 47
Editorial: Extinction of the British Large Blue Butterfly. Mattoni. 1 8( 1 ): 1 ,3
44
FAMILY/GENUS INDEX
J . Res. Lepid.
Lycaenidae (continued)
Notes: An interfamilial courtship (Lycaenidae - Pieridae). Shapiro. 20(1):54
International Nepal Himalaya Expedition for Lepidoptera Palaearctica
(INHELP) 1977, Report No. 1: Introduction and Lycaenidae.
Shields 20(2):65-80
Notes: Three intersubfamilial matings in nature (Lycaenidae).
Mattoni 24(l):86-87
Notes: An intersubfamilial courtship (Lycaenidae). Shapiro .... 24(2):195
Notes: Aberrant Polymmatinae (Lycaenidae) from Ohio and Florida.
Calhoun 26(l-4):264-266
A survey of the last instar larvae of the Lycaenidae (Lepidoptera) of
California. Ballmer and Pratt 27(1 ): 1 -8 1
Lycaenidae/ Agriades
Notes on a little known ecologically displaced blue, Agriades pyrenaicus
ergane Higgins (Lycaenidae). Pljushtch 2?(2):129-134
Lycaenidae/ Apodemia
Apodemia palmerii (Lycaenidae: Riodininae): Misapplication of names, two
new subspecies and a new allied species. Austin 26( 1 -4): 1 25- 1 40
Lycaenidae/Brephidium
Habitat: Brephidium pseudo fea (Lycaenidae). Heppner 13(2):99-100
Notes: Lateral perching in Brephidium exilis (Boisduval) (Lycaenidae) in
Texas. Johnson 23( 1 ): 1 04- 1 06
Notes: Notes on the biology of Brephidium exilis (Boisduval) (Lycaenidae).
Haeger 26(l-4):254-255
Lycaenidae/Callophrys
An analysis of the North American species of the genus Callophrys.
Tilden l(4):281-300
Callophrys (Lycaenidae) from the northwest. Clench 2(2): 151-1 60
A new subspecies of Callophrys dumetorum from Washington and Oregon.
Gorelick 7(2):99-104
Larva and habitat of Callophrys fotis bayensis. Brown 8(2):49-50
Intergradation between Callophrys dumetorum oregonensis and Callophrys
dumetorum af finis in northwestern U.S. (Lycaenidae). Scott and
Justice 20(2):8 1-85
Chromatic polymorphism in Callophrys mossii bayensis larvae
(Lycaenidae): Spectral characterization, short-term color shifts, and
natural morph frequencies. Orsak and Whitman 25(3): 1 88-20 1
Lycaenidae/Celastrina
Notes: Celastrina ladon (Lycaenidae) female ovipositing on Sambuscus
canadensis , a plant unsuitable for larval development. Oliver . 20(1):54
Notes: A bilateral gynandromorph Celastrina ebenina (Lycaenidae). Shuey
and Peacock 24(2):195-196
Lycaenidae/Chlorostrymon
Notes on the life history and Baja California distribution of
Chlorostrymon simaethis sarita (Skinner) (Lepidoptera: Lycaenidae).
Brown 20(4):207-213
Lycaenidae/Collophrys
Callophrys (Mitoura) spinetorium and C. (M.) johnsoni : Their known range,
habits, variation, and history. Shields 4(4):233-250
J. Res . Lepid.
FAMILY/GENUS INDEX
45
Lycaenidae/Euphilotes
Conservation and management of the endangered Smith’s Blue Butterfly,
Euphilotes enoptes smithi (Lepidoptera: Lycaenidae).
Arnold 22(2): 1 35- 1 53
The phenetics and comparative biology of Euphilotes enoptes (Boisduval)
(Lycaenidae) from the San Bernadino Mountains. Pratt and
Ballmer . . 25(2): 1 2 1 - 1 35
The Euphilotes battoides complex: recognition of a species and description
of a new subspecies. Mattoni 27(3): 173-185
Lycaenidae/Everes
Morphology of the immature stages of Everes comyntas Godart. Lawrence
and Downey 5(2):61-96
Lycaenidae/Glaucopsyche
Habitat: General type locality, Glaucopsyche lygdamus xerces, Plebejus
icariodes pheres. Hovanitz 7(2): 126
A name for Glaucopsyche lygdamus behrii auct., not Edwards 1862.
Tilden 1 2(4):2 1 3-2 1 5
Notes: An aberration of Glaucopsyche lygdamus (Lycaenidae) with a
complete Scolitantidine dorsal pattern. Shapiro 20(4):240
Lycaenidae/Hypaurotis
Notes: Records of Hypaurotis crysalus (Ed) (Lycaenidae) from western
Mexico. Brown 27(2): 135
Lycaenidae/Icaricia
Specific entities of the subgenus Icarica Nabokov (Lycaenidae).
Tilden 12(1):1 1-20
Lycaenidae/ Juditha
Notes on the immature biology of two Myrmecophilous Lycaenidae:
Juditha molpe (Riodininae) and Panthiades bitias (Lycaeninae).
Callaghan 20(l):36-42
Lycaenidae/Lycaeides
The genus Lycaeides in the Pacific Northwest. Sheppard 3(l):25-36
Plebeian courtship revisited: Studies on the female-produced male
behavior- eliciting signals in Lycaeides idas courtship (Lycaenidae).
Pellmyr 2 1 (3): 1 47- 1 57
Lycaenidae/Lycaena
The distribution of an endemic butterfly Lycaena hermes.
Thorne 2(2):143-150
The synonymy, variability and biology of Lycaena nivalis.
Newcomer 2(4):27 1-280
A gynandromorph of Lycaena gorgon. Opler 5(4):230
The distribution and bionomics of arctic-alpine Lycaena phlaeas subspecies
in North America. Shields and Montgomery 5(4):231-242
Appendix to the distribution of Lycaena phlaeas. Shields and
Montgomery 5(4):265-266
Fixation of the type locality of Lycaena phlaeas hypophlaeas and a
foodplant correction. Shields 6(1):22
Biochemical studies of the larval hosts of two species of Lycaena Fabricius
(Lycaenidae). Ferris 17(1 ):27-32
The identity of the Rocky Mountain Lycaena dorcas-helloides complex.
Scott 1 7(l):40-50
Habitat: Lycaena heteronea clara (Lepidoptera: Lycaenidae). Orsak and
Miller 1 7(3):204-206
46
FAMILY/GENUS INDEX
J. Res. Lepid.
Lycaenidae/Lycaena (continued)
Geographic variation in Lycaena xanthoides. Scott 18(1 ):50-59
Life history and morphology of the immature stages of the Bog Copper
butterfly Lycaena epixanthe (Bsd. & Le C.) (Lepidoptera: Lycaenidae).
Wright 22(1):47-100
A new subspecies of Lycaena editha (Mead) (Lycaenidae) from Nevada.
Austin 23(l):83-88
Notes: A replacement name for Lycaena editha nevadensis Austin
(Lycaenidae). Austin 27(3):266
Lycaenidae/Mitoura
A new species of Mitoura Scudder from southern California (Lepidoptera:
Lycaenidae). Brown 21(4):245-254
Lycaenidae/Panthiades
Notes on the immature biology of two Myrmecophilous Lycaenidae:
Juditha molpe (Riodininae) and Panthiades bitias (Lycaeninae).
Callaghan 20(l):36-42
Lycaenidae/Philotes
Techniques in the study of population structure in Philotes sonorensis.
Mattoni and Seiger l(4):237-244
Distribution and pattern of variation in Philotes rita. Mattoni . 4(2):8 1-101
Natural habitats - Philotes sonorensis. Hovanitz 6(3): 199-202
A previously unrecognized subspecies of Philotes speciosa.
Tilden 6(4):28 1-284
Studies on North American Philotes (Lycaenidae) Part V. Taxonomic and
Biological Notes (continued). Shields 16(1 ): 1 -67
Notes: A melanic aberration of Philotes sonorensis (Lycaenidae) from
California. Priestaf 27(3):265-266
Lycaenidae/ Plebejinae
Two new forms of Plebejinae from Wyoming. Ferris 8(3):9 1 -93
Lycaenidae/Plebejus
Habitat: Specific type locality, Plebejus icariodes missionensis.
Hovanitz 7(2): 1 22
Habitat: General type locality, Glaucopsyche lygdamus xerces , Plebejus
icariodes pheres. Hovanitz 7(2): 126
The biology of Plebejus ( Icaricia ) shasta in the Western United States
(Lycaenidae). Emmel and Shields 1 7(2): 1 29- 1 40
Lycaenidae/Polyommatus
The role of intra- and interspecific male:male interactions in Polyommatus
icarus Rott. and some other species of blues (Lycaenidae).
Lundgren 16(4):249-264
Lycaenidae/Riodininae
A study of isolating mechanisms among Neotropical butterflies of the
subfamily Riodininae. Callaghan 2 1 (3): 1 59- 1 76
Notes on the biology of three Riodinine species: Nymphidium lisimon
attenuatum , Phaenochitonia sagaris satnius , and Metacharis ptolomaeus
(Lyceanidae: Riodininae). Callaghan 27(2): 1 09- 114
Lycaenidae/Sandia
Notes: Laboratory rearing of Sandia xami xami (Lycaenidae: Eumaeini).
Jimenez and Soberson 27(3):268-271
Lycaenidae/Satyrium
Life history of Satyrium sylvinas dryope. Emmel and Emmel . 7(2): 123- 125
J . Res. Lepid.
FAMILY/GENUS INDEX
47
Lycaenidae/Satyrium (continued)
Polymorphism in Satyrium calanus (Huebner) from Wyoming and Colorado
(Lepidoptera: Lycaenidae: Theclinae). Ferris 2 1(3): 188-1 94
A second phenotype of Satyrium calanus (Heubner) from Wyoming
(Lepidoptera: Theclinae). Ferris 23(4):297-302
Notes: The origin of Satyrium calanus albidus. Scott 23(4):334
Lycaenidae/Scolitantidini
The Scolitantidini I: Two new genera and a generic rearrangement
(Lycaenidae). Mattoni 16(4):223-242
Lycaenidae/Shijimiaeoides
Distribution of Shijimiaeoides rita, especially S. r. rita and S. r.
color adensis (Lycaenidae). Shields 16(3): 162- 172
Lycaenidae/Stalachtis
Notes on the biology of Stalachtis susanna (Lycaenidae: Riodininae) with a
discussion of Riodinine larval strategies. Callaghan 24(3):258-263
Lycaenidae/Strymoninae
The heathii- white banding aberration in the Strymoninae (Lycaenidae).
Fisher . 1 5(3): 1 77- 1 8 1
Lycaenidae/Theclinae
Parallel albinism in two Theclines (Lycaenidae). Holland 2 1 (3): 158
L YC AENID AE/T OMARES
Notes: Notes on Tomares mauretanicus (Lycaenidae) in Morocco.
Courtney 21(3):205-206
Lycaenidae/Turanana
The Scolitantidini II. The World’s smallest butterfly? Notes on Turanana ,
and a new genus and species from Afghanistan (Lycaenidae).
Mattoni 18(4):256-264
Lymantriidae
Response to J. C. E. Riotte’s Review of the Lymantriid fascicle.
Ferguson 17(4):265-267
Lymantriidae/Gynaephora
Gynaephora rossii (Curtis) on Mt. Katahdin, Maine, and Mt. Daisetsu,
Japan, and comparisons to records for populations from the Arctic
(Lymantriidae). Schaefer and Castrovillo 1 8(4):24 1 -250
L YM ANTRIID AE/ L YMANTRIA
A black-backed larval mutant of Lymantria dispar (L.) (Lepidoptera:
Lymantriidae) in Japan. Schaefer and Furuta 1 8(3): 1 67- 1 70
Notes: Incidence of the black backed larval mutant of Lymantria dispar
(L) (Lepidoptera: Lymantriidae) in Ukrainian SSR. Schaefer et
al . . 23(1):103-104
Lymantriid ae/Orgyia
Investigation of selected species of the genus Orgyia (Lymantriidae) using
isoelectrofocusing in thin layer polyacrylamide gel. Chua et
al 15(4):2 15-224
Megathymidae
Type localities of the Megathymidae. Freeman 2(2): 137-141
The effects of pH on the distribution of the Megathymidae.
Freeman 3( 1 ): 1 -4
Early work on the Megathymidae. Comstock 14(2):98-99
Megathymidae/Agathymus
Bionomics of Agathymus (Megathymidae). Roever 3(2): 1 03- 1 20
Larval habits of Agathymus mariae. Freeman 3(3): 145- 147
48
FAMILY/GENUS INDEX
J. Res. Lepid.
Megathymidae/Agathymus (continued)
Instar determination of Agathymus larvae. Roever 3(3): 1 48- 1 50
Speciation in the Agathymus (Megathymidae). Freeman 5(4):209-214
Megathymidae/Megathymus
Additional notes on the distribution and foodplant preferences of
Megathymus coloradensis navajo. Wielgus et al 9(3): 169- 174
Some techniques for the rearing of Megathymus larvae. Wielgus and
Wielgus 1 1(4):245-250
Acceptance of artificial diet by Megathymus streckeri (Skinner). Petterson
and Wielgus 1 2(4): 1 97-1 98
Artificial Diet: The key to the mass rearing of Megathymus larvae.
Wielgus 1 3(4):27 1-277
N EPTICULID AE/NEPTICULA
A new species of Nepticula on bur oak in Ontario (Nepticulidae).
Freeman 6( 1 ): 1 9-2 1
Noctuidae
Karyotypes of some Indian Noctuid moths (Lepidoptera). Mohanty and
Nayak 22(4):238-248
A new genus and species from the southwestern United States (Noctuidae:
Acontiinae). Brown 25(2): 1 36- 1 45
N OCTUIDAE/ ACRONICTA
The larva of Acronicta spinigera Guenee (Noctuidae). McCabe. 1 7(3): 173-1 79
N OCTUIDAE/ ACRONICTINAE
New species and new nomenclature in the American Acronictinae
(Lepidoptera: Noctuidae). Ferguson 26( 1 -4):20 1-218
Noctuid ae/Agrotis
Diel egg-laying activity of Agrotis exclamationis (Noctuidae).
Persson 10(4):225-260
N OCTUIDAE/ A NNAPHILA
The Annaphila astrologa complex, with descriptions of three new species.
Sala 2(4):289-301
Collecting of Annaphila spila with notes on the "crimson winged" group of
the genus. Buckett 2(4):303-304
Review of the depicta group of the genus Annaphila. Buckett and
Bauer 3(2):95-101
Studies in life histories of North American Lepidoptera, California
Annaphilas. Comstock and Henne 3(3): 173-191
The Noctuid moth Annaphila baueri with notes on its habits.
Buckett 4(3): 185-1 89
A reevaluation of Annaphila casta (Noctuidae). Buckett ..... 4(3):199-204
Studies in the life histories of North American Lepidoptera, California
Annaphila II. Comstock and Henne 5( 1 ): 1 5-26
Rediscovery of Annaphila casta Hy. Edw. in California (Noctuidae).
Buckett 5(l):37-38
Discovery of a larval hostplant for Annaphila lithosina with notes on the
species (Noctuidae: Amphipyrinae). Buckett 5(4):262-264
Life history studies on the lithosina-miona-casta complex in the genus
Annaphila. Henne 6(4):249-256
Studies in the life histories of North American Lepidoptera, California
Annaphila III. Comstock and Henne 6(4):257-262
J. Res. Lepid .
FAMILY/GENUS INDEX
49
N octuidae/Basilodes
A new species of Basilodes from the southwestern United States
(Noctuidae). Hogue 4(4):275-280
Noctuidae/Behrensia
Revision of the North American genus Behrensia. Buckett . . . 3(3): 1 29- 1 44
N octuid ae/Bellura
Taxonomic and biological notes on Bellura gortynoides Walker
(Noctuiidae). Heitzman and Habeck 18(4):228-235
N OCTUID AE/C ALLOGRAMA
Studies on the excretory system of the fully grown larva of Callograma
/ estiva Donov. (Noctuidae). Mohamed and Murad 16(4):2 13-221
N octuid ae/Euclidea
Notes: Mating confusion between a mimic and its model: Erynnis
(Hesperiidae) and Euclidea (Noctuidae). Shapiro 24(l):79-80
N octuid ae/Catocala
Field studies of Catocala behavior. Keiper 7(2): 1 13-121
Two new California Catocala subspecies (Noctuidae).
Johnson 20(4):245-248
The immature stages of Catocala erichi Brower (Lepidoptera: Noctuidae).
Johnson and Walter 23(3):23 1 -235
The immature stages of six California Catocala (Lepidoptera: Noctuidae).
Johnson 23(4):303-327
N octuid ae/Euxoa
The little known moth Euxoa sculptilis (Harvey) in Arizona, with
descriptions, illustrations, and notes on Euxoa violaris (Grote and
Robinson) (Noctuidae). Buckett 5(4):255-261
Species in the genera Polia and Euxoa. Buckett 7(2):87-94
N octuid ae/Exyra
Daytime vision by the moth, Exyra ridingsi. Keiper 7(2): 131-1 32
N OCTUID AE/FARONTA
A new species of armyworm - genus Faronta. Buckett 6(4):268-274
Noctuidae/Feralia
A new species of Feralia. Buckett 6(1):43-51
N OCTUID AE/HELIOSEA
Identity of Heliosea celeris melicleptroides. Buckett 4(l):79-80
N octuidae/Lithophane
Rediscovery and redescription of the moth Lithophane vanduzeei (Barnes).
Buckett and Leuschner 4(4):281-286
N octuidae/Loxagrotis
Identity of the moth Loxagrotis pampolycala from the southwestern US
and Mexico (Noctuidae). Buckett 8(3): 118-1 28
N octuidae/Luperina
The little known species Luperina venosa. Buckett and
Lounibos 4(4):227-232
N octuidae/N ephelodes
A new species of Nephelodes Guenee for the Great Basin.
Buckett 1 1(4):260-268
N octuid ae/Oncocnemis
A new species of Oncocnemis from the western United States (Noctuidae:
Cuculliinae). Buckett and Bauer 5(4):197-208
Identity of the moth Oncocnemis semicollaris J.B. Smith.
Buckett 10(3):248-254
50
FAMILY/GENUS INDEX
J. Res. Lepid.
N octuidae/ Petaluma
Petaluma , a new genus. Buckett and Bauer 3(3): 1 93- 1 96
Homonymy of the "new genus” Petaluma and proposal of the name
Petalumaria. Buckett and Bauer 6(1):52
N OCTUIDAE/ P OLIA
A new species of Polia Ochsenheimer from California and notes on Polia
discalsis (Grote) (Noctuidae: Hadeninae). Buckett and
Bauer 5(4):221-228
Species in the genera Polia and Euxoa. Buckett 7(2):87-94
N OCTUIDAE/SCOTOGRAMMA
Notes on the first-instar and two parasites of the clover cutworm
Scotogramma trifollii (Noctuidae: Hadeninae). Santiago-Alvarez and
Federici 17(4):226-230
N octuidae/Sesamia
Digestive enzymes of sugarcane pink borer, Sesamia inferens Walker
(Noctuidae). Agarwal 1 5(3): 1 53- 1 62
N octuidae/Stretchia
Identity of the moth "Stretchia" behrensiana with new synonymy
(Noctuidae). Buckett 7(l):57-63
N octuidae/Sunira
Polymorphism in Sunira bicolorago (Noctuidae). Shapiro 4( 1 ): 1 -5
N OCTUIDAE/T ARACHE
On the supernumerary chromosomes of Tarache tropica Guen.
(Lepidoptera: Noctuidae). Mohanty and Nayak ......... 20(3): 1 70- 173
N OCTUIDAE/X YLOMIGES
Description of a new species of Xylomiges from California.
Buckett 6(l):23-30
Nolidae
Retention of cast head capsules by some Nolid immatures in four Old
World countries. McFarland 17(4):209-217
Nymphalidae
Tertiary Nymphalid butterflies and some phylogenetic aspects of
systematic lepidopterology. Nekrutenko 4(3): 1 49- 158
Comparitive speciation in two butterfly families: Pieridae and
Nymphalidae. Petersen 5(2): 113-1 26
An interfamilial courtship (Nymphalidae, Pieridae). Shapiro. 1 1(3): 1 97-1 98
Notes: Occurrence of homosexual mating pairs in a checkerspot butterfly.
Shaw etal 24(4):393
Stratification of fruit-feeding nymphalid butterflies in a Costa Rican
rainforest. DeVries 26(l-4):98-108
Notes: Courtship of a model (Nymphalidae: Adelpha) by its
probableBatesian mimic (Nymphalidae: Limenitis).
Porter 26(l-4):255-256
Nymphalidae/ Adelpha
A new species of Adelpha (Nymphalidae) from Parque Nacional Braulio
Carrillo, Costa Rica. DeVries and Gamboa 20(2): 1 23- 1 26
Nymphalidae/ A graulis
The generic, specific and lower category names of the Nearctic butterflies.
Part 8 - the genus Agraulis. McHenry 7(2): 127- 1 30
Notes: A homoeotic Agraulis vanillae incarnata (Nymphalidae).
Dimock 23(4):332
J. Res. Lepid.
FAMILY/GENUS INDEX
51
Nymphalidae/Agraulis (continued)
Hidden genetic variation in Agraulis vanillae incarnata (Nymphalidae).
Dimock and Mattoni 25( 1 ): 1 - 1 4
Nymphalidae/Anartia
The Neotropical Nymphalid butterfly, Anartia amalthea.
Fosdick 1 1(2):65-80
Notes: On the behavior and flight patterns of the neotropical butterfly,
Anartia fatima Fab (Nymphalidae). Thomas and
Cheverton 21(3):202-204
Nymphalid ae/Argynnids
The generic, specific and lower category names of the Nearctic butterflies.
Part 3 - Argynnids. McHenry 3(4):231-268
N ymphalidae/ Argynnis
Variations in the silvering of Argynnis (Speyeria) callippe in the interior
mountain area of south central California. Sette 1(1 ):3-20
Argynnis and Speyeria. Hovanitz 1(1 ):95-96
The Argynnis populations of the Sand Creek area, Klamath Co., Oregon,
Part 1. Tilden 1(2):109-1 13
Geographical distribution and variation of the genus Argynnis. I.
Introduction. II. Argynnis idalia. Hovanitz 1 (2): 1 17-123
Geographical distribution and variation of the genus Argynnis. III.
Argynnis diana. Hovanitz 1(3):20 1-208
Generic or subgeneric names closely related to Argynnis.
McHenry 2(3):229-239
Ecological color variation in some Argynnis of the western United States.
Hovanitz 6(3):197-198
Habitat - Argynnis callippe laurina. Hovanitz 7(1 ):50
Habitat - Argynnis nokomis. Hovanitz 8(1):20
On the Gunder collection of Argynnids. Grey 8(2):55-64
Habitat - Argynnis adiaste. Hovanitz 9(3): 1 68
N YMPHALIDAE/ A STEROCAMPA
The biology and morphology of the immature stages of Asterocampa idyja
argus (Bates) (Lepidoptera: Nymphalidae). Friedlander . . 24(3):209-225
Egg mass design relative to surface-parasitizing parasitoids, with notes on
Asterocampa clyton (Lepidoptera: Nymphalidae).
Friedlander 24(3):250-257
Taxonomy, phylogeny and biogeography of Asterocampa Rober 1916
(Lepidoptera: Nymphalidae: Apaturinae). Friedlander . . . 25(4):215-338
Male mate-locating behavior in the desert hackberry butterfly,
Asterocampa leilia (Nymphalidae). Rutowski and Gilchrist . 26( 1 -4): 1-12
Nymphalid ae/Boloria
A new subspecies of Boloria eunomia from Wyoming. Ferris and
Groothuis 9(4):243-248
Polymorphism in two species of Alaskan Boloria. Ferris .... 1 1(4):255-259
A critique of the genus Boloria (Nymphalidae) as represented in "The
Butterflies of North America", with corrections, additions and a key to
species. Pike 1 8(3): 153- 1 66
Nymphalidae/Brassolis
Notes: Natural history notes on Brassolis isthmis Bates (Lepidoptera:
Nymphalidae: Brassolinae) in northeastern Costa Rica.
Young 24(4):385-392
52
FAMILY/GENUS INDEX
J. Res. Lepid.
N YMPHALID AE/B YBLIA
Note on the chromosomes of Byblia ilithyia (Drury) (Nymphalidae). Murty
and Rao 1 5(3): 1 29- 1 3 1
N YMPHALID AE/CALIGO
Notes on Caligo memmon Felder and Caligo atreus Kollar (Lepidoptera:
Nymphalidae: Brassolinae) in Costa Rica and El Salvador. Young and
Muyshondt 24(2): 1 54- 1 75
N YMPHALID AE/CHARIDRY AS
Chciridryas flavula Barnes and McDunnough (Nymphalidae): A question of
identity. Ferris and Fisher 1 6(3): 133-1 40
N YMPHALID AE/CHLOSYNE
History of scientific study on a larval color polymorphism in the genus
Chlosyne (Nymphalidae). Neck 14(1 ):4 1 -48
Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) Nymphalidae
II. Additional larval food plant data. Neck 16(2):69-74
Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) (Nymphalidae)
III. Adult resources. Neck 1 6(3): 1 47- 1 54
N YMPHALID AE/CLOSSIANA
Notes: Field notes on Clossiana improba harryi Ferris (Lepidoptera:
Nymphalidae). Ferris 25( 1 ):7 1-72
N ymphalidae/ Cynthia
Type locality and habitat - Cynthia annabella. Dimock 10(4):265-266
Natural and laboratory occurrence of "Elymi" phenotypes in Cynthia
cardui (Nymphalidae). Shapiro 13(1 ):5 7-62
Nymphalidae/Dione
Notes: Dione moneta poeyii Butler (1873) in New Mexico (Lepidoptera:
Nymphalidae). McCaffrey 23( 1 ): 1 06- 107
N YMPHALID AE/DRYADULA
The generic, specific and lower category names of the Nearctic butterflies.
Part 7 - the genus Dryadula. McHenry 7(2): 112
N YMPHALID AE/DRYAS
The generic, specific and lower category names of the Nearctic butterflies.
Part 6 - the genus Dryas. McHenry 6(4):263-265
N YMPHALID AE/ERGOLIS
Chromosome numbers in two species of Ergolis (Lepidptera: Nymphalidae).
Murty and Rao 15(1 ):23-26
N YMPHALID AE/EUPHYDRYAS
Habitat: Euphydryas editha wrighti. Thorne 7(3): 167- 168
A new subspecies of Euphydryas from Wyoming (Nymphalidae).
Ferris 9(l):17-20
Euphydryas editha gunnisonensis, a new subspecies from western Colorado.
Brown 9( 1 ):2 1 -23
Two new subspecies of Euphydryas chalcedona from the Mojave desert of
Southern California. Emmel and Emmel 1 1 (3): 141-1 46
A new subspecies of Euphydryas editha from the Channel Islands of
California. Emmel and Emmel 1 3(2): 131-1 36
A survey of valvae of Euphydryas chalcedona , E. c. colon , and E. c. anica.
Scott ’ 17(4):245-252
On the status of Euphydryas editha baroni with a range extension of E.
editha luestherae. Murphy 2 1(3): 194- 198
Biosystematics of the Euphydryas of the central Great Basin with the
description of a new subspecies. Murphy and Ehrlich . . . 22(4):254-261
J . Res. Lepid.
FAMILY/GENUS INDEX
53
Nymphalidae/Euphydryas (continued)
Interactions of parasitoids and checkerspot caterpillars Euphydryas spp.
(Nymphalidae). Stamp 23( 1 ):2- 1 8
Some observations on spatial distribution in a montane population of
Euphydryas editha. Ehrlich and Wheye 23(2): 143-1 52
Genetic differentiation between subspecies of Euphydryas phaeton
(Nymphalidae: Nymphalinae). Vawter and Wright 25(l):25-29
Pupal mortality in the Bay checkerspot butterfly (Lepidoptera:
Nymphalidae). White 25(l):52-62
Euphydryas anicia and E. chalcedona in Idaho (Lepidoptera: Nymphalidae).
Ferris 26(1 -4): 109-1 15
Notes: A bibliography of Euphydryas. Murphy and Weiss . 26(l-4):256-264
N YMPHALID AE/EUPTOIETA
The generic, specific and lower category names of the Nearctic butterflies.
Part 4 - the genus Euptoieta. McHenry 4(3):205-208
A dwarf form of Euptoieta claudia. Rahn 11(3): 174
Nymphalidae/Hamadryas
Evidence for lack of territoriality in two species of Hamadryas.
Ross 2(4):24 1-246
N YMPHALID AE/HELICONIUS
A little-recognized species of Heliconius butterfly. Turner . . . 5(2):97- 112
Correction to "A little-recognized species of Heliconius butterfly".
Turner 5(4):267
The generic, specific and lower category names of the Nearctic butterflies.
Part 5 - the genus Heliconius. McHenry 6(l):65-68
Heliconius cydno in Venezuela with descriptions for two new subspecies.
Masters 10(4):267-272
Concerning Heliconius cydno aberration "larseni" Niepelt.
Masters 1 1(4):251-254
Semispecies relationships between Heliconius erato cyrbia Godt. and H.
himera Hew. in southwestern Ecuador. Descimon and de
Maeght 22(4):229-237
Evidence for host plant preferences in Heliconius erato phyllis from
southern Brazil (Nymphalidae). Menna-Barreto and Araujo . 24(l):41-46
Correlations of ultrastructure and pigmentation suggest how genes control
development of wing scales of Heliconius butterflies. Gilbert et
al 26( 1 -4): 141-1 60
N YMPHALID AE/J UNONIA
Junonia and Precis. A correction. Tilden 1 2(4):2 1 6
The identification of two species of Junonia Hubner (Lepidoptera:
Nymphalidae) : J. evarete and J. genovena in Jamaica. Turner and
Parnell 24(2): 1 42- 1 53
N YMPHALID AE/LlMENITIS
A hybrid Limenitis from New York. Shapiro and Biggs 7(3): 1 49- 152
A new subspecies of Limenitis archippus. Herlan 9(4):2 17-222
On the occurrence of Limenitis archippus X L. lorquini hybrids. Perkins
and Gage 9(4):223-226
A bilateral gynandromorph of Limenitis weidemeyerii lati fascia
(Nymphalidae). Perkins and Perkins 1 1 (3): 1 95- 1 96
Limenitis weidemeyerii angusti fascia X L. astyanax arizonensis = (?) ab.
doudoroffi (G under) 1934. Perkins and Garth 1 1(4):229-234
54
FAMILY/GENUS INDEX
J. Res. Lepid.
Nymphalidae/Limenitis (continued)
The correct name for the subspecies of Limenitis weidemeyerii occurring in
Arizona (Nymphalidae). Dos Passos 1 2( 1 ):2 1 -24
Records of Limenitis hybrids from Colorado. Simpson and
Pettus 1 5(3): 1 63- 1 68
A new Limenitis weidemeyerii W. H. Edwards from southeastern Arizona
(Nymphalidae). Austin and Mullins 22(4):225-228
Stubby-winged mutants of Limenitis (Nymphalidae) - Their occurrence in
relatrion to photoperiod and population size. Platt 23(3):2 1 7-230
"Black-light" induction of photoperiod-controlled diapause responses of
the viceroy butterfly, Limenitis archippus (Nymphalidae). Platt and
Harrison 26( 1 -4): 1 77- 1 86
Nymphalidae/Nymphalis
Why do California tortoiseshells migrate? Shapiro 14(2):93-97
Canalization of the phenotype of Nymphalis antiopa (Lepidoptera:
Nymphalidae) from subarctic and montane climates. Shapiro. 19(2):82-87
N YMPHALID AE/PHYCIODES
Concerning the names and status of certain North American members of
the genus Phyciodes. Tilden 8(3):94-98
Early stages and biology of Phyciodes orseis. Scott 12(4):236-242
Early stages of Phyciodes pallida , P. orseis , and P. mylitta (Nymphalidae).
Scott 14(2):84
Field study of Phyciodes batesii (Reakirt) and P. tharos (Drury) from a site
in the Black Hills, South Dakota (Lepidoptera: Nymphalidae:
Melitaeinae). Ferris 20(4):235-239
N YMPHALID AE/POLYGONIA
A review of Polygonia progne (oreas) and P. gracilis (zephyrus)
(Nymphalidae) including a new subspecies from the southern Rocky
Mountains. Scott 23(3): 1 97-2 1 0
N YMPHALID AE/PRECIS
Controlled environment experiments with Precis octavia Cram.
McLeod 7(1):1-18
Controlled environment experiments with Precis octavia Cram.
McLeod 8(2):53-54
Comments on the Nearctic members of the genus Precis Huebner.
Tilden 9(2):101-108
Junonia and Precis. A correction. Tilden 1 2(4):2 1 6
Variability of courtship of the buckeye butterfly, Precis coenia
(Nymphalidae). Scott 1 4(3): 1 42- 1 47
Development of the wing margin in Precis coenia (Lepidoptera:
Nymphalidae). Dohrmann and Nijhout 27(3): 151-1 59
N ymphalidae/Speyeria
Argynnis and Speyeria. Hovanitz 1(1 ):95-96
Speyeria cybele in Mississippi (Argynninae: Argynnis). Mather. 5(4):252-253
A search for Speyeria nokomis coerulescens (Holland) (Nymphalidae) in
southern Arizona. Wielgus 1 1(3): 187-1 94
Speyeria id alia. McCabe 16(1 ):68
Early stages of Speyeria nokomis (Nymphalidae). Scott and
Mattoon 20( 1 ): 1 2- 1 5
An apparent Interspecific Fx Hybrid Speyeria (Nymphalidae).
Scott 20(3):174-175
J. Res. Lepid.
FAMILY/GENUS INDEX
55
Nymphalidae/Speyeria (continued)
The colonization of violets and Speyeria butterflies on the ash-pumice
fields deposited by Cascadian volcanoes. Hammond 20(3): 1 79- 191
Speyeria atlantis phenotypes in the southern Rocky Mountains
(Lepidoptera: Nymphalidae: Argynninae). Ferris 22(2): 101-1 14
The decline and extinction of Speyeria populations resulting from human
environmental disturbances (Nymphalidae: Argynninae). Hammond
and McCorkle 22(4):2 1 7-224
Reproductive diapause in Speyeria (Lepidoptera: Nymphalidae).
Sims 23(3):21 1-216
Opinion: A rebuttal to the Arnold classification of Speyeria callippe
(Nymphalidae) and defense of the subspecies concept.
Hammond 24(3):197-208
N YMPHALIDAE/ V ANESSA
General characteristics of the movements of Vanessa cardui.
Tilden l(l):43-49
Territorial behavior of the red admiral, Vanessa atalanta (L.) (Lepidoptera:
Nymphalidae). Bitzer and Shaw 18(1 ):36-49
A new record of Vanessa virginiensis "ab. ahwashtee" from northern
California (Lepidoptera: Nymphalidae). Shapiro 20(3): 1 76- 178
Occurrence of the "Elymi" aberrant phenotype in Vanessa carye (Huebner)
(Nymphalidae). Lamas 22(2):1 15-117
Notes: Six homoeotic Vanessa atalanta rubria (Nymphalidae).
Dimock 23(2):176
Notes: Culture maintenance of Vanessa atalanta rubria (Nymphalidae).
Dimock 23(3):236-240
The mating system of Vanessa kershawi: males defend landmark territories
as mate encounter sites. Alcock and Gwynne 26( 1 -4): 116-1 24
N YMPHALIDAE/ Y RAMEA
Habitat - Yramea cytheris. Hovanitz 9(2): 126
Oecophoridae/Odonna
Immature stages of Odonna passi florae Clark (Lepidoptera: Oecophoridae):
Biology and morphology. Chacon and de Hernandez 20(l):43-45
A new genus and two new species of Oecophoridae from Columbia
(Lepidoptera). Clark 20(l):46-49
Olethreutidae/Epinotia
A new species of Epinotia Hubner from British Columbia (Olethreutidae).
Freeman 5( 1 ):1 3-1 4
Olethreutidae/Zeiraphera
The North American species of the genus Zeiraphera. Mutuura and
Freeman 5(3):153-176
Oxytenidae/Oxytenis
A tropical caterpillar that mimics faeces, leaves and a snake (Lepidoptera:
Oxytenidae: Oxytenis naemia). Nentwig 24(2): 1 36- 141
Papilionidae
Latitudinal gradients in species diversity of the new world swallowtail
butterflies. Slansky 1 1(4):201-217
Hibernal diapause of North American Papilionoidea and Hesperioidea.
Scott 1 8(3): 171 -200
Seasonality of the butterfly fauna in southeastern Vietnam (Papilionidae).
Spitzer 22(2):126-130
56
FAMILY/GENUS INDEX
J. Res. Lepid.
Papilionidae/Lamproptera
Notes on the biology of Lamproptera curius Walkeri Moore (Lepidoptera:
Papilionidae). Howarth 15(1 ):27-32
Papilionidae/Papilio
Larval food-plant records for six western Papilios. Emmel and
Emmel 1(3):191-193
Life histories of Papilio indra and Papilio oregonius. Newcomer . 3(l):49-54
Hybrids between Papilio memnon and Papilio protenor. Ae 3(l):55-62
Genetic relationships of Papilio indra and Papilio polyxenes. Emmel and
Emmel 3(3):157-158
Oxygen consumption and metabolic rate of Papilio zelicaon pupae in a
state of delayed eclosion. La Due 3(4):197-206
An additional food plant record for Papilio thoas autocles R. & J,
Comstock 5(4):220
Further observations on "hilltopping" in Papilio zelicaon.
Guppy 8(3): 1 05- 1 17
A rearing of Papilio indra kaibabensis. Wielgus 8(4): 177-181
A new species of Papilio from the eastern United States (Papilionidae).
Heitzman 1 2( 1 ): 1 - 1 0
Melanie Papilio machaon larvae. Gardiner 15(3): 184
The use of alpha-eedysone to break permanent diapause of female hybrids
between Papilio glaucus L. female and Papilio rutulus male. Clarke and
Willig 16(4):245-248
Larval foodplant records for Papilio zelicaon in the western United States
and further evidence for the conspecificity of P. zelicaon and P.
gothica. Emmel and Shields 17(1 ):56-67
An aberrant Oregon Swallowtail, Papilio oregonius Edwards from Oregon.
Wescott 18(4):255
A melanic male aberration of Papilio glaucus canadensis from northern
Wisconsin. Scriber and Lintereur 2 1 (3): 1 99-20 1
Notes: Abnormal chrysalis of Papilio zelicaon (Papilionidae).
Priestaf 21(4):270
Notes: Type locality of Papilio indra pergamus (Lepidoptera: Papilionidae).
Miller . 23(2): 175
An exceptional case of paternal transmission of the dark form female
trait in the tiger swallowtail butterfly, Papilio glaucus (Lepidoptera:
Papilionidae). Scriber and Evans 25(2): 110-1 20
The effect of temperature on expression of the dark phenotype in Papilio
glaucus (Papilionidae). Ritland 25(3): 179-187
The mating behavior of Papilio glaucus (Papilionidae). Krebs. 26(l-4):27-31
A new heritable color aberration in the tiger swallowtail butterfly, Papilio
glaucus (Papilionidae: Lepidoptera). Scriber and Evans . . 26(l-4):32-38
Bilateral gynandromorphs, sexual and/or color mosaics in the tiger
swallowtail butterfly, Papilio glaucus (Lepidoptera: Papilionidae).
Scriber and Evans 26(l-4):39-57
Suppression of the black phenotype in females of the P. glaucus group
(Papilionidae). West and Clarke 26( 1 -4): 1 87-200
Hand-pairing of Papilio glaucus glaucus and Papilio pilumnus
(Papilionidae) and hybrid survival on various food plants. Scriber and
Lederhouse 27(2):96-103
Genetic experiments with a calverleyi-Mke. mutation isolated from Papilio
bairdi oregonius (Papilionidae). McCorkle and Hammond . 27(3): 1 86- 191
J. Res. Lepid.
FAMILY/GENUS INDEX
57
Papilionidae/Papilio (continued)
Hybridization of the Mexican tiger swallowtail, Papilio alexiares garcia
(Lep: Pap) with other P. glaucus group species and survival of pure and
hybrid larvae on potential host plants. Scriber et al 27(3):222-232
Papilionidae/Parnassius
A new Parnassius eversmanni race from northeast Siberia (USSR).
Weiss 9(4):2 1 5-2 1 6
A proposed revision of non-Arctic Parnassius phoebus Fabricus in North
America (Papilionidae). Ferris 1 5( 1 ): 1 -22
A note on the subspecies of Parnassius clodius Menetries found in the
Rocky Mountains of the United States (Papilionidae). Ferris. 15(2):65-74
A mutant affecting wing pattern in Parnassius apollo (Linne) (Lepidoptera
Papilionidae). Descimon and Vesco 26(1-4):161-172
Papilionidae/Protesilaus
A study of Protesilaus microdamas (Burmeister) and the little-known P.
huanucana (Varea de Luque) (Papilionidae). Johnson et al. . 27(2):83-95
Papilionidae/Troidine
Troidine swallowtails (Lepidoptera: Papilionidae) in southeastern Brazil:
natural history and foodplant relationships. Brown et al. . 1 9(4): 1 99-226
The biology of seven Troidine swallowtail butterflies (Papilionidae) in
Colima, Mexico. Spade et al 26( 1 -4): 1 3-26
Pericopidae/Gnophaela
Notes: Mate locating behavior of Gnophaela latipennis vermiculata G. & R.
(Pericopidae). Scott 20(1):51
PlERIDAE
Comparitive speciation in two butterfly families: Pieridae and
Nymphalidae. Petersen 5(2): 113-1 26
Origin of autumnal "false broods" in common Pierid butterflies.
Shapiro 6(3): 181-1 93
The effect of pterin pigments on wing coloration of four species of
Pieridae. Pfeiler 7(4):183-189
An interfamilial courtship (Nymphalidae, Pieridae). Shapiro. 1 1 (3): 1 97-1 98
The role of watercress, Nasturtium officinale as a host of native and
introduced pierid butterflies in California. Shapiro 1 4(3): 158-1 68
Enzyme electrophoretic studies on the genetic relationships of Pierid
butterflies (Lepidoptera: Pieridae) I. European taxa.
Geiger 1 9(4): 1 8 1 - 1 95
Notes: An interfamilial courtship (Lycaenidae - Pieridae). Shapiro. 20(1):54
The biological and systematic significance of red fecal and meconial
pigments in butterflies: A review with special reference to the Pieridae.
Shapiro 20(2):97-102
Polyphenism, phyletic evolution, and the structure of the Pierid genome.
Shapiro 23(3): 1 77- 1 96
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera). Lorkovic 24(4):334-358
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera) - The case for enzyme electrophoresis.
Geiger 26(l-4):64-72
Pierid ae/Anthocharis
A field captured scale-deficient mutant of Anthocharis sara.
Dornfield 9(l):25-28
Recent captures of Anthocharis cethura catalina Meadows. Orsak. 14(2):85-89
58
FAMILY/GENUS INDEX
J. Res. Lepid.
Pieridae/Anthocharis (continued)
Another Anthocharis lanceolata X A. sara hybrid. Shields and
Mori 17(l):53-55
The impact of Pierid feeding on seed production by a native California
crucifer. Shapiro 24(2): 191-1 94
Electrophoretic evidence for speciation within the nominal species
Anthocharis sara Lucas (Pieridae). Geiger and Shapiro .... 25(0-15-24
Pierid ae/ A scia
Systematics of Ascia ( Ganyra ) (Pieridae) populations in the Sonoran Desert.
Bailowitz 26(l-4):73-81
Pierid ae/Catopsilia
Notes: An early season migration of Catopsilia pomona (Lepidoptera:
Pieridae) in Java, Indonesia. New 24(0-84-85
Pierid ae/Colias
The male genitalia of some Colias species. Petersen 1 (2): 135-1 56
Colias philodice in Chiapas, Mexico. Emmel 1(3): 194
The generis, specific and lower category names of the Nearctic butterflies.
Part 2 - the genus Colias. McHenry 1(3):209-221
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 1 (4):26 1 -274
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 2(3):205-223
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 3(0-37-44
New gynandromorph of Colias philodoce from Colorado. Emmel 3(0-63-64
A Colias Christina gynandromorph. Hovanitz 4( 1 ):4 1
Colias christina-alexandra intergradation. Cover illust. Hovanitz . . 4(1):42
Vital staining of Colias philodoce and C. eurytheme. Kolyer . . 5(3): 137-1 52
Man-made habitat for Colias eurytheme. Hovanitz 6(4):267
Scanning electron microscopy on the wing scales of Colias eurytheme.
Kolyer and Reimschuessel 8( 1 ): 1 - 1 5
Habitat - Colias philodice eriphyle and Colias eurytheme. Hovanitz . 8(4): 182
Concerning Colias eurytheme alberta Bowman (Pieridae). Masters. 9(2):97-99
Habitat - Colias vautieri. Hovanitz 9(2): 100
Concerning Colias Christina mayi. Masters 9(4):227-232
Lack of melanism in Colias (Cover illustration). Masters 1 1 (4):2 1 8
Cover Illustration: Variation in Colias nastes of Lapland.
Hovanitz 1 2(3): 1 80
Variation in Colias alexandra Christina Edwards (Pieridae) in southwest
Manitoba. Masters 1 4(3): 148-157
A new subspecies of Colias palaeno (Linnaeus) from Baffin Island, N.W.T.,
Canada (Pieridae). Ebner and Ferris 1 6(3): 155-161
Notes: On Colias hecla Lefebvre re a recent paper by Oosting and Parshall
(Lepidoptera: Pieridae). Ferris 20(l):52-53
On the nomenclature of Colias alfacariensis Berger 1948 (Lepidoptera:
Pieridae). Kudrna 20(2): 1 03- 110
An apparent "intersexual" Colias eurytheme (Pieridae). Shapiro . 20(4):244
Notes: Further notes regarding Colias hecla Lefebvre (Lepidoptera:
Pieridae) at Churchill, Manitoba. Parshall 20(4):250
Colias alexandra : A model for the study of natural population of
butterflies. Hayes 23(2): 113-1 24
J. Res. Lepid .
FAMILY/GENUS INDEX
59
Pieridae/Colias (continued)
Protein and lipid composition of Colias philodice and C. eurytheme
spermatophores and their changes over time. Marshall .... 24( 1 ):2 1 -30
Notes: A melanic Colias euxanthe stuebeli from Peru (Pieridae).
Shapiro 24(1):87
Pieridae/Colotis
The life histories of South African Colotis erone , C. ione , C. vesta and
Leptosia alcesta (Pieridae). Clark and Dickson 6( 1 ):3 1 -42
Pieridae/Eucheira
Notes: Homosexual pseudocopulation in Eucheira socialis (Pieridae).
Shapiro 27(3):262
Pieridae/Euchloe
Studies on the Nearctic Euchloe. Parts I, II. Opler 5(l):39-50
Studies on the Nearctic Euchloe. Part 3. Complete synonymical treatment.
Part 4. Type data and type locality restrictions. Opler .... 5(3): 185-1 95
Studies on Nearctic Euchloe. Part 5. Distribution. Opler 7(2):65-86
Habitat - Euchloe hy antis and rew si. Hovanitz 8( 1 ): 1 6- 1 7
Studies of Nearctic Euchloe. Part 6. Systematics of adults.
Opler 8(4): 153-1 68
Studies on Nearctic Euchloe - Part 7. Comparative life histories, hosts and
the morphology of immature stages. Opler 1 3( 1 ): 1 -20
Pieridae/Eurema
The life history of two species of South African Eurema. Clark and
Dickson 4(4):252-257
South African Eurema. Clark and Dickson 8( 1 ): 1 8- 1 9
Pieridae/Gonepteryx
The hidden wing-pattern of some Palearctic species of Gonepteryx and its
taxonomic value. Nekrutenko 3(2):65-68
Three cases of gynandromorphism in Gonepteryx. Nekrutenko. 4(2): 1 03- 1 07
A new subspecies of Gonepteryx amintha (Pieridae) from Yunnan,
mainland China. Nekrutenko 1 1(4):235-240
Pieridae/ Ixias
A study of the meiotic chromosomes of Ixias marianne (Cramer) (Pieridae).
Rao 1 7(3): 1 70- 172
Pieridae/Leptosia
The life histories of South African Colotis erone , C. ione , C. vesta and
Leptosia alcesta (Pieridae). Clark and Dickson 6( 1 ):3 1 -42
Pieridae/Mathania
Oviposition by the mistletoe-feeding Pierid butterfly Mathania leucothea
(Mol.) in Chile. Courtney 24(3):264-270
Pieridae/ Nathalis
Courtship behavior of the dainty sulfur butterfly, Nathalis iole with a
description of a new, faculative male display (Pieridae).
Rutowski 20(3): 161-1 69
Pierid ae/Perrhybris
Observations on the apparent Lek behavior in Costa Rican rainforest
Perrhybris pyrrha Cramer (Pieridae). DeVries 17(3): 142- 144
Pierid ae/Phoebis
Observations on Phoebis sennae (Pieridae). Brown 17(3)168-169
Courtship leading to copulation in the cloudless sulphur, Phoebis sennae
(Pieridae). Rutowski 22(4):249-253
60
FAMILY/GENUS INDEX
J. Res. Lepid.
Pieridae/Pierini
Photoperiod and temperature in phenotype determination of Pacific slope
Pierini: biosystematic implications. Shapiro 1 6(4): 1 93-200
Pieridae/Pieris
The effect of various food plants on survival and growth rate of Pieris.
Hovanitz and Chang 1(1 ):2 1 -42
Three factors affecting larval choice of food plant. Hovanitz and
Chang 1(1 ):5 1-61
The generic, specific and lower category names of the Nearctic butterflies.
Part 1 - the genus Pieris. McHenry 1(1 ):63-7 1
The distribution of the species of the genus Pieris in North America.
Hovanitz 1(1 ):73-83
The relation of Pieris virginiensis Edw. to Pieris napi L. species formation
in Pieris ? Hovanitz 1 (2): 1 24- 1 34
The effect of hybridization of host-plant strains on growth rate and
mortality of Pieris rapae. Hovanitz and Chang 1 (2): 157-1 62
Change of food plant preference by larvae of Pieris rapae controlled by
strain selection and the inheritance of this trait. Hovanitz and
Chang 1 (2): 1 63- 1 68
Selection of allyl isothiocyanate by larvae of Pieris rapae and the
inheritance of this trait. Hovanitz and Chang 1 (3): 1 69- 1 82
The effectiveness of different isothiocyanates on attracting larvae of
Pieris rapae. Hovanitz et al l(4):249-259
A method for breeding Pieris napi and Pieris bryoniae.
Petersen l(4):275-279
Quantitative analysis of certain wing and genitalia characteristics of
Pieris in western North America. Chang 2(2):97- 125
Genetic and environmental variation in Pieris brassicae.
Gardiner 2(2):127-136
Oviposition preference tests with Pieris. Hovanitz and Chang . 2(3): 1 85-200
Comparison of the selective effect of two mustard oils and their
glucosides on Pieris larvae. Hovanitz and Chang 2(4):28 1 -288
The southern limits of the range of Pieris napi and P. virginiensis.
Mather 3(l):45-48
Adult oviposition responses in Pieris rapae. Hovanitz and
Chang 3(3): 1 59- 172
The alteration of host plant specificity in larvae of Pieris rapae by
induction. Hovanitz and Chang 4( 1 ): 1 3-2 1
The feeding of coloring matters to Pieris rapae larvae. Kolyer.4(3):159-172
A melanic form of Pieris rapae. Donahue 6(4):266
Habitat - Pieris beckeri. Hovanitz 7(1 ):56
Development of the markings on the pupal wing of Pieris rapae (Pieridae).
Kolyer 8(3):69-90
Records of Colias gigantea from southwest Manitoba and Minnesota.
Masters 8(3):129-132
Dispersal in cosmopolitan butterfly species ( Pieris rapae) having open
population structure. Emmel ll(2):95-98
Gynandromorphism in Pieris brassicae L. Gardiner 1 1 (3): 1 29- 1 40
Vital staining as evidence for wing circulation in the cabbage butterfly
Pieris rapae. Kolyer 1 1(3): 161-173
J . Res. Lepid.
FAMILY/GENUS INDEX
61
Pieridae/Pieris (continued)
The genetics of subspecific phenotype differences in Pieris occidentalis
Reakirt and of variation in P. o. nelsoni W. H. Edwards (Pieridae).
Shapiro 14(2):61-83
Habitat: Pieris occidentalis Nelsoni (Pieridae). Shapiro 1 5(2): 1 03- 1 05
Habitat: Pieris occidentalis (Pieridae). Shapiro 1 5(3): 182-183
Weather and the liability of breeding populations of the checkered white
butterfly, Pieris protodice Boisduval and Le Conte. Shapiro . 17(1):1-16
The depredations of the large white butterfly ( Pieris brassicae ) (Pieridae).
Feltwell 17(4):2 18-225
Diapause in various populations of Pieris napi L. from different parts of
the British Isles. Lees and Archer 19(2):96-100
Two homoeotic Pieris rapae of Mexican origin (Pieridae).
Shapiro 20(4):242-244
Allozyme variation in a colonizing species: The cabbage butterfly Pieris
rapae (Pieridae). Vawter and Brussard 22(3):204-216
A critical review of "Systematische Untersuchungen am Pieris
napi-bryoniae- Komplex (s.l.)" Lepidoptera: Pieridae) by Ulf
Eitschberger. Kudrna and Geiger 24(l):47-60
Oviposition on peripheral hosts by dispersing Pieris napi (L.) (Pieridae).
Courtney 26(l-4):58-63
On Pieris ( Artogeia ) marginalis macdunnoughii Remington (Pieridae).
Bowden 26(l-4):82-88
Notes: A significant new host plant record for Pieris virginiensis (Pieridae).
Shuey 27(3):259-260
Pieridae/ Pontia
Notes: A complex gynandromorph of Pontia daplidice (Pieridae).
Shapiro 23(4):332-333
Electrophoretic confirmation of the species status of Pontia protodice and
P. occidentalis (Pieridae). Shapiro and Geiger 25(l):39-47
Pieridae/Tatochila
Notes: A reared gynandromorph of Tatochila (Pieridae).
Shapiro 20(4):240-242
Notes: A recessive lethal "wingless" mutation in Tatochila (Pieridae).
Shapiro 22(4):262-263
Pieridae/Zerene
Habitat - Zerene caesonia eurydice. Hovanitz 7(4): 182, 190
Psychidae/Pteroma
Variations in the wing venation of Pteroma plagiophleps Hampson
(Lepidoptera: Psychidae). Mathew 24(4):359-363
Pyralidae
Chromosome studies in sixteen species of Indian Pyralid moths
(Pyralidae). Mohanty and Nayak 20(2):86-96
Riodinidae/Anatole
A new species of Rioninidae from Mexico. Clench 3(2):73-79
Life history studies on Mexican butterflies. II, Anatole rossi.
Ross 3(2):8 1-94
Riodinidae/Apodemia
Description and taxonomic implications of an unusual Arizona population
of Apodemia mormo (Riodinidae). Forbes 1 8(3):20 1 -207
Riodinidae/Calephelis
The butterfly genus Calephelis. McAlpine 1 0( 1 ): 1 - 1 25
62
FAMILY/GENUS INDEX
J. Res . Lepid.
Riodinidae/Lasaia
A review of the genus Lasaia (Riodinidae). Clench 1 0(2): 1 49- 1 80
Saturniidae/Actias
Note on vital staining of Actias lima silk. Kolyer 7(1 ):29-30
Saturniidae/Anisota
A revision of the American Genus Anisota (Saturniidae). Riotte and
Peigler 1 9(3): 1 0 1 - 1 80
Saturniidae/ A ntheraea
Susceptibility of eggs and first-instar larvae of Callosamia promethea and
Antheraea polyphemus to Malathion. Miller et al 25(1):48-51
Notes: Effect of refrigeration on hatching of eggs of the tasar silk moth
Antheraea mylitta (Saturniidae). Dash and Nayak 27(3):263-265
Saturniidae/ Attacus
Ocellus variation and wingspan in Attacus atlas Linnaeus, Is there a
relationship? Hadley 1 6(3): 1 4 1 - 145
Saturniidae/ Automeris
The life history of Automeris zephyria (Saturniidae). Tuskes and
Smith 27(3):192-196
Saturniidae/Callosamia
Demonstration of reproductive isolating mechanisms in Callosamia
(Saturniidae) by artificial hybridization. Peigler 19(2):72-81
Susceptibility of eggs and first-instar larvae of Callosamia promethea and
Antheraea polyphemus to Malathion. Miller et al 25(1):48-51
Saturniidae/Calosaturnia
Type locality for Calosaturnia walterorum Johnson (Saturniidae).
Orsak 15(4):214
Saturniidae/Coloradia
A new species of Coloradia in California (Saturniidae, Hemileucinae).
Johnson and Walter 18(1 ):60-66
S ATURNIID AE/ DlRPHIA
The early stages of various species of the genus Dirphia (Saturniidae).
Gardiner 13(2):101-1 14
Saturniidae/Dirphiopsis
The rearing of Dirphiopsis eumedide (Saturniidae). Gardiner . 4(4):287-291
Saturniidae/Eacles
Notes on Eacles penelope (Saturniidae). Gardiner 5(3): 177-1 80
Saturniidae/Euleucophaeus
Rearing Euleucophaeus rubridorsa and E. Lex. Gardiner 6(l):53-58
Saturniidae/Hemileuca
The life history of Hemileuca magnifica (Saturniidae) with notes on
Hemileuca hera marcata. Stone et al 26(1 -4):225-235
Saturniidae/Hyalophora
Notes: Weights and dimensions of Hyalophora euryalis and pupae
(Lepidoptera: Saturniidae). Miller 24(l):83-84
Saturniidae/Leucanella
The early stages of Leucanella memusae ssp. gardinerii Lemaire
(Saturniidae). Gardiner 15(4):201-205
Rectification of a recent paper on Leucanella memusae gardineri.
Peigler 16(4):222
Saturniidae/Ormiscodes
A new species of Ormiscodes (Dirphiella) from Mexico (Saturniidae:
Hemileucinae). Donahue and LeMaire 1 3(2): 1 23- 1 30
J. Res. Lepid.
FAMILY/GENUS INDEX
63
Saturniidae/Paradirphia
Three new species of Paradriphia (Saturniidae: Hemaleucinae) from
Mexico and Central America with notes on the immature stages.
LeMaire and Wolfe 27(3): 1 97-2 1 2
Saturniidae/Philosamia
Supernumerary chromosomes in the domesticated eri-silkmoth, Philosamia
ricini (Saturniidae: Lepidoptera). Padhy and Nayak 20( 1 ): 1 6- 1 7
Chromosome aberrations in the holocentric chromosomes of Philosamia
ricini (Saturniidae). Padhy 25(l):63-66
Saturniidae/Samia
Samia watsoni Obertheur color plate. LeMaire and Peigler ...... 18(1 ):67
Saturniidae/Saturnia
Systematics and life history of Saturnia ( Calosaturnia ) albofasciata in
California (Saturnidae). Hogue et al 4(3): 1 73- 1 84
Saturniidae/Sonthonnaxia
Chromosome studies including a report of B-chromosome in a wild
silkmoth, Sonthonnaxia maenas (Doubleday) (Saturniidae: Saturniinae).
Narang and Gupta 1 8(3):208-2 1 1
Satyridae
On Mexican Satyridae with description of a new species. Miller .7(1 ):5 1-55
Notes: Moss feeding by a Satyrine butterfly. Singer and Mallet . 24(4):392
Satyridae/Calisto
A new species of Calisto from Hispaniola with a review of the female
genitalia of Hispaniolan congeners (Satyridae). Johnson et
al 25(2):73-82
Satyridae/Cercyonis
Estimation of natural mutation rates for albinism in two species of
Satyrid genus Cercyonis. Emmel 8(2):65-68
Satyridae/Coenonympha
W. H. Edwards’ life histories of North American Coenonympha.
Brown 3(2):121-128
Comments on the genus Cercyonis Scudder, with figures of types
(Satyridae). Brown 4(2): 131-1 48
Notes: Description of the larvae of Coenonympha haydeni Edwards
(Lepidoptera: Satyridae). Rosenberg 24(4):394-395
Satyrid ae/Erebia
Ecological and distribution notes on Erebia disa (Satyridae) in central
Canada. Masters 7( 1 ): 1 9-22
Ecological and distributional notes on Erebia discoidalis (Satyridae) in the
north central states. Masters 9( 1 ): 1 1-16
A review of the Erebia dabanensis complex (Lepidoptera: Satyridae), with
descriptions of two new species. Troubridge and Philip . . 2 1 (2): 107- 1 46
Notes: Notes on Erebia occulta (Lepidoptera: Satyridae). Philip and
Roos 24(l):81-82
Satyridae/ Euptychia
Population biology of the Neotropical Satyrid butterfly, Euptychia hermes.
1. Interpopulation movement, etc. Emmel 7(3):1 53-165
A population study of Euptychia hermes in northern Florida.
Kilduff 1 1(4):219-228
Satyridae/ H ipp archia
Migration of Hipparchia semele L. Feltwell 1 5(2):83-9 1
64
FAMILY/GENUS INDEX
J. Res. Lepid.
Satyridae/Hipparchia (continued)
Notes: Further migration of Hipparchia semele (L.) in 1976 and 1980.
Feltwell and Ducros 20(1 ):53
Hipparchia azorina (Strecker, 1899) (Satyridae) biology, ecology and
distribution on the Azores Islands. Oehmig 20(3): 1 36- 1 60
Description of the female genitalia of Hipparchia fagi Scopoli, Hipparchia
semele Linnaeus (Satyridae) and their related taxa.
Coutsis 22(3):161 -203
Satyridae/Oeneis
Population structure of Oeneis melissa semidea (Satyridae) from the
Presidential Range, New Hampshire. Anthony 7(3): 1 33- 148
Habitat - Oeneis chryxus Stanislaus. Hovanitz 8(4): 194
Habitat: Oeneis macounii Edwards. Masters 1 0(4):30 1-302
Habitat: Oeneis jutta ascerta Masters & Sorenson. Masters 1 1 (2):94
A note on Oeneis Melissa (Fabricius) in the western United States
(Satyridae). Ferris 1 4(4):2 1 3-2 1 5
A note on Oeneis jutta harperi , its author and date of publication
(Satyridae). Dos Passos 1 5(4):2 1 1 -2 1 3
Satyridae/Pyronia
Notes: An effect of the colony edge on gatekeeper butterflies Pyronia
titonus L. (Satyridae). Thomas 21(3):206-207
Sesiidae/Melittia
A new squash borer from Mexico (Lepidoptera: Sesiidae).
Friedlander 24(4):277-288
Sphingidae
Chromosomes of seven species of Indian Sphingid moths. Mohanty and
Nayak 21(4):238-244
Sphingid ae/Darapsa
New food plant for Darapsa pholus (Cramer). Riotte 12(4):209-210
New food plant for Darapsa pholus (Cramer). Riotte 13(4):247-248
Sphingid ae/Euproserpinus
Overcoming difficulties with the pupae of Euproserpinus phaeton mojave.
McFarland 5(4):249-252
Sphingid ae/Hyles
Larval migration of Hyles lineata (Fab.). Wells and Brown 13(4):246
Sphingid ae/Pholus
An evaporative cooling mechanism in Pholus achemon (Sphingidae).
Adams and Heath 3(2):69-72
Tortricidae
Discovery of two new species and genera of Shaggy Tortricids related to
Synnoma and Niasoma (Tortricidae: Sparganothini). Powell 24( 1 ):6 1-71
New host records and morphological notes on four Tortricines
(Tortricidae). Sandberg and Passoa 27(2):104-108
T ortricidae/Gnathmocerodes
Notes on Gnathmocerodes petri fraga Diakonoff 1967 (Lepidoptera:
Tortricidae) associated with Barringtonia trees. Spitzer and
Jaros 24(2):187-190
T ortricidae/Rhyacionia
Nantucket Pine Tip Moth, Rhyacionia frustrana , in Kern County,
California: Integrated control and biological notes (Lepidoptera:
Tortricidae, Olethreutinae). Poore 19(2):65-67
J. Res. Lepid.
FAMILY/GENUS INDEX
65
Zygaenidae
An annotated catalogue of the Burnets and Foresters (Lepidoptera:
Zygaenidae) named by Roger Verity. Balletto and
Kudrna 24(3):226-249
GEOPOLITICAL INDEX
[Volumes 1-27, 1962-1988(89)1
Journal of Research on the Lepidoptera
AFRICA
Morocco
Notes: Notes on Tomcires mauretanicus (Lycaenidae) in Morocco.
Courtney 21(3):205-206
Notes: Oviposition records and larval foodplants of butterflies in the
Atlas Mountains of Morocco. Thomas and Mallorie 24(l):76-79
Nigeria
The butterfly fauna of a secondary bush locality in Nigeria. Larsen et
al 18(1 ):4-23
South Africa
The life history of two species of South African Eurema. Clark and
Dickson 4(4):252-257
The life histories of South African Colotis erone, C. lone , C. vesta and
Leptosia alcesta (Pieridae). Clark and Dickson 6( 1 ):3 1 -42
South African Eurema. Clark and Dickson 8( 1 ): 1 8- 1 9
ASIA (north of Himalayas)
Afghanistan
The Scolitantidini II. The World’s smallest butterfly? Notes on Turanana ,
and a new genus and species from Afghanistan (Lycaenidae).
Mattoni 18(4):256-264
China
A new subspecies of Gonepteryx amintha (Pieridae) from Yunnan,
mainland China. Nekrutenko 1 1(4):235-240
Siberia
A new Parnassjus eversmanni race from northeast Siberia (USSR).
Weiss 9(4):2 1 5-2 1 6
CARIBBEAN ISLANDS
Antillean Islands
Affinities and distribution of Antillean Ithomiidae. Fox .... 2(3): 173-1 84
A list of Antillean butterflies. Scott 9(4):249-256
Hispaniola
A new species of Calisto from Hispaniola with a review of the female
genitalia of Hispaniolan congeners (Satyridae). Johnson et
al 25(2):73-82
Jamaica
The identification of two species of Junonia Hubner (Lepidoptera:
Nymphalidae) : J. evarete and J. genovena in Jamaica. Turner and
Parnell 24(2): 1 42- 1 53
Providencia
The butterfly faunas of San Andres and Providencia Islands in the
western Caribbean. Emmel 14(1 ):49-56
66
J. Res. Lepid.
GEOPOLITICAL INDEX
67
San Andreas
The butterfly faunas of San Andres and Providencia Islands in the
western Caribbean. Emmel 14(1 ):49-56
Virgin Islands
Variation of Uthetheisa ornatrix (Arctiidae) including a new species from
St. Croix, Virgin Islands. Pease 10(4):261-264
Butterflies of St. Croix. Leek 1 2(3): 161-1 62
West Indies
A review of the West Indian "C ho ran thus". Miller 4(4):259-274
CENTRAL AMERICA & MEXICO
Three new species of Paradriphia (Saturniidae: Hemaleucinae) from
Mexico and Central America with notes on the immature stages.
LeMaire and Wolfe 27(3): 1 97-2 1 2
Costa Rica
Population biology of the Neotropical Satyrid butterfly, Euptychia hermes.
1. Interpopulation movement, etc. Emmel 7(3): 153-1 65
Observations on the apparent Lek behavior in Costa Rican rainforest
Perrhybris pyrrha Cramer (Pieridae). DeVries 17(3): 142- 144
A new Tortyra from Cocos Island, Costa Rica (Lepidoptera: Choreutidae).
Heppner 1 9(4): 1 96- 1 98
Illustrations and descriptions of some species of Pyrrhopyginae from
Costa Rica, Panama and Columbia (Hesperiidae). Nicolay and
Small 19(4):230-239
A new species of Adelpha (Nymphalidae) from Parque Nacional Braulio
Carrillo, Costa Rica. DeVries and Gamboa 20(2): 1 23- 1 26
Hostplant records and natural history notes on Costa Rican butterflies
(Papilionidae, Pieridae & Nymphalidae). DeVries 24(4):290-333
Notes: Natural history notes on Brassolis isthmis Bates (Lepidoptera:
Nymphalidae: Brassolinae) in northeastern Costa Rica.
Young 24(4):385-392
Notes: Moss feeding by a Satyrine butterfly. Singer and Mallet . 24(4):392
The mating system of three territorial butterflies in Costa Rica.
Alcock 26(l-4):89-97
Stratification of fruit-feeding nymphalid butterflies in a Costa Rican
rainforest. DeVries 26( 1 -4):98- 1 08
Mexico
Life history studies on Mexican butterflies. I. Ross 3( 1 ):9- 1 7
A new species of Rioninidae from Mexico. Clench 3(2):73-79
Life history studies on Mexican butterflies. II, Anatole rossi.
Ross 3(2):81-94
Life history studies on Mexican butterflies. III. Nine Rhopalocera from
Ocotal Chico, Vera Cruz. Ross 3(4):207-229
New skipper records for Mexico. Freeman 5(l):27-28
Remarks on the genus Zera Evans in Mexico with a new record.
Freeman 5(3): 181-1 84
The status of some Hesperiidae from Mexico. Freeman 6(l):59-64
Polyctor polyctor in Mexico. Freeman 6(3): 1 95- 1 96
On Mexican Satyridae with description of a new species. Miller. 7(1 ):5 1-55
Butterflies of middle and southern Baja California. Holland. 1 1 (3): 1 47-1 60
A new species of Ormiscodes (Dirphiella) from Mexico (Saturniidae:
Hemileucinae). Donahue and LeMaire 1 3(2): 1 23- 1 30
68
GEOPOLITICAL INDEX
J. Res. Lepid .
Mexico (continued)
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico. Ross 14(2): 103- 124
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued). Ross 14(3): 169- 188
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued). Ross 14(4):233-252
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued). Ross 15(1 ):4 1 -60
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued). Ross 15(2): 109- 128
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued ). Ross 15(3): 185-200
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (continued). Ross 15(4):225-240
An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz,
Mexico (concluded). Ross 1 6(2):87- 1 30
A new squash borer from Mexico (Lepidoptera: Sesiidae).
Friedlander 24(4):277-288
The biology of seven Troidine swallowtail butterflies (Papilionidae) in
Colima, Mexico. Spade et al 26( 1 -4): 1 3-26
Notes: Records of Hypaurotis crysalus (Ed) (Lycaenidae) from western
Mexico. Brown 27(2): 135
The butterflies of Isla de Cedros, Baja California Norte, Mexico. Brown
and Faulkner 27(3):233-256
Panama
Seasonal changes in organization of tropical rain forest butterfly
populations in Panama. Emmel and Leek 8(4): 133-1 52
Illustrations and descriptions of species of some Pyrrhopyginae from
Panama (Hesperiidae). Nicolay 1 3(3): 181-1 90
Illustrations and descriptions of some species of Pyrrhopyginae from
Costa Rica, Panama and Columbia (Hesperiidae). Nicolay and
Small 19(4):230-239
EUROPE
Azores Islands
Hipparchia azorina (Strecker, 1899) (Satyridae) biology, ecology and
distribution on the Azores Islands. Oehmig 20(3): 1 36- 1 60
British Isles
The nomenclature in an important British check list (1972). Part I.
Paclt 12(4):21 1-212
The nomenclature in an important British check list (1972) Part 2:
Corrections of family-group names for Geometridae (lepidoptera).
Paclt 1 3(3): 179-1 80
The nomenclature in an important British check list (1972). Part 3. Correct
gender for generic names derived from classical without change of
termination. Paclt 13(4):267-270
The nomenclature in an important British check list (1972) Part 4: Correct
gender for some other generic names. Paclt 17(1 ):24-26
Diapause in various populations of Pieris napi L. from different parts of
the British Isles. Lees and Archer 1 9(2):96- 1 00
J . Res. Lepid.
GEOPOLITICAL INDEX
69
Spain
Diversity and species richness of butterflies and skippers in central Spain
habitats. Montesinos 24(4):364-371
INDO- AUSTRALIA (south of Himalayas)
Australia
Egg photographs depicting 40 species of Southern Australian moths.
McFarland 10(3):2 1 5-247
Biology and immature stages of Australian Ethmiid moths (Gelechioidea).
Powell 20(4):214-234
India
Chromosome studies in sixteen species of Indian Pyralid moths
(Pyralidae). Mohanty and Nayak 20(2):86-96
Chromosomes of seven species of Indian Sphingid moths. Mohanty and
Nayak 2 1 (4):238-244
Karyotypes of some Indian Noctuid moths (Lepidoptera). Mohanty and
Nayak 22(4):238-248
Indonesia
Notes: An early season migration of Catopsilia pomona (Lepidoptera:
Pieridae) in Java, Indonesia. New 24(l):84-85
Nepal
International Nepal Himalaya Expedition for Lepidoptera Palaeartica
(INHELP) 1977, Report No. 1: Introduction and Lycaenidae.
Shields 20(2):65-80
Vietnam
Seasonality of the butterfly fauna in southeastern Vietnam (Papilionidae).
Spitzer 22(2): 1 26- 1 30
Notes on Gnathmocerodes petri fraga Diakonoff 1967 (Lepidoptera:
Tortricidae) associated with Barringtonia trees. Spitzer and
Jaros 24(2):187-190
NORTH AMERICA (north of Mexico)
Canada
New Canadian species of leaf-mining lepidoptera of conifers.
Freeman 4(3):209-220
A new species of Epinotia Hubner from British Columbia (Olethreutidae).
Freeman 5( 1 ): 1 3- 1 4
A new species of Nepticula on bur oak in Ontario (Nepticulidae).
Freeman 6( 1 ): 1 9-2 1
Ecological and distribution notes on Erebia disa (Satyridae) in central
Canada. Masters 7( 1 ): 1 9-22
On the distribution of some Skippers in Ontario. Riotte 1 1 (2):8 1 -82
On the origin of austral elements in the moth fauna of south-eastern
Ontario, including a number of species new for Canada. Harmsen et
al 1 2(3): 1 27- 1 34
Checklist of the Macroheterocera of south-eastern Ontario. Ward et
al 13(l):23-42
Significant additions to the Lepidopterous fauna of southeastern Ontario.
Riotte 1 5(2): 101-1 02
Butterfly collecting in Labrador and Newfoundland. Ferris . 1 5(2): 1 06- 1 08
A new subspecies of Colias palaeno (Linnaeus) from Baffin Island, N.W.T.,
Canada (Pieridae). Ebner and Ferris 1 6(3): 155-161
70
GEOPOLITICAL INDEX
J. Res. Lepid.
Canada (continued)
Ecological notes on the butterflies of the Churchill region of Northern
Manitoba. Oosting and Parshall 1 7(3): 1 88-203
Notes: Further notes regarding Colias hecla Lefebvre (Lepidoptera:
Pieridae) at Churchill, Manitoba. Parshall 20(4):250
A list of the butterflies and skippers of Mount Revelstoke and Glacier
National Parks, British Columbia, Canada. Threatful . . . 27(3):2 1 3-22 1
Records of Colias gigantea from southwest Manitoba and Minnesota.
Masters 8(3): 1 29- 1 32
United States
A new species of Nephelodes Guenee for the Great Basin.
Buckett 1 1(4):260-268
Post Pleistocene environments and montane butterfly relicts on the
western Great Plains. Johnson 1 4(4):2 1 6-232
Lowland riparian butterflies of the Great Basin and associated areas.
Austin 24(2):1 17-131
Arizona
The little known moth Euxoa sculptilis (Harvey) in Arizona, with
descriptions, illustrations, and notes on Euxoa violaris (Grote and
Robinson) (Noctuidae). Buckett 5(4):255-261
A search for Speyeria nokomis coerulescens (Holland) (Nymphalidae) in
southern Arizona. Wielgus 1 1(3):187-1 94
The correct name for the subspecies of Limenitis weidemeyerii occurring in
Arizona (Nymphalidae). Dos Passos 1 2( 1 ):2 1 -24
Description and taxonomic implications of an unusual Arizona population
of Apodemia mormo (Riodinidae). Forbes 1 8(3):20 1 -207
A new Limenitis weidemeyerii W. H. Edwards from southeastern Arizona
(Nymphalidae). Austin and Mullins 22(4):225-228
Census of the butterflies of the National Audubon Society’s
Appleton-Whittell Research Ranch, Elgin, Arizona.
Bailowitz 27(2): 1 20- 1 28
California
Variations in the silvering of Argynnis (Speyeria) callippe in the interior
mountain area of south central California. Sette 1(1 ):3-20
Further evidence of the distribution of some boreal Lepidoptera in the
Sierra Nevada. Ericksen 1(1 ):89-93
Composition and relative abundance in a temperate zone butterfly fauna.
Emmel and Emmel 1 (2):97- 1 08
Notes on the early stages of two California geometrids.
Comstock l(3).T95-200
Early stages of a southern California Geometrid moth, Drepanulatrix hulsti
hulsti (Dyar). Comstock l(4):245-248
Yosemite butterflies: An ecological survey of the butterflies of the
Yosemite sector of the Sierra Nevada, California. Garth and
Tilden 2( 1 ): 1 -96
The distribution of an endemic butterfly Lycaena hermes.
Thorne 2(2):143-150
Studies in life histories of North American Lepidoptera, California
Annaphilas. Comstock and Henne 3(3): 173-191
Petaluma , a new genus. Buckett and Bauer 3(3): 1 93- 1 96
The moths (Macroheterocera) of a chaparral plant association in the Santa
Monica Mountains of southern California. McFarland 4(l):43-73
J. Res. Lepid.
GEOPOLITICAL INDEX
71
California (continued)
Systematics and life history of Saturnia ( Calosaturnia ) albofasciata in
California (Saturnidae). Hogue et al 4(3): 173-1 84
California coastal Eupithecia with description of new species
(Geometridae). Leuschner 4(3): 191-1 97
A population study of a hibernal roosting colony of the monarch butterfly
( Danaus plexippus ) in northern California. Urquhart et al. 4(4):221-226
Studies in the life histories of North American Lepidoptera, California
Annaphila II. Comstock and Henne 5(1): 15-26
Rediscovery of Annaphila casta Hy. Edw. in California (Noctuidae).
Buckett 5(l):37-38
The butterfly fauna of a yellow pine forest community.
Shields 5(2): 1 27- 1 28
The eggs and first instar larvae of three California moths.
Comstock 5(4):2 1 5-2 1 9
A new species of Polia Ochsenheimer from California and notes on Polia
discalsis (Grote) (Noctuidae: Hadeninae). Buckett and
Bauer 5(4):221-228
Description of a new species of Xylomiges from California.
Buckett 6(l):23-30
A new species of Feralia. Buckett 6(1):43-51
Studies in the life histories of North American Lepidoptera, California
Annaphila III. Comstock and Henne 6(4):257-262
A previously unrecognized subspecies of Philotes speciosa.
Tilden 6(4):281-284
Variation in color and maculation in Nemoria pulcherrima from the Sierra
Nevada of California. Lepidoptera: Geometridae. Buckett and
Sears 7(2):95-98
A continuously breeding population of Danaus plexippus in southern
California. Urquhart et al 7(4): 1 69- 181
Seasonal distribution of "Macrolepidoptera" in Santa Clara County,
California. Opler and Buckett 9(2):75-88
Synaxis mosesiani Sala; a new Synaxis from southern California.
Sala 9(3): 185-191
The distribution of Paratrytone melane and its spread into San Diego
County. Heppner 10(4):287-300
Two new subspecies of Euphydryas chalcedona from the Mojave desert of
Southern California. Emmel and Emmel 1 1(3): 141-1 46
Altitudinal migration of butterflies in the central Sierra Nevada.
Shapiro 1 2(4):23 1 -235
The butterfly fauna of the Sacramento Valley, California.
Shapiro 13(2):73-82
Extended flight periods of coastal and dune butterflies in California.
Langston 13(2):83-98
A new subspecies of Euphydryas editha from the Channel Islands of
California. Emmel and Emmel 1 3(2): 131-1 36
Altitudinal migration of central California butterflies.
Shapiro 1 3(3): 1 57- 1 6 1
Butterflies of the Suisun Marsh, California. Shapiro 1 3(3): 191 -206
Recent captures of Anthocharis cethura catalina Meadows. Orsak. 1 4(2):8 5-89
Why do California tortoiseshells migrate? Shapiro 14(2):93-97
72
GEOPOLITICAL INDEX
J . Res. Lepid.
California (continued)
Supplementary records of the butterflies in the Sacramento Valley and
Suisun Marsh, lowland central California. Shapiro 1 4(2): 1 00- 1 02
The role of watercress, Nasturtium officinale as a host of native and
introduced pierid butterflies in California. Shapiro 1 4(3): 158-1 68
Urbanus simplicus (Stoll), a new record for California. Tilden . . . 15(1 ):40
Supplementary notes on the distribution of Epargyreus clarus in southern
California (Hesperiidae). Miller 15(4):206-207
Paratrytone melane in San Luis Obispo County, California (Hesperiidae).
Miller 1 6(2): 1 3 1 - 1 32
Autumnal false broods of multivoltine butterflies at Donner Pass,
California. Shapiro 16(2):83-86
The Rhopalocera of Santa Cruz Island, California. Langston . 18(1 ):24-35
A new species of Coloradia in California (Saturniidae, Hemileucinae).
Johnson and Walter 18(1 ):60-66
The ecology and biogeography of the butterflies of the Trinity Alps and
Mount Eddy, Northern California. Shapiro et al 1 8(2):69- 151
Nantucket Pine Tip Moth, Rhyacionia frustrana, in Kern County,
California: Integrated control and biological notes (Lepidoptera:
Tortricidae, Olethreutinae). Poore 19(2):65-67
Notes: A recondite breeding site for the monarch ( Danaus plexippus
Danaidae) in the montane Sierra Nevada. Shapiro 20(1):50-51
Two new California Catocala subspecies (Noctuidae).
Johnson 20(4):245-248
A new species of Mitoura Scudder from southern California (Lepidoptera:
Lycaenidae). Brown , 21(4):245-254
Butterflies of the California Channel Islands. Miller 23(4):282-296
The immature stages of six California Catocala (Lepidoptera: Noctuidae).
Johnson 23(4):303-327
The impact of Pierid feeding on seed production by a native California
crucifer. Shapiro 24(2): 191-1 94
The phenetics and comparative biology of Euphilotes enoptes (Boisduval)
(Lycaenidae) from the San Bernadino Mountains. Pratt and
Ballmer 25(2): 12 1- 1 35
A survey of the last instar larvae of the Lycaenidae (Lepidoptera) of
California. Ballmer and Pratt 27( 1 ): 1 -8 1
The Euphilotes battoides complex: recognition of a species and description
of a new subspecies. Mattoni 27(3): 173-185
Colorado
Euphydryas editha gunnisonensis , a new subspecies from western Colorado.
Brown 9( 1 ):2 1 -23
Ecology and distribution of the butterflies of southern central Colorado.
Scott and Scott 1 7(2):73- 1 28
Florida
A population study of Euptvchia hermes in northern Florida.
Kilduff ' 1 1 (4):2 1 9-228
Georgia
A new species of Narraga (Geometridae, Ennominae) from Georgia, with
biological notes. Coveil et al 23(2): 161-1 68
Hawaii
Butterflies of the Hawaiian Islands according to the stand of late 1976.
Riotte and Uchida 17(1 ):33-39
J. Res. Lepid.
GEOPOLITICAL INDEX
73
Idaho
Euphydryas anicia and E. chalcedonci in Idaho (Lepidoptera: Nymphalidae).
Ferris 26(1-4):109-1 15
Indiana
Habitat associations of wetland butterflies near the Glacial Maxima in
Ohio, Indiana, and Michigan. Shuey 24(2): 1 76- 1 86
Illinois
Euphyes dukesi and other Illinois Herperiidae. Irwin 8(4): 183-1 86
Further notes on Euphyes dukesi. Irwin 1 0(2): 185-188
Kansas
Spring moths of a natural area northeastern Kansas. McFarland . 6( 1 ): 1 - 1 8
Maryland
Eighteen new or scarse butterflies for the state of Maryland. Simmons
and Andersen 9(3): 175-1 84
Notes on Maryland Lepidoptera No. 7, No. 8, and No. 9. Simmons and
Andersen 17(4):253-259
Notes: Notes on Maryland No. 10: Three new butterfly records for the
state of Maryland. Simmons et al 20(4):249
Notes: Notes on Maryland Lepidoptera No. 11: Six new butterflies for the
state of Maryland. Simmons and Andersen 23( 1 ): 1 02- 1 03
Michigan
Habitat associations of wetland butterflies near the Glacial Maxima in
Ohio, Indiana, and Michigan. Shuey 24(2): 176-1 86
Minnesota
Records of Colias gigantea from southwest Manitoba and Minnesota.
Masters 8(3):129-132
Missouri
The habits and life history of Amblyscirtes nysa (Hesperiidae) in Missouri.
Heitzman 3(3):154-156
The life history of Amblyscirtes belli in Missouri. Heitzman . . . 4(l):75-78
An annotated checklist of the Missouri Geometridae.
Heitzman 1 2(3): 1 69- 1 79
Male genitalic illustrations and notes on the Larentiinae (Geometridae) of
Missouri. Heitzman and Enns 1 7(3): 1 45- 1 67
Nebraska
The Butterflies of Nebraska. Johnson 1 1 ( 1 ): 1 -64
Nevada
A new subspecies of Limenitis archippus. Herlan 9(4):2 17-222
Butterflies of Clark County, Nevada. Austin and Austin 1 9( 1 ): 1 -63
A new subspecies of Lycaena editha (Mead) (Lycaenidae) from Nevada.
Austin 23(1 ):83-88
New Hampshire
Population structure of Oeneis melissa semidea (Satyridae) from the
Presidential Range, New Hampshire. Anthony 7(3): 133-1 48
New Jersey
Is air pollution responsible for melanism in lepidoptera and for scarcity of
all orders of insects in New Jersey? Muller 1 0(2): 189-1 90
New York
The ecological associations of the butterflies of Staten Island. Shapiro
and Shapiro 12(2):65-126
North Dakota
North Dakota butterfly calendar. McCabe and Post
1 5(2):93-99
74
GEOPOLITICAL INDEX
J . Res. Lepid.
Ohio
An annotated list of the butterflies for northwestern Ohio.
Porter 4(2):109-1 12
Habitat associations of wetland butterflies near the Glacial Maxima in
Ohio, Indiana, and Michigan. Shuey 24(2): 176-1 86
Oregon
The Argynnis populations of the Sand Creek area, Klamath Co., Oregon,
Part 1. Tilden 1(2):109-1 13
A new subspecies of Callophrys dumetorum from Washington and Oregon.
Gorelick 7(2):99-104
The butterflies of Crater Lake National Park, Oregon. Tilden and
Huntzinger 1 6(3): 176-1 92
Pennsylvania
Melanie tendencies in Phalaenid and Geometrid moths in eastern
Pennsylvania. Shapiro 3( 1 ): 1 9-24
South Dakota
Field study of Phyciodes batesii (Reakirt) and P. tharos (Drury) from a site
in the Black Hills, South Dakota (Lepidoptera: Nymphalidae:
Melitaeinae). Ferris 20(4):235-239
Texas
The genus Panoquina occurring in Texas. Tilden 4(l):37-40
Rhopalocera collected at light in Texas. Kendall and Glick . 10(4):273-283
Lepidopteran foodplant records from Texas. Neck 15(2):75-82
A new subspecies of Hemileuca mala from central Texas (Attacidae,
Hemileucinae). LeMaire 1 8(3):2 1 2-2 1 9
Role of an ornamental plant species in extending the breeding range of a
tropical Skipper to subtropical southern Texas (Hesperiidae).
Neck 20(3): 1 29- 133
Utah
A checklist of the Utah butterflies and skippers. Callaghan and
Tidwell 1 0(3): 191 -202
Addition to: "A checklist of Utah butterflies and skippers". Callaghan and
Tidwell 1 1 (3): 1 99-200
Washington
A new subspecies of Callophrys dumetorum from Washington and Oregon.
Gorelick 7(2):99-104
Wisconsin
Butterfly records for three northwest Wisconsin counties.
Masters 1 1 (3): 175-1 82
A list of the butterflies of the Willow River State Park, Wisconsin.
Masters 14(1 ):57-59
Wyoming
Two new forms of Plebejinae from Wyoming. Ferris 8(3):9 1 -93
A new subspecies of Euphydryas from Wyoming (Nymphalidae).
Ferris 9(l):17-20
A new subspecies of Boloria eunomia from Wyoming. Ferris and
Groothuis 9(4):243-248
SOUTH AMERICA
Brazil
Troidine swallowtails (Lepidoptera: Papilionidae) in southeastern Brazil:
natural history and foodplant relationships. Brown et al. . 1 9(4): 1 99-226
/ . Res. Lepid.
GEOPOLITICAL INDEX
75
Brazil (continued)
Distribution and abundance of butterflies in the urbanization zones of
Porto Alegre, Brazil. Ruszczyk 25(3): 1 57- 178
Chile
Oviposition by the mistletoe-feeding Pierid butterfly Mathania leucothea
(Mol.) in Chile. Courtney 24(3):264-270
Columbia
Illustrations and descriptions of some species of Pyrrhopyginae from
Costa Rica, Panama and Columbia (Hesperiidae). Nicolay and
Small 19(4):230-239
A new genus and two new species of Oecophoridae from Columbia
(Lepidoptera). Clark 20(l):46-49
Venezuela
Heliconius cydno in Venezuela with descriptions for two new subspecies.
Masters 10(4):267-272
MISCELLANEOUS SUBJECT INDEX
[Volumes 1-27, 1962-1988(89)1
Journal of Research on the Lepidoptera
Aberrants, Gynandromorphs, Homoeotics, Melanics, Mutants, etc.
New gynandromorph of Colias philodoce from Colorado. Emmel. 3(l):63-64
A Colias Christina gynandromorph. Hovanitz 4(1):41
Three cases of gynandromorphism in Gonepteryx. Nekrutenko 4(2): 103-1 07
A gynandromorph of Lycaena gorgon. Opler 5(4):230
A melanic form of Pieris rapae. Donahue 6(4):266
A field captured scale-deficient mutant of Anthocharis sara.
Dornfeld 9(l):25-28
Gynandromorphism in Pieris brassicae L. Gardiner 1 1 (3): 1 29- 1 40
A dwarf form of Euptoieta claudia. Rahn 1 1(3): 174
A bilateral gynandromorph of Limenitis weidemeyerii lati fascia
(Nymphalidae). Perkins and Perkins 1 1 (3): 1 95- 1 96
Lack of melanism in Colias (Cover illustration). Masters 1 1 (4):2 1 8
Concerning Heliconius cydno aberration "larseni" Niepelt.
Masters . 1 1(4):251-254
Natural and laboratory occurrence of "Elymi" phenotypes in Cynthia
cardui (Nymphalidae). Shapiro 13(1 ):57-62
Aberrant species of New Jersey Lepidoptera. Muller 1 5(3): 1 44- 1 45
The heathii-whitc banding aberration in the Strymoninae (Lycaenidae).
Fisher 1 5(3): 177-181
Melanic Papilio machaon larvae. Gardiner 1 5(3): 1 84
Speyeria idalia. McCabe 16(1 ):68
Gynandromorphic Polites skippers (Hesperiidae). Nielsen ... 16(4):209-21 1
Gynandromorphs in Hawaiian butterflies and moths. Riotte . 17(1):17-18
A black-backed larval mutant of Lymantria dispar (L.) (Lepidoptera:
Lymantriidae) in Japan. Schaefer and Furuta 1 8(3): 1 67-1 70
An aberrant Oregon Swallowtail, Papilio oregonius Edwards from Oregon.
Wescott 18(4):255
Aberrant New Mexican butterflies. Holland 19(2):88-95
A new record of Vanessa virginiensis "ab. ahwashtee" from northern
California (Lepidoptera: Nymphalidae). Shapiro 20(3): 1 76- 178
Notes: An aberration of Glaucopsyche lygdamus (Lycaenidae) with a
complete Scolitantidine dorsal pattern. Shapiro 20(4):240
Notes: A reared gynandromorph of Tatochila (Pieridae).
Shapiro 20(4):240-242
Two homoeotic Pieris rapae of Mexican origin (Pieridae).
Shapiro 20(4):242-244
An apparent "intersexual" Colias eurytheme (Pieridae). Shapiro . 20(4):244
Parallel albinism in two Theclines (Lycaenidae). Holland 2 1 (3): 1 58
A melanic male aberration of Papilio glaucus canadensis from northern
Wisconsin. Scriber and Lintereur 2 1 (3): 1 99-20 1
Notes: Abnormal chrysalis of Papilio zelicaon (Papilionidae).
Priestaf 21(4):270
A compilation of data on wing homoeosis in Lepidoptera.
Sibatani 22( 1 ): 1 -46
76
J. Res. Lepid .
MISC. SUBJECT INDEX
77
Aberrants, Gynandromorphs, etc. (continued)
Occurrence of the "Elymi" aberrant phenotype in Vanessa carye (Huebner)
(Nymphalidae). Lamas 22(2): 115-117
Compilation of data on wing homoeosis on Lepidoptera: Supplement I.
Sibatani 22(2):1 18-125
Notes: A recessive lethal "wingless" mutation in Tatochila (Pieridae).
Shapiro 22(4):262-263
Notes: Incidence of the black backed larval mutant of Lymantria dispar
(L) (Lepidoptera: Lymantriidae) in Ukrainian SSR. Schaefer et
al 23( 1 ): 1 03- 1 04
Notes: Six homoeotic Vanessa atalanta rubria (Nymphalidae).
Dimock 23(2):176
Stubby-winged mutants of Limenitis (Nymphalidae) - Their occurrence in
relation to photoperiod and population size. Platt 23(3):217-230
Notes: A homoeotic Agraulis vanillae incarnata (Nymphalidae).
Dimock 23(4):332
Notes: A complex gynandromorph of Pontia daplidice (Pieridae).
Shapiro 23(4):332-333
Notes: A melanic Colias euxanthe stuebeli from Peru (Pieridae).
Shapiro 24(1):87
Notes: A bilateral gynandromorph Celastrina ebenina (Lycaenidae). Shuey
and Peacock 24(2):195-196
A new heritable color aberration in the tiger swallowtail butterfly, Papilio
glaucus (Papilionidae: Lepidoptera). Scriber and Evans . . 26(l-4):32-38
Bilateral gynandromorphs, sexual and/or color mosaics in the tiger
swallowtail butterfly, Papilio glaucus (Lepidoptera: Papilionidae).
Scriber and Evans 26(l-4):39-57
A mutant affecting wing pattern in Parnassius apollo (Linne) (Lepidoptera
Papilionidae). Descimon and Vesco 26( 1 -4): 161-172
Notes: Aberrant Polymmatinae (Lycaenidae) from Ohio and Florida.
Calhoun 26(l-4):264-266
Notes: A melanic aberration of Philotes sonorensis (Lycaenidae) from
California. Priestaf 27(3):265-266
Anatomy, Biology, Physiology, etc.
An evaporative cooling mechanism in Pholus achemon (Sphingidae).
Adams and Heath 3(2):69-72
Forelegs of butterflies I. Introduction: Chemoreception. Fox . . . 5( 1 ): 1 - 1 2
Nomenclature of wing veins and cells. Miller 8(2):37-48
Pupal sound production of some Lycaenidae.
Hoegh-Guldberg 1 0(2): 1 27- 147
Survey of ultraviolet reflectance of Neartic butterflies. Scott. 1 2(3): 151-1 60
Lifespan of Butterflies. Scott 12(4):225-230
Antennal sensilla of some Crambinae. Kamm 16(4):201-207
The assumption of adaptivity in genital morphology. Shapiro . 17(1 ):68-72
Ecological color variation in a butterfly and the problem of "protective
coloration". Hovanitz 1 7(S): 1 0-25
Parallel ecogenotypical color variation in butterflies. Hovanitz. 17(S):26-65
The biological and systematic significance of red fecal and meconial
pigments in butterflies: A review with special reference to the Pieridae.
Shapiro 20(2):97-102
Protein and lipid composition of Colias philodice and C. eurytheme
spermatophores and their changes over time. Marshall .... 24( 1 ):2 1 -30
78
MISC. SUBJECT INDEX
J. Res. Lepid.
Behavior, including Courtship and Mating
Evidence for lack of territoriality in two species of Hamadryas.
Ross 2(4):24 1-246
Butterfly aggregations. Reinthal 5( 1 ):5 1 -59
Hilltopping: An ecological study of summit congregation behavior of
butterflies on a southern California hill. Shields 6(2):69- 178
Hilltopping as a mating mechanism to aid survival of low density species.
Scott 7(4): 191 -204
Further observations on "hilltopping" in Papilio zelicaon.
Guppy 8(3): 1 05- 1 17
Mating of butterflies. Scott 1 1 (2):99- 1 27
An interfamilial courtship (Nymphalidae, Pieridae). Shapiro. 1 1(3):197-1 98
A review of carrying pair behavior and mating times in butterflies.
Shields and Emmel 12(l):25-64
Some observations on the eggs of moths and certain aspects of first instar
larval behavior. McFarland 1 2(4): 1 99-208
Mate-locating behavior of the western North American butterflies.
Scott 14(l):l-40
Courtship and mating behavior of the fiery skipper, Hylephila phylaeus
(Hesperiidae). Shapiro 14(3): 125-1 41
Variability of courtship of the buckeye butterfly, Precis coenia
(Nymphalidae). Scott 1 4(3): 1 42- 1 47
The role of intra- and interspecific male:male interactions in Polyommatus
icarus Rott. and some other species of blues (Lycaenidae).
Lundgren 16(4):249-264
Observations on the apparent Lek behavior in Costa Rican rainforest
Perrhybris pyrrha Cramer (Pieridae). DeVries 17(3): 142- 144
Attempted mating between male monarchs. Tilden 18(1 ):2
Territorial behavior of the red admiral, Vanessa atalanta (L.) (Lepidoptera:
Nymphalidae). Bitzer and Shaw 18(1 ):36-49
Notes: Mate locating behavior of Gnophaela latipennis vermiculata G. & R.
(Pericopidae). Scott 20(1):51
Notes: An interfamilial courtship (Lycaenidae - Pieridae). Shapiro. 20(1):54
Courtship behavior of the dainty sulfur butterfly, Nathalis iole with a
description of a new, faculative male display (Pieridae).
Rutowski 20(3):161-169
Plebeian courtship revisited: Studies on the female-produced male
behavior- eliciting signals in Lycaeides idas courtship (Lycaenidae).
Pellmyr 2 1 (3): 1 47- 1 57
Mate-locating behavior of western North American butterflies. II. New
observations and morphological adaptations. Scott 2 1 (3): 177-187
Notes: An effect of the colony edge on gatekeeper butterflies Pyronia
titonus L. (Satyridae). Thomas 21(3):206-207
Notes: Hide and/or seek. Rutkowski 21(3):207
Courtship leading to copulation in the cloudless sulphur, Phoebis sennae
(Pieridae). Rutowski 22(4):249-253
Notes: Lateral perching in Brephidium exilis (Boisduval) (Lycaenidae) in
Texas. Johnson 23( 1 ): 1 04- 1 06
Sexual selection and the evolution of butterfly mating behavior.
Rutowski 23(2):125-142
Butterfly thermoregulation: Organismic mechanisms and population
consequences. Kingsolver 24( 1 ): 1 -20
J. Res. Lepid.
MISC. SUBJECT INDEX
79
Behavior, including Courtship and Mating (continued)
Notes: Mating confusion between a mimic and its model: Erynnis
(Hesperiidae) and Euclidea (Noctuidae). Shapiro 24(l):79-80
Notes: Three intersubfamilial matings in nature (Lycaenidae).
Mattoni 24(l):86-87
Notes: An intersubfamilial courtship (Lycaenidae). Shapiro .... 24(2):195
Notes: Occurrence of homosexual mating pairs in a checkerspot butterfly.
Shaw et al 24(4):393
Male mate-locating behavior in the desert hackberry butterfly,
Asterocampa leilia (Nymphalidae). Rutowski and Gilchrist . 26( 1 -4): 1-12
The mating behavior of Papilio glaucus (Papilionidae). Krebs. 26(l-4):27-31
The mating system of three territorial butterflies in Costa Rica.
Alcock 26(l-4):89-97
The mating system of Vanessa kershawi : males defend landmark territories
as mate encounter sites. Alcock and Gwynne 26( 1 -4): 1 16-124
Notes: Courtship of a model (Nymphalidae: Adelpha) by its probable
Batesian mimic (Nymphalidae: Limenitis). Porter 26(l-4):255-256
Notes: Homosexual pseudocopulation in Eucheira socialis (Pieridae).
Shapiro 27(3):262
Bibliographies
A partial bibliography of the world distribution and zoogeography of
butterflies. Shields 1 3(3): 1 69- 1 78
Moths of North America north of Mexico, Supplemental literature: 1.
Riotte 19(2):68-7 1
Some little-known U. S. publications on Lepidoptera I. Dos
Passos 20(2):1 11-122
Japanese literature. Nakamura 20(2): 127- 128
Japanese literature. Nakamura 20(3): 1 34- 135
Moths of America north of Mexico, supplemental literature: II.
Riotte 22(2): 131-1 34
Bibliography 1982 No. 1. Murphy 23(4):328-331
Bibliography 1982-1983 No. 2. Murphy 24(l):72-75
Bibliography 1983-1984 No. 3. Murphy 24(3):271-275
Bibliography 1984-1985 No. 4. Murphy 26(l-4):248-253
Notes: A bibliography of Euphydryas. Murphy and Weiss . 26( l-4):256-264
Book Reviews
Fox and Fox: Introduction of Comparative Entomology. Hovanitz . 3(1):8
C. F. dos Passos: A Synonymic List of the Neartic Rhopalocera.
Hovanitz 3( 1 ): 1 8
Dos Passos: Synonymic list of Neartic Rhopalocera. Hovanitz . 3( 1 ): 1 9-24
Ronald W. Hodges: The Moths of North America Fasc. 21 Sphingoidae.
Hovanitz 9( 1 ): 1 0
Malcolm Barcant: Butterflies of Trinidad and Tobago. Hovanitz . . 9(1):24
Brown and Heineman: Jamaica and its Butterflies. Hovanitz . . . 1 0(2): 1 48
Harris: Butterflies of Georgia. Urquhart 1 1 (2): 1 28
Annotated check list of the butterflies of Illinois by R. R. Irwin and J. C.
Downey. Funk 12(4):243-244
Butterflies of the World by H. L. Lewis. Ferris 13(4):278-280
Beattie: Rhopalocera directory. Brown 1 5(3): 173-1 75
Bradley, Tremewan and Smith: British Torricoid moths, Cochylidae and
Tortricidae: Tortricinae. Heppner 15(4):208-210
80
MISC. SUBJECT INDEX
J. Res. Lepid.
Book Reviews (continued)
Gilbert: A Compendium of the Biographical Literature on Deceased
Entomologists. Miller 17(3):207
Singh: Artificial Diets for Insects, Mites and Spiders. Morton. 17(3):207-208
Ferguson: The Moths of America North of Mexico. Riotte . . 17(4):260-264
Dornficld: The Butterflies of Oregon. Mattoni 18(1 ):68
Cracraft and Eldredge (ed.): Phylogenetic Analysis and Paleontology.
Shapiro 18(3):220
Herbivores: Their interaction with secondary metabolites. Edited by
Gerald A. Rosenthal and Daniel H. Janzen. Shapiro 19(1 ):64
Miller and Brown: A Catalogue/Checklist of the Butterflies North of
Mexico. Austin 19(4):241-243
Miller and Brown: A Catalogue/Checklist of the Butterflies North of
Mexico. Mattoni 19(4):243-244
Pyle: The Audubon Society Field Guide to North American Butterflies.
Shields 20(1):55
Pyle: The Audubon Society Field Guide to North American Butterflies.
Scott 20(l):55-58
Ferris and Brown eds.: Butterflies of the Rocky Mountain States.
Scott 20(l):58-64
Dowdeswell: The Life of the Meadow Brown. Kudrna 20(3):192
Nelson and Platnick: Systematics and Biogeography: Cladistics and
Vicariance. Shields 21(3):208-209
Hollis (ed.): Animal Identification: A Reference Guide. Vol 3 Insects.
Miller 21(3):209-210
Larsen and Larsen: Butterflies of Oman. Kudrna 2 1 (3):2 1 0
Heath: Threatened Rhopalocera (butterflies) in Europe.
Kudrna 22(2):159-160
Dabrowski: Ginace i zagrozone gatunki motyli (Lepidoptera) w. faune
Polski. Kudrna 22(2):160
Arnold: Ecological studies of six endangered butterflies (Lepidoptera,
Lycaenidae): Island biogeography, patch dynamics, habitat
preservation. Murphy 22(4):267-269
Brooks and Knight: A Complete Guide to British Butterflies.
Kudrna 23(1):108
Whalley: The Mitchell Beazley Pocket Guide to Butterflies.
Kudrna 23( 1 ): 1 08- 1 09
Bjorn: The Butterflies of Northern Europe. Kudrna 23( 1 ): 1 09- 110
Barlow: An Introduction to the Moths of South East Asia.
Peigler 23(1):1 10-111
Eliot and Kawazoe: Blue Butterflies of the Lycaenopsis - Group.
Mattoni 23( 1 ): 1 1 1-112
Young: Population Biology of Tropical Insects. Shapiro 23( 1 ): 1 1 2
Blab and Kudrna: Hilfsprogramm fur Schmetterlinge. Okologie und
Schutz von Tagfaltern und Widderchen. Murphy 23(2): 1 69- 170
D’Abrera: Butterflies of South America. Emmel 23(2): 171-172
D’Abrera: Butterflies of South America. Shapiro 23(2): 172-1 74
Futuyma: Coevolution. Shields 23(2): 174
Cater (ed): Love Among the Butterflies: The Travels and Adventures of a
Victorian Lady. Murphy 23(4):335-336
J. Res. Lepid.
MISC. SUBJECT INDEX
81
Book Reviews (continued)
A critical review of "Systematische Untersuchungen am Pieris
napi-bryonicie- Komplex (s.l.)" Lepidoptera: Pieridae) by Ulf
Eitschberger. Kudrna and Geiger 24(l):47-60
Miyata: Handbook of Moth Ecology. Mattoni 24(1):88
King and Leppla: Advances and Challenges in Insect Rearing.
Mattoni 24(1):88
Hafernik: Phenetics and Ecology of Hybridization in Buckeye Butterflies
(Lepidoptera: Nymphalidae). Shapiro 24(1):89-91
Opler and Krizek: Butterflies East of the Great Plains: An Illustrated
Natural History. Shapiro 24( 1 ):9 1-93
Larsen: Butterflies of Saudi Arabia and its Neighbors. Kudrna. 24(l):93-94
Kock: Wirbestimmen Schmetterlinge. Kudrna 24(1):94
Hodges et al.: Checklist of the Lepidoptera North of Mexico.
Murphy 24(l):95-96
Landing: Factors in the Distribution of Butterfly Color and Behavior
Patterns - Selected Aspects. Kudrna 24(4):375-376
Landing: Factors in the Distribution of Butterfly Color and Behavior
Patterns - Selected Aspects. Murphy 24(4):375-376
Bridges: Lepidoptera: Hesperiidae. Notes on Species - Group Names. De
Jong 24(4):379-381
Pyle: The Audubon Society Handbook for Butterfly Watchers. Ehrlich
and Murphy 24(4):381-382
Sbordioni and Forestiero: The World of Butterflies and II Mondo delle
Farfalle. Kudrna 24(4):382-383
Tarmann: Generic revision of the American Zygaenidae, with descriptions
of New Genera and Species. Heppner 24(4):383-384
Vane-Wright and Ackery: The Biology of Butterflies. Gall . . 25(2): 1 49- 155
Kudrna: Butterflies of Europe. Ferris 25(2): 155
Tekulsky: The Butterfly Garden. Mattoni 25(2): 1 56
Menke and Miller: Entomology of the California Channel Islands.
Mattoni 25(2): 1 56
Scott: The Butterflies of North America. A Natural History and Field
Guide. Opler 26(l-4):267-270
Scott: The Butterflies of North America. A Natural History and Field
Guide. Gall 26(l-4):270-275
Scott: The Butterflies of North America. A Natural History and Field
Guide. Murphy and Baughman 26(l-4):275-278
Tilden and Smith: A Field Guide to Western Butterflies.
Austin 26(l-4):278-283
Mani: Butterflies of the Himalaya. Shapiro 26(1 -4):283-285
Friedrich: Breeding of Butterflies and Moths. Kudrna . . . 26(l-4):285-286
Erhardt: Wiesen und Brachland als Lebensraum fur Schmetterlinge.
Shapiro 26(l-4):286-287
Erlich: The Machinery of Nature. Mattoni 26(l-4):287-288
Weidemann: Tagfalter 1. H-J. Kudrna 26(l-4):288
McKibben: The End of Nature. Mattoni 27(3):271
McFarland: Portraits of South Australian Geometrid Moths.
Mattoni 27(3):272
Emmet and Heath: The Moths and Butterflies of Great Britian and
Ireland. Vol 7, Part 1, Hesperiidae - Nymphalidae, The Butterflies.
Mattoni 27(3):272-274
82
MISC. SUBJECT INDEX
J. Res. Lepid.
Book Reviews (continued)
Nielsen and Kristensen: Primitive Ghost Moths. Mattoni 27(3):274
Schwartz: The Butterflies of Hispaniola. DeVries 27(3):274-276
Climatological Considerations
The climatological tool in lepidoptera research. Crowe 4(l):23-36
Present and Ice Age Life Zones and distributions. Hovanitz . . 7( 1 ):3 1 -34
Spring moth activity in relation to locality, temperature, and air pressure.
Selman and Barton 9( 1 ): 1 -9
Extended flight periods of coastal and dune butterflies in California.
Langston 13(2):83-98
Conservation and related issues
Is air pollution responsible for melanism in lepidoptera and for scarcity of
all orders of insects in New Jersey? Muller 1 0(2): 1 89- 1 90
Editorial: Extinction of the British Large Blue Butterfly. Mattoni. 1 8( 1 ): 1 , 3
Conservation and management of the endangered Smith’s Blue Butterfly,
Euphilotes enoptes smithi (Lepidoptera: Lycaenidae).
Arnold 22(2): 1 35- 1 53
The decline and extinction of Speyeria populations resulting from human
environmental disturbances (Nymphalidae: Argynninae). Hammond
and McCorkle 22(4):217-224
Pupal mortality in the Bay checkerspot butterfly (Lepidoptera:
Nymphalidae). White 25(l):52-62
Opinion: Are we studying our endangered butterflies to death?
Murphy 26(l-4):236-239
Diapause
The relationship between migration and diapause during phylogeny and
ontogeny of some Lepidoptera. Novak and Spitzer 1 0(2): 181-1 84
A proposed terminology for the types of diapause occurring in the order
Lepidoptera. Tilden 15(1 ):33-39
Hibernal diapause of North American Papilionoidea and Hesperioidea.
Scott 1 8(3): 1 7 1 -200
Diapause in various populations of Pieris napi L. from different parts of
the British Isles. Lees and Archer 19(2):96-100
Records of prolonged diapause in Lepidoptera. Powell 25(2):83-109
Evolution, Taxonomy, Nomenclature, etc.
Tertiary Nymphalid butterflies and some phylogenetic aspects of
systematic lepidopterology. Nekrutenko 4(3): 1 49- 158
Comparitive speciation in two butterfly families: Pieridae and
Nymphalidae. Petersen 5(2): 113-1 26
Fossil butterflies and the evolution of Lepidoptera. Shields . 1 5(3): 1 32- 1 43
Butterfly nomenclature: A critique. Ehrlich and Murphy 20( 1 ): 1 - 1 1
Butterfly taxonomy: A reply. Miller and Brown 20(4):193-198
Nomenclature, taxonomy, and evolution. Ehrlich and
Murphy 20(4):199-204
Taxonomic uncertainty, the biological species concept, and the Neartic
butterflies: a reappraisal after twenty years. Shapiro .... 2 1 (4):2 1 2-2 1 8
Butterflies and biospecies. Ehrlich and Murphy 21(4):219-225
The biological species concept and the aims of taxonomy. De
Jong 21(4):226-237
Commentary on Miller and Brown vs. Erhlich and Murphy et al .: Pluralism
in systematics and the worldwide nature of kinship groups. Johnson
and Quinter 21(4):255-269
J. Res. Lepid.
MISC. SUBJECT INDEX
83
Evolution, Taxonomy, Nomenclature, etc. (continued)
Opinion: Crows, bobs, tits, elfs and pixies: The phoney "common name"
phenomenon. Murphy and Ehrlich 22(2): 1 54- 158
On butterfly taxonomy. Murphy and Ehrlich 23( 1 ): 1 9-34
Opinion: Rebuttal to Murphy and Ehrlich on common names of
butterflies. Pyle 23(l):89-93
The phylogeny of butterflies (Papilionoidea and Hesperioidea).
Scott 23(4):241-281
A butterfly-moth (Lepidoptera Castniidae) from the Oligocene shales of
Florissant, Colorado. Tindale 24( 1 ):3 1 -40
Opinion: A rebuttal to the Arnold classification of Speyeria callippe
(Nymphalidae) and defense of the subspecies concept.
Hammond 24(3):197-208
Opinion: Parallelism and phylogenetic trees. Scott 27(3):257-258
False Broods
Origin of autumnal "false broods" in common Pierid butterflies.
Shapiro 6(3):181-193
Autumnal false broods of multivoltine butterflies at Donner Pass,
California. Shapiro 16(2):83-86
Flight Periods
The flight periods of several sibling species of moths.
Selman 12(4):217-224
Food Plants
Three factors affecting larval choice of food plant. Hovanitz and
Chang 1(1 ):5 1-61
The effect of hybridization of host-plant strains on growth rate and
mortality of Pieris rapae. Hovanitz and Chang 1 (2): 157-1 62
Butterfly larval foodplant records and a procedure for reporting
foodplants. Shields et al 8( 1 ):2 1 -36
Botanical names in entomological papers and habitat studies.
McFarland 9(2):89-96
Larval foodplant records for North American Rhopalocera. Part 2. Emmel
et al 9(4):233-242
The role of watercress, Nasturtium officinale as a host of native and
introduced pierid butterflies in California. Shapiro 1 4(3): 1 58- 1 68
Biochemical studies of the larval hosts of two species of Lycaena Fabricius
(Lycaenidae). Ferris 17(1 ):27-32
Role of an ornamental plant species in extending the breeding range of a
tropical Skipper to subtropical southern Texas (Hesperiidae).
Neck 20(3):129-133
Notes: Oviposition records and larval foodplants of butterflies in the
Atlas Mountains of Morocco. Thomas and Mallorie 24(l):76-79
The impact of Pierid feeding on seed production by a native California
crucifer. Shapiro 24(2): 191-1 94
Notes: Moss feeding by a Satyrine butterfly. Singer and Mallet . 24(4):392
Genetics, including Electrophoretic Studies
Methods for studying the chromosomes of lepidoptera. Emmel . 7(1 ):23-28
Genetic control of maculation and hindwing color in Apantesis phalerata.
Bacheler and Emmel 13(l):49-56
The chromosomes of Apantesis phalerata , A. radians , and their hybrid in
Florida populations (Arctiidae). Bacheler and Emmel . . . 1 3(3): 1 62- 1 68
84
MISC. SUBJECT INDEX
J . Res. Lepid.
Genetics, including Electrophoretic Studies (continued)
The genetics of subspecific phenotype differences in Pieris occidentalis
Reakirt and of variation in P. o. nelsoni W. H. Edwards (Pieridae).
Shapiro 14(2):61-83
Chromosome numbers in two species of Ergolis (Lepidptera: Nymphalidae).
Murty and Rao 15(l):23-26
Note on the chromosomes of Byblia ilithyia (Drury) (Nymphalidae). Murty
and Rao 1 5(3): 1 29- 1 3 1
Investigation of selected species of the genus Orgyia (Lymantriidae) using
isoelectrofocusing in thin layer polyacrylamide gel. Chua et
al 1 5(4):2 1 5-224
On the meiotic chromosomes of Argina stringa Cram (Arctiidae). Rao and
Lakshmi . 17(l):51-52
A study of the meiotic chromosomes of Ixias marianne (Cramer) (Pieridae).
Rao 1 7(3): 170-172
Asynaptic meiosis in three species of Lepidopteran males.
Nayak 17(4):240-244
Chromosome studies including a report of B-chromosome in a wild
silkmoth, Sonthonnaxia maenas (Doubleday) (Saturniidae: Saturniinae).
Narang and Gupta 1 8(3):208-2 1 1
Enzyme electrophoretic studies on the genetic relationships of Pierid
butterflies (Lepidoptera: Pieridae) I. European taxa.
Geiger 1 9(4): 1 8 1 - 1 95
Karyology of three Indian Lasiocampid moths (Lepidoptera). Mohanty
and Nayak 19(4):227-229
Chromosome studies in sixteen species of Indian Pyralid moths
(Pyralidae). Mohanty and Nayak 20(2):86-96
On the supernumerary chromosomes of Tarache tropica Guen.
(Lepidoptera: Noctuidae). Mohanty and Nayak 20(3): 170-173
Chromosomes of seven species of Indian Sphingid moths. Mohanty and
Nayak 21(4):238-244
Allozyme variation in a colonizing species: The cabbage butterfly Pieris
rapae (Pieridae). Vawter and Brussard 22(3):204-216
Karyotypes of some Indian Noctuid moths (Lepidoptera). Mohanty and
Nayak 22(4):238-248
Polyphenism, phyletic evolution, and the structure of the Pierid genome.
Shapiro 23(3): 1 7 7- 1 96
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera). Lorkovic 24(4):334-358
Electrophoretic evidence for speciation within the nominal species
Anthocharis sara Lucas (Pieridae). Geiger and Shapiro .... 25( 1 ): 1 5-24
Genetic differentiation between subspecies of Euphydryas phaeton
(Nymphalidae: Nymphalinae). Vawter and Wright 25(1 ):25-29
Electrophoretic confirmation of the species status of Pontia protodice and
P. occidentalis (Pieridae). Shapiro and Geiger 25(l):39-47
Enzyme electrophoresis and interspecific hybridization in Pieridae
(Lepidoptera) - The case for enzyme electrophoresis.
Geiger 26(l-4):64-72
Correlations of ultrastructure and pigmentation suggest how genes control
development of wing scales of Heliconius butterflies. Gilbert et
al 26( 1 -4): 141-1 60
J . Res. Lepid .
MISC. SUBJECT INDEX
85
Genetics, including Electrophoretic Studies (continued)
Notes: A chromosome study of Brahmaea japonica Butler (Lepidoptera:
Brahmaeidae). Trentini and Marini 27(2): 1 36- 138
Genetic experiments with a calverleyi-Vikc mutation isolated from Papilio
bairdi oregonius (Papilionidae). McCorkle and Hammond . 27(3): 1 86- 191
Historical Interest Items
On the Gunder collection of Argynnids. Grey 8(2):55-64
Old Timers. Comstock 14(2):90-92
Dr. William Hovanitz, 1915-1977. Mattoni 17(S):l-6
An annotated catalogue of the butterflies (Lepidoptera: Paplionoidea)
named by Roger Verity. Kudrna 21(1):1-1 06
An annotated catalogue of the Skippers (Lepidoptera: Hesperiidae) named
by Roger Verity. Kudrna and Balletto 23(l):35-49
An annotated catalogue of the Burnets and Foresters (Lepidoptera:
Zygaenidae) named by Roger Verity. Balletto and
Kudrna 24(3):226-249
Hybrids and Hybridization
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 1 (4):26 1-274
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 2(3):205-223
The origin of a sympatric species in Colias through the aid of natural
hybridization. Hovanitz 3(l):37-44
Hybrids between Papilio memnon and Papilio protenor. Ae 3(l):55-62
A hybrid Limenitis from New York. Shapiro and Biggs 7(3): 1 49- 1 52
A possible new hybrid copper. Crowe 8(2):5 1 -52
On the occurrence of Limenitis archippus X L. lorquini hybrids. Perkins
and Gage 9(4):223-226
Limenitis weidemeyerii angusti fascia X L. astyanax arizonensis = (?) ab.
doudoroffi (Gunder) 1934. Perkins and Garth 1 1(4):229-234
Records of Limenitis hybrids from Colorado. Simpson and
Pettus 1 5(3): 1 63- 1 68
The use of alpha-ecdysone to break permanent diapause of female hybrids
between Papilio glaucus L. female and Papilio rutulus male. Clarke and
Willig 16(4):245-248
Another Anthocharis lanceolata X A. sara hybrid. Shields and
Mori 17(l):53-55
An apparent Interspecific Fj Hybrid Speyeria (Nymphalidae).
Scott 20(3): 174-175
Hand-pairing of Papilio glaucus glaucus and Papilio pilumnus
(Papilionidae) and hybrid survival on various food plants. Scriber and
Lederhouse 27(2):96-103
Hybridization of the Mexican tiger swallowtail, Papilio alexiares garcia
(Lep: Pap) with other P. glaucus group species and survival of pure and
hybrid larvae on potential host plants. Scriber et al 27(3):222-232
Migration
The relationship between migration and diapause during phylogeny and
ontogeny of some Lepidoptera. Novak and Spitzer 1 0(2): 181-1 84
Altitudinal migration of butterflies in the central Sierra Nevada.
Shapiro 1 2(4):23 1 -235
Altitudinal migration of central California butterflies.
Shapiro
1 3(3): 1 5 7- 1 6 1
86
MISC. SUBJECT INDEX
J. Res. Lepid.
Migration (continued)
Toward a theory of butterfly migration. Shields 1 3(4):2 17-238
Larval migration of Hyles linecita (Fab.). Wells and Brown 13(4):246
Why do California tortoiseshells migrate? Shapiro 14(2):93-97
Migration of Hipparchia semele L. Feltwell 15(2):83-91
Notes: Further migration of Hipparchia semele (L.) in 1976 and 1980.
Feltwell and Ducros 20(1):53
Notes: An early season migration of Catopsilia pomona (Lepidoptera:
Pieridae) in Java, Indonesia. New 24(l):84-85
Mimicry
Antepione thisaria and Xanthotype : A case of mimicry. Shapiro . . 4( 1 ):6- 1 1
Notes: Mating confusion between a mimic and its model: Erynnis
(Hesperiidae) and Euclidea (Noctuidae). Shapiro 24(l):79-80
A tropical caterpillar that mimics faeces, leaves and a snake (Lepidoptera:
Oxytenidae: Oxytenis naemia). Nentwig 24(2): 1 36- 141
Mimicry by illusion in a sexually dimorphic, day-flying moth, Dysschema
jansonis (Lepidoptera: Arctiidae: Pericopinae). Aiello and
Brown 26( 1 -4): 1 73- 1 76
Notes: Courtship of a model (Nymphalidae: Adelpha) by its
probableBatesian mimic (Nymphalidae: Limenitis).
Porter 26(l-4):255-256
New Taxa Descriptions
The Annaphila astrologa complex, with descriptions of three new species.
Sala 2(4):289-301
A new species of Rioninidae from Mexico. Clench 3(2):73-79
Petaluma , a new genus. Buckett and Bauer 3(3): 1 93- 1 96
California coastal Eupithecia with description of new species
(Geometridae). Leuschner 4(3): 1 9 1 - 1 97
New Canadian species of leaf-mining lepidoptera of conifers.
Freeman 4(3):209-220
A new species of Basilodes from the southwestern United States
(Noctuidae). Hogue 4(4):275-280
A new species of Epinotia Hubner from British Columbia (Olethreutidae).
Freeman 5( 1 ): 1 3- 1 4
A new species of Oncocnemis from the western United States (Noctuidae:
Cuculliinae). Buckett and Bauer 5(4):197-208
A new species of Polia Ochsenheimer from California and notes on Polia
discalsis (Grote) (Noctuidae: Hadeninae). Buckett and
Bauer 5(4):221-228
A new species of Nepticula on bur oak in Ontario (Nepticulidae).
Freeman 6( 1 ): 1 9-2 1
Description of a new species of Xylomiges from California.
Buckett 6(l):23-30
A new species of Feralia. Buckett 6(1):43-51
A new species of armyworm - genus Faronta. Buckett 6(4):268-274
A previously unrecognized subspecies of Philotes speciosa.
Tilden 6(4):281-284
On Mexican Satyridae with description of a new species. Miller. 7(1 ):5 1-55
Identity of the moth "Stretchia" behrensiana with new synonymy
(Noctuidae). Buckett 7(1 ):57-63
A new subspecies of Callophrys dumetorum from Washington and Oregon.
Gorelick 7(2):99-104
J. Res . Lepid.
MISC. SUBJECT INDEX
87
New Taxa Descriptions (continued)
Two new forms of Plebejinae from Wyoming. Ferris 8(3):9 1 -93
A new subspecies of Euphydryas from Wyoming (Nymphalidae).
Ferris 9(l):17-20
Euphydryas editha gunnisonensis , a new subspecies from western Colorado.
Brown 9(l):21-23
Revision of the Neartic genus Philtraea Hulst with notes on biology and
description of new species (Geometridae). Buckett 9(l):29-64
Synaxis mosesiani Sala; a new Synaxis from southern California.
Sala 9(3): 185-191
A new Parnassius eversmanni race from northeast Siberia (USSR).
Weiss 9(4):215-216
A new subspecies of Limenitis archippus. Herlan 9(4):2 17-222
A new subspecies of Boloria eunomia from Wyoming. Ferris and
Groothuis 9(4):243-248
Variation of Uthetheisa ornatrix (Arctiidae) including a new species from
St. Croix, Virgin Islands. Pease 1 0(4):26 1 -264
Heliconius cydno in Venezuela with descriptions for two new subspecies.
Masters 10(4):267-272
Two new subspecies of Euphydryas chalcedona from the Mojave desert of
Southern California. Emmel and Emmel 1 1 (3): 141-1 46
Variations of Parasemia parthenos. Brower 1 1 (3): 1 83- 1 86
A new subspecies of Gonepteryx amintha (Pieridae) from Yunnan,
mainland China. Nekrutenko 1 1(4):235-240
A new species of Papilio from the eastern United States (Papilionidae).
Heitzman 12(1):1-10
A new species of Hypagyrtis (Geometridae). Heitzman 13(1 ):43-48
A new species of Ormiscodes (Dirphiella) from Mexico (Saturniidae:
Hemileucinae). Donahue and LeMaire 13(2):123-130
A new subspecies of Euphydryas editha from the Channel Islands of
California. Emmel and Emmel 1 3(2): 131-1 36
Descriptions of a new species of Eupithecia and the male of E. cocoata
Pearsall (Geometridae). Heitzman and Enns 16(2):75-82
A new subspecies of Colias palaeno (Linnaeus) from Baffin Island, N.W.T.,
Canada (Pieridae). Ebner and Ferris 1 6(3): 155-161
The Scolitantidini I: Two new genera and a generic rearrangement
(Lycaenidae). Mattoni 16(4):223-242
A new species of Coloradia in California (Saturniidae, Hemileucinae).
Johnson and Walter 18(1 ):60-66
A new subspecies of Hemileuca maia from central Texas (Attacidae,
Hemileucinae). LeMaire 1 8(3):2 1 2-2 1 9
A new species of Automeris cecrops (Attacidae: Hemileucinae).
LeMaire 18(4):236-240
The Scolitantidini II. The World’s smallest butterfly? Notes on Turanana ,
and a new genus and species from Afghanistan (Lycaenidae).
Mattoni 18(4):256-264
A new Tortyra from Cocos Island, Costa Rica (Lepidoptera: Choreutidae).
Heppner 1 9(4): 1 96- 1 98
A new genus and two new species of Oecophoridae from Columbia
(Lepidoptera). Clark 20(l):46-49
A new species of Adelpha (Nymphalidae) from Parque Nacional Braulio
Carrillo, Costa Rica. DeVries and Gamboa 20(2):123-126
88
MISC. SUBJECT INDEX
J. Res. Lepid.
New Taxa Descriptions (continued)
Two new California Catocala subspecies (Noctuidae).
Johnson 20(4):245-248
A review of the Erebici dabanensis complex (Lepidoptera: Satyridae), with
descriptions of two new species. Troubridge and Philip . . 2 1 (2): 1 07- 1 46
A new species of Mitoura Scudder from southern California (Lepidoptera:
Lycaenidae). Brown 21(4):245-254
A new Limenitis weidemeyerii W. H. Edwards from southeastern Arizona
(Nymphalidae). Austin and Mullins 22(4):225-228
Biosystematics of the Euphydryas of the central Great Basin with the
description of a new subspecies. Murphy and Ehrlich . . . 22(4):254-261
A new subspecies of Lycaena editha (Mead) (Lycaenidae) from Nevada.
Austin 23(l):83-88
A new species of Narraga (Geometridae, Ennominae) from Georgia, with
biological notes. Coveil et al 23(2): 161-1 68
A review of Polygonia progne (oreas) and P. gracilis ( zephyrus)
(Nymphalidae) including a new subspecies from the southern Rocky
Mountains. Scott 23(3):197-210
Discovery of two new species and genera of Shaggy Tortricids related to
Synnoma and Niasoma (Tortricidae: Sparganothini). Powell . 24( 1 ):6 1-71
A new squash borer from Mexico (Lepidoptera: Sesiidae).
Friedlander 24(4):277-288
A new species of Calisto from Hispaniola with a review of the female
genitalia of Hispaniolan congeners (Satyridae). Johnson et
al 25(2):73-82
A new genus and species from the southwestern United States (Noctuidae:
Acontiinae). Brown 25(2): 1 36- 1 45
Apodemia palmerii (Lycaenidae: Riodininae): Misapplication of names, two
new subspecies and a new allied species. Austin 26(1-4):125-140
New species and new nomenclature in the American Acronictinae
(Lepidoptera: Noctuidae). Ferguson 26( 1 -4):20 1-218
The morpho-species concept of Euphyes dion with the description of a new
species (Hesperiidae). Shuey 27(3): 1 60- 172
The Euphilotes battoides complex: recognition of a species and description
of a new subspecies. Mattoni 27(3):173-185
Three new species of Paradriphia (Saturniidae: Hemaleucinae) from
Mexico and Central America with notes on the immature stages.
LeMaire and Wolfe 27(3): 1 97-2 1 2
Opinions
Did the caterpillar exterminate the giant reptile? Flanders . . . 1(1 ):85-88
Caterpillar versus dinosaur? Eaton 1 (2): 1 14-1 16
Kloet and Hincks’ Check list of British Lepidoptera Insects (Lepidoptera)
Edn. 2. A reply to criticisms. Bradley et al 13(4):265-266
Editorial. Mattoni 16(4):243-244
Opinion: Patronyms in Rhopaloceran nomenclature. Dimock . 23(1 ):94- 101
Opinion: Rebuttal to Murphy on factors to the distribution of butterfly
color and behavior patterns - selected aspects. Landing . . . 25(l):67-70
Opinion: The trouble with butterflies. White 25(3):207-212
A response to Landing: On factors in the distribution of butterfly color
and behavior. Murphy 25(3):2 1 3-2 1 4
Opinion: Reply to Scott’s criticism. Brock 26(l-4):240-247
J. Res. Lepid.
MISC. SUBJECT INDEX
89
Population Study Techniques
Techniques in the study of population structure in Philotes sonorensis.
Mattoni and Seiger l(4):237-244
Estimating the density of an animal population. Hanson .... 6(3):203-247
Trials of several density estimators on a butterfly population. Hanson
and Hovanitz 7(l):35-49
The study of populations of Lepidoptera by capture-recapture methods.
Sheppard and Bishop 1 2(3): 135-1 44
An area census method for estimating butterfly population numbers.
Douwes 1 5(3): 1 46- 152
Colias alexandra : A model for the study of natural population of
butterflies. Hayes 23(2): 1 13-124
Measuring the size of Lepidopteran populations. Gall 24(2):97- 116
Predation
Note on damaged specimens. Kolyer 7(2):105-111
Observations of predation on Lepidoptera in Alaska. Tilden . . . 1 5(2): 1 00
Interactions of parasitoids and checkerspot caterpillars Euphydryas spp.
(Nymphalidae). Stamp 23( 1 ):2- 1 8
Egg mass design relative to surface-parasitizing parasitoids, with notes on
Asterocampa clyton (Lepidoptera: Nymphalidae).
Friedlander 24(3):250-257
Techniques, including Collecting, Rearing, Laboratory, etc.
A standard method for mounting whole adult lepidoptera on slides
utilizing polystyrene plastic. Hogue l(3):223-235
A rubber stamp method for producing specimen labels. Adams. 2(3):225-228
Decapitation-initiated oviposition in Crambid moths. Crawford . 3(l):5-8
The feeding of coloring matters to Pier is rapae larvae. Kolyer.4(3):159-172
A moth sheet. McFarland 5(l):29-36
Laboratory techniques for maintaining cultures of the monarch butterfly.
Urquhart and Stegner 5(3): 1 29- 1 36
Vital staining of Colias philodoce and C. eurytheme. Kolyer . . 5(3): 137- 152
Methods for studying the chromosomes of lepidoptera. Emmel 7(1 ):23-28
Rearing techniques for speeding up larval stages of some root or
stem-boring Lepidoptera. McFarland 7(3): 1 66
Notes on describing, measuring, preserving and photographing the eggs of
lepidoptera. McFarland 10(3):203-214
Notes on Artie and Subartic collecting. Ferris 13(4):249-264
A rapid method for producing insect labels. Adams 1 5(3): 1 69- 1 72
Rearing butterflies on artificial diets. Morton 1 8(4):22 1 -227
Portable apparatus for photographing genitalic dissections.
McCabe 27(2):1 15-119
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THE JOURNAL
OF RESEARCH
ON THE LEPIDOPTERA
Volume 28
Number 1-2
Spring/Summer 1989 (1990)
THE JOURNAL OF RESEARCH
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Journal of Research on the Lepidoptera
28(1-2):1-13, 1989(90)
Mating Behavior and Male Investment in Euphydryas
anicia (Lepidoptera: Nymphalidae)
Francois J. Odendaal,1,4
Kristina N. Jones,2
and
Frank R. Stermitz3
Abstract. The size of male ejaculates in butterflies have often been
suggested to play a key role in shaping the characteristics of butterfly
mating systems. Females might choose males with larger ejaculates
(greater nutrient investment), but a large ejaculate may be more costly
for the male to produce. It remains unclear how various factors may
interact to determine the size of male ejaculates. Rutowski (1984)
called for tests over a range of species, including ones with unusually
large or small nutrient investments. We studied mating behavior in
Euphydryas anicia , which has an unusually small spermatophore
(<2% of body weight), and suggest that spermatophore size relates
strongly to many aspects of the mating system: male and female
choosiness, female mating frequency, and the length and complexity of
courtship.
Introduction
In some butterflies males transfer sperm and nutrients through
mating to females (eg. Boggs and Gilbert 1979, Engebretson and Mason
1980, Walker 1980, Boggs 1981, Boggs and Watt 1981). A range of sizes
exists for male ejaculates among species (Rutowski et al. 1983), and it
has been suggested by many authors (for review, see Thornhill and
Alcock 1983) that the size of the male nutrient investment plays a key
role in shaping the characteristics of the mating system. Females would
presumably perfer males with larger nutrient investments. Rutowski
(1984) cautions that “untested assumptions underlie this conclusion”
and that “students of the mating behavior of butterflies and moths
should pay special attention to species whose males . . . produce [un]-
usually large or small nutrient investments”. One major untested
assumption, for instance, is that offspring benefit from the nutrients
transferred at mating. In studies on a moth (Greenfield 1982) and a
butterfly (Jones et al. 1986), it was found that female fecundity was in
department of Zoology, Duke University, Durham, North Carolina 27706
department of Biology, University of California, Davis, California
department of Chemistry, Colorado State University, Fort Collins, CO 80523
4Present address: Zoology Department, University of Cope Town, Rondebosch, South
Africa 7700
2
J. Res. Lepid.
fact not a function of the size of male ejaculate weight. Without a
positive correlation between male ejaculate size and female fecundity, it
is difficult to argue that males with larger spermatophores are more
attractive to females.
Male ejaculate size may also affect the mating system through
male choice. Recent evidence suggests that males incur non-trivial
costs in producing ejaculates, that the number of ejaculates they can
produce is limited, and therefore some discrimination should be ex-
pected with respect to their patterns of allocating ejaculates (eg.
Marshall 1982, Dewsbury 1982, Svard 1985, Svard and Wilklund
1986). Rutowski (1985) studied choice in Colias eurytheme males which
pass about 6% of their body weight (cf. Rutowski et al. 1983) to females
at mating, and found that the size of females accepted by males is less
variable than that of rejected females, implying choice by males. We
studied the behavior of a butterfly with a very small spermatophore
(<2% of male body weight), Euphydryas anicia. By following individual
Euphydryas anicia in the field we were able to collect data on mate
location behavior, mate discrimination, and courtship complexity and
duration. Dissection of mated females gave information on sperma-
tophore size and mating frequency. Our results support the prediction
by Svard (1985) that males of monandrous species will tend to have
small spermatophores, and may be compatible with those of Rutowski
(1985) because of the difference in the size of the investment by the
males of the two species studied. Like Rutowski’s study, ours may
support the notion that the size of the investment may be reflected in
the degree of male choosiness. We discuss possible reasons for the small
spermatophore in E. anicia , and discuss how a small spermatophore
relates to other characteristics of the mating system.
Materials and Methods
Euphydryas anicia Doubleday and Hewitson occurs in scattered populations
over the western half of the United States (White 1979, Cullenward et al. 1979,
Ferris and Brown 1981). Near Red Hill Pass, 11 km east of Fairplay, Park
County, Colorado, a population occurs in a part of a very flat, high altitude
(2,900 m) intermontane plain (Odendaal et al. 1988). This site is an approximately
700 x 1300 m area containing a relatively localized population of E. anicia. The
flight season is very short. Several years of site observations show that after snow
has melted (early to mid-May) postdiapause larave feed on Besseya plantaginea and
Castilleja integra (Scrophulariaceae), pupate, and emerge as adult butterflies in
mid- June. About four to five weeks pass from first emergence to the last
butterfly. For the first several days there are only males as occurs with other
Euphydryas (Iwasa et al. 1983). Butterflies from this site contain bitter iridoid
glucosides sequested during larval consumption of the host plants (Stermitz et
al. 1986, Gardner and Stermitz 1988). Host and nectar plants are distributed
throughout the site, although patchily in some years. All plants on the site are
low-growing and herbaceous, with consequent excellent visibility for observing
and censusing larvae, pupae and adult butterflies. During the 1985 flight season
we conducted a mark-release-recapture program, capturing and marking 1260
28(1-2):1-136, 1989(90)
3
males and 671 females. The first males were observed on June 13 and the first
females on June 17, with an approximate 50:50 ratio reached on June 27.
Behavior was observed during the entire flight season and some additional
observations were conducted during the 1986 season.
MALE MATING BEHAVIOR
Individual field-collected males were marked and followed by two researchers,
one recording behavioral events with a hand-held Radio Shack TRS 80
computer while the other marked landing spots. Males were recorded as
collecting nectar, sitting, flying and chasing virgin females, previously mated
(plugged) females, other males or other species.
FEMALE MATING BEHAVIOR
Because lab-rearing or tethering virgins may alter their behavior (Odendaal,
unpublished data; M. Singer, personal communication), we followed unre-
strained field-hatched virgins. Thirty-one virgins were encountered flying or
just emerging from pupae in the field. Some were followed immediately, others
were caught and released later the same day, while still others were kept for up
to two days in a cage and fed with a mixture of honey and water (see Jones et al.
1986). One or two observers followed each virgin, noting and timing each type of
behavior, and marking each landing spot with a numbered flag when possible. As
a measure of relative local male density at various landing spots, all males
passing within 1.5 m of a sitting virgin were counted. The same procedure was
repeated with twelve field-collected plugged females.
Behavioral data for females were analyzed with an ANOVA to show whether
a significant amount of variance in each of the parameters (flight frequency,
average flight distance, local male density, number of male chases and chase
distance) could be explained by female type (previously mated or plugged
females, virgins that mated as a result of a male chase, and virgins that did not
mate). Fischer’s Least Significant Difference Test was performed for each
parameter to determine whether the means of the female types differed
significantly from one another.
SPERMATOPHORE DATA
Virgins that were mated were dissected within several hours of mating to
obtain spermatophores, or were kept in a refrigerator, until dissection a few
days later. Refrigeration slows down absorption of the spermatophore (C. Boggs,
personal communication), and a truer assessment of the original spermatophore
condition and weight is obtained. Forty-two field collected females were dis-
sected for comparison. Spermatophores were described as full, half full and
empty (when only a crust remained). Excised spermatophores and butterflies
were dried at ambient lab temperature and low humidity. Our data using dry
weight spermatophore to dry weight butterfly comparisons correlate extremely
well with recent data for E. editha and E. chalcedona (Jones et al. 1986), where
wet weight comparisons were used, presumably because both spermatophores
and butterflies are approximately 75-85% water. The use of wet weights was
suggested (Rutowski et al. 1983) to be of value since water has been considered a
“nutrient” for female eggs (Marshall 1982b). This is not likely for Euphydryas
species where water contributed from the spermatophore would represent an
4
J. Res. Lepid.
insignificant portion of that present in the egg and where water has been shown
to play no direct role in enhancing fitness in females (Murphy et al. 1983).
SIZE AND AGE OF INDIVIDUALS
The length of the forewing was used a size measurement. Wing wear, as an
estimate of age, was determined for all females using a five point rating system
(1 = extremely fresh, 2 = fresh, 3 = medium, 4 = old or worn, 5 = extremely
worn; see Iwasa et al. 1983). Wing length and wear estimates were also obtained
for males from field-captured mating pairs, and for 35 field-caught males.
WEATHER DATA
The condition of the weather was recorded about every fifteen minutes, or
whenever it changed, by using the simple denotations fair (sunny, with little or
no wind), marginal (largely cloudy with little sun or windy) and bad (heavy
clouds or very windy). At least some butterflies were active during all three
weather classes. When the weather became very bad, such as when wind was
extremely strong or it was raining, no butterflies were active, and few were ever
active if the sun was obscured for any length of time.
Results
MALE BEHAVIOR (Table 1)
Males tend to chase all flying objects vaguely the size of a Euphydryas
butterfly, including other insects such as grasshoppers (for male behavior
see also Odendaal et al. 1988, 1989). Males often chase other males,
engaging in frantic erratic flights during which the original object of the
chase is often replaced by another. Some of these flights included mating
attempts. One extreme male-male chase lasted 22 minutes and involved
numerous mating attempts, during which the pursuing role often
switched and brief copulations were achieved three times.
From a total of 1 19 male chases recorded (Table 1), virgins were found to
be chased longer than plugged females (p<.005, Mann-Whitney U-test),
and plugged females were chased longer than males (t = -2.4488,
p<0.02, Mann-Whitney U-test for large sample sizes see Siegel 1956);
males were not chased significantly longer than other species (t =
1.5088, 0.1<p<0.2, Mann-Whitney U-test). The extreme 22 minute
male-male chase was not included in these data as males often switched
pursuing roles, which made it difficult to time individual chases.
Males responded readily only to flying objects. They often flew within
centimeters of sitting virgins or plugged females or even walked around
on the same inflorescence with such females without reacting to them.
Only when the female took flight would the male chase her.
FEMALE BEHAVIOR (Table 2)
Virgins that mated - — Twelve of the 31 virgins mated in an average of
31 minutes after release. Of these, nine mated after an aerial pursuit by
a male. In a typical case of these nine, a virgin sat for a while on a plant
28(1-2):1-136, 1989(90)
5
Table 1. Summary of the duration of different types of male chases: males
chasing non-specifics, males chasing males, males chasing
females, and males chasing virgins.
Object of
male chase:
heterospecific male
mated female virgin female
n
60
51
25
2
ave(sec)
2.86
3.72
11.96
98
S.D.
2.41
3.63
12.18
26.87
Table 2.
Behavior data for females and ANOVA showing whether a
significant amount of the variance in each of the parameters can be explained
by female type. Asterisked entries have means that do not differ significantly
(at.05 level) according to the Fisher's Least Significant Difference Test. Male
density =
Number of males passing within 1 .5 m of a female per minute.
Behavior
FEMALES TYPE
Parameters
Plugged
Virgins
Virgins
Females
that Mated
not Mated
F
df P
(n = 12)
(n = 12)
(n = 19)
Flights
15.50*
14.17*
2.79
4.02
2,35 ns
per hour
± 9.99
±16.42
± 3.31
Flight
20.91*
14.91*
12.00*
4.83
2,25 ns
Distance(m)
±16.20
± 9.64
±13.59
Male
1.12
9.52*
10.23*
7.15
2,35 p<.01
density
± 1.98
± 9.86
± 7.73
Male
0.83*
0.83*
0.00
1,22 ns
chases/min
± 1.03
± 0.58
Chase
5.30*
20.20*
4.27
1,18 ns
Distance(m)
± 8.87
±21.01
or on the ground and then made a short flight. Almost immediately a
male gave chase. They landed, he crawled behind her, bent his abdomen,
and they mated, achieving the straight back-to-back position within
one minute of landing. Of the remaining three virgins, one was crawling
on the ground, a second was collecting nectar and the third was
hardening her wings after eclosion when mated. In these cases males
landed virtually on top of the females, seemingly by accident. All
virgins were mated by the first male that attempted mating and only
one seemed to resist briefly by walking away a few centimeters prior to
being mated.
During the 1986 season, two additional females were observed from
eclosion to mating. Both had wings that had not hardened yet and were
6
J. Res. Lepid.
sitting a few centimeters above empty pupal cases when encountered in
the field. One mated 98 minutes and the other 101 minutes after
observation began. In the first case, 68 males passed within 1.5 m and
several less than ten cm from the virgin without reacting to her. Finally
one male landed virtually on top of the female and they mated immedi-
ately. In the second case, 105s male passed within 1.5 m. Four short
flights of less than 20 cm each were taken when no males were present,
followed by a 10 m flight, during which one male and then a second
chased her. She alighted and both males attempted to mate her, with
one succeeding after about one minute.
As an additional measure of female choice we recorded wing size and
condition of males. These factors did not differ significantly between 12
males that mated and 210 males collected during the same period (size: t
= 1.9178, p<0.05; condition: t = 0.6723, p<0.05).
Virgins that did not mate — Nineteen out of 31 virgins did not mate
during the time of observation (approximate one hour cut-off, usually
due to unfavorable weather). Sixteen of these were never chased by
males. Those that flew did so significantly less often and for shorter
distances than virgins that mated (Table 2). Seven did not fly at all as
opposed to only one of those that mated. Three virgins were lost from
observation while being chased by males. One was pursued simul-
taneously by four males who also scrambled trying to mate with one
another during the chase. The number of males that passed within 1.5 m
of the virgins (relative male density) did not differ significantly between
virgins that mated and those that did not (Table 2). In the cases of five
non-mated virgins, males literally crawled over them or were side-by-
side while obtaining nectar.
Previously mated (plugged) females — Plugged females flew greater
distances and more frequently than virgins, but this was significant
only in comparison with virgins that did not mate (Table 2). When
pursued by males in flight, plugged females continue flying, trying to
evade the males. If unsuccessful, they land and energetically flutter
their wings during mating attempts. Six of the 12 plugged females were
chased by males, two of them twice and one three times. None mated.
(This nonreceptivity was observed many additional times during the
1986 season when we followed thirty-one plugged females in a detailed
study of their behaviour and movement; see Odendaal et al., 1989.
Plugged females fly differently from virgins. Once they have begun
laying eggs, their abdomens become noticeably slimmer and their flight
is smoother and stronger. Male density around plugged females was
significantly less than around virgins (p<.01), but males chased both
groups equally readily.
Spermatophore weights and condition (Table 3)— All twelve freshly
mated females were dissected and classified as having full sperma-
tophores. Dissection of 42 plugged females of various ages collected in
the field over the season yielded only three that had mated twice, each
28(1-2):1-136, 1989(90)
7
having two spermatophores. Of the 39 females that mated once, five had
very old, shriveled spermatophores and near-empty bursas, whereas
the remaining 34 had full or half full spermatophores. Of the three twice-
mated females, two had one old and one fresher spermatophore and the
third had two freshly deposited spermatophores. The ages of the three
twice-mated females (x = 2.33) did not differ significantly from that of
the 39 females (x = 2.76) that mated only once (t = 0.5617, p<0.5, Mann-
Whitney U-test for large sample sizes; see Siegel 1956).
Virgins had a condition of 1 which differed significantly from the
average condition of 2.69 of field caught females (t — 6.1984, p<0.001).
Female body weight also differed significantly between the two groups
(t = 4.0572, pCO.OOl), and body weight regressed significantly on
condition (age), F = 49.8804, p<0.001 . Spermatophore weight or % dry
weight did not differ between the groups, and neither variable regressed
significantly on condition (age).
A group of 43 field collected males were dried and weighed, with
average weight being 27.7±5.1 mg. The range was 16.2 to 38.0 mg. The
average spermatophore weight (Table 3) thus represents about 2% of
the male body weight.
Effect of weather — The weather had a profound effect on behavior.
When it was raining, or cold and cloudy, no butterflies were active.
Butterflies were clinging to plants when it was too cold or windy to fly
and could easily be picked up by observers. A few butterflies still showed
sporadic activity when we classified weather as bad, and more when we
considered the weather as marginal. We estimated about 56 percent of
all potentially available time (daylight hours) to be good, about 29
percent to be marginal, about 3 percent to be bad, and about 12 percent
of the time as too bad for any activity at all. To illustrate the effect of
Table 3. Summary of spermatophore data that were obtained from
dissections of field-caught plugged females and lab-mated virgins.
Age
(days)
Body Weight
(mg)
Spermatophore % Dry Weight
Weight (mg)
Field-caught
females
X
2.69
36.3
0.48
1.3
(n=42)
S.D.
0.84
9.5
0.28
0.83
Lab-mated
virgins
X
1
48.8
0.64
1.3
(n = 1 2)
S.D.
0
9.2
0.37
0.66
t-value
Significance
level
-4.0572
1.6700-1
-1.6501
0.01049
ns
0.010795
0.9144
ns
8
J. Res . Lepid.
slight weather changes on male butterfly activity, we plotted the
number of male butterflies chasing females, other males or non-specifics/
unit time for good and marginal weather (Figure 1).
Discussion
There remains a “specific lack of field data to understand the selective
advantages behind the evolution and maintenance of large nutritive
spermatophores” (Wickler 1986). This study shows that spermatophores
in Colorado Euphydryas anicia are very small compared to those species
investigated by Rutowski et al. (1983). What are the possible reasons for
the small spermatophore size in this species?
Svard (1985) showed that male Pararge aegeria invested only about
1.4% of their weight under laboratory conditions, and explained this in
terms of P. aegeria being a monandrous species with the male supplying
only enough sperm to fertilize all the eggs of the female. The problem
with this hypothesis is that the males in that study still produced 74% of
material other than sperm. However, our data on female mating
frequency do support the Svard explanation in that females in our field
% males
100
90
80
70
60
50
40
30
20
10
a b c
Fig. 1 Comparison of male behavior in good and marginal weather.
Thirty-three Euphydryas males were followed in good weather for a
total of 527 minutes and fifteen males in marginal weather for a total
of 520 minutes. It was recorded whether they chased or ignored other
males, females, and non-specifics that flew within a radius of 1.5 m
past them. Clear bars indicate percentage of (a) other males, (b)
females, and (c) non-specifics that were chased in good weather and
solid bars percentage of passerby butterflies chased in marginal
weather.
28(1-2):1-136, 1989(90)
9
study showed a very low degree of polyandry (cf. Ehrlich and Ehrlich
1978).
The length of the breeding season may be one link between the
environment and mating system characteristics (Odendaal et al. 1985a).
In species with very short breeding seasons, females may be limited by
the availability of time for repeated matings, host plant search and
opposition and should produce eggs mainly from resources acquired
during the larval stage. The general picture in Euphy dry as, which has a
short breeding season (Iwasa et al. 1983), appears to support this. Singer
and Ehrlich (1979) showed that only the offspring of the first egg batches
have time to reach diapause in California E. editha before host plants
senesce, and Murphy et al. (1983) showed that nutrient substances
ingested by adults only slightly benefit later egg batches. Further-
more, Jones et al (1986) showed that the size of the male investment
does not affect female reproductive output in E. editha. In contrast,
female Colias butterflies did show reduced reproductive output with
reduced male investment (Rutowski et al. 1987). On a high altitude
plain in Colorado, time for locating and choosing host plants may be
even more limited for E. anicia than for California Euphy dry as because
of lower temperatures and the frequently unfavorable mountain
weather. Monandry may be advantageous. Male mating success is
largely determined by access to fertilizable females (cf. Iwasa et al.
1983, Odendaal et al. 1985a) and if females tend to be monandrous,
rapid mate acquisition may be crucially important to males. Viewed in
this background, several points emerged from this study that may be
related to a small male ejaculate:
(1) Low level of mate discrimination for males and females— Very
limited time for locating and choosing host plants may have led to
monandry in Euphy dryas females. Females on this site use considerable
time locating hostplants, then sometimes as much as an hour inspecting
various plants and up to another hour laying eggs. This process is often
interrupted by unfavorable weather. Monandry and a low level of
female discrimination will place a high premium on rapid mate acquisi-
tion by males which may lead to a low level of male discrimination. Male
E. anicia pursue any flying object that might possible be a conspecific
female. Males also chase and try mating with one another. That these
are true mating attempts is corroborated by the observation of brief
amplexes between males. Males generally do not approach sitting
males, females or even virgin butterflies but chase almost any insect
that flies, presumably to maximize their chances of encountering
fertilizable females. The data indicate that males follow virgins more
persistently than plugged females, which in turn are followed more
persistently than males. Odendaal et al. (1985b) suggested that there
may be a close contact pheromone for identification of sexual partners,
but since males also try to mate with males and plugged females it is
10 J. Res. Lepid.
highly unlikely that males would discriminate among individual virgins
in the field.
Similarly, females of E. anicia can hardly be regarded as discri-
minatory. All twelve virgins in 1985 and the two observed in 1986
accepted the first male almost immediately. These males were not
larger than average. Rutowski (1984) builds a strong case for mate
choice in the lepidoptera on the basis of nutrients passed from males to
females in many species (eg., Boggs 1981; Rutowski 1982; Marshall
1982b). Females should select among males on the basis of traits (such
as size) which indicate that they can provide a large nutrient invest-
ment (Thornhill 1976). This apparently is not true of E. anicia , pre-
sumably because of the small spermatophore produced by males, or
because females acquired sufficient nutrients for egg production while
in the larval stage (cf. Murphy et al. 1983). Jones et al. 1986 showed
no relation between spermatophore size and reproductive output in
Euphydryas editha and E. chalcedona.
(2) Males exhibit scramble competition— Time-constrained males of
explosive breeders evaluate the quality of mates quickly if at all, and
sometimes males seem unable to discriminate visually between males
and females (Odendaal et al. 1985a). In four of the twelve virgins we
observed mating, more than one male followed the virgin, and in three
cases males scrambled intensely for her. One of the virgins we lost
escaped when four males who followed her scrambled and tried to mate
with one another. In 1986 we observed eight or nine males scrambling
for a single virgin. Similar frenzied mating attempts were also observed
with caged males (Odendall and Ehrlich, unpublished data).
(3) Spermatophore size and female mating frequency — As predicted by
Svard (1985) for monandrous species, E. anicia spermatophores are very
small relative to their body size. They are smaller than any recorded in
the available literature (Rutowski et al., 1983). Females of the study
population also remated very infrequently, with only three (7%) of field-
caught females having more than one spermatophore. Furthermore,
one of these females contained two fresh spermatophores and this could
have been the result of scramble competition rather than a female
tendency to remate. In a laboratory experiment Odendaal (unpublished
data) once observed a virgin copulating with two males. Males often try
to displace a mating male, and this may result in two spermatophores as
the plug of the first one may not have hardened yet (Labine 1964).
Percentage of the body weight made up by spermatophores did not differ
between randomly collected mated females and freshly mated females,
suggesting that females did not gradually use substantial amounts of
the spermatophore for nutrition.
(4) Length and complexity of courtship— Rutowski (1984) states that
nutrient investment made by male butterflies appears to have given
rise to selection pressures that have shaped courtship behavior of males
and perhaps females. Supportive data comes primarily from pierids (see
28(1-2):1-136, 1989(90)
11
Rutowski 1984). Because of the small male investment in E. anicia, it
may not be surprising that courtship is essentially non-existent in this
population. All copulations were closely observed, sometimes from less
than a meter away, and a successful mating merely involved the curling
of the male’s abdomen, sometimes a second or two searching for the
female genital aperture with his genitalia, and copulation, which we
regarded as complete when the couple achieved the straight back-to-
back position. A further twenty copulations of caged butterflies obtained
from this site and filmed on video tape yielded essentially the same
results (Odendaal and Ehrlich, unpublished data).
Our work deviates somewhat from data on the Euphydryas species.
The female remating frequency at Red Hill was considerably less than
that for E. editha in California (Labine 1964) and there was a striking
difference between relative spermatophore weights in the present work
(<2%) as compared to that reported (10.8%) for three E. chalcedona
individuals in Arizone (Rutowski et al., 1983). A detailed comparison of
mating behavior of species within the genus Euphydryas would provide
an interesting test of predictions on how spermatophore size may affect
mating systems.
Acknowledgements. We thank Dale R. Gardner for data on butterfly and
spermatophore weights and for assistance in field observations. Peter Turchin
and Guy H. Harris also provided field assistance. We thank Mark Rausher for
use of the field computers. The research was funded by National Science
Foundation grant BSR-8506064.
Literature Cited
BOGGS, C.L. 1981. Selection pressure affecting male nutrient investment at
mating in heliconiine butterflies. Evolution 35: 931-940.
BOGGS, C.L. and L.E. GILBERT. 1979. Male contribution to egg production in
butterflies: evidence for transfer of nutrients at mating. Science 206: 83-84.
BOGGS, C.L. and W.B. watt. 1981. Population structure of pierid butterflies. IV.
Genetic and physiological investment of offspring by male Colias. Oecologia
50: 320-324.
CULLENWARD , M.J., P.R. EHRLICH, R.R., WHITE, and C.E. HOLDREN. 1979. The ecology
and population genetics of an alpine checkerspot butterly, Euphydryas
anicia. Oecologia 38: 1-12.
DEWSBURY, D.A. 1982. Ejaculate cost and male choice. Amer. Nat 119: 601-610.
EHRLICH, A.H. and P.R. EHRLICH. 1978. Reproductive strategies in the butterflies: I.
Mating frequency, plugging, and egg number. J. Kansas Ent. Soc. 51:
666-697.
ENGEBRETSON, J.A. and W.H. MASON. 1980. Transfer of Zn at mating in Heliothis
virescens. Environ. Entomol. 9: 119-121.
FERRIS, C D. and F.M. BROWN 1981. Butterflies of the Rocky Mountain States.
University of Oklahoma Press, Norman, Oklahoma, p. 331-332.
GARDNER, D.R. and F.R. STERMITZ 1988. Hostplant utilization and iridoid glycoside
12
J. Res. Lepid.
sequestration by Euphydryas anicia (Lepidoptera: Nymphalidae). J. Chem.
Ecol. 14: 2145-2166,
GREENFIELD, M.D. 1982. The question of parental investment in Lepidoptera:
male-contributed proteins in Plodia inter punctella. Int. J. Invertebrate
Reprod. 5: 323-330
IWASA, Y., F.J. ODENDAAL. D.D. MURPHY, P.R. EHRLICH and A.L. LAUNER. 1983.
Emergence patterns in male butterflies: a hypothesis and a test. Theor. Pop.
Biol. 23: 363-379.
JONES, K.N., F.J. ODENDAAL and P.R. EHRLICH. 1986. Evidence against the sperma-
tophore as paternal investment in checkerspot butterflies ( Euphydryas :
Nymphalidae). Amer. Midi. Nat. 116(1): 1-6.
LABINE, P.A. 1964. Population biology of the butterfly, Euphydryas editha. I
Barriers to multiple inseminations. Evolution 18: 335-336.
MARSHALL, L. 1982a. Male courtship persistence in Colias philodlice and C.
eurytheme (Lepidoptera: Pieridae). J. Kansas Ent. Soc. 55: 729-736.
MARSHALL, L. 1982b. Male nutrient investment in the Lepidoptera: What
nutrients should males invest? Amer. Nat. 120: 273.
MURPHY, D.D., A.L. LAUNER and P.R. EHRLICH. 1983. The role of adult-feeding in
egg production and population dynamics of the checkerspot butterfly,
Euphydryas editha. Oecologia (Berl.) 56: 257-263.
ODENDAAL. F.J., Y. IWASA and P.R. EHRLICH. 1985a. Duration of female availability
and its effect on butterfly mating systems. Amer. Natur. 125(5): 673-678.
ODENDAAL. F.J., P.R. EHRLICH and F.C. THOMAS 1985b. Structure and function of
the antennae of Euphydryas editha (Lepidoptera: Nymphalidae). J. Morph.
184: 3-22.
ODENDAAL. F.J., P. TURCHIN, and F.R. STERMITZ. 1988. An incidental-effect
hypothesis explaining aggregation of males in a population of Euphydryas
anicia (Nymphalidae), Amer. Natur. 132: 735-749.
ODENDAAL, F.J., P. TURCHIN, and F.R. STERMITZ. 1989. Male harassment, host plant
spatial availability, and the distribution of female Euphydryas anicia ,
Oecologia. 78: 283-288.
RUTOWSKI, R.L. 1982. Mate choice and lepidopteran mating behavior. Florida
Ent. 65: 72-82.
RUTOWSKI, R.L., M. NEWTON and J. SCHAEFER. 1983. Interspecific variation in the
size of the nutrient investment made by male butterflies during copulation.
Evolution 37(4): 708-713.
RUTOWSKI, R.L. 1984. Sexual selection and the evolution of butterfly mating
behavior. J. Res. Lep. 23: 125-142.
RUTOWSKI, R.L. 1985. Evidence for mate choice in a sulphur butterfly ( Colias
eurytheme). Z. Tierpsychol. 70: 103-114.
RUTOWSKI, R.L., G.w. GILCHRIST, and B. terkanian 1987. Female butterflies mated
with recently mated males show reduced reproductive output. Behav. Ecol.
Sociobiol. 20: 319-322.
SIEGEL. S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-
Hill, New York, 312 pp.
SINGER, M. and P.R. EHRLICH 1979. Population dynamics of the checkerspot
butterfly Euphydryas editha. Fortschr. Zool. 25: 53-60.
STERMITZ, F.R., D.R. GARDNER, F.J. ODENDAAL, and P.R. EHRLICH 1986. Euphydryas
anicia (Lepidoptera: Nymphalidae) utilization of iridoid glycosides from
28(1-2):1-136, 1989(90)
13
Castilleja and Besseya (Scrophulariaceae) host plants. J. Chem. Ecol. 12:
1459-1468.
SVARD, L. 1985. Paternal investment in a monandrous butterfly, Pararae
aegeria. Oikos 45: 66-70.
SVARD, L. and C. WIKLUND. 1986. Different ejaculate delivery strategies in first
versus subsequent matings in the swallowtail butterfly Papilio machaon. L.
Behav. Ecol. Sociobiol. 18: 325-330.
THORNHILL, R. 1976. Sexual selection and paternal investment in insects. Amer.
Natur. 110: 152-163.
THORNHILL, R. and J. ALCOCK. 1983. The evolution of insect mating systems.
Harvard University Press, Cambridge.
WALKER, W.F. 1980. Sperm utilization in nonsocial insects. Amer. Natur. 115:
780-799.
WHITE, R.R. 1979. Foodplant of alpine Euphydryas anicia (Nymphalidae). J. Lep.
Soc. 33: 170-173.
WICKLER, w. 1986. Mating costs versus parental investment: a reply to Gwynne.
Ethology 71: 78-79.
Journal of Research on the Lepidoptera
28(l-2):14-25, 1989(90)
The Biology of Colias blameyi^ Pieridae), the “Green
Sulphur” of the Argentine Puna
Arthur M. Shapiro
Department of Zoology, University of California, Davis, California 95616
Abstract. Colias blarney i Joergensen is a high-altitude endemic found
in the puna of the Provinces of Catamarca, Tucuman, Salta and Jujuy
in northwestern Argentina. It is probably double-brooded, with a
seasonal up-and-downslope migration tracking the availability of its
host plants, Astragalus spp. (Leguminosae). The early stages, reared
on alfalfa ( Medicago satiua L.) are described. The relationship of C.
blameyi to other taxa of high-altitude Sulphurs in the central Andes
remains problematical.
Introduction
The biogeography of the high- Andean butterfly fauna is receiving
renewed attention in the context of attempts to reconstruct Quaternary
climate dynamics and their impact on biotic diversity in tropical
America (Brown 1987, Descimon 1986, Shapiro 1989). For most groups,
both distributional and biological data are still inadequate for a proper
analysis to be done. In many cases the sister-groups of endemic Andean
taxa are unknown. The genus Colias has its greatest diversity in the
Holarctic, but has undergone considerable adaptive radiation in the
cold and temperate parts of South America. The systematic position of
the Andean Colias is far from resolved. Descimon (1986) considers them
a monophyletic group, while Berger (1986), in a global revision at the
subgeneric level, does not. The characters used by Berger to delimit
subgenera are superficial and poorly if at all rationalized, while
Descimon’s discussion is informal and his bases for judgment are
inexplicit. Speculation must be replaced by data if progress is to occur.
Descimon has reared several Andean Colias and indicates (loc. cit.) that
their life-histories will be published. One which he has not reared is
Colias blameyi Joergensen, whose life-history is reported here.
In 1916 Pedro Joergensen, one of the three founders of Argentine
Lepidopterology (with Eugenio Giacomelli and Carlos Berg), published
a landmark monograph on the Argentine Pieridae which included the
description of a new Colias from the Sierra de Aconquija, Provinces of
Catamarca and Tucuman. The Sierra de Aconquija and Cumbres
Calchaqules form a major eastern outlier of the Andes proper, reaching
altitudes over 5000 m (Nevado del Candado, near the southern end of
the Aconquija range, reaches 5450 m), and separated from the Andes by
a deep trough, the Valles Calchaqules. Moisture-bearing winds strike
28(1-2):1-136, 1989(90)
15
the range from the east, so that there is a very pronounced rain shadow
west of the crest. The seasonally wet climates east of the crest provide
the last refuge for many b timid- N eo tropical biotic elements of the mid-
elevation forest zone, while the subalpine and alpine zones shelter many
elements of the central Andes (Peru, Bolivia) which drop out in the
Andes themselves south of the Province of Salta. Joergensen was able to
mount several expeditions into the Aconquija range with the help of his
friend Joel Blarney of Huazan, after whom he named his new Sulphur.
There was a burst of description of new Andean Colias around this time,
but as usual they were published in European journals and Joergensen
was unaware of them; for him the dusky green phenotype of Coiias
blameyi was something entirely new for the continent, and it immedi-
ately reminded him of various boreal Holarctic species: “The male of
this new little species . . . cannot be confused with any of its South
American congeners, but it has the size and the dark glaucous green
ground color of the species nastes Boisduval and behri Edwards, the
former from the North American Arctic (Labrador, Greenland, Alaska
and British Columbia), the latter from the mountains of California; but
the patterns are different.” (Translation by A.M.S.) He provided an
excellent, detailed description of both sexes and a brief summary of
what he knew of its biology. Since then this striking insect has been
collected occasionally by travelers, but nothing further on its biology
has appeared. If Kenneth Hayward knew any more about it, his
information apparently died with him, as the projected Fiend volume in
Hayward’s monograph of the Argentine butterflies never appeared and
no manuscript has been found. Although several South American
Colias have been reared, the only species for which published informa-
tion on the life-history and early stages is available is C. lesbia
Fabricius, which is a serious alfalfa pest in several countries. It has been
monographed by Biezanko (1954), Freiberg (1947), and Reed (1922).
Although C. blameyi is common in the proper habitat in season,
specimens are rare in collections — even in Argentina itself.
Biology of Adults
Joergensen (1916, p.510) states that C. blameyi “is common on the
summits of the grassy mountains: Cerro La Tambilla, 3700 m; Cerro
Medio, 3750 m; Cerro Yutoyaco, 3500; Cerro Negro, 3500, and Cerro La
Ensenada, 3200. All these localities are east and southeast of the snowy
summits of the Aconquija range. There it flies from the end of January
until the end of April. Its flight is not very high, but in good weather it
seems constantly in motion, (though) often settling on the ground or on
flowers such as Gutierrezia repens Gr., Hypochaeris meyeniana Wolp.
(Compositae), Verbena microphylla H.B.K. (Verbenaceae), Malvastrum
capitatum Gr, and M. parnassifolium Hook. (Malvaceae). When the
hard freezes begin on the heights in late March, it descends ... to more
protected valleys, for example at La Olla da (3100 m), where it is never
16 J. Res. Lepid.
found in summer.” Joergensen was an excellent observer and is correct
on all points.
There is no amplification of the range, as stated by Joergensen, in any
subsequent literature. Because the closely related entity C. weberbaueri
Strand, which differs from c. blameyi primarily in its lack of an
androconial patch in the male, is the only taxon of the group recorded in
Bolivia, it is important to note that C. blameyi is not confined to the
Sierra de Aconquija. Within the Province of Tucuman it extends north
an indeterminate distance well into the Cumbres Calchaqules, which
extend to the NE of Abra Infiernillo (3800 m), where Highway 307
(Monteros-Amaicha del Valle) crosses the range. In the true Andean
puna it, or an entity transitional from it to C. weberbaueri , occurs
abundantly in the Provinces of Jujuy (Abra Pampa, Tres Cruces,
Esquinas Blancas, 3893-3875 m, all along Highway 9 above Humahuaca)
and Salta (Cuesta del Obispo, Abra Molina, Cerro Zapaiiar, Valle
Encantado, all along or near Highway 33). There is great individual and
interpopulational variability in both sexes (figs. 1,2). Puna animals
average lighter than Tucuman and Catamarca ones, but nearly all have
well-developed androconial patches. There is no obvious tendency for
them to be smaller, or more frequently reduced, in the puna than in the
Aconquija-Cumbres Calchaqules populations.
Both sexes fly from roughly 1000 to 1500 daily in good weather. Flight
initiation in the morning occurs with air temperatures of roughly LO O
with light wind and strong sunshine. Flight may be terminated early by
cloudiness or, even under clear sky, by strong and turbulent upslope
afternoon winds which often develop on the eastern slopes of the
Tucuman-Catamarca ranges and at the head of the Quebrada de
Humahuaca. A few animals continue to fly in the lee of ridges or hills
until the sun fails to reach them.
Males patrol linear habitats such as roadsides, streamsides and
gullies, and below the crests of ridges, but do not hilltop. They can often
be seen coursing back and forth over alpine grassland and rock gardens
(as in fig. 3) about 1 m above the ground. All-male aggregations occur on
moist earth, at puddles and along streambanks. Up to 30 animals have
been seen puddling together, mainly after 1330. Females occur singly
and are seldom seen where males are patrolling. Most of my observa-
tions of females have been at or near summits, where host plants grow
among rocks, or around shrubs, where they often grow within the drip
line. Oviposition occurs singly, usually on the underside of a leaf, and
females will frequently proceed in more or less of a straight line, laying
one egg on each plant they encounter.
Host Plants
Three definite hosts have been identified (by E. Barneby, New York
Botanic Garden). All are based on many (> 10) oviposition records/each.
28(1 2) 1-136, 1989(90)
17
They are Astragalus garbancillo Cav. and A. micranthellus Wedd., both
at Tres Cruces, Jujuy, 3800 m±, and A. hypsogenus LM. Johnston, at
both the summit of Cerro Zapallar, ca. 4200 m, Salta, and on several
summits near Abra Infiernillo, Tucuman, ca. 3500 m (all Leguminosae).
These three species are disparate in both facies and phylogenetic
affinities. Their differences imply that C. blameyi is a generalist at least
within the genus Astragalus. Astragalus is very well-developed and
diverse in the Andes (Johnston 19 47). A. garbancillo is “the most widely
distributed and most commonly collected South American Astragalus’
(Johnston 1947, p. 384). It is erect and ascending in habit. In Argentina
it is largely confined to moist and dissected areas at the periphery of the
puna. It is a common species in much of the range of C. weberbaueri in
Bolivia and Peru and should be considered a probable host. It is
taxonomicaily isolated within the genus Astragalus.
Astragalus micranthellus is depressed, prostrate to tufted and much
less leafy and conspicuous than A. garbancillo. Its range includes
altiplano and puna in Peru and Bolivia, extending in Argentina only as
far south as the Sierra de Aconquija (Johnston, p. 391).
Astragalus hypsogenus is a small, tufted plant with tiny leaves but
very showy purple flowers, reminiscent in habit of some of the alpine
Lupinus. It is one of the aspect dominants of alpine rock garden habitats
on the wetter summits in both Salta and the Aconquija-Cumbres
Calchaqules and grows in the sites shown in both figs. 3 and 4. Its range
includes Bolivia and northern Argentina, again not south of the Sierra
de Aconquija. It forms a compact, isolated species-group along with A.
confinis LM. Johnston and A. crymophilus I.M. Johnston.
Both A. garbancillo and A. micranthellus are confirmed wild hosts of
the pierine Tatochila distincta distincta Joergensen, which can however
be reared on Crucifers in the laboratory (Shapiro 1986). This butterfly
co-occurs with A. hypsogenus in both Salta and Tucuman, as does the
very rare and as yet unreared T. inversa Hayward.
Early Stages
No significant differences have been observed in material from Salta,
Jujuy and Tucuman. Rearing was done at Davis on alfalfa ( Medicago
Fig. 1. Co/ias blameyi from these disjunct populations in northwestern
Argentina, males at left. A: Quebrada Carapunco, Province of
Tucuman, 20. i. 1986. B: Esquinas Blancas, Province of Jujuy,
7.ii.1984. C: Summit of Cerro Zapallar, Province of Salta, 22. i. 1986.
The Tucuman populations are essentially topotypical. The Salta and
Jujuy populations resemble C. weberbaueri from Bolivia but have
well-developed androconial patches in the males.
18
J. Res. Lepid.
28(1-2):1-136, 1989(90)
19
20
J. Res. Lepid.
sativa L.) cuttings; Vicia benghalensis L. was eaten but no larvae
survived beyond the third instar on it. Larvae were kept in plastic Petri
dishes under 14L:10D, 23.9°/12.8°C. Preserved early stages have been
retained at Davis. All color descriptions are from life. Those in paren-
theses refer to the color-standards system of Kornerup and Wanscher
(1978).
Egg (fig. 5). — Erect, fusiform, strongly tapered at both ends, 1.1 x 0.3
mm, the chorion sculptured as figured with about 16-17 and 42-52
vertical and horizontal ribs. Madder red (9A7) when laid, becoming
translucent about 12 hr before hatching. Laid singly, usually on lower
leaf surfaces. Newly-hatched larvae do not eat their eggshells. Time to
hatch, 7-8 days.
Larva : First Instar (fig. 6). — At hatch 1.15 mm, grayish to brownish
orange, the head much darker; head and body with pale, mostly
glandular hairs disposed as below. After feeding grayish green (1D7)
dorsally, grayish yellow (1B3) vent rally, a darker shade immediately
below the spiracles and above the prolegs, gradually lightening toward
the venter. First thoracic segment with a transverse fold and 9-10
glandular hairs in a single row. Second and third segments with folds
dividing them into five annulae, the fourth (counting caudad) bearing 8
glandular hairs in a single transverse row. First two abdominal seg-
ments each with four annulae, of which the first and fourth each bear
two glandular hairs. Other abdominal segments with five annulae, the
first and fifth of which each bear two glandular hairs, except the eighth
and ninth with three annulae, two hairs each on first and third; and the
last with a darkened sclerotized shield bearing several dark, non-
glandular setae, plus six glandular hairs anterior to the shield. Exca-
vates strips of parenchyma; feeds by day and night and rests along the
midrib. Lenght of instar, 3-4 days.
Second Instar. — After molt 3.8 mm. Similar, with annulae disposed as
follows: five annulae per segment except the following abdominal
segments with six: second, fourth, fifth, seventh; third with seven; ninth
and tenth apparently unitary. This arrangement is continued in later
instars, with intercalation of annulae on some segments especially near
the front of the abdomen. Dorsal and lateral surfaces densely covered
with small dark tubercles in two sizes, each surmounted by either a
Fig. 2. Habitats of C. b/ameyi in wet season, during the flight period. A:
Summit of Cerro Zapallar, Salta, looking toward Valle Encantado
below. Both sexes are common here, flying over alpine rock gardens.
B: Rocky summit at about 3900 m in the Cumbres Calchaquies near
Abra Infiernillo, Tucuman. Females occur here and oviposit on
Astragalus hypsogenus among the rocks. The yellowish cushion
plant is Azore/la (Umbelliferae, “yareta"), a characteristic alpine plant
in the region.
28(1-2):1-136, 1989(90)
21
Figs. 5-12. Life history of C. blarney/. 5, egg; 6, first-instar larva, lateral view;
7, fifth-instar larva, dorsal view; 8, same, lateral view; 9, same,
head capsule; 10, pupa, lateral view; 11, same, dorsal; 12, same,
ventral.
22 J. Res. Lepid.
glandular or a simple hair. Head darker than body, densely tuberculate.
No change in habits. Duration, 3-4 days.
Third Instar. — After molt 5.25 mm. Similar, tubercles densely and
rather evenly distributed over dorsal and lateral surfaces of body and
even more densely on head, the larger ones darker and bearing mostly
dark hairs, the smaller either darker or concolorous and bearing either
dark or pale hairs. Head capsule scarcely darker than body: ocelli black.
A vague pale line on each side incorporating the spiracles; directly
below it a very dark shade of the ground color, grading insensibly into
the paler venter. Third instars consume epidermis as well as paren-
chyma, Length of instar, 4-5 days.
Fourth Instar. — After molt 8.5 mm. Similar, with a decidedly granular
appearance due to the very numerous tubercles and hairs. Rests
lengthwise on the petiole when not feeding. Length of instar, 5 days.
Fifth Instar (figs. 7, 8, 9). - After molt 14 mm, reaching 23 mm at
maturity. Head and body above olive (2E6) with numerous tubercles,
both dark and concolorous, over the dorsal and lateral surfaces; hairs
both light and dark, between 50-100 per annulus. Spiracles not con-
trasting, but incorporated in an ill-defined pale line (grayish yellow,
1B3) not enclosing any red or pink color; below this a darker shade of the
ground color, passing into dull olive (2D4) just above the bases of the
legs, which are concolorous with the venter. Crochets black. Dorsal
midline slightly darker than ground; the entire dorsum slightly paler
than the sides, the pale hue ending abruptly where the subdorsal pale
stripes would be if present. Head dark olive, densely tuberculate, the
tubercles bearing dark hairs; ocelli brownish-black.
The mature larva feeds by day and night in the lab, resting on stems.
If disturbed, it drops to the ground in a coil, reascending the plant 10-15
min later. This is a stereotyped defensive reaction in all Colias I have
reared. The day before pupation the larva turns grayish with a slightly
purple tinge, leaves the plant and wanders for several hours before
spinning a mat of silk in preparation for the molt. Duration of instar,
7-10 days.
Prepupa. — Formed vertically, head up, pendant by the silken girdle
and attached at the cremaster, appearing greasy and grayish-yellow-
green. Length of prepupal period 20-36 hr.
Pupa (figs. 10, 11, 12). — Typical Colias form, chunky, the wing-cases
not particularly inflated and the frontal prominence short and broad;
length 14-15 mm, width at base of abdomen 3.3-4 mm. Dorsal surface
olive yellow (2C7); ventral, including wing cases, canary yellow (2B7).
Proboscis not reaching tips of wing cases. Wings with a black dot
corresponding to the discocellular spot of the adult and black dots at the
vein-tips. Brownish-red (10C7) shading as follows: on dorsal surface of
the frontal prominence; along hind margin of wing cases; on the dorsal
thoracic keel; above the spiracles; and two parallel rows of blotches on
the ventral abdomen, one on either side of the midline. Spiracles
28(1-2):1-136, 1989(90)
23
enclosed in a yellowish-white, moderately contrasting line. Eyes, wings
and body becoming pigmented in that order the day before eclosion, the
wings of both sexes initially yellowish-white, those of the males sub-
sequently turning dark (black pigment laid down several hr after
white). First meconium dull rose pink, second colorless. Time to hatch,
12-17 days.
Diapause. — Colias usually diapause as third-instar larvae. Several
larvae indeed stopped feeding in the third instar and survived 2-4 wk
thereafter, but there was so much disease mortality that I cannot say
with confidence that they were attempting to diapause. Altitudinal
migration, such as between La Ollada and the Sierra de Aconquija or
between Valle Encantado and the summit of Cerro Zapallar, may be a
seasonal strategy to avoid severe cold and to track host plant availability.
In many insects such migration substitutes for diapause as a mechan-
ism to avoid seasonal stress. Colias blameyi , however, disappears
altogether for more than half the year and diapause is thus very likely.
Descimon (1986) echoes other authorities in recording the entire
assemblage of pale and green puna species as flying only in rainy season
(“March- April, or December in the Arequipa region”).
Altitudinal migration appears to be very common in the butterfly
fauna of the northwestern Argentine highlands. Several species of
Tatochila, including T. sterodice macrodice Stgr., T. stigmadice Stgr.
and T. orthodice Weymer, which fly with C. blameyi in January and
February, can be found at much lower elevations in the Provinces of
Salta and Tucuman in November. The member of the Phulia nymphula
Blanchard complex ( aconquijae Joerg.) found in the Aconquija and
Cumbres Calchaquies parallels C. blameyi in its winter retreat to the
level of La Ollada.
Comparisons to C. lesbia. — Colias lesbia is larger (except in cold-
weather broods) throughout its development, and both the larva and
pupa are more slender. The disposition of annulae and glandular hairs
on the first-instar larva is very similar, but the number of annulae
diverges in later instars. The larva of C. lesbia is bright “alfalfa green”
rather than dull or olivaceous green as in C. blameyi , and has a pink
spiracular line. The pupa is brighter green and has a bolder spiracular
line with silvery reflections. Both morphology and pattern are very
conservative in Colias immatures, as noted by Descimon (1986). Until a
detailed morphological study is done of representative members of
various species-groups, isolated rearings will cast little light on the
Holarctic sister-group of the Andean group to which C. blameyi belongs;
the relevant information is still largely lacking for the Holarctic taxa as
well. There is nothing in these descriptions which would lead one to
question the joint membership of C. lesbia and C. blameyi in a mono-
phyletic Andean group, but the point is moot until more descriptions are
available. All of the Andean species reared so far are Legume feeders,
though some are presently known only from naturalized European
clovers.
24
J. Res. Lepid.
Discussion
The group of taxa embracing C. blarney i\ C. weberbaueri , C. erika
Lamas, and C. mossi Rothschild (including “form” nigerrima Fassl), all
from Peru; and C. flaveola from Chile (and northwestern Argentina,
Shapiro, unpublished), is badly in need of revision. Of the heavily
melanized taxa only weberbaueri is recorded in Bolivia, but blameyi is
now recorded within about 25 km of the Argentine-Bolivian border in
the puna of Jujuy, and flaveola is now known to cross the Andean crest
and penetrate the eastern slope. Berger (1986) treats blameyi as a
subspecies of mossi but weberbaueri becomes a separate species by
virtue of its lack of an androconial patch. Descimon (1986, p. 506) states
that this is a fluctuating character in some populations, even of flaveola.
Shapiro (1985) has published a figure of a melanic aberration of C.
euxanthe stuebeli Reiss, from the Department of Cusco, Peru. The type of
melanization displayed is quite different from that of melanic aberra-
tion of the Nearctic C. philodice Godt. and C. eury theme Bdv. but agrees
perfectly with that seen in the puna complex, underscoring the point
that phenotypic similarity within this group could easily have arisen by
parallelism. It cannot be assumed automatically that all the “green”
taxa are more closely related among themselves than they are to the
non-“green” ones.
Acknowledgements. This research was supported by National Science Foun-
dation grant BSR-8306922 (Systematic Biology Program). It would not have
been possible without the help of Sr. Robert Eisele of Yerba Buena (Tucuman)
and Lie. Estela Neder de Roman and Lie. Martha Arce de Hamity of the Institute
de Biologia de la Altura, S.S. de Jujuy. The specimen photographs are by Samuel
W. Woo and the drawings by Karen English-Loeb and Adam H. Porter. I also
thank Dr. Henri Descimon for his encouragement and much stimulation in
discussions of the evolution of the high- Andean fauna, and R. Barneby and J.
McCaskill for help with the plants.
Literature Cited
BERGER, L A. 1986. Systematique du genre Colias F. (Lepidoptera-Pieridae),
Lambillionea 86: Supplement, 68 pp.
BIEZANKO, C.M. 1954. Colias lesbia pyrrhoihea Huebn, 1823 (Lepidoptera,
Rhopalocera, Pieridae) inimigo de alfalfa . . . Bol. Esc. agron. Eliseu Maciel,
Pelotas, Brasil, #29: 7-23.
BROWN, R.S. JR. 1987. Biogeography and evolution of neotropical butterflies,
pp. 66-104. In T.C. Whitmore and G.T. Prance, eds., Biogeography and
Quaternary History in Tropical America. Clarendon Press. Oxford.
DESCIMON, H. 1986. Origins of Lepidopteran faunas in the high tropical Andes,
pp. 500-532. In F. Vuilleumier and M. Monasterio, eds., High Altitude
Tropical Biogeography. Oxford University Press, New York and Oxford.
FREIBERG, M.A. 1947. La oruga de la alfalfa en la Argentina ( Colias lesbia
Fabricius, Lep. Pier.). Bol. Inst. Sanidad Veg. Ser. A, #36:1-32.
JOERGENSEN, P. 1916. Las mariposas argentinas, familia Pieridae, Anales Mus.
Nac. Hist. Nat. Bs. As. 28:427-520
28(1-2):1"136, 1989(90)
25
JOHNSTON, I.M. 1947. Astragalus in Argentina, Bolivia and Chile. J. Arnold
Arboretum 28:336-408.
KORNERUP, A. & J.H. WANSCHER. 1978. Methuen Handbook of Colour. 3rd Edition.
Methuen, London. 252 pp.
REED, C.S. 1922, La cuncuna o isoca de los alfalfares de Mendoza ( Colias lesbia
Fabr.). Min. Ind. Obr. Publ. Prov. Mendoza, 20 pp.
SHAPIRO, A.M. 1985. A melanic Colias euxanthe stuebeli from Peru (Pieridae). J.
Res Lepid. 24:87.
- — - — —1986. The life history of Tatochila distincta distincta , a rare butterfly
from the puna of northern Argentina (Lepidoptera: Pieridae). J N.Y.
Entomol, Soc. 94; 526—530
— 1989. Ignorance in high places. Paleobiology 15: 61-67.
Journal of Research on the Lepidoptera
28(l-2):26-36, 1989(90)
The Early Stages of Doa dora Neumoegen and Dyar
(Lepidoptera: Noctiioidea: Doidae) in Baja Califor-
nia, Mexico
John W. Brown1
Department of Entomology, National Museum of Natural History, Washington, D. C.
20560
Abstract. The early stages of Doa dora Neumoegen and Dyar from
Baja California, Mexico, are described and illustrated. Adults were
reared on Euphorbia misera Bentham (Euphorbiaceae) from eggs
deposited by females collected on Isla de Cedros. The unique combina-
tion of larval characters possessed by the doids, i.e., small head,
hump-backed thorax, biordinal crochets in a homoideous mesoseries,
and integumental spicules, contradict traditional assignments to
families of similar adult morphology (i.e., Lymantriidae, Hypsidae,
Pericopidae, Artiidae). It is likely that specimens from the northern
part of the range of D. dora (i.e., Baja California and Sonora, Mexico)
represent an undescribed species. Although adults have not been
collected in the United States, larvae have been taken in San Diego,
California.
Introduction
The genus Doa Neumoegen and Dyar (1894) has traditionally defied
attempts at familial assignment. Its long history of taxonomic uncer-
tainty includes placement in the Lymantriidae (Dyar 1903; Barnes
and McDunnough 1917; Holland 1903; Bryk 1934), Hypsidae (Walton
1912), Pericopidae (Schaus 1927; McDunnough 1938; Peterson 1948),
and Arctiidae (Franclemont 1983). Most recently, Doa and its sister
genus, Leuculodes Dyar, have been treated as a distinct family — the
Doidae (Donahue and Brown 1987). However, the phylogenetic rela-
tionship of Doa to other noctuoid families is uncertain, and elevation to
family level probably represents only an interim solution. It is likely
that the early stages will provide characters useful in illuminating
relationships among the doids and other noctuoid clades.
Dyar (1911, 1912) provided superficial descriptions of the early stages
of Doa ampla (Grote) and Doa raspa (Dyar). However, features of the
chaetotaxy and crochet arrangement have been presented only recent-
ly (Donahue and Brown 1987). The purpose of this paper is to provide
1 Research Associate, Entomology Department, San Diego Natural History Museum,
P. O. Box 1390, San Diego, CA, 92112.
28(1-2):1-136, 1989(90)
27
descriptions and illustrations, of the larva, pupa, and adult, and notes
on the biology of Doa dora Neumoegen and Dyar in Baja California,
Mexico. It is not my intention to draw conclusions regarding the
phylogenetic position of the Doidae, but to make available specific life
history information that has accumulated.
Materials and Methods
A single female Doa dora was collected at blaeklight (LTV) on the
north end of Is I a de Cedros, Baja California, Mexico, 31 March 1983. A
second female was collected the following morning, while it perched on
a large bush of Euphorbia misera Bentham (Euphorbiaceae). On 2
April 1983, a fourth and a fifth instar larva were collected on E. misera
by D. K. Faulkner, near El Pueblo, in the southeastern portion of the
island.
The adult females were confined together in a plastic bag with a fresh
cutting of E. misera. They readily oviposited on the leaves and stems of
the plant material. I estimated that between 50 and 75 eggs had been
deposited by the evening of 5 March. The eggs were taken to San
Diego, California, where the larvae were reared to maturity on local E.
misera. As the eggs hatched, larvae were transferred in small groups
to 4.5 ounce 'glass jars with small pieces of netting for lids. When the
larvae reached the third instar, they were transferred to a cylindrical,
ha if gal Ion, cardboard container, where they continued to feed and
eventually pupated. Rearing was done indoors at ambient temperature
(65-77°F).
Upon emergence, most adults were removed. However, the last 5 or 6
were left in the container. Mating took place within 1 to 3 days of
eclosion; females readily oviposited on the dry plant material remain-
ing in the container. A second generation was reared from these eggs.
Insufficient host material resulted in a brood of dwarfed adults. All
specimens are deposited in the collection of the San Diego Natural
History Museum (SDNHM).
Description of Early Stages
Morphological terminology and homology of setae follow Stehr (1987);
terminology and homology of pupal characters follow Mosher (1918).
Egg. Flattened, oblong, oval; width ea. 0.6 mm, length ca. 0.85 mm;
chorion with fine punctations; light yellow -when first laid, becoming
conspicuously collapsed as embryo develops; becoming transparent 2-
3 days prior to hatching, revealing gray larva,- with dark gray or blue-
gray spot representing head. Last Instar Larva . General (Fig. 1): Total
length 18.0-22.0 mm. Head small, smooth, shiny, without secondary
setae. Thorax inflated, larva appearing slightly humpbacked
(although not as pronounced as in Doa ampla). Integument with dense
28
J. Res. Lepid.
Fig. 1 . Last instar of Doa dora
Fig. 2. Integument of last instar larva showing spicules.
Fig. 3. Integumental spicules at higher magnification.
Fig. 4. Crochets of abdominal proleg of segment VI.
Fig. 5. Mouthparts of last instar.
28(1-2):1-136, 1989(90)
29
spicules (Figs. 2-3); all setae simple; pinacula small or absent. All
prolegs equal in size; crochets biordinal, in homoideous mesoseries
(covering approximately 0.60 perimeter of planta) (Fig. 4). Spiracles
small, elliptical, peritreme well sclerotized, uniform in size on A1-A7,
those on T1 and A8 larger.
Head : As in Figs. 5-7. Width 1. 8-2.0 mm. Height of frons approxi-
mately 0.8 mm. Length of epicranial suture approximately 0.75 x
height of frontoclypeus. Frontoclypeal height slightly greater than its
basal width. PI setae about twice as far apart as P2s, P2s located
dorsad of juncture of adfrontal line; A2 dorsoanterad of Al; LI nearly
in a straight line with Al and A2; L2 posteroventrad of LI. Six
stemmata (Fig. 7), 1 and 6 similar in size, larger than 2-5; stemmata
1-4 nearly equally spaced in an arc; 5 and 6 approximately equidis-
tant from 4. Seta S2 below stemma 1; SI below stemma 6. Labrum with
a broad, u-shaped, ventral notch. Mandible (Fig. 10) nearly square,
with two lateral setae; inner surface with 3 triangular teeth.
Thorax : As in Fig. 8. Segment Tl: Cervical gland absent; prothoracic
shield greatly reduced, bearing only XD1 and XD2. Dls closer to
meson than XDls; XD2s slightly further apart than D2s; SD1 dorsad
of spiracle; SD2 small, between XD2 and spiracle; L group bisetose,
anterad and slightly ventrad of spiracle; SV group bisetose. Segments
T2-T3: D2 closer to meson than Dl; SD2 directly ventrad of D2; SD1
antero ventrad of SD2; LI unisetose, in line with spiracles; L2 antero-
ventrad of LI; L3 dorsoposterad of LI, in nearly straight line with L2
and LI; an extra seta directly posterad of L2 and ventrad of L3; SV
group bisetose. Legs: Femur with 2 mesal setae; tibia with 6 setae in
ring around circumference; tarsus with 3 setae.
Abdomen : As in Fig. 8. Distance between D2s approximately 2 x
distance between Dls. A 1—8 with extra seta dorsad of D2, giving
appearance of 4 (total) equally spaced D2 setae in transverse line
across dorso-meson. SD1 dorsad of spiracle, SD2 greatly reduced. LI
unisetose, posterad of spiracle; L2 and L3 approximately halfway
between spiracle and SV1, about one spiracle height apart, L3 slightly
ventrad to L2. SV1 unisetose on Al-2 and A7-9, absent on A3 -6.
SV2 bi- or trisetose on A1-A6 (variable on opposite sides of same
segment), unisetose on A7-9. SV3 unisetose on Al-2, absent on A3-
9. A9 with D2, SD1, SD2, and LI on nearly contiguous pinacula in a
diagonal line. A10 (Fig. 9) with 20-24 setae irregularly arranged.
Prolegs with 12-15 lateral setae; planta with 20-24 biordinal crochets
in homoideous mesoseries.
Color : Head shiny brick red; a prominent black patch at stemmata;
clypeus and bases of antennae white; labrum black. Body with a series
of longitudinal stripes from meson to prolegs arranged as follows:
black at middorsum, bordered by white, black, yellow, black, white,
black, yellow with two black dots on each abdominal segment (anterior
one larger, including spiracle), black, white with two black dots per
30
J. Res. Lepid.
o
o
C\J
Fig. 6. Flead of last instar; anterior view.
Fig. 7. Arrangement of stemmata; lateral view; anterior at right.
abdominal segment, black; A- 10 brick red. Thoracic legs brick red;
tarsi black. Entire dorsal surface smooth and rather shiny; ventral
surface mostly black with diffuse yellow bands laterally between pairs
of legs.
Although D. dora is most similar to D. ampla in both superficial facies
and genital morphology, the larvae are remarkably different in colora-
tion.
Pupa : As in Figs. 11-12. Total length 14.5 mm. All appendages closely
appressed; setae sparse, similar to last instar. Head: Vertex simple,
rounded; epicranial suture indistinct. Antennae well defined, filiform,
extending nearly to caudal margin of wings. Labrum well defined,
square, with rounded corners; mandibles represented by triangular,
rounded regions adjacent to, and caudo-laterad of labrum; labial pal-
pus narrow, attenuate, ca 1.2 “x” as long as labrum; maxillae well
developed, extending ca 0.33 from eyes to caudal margin of wings.
28(1-2):1-136, 1989(90)
31
Fig. 8. Seta I map of last instar, T1 -2, A1 -2, A6-9; lateral view, anterior at
left.
Fig. 9. Seta I map of A9-10; dorsal view.
Fig. 10. Left mandible, mesal view.
Thorax: Prothorax dorsally a narrow collar; ventrally with legs well
defined, extending slightly less caudad than antennae. Mesothorax
dorsally broad with moderate, mesal, longitudinal, sclerotized hump;
ventrally with legs well defined, extending slightly caudad of anten-
32
J. Res. Lepid.
Fig. 1 1 . Pupa of Doa dora ; ventral view.
Fig. 12. Pupa of Doa dora ; dorsal view.
nae. Metathorax dorsally a moderate transverse band, with strongly u-
shaped margin anteriorly; margin of hindwings conspicuous along
entire latero-dorsum. Abdomen : Spiracles 1 -2 concealed beneath
wings; spiracles 3-8 with strongly sclerotized peritreme. Cremaster
indistinct with numerous long, distally-hooked bristles. The entire
pupa is brown, translucent, and shiny.
The cocoon is an unusual, single layered, wiry, open mesh, nearly
twice the volume of the pupa; the cast larval skin and head capsule are
included within the cocoon.
28(1-2):1-136, 1989(90)
33
Fig. 13-16. Adults of Doa dora: 13) Male from Baja California; 14) Female
from Baja California; 15) Male from Colima; 16) Female from
Nayarit.
Biology and Ecology
Doa dora (TL: Guadalajara, Jalisco, Mexico) (Figs. 13-16) is wide-
spread throughout northwestern Mexico, ranging from Baja California
to Tamaulipas, and as far south as Colima and Cuernavaca (label
data). Although adults have not been taken in the United States,
larvae have been collected in San Diego, California (Oceanside, 27-XT
76, on Wandering Jew [Commelinaceae], D. K. Faulkner, SDNHM).
Specimens from Baja California and Sonora may represent a closely
related, undescribed species. The same is likely for specimens from
Tamaulipas. Females from northwestern Mexico are distinguished
from typical D. dora by a more uniform gray forewing; males possess a
large, round, sclerotized region at the base of the valva lacking in D .
dora. In addition to D. dora , the genus includes D. ampla, D. raspa, D.
34
J. Res. Lepid.
cubana Schaus, D. translucida Dognin, and several undescribed spe-
cies from Mexico and Costa Rica. The relationship of Leuculodes to Doa
has not been examined in a phylogenetic context. The two appear to
represent sister taxa, although it is posible that they represent a single
genus. The group is in need of systematic revision.
In captivity, eggs of Doa dora are laid in irregular, contiguous,
parallel rows on the leaves and stems of the host. On Isla de Cedros,
the larval host is Euphorbia misera. Owing to the limited distribution
of E. misera (Munz 1974; Wiggins 1980), other euphorbiaceous plants
also must serve as larval hosts for Doa dora. Early instars live and feed
within a loose communal nest, dispersing and feeding externally on
the leaf surface in later stages. Early instar larvae will drop by a line
of silk when disturbed. In the laboratory, pupation occurred in debris
at the base of the host material. Developmental periods were as
follows: 10-12 days as ovum; 30-35 days as larva; 15-18 days as
pupa.
Females of D. dora appear to be partially diurnal, males appear to be
more so. The flight is weak and fluttering, similar to Ctenucha species.
Both sexes are attracted to blacklight (UV).
Discussion
The doids traditionally have been shuffled from family to family by
various authors who have based their hypotheses on adult morphologi-
cal characters. In the most recently proposed classification of the
nearctic Lepidoptera, Franclemont (1983) erected the tribe Doaini in
the Pericopinae (considered a subfamily of the Arctiidae), to accommo-
date the genera Doa and Leuculodes. However, characters of the larvae
contradict this placement.
According to Habeck (1987), pericopid and arctiid larvae, respective-
ly, are characterized by the presence of 3 and 4 verrucae above the
coxae on T2 and T3, and heteroideous crochets (except for some
lithosiines); the head is moderate in size, and integu mental spicules
are absent. In contrast, doid larvae lack verrucae on the thoracic coxae,
have homoideous crochets, the head is very small, and the integument
is covered with spicules.
Doid larvae share no uniquely derived characters with lymantriid
larvae. Symplesiomorphies include typical noctuoid chaetotaxy and
hypognathous head, elliptical spiracles, homoideous crochets, and fully
developed abdominal prolegs. Doids lack the abundant secondary
setae, which are responsible for the superficial similarity between
lymantriids and arctiids, have biordinal as opposed to uniordinal
crochets, and lack the fleshy, eversible middorsal gland on A7, which
appears to represent an autapomorphy for the Lymantriidae.
Notodontids, likewise, share many noctuoid symplesiomorphies with
the doids, but notodontid larvae can be distinguished from doids by
28(1-2):1~136, 1989(90)
35
their modified A10 prolegs (sometimes reduced to peg-like structures),
and the presence of two MD setae on T3 of the first instar (Hinton
1946), which appears to represent an autapomorphy for the Notodonti-
dae.
The dioptids, which probably represent a specialized group within the
Notodontidae, can be distinguished from doids by their larger head
(larger than prothorax) and uniordinal crochets. Although the larva of
Phryganidia California Packard is similar in general facies to that of
Doa dor a, and possess integumental spicules that are remarkably
similar to those of doids, features of the chaetotaxy, crochet arrange-
ment, and mandibular configuration suggest that the two are not
closely related.
The unique combination of larval characters possessed by doids
appears to contradict traditional assignments of this group to lepidop-
terous families of similar adult morphology. It is likely that an in-
creased knowledge of the distribution and significance of larval and
pupal characters among the various clades of the Noctuoidea may lead
to a greater understanding of phylogenetic relationships within this
superfamily.
Acknowledgements. I thank J. S. Miller, American Museum of Natural His-
tory, New York, New York, and J. E. Rawlins, Carnegie Museum of Natural
History, Pittsburgh, Pennsylvania, for their patience and for numerous helpful
comments on the manuscript; R. Poole, National Museum of Natural History,
Washington, D. C., for discussions and literature; J. P. Donahue, Natural
History Museum of Los Angeles County, for suggestions; D. K. Faulkner, San
Diego Natural History Museum, San Diego, California, for assistance in field
work and the loan of material in his care, and Victor Krantz, National
Museum of Natural History, Washington, D. C., for photographs of the adult
moths.
Literature Cited
BARNES, w. & J. mcdunnough. 1917. Check list of the Lepidoptera of Boreal
America. Herald Press, Decatur, 111. 392 pp.
BRYK, A. 1934. Lymantriidae, in Strand, Lepid. Cat. 62:356.
DONAHUE, J. P. & J. W. BROWN. 1987. The family Doidae, in Stehr, F. (ed.),
Immature Insects, vol. 1:534-536. Kendall/Hunt Publ. Co., Dubuque,
Iowa.
DYAR, H. G. 1903. A list of North American Lepidoptera. Bull. U. S. Natl. Mus.
52:261.
1911. Descriptions of the larvae of some Mexican Lepidoptera. Proc.
Entomol. Soc. Wash. 13:227-232.
1912. [Note describing early stages of Doa ampla .] in Walton, Notes
on certain species of flies. Proc. Entomol. Soc. Wash. 14:14-15.
FRANCLEMONT, J. G. 1983. The family Arctiidae, in Hodges, R. (ed.), Check List
of the Lepidoptera of America north of Mexico, pp. 114-119. E. W.
Classey Ltd., Wedge Entomol. Res. Found., London.
36
J. Res. Lepid.
HABECK, D. 1987. The families Arctiidae and Pericopidae, in Stehr, F. (ed.),
Immature Insects, vol. 1:536-542. Kendall/Hunt Publ. Co., Dubuque,
Iowa.
HINTON, H. E. 1946. On the homology of and nomenclature of the setae of
lepidopteraous larvae, with some notes on the phylogeny of the Lepidop-
tera. Trans. Royal Entomol. Soc. London 97:1-37.
HOLLAND, W. J. 1903. The Moth Book. Doubleday, Page, and Co., New York. pp.
309-310.
MCDUNNOUGH, J. 1938. Checklist of the Lepidoptera of Canada and the United
States of America. Part 1. Macrolepidoptera. Mem. So. Calif. Acad. Sci.
1:132.
MOSHER, E. 1916. A classification of the Lepidoptera based on characters of the
pupa. Bull. Illinois State Lab. Nat. Hist. 12:17-159 “+” plates.
MUNZ, P. A. 1974. A flora of southern California. Univ. Calif. Press, Berkeley,
Los Angeles, London. 1086 pp.
NEUMOEGEN, B. & H. DYAR. 1894. A preliminary revision of the Bombyces of
America north of Mexico. J. New York Entomol. Soc. 2:147-174.
PETERSON, A. 1948. Larvae of Insects, part 1. Edwards Brothers, Inc., Ann
Arbor, Michigan.
SCHAUS, w. 1927. Pericopidae, in Seitz, Gross-Schmett. Erde 6:563.
STEHR, F. 1987. Immature Insects, vol. 1:289-305. KendalPHunt Publ. Co.,
Dubuque, Iowa.
WALTON, w. R. 1912. Notes on certain species of flies. Proc. Entomol. Soc. Wash.
14:13-19.
WIGGINS, I. L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford,
Calif. 1025 pp.
Journal of Research on the Lepidoptera
28(l-2):37-74, 1989(90)
The Lepidoptera of a central florida sand pine scrub
community
Dennis Profant
Biology Department, Central Michigan University Mt. Pleasant, MI. 48859
Abstract. A Lepidoptera survey was conducted between September
1982 and April 1985 in the Sand Pine Scrub area of Blue Spring State
Park,* Volusia County, Florida. A total of 633 species comprising 43
families was recorded, including at least 12 undescribed species and
one verified state record. Abundance and monthly distribution records
are listed for moths. A floristic study of the scrub was also conducted.
Introduction
Blue Spring State Park is located in the west-central portion of
Volusia County just outside of Orange City, Florida. (Fig. 1). The area
consists of 590 hectares (1459 acres) of scrub, flatwoods, hammock,
swamps, marshes, and riverine environments, and has a subtropical
maritime climate. Volusia County has a mean temperature of about
21°C, and the mean annual rainfall is 1250mm. About 60% of the
annual rainfall occurs between the first of June and the middle of
October (USD A 1980). Volusia County sits within the lower Atlantic
Coastal Plain. The surface is covered with sandy marine sediments from
the late Pleistocene to Recent Age. Blue Spring is located on the extreme
western edge of the Deland Ridge, an ancient sand dune formed during
an interglacial period approximately 125,000 years ago.
With the cooperation of the Florida State Park Service, professional
and amateur lepidopterists have begun to accumulate much-needed
data on Florida Lepidoptera. Extensive surveys are being conducted in
north and south Florida at Torreya and Collier-Seminole State Parks,
respectively. The present study was done to provide additional distribu-
tion records for Lepidoptera, with a emphasis on moths, in the north-
central region of Florida by concentrating on one specific, and little
studied but important endemic plant community, the Sand Pine Scrub.
Monthly distribution and abundance figures for all moth species were
compiled, along with a floristics survey of the scrub.
Methods & Materials
Lepidoptera were collected an average of five times per week from September
1982 to April 1985. Collecting permits were issued annually from the Florida
Department of Agriculture and Deparment of Natural Resources. Butterflies
were recorded by collecting or by field sightings, but all moths were recorded
38
J. Res. Lepid.
Fig. 1. Study site and distribution of sand pine scrub.
only by collecting. Moths were collected at all hours except 0300 to 0600. Ten
existing mercury vapor lights on various park buildings were the primary
source for moths. Occasionally, filtered black lights were also used. A portable
generator was used in areas inaccesible to electricity. A bait of molasses, sugar,
and stale beer was brushed on tree bark, primarily to catch members of the
genera Catacola and Zale. The pheromone 3, 13- octadecadien- 1 -OL acetate
(ZZ-ODDA) was used to collect 3 of the 4 species of Sesiidae. Macrolepidoptera
were collected in cyanide and ethyl acetate killing jars. Microlepidoptera were
collected in small vials and frozen to prevent damage. Several species appeared
for only one or two months but were found in higher numbers than other species
recorded for five or six months. Therefore, monthly distribution was not
considered in determining abundance of each species. Abundance was deter-
mined by the total number of specimens observed during the 32 month
collecting period. The following criteria were used: uncommon (1-5 specimens),
occasional (6-20), common (21-50), abundant (51+). New species are indicated
in the checklist as n. sp. A question mark preceding a generic or specific name
indicates an uncertain determination.
Approximately one-third of the Lepidoptera were identified through the
taxonomic literature. Those references included Blanchard (1979), Blanchard &
Knudson (1983), Cashatt (1984), Coveil (1984), Eichlin & Cunningham (1978),
Hodges et al. (1983), Hodges (1986), Holland (1968), Howe (1975), Kimball
(1965), Klots (1951), Maxwell (1981), Mitchell & Zim (1977), Rockburne &
28(1-2):1-136, 1989(90)
39
SCRUB
HAMMOCK
MARSH
RIVER SWAMP
N
Fig. 2. The vegetation of Blue Spring State Park. Collecting sites are indicated
by circles.
Lafontaine (1976), and the USDA (1975). Approximately one-third were ident-
ified through the use of a comparative collection at the Florida Department of
Agriculture, Division of Plant Industry, Gainesville, Florida. The final third
were identified by D. Baggett, L. Dow, & J. Heppner. Forty species of microlepi-
doptera were deposited in the Division of Plant Industry collection (FSCA),
while all others remained in the private collection of the author.
A survey of the plants found in the scrub of Blue Spring was conducted
between April and August 1986. Plants were prepared with a standard leaf
press, then identified, mounted, and labeled. Voucher specimens of all vascular
plants collected are on deposit in the Florida State Museum’s Herbarium
(FLAS), University of Florida, Gainesville. References used for plant identifica-
tion included Cronquist (1980), Duncan (1967), Duncan & Foote (1975), Grimm
40
J. Res. Lepid.
(1966), Kartesz (1980), Kurz & Godfrey (1962), Radford et al. (1968), Tarver et
al. (1979), USD A (1982), and Wunderlin (1982).
Description of Study Area
Some moths not normally associated with a scrub environment were col-
lected. Since Lepidoptera may fly from one area to another, plant species in
several other plant communities surrounding the scrub may be serving as larval
food hosts. Therefore, the common vegetation of these communities was also
included in this study (FIG. 2).
HAMMOCK
Bordering the scrub throughout the park is a mesic mixed hardwood
hammock. The dominant species include Sabal palmetto (Walt.) Lodd. ex
Schult., Quercus virginiana Mill., Q. laurifolia Michx., Liquidamber styraciflua
L., and Magnolia grandiflora L. Common understory species include Quercus
nigra L., Carya glabra (Mill.) Sweet, Arilia spinosa L., Asimina paruiflora
(Michx.) Dunal, Callicarpa americana L., and Gelsemium semperuirens (L.) St.
J. H. Hil. Other common plants include Phlebodium aureum (L.) Small, Poly-
podium polypoidioides (L.) Watt, Vittaria lineata (L.) J. Smith, Mitchella repens
L., Epidendrum conopseum R. Br., Ruellia caroliniensis (J.F.Gmel.) Steud.,
Salvia lyrata L., and Elaphantopus elatus Bertol.
FLATWOODS & BAYHEAD
A major part of the flatwoods is dominated by Pinus elliottii Engelm. with a
thick understory of Serenoa repens (Bartr.) Small. Other important shrubs
include Ilex glabra (L.) A. Gray, Lyonia fruticosa (Michx.) G.S. Torr., L. lucida
(Lam.) K. Koch and Asimina reticulata Shuttlew. ex Chapm. Herbaceous plants
include Liatris tenuifolia Nutt., Sabatia brevifolia Raf., Polygala nana (Michx.)
DC, P. lutea L., Eriocaulon compressum Lam., and Lachnocaulon anceps (Walt.)
Morong. In the more poorly drained sites the dominant pine is typically Pinus
serotina Michx. Herbaceous plants in this area include Pinguicula pumila
Michx., Drosera sp., Utricularia sp., and Hypoxis sp. These soils become even
further saturated as a flatwoods depression forms a small bayhead on the south
edges of the park. The characteristic trees of this area are Taxodium distichum
(L.) L. Rich., Persea palustris (Raf.) Sarg., Gordonia lasianthus (L.) Ellis, and
Magnolia virginiana L. Understory plants include Smilax glauca Walt., Wood-
wardia areolata (L.) Moore, Osmunda cinnomomea L., and O, regalis L.
FLOODPLAIN FORESTS
Also known as river swamps, these areas border the St. Johns River and are
constantly inundated. These deciduous hardwood swamps consist of Sabal
palmetto , Taxodium distichum, Carya aguatica (Michx.) Nutt, ex Ell., Nyssa
biflora (Walt.) D. Sarg., Acer rubrum L.,Fraxinus caroliniana Mill., and Cornus
foemina Mill. Common herbaceous plants include Saururus cernuus L., Thalia
geniculata L., Crinum americanum L., and Aster caroliniana Walt.
28(1-2):1-136, 1989(90)
41
AQUATIC ENVIRONMENTS
These areas include the spring run, lagoon, freshwater marsh, and stream-
banks. The marshes are dominated either by Spartina bakeri Merr. or Panicum
hemitomon Schult. Commonly scattered along marsh edges are woody species
such as Salix caroliniana Michx., Sambucus canadensis L., and Cephalanthus
occidentalis L.
The open areas of the river, lagoon, and spring run include plants such as
Pistia stratiotes L., Eichhornia crassipes (Mart.) Solms, Nuphar luteum (L.)
Sibth. + J.E. Smith, Ceratophyllum demersum L., and Salvinia minima Baker.
Many plants found along the banks of these waters occur naturally or were
washed in from the river. Common species along the waters edge include
Sagittaria latifolia Willd., Alternantha philoxeroides (Mart.) Griseb., Ponte -
deria cordata L.,Kosteletzkya virginica (L.) Presl ex A. Gray. Hibiscus coccineus
(Medic.) Walt., Amaranthus australis (A. Gray) Sauer, Vigna luteola (Jacq.)
Benth., Lythrum salicaria L., and Paspalum repens Berg.
SCRUB
Several times during Florida’s history, the sea levels were higher than they
are today and the coastline was much further inland. Sand dunes formed along
these ancient shorelines and still persist today. These are the natural sites of the
Sand Pine Scrub community in Florida (DNR 1975). With the exception of a few
locations in Alabama, the Sand Pine Scrub is restricted to the state of Florida
(Laessle 1958). The scrub consists of well-drained, fine white siliceous sands and
is composed almost entirely of thick growths of broad-leaved evergreen shrubs.
Because of the sterile soils, there is very little diversity among the herbaceous
plants. Although a fire-dependent community, ground cover is sparse and leaf
litter accumulates very slowly. Therefore, fires are infrequent, perhaps every 20
to 40 years. When a fire does occur, it will burn hot enough to allow the
serotinous cones of the Sand Pine to open and begin dropping seeds. If a scrub is
not exposed to fire, it will most likely succeed into a xeric hammock (Monk
1968). Due to their dry upland locations, scrub environments are rapidly being
lost to real estate development, and therefore are considered highly endangered
areas (DNR 1975).
The Sand Pine Scrub of Blue Spring is part of a much larger scrub which
extends south and east through Orange City and Deltona. Due to the growth of
the area, especially in Deltona, this scrub is disappearing. The scrub within the
boundaries of Blue Spring consists of approximately 202 hectares (500 acres),
situated on soils of Daytona and Paola fine sand (USD A 1980). North and east of
the park, nearly 200 more hectares continue to occur on Apopka fine sands until
they meet a Longleaf Pine/Turkey Oak Sandhill area.
The overstory of the Blue Spring scrub is dominated by sand pine, {Pinus
clausa (Chapm. ex Engelm.) Vasey ex Sarg.). The understory consists of three
dominant scrub oaks: sand-live oak {Quercus geminanta Small), myrtle oak (Q.
myrtifolia Willd.), and chapman oak (Q. chapmanii Sarg.). Other important
shrubs include devilwood or wild olive ( Osmanthus americana (L.) Benth. &
Hook. f. ex Gray), scrub holly {Ilex opaca Ait. var. arenicola (Ashe) Ashe),
Carolina holly {I. ambigua (Michx.) Torr.), saw palmetto ( Serenoa repens ),
silkbay (Persea humilis Nash), and rusty lyonia {Lyonia ferruginea (Walt.)
Nutt.). The ground cover includes small leaved blueberry {Vaccinium myrsi-
42
J. Res. Lepid.
nites Lam.), gopherapple ( Licania michauxii Prance), and scattered lichens
Cladina spp. Occasionally a scrub will lack sand pine all together, yet the
understory will have the same species composition. This situation is found in a
20 hectare section of the park scrub.
Many of the sand pines in the park are beginning to degenerate. By 50 years of
age, heartrot is a common occurrence. With such a dense understory, competi-
tion has made it difficult for sand pine to regenerate. Only in the highly
disturbed areas such as old fire roads and borrow pits are the sand pine seedlings
growing successfully. Due to the disturbed nature of this scrub, many succes-
sional plant species have invaded the area and this is resulting in a faster
accumulation of leaf litter. Because of the campground and cabins, the high
recreational use of the area makes it unfeasible for prescribed burning. With
each passing year, the scrub accumulates large amounts of herbaceous and
woody litter, both on the soil surface and in the trees. This suggests that the
probability of a wildfire is greatly increased (Veno 1976).
Annotated List of Scrub Plants
The following is a list of plants recorded from the scrub of Blue Spring.
Vascular plant nomenclature follows that of Wunderlin (1982). Genera and
species within the families are arranged alphabetically.
PINACEAE
Pinus clausa (Chapm. ex Engelm.)
Vasey ex Sarg.
POACEAE
Andropogon glomeratus (Walt.)
BSP.
var .glaucopsis (Ell.) Mohr.
Eustachys neglecta (Nash) Nash
Panicum ciliatum Ell.
P. commutatum Schult.
P. miliaceum L.
Paspalum notatum Fluegge.
Setaria geniculata (Lam.) Beauv.
CYPERACEAE
Rhynchospora megalocarpa A. Gray
ARECACEAE
Serenoa repens (Bartr.) Small
XYRIDACEAE
Xyris caroliniana Walt.
JUNCACEAE
J uncus scirpoides Lam.
SMILACEAE
Smilax auriculata Walt.
S.glauca Walt.
S.pumila Walt.
AGAVACEAE
Y ucca flaccida Haw.
MYRICACEAE
Myrica cerifera L.
FAGACEAE
Quercus chapmanii Sarg.
Q. geminata Small
Q. laurifolia Michx.
Q. myrtifolia Willd.
ULMACEAE
Ulmus americana L.
POLYGONACEAE
Polygonella gracilis (Nutt.) Meisn.
AMARANTHACEAE
Froelichia floridana (Nutt.) Moq.
MAGNOLIACEAE
Magnolia grandiflora L.
ANNONACEAE
Asimina ohovata (Willd.) Nash
LAURACEAE
Cinnamomum camphora (L.) Presl
Persea humilis Nash
BRASSICACEAE
Lepidium virginicum L.
ROSACEAE
Prunus serotina Ehrh.
CHRYSOBALANACEAE
Licania michauxii Prance
FABACEAE
28(1-2):1-136, 1989(90)
43
Amorpha fruticosa L.
Desmodium incanum DC.
D. tortuosum (Sw.) DC.
Galactia elliottiiNutt.
G. floridana Ton*. & Gray
G. ? regularis (L.) BSP
Medicago lupulina L.
EUPHORBIACEAE
Chamaesyce hyssopifolia (L.) Small
Cnidoscolus stimulosus (Michx.)
Engelm. & Gray
Croton glandulosus L.
EMPETRACEAE
Ceratiola ericoides Michx.
ANACARDIACEAE
Rhus copallina L.
AQUIFOLIACEAE
Ilex ambigua (Michx.) Torr.
I. opacaAit. var. arenicola (Ashe)
Ashe
VITACEAE
Ampelopsis arborea (L.) Koehne
Parthenocissus quinque folia (L.)
Planch.
Vitis aestivalis Michx.
V. rotundifolia Michx. ( munsoniana
Simpson of some authors)
CLUSIACEAE
Hypericum hypericoides (L.)
Crantz.
H. reductum P. Adams
CISTACEAE
Helianthemum corymbosum Michx.
Lechea mucronata Raf.
PASSIFLORACEAE
Passiflora incarnata L.
CACTACEAE
Opuntia humifusa (Raf.) Raf.
ONAGRACEAE
Gaura angustifolia Michx.
Oenothera laciniata Hill
APIACEAE
Apium leptophyllum (Pers.) Muell.
ERICACEAE
Befaria racemosa Vent.
Gaylussacia dumosa (Andrz.)
T.&G.
G. tomentosa (A. Gray) Small
Lyonia fer rug inea (Walt.) Nutt.
L. lucida (Lam.) K. Koch
Vaccinium myrsinites Lam.
V. stamineum L.
SAPOTACEAE
Bumelia tenax (L.) Willd.
EBENACEAE
Diospyros virginiana L.
OLEACEAE
Osmanthus americana (L.) Benth. &
Hook. f. ex Gray
LOGANIACEAE
Polypremum procumbens L.
ASCLEPIADACEAE
Asclepias tomentosa Ell.
CONVOLVULACEAE
Ipomoea pandurata (L.) G.F. W. Mey
Merremia dissecta (Jacq.) Hall. f.
POLEMONIACEAE
Phlox drummundii Hook.
VERBENACEAE
Callicarpa americana L.
LAMIACEAE
Hyptis mutabilis (A. Rich) Briq.
Monarda punctata L.
Salvia lyrata L.
Teucrium canadense L.
Trichostema dichotomum L.
SCROPHULARIACEAE
Gratiola hispida (Benth.) Pollard
Linaria canadensis (L.) Dum.
Seymeria pectinata Pursh.
BIGNONIACEAE
Campsis radicans (L.) Seem, ex
Bureau
RUBIACEAE
Diodia teres Walt.
Richardia brasiliensis (Moq.)
Gomez
ASTERACEAE
Baccharis halimifolia L.
Berlandiera subacaulis (Nutt.)
Nutt.
Bidens alba (L.) DC.
Carphephorus corymobsus (Nutt.)
Torr. & Gray
C. odoratissimus (J.F.Gmel.) Herb.
Erigeron strigosus Muhl.
Eupatorium compositifolium
Walt.
Garberia heterophylla (Bartr.)
Merr. & Harp.
44
J. Res. Lepid.
Gnaphaliumfalcatum Lam.
Heterotheca subaxillaris (Lam.)
Nutt.
Pterocaulon virgatum (L.) DC.
Pyrrhopappus carolinianus (Walt.)
DC.
Britt. & Rusby
Hieracium megacephalon Nash.
Krigia virginca (L.) Willd.
Lactuca gramini folia Michx.
Pityopsis graminifolia (Michx.)
Solidago sp.
V ernonia gigantea (Walt.) Trel. ex
Branner & Coville
Results & discussion
A total of 633 species of Lepidoptera were recorded, consisting of 591
moths and 42 butterflies in 43 families. Families with the most species
recorded were Noctuidae (172), Pyralidae (100), and Tortricidae (76).
The average monthly distribution curve shows that the greatest species
diversity occurred in the spring, and the least diversity in the summer
(Fig. 3). The highest total was in March (201 species) and the lowest was
in July (32 species). The fall, winter, and spring months were surveyed
for three years, while the summer months were surveyed for only two
years. Thus, sampling time may account in part for the lower number of
species throughout June, July, and August.
#226, previously known from Florida as E. poaphilodes, is now listed
as E. fergusoni (Solis 1986). #197 is unconfirmed as being collected
within the boundries of Blue Spring. #555 is unconfirmed as the
specimen is missing. #506 was identified from the casings. #485 was
collected at light and not with the Sesiidae pheromone. Heppner
(personal communication) indicated that the collection of #134 Phyl-
lonorycter fitchella (Gracillariidae) represented the first report of this
species in Florida. Twelve other microlepidoptera were determined by
Heppner as being new species, most or all of which should be state
records upon their description. Doug Ferguson, of the Smithsonian
Natural History Museum, and David Baggett (personal communi-
cation), indicated that #129 is probably a worn specimen of frondaria or
N. bifiliata , and its very faded condition makes a final determination
unlikely. Synchlora aerata has yet to be recorded this far south. Baggett
indicates that #187 identified here as Arugisa latiorella , may be A.
watsoni Richards. Baggett also mentioned that #257, #258, and #559
may be state records and, upon future examination by taxonomists, the
list should contain other state records as well as hundreds of county
records. This study has facilitated a better understanding of the
distribution of Florida Lepidoptera, and should also provide the basis for
further investigations into host plant relationships and possible ende-
mic lepidopterans of scrub environments.
28(1-2):1-136, 1989(90)
45
Fig. 3. Average monthly distribution of moth species between September
1982 and April 1985.
Acknowledgements. I wish to than K Jim Stevenson, Doug Carter, Charles
DuToit, Walt Young, and Nick Robbins of the Florida Department of Natural
Resources for their cooperation. Linwood “Woody” Dow of Largo Florida, H.
David Baggett, research associate (FSCA) Gainesville, Florida, and John
Heppner, Division of Plant Industry (FSCA), Gainesville, provided valuable
help in Lepidoptera identification. From Central Michigan University, Ray-
mond Hampton & Roger Bland assisted in photography and the writing of this
paper, respectively. At the University of Florida, Kent Perkins, David Hall
& Walter Judd helped in plant identification. I especially thank Thomas C.
Emmel, University of Florida, for his initial support of this project, continued
advice, and review of the manuscript.
Literature Cited
BLANCHARD, A., 1979. Five new species of the tribe Eucosmini (Tortricidae). J.
Lepid. Soc. 33:209215.
BLANCHARD, A. & E. C. KNUDSON, 1983. Two new species of the tribe Eucosmini
(Tortricidae) closely related to Phaneta granulatana (Kearfott). J. Lepid.
Soc. 37:140-145
46
J. Res. Lepid.
CASHATT, E. D., 1984, Revision of the genus Parachma Walker (Pyralidae:
Chrysauginae) of America north of Mexico, with a description of a new
genus. J. Lepid. Soc. 38:268-280
COVELL, C. V. Jr., 1984. A Field Guide to the Moths of Eastern North America.
Houghton Mifflin Co., Boston. 496p.
CRONQUIST. A., 1980. Vascular Flora of the Southeastern United States. Vol.l
Asteraceae. Univ. of North Carolina Press, Chapel Hill, 261p.
DEPARTMENT OF NATURAL RESOURCES, 1975. Florida Environmentally Endan-
gered Lands Plan. Division of Recreation & Parks, Tallahassee. 143p.
DUNCAN, w. H., 1967. Woody vines of the southeastern states. SIDA Cont. to Bot.
3:1-76
DUNCAN, W. H. & L. E. FOOTE, 1975. Wildflowers of the Southeastern United States.
Univ. of Georgia Press, Athens. 296p.
EICHLIN, T. D. & H. B. CUNNINGHAM, 1978. The Plusiinae (Lepidoptera: Noctuidae)
of America north of Mexico, emphasizing genitalic and larval morphology.
USDA Tech. Bull. #1567. Washington, D. C. 122p.
GRIMM, w. C., 1966. Recognizing Native Shrubs. Stackpole Books, Harrisburg,
PA. 319p.
HODGES, R. W., et al., 1983. Checklist of the Lepidoptera of America North of
Mexico. Univ. Press, Cambridge. 284p.
HODGES, R. W., 1986. Gelechioidea: Gelechiidae (in part) in Dominick, R. B., et al.,
The Moths of America North of Mexico, fasc. 7.1 Allen Press Inc. Lawrence,
Kansas. 195p.
HOLLAND, W. J., 1968. The Moth Book. Dover Publ. New York. 479p.
HOWE, w. H., 1975. The Butterflies of North America. Doubleday & Co. Inc.,
Garden City, N. Y. 633p.
KARTESZ, J. T. & R. KARTESZ, 1980. A Synonymized Checklist of the Vascular Flora
of the United States, Canada, and Greenland. Univ. of North Carolina
Press, Chapel Hill. 498p.
KIMBALL, C. P., 1965. Lepidoptera of Florida. Div. of Plant Industry, Florida Dept.
Of Agr., Gainesville. 311p.
KLOTS, A. B., 1951. A Field Guide to the Butterflies of North America, East of the
Great Plains. Houghton Mifflin Co., Boston. 349p.
KURZ, H. & R. K. GODFREY, 1962. Trees of North Florida. Univ. of Florida Press,
Gainesville. 31 Ip.
LAESSLE, A. M., 1958. The origin and successional relationships of sandhill
vegetation and sand pine scrub. Ecol. Monogr. 28:361-387
MAXWELL, L. S., 1959. Florida Insects. L.S. Maxwell Publ., Tampa. 120p.
MITCHELL, R. T. & H. S. ZIM, 1962. Butterflies and Moths. Golden Press. New York.
160p.
MONK, C. D., 1968. Successional and environmental relationships of the forest
vegetation of north central Florida. Am. Midi. Nat. 79:441-457
RADFORD, A. E„ H. E. AHLES & C. R. BELL, 1968. Manual of the Vascular Flora of the
Carolinas. Univ. of North Carolina Press, Chapel Hill. 1183p.
ROCKBURNE, E. W. & J. D. LaFONTAINE, 1976. The Cutworm Moths of Ontario and
Quebec. Canada Dept, of Agriculture, Ottawa. 164p.
SOLIS, M. A., 1986. A new species of Epidromia (Noctuidae) from Florida. J. Lepid.
Soc. 40:8-19
TARVER, D. P., J. A. RODGERS, M. J. MAHLER, R. L. LAZOR, 1979.
28(1-2):1-136, 1989(90)
47
Aquatic and Wetland Plants of Florida. Florida Dept, of Natural Resources,
Tallahassee. 127p.
U.S. department OF AGRICULTURE (USDA)., 1975. Seed and Cone Insects of
Southern Pines. U.S. Forest Service. General Technical Report SE-8.
Asheville. 41p.
— , (USDA). 1980. Soil Survey of Volusia County, Florida. Soil Conserva-
tion Service, IF AS, Gainesville.
— -, (USDA). 1982. National List of Scientific Plant Names. Soil Conserva-
tion Service, TP-159. Washington, D.C. 416p.
VENO, P.A., 1976. Successional relationships of five Florida plant communities.
Ecol. 57:498-508.
WUNDERLIN, R. P., 1982. Guide to the Vascular Plants of Central Florida. Univ.
Presses of Florida, Tampa. 472p.
48
J. Res. Lepid.
Table 1. Complete listing of Lepidoptera of Blue Spring — Abundance
indicated by C = common, O = occasional, U = uncommon,
A = abundance
Abundance Month
APATELODIDAE
1 Olceclostera indistincta (Hy. Edw.) C - F M ------
ARCTIIOAE
2 Afrida ydatodes Dyar. A JFM-----S0N-
3 Cisseps ful vicol 1 is (Hbn.) U ^
4 Cisthene packardi (Grt.) C * F M - N -
5 Cisthene striata Ottol . A --MAM-----N-
6 Cisthene subjecta Wlk. A J-MAM ■ 0 N -
? Cisthene tenui fascia Harv. 0 ---A--------
8 Clemensia a'lbata Pack. A J - M A — ■
9 Cosmosoma my r odor a Dyar 0 - F ---NO
10 Crambidia 1 ithosioides Dyar 0 ----------n-
11 Ecpantheria scribonia (Stoll) 0 - - M
12 Estigmene acrea (Drury) U --M---------
13 Euerythra phasma Harv. 0 -FM--J------
14 Grammia nais (Drury) C JF-----AS---
15 Halysidota tessellaris (J.E. Smith) C -F--MJ---S---
16 Holomelina aurantiaca (Hbn.) C JFMA
17 Holomelina ferruginosa (Wlk.) C - F M ------
18 Holomelina laeta (Guer .-Meneville) U — _ _ _ q
19 Holomelina opella (Grt.) U -FM---------
20 Holomelina rubicundaria (Hbn.) D ---A--------
21 Hyphantria cunea (Drury) A JFM-----S---
22 Hypoprepia miniata (Kby.) A AM----0--
23 Leucanopsis longa (Grt.) U - - -----q
24 Pyrrharctia Isabella (J.E. Smith) 0 - F
25 Spi losoma congrua Wlk. U JF
26 Utetheisa bell a (L.) C J--- • - - - N -
8LAST08ASIDAE
27 Glyphidocera lactif losella (Cham.) C -------- s 0 - -
28 Holcocera ? lepidophaga Clarke C --M A - -- -- -- -
29 Valentinia glandulella (Riley) C ■ " 0 - -
30 sp. U - - M A - - - - -
COCHYLIDAE
31 Aethes sp. U --M-
32 Aethes sp. U 0
33 Aethes sp. U -----------o
34 Aethes sp. U
35 Aethes sp. 0 --MA----SQN-
36 Carol el 1 a bimaculana (Rob.) C ---AMJJ--0N-
37 Carolella erigeronana (Riley) 0 - - - A - - J - -
28(1-2):1-136, 1989(90)
49
Table 1 (Continued)
38 Carolella sartana (Hb.)
39 Hysterosia argent ili mi tana Rob.
COLEOPHORIOAE
40 Homaledra sabalella (Cham.)
C0SM0PTERIGIDA6
41 Cosmopterix prob. gemmi fere'll a Clem.
42 Euclemensia bassettella (Clem.)
43 Perimede erransella Cham.
COSSIOAE
44 Givi ra francesca (Dyar)
45 Prionoxystus robin iae (Peck)
OREPANIDAE
46 Eudeilinea luteifera Oyar
ERIOCRANIOAE
47 Eriocraniella mediabulla Davis
GELECHI IDAE
48 Anacampsis coverdalella Kft.
49 Aristotelia roseosuffusella (Clem.)
50 Aristotelia sp.
51 Aroga coloradensis (8sk.)
52 Die homer is ? georgiella (Wlk.)
53 Evippe prunifoliella Cham.
54 Exoteleia pinifoliella (Cham.)
55 Polyhymno luteostrigella Cham.
56 ?Sinoe sp.
57 Stegasta bosqueella (Cham.)
58 Telphusa sp.
59 Dichomeris ? xanthoa Hodges
60 sp.
61 sp.
62 Dichomeris ? aglaia Hodges
63 sp.
64 sp.
GEOMETRIDAE
65 Anacamptodes defectaria (Gn.)
66 Anacamptodes vellivolata (Hulst)
67 Anavitrinella pampinaria (Gn.)
68 8esma quercivoraria (Gn.)
69 Caripeta aretaria (Wlk.)
70 Chlorochlamys chloroleucaria (Gn.)
71 Chloropteryx tepperaria (Hulst)
Abundance Month
0 A
0 J F - A
U - - - A
U - p M ------ - —
U ------- A S - - -
U - - M
U - - - A — -
y - - M - -
U s
U - - - A
0 M -
0 J-M-M —
U - F --ND
A J - - - N -
U - F
0 - F M
0 - F H -
0 - - M -
0 J F-
C AM SO —
U j - M - -- -- -- - -
U - - - - ,() —
U A - -
U - F
U D
U - - - a ------- -
U - F
U j f - - M - -
U - - - A - D
C - F M - M - - N -
U - p M -------- -
0 J F M
C -FMAM-------
C -FMA SO —
50
J. Res. Lepid.
Table 1 (Continued)
Abundance Month
72 Cyclopbora myrtaria (6n.) U
73 Cyiatophora approximaria Hbn. C
74 Olchorda Irldaria Tati pennis (Hulst) 0
75 Disci isioppocta stellate (Gn.) U
78 Dyspteris abortivaria H.-S. 0
77 Epiiecis hortaria (F. ) C
78 Episeiasia soli tana (Ilk.) U
79 Euchlaena aioenaria astylusaria (Ilk.) 0
80 Euchlaena deplanaria (Ilk.) 0
81 Eulithis diversilineata (Hbn.) C
12 Eupithecia liserulata Grt. 0
83 Eusarca eonfusaria Hbn. 0
84 Eusarca fundaria (Gn.) U
85 tut rape! a clemstaria ( J * E . Smith) A
86 Glenoides texanaria (Hulst) A
S? Hethemia pistasciaria insecutata (Ilk.) U
88 Bydr i often a pluviata meridianata McD. 0
8§ Hypagyrtis esther (Barnes) C
§0 Byponticis umbrosaria (Hbn.) U
91 Idaea demissaria (Hbn.) C
92 Idaea ereraiata (Hulst) U
93 Idaea ostentaria (Ilk.) U
94 Idaea tacturata (Wlk.) 0
§5 lambdina pultaria (Gn.) U
98 Leptostales pannaria (Gn.) 0
97 lobocleta peralbata (Pack.) U
98 Lophosis labeculata (Hulst) A
99 lychrwsea interaicata (Ilk.) U
108 lyeia ypsilon carlotta (Hulst) C
101 Melanolophia canadaria (Gn.) 0
102 $et arrant his homuraria (G.& R.) U
103 He tar ran this obfimaria (Hbn.) U
104 Nacophora quernaria (J.E. Smith) C
105 Neiatocarapa 1 imbata (Haw.) 0
108 Nemoria b. bifiliata (Wlk.) C
107 Neioria catachloa (Hulst) C
108 Nemoria elfa Fgn. C
109 Neioria lixaria (Gn.) A
110 Nemoria saturiba Fgn. U
111 Nepheloleuca floridata (Grt.) U
112 Orthonama centrostpigaria (H.-S.) C
113 Orthonaia obstipata (f.) U
114 Patalene olyzonaria (Wlk.) C
- - - - ■ - - - N -
J F--M-- — 0--
-F M' - -
-FM - -- -- -- --
--HA. •-
-F------S---
-FH-----S0N-
----M----0N-
J-M-------ND
-FM--
-FH
JFM 0
--M---------
--MA-
JF-AH-J--0N-
---AM---S0--
---A--------
---A----S0N-
JF--------ND
JFMA----S-N-
---A--------
-FM---------
--MA--------
-FM---------
-FM---------
---AM-------
-FMAM-JA-Q--
-FMAM-J-S-N-
-F-------0ND
J-MAMJJ-SO-D
---AM-------
------- A - - - -
JF-A-------D
-F--- - - -- -- -
28(1-2):1-136, 1989(90)
51
Table 1 (Continued)
Abundance Month
115 Phigalia strigataria (Minot) U - - M
116 Phrudocentra centrifrugaria (H.-S.) U - - - . - - - o - -
117 Pleuroprucha insulsaria (Gn.) U - - - 0 N D
118 Prochoerodes transversata incurvata (Gn.) 0 - M
119 Protoboarmia porcelaria (Gn.) 0 -'EM--
120 Scopula aemulata (Hulst) U -F-*
121 Scopula compensata (Wlk.) U J N -
122 Scopula lautaria (Hbn.) 0 JFMA - “ D
123 Scopula timandrata (Wlk.) U - - M - M -
124 Semiothisa bicolorata (F.) 0 J - M - M ~ ~ N D
125 Semiothisa distribuaria (Hbn.) U ----M----
126 Semiothisa gnophosaria (Gn.) U J - - A *
127 Semiothisa sanfordi Rindge C JFMAM SON-
128 Stenaspilatodes antidiscaria (Wlk.) U - F - -
129 Synchlora looks like aerata (F.) U ---------o--
130 Synchlora frondaria Gn. C JFMA-J--S-N-
131 Synchlora gerularia (Hbn.) C -FMA--J---N0
132 Tornos scolopacinarius spodius Rindge C - F M A
GLYPHIPTERIGIDAE
133 Diploschizia sp. U ---A-----
GRACILLARIIOAE
134 Phyllonorycter fitchella (Clem.) U - - - 0 - -
135 sp. U - - M
136 sp. U --------- o - -
137 sp. U - - - a - - - -
INCURVARIIDAE
138 Adela caeruleella Wlk. U - - M
LASiOCAMPIOAE
139 Artace cribraria (Ljungh) C
140 Malacosma americana (F.) A - - m A -
141 Malacosma disstria Hbn. A - M - -
142 Phyllodesma americana (Harr.) C -FMA---
143 Tolype minta Dyar U S
144 Tolyoe not ial is Franc. C J-MA-J--S-ND
LIMACODIDAE
145 Adoneta spinuloides (H.-S.) U ---S'
146 Apoda Y-inversa (Pack.) U ~ - M
147 Apoda rectilinea (G.& R.) C ---A-J--S
148 Euclea delphinii (8dv.) A --MAMJ----ND
149 Isa textula (H.-S.) A -------- s 0 N D
150 Isochaetes beutenmulleri (Hy.Edw.) U - - - - A-
151 Lithacodes gracea Dyar 0 A--J
152 Monoleuca erect i fascia Dyar U J -
52
J. Res. Lepid.
Table 1 (Continued)
153 Monoleuca near semifascia (WHO
154 Monoleuca subdentosa Dyar
155 Natada nasoni (Grt . )
156 ProJImacodes bad la (Hbn.)
157 Si bine stimulea (Clem.)
LYMANTRI IDA6
158 Dasychira leucophaea (J.E. Smith)
159 Dasychira manto (Stkr.)
160 Dasychira tephra Hbn.
MEGAL0PY6 I DAE
161 Lagoa lacyi 8.& McD.
162 Megalopyge opercular is (J.E. Smith)
MIMALLONIDAE
163 Cicinnus melsheimeri (Harr.)
MOMPHIDAE
164 Mompha eloisella (Clem.)
NEPTICULIDAE
165 ?Ectodemia sp.
NOCTUIDAE
166 Ababletnma brimleyana (Dyar)
167 Acronicta afflicta Grt.
168 Acronicta americana (Harr.)
169 Acronicta brumosa Gn.
170 Acronicta hasta Gn.
171 Acronicta impleta Wlk.
172 Acronicta lanceolaria (Grt.)
173 Acronicta oblinita (J.E. Smith)
174 Acronicta tritona (Hbn.)
175 Acronicta vinnula (Grt.)
176 Agrotis subterranea (F.)
177 Alypia wittfeldi Hy.Edw.
178 Amolita fessa Grt.
179 Amolita obi i qua Sm.
180 Anicla infecta (Ochs.)
181 Anomis erosa Hbn.
182 Anomis flava fimbriago (Steph.)
183 Anomogyna elimata (Gn.)
184 Anticarsia gemmatilis Hbn.
185 Argyrogramma basigera (Wlk.)
186 Argyrostrotis quadrif i laris (Hbn.)
187 Arugisa ? latiorel la (Wlk.)
188 Bagisara repanda (F.)
Abundance Month
l) - J ■ -
A * A-JJ-S
U -----j--
A A-JJ-S
C J-SON-
0 ---A-
C J-M-M-----N-
0 - — - M — - - 0 - -
C MJJ
A MJJ-S
U ----M-------
0 A -----
U ---------- - D
0 — MA — 0 N -
U - - S
U — M
U - - m A ------- -
y -p----------
U - F M ------
U - - M
U J - - -
0
0 JF-----AS-N-
U - - -v -------- -
U
U - — a — - - - 0 - -
0 J F M A - -
U - F --------- -
U ------ N -
U - - - - N -
U ---------- N -
0 --------SOND
U ----M-----N-
U --M-M---
U --M------
U ----------- D
28(1”2):1-136, 1989(90)
53
Table 1 (Continued)
Abundance Month
189 Bellura gortynoides Wlk.
U
-
- - A - -
-
A
-
0
-
-
190 Bellura obi i qua (Wlk.)
U
-
F M
191 Bleptina caradrinalis 6n .
U
-
F M
192 8omolocha bal timoral is (Gn.)
U
-
- M A - -
-
-
-
-
~
-
193 Caenurgia chloropha (Hbn.)
U
J
- M - - -
194 Callopistria cordata (Ljungh)
U
-
- - A —
-
-
-
-
-
-
195 Callopistria granitosa (6n.)
U
-
-
-
S
-
-
-
196 Callopistria moll issima (Gn.)
U
-
--AM-
-
-
~
-
“
-
197 ? Catocala arnica (Hbn.)
0
J
198 Catocala andromedae Gn.
A
M -
199 Catocala cara Gn.
U
- - - - J
200 Catocala cl in ton ii Grt.
u
-
--AM-
-
-
-
-
-
-
201 Catocala connubial is Gn.
u
M -
202 Catocala consors (J.E. Smith)
u
J
203 Catocala ilia (Cram.)
A
-
- - - M J
-
-
-
-
-
-
204 Catocala jair Stkr.
U
- - - - J
205 Catocala louiseae J. Bauer
c
- - - M -
206 Catocala micronympha Gn.
0
-
M J
-
-
-
-
-
-
207 Catocala muliercula Gn.
c
-
- - - M J
-
~
-
-
-
-
208 Catocala similis Edw.
A
- - - M -
209 Catocala ultronia (Hbn.)
C
- - - M -
210 Chaetaglaea tremula (Harv.)
0
J
F - - - -
D
211 Charadra deridens (6n.)
0
-
F M - - J
J
-
S
-
-
-
212 Cissusa spadix (Cram.)
0
-
F M A - -
213 Copipanolis styracis (Gn.)
A
J
F - - - -
214 Cryphia nanoides Franc. & Todd
c
-
-MAM-
-
-
-
0
-
-
215 Cutina albopunctella Wlk.
u
- M
216 Cutina distincta (Grt.)
u
- M
217 Cutina sp.
0
-
F - A - -
218 Cutina sp.
0
-
F
219 Derrima stellata Wlk.
u
-
- - - - -
-
-
s
-
-
-
220 Dyspyralis n. sp.?
u
(
—
-
-
-
0
-
-
221 Egira alternans (Wlk.)
u
- M - - -
222 Elaphria chalcedonia (Hbn.)
u
- - A - -
0
-
0
223 Elaphria exesa (Gn.)
u
- M - - -
224 Elaphria festivoides (Gn.)
c
-
- M A - -
-
-
-
-
N
0
225 Elaphria versicolor (Grt.)
0
-
F - - - -
D
226 Epidromia fergusoni Solis
u
-
-
-
-
0
-
-
227 Euclida cuspidea (Hbn.)
u
- - A - -
228 Eucoptocnemis dapsilis (Grt. )
u
N
-
229 Eudryas grata (F.)
u
- - - - J
230 Eudryas unio (Hbn. )
u
- M
54
J. Res. Lepid.
Table 1 (Continued)
Abundance Month
231 Eumicremma minima (Gn.)
232 Eutolype rolandi Grt.
233 Feltia geniculata G.& R.
234 Galgula partita Gn.
235 Harrisimemma trisignata (Wlk.)
236 Heliothis turbatus (Wlk.)
237 Heliothis virescens (F.)
238 Hemeroplanis habital is (Wlk.)
239 Himella intractata (Morr.)
240 Homophoberia cristata Morr.
241 Hormisa orciferalis Wlk.
242 Hormoschista latipalpis (Wlk.)
243 Hypenula cacuminal is (Wlk.)
244 Hypsoropha hormos Hbn.
245 Hypsoropha monilis (F. )
246 Idia aemula Hbn.
247 Idia americalis (Gn.)
248 Idia lubrical is (Gey. )
249 Iodopepla u-album (Gn.)
250 Isogona tenuis (Grt.)
251 Lacinipolia laudabilis (Gn.)
252 Lascoria ambigualis Wlk.
253 Ledaea perditalis (Wlk.)
254 Lesmone detrahens (Wlk.)
255 Lesmone hinna (Gey.)
256 Leucania scirpicola Gn.
257 Lithophane looks like innominata
258 Lithophane viridipallens Grt.
259 Lithophane sp.
260 Litoprosopus futilis (Grt. & Rob
261 Marathyssa basal is Wlk.
262 Marathyssa inficita (Wlk. )
263 Meganola minuscula (Zell . )
264 Mel i pot is jucunda Hbn.
265 Meropleon cosmion Oyar
266 Metalectra quadrisignata (Wlk.)
267 Metalectra sp.
268 Metria amelia (Gn.)
269 Mocis disseverans (Wlk.)
270 Mocis latipes (Gn.)
271 Mocis marcida (Gn.)
272 Mocis texana (Morr.)
273 Morrisonia confusa (Hbn.)
U so-
il - . - M - - - - -
U ON-
U - F M -
U - - - S
u - - - o - -
U - - - A — - - S — -
* C
U - - M - - -- -- -- -
y - - - A
U - F M J A — - -
0 A A S - N -
U AM
0 --M-M-------
U - - M - - —
0 J - M - - - -
D - - - A
C -M-------
U J ---------- -
U ------ - A - - --
0 J F M A 0 - -
U - F --------- -
0 - F M A
U - - - A
U
U - F - A - -
(Smith) U J -
U J - - - -
U - F - - -
) U - - - - - J ----- -
0 J - M - - -
0 --MAM-
0 - F - - M 0 - -
0 ---AM
U - - - -NO
U - - M - - -- -- -- -
U ----M-------
U - - M A
0 - - 0 - -
c - - 0 - -
U - F --------- -
u --------- o - -
C - - M A ------- -
28(1-2):1-136, 1989(90)
55
Table 1 (Continued)
Abundance Month
274 Mop pi sonia mucens (Hbn.) A -FMA
275 Nigetia formosalis Wlk. C -FMAM---S0--
276 Nola sorghiella Riley U A S
277 Ogdoconta cinereola (Gn.) U - - M
278 Oligia fracti Tinea (Grt . ) U ---------o--
279 Ophiuche minual is (Gn.) U ----- ■ - N -
280 Oruza albocostaliata (Pack.) 0 ---A-JJ-----
281 Oxycilla prob. mi tographa (Grt.) U S
282 Paectes abrostoloides (Gn.) U S - - -
283 Pal this angulal is (Hbn.) U
284 Pal this asopialis (Gn.) U J F ~ - M --D
285 Pangrapta decoral is Hbn. 0 - - M A M
286 Panopoda repanda (Wlk.) U - - M
287 Panopoda rufimargo (Hbn.) U
288 Panthea furcilla (Pack.) C J-M-------ND
289 Parallel ia bistriaris Hbn. U ----M---S---
290 Phalaenostola larentioides Grt. U ---a--
291 Phoberia atomaris Hbn. 0 - - M
292 Phosphila miseloides (Gn.) U -FM-------N-
293 Phosphila turbulenta Hbn. 0 -F------S0--
294 Phyprosopus callitrichoides Grt. U -F--M-------
295 Phytometra rhodarialis (Wlk.) U - ~ M - ~ J -
296 Plathypena scabra (F.) U - F
297 Platysenta mobi 1 is (Wlk.) U - - D
298 Platysenta sutor (Gn.) 0 - F - - - J NO
299 Platysenta videns (Gn.) U - - M
300 Polygrammate hebraeicum Hbn. 0 - - - - A
301 Prorobl emma testa B.& McD. 0 A - J NO
302 Psaphidia resumens Wlk. U J F -
303 Pseudanthracia coracias (Gn.) U - F - - -
304 Pseudoplusia includens (Wlk.) 0 J ----- 0-0
305 Ptichodis herbarum (Gn.) U ----M-J-S---
306 Ptichodis vinculum (Gn.) U ---AM
307 Redectis vitrea (Grt.) U A -
308 Renia salusalis (Wlk.) U -FM
309 Schinia bina (Gn.) U S
310 Schinia gaurae (J.E. Smith) U - M -
311 Schinia nubila (Stkr.) U S
312 Schinia nundina (Orury) U - S
313 Schinia rivulosa (Gn.) U S
314 Schinia saturata (Grt.) C - - A S 0 - -
315 Schinia scissoides (8enj.) U ---------o--
316 Schinia siren (Stkr.) U - - -■ S
56
J. Res. Lepid.
Table 1 (Continued)
Abundance Month
317 Schinia trifascia Hbn.
0
SON"
318 Schinia tuberculum (Hbn.)
U
-------- - 0 - ~
319 Scolecocampa liburna (Gey.)
U
- - M - M ------ -
320 Selenisa sueroides (Gn.)
0
0 - D
321 Sigela prob. eoides (B.& McD.)
U
- F - D
322 Spodoptera eridania (Cram.)
U
----- j - - - o - -
323 Spodoptera latifascia (Wlk.)
0
M ND
324 Spragueia onagrus (Gn.)
c
---A-JJ-S---
325 Tarachidia candefacta (Hbn.)
0
— M AS---
326 Tarachidia semi f lava (Gn.)
u
MJ — S
327 Thioptera nigrofimbria (Gn.)
u
— MA S
328 Trichoclea vindemialis (Gn.)
u
- - M
329 Xystopeplus rufago (Hbn.)
u
- F -
330 Zale aeruginosa (Gn.)
c
J F M - M J ----- -
331 Zale buchholzi McD.
c
J F M - - N -
332 Zale declarans (Wlk.)
c
- F M A - - - -
333 Zale horrida Hbn.
u
- - M - -
334 Zale lunata (Drury)
0
- - - - M J -■- -
335 Zale lunifera (Hbn.)
0
- F M - - - - -
336 Zanclognantha minora! is Sm.
u
- F - - - -
337 ? Cyathissa n. sp.
c
-FMA---AS0N-
NOTODONTIDAE
338 Dasylophia anguina (J.E. Smith)
0
-FMA S
339 Datana angusii G.& R.
0
— s —
340 Datana integerrima 6. & R.
u
----M----
341 Datana major G.& R.
0
-----jjAS---
342 Datana modesta Beutenmuller
u
no date
343 Datana near ranaeceps (Guer.-Meneville)
u
------- A - - - -
344 Datana robusta Stkr.
u
- A - *
345 Furcula cinerea (Wlk.)
u
- - - - s - - -■
346 Heterocampa astarte Doubleday
u
- - M A A - - - -
347 Heterocampa biundata Wlk.
0
-F-A 0 N -
348 Heterocampa umbrata Wlk.
0
- - M A - - D
349 Heterocampa varia Wlk.
0
A AS
350 Hyparpax perophoroides (Stkr.)
u
-F-A
351 Hyperaeschra georgica (H.-S.)
u
— MA A
352 Lochmaeus bilineata (Pack.)
u
- F - - - -
353 Lochmaeus manteo Doubleday
D
-J--"--
354 Macrurocampa marthesia (Cram.)
0
- - S 0 N D
355 Nadata gibbosa (J.E. Smith)
c
-FM--JJ-S
356 Oligocentria lignicolor (Wlk.)
u
-----S---
357 Per idea angulosa (J.E. Smith)
u
j --------- N -
358 Schizura i pomoeae Doubleday
u
-"-----"SO--
28(1-2):1-136, 1989(90)
57
Table 1 (Continued)
Abundance Month
359 Schizura unicornis (J.E. Smith) 0 - - M A - - N -
360 Symmerista albifrons (J.E. Smith) 0 J F - - -
0EC0PH0RI DAE
361 Antaeotricha leuci liana (Zell.) A JFM„ A-----0-D
362 Antaeotricha osseella (Wlsm.) U 0 - -
363 Antaeotricha vestal is (Zell.) C M--AS---
364 Cal lima nathrax Hodges 0 - S
365 Decantha boreasella (Cham.) 0 - F
366 Inga sparsiciliella (Clem.) C A--J - -
PLUTELLIDAE
367 Plutella xylostella (L.) 0 - - ~ - 0 - -
PSYCH I DAE
368 Cryptothelea gloverii (Pack.) 0 M 0 - -
369 Thyridopteryx ephemeraeformis (Haw.) C ---J
PTER0PH0RIDAE
370 Geina ? periscelidactyla (Fitch) U - - - D
371 Oidaemotophorus balanotes (Meyr.) A - - M - - S 0 N 0
372 Stenoptilia parva Wlsm. U N -
PYRALIDAE
373 Acrobasis grossbecki (B.& McD.) U M -
374 Adel phia petrel la (Zell.) 0 - F M
375 Aglossa cuprina Zell. U A -
376 Anageshna primordial is (Dyar) 0 - - M
377 Apogeshna stenialis (Gn.) U --.M---J-S
378 Argyria lacteella (F.) 0 J - - 0 - -
379 Arta sp. U 0 - -
380 At he 1 oca subrufella (Hulst) U J - -
381 Basacallis tarachodes Dyar 0 J - M - - - - - D
382 81epharomastix ranalis (Gn.) 0 ---AM----
383 Chrysendeton imitabilis (Dyar) U A
384 Clydonopteron tecomae Riley U 0
385 Conchylodes concinnalis Hamp. 0 JAS
386 Crambus praefectellus (Zinck.) U AM
387 Crambus quinquareatus Zell. U - -M - - 5 - - -
388 Crambus sanfordellus Klots 0 --M 0
389 Crambus satrapellus (Zinck.) C JFMAM 0 - D
390 Desmia funeral is (Hbn.) 0 - F M - M - -
391 Diacme ? adipaloides (G.& R.) U JF - N -
392 Diasemiopsis leodocusalis (Wlk.) U 0 - -
393 Diatraea lisetta (Dyar) U 0 - -
394 Dicymolomia julianalis (Wlk.) 0 - - M A -----
395 Dioryctria abietivorella (Grt.) U - - M
396 Dioryctria amatella (Hulst) C AM----0N-
58
J. Res. Lepid.
Table 1 (Continued)
Abundance Month
397 Dioryctria clarioralis (Wlk.) 0 --MAM-J-----
398 Oonacaula maxima] la (Fern.) 0 ---AM-
399 Donacaula prob. melinella (Clem.) U - - S
400 Oonacaula nitidella (Oyar) U --MA--J-
401 Donacaula roscidella (Dyar) U -M S 0 - -
402 Donacaula sordidel la (Zinck . ) U ---A--------
403 Elasmopalpus lignosellus (Zell.) U ------- - --nd
404 Eoparargyractis irroratalis (Dyar) C ---A ON-
405 Epipagis huronalis (Gn.) U - - ON-
406 Epipaschia superatalis Clem. U ----M-------
407 Eudonia strigalis (Dyar) 0 J F - - N -
408 Eustixia pupula Hbn. U A
409 Fissicrambus ?hemiochrellus (Zell.) 0 J--A-----0N-
410 Fissicrambus mutabi 1 is (Clem.) U 0 - -
411 Galasa nigrinodis (Zell.) U ---A
412 Glaphyria basif laval is 8.& McD. U - - M -
413 Glaphyria fulminalis (Led.) U ---A---A----
414 Glaphyria glaphyral is (Gn.) U AM
415 Glaphyria sesquistrialis Hbn. U - * M - -
416 Glyphodes sibillalis Wlk. U - S 0 - -
417 ? Hahncappsia mancalis (Led.) U - M - - -
418 Hellula rogatalis (Hulst) U ~ F ~ A
419 Herculia binodulalis (Zell.) 0 - - - A - - ■ - N -
420 Herculia sordidal is 8.& McD. U - - - • N -
421 Hydriris ornatalis (Dup.) U --N-
422 Hymen i a perspectalis (Hbn.) U - - - - ■ NO
423 Jocara incrustalis (Hulst) 0 JF--M--AS
424 Laetilia coccidivora (J .H. Comstock) U -F - 0
425 Lepidomys irrenosa Gn. C --MAM--AS---
426 Lineodes fontella Wlsm. U - • J - - S
427 Lipocosmodes fuliginosalis (Fern.) U • - - - - S
428 Marasmia cochrusalis (Wlk.) U -F -------
429 Melitara prodenial is Wlk. LJ ---A 0--
430 Mesolia incertella (Zinck.) U 0 - -
431 Microcausta f lavi punctal is 8.+McD. 0 -FM
432 Microcrambus biguttel lus (Fbs.) U no date
433 Microcrambus elegans (Clem.) C --HAH
434 Microtheoris ophionalis (Wlk.) U - - ■ j ----- -
435 Moodna ostrinella (Clem.) C JFMA - D
436 Munroessa gyralis (Hulst) U -------A-0--
437 Munroessa icciusalis (Wlk.) U N -
438 Munroessa nebulosalis (Fern.) U ---------o--
439 Neargyractis slossonalis (Dyar) U - - - - - - N -
28(1-2):1-136, 1989(90)
59
Table 1 (Continued)
Abundance Month
440 Nomophila nearctica Mun. U • S
441 Oenobotys vinotinctalis (Hamp.) U - - M - - •
442 Omphalocera munroei Martin U M 5 - ~ ~
443 Palpita magniferalis (Wlk.) U J--A---A-0--
444 Palpita cincinnatalis Mun. U - F M
445 Parachma ochracealis Wlk. 0 - - ‘ A - - - A S
445 Parapediasia decorella (Zinck.) U A - 0 - -
447 Paraponyx all ioneal is Wlk. C -F-A-J-----D
448 Paraponyx obscuralis (Grt.) U - ~ ~ ~ D
449 Peoria approximella (Wlk.) U - S
450 Phycitinae (sp.?) U 0 - -
451 Pleuroptya penumbralis (Grt.) U - ~ N ~
452 Prionapteryx achatina Zell . U J 0 - -
453 Prionapteryx serpentella (Kft.) U J ----- -
454 Pyrausta tyralis (Gn.) 0 -F A - ON ~
455 Raphiptera argillaceella mimimel la (Rob.) 0 - f M A - - • - N -
456 Salebriaria fructetella (Hulst) U - M
457 Samea ecclesialis Gn. C J .-NO
458 Samea mult i pi ical is (Gn.) A - - N D
459 Scirpophaga perstrialis (Hbn.) U - - S
460 Synclita obi iteral is (Wlk.) 0 • - - 0 - D
461 Synclita tinealis Mun. U
462 Tampa dimediatel 1 a Rag. U -A--------
463 Tetralopha melanogrammos Zell. 0 -FMAM--A
464 Tetralopha robustella Zell. 0 AMJ-A-0--
465 Tetralopha scortealis (Led.) 0 --MA
466 Thaumatopsis edonis (Grt.) 0 ----- - Q N -
467 Tulsa finitella (Wlk.) U J
468 Ufa rubedinella (Zell .) C - - M - - - OND
469 Uresiphita reversalis (Gn.) U - - 0 - -
470 Urola nivalis (Drury) C JFMA ■ - - - N -
471 Xanthophysa psychial is (Hulst) U '--J
472 Xubida linearella (Zell.) 0 --MA-J------
SATURNIIDAE
473 Act las luna (L.) 0 --M----A----
474 Anisota consularis Dyar U -----A----
475 Anisota virginiensis pel lucida (J.E. Smith) A ~ - J" A 5
476 Antheraea polyphemus (Cram.) C 0 - -
477 Automeris io (F.) A - J-AS0-"
478 Citheronia regal is (F.) U - - A - - - -
479 Dryocampa rubicunda (F.) A M-JAS
480 Eacles imperial's Orury 0 --AS
481 Hemileuca maia (Drury) 0 J
60
J. Res. Lepid.
Table 1 (Continued)
Abundance Month
SCYTHRI 0IDA6
482 Scythris n. sp. U 0 - -
483 Scythris sp. U - 0
SESIIDAE
484 Carmenta texana (Hy.Edw.) U - - SO--
485 Synanthedon alleri ( Engel h.) U - 0 - -
488 Synanthedon exitiosa (Say) A S 0 - -
487 Synanthedon sapygaeformis (Wlk.) A - - - S0--
SPHINGI DAE
488 Ceratomia catalpae (Bdv.) U - - A -
489 Darapsa myron (Cram.) C --MA----S---
490 Deidamia inscripta (Harr.) C --MA--------
491 Oolba hyloeus (Orury) 0 ---A-J------
492 Enyo lugubris (L.) C - . - - - q - -
493 Eumorpha fasciatus (Sulz) U ----M-------
494 Laothoe jug land is (J.E. Smith) C - - M - ■ S - - -
495 Lapara coniferarum (J .E. Smith) 0 ---A-J--S---
496 Paonias excaecatus (J.E. Smith) U --AS
497 Xylophanes tersa (L.) 0 - - * • - - 0 - -
TINEI0A6
498 Acrolophus arcanella (Clem.) U - - - - - S
499 Acrolophus plumifrontella (Clem.) C -----j------
500 Acrolophus propinquus (Wlsm.) U J ~ - S
501 Acrolophus texanella (Cham.) U A -0
502 Acrolophus near variabi 1 is (Wlsm.) U -----J------
503 Acrolophus sp. 0 - - M
504 Acrolophus n. sp. II - - - - M
505 Nemapogon rileyi (Dietz) C - F - A - - - N -
506 Phereoeca walsinghami (Busk. ) C no date
507 Tinea apicimaculella Cham. U -F
508 Xylesthia pruniramiella Clem. U -F-A-----
509 sp. U - - M
510 sp. U N -
511 sp. U J - M - - - 0
TORTRICI DAE
512 Amorbia humerosana Clem. A J---0ND
513 Ancylis comptana (Frol ich) 0 - F - - M - - 0
514 Arc hips argyrospila (Wlk.) A ---AM
515 Archips georgiana (Wlk.) A ---AM * - -
516 Archips ? grisea (Rob.) U -.--A-----
517 Archips infumatana (Zell.) U ----M---
518 Archips semiferana (Wlk.) A ---AM
519 Argyrotaenia n. sp. 0 - F - - - --0
28(1-2):1-136, 1989(90)
61
Table 1 (Continued)
520
521
522
523
524
525
526
527
528
529
530
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
Abundance Month
Argyrotaenia ivana (Fern.) U
Argyrotaem’a quercifo liana (Fitch) C
Argyrotaenia tabu! ana Free. A
Cacocharis cymotoma (Meyr.) U
Chimoptesis pennsylvaniana (Kft.) A
Chimoptesis n. sp. 0
Choristoneura obsoletana (Wlk.) II
Choristoneura rosaceana (Harr.) A
Coelostathma discopunctana Clem. 0
Crocidosema plebejana Zell. U
Croesia semipurpurana (Kft.) A
Cydia ingens (Heinr.) U
Cydia n. sp. 0
Cydia n. sp. 0
Cydia n. sp. U
Ecdytolopha punctidiscanum (Oyar) U
Endopiza prob. liriodendrana (Kft.) 0
Endopiza spiraeifoliana (Heinr.) U
Endothenia hebesana (Wlk.) U
Epiblema scudderiana (Clem.) 0
Epiblema strenuana (Wlk.) U
? Epiblema sp. U
? Epinotia sp. U
Episimus argutanus (Clem.) U
Episimus tyrius Heinr. U
Eucosma adamantana (6n. ) C
Eucosma circulana Hbn. U
Eucosma cocana Kft. A
Eucosma gigantica (Riley) U
Eucosma guttalana 81anchard 0
Eucosma robinsonana (6rt.) A
? Eucosma n. sp. 0
Eumarozia malachitana (Zell.) 0
Gretchena bol liana (Slingerland) 0
Melissopus latiferreanus (Wlsm.) 0
?01ethreutes devotana Kft. ?
Olethreutes near hippocastana (Kft.) U
Petrova gemistrigulana (Kft.) C
Phaecasiophora niveiguttana (Grt.) 0
Phaneta ?argutipunctana 81anch. & Knudson 0
Phaneta raracana (Kft.) 0
Phaneta sp. U
? Phaneta sp. U
- F
---AM- - -
J F M A - - - - S 0 - 0
- - M ------- N -
JFM
- F M
- F - A M - -
-FM-
J F M -------- -
AH
A K -
- F
JFM------
- F
- - — M - 0 —
- F M
- - M - - - -
A - - - -
--MAMJ- -
--MA —
- F -
-FMAM - - - -
---------- N 0
J -
- F M A M - ~ - -
_ „ _ . j „ . _ _ _ „
----M--AS-N-
- - M A M - J
JFM -
-F- ---ON-
-F-AM---S — 0
A-ON-
- M
- - - A M ------ -
- - M A
s
--M - — - - S 0 - -
J - S - - 0
D
62
J. Res. Lepid.
Table 1 (Continued)
563 Platynota exasperatana (Zell.)
564 Platynota flavedana Clem.
565 Platynota idaeusalis (Wlk.)
566 Platynota rostrana (Wlk.)
567 Pseudexentera haracana (Kft.)
568 Pseudexentera spoliana (Clem.)
569 Pseudexentera sp.
570 Pseudogalleria inimicella (Zell.)
571 Ptycholoma peri tana (Clem.)
572 Rhopobota near finitimana Heinr.
573 Rhyacionia busckana Heinr.
574 Rhyacionia frustrana (Comstock)
575 Rhyacionia n. sp.
576 Sonia constrictana (Zell .)
577 Sonia sp.
578 Sparganothis caryae (Rob.)
579 Sparganothis n. sp.
580 Strepsicrates smithiana (Wlsm.)
581 Suleima sp.
582 ? Suleima sp.
583 Zomaria andromedana (B.& McD. )
584 Zomaria interruptolineana (Fern.)
585 Zomaria rosaochreana (Kft.)
586 n. sp.
587 sp.
YPONOMEUT I DAE
588 Atteva punctella (Cram.)
589 Urodus parvula (Hy.Edw.)
ZYGAENIDAE
590 Acoloithus falsarius Clem.
DANA I DAE
591 Danaus gilippus berenice (Cram.)
592 Danaus p. plexippus (L. ) Monarch
HESPERI I DAE
Abundance Month
C JFM - 0 - -
A JFH-AH SOND
U - - M - - ■
U - p -------- - -
U - F M -
U - F - - -
0 - F M - - - -
0 - - M A 0 - -
C - F M A - - N D
C - F M S - - -
C - F - - - N D
U - f -------- - -
U - F
C J - M A 0 N -
y ---------o--
U - - — MJ - -
0 no date
0 AM-J
U - F
A - F M
C -FM SON-
C -FM SO —
0 A-O-D
U - - - A
U --------S---
c M - — S
0 -FM---------
0 SO —
Queen
593 Calpodes ethlius (Stoll) Brazilian Skipper
594 Copaeodes minimus (Edw.) Southern Skipper ling
595 Epargyreus c. clarus (Cram.) Silver-spotted Skipper
596 Erynnis horatius (Scud. & 8urg.) Horaces Dusky-wing
597 Lerema accius (J.E. Smith) Clouded Skipper
598 Oligoria maculata (Edw.) Twin-spotted Skipper
599 Panoquina ocola (Edw.) Ocola Skipper
600 Polites v. vibex (Gey.) Whirlabout
601 Urbanus p. proteus (L.) Long-tailed Skipper
28(1-2):1-136, 1989(90)
63
Table 1 (Continued)
Abundance Month
PAPILIONIOAE
602 Battus p. philenor (L.) Pipevine Swallowtail
603 Eury tides marcel lus (Cram.) Zebra Swallowtail
604 Papilio c. cresphontes Cram. Giant Swallowtail
605 Papilio glaucus australis Maynard Tiger Swallowtail
606 Papilio palamedes Drury Palamedes Swallowtail
607 Papilio polyxenes asterius Stoll Black Swallowtail
608 Papilio t roil us ilioneus J.E. Smith Spicebush Swallowtail
PIERIDAE
609 Ascia monuste phi 1 eta (F.) Great Southern White
610 Eurema d. daira (Godt.) 8arred Sulpher
611 Eurema 1. lisa Bdv. & Leconte Little Sulpher
612 Eurema nicippe (Cram.) Sleepy Orange
613 Phoebis sennae eubule (L.) Cloudless Sulpher
614 Zerene c. cesonia (Stoll) Dogface Sulpher
LYCAENIDAE
615 Calycopis cecrops (F.) Red-banded Hairstreak
616 Euristrymon favonius (J.E. Smith) Southern Hairstreak
617 Hemiargus ceraunus antibubastus Hbn. Ceraunus Blue
618 Parrhasius m-album (8dv. & Leconte) White-m Hairstreak
619 Strymon m. melinus Hbn. Gray Hairstreak
NYMPHALIDAE
620 Agraulis vanillae nigrior Michener Gulf Fritillary
621 Anartia jatrophae guantanamo Mun. White Peacock
622 Asterocampa celtis (Bdv. & Leconte) Hackberry 8utterfly
623 Basilarchia archippus floridensis (Stkr.) Vicery
624 Basilarchia arthemis astyanax (F.) Red-spotted Purple
625 Heliconius chari torn* us tucker i Comstock Zebra
626 Junonia coenia (Hbn.) Buckeye
627 Phyciodes phaon (Edw. ) Phaon Cresent
628 Phyciodes t. tharos (Drury) Pearl Cresent
629 Vanessa atalanta rubria (Fruhstorfer) Red Admiral
630 Vanessa virginiensis (Drury) Am. Painted Lady
SATYRIDAE
631 Hermeuptychia sosybius (F.) Carolina Satyr
632 Meg is to cymela viola (Maynard) Little Wood Satyr
additions;
633 TINEIOAE: Acrolophus sp.
64
J. Res. Lepid.
In the following plates, the figure number is equivalent to the species list
number for each species in table 1.
28(1-2):1-136, 1989(90)
65
66
J. Res. Lepid.
'
271
-
|HMm|
A**
28(1-2):1-136, 1989(90)
67
68
J. Res. Lepid.
18§lf||§tS
1L "
I llfl| HBHI
! iggf g l|i leǤS
4
gilSii
W
429
•JHB
tjj8|jf
111111
vi':**- I
■
1
28(1-2):1-136, 1989(90)
69
70
J. Res. Lepid.
28(1-2):1-136, 1989(90)
71
72
J. Res. Lepid.
28(1-2):1-136, 1989(90)
73
74
J. Res. Lepid.
Journal of Research on the Lepidoptera
28(l-2):75-83, 1989(90)
New Records of Lepidoptera for New York and New
Hampshire (Nymphalidae, Noctuidae)
Tim L. McCabe
Biological Survey, State Education Department, New York State Museum, Albany, New
York 12230
Abstract. Recent collecting in the Northeast has added the following
Lepidoptera species to our regional lists: Aglais urticae (Linneaus)
[Nymphalidael, Syngrapha abstrusa Eichlin & Cunningham,
Syngrapha montana Packard, Syngrapha microgamma (Hubner),
Autographa rubida Ottolengui, Papestra quadrata (Smith), Anarta
cordigera (Thunberg), Pachypolia atricornis Grote, Gabara sub-
nivosella Walker, Xestia atrata (Morrison), Anomogyna rhaetica
(Staudinger), Anomogyna fabulosa Ferguson, Apamea commoda
(Walker), Oligia obtusa (Smith), Eutricopis nexilis Morrison, Bagisara
rectilinea (Grote), Sympistis heliophila (Paykull), Sympistis funesta
(Paykull), Macrochilo hypocritalis Ferguson [all Noctuidae]. Dates,
localities and life history notes are given and the species are illustrated.
Chamaedaphne calyculata (L.) Moench [Ericaceae] is reported as a host
for Syngrapha microgamma .
Introduction
Several summers spent rearing Lepidoptera in Albany’s Pine Bush
and in the Adirondack Mountains of New York, supplemented by
several trips to Mount Washington in New Hampshire, has resulted in
the discovery of several species that appear to represent new distribu-
tion records for the region. Some appear to be new to the continental
United States or even North America. Detailed locality information is
given in the legend below the plates. These records may not represent
the first individuals ever collected at these localities, but are the first
published report of which I am aware and represent noteworthy range
extensions. Several papers appeared on the White Mountain Lepi-
doptera in the first issue of the journal Psyche. Of particular note is a
paper by Morrison (1875) that lists Noctuidae. Forbes (1954) gives a
recent synopsis of the Lepidoptera fauna for our region and makes many
references to the higher elevations. The new records are as follows.
Nymphalidae
Aglais urticae (Linneaus). [Fig. 1]. A colleague, Charles Sheviak,
discovered this butterfly flying on the grounds of the New York State
Museum in downtown Albany, directed me to the spot, and I netted it
after watching it frequent bare spots of ground where it would bask.
76
J. Res. Lepid.
Emmons (1854) reported this as having been collected in the vicinity of
Albany in 1853. However, no North American specimens were found in
the New York State Museum’s collection and Emmons’ work was not
well received. Emmons was a geologist and criticisms of his work
included this comment by Schwarz (1892): “There are several instances
on record where useless books have been printed at the public expense,
but there has never been a more striking illustration of waste of money.”
In correspondence, Asa Fitch wrote, “Like his volume on Fruits, this on
Insects, I think, must fall still born from the press” (Barnes, 1984).
Possibly as a consequence of the lack of authority, catalogors have
ignored Emmons report of Aglais urticae . Indeed, Emmons may well
have meant to illustrate Nymphalis milberti inasmuch as no mention is
made of that species’ occurrence in New York. Also, specimens of A.
urticae from Europe were apparently available to Emmons. All this
notwithstanding, the present findings lends credence to the 1853
records.
The butterfly’s occurrence, by any means, is not natural. Albany is an
important port of commerce. The butterfly is univoltine at this latitude
and the adult overwinters. It should be looked for very late and very
early in the season to determine if it has become established. The
example figured is clearly the nominate race and is found throughout
western Europe, across Russia and Asia, and east to the Pacific coast of
the Palearctic region.
Noctuidae
Syngrapha abstrusa Eichlin & Cunningham. [Fig. 2]. A distribution
map is provided in the original description (1978). Its discovery in the
Adirondacks was not unexpected.
Syngrapha montana Packard. [Fig. 3]. Described from Mt. Washing-
✓ ton, I have taken a specimen at ultraviolet light at Lake Tear well below
tree line on Mt. Marcy. In addition, I have taken a specimen at flowers in
the daytime on Mt. Washington. It is decidedly rare in the East.
Ferguson (1955) cites additional records.
Autographa rubida Ottolengui. [Fig. 4]. This boreal species was taken
in the Adirondacks on Apocynum blossoms at night. Eichlin and
Cunningham (1978) show a distribution dot for the Adirondacks, but I
have been unable to locate the specimen.
Syngrapha microgamma (Hubner). [Fig. 5]. Ferguson (1955) has
described the North American population as race nearctica. I swept two
mature larvae (Fig. 20) from Chamaedaphne calyculata (L.) Moench
[Ericaceae] on a bog near Raquette Lake in the Adirondacks. One was
reared to adult and I have several light-trapped specimens from the same
bog as well as from Bloomingdale bog in Franklin County, New York.
Papestra quadrata (Smith). [Fig. 6]. McCabe (1980) gives a distribu-
tion map of this species. One specimen was taken above tree line on Mt.
Washington in New Hampshire at ultraviolet light.
28(1-2): 1-136, 1989(90)
77
Anarta cordigera (Thunberg). [Fig. 7]. This day-flying, bog-inhabiting
species has been recorded from nearby Hawley bog in Massachusetts
and is much more general to our north. 1 have it from three different
bogs near the Browns Tract Ponds in the Adirondacks (all Hamilton
County).
Pachypolia atricornis Grote. [Fig. 8]. This moth occurs later in the
season than when most people collect. I have taken it in late September
in the Adirondacks. I recently identified one for John Glaser which he
had collected on Warrion Mt., Allegany Co., Maryland, on November
3rd, 1987, so it may prove to be much more widespread in the East than
formerly recognized.
Gabara subnivosella Walker. [Fig. 9]. I have taken several specimens
in Albany's Pine Bush. They represent a coastal form known as bipuncta
(Morrison). According to Richards (1942) this form is most common in
salt marshes, but also occurs in some inland marshes and he reported in
from Long Island. I have it from dry, sandy barrens in Albany's Pine
Bush.
Xesiia atrata (Morrison). [Fig. 10]. Found just above tree line and in
the krummholz on Whiteface Mt. It is known from Mt. Washington in
New Hampshire and from numerous Canadian localities. Lafontaine, et
al., (1987) give a distribution map. The example illustrated represents
the nominate race.
Anomogyna rhaetica (Staudinger). [Fig. 11]. This species is the
sincera mentioned by Forbes (1954) as being from Glens Falls. The
Glens Falls locality seems suspect as this is a krummholz species and I
have recorded it from Whiteface Mt. in the Adirondacks. The moth has
also been associated with the name Anomogyna homogena conditoides
Benjamin (see Ferguson, 1965).
Anomogyna fabulosa Ferguson. [Fig. 12]. This recently described
species has been recorded from the White Mountains of New Hampshire
and in Canada. I took it in the krummholz on Whiteface Mt. and also at
Lake Tear on Mt. Marcy.
Apamea commoda (Walker). [Fig. 13]. Walker's type locality is not
known for certain and Forbes (1954) suggests Trenton Falls, N.Y. Three
specimens were taken in the krummholz on Whiteface Mt.
Oligia obtusa (Smith). [Fig. 14]. This moth may be utilizing roadside
Rumex. It has been recorded from Albany’s Pine Bush. I kept the
solitary female specimen alive in the hopes of obtaining eggs (un-
successfully) as a consequence the specimen figured has become very
rubbed.
Eutricopis nexilis Morrison. [Fig. 15]. I took adults, which are diurnal,
and (later in the season) larvae (Fig. 22) on the blossoms of Antennaria
■canadensis Greene in the Adirondacks. Hardwick (1970) describes the
biology of the species.
Ragisara rectifascia (Grote). [Fig. 16]. This has been collected on
Albany's Pine Bush and may be a new arrival. Increased use of
ornamental Malvaceae may account for its recent occurrence.
78
J. Res. Lepid.
Sympistis heliophila (Paykull) (= melaleuca (Thunberg)). [Fig. 17].
This species and the next occur above tree line and are day-flying and
extremely difficult to catch. I collected a larvae (Fig. 21) of this species
on Vaccinium uliginosum L., but it eventually died. A photograph of the
larva was identified. Adults were also taken on Mt. Washington. They
fly mid-morning on sunny days and can be taken when they bask on
rocks.
Sympistis funesta (Paykull). [Fig. 18]. Douglas Ferguson [pers. comm.)
has also recorded this moth from Mt. Washington. Its adult habits are
similar to the former species.
Macrochilo hypocritalis Ferguson. [Fig. 19]. Members of this genus
are found in wet places. The Black Creek, Albany Co., N.Y. locale has
provided many swamp species including four other Macrochilo species.
Acknowledgements. I thank Charles Sheviak for calling my attention to the
Aglais on the Museum’s grounds. Kauri Mikkola, acting on my behalf, for-
warded my photograph of the larva of Sympistis heliophila where it was kindly
identified by E.O. Peltonen. All specimens were collected by the author.
Christopher Supkis prepared the photographs. Douglas Wolfe of the Atmo-
spheric Sciences Research Center extended every courtesy towards my col-
lecting efforts on Whiteface Mt. Both James R. Jordan, Forest Supervisor, and,
more recently, Gary W. Carr, District Ranger, of the White Mountain National
Forest, granted permission for collecting moths on Mt. Washington. To all the
above I am most grateful. Contribution number 572 of the New York State
Science Service.
Literature Cited
BARNES J.K., 1984. The Membracidae and other Homoptera described by Asa
Fitch, 1851, and Ebenezer Emmons, 1855: Historical perspective and
analysis. N.Y. Entomol. Soc. 92:27-34.
EICHLIN, T.D. & H.B. CUNNINGHAM, 1978. The Plusiinae (Lepidoptera: Noctuidae)
of America North of Mexico, emphasizing genitalic and larval morphology.
U.S. Dept. Agr. Tech. Bull. 1567. 115pp.
EMMONS, E., 1854. Agriculture of New- York. vol. V. Insects of New- York.
Albany. 272pp., 47 pis.
FERGUSON, D C., 1955. A Nearctic race of Syngrapha microgamma Hubner, with
remarks on the status of montana Packard (Lepidoptera: Phalaenidae). Bull.
Brooklyn Entomol. Soc., 50(1):2327.
FERGUSON, D C., 1965. A new North American noctuid of the genus Anomogyna
(Insecta, Lepidoptera). Postilla 89: 111.
FORBES, W.T.M., 1954. Lepidoptera of New York and neighboring states. Pt. 3.
Cornell Univ. Agr. Expt. Sta. Bull. 329, 433pp.
HARDWICK, D.F., 1970. The life history of Eutricopis nexilis (Noctuidae). Journ.
Lep. Soc. 24: 151156.
LAFONTAINE, J.D., K. MIKKOLA & V.S. KONONENKO, 1987. A revision of the genus
Xestia subg. Pachnobia (Lepidoptera: Noctuidae), with descriptions of two
new subspecies. Entomologica Scandinavica 18:305331
28(1-2):1-136, 1989(90)
79
McCABE, T.L., 1980. A reclassification of the Polia complex for North America.
N.Y. State Mus. Bull. 432, 141 pp.
MORRISON, H.K., 1875. Notes on White Mountain Noctuidae. Psyche 1(10):4143.
RICHARDS, A.G., 1942. A revision of the species of Gabara of eastern United States
(Lepidoptera: Phalaenidae). Trans. Am. Entomol. Soc. 68:110.
SCHWARZ, E.A., 1892. North American Publications on Entomology. Proc.
Entomol. Soc. Wash. 2:523.
Fig. 1. Ag/ais urticae. Albany, Albany Co., N. Y., 19 October 1987, elev. 100
meters;
Fig. 2, Syngrapha abstrusa. South Inlet, Raquette Lake, Hamilton County,
N.Y., lat. 43.48.16 long. 74.36.30, 17 July 1977, elev. 555 meters;
Fig. 3. Syngrapha montana, Lake Tear, Essex Co., N.Y., lat. 44.06.25 long.
73.56.05, 10 July 1980, elev 1310 meters;
Fig. 4. Autographa rubida, 10 kilometers east of Indian Lake, Hamilton Co.,
N.Y., lat. 43.45.30 long. 74.10.14, 10 June 1977, 555 meters;
Fig. 5. Syngrapha microgamma, 10 kilometers east of Indian Lake,
Hamilton Co., N.Y., lat. 43.45.30 long. 74.10.14, 4 June 1977, 555
meters;
Fig. 6. Papestra quadrata, Mt. Washington, Coos County, N.H., lat. 44.16.13
long. 71.18.02, 11 July 1985, elev. 1856 meters;
Fig. 7. Anarta cordigera, Browns Tract Pond, Hamilton Co., N.Y., lat.
43.48.00 long. 74. 42. 17, 20 May 1980, elev. 555 meters;
Fig. 8. Pachypoiia atricornis, 10 kilometers east of Indian Lake, Hamilton
Co., N.Y., lat. 43.45.30 long. 74.10.14, 30 September 1980, 555
meters;
Fig. 9. Gabara subnivose/ia, Pine Bush, Albany Co., N.Y., lat. 42.42.19 long.
73.53.17, 9 July 1978, elev. 100 meters.
28(1-2):1-136, 1989(90)
81
Fig. 10. Xestia strata , Whiteface Mt., Essex Co., N.Y., lat. 44.22.58 long.
73.54.15, 6 July 1986, elev. 1065 meters;
Fig. 11. Anomogyna rhaetica, Whiteface Mt., Essex Co., N.Y., lat. 44.22.58
long. 73.54.15, 6 July 1986, elv. 1065 meters;
Fig. 12. Anomogyna fabulosa, Whiteface Mt., Essex Co., N.Y., lat. 44.22.58
long. 73.54.15, 6 July 1986, elev. 1065 meters;
Fig. 13. Apamea commoda, Whiteface Mt., Essex Co., N.Y., lat. 44.22.58 long.
73.54.15, 6 July 1986, elev. 1065 meters;
Fig. 14. O/igia obtusa, Pine Bush, Albany Co., N.Y., lat. 42.42.19 long.
73.53.17, 24 July 1987, elev. 100 meters;
Fig. 15. Eutricopis nexilis, 6 miles east of Indian Lake, Hamilton Co., N.Y., lat.
43.45.30 long. 74.10.14, 29 May 1980, 555 meters;
Fig. 16. Bagisara recti tinea, Pine Bush, Albany Co., N.Y., lat. 42.42.19 long.
73.53.17, 17 July 1986, elev. 100 meters;
Fig. 17. Symistis heliophi/a, Mt. Washington, Coos County, N.H., lat. 44.16.13
long 71.18.02, 17 July 1984, elev. 1856 meters;
Fig. 18. Sympistis funesta, Mt. Washington, Coos County, N.H., lat. 44.16.13
long 71.18.02, 17 July 1984, elev. 1856 meters;
Fig. 19. Macrochi to hypocritalis, Black Creek, Albany Co., N.Y., lat. 42.39.53
long. 74.58.01, 3 July 1984, elev. 100 meters.
82
J. Res. Lepid.
Fig. 20. Syngrapha microgamma, South Inlet, Raquette Lake, Hamilton
County, New York, lat. 43.48.16 long. 74.36.30, 17 May 1980, elev. 555
meters;
Fig. 21 . Sympistis heliophila, Mt. Washington, Alpine Gardens, Coos County,
New Hampshire, lat. 44.16.13 long. 71.18.02, 15 July 1981, elev. 1856
meters.
28(1-2):1-136, 1989(90)
83
Fig. 22. Eutricopis nexi/is, 10 kilometers east of Indian Lake, Hamilton
County, New York, 12 June 1980, lat. 43.45.30 long. 74.10.14, elev.
555 meters.
Journal of Research on the Lepidoptera
28(l-2):84-87, 1989(90)
Suppression of the black pigment in female hybrids
of Papilio glaucus and P. multicaudatus : further
evidence of the value of ecdysone in breaking pupal
diapause.
Sir Cyril A. Clarke
Department of Genetics and Microbiology, University of Liverpool, Liverpool L69 3BX,
England.
H. H. Rees
Department of Biochemistry, University of Liverpool, Liverpool L69 3BX, England.
David A. West
Department of Biology, Virginia Polytechnic Institute and State University,
Blacksburg, VA 24061, U.S.A.
Abstract. Using injections of ecdysone it was possible to break
indefinite pupal diapause in female hybrids between a black Papilio g.
glaucus female and a male P. multicaudatus. The insects demonstrate
well-marked suppression of the black glaucus pigment.
In previous papers (Clarke, Sheppard and Willig, 1972; Clarke and
Willig, 1977 and West and Clarke, 1988) we showed that the black
pigment of Papilio glaucus females could be suppressed in varying
degrees in hybrids using males of Papilio rutulus, Papilio glaucus
canadensis and Papilio eurymedon, and probably also with Papilio
multicaudatus. However, there was some doubt about the last (see
West and Clarke, 1988, page 197) as the two yellow females recorded
in a back-cross brood (female black glaucus X male FI glaucus X multi-
caudatus) were lost from our collection. The present paper remedies
this carelessness.
Materials and Methods
On 11. VII. 1987, using the hand-pairing technique (Clarke, 1952) we
crossed a black female P. glaucus (ex 1986 stock obtained from Pasade-
na, Maryland, U.S.A.) with a P. multicaudatus male (obtained from
pupae sent by David V. McCorkle and originating from Klickitat Co.
Washington). The caterpillars (brood 19052) were fed on Liriodendron ,
and produced 13 male butterflies by 1. IX. 1987. A further two male
pupae overwintered but only produced deformed insects in April, 1988,
i.e. 15 males in all.
We scored 13 of the pupae as being female but none of them eclosed.
28(1-2) 1-136, 1989(90)
85
Seven were therefore given cold/hot shock treatment (see footnote)
some by Mr. Karl Bailey and some by Mr. George Beccaloni, but this
was unsuccessful and by 30. III. 1989 we only had six (untreated)
pupae left alive. We therefore decided to try the use of ecdysone
preparations and one of us (HHR) carried out the injections. The six
pupae were divided into two groups of three, one batch being injected
with ecdysone and the other with 20-hydroxyecdysone, but the latter
was unsuccessful, all the pupae dying, possibly because the action of
the hormone was too rapid. To the other three pupae six ecdysone
injections, each of 170 ng, were given (as in Clarke and Willig, 1977)
on 3, 6, 8, 10, 12 and 14, IV. 1989.
Results
Three females emerged between 29. IV. 1989 and 2. V. 1989 and are
shown in Figure 1. It will be seen that only one (No. 1) grew fully, but
this is clearly primarily a yellow insect though with some “sooty”
features. (Black females normally always produce black daughters and
yellow females yellow ones.) The second hybrid (No. 2) failed to grow,
but it seems likely that had it done so it would have resembled No. 1.
No. 3 is more problematical — it too did not grow and is a good deal
blacker, but there is some yellow pigment in the hindwing. Suppression
of black therefore is probably variable.
We can therefore safely conclude that the male multicaudatus does
carry a suppressor of black, probably autosomally controlled (see West
and Clarke, 1988) though the expression is variable.
Discussion
It is easy to surmise that the presence of a suppressor would protect
the insect in areas where the model Battus philenor does not fly and a
black mutant would then be disadvantageous. However from an evo-
lutionary point of view these “anticipatory” theories are troublesome,
the rather lame explanation being that the “waiting” gene must have
been doing “something else”. The same problem occurs in Man with
certain drugs, for example isoniazid — where there is a precise dimor-
phism for the rates at which the drug is metabolised. However the gene
controlling this must have been present millions of years before isoni-
azid was synthesised.
Acknowledgements. We thank Mr. Karl Bailey and Mr. George Beccaloni for
carrying out the temperature experiments and are grateful to Professor B.
Surholt and Herr H. Brockhoff for useful discussions. We are indebted to the
Nuffield Foundation for its generous support.
Literature Cited
CLARKE, C. A., SHEPPARD, P. M. & WILLIG, A., 1972. The use of ecdysone to break a
80
J. Res . Lepid .
Fig. 1. FI hybrid females of the cross $ black P. giaucus Xc f P. multicauda -
tus (see text).
28(1-2):1-136, 1989(90)
87
two and a half year pupal diapause in Papilio glaucus female x Papilio
rutulus male hybrids. The Entomologist, 105 : 137-8
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
WEST, D.A., & CLARKE, SIR CYRIL A., 1988. Suppression of the black phenotype in
females of the P. glaucus group (Papilionidae). Journal of Research on the
Lepidoptera. 26: 187-200
CLARKE, C. A., 1952. Hand pairing of Papilio machaon in February. Entomo-
logist’s Record. 64: 98-100
Footnote
Details of Mr. Karl Bailey’s cold/hot shock therapy.
On 17. XI. 1987 the hybrid pupae were refrigerated at -4.5°C ± .5°C and left
for 8 weeks. They were then transferred to an incubator at 26°C in the hope
that the diapause would be broken. This did not occur and the pupae died.
Details of Mr. George Beccaloni’s cold/ hot shock therapy.
Pupae received on 8. II. 1989 and placed in a refrigerator at 0°C where they
were kept for three months. They were removed on 7. V. 1989 and subsequent-
ly kept at room temperature. They did not emerge and were given our standard
ecdysone treatment beginning on 9. VI. 1989. They were then kept at room
temperature but all died and no insects had formed up inside them.
Journal of Research on the Lepidoptera
28(l-2):88-96, 1989(90)
Studies on Spatial Distribution in the Teak
Carpenterworm Cossus cadambae Moore (Lepidoptera,
Cossidae)*
George Mathew1
P. Rugmini2
and
K. Jayaraman2
Kerala Forest Research Institute, Peechi 680 653 kerala, India
Abstract. Cossus cadambae Moore (Lepidoptera, Cossidae) is a rela-
tively new insect pest of teak in India and it has recently assumed
major pest status in several plantations in Kerala, Tamil Nadu and
Karnataka States. Caterpillars of this insect characteristically bore in
the wood of standing trees leading to deterioration of timber. C.
cadambae has annual generations with an exceptionally prolonged
larval stage. At Palappilly, Kerala, where the study was carried out,
the generations were continuous and overlapping. The progression of
infestation intensity among the trees was studied here. The intensity
of attack was studied by scoring the affected trees visually into
the following score classes viz., 0 = healthy tree; 1 = low level
infestation with a few scattered borer holes; 2 = medium level of
infestation with borer holes confined to small groups; 3 = heavy
infestation with numerous borer holes in large patches all over the
stem and 4 = tree dead as a result of heavy infestation. The results
show that during the initial phase of infestation a considerable number
of trees in a plantation get affected. The intensity of infestation during
this stage remains at a low level (score 1) and usually goes unnoticed
since the feeding scars are not often easily detectable. During the
subsequent phase of infestation there is a tendency for the already
affected trees to get reinfested, besides fresh attack to the unaffected
trees in the plantations. As a result there is slow transfer of trees of low
intensity score to high intensity classes (score 2, 3, 4) and the affected
trees generally occur in distinct patches. As the infestation progresses,
further recruitment of attacked trees to higher scores of infestation
intensity take place. This phase is characterised by large-scale mor-
tality of the heavily affected trees (score 4). As more and more healthy
trees get affected, the infestation becomes more or less uniformly
distributed throughout the plantation obliterating the original patchy
infestation.
*KFRI Scientific Paper No. 158
Entomology Division, 2Management Division
28(1-2):1-136, 1989(90)
89
Introduction
Teak ( Tectona grandis Lin.f.) is an important forest tree species
raised in extensive plantations in several parts of India, notably
in Madhya Pradesh, Andhra Pradesh, Karnataka, Tamil Nadu and
Kerala States. In Kerala alone, there are over 60,000 ha. of area planted
with this species.
About 140 species of insects are known to attack this tree in the
Indian sub region (Nair & Kumar, 1986) although only a few have been
reported to cause major damage in plantations. This includes 2 species
of foliage feeders viz., Hyblaea puera Cram. (Hyblaeidae) and Eutectona
machaeralis Wlk. (Pyraustidae) and a sapling borer Sahyadrassus
malabaricus Moore (Hepialidae). Recently, outbreaks of a wood boring
insect viz., Cossus cadambae (Cossidae) have been noticed in several
teak plantations in Kerala, Tamil Nadu and Karnataka States in India
(Mathew, 1988).
C. cadambae mostly attacks mature teak trees. The caterpillars of
this insect initially get established in the bark surrounding a wound
caused by mechanical injury or in the axillary region of branches (Fig.
1). Subsequently they bore into the sap wood and then into the heart-
wood. The larval stage lasts for about 8 months and the fully mature
larvae measure about 5 cm in size. One caterpillar makes only a single
borer hole although heavy infestation over several years can lead to the
formation of numerous holes on the wood.
In Kerala, infestation by C. cadambae was observed mostly in
plantations adjacent to human habitations. Trees growing in such areas
were frequently subjected to mechanical damages such as lopping of
branches, plucking of leaves etc. Usually, trees along the borders or
those standing along the sides of forest tracks were repeatedly subjected
to this type of damage. Such trees were often found to be heavily
attacked while those occurring in the interior or inaccessable areas
were not affected. Mechanical injury leads to the formation of callus
growth over wounds or profuse growth of coppices which offer conditions
favourable for the establishment of this insect. Beeson (1941) also
considered mechanical damage as a factor promoting cossid infestation.
Presently there is no information available on the establishment
pattern of this insect in plantations over space and time. Such informa-
tion is needed for development suitable management strategies and
therefore an attempt was made to study this aspect in selected planta-
tions of varying intensity of infestation.
Materials and Methods
Data for this study were collected from three borer-infested teak plantations
in Kerala, which were selected to represent three distinct phases in the
establishment of this insect. These phases include (i) an initial stage when the
90
J. Res . Lepid.
Fig. 1
Trunk of teak showing damage by C. cadambae
Fig. 2(a) Larva of C. cadambae ; 2(b) moth
affected trees show only low level infestation, with a few borer holes on the
trunk, (ii) an advanced stage of infestation characterised by the formation of
several holes on the stem and (iii) a later stage when the damage intensity has
substantially increased resulting in many holes on the wood, rendering it unfit
for any commercial use. Plantations having the above situations were selected
at Parambikulam, Thattakad and Palappilly respectively, based on a sampling
survey taking into account the number of affected trees as well as their damage
28(1-2):1-136, 1989(90)
91
intensity (Table 2). The intensity of attack was rated by scoring the affected
trees visually as given below.
0 — healthy tree
1 = low level of infestation ie., with few scattered borer holes
2 = medium level of infestation with several borer holes usually confined to
samll groups
3 = heavy infestation with numerous borer holes in large patches all over the
stem
4 = tree dead as a result of heavy infestation
In each of the plantations selected for study, a series of rectangular plots
(Table 1) of size 20 m x 8 m were taken linearly extending from one boundary to
the other. Each plot contained 34 trees depending on the terrian as well as the
extent of disturbance due to various factors like illicit felling, windfall etc. The
number of healthy and affected trees as well as the intensity of attack on each of
the affected trees was recorded. A negative binomial distribution was fitted to
the data on the number of trees affected per plot, in the Parambikulam and
Thattaked plantations. Similarly a binomial distribution was fitted to the data
on the number of trees affected in the Palappilly plantations. The probability
density function of the negative binomial is
where P(x) is the probability of x individuals of a given attribute in the
sampling unit
[i is the location parameter,
k is the dispersion parameter
and that of the binomial is
where P is the proportion of the population that shows the attribute q is (1 — p)
and n is the maximum number of individuals in the sampling unit.
The parameters in both the cases were estimated through the method of
maximum likelihood. The methods given in Bliss and Fisher (1953) were
followed in fitting the negative binomial distribution.
Results and Discussion
Data on the infestation status at the three localities studied herein
are given in Table 1. The percentage of affected trees was comparatively
low at Parambikulam (19.08%) and Thattaked (17.64%) as compared to
that of Palappilly (83.84%). Although the percentage of affected trees in
the first two localities were more or less the same, the intensity of
infestation in each of these plots was found to vary.
At Parambikulam all the affected trees belonged to a single intensity
score class (score 1) while at Thattakad the affected trees belonged to all
the four intensity classes. The data for the third locality Palappilly,
showed a marked increase in the percentage of affected trees (83.84%).
92
J. Res. Lepid.
Table 1 . Basicfeatures of the data gathered from 3 localities
Locality
Number
of plots
Total
No. of
trees
0
% of trees affected underthe
various score classes
1 2 3
4
Parambikulam
90
414
80.92
19.08
_
__
_
Thattakad
90
323
82.35
5.57
4.02
5.57
2.48
Palappilly
73
167
16.16
6.59
11.98
26.35
38.92
Table 2.
Estimates of the parameters for the fitted distributions
and goodness of fit
Locality
Variable
Binomial
distribution
P x2
Negative binomial
distribution
k X2
Parambikulam
V,
0.76
3.80 (ns)
Thattakad
v.
0.72
26.18 (**)
Palappiily
v.
0.46
7.79 (ns)
Parambikulam
v2
0.87
11.15
6.34 (ns)
Thattakad
v2
0.62
2.62
5.77 (ns)
Palappilly
v2
0.39
5.85 (ns)
V-i = Number of trees present per plot
V2 = Number of trees affected per plot
ns = non significant
** = significant at 1%
When the relative numbers of affected trees belonging to the various
score classes at Thattakad and Palappilly were examined, we found that
there was a transformation of the affected trees from the low intensity
score to the higher scores with a certain extent of mortality of trees.
That is, at Thattakad, out of 17.64% affected trees, only 5.57% belonged
to score 1 and the remaining trees belonged to the other score classes
(Table 1). Similarly at Palappilly there was a tendency for increase in
the number of trees belonging to the high score classes (score 2 =
1 1.98%, score 3 = 26.35%). The site was also characterised by a high rate
of tree mortality (score 4 = 38.92%) and thus only a small percentage of
trees belonged to score 1 (score 1 = 6.59%).
The infestations at each of the above localities were very characteristic
and illustrated the various phases in the establishment of this insect.
The situation observed for Parambikulam represents the initial phase,
when the trees show only minimum damage with few borer holes on the
trunk. The situation at Thattakad represents the second phase, when
the already affected trees were subjected to reinfestation in the sub-
sequent years when the damage became more pronounced often leading
to mortality of some trees. In the third phase as represented at
Probability Probability Probability
28(1-2):1-136, 1989(90)
93
Fig. 3a Para mbik^u lam
0-4
0.3
0.2
O.t
□ Observed probability
Binomial distribution
. — i^i
2 3 4
No. of trees present /plot
Fig ,4a Thattakkad
No. of trees present /plot
Fig. 3a, 4a, 5a Distribution of number of trees present/plot in Parambikulam,
Thattakad and Palappilly.
Probability Probability Probability
94
J. Res. Lepid.
Fig. 3b parambikulam
Fig. 5 b Palappi lly
0.4
0.3
0.2
0.1
0
I ] Observed probability
Binomial distribution
0 1 2 3 4 5
No- of trees affected/ plot
Fig. 3b, 4b, 5b Distribution of number of trees affected/plot in Parambikulam,
Thattakad and Palappilly.
28(1-2): 1-136, 1989(90)
95
Palappilly, there was a marked increase in the number of affected trees
belonging to higher intensity scores besides fresh attack on the un-
affected trees over years resulting in extensive riddling of the timber
and subsequent large scale tree mortality.
We studied the distribution of two variables, number of trees present
per plot and the number of trees affected per plot, in each of the three
localities. The test of independence between the two variables in
different plots showed them to be dependent in the case of Palappilly (x2
= 144.73) and independent in the cases of Thattakad (x2 = 5.81) and
Parambikulam (x2 = 11.86). Therefore it was necessary to study the
distribution of variables in the three localities separately. The esti-
mates of the parameters in the fitted distributions are given in Table 2.
It indicates that the variable, the number of trees affected per plot,
follows negative binomial distribution in the cases of Parambikulam
and Thattakad and binomial distribution in the case of Palappilly (Fig.
3b, 4b, 5b). That is, the affected trees occur in definite patches or clusters
in Parambikulam and Thattakad, while at Palappilly the affected trees
were uniformly distributed. The reason for the uniform distribution of
affected trees at Palappilly might be due to the patches of affected trees
becoming confluent with the progression in the infestation level over a
period of time. The distribution of the number of trees present per plot,
was found to be binomial in the case of Parambikulam and Palappilly
(Fig. 3a, 5a). However, in the case of Thattakad it was not so (Fig. 4a)
probably due to irregularities in the actual frequencies of trees per plot.
The time taken for transformation from one phase of infestation to the
other could not be arrived at but it seems that this change is a slow one
perhaps requiring several years due to the biology of this insect.
The study has shed some light on the distribution pattern of C.
cadambae in teak plantations in Kerala under varying levels of infesta-
tion status. During the initial phase of attack, the infestation usually
goes unnoticed since the feeding scars are usually not very prominent.
At this stage a considerable number of trees get attacked. The second
phase is characterised by further deterioration of the already affected
trees due to reinfestation in the subsequent years leading to slow
mortality of trees and by a slow spread of attack to the healthy trees in
plantations. During these two phases the affected trees are usually
confined to distinct patches. C. cadambae being highly mobile organ-
isms can fly to the other parts of the plantation or even to other
plantations in the vicinity, resulting in the spread of attack. During the
last phase the infestation spreads at a faster pace leading to a high
mortality of affected trees and a more or less uniform distribution of
attack throughout the plantation.
Due to the behavioural characteristics of this insect, specialised pest
management strategies need to be developed for its control in planta-
tions. Since its attack is more clustered in the initial phase, manage-
96 J. Res. Lepid.
ment operations need be confined to the affected patches only, rendering
control operations more economical.
Acknowledgement. We are extremely grateful to Dr. C.T.S. Nair, Director,
and Dr. K.S.S. Nair, Scientist in Charge, for their keen interest in this study. We
thank Mr. Unnithan (Professor, Department of Statistics, Kerala Agricultural
University, Vellanikkara) for useful suggestions and discussions during the
course of this study and Dr. Paul D. Syme, Forest Insect and Disease Survey
Unit, Great Lakes Forestry Centre, Ontario, Canada for critically reviewing
this paper. Thanks are also due to Mr. E.P.S. Nair for neatly typing out the
manuscript.
Literature Cited
BEESON, C.F.C., 1941. The ecology and control of forest insects of India and
neighbouring countries. (1961 Reprint), New Delhi, Government of India,
767 pp.
BLISS, C.l. and FISHER, R.A. 1953. Fitting the negative binomial distribution to
biological data and a note on the efficient fitting of the negative binomial.
Biometrics, 9: 176-200.
MATHEW, G. 1989. Biology and ecology of the teak trunk borer, Cossus cadambae
Moore and its possible control, KFRI Research Report (In Press).
NAIR, K.S.S. and SUDHEENDRA KUMAR, V.V., 1986. Population dynamics of teak
defoliators, Proc 18th IUFRO World Congress. Vol. 2: 673-684.
Journal of Research on the Lepidoptera
28(l-2):97-99, 1989(90)
A New Species of Argyrotaenia from Arizona
(Lepidoptera: Tortricidae)
J. F. Gates Clarke
Department of Entomology Smithsonian Institution Washington, D. C., 20560
Abstract. A new species of Argyrotaenia is described, figured and
compared with A. cockerellana (Kearfott) from which it differs by the
much darker fore wing and longer aedeagus.
Mr. Ronald W. Wielgus has been studing and contributing to the
knowledge of the lepidoptera of Arizona for many years. Among his
recent discoveries, near his home, is a species of Argyrotaenia for
which he needs a name, and which is described below. In addition to
the specimens submitted by Mr. Wielgus, there is a series from Texas,
which came to the National Collection in the gift of the late Andre
Blanchard. Dr. John G. Franclemont of Cornell University and Dr.
Ronald W. Hodges, U. S. Department of Agriculture, have given a
series, also from Arizona, which is listed with the paratypes.
Argyrotaenia wielgusi, Clarke, new species (Figure 1)
Alar expanse 24-28 mm.
Labial palpus russet; third segment brown. Antenna russet. Head,
thorax, and forewing ground color vinaceous-cinnamon; on basal half
of costa four short brown bars and on apical half of costa four brown
spots; on dorsal edge similarly colored but somewhat suffused mark-
ings; around termen four or five brown spots; from base to termen,
following fold, a slender, dark brown line, broken about middle and
overlain in part by dark brown blotches; from near base to near termen
of fore wing a series of irregular dark brown markings; near base of fold
a small white mark; on costal edge of cell two conspicuous, white
marks, the outer one split by a slender, transverse dark brown line;
cilia vinaceous-cinnamon. Hindwing whitish, suffused grayish to pale
gray, the veins clearly outlined darker gray; cilia sordid white, suffu-
sed grayish with a gray subbasal line. Foreleg tibia white on inner
surface, vinaceous-cinnamon on outer surface; tarsal segments brown
narrowly annulated buff distally; midleg similar; hindleg buff; tarsal
segments lightly suffused grayish. Abdomen gray with some buff
scales ventrally.
Male genitalia slides No. 28358, 26366, 26683, 26684. Harpe broad;
costal half cupshaped, cucullus bluntly pointed; sacculus thickened
and produced ventrally. Gnathos a long, slender, slightly curved pro-
cess. Uncus stout, curved, slightly dilated distally. Vinculum a narrow
98
J. Res. Lepid.
semicircle. Tegumen strongly sclerotized, longer than broad. Anellus
triangular, deeply cleft posteriorly. Aedeagus longer than tegumen,
slender, strongly curved; extension of the phallobase unusually develo-
ped.
Female genitalia slides No. 26359, 26388, 26389. Ostium very broad,
funnelshape. Antrum narrow, strongly sclerotized. Inception of ductus
seminalis from right side at junction of antrum and membranous
portion of ductus bursae. Ductus bursae membranous. Bursa copulat-
rix membranous with little or no ornamentation. Signum a long,
slender, hook.
A. wielgusi is very closely related to A. cockerellana (Kearfott), but is
a much larger and darker species. A. wielgusi measures 24-28 mm.,
and the ground color of the forewing is vinaceous-cinnamon; cockerel-
lana measures 14-22 mm., and the forewing ground color is light
cinnamon-buff. The length of the aedeagus of cockerellana is about
two-thirds the length of that of wielgusi and is not so strongly curved.
It gives me great pleasure to name this handsome moth for Mr.
Wielgus in recognition of the many contributions to the knowledge of
the lepidoptera of Arizona which he has made.
I thank Mr. Victor Kranz, Smithsonian staff, for the photographs of
the moth wings and genitalia. Also, I am indebted to Mrs. Nancy L.
McIntyre for typing the manuscript.
Holotype: United States National Museum of Natural History.
Type locality: Arizona, Cochise Co., Huachuca Mts., Pueblo del Sol.
Distribution: Arizona and Texas.
Food plant: Unknown.
Described from the 9 holotype from Arizona: Cochise Co., Huachuca
Mts., Pueblo del Sol (6. XI. 1986, R. S. Wielgus), one 9 paratype with
same data except 5. XI. 1985; one C? paratype, Cochise Co., Huachuca
Mts., Ash Canon (15. XI. 1985, Noel McFarland [found in water]); one
Cf paratype, Cochise Co., 5 mi. W. Portal, 5400’, 17. X. 1964, V. Roth;
3cfd\ 5J9 paratypes, Santa Cruz Co., Santa Rita Mts., Madera
Canon, 4880’, 22.X. to 7. XI. 1959, J. G. Franclemont; 499 paratypes,
same locality, 25-28.X.1959, R. W. Hodges. Texas: 7c?C? paratypes,
Davis Mts., Mt. Locke, 6700’, 21.X.1973; 2 C?C? paratypes, Jeff Davis
Co., Fort Davis, 23. X. 1973, all A. and E. Blanchard.
There are two cfcT specimens from Montana, considerably smaller
(20 mm.) than the Arizona and Texas specimens, that are not included
in the type series, but the genitalia of which are indistinguishable. The
Arizona specimens have a more violaceous tinge to the ground color,
but otherwise are inseparable.
/V
28( 1-2): 1-136, 1989(90)
99
Fig. 1. a, right wings; b, ventral view of male genitalia with aedeagus
removed; c, aedeagus; d, ventral view of female genitalia; e, en-
larged view of papillae anales and ostium; f, enlarged bursa coplatrix
and signurn.
Journal of Research on the Lepidoptera
28(1-2):100-104, 1989(90)
A new subspecies of Satyrium auretorum
(Lycaenidae) from the Santa Monica mountains of
southern California
John F. Emmel
26500 Rim Road
Hemet, CA 92344
and
Rudolf H.T. Mattoni
9620 Heather Road
Beverly Hills, CA 90210
Abstract. A new subspecies of hairstreak butterfly, Satyrium aure-
torum fumosum, is named to designate the differentiated endemic
populations of the species that are restricted to the western end of the
Santa Monica mountains in California.
Satyrium auretorum (Boisduval) is a widespread, but local, hair-
streak species found throughout the foothills and lower mountain
slopes of much of California. The nominotypical subspecies was de-
scribed from a single male (Boisduval 1852), probably taken in the
Feather River drainage in the northern Sierra Nevada foothills. For
many years the species was considered a great rarity and even Com-
stock (1927) was unable to illustrate it in color due to the lack of
specimens. Since then, however, collectors have taken it in numerous
locations across the coast ranges and the Sierra Nevada foothills.
In 1881 Henry Edwards described a southern California subspecies,
spadix (type locality, Tehachapi pass, California), which he charac-
terized by a lighter ventral surface and more extensively developed
fulvous scaling on the dorsal surface of females. This subspecies has
subsequently been more frequently collected than the nominotypical
one, a function of the greater concentration of collectors near its
habitat. From all available information, the species is restricted to
scrub oak chaparral and is wholly found within the California floral
province.
In 1973 Emmel and Emmel made brief reference to an undescribed
subspecies of S. auretorum , from the Santa Monica mountains of
southern California, which they characterized by a phenotype darker
than known from any other population. We now describe this distinct
segregate as follows:
28(1-2):1-136, 1989(90)
101
Fig. 1. The subspecies of Satyrium auretorum. Left half of each figure
dorsal, right half of each figure ventral, surface. Left column males,
right column females. Top row, S. auretorum auretorum, male, 4 mi.
N. Camptonville, Sierra Co. CA. 29 June 1964; female, Capell creek,
Napa Co., CA. 3 June 1966. Middle row S. auretorum spadix. NE
slope San Gabriel Mts., Los Angeles Co., CA. male 6 June 1974;
femaleB June 1974. S. auretorum fumosum, male, (holotype) Malibu
Lake, Los Angeles Co., CA. 6 June 1948; female (allotype) same
locality 16 June 1948.
102
J. Res. Lepid.
Fig. 2. Distribution map of Satyrium auretorum fumosum in the western
section of the Santa Monica mountains. The entire known distribu-
tion of the subspecies is shown.
Satyrium auretorum fumosum Emmel and Mattoni new
subspecies
MALE. Forewing length 12.5-13.5 mm, mean 13.1 mm (N—13).
Dorsal Surface. Forewing: Ground color dark brownish gray. Outer
margin with a thin dark brown border, diffused basad into ground
color. Fringe pale tannish gray. Androconial scales pale gray, standing
out in greater contrast against the ground color than in either aure-
torum or spadix. Hindwing: Ground color, outer margin and fringe as
in forewing. Anal area pale tannish gray. Tail black with white scaling
at tip.
Ventral Surface. Forewing: Ground color dull brown, darker than
the fulvous brown seen in nominotypical auretorum or spadix. Pale
gray overscaling present in post discal and submarginal areas, render-
ing these areas lighter than the basal half of the wing. Dark brown
rectangular macule at distal end of discal cell enlarged, approximately
two to three times as wide as seen in nominotypical auretorum or
spadix. Postmedian series of dark brown macules crescent-shaped,
enlarged over those seen in nominotypical auretorum (Usually about
twice the width), and edged distally with pale gray scaling. Submar-
ginal series of dark brown macules obsolescent, but more developed
than in spadix, in which they are usually absent. In nominotypical
auretorum the submarginal series are usually very well developed.
Outer margin edged with a thin brown line, fringe pale tan. Hindwing:
28(1-2): 1-136, 1989(90)
103
Ground color, macules, outer margin and fringe as forewing, except
that postmedian series of macules are ovoid to rhomboid in shape. Pale
orange “eyespot” mark in cell CUi CU2 less developed than nominoty-
pical auretorum , more prominent than in spadix , in which it is often
obsolescent.
FEMALE. Size: forewing length 13.5-14.5 mm, mean 13.9 mm
(N=6).
Dorsal surface. Fore wing: Ground color dark brownish gray with a
small area of dull fulvous scaling in the center of the wing. Fulvous
scaling markedly reduced in extent from that seen in both other
subspecies. In spadix the scaling often covers over one third of the wing
and has relatively discrete borders, in nominotypical auretorum the
scaling is usually extensive, but more diffused into the dark brown-
gray ground color. Outer margin and fringe as in male. Hindwing:
Ground color as forewing. Fulvous scaling absent, or present in small
diffuse patch in the posterior half of the submarginal area. In spadix
and nominotypical auretorum the fulvous scaling is usually present
and more extensively developed. Ventral surface. Forewing and hind-
wing: Ground color and marking as in male.
TYPES. Holotype male: California, Los Angles County, Malibu
Lake, 6 June, 1948, leg. Wm. T. Meyer. Allotype female: same data as
male except 13 June, 1948. Paratypes (12 males, 5 females): 2 males
same data as holotype, 1 male and 1 female same data except 16 June,
1948, 4 males same data except 13 June 1948, 2 males and 1 female
same data except 17 June, 1948; 3 males, Malibu, 31 May, 1950, leg.
E. R. Hulbirt; 1 female, Seminole Hill (Santa Monica mountains) 15
June, 1941, no leg; 1 female, No. of Hyw. 101, 1-1.5 mi. from Brent’s
Junction, 27 April 1989, leg. Robert Allen.
DEPOSITION OF TYPES. The type series except for the specimens
of Allen and Pasko are in the collection of the Natm ?1 History
Museum of Los Angeles county. The other paratypes will be placed in
the Smithsonian Institution.
ETYMOLOGY. The name fumosum is derived from the latin root for
smoke, in reference to the darkened, “smoky” appearance of this
subspecies in contrast to both S. a. auretorum and S. a. spadix. The
suggested common name for this butterfly is the Santa Monica Moun-
tains hairstreak as all information indicate it is an endemic restricted
to that range.
DISTRIBUTION AND PHENOLOGY. Satryium auretorum fumo-
sum is thus far known only from the northern slopes and plateau of the
western Santa Monica Mountains, where it presumably flies in a
single brood from late April to June. The known distribution is illu-
strated in figure 2. The eastern part of the mountains have been
intensively collected since the 1940’s, including the detailed records of
McFarland, without any evidence of the species. Scanty available
information suggests flight usually occurs in June. The April record of
the single female taken by Allen may reflect an adaptive response to
104
J. Res. Lepid.
the early spring hot spell of 1989. This specimen was taken in a valley
oak savannah at least one mile from potential foodplant, scrub oak,
Quercus dumosa, which is the known foodplant of the subspecies
spadix. The scrub oak is present in the other known localities of
fumosum and is its likely foodplant. Scott (1986) cites two other oaks,
Q. lobata and Q. wislizenii as hostplants.
A single female which is intermediate between fumosum and spadix
was taken by John Pasko at Wildwood Park, Thousand Oaks, Ventura
county, 5 June, 1980. The status of the population this specimen
represents is insufficiently known.
DIAGNOSIS AND DISCUSSION. This subspecies is the darkest of
the S. auretorum segregates and may represent an adaptive response
to a moist coastal climate. It is readily distinguished from both aure-
torum and spadix by the dark ground color both dorsally and ventrally
and by the more prominent series of ventral postmedian macules. A
sample of all three subspecies are illustrated in figure 1, which permits
comparison of the ventral shading and maculation character states
among these segregates.
A somewhat similar phenotype is known from the Santa Ana moun-
tains of Orange County. Three males specimens were examined, Sil-
verado Canyon, 4 and 9 June, 1981, leg. Bob Iwahashi, collection of
LACM. Several additional records from the same locality are given in
Orsak (1977). These specimens were not examined. Because interven-
ing habitat, the foothills of the San Gabriel mountains, are occupied by
spadix , it is unlikely that the Santa Ana populations are monophyletic
with fumosum even if they prove morphologically similar. Further
systematic work is called for to clarify the matter, since this is the only
known Santa Monica mountain endemic butterfly. Such research is
urgent because of the great rate of land conversion in the area of both
segregates and increasing fragmentation across their entire ranges.
The limited distribution of fumosum in a rapidly changing urban
area indicate the subspecies should be considered for listing as threate-
ned or endangered.
Acknowledgements . Loan of the specimens of Satyrium from the .Los Angeles
County Museum collection was provided by Julian Donahue. Sterling Mattoon
provided information on auretorum populations.
Literature Cited
BOISDUVAL, J. 1852. Ann. Soc. Ent. France. (2) 10:288.
COMSTOCK, J. A. 1927. Butterflies of California. Publ. by author. Los Angeles.
EDWARDS, H. 1881. Papilio, 1:53.
EMMEL, T. C. & J. F. EMMEL, 1973. The butterflies of Southern California. Nat.
Hist. Mus. Los Angeles. Sci. Publ. no. 26.
mattoni, R. 1990. Butterflies of greater Los Angeles. Lep. Res. Found.
Beverly Hills, CA.
ORSAK, L. 1977. The butterflies of Orange County, California. Univ. Calif.
Irvine, Center for Pathobiology, Misc, Public, no. 3.
SCOTT, J. A. 1986. he butterflies of North America. Stanford. Palo Alto.
Journal of Research on the Lepidoptera
28(1-2):105-111, 1989(90)
Potential host range of Spilosoma dalbergiae
(Moore) n. ssp. (Lepidoptera: Arctiidae) in India
S. N. Tiwari*
and
N. P. Kashyap
Department of Entomology — Apiculture, Himachal Pradesh Agricultural University, •
Palampur- 1 7 6062 INDIA
Abstract. The potential host range of Spilosoma dalbergiae (Moore)
n. ssp. was determined by measuring feeding damage to pieces cut
from leaves of 67 plant species and varieties, and was compared with
literature reports of host range of other Spilosoma species. These tests
indicated that dalbergiae is polyphagous, and has both horticultural
and agricultural pest potentialities.
Introduction
A number of species of Spilosoma are engaged in the defoliation of a
large variety of plant species of economic importance in North Amer-
ican, Afro-Asian, and European countries. In the oriental region, S.
ohliqua has been reported to cause serious damage to many crops
(Patel, 1944, Anonymous, 1969; Prasad and Premchand, 1980). The
taxa Spilosoma dalbergiae , S. todara and S. bifascia have been re-
garded as synonyms of obliqua (Lall, 1964). There are now reasons to
believe that ohliqua and dalbergiae are separate species (W. Thomas,
personal communication), and in some instances damage attributed to
obliqua might have been caused by dalbergiae.
The feeding behavior of larvae of S. dalbergiae n. ssp. is similar to
obliqua. During early instars (I and II) larvae of dalbergiae n. ssp. feed
gregariously on the leaf surface and do not move from plant to plant. In
their advanced instars they move from plant to plant and from field to
field, feeding on many plant species. The larvae may not restrict
themselves to a particular habitat in later instars. The damage is done
mainly by 3rd to 7th instar larvae, which skeletonize the plant.
There are several records of plants susceptibilities to S. obliqua , but
most are based on observations of feeding damage. Bhattacharya and
Rathore (1977) have published the host list of S. obliqua which con-
tains 94 actual or potential hosts. However, host lists based on visual
signs of damage provide no data concerning the influence of host plants
* Present address: Department of Entomology, College of Agriculture, G. B. Pant
University of Agriculture and Technology, Pantnagar-263145, INDIA.
106
J. Res. Lepid.
on the feeding behavior of insect (Ladd, 1987). Many researchers have
carried out feeding tests to obtain information on these aspects (Kogan
and Goeden, 1970; Bhattacharya and Rathore, 1977).
The information on the host range of species can be utilized in the
planning of the ecosystem for the establishment of an insect pest
management programme. In the majority of ecosystems, agricultural
and horticultural crops are surrounded by large tracts of uncultivated
land. These areas harbor many plants which constitute a vital part of
ecological niche of both pests and beneficial insects. In addition to
providing protected sites during adverse conditions, such areas provide
food to the insects when the fields are crop-free (Price and Waldbauer,
1982).
The purpose of this study was to determine the range of potential
hosts of S. dalbergiae n. ssp. so that feeding behavior and pest poten-
tiality could be quantified and susceptibility of ecosystem to it determi-
ned.
Materials and Methods
Taxonomy : The adults of S. dalbergiae and S. obliqua have been
described by Hampson (1894) and Ahmad and Ahmad (1976) respec-
tively. A complete description of the adults of this new subspecies will
be published by Dr. W. Thomas (West Germany). Nominate S. dalber-
giae is distributed in Kangra, Sikhim, Kasis and Nagas (Hampson,
1894), this new subspecies has been observed in Kangra and Pant-
nagar. *We provide here enough information to distinguish the imma-
ture stages of the two species.
The egg clusters of dalbergiae n. ssp. are not as compact as those of
obliqua. The newly-hatched larvae of dalbergiae n. ssp. are yellowish,
and the body is covered with tiny hair. A pink ring on the first
abdominal segment, and pink spots on the last three segments, appear
on the 2nd day. A dark black band on the first abdominal segment is
apparent in the 2nd and 3rd instars, on a ground color of filthy yellow.
The 4th, 5th and 6th instars are a dark and dirty yellow and possess a
black band on the 1st abdominal segment. The dorsal part of the
posterior segment is also black. The color of the 7th instar becomes
dark and rusty.
In case of S. obliqua , the color of 1st instar is pale yellow and there
are no pink bands or spots on the body. The second instar larva
becomes yellowish with a greenish tinge, and there is no black band on
the 1st abdominal segment. The color further darkens through the
fourth instar. The thoracic segments and the three posterior abdomi-
nal segments develop a blackish tinge in the 5th instar, which remains
until the last instar (Goel and Arun Kumar, 1983). The black band on
the 1st abdominal segment of S. dalbergiae is not found in S. obliqua at
any stage.
28(1-2):1-136, 1989(90)
107
Culture: Adults were collected in a light trap. Paired insects were
held for egg laying in plastic jars (diameter 9 cm; height 10.5 cm), the
walls of which were lined with white paper. Adults were provided with
a 10 percent sucrose solution soaked in cotton as food. The eggs
obtained were kept in petridishes for hatching, and newly emerged
larvae were reared under laboratory conditions (temperature: mean T
min = 17.3°C; mean T max = 26.3°C; relative-humidity: mean RH
min = 58.1% mean RH max = 93.7%) in glass jars (22 X 30 cm). The
larvae were reared to fourth or fifth instar on the leaves of cowpea
(' Vigna unguiculata (L.) Walp.), which is known to be a good medium
for rearing S. obliqua.
Host range test: The experiment was conducted in glass petridishes
(12.25 cm diameter), the bottoms of which were filled with a 1 cm thick
layer of wax. The surface of the wax was covered with blotting paper.
Sixty seven cultivated or wild plant species were selected from various
habitats (forest, horticultural, and agro-ecosystem) in Palampur
(Himachal Pradesh). Each host was tested simultaneously in two
petridishes. Four 1 cm square pieces were cut from the leaves of test
plants and were fixed equidistant in the perimeter of each petridish
with micro-entomological pins, 3 to 4 mm above the surface of blotting
paper, so that larvae could feed freely on them. Before experimenta-
tion, the larvae were starved for 20 hours. One larva was released in
the center of each petridish and was allowed to feed for two hours. The
area of each leaf piece eaten by the larva was estimated using graph
paper. Percent feeding on each plant species was calculated by taking
the average of the 8 leaf pieces.
Results and Discussion
Feeding varied from 0 to 100 per cent among plant species (Table 1).
In our opinion, the plants on which feeding was less than 5 per cent
should not be included in the host spectrum of the species, since such
low feeding rates may result from test bites by the insect, which may
be taken even on non-host plants. So, the plants D. gyrans, C. cajan , M.
charantia, M. cochinchinensis, Z. mays , S. vulgare , L. esculentum , O.
sativa , S. officinarum, P. purpureum , C. sinensis, J. regia, C. rotundas,
C. nobilis x C. deliciosa, L. ehinensis, Musa sp., C. medica v. galgal, E.
japonica, O. europaea, M. indica, R. religiosa, and E. globulus should
not be included in the host list of S. dalbergiae.
Significant differences were noticed in the degree of feeding on
different varieties of G. max; Punjab- 1 was eaten more as compared to
Bragg and Lee. Other Leguminous crops (i.e., D. biflorus, P. sativum,
P. vulgaris, P. mungo and V. unguiculata ) did not show any difference
in percent feeding. Among Cucurbitaceous vegetables, L. cylindrica
and C. sativus were eaten appreciably. As compared to other Solana-
108
J. Res. Lepid.
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28(1-2):1-136, 1989(90)
109
ceous vegetables tested, S. melongena was eaten more. No significant
difference was seen in feeding among Cruciferous vegetables.
Temperate fruits were consumed more as compared to sub-tropical
fruits; however, no significant differences existed among different
pome, stone, or nut fruits. Other temperate fruits like F. ananassa and
P. granatum were also eaten appreciably while O. europaea was re-
jected. Percent feeding was very low on all the sub-tropical fruits tested.
Appreciable feeding occured on some other plants of this group also.
Maximum feeding occured on M. alba followed by B. variegata and
L. camara.
Comparison of the host range of different species of Spilosoma indi-
cates that various plant species such as A. esculentus, B. campestris
var. sarson , C. cajan, C. saliva, C. sativus, D. hiflorus , L. camara, L.
cylindrica, P. mungo, S. oleracea and V. unguiculata are eaten by both
the Oriental species, obliqua and dalbergiae. S. dalbergiae also has
some hosts like C. pepo, P. vulgaris, P. domestica and P. persica
common with the North American species S. virginica. G. max , P.
sativum, R. sativus, R. communis, S. melongena, S. tuberosum, and B.
oleracea var. capitata are hosts of S. dalbergiae, S. obliqua and S.
virginica. M. alba has been found to be a host of many Spilosoma
species, including dalbergiae, obliqua, imparilis, lubricipeda, mori and
subcarnea (Maki, 1916; Fenton, 1937; Golanski, 1967; Tietz, 1972;
Bhattacharya and Rathore, 1977; Roberts et al., 1977; Hondo, 1981).
S. dalbergiae and S. obliqua (Bhattacharya and Rathore, 1977) sho-
wed several similarities and differences in their host preference. Both
these species rejected M. charantia, M. indica, F. religiosa, O. saliva
and S. officinarum. S. dalbergiae ssp. n. accepted the plants P. grana-
tum, S. tuberosum and F. carica, which are rejected by S. obliqua. This
species rejected C. rotundas and Z. mays, which are accepted by S.
obliqua. The plants C. sativus and S. oleracea were found to be good
host plants for both species. Neither species preferred P. guajava.
The wide range of acceptable hosts clearly indicates that the new
subspecies of dalbergiae is polyphagous. The tendency of the insect to
feed on temperate fruits (e.g., almond, apple, pear, peach, plum, and
strawberry), vegetables (e.g., pumpkin, vegetable sponge, cucumber,
brinjal, potato, radish, cabbage, mustard, lady-fingre, and spinach),
and legumes (e.g., pea, black gram, French bean and soybean) indi-
cates that the species has both horticultural and agricultural pest
potentialities. It also readily accepted many forest and ornamental
plants. We cannot rule out the possibility that some of the out-breaks
recorded for obliqua (Lall, 1964) might have been caused by dalber-
giae. The tendency of dalbergiae to feed on a wide variety of plant
species indicates that the insect can be a pest in many ecosystems.
The spectrum of potential host plants of early and advanced instars of
larvae may vary. Generally, the spectrum of potential host plants have
been found to be wider for early instars than for old larvae (Wiklund,
110
J. Res. Lepid.
1973). The host range of this new subspecies indicates that it can
survive well in agricultural, forest, or mixed systems. The ability of
the species to feed on seasonal, annual, biennial, perennial, herbs,
shrubs, or trees indicate its substantial potential as a pest species.
Acknowledgements. The authors are thankful to K. M. Harris and J. D.
Holloway, C. A. B. International Institute of Entomology, London, for provid-
ing identifications of these species. Thanks are also due to W. Thomas, West
Germany, for the identification of subspecies. We are very grateful to Adam
Porter, Zoology Department, University of California, Davis, for his valuable
suggestions to improve the manuscript. The financial assistance provided by
the Department of Environment, Government of India is gratefully acknow-
ledged.
Literature Cited
AHMAD, M. & AHMAD, G. 1976. Morphological studies on the adult of Diacrisia
obliqua Walker (Lepidoptera: Arctiidae). Pakist. J. Zool., 8, 1-11.
ANONYMOUS, 1969. Annual schedule of control for Bihar Hairy caterpillar
(. Diacrisia obliqua Walker) in Nainital Tarai. Agri. Ext. Ser. Bull. G. B.
Pant University of Agriculture and Technology, Pantnagar, 8 p.
BHATTACHARYA, A. K. & RATHORE, Y. S. 1977. Survey and study of the bionomics
of major soybean insects and their chemical control. Research Bulletin
No. 107, G. B. Pant University of Agriculture and Technology, Pantnagar,
324 p.
FENTON, F. A. 1937. The insect record for Oklahoma 1935-1936. Proc. Okla.
Acad. Sci., 17, 29-31.
GOEL, S. C. & ARUN KUMAR, 1983. Studies on the morphology of larval instars of
Diacrisia obliqua (Wlk.) (Lepidoptera: Arctiidae). Ann. Soc. ent., Quebec,
28, 2-12.
GOLANSKI, K. 1967. Observations on the occurence of pests of white mulberry
(Af. alba) in Poland. Polskie Pismo ent., 37, 193-199.
HAMPSON, G. F. 1894. The fauna of British India. Moth-II. Dr. W. Junk b. v.
Publisher, the Hague, 609 p.
HONDO, M. 1981. Mortality factors and mortality processes of cononial stages of
the mulberry tiger moth, Spilarctia imparilis Butler (Lepidoptera:
Arctiidae). Jap. J. appl. Ent. Zool., 25, 219-228.
KOGAN, M. & GOEDEN, R. D. 1970. The host plant range of Lema trilineata
daturaphila (Coleoptera: Chrysomelidae). Ann. ent. Soc. Am., 63, 1175-
1180.
LADD, T. L. JR., 1987. Japanese beetle (Coleoptera: Scarabaeidae): influence of
favoured food plants on feeding response. J. econ. Ent., 80, 1014-1017.
LALL, B. S. 1964. Vegetable pests. In “Entomology in India” (Pant, N. C. ed.),
Entomological Society of India, Delhi. 187 p.
MAKI, M. 1916. Report on injurious insects of the mulberry trees in Formosa.
Formosan Government Agricultural Experiment Station, Publ. No. 90,
265 p.
PATEL, J. S. 1944. Indian Central Jute Committee. Annual Report of the
Agricultural Research Scheme for the year 1942-43, 51 p.
28(1-2):1-136, 1989(90)
111
PRASAD, D. & premchand, 1980. Some observations on moulting behaviour and
larval mortality of Diacrisia obliqua Walker on different foods. Indian J.
Ent., 42, 826-828.
PRICE, P. w. & waldbauer, G. P. 1982. Ecological aspects of pest management, pp.
33-68. In “Introduction to Insect Pest Management” Metcalf, R. L. and
Luckman W. H. eds. John Wiley & Sons, New York, 577 p.
ROBERTS, s. J.; MELLORS, w. K. & ARMBRUST, E. J. 1977. Parasites of lepidopterous
larvae in alfalfa and soybeans in central Illinois. Great Lakes Entomolog-
ist, 10, 87-93.
TIETZ, H. M. 1972. An index to the described life histories, early stages and hosts
of the macrolepidoptera of the continental United States and Canada. The
Allyn Museum of Entomology, Sarasota, Florida.
wiklund, c. 1973. Host plant suitability and the mechanism of host selection in
larvae of Papilio machan. Entomologia exp. appl., 16., 232-242.
Journal of Research on the Lepidoptera
The life-history of Tomares ballus (Fabricius, 1787)
(Lepidoptera: Lycaenidae): phenology and host plant use
in southern Spain
D. Jordan©, J. Fernandez Haeger
and
J. Rodriguez Gonzalez
Departamento de Biologia Vegetal y Ecologia. Facultad de Ciencias. Universidad de
Cordoba. 14071 Cordoba. Spain.
Abstract. The life-history of Tomares ballus in southern Spain is
described. In Sierra Morena T. ballus is monophagous and feeds on
flowers and fruits of Astragalus lusitanicus (Lam., 1783) (Fabaceae),
despite the availability of other potential host plants. The phenological
coupling between T. ballus and A. lusitanicus is considerable in Sierra
Morena, whereas the flowering period of the remaining potential host
plant species is approximately one month later. In the Guadalquivir
Valley butterflies show preferences for Medicago polymorpha (L.,
1753). Other aspects of the life-history of T. ballus are discussed in
relation to the morphological and productive features of A. lusitanicus.
Introduction
Tomares ballus is a Mediterrranean biogeographical component of
the Palaearctic butterfly fauna. It is frequently found in some parts of
North Africa, the southeastern half of the Iberian Peninsula and a small
region in southeastern France, feeding mainly on Lotus hispidus (D.C.,
1805) (Higgins & Riley, 1980). The other species of the same genus
flying in Europe is T. nogelli (Henrich-Schaffer, 1851),. which flies in
Rumania and uses Astragalus ponticus (Pallas, 1800) as host plant
(Higgins & Riley, 1980). In northern Africa the range of T. ballus
overlaps that of T. mauretanicus (Lucas, 1849) which feeds on Hippo -
crepis multisiliquosa (L., 1753) (Higgins & Riley, 1980; Courtney, 1983).
Despite the recent work by Descimon and Nel (1986), T. ballus is not a
well known species. The present work focuses on different aspects of the
life history of T. ballus and its host plants in southern Spain.
Study Area
Field work was carried out in areas located on both sides of the middle of the
Guadalquivir River in southern Spain (Cordoba province) (Fig. 1). In the
northern zone (Sierra Morena Mountains, SM), soils are mainly siliceous and
poorly-developed. A varying degree of human management has affected the
composition and structure of the original vegetation. The region is a mosaic of
different units, where areas of oak forest (Quercus rotundifolia and Q. suber)
28(1-2):1-136, 1989(90)
113
0 20Km
Map of the area showing
the location of the study
sites.
alternate with scrubland or therophytic pastureland. Scrubland is mainly
composed by Cistaceae ( Cistus ladanifer, C. monspelliensis, C. saluifolius ),
Labiatae ( Rosmarinus officinalis , Lavandula stoechas) and Ericaceae ( Arbutus
unedo, Erica arborea). A more detailed description can be found in Gonzalez
Bemaldez et al. (1976). Seven different study sites (1-7) were choosen in this
area (Fig. 1).
The deep and fertile calcareous soils of the southern zone, located in the
Guadalquivir Valley (Campina, CA), have allowed an intensive agricultural
exploitation; The original vegetation has been completely removed and replaced
by herbaceous crops (wheat and sunflowers) and olive groves. Only one site (8)
was choosen in this area (Fig. 1).
114
J. Res. Lepid.
The study area has a Mediterranean climate (Ashmann, 1973) which in-
fluences the phenology and growth rhythm of both vegetation and fauna. In the
year of our field research (1986) the annual rainfall was 532.5 mm, a figure well
below the long-term average annual rainfall for the area (x — 657 mm, n = 21
years).
Methods
Most of the data concerning the life-history of the species were obtained from
larvae collected in the field and reared in the laboratory or hatched from eggs
laid by females in insectaries (net-cages).
Different patches of A. lusitanicus (located in sites (1-6) were selected for field
measurements. A total of 108 A. lusitanicus plants were randomly tagged in
early February, when they were just resprouting. These plants were examined
weekly; their height and number of stems, inflorescences and buds were
recorded. At the end of the growing season (June), the numbers of fruits and
viable seeds were also counted. To evaluate the other potential food plants for
their frequencies, flowering phenology and flower availability in the first zone
(SM), we established five permanent line transects, each one with six quadrats
of 50 x 50 cm spaced at 1.50 m intervals. The number of inflorescences for each
herbaceous plant species were counted in the 30 quadrats every fifteen days
from February to June.
Results and Discussion
1. Use of host plants.
Since T. hallus uses a wide range of species belonging to Fabaceae as
host plants throughout its distribution range (Jordano & Rodriguez,
1988), this butterfly can be classified as oligophagous (Slansky, 1974;
Wiklund & Ahrberg, 1978; Scriber & Slansky, 1981). For example,
Descimon & Nel (1986) have reported five different foodplants in the
same area in SE France (. Medicago truncatulata, Anthyllis tetraphylla,
Hippocrepis unisiliquosa , Onohrychis caput-gallii and Lotus ornithopo-
dioides).
In the Campiha area (CA) and other areas of the Guadalquivir River
Valley where A. lusitanicus is absent, M. polymorpha is the host plant
for T. hallus. This species is a common papilionaceous legume growing
in ditches, river banks and abandoned olive groves. This area (CA)
contains a lower number of potential host plant species than pasture-
lands of the Sierra Morena Mountains (SM).
All oviposition records during 1986 from the CA population were on
M. polymorpha , despite the fact that M. sativa L. was also available, but
less abundant. At the time of maximum butterflies density, no M. sativa
plants with flowers or buds were found.
Eggs are laid on the leaves of M. polymorpha usually on the upperside
(80%, n = 24), and in many cases on plants still without flowers. Despite
the abundance of M. polymorpha in this zone, where it forms extensive
dense patches, considerable overspread egglaying is exhibited (distance
between two successive egglayings x = 12.3 ± 14.5 m, max. = 49.5, min.
28(1-2): M36, 1989(90)
115
Spp Fig. 2
Frequency of appearance
(%) of potential host
plants species in thero-
phytic pastureland in SM.
Measurement was made
using thirty 50 x 50 cm
quadrats in sites 1 - 3.
Abbreviations: TS = Tri-
folium striatum. TC = T.
campestre, MR = Medi-
cago po/ymorpha , TCH =
T. chert eri, TG ~ T. g/o -
meratum, IT - T. tomen-
tosum, BP = Biserrula
pe/ecynus , LP = Lotus
parvi floras , OC = Orni-
thopus compressus, TB =
T. bocconei, TSB = T.
subterraneum, TA = T.
arvensis , TST = T. stet
latum , CR = Coronilla
repanda, LC = Lotus
conimbricensis, MO =
Medicago orbicularis,
TAN = T. angustifolia, L =
Lathy rus sp., AV = Anthyl-
lis vulneraria, AH = A.
hamosus, AL = Astragalus
lusitanicus, GH = Genista
hirsuta and LA = Lupin us
angustifo/ius, + = species
present in the study area
but not in the quadrats.
The last three species are
perennial herbaceous or
woody plants with a bushy
appearance. Its quantifica-
tion requires a different
sampling method.
= 2, n = 15). Moreover, eggs are usually laid singly. Only one
oviposition event of three eggs on a single plant and another of two eggs
were observed. This isolated egg-laying pattern is consistent with the
one reported by other authors (Nel, 1984; Descimon and Nel, 1986).
The colonies of T. ballus in Sierra Morena (SM, study sites 1-7) are
restricted in host plant usage, feeding on A. lusitanicus despite the
availability of different potential host plants (Fig. 2). In 1986, a total of
1962 eggs were counted on A. lusitanicus and only one on Ornithopus
compressus.
Most eggs are laid between the buds of incipient inflorescences
(91.13%, n = 1962), but females occassionally lay them on apical or
mature leaves or on the main stems of plants. Several are often found on
116
J. Res. Lepid.
the same inflorescence, but not in clusters as does T. mauretanicus
(Courtney, 1983). Eggs of different age (recognizable through the
variable color of them, from pale green to dark grey) are frequently
found on the same inflorescence. Thus, plants can receive considerable
egg loads, up to a maximum of 38 eggs recorded for a single stem. This
egg-laying behaviour differs from the single egg opposition, quite
common in Lycaenidae, and is surprising for a species whose cater-
pillars have been described as cannibalistic (Nel, 1984; Descimon & Nel,
1986).
A. lusitanicus is a perennial herbaceous plant whose distribution
includes the southwest of the Iberian Peninsula and the northwest of
Africa. Plants show a variable number of vigorous and erect stems (x =
7.0 ± 5.02, n = 108) which grow up to 90 cm high (x = 55.8 ± 14.1 cm, n =
108). Stems resprout yearly from a woody root. Leaves are pinnate and
8-12 cm long. Stems produce numerous conspicuous inflorescences (x =
6.5 ± 2.0, n — 108) of dense white flowers in racemes. The average
number of floral buds per inflorescence is 17.7 ± 13.2 (n = 108). Fruits
are legumes approximately 10 cm long (Polunin, 1982) and dehisce
when ripening. Each stem produce an average of 7.5 ± 10.0 (n = 108)
mature fruits. Ripe fruits produced an average of 11 seeds (n = 100),
with only 22.4% being viable and the remainder being aborted. The
appearance of A. lusitanicus is quite different from that of the re-
maining potential host plants, since all of them are procumbent small
herbaceous plants.
This Astragalus species is toxic for livestock (Gonzalez Rodriguez,
1980) especially for sheep (Moyano, 1985) but the compound responsible
of this toxicity is unknown (Infante et a!., 1964; Poyato, 1968; Baraibar,
1982). Thus, plants are avoided by vertebrate herbivores ( Cervus
elaphus , Oryctolagus cuniculus ), which clearly benefits T. ballus survi-
vorship in the SM population, whereas grazing and trample may cause
high mortality to larvae feeding on M. polymorpha in the CA popula-
tion. The effects of A. lusitanicus alellochemicals on SM T. ballus
population remain unassessed.
A. lusitanicus grows vigorously following particular shrub vegetation
management practices (e.g. fires, plowing). On the other hand, it is
scarce in areas with considerable shrub and tree cover. This fact
suggests that the suitability of A. lusitanicus patches to T. ballus
colonies may decrease through the years and finally disappear if
additional disturbance does not occur; that is, A. lusitanicus is a
successional or even a fugitive species.
2. Life history of T. ballus.
Eggs are roughly spherical with average diameter of 0.54 ± 0.01 mm
(n = 115). The average weight is 0.0007 ± 0.0005 g (n = 151). Our data
does not allow to establish any relationship between egg weight and
female age, as happens in different satyrid butterflies (Wiklund and
28(1-2):1-136, 1989(90)
117
Karlsson, 1984; Karlsson and Wiklund, 1985). Hatching usually occurs
ten days after oviposition. New caterpillars make a hole in the bud
where the egg was laid or across the petals (in the case of an unopened
flower). Feeding occurs inside the bud. Caterpillars, described in detail
by Chapman (1904), are adapted to an endophytic life, feeding on
anthers and ovaries during their early stages. Sometimes several
caterpillars have been found in the same flower.
As flowering progresses, caterpillars feed on the developing seeds
within the fruits. In order to reach the seeds, they make a hole through
the fruit valve which is plugged with silk from the inner side. In this
manner, caterpillars can consume several fruits before finishing their
development. The high number and size of inflorescences and fruits of A .
lusitanicus allow the development of several caterpillars on the same
plant without apparent problems of competition or cannibalism
(Jordano, 1987).
Some authors have pointed out a possible mutualistic relationship
between this species and ants (Martin Cano, 1982). T. ballus cater-
pillars have three types of specialized organs: porous cupola glands,
eversible tentacles and Newcomer’s gland (Martin Cano, 1982). Obser-
vations made with a scanning electron microscope confirm the presence
of these organs in T . ballus larvae (Jordano, unpubl.). However, we have
never observed any type of interactions between this species and ants in
the study area. This supports the observations made by Descimon and
Nel (1986). In spite of this, caterpillars and pupae introduced in
artificial nests of Cataglyphis hispanica, a carnivorous ant species, were
not attacked by the ants (Jordano, unpubl.), as happens with Artogeia
rapae larvae.
During 1985, twenty five third and fourth instars larvae were
collected in the field. Of them, 36% showed evidence of attack by
parasitoids. Every parasitized larvae contained a considerable number
of parasitoids (x = 16.8 ± 6.2, n = 8) belonging to an undescribed species
of a small wasp of the genus Cotesiia (Braconidae, Microgastrinae,
Cotesiini). Parasitoids killed the larvae within 4-7 days. During this
time, larvae lost on the average 48.8% of their weight (n = 9). The
endophytic habits of larvae and their specialized feeding behavior,
likely provide some protection against parasitoids. The larvae are
exposed to parasitoid attacks only when they have to move from one
consumed flower or fruit to another.
The larvae of the wasps parasitoid come outside the caterpillars to
pupate in small cocoons and imagos emerged during the same spring.
Therefore they have more than one generation per year and probably
are not specific to T. ballus.
Larval development of T. ballus lasts about two months in laboratory
conditions with no precise light or temperature regulation (x = 58.1 ±
3.5 days, n = 18). Males and females have development periods of
similar lengths.
118
J. Res. Lepid.
Caterpillars reach a maximum weight over 0.30 g (female max.
weight = 0.40 g; male max. weight = 0.35 g). However, weight losses
occur after reaching maximum weight during prepupal phase, during
which larvae stop feeding. In the prepupal phase caterpillars leave the
plant on which they have developed, searching for a place to pupate. As
a result, the final weight of caterpillars before pupation is considerably
lower than their maximum weight, ranging between 0.20-0.25 g (females
max. final weight = 0.26 g; males max. final weight = 0.24 g). Weight
loss during the prepupation phase was about 30% of maximum weight (x
= 30.4% ± 7.6, n = 12).
Pupation occurs in the soil, generally partially buried or under stones.
Pupae are brown without rugosities or maculations. We did not find any
myrmicophilous or sound producing organs. In a random sample of
pupae obtained in the laboratory, females were found to weigh signifi-
cantly more than males (females, x = 0.17 ± 0.02 g, n = 27; males,
x = 0.14 ± 0.03 g, n = 27; F = 10.94, p < 0.01).
Diapause occurs in the pupal stage and imagos emerge the following
year. Notwithstanding, some pupae did not produce imagos, continuing
in diapause and giving imagos two years later. This fact suggests that a
cold exposure for interrupting diapause might be necessary, as has been
reported for other species (Templado & Alvarez, 1985).
Female butterflies were larger (CVL mean = 15.85 ± 1.07, n = 8; WS
mean = 29.30 ± 1.60 mm, n = 6) than males (CVL mean = 14.38 ± 0.67
mm, n = 9; WS = 27.70 ± 1.32, n = 9) (CVL - costal vein length, WS =
wing span).
Longevity of imagos in the field has not been assessed, but it ranged
between 18 and 22 days in insectaries.
Potential female fertility was approached by dissecting the abdomen
of virgin butterflies just emerged from the pupae and counting their
eggs and oocytes (Dunlap Pianka et al. 1977; Ehrlich & Ehrlich, 1978;
Dunlap-Pianka, 1979). Values ranged between 317 and 584 eggs per
female (x = 456.2 ± 102.0, n = 5).
3. Phenology.
T. ballus is univoltine and one of the first species which can be seen
every year at the study area, excluding those which spend the winter as
imagos (eg. Gonepteryx rhamni, G. cleopatra and Nymphalis poly-
chloros ). It is on the wing from early February until the end of April,
with a peak flight around mid-March. Slight between-year variations
are observed, depending on weather conditions. Isolated individuals
have been observed as early as the end of January and as late as May.
Data of study sites 1-3 displays the flowering phenology of the
potential host-plant species living in the same zone as A. lusitanicus
(Fig. 3). The flowering period of the latter species occurs earlier than
that of the remaining ones except for Trifolium subterraneum (whose
flowering and underground ripening characteristics excludes it as a
suitable host plant).
28(1-2):1-136, 1989(90)
119
Fig. 3 Flowering phenology of potential T. ba/lus host plants (Fabaceae)
growing in the study area. Inflorescences were counted in thirty 50 x
50 cm quadrats every fifteen days. The central line on the graphs show
T. ba/lus egg laying peak. Notice the phenologycal coupling between
T. ba/lus egg laying peak. Notice the phenologycal coupling between
T. ba/lus and A. lusitanicus and in a lesser extent with T. subterraneum.
Abbreviations as in Fig. 2.
Figure 4 shows the comparative phenology of T. ballus and of A.
lusitanicus at the study area (SM). The flight period of T. ballus and,
more specifically, its maximum egg laying activity, is coincident with
the flowering peak of A. lusitanicus (Jordano, 1987), whereas the
flowering peak of the remaining potential host plants occurs at least one
month later (Fig. 3). This is the case oiAnthyllis tetraphylla (the main
120
J. Res. Lepid.
ZZZZZZZZ2
ADULTS
T.baltus
tzzzzzzzzzzzz
PUPAE
EGGS
g^ZZTyZTZ/ZZZZZTZZZZ//?/^
iyyvy7-7yyyy///////xyy7^CATERPILLARS
ZZZZZZZZZZZZ2ZZZZZZZZZZZZZZZZZZZZ l
inf IEZZZZZZZZZZZZZZZZZZ23 PLANT GR°W™
A. lusitanicus
mf \\uzzzzzzzzzzzzzzzzn
Inf wWZZZZZZZZZZZZZZl
inf IV EZZZZZZZZZZZZZZ3
J
M
My DATE
Fig. 4 7. ballus and >4. lusitanicus phenology. Plant growth includes from
plant resprouting until the end of the biomass increase. Inf I: Inflore-
scences with floral buds (petals not visible). Inf II: Inflorescences with
floral buds and closed flowers (petals visible). Inf III: Inflorescences
with open flowers. Inf IV: Inflorescences with fruits.
host-plant in SE France) and M. polymorpha (the host plant for the
Guadalquivir Valley population, CA) (Fig. 3). These time lags account
for the restricted monophagous strategy of T. ballus at the SM popula-
tion, since the caterpillars feed on flowers during their earlier instars.
At Sierra Morena, (SM), T. ballus practices monophagy with relevant
behavioral and ecological relationships with its host plant, A. lusi-
tanicus. The patches of this species are easy to locate for ovipositing
females due to their spatial and temporal predictability. At the same
time, its high flower and fruit production allow the successful develop-
ment of several T. ballus larvae on a single plant. Therefore, it can be
considered as the most suitable host plant used by the butterfly locally.
In this way, herbaceous terophytic legumes could be considered as
secondary hosts in areas where A. lusitanicus is absent.
Acknowledgements. We wish to thank H. Descimon, A.M. Shapiro, P. Jordano
and two anonymous referees for helpful suggestions on an earlier draft of this
work. M. and R. Zamora provided facilities to work in “Villa Alicia”. Financial
support was provided by grant 3126/83 of the Caicyt to J.F.H.
Literature Cited
ASCHMANN, H., 1973. Distribution and peculiarity of Mediterranean eco-
systems. In: di Castri, F. & Mooney, H. A. (ed), Mediterranean type ecosystems.
Springer Verlag, Berline, pp. 11 — 19.
BARAIBAR, c., 1982. Contribucion alestudio fitoquimico y toxicolpgico del Astra-
galus lusitanicus. Tesina de Licenciatura, Facultad de Farmacia, Univer-
sidad Complutense, Madrid.
28(1-2):1-136, 1989(90)
121
CHAPMAN, T.A, 1904. Notes toward a life-history of Thestor ballus Fab. The
Entomologist 26:238 —240.
COURTNEY, S.P., 1983. Notes on Tomares mauretanicus (Lycaenidae) in Morocco.
J. Res. Lepid. 22:205-206.
DESCIMON, H. & J. NEL, 1986. Tomares ballus F. est-il une espece vulnerable en
France? (Lepidoptera Lycaenidae). Alexanor 14:219-231.
DUNLAP-PIANKA, H.L., 1979. Ovarian dynamics in Heliconius butterflies: correla-
tion among daily oviposition rates, egg weights, and quantitative aspects of
oogenesis. J. Insect Physiol. 25:741-749.
DUNLAP-PIANKA, H.L., BOGGS C.L., & L.E. GILBERT, 1977. Ovarian dynamics in
Heliconiinae butterflies: programed senescence versus eternal youth.
Science 187:487-490.
EHRLICH, P R. & A.H. EHRLICH, 1978. Reproductive strategies in butterflies. I.
Mating frequency, plugging, and egg number. J. Kansas Ent. Soc. 51:666-
697.
GONZALEZ BERNALDEZ, F„ POU ROYO, A., RAMIREZ DIAZ, L. & F. SANCHO ROYO, 1976.
E studios ecologicos en Sierra Morena. Monografia no 8. Servicio de Publica-
ciones del Ministerio de Agricultura, Madrid, 80 pp.
GONZALEZ RODRIGUEZ-CORDOBA, J.M., 1980. Toxicologia pratense. Nuestra
Cabana Julio: 35—39.
HIGGINS, L.G. Sc N.D. RILEY, 1980. Guta de campo de las mariposas de Espaha y de
Europa. Omega, Barcelona, 452 pp.
INFANTE, F., PEINADO, E. & M. MEDINA, 1964. Aportacion al estudio de la composi-
cion quimica del Astragalus lusitanicus LAM. Ill Semana Nacional Veter-
inaria : 549-560.
JORDANO, D., 1987. Estudio ecologico de las relaciones entre mariposas y plantas:
interacciones de Tomares ballus ( Lycaenidae ) y Astragalus lusitanicus
(Leguminosae) . Tesis Doctoral. Fac. de Ciencias, Sec. Biologicas, Univ. de
Cordoba, 197 pp.
JORDANO, D. & J. RODRIGUEZ, 1988. Nuevas citas de plantas nutricias para tres
especies de ropaloceros. Shilap Rev. Lepid. 16 (62):89-95.
KARLSSON, B. & C. WIKLUND, 1985. Egg weight variation in relation to egg
mortality and starvation endurance of newly hatched larvae in some satyrid
butterflies. Ecol. Entomol. 10:205-211.
MARTIN CANO, J., 1982. La biologla de los licenidos espanoles (Lep. Rhopalocera).
Miscelanea Conmemorative UA.M 1003-1020.
MOYANO, M.R., 1985. Aspectos toxicologicos del Astragalus lusitanicus LAM. en
ovinos: intoxicacion experimental. Tesis Doctoral. Fac. de Veterinaria, Univ.
de Cordoba, 149 pp.
NEL, J., 1984. Sur la plasticite ecologique et la biologie de quelques lepidopteres
(Rhopalocera) du sud~est mediterraneen de la France . These. Faculte des
Sciences et Techniques de Saint-Jerome, Universite d’Aix-Marseille, 124 pp.
POLUNIN, O., 1982. Guta de campo de las flares de Europa. Omega, Barcelona, 796
PP-
POYATO MONTES, J., 1968. Estudio fitoquimico, toxicologico y farmacologico del
Astragalus lusitanicus. Serv. Contrast.: 67-118.
SCRIBER, J.M. & F. SLANSKY, 1981. The nutritional ecology of inmature insects.
Ann. Rev. Entomol. 26:183-211.
SLANSKY, F., 1974. Relationship of larval food-plants and voltinism patterns in
temperate butterflies. Psyche 81:243-253.
122
J. Res. Lepid.
templado, J. & J. ALVAREZ, 1985. Observaciones sobre Zegris eupheme (Esper,
1800) (Lepidoptera, Pieridae). Bol. Est. Cent. Ecol. 28:81-86. .
WIKLUND, C. & C. AHRBERG, 1978. Host plants, nectar source plants, and habitat
selection of males and females of Anthocharis cardamines (Lepidoptera).
Oikos 31:169-183.
WIKLUND, C. & B. KARLSSON, 1984. Egg size variation in satyrid butterflies:
adaptive vs historical, “Bauplan”, and mechanistic explanations. Oikos 43:
391-400.
28(1-2):123-128, 1989(90)
Notes
Temporary breeding populations of Phoebis sennae eubule (L.)
(Lepidoptera: Pieridae) in Ohio and West Virginia
The cloudless sulphur, Phoehis sennae (L.) is primarily a tropical and
subtropical butterfly. In eastern North America, the subspecies P. s: eubule (L.)
is a permanent resident generally south of North Carolina, Tennessee and
Arkansas. The species regularly emigrates northward in late summer ana
early autumn, occasionally reaching southern Canada (Opler and Krizek,
1984; Scott, 1987). Northward, the species occurs in the Mississippi Valley to
central Illinois (Irwin and Downey, 1973) and along the Atlantic Coastal Plain
to New Jersey (Stone, 1903; Shapiro, 1966). In these areas, it is known to re-
produce as far north as west-central Illinois (Sedman and Hess, 1985) and
Virginia (Clark and Clark, 1951). Northern breeding records of P. s. eubule
from outside the Mississippi Valley and Atlantic Coastal Plain are virtually
nonexistant.
In 1987, much of the eastern United States witnessed a large and previously
unsurpassed flight of P. s. eubule . Numbers were observed as far north as
Wisconsin and New York. It was common in portions of Missouri, Illinois and
Kentucky and frequently encountered in Indiana, Ohio and West Virginia.
The species was unusually common even in the extreme southeastern states
where it is a permanent resident.
In Ohio, historical records of P. s eubule were limited to only eight counties
following over 130 years of collecting. During the exceptional flight of 1987,
the species was recorded in 15 additional counties and observed in at least
eight others. In West Virginia, P. s. eubule was recorded in seven counties,
representing the first records in the state. Individuals were observed and
captured as early as April and May in this region. Prior to 1987, the earliest
date of capture in Ohio was 29 July (1931).
Several temporary breeding populations of P. s. eubule were found in Ohio
and West Virginia in 1987. Many of these populations were located directly
adjacent to rivers and streams. Valleys of rivers and major streams appear to
serve as primary corridors of dispersal for this species in the region. A riparian
affinity has also been noted in Illinois (Sedman and Hess, 1985). In West
Virginia, P. s. eubule was observed at the summit of Cheat Mountain (approx.
1140 m.), thus it is possible that the species also follows mountain ridgetops.
Two color forms of P. s. eubule larvae were found and reared on wild senna
( Cassia hebecarpa Fern.) and partridge-pea ( Cassia fasciculata Michx.). Where
the butterfly utilized wild senna in Ohio, green larvae were found feeding on
both the leaves and yellow flowers of the plants. Conversely, yellow larvae were
nearly always found feeding on the flowers. Because of these color associations,
even large mature larvae were inconspicuous. Pupal periods of the species in
Ohio and West Virginia ranged from five to nine days, in contrast to the re-
ported pupal periods of P. s. eubule in Georgia (10-12 days) (Scudder, 1889)
and P. s. marcellina in Brazil (nine-13 days) (Brown and Heineman, 1972).
The finding may suggest that development is more rapid where the species is
not a permanent resident.
At least two broods of P. s. eubule were produced in Ohio and West Virginia in
124
J. Res. Lepid.
1987. The populations persisted until the hostplants began to dry in late
summer. Adults, especially males, continued to be observed into late September.
No southward autumn migration was reported, though such an occurrence was
noted in Tennessee in September (L. Martin, pers. comm.). Because the flight
of P. s. eubule is typically rapid and unflagging, many more individuals were
observed than collected. However, males are strongly attracted to bright
yellow objects and many were captured using decoys of yellow paper and dried
specimens of the species.
Exactly what caused this unprecedented explosion of P. s eubule is unknown.
Surely, no single factor can be attributed, but climatic conditions probably
played a major role. In much of the east, the winter of 1986” 87 was mild and
followed by an early spring. This may have allowed P. s. eubule to begin
reproducing and emigrating prematurely, thereby reaching northern areas
early in the season and establishing breeding populations. An early arrival in
the north may also have been augmented by overwintering individuals. If P. s.
eubule adults have the ability to overwinter, as suggested by Scudder (1889), it
could have survived far to the north of its permanent range during the mild
winter of 1986™ 87. These factors alone, however, do not explain the overall
abundance throughout much of the east.
It was also noted in 1987 that the hostplants, especially partridge-pea,
experienced a very productive year in Kentucky, Ohio and West Virginia. If
the situation was widespread in the east, it could have provided the necessary
catalyst for the rapid spread and increased productivity of P. s. eubule in this
portion of the country. An abundance of partridge-pea was observed in connec-
tion with a local outbreak of P. s. eubule in Kansas in 1987 (Howe, 1987).
Acknowledgements. Thanks are extended to the following individuals for
providing regional information: Thomas W. Carr, John F. Cryan, Les A. Ferge,
Loran D. Gibson, Dana M. Gring, Leland L. Martin, Lee D. Miller and Harry
Pavulaan.
Literature Cited
BROWN, F. M. & B. HEINEMAN. 1972. Jamaica and its Butterflies. E. W. Classey
Ltd., London, xvi + 478 pp.
CLARK, A. H. & L. F. CLARK. 1951. Butterflies of Virginia. Smithsonian Misc. Coll.
116(7): vii + 239 pp.
HOWE, w. H., 1987 Outbreak of cloudless sulfurs in Kansas. News Lepid. Soc.
No. 6 (Nov./Dec.): 80-81.
IRWIN, R. R. & J. C. DOWNEY. 1973. Annotated checklist of the butterflies of
Illinois. Illinois Nat. His. Survey. Biol. Notes No. 81. 60 pp.
OPLER, P. A. & G. o. krizek. 1984. Butterflies east of the Great Plains: an illustra-
ted natural history. Johns Hopkins Univ. Press, Baltimore, xvii + 294 pp.
SCOTT, J. A., 1986. The butterlies of North America: a natural history and field
guide. Stanford Univ. Press, Stanford, CA. xvi + 583 pp.
SCUDDER, S. H., 1889. The butterflies of the eastern United States and Canada
with special reference to New England. Cambridge; publ. by the author.
3 vols. xxiv, xi, vii + 1958 pp.
SEDMAN, Y. & D. F. HESS. 1985. The butterflies of west central Illinois. Western
Illinois Univ., Ser. Biol. Sci. No. 11. 120 pp.
28(1-2):1-136, 1989(90)
125
SHAPIRO, A. M., 1966. Butterflies of the Delaware Valley. Amer, Ent. Soc.,
Special Publ. vi + 79 pp.
STONE, w., 1903. Callidryas eubule in New Jersey and Pennsylvania. EntomoL
News 14(9): 304.
John V. Calhoun, 369 Tradewind Ct. Westerville , OH 43081
Thomas J. Allen , West Virginia Dept, of Nat. Res. Box 67 -Ward Rd. Elkins ,
WVA 26241
David C. I finer, 2161 Heatherfield Ave. Worthington , OH 43085
Thaiiatosis in the Neotropical Butterfly Caligo illioneus
(Nymphalmae: Brassolinae)
During the course of studies on flight kinematics in Neotropical butterflies,
thanatosis (death-feigning behavior) was noticed in a female Caligo illioneus.
The following observations were made in July 1987 in a small screened
insectary on Barro Colorado Island, Republic of Panama. Mass of the insect
was 1.84 g, the ambient air temperature was 27°C, and the relative humidity
88%. Ambient air motions in the insectary were negligible. Observations
began with the butterfly at rest on an insectary wall with the wings folded
together dor sally . When grasped by the base of the folded fore wings and
removed from the wall, the butterfly entered a thanatonic condition, characte-
rized by complete absence of wing or leg motion, with the legs tucked against
the body as in flight. When then placed upon (and with wings parallel to) a
horizontal surface, the insect remained motionless. While in this condition,
tactile stimulation of the wings, legs and body produced no behavioral respon-
se. Eventually the insect righted itself and flew away; in eight consecutive
trials each separated by several minutes, the mean time to self-righting was 55
seconds (s.d. = 49 s). Dropping the thanatonic insect upside-down from a
height of several meters resulted in a short vertical drop followed by wing-
spreading and active flapping flight or gliding to the walls of the insectary.
Thanatosis has been observed in a variety of animals, including numerous
beetles (Bleich, 1928), mantids (Edmunds, 1972), moths (Blest, 1964), snakes
(Gehlbach, 1970; see also Greene, 1988) and mammals (e.g. Francq, 1969;
Ewer, 1966). It has been suggested that thanatosis induces relaxation of
predator attention, possibly allowing for active escape of the prey during the
handling phase of a predatory event (Ratner & Thompson, 1960). Butterflies
are frequently attacked by insectivorous birds. Chai (1986) reported that the
jacamar Galbula ruficauda , a specialized avian predator of flying insects,
required on average 9 minutes (maximally 40 minutes) to strip large butter-
flies of their wings prior to consumption of the body. Wing-stripping by
jacamars occurs upon the cessation of struggle by the butterfly. If thanatosis on
the part of butterflies does induce momentary inattention during the wing-
stripping procedure, possibilities for escape are heightened. Additionally,
death-feigning may be an advantageous defense, following an initial unsuc-
cessful attack, against predators which only attack moving prey. Curiously,
thanatosis involves an inhibition of wing flapping concurrent with an absence
126
J. Res. Lepid.
of tarsal contact with a substrate. In general, loss of tarsal contact initiates
wing flapping responses in insects (see Chapman, 1971). Neural reflexes which
ordinarily would initiate flight must therefore be facultatively suppressed
during the thanatonic condition.
It was unfortunately not possible to obtain additional specimens of Caligo
illioneus to evaluate in detail various physiological and behavioral aspects of
thanatosis. However, the present results are unlikely to be anomalous, as
DeVries ( pers . comm.) has observed thanatosis in three papilionid species
C Parides areas, P. childrenae, and P. erithalion), in a Ly corea sp. (Danainae),
and in several ithomiine genera. Thanatosis may thus be a widespread anti-
predatory defense in tropical butterflies.
Literature Cited
BLEICH, O. E. 1928. Thanatose und Hypnose bei Coleopteren. Z. wiss. Biol. (A).
10: 1-61.
BLEST, A. D. 1964. Protective display and sound production in some New World
arctiid and ctenuchid moths. Zoologica 49: 161-162.
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.
CHAPMAN, R. F. 1971. The Insects: Structure and Function. New York, Elsevier.
xii+ 819 pp.
EDMUNDS, M. 1972. Defensive behaviour in Ghanaian praying mantids. Zool. J.
Linn. Soc. 51: 1-32.
EEWER, R. F. 1966. Juvenile behavior in the African Ground Squirrel, Xerus
erythopus (E. Geoff). Z. Tierpsychol. 23: 190-216.
FRANCQ, E. 1969. Behavioral aspects of feigned death in the opossum Didelphis
marsupialis. Amer. Midi. Natur. 81: 556-567.
GEHLBACH, F. R. 1970. Death-feigning and erratic behavior in leptotyphlopid,
colubrid, and elapid snakes. Herpetologica 26: 24-34.
GREENE, H. w. 1988. Antipredator mechanisms in reptiles. In Biology of the
Reptilia, Vol. 16. Alan Liss, New York, xi+659 pp.
RATNER, S. C. & R. W. THOMPSON 1960. Immobility reactions (fear) of domestic
fowl as a function of age and prior experience. Anim. Behav. 8: 186-191.
Robert Dudley, Smithsonian Tropical Research Institute Box 2072 Balboa
Republic of Panama
A New Specimen of Vanessa braziliensis “ab. dallasi”
(Nymphalidae) from Argentina
Vanessa braziliensis (Moore) “ab. dallasi ” was described and figured by
Koehler (1945, p. 256; pi. 20, fig. 2). The “cotypes” (apparently at least two
specimens) are stated to be from the Sierra de Ambato, Province of Catamarca,
Argentina, at 2000 meters. In addition to the color plate, I have examined the
“cotype” in the Breyer collection at the Museo de La Plata. The labels of the
“cotypes” of this and Vanessa carye “ab. bruchi,” described in the same paper,
28(1-2):1-136, 1989(90)
127
Fig. 1 . Vanessa brazi/iensis: " ab . da I Iasi" (above) and normal (below), upper
and lower surfaces, San Miguel de Tucuman, Argentina, 29. XI. 1989.
appear to have become interchanged at some point, but both specimens bear
the same label data, “Los Angeles (Capayan), 1800 m, 1.1941.” Los Angeles and
Capayan are two towns a few km NW and S of San Fernando del Valle de
Catamarca, the provincial capital, respectively.
It is evident that “ab. dallasi ” is extremely similar to “ab. ahwashted ’ Gunder
of the closely-related V. virginiensis Drury, and “ab. bruchi ” is nearly identical
to one phenotype in the “ letcheri-muelleri ” series of aberrations in V. annabel-
la Field. Just as these aberrations are not very rare in V. annabella, similar
aberrant specimens of V. carye exist in most major Argentine collections and I
have taken several myself. In V. virginiensis , specimens of uahwashtee ” are
very rare (Shapiro 1983). Likewise, other than the “cotypes,” no captures of V.
braziliensis “ dallasi ” have been reported or located in Argentine collections.
A male nearly identical to the La Plata “cotype” was captured by me on 29
November 1989 in a vacant lot in San Miguel de Tucuman, Argentina in the
company of numerous normal specimens (fig. 1).
Koehler was familiar with the European literature of temperature-induced
aberrations, and confidently attributes all the cases of “melanism” described in
his paper to “passing strong nocturnal cooling. . . . Some frost of near frost,
acting on the young chrysalids has produced the specimens in question.”
128
J, Res. Lepid.
Koehler’s explanation may be valid (Shapiro 1973, 1975), but it is extremely
unlikely that several similar aberrations of two different species would be
collected in the same locality on the same day. The possibility that the
specimens were actually produced experimentally cannot he ruled out. Indeed,
such practices ultimately gave the study of shock phenotypes a bad reputation
in Europe.
The Tucuman specimen was collected in the subtropical lowlands, where no
local cold shocks were likely. The weather records for Tucuman do not indicate
any unusual temperatures in the previous two months. However, V. brazilien-
sis is highly mobile and like V. carye in the same region appears to undergo
regular seasonal altitudinal migration. At the time of this collection, indi-
vidual immigrant V. braziliensis could be seen moving upslope and colonizing
the Sierra de Aconquija and Cumbres Calchaquies west of Tucuman, up to
4000 m. This movement coincides with the first seasonal rains of the “Bolivian
winter,” before which host plants are not available in the highlands. The
seasonality thus argues strongly against this individual having bred in the
cold mountains and descended to the lowlands, unless it had taken part in the
downslope migration five or six months before.
This is the first aberration I have seen among many hundreds of V. brazilien-
sis in 12 years of field work in Argentina. It is a very powerful testimony to the
conservatism of this series of apparently homologous aberrations, which seem
to occur throughout not only Vanessa but the closely allied genera as well.
I thank the Museo de La Plata for providing access to its collections. The
photographs are by S.W. Woo.
Literature Cited
KOEHLER, P. 1945. Melanismos naturales en Lepidopteros argentinos. Rev. Soc.
Ent. Arg. 12: 253=256.
SHAPIRO, A. M. 1973. Recurrent aberration in Cynthia annabella : a reivew with
four new records (Lepidoptera-Nymphalidae). Pan-Pac. Entomol. 49: 289 -
293.
— — — 1975. Natural and laboratory occurrence of “c/yrai” phenotypes in
Cynthia cardui (Nymphalidae). J. Res. Lepid. 13: 57=62.
— — 1983. A new record of Vanessa virginiensis “ab. ahwashtee” from
northern California (Nymphalidae). J. Res. Lepid. 20: 176-177.
Arthur M. Shapiro, Department of Zoology, University of California, Davis, CA
95616.
28(1-2):129-136, 1989(90)
Book Reviews
LEPIDOPTERAN ANATOMY. Eaton, J. L. 1988. 257 pp. W iley-Interscience ,
New York ISBN 0*471-05862-9. $49.95.
This is yet another overpriced lousy book. To make matters worse, it is
misleadingly titled: the title should be The Anatomy of Manduca sexta,
because that is the only species illustrated. The book is mainly a collection and
republication of Eaton’s work on that species, which is of course very impor-
tant in physiological and toxicological studies. Perhaps there will be follow-ups
on other very important species, such as Pieris brassicae, Galleria , or Bombyx
mori — but what can they be called if this book has co-opted the name of the
entire order?
The illustrations are usable but remarkably crude. Many of them resemble
casual preliminary sketches rather than publishable, definitive illustrations.
Not unsurprisingly, they suffer from occasional errors which will not trouble
most users but could lead graduate students, for example, astray. The most
egregious of these are identified by neurobiologist Nicholas J. Strausfeld in his
review of the book in Quarterly Review of Biology 64: 206-207, which should
be pasted into every copy at large. The book also suffers from sloppy editing,
many typos, no glossary (and many terms are quite unfamiliar, even to those
with a passing knowledge of insect morphology), and a form of organization
which makes the book needlessly difficult to use as a reference.
There is no attempt to place Manduca in any kind of ecological or phylogene-
tic context. This is inexcusable, especially since most of the really exciting
work on Lepidopteran anatomy is being done in exactly that context, particu-
larly by Niels Kristensen. Fortunately, Kristensen is heading up a synthetic
two-volume work on Lepidoptera as part of the Handbuch der Zoologie which
will, once it appears, become the standard (we are assured it will be published
in English) and relegate this work to deserved obscurity. It will, however, be
“institutionally priced” beyond any shadow of a doubt.
Study of the neuromuscular anatomy has the potential to provide a plethora
of new and important taxonomic characters for phylogenetic reconstruction, if
we can convince Lepidopterists to pickle specimens and then cut ’em up. This
book could have had a valuable role in encouraging such developments, had its
author and publisher taken a broader view. Instead, it is basically a “how-to-
cut-up-a -Manduca" manual and might be useful for insect morphology courses
in places where that beast is a pest and readily available. Publishers who issue
misleading blurbs for new books, like the one put out by Wiley for this, deserve
worse than contempt. They deserve to be boycotted.
Arthur M. Shapiro , Department of Zoology, U. C. Davis, Davis, CA 95616.
BIOGEOGRAPHY AND QUATERNARY HISTORY IN TROPICAL LATIN
AMERICA. T. C. Whitmore and G. T. Prance (Eds.). Oxford University Press,
Oxford. 214 pp. ISBN 0-19-854546-0. Price £45.00 (hardback).
During the early 1970s, many scientists working on the biogeography of
Neotropical organisms, fell under the spell of what was to be called afterwards
the “Pleistocene Refuge Theory”. The simplicity and elegance of the evolution-
130
J. Res. Lepid.
ary model, first proposed for South America by Haffer, and Vanzolini &
Williams, rapidly gained many adepts. Unfortunately, a number of workers
were misled by the oversimplification and popularization of the model, and
applied it indiscriminately to several groups of organisms for which a poor or
weak database was available. It is not suprising then, that by the end of the
decade, several scientists were questioning strongly the model, pointing out its
inconsistencies, and proposing alternative views.
The book under review has had a very long gestation period. I have been
privileged to be allowed examination of preliminary drafts of portions of it, and
have been witness to its development. I am under the impression that its publi-
cation would have had more impact a few years ago, as the subject is now con-
sidered somewhat demode , having been incorporated uncritically into several
textbooks.
Nevertheless, the different chapters included, are the best introduction avail-
able at present, on this fascinating interpretation of the biogeography of
tropical South America which, as indicated in the Introduction, states:
“. . . there are in the lowlands of tropical South America centres rich in species
or characters separated by regions with a poorer or more mixed fauna. These
centres were interpreted to coincide with refugia to which the rain forest was
restricted at past times of drier, more seasonal climate.”
The book is divided into seven chapters by four authors; their quality is
uneven, but this should not be interpreted as implying that some authors are
less knowledgeable than others. In fact, all four authors are respected author-
ities, with foremost experience of their specialities. Rather, the unevenness is
a reflection of the widely differing amounts (and quality) of information
available for the groups analysed (plants, by Prance; butterflies, by Brown;
birds, by Haffer; and early man, by Meggers). It is immediately obvious that
the paucity of data afflicts severely the contributions by Prance and Meggers;
much finer analyses can be made with the wealth of information gathered for
birds and butterflies. But even such a rich database has its interpretation
problems; how can the superspecies-allospecies nomenclature of the ornitho-
logists be reconciled with the more “orthodox” species-subspecies nomencla-
ture of the lepidopterists? Haffer goes even as far as to recognize “first-order
superspecies” and “second-order superspecies” (= megasuperspecies), while at
the same time rejecting the “unqualified” use of subspecies, which he regards
as possibly based on varying subjective criteria.
It appears to me that many of the “allospecies” recognized by ornithologists,
are based on negative evidence (“absence or near-absence of phenotypic indica-
tions for hybridization in the populations along the contact zone”), while
recognition of subspecies among butterflies is effected more as an extrapola-
tion from a few observations made on contiguous natural populations hybridiz-
ing freely along narrow or very narrow contact zones (e.g. among Heliconius).
It should be understood then, that what ornithologists call “species” (or
allospecies) does not mean necessarily the same for the lepidopterists; actually,
there is a complete continuum between the case of two fully developed, closely
related (sister) “species”, completely sympatric, which never hybridize, and
that of two sets of distinguishable (by whatever means, morphological, statisti-
cal, biochemical, caryological, etc.) parapatric populations, connected by a
broad intergradation (“hybridization”) zone. It is easy to recognize the former
as an instance of two evolutionarily independent “species”, but what shall (or
could) the latter sets be named? One may decide to call them subspecies, and
28<1-2):1-136, 1989(90)
131
that might be an appropriate and useful action, but where does the limit lie
then between “species” and “subspecies”? At what hybridization rate do you
stop calling them “species” and start recognizing them as “subspecies”?
This problem could be circumvented by using a “phylogenetic species” (e.g.
diagnosed by unique combinations of characters) concept, instead of the more
traditional “biological species” concept, in order to be able to prepare phyloge-
netic analyses of taxa. These in turn would be compared to area cladograms,
searching for congruences, following the rules of vicariance biogeography. This
approach has recently been applied by Cracraft & Prum (1988. Patterns and
processes of diversification: Speciation and historical congruence in some
Neotropical birds. Evolution 42: 603-620), obtaining some thought-provoking
preliminary conclusions.
The above considerations may seem somewhat out of place in this review, but
I believe they are fundamental to the main conclusions of this book: There are
centres of endemism for groups of organisms in the humid tropical lowlands
of Central and South America (although statisticians may frown about the
methods employed in delimiting them), and they seem to require more than
just an ecological explanation.
What is particularly worrisome, is that most recent criticisms directed
against the “refuge theory,” focus principally on the early data and prelimin-
ary hypotheses, presented in the first discussions on the subject, without
taking into account the great deal of information gathered after the sympo-
sium at Macuto in 1979 (papers presented at that meeting were edited by
Prance, 1982, Biological Diversification in the Tropics. New York, Columbia
Univ. Press). For instance, two commonplace (and erroneous) generalizations,
often attacked by the opponents of the “refugialists” are, first, that large
portions of the humid tropical forest were replaced by savanna during glacial
times and, second, that endemism centers are areas of maximal species
diversity.
As regards the first argument, it has been shown that there are several more
kinds of forest types than previously imagined, each with its own retinue of
more or less specialized residents. It can be understood then that some
organisms may dwell only in certain forest types, while others might be more
tolerant. Thus, it is not necessary to invoke large-scale replacement of “forest”
by “savanna” to produce vicariances in the distributions of highly specialized
organisms: it might be enough to substitute for example a humid forest for a
drier kind of forest to produce isolation of populations. This would explain why
certain groups of (ecologically specialized) organisms are split into many more
endemism centers than other (generalized) groups.
Another myth which must be abandoned (and this might produce some
discomfort among conservationists who have used the refuge theory to propose
areas for preservation of genetical resources), is that core areas represent
centers of highest diversity; instead, diversity is maximal in transition areas
between centers of endemism, where natural disturbances (by wind, fire, river
dynamics, etc.) appear to be strongest.
This book must be read carefully by everyone interested in the subject; it is by
far the most complete and authoritative treatment on the controversial ideas
surrounding modern thoughts on the biogeography of tropical America,
although obviously favoring a model of allopatric differentiation.
Gerardo Lamas, Museo de Historia Natural, Universidad Nacional Mayor de
San Marcos, Apartado 14-0434, Lima-14, PERU .
132
J. Res. Lepid.
THE ECOLOGY AND CONSERAVATION OF THE PURPLE EMPEROR
BUTTERFLY ( Apatura iris).
K. J. Willmott. 1987. Published by the author, London. 140 + 8 + 6 + 18 +
18 pp., ill., 3 pull out maps. Price not stated.
The publication under review is a report on a research project carried out in
England by the author over three consecutive years (1981-1984) and taking
into account his previous experience with the species. The publication was pro-
duced in a small number of copies. As it contains confidential information on
localities and their owners, it is not offered for sale in a general way. The
research and publication of the report were sponsored by “Associated Tyre
Specialists” of Harrow (Middlesex). The author is not a professional ento-
mologist, but has had some 20 years of lepidopterological experience when he
started on the project. Topics from the contents include: Status and distri-
bution; ecology of adults; Territories; Pairing; Ovipositing; Larval ecology
instar by instar; The pupa; Behaviour; Conservation; The ideal habitat; Effect
of Climate; Habitat management. The whole work is richly illustrated by
photographs, line drawings and maps. The research has been carried out in the
field and the danger of substituting captive breeding for extensive field work is
pointed out. Some of the reasons for the fluctuation of A. iris populations are
demonstrated including climatic effects during critical stages of the species’
life-cycle. I would have wished to find more information on the methods and
techniques used, on the population size, structure and dynamics (a very diffi-
cult task indeed with this species!) as well as a comparison between the
assumptions of previous authors and the results of the present research. This is
a minor criticism, outweighted by the excellent line drawings showing various
aspects of the A. iris' ideal habitat, detailed maps of selected habitats, and dia-
grams warning of possible forms of habitat destruction and the instructive
photographs. All in all, this is one of the best applied ecological studies for con-
servation purposes of a single butterfly species. It is to be hoped that the author
will be given the opportunity both to continue his study of A. iris on a long-
term basis as well as to expand his research in other areas and species; why not
give him an opportunity to study all three European Apatura species on the Con-
tinent? On the work he has accomplished he fully deserves our compliments.
Otakar Kudrna, Karl- Straub- Str. 21, D-8740 Bad Neustadt — Salz ( Germany).
TAGF ALTER 2. Hans-Josef Weidemann. 1988. Neumann-Neudamm Verlag,
Melsungen (Germany). 372 pp., col. ill.; ISBN 3-7888-0509-9. Price 48, — DM
hardback.
This small pocket book is the conclusion of the book reviewed in J. Res. Lepid.
26:288. Generally speaking, almost everything said in that review is relevant
also to this volume. It deals with the “remainder” of the Lycaenidae and the
families of Riodinidae, Nymphalidae, Satyridae and Hesperiidae, as recognized
by the author. It is introduced by a general part dealing with butterfly biology
including the early stages, butterfly conservation (the counterproductiveness
of typical German contemporary alibi-legislation forbidding the collecting of
28(1-2): 1-136, 1989(90)
133
and research on nearly all European butterflies without a special permit given
from case to case at the will of bureaucrats lacking knowledge of the problems
is pointed out), key to larvae and a table of biological data. The systematic part
is arranged in the same way as in the first volume; the word “Verhalten” (i.e.
behaviour) is also missapplied in this volume to the time of appearance, voltin-
ism and occasionally other aspects of adult biology and very rarely used in its
correct sense. A systematic list of taxa with, in Weidemann’s opinion, valid
names and some synonyms as well as German vernacular names (often new), a
bibliography and an index conclude the book. Some chapters were written
chiefly by the Dutch butterfly ecologist F.A. Bink. The small size of about 11 x
18 cm is not to be recommended for this type of book. The book is richly illusj
trated by colour photographs, mostly taken by the author in captivity. Of 87
references cited by Weidemann 25 (30%) refer to his own papers, whereas
numerous standard works are missing.
As in the case of the first volume, the most important parts of this book are
interesting (representative?) observations on the biology of many species,
whereas the systematic and related parts are much less worthy of attention.
Unfortunately, the biological observations mostly lack information on the
methods and techniques used to obtain them. Like in the first volume, the
author split butterflies into two categories, K-strategists and r-strategists
without ever having counted their populations, necessary information for
making such judgement, assuming that there really are K-strategists among
the butterflies (we know that all butterfly species are r-strategists!). Further,
in the table of biological data, information is presented as fact without a state-
ment as to how it has been obtained. Curiously, some of these “data” are
quantified, like the egg load of all species, stated in numbers. As Weidemann’s
book is said to deal with the butterflies of Germany (the Alps excepted), I
wonder why some Mediterranean (s.l.) species (in vol. 1 Archon apollinus,
Papilio hospiton, P. alexanor etc.) have been included and Erebia euryale has
been excluded. Most regretable is the misidentification of the figures of some
species: the butterfly on p. 295 (top) is Coenonympha tullia (not C. glycerion )
and of the six species figured on p. 307 only Pyrgus malvae and P. cirsii are
correctly identified. Weidemann’s comments on zoological nomenclature and
the instability of scientific names are deplorable and naive. His remedy, the
utilisation of “stable” German vernacular names, contradicts his changing
probably a half of them around or replacing them by new original creations of
his own. Like the first volume, this one is also full of statements that could be
arrived at only after years of painstaking research. The study of distribution
and population dynamics of Carcharodus alceae (cf. p. 324) would be methodi-
cally very difficult, if at all possible. The majority of Weidemann’s readers will
be amateurs unlikely to recognize these shortcomings, which makes them even
more dangerous. I cannot explain Weidemann’s obsession of enclosing many
terms in inverted commas; I suspect his uncertainty in matters of terminology
is the reason of this bad form.
It is difficult to pass a definitive judgement upon this book, praised by many
reviewers. On the one hand it is far more than the simple small pocket book it
really is, and should probably deserve much praise as such, had it been better
written. On the other hand, it must be judged as a serious attempt at a natural
history of German butterflies, and as such it leaves much to be desired. I would
prefer to judge it just as a first draft of a manuscript presented for discussion
134
J. Res. Lepid.
before being finalized. I would surely congratulate its author under such
circumstances.
Otakar Kudrna, Karl- Straub- Str. 21, D-8740 Bad Neustadt — Salz (Germany .
LISTE INVENT AIRE SYSTEMATIQUE ET SYNONYMIQUE DES
LEPIDOPTERES DE CORSE. Charles E. E. Rungs. 1988. Alexanor (Suppl.)
15:[l]-[86]. ISBN 2-903273-02-2. Price not stated.
The publication under review is a supplement to the popular Leraut’s cata-
logue of the Lepidoptera of France and Belgium and deals with all families,
including the “Microlepidoptera”, listing a total of 1386 species. The species
are listed in systematic order, some with the usual synonyms added to the
valid name. Apart from the systematic list of taxa, there are an extensive biblio-
graphy (around 400 titles); explanatory notes and comments on the taxonomy,
nomenclature, occurrence in Corsica and other aspects concerning 77 taxa; an
index of abbreviations of author’s names; an alphabetic index of scientific
names and an appendix listing species to be excluded from the fauna of Corsica
mostly for being misidentified by previous authors. Species entries are num-
bered; each entry includes the author’s name in full and the year of publication,
as well as the relevant subspecies or “subspecies” name. All entries are cross-
referenced to Leraut’s catalogue and some to the numbered annotations at the
end of the systematic part.
Rungs is not quite as generous as Leraut in his recognition of generic status
of some of at best subgeneric names (e.g. he does not recognize Leraut’s “genus”
Ly sandra), but he certainly is a splitter in my view, having recognized weak
“genera” like Heodes, Fabriciana and Cynthia. He is certainly very generous in
attributing the rank of subspecies to some unequivocally infrasubspecific
names: Anthocharis cardamines cardamines meridionalis (Verity, 1908),
originally proposed for an infrasubspecific race from Florence is treated as a
Corsican subspecies of this species, and Verity’s seasonal form calidogenita
described from Italy: Toscana is treated as the valid name for the Corsican
subspecies of Celastrina argiolus (Linnaeus, 1758). As Rungs correctly listed
all authors’ names in full, I fail to follow why he published the list of their
abbreviations. The omission of parenthesis for the names of authors of species-
group taxa listed in combination with a generic name other than that forming
the original combination can be accepted in a faunistic and similar publication;
it is regretable in a systematic catalogue like this. It would have been better to
place the page numbers in parenthesis, to distinguish them from unbracketed
pages of the same volume, than to employ for the purpose square brackets
which have a different specific purpose in taxonomic publications. In spite of
these criticisms, I am sure that the publication under review can be of interest
to any serious student of the Lepidoptera of Corsica.
Otakar Kudrna, Karl- Straub -Str. 21, D-8740 Bad Neustadt-Salz ( Germany).
28(1-2):1-136, 1989(90)
135
LEPIDOPTERA. II. Rhopalocera, Hesperiidae, Bombyces, Sphinges,
Noctuidae, Geometridae. W. Mack. 1985. In: H. Franz: DIE NORDOST-
ALPEN IM SPIEGEL IHRER LANDTIERWELT. Band V. 484 pp.;
U ni ver sitatsverlag Wagner, Innsbruck (Austria). Price cca 240, — DM. (In
German).
The first part of the fifth volume of the monumental monograph of the north-
eastern (Austrian) Alps was devoted to the “Microlepidoptera”; the second
part, here under review, is fully devoted to the “Macrolepidoptera”. It deals
with the Rhopalocera (inch superfamilies Papilionoidea and Hesperioidea) and
Heterocera (the “Bombyces”, “Sphinges”, Noctuoidea and Geometroidea). The
author of this book, who died before the manuscript went to press, was little
known outside his own “home range”; he spent most of his life as a natural
science teacher at a grammar school (“Gymnasium”) in Gmunden, lepidoptero-
logy being his lifelong hobby, although he was a naturalist by education. The
book under review is little known by European lepidopterists, but constitutes
one of the most painstaking compilations of faunistic data I have ever seen.
Apart from general information on the range, ecology and other aspects, it lists
all records of the species from the area studied, divided into 11 smaller dis-
tricts. Each record provides information so far as is known on the collector or
author, locality, date, method of capture, altitude and source of reference. The
large-size (A4) book is concluded by an extensive bibliography and an alpha-
betical index to scientific names. It is difficult to review a book of this type. It is
a “boring” and expensive book lacking illustrations, practically an enormous
collection of records, as they are seldom found. I would wish to have someone
take up the records collected on the about 190 butterfly species recorded in the
northeastern Alps, as contained in this book, bring them up to date, analyze
them and utilize them for the conservation of the rich butterfly fauna of the
area. I recommend the book to anyone interested in the Lepidoptera of Austria
and the Alps; it is well worth paying the high price the publishers ask for it.
Otakar Kudrna, Karl- Straub -Str. 21, D-8740 Bad Neustadt — Salz (Germany).
DECLINE AND CONSERVATION OF BUTTERFLIES IN JAPAN I.
Hama, E., M. Ishii, & A. Sibitani. 1989. Lepidopterological Society of Japan,
Osaka. 145 pages + x, 14 color plates. 1500 Yen. In Japanese, English
summaries
Although the Japanese as a nation are widely viewed as unrestrained
environmental despoilers, this fine volume indicates there is a sensitive cadre
of individuals with great concern to what is happening to their homeland. A
series of papers herein show the conservation situation in Japan is strikingly
similar to that in most other industrialized nations, with butterflies, as key
indicators of ecosystem fitness, declining under many circumstances.
The three editors combined the efforts of 19 contributors to round out the
volume. It is organized as a series of case histories of 22 butterfly species which
together are found across a wide variety of habitat types of the peninsula. Each
case gives the details for that species in a specific area for which long term (at
136
J. Res. Lepid.
least ten years) information are available, including facts relating to the
species decline or extirpation. All families are covered, and most species are
local sedentary animals. Unfortunately, these accounts are in Japanese so
the information was only accessible to me from the brief English summary of
each. The tables, maps, and photographs accompanying each account imply
the treatments are descriptively thorough, but frustrating for an English
limited person.
A 7 page introductory essay by Sibatani gives an outstanding overview of the
conservation status of the butterfly fauna of the country. The first legislation
to protect butterflies dates from the mid-1930’s, when special populations at
specific sites were designated Tennen Kinenbutu or “natural monument things”.
The protection was in the form of collecting prohibition. The trend of protection
continues to the present, with 37 species now “monuments”. As is well known,
this action has almost no effect in bringing about the advertised protection.
Thus Japan repeats the German fallacy. Sibatani gives a tabular summary of
the 22 treated species with years of extinction or decline, year of protective
action, and processes and possible causes of decline cited.
The Environmental Agency of Japan is making a survey of the biota of the
county, but no results are yet in. Clearly there is growing awareness of
conservation issues. We must applaud the Lepidopterological Society of Japan
for taking a leadership role in calling attention to the plight of the rich
endemic biota of its area of expertise. This is an historically important position
paper. We strongly encourage joint exchanges of information and moral
support to the movement to conserve what we can of the natural world. We are
applying to the Society to permit us to reprint Prof. Sibatani’s paper in order
that a wide audience of non- Japanese language workers can share in this
distressing situation.
R. H. T. Mattoni, 9620 Heather Road, Beverly Hills, CA 90210 USA
INSTRUCTIONS TO AUTHORS
Manuscript Format: Two copies must be submitted (xeroxed or carbon papered),
double-spaced, typed, with wide margins. Number all pages consecutively and put
author’s name at top right corner of each page. Underline all words where italics are
intended. Footnotes, although discouraged, must be typed on a separate sheet. Do not
hyphenate words at the right margin. All measurements must be metric. Metric altitudes
and distances should include imperial equivalents in parenthesis. Time must be cited on a
24-hour basis, standard time. Abbreviations must follow common usage. Dates should be
cited as example: 4. IV. 1979 (day-arabic numeral; month-Roman numeral; year-arabic
numeral). Numerals must be used before measurements (5mm) or otherwise up to number
ten e.g. (nine butterflies, 12 moths).
Title Page: All papers must have the title, author’s name, author’s address, and any titular
reference and institutional approval reference, all on a separate title page. A family
citation must be given in parenthesis (Lepidoptera: Hesperiidae) for referencing.
Abstracts and Short Papers: All papers exceeding two typed pages must be accompanied
by an abstract of no more than 300 words. An additional summary is not required.
Name Citations and Systematic Works: The first mention of any organism should include
the full scientific name with unabbreviated author and year of description. New
descriptions should conform to the format: description of male and/or female, type data,
diagnosis, distribution, discussion. There must be conformity to the current International
Code of Zoological Nomenclature. We strongly urge deposition of types in major museums,
all type depositories must be cited.
References: All citations in the text must be alphabetically listed under Literature Cited
in the format given in recent issues. Abbreviations must conform to the World List of
Scientific Periodicals. Do not underline periodicals. If four or less references are cited,
please cite in body of text not in Literature Cited. Journals and serials not listed in the
World List are to be abbreviated according to the Serial Publications on the British
Museum (NH), 3rd edition (1980) or given in full.
Tables: Tables should be minimized. Where used, they should be formulated to a size
which will reduce to 11 x 19 cm (or 4 ¥2 x 7V2 inches). Each table should be prepared as a
line drawing or typed with heading and explanation on top and footnotes below. Number
with Arabic numerals. Both horizontal and vertical rules may be indicated. Complex tables
may be reproduced from typescript.
Illustrations: Color can be submitted as either a transparency or print, the quality of
which is critical. Black and white photographs should be submitted on glossy paper, and, as
with line drawings, must be mounted on stiff white cardboard. Authors must plan on
illustrations for reduction to page size. Allowance should be made for legends beneath,
unless many consecutive pages are used. Drawings should be in India ink at least twice the
final size. Include a metric scale or calculate and state the actual magnification of each
illustration as printed. Each figure should be cited and explained as such. Each illustration
should be identified as to author and title on the back, and should indicate whether the
illustration be returned, which will be at the authors expense.
Legends should be separately typed on pages entitled “Explanation of Figures”. Number
legends consecutively with separate paragraph for each page of illustrations. Do not attach
to illustration. Retain original illustrations until paper finally accepted.
Review: All papers will be read by the editor(s) & submitted for formal review to two
referees.
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 28 Number 1-2 Spring/ Summer 1989(1990)
IN THIS ISSUE
Date of Publication: July 25, 1990
Mating Behavior and Male Investment in Euphydryas anicia 1
(Lepidoptera: Nymphalidae)
Francois J. Odendaal, Kristina N. Jones & Frank R. Stermitz
The Biology of Colias blameyi (Pieridae), the “Green Sulphur” 14
of the Argentine Puna
Arthur M. Shapiro
The Early Stages of Doa dor a Neumoegen and Dyar 26
(Lepidoptera: Noctuoidea: Doidae) in Baja California, Mexico
John W. Brown
The Lepidoptera of a central florida sand pine scrub community 37
Dennis Profant
New Records of Lepidoptera for New York and New Hampshire 75
(Nymphalidae, Noctuidae)
Tim L. McCabe
Suppression of the black pigment in female hybrids of Papilio 84
glaucus and P. multicaudatus : further evidence of the
value of ecdysone in breaking pupal diapause
Sir Cyril A. Clarke, H.H. Rees & David A. West
Studies on Spatial Distribution in the Teak Carpenterworm 88
Cossus cadamhae Moore (Lepidoptera, Cossidae)
George Mathew, P. Rugmini & K. Jayaraman
A New Species of Argyrotaenia from Arizona (Lepidoptera: 97
Tortricidae)
J.F. Gates Clarke
A new subspecies of Satyrium auretorum (Lycaenidae) from 100
the Santa Monica mountains of southern California
John F. Emmel & Rudolf H. T. Mattoni
. Potential host range of Spilosoma dalhergiae (Moore) n.ssp. 105
(Lepidoptera: Arctiidae) in India
S.N. Tiwari & N.P. Kashyap
The life-history of Tomares ballus (Fabricius, 1787) 112
(Lepidoptera: Lycaenidae): phenology and host plant
use in southern Spain
D. Jordan, J. Fernandez Haeger & J. Rodriguez Gonzalez
Notes 123
Book Reviews 129
Cover Illustration: Hybrids of Papilio glaucus x P multicaudatus. See
Clarke, Rees & West, pages 97=99
'lume 28
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Founder:
The Lepidoptera Research Foundation, Inc.
9620 Heather Road
Beverly Hills, California 90210
(213) 274-1052
William Hovanitz
Editorial Staff: Rudolf H. T. Mattoni, Editor
Scott E. Miller, Assistant Editor
Associate Editors: Emilio Balletto, Italy
Henri Descimon, France
Philip DeVries, U.S.A.
Thomas Emmel, U.S.A.
Lawrence Gall, U.S.A.
Hansjuerg Geiger, Switzerland
Otakar Kudrna, Germany
Ichiro Nakamura, U.S.A.
Arthur Shapiro, U.S.A.
Atuhiro Sibatani, Japan
Manuscripts and Notices Material may be sent to the Editor at:
9620 Heather Road, Beverly Hills, CA 90210 (213) 274-1052
The JOURNAL is sent to all members of the FOUNDATION.
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Journal of Research on the Lepidoptera
28(3):137-238, 1989(91)
The Zoogeography and Systematics of the Argentine
Abstract. The geographic distributions and systematics of the
Pieridae of the Andean highlands and Patagonia, Argentine Republic,
are reviewed. Previously unpublished records based on twelve years’
field work by the author and collections in Argentina, the United
States, the United Kingdom, etc. are provided to clarify zoogeographic
and systematic problems in this fauna. Tatochila blanchardii is
reduced to a subspecies of T. autodice . The taxa T. arctodice, macro-
dice, sterodice, vanvolxemii and fueguensis all become subspecies of
the widespread polytypic species T. mercedis. The possibility that
Hypsochila wagenknechti and H. galactodice are conspecific is dis-
cussed. Records of putative galactodice from the puna of Jujuy are
presented. The taxonomic history of Phulia nymphula in Argentina is
clarified, and its range extended south to western Neuquen. Precise
localities are given for the first time for the Chilean taxa Eroessa
chiliensis and Mathania leucothea in Argentine Patagonia. Colias
flaveola is* reported for the first time in Argentina. Colias mendozina,
rediscovered in the highlands of Mendoza, is treated as a full species.
Zoogeographic relations among the puna , the pampean ranges
(Cumbres Calchaqufes — Sierra de Aconquija), the main cordillera ,
the Cordon del Viento, and Patagonia are clarified by the Pierid data.
Previous work has overemphasized the distinctness of the Fuegian
fauna, whose only important Pierid endemic is the possibly-extinct
Colias ponteni.
Subspeciation in most genera is probably of Quaternary origin, but
speciation and certainly the origin of endemic genera or species-
groups antedates the Pleistocene. The existing Pierid fauna repre-
sents the current stage in a long and complex process involving
dispersal, vicariance, and evolution, the outlines of which are only
beginning to emerge. Current distributions reflect both contemporary
ecological phenomena and the long-term sequelae of past faunal
movements in response to geoclimatic change.
Resumen. La fauna de mariposas Pieridae de los sectores altoandi-
nos y patagonico-fueguinos de la Republica Argentina fue analizada
del punto de vista zoogeografico — historico, utilizando muestreos
basados en 12 anos de colecta por el autor y los registros confiables de
diversos museos y colecciones en dicho pais, Estados Unidos y Ing-
laterra, entre otros. La Tatochila blanchardii por razones de intergra-
dacion espontanea ya se halla subespecie de T. autodice. Las taxa T.
arctodice , macrodice , sterodice , vanvolxemii y fueguensis cambian en
subespecies de la especie politipica muy ampliamente difundida T.
mercedis. La ocurrencia de fenotipos muy parecidos a la Hypsochila
138
J. Res. Lepid.
galactodice en la puna argentina indica la posibilidad de simpatria
con la H. wagenknechti sulfurodice en dicha zona, a pesar de la
ambiguedad de la relacion de estas “especies” mas al sur. Se aclara la
historia muy confusa de la taxonomia del genero Phulia en el Cono
Sur, y se presenta registros mas australes que los anteriormente
conocidos. Tambien se ofrece registros minuciosos de las mariposas
chilenas Eroessa chiliensis y Mathania leucothea en la Patagonia
argentina y de Colias flaveola en la cordillera de San Juan por
primera vez. La Colias mendozina , recien descubierta en la cordillera
de Mendoza, se coloca como especie. Se aclaran las relaciones entre la
puna, las Sierras Pampeanas, la cordillera, el Cordon del Viento, la
Patagonia y la Tierra del Fuego, mostrando entre cosas una enfasis no
justificada en el tratamiento de la piendofauna fueguina, cuya unica
especie endemica es la Colias ponteni , “perdida” desde 1852.
Para explicar los patrones de especiacion y distribucion de dicha
fauna hay que desarrollar un esquema historico tomando en cuenta
los cambios geofisicos y climaticos del continente y vinculando los
ambientes tropicales con los extratropicales y los altos con los bajos.
Segun parece la especiacion y seguramente el origen de generos y
subgeneros o grupos de especies ha sido anterior a la Pleistocena,
aunque la subespeciacion, como en la T. mercedis, sea fenomeno
pleistoceno y/o holoceno. El estado actual de la fauna refleja una
historia muy complicada de dispersion y migracion, mezclada con
procesos de vicariancia tal que no se puede atribuir todo a ningiin solo
proceso evolutivo y/o faunlstico.
El cuadro que doy al final de este articulo, sin contener casi nada de
nuevo, excepto quiza algunos datos biologicos de poca importancia,
tiene por objeto mostrar de un solo golpe de vista ese laberinto en
parte desembrollado . . . demostrar que desgraciadamente nuestros
conocimientos sobre las especies del genero Tatochila son aun muy
incompletos .... Surge inmediatamente la vieja cuestion: ^son
especies o son variedades? Sabido es que este termino especie es por
demas elastico ... 1
Eugenio Giacomelli, 1915
Introduction
Twenty years ago the lowland tropics were commonly viewed by
biologists as a sort of museum of living fossils, persisting untouched
through the ages while the biota of higher latitudes was decimated and
reinvigorated by the geoclimatic catastrophes of the Quaternary.
Academic discussions of the causes of tropical biotic diversity — which
1 “The chart which I give at the end of this article, without containing anything much
that is new except a few biological data of little importance, has as its object to
demonstrate at a glance this partly-disentangled labyrinth. . .to demonstrate that,
unfortunately, our knowledge of the species of the genus Tatochila is yet very incom-
plete . . . The old question immediately arises: are they species or varieties? It is known
that this term species is, moreover, elastic ...”
28(3):137-238, 1989(91)
139
typically meant tropical lowland rainforest biotic diversity — were
very popular in the 1960s, In this context, the “geologic time hypo-
thesis” claimed that a vast flow of time uninterrupted by disaster had
allowed biotic richness to accumulate unencumbered in low latitudes.
That view was later stood precisely on its head. By the 1980s the
“refugial hypothesis” had become the conventional wisdom: the tropi-
cal forest had been fragmented repeatedly in the Quaternary, dividing
populations and promoting speciation. Instead of stasis, the tropics
were now seen to be in a state of perpetual dynamism; instead of
antiquity, species were now seen as manifestations of the recent past
— an equally erroneous view, as subsequent events have shown.
The locations of proposed Quaternary refugia in the tropical lowlands
are inferred from contemporary biological phenomena, such as the
geographic distribution of endemism and of zones of intergradation or
hybridization (the two are not necessarily the same). Extremely little
paleoclimatic information is available for the lowland tropics, pri-
marily because suitable depositional environments for pollen and
other fossils are rare (cf. Liu and Colinvaux 1985, 1988). To a degree
which can be embarrassing to some of its practitioners, the refugial
theory is pegged to climatic reconstructions in the Andean highlands
(where excellent palynological records abound) (Shapiro 1989a). At
present there are no satisfactory dynamic climate models incorporat-
ing orographic effects which relate the highlands to the tropical
forests, permitting a test of refugial scenarios derived from present
distributions. Progress is being made in refining the existing crude
models (Manabe and Hahn 1977, Rind and Peteet 1985), but until they
are much better this will remain an important structural weakness of
refugia as a purported synthetic theory.
The asymmetry in paleocl i m atology is mirrored in biogeography:
there have been few in-depth studies of highland organisms, and our
knowledge of their distribution is proportionately poorer than in the
lowlands. Published distribution maps create a false impression of
good coverage. On closer inspection one typically finds the dots are
distributed along the major trans-Andean highways. One of the first
efforts to correlate animal speciation and distribution in the Andes
with Quaternary climatic dynamism was by Adams (1973, 1977) and
Adams and Bernard (1977, 1979, 1981), working on the pronophiline
Satyrid butterflies of the Colombian and Venezuelan Andes and near-
by ranges. But Adams merely suggested a mechanism for speciation —
a plausible and quite conventional one - and was unable to correlate
specific species with specific geohistorical events, since most of the
relevant palynology had not been published yet. Indeed, there is
nothing in Adams’ work which enables us to differentiate between
geoclimatic dynamism and contemporary ecological forces to account
for the pattern of narrow altitudinal ranges and serial species replace-
ment along transects which is characteristic not only of Pronophilini
but of tropical organisms generally (Stevens 1989). A decade has
140
J. Res. Lepid.
passed, but little progress has been made; most of what is being said
about high- Andean organisms remains at the arm- waving stage.
Nonetheless, the importance of the problem of high-altitude tropical
biogeography has been recognized. It has even led to the publication of
not one but two attempts at synthesis, however premature (Whitmore
and Prance 1987, Vuilleumier and Monasterio 1986).
Basically we are asking whether it is possible to find an unmistake-
able footprint of the Quaternary in the geography of the high-altitude
biota, when we have paleoclimatic reconstructions from the immediate
neighborhood. If not, the prospects of validating lowland scenarios
with highland paleoclimatology look especially bleak. But we are also
asking whence comes the highland biota and how old it is. This is a
different and largely independent set of questions, because taxonomi-
cally the high-Andean biota has remarkably little connection with the
tropical lowlands nearby.2 Indeed, the taxonomic decoupling of high
and low altitudes in the Neotropics is such a striking phenomenon that
it seems as if it must bear on the geologic evolution of the region and
have important implications for how we look at the problem of divers-
ity — even if no one has any idea what they might be. Chabot and
Billings (1972) addressed the question of how high-altitude floras
originate, using the geologically young and relatively isolated Sierra
Nevada of California as their example. They found a small floristic
contribution from circumpolar-boreal sources. The principal source
was the adjacent Great Basin, and they were able to identify physio-
logical adaptations to the Great Basin climates which preadapted
species to enter the alpine as it appeared. But the principal geographic
sources of the high-Andean biota are in many cases not in adjacent
climates but in another hemisphere, raising special and very interest-
ing problems (Van der Hammen and Cleef 1987, p. 159). Raven and
Axelrod (1974) attempted to assess probable antiquity of the various
Andean plant families (and make Crucifers, hosts of most Pierini,
quite late — late Miocene-Pliocene — though Tropaeolaceae were j
clearly there earlier and probably evolved there).
Among the Andean butterflies the Pierids have distinct advantages
for biogeographic study. Their diversity is great enough to be interest-
ing and quantifiable, but not so great as to be overwhelming. (Ulti-
2 Descimon, 1986, p. 526 states categorically that “ ... it is clear that the Neotropical and
southern temperate regions contributed little (or nothing) to the oreal butterfly fauna of
the Andes. Its affinities lie instead with the Holarctic realm.” The dangers in such
generalizations have been demonstrated in recent revisionary work on the Argentine
Theclini by Kurt Johnson and his coworkers. Johnson finds that the “Andean 7«cisa/ia”
( Theda culminicola group) are “a sister-group of the diverse and primarily montane
Neotropical . . . groups . . . not an immediate southern relative of cryptically-marked I
Nearctic Incisalia (Johnson in litt 7 April 1989). As a cautionary note I mention that i
Tatochila and the Neotropical genus Ascia show striking electrophoretic similarities, as
well as resemblances in anatomy of the early stages (Shapiro and Geiger, unpublished).
If this relationship is real, however, the direction of evolution has not been established.
28(3):137-238» 1989(91)
141
mately the Andean Theelines, with their high diversity, may be the
most informative butterfly group. However even their generic-level
taxonomy remains fluid, many taxa are undescribed, and distributio-
nal data are fragmentary at best.) The Pierids are medium-sized,
conspicuous, often easily reared, and not prone to cryptic speciation.
They are the most conspicuous element of most high- Andean butterfly
faunas: most of the common, widespread species are Pierids (and they
were commented upon by many early travelers, such as Whymper
(1892)). Their colors and patterns, as noted by Descimon (1986), are
comfortably familiar to biologists of Holarctic origin, the product either
of common ancestry or striking convergence — as usual, posing the
problem of telling the difference. Most of these factors were noted by
the end of the last century. Unsatisfactory as it is, it is not accidental
that the Andean Pierid literature is better (richer) than that of any
other high- Andean butterfly group. Descimon (1986) emphasizes the
Pierids for all these reasons when he attempts a synthesis of high-
Andean butterfly biogeography.
Another ongoing problem of refugial theory has been its failure to
integrate its scenarios for both climate and biogeography in northern
South America with the large, powerful and growing body of paleodi
matic evidence from farther south. Among the best- studied regions on
earth from a palynologicabpaleoclimatic standpoint are Patagonia and
Fuegia. Moreover, some of the information emerging from them is
strongly at variance with the currently conventional wisdom on the
geohistory of the continent, particularly the timing of the most impor-
tant uplift (Clapperton 1983, Mercer and Sutter 1981). The Patagonian
and Fuegian butterfly faunas are taxonomically unbalanced (domin-
ated by Satyrids) but not especially impoverished. They are instructive
in their apparently very close ties to the faunas of the high central
Andes and also in the character and level of their endemisms. They are
highly relevant to the interpretation of the Andean faunas as a whole.
The group of endemic pierine genera including Tatochila and Phulia
has been monographed by Field (1958), Herrera and Field (1959), Field
and Herrera (1977) and Ackery (1975). This body of work has made the
Andean pierines attractive for biogeographic interpretation, as by
Brown (1987) and Descimon (1986). But if the data for Pierini are,
relatively, unusually good, they are still not very good in the absolute
sense. It is possible to extract meaningful inferences from fragmen-
tary data, and perhaps even to arrive at correct scenarios based on
them. But one should not count on it Even for Pierids, almost every
field trip to the high Andes or Patagonia turns up important surprises;
for Lycaenidae or Hesperiidae, such surprises are guaranteed.
This paper draws on both published and unpublished data in an
attempt to pull together and analyze the zoogeography and systema-
tics of the Andean and Patagonian Pierid fauna of the Argentine
Republic. It calls on field trips undertaken over 12 years, comprising
over 18,000 miles of travel on the ground, from the Bolivian border to
142
J. Res. Lepid.
Ushuaia, Tierra del Fuego — as well as on the accumulated experience
of previous butterfly workers in that country. Argentina comprises an
array of biomes ranging from tropical rain forest to cold steppe and
periglacial tundra. Northwest Argentina is at the margin of the great
Andean altiplano , the center of diversity for the miniature Pierids of
the Phulia lineage, and of the dissected jalca or “ paramo ” where
Tatochila achieves its greatest diversity. The altiplano is manifested in
the puna of Salta and Jujuy and the highest elevations of the Sierras
Pampeanas , while the updramo ” can be seen in, for example, the
Cuesta del Obispo and the mid-reaches of the Sierras Pampeanas . If
Pierid diversity does not quite match Arequipa or Cusco, it is still
sufficient to be informative. More importantly, the geography is singu-
larly favorable for tracing the rarefaction of northern (i.e., tropical)
lineages as one proceeds southward into temperate climates, and vice
versa . In so doing, one connects Patagonia and Fuegia to the central
Andes at last, within the taxonomic limits set below.
Of the Andean countries, Argentina may not be the most critical in
terms of its overall potential contribution to historical biogeography,
but it is certainly not the least and it is definitely the easiest. Working
out Argentine Pierid faunistics is logistically feasible. Doing the same
for Peru or Bolivia, which may have more to say about coupling the
highlands and the lowland tropics, is a much more daunting project
(barely begun for Peru by Lamas, 1982): the faunas are much richer,
there is much less of a data base, and field work is much more difficult.
Repeating the task attempted here for Argentina for other countries
and other taxa will be a slow, agonizing process, but there is no
substitute for it. Perhaps the next attempt at a synthesis of high-
altitude tropical biogeography will be shorter on speculation and much
longer on hard data where the butterflies are concerned. Perhaps some
day there will even be an integrated overview of continental paleo-
climates and paleobiogeography for South America, and the butterflies
will be a significant group in its preparation.
Defining the Target Fauna
Breyer (1939) provided the first systematic list of Pieridae from the
entire Argentine Republic. It reveals a large Neotropical, lowland —
lower montane element distributed in the Northeast, the Chaco and
the Yungas. Because the affinities of this fauna, as well as its south-
and westward limitSj are so clear, I have not treated it in this paper. A
study like Larsen’s (1984) for the Arabian Peninsula, which seeks to
identify the relative contributions of different source regions to a
“crossroads” fauna, is particularly interested in such things as the
tropical-temperate transition alluded to here. But the taxa of Phoebis,
Eurema , Zerene, Anteos , Leptophobia, Perrhybris , Pereute , Appias,
Catasticta, Hesperocharis , Dismorphia , Pseudopieris . . .barely if at all
impinge on the objectives of this paper. Of the group of long-range
28(3):137-238, 1989(91)
143
migrants, Eurema deua Dbl. and Colias lesbia Fabr. have been in-
eluded because they penetrate (and breed seasonally) so far into the
middle latitudes, while Phoebis sennae L. and Ascia monuste automate
Burm. have been excluded because they rarely penetrate south of
Buenos Aires. Theochila maenacte Bdv. poses a unique problem. It does
not appear in the analyses because it is confined to lowlands in the
Northeast, south to greater Buenos Aires. But it is discussed in the
text because of its close phylogenetic relationship to Tatochila and its
broader biogeographic import. Teriocolias also poses a problem. It is a
montane insect whose seasonal cycle includes regular up- and down-
slope movements which make it a component of the highland fauna.
On that basis it has been included in the analyses, and a few new data
are reported (cf. Schaefer and Breyer 1941).
After all these somewhat arbitrary decisions, the fauna comprises: all
the Argentine taxa of the genera Phulia , Hypsochila , Tatochila ,
Eroessa , Colias and Teriocolias , plus Mathania leucothea Mol. and E.
deva. The concluding analysis incorporates Chilean data, adding a few
taxa of the above genera plus Infraphulia ilyodes Ureta, Pierphulia
rosea Ureta (two subspecies) and P. isabela Field and Herrera — any of
which might still turn up in the poorly-collected puna of far NW
Argentina.
Phytogeographical Analyses of Argentina: an Overview
Argentina has a venerable tradition of ecology and natural history
and an extensive domestic literature in Spanish, which is largely un-
known outside the country. There have been several comprehensive
attempts to schematize the ecobiogeography of Argentina, one of which
(Shannon 1927) actually emphasized entomological considerations.
The majority, however, are concerned with climate and plant com-
munities (taxonomically defined) or formations (physiognomically).
In 1953(a) Cabrera summarized and attempted to synthesize these. He
updated this work in 1971; what follows is largely abstracted from that
paper (Fig. 2), but with some input from Castellanos and Perez Moreau
(1945), Sarmiento (1975), Hueck and Seibert (1972), Cabrera and
Willink (1980), Davis (1986) and Irwin and Schlinger (1986) with
additional from Gomez Molina and Little (1981) on mountainous
regions and Correa Luna et al. (1977) for the National Parks. Figs. 3=5
are from Madsen et al ., 1980, redrawn and should be used in conjunc-
tion with Cabrera. Given the montane-austral focus of this paper, the
lowland tropics and subtropics are not discussed below.
Cabrera divides Argentina into two regions, five domains, and 13
provinces, as follows; those marked * are discussed in more or less
detail in this paper:
I. Neotropical Region
A. Amazonian Domain
1. Yungas Province*
144
J. Res. Lepid.
2. Parana Province
B. Chaco Domain
3. Chaco Province
4. Espinal Province
5. Prepuna Province*
6. Monte Province*
7. Pampa Province
C. Andean — Patagonian Domain
8. High- Andean Province*
9. Puna Province*
10. Patagonian Province*
II. Antarctic Region
A. Subantarctic Domain
1. Subantarctic Province*
2. Insular Province
B. Antarctic Domain
Many of the habitats described below are illustrated in Plates I -IV.
Y ungas Province. The Yungas occupy a narrow altitudinal belt on the
eastern slopes of the mountains lying in or just barely beyond the
tropics in northern Argentina, from roughly 500 to 2500 m above sea
level. They extend far N into Bolivia, and S only to the north of
Catamarca in the Pampean Sierras (Sierra de Aconquija), but includ-
ing all the ranges in Tucuman. The climate is warm and humid, with
most of the rain falling in summer. Precipitation ranges locally from
700 to 2500 mm or more, with very pronounced orographic effects.
Mean annual temperature also varies greatly: from 14 to 26°C. A great
variety of microclimates thus exists. Winter frosts occasionally occur
at lower elevations and are common higher, where heavy snow may
fall several times a year. In the tropics there is no true winter, and any
snow that falls at these altitudes does not persist.
The predominant vegetation is cloud forest, with trees reaching 30 m
in height and abundant lianas, epiphytes, and a dense understory of
herbs and shrubs. Various transitions from such vegetation to desert,
steppe, shrub-steppe, and temperate deciduous forest occur; ecotones
may be gradual or abrupt, the latter usually reflecting rain shadows or
the altitudes of semi-permanent inversions. Some of the most dramatic
vegetational transects in South America may be made here, parti-
cularly in Tucuman. Cabrera divides the Province into three Districts,
viz.:
(a) Transitional Forest. This is the transition from Yungas to Chaco,
occurring from Pocitos to Oran in northern Salta; on the E flanks of the
Sierras Maiz Gordo, Centinela and Santa Barbara, etc. in Jujuy, the
valley of Lerma and the mountains of Metan and Rosario de la
Frontera again in Salta, the lower slopes of the Sierra de Medina and
the Aconquija-Calchaqules ranges in Tucuman. Temperatures are
28(3):137-238, 1989(91)
145
high but precipitation relatively low (700-1000 mm), and deciduous
trees, especially rather small Leguminosae, predominate.
(b) Mountain Forest. This narrow belt occurs above the preceding,
reaching up to 1300 to 1800 m. It is very rich floristically and faunisti-
cally and commonly includes the southernmost populations of many
lowland-tropical organisms on the continent; this is the so-called selva
Tucumana. Precipitation is from 1500-2500 mm or locally higher;
temperatures within the forest are cool in the intense shade and,
frequently, in cloud and mist. Characteristic of this forest is the horco
molle tree, Blepharocalyx gigantea, but many other conspicuous tree
species occur. Butterfly species richness here is similar to that
observed in the warm, humid far NE of Argentina.
(c) Mountain Woodland. This is the uppermost stratum of the Yungas
and marks the transition from Mountain Forest to the treeless forma-
tions of the high mountains. Some characteristic woody species are
Podocarpus parlatorei , alder ( Alnus jorullensis var. spachii ), elder-
berry ( Sambucus peruviana ), Schinus gracilipes, etc. with local out-
liers of quenoa ( Polylepis australis) generating a spotty second tree-
line above 2500 m in some locations (Fernandez 1970). The woody flora
includes both Holarctic and Subantarctic elements. The herbaceous
flora is very rich and includes many showy wildflowers, largely of
Holarctic origin. The higher meadows — as between Tafi del Valle and
Abra Infiernillo — include many perennial bunchgrasses. Above
2000 m trees are often confined to creek bottoms. Meadow and brush
vegetation may occur much lower, however, as on the Cumbre de San
Javier near San Miguel de Tucuman, and it is unclear whether this
reflects the well-known “telescope effect” found in tropical mountains
or the medium- to long-term consequences of land use. Similar vegeta-
tion occurs at the highest elevations of the Sierras of Cordoba, repre-
senting a far-southern outlier of the Yungas.
Prepuna Province. This is the vegetation of the dry slopes and canyons
from Jujuy to La Rioja, mainly between 2000 and 3400 m, but locally
down to 1000 m in certain microclimates. It intervenes between the
uppermost Yungas and the true Puna , but also between the Chaco and
the Puna or the Monte and the Puna. The climate is warm and dry,
with rain exclusively in summer thunderstorms. The vegetation is
xerophytic shrub-steppe with aspect dominance by columnar cacti, of
which Trichocereus pasacana and T. terscheckii are typical in the N
and S respectively. Terrestrial Bromeliads and shrubby Leguminosae
abound. Common genera include Adesmia , Azorella , Junellia, Muli-
num, Nassauvia , Parastrephia and Senecio. Riparian woodland occurs
in canyon bottoms, with molle ( Schinus areira ) and chilca ( Baccharis
salicifolia) . Because of its complex geography, prepuna has little dis-
tinctive butterfly fauna and indeed is often rather depauperate, though
it occasionally serves as a corridor to bring higher-altitude species
146
J. Res. Lepid.
lower and vice versa , in butterflies as in plants. A major reference is
Hunziker (1952).
Monte Province. The Monte begins in the valley of the Rio Santa Marfa
in the Valles Calchaqufes, extending S through the W of Tucuman and
La Rioja, through Mendoza and San Juan and thence into Neuquen
and the E of Rfo Negro and in dilute form even to far NE Chubut.
Precipitation varies from 80-250 mm (locally higher), mean tempera-
ture from 13-17. 5°C, with pronounced climatic gradients both E W
and N-S. The vegetation is desertic and diverse, but dominated
throughout by creosote bush ( Larrea spp.) and mesquite ( Prosopis spp.)
except in alkali sinks, where Distichlis and Chenopods predominate.
Common genera include Bougainvillea , Bulnesia, Caesalpinia, Cassia,
Cercidium, Mimosa , Trichocereus and Zuccagnia and in gullies Acacia
and Celtis. The Monte corresponds ecologically to the Sonoran Desert of
North America, and like it segues almost imperceptibly into adjacent
colder formations. Large-scale comparisons of community organization
and function were made between the Monte and the Sonoran Desert as
part of the International Biological Programme and are summarized in
Orians and Solbrig, 1977.
Pampa Province. This is the bunchgrass prairie ofArgentina, much of
it now degraded and invaded by exotic weeds. It occurs at low to
moderate elevation in flat to rolling country in the E, roughly between
31-39°S. There are pronounced climatic gradients, particularly in
precipitation, from 1100 mm in the NE (“subhumid pampa ”) to 600 mm
in the SW (“subarid pampa ”). The various floristic subdivisions are
unimportant for this paper. The butterfly fauna has surprisingly low
endemism.
The Andean — Patagonian Domain : General Considerations. The fol-
lowing translation from Cabrera (1971, p. 29 ff.) captures the character
of the Domain well: “In the Argentine Republic this Domain extends
all along the extreme W of the country, covering the puna and the
Andean cordillera from the border with Bolivia to the S of Mendoza.
There it begins to reach out to the E on the mesetas and the Patago-
nian ranges, reaching the Atlantic in Chubut and Santa Cruz. Its
climate is cold and dry, with frosts almost all year and snow in winter.
The Andean-Patagonian Domain is characterized by the scarcity of
endemic families (of plants), only the Malesherbaceae and Nolanaceae
being exclusive to it, and in turn a great richness of endemic genera of
the most diverse groups. The families of greatest importance for their
richness in genera and species are the Composites, Grasses, Verbena-
ceae, Solanaceae, Cruciferae, etc. The Legumes are represented by few
genera, principally Papilionaceous, but at times with numerous spe-
cies, as in Adesmia and Astragalus. The Zygophyllaceae and Mimosoid
Legumes, so abundant in the Chaco Domain, are almost completely
lacking. The dominant vegetation is shrub- or herbaceous steppe, with
28(3):137-238, 1989(91)
147
extreme forms of adaptation to wind and drought. . .Where springs
exist or water accumulates, inundated meadows form, called vegas in
the cordillera and mollifies in Patagonia.” Cabrera gives a detailed
subdivision of the Domain.
High- Andean Province . Again from Cabrera 1971, p. 30: “From the
border with Bolivia to Tierra del Fuego. In Jujuy and Salta found
approximately above 4400 m; in Mendoza above 3000, in Neuquen and
Rio Negro above 1600 m and in Tierra del Fuego above 500 m. . . .
Immature, rocky or sandy soils, with a high-montane climate, cold and
dry (except in the S), with snow or hail at any time of year. The mean
temperature is low, below 8°C (— 1.5°C at Cristo Redentor, Mendoza, at
3829 m). The vegetation is very poor and is formed by grassy steppe or
steppe of cushion chamaephytes ...” Three Districts are recognized,
which are very important for understanding the butterfly fauna:
Quick ua District. This is the true high-Andean vegetation, occupying
the Eastern, or Royal Cordillera S to the N of San Juan from 4300-
5600 m. It is thus tropical and subtropical. Precipitation' is higher in
the N and E, where the District contacts the Yungas and Chaco, than
in the S and W where it contacts the Monte. It is strongly seasonal,
concentrated in afternoon thunderstorms during the “Bolivian Winter”
from November to February caused by the S-ward migration of the
Intertropical Convergence Zone in response to heating in the continen-
tal interior. The most common climax formations recognized by Cab-
rera are: Festuca orthophylla — F. chrysophylla — Poa gymnantha
steppe, with such associated spp. as Stipa and Deyeuxia , Baccharis
incanum , Senecio graveolens , Werneria poposa , Adesmia caespitosa
and patancana, and the remarkable yareta, Azorella compacta (Umbel-
lifer ae), a cushion plant formed of thousands of tiny terminal rosettes;
also smaller, delicate wildflowers including Perezia ciliosa, Silene
friesii , Cajophora coronata , Calceolaria glacialis, Valeriana spathu -
lata , Nototriche anthemidi folia, etc.; steppe of vizcacha grass ( Stipa
frigida ) with shrubs such as Artemisia copa and Senecio viridis ; Bitter
Coiron ( Stipa chrysophylla ) • vizcacha grass steppe associated with
the dwarf Leguminous shrubs Adesmia glanduligera and A. nanolig -
nea; and others. Some additional characteristic or indicator species
are: among the cushion plants, Oxalis compacta , Senecio algens and
Pycnophyllum molle ; very high-altitude Crucifers ( Aschersoniodoxa
mandoniana , Parodiodoxa chionophylla , both reaching above 5000 m);
sedges of inundated vegas ( Oxychloe andina , Carex incurva, Scirpus
atacamensis ), often with grasses ( Deyeuxia spp., Festuca scirpi folia);
the dwarf rush Distichia muscoides , and showy wildflowers including
Gentiana prostrata , Gentianella punensis , Calandrinia acaulis , Wer-
neria pygrnaea. and many Astragalus spp. This is a flora closely allied
to that of the highlands of southern and central Peru (Weberbauer
1945).
148
J. Res. Lepid.
Cuyo District. This is the vegetation of the high Andes of San Juan,
Mendoza and the N of Neuquen, from 4500 to 2200 m descending
southward. The winters are very snowy; total preciptation is highly
variable but always includes summer thunderstorms. The District is
entirely outside the tropics and beyond the influence of the “Bolivian
Winter,” being dominated instead by the interplay of tropical and
subantarctic air masses with traveling storms and fronts. Coiron
( Stipa spp.) is a climax aspect dominant in much of the District, often
associated with perennial Festuca and Poa. The dominant species of
Stipa varies geographically. On steep slopes shrubs are common,
including Adesmia pinifolia, Ephedra andina, Berheris empetrifolia,
Senecio uspallatensis , Mulinum ovalleanum, etc. Typical herbaceous
species include Menonvillea cuneata, Nassauvia lagascae , Tropaeolum
polyphyllum , and, in vegas , the dwarf rush Andesia hisexualis , Plant -
ago barhata, and Senecio breviscapus. Streams are often bordered by
Cardamine nivalis and there may be a turf of Hordeum secalinum or
Agrostis glabra with Hypsela oligophylla and Werneria pygmaea. The
classic description is by Hauman (1919). A useful modern treatment of
the flora is Wingenroth and Suarez (1983).
There is a remarkably rapid floristic and vegetational gradient from
N to S in the Cuyo. The northern part of the Cuyo forms a transition
from the tropical climates to the N, dominated by rainfall seasonality
but with no true winter, to the strongly temperate-seasonal climates in
Mendoza with a pronounced winter snow pack. By the latitude of Las
Lenas and Cerro Sosneado the climates and aspect have become
decidedly more Patagonian. These gradients are reflected from the
high desert to the high alpine in the presence of species-level endemics
among flowering plants and butterflies. Because of its easy accessibil-
ity, the vicinity of the Paso Bermejo is often taken as “typical” of the
montane Cuyo , thereby underestimating the high diversity within the
District.
Austral District. From C and SW Neuquen southward at ever-
decreasing elevation, the high-Andean flora descends and mixes with
Gondwanaland (subantarctic) elements, especially in the more humid
areas. Steppes are often dominated by Poa obvallata and Festuca
weberbaueri or F. monticola , the marshy meadows by Deyeuxia , De-
schampsia, and Poa , and especially in the far S, crowberry ( Fmpetrum
rubrum ) becomes conspicuous or dominant on poor (sandy, sterile, and
boggy) sites. The communities of Tierra del Fuego, which partake of
this District but also of the Subantarctic Domain, are well-described in
Moore (1983).
Puna Province. The Argentine puna occurs from 3400 to 4500 m, from
the Bolivian border to NE Mendoza, where it can be found in dilute
form to 2000 m. The climate is dry and cold, with mean annual
temperature of 7.5-9. 9°C and annual precipitation from 324 mm at La
28(3): 137-238, 1989(91)
149
Quiaca to 103 mm at San Antonio de los Cobres, to nearly zero at the
Chilean border.
The puna is a xerophytic high-altitude shrub-steppe dominated by
tola ( Parastrephia lepidophylla ) and river tola (P. phylicaeformis )
where ground- water levels are fairly high. Many other species occur
( totilla , Fabiana densa; chijua , Psila boliviensis; anagua , Adesmia
horridiuscula ; etc.) along with dwarf cacti ( Opuntia soerensii , Oreocer-
eus trollii , etc.). Some other characteristic species are Pennisetum
chilense , Scirpus atacamensis , Juncus depauperaius , Plantago tubulo-
sa, Hypsela oligophylla , Festuca scirpifolia, Bouteloua simplex ,
Muhlenhergia fastigiata , Trifolium amabile , Astragalus bustillosi ,
Ipomoea minuta , etc. Farther W, in Chile, extensive flat bogs occur
within the puna. The classic description is Cabrera’s (1958) mono-
graph.
Comment on the High- Altitude Vegetation in Salta, Jujuy and Tucumdn.
— The Argentine NW is vegeta tionally very complex, as Figs. 6 and 7
demonstrate. The upper reaches of the Yungas grade into prepuna,
puna , and Quichua high Andean vegetation and the formation is
exquisitely dependent on slope, exposure and in some cases, substrate.
As noted below, many of the high-altitude butterfly species appear to
have very specific requirements within this complex mosaic, but may
be widespread where their particular microclimates occur. This is
characteristic of tropical organisms, while temperate species tend to be
more broadly distributed both ecologically and geographically (Stevens
1989). A notable exception is Tatochila mercedis macrodice , which
seems to transcend formation or association boundaries, occurring
nearly everywhere at high altitude.
Patagonian Province. The first hints of Patagonian vegetation appear
in the precordillera of Mendoza. The transition from the Monte and
Cuyo to Patagonia occurs gradually from Chos Malal, Neuquen S
through Loncopue to Zapala. In the N of this transition zone the
mallines are nearly or quite pure Patagonian while the nearby hills
have an increasing proportion of Patagonian species with both latitude
and altitude.
The traditional “political” boundary of Patagonia is the Rio Negro,
but Patagonian floristic elements occur N in the W of the region, while
elements of the Monte reach even beyond Comodoro Rivadavia on the
Atlantic shore. The precise location of the boundary of the Patagonian
Province is controversial (cf. Soriano 1949, 1956, Ragonese and Pico
nini 1969). Soils are sandy or gravelly — much of the Province is
covered with a thin pavement of gravel washed out from the glaciers
and the climates are cold and dry with winter snow, frosts much of the
year, and very persistent strong winds which are perhaps the most
characteristic feature. The temperature varies from an annual mean of
13.4° at Chos Malal to about 5° at Rio Grande, NE Tierra del Fuego;
150
J. Res. Lepid.
precipitation ranges from 100-270 mm over the Patagonian steppe,
but rises very rapidly to the W as the continental divide (= border with
Chile) is approached. The dominant plant communities are shrub-
steppe and bunchgrass steppe. Cabrera recognizes six Districts (but see
Boelcke 1957):
Payunia District. This is the Monte- Patagonia ecotone in the far N,
characterized by solupe C Ephedra ochreata ), the composite Chuquiraga
rosulata, Lycium chilense, Grindelia chiloensis, Junellia seriphioides
and, locally in cool sites, neneo ( Mulinum spinosum ), the most distinc-
tively Patagonian element. The unusual Umbellifer Hydrocotyle , of
subtropical origin, occurs along slow streams — its only occurrence in
the Province.
Western District. This is a narrow fringe of shrub-steppe, from north-
ern Neuquen to NE Santa Cruz, dominated by Mulinum spinosum ,
Trevoa patagonica , Colliguaya integerrima and Nassauvia axillaris ,
with local coironales (bunch-grass prairies) of Stipa humilis, neaei and
speciosa often containing Poa huecu , Bromus macranthus , Festuca
argentina , etc., and more or less saline seeps or marshes with Distichlis
scoparia and D. spicata and bulrush marshes ( Scirpus californicus) .
Pampas-grass ( Cortaderia spp.) reaches its southern limits here, often
in overgrazed bottomlands just upslope a few cm from mallines of
Distichlis.
Central District. This is the most arid part of Patagonia, from the
center of Rio Negro through most of Santa Cruz. The predominant
steppe species are quilenbai ( Chuquiraga avellanedae) , colapiche ( Nas-
sauvia glomerulosa) , and coiron amargo ( Stipa humilis , neaei or spe-
ciosa); associates include Ameghinoa patagonica , Nardophyllum obtu-
sifolium , and Brachyclados caespitosus. In the far S, quilenbai drops
out and is replaced by mata negra, Junellia tridens (Verbenaceae, not
to be confused with other shrubs having the same common name in
other regions). Saline marshes are dominated by Atriplex lampa. In
moist gullies larger shrubs such as Anarthrophyllum rigidum , calafate
( Berberis cuneata ), Senecio filaginoides and Lycium chilense occur.
San Jorge District. This is a “warm pocket” along the Gulf of San
Jorge in the vicinity of Comodoro Rivadavia, where Monte elements
including creosote bush can be found. Characteristic dominants are
Trevoa patagonica , Colliguaya integerrima , Stipa humilis , Mulinum
spinosum , Adesmia campestris, Anarthrophyllum rigidum , Festuca
pallescens and argentina , etc.
Subandean District. This is the lower fringe of the Patagonian Andes
proper, widening out S of 51° to reach the Atlantic. Annual precipita-
tion is 200-350 mm; the climate is humid at least part of the year,
even the dry season is rather cloudy due to high-altitude moisture
transported over the Andes by the prevailing westerlies and formed
28(3):137-238, 1989(91)
151
into extensive wave altocumulus in their lee, and the soils are richer
and more mature than in areas further NE. The climax vegetation is
Festuca pallescens steppe, with a very long list of associates including
Poa ligularis , Bromus macranthus , Elymus patagonicus , Calceolaria
polyrhiza , Acaena pinnatifida , Viola maculata , Lepidophyllum cupres -
siforme , etc.
Fuegian District . This is the NE of Tierra del Fuego, extending to the
transition from steppe to Nothofagus forest SW of Rio Grande. Festuca
gracillima is the dominant steppe grass. Many characteristic and
showy wildflowers occur, of which Primula magellanica and Oxalis
enneaphylla and fueguensis are characteristic.
Subantarctic Domain and Province . Characterized by the dominance
of south-end-of-the-world (Gondwanaland) taxa of great antiquity,
adapted to cool and moist, temperate climates — Nothofagus , Dacry
dium, Fitzroya , Myzodendraceae, Desfontainaceae, Tetrachondraceae,
Donatiaceae, etc. This Domain is mainly W of the Andean crest in
Chile. Four distinct Districts are recognized by Cabrera; their relation-
ships are clearer when the vegetation map of Chile is consulted along
with that of Argentina.
Pehuenia. The relict forests of Pehuen, Araucaria araucaria, from
Volcan Copahue to Lake Lolog in western Neuquen, from 900-1800 m,
associated with Nothofagus pumilio , Chusquea culeou , Berberis buxi-
folia , Pernettya mucronata , Maytenus disticha, Ribes magellanica ,
Escallonia virgata , Nardophyllum obtusifolium , Cortaderia pilosa ,
Chlorea alpina , Acaena pinnatifida , etc. — marking the N limits of
many species, and diluting gradually to the E in the San Marl in-
Jonin -Alumine region until only occasional Araucaria are superim-
posed on bunchgrass steppe; such sites are often called “Primeros
Pinos.”3
Deciduous Forest District . The forests of Nothofagus pumilio and
antarctica and of Austrocedrus chilensis extend to tree-line above and
steppe below, with a complex structure of communities or associations.
In the far N Nothofagus procera and obliqua also occur. The coligiie
bamboo, Chusquea culeou , Is locally very abundant, as on Cerro Gated-
ral near Rariloche. Wildflowers are numerous and include amancay
3 The easternmost stand of pehuenes is located on the Espinazo del Zorro along Highway
46, SW of the Laguna Blanca National Park in Neuquen. It very clearly indicates a
much more mesic climate eastward within the past 1000 yr or so. The genus Araucaria
was formerly much more widespread, both globally and in South America, and has been
in decline since the early Tertiary. The Olsacher Museum of Geology in Zapala,
Neuquen has an excellent collection of fossil Araucaria material from the region. The
complete lack of a distinctive butterfly fauna associated with the pehuen vegetation is
striking.
152
J. Res. Lepid.
(. Alstroemeria aurantiaca), Codonorchis lessonii, Mutisia spinosa and
decurrens, etc. Like the preceding, this District is defined by relicts
whose distributions reflect very local, topographically-mediated mic-
roclimates E of the crest.
Valdivian District. This is the true cool-temperate Tertiary rainforest
of S Chile, floristically extremely rich but dilute on the Argentine side
of the border where it penetrates only in the far W of the Lanin,
Nahuel Huapi, and Los Alerces National Parks. Precipitation in some
places reaches 4000 mm/yr. It has two characteristic butterflies: the
Pierid Eroessa chiliensis and the Hesperiid Argopteron aureipennis
Butl. See Ringuelet (1955).
Magellanic District. The southern beech forests ( Nothofagus ) and
associated Sphagnum- bog ( turhal ) habitats of Tierra del Fuego (Moore
1983).
Comment on the Vegetation of Patagonia and Fuegia. These parts of
Argentina have engendered an immense literature. Besides Cabrera,
an excellent overview in English, with comparisons to North America,
is to be found in Beetle (1943). Dimitri (1962) reviews the flora, and
(1972) sets it in a regional and physiological context.
Cabrera’s classification and mapping are necessarily somewhat typo-
logical. Even so, it is evident that the Districts are not very well-
defined and the lines demarcating them are at best approximations of
statistical changes in community composition. The intimate interdigi-
tation of subantarctic and Patagonian elements makes line-drawing in
the far S quite arbitrary. My experience suggests that the distribution
of Satyridae in Patagonia may be studied profitably with respect to
plant community patterns. The Patagonian and Fuegian Satyrids have
been monographed by Heimlich (1972).
Among numerous accounts of Patagonia, Lista (1896a, b) and Willis
(1914) are particularly instructive in enabling us to assess the amount
of vegetational change due to the activities of the white man. It is quite
evident, especially from Willis, that the extent of Nothofagus forest
has been greatly reduced by cutting, burning and conversion to pastur-
age. Given that many evolutionarily interesting and important pheno-
mena are today occurring in zones where such conversion occurred in
the last century, it is important to keep this in mind. The W of
Neuquen and Rio Negro and the vicinity of the Gulf of San Jorge rank
with the W of Salta and the Sierras Pampeanas as regions of special
concern for butterflies; the mainland-Fuegian transition, in particular
the littoral from Comodoro to Rio Grande, will also require further
attention. Very rapid urban growth in the far S since 1970 is already
impacting butterfly habitats.
28(3):137-238, 1989(91)
153
Systematic Treatment
Collection abbreviations are identified in the Acknowledgments
section.
Genus Theochila Field
Theochila maenacte Boisuval
Theochila is probably the sister-genus of the non-Crucifer-feeding
part of the large genus Tatochila , which will almost certainly be
divided in two or three genera once sufficient species have been reared.
Theochila is defined by a variety of odd autapomorphies (Field 1958).
The single species maenacte is divided into two allopatric subspecies.
Its distribution (Brown 1987, Fig. 4.18) is part of a repeating pattern of
faunal and floral connections between the Andes and Serra do Mar/SE
Brazil, which “leaves no doubt that links of suitable vegetation and
humid temperate climate existed across northern Argentina in the
distant past” (Brown 1987, p. 95; O. Mielke, pers. comm.).
The nominate subspecies T. m. maenacte differs from the Brazilian T.
m. itatiayae Foetterle primarily in slightly smaller size and a drastic
reduction in the dark wing-pattern in the male, producing an effect not
unlike Pieris rapae L. Herrera and Field (1959) had little material
available and apparently did not recognize that T. m. maenacte is
seasonally diphenic, with a winter form nearly indistinguishable from
itatiayae and a very pale summer form on which their redescription
was based. The BM contains a very extreme winter male, unfortunate-
ly without date, labeled “Buenos Ayres, Elwes 1920.” It also has a long
series of more or less typical summer specimens labeled “Buenos Aires
(Belgrano), 15. XII. 1889.” The very pronounced seasonal polyphenism
of the male and lack thereof in the female (in which the denser thoracic
and abdominal pelage, and richer black color in the wing pattern are
the only manifestations) parallels the situation in Tatochila uanuolx-
emii = T. mercedis vanvolxemii Capronnier. It would be of great
interest to determine whether photoperiod or temperature or both
influence the polyphenism.
Theochila m. maenacte was formerly common in riparian and marsh
habitats in and around Buenos Aires (Riachuelo; Avenida de los
Italianos — Zona Portuaria, etc.) but has disappeared or diminished in
many sites in the past decade. Until slum clearance and construction of
the central bus station destroyed its habitat, it was common behind the
Retiro railroad station several blocks from the center of the city, as late
as the mid-1970s. It was not found in the “Vida Silvestre” preserve in
the Zona Portuaria in XI. -XII. 1989. It is still common in marshy sites
in Quilmes and south to the vicinity of La Plata, which seems to be its
southern limit. The Museo de La Plata contains much local material,
e.g. series from La Plata, 27. III. 1927; 3J, 2. XII. 1928; an extreme
154
J. Res. Lepid.
winter form J, 28. X. 1928, and a dozen from the northern suburb of
Punta Lara, II. 1928, leg. R. Maldonado. It is also still locally common
in Tigre and the Parana Delta north of the capital, and should be
looked for in the marshes near Ezeiza (the old “Belgrano” locality?)
Genus Tatochila Butler
As noted under Theochila , this genus appears to be polyphyletic and
will probably need to be divided. At present it is the largest Pierine
genus on the continent. Since the lines of the division are not yet clear,
I am opting for taxonomic conservatism by retaining all of Herrera and
Field’s (1959) species-group designations within Tatochila , whose
type-species is autodice Hiibner.
Tatochila theodice theodice Boisduval (Fig. 8A, D).
The N and S limits of the nominate subspecies have been poorly
defined in Argentina. Herrera and Field (1959, p. 478) had little
Argentine material available.
3cf Loncopue, Neuquen, 8.XI.1988 ( AMS)
lef Catan Lil, Neuquen, 840 m, 16.1.1977 (MG)
The Loncopue record (38°04'S, 70°37'W) probably defines the N limit
of both the subspecies and species as it is unrecorded at Chos Malal,
which I have collected very thoroughly. The Catan Lil record may
represent the farthest E and downslope penetration of the high desert
in river bottoms, to which this mesic species is largely restricted in the
N of its range. It is clearly resident at Loncopue, and the specimens are
of the early spring (post-diapause) phenotype.
The S limit is defined by the transition to ssp. gymnodice Staudinger, |
which is accomplished in gradual, clinal fashion, as demonstrated by
the following material:
2c? 3J Lago Argentino, Peninsula Magallanes, Santa Cruz, loc. #28,
11.1.1979 (DE)
6c? Tecka, Corcovado, Chubut, 750 m, loc. #47, 17.11.1979 (DE)
lcf La Esperanza, 130 km NW Rio Gallegos, Santa Cruz, loc. #30,
15.1.1979 (DE)
Herrera and Field had no material between the Nahuel Huapl Na-
tional Park, Rio Negro (nominate theodice ) and Rio Tunel, Santa Cruz
( gymnodice ), creating a false impression of disjunction. In the Lago
Argentino series the females are somewhat more gymnodice-\ike than
the males. See Fig. 8B, E.
Tatochila theodice staudingeri Field
Herrera and Field (1959) divide the Fuegian theodice into two sub-
species, recognizing gymnodice from Porvenir, Magellanes (Chile) and
staudingeri from Puerto Harberton (the type locality) and Ushuaia,
both in Argentine Tierra del Fuego, as well as from Isla Navarino,
28(3):137-238, 1989(91)
155
Magallanes. Again, when sufficient material is assembled, the dis-
tinctness of these subspecies disappears.
2 c? 2 J Cabo Penas, Depto. Rio Grande, T. del F., 17.XII.1983 (ML)
56c? 30J Rio Grande, T. del F., 25.XI.1988 (AMS) (Figs. 8C, F)
2c? Estancia Marla Cristina, Route 3, T. del F., 27. XL 1988 (AMS)
lC? 19 Base of Monte Susana, Parque Nacional, T. del F., 18.1.1979
(AMS)
A cline exists from NE to SW across Isla Grande de la Tierra del
Fuego, corresponding to both precipitation and' vegetation gradients.
Staudingeri phenotypes are increasingly frequent to the SW, but occur
even at Rio Grande in the extreme NE where populations are immense
and variation very pronounced. Herrera and Field fail to note that
Fuegian females are dimorphic in ground color: yellow, or white like
the males. This is a genetic trait, as established by rearing.
Nominate theodice from Rio Negro and gymnodice/ staudingeri from
Rio Grande, T. del F., have been reared ex ovo and descriptions are in
preparation. There are subspecific differences in the early stages.
Tatochila autodice Hiibner (including T. blanchardii Butler)
Shapiro (1986a) demonstrated that T. autodice and T. blanchardii
blanchardii intergrade in NW Patagonia, from Chos Malal, far N
Neuquen, to C Chubut, mainly in the ecotone from the forested
Patagonian Andes to bunchgrass — shrub steppe. Detailed records are
presented there. The farthest S record for apparently pure autodice is
Puerto Deseado, Santa Cruz, lcf, 16.1.1967 (A. Willink) (ML). In 1989
I found T. autodice common in eastern Patagonia, viz. 19 Trelew,
Chubut, 7. XII; 2c? Las Plumas, Chubut, 7. XII; 3c? Parada Uzcudun,
Chubut, 7. XII; 19 El Tordillo, Chubut, 8. XII; 19 Pampa del Castillo,
Chubut, ll.XII; 1c? 29 Caleta Olivia, Santa Cruz, 9. XII; and 19 Fitz
Roy, Santa Cruz, 9. XII. Both Caleta Olivia 9 have an unusual yellow
ground color but are otherwise typical. None shows any trace of
blanchardii influence. Three of these localities represent tiny pockets
of weedy Crucifers in extensive shrub-steppe, demonstrating the high
dispersal capability of this species.
The farthest SE that blanchardii influence has been recognized is El
Trebol, Chubut, 19, 12.11.1967 (A. Willink) (ML). Some intermediate
phenotypes from Rio Negro and Chubut are shown in Fig. 9. As noted
in Shapiro (1986), the intergrading populations in the Lake District
are unusually sparse. This is consistent with the now widely-
recognized phenomenon of the “hybrid sink” (Barton and Hewitt 1985),
although the intergrading population as Esquel is much larger; it is
also in a drier climate and has a stronger autodice component. It
should be noted that the usual host plants of the Lake District popula-
tions are native Tropaeolum polyphyllum rather than introduced
weedy Crucifers, based on censuses done in 1988. At least some of the
area occupied by these populations is mapped as recently (turn-of-the-
156
J. Res. Lepid.
century) deforested in Willis (1914).
The La Plata collection contains numerous apparently typical blan~
chardii from northwestern Patagonia, viz.: lcf Lago Lacar, Pucara,
Neuquen, 1.1958; 1J Lago Hermoso, Neuquen, 1.1958; 1$ Ruca Malen,
Neuquen, undated; 1 J Isla Victoria, Lago Nahuel Huapi, Rio Negro,
1.1960; 1J Quila Quina, Lago Lacar, Neuquen, 1.1958; 2J San Martin
de los Andes, Neuquen, 1.1958; and lj Lago Curahue, 950 m, Neu-
quen, 1.1958. There is also one obvious c? intergrade, labeled “Rio
Negro,” 1.1936 (#2646).
On the basis of this rather extensive if mosaic intergradation I
propose treating Herrera and Field’s (1959) “ autodice species group
(Group B)” as a polytypic species, viz.:
Tatochila autodice Hiibner 1818
Tatochila autodice blanchardii Butler 1881, new combination
Tatochila autodice ernestae Herrera 1954, new combination
Because nominate autodice and ernestae may be parapatric and altitu-
dinally stratified in the Bolivian yungas , autodice may be a “ring
species” (circular overlap, Mayr 1963, p. 507 ff.).
Tatochila mercedis Eschscholtz (including T. sterodice
Staudinger, T. fueguensis Field, T. macrodice Staudinger, T.
arctodice Staudinger, and T. vanvolxemii Capronnier)
This is the umicrodice species group (Group C)” of Herrera and Field
(1959). The incorrect use of the name microdice was corrected by
Ackery (1975). Thereafter the name sterodice was used for a polytypic
species embracing the taxa fueguensis , macrodice and arctodice —
mercedis and vanvolxemii being treated as separate species. However,
all the taxa of this group, from Colombia to northern Fuegia at least,
are interfertile in the laboratory and intergrade through fairly
sharply-defined hybrid zones wherever they come into contact afield
(Shapiro 1979, 1984, 1986b and unpublished). The oldest name in the
group is mercedis and all the other taxa may be considered subspecies
of it. This action was initiated by Lamas and Perez (1983) in listing
macrodice as a subspecies of mercedis. “Group C,” the “ sterodice
species-group” of my earlier papers, thus becomes:
Tatochila mercedis Eschscholtz 1821
Tatochila mercedis macrodice Staudinger 1898
Tatochila mercedis arctodice Staudinger 1898, new combination
Tatochila mercedis vanvolxemii Capronnier 1874, new combination
Tatochila mercedis sterodice Staudinger 1898, new combination
Tatochila mercedis fueguensis Field 1959, new combination
Because of intrinsic problems with the subspecies as a taxonomic
category, and because one does not expect to find clear-cut taxonomic
situations in groups as evolutionarily active as this one, a certain
degree of arbitrariness seems inevitable in ranking the taxa. Cracraft
28(3): 137-238, 1989(91)
157
(1989), writing from one cladistic standpoint, denounces the ontologi-
cal vacuity 'of polytypic species, a legacy of the Biological Species
Concept, which he rejects. Nonetheless, the reproductive, genetic and
geographic data available all appear to argue for polytypic species
status for this group, despite the morphological distinctness of some of
its members; I accept the primacy of data over ideology.
The following, hitherto-unpublished data amplify the known Argen-
tine distributions of these taxa and fill in gaps on previously -published
maps (Figs. 10A, 11 A).
Tatochila mercedis macrodice
Reported occurrences of this taxon in Argentina fall into three geo-
graphic regions; the puna , the Quichua District, and the Cuyo District.
(i) The puna of Jujuy and Salta
R. Eisele states {in litt 26.1.1978): “. . .according to Hayward. . .has
been found as far S as Mendoza. In the last few years I have got a
number of good series of this in Jujuy and Salta. All come from
altitudes from 2450 to 4000 m and most common above 3000 m, at
which altitude it is the most common Tatochila ”
Some specific records:
2c? Abra de Fives, Jujuy, 3900 m, 3.II.1969 (ML)
18c? 132 Abra Pampa, Jujuy, 7. XL 1984 (AMS)
l ie? Esquinas Blancas, Jujuy, 7 II 1984 (AMS)
2c? 31 km N Humahuaca, top Azul Pampa, Jujuy, 121.1978 (RE)
3d Abra Azul Pampa, Jujuy, 7,11 1984 (AMS)
6c? Altos de Abra Munano, Salta, 4165-4780 m, 21.1.1983 (AMS)
(ii) The Quichua District
This entity occurs in the puna-jalca alpine vegetation above the
yungas . It is recorded in two geographic subdivisions:
(a) Salta highlands
3C? Abra Molina, Salta, 221.1986 (AMS), lef 22 28.XI.1989 (AMS)
5c? Cerro Zapallar, Salta, 22.L1986 (AMS)
Id1 Valle Encantado, Salta, 3550 m, 14.XII.1976 (ML)
(b) The Sierras Pampeanas (Cumbres Calchaquies, Sierra de
Aconquija)
12 Cerro de la Mina, Depto. Tali, Tucuman, IV. 1933 (K. Hayward)
(ML)
Id “Aconquija”, 1 LIU 1917 (P. Jorgensen) (BM)
12 Esquina Grande, Catamarca, 1640 m, 30.XI.1915 (BM)
22 “Tucuman, 2000 m,” III. 1905 (J. Steinbach) (BM)
2d 12 Tafi del Valle, Tucuman, 2150 m, 30.XL1977 (AMS)
3d Abra Infiernillo, Tucuman, 20.1.1986 (AMS), 2d 26.XI.1989
(AMS)
Specimens from the Sierras Pampeanas appear to average slightly
smaller and more heavily marked than the others. Except for a
158
J. Res. Lepid.
genetically-determined ground-color polymorphism (white vs. yellow,
white dominant --- AMS, unpublished data) found in most populations,
macrodice is extremely constant over its entire range in Peru, Bolivia,
Chile and Argentina.
Like many butterflies of the puna and upper yungas , T. m. macrodice
appears to migrate altitudinally with the season. It is present at lower
altitudes in dry season than at other times, and disappears entirely
from the vicinity of Tafi del Valle in January. However, as the 1989
data show, it emerges immediately in the highlands of both Salta and
Tucuman with the onset of the rainy season in November, flying even
before the vegetation has greened up. It thus must be 2- or more likely
3-brooded in the highlands, passing the dry season as a pupa, but no
trace of diapause exists under laboratory conditions.
(iii) The Cuyo District
I have not found the documentation for the Hayward citation men-
tioned by Eisele. He did not include Mendoza for this entity in his
(1950) Catalogo Sinonimico . . .but did in the posthumously-published
1973 catalog (p. 114). This reference also lists San Luis, a lowland
Province whose highest point (in a pre- Andean range) is under 2000 m
and which is unlikely to have any macrodice habitat at all. This
suggests some kind of confusion existing in Hayward’s mind late in his
life concerning the identity of macrodice , and it is not hard to find
evidence for this in museums. Herrera and Field (1959) record a
specimen of macrodice from “Funes, Mendoza, January” which is the
only precise locality ever published from Mendoza, but I have been
unable to trace either the specimen or the locality. There is no “Funes”
(or “Dean Funes”) in Mendoza in available maps (Automovil Club
Argentine, sections 3 and 5, Carta Turlstica, 1969-72) or gazzetteers
(US Department of Interior 1968).
ML contains a small female vanvolxemii from Potrerillos, Mza.,
1500 m, 1947 (Hayward and Willink), misidentified as macrodice. La
Plata has much more. Under macrodice (box 297) are 7cf 4J from
Potrerillos, X.1951. In addition, likewise identified as macrodice by
Hayward, one finds a cf from Los Corrales, La Rioja, 1935 (lot #2643)
(box 312) and a female without locality, “Junio 1925”. All of these are
vanvolxemii. Most are heavily marked, winter-spring forms. The van-
volxemii at Potrerillos may have some mercedis introgression and are
undersized in summer. Also in La Plata are lef 1$ from Potrerillos
(also X.1951) which were determined by Field as vanvolxemii but bear
a handwritten label: “Opina Field que son intermedio de microdice
macrodice o hibridos de estos” (“Field opines that these are inter-
mediate to macrodice or hybrids with it”). (See Porter and Shapiro
1990 for discussion of the Potrerillos population.) There is no current
basis for recording macrodice in the Cuyo.
I have a single, highly unusual male from the highlands of San Juan
which is in some ways intermediate between macrodice and sterodice
28(3):137-238, 1989(91)
159
or mercedis. Its data are: Arroyo de Ago a Negra, 3300 in, 3. XL 1988
(AMS). The pattern is more or less midway between macrodice and
mercedis (Fig, 12); the scales behind the eyes are white (as in mercedis
or sterodice ) rather than orange (as in macrodice ); its compound eyes in
life were gray-green, intermediate between the gray-blue of sterodice
or mercedis and the bright chartreuse-green of macrodice , and unlike
any other Tatochila I have examined alive in nature; and its genitalia,
examined only externally, appeared to be arranged as in sterodice or
macrodice rather than as in mercedis, I interpret this individual as
possible evidence for a transition from macrodice to mercedis in the
very-poorly-collected zone of transition from the Quichua to the Cuyo
District.
Given the amount of collecting done in the vicinity of the Paso
Bermejo (Las Cuevas, Punta de Vacas, Puente del Inca) in the past
century, if macroUice occurs in that easily accessible locality in the
cordillera of Mendoza it must be either at very low density or at
altitudes above 4200 m, which are less well-known. I did not find it
there (up to 4350 m) in six days in XI— XII. 1989.
Tatochila mercedis vanvolxemii
The winter form of this widespread and common entity shows the
complete sterodice pattern in the male. As only reared (experimental)
examples have been figured heretofore, and then not in systematic or
biogeographic works, I provide (Fig. 13) wild-collected ones from the
“core” range where introgression from other subspecies is highly un-
likely.
The NW limits of vanvolxemii have been in question. It has often been
cited from “Tucuman,” which is usually (and often wrongly) inter-
preted as referring to the capital, San Miguel de Tucuman, where it
definitely does not occur (and has not in historic times). The only
definite records I have from the Province of Tucuman are from
Amaicha del Valle: 4cf 3$, 4.II.1984 and 1$, 26.XI.1989 (all AMS).
Amaicha, on the dry side below Abra Infiernillo en route to the Valles
Calchaquies, is a high-desert locality. More or less similar habitat
occurs up the Valles Calchaquies (Rio Santa Maria drainage, the N-
most extent of the monte) as far as the vicinity of Cachi, Salta where I
have collected only T autodice and Ascia monuste automate Burmeis-
ter. Thus we may infer that vanvolxemii follows the monte to near its N
limit but does not extend into the prepuna. If T. m. vanvolxemii is a
permanent resident at Amaicha it is spatially very close to T. m.
macrodice , though altitudinally and ecologically segregated. Our
samples from Amaicha are very small to detect introgression from
macrodice. Two of the females, are unusually heavily-marked, and one
is the darkest vanvolxemii I have seen. The XL 1989 female is old and
worn but vaguely suggestive of macrodice influence. It does seem cer-
160
J. Res. Lepid.
tain that there is no extensive intergradation, such as occurs farther
south in this polytypic species.
Populations of apparently pure vanvolxemii occur at low-elevation,
desertic sites quite close to the zone of intergradation with sterodice
and mercedis in NW Patagonia. Examples are:
2 c? Covunco, 820 m, Neuquen, 12. XI. 1966 (MG)
2c? 1? Estancia Corral de Piedra, Collon Cura, Neuquen, 650 m,
3. XII. 1969 (MG)
lC? Las Lajas, Neuquen, 730 m, 15.1.1980 (MG)
The Collon Cura series includes both extreme winter forms and
intermediates to the summer form. A similarly mixed series was taken
at Zapala, Neuquen, 9-11.XI.1988 (AMS). This is also a pure popula-
tion. The occurrence of winter forms as late as early December at low
altitude is fairly unusual; they normally occur only in the first brood.
Control of the polyphenism is discussed in Shapiro (1980a).
Typical vanvolxemii (averaging small) are abundant in eastern Pata-
gonia as far south as Trelew (18c? 5?, 7. XII. 1989), Las Plumas
(43°43'S, 67°15'W, 11c? 6$, 7.XII.1989), and Parada Uzcudun
(44°13'S, 66°09'W, 17c? 17?, 7.XII.1989) (all Chubut, all AMS). At
Trelew oviposition was observed on Cardaria draba (L.) Desv. (Cruci-
ferae), usually a disfavored Pierid host.
Records of T. m. vanvolxemii from “Buenos Aires” have always been
ambiguous due to confusion of the city and the Province. On
23. XI. 1989 it was common in the city (Costanera Sur, Avenida de los
Italianos, Zona Portuaria, vacant lots in La Boca), the first time I have
seen it there.
Hybrid Zones involving T. m. vanvolxemii
2c? 13? Loncopue, Neuquen, 8. XL 1988 (AMS)
This remarkable series marks a new N limit (38°04'S) for the zone of
intergradation, or hybrid zone, between vanvolxemii and other enti-
ties. Although very close to Alumine (39°13', 70°57'W), this population
is phenotypically very different. The Alumine population shows no
significant vanvolxemii component, while that at Loncopue is mostly
vanvolxemii (by phenotype). The females are most like vanvolxemii ,
with variable sterodice influence. The two males differ astonishingly
from each other: one resembles the putative sterodice -vanvolxemii
hybrid from Bariloche figured by Shapiro, 1980a (which in turn resem-
bles series of this cross, reared in the laboratory under summer photo-
period/temperature regimes); the other resembles a winter form of
vanvolxemii below and a sterodice -mercedis intergrade with strong
sagittate pattern above. All of these (Fig. 14) were taken flying
together in one pasture. The host plant is Lepidium sp. (Cruciferae).
Loncopue is, as noted above, the N-most locality known for T. theodice
28(3): 137-238, 1989(91)
161
in Argentina. It is also the N most locality for the skipper Hylephila
signata Butler and the third most N for Colias vauthierii Guerin,
discussed later. Both of these are typical Patagonian mallin species
which rarefy progressively as one moves N into the transition to the
cuyo.
22 ( f 11J Barrio Prosper© Palazzo, Comodoro Rivadavia, Chubut.
19.XI.1988 (AMS) Several previous collections from Comodoro Riva-
davia (San Jorge District) (15°52'S, 67°30'W) were made in high
summer and had produced the erroneous impression that the popula-
tion there was pure vanvolxemii. As noted above, summer vanvolx -
emii lose most or all of the black pattern in the male, and this is the
summer phenotype at Comodoro. This large series of the first (ex dia
pause), spring generation — in which the pattern is fully expressed
— reveals a strong apparent admixture of sterodice genes (most
apparent in the dotted DFW pattern, which is almost never seen in
vanvolxemii in its “core” range far from hybrid zones). A few of these
Comodoro males would unhesitatingly receive sterodice labels if cons-
idered out of context.
In 1989 a systematic attempt was made to map the distribution of
intergrading populations in eastern Patagonia. Populations very simi-
lar to that at Comodoro were found at: El Tordillo (45°53'S, 67°57'W,
6cf 5$, 8. XII); Pampa del Castillo (45°48'S, 68°65'W, 13d1 72, ll.XII),
both Chubut; Caleta Olivia (46°26'S, 67°32'W, 9d 62, 9. XII) and Fitz
Roy (47°02'S, 67°15'W, 25d 72, 9.XII), both Santa Cruz (all AMS).
The seasonal polyphenism of vanvolxemii is expressed in the San
Jorge region superimposed on a mixed genetic background. This
phenomenon, as noted above, is best detected in the spring brood and it
remains to be seen if these populations differ among themselves in the
strength of the polyphenism. The occurrence of the intermediates here
is consistent with the anomalous climatic and vegetational character
of the San Jorge District, reflected also in the abundance of T. a.
autodice, Colias lesbia, and other species much farther south than they
occur in the interior. Between Fitz Roy and Comandante Luis Piedra-
buena all trace of vanvolxemii phenotype disappears. Although I have
visited both San Julian (49°18;S, 67°13'W) and Puerto Deseado
(47°45'S, 65°54'W), I have collected no Tatochila due to bad weather.
ML, however, contains a Puerto Deseado male which if anything shows
a transition from sterodice to fueguensis phenotype, certainly nothing
of vanvolxemii. MLP contains 3 San Julian specimens which appear to
be pure sterodice . All trace of vanvolxemii also disappears between
Pampa del Castillo and Sarmiento (45°36'S, 69°G5'W).
Fig. 15 demonstrates the variation in the first brood at Comodoro.
MLP also contains a 2 from San Martin de los Andes, Neuquen,
2.1.1958, which resembles closely a lab Fx hybrid between mercedis
and vanvolxemii (not involving sterodice ).
162
J. Res. Lepid.
Tatochila mercedis sterodice
Populations of T. m. sterodice are extremely variable even when not
plainly involved in intergradation with other taxa; indeed, there are
few characters which are sufficiently constant throughout its range as
to be diagnostic. Thus, singletons or short series are of limited use in
characterizing geographic patterns of variation. The range of variation
is, however, broader still in zones of contact with other taxa, and
numerous novel phenotypes not seen elsewhere occur there. When in
contact with nominate mercedis in NW Patagonia, sterodice displays
lability in genitalic morphology as well as in wing pattern. Elsewhere,
its genitalia are constant (Porter and Shapiro 1989).
The following short series all appear to fall within the “normal
variation” displayed by “pure sterodice .” The Valle Lago Blanco series
is especially variable, and one specimen from there (reported below)
appears to be a hybrid with mercedis. The neotype of sterodice designa-
ted by Ackery (1975) is singularly unfortunate in being at the heavily-
marked end of the “normal variation” spectrum. Moreover, the entire
type-series of sterodice came from within the zone of intergradation to
fueguensis, in the far S of the range of sterodice. The synonym allodice
Bryk, from LLau-LLau (near Bariloche) would make better geographic
sense!
lcf Estancia Huechahue, Neuquen, 14. XL 1988 (AMS)
2cT Chapelco, 1750 m, Neuquen, 20.11.1973 (MG)
1$ Arroyo Chapelco Grande, Neuquen, 900 m, 15. XII. 1970 (MG)
lcf Cordon Chapelco, Portezuelo Trahunco, 1750 m, Neuquen,
27.XII.1978 (MG)
3cf Refugio Graeff, Parque Nacional Lanin, Neuquen, 1750 m,
12. 111. 1980 (MG)
lcf Lago La Kika, Neuquen, 1750 m, 24.1.1979 (MG)
lcf Quila Quina, Lago Lacar, Neuquen, 1.1958 (MLP)
4cf Pulmari, Rio Alumine, Neuquen, 4500', Feb. 1902 (H. J. Elwes)
(BM)
lcf Bariloche, Rio Negro, III. 1948 (Williamson & Martinez Fontes)
(MR)
lcf Parque Nacional Nahuel Huapi, Dec. 1912 (BM)
lcf Puerto Blest, Lago Nahuel Huapi, Rio Negro, 770 m, loc. #8,
1. 111. 1979 (DE)
lcf Canada Leon, Chubut, no date (MLP)
lcf Alto Rio Senguerr, Chubut, 18. XI. 1988 (AMS)
lcf Colonia Sarmiento, Chubut, 600 m, loc. #46, 15.11.1979 (DE)
4cf Tecka, Gobernador Costa, Chubut, 600 m, loc. #23, 7.1.1979 (DE)
Ij Tecka, Chubut, 3000', Jan. -Feb. 1920 (BM)
2cf 19 Valle Lago Blanco, Chubut, “Thursby 1904-26” (BM)
lcf 19 Glen Kross, Santa Cruz, 11.1938 (MLP)
lcf Lago Argentino, Peninsula Magallanes, Santa Cruz, loc. #28,
1.II.1979 (DE)
28(3): 137-238, 1989(91)
163
3<j San Julian, Santa Cruz, no date (MLP)
tcj Cte. Luis Piedrabuena, Santa Cruz, 20. XL 1988 (AMS)
lcT Lago Onelli, Santa Cruz, 11.1953 (MLP)
Tatochila mercedis fueguensis
Herrera and Field (1959, p. 488) state that this is “probably most
distinct” among the subspecies of a microdice™ ( =sterodice ). Once again
this impression was an artifact of lack of far-S material. Such material,
once assembled, demonstrates an unequivocal and relatively even
cline from sterodice to fueguensis phenotypes. The following records
represent elements of that cline. Although some individuals are “typi-
cal” fueguensis , none of the longer series is, and some of the La
Esperanza material from the mainland is pi lenoty picall y indisting-
uishable from topotypical fueguensis (Fig. 18).
2d 4J La Esperanza, Santa Cruz, 130 KM NW Rio Gallegos, loc. #30,
15.1.1979 (DE)
1.CT Perito Moreno, Rio Fenix, Santa Cruz, XII. 1982 (J. Carreras) (MR)
3d Estancia La Cristina, near Lago Argentine, Santa Cruz, 8. II. 1953
(A. Willink) (ML)
lCf Puerto Deseado,' Santa Cruz, 16 J. 1967 (A. Willink) (ML)
1J Canterla Masci, Rio Gallegos, Santa Cruz, 22.XI.1988 (AMS)
3c; 7 Rio Gallegos, Santa Cruz, 1011.1979, loc. #31 (DE)
3d 6 J Rio Gallegos, Santa Cruz, 23 J. 1979 (AMS)
2c f Lago Fagnano, Tierra del Fuego, 100 m, loc. #33, 19.1.1979 (DE)
Tatochila mercedis mercedis
The location of the zone of intergradation between sterodice and
mercedis has been documented previously (Shapiro, loc. cit .) but the
following hitherto unpublished records are of interest because they fill
in gaps, or because they demonstrate that hybridization has been in
progress for at least several decades.
(a) Apparent hybrids or intergrades
lu Cerro Malo, 1700 m, Neuquen, 15.11.1954 (S. Schajovskoy) (MR)
lCf 1? San Martin de los Andes, Neuquen, XII. 1952 (S. Schajovskoy)
(MR)
lCf Pucara, Neuquen, 13.111.1960 (S. Schajovskoy) (MR)
IJ Pucara, Neuquen, 19. HI. 1960 (S. Schajovskoy) (MR)
lcT Refugio Graeff, Parque Nacional Lanin, Neuquen, 1750 m,
2.1.1980 (MG)
lCf San Martin de los Andes, Neuquen, 5 XL 1979 (MG)
IJ San Martin de los Andes, Neuquen, 640 m, 6.1.1978 (MG)
1J San Martin de los Andes, Neuquen, extreme winter form,
15.XI.1969 (MG)
Id' San Martin de los Andes, Neuquen, 1.1958 (MLP)
IJ San Martin de los Andes, Neuquen, 1. II. 1939 (MLP)
164
J. Res. Lepid.
lCf 1? Lago Lacar, Pucara, Neuquen, 750 m, loc. #9, 1. XII. 1978 (DE)
3cf 1$ Quila Quina, Lago Lacar, Neuquen, 1.1958 (MLP)
lCf Alumine, Neuquen, 1200 m, loc. #57, 14.III.1979 (DE)
Id" 22 Estancia Aschieri, below Primeros Pinos, Neuquen, 1130 m,
9. XI. 1988 (AMS)
lcf El Bolson, Lago Puelo, Rio Negro, Loc. #13, 26.11.1979 (DE)
Id" “Chile (sic), El Bolson,” no date (A. Kovacs) (BM)
lCf Valle Lago Blanco, Chubut, “1904-26”, #10316 (BM)
(b) Indistinguishable from Chilean mercedis
Id" Pucara, Neuquen, 22.1.1958 (S. Schajovskoy) (MR)
Id" 12 Pucara, Neuquen, 15.11.1956 (S. Schajovskoy) (MR)
Id1 Caviahue, Neuquen, 1500 m, 25.11.1962 (S. Schajovskoy) (MR)
Id" Lago Lacar, Pucara, Neuquen, 750 m, loc. #9, 1.XII.1978 (DE)
12 Quila Quina, Lago Lacar, Neuquen, 1.1958 (MLP)
Id" 12 San Martin de los Andes, Neuquen, 11.1939 (MLP)
Id" San Martin de los Andes, Neuquen, 1.1958 (MLP)
12 San Martin de los Andes, Neuquen, III. 1952 (S. Schajovskoy)
(MLP)
(c) Miscellaneous
The following specimen appears to be a complex hybrid involving
vanvolxemii, mercedis and sterodice; it matches certain laboratory-
reared hybrids of that composition almost exactly:
12 Pulmarl, Rio Alumine, Neuquen, 925 m, 27.11.1978 (MG)
Interspecies Hybrids
The following may be a unique hybrid of T. a . autodice and T. m.
vanvolxemii, showing a mix of characters of both. Both putative
parents occur at the site. They are in fact sympatric over nearly half
the country, so hybrids must be extremely rare since this is the only
suspected one yet found. This hybrid has not been produced in the
laboratory.
Id" Las Lajas, Neuquen, 730 m, 15.1.1980 (MG)
“Group D” Tatochila (the “ orthodice group”)
This is a heterogeneous and possibly not monophyletic group.
Ackery’s (1975) work has increased our knowledge of the central-
Andean members of this group, but species limits remain very poorly
defined and some of his assignments of subspecies to species may be
incorrect. The entire group may be Legume rather than Crucifer
specialists. In Argentina they extend only barely to Cordoba; most
species are found in the Y ungas and Quichua District.
Tatochila inversa Hayward (Fig. 17A, B, Plate IV)
Herrera and Field (1959) figured as the male of this species some-
thing from the Department of Cusco, Peru. Whether this is really
28(3):137-238, 1989(91)
165
inuersa remains uncertain, but true inversa males are now available
from various localities in northwestern Argentina. They differ
phenotypically among populations and perhaps between broods as well
(Fig. 17, from the puna; PI. IV, from the Sierras Pampeanas ) but differ
from Herrera and Field’s specimen in a number of details. The species
is not rare on summits within the puna. The type locality — Quebrada
Carapunco — is near Abra Infiernillo in the Sierras Pampeanas of
Tucuman. Animals from these ranges average larger than those from
elsewhere. If a subspecies name is ultimately necessary it would apply
to the smaller phenotype from Salta and Jujuy.
Eisele writes (in litt ., 26.1.1978): “Just got my first female ( inversa )
from the mountains in Salta. I had previously got a number of males
from Jujuy.” One of these is figured: Huacalera, N end Cerro Amarillo,
Jujuy, 3250 m, 4 J. 1980 (RE). Other records: Abra Infiernillo area,
Tucuman, lcf, 26. XI. 89, 3$, 20.1.1986 (AMS); lj Cerro Zapallar,
3720 m, Salta, 22.1.1986 (AMS); 1J Abra Azul Pampa, Jujuy,
23.1.1986 (AMS).
The Infiernillo records suggest that this species may be multiple-
brooded. The 26. XI. 89 male was taken at the very beginning of the
rains, before the vegetation had resumed growth.
Tatochila orthodice Weymer
This is a well-known and common species of N Argentina and adja-
cent Bolivia, essentially restricted to the yungas. Most records are low-
to-mid montane. It is not generally recognized as a component of the
high- Andean fauna but in fact occurs there seasonally in January,
flying with T. m. macrodice , T. inversa , T. distincta, etc. This altitudi-
nal migration occurs at the yungas-puna and yimgas-Quichua District
interfaces, including the Sierras Pampeanas. Some high-altitude re-
cords: 1J Abra Infiernillo, summit near 3800 m, 20.1.1986 (AMS); 2c?
19 Valle Encantado, 9725', Salta, 22.1.1986 (AMS); 29 Cerro Zapal-
lar, 3700 m, Salta, 22.1.1986 (AMS). At the time these were collected,
no orthodice were flying in the foothills just above San Miguel de
Tucuman, where the species is typically abundant in spring (Anta
Muerta, 26.XI.1977; El Siambon and Sala de San Javier, 26.XI.1977,
all AMS). I have a mid-elevation record seasonally inbetween these
(Tafi del Valle, 2200 m, 26.XII.1977, AMS).
This species was common at La Vina, Catamarca, 29. XI. 1989 (AMS).
Tatochila stigmadice Staudinger
Also best-known as a foothill yungas species, T. stigmadice occurs
occasionally in the high country in summer and also as low as the city
of San Miguel de Tucuman. My extreme records are: 19 Cerro Zapal-
lar, 3600 m, Salta, 22.1.1986 (AMS); 2c? Barrio Fray Usquiu, S.M. de
Tucuman, 450 m, 28. XL 1977 (AMS); lcf San Miguel de Tucuman
166
J. Res. Lepid.
(centro, along R.R. track), 29.XI.1989 (AMS). It was also flying at San
Javier, Tucurnan and La Vina, Catamarca the same day.
The southernmost record of this species is apparently an unusual,
white female from Yacanto, Cordoba, no date (Breyer) (MLP). The
Sierras de Cordoba have a dilute yungas element.
“Group E” Tatochila (the “xanthodice group”)
Although this group contains only two species, they are united only
by genitalic morphology. Their wing patterns, biogeography, and host
plants are different enough to raise serious doubts as to their true
affinity, and the morphology of the early stages is also somewhat
equivocal.
Tatochila distincta Jorgensen (Fig. 17C, D)
The La Plata collection contains two Jorgensen specimens: d\ Cerro
Ensenada (Catamarca), 22.11.1915 (#2651) and J, Cerro Negro, same
date (no number). These were selected in 1971 by L.E. Pena as
“hololectotipo” and “allolectotipo” respectively, and so labeled. Both
are small and dark.
Not uncommon between 3000-4000 m in dissected puna , and in the
Quichua District including the Sierras Pampeanas, flying with T.
inversa and T. m. macrodice. The life-history of this species has been
published (Shapiro 1986d). It is apparently an Astt'agalus-feeder
(Leguminosae) in nature, but can be reared on Crucifers. There are
phenotypic differences between the Salta and Jujuy populations on one
hand and the (topotypical) material from the Sierras Pampeanas
(Cumbres Calchaquies — Aconquija), which may ultimately justify
naming subspecies within Argentina. Sexual dimorphism is somewhat
reduced in this species.
2c? 20 km N Humahuaca, Salta, 3700 m, 12.1.1978 (RE)
8c? 3J Tres Cruces, Jujuy, 3800 m, 23.1.1986 (AMS)
1C? l9Abra Infiernillo, Tucurnan, 20.1.1986 (AMS)
29 Cerro Zapallar, Salta, 3720 m, 22.1.1986 (AMS)
Genus Hypsochila Ureta (Figs. 18, 19; ranges 10B, 11B)
In early stages as well as adults, one group of species of Hypsochila
appears to be the sister-group of the Tatochila mercedis and autodice
(Crucifer-feeding) complexes. Unfortunately, the genus Hypsochila
itself is in some disarray despite work by Ureta (1955, 1963) and a
revision by Field and Herrera (1977). The latter was regarded by its
authors as very preliminary, and was based largely on Herrera’s
Chilean material. The species limits were poorly defined, and the
authors went so far as to state (p. 5): “Five of the six species . . . are very
closely related and could be considered subspecies of a single widely
distributed species. However. . .two of these species are known to fly at
28(3):137-238, 1989(91)
167
the same time in at least two of the same localities ...” The data
reported here demonstrate that Field and Herrera were wise in con-
tinuing to treat these taxa as species, though tantalizing ambiguities
remain. The Argentine species appear to fall into two groups, based on
both adult and immature characters. Because the type-species is H.
wagenknechti Ureta, the group to which it belongs (comprising in
addition the taxa sulfurodice Ureta and galactodice Ureta) will retain
the generic name, should the genus be divided as seems likely. This
group has close affinities to Tatochila as noted above. The other group,
comprising the taxa argyrodice Staudinger, microdice Blanchard and
huemul Pena, has many unusual derived character states and is
farther removed from Tatochila. All of its taxa are austral in distribu-
tion. No name is currently available for the group, should it be
formally raised to subgeneric or generic status.
Hypsochila argyrodice (Fig. 18C)
lcT Cabo Penas, Depto. Rio Grande, Tierra del Fuego, 17. XII. 1983
(ML)
lC? Fitz Roy, Santa Cruz, loc. #26, 11.11.1979 (DE)
Hypsochila microdice (Fig. 18G, H)
37c? 11? Rio Grande, Tierra del Fuego, 25.XI.1988 (AMS)
1CT Estancia Maria Cristina, Route 3, Tierra del Fuego, 27. XI. 1988
(AMS)
lC? 1$ Foot of Glaciar Martial, Cordon Martial above Ushuaia, T. del
F., 29-30.XI.1988 (AMS)
These combined records demonstrate that the ranges of the two
southernmost Hypsochila interdigitate and they cannot be conspecific.
The Danish Expedition argyrodice from Fitz Roy appears to be the first
mainland specimen with a precise locality. Breyer (1939) never saw it
at all, even a specimen, but cites “Ushuaia, Rober.” Hayward (1950, p.
92) records “Chubut” without data. I searched unsuccessfully for it in
marginal weather at Fitz Roy on 20. XI. 1988. H. microdice is abundant
at Rio Grande and its life-history is in preparation. The phenotype is
quite variable, but not easily confused with anything else in the
region. The two Cordon Martial specimens are large, especially the
male, but still smaller than argyrodice. Microdice is a Legume-feeder
and given the very close morphological affinities, the others are likely
to be as well. I have no new data to report on H. huemul.
Hypsochila galactodice (Fig. 18D, E; 19C, F, G, H)
1J Rio Agrio, Neuquen, 10. IV. 1932 (MLP)
2c? Huacalera, N end Cerro Amarillo, 3250 m, Jujuy, 4.1.1980 (RE) (*)
lC? Cordon del Viento, Neuquen, 3000 m, 28.1.1985 (AMS) (*)
29 Lago Meliquina, Neuquen, loc. #10, 12. XI. 1979 (DE)
168
J. Res. Lepid.
lcf 1$ Alumine, Neuquen, 15-16.1.1981 (AMS)
lcT Jumn de los Andes, Neuquen, 13.XI.1988 (AMS)
12 Loncopue, Neuquen, 8. XI. 1988 (AMS) (*)
lcf 62 San Carlos de Bariloche, Rio Negro, 15.XI.1988 (AMS)
13cf 42 Esquel, Chubut, 17. XI. 1988 (AMS)
(plus reared diapaused and non-diapaused material from Bariloche
and Esquel)
Hypsochila wagenknechti wagenknechti (Figs. 181, 19A, D)
6cf 22 Las Cuevas, Mendoza, 31.X-1.XI.1988 (AMS)
2 lcf 112 Arroyo de Agua Negra, above 3200 m, San Juan, 3. XI. 1988
(AMS)
Hypsochila wagenknechti wagenknechti “spring form”
lcf 12 Arroyo Chapelco Grande, 900 m, Neuquen, 15. XII. 1979 (MG)
lcf Chapelco, Neuquen, 1700 m, 24.11.1952 (S. Schajovskoy) (ML)
Hypsochila wagenknechti sulfurodice (Fig. 18A, B, F)
lcf Huacalera, N end Cerro Amarillo, 3250 m, Jujuy, 4.1.1980 (RE)
3cf Altos de Abra Munano, 4165-4780 m, Salta, 21.1.1983 (AMS)
8cf Tres Cruces, 3800 m, Jujuy, 7. II. 1984 (AMS)
This group of taxa is so difficult that some critical determinations are
provisional (marked *). The life-histories of H. w. wagenknechti and H.
galactodice are fully known (Courtney and Shapiro 1986a, b; Shapiro,
in preparation). They differ about as much as the adults: wagenknechti
feeds in nature on Crucifers, galactodice on Tropaeolum. Brown (1987,
p. 102) maps these two species as either allopatric or parapatric at
roughly 32°S. Field and Herrera have galactodice only from Epulaf-
quen, Neuquen on the Argentine side and wagenknechti only from Las
Cuevas and the directly adjacent Quebrada de los Horcones, Mendoza
— widely-separated localities, falling in the Western Patagonian Dis-
trict and the Cuyo District respectively. The Loncopue female and the
male from Cordon del Viento, Neuquen are both virtually undetermin-
able, and both are geographically feasible as zones of primary (not
secondary?) intergradation if these two taxa are biologically con-
specific. To complicate matters, a second female Hypsochila taken the
same day at Loncopue is quite different and does not agree with any
described taxon.
The pair of apparent galactodice from Cerro Amarillo, Jujuy is extre-
mely problematical. Although matching the description of galactodice ,
they differ from Patagonian specimens in exactly the same ways
sulfurodice differs from nominate wagenknechti. If H. galactodice
actually occurs some 2000 km N of its previously-known range, in a
different biome and sympatrically with a subspecies of wagenknechti ,
there can be no question of conspecificity with that species. However,
28(3):137-238, 1989(91)
169
the series is too short to rule otit sampling error disguising continuous
population variation at Cerro Amarillo, from a usual sulfurodice to a
galactodice- like phenotype. My long series of wagenknechti wagenk-
nechti from Mendoza and San Juan includes individuals which
approach galactodice in phenotype, underscoring the variability of
these animals and the potential unreliability of small samples. Unfor-
tunately, females are rarely encountered and males are concentrated
on relatively inaccessible hilltops, where they are difficult to catch. In
the absence of any reliable, diagnostic morphological character this
problem is insoluble at this time. Absolutely no tendency to resemble
galactodice has been seen in any other sulfurodice (i.e. from localities
other than Cerro Amarillo).
In the Cordon Chapelco near San Martin de los Andes a small, very
dark form of (?) wagenknechti occurs in which the dorsal apical mark-
ings tend to fuse; the reflective gloss at the base of the wings is more
pronounced; and the VHW pattern is extremely heavy. The pattern is
vaguely suggestive of the next species. I am treating these as spring
forms of wagenknechti because I have very similar specimens from
Chile: Cumbres de La Parva, Prov. Santiago, 24.XI.1982 and Los
Libertadores, Prov. Los Andes, 3900 m, 27-28.1.1983 (all AMS), taken
right at melting snow-line in an area (just across the Paso Bermejo
from Las Cuevas) where no other taxon is at issue. However, seeming-
ly typical galactodice occurs at Junfn de los Andes (first brood, XL 13),
quite close to but lower than the Cordon Chapelco.
Near Las Lenas in southern Mendoza, 3d" Hypsochila were collected
on an altitudinal transect up Cerro de los Fosiles, 3. XII. 1989: a typical
galactodice at 2100 m, an intermediate specimen at about 3000 m and
a typical wagenknechti at 3300 m. All are figured on Plate IV. Again,
this suggests altitudinal stratification and intergradation (but the
sample size is very small). Strikingly, the two higher specimens were
hilltopping while the low-altitude one was visiting a dandelion on a
vega , behaving much as a galactodice “should.”
Problematic specimens of this group of taxa, including a striking
aberration of wagenknechti , are shown in Figs. 18-19.
Hypsochila penai Ureta
So far known only from Chile. A cf in ML is labeled “Alto de Puripica,
4600 m.” The country is not given. This locality is in Chile (22°30'S,
68°07'W) quite close to the Argentine border.
Genus Phulia Herrich- Schaeffer (Fig. 20; ranges 10C, 11C)
This is a classically high-Andean genus, restricted to puna and
altiplano from C Peru through Bolivia to NE Chile and NW Argentina.
All the Argentine populations known are treated by Field and Herrera
(1977) under the name P. nymphula Blanchard. Genetically and ecolo-
gically these populations are somewhat diversified. Life-history and
170
J. Res. Lepid.
electrophoretic data will be published (Shapiro, Courtney, Descimon &
Geiger, in preparation).
In Argentina “P. nymphula ” is distributed in four geographic regions.
Rather than multiply names, I present representative data and dis-
cuss the status of available names potentially applicable to these
populations.
(i) The puna of Jujuy and Salta
26c? 13J Tres Cruces, Esquinas Blancas, Jujuy, 7. II. 1984 (AMS),
23.1.1986 (AMS)
2cf Abra de Fives, 4200 m, Jujuy, 29.1.1969 (ML)
1C? near Turilari, Jujuy, 4000 m, 5.IX.1968 (ML)
12 Rosario de Coyahuayma, Salta, 4400 m, 11 JX. 1968 (ML)
ICf Rio Cincel, Jujuy, 3800 m, 3. IX. 1968 (ML)
Plate I. Pierid habitats in montane northwestern Argentina. A, view of puna
dominated by tola, looking N from a summit near Abra Muhano,
Salta; habitat of Hypsochila wagenknechti sulfurodice. ii. 1 983. B,
Dissected puna at the head of the Quebrada de Humahuaca, Jujuy,
near Abra Azul Pampa. Habitat of H. w. sulfurodice, TatochHa dis-
tincta, T. mercedis macrodice, Phu/ia nymphula, and Co/ias b/ameyi.
ii. 1 985. C, Yungas in Salta, below Valle Encantado, showing deeply
dissected topography. Habitat of TatochHa orthodice and stigmadice
and Terioco/ias riojana. i. 1 986. D, Valle Encantado, Salta, in the
uppermost yungas, at the height of wet season; Co/ias blarney i,
TatochHa orthodice, T. stigmadice, Terioco/ias riojana. i.1986. E,
Rocky summit in the Cumbres Calchaquies, Tucuman (quichuan
vegetation with large Azorei/a in foreground). Habitat of TatochHa
inversa, distincta, mercedis macrodice, and Co/ias b/ameyi. i. 1 986. F,
Dry subalpine shrub-steppe above Amaicha del Valle, Tucuman,
looking toward the Valles Calchaquies. This belt forms the apparent
barrier between TatochHa mercedis macrodice above and T. m.
vanvo/xemii below, i. 1 986. All photos by AMS.
Plate II. Pierid habitats in montane west-central Argentina. A, prepuna with
columnar cacti near Cachi, Salta. No endemic Pierid fauna, but
TatochHa autodice and (seasonally) Ascia monuste automate com-
mon. i. 1 986. B, Riparian vegetation in wet season in the monte,
Valles Calchaquies between Fuerte Quemado and Cachi. TatochHa
mercedis vanvo/xemii occurs in similar vegetation farther S but is
not known N of Amaicha del Valle at this time. i. 1 986. C, Precordil-
lera near Potrerillos, Mendoza. T. m. vanvo/xemii with possible
introgression from T. m. mercedis are abundant here, x.1988. D,
Vega at the head of the Arroyo de Agua Negra, San Juan, near the
Chilean border. The only known locality for Co/ias f/aveo/a in Argen-
tina. Nearby occur Hypsochila w. wagenknechti, Phu/ia nymphula
and other characteristic cordilieran taxa. xi.1988. E, Sparse alpine
steppe in the Aconcagua Provincial Park, Mendoza, in early spring;
Phu/ia nymphula very abundant, xi.1988. F, Rockslides among the
summits overlooking the Paso Bermejo, Mendoza. Hypsochila w.
wagenknechti abundant, xi.1988. All photos by AMS.
28(3):137-238, 1989(91)
171
172
J. Res. Lepid.
28(3):137-238, 1989(91)
173
174
J. Res. Lepid.
28(3):137-238, 1989(91)
175
Plate III. Pierid habitats in southern Argentina. A, partially inundated mal-
lines at Loncopue, Neuquen, site of a hybrid Tatochila me reed is
sterodice X vanvolxemii population. Also occurring here are T.
theodice, Hypsochila galactodice, Co/ias vauthieri, and various
other Patagonian taxa near or at their N limits. xi.19S8. B, Mosaic of
Nothofagus forest, subalpine steppe, and Patagonian grassland/
mallm E of San Martin de los Andes, Neuquen, a major ecotone.
Hybrid Tatochila mercedis mercedis X T. m. sterodice with slight
T. m. vanvolxemii influence eastward common in the valley;
Hypsochila Iwagenknechti occurs above tree line and H. Igalacto-
dice in the valley at nearby Junin de los Andes. Eroessa chi/iensis
and Mathania leucothea fly further west in the Valdivian forest.
xi.1988. C, Shrub-steppe with a few small Nothofagus near San
Carlos de Bariloche, Rio Negro, in the deforested steppe-forest
ecotonal zone. Now dominated by neneo { Mulinum , Umbelliferae).
Tropaeo/um polyphyllum abundant. This is the habitat of Tatochila
autodice autodice X T. a. blanchardii intergrades and of Hypsochila
galactodice. xi.1988. D, Patagonian shrub-steppe at Fitz Roy, Santa
Cruz, the only definite mainland locality for Hypsochila argyrodice.
xi.1988. E, Windswept bunchgrass steppe in the Department of Rio
Grande, Tierra del Fuego; habitat of Hypsochila microdice and
Tatochila theodice near staudingeri. xi.1988. F, alpine shrub-steppe
and Nothofagus krummho/z in the Cordon Martial, Tierra del
Fuego. Hypsochila microdice and a dwarf race of Yramea cytheris
(Nymphalidae) occur here, xii.1988. All photos by AMS.
Plate IV. Zoogeographically important Argentine Pieridae and some of their
habitats. A, upper and B, lower surfaces: 1. Co/ias mendozina,
Aconcagua Provincial Park, Quebrada de los Horcones, Mendoza
(across top: left, male, 30. XI. 1989; center, yellow female and right,
whitish female, 15. XII. 1989). 2. Male Hypsochila from vicinity of
Las Lenas, Mendoza, 3. XII. 1989, showing apparent intergradation
from wagenknechti to galactodice phenotypes. The two upper
specimens are from >3000 m on Cerro de los Fosiles; the lower
one is from a wet vega at 2100 m. 3. Phulia nymphula from the
southernmost locality known in the main Andean cordillera, male
(above) and female; Cerro de los Fosiles, near Las Lehas, Mendoza,
3100 m, 3. XII. 1989. 4. Co/ias vauthierii from the northernmost
known locality, Arroyo El Deshecho, near Las Lenas, Mendoza,
2100 m, 5. XII. 1989, male (above) and female. 5. Male Tatochila
in versa, Abra Infiernillo, Tucuman, 26. XI. 1989, the only cT specimen
of the first brood known to me. C, D, Habitats of Co/ias mendozina
in Quebrada de los Horcones, Aconcagua Provincial Park, Mendo-
za. Phulia nymphula and Hypsochila wagenknechti wagenknechti
also fly here; 3300-3900 m. XII. 1989. E, Habitat of Hypsochila near
galactodice, near Las Lenas, Mendoza, 3000 m, XII. 1989. F, Alpine
steppe near summit of Cerro de los Fosiles, near Las Lehas,
Mendoza, 3400 m, habitat of Hypsochila near wagenknechti and of
Phulia nymphula. XII. 1989. (Figs. A and B by S. W. Woo; remainder
by AMS.)
176
J. Res. Lepid.
lCf Tolar Grande, Salta, 3525 m, 15.L1978 (M. Vargas) (ML)
(ii) The Quichua District
(a) Salta highlands
3d" Cerro Zapallar, Salta, 28. XL 1989 (AMS)
(b) The Sierras Pampeanas
2J El Manchal, Catamarca, 4000 m, 1.1959 (ML)
IJ Cerro Munoz, Tucuman, 4000 m, 1.1959 (ML)
lcf Summit above Abra Xnfiernillo, 3450 m, 26.XI.1989 (AMS)
6cT 59 Huaca Huasi, Tucuman, 4150-4250 m, 2. III. 1977, 4.IV.1977,
14.VII.1977, 1. IX. 1978, 23.IX.1978, 28.IX.1978 (all S. Halloy and E.
Dominguez) (ML)
(iii) The Cuyo District
25 cT 59 + 1 mosaic gynandromorph (Fig. 21), Las Cuevas to Puente
del Inca, Mendoza, 31.X-1.XI.1988 (AMS)
16c? 209 Arroyo de Agua Negra, San Juan, above 3200 m, 3. XI. 1988
(AMS)
+ 28 additional specimens from the Paso Bermejo, Mendoza (MR, ML,
BM, AMS)
4 cT 19 Cerro de los Fosiles, 3100 m, near Las Lenas, Mendoza,
3. XII. 1989 (AMS)
3d1 Cerro de los Fosiles, 3300-3400 m, Mendoza, 3.XII.1989 (AMS)
(iv) The Cordon del Viento
Id" Cordon del Viento, Neuquen, 3700 m, 28.1.1985 (AMS)
The Problem of Phulia aconquijae Jorgensen
Since its original description (1916), Phulia aconquijae has generally
been synonymized to nymphula — beginning with Breyer (1939), who
used it in a subspecific sense. Field and Herrera (1977) state that “a
study of the original description and of topotypes show that this name
is a junior synonym of Phulia nymphula nymphula .” In taxonomy as in
jurisprudence, however, the reasoning leading to a conclusion of fact
must be spelled out before the conclusion can be accepted, and in this
case it was not. Field and Herrera were unable to locate any definite
material from the type-series, but state that “Topotypes are present in
the collection of the National Museum of Natural History ... and
indeed . . . may represent syntypes (although they are not so labeled). A
lectotype designation is not needed at the present time.’5 The reader
cannot reconstruct how Field and Herrera dealt with the differences
identified by Jorgensen between his “new” species and P. nymphula .
Jorgensen was an unusually meticulous observer, and the question of
how he came to erect a synonym of the well-known, widely-distributed
P. nymphula has never been addressed. The clue lies in the description.
Jorgensen knew, or thought he knew, P. nymphula from Bolivia, and
used it as a comparison in his diagnosis of aconquijae. Although he
gives a very precise description of the wing phenotype, it is clear that
for him the critical character was the venation. He says (1916, p. 517):
28(3):137-238, 1989(91)
177
“This new species much resembles P. nymphula Stgr. from Bolivia,
but distinguishes itself beyond differences of color and pattern princi-
pally in that the second radial vein (as in the genus Andina Stgr. =
Piercolias Stgr., AMS) issues directly from the subcostal and not, as in
the other species, united for a while with it (in one example of the
female they are united a short distance).”
This makes no sense if one has learned only the Comstock-Needham
system of naming the wing veins. But Jorgensen is using an antique
terminology which employs some of the same names as Comstock-
Needham but for different veins . The “second radial vein” as used here
means our M2, which is entirely free in Piercolias and in P. nymphula
but anastomosed with R3+4+5 in Phulia paranympha Stgr. from Boli-
via. The “subcostal” is not our Sc, but the combined radials. Thus we
may infer that Jorgensen had Bolivian paranympha misidentified as
nymphula , and concluded that his material — true nymphula , in fact
— was different and new. The source of this confusion was traced by
Breyer (1939) and echoed by Field and Herrera (1977) but only in
reference to the name Phulia reedi Giacomelli (1918). Giacomelli’s
type series was from Las Cuevas in the Paso Bermejo, Mendoza, that
most accessible of high-cordilleran sites. Staudinger (1894) had mis-
identified Chilean material as his own paranympha , when it was
actually nymphula. Giacomelli reasoned that his material from Men-
doza was not paranympha , hence had to be something new! In 1924
Rober proposed the replacement name joergenseni for the Bolivian
insect, if in fact Jorgensen’s insect aconquijae were synonymous with
nymphula (type locality Coquimbo, Chile): after all, wouldn’t that
leave the Bolivian species nameless?
In fact, aconquijae , that is, Phulia from the Sierras Pampeanas , like
some of the sympatric Tatochila is perhaps marginally taxonomically
recognizable on the basis of the very small size of September -October
material, electrophoretic and early-stage characters. The populations
in areas i-iii all differ at this (subspecific) level, but adult wing
phenotyes are extraordinarily variable in all of them. (The Cordon del
Viento population is known only from my specimen and an indepen-
dent collection by MG, which I have not seen.) If it is ultimately
desirable to name these, nymphula is the correct name for the Cuyo
District populations, aconquijae remains available for the Calcha-
quies-Aconquija animal, reedi is unambiguously a synonym of nym-
phula sensu stricto , and there is no name available (unambiguously)
for the puna populations.
Field and Herrera present the type-locality information for acon-
quijae in a confusing way. On p. 19 they quote Jorgensen’s list of
localities (from his p. 517; Cerro Medio through Cerro Negro) without
mentioning La Ollada, which Jorgensen finally refers to 16 lines later.
Their list of material examined (p. 20) includes among localities only
La Ollada as possible topotypes, as referred to in the text. Breyer
(1939, p. 46) refers to Jorgensen’s “types”: “Ein Vergleich unserer Tiere
178
J. Res. Lepid.
mit der Typen Joergensens ...” Breyer’s collection is at La Plata,
where there are five Jorgensen Phulia, none identified as the type of
aconquijae. They are in drawer 83 of the “Petrowsky collection” cabinet
and are not listed as types in the type file. All have Jorgensen MS
labels and they are from: Cerro Medio, 2 c? 1$, 13.11.1915 and La
Ollada, Icf 28.XXI.1916, 1? 17.III.1916. In MR are several more
Jorgensen specimens bearing MS labels “aconquijae n. sp.” and “La
Ollada/Catamarca,” almost certainly syntypes. All of these are normal
Phulia nymphula from the Sierras Pampeanas , not a new species.
Jorgensen himself notes variation in his venation character. The
venation of several Phulia sensu lato appears quite labile. Very small
P. nymphula , such as occur at Huaca Huasi in September, may have
R3+4+5 crowded at the very apex of the FW, where it diverges from Mx,
looking like it is about to be pushed off the apex altogether. This in fact
has happened in Infraphulia madeleinea Field and Herrera from Peru,
but Shapiro (1985) reported a female from the Department of Junln
(above Lima) which had conserved the “lost” vein on both FW, and
Lamas (in lilt ., 15. XIX. 1986) reported a male from Pampa Galeras,
Ayacucho with a 3-branched radial on one FW and 2-branched on the
other.
The S extent of P. nymphula remains conjectural. The Cordon del
Viento population, at 37°S (70°30'W), is considerably S of the farthest
S record in the main cordillera (35°04'S). We have not found P.
nymphula in seemingly suitable habitat in the Maule district, Chile
(36°S, S.P. Courtney) or at Copahue, Argentina, above Chos Malal
(37°45'), where T. m. mercedis leaks over on to the E slope.
Genus Eroessa Doubleday
Eroessa chiliensis Guerin
This remarkable species barely enters Argentina at the extreme W
ends of the Nahuel Huapi (Rio Negro) and Lanin (Neuquen) National
Parks. Pena (1975) records it simply from Neuquen, but its potential
habitats occupy less than 1% of the area of the Province. I have seen it
between Puerto Blest and Laguna Frias in the Nahuel Huapi park.
Jorgensen (1916) and Breyer (1936, 1939, 1945) were apparently
unaware of its occurrence in Argentina and Hayward (1950) as usual
cites only the Province, Neuquen. Schajovskoy, who was resident
naturalist in the Lanin Park, knew it well and found it at Quechu-
quina (40°10'S, 71°35'W). MR contains a seemingly reared Schajovs-
koy male from there, dated 5. XI. 1952. MLP has a male from San
Martin de los Andes, Nqn., 27.1.1941 (R. P. Bilardi). The life history
has been published (Angulo and Weigert 1974); Wagenknecht (1968)
offered notes on the behavior and ecology of the adult. The host plant
has been reported in print only by Pena (1975), who identifies it as
Flotouia (= Dasyphyllum) diacanthoides Cabr., a Composite of the
28(3):137-238, 1989(91)
179
primitive endemic tribe Mutiseae. It is a shrub or small tree, known
locally as Palo Santo , and is typical of the Valdivian Tertiary relict
rain forest (Ringuelet 1955) — one of the oddest hosts recorded for any
Pierid in the world and thus seemingly underscoring the antiquity and
taxonomic isolation of Eroessa.
Genus Mathania Oberthiir
Mathania leucothea Molina
Another essentially Chilean species, in this case not restricted to the
Valdivian forest but extending far north in matorrol in the precordil-
lera. In Argentina it occurs in the same areas as Eroessa but pene-
trates somewhat farther eastward — its range may have contracted as
a result of 19th-century deforestation in the Lake District. Schajovs-
koy collected it frequently at Pucara in the Lanin National Park
(5.XI.1958, 30.XI.1960, XII.1950, etc., MR) and I have seen it at Cerro
Catedral near Bariloche as well as further W, and once in the hills just
SW of the city limits. The hosts are mistletoes (“Quintral,” Phrygilan-
thus = Tristerix tetrandus Ruiz and Pavon, Loranthaceae), which are
common near Pucara. For aspects of its biology, see Courtney (1986).
Genus Colias Fabricius
Colias ponteni Wallengren = C. imperialis Butler
This biologically very important species remains “lost” since the
original series was collected, and is discussed here because of the
possibility it may yet turn up in Fuegia. G. Lamas writes (in litt .,
13.V.1981): “Ponteni’s type locality is ‘Honolulu,’ (collected by the
Eugenies Expedition; a gross mistake), and the type locality of im-
perialis is ‘Port Famine’ (collected by P. King?). Port Famine is Puerto
del Hambre, Magallanes, Chile (53°38'S, 70°56'W), which is S of Punta
Arenas; the frigate ‘Eugenies’ was in Port Famine from 31.1 to
2. II. 1852, and ponteni was probably collected on an excursion the
scientists aboard made to Mt. Tarn.” Cerro Tarn (819 m) is right by the
coast just S of Puerto Hambre. Apparently no one has seen Colias
ponteni alive in over 137 years! Nor has anyone visited the presump-
tive type locality to look for it, though I have assiduously searched
meadows with clover and vetch in various parts of Argentine Tierra
del Fuego. Any Lepidopterist visiting the region should look for it. It is
undoubtedly the morphologically most primitive Colias known, a
living — or recently extinct — fossil (Petersen 1963, Berger 1986).
H. Descimon has raised (in litt.) the intriguing possibility that the
Hawaii-Fuegia confusion arose over the South Sandwich Islands, one
of the Falkland Islands Dependencies in the South Atlantic, — “Sand-
wich Islands” being an antique English name for Hawaii. These
islands are so remote and isolated (latitude 56° to 59°S, longitude
180
J. Res. Lepid.
26°15'W) and have such a harsh climate that the occurrence of any
butterfly there would be extremely remarkable.
Colias vauthierii Guerin
This widespread and common Patagonian and Chilean species ex-
tends N into the transition zone between Patagonia and the Cuyo.
Hovanitz (1970) claimed a new N record in agriculturalized mallines at
Plottier in the valley of the Rio Limay, Neuquen, 8. XII. 1970. (There
are no Plottier specimens in the Hovanitz collection at CAS: the only
Argentine material is a series from “Coyaique,” 14.1.1967, which could
be either Coy Aike, Santa Cruz, or Coihaique in the pass between
Santa Cruz and Aisen (Chile).) Breyer (1939) quotes Kohler for “Rio
Agrio” (p. 50). In Neuquen I have three N collections of this species:
Alumine (16.1.1981), Loncopue (8. XI. 1988), and Chos Malal
(28.1.1985). Loncopue is in the Rio Agrio drainage.
Hayward (1973, p. 123) records this species from “Mendoza.”
Although no substantiating specimen has turned up, on 3. XII. 1989 I
found C. vauthierii common on vegas in the Valle de Las Lenas in
southern Mendoza at 2100 m. This is a remarkable range extension
which graphically illustrates the interdigitation of the high-Andean
and Patagonian biota in the Cuyo. Two specimens from this population
are shown on Pi. IV; they are completely “typical.”
C. vauthierii and C. lesbia Fabr. co-occur at Chos Malal with no trace
of interbreeding and apparently have a fluctuating zone of overlap
across central Patagonia. Strays of C. lesbia occur regularly S to San
Martin de los Andes, Neuquen and San Carlos de Bariloche, Rio Negro
and somewhat less often to Esquel, Chubut. C. lesbia is abundant in
eastern Chubut (Trelew, Rawson, Puerto Madryn), breeding and
perhaps overwintering at Comodoro Rivadavia since it has been taken
there in spring.
Colias flaveola Blanchard (Fig. 22; range 10D, 11D)
Nominate flaveola has been considered a Chilean endemic, and no
supposed subspecies are reported from Argentina. On 3.XI.1988 I
collected 18c? 4J flaveola above 3350 m in the Arroyo de Agua Negra,
San Juan (30°12'S, 69°51'W) in typical habitat — high-altitude vega
(sedgy stream bottom) (PL II -D). This locality is directly across the
crest from the classic localities in the Province of Coquimbo, Chile. The
Argentine material does not differ phenotypically from series from
Banos del Toro, Coquimbo, 3800 m, 7.1.1972 (Hovanitz) and Rio Seco,
11. III. 1936 and 20.11.1937 (E. P. Reed) (all CAS). The Chilean skipper
Hylephila isonira mima Evans (Hesperiidae) was taken in the Arroyo
de Agua Negra the same day.
28(3):137-238, 1989(91)
181
Colias mendozina Breyer (Plate IV; range Figs. 10B, 11D, 24)
Breyer (1939, p. 52) described this entity as C. blameyi f. mendozina.
The text of his description follows: “1st eine Abart, die sich durch
starke Verbreitung der schmutzigen Gelbfarbung auszeichnet. Vor-
derflugel mit breitem schwarzen Apex und Aussenrand, von dem aus
die breit geschwarzten Adern nach innen ziehen und mit dem Dis-
kalfleck verfliessen. Wurzel hinten und Innenteil des Innenrandes
intensiv schwarz. Hinterflugel am vorderen Tornus breit geschwarzt;
Diskalpunkt rein gelb. Wurzel und Zellbasis tief schwartz. Unterseite
zeichnungslos; Hinterflugel-Diskalfleck gelb und Wurzelfeld ver-
dunkelt. — Typus und Paratypus in unserer Sammlung. — Habitat:
Mendoza auf 3000 m Hohe, leg. Breyer.” (“A variety, distinguished by
the strong diffusion of the dirty yellow color. Forewing with broad
black apex and outer border, from which the broad blackish veins
reach inward and blend with the discal spot. Base and inner border
intense black. Hindwing with front angle broadly blackened; discal
spot pure yellow. Base of wing and cell deep black. Underside without
pattern; hindwing discal spot yellow and basal portion darkened. —
Type and paratype in our collection. — Habitat: Mendoza at 3000 m,
leg. Breyer.”)
The existence of this entity has been studiously ignored, e.g. by
Berger (1986). On biogeographic grounds the occurrence of a “variety”
or subspecies of C. blameyi Jorg. in the highlands of Mendoza seems
unlikely, since the entire group to which that taxon belongs is tropical
or subtropical except C. flaveola, which occurs S of the range of blameyi
itself in Argentina and thus would apparently intervene between it
and any putative subspecies in Mendoza. The description is barely
adequate to allow one to visualize the animal. At the end of November
1989 I examined the three specimens of mendozina in the Breyer
collection at MLP. The two types are faded and show evidence of
having been mildewed. Both are labeled “Argentina Prov. Mendoza”
and one bears a pink “TYPUS,” the other a green “PARATIPUS” label.
The third specimen is much brighter and fresher-looking, but has been
broken and glued with a heavy, opaque material. It is labeled (in
English) “Las Cuevas F.C.T. 15.I.1904/W.M.B. Seen also at Puente del
Inca.” F.C.T. presumably means “Ferrocarril Transandino.” These
specimens are clearly distinct from any other Andean Colias and not
particularly close to C. blameyi. Only a few days after examining them
I found C. mendozina flying in the Mendoza highlands! At present I
have 9 specimens, all from Quebrada de Los Horcones (Lower Horcones
to Confluencia), Parque Provincial Aconcagua: lef 30. XL 1989 and 4 cf
49 15. XII. 1989, all AMS. A male and both color forms of the female
are shown on Plate IV.
C. mendozina is easily told from other Andean Colias by its odd,
almost mustard-yellow ground color (even the pale females are of a
different hue than other Andean ones), small size, and short, almost
182
J. Res. Lepid.
truncate, fore wing. The dorsal black suffusion is weak and less exten-
sive than in other taxa, scarcely more than in C. flaveola. The discal
spot is large and rounded on males and variable on females. The large
blackish area at the apex of the hindwing above and the brownish
basal suffusion below are also distinctive. Oviposition was observed on
a small, tufted, blue-flowered Astragalus ( arnottianus (Gill.) Reiche)
(det. R. Barneby).
The known localities for C. mendozina are on the so-called “normal
route” used by climbers to access Cerro Aconcagua. Fig. 24 reproduces
a map of the Parque Provincial with known habitats of appropriate
type indicated, that is, vegas. I suspect C. mendozina will eventually be
found on all such vegas up to at least 4000 m. Conversations with
climbers and guides indicate many of them are familiar with the
animal.
Whether C. mendozina will eventually be found outside the Aconca-
gua Park is highly doubtful. Although the vegetation along Highway 7
is commonly taken as typical of the montane Cuyo (e.g. Wingenroth
and Suarez 1983), there is actually a rapid turnover in both floristics
and community composition between the C. flaveola locality in San
Juan and the vicinity of Las Lenas in southern Mendoza. C. mendozina
could thus easily be a narrow endemic (like C. flaveola ). Since it
formerly and perhaps still could be found at Las Cuevas it should be
looked for on vegas across the border in high-altitude Chile.
A formal redescription will be published at a later date.
The evolutionary and biogeographic history of this animal should be
of great interest. If it is truly related to C. blameyi , it must represent
an Interglacial stranding of.an essentially tropical stock which man-
aged to adapt to the strong temperate seasonality (including persistent
snow for 5-7 months) of the Paso Bermejo. Its existence under our
noses in the most accessible high-altitude locality in the Argentine
Andes is a potent reminder of our extreme ignorance.
Colias blameyi Jorgensen — C. weberbaueri Strand (Fig. 10D,
11D)
MLP contains an original Jorgensen specimen — a female dated
13.11.1915 from Cerro Medio, as well as later topotypes (very dark c?
and J, Cerro de la Mina, Depto. Tafl, Tucuman, IV. 1933, no collector).
The life-history of this species has been described by Shapiro (1989b).
Its distribution resembles the N part of that of Phulia nymphula or the
high-altitude Tatochilas — puna , Quichua District and Sierras Pam-
peanas. Some representative data: Quebrada Carapunco, Abra In-
fiernillo, Pcia. Tucuman, 12c? 3? 26.XI.1989, 35c? 10$ 20.1.86 (all
AMS); Abra Molina, Cerro Zapallar, Salta, 18c? 7$ 28.XI.1989, 5c? 7$
22.1.1986 (all AMS); 1c? 2$ Valle Encantado, Salta, 22.1.1986 (AMS);
13c? 3$ Esquinas Blancas, Jujuy, 7. II. 1984 (AMS); lc? Abra Pampa,
Jujuy, 7. II. 1984 (AMS). The Abra Infiernillo and Abra Molina data
28(3): 137-238, 1989(91)
183
unambiguously indicate at least two, perhaps three broods, the first
beginning immediately upon the onset of the rains (in November the
Astragalus are just beginning to grow and bud).
Variation among populations is quite noticeable. All the Argentine
material I have seen has at least traces of an androconial patch in the
male, but material from the puna is colored more like Bolivian C.
weberbaueri than is topotypical blameyi from the -Sierras Pampeanas .
CAS has long series of weberbaueri . In a group of 9 cT from “50 km S
Oruro, 3700 m, 14.1.1972, W. Hovanitz,” 7 have no androconial patch, 1
has traces, and 1 has a well-developed patch and is indistinguishable
from Jujuy males. This site is roughly 475 km NNW of Abra Pam pa.
Further study will almost certainly demonstrate the conspecificity of
these taxa. It is quite unusual to find such variation in a secondary
sexual character and potentially of great evolutionary interest. Pre-
sence of androconia appears primitive relative to absence in Co lias.
This, however, does not necessarily imply S-to-N movement in the
history of the green complex.
Genus Eurema Hubner
Eurema deva Doubleday
A common, weedy, highly dispersive species in N Argentina, reaching
far S as strays in much the same manner as E, lisa Bdv. & LeC. in
North America migrates N ward in summer. My S-most records are
Bariloche, Rio Negro, 17.1.1984 and Caleta Olivia, Santa Cruz,
9. XII. 1989. It also reaches high elevations, such as Abra Infiernillo,
Tucuman, 3300 m, 20. 1. 1986. All of these were females.
Genus Teriocolias Rober
TeHocolias riojana Giacomelli
This altitudinal disperser apparently reaches the summits in the
Sierras Pampeanas remarkably early in the rainy season, as single
females were taken at Abra Infiernillo, 26.XI.1989 (Tucuman) and
Cerro Zapallar, 28. XL 1989 (Salta) (both AMS). No known host plants
were available in either site so early in the year.
Discussion
The raw data for an analysis of the zoogeography of the faunas and
taxa reviewed here are based on collection records. Tables 1-2 present
the species compositions of 25 selected faunas in Argentina and Chile.
They are variably well-studied; richness is not pro-rated by area. Thus
the term “Puna de Atacama” covers a vast and heterogeneous area —
partly puna and partly cordilleran — which overall has been visited by
perhaps ten collectors and is too poorly known to be subdivided yet,
although it contains great riches and much endemism. Chos Malal,
184
J. Res. Lepid.
Alumine and Loncopue are dots on the map which have been visited by
only one or two collectors, but frequently and are thus well-known.
Much of the Argentine data used here is explicitly presented in the
previous section. That which is not is based on precise and reliable
sources, including the collections enumerated in the Acknowledg-
ments, the author’s collections at U.C. Davis (=AMS), and a handful of
publications. By and large, the Argentine literature is much too vague
and full of misidentifications to be credible for use in analytical
biogeography. The Chilean data are from Herrera and Field 1959,
Field and Herrera 1977, Herrera 1953 and 1970, and unpublished data
provided by Herrera, S. P. Courtney, and AMS. Chilean biogeographic
concepts are from Artigas (1975), Pena (1966), Davis (1986), Irwin and
Schlinger (1986), and Hueck and Seibert (1972). Unsatisfactory as the
data may be in scope and repeatability, they are far better than for any
other part of the Andean region except for the Satyrid faunas studied
by Michael Adams in Colombia and W. Heimlich in the Southern
Cone.
Many indices of faunal similarity exist, which allow one to compare
areas and to construct a hierarchical “classification,” or dendrogram, of
faunas based on shared elements (species, subspecies). Two recent
reviews (Janson and Vegelius 1981, Hubalek 1982) have identified
Sprensen’s (1948) coefficient as one of the best if not the best in terms of
objectivity and properties. (Although Sanchez and Lopez-Ortega, 1988,
disagree, their objections do not apply to these faunas.) From the
values in Table 3 a dendrogram can be derived by cluster analysis (Fig.
23). This approach allows an easy visualization of overall faunal
affinities, but its use for historical reconstruction is limited, as in all
phenetic methods.
The dendrogram shows few surprises, but does offer some useful
insights. It emphasizes the distinctness of the central- Andean high-
altitude faunas of the N from the Patagonian faunas, at least at the
taxonomic level used here. (The extremity would have been softened
had subspecies not been weighted equally with full taxonomic species
in the analysis.) The only northern fauna falling outside this cluster is
the Valles Calchaquies, the arid depression lying W of and in the rain
shadow of the Sierras Pampeanas, which represents the N-most ex-
tremity of the monte both floristically and faunistically (Hayward
1955b) — its fauna is small and depauperate and quite similar to that
of the Uspallata Valley (Precordillera Mendocina) in the Cuyo District;
none of the highland entities, even the altitudinal migrants, descend
into it. (If Ascia monuste had been counted, it would count as a regular
migrant crossing both highlands and valleys, cf. Hayward 1931.) These
two localities in turn cluster with the Gulf of San Jorge (Comodoro
Rivadavia), which has the rarefied desertic fauna at the opposite end of
the monte. The analysis does not include the prepuna , but its fauna is
virtually identical to that of the Valles Calchaquies. This fauna thus
wraps around the seasonally arid lower reaches of all the highlands in
28(3):137-238, 1989(91)
185
western Salta and Jujuy and in Catamarca and La Rioja.
Farther S, the N-Patagonian localities (Cordon del Viento, Loncopue,
Alumine, Bariloche, San Martin) cluster stair-step fashion. The se-
quence is reversed in the S because of the heavier Chilean influence at
San Martin. This emphasizes that faunal turnover is not a simple
function of distance from sources. In the Patagonian Andes the passes
provide variably effective access for both low-altitude moisture and the
Valdivian biota. From Bariloche south the Chilean contribution is less,
not because the passes are more difficult but because the butterfly
fauna becomes sparse in the very wet W-slope climates.
One of the more interesting clusters is of Chos Malal with the Chilean
Central Valley; the latter has a conspicuously Patagonian element in
the form of Colias vauthierii (and Yramea cytheris , etc.) not easily
predictable from climatic data. This is explicable under the dispersal
model of Caviedes and Iriarte (1989), discussed later.
In order to attempt reconstructions of specific histories, one need take
into account the identities of individual taxa shared and not shared
among faunas, the geographic distributions of individual taxa and the
relationships of the ranges among related taxa, the degree of taxono-
mic differentiation and endemism in different regions, and the re-
peating patterns of distribution of different lineages, corresponding
roughly to the “generalized tracks” of Croizat (1964; see also Craw
1982). Ultimately the phylogeny of the individual genera and of the
Andean Pierini will be resolved, but so ambitious an undertaking must
await much more data on “biology.” Within the area covered by this
study, however, patterns are already evident and in some cases their
causation may be inferred. This returns us to the objectives enumer-
ated in the Introduction.
The Uniqueness of the Puna . The high-altitude puna of NE Chile has
a remarkably large and diverse fauna of Pierini, much of it endemic at
the species level, which appears to dilute rapidly once one crosses the
crest into Argentina. Far NW Argentina is very poorly collected due to
its inaccessibility, difficult topography and harsh climate. Large areas
are not only roadless but essentially uninhabited. To my knowledge no
one has ever collected in the Sierra de Calalaste or farther W in N
Catamarca, or even from Highways 27 or 17 in W Salta or from
Highway 70/b in Jujuy. This is very bleak, barren terrain with many
salt flats. The interesting Pierini are to be expected on bogs and rocky
summits, not on the tola-co vered flats through which most of the roads
go. Thus, off-road vehicles or pack animals are required. When we do
this work, I imagine we will extend the ranges of at least part of the
Chilean puna fauna, perhaps including Infraphulia ilyodes Ureta,
Pierphulia rosea Ureta, Hypsochila penai, Tatochila mariae Herrera,
and T. distincta fieldi Herrera — just as the range of the genus
Hypsochila will surely be extended into Bolivia, where there are no
records today.
186
J. Res. Lepid.
The very rich Chilean puna fauna is a S-ward extension of that of the
Peruvian altiplano and seasonally dry high Sierra. The greatest faunal
diversity of Andean Pierini is between the Peruvian Departments of
Jumn and Cusco, with perhaps the maximum diversity of altiplano
taxa in Arequipa and of Tatochila in the complex ecogeography incor-
porating y ungas, jalca~“ paramo” in the Peruvian sense (which is not
the same as its use in Colombia, Venezuela and perhaps Ecuador), and
altiplano within short (air) distances. These faunas appear to rarefy to
the E in Bolivia, but collecting has been so spotty that little can really
be said. Within the puna and altiplano the small Pierini (except
perhaps Phulia nymphula ) are restricted to bogs or bog margins at
least in dry season (Shapiro 1985, 1986a, Shapiro and Courtney 1986).
This limits their dispersal and seems to have promoted local differen-
tiation and subspeciation. Infraphulia madeleinea females are very
poor fliers — I. ilyodes are somewhat better — and electrophoretic as
well as morphological data suggest that different allopatric popula-
tions of small pierines are evolving in isolation from one another.
Despite seasonal variability, many of the bogs are certainly of Pleisto-
cene age and, like Nearctic bogs supporting relict butterflies, have
been able to sustain isolated populations for millenia. The bog illus-
trated by Weberbauer (1945, pi. XIII, p. 391) at Morococha (4500 m)
has an exceptionally rich fauna and had not changed visibly in the 80-
odd years since the picture was taken; pasturage does not appear
harmful.
Phulia nymphula is the most ecologically versatile and dispersive of
the small Pierini, in Peru (Perez 1982, Lamas and Perez 1983) as
elsewhere; it is not surprising that it is so widely distributed in N
Argentina (see below).
The Quichua District and the Sierras Pampeanas. There is great
similarity between the high-altitude faunas of Salta and Jujuy and
that of the Pampean Sierras ( Cumhres Calchaqmes — Sierra de Aeon-
quija ) in Tucuman and Catamarca. The affinities of these high-
altitude faunas, in turn, are with the puna. Phulia nymphula ( aconqui -
jae) is very instructive. In the puna above the Quebrada de Huma-
huaca, P. nymphula flies at the same elevation (albeit often in different
habitats) as Tatochila m. macrodice, T. d. distincta, T. inversa, and C.
blameyi. In the Sierras Pampeanas it occurs mostly above those species
— mostly above 4000 m, though descending in autumn (March-April)
as low as 3100 m (as noted by Jorgensen, 1916). It is not clear whether
this altitudinal stratification in the Sierras Pampeanas reflects clima-
tic differences or merely the occurrence of Phulia habitat — sandy
alpine grassland-steppe with only very low plants — mainly above the
Tatochila concentrations, which occur on rocky substrates below the
glacial-pothole landscapes. As noted under that species, Pampean
Sierra P. nymphula might be recognized as a subspecies. The same is
true of T. inversa , T. distincta and perhaps even T. m. macrodice , and,
28(3):137-238, 1989(91)
187
as noted above and in Shapiro (1989b), topotypieal Colias blameyi seem
to grade into C. weberbaueri both NW and N of the Sierras Pampeanas
in the Quichua District. The distances are not great (100-150 km)
between the highlands of Salta and Jujuy and the Sierras Pampeanas ,
but the ecological barriers are formidable. These barriers are traversed
annually by Ascia monuste automate and perhaps by Teriocolias , as
well as by many other seasonally migrant Lepidoptera. It seems
unlikely that any of the true high-altitude Pierids are presently
moving between the ranges. This might have been much easier in the
Pleistocene. Halloy (1978, 82, 83) provides the most in-depth descrip-
tion and analysis of the Cumbres Calchaquies from ecobiogeographic,
physiological, and paleoclimatic perspectives. He finds that from a late
Pleistocene glacial maximum, they are continuing to re warm, and the
snow line is still regressing upslope. If we know little of the regional
biota before the Quaternary, we can still infer that the differences
between populations in the Sierras Pampeanas and the main cordillera
and puna are unlikely to antedate the later Quaternary. Using this
inference, it may be possible to calibrate “molecular clocks” for the
Andean Pierids more precisely than has been done heretofore. All
these mountains attained alpine heights only in the Plio Pleistocene
Are the insects conceivably any older?
It is very striking that the genus Hypsochila appears to be absent in
the well-collected Sierras Pampeanas , eluding Giacomelli, Jorgensen,
Hayward, Halloy, Dominguez, and me. Hypsochila is also unrecorded
from Cerro Zapallar (Cuesta del Obispo). In fact, there are no records
to connect up. H. w. wagenknechti of the temperate cordillera of the
Cuyo District with H. w. sulfurodice of the puna (San Antonio de los
Cobres, Quebrada de Humahuaca). This is probably significant. It also
casts doubt on the true conspecificity of the galactodice- like specimens
from the puna and those from the main range of that entity far to the S
in Patagonia. Indeed, it is by no means certain that wagenknechti and
sulfurodice are conspecific. The zone where no Hypsochila occur is the
zone of heaviest seasonal precipitation in the uplands, precisely where
the tropical northeasterlies are wrung out at the head of the y ungas.
This may well be a limiting factor on Hypsochila , which is happy in the
extreme aridity of the summits above Abra Mariano. It may also
contribute to the upslope displacement of Phulia in the Cumbres
Calchaquies.
The seasonal presence of Tatochila orthodice and stigmadice in these
same high-precipitation areas reinforces the impression that contem-
porary climate exerts strong control over contemporary distributions.
These are true y ungas species, which do occur in the wetter parts of the
true puna close to the tops of the big canyons, without penetrating into
tolar es.
In summary: the wet alpine reaches of the Sierras Pampeanas share
their fauna of Tatochila and Colias with the wetter parts of the
Quichua District in Salta and Jujuy; the altitudinal distribution of
188
J. Res. Lepid.
Phulia is somewhat skewed; and Hypsochila is absent from both,
though common in the drier parts of the Quichua District and in the
puna. These patterns were probably created in the late Pleistocene and
are controlled today by micro- and mesoclimate distributions.
Connections Between the Cuyo and the Quichua District and Puna.
The fauna of the high cordillera in Mendoza and San Juan is depauper-
ate. I have not included Colias mendozina in the analysis only because
it was rediscovered as this paper was about to go to press. It is the most
unusual faunistic element in the Paso Bermejo but at present we
cannot say whether it is more closely related to C. blameyi of the puna
or to C. flaveola. (It is striking that if C. blameyi , flaveola and men-
dozina are combined, their range approximates that of Phulia nym-
phula sensu lato which, as noted previously, is differentiated into
several segregates adapted to either tropical or temperate seasonality.)
The only true puna element found in the alpine elevations of the cuyo
is Phulia nymphula. Hypsochila wagenknechti is present as a presump-
tive sister-subspecies of the puna sulfurodice , and there may be either
T. m. mercedis — T. m. macrodice hybrids or intergrades in San Juan.
Colias flaveola seems limited to a very narrow latitudinal band on both
sides of the Andes. Although Descimon (1986) treats it as potentially
conspecific with blameyi , weberbaueri, etc., there is little reason to do
so. Thus, the very rich puna Pierid fauna rarefies very rapidly once the
regime of the tropical “Bolivian Winter” is replaced by the true cordil-
leran winter with a more or less continuous snow pack which forces a
several-month interruption in butterfly activity. Since Phulia nym-
phula has adapted its life-cycle to the mendocino winter it is unclear
what limits its range farther S, but it seems to drop out before the
Maule district. H. w. wagenknechti continues S to Laguna del Maule
and seems to segue into H. galactodice in NW Patagonia in a manner
which remains unclear (see below). The cordillera was very heavily
glaciated in the cuyo , and at present it cannot be told whether P.
nymphula reinvaded from the NW after deglaciation or had been
depressed to lower elevations, such as the precordillera above Uspah
lata, and reinvaded upslope.
The Las Lenas fauna (35°04'S, 70°02'W) is very unusual and interest-
ing in presenting an altitudinally-stratified mix of northern and south-
ern elements, including P. nymphula and H. wagenknechti above
3000 m and H. galactodice and Colias vauthierii below. The same phen-
omena are reproduced in the Lycaenid genus Itylos (Shapiro, unpub-
lished data). The habitats of P. nymphula on Cerro de los Fosiles are
very local and specialized in aspect and vegetation. One might expect a
puna species to occur at ever-lower altitudes to its S limit outside the
tropics, but the known P. nymphula sites on Cerro de los Fosiles are all
higher than its lowest sites in the Paso Bermejo further N, and it is not
a vega species as it often is in the Paso Bermejo. Indeed, its habitats
28(3): 137-238, 1989(91)
189
near Las Lenas are more like those where it has been taken in the
Cordon del Viento.
The Cordilleran - Cordon del Viento Connection . This isolated, high,
arid pre-Andean range, dominated by the 4710 m Volcan Bomuyo, has
the S-most known population of Phulia nymphula and also has a
Hypsochila , unfortunately thus far known from only one male, which
may be galociodice or wagenknechti or something in be tween The
distance to the nearest Phulia known in the cordillera is between 175
and 225 km. The presence of Hypsochila here is especially interesting
in view of its absence from precordilleran ranges further N. Again,
either these taxa were depressed to low elevations during the Pleisto-
cene or colonized in the 'past 10 -15 .000 years. The distances involved
imply the former as more likely, especially for Phulia . Thus, these
high-altitude taxa may at one point have entered quite low parts of far
NW Patagonia (the Payunia district), where we now find a mosaic of
Patagonian and monte elements segregating by micro- and meso-
habitat.
Geography of the Tatochila mercedis complex . It is quite clear that T.
m. macrodice is the true central Andean representative of this polyty-
pic species, and most closely related to T. m. arctodice of the far N. In N
Argentina as well as in Peru and Bolivia it has a remarkable ecological
amplitude, occurring in both wet and very dry climates at high alti-
tude. Its great vagility undoubtedly contributes to this, and it migrates
elevationally with season in much of its range. Both it and arctodice
are adapted to tropical seasonality, and its absence from the cordillera
in the Cuyo District (despite the queried Mendoza record, not in
Hayward 1950) is consistent with this: it has no place to go in winter
and (in the laboratory) does not seem to be able to diapause. Diapause
was, however, evolved as the complex invaded the temperate climates
of the Southern Cone. The non-diapausing macrodice comes spatially
very close to T. m, vanvolxemii , which diapauses in winter, along
Highway 307 between Abra Infiernillo and Amaicha, but no actual
contact has been found and the two are separated by a zone in which
neither seems to occur.
Otherwise, T. m. vanvolxemii contacts and exchanges genes with T.
m. sterodice and T. m sterodice — mercedis hybrid populations along
the W and SW edges of its range in Neuquen and Rio Negro; its
populations in the Mendoza precordillera (e.g., Potrerillos) and below
Copahue and W of Chos Malal show signs of past or perhaps present
introgression from m. mercedis , though the populations at both
Mendoza and Chos Malal themselves appear pure; and the population
in the Gulf of San Jorge warm pocket (Comodoro Rivadavia) presents a
normal summer vanvolxemii phenotype but its cold-season brood dis-
plays strong sterodice influence. Intergrading populations in the San
190
J. Res. Lepid.
Jorge District are clearly dependent on local populations of adventive
Cruciferous weeds and cannot be very old — indeed, their ruderal
character argues that the intergradation itself is likely to be secon-
dary, and of recent origin.
True T. m. mercedis enters Argentina from Chile in NW Patagonia,
but it does not seem to occur anywhere where it does not contact and
hybridize with T. m. sterodice. Its range in Chile is basically limited to
the Mediterranean climate belt, from Copiapo to Valdivia, and it does
not occur in the “pampas” of NE — C Chile or approach T. m.
macrodice closely anywhere except perhaps in Coquimbo and San
Juan. The oldest museum specimens of T. m. mercedis from NW
Patagonia date from 1939, with intergrades already present — so the
phenomenon has a minimum age of some 50 yr, or 150 generations, but
deforestation and land-use patterns suggest it could be at least twice
that age if it in fact is a recent, man-influenced phenomenon.
Of all the Argentine Pierids, this complex most accurately mirrors
the phytogeography. It also has the most extensive distribution of any
Pierid, and probably any butterfly, on the continent — from central
Colombia to Ushuaia, though avoiding forested and lowland-tropical
habitats.
The Relationships of Fuegia and their Evolutionary Implications.
Although Fuegia is recognized as a separate phytogeographic entity,
its butterfly fauna is less distinct than heretofore supposed — though it
has more Pierid taxa than the Cuyo District! The allegedly endemic
Tatochila and Hypsochila are either also found on the mainland
(■ argyrodice , microdice ) or are clinal, with no sharp step at the Straits of
Magellan ( theodice ssp., sterodice /fueguensis) . These facts argue
against postglacial higher sea levels as responsible for isolation and
differentiation-by-vicariance in the far S. But interesting evolutionary
problems remain.
The Quaternary history of Tierra del Fuego is exceptionally well-
known (Auer 1956, 1958, 1965, 1966, 1970). All of the mountains were
glaciated; glacial retreat began some 16,000 BP. The entire vegetation
of Fuegia can be assumed to have developed in this period, and it is not
surprising that only 3% of the Fuegian flora of 545 vascular plant taxa
is endemic; some 64% of the flora occurs N up the Andes on both slopes,
2% only on the drier Argentine side, 4% only in Patagonia E of the
Andes and 8% N in Chile only (Moore 1983). The Fuegian Pierid fauna
is overwhelmingly a steppe fauna; all the species occur on the Pata-
gonian mainland in steppe, though two ( Tatochila theodice and T.
sterodice) also occur in the subhumid Patagonian Andes. T. t. theodice
is recorded in Chile S to Victoria, at about the same latitude as
Alumine, but the species apparently continues S along the E slope of
the Andes, grading into ssp. gymnodice and thence into ssp. standing -
eri in Fuegia. The type locality of gymnodice is Punta Arenas, Magal-
lanes, which further confuses the situation! T. m. sterodice is not
28(3):137-238, 1989(91)
191
recorded unambiguously in Chile at all, though again the Magallanes
populations are part of a cline between it and fueguensis. None of this
should be surprising, were the 10,000-yr-old Straits of Magellan not
taken unduly seriously as a barrier. Moore (op. at., p. 33) says: “The
four principal climatic and vegetation zones described in passing from
N and E to S and W Fuegia . . . parallel rather closely the sequence seen
N of the Estrecho de Magallanes in traversing cool temperate Argen-
tina and Chile from the Atlantic to the Pacific Oceans. Since all plant
species can only occupy areas with climatic and ecological conditions
within their range of tolerance it is not surprising that some species
restricted to the drier steppe areas of NE Tierra del Fuego extend N-
wards in the drier parts E of the Andes ...” The only question is
whether the Pierids crossed into Tierra del Fuego before or after there
was a water barrier. In any case, the taxa are both weak and young
and the climate unstable (Markgraf 1985).
More interesting is the hostplant specialization of those taxa. All
three subspecies of Tatochila theodice feed on Legumes. So does
Hypsochila microdice (and probably also H. argyrodice and H. huemul ,
which seem very closely related). How are we to interpret this oddity in
a Crucifer-feeding lineage? Of all Tatochila , distincta is closest to
Hypsochila in genitalia (Field 1938, Herrera and Field 1959, Field and
Ferrera 1970). This has not been resolved electrophoretically (Shapiro
and Geiger, unpublished). If they were sister-taxa, one could then treat
Legume-feeding as a symplesiomorphy, and Crucifer- Tropaeo/wm feed-
ing would then have arisen independently in both lineages, and repre-
sent convergence. But this is very improbable. The fact that most
pierines globally feed on mustard-oihcontaining plants argues against
it. So does the remarkably close resemblance of the early stages of the
Crucifer- Tropaeolum feeding branch of Hypsochila , which includes
galactodice and wagenk nenchti , to the Crucifer-feeding Tatochila of
the autodice and mercedis complexes. (T. theodice is highly divergent
in its early stages, and its affinities may actually lie with the
orthodice s tigmadice end of the genus, a notion supported by certain
pattern characters and wing pigments of the adults, and karyotypes
(deLesse, 1967; Shapiro, unpublished).) If Crucifer-feeding is the sym-
plesiomorphy, Legume-feeding has originated separately as a deriva-
tive condition in both genera. It is not clear whether it arose in
sympatry, however, Legume-feeding may have arisen more than once
— perhaps three times — in Tatochila . These conundrums can be
resolved only when the entire group of genera can be subjected to a
thorough cladistic analysis incorporating early-stage data.
Some Legume-feeders have austral distributions. If they were derived
from more northerly Crucifer-feeders this makes sense: although Cru-
cifers extend at least as far S in Fuegia as Legumes and are actually
more numerous, their distribution is very patchy and their biomass is
much less than that of small vetches on the steppe. (Introduced weedy
Crucifers are now extremely abundant around towns, but this is a
192
J. Res. Lepid.
recent condition.) Herrera and Covarrubias (1983) claim that the
Andean-Patagonian Pierini are of Gondwanaland origin, however —
suggesting that Legume-feeding (discovered since 1983) would repre-
sent the primitive condition. Is there any independent support for the
Gondwanaland claim? The only suggestive datum is the bizarre pro-
venance of Colias ponteni.
Biogeographers have failed to reach consensus over the interpreta-
tion of “centers of origin,” a fact which has exposed the entire field to
ridicule (Cain 1944, Croizat et al. 1974). The location of an endemic
primitive taxon is not a reliable indicator of a center of origin, because
primitive forms often survive in isolated regions far from where they
originated (supposedly because of a lack of competitors). Any biogeo-
graphy text gives several examples, the Tuatara ( Sphenodon ) being
one of the commonest citations.4 Colias ponteni is, genitalically, easily
the most primitive living (or recently extinct?) member of its large
genus. (It was classified by Peterson, 1963 in its own genus, Proto -
colias.) Is its inferred endemism in Tierra del Fuego an indicator that
the entire genus is of Gondwanaland origin? Its nearest relative seems
to be vauthierii , which occurs in Patagonia and the Central Valley of
Chile. Electrophoretically all the South American Colias seem to
cluster together, including vauthierii (Descimon and Geiger, pers.
comm.). There is no indication in these data that vauthierii is the stem-
species of the monophyletic Andean group inferred by Descimon
(1986), or the group’s sister-taxon. Nor do the global data on Colias
support a Gondwanaland origin for the genus: the only other austral
Colias , C. electo L., has a spotty relict distribution in east, central and
southern Africa and belongs to a Palearctic species-group.
The Limits of Patagonia. The traditional political limit of Patagonia is
the Rio Negro, but there has never been a formal jurisdiction named
“Patagonia” and in any case the biota on one bank of the Rio Negro does
not differ from that on the other. The Argentine phytogeographical
literature attempts to define boundaries in the low, undulating relief of
N Patagonia where it contacts and intergrades to the monte and pampa
(Morello 1958, Ragonese and Piccinini 1969). From a pierid stand-
point, Patagonia extends into the river bottoms of NW Neuquen (the
Payunia district) which form the N limits of Colias vauthierii and some
4 This concept was articulated at least as early as 1752 by Maupertuis, who wrote with
reference to the alleged occurrence of “giants” in Patagonia (Tehuelches or Ona) and
“dwarves” in the north-polar regions (Eskimos): “If there is truth in what the travelers ;
tell us of the Strait of Magellan and the lands of the far North, the races of giants and
dwarves settled there because of the fitness of the climate or, what is more likely,
because. . .they were driven to those regions by other men, who feared the giants or
scorned the pygmies. . ( Origine des Animaux, p. 266) Although the Patagonian giants
were ultimately debunked (Adams 1962), the existence of Colias ponteni in the fairly
recent past is supported by a handful of specimens.
28(3):137-238, 1989(91)
193
Lycaenids and Hesperiids as well, and form part of the Tatochila
mercedis-sterodice-vanvolxemii intergradation zone. In the reciprocal
sense, the monte and pampa penetrate Patagonia as far as the genetic
influence of T. m, vanvolxemii extends, i.e. the Gulf of San Jorge. The S
limits of Colias lesbia (as an occasional seasonal breeder) and Eurema
deva (as an immigrant) are in the same area. It would be instructive to
use Satyr ids to define the limits in these same areas, as they are
grassland-associated while the Pierids are not necessarily so.
Quaternary Climate Dynamics and Dispersal Routes. During the Qua-
ternary the extreme aridity which characterizes the coastal deserts of
Peru and northern Chile developed, serving as a very effective barrier
to contain the biota of the Andean highlands. At the same time, there
is reason to believe that precipitation waxed and waned repeatedly in
temperate Chile, resulting in pulsations of N- ward migration by the
Valdivian rain forest. Relict elements of that forest exist today in such
areas as Fray Jorge, Talinay and Mantagua in Coquimbo at roughly
30°S, near the N limits of many organisms of central Chile including
the butterflies Tatochila mercedis and Colias vauthierii. The best
butterfly indicator of the Valdivian forest, Eroessa chiliensis , seems to
stop at about 35°S (Constitution). Caviedes (1990) provided a predic-
tive model of precipitation which generates up to 9-fold increases in
rainfall in central Chile with T depressions of 3°C or less. If these
calculations are valid, the gradient between the desertic Atacama
climate and the much wetter central Chilean climate must have been
extremely steep at times in the Quaternary (Paskoff, 1977). Caviedes
and Iriarte (1989) have used these projections to develop a verbal
model which accounts for the distribution of faunal richness of Cricetid
rodents and possibly other mammals in Chile and Argentina. They
conclude that the Atacama barrier would have held in the rich, diverse
and highly endemic Cricetid faunas (cf. Pierini), forcing whatever
dispersal occurred to have been down the eastern (Argentine) side of
the mountains in the relatively mesic climates influenced by _ the
tropical flow from the NE. During the periodic episodes when Valdi-
vian vegetation migrated N-ward to the vicinity of 30° in Chile, many
of the trans-Andean passes would have been available for faunal
migration from E to W. The limited Chilean fauna S of the Atacama
can be derived from such movements.
Cricetids resemble butterflies in having greater faunal richness on
average E of the Andes, but the resemblances break down on detailed
examination. Pierids (except Eroessa and Mathania) are not forest
species and indeed would be excluded from extensively-forested areas.
There is no Pierid fauna in archipelagic Chile, and little butterfly
fauna at all. The same phenomenon can be seen in the North American
Pacific NW rainy belt. For the Cricetid model to account for the
Patagonian character of the Chilean Central Valley fauna, the neces-
sary assemblage would have to have been far enough N on the Argen-
194
J. Res. Lepid.
tine side to have crossed the passes between 33-30°S (including the
Bermejo) when they were sufficiently mesic to be good corridors. There
is no concrete evidence bearing on this. See also Heusser, 1983 and
Veblen et al. 1981.
What we do know of butterflies crossing the Andes is based largely on
the passes in the Lake District, which today are mesic and penetrable
but were icebound in the Pleistocene. All the movement we can detect
has been from W to E in these passes, following climate and prevailing
winds. Thus, the hybrid-zone phenomena described for Tatochila
mercedis and autodice are presumably post-Pleistocene in origin. It is
striking, however, that populations of T. m. vanvolxemii from further
N (Copahue and Paso Bermejo-Potrerillos) show signs of previous gene
flow from nominate mercedis over the crest, though the genitalic
morphology that defines such contacts farther S has disappeared. This
presupposes the existence of mercedis in Chile at least during one
fairly recent more mesic interval, and thus implies latitudinal oscilla-
tion of ranges. The stranded high- Andean taxa in the Cordon del
Viento imply a retreat N-ward with recent drying, as do the ranges
of the high-altitude taxa in Salta, Jujuy and Tucuman. (Cei, 1980
discusses some possible biogeographic roles of the yungas in the
Quaternary.)
“The Argentine provinces of Mendoza and San Juan report 2.5 times
as many mammal species as the Chilean provinces at equivalent
latitudes” (Caviedes and Iriarte 1989). These provinces are, however,
very depauperate insofar as Pierid butterflies are concerned. Although
both the timing of S-ward movement and the general routes of dis-
persal may be similar for Pierids and Cricetids, it is still necessary to
account for such striking contradictions in their distributions.
Perspectives on Faunistics. Hayward (1955c) noted that high- Andean
elements penetrated the cuyo in the Uspallata Valley. Despite the
unusually clear overall phytogeographic patterns defined by climate,
any attempt to define precise boundaries for the Argentine floras and
faunas is bound to be frustrated by the individualistic nature of species
distributions. Both the high- Andean and Patagonian floras show abun-
dant evidence of relative youth and evolutionary dynamism, while the
Valdivian forest is clearly relictual and ancient. The butterfly faunas
treated in this paper have little connection to the Valdivian forest
(except Eroessa chiliensis ), and no butterfly in the region — Pierid or
otherwise — is tied by identified sister-group relationships to any
other austral region so as to suggest a Gondwanaland-based history.
The nearest relative of Eroessa is by no means clear. Klots (1932)
correctly identifies it as exceedingly primitive in genitalia, venation
and form of palpus. Electrophoretically it comes closest to the east-
Asian and Indian genus Hehomoia Hbn., which shares one probable
apomorphic trait with Hesperocharis Felder (including Mathania ); few
taxa have been examined in this group of genera (Geiger and Shapiro,
i
28(3):137-238, 1989(91)
195
unpublished). There are, however, a variety of suggestive trans -Pacific
linkages in Pieridae,. which if substantiated would imply considerably
greater antiquity for the modem butterflies than has been established
in the fossil record; these will be discussed elsewhere.
The traditional geochronology of the Andes has the cold-adapted
high-altitude biota, which shows a dominance of circumboreal taxa at
the family level (Van der Hammen and Cleef 1987, Raven and Axelrod
1974), entering late in the Pliocene when the latest uplift reached
adequate heights to support such a biota. By this view, these organ-
isms have undergone extensive adaptive radiation since the initial
phases of the Great American Interchange (Stehli and Webb 1985). It
is increasingly clear that this scenario is too simple, and probably
largely wrong. More groups have been shown in the fossil record to
have arrived in South America from elsewhere before the Interchange,
and, more importantly, the amount of differentiation in some high-
Andean groups appears too great to be accommodated in so little time.
This very large literature is summarized by Briggs (1987), who empha-
sizes the complexity of patterns observed in different taxonomic groups
and their inferred differential antiquity on the continent. The various
genera of Andean pierines seem unlikely to have originated and
diversified within only 2 or 3 MY — especially since our estimates of
the antiquity of speciation based on electrophoretic differences tend to
fall close to that number (Geiger and Shapiro, unpublished; compare
Shapiro and Geiger, 1989 for Vanessa across the Isthmus of Panama).
But how could cold-adapted genera antedate cold environments? It is
of course possible that they arrived on the suggested schedule, but
already generically differentiated elsewhere. This requires multiple
colonizations by small and weak-flying animals over very great dis-
tances. Given the strong morphological affinities to the Asiatic alpine
genus Baltia Moore, affinities which appear real and not convergent
(Field 1958, though the electrophoretic data, such as they are, are
ambiguous — Geiger, Michel and Shapiro, unpublished data), some
kind of Matthew (1915) “Camelid scenario” appears required. By this
view the Asiatic and South American genera were linked across North
America and might even have originated there, but subsequently went
i extinct there. We are unlikely to get fossils to vindicate this scenario,
as they have for the Camelidae. But recent evidence (Mercer and
Sutter 1982, Clapperton 1983) strongly suggests that Patagonia
underwent episodes of glaciation some 7 and 4.6 MYA, before the Ice
Ages began in the N Hemisphere (except perhaps in Alaska). This
lengthens the potential time-line for the evolution of adaptation to cold
in South America, without clarifying the question of where the Pier ini
came from or who their ancestors might have been.
The genus Tatochila is apparently evolving very rapidly right now
throughout the Andes, with all sorts of speciation-related phenomena
visible, a “ferment of variability” as Dunbar (1968) described the
situation in Arctic crustaceans. For a North American butterfly
196
J. Res. Lepid.
worker it is strikingly reminiscent of the situation in the machaon L.
group of Papilio — the entities are often not fully speciated, intergrade
in complex ways, form hybrid zones, produce local subspecies in some
areas and not others, and in general look like sequelae of the Pleisto-
cene (compare Sperling 1987). The data for Hypsochila are only begin-
ning to accumulate, but clearly suggest a group in similar if less
extensive ferment. The Phulia group of taxa are perplexing in that
morphospecies tend to appear older than in Tatochila, but local popula-
tions of the Phulia nymphula complex are clearly evolving in a variety
of directions, and the various Infraphulia and Pierphulia look similar.
For all these taxa, the genera and subgenera certainly antedate the
Pleistocene, speciation may often also, but subspeciation seems to have
been working overtime in the Quaternary.5
The ultimate interpretation of these radiations will depend on our
ability to develop a convincing reconstruction of pierine phylogeny and
to determine whence came the stem-species of the various radiations.
At present we can say reasonably confidently that Tatochila , in its
current broad sense, has undergone several radiations, all of which
may still be in progress. Certainly the two southernmost (mer cedis and
theodice ) are, and the next southernmost 0 autodice ) is clearly quite
active too. The central- Andean Jy ungas taxa are so beset with taxono-
mic problems that we may infer activity there too, even if we do not
understand it. In the far N, T. xanthodice Lucas is actively differentiat-
ing in different ranges (Ackery 1975). Although we are still not in a
position to interpret the extreme rarefaction of the Andean pierines in
the N, we can say it is not de facto evidence of Gondwanaland origins
for the group. Rather, it suggests spread both N and S from a central-
Andean center for several genera, though that “center” may not be
where they originated.
The Fuegian and far-S Patagonian taxa range from the probable
ancient relict Colias ponteni , to recently-derived and still only weakly-
differentiated subspecies in Tatochila which as noted above may ante-
date the inundation of the Straits of Magellan, but perhaps not by
much. The mere fact of remoteness in far-southern South America may
5 The maximum antiquity of T. m. vanvolxemii and other taxa of the monte can be
inferred from recent discussions of the evolution of aridity in Argentina. Volkheimer
(1971) inferred humid climates in Patagonia except perhaps semiarid in the modern
pampa region in the early Tertiary, and again in the Miocene. Axelrod (1979) and
Sarmiento (1975) both consider the monte and the Patagonian steppe as consequences of
the Plio-Pleistocene elevation of the Andes, disagreeing with Solbrig (1976), who feels
that arid climates might have existed longer. Since vanvolxemii is highly derivative vis-
a-vis its relatives, it almost certainly has originated within this time frame. Axelrod ( loc .
cit.) accepts the spotty evidence from Bolivia of an emerging xerophy tic- microphy 1 lous
vegetation there perhaps as early as the late Eocene, but in any case well-defined by the
Miocene. The Phulias could thus conceivably go back that far, but whence came their
ancestors?
28(3):137-238, 1989(91)
197
have preserved Colias ponteni and Eroessa chiliensis — which once
may have ranged much more widely, and may indeed have originated
elsewhere.
Biogeographers tend to believe what geoscientists tell them about the
antiquity of continents and orogenies, a dangerous tendency (though
some biogeographers championed continental drift when the geoscien-
tists mainly considered it impossible). In interpreting the distributions
and phylogenies of Andean organisms we have always attempted to
ram everything into the short time since the late Pliocene, because
geoscientists said we had to. Now some glaciologists (Clapperton,
Mercer) hold out the possibility of a considerably longer time-line (at
least 4, perhaps 6 or 7 million years). Just what this implies for us is
still unclear. Nothing in the butterfly data yet requires a longer
Andean time-line; thanks to the Baltia connection we can always make
up stories about generic differentiation having occurred elsewhere.
And it is clear that much of the evolutionary activity in the Andean-
Patagonian Pieridae is very young, certainly of Holocene origin. Table
4 summarizes what we might know at the moment, which in terms of
data is far greater than what Giacomelli (1915) knew when he pub-
lished his concluding table, yet not much more definitive!
There is one further approach to these problems, which has not been
addressed here. It is strictly ecological and ahistorical and in the
“MacArthur tradition,” exemplified by the analysis of the Argentine
passerine bird fauna by Rabinovich and Rapoport (1975). Neither the
size of the Pierid fauna nor the completeness of ground coverage
justifies such an analysis at this time. It may ultimately be useful, but
it should be obvious that such an approach ignores many of the most
interesting and evolutionarily exciting phenomena of faunistics.
Now that we have learned a little more, we can join Giacomelli in
lamenting the profundity of our ignorance. Giacomelli wrote even as
his compatriots Florentino Ameghino and Francisco P. Moreno were
beginning the process of opening up the inescapably complicated
geohistorical context in which the Argentine Pierid fauna is imbedded.
The ultimate fruits of their labor can be seen in the informed nature of
our modern, profound uncertainty.
Acknowledgments. This work has been supported by the National Science
Foundation (USA) through grants DEB-76-18611 and BSR-83-06922: by a
grant from NSF (International Programs) for a scientific visit to Argentine
institutions; by the National Geographic Society through grant 2263-80; by
grant OPER-46 to A. M. S. and D. Helgren from the Institute of Ecology, U. C.
Davis; and by the Department of Zoology, U.C.D. It has benefitted from the
help, advice and insights of colleagues and friends, including (in alphabetical
order): Dr. Alberto Anziano, Lie. Martha Arce de Hamity, Dr. Francisco
J. Ayala, Dr. Keith S. Brown, Jr., Dr. Michael M. Collins, Dr. Steven P.
Courtney, Dr. Henri Descimon, Dr. Eduardo Dominguez, Rev. Robert C.
Eisele, Sr. Andres Garcia Hidalgo, Dr. Hansjurg Geiger, the entire Mario
198
J. Res. Lepid.
Gentili family, Dr. Stephan Halloy, Dr. Jose Herrera G., Dra. Mary Kalin
Arroyo, Dr. Clinton V. Kellner, Dr. Gerardo Lamas M., Dr. Frangois Michel,
Dr. Enrique Monaglio, Dr. Roberto Murua, Lie. Estela Neder de Roman, Dr.
Oliver Pearson, Dr. Adam H. Porter, Dr. Timothy Prout, Dr. Eduardo Rapo-
port, Sr. Jose Maria Saenz Cabezon, Dr. Ebbe Schmidt Nielsen, Ms. Margaret
J. Stern, Dr. Michael Turelli, Dr. Philip Ward, and Dr. Abraham Willink. The
California Department of Food and Agriculture and the U.S. Department of
Agriculture kindly expedited permits to import livestock for genetic and
physiological study. Specimen photographs are by Samuel W. Woo and line
drawings by Adam H. Porter. Logistic support was kindly provided by Mr.
Robert O. Schuster of the Bohart Museum of Entomology at U.C.D. The
directors and curators of the following institutions extended their hospitality
and permission to work in their collections: Zoological Museum, University of
Copenhagen (DE); British Museum, Natural History (BM); California Aca-
demy of Sciences, San Francisco (CAS); Instituto Patagonico de Ciencias
Naturales, San Martin de los Andes (MG); Museo Nacional de Ciencias
Naturales “Bernardino Rivadavia,” Buenos Aires (MR); Instituto Miguel Lillo,
San Miguel de Tucuman (ML); Universidad Nacional de Jujuy, Instituto de
Biologia de la Altura; Fundacion Bariloche; Universidad Nacional de La Plata,
Museo de La Plata (MLP). Most of the remaining referenced data are from the
author’s collection at U.C.D. (AMS) and the private collection of Rev. R. C.
Eisele (RE). None of my work would have been possible without the continuous
support of Adrienne R. Shapiro; the intellectual challenges constantly posed by
my undergraduate and graduate students and postdocs, and the warmth and
kindness of the Argentine people, who have never failed to make me feel at
home, from La Quiaca to Ushuaia.
Figures 3, 4 and 5 are reproduced from Geografisk Tidsskrift by permission of
the Editor, Dr. N. Kingo Jacobsen. R. Barneby and J. McCaskill assisted with
plant determinations.
Literature Cited
ACKERY, P. R. 1975. A new pierine genus and species with notes on the genus
Tatochila (Lepidoptera: Pieridae). Bull. Allyn Mus. 30:1-9.
ADAMS, M. J. 1973. Ecological zonation and the butterflies of the Sierra Nevada
de Santa Marta, Colombia. J. Nat. Hist. 7:699-718.
, 1977. Trapped in a Colombian Sierra. Geogr. Mag. 49:250-254.
, & G. I. BERNARD. 1977. Pronophiline butterflies (Satyridae) of the
Sierra Nevada de Santa Marta, Colombia. Syst. Entomol. 2:263-281.
, 1979. Pronophiline butterflies (Satyridae) of the Serrama de Valle-
dupar, Colombia — Venezuela border. Syst. Entomol. 4:95-118.
, 1981. Pronophiline butterflies (Satyridae) of the Cordillera de
Merida, Venezuela. Zool. J. Linn. Soc. 71:343-372.
ADAMS, P. G. 1962. Travelers and Travel Liars 1660-1800. University of
California Press, Berkeley. 292 pp.
ANGULO, A. O. & G. T. WEIGERT. 1974. Estados postembrionales de Eroessa chilen-
sis (Guerin) (Lepidoptera: Pieridae). Bol. Soc. Biol. Concepcion 47:49-56.
ARTIGAS, J. N. 1975. Introduccion al estudio por computacion de las areas
zoogeograficas de Chile continental, basado en la distribucion de 903
especies de animales terrestres. Gayana Miscelanea 4:3-25.
28(3): 137-238, 1989(91)
199
AUER, V. 1956. The Pleistocene of Fuego-Patagonia. I. The ice and interglacial
ages. Ann. Acad. Set. Fenn. A (III), #45.
; 1958. The Pleistocene of Fuego-Patagonia. II. The history of the flora,
and vegetation. Ann. Acad. Sci. Fenn. A (III), #50.
, 1965. The Pleistocene of Fuego-Patagonia. IV. Bog profiles. Ann.
Acad. Sci. Fenn. A (III), #80.
, 1966. Climatic variations in Fuego-Patagonia. pp. 37-55 in D. L
Blumenstock, ed., Pleistocene and Post-Pleistocene Climatic Variations in
the Pacific Area. Bishop Museum Press, Honolulu.
. 1970. The Pleistocene of Fuego-Patagonia. V. Quaternary problems
of southern South America. Ann. Acad. Sci. Fenn. A (III), #100.
AUTOMOVIL CLUB ARGentinq. 1969. Carta Turistica. Hoja de Zona 5 (Mendoza,
La Parnpa, Neuquen y Rio Negro). Publ. #289, Buenos Aires.
, 1972, Carta Turistica. Hoja de Zona 3 (Catamarca, Santiago del
Estero, La Rioja, San Juan, San Luis, Cordoba, Mendoza). Publ. #407,
Buenos Aires.
AXELROD, D. I. 1979. Desert vegetation, pp. 1 = 72 in J. R. Goodin and D. K.
Morthington, eds. . Arid Land Plant Resources. Lubbock, Texas.
BARTON, N. H, & G. M. HEWITT. 1985. Analysis of hybrid zones. Ann. Rev. Ecol.
Syst. 16:113-148.
BEETLE, A. A. 1943. Phytogeography of Patagonia. Bot. Rev. 9:667-679.
BERGER, L. A. 1986. Systematique du genre Colias F. (Lepidoptera — Pieridae).
Lambillionea 86 suppl.:l— 88.
boelcke, o. 1957. Comunidades herbaceas del norte de la Patagonia y sus
relaciones con la ganaderia. Rev. Invest. Agric 11:5-98.
BREYER, A. 1939. Uber die Argentinischen Pieriden (Lepidoptera Rhopalocera).
Verhandl. VII int. Kongr. Ent. 1:26-55.
BRIGGS, J. C. 1987. Biogeography and Plate Tectonics. Developments in Paleon-
tology and Stratigraphy #10. Elsevier, Amsterdam. 193 pp.
BROWN, K. S. 1987. Biogeography and evolution of Neotropical butterflies,
pp. 66-104 in T. C. Whitmore and G. T. Prance, eds. Biogeography and
Quaternary history in Tropical America. Clarendon Press, Oxford.
CABRERA, A. L. 1953a. Esquema fitogeografica de la Republica Argentina. Rev.
Mus. La Plata (N. S.) (Bot.) 8:87-168.
1958. La vegetation de la Republica Argentina. VI. La vegetation
de la puna Argentina. Serie fitogeografica #6. Rev. Invest. Agric. 11:
317-412.
— , 1971. Fitogeografia de la Republica Argentina. Bob Soc. Arg. Bot.
14:1-42.
, & A. WILL If IK. 1980. Biogeografia de America Latina. Organization
de los Estados Americanos, Washington, D. C. 122 pp.
CAIN, S.A. 1944. Foundations of Plant Geography. Harper, New York. 556 pp.
CASTELLANOS, A. & R A. PEREZ MOREAU. 1945. Los tipos de vegetation de la
Republica Argentina. Monogr. Inst. Estud. Geogr. Univ. Nac. Tucuman
4:1-154.
CAVIEDES, C. 1989. Rainfall, snowline depression, and vegetational shifts in
Chile during the Pleistocene. Climatic Change :
, & A. IRIARTE W. 1989. Migration and distribution of rodents in central
Chile since the Pleistocene: the paleogeographic evidence. J. Biogeog.
16:181-187.
CEI, J. M. 1980. Reliquias y refugios, al sur del tropico, de la herpetofauna
200
J. Res. Lepid.
austral Pleistocena sudamericana. pp. 213-227 in P. Salinas, ed., Zoologia
Neotropical: Actas del VIII Congreso Latinoamericano de Zoologia,
Merida, Venezuela.
CHABOT, B. F. & w. D. BILLINGS. 1972. Origins and ecology of the Sierran alpine
flora and vegetation. Ecol. Monogr. 42:163-199.
CLAPPERTON, C. M. 1983. The glaciation of the Andes. Quat. Sci. Reviews 2:
83-155.
CORREA LUNA, H., R. C. TOLOSA & E. MONAGLIO. 1977. La conservation de la
naturaleza: parques nacionales argentinos. Ministerio de la Economia,
Buenos Aires. 169 pp.
COURTNEY, S. P. 1986. Oviposition by the mistletoe-feeding Pierid butterfly
Mathania leucothea (Mol.) in Chile. J. Res. Lepid. 24:264-270.
, & A. M. SHAPIRO. 1986a. The life-history of Hypsochila wagenknechti
wagenknechti, a scarce butterfly from the Andes of temperate Chile
(Lepidoptera: Pieridae). J. New York Entomol. Soc. 94:531-535.
, 1986b. The ecology and behavior of the high-Andean butterfly
Hypsochila wagenknechti (Lepidoptera: Pieridae). Stud. Neotrop. Fauna
Envt. 21:169-188.
CRACRAFT, J. 1989. Speciation and its ontology: the empirical consequences of
alternative species concepts for understanding patterns and processes of
differentiation, pp. 28-59 in D. Otte and J. A. Endler, eds., Speciation and
its Consequences Sinauer, Sunderland, Mass.
CRAW, R. C. 1982. Phylogenetics, areas, geology and the biogeography of
Croizat: a radical view. Syst. Zool. 31:304-316.
CROIZAT, L. 1964. Space, Time, Form: The Biological Synthesis. Published by
Author, Caracas, Venezuela. 881 pp.
, G. NELSON & D. E. ROSEN. 1974. Centers of origin and related concepts.
Syst. Zool. 23:265-287.
DAVIS, D. R. 1986. A new family of monotrysian moths from Austral South
America (Lepidoptera: Palaephatidae), with a phylogenetic review of the
Monotrysia. Smithson. Contrib. Zool. 434:1-202.
DELESSE, H. 1967. Les nombres de chromosomes chez les Lepidopteres Rhopa-
loceres Neotropicaux. Ann. Soc. Ent. France (N.S.) 3:67-136.
DESCIMON, H. 1986. Origins of Lepidopteran faunas in the high tropical Andes,
pp. 500-532 in F. Vuilleumier and M. Monasterio, eds., High Altitude
Tropical Biogeography. Oxford University Press, New York.
DIMITRI, M. J. 1962. La flora andino-patagonica. Anales de Parques Nacionales
9:1-115.
, 1972. La region de los bosques andino-patagonicos: sinopsis general.
Coll. Cientif. INTA #10. Buenos Aires. 254 pp.
DUNBAR, M. J. 1968. Ecological Development in Polar Regions: A Study in
Evolution. Prentice-Hall, Englewood Cliffs, N. J. 119 pp.
FERNANDEZ, j. 1970. Polylepis tomentella y orogenia reciente. Bol. Soc. Arg.
Bot. 13:14-30.
field, W. D. 1958. A redefinition of the butterfly genera Tatochila, Phulia,
Piercolias, and Baltia, with descriptions of related genera and subgenera.
Proc. U. S. Nat. Mus. 108:103-131.
., & J. HERRERA. 1977. The pierid butterflies of the genera Hypsochila
Ureta, Phulia Herrich-Schaffer, Infraphulia Field, Pierphulia Field, and
Piercolias Staudinger. Smithson. Contrib. Zool. 232:1-64.
28(3):137-238» 1989(91)
201
GIACOMELLI, E. 1915. El genero Tatochila But!.: lo que sabemos y lo que
ignoramos de el. Anales Mus. Nac. Hist. Nat. Buenos Aires 26:403-415.
GOMEZ MOLINA, E. & A. V. LITTLE. 1981. Geoecology of the Andes: the natural
science basis for research planning. Mountain Research and Development
1:115-144.
HALLOY, S. 1978. Contribucion al estudio de la zona de Huaca-Huasi, Cumbres
Calchaquies (Tucuman, Argentina). I. La biota actual en relation con la
geologla historica. Unpublished MS, Fac. Gs. Nat., Univ. Nac. Tucuman.
, 1982. Contribucion al estudio de la zona de Huaca-Huasi, Cumbres
Calchaquies (Tucuman, Argentina). II. Thesis, Univ. Nac. Tucuman. 839
PP-
. 1983. Algunos dates ecologicos para Nototriche caesia Hill, Malvacea
altoandina, en la's Cumbres Calchaquies, Tucuman. Lilloa 36:85-104.
HAUMAN, L. 1919. La vegetation des hautes cordilleres de Mendoza. Anal. Soc.
Cient. Arg. 86:121-188, 225-348.
HAYWARD, K. J. 1931. Notas sobre una migration de Pieris phileta automate
Burm. Rev. Soc. Entomol. Arg. 3:225-232.
1950. Catalogo sinonimico de los Ropaloceros argentinos excluyendo
Hesperiidae. Acta Zool. Lilloana 9:85-282.
. , 1955b. An entomologist in Argentina. XIII. The Calchaqui valleys.
Ent. Rec. J. Var. 67:165-169.
1955c. An entomologist in Argentina. XIV. Mendoza. Ent. Rec. J.
Var. 67:223-226.
1973. Catalogo de los Ropaloceros- argentinos. Opera Lilloana 23:1 —
318.
HEIMLICH, W. 1972. Satyridae der sudlichen Neotropis und Subantantarktis.
Beitrag Entomol. 22:149-197.
HERRERA, J. 1953. Lepidopteros nuevos para Chile. Rev. Chil. Entomol. 3:
140-147.
, 1970. Una especie y dos subespecies nuevas de Tatochila colectadas
en la alta cordillera de Antofagasta. Publ. Centro de Estudios Entomol.
10:5-18.
, & R. COVARRUBIAS. 1983. Distribution biogeografica del grupo Tato-
cheila (sic) Pkulia (Lepidoptera: Pieridae). Res. Comm. Sci. IX Congr.
Latinoamer. Zool., Arequipa, Peru 9-15 Oct. 1983. p. 213.
& W. D. FIELD. 1959. A revision of the butterfly genera Theochila and
Tatochila (Lepidoptera: Pieridae). Proc. U. S. Natl. Mus. 108:467-514.
HEUSSER, c. J. 1983. Quaternary pollen record from Laguna de Tagua Tagua,
Chile. Science 219:1429-1432.
HOVANITZ, w. 1970. Habitat: Celias vautieri . J. Res. Lepid. 9:100.
HUBAlek, Z. 1982. Coefficients of association and similarity based on binary
(presence-absence) data: an evaluation. Biol. Rev. 57:669-689.
HUECK, K. & P SEIBERT. 1972. Vegetationskarte von Sudamerika. Verlag Gustav
Fischer, Stuttgart. 69 pp.
HUNZIKER, J. H. 1952. Las comunidades vegetales de la cordillera de La Rioja.
Rev. Invest. Agrie, 6:167-196.
IRWIN, M. E. & E. I. SCHLINGER. 1986. A gazetteer for the 1966-67 University of
California — Universidad de Chile arthropod expedition to Chile and
parts of Argentina. Occ. Papers Calif. Acad. Sci. 144:1-11.
JANSON, S. & J. VEGELIUS. 1981. Measures of ecological association. Oecologia
202
J. Res. Lepid.
49:371-376.
JORGENSEN, P. 1916. Las mariposas argentinas (Lepidoptera), familia Pieridae.
Anal. Mus. Nac. Hist. Nat. Buenos Aires 28:427-520.
KLOTS, A. B. 1932. A generic revision of the Pieridae (Lepidoptera) together with
a study of the male genitalia. Entomol. Americana 12 (N. S.):139-242.
LAMAS, G. 1982. A preliminary zoogeographical division of Peru based on
butterfly distributions (Lepidoptera, Papilionoidea). pp. 336-357 in G. T.
Prance, ed., Biological Diversification in the Tropics. Columbia University
Press, New York.
, & J. PEREZ R. 1983. Las mariposas del Parque Nacional Huascaran
(Ancash, Peru) y su medio ambiente: sector Llanganuco. Boletln de Lima
30:27-37.
LARSEN, T. B. 1984. The zoogeographical composition and distribution of the
Arabian butterflies (Lepidoptera: Rhopalocera). J. Biogeog. 11:119-158.
LISTA, R. 1896a. Viaje a los Andes australes. Anal. Soc. Cient. Arg. 41:5-138,
145-170.
, 1896b. Los Andes patagonicos. Anal. Soc. Cient. Arg. 41:286-296.
LIU, K.-B. & P. A. COLINVAUX. 1985. Forest changes in the Amazon basin during
the last glacial maximum. Nature (London) 318:556-557.
, 1988. A 5200-year history of Amazon rain forest. J. Biogeog. 15:231-
248.
MADSEN, H. B., E. SCHMIDT-NIELSEN & s. 0DUM. 1980. The Danish scientific
expedition to Patagonia and Tierra del Fuego 1978-79. Geografisk Tids-
skrift 80:1-28.
MANABE, S. & D. G. HAHN. 1977. Simulation of the tropical climate of an ice age. J.
Geophys. Res. 82:3889-3911.
MARKGRAF, V. 1985. Late Pleistocene faunal extinctions in southern Patagonia.
Science 228:1110-1112.
MATTHEW, W. D. 1915. Climate and evolution. Ann. New York Acad. Sci.
24:171-318.
MAYR, E. 1963. Animal Species and Evolution. Harvard University Press,
Cambridge, Mass. 797 pp.
MENENDEZ, C. A. 1971. Floras terciarias de la Argentina. Ameghiniana 8:
257-268.
MERCER, J. H. & J. F. SUTTER. 1981. Late Miocene-earliest Pliocene glaciation in
southern Argentina: implications for global ice sheet history. Paleogeog.,
Paleoclimat., Paleoecol. 38:185-206.
MOORE, D. M. 1983. Flora of Tierra del Fuego. Missouri Botanic Garden, St.
Louis. 396 pp.
MORELLO, J. 1958. La provincia fitogeografica del Monte. Opera Lilloana 2:
5-155.
ORIANS, G. H. & O. T. SOLBRIG, EDS. 1977. Convergent Evolution in Warm Deserts.
Dowden, Hutchison and Ross, Stroudsburg, Penna. 333 pp.
PASKOFF, R. 1977. Quaternary of Chile: the state of research. Quat. Rev. 8:2-
31.
PENA, L. E. 1966. A preliminary attempt to divide Chile into entomofaunal
regions based on the Tenebrionidae (Coleoptera). Postilla 97:1-17.
, 1975. Guia Para Reconocer Mariposas. Editorial Gabriela Mistral,
Santiago. 120 pp.
PEREZ R., J. E. 1982. Ecologla de las mariposas diurnas (Lepidoptera, Rhopalo-
28(3): 137-238, 1989(91)
203
cera) del Sector Llanganuco, ,Parque Nacional Huascaran, Ancash, Peru.
Thesis, Univ. Partic. Ricardo Palma, Lima. 80 pp.
PETERSON, B. 1963. The male genitalia of some Colias species. J. Res. Lepid.
1:135-156.
PORTER, A. H. & A. M. SHAPIRO. 1990. Genitalia and arthropod taxonomy: lack of
mechanical isolation in a butterfly hybrid zone (Lepidoptera: Pieridae).
Ann. Rntomol. Soc. Amer. 83:107-114.
RABINOVICH, J. E. & E. H. RAPOPQRT. 1975. Geographical variation of diversity in
Argentine passerine birds. J. Riogeog. 2:141-157.
RAGONESE, A. E. & B. c. PICCININI. 1969. I. unite entre el Monte y el semidesierto
patagonico en las provincial de Rio Negro y Meuquen. Bob Soc. Arg. Bot.
11:299-302.
RAVEN, P. H & D. I AXELROD. 1974. Angiosperm biogeography and past continen-
tal movements. Ann. Missouri Bot. Gard. 61:539-673.
RIND, D. & D. PETEET. 1985. Terrestrial conditions at the last glacial maximum
and CLIMAP sea-surface temperature estimates: are they consistent?
Quat. Res. 24:1-22.
RINGUELET, R. A. 1955. Vmculaciones faunlsticas de la zona boscosa del Nahuel
fluapi y el Dominio Zoogeografico Australcordillerano. Notas Museo Univ.
Nac. Eva Peron 18:81-121.
ROBER, J. 1908-1924. (South American Pieridae.) pp. 53, 55-56, 97-98, 1015,
1016, 1024 in vol. 5, The American Rhopalocera, A. Seitz, ed., The
Macrolepidoptera of the World. Pla tes 18, 28, 192, 194. Verlag A. Kernen.
Stuttgart.
SANCHES, O. & G. LOPEZ ORTEGA 1988. A theoretical analysis of some indices of
similarity as applied to biogeography. Folia Entomol. Mexicana 75:119 —
145.
SARMIENTO, G. 1975. The dry plant formations of South America and their
floristic connections. J. Biogeog. 2:233-251.
SCHAEFER, B. & A. BREYER. 1941. Lista de Lepidopteros de Catamarca, y algunas
observaciones. Rev. Soc. Ent. Arg. 11:221-228.
SHANNON, R. C. 1927. Contribucion a los estudios de las zonas biologicas de la
Republica Argentina. Rev. Soc. Entomol. Arg. 1 #4, pp. 1-14.
SHAPIRO, A. M. 1979. The life-histories of the autodice and sterodice species-
groups of Tatochila (Lepidoptera: Pieridae). J. New York Entomol. Soc.
87:236-255.
, 1980a. Convergence in Pierine polyphenisms. J. Nat. Hist. 14:781 —
802.
, 1984. Experimental studies on the evolution of seasonal polyphen-
ism. pp. 297-307 in Vane-Wright and P. Acker y, eds., The Biology of
Butterflies. Academic Press, New York.
, 1985. Behavioral and ecological observations of Peruvian high
Andean Pierid butterflies. Stud. Neotrop. Fauna Envt. 20:1-14.
, 1986a. Intergradation of Tatochila autodice and T. blanchardii
(Lepidoptera: Pieridae) in northwest Patagonia. J. Nat. Hist. 20:1309-
1320.
1986b. The genetics of polyphenism and its role in the phylogenetic
interpretation of the sterodice species-group in the Andean-Neantarctic
region. J. Res. Lepid. 24, suppl.:24-31.
, 1986c. r-K selection at various taxonomic levels in the pierine
204
J. Res. Lepid.
butterflies of North and South America, pp. 135-152 in F. Taylor and R.
Karban, eds., The Evolution of Insect Life Cycles. Springer- Verlag, New
York.
, 1986d. The life-history of Tatochila distincta distincta, a rare but-
terfly from the puna of northwestern Argentina. J. New York Entomol.
Soc. 94:526-530.
, 1989a. Ignorance in high places. Paleobiology 15:61-67.
, 1989b. The biology of Colias blameyi (Pieridae), the “green sulphur”
of the Argentine puna. J. Res. Lepid. : - .
, & S. P. COURTNEY. 1986. The life-history of Pierphulia rosea an-
namariea, an unusual butterfly from the Peruvian altiplano. J. New York
Entomol. Soc. 94:536-541.
, & H. J. GEIGER. 1989. Electrophoretic comparisons of vicariant
Vanessa : genetic differentiation between V. annabella and V. carye (Nym-
phalidae) since the Great American Interchange. J. Lepid. Soc. 43:81-92.
SOLBRIG, O. T. 1976. The origin and floristic affinities of the South American
temperate desert and semidesert regions, pp. 7-49 in D. W. Goodall, ed.,
Evolution of Desert Biota. University of Texas Press, Austin.
SORENSEN, T. 1948. A method of establishing groups of equal amplitude in plant
sociology based on similarity of species content and its application to
analyses of the vegetation on Danish commons. Kongelige Danske Videns-
kabernes Selskab, Biologiske Skrifter 5:1-34.
SORIANO, A. 1949. El Ifmite entre las provincias botanicas patagonica y central
en el Territorio del Chubut. Lilloa 20:193-202.
, 1956. Los distritos florlsticos de la Provincia Patagonica. Rev. Invest.
Agric. 10:323-347.
SPERLING, F. A. H. 1987. Evolution of the Papilio machaon group in western
Canada (Lepidoptera: Pieridae). Quaest. Entomol. 23:198-315.
STAUDINGER, O. 1894. Hochandine Lepidopteren. Deutsche Entomol. Zeitschr.
“Iris” 7:43-100.
STEHLI, F. G. & S. D. WEBB. 1985. The Great American Biotic Interchange. Plenum
Press, New York. 532 pp.
STEVENS, G. C. 1989. The latitudinal gradient in geographical range: how so
many species coexist in the tropics. Am. Nat. 133:240-256.
UNITED STATES DEPARTMENT OF THE INTERIOR. 1968. Argentina. Official stan-
dard names approved by the U. S. Board on Geographic Names. Gazetteer
#103. Office of Geography, Washington, D. C. 699 pp.
URETA ROJAS, E. 1955. Nuevas especies de Pieridae (Lepidoptera, Rhopalocera)
de Chile y Argentina. Bol. Mus. Nac. Hist. Nat. 26:57-71.
, 1963. Lepidopteros de Chile. Bol. Mus. Nac. Hist. Nat. 28:63-140.
VAN DER HAMMEN, T. & A. M. CLEEF. 1986. Development of the high-Andean
paramo flora and vegetation, pp. 153-201 in F. Vuilleumier and M.
Monasterio, eds., High Altitude Tropical Biogeography. Oxford University
Press, New York.
VEBLEN, T. T., C. DONOSO Z., F. M. SCHLEGEL & B. ESCOBAR R. 1986. Forest dynamics
in south-central Chile. J. Biogeog. 8:211-247.
VOLKHEIMER, W. 1971. Aspectos paleoclimatologicos del Terciario argentino.
Rev. Mus. Arg. Cs. Nat. “Bernardino Rivadavia”, ser. Paleontol. 1:243-
262.
VUILLEUMIER, F. & M. MONASTERIO, EDS. 1986. High Altitude Tropical Biogeo-
graphy. Oxford University Press, New York, 649 pp.
28(3):137-238, 1989(91)
205
WAGENKNECHT H., R. 1968. Observaciones biologicas sobre la Eroessa chilensis
(Guerin, 1829), Pierididae (Rhopalocera). An. Mus. Hist. Nat. Valparaiso
1:173-177.
WEBERBAUER, A. 1945. El Mundo Vegetal de los Andes Peruanos. Ministerio de
Agricultura, Lima. 776 pp.
WHITMORE, T. C. & G. T. PRANCE, EDS. 1987. Biogeography and Quaternary
History in Tropical America. Clarendon Press, Oxford. 214 pp.
WHYMPER, E. 1892. Travels Amongst the Great Andes of the Equator. Facsimile
reprint (1987), Peregrine Smith Books, Salt Lake City. 456 pp.
WILLIS, B. 1914. Northern Patagonia: Character and Resources. Ministry of
Public Works, Argentine Republic. Scribner’s, New York. 464 pp.
WINGENROTH, M. & J. SUArez. 1983. .Flores de los Andes: Alta Montana de
Mendoza. Institute Argentino de Nivologia y Glaciologla, Mendoza.
Supplementary Bibliography
The following references are not cited specifically in the text, but were
consulted in the preparation of this study. Designations (e.g., 1983a, b, c) are
coordinated with the Literature Cited above.)
BERG, C. 1877. Contribqcion al estudio de la fauna entomologica de la Patago-
nia: Rhopalocera. Anal. Soc. Cient. Args 4:87-95.
, 1895. Revision et description des especes Argentines et Chiliennes
du genre Tatochila Butl. Anal. Mus. Nac. Buenos Aires 4:217-255.
BREYER, A. 1931. Lepidopteros de Yacanto (Cordoba). Rev. Soc. Entomol. Arg.
3:169-173.
, 1936. Lepidopteros de la zona del lago Nahuel Huapi, Territorio del
Rio Negro (Republica Argentina). Rev. Soc. Entomol. Arg. 8:61-63.
, 1945. Los Lepidopteros del sur de la Republica Argentina. Anales
Museo de la Patagonia 1:237-241.
BROWN, F. M. 1974. Entomological bibliography of Kenneth John Hayward
(1891-1972). J. Lepid. Soc. 28:332-343.
CABRERA, A. L. 1953B. Manual de la Flora de los Alrededores de Buenos Aires.
Editorial Acme, Buenos Aires. 585 pp.
, 1980. Ecologia vegetal de la puna: geoecologia de las regiones
montanosas de las Americas tropicales. Proc. Simposio UNESCO, Mexico
1966:91-116.
CASTELLANOS, A. & R. A. PEREZ-MOREAU. 1941. Contribucion a la bibliografia
botanica argentina. Cruciferae. Lilloa 7:65-73.
CAVIEDES, C. & R. PASKOFF. 1975. The Quaternary glaciations of north central
Chile. J. Glaciol. 14:155-170.
GARCIA, C. V. 1970. El clima de Bariloche y sus alrededores. Gaea (Buenos
Aires) 14:55-70.
GEMIGNANI, E. v. 1939. La seccion entomologica del Museo Argentino de Cien-
cias Naturales. Verhandl. VII int. Kongr. Ent. 1:133-143.
GIACOMELLI, E. 1916. Sinopsis de los Lepidopteros chilenos del genero Tato-
chila. Rev. Chil. Hist. Nat. 20:41-57.
, 1917. Nuevos estudios y observaciones sobre Pieridae argentinos.
Physis 3:370-385.
, 1923. Enumeracion sistematica y anotaciones sobre los Lepidopteros
206
J. Res. Lepid.
de Villa Cabrera, Cordoba. Rev. Univ. Nac. Cordoba 10:174-188.
GOMEZ, S. A. 1953. Caparidaceas Argentinas. Lilloa 26:279-341.
HAYWARD, K. J. 1935. The geographical aspects of Argentine entomology. Ent.
Rec.J.Var. 47:109-110.
, 1955a. An entomologist in Argentina. XII. Neuquen. Ent. Rec. J.
Var. 67:116-121.
, 1966. Los ropaloceros de Cafayate (Salta). Rev. Soc. Entomol. Arg.
28:65-70.
HERRERA, J. (Undated; apparently the MS form of Herrera and Covarrubias,
1983.) Zoogeografia de las mariposas sudamericanas de la rama
Tatocheila-Phulia. Unpublished MS.
HEUSSER, C. J. & S. S. STREETER. 1980. A temperature and precipitation record of
the past 16,000 years in southern Chile. Science 210:1345-1347.
JAMES, D. E. 1973. The evolution of the Andes. Sci. Amer. 229 (2):60-69.
KOHLER, P. 1923. Fauna Argentina. I. Teil. Lepidoptera e collectione Alberto
Breyer. Sonderbeilage der Zeitschrift fur Wissenschaftliche Insektenbiolo-
gie 18, #12.
, & E. GEMIGNANI. 1932. Los Lepidopteros del Museo Nacional de
Historia Natural de Buenos Aires “Bernardino Rivadavia.” Rev. Soc.
Entomol. Arg. 21:67-72.
KREBS, C. A. 1959. Observaciones fenologicas sobre plantas indigenas del Parque
Nacional Nahuel Huapi. Anales de Parques Nacionales 8:127-133.
LAMAS, G. 1981. Notes on Peruvian butterflies (Lepidoptera). VI. Twelve new
Pieridae. Rev. de Ciencias, Univ. Nac. Mayor de San Marcos 73:44-55.
MAHAN, S. 1941. Observaciones biologicas sobre la Tatochila blanchardii (Lepi-
doptera, Pieridae). Rev. Chil. Hist. Nat. 44:63-67.
MARGHERITIS, A. E. & H. F. E. RIZZO. 1965. Lepidopteros de Interes Agricola.
Editorial Sudamericana, Buenos Aires. 193 pp.
MARSHALL, L. G. & B. PATTERSON. 1981. Geology and geochronology of the
mammal-bearing Tertiary of the Valle Santa Maria and Rio Corral
Quemado, Province of Catamarca, Argentina. Fieldiana Geolog. (N. S.)
9:1351-1357.
MELLINI, E. & M. VERENINI. 1986a. Note di biologia sugli insetti delle aree
semiaride dell’Altiplano Boliviano durante la primavera. Boll. 1st. En-
tomol. “Guido Grandi” Univ. Bologna 60:121-149.
, 1986b. Note di biologia sugli insetti delle aree semiaride dell’Altipla-
no Boliviano all’inizio della stazione estiva. Boll. 1st. Entomol. “Guido
Grandi” Univ. Bologna 60:185-214.
MERCER, J. H. 1965. Glacier variations in southern Patagonia. Geogr. Rev.
55:390-413.
, 1968. Variations of some Patagonian glaciers since the late glacial. I.
Amer. J. Sci. 266:91-109.
, 1970. Variations of some Patagonian glaciers since the late glacial.
II. Amer. J. Sci. 269:1-25.
, 1976. Glacial history of southernmost South America. Quat. Res.
6:125-166.
, 1982. Holocene glacier variations in southern Patagonia. Striae
18:35-40.
, 1983. Cenozoic glaciation in the Southern Hemisphere. Ann. Rev.
Earth and Planetary Sci. 11:99-132.
28(3):137-238, 1989(91)
207
, & O. PALACIOS. 1977. Radiocarbon dating of the last glaciation in
Peru. Geology 5:600-604.
NEDER DE romAn, L. E. & M. ARCE DE H amity. 1983. Desarrollo ontogenetico,
morfologla y biologia de Leptophohia aripa Bsd. (Lepidoptera: Pieridae).
Acta Zool. Lilloana 37:77-85.
PETRIELLA, B. 1972. Estudio de maderas petrificadas del terciario inferior del
area central de Chubut (Cerro Bororo). Rev. Museo La Plata (N. S.)6,
Paleontologla #41:159-254.
REED, E. C. 1877. Monografla de las mariposas chilenas. Anales Univ. (San-
tiago) ano 1877:647-736.
SERVicio meteorolOgico nacional. 1951-1980. Estadlsticas climatologicas.
SHAPIRO, A. M. 1980b. Physiological and developmental responses to photoperiod
and temperature as data in phylogenetic and biogeographic inference.
Syst. Zool. 29:335-341.
, 1989. Observaciones acerca de las mariposas fueguinas. Praxis
(Ushuaia) 1:43-45.
SIGONIER, P. & V. CARVAJAL. 1975. Errores y confusiones en la identificacion de
las mariposas chilenas del genero Tatochila en el Siglo XIX (1818-1897).
Thesis, Universidad de Chile (Depto. Biologia). 44 pp.
SORIANO, A. 1950. La vegetacion del Chubut. Rev. Arg. de Agronomla 17:30-
66.
TORRES BRUCHMANN, E. 1971. Mesoclimas de la Provincia de Tucuman. Rev.
Agron. Norest. Arg. 9:527-544.
URETA ROJAS, E. 1936-37. Lepidopteros de Chile. Rev. Chil. Hist. Nat. 40:343-
380, 41:237-287.
J, Res. Lepid.
I. ilyodes X
p. r. rosea X
P. r . maria X
P. isabela X
P . nyrouhula X
H. penai X
H. w. sulfurodice X
F. w. wagenknechti 0
K. galactodice 0
E. huemul 0
H. microdice 0
E. argyrodice 0
T. t. theodice 0
T. t. gymnodice 0
T. t. staudingeri 0
T. a. autodice 0
T. a. blanchardii 0
T. a. ernestae X
T. m. mercedis 0
T. m. macrodice X
T. m. vanvolxemii 0
T. m. sterodice 0
T. m. fueguensis 0
T. inversa 0
T. orthodice 0
T. stigmadice 0
T. mariae X
T. d. distincta 0
T. d. fieldi X
E, chilensis 0
M. leucothea 0
C. lesbia X
C» weberbaueri X
C. blameyi 0
C. flaveola 0
C. vauthieri 0
C. ponteni 0
F . deva X
T. rioiana X
00000000
00000000
00000000
00000000
XXXXXXOO
00000000
xooooooo
oooxxxoo
xooooooo
00000000
00000000
00000000
000000X0
OdOOQOOO
00000000
00000000
oooooxxx
00000000
oooooxxx
XXX0000 Q
00000000
00000000
00000000
xxxooooo
xxxooooo
xxxooooo
00000000
xxxooooo
00000000
00000000
000000X0
xxxooooo
0 0 000000
xxxooooo
000X0000
oooooxxx
00000000
xxxooooo
xxxooooo
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 X 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 X
0 0 X 0 X X 0
0 0 0 0 0 0 X
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 X X X
0 0 0 0 0 0 0
0 0 0 0 0 0 0
X X X X X 0 X
0 0 0 0 0 X X
0 0 0 0 0 0 0
o o o a o x x
0 0 0 0 0 0 0
X X 0 X X 0 0
0 0 0 0 X X X
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 X
0 0 0 0 0 0 X
X X X X X X 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 X X X X
0 0 0 0 0 0 0
0 0 0 0 0 0 0
0 0 0 0 0 0 0
00000000
00000000
00000000
00000000
00000000
00000000
00000000
00X00000
xxoooooo
00X00000
oooooxxx
oooxooxx
xoxoxooo
000000X0
ooooooxx
xxoxoooo
xxxooooo
00000000
00X0X000
00000000
000X0000
xxoooxoo
oooooxxx
00000000
00000000
00000000
00000000
00000000
00000000
xoooxooo
xoooxooo
xooxoooo
00000000
00000000
00000000
XXXXXXOO
00000000
xooxoooo
00000000
# Of species: 15 12 10 10 3 2 5 5 3 3 3 4 4 7 7 10 10 5 6 6 5 4 5 4
Table 1 . Occurrence of selected pierine species in selected areas of Argenti-
na and Chile.
Co/ias mendozina from the Paso Bermejo is omitted from this and
subsequent analyses.
28(3): 137-238, 1989(91)
209
Puna de Atacama X6551110
Puna de Jujuy/ Salta X 10 10 1 1 1 0
Cuesta del Obispo X 10 1 1 1 0
Sierras Pampeanas X 1 1 1 0
Arroyo de Agua Negra X 2 2 0
Paso Bermejo X 2 0
Cordon del Cepo X 3
Precordillera Chilena X
Valle Central (Chile)
Precordillera Mendocina
Valles Calchaaules
Cordon del Viento
Chos Malal
Loncopue
Alumine
San Martin de los Andes - Parque Nacional
San Carlos de Bariloche - Parque Nacional
Esquel - Parque Nacional Los Alerces
Laguna del Maule
Comodoro Pivadavia - Fitz Roy
Valdivia Region
Rio Gallegos
Rio Grande
Ushuaia - Cordon Martial
Chilean Magallanes
0112111010010000
011312 2021010000
0112111010010000
0112111010010000
0001000000000000
1001000000000000
30011 1 3433211110
3000124544213100
X000313322312100
X323311 21030000
X2331122030000
X233132020000
X42223142100
X544424 22 10
X564423210
Lanin X74525210
Nahuel Huapi X5344200
X 2 2 1 2 0 0
X 1 3 1 0 0
X 1 0 1 0
X 1 0 0
X 2 2
X 2
X
0
0
0
0
0
0
1
1
1
0
0
0
1
2
2
2
2
2
1
2
4
4
2
4
X
Table 2
Numbers of species shared by all possible pairs of regions. Data
from table 1.
Ushuaia - Cordon Martial
500 Ushuaia - Cordon Martial
+ Chilean Magallanes
210
J. Res. Lepid .
Puna de Atacama (Chile)
Puna de Jujuy/Salta
Cuesta del Obispo/Co. Zapallar
Sierras Pampeanas
Arroyo de Agua Negra
Paso Bermejo
Cordon del Cepo
Precordillera Chilena
Valle Central (Chile)
Precordillera Mendocina
Valles Calchaquies
Cord<5n del Viento
Chos Malal
Loncopue
Alumine
San Martin - P.N. Lanin
Bariloche - P.N. Nahuel Huapi'"
Esquel - P.N. Los Alerces
Laguna del Maule
Comodoro Rivadavia - Fitz Roy
Valdivia Region
Rio Gallegos
Rio Grande
Ushuaia - Cord«5n Martial
Chilean Magallanes
Table 3. Sprensen's index of similarity used to compare all possible pairs of
faunas from Table 2. These data are presented as a dendrogram in
Fig. 23.
28(3):137-238, 1989(91)
211
EPOCH
GEOHISTORY
BIOTA
PIERIDAE
Glaciers retreating in Mendoza and Tucuman; - —
much fluctuation in glacial advances and retreats
in Patagonia
warm interval — — — — ~ — -
Deglaciation of Fue go -Patagonian
lowland commences
ULTIMATE MAJOR
GLACIATION
"L
W->E Goss-Andean migrations
in NW Patagonia; origin of
hybrid zones in Tatochila;
weak subspeciation in NW
Tatochila taxa
Standings in Corddn del Viento; High Andean
fauna in NW assumes present configuration
Colonization of Fuegia from Patagonian mainland
Patagonian faunal refugia along coastline and on
continental shelf; Andean fauna in NW shifted
downslope
L PENULTIMATE
GLACIATION
0.13-0.17 MY A
GREA TEST PA TA GONIAN
GLACIATION
(Condor Cliff— Initio glacial —
Pic hi Leufu)
Repeated N-S
biotic migrations
on both sides
of cordillera
Origin of Atacama desert barrier •
to dispersal; origin of Argentine
monte desert
E->W cross-Andean
migrations into Chile
at latitude of
Mendoza & San Juan;
subspecific
differentiation in
Tatochila mer cedis
complex?
— Origins of Andean
Pierine species?
- 2.0
2.5
-3.0
3.5
-4.0
■4.5
^
^ ^ ^
5 '•J Cjj
■8£|-
-
? 5 n-
Usual assumed earliest date of
Andean uplift to alpine heights
(First buildup of mid- latitude
Northern hemisphere ice sheets)
First "definite" Patagonian glaciation;
inference (Clapperton) that Andes
already near present heights
Origins of Andean
Pierine genera?
Beginning of ancestral Andean
First evidence of Bolivian altiplano
orogeny
vegetation in Miocene (or perhaps
Paleocene — Eocene
Eocene?)
Table 4. Attempted cross-correlation of geohistorical and biotic events in
the southern Andes and Patagonia with development of the Pierid
butterfly fauna. KYA = thousands of years ago; MYA = millions of
years ago. Note discrepancy between the usual Plio-Pleistocene
estimate for the origin of alpine habitats in the Andes and the much
older estimate inferred by Mercer and Clapperton from alleged pre-
Pleistocene tills, and the older-still earliest paleobotanical evidence
for xerophytic vegetation in Bolivia. Very detailed paleoclimatic
sequences are available for the Holocene in much of southern
South America, but the quality and quantity of evidence diminish
with time. Pre-Pleistocene events are in part inferred by cross-
correlation with evidence bearing on marine temperatures near
New Zealand (see Clapperton reference).
212
J. Res. Lepid .
Fig, 1, PoSiticaS map of Argentina.
28(3): 137-238, 1989(91)
213
m
Phytogeography
AMAZONIAN DOMAIN
Yungas Province
Parana Province
■d
CHACO DOMAIN
Chaco Province
ES3
Espinal Province
■
Prepuna Province
Monte Province
□
Pampa Province
ANDEAN-PAT AGONI AN DOMAIN
■
High-Andean Province
III
Puna Province
EH
Patagonian Province
EH
SUBANTARCTIC DOMAIN
Subantarctic Province
■
Insular Province
Fig. 2. Phytogeography of Argentina, redrawn from Cabrera, 1971. Fig. 1.
214
J. Res. Lepid.
Fig. 3. Distribution of altitude above sea level in Argentina and Chile, from
Madsen et a/., 1980.
fill
28(3):137-238, 1989(91)
215
Fig. 4. Climate types of Argentina and Chile according to the Koppen
system, from Madsen et a/., 1980.
216
J. Res. Lepid.
Fig. 5. Phytogeography of Argentina and Chile as mapped by Madsen et a!.,
1980. Key: 1. Subtropical forest; 2. Subantarctic (mostly Nothofagus)
forest; 3. Pam pa; 4. Xerophytic woodland and scrub (Espinal);
5. Patagonian steppe; 6. Puna; 7. Andean boreal and nival zones;
8. Chilean Mediterranean scrub (Matorral) and valley grassland;
9. Atacama Desert; 10. Monte Desert and arid montane (Prepuna).
Compare Fig. 2.
28(3):137-238, 1989(91)
217
Fig. 6. Generalized distribution of vegetation types in NW Argentina, reflect-
ing orographic influences on precipitation. Redrawn from Cabrera,
1971.
Phulia nymphula > 4000 r
ABRA
INFIERNILLO
Phoebis sennae
Eurema deva
Tatochila stigmadice
Tatochila autodice
distance (km)
Fig. 7. Transect across the Sierras Pampeanas in Tucuman, showing vege-
tation types, altitudinal distribution of Pieridae, and relationship of
Precipitation (P) to Potential Evapotranspiration (Ep) (annualized).
Phulia nymphula occurs only on the peaks over 4000 m, not reaching
the level of Abra Infiernillo. Partially redrawn after Madsen et aL
(1980).
meters above sea level
218
J. Res. Lepid.
Fig. 8. Tatochila theodice along a latitudinal cline. A-C, males; D- F,
females. A, Loncopue, Neuquen, 8. XI. 1988 (first brood) (N-most
known population of nominate theodice in Argentina). B, La Esper-
anza, Santa Cruz, 15.1.1979 (DE) (2nd brood) (± gymnodice). C, Rio
Grande, Tierra del Fuego, 25.XI.1988 (1st brood) (± staudingeri). D,
Puerto Blest, Parque Nacional Nahuel Fluapi, Rio Negro, 28.11.1979
(DE) (2nd brood?) [theodice). E, Lago Argentino, Santa Cruz, 1 1.1.1979
(DE) (2nd brood?) (± gymnodice). F, Rio Grande, Tierra del Fuego,
25. XI. 1988 (1st brood) (± staudingeri).
28(3):137-238, 1989(91)
219
Fig. 9. Tatochi/a autodice — blanchardii intergrades reared from wild ova
collected in the NW Patagonian hybrid zone, XI. 1988. A-C, males;
D-F, females. A, D, E from Esquel, Chubut; B, C, F from San Carlos
de Bariloche, Rio Negro. Specimens A and F are within the range
of variation of pure Chilean blanchardii. See Shapiro, 1986a for
characters.
220
J. Res. Lepid.
Tatochila mercedis subspecies
■
T. m. mercedis
▲
T. m. macrodice
A
T. m. vanvolxemii
•
T. m. sterodice
O
T. m. fueguensis
Hybrid populations
*
Complex ancestry involving
mercedis, vanvolxemii, & sterodice
★
sterodice & vanvolxemii
☆
T. m. vanvolxemii with
mercedis influence
K
Possible mercedis x macrodice
specimen
Figs. 10A-D. Distributions of selected taxa, superimposed on political map
of Argentina. A: Ta to chi la mercedis subspecies. B: Hyp
sochila, except H. penai and H. huemul, for which no new data
are reported. C: Phulia nymphu/a. D: Andean Col / as. Chilean
data mostly from Field and Flerrera, Flerrera and Field, and
Flerrera references in text.
221
B
Hypsochila populations
■
H. wagenknechti wagenknechti
□
H. wagenknechti sulfurodice
A
H. galactodice
A
H. (?) galactodice
o
H. microdice
★
H. ar gyro dice
J. Res. Lepid.
C
Phulia populations
% P. nymphula
28(3):137-238, 1989(91)
223
224
J. Res. Lepid.
••• •
Tatochila mercedis subspecies
■
T. m. mercedis
A
T. m. macrodice
A
T. m. vanvolxemii
•
T. m. sterodice
O
T. m.fueguensis
Hybrid populations
*
Complex ancestry involving
mercedis, vanvolxemii, & sterodice
★
sterodice & vanvolxemii
☆
T. m. vanvolxemii with
mercedis influence
*
Possible mercedis x macrodice
specimen
Fig.llA-D. Distributions of selected taxa, superimposed on phytogeo-
graphy as mapped by Cabrera, 1971 (cf. Fig. 2). A: Tatochila
mercedis subspecies. B: Hypsochila, except H. penai and
H. huemul. C: Phulia nymphula. D: Andean Co/ias. Note the
absence of Hypsochila from the Sierras Pampeanas.
28(3):137-238, 1989(91)
225
226
J. Res. Lepid.
28(3):137-238, 1989(91)
227
228
J. Res. Lepid.
Fig. 12. Unusual male (mercedis — macrodice intergrade?) not fitting any
named entity in polytypic species Tatochila mercedis. Arroyo de
Agua Negra, San Juan, 3. XI. 1988.
Fig. 13. Wild-collected first-brood males (A, B) and female (C) of Tatochila
mercedis vanvolxemii from Zapala, Neuquen, 7. XI. 1988. Note vari-
able loss of apical FW markings and tendency of these to form a
continuous line if present.
28(3):137-238, 1989(91)
229
Fig. 14. Tatochila mercedis vanvolxemii — stem dice intergrades from
Loncopue, Neuquen, 8. XI. 1988, all collected in the same field. A, B
males; remainder females; first brood.
230
J. Res. Lepid.
Fig. 15. Tatochila mercedis vanvolxemii — sterodice intergrades from
Barrio Prospero Palazzo, Comodoro Rivadavia, Chubut, 19. XI. 1988,
all collected in close proximity. A- K, males; l-R, females. Most of
these phenotypes are outside the range of variation of "pure"
vanvolxemii ; note tendency of apical FW markings in males to form
discrete dots. First brood.
28(3):137"238, 1989(91)
231
Fig. 16. Intergradation of Tatochila mercedis sterodice and T. m. fueguen-
sis. A, B, C, E, males; D, F, G, FI females. A, Lago Fagnano, Tierra del
Fuego, 19.1.1979 (DE) (intermediate). E, Cte. Luis Piedrabuena, Santa
Cruz, 20.XI.1988 ( sterodice ). All others from La Esperanza, Santa
Cruz, 130 km NW Rio Gallegos, 15.1.1979 (DE) (complete intergrada-
tion).
232
J. Res. Lepid.
Fig. 17. High-altitude Tatochi/a from NW Argentina. A, T. inversa male,
Cerro Amarillo, Jujuy, 4.1.1980 (RE). B, T. inversa female, Cerro
Zapallar, Salta, 22.1.1986. C, T. distincta distincta male, Abra In-
fiernillo, Tucuman, 20.1.1986. D, T. distincta, female, Cerro Zapallar,
Salta, 22.1.1986. The male inversa is the first figured from Argentina.
28(3):137”238? 1989(31)
233
Fig. 18. Various Hypsochi/a. AH males except H, female. A, H. wagenknechti
suifurodice , Chile, Mamina, Tarapaca, 30. IX. 1974 (J. Herrera). B,
same, Tres Cruces, Jujuy, 7.11.1984. C, H. argyrodice , Fitz Roy, Santa
Cruz, 11.11.1979 (DE). D, E, F all Cerro Amarillo, Jujuy, 4.1.1980 (RE):
D and E resemble H, galactodice, F resembles H. w. suifurodice. G,
H, //. microdice, Rio Grande, Tierra del Fuego, 25.XI.1988. I, H. w.
wagenknechti , aberration, Arroyo de Agua Negra, San Juan,
3. XI. 1988 (first brood).
234
J. Res. Lepid.
Fig. 19. Various Hypsochi/a. A, B, C, D, G, I males; E, F, H females. A, H. w.
wagenknechti, Chile, Los Libertadores, Prov. Los Andes, 27-
28.1.1983 (second brood). B, E, same, Arroyo de Agua Negra, San
Juan, 3.XL1988 (first brood). C, F, H. ga/actodice, Esquel, Chubut,
17.XI.1988 (first brood). D, H. w. wagenknechti, Chile, La Parva,
Prov. Santiago, 24.XL1982 (first brood). G, H, H. ga/actodice, bred
without diapause ex Esquel, Chubut, emerged 23-25. XII. 1988. I,
Species uncertain ( wagenknechti ? ga/actodice ?), Cordon del Viento,
Neuquen, 28.1.1985.
28(3):137-238, 1989(91)
235
Fig. 20. Phulia nymphula. A-D, males; E-H, females. A, B, E, F, G, all Paso
Bermejo, Mendoza, 31 .X-1 .XI.1988 (1st brood; G is a white, male-
like female, E is phenotypically similar to Pierphulia above). C,
Cordon del Viento, Neuquen, 28.1.1985. D, H, Esquinas Blancas,
Jujuy, 7.11.1984. D and FI are quite typical of puna specimens, with
longer wings, heavier female pattern, and a more straw, less pinkish
color below.
236
J. Res. Lepid.
Fig. 21 . Mosaic gynandromorph Phulia nymphu/a from Paso Bermejo, Men-
doza, 31.X-1.XI.1988. The left FW is apparently all male, the right
FW all female, both HW mostly female (left 7/8, right 2/3); the
external genitalia are malformed and intermediate. This is the first
gynandromorph reported in nature for any Andean pierine.
Fig. 22. Co/ias flaveola from Arroyo de Agua Negra, San Juan, 3. XI. 1988,
males at left in both views.
28(3):137-238, 1989(91)
237
SORENSEN S INDEX OF FAUNAL SIMILARITY
0 0.1 0.2 0.3 0.4 Oi 0.6 0.7 0.8 0.9 1.0
I I I I I I _l I I I
i Cuesta del Obispo - Co. Zapallar
___ * Sierras Pampeanas
Puna de Jujuy - Salta
Puna de Atacama (Chile)
Arroyo de Agua Negra
i Paso Bermejo
| Cordon del Cepo
Precordillera Chilena
Esquel - P.N. Los Alerces
Cordon del Viento
Loncopue
A I limine
Bariloche - P.N. Nahuel Huapi
San Martin - P. N. Lanin
Valle Central (Chile)
Chos Malal
Precordillera Mendocina
Valles Calchaquies
Comodoro Rivadavia - Fitz Roy
Valdivia Region
Laguna del Maule
Rio Gallegos
Chilean Magallanes
Ushuaia - Cordon Martial
Rio Grande
i i 1 1 1 1 1 1 1 i r
0 0.1 0.2 0.3 0.4 Oi 0.6 0.7 0.8 0.9 1.0
SORENSEN S INDEX OF FAUNAL SIMILARITY
Fig. 23. Dendrogram generated by cluster analysis (UPGMAA) showing
relationships among selected faunas compared in Tables 1-3. (For
purposes of calculation, species and subspecies were treated
equally.)
238
J. Res. Lepid.
Fig. 24. Argentine sketch map of the Parque Provincial Aconcagua,
Mendoza, showing (stars) locations of known suitable vegas for the
occurrence of Colias mendozina. Actual captures are from Que-
brada de los Horcones, just NW of Puente del Inca near the middle
of the map.
r
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THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 28 Number 3 Fall 1989(1991)
IN THIS ISSUE
Date of Publication: September 15, 1991
The Zoogeography and Systematics of the Argentine 187
Andean and Patagonian Pierid Fauna
Arthur M. Shapiro
Cover Illustration: Female Tatochila m. mercedis ovipositing on high-
altitude rosette Crucifer near the Argentine-Chilean border in the Cordillera
Real. Photograph by Arthur M. Shapiro.
05
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 28
Number 4
Winter 1989(1991)
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
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Journal of Research on the Lepidoptera
28(4):239-257, 1989(91)
Konrad Fiedler
Zooiogisches Institut II der Universitat Wurzburg, Rontgenring 10, D
Wurzburg, Fed, Rep. Germany
European and North West African
(Lepidoptera) and their associations
Abstract. The information about ant-associations of European and
North West African lycaenid caterpillars is summarized. A tentative
classification of the different degrees of myrmecophily is proposed.
More than 75 % of the species considered are myrmecophilous. In the
Poiyommatini, even more than 90 % are ant-associated. An apparent
correlation between the ant-association of lycaenids and their system-
atic position is discussed. This is contradictory to a recent work that
suggested a much lower proportion of myrmecophilous species in the
Western Palaearctic region and rejected any connections between
lycaenid phylogeny and myrmecophily.
Introduction
The association of lycaenid larvae and ants, termed myrmecophily, has
been the subject of extensive research. During the last decade, in
particular, a number of papers concerned with the physiology, ecology,
and evolution of lycaenid myrmecophily have been published (e.g. Fierce,
1983, 1985, 1987; Pierce and Mead, 1981; Pierce and Eastseal, 1988;
Henning, 1983 a & b; Cottrell, 1984; Fiedler and Maschwitz, 1988 a & b,
1989 a & b). However, a comprehensive compilation of all information
regarding lycaenid myrmecophily from a single biogeographical area has
not yet been undertaken. In 1969, Malicky listed the ant-associations of
European species recorded until then, but in the past 20 years consider-
able progress has been made in the investigation of larval life-histories,
adding numerous records of ant-larvae associations. Thus, it seems
, justified to summarize the current knowledge of Western Palaearctic
i lycaenids and their ant-associations, to point out the still significant gaps
! in our knowledge, and to discuss the results in the light of the recent
research on myrmecophily and, in particular, the biogeographical and
evolutionary hypotheses of Pierce (1987).
In the present paper I have attempted to gather all available, but
scattered information on ant-associations and the presence of myrme-
i cophilous organs in European and North West African lycaenid species.
Important sources of data were the most useful review papers of Warnecke
j (1932/33), Hinton (1951), Malicky (1969), and Kitching and Luke (1985).
! Further information was derived from the books of Weidemann (1986,
1 1988) and SEN (1987), but special efforts were devoted to sample data
! from original sources, e.g. numerous journal papers and observations of
several colleagues. Despite my efforts to approach completeness, the
240
J. Res. Lepid.
following list is certainly still far from being complete, and further
additions will be welcome.
Methods
In the present study I have considered all lycaenid species known from Europe
and North West Africa north of the Sahara desert as listed in Higgins and Riley
(1978) and Kudrna (1986), and the available information regarding the presence
of myrmecophilous organs and ant-associations was compiled. The systematic
arrangement basically follows Scott and Wright (1990). Thus, the “Theclinae”
sensu Eliot (1973) are regarded as paraphyletic and are replaced by the (more
likely monophyletic) tribes Aphnaeini, Theclini, and Eumaeini. The former
subfamilies Lycaeninae and Polyommatinae are likewise downgraded to tribal
level, i.e. Lycaenini and Polyommatini, respectively. The phylogenetic relation-
ships of these tribes to each other, as well as to the other lycaenid subfamilies
(Poritiinae including Liptenini; Miletinae including Liphyrini; Curetinae) are
not yet sufficiently established. Indeed, some of the taxa considered here strongly
need further confirmation as monophyletic groups. Because a more satisfactory
phylogenetic higher classification of the Lycaenidae is not yet available, the
arrangement adopted here is necessarily tentative.
The Riodinidae (with the single European species Hamearis lucina L.) are
treated as a separate family, their associations with ants being based on the
convergent evolution of structurally different ant-organs (see Cottrell, 1984;
DeVries, 1988 & 1990) which occur only in the most apomorphic tribes Eurybiini,
Lemoniini, and Nymphidiini (Harvey, 1987). It is important to stress that
riodinid myrmecophily should be discussed as a phylogenetically separate and
functionally convergent phenomenon when compared with ant-associations of
the Lycaenidae (DeVries, 1990).
Nomenclature and taxonomy largely follow Kudrna (1986) with only minor
deviations. The subgenera of the genus-groups Plebejus and Polyommatus are
basically treated as in the paper of Zhdanko (1983). Exceptions are the taxa
Eumedonia (here included in Aricia ), Lysandra (including Plebicula sensu
Higgins and Riley, 1978) and Meleageria (both as subgenera of Polyommatus ).
The generic classification within most groups still strongly requires a phyloge-
netic analysis.
The species concept is in most cases adopted from Higgins and Riley ( 1978) and
Kudrna (1986). Exceptions are, for example, the Plebejides and Lysandra groups
(after Schurian, 1989, Balint & Kertesz 1990) and a few taxa of Agrodiaetus. In
the latter subgenus, a number of taxa has been described from the Mediterranean
area solely based on chromosome studies, but their taxonomic status needs
further investigations. In general, subspecies and local forms are not considered
separately, but highly isolated endemic forms (e.g. from several southern Euro-
pean mountain areas) are treated as distinct species (instead of vicariant
subspecies) because of their genetic separation.
Based on the records available, I have tentatively assigned the degree of larval
myrmecophily to each species where possible. This assignment is based primarily
on field records, laboratory experiments are considered only exceptionally. The
scoring is as follows:
0: not ant associated: myrmecoxenous.
1: very few ant-associations reported, stable ant-associations formed only
exceptionally: weakly myrmecophilous.
28(4):239-257, 1989(91)
241
2: a varying proportion of larvae attended by ants, intermediate between 1 and
3: moderately myrmecophilous.
3: most if not all mature larvae ant-associated: steadily myrmecophilous.
4: larvae dependent on ants as commensales ( Cigaritis ) or parasites (. Maculinea ):
obligately myrmecophilous.
Usually the scorings in Table 1 refer to the larvae. Where the pupae are known
to have a different stage of myrmecophily, this is indicated separately.
The presence of larval myrmecophilous organs is represented in Table 1 as
follows: two asterisks (**) indicate a complete set of ant-organs (a dorsal nectary
organ (DNO) plus a pair of eversible tentacle organs (TOs)); one asterisk means
that only a DNO is present (sometimes only rudimentary, e.g., Gallop hrys:
Fiedler 1990b). Species without an asterisk possess only pore cupola organs
(PCOs) (see Cottrell, 1984 and Fiedler, 1988 for terminology, details and
references).
When the figures indicating the degree of myrmecophily, or the asterisks
referring to the equipment with ant-organs are bracketed, the respective assign-
ments are hypothetical. In these cases the assignment is based on the status of
closely related species (example: in Agrodiaetus the early instars of only a few
species are well-known and all are highly ant-associated; judging from the close
affinity between these taxa, it seems very likely that all Agrodiaetus caterpillars
possess a full set of myrmecophilous organs and are steadily myrmecophilous).
Where possible from the records available, the ant genera and/or species
involved in myrmecophilous interactions are listed. It has to be emphasized that
many determinations were not checked by ant specialists, thus several old
records are reliable only on genus level. For example: Lasius “niger”, aalienus”,
and “flavus” all are complexes of several closely related ant species which are
nearly indistinguishable for the non-myrmecologist (e.g. Seifert, 1988 and pers.
comm.). The term “indet.” means that ant-associations have been observed but
the ants involved were not determined. A question mark (?) indicates that ant-
associations are likely, but no certain information is available. The dash (-)
means that ant-associations have never been reported for the respective lycaenid
species.
Regarding the sources of data, I have listed the review papers and books as
references where appropriate to faciliate use. Special journal articles and
personal communications are only cited when the information in the major
references is incomplete or even incorrect. Purely descriptive papers (e.g. rearing
records) are excluded except when they yield the only information about the
presence or absence of ant-organs. Similarly, laboratory observations on
myrmecophily are only considered when no sufficient field data are available. For
most species the knowledge is still far from being complete. In several cases
(species from Southern Europe) only information from outside Europe (e.g.
Africa) is available, and for a few taxa there is no information regarding the larval
biology at all (see footnotes to Table 1).
The information sampled in Table 1 is used for a quantitative analysis to
determine the number and proportion of myrmecophilous species in the Western
Palaearctic fauna. In Table 2a only those species are considered where appropri-
ate information on the larval biology, including direct (positive or negative)
evidence concerning myrmecophily, is present. In Table 2b, in addition, all
species are included where at the current stage of our knowledge a reasonable
242
J. Res. Lepid.
hypothetical assignment can be made (based on the presence of myrmecophilous
organs and/or the state of the closest relatives). In accordance with experimental
work (e.g. Fiedler and Maschwitz, 1989a, Fiedler, 1990a), the presence of a
functional DNO is considered to indicate a facultative ant-association at least,
while in species without ant-organs (DNO and TOs) such associations are
supposed to be non-existent or weakly developed at most.
Results and Discussion
In Table 1 the European and North West African lycaenid species are
listed together with the information about their ant-associations. The
first column contains the species name, the second column gives the
tentative assignment of the degree of larval myrmecophily. In the third
column the ant genera or species involved are given, and the last column
contains the source of data. Table 2 summarizes the absolute numbers
and percentages of myrmecophilous species.
It is apparent from the Tables 1 and 2 that, in the Western Palaearctic
region, the vast majority (more than 75 %) of lycaenid species are
myrmecophilous at least towards the end of the larval stage. This
conclusion can be drawn from either direct evidence (Table 1, 2a) or
hypothetical assignments (Table 2b), which both yield almost identical
figures. Thus there is a marked contrast to the results of Pierce (1987),
who stated that only 30 % of the European species (32 % of the genera)
are myrmecophilous. The reason for this difference is the incomplete
evaluation of literature records: Pierce’s data are derived solely from the
review papers of Malicky (1969), Batching and Luke (1985), and identi-
fication guides like Higgins and Riley (1978).
On the grounds of her data, Pierce (1987) concluded that there is a
difference in the proportion and obligateness of ant-associated lycaenids
between the northern and southern hemisphere, the latter having 70-90
% myrmecophilous species, the former only 20-40 %. From the results
presented here it becomes obvious that this disparity does not exist for,
at least, the proportion of myrmecophily in Europe; instead, the figures
in Table 2 are close to those for the southern hemisphere given by Pierce.
In Japan, the proportion of myrmecophilous species is about 56 %, but
this rather low figure is mainly due to the preponderance of
myrmecoxenous Thecliti there (Fiedler, 1990 a). The systematic struc-
ture of the Japanese lycaenid fauna is thus not representative for the
whole Eastern Palaearctic region. From the Nearctic region a consider-
ably smaller proportion of myrmecophilous species has been definitely
recorded (about 30 %: Fiedler, 1990 a), but this awaits further confirma-
tion (see Ballmer and Pratt, 1988).
Whether the degree of myrmecophily does show a north-south dispar-
ity remains to be clearly demonstrated. In the Western Palaearctic
region only few species in the genera Maculinea (Thomas et al. , 1989) and
Cigaritis (Rojo de la Paz, 1990) are known to be obligately myrmecophi-
lous. Recent observations on Plebejus argus and P. (Lycaeides) idas in
28(4):239-2S7} 1989(91)
243
Europe, however, suggest that both are obligatorily and specifically
associated with certain ants (. Lasius species from the niger and alienus
groups in the case ofP. argus , species of the Formica cinerea group in the
case ofP. idas; see Mendel and Parsons, 1987, Jutzeler, 1989c & d, 1990,
Ravenscroft, 1990). In the tropics detailed studies on lycaenid-ant inter-
actions are still rather sparse. Only from South Africa (e.g. Clark and
Dickson, 1971, Henning, 1983a, b) and from 'Australia (Common and
Waterhouse, 1981) sufficient information is present on a larger number
of species, while in South Asia most records are merely anecdotal (cf.
Corbet and Pendlebury, 1978), and the life-histories of Neotropical
Lycaenidae are largely unknown. The data from South Africa indeed
suggest a high proportion (about 50 %) of obligatorily myrmecophilous
lycaenid species, largely due to the great diversity of the tribe Aphnaeini
( Aloeidesy Poecilmitis etc.) and the genus Lepidochrysops in Africa. In
Australia, the Lucia and Zesius sections of the Theclini contain a rather
high number of obligate myrmecophiles (> 30 % of all Australian
lycaenids), whereas the situation in South Asia appears to be intermedi-
ate (10-20 % obligate myrmecophiles; Fiedler, 1990 a). Thus, the current
stage of our knowledge does not conclusively support Pierce’s hypothesis
concerning the general north-south disparity in the obligateness of
lycaenid-ant interactions. Rather the high proportion of obligate myrme-
cophiles among the lycaenids of South Africa and Australia may reflect
the peculiar history of the latter 2 areas (Fiedler, 1991).
In Europe, most lycaenids are associated non- specifically with a vari-
ety of ant species, often from different subfamilies. Only about 10 species
from the genera Cigaritis , Maculinea and Plebejus maintain species- or
at least genus-specific relationships with ants (see above). As was
already pointed, out by Malicky (1969), the dominance structure of the
ant fauna in the larval habitats is decisive for which ant species actually
tends a lycaenid, caterpillar. In fact, members of any ant genera that
maintain trophohiotic relationships with other organisms producing
nutritive liquids (e.g. homopterans, myrmecophytes) are likely to attend
myrmecophilous lycaenid larvae (DeVries, 1991). Because of the general
dominance of FormMnae ants in temperate regions (e.g. Seifert, 1986;
Fellers, 1987, 1989) it is not surprising that the dominant trophohiotic
formicine genera Lasius (recorded with 23 lycaenid species), Formica (14
species), Camponotus (10 species) and Plagioiepis (10 species), as well as
Tapinoma (Dolichoderinae; with 12 lycaenid species), Myrmica
(Myrmicinae; 20 species) and Crematogaster (10 species) are mentioned
most often in Table 1.
Although the higher classification of the Lycaenidae is not yet resolved,
another pattern is apparent from the results above: There is a strong
correlation between systematic position and myrmeeophily (Fiedler,
1990a). Most members of the Lycaenini, for example, have no ant-
associations, presumably due to the absence of a dorsal nectary organ.
Only for one European species, Lycaena dispar , there exist old records of
244
J. Res. Lepid.
ant-associations, while the remaining 12 European species appear to be
myrmecoxenous. In North America only 4 out of 15 species of the genus
Lycaena are with certainty known to associate with ants with the help of
specialized dendritic setae (Ballmer and Pratt, 1988). Ant-associations
are unknown from African and New ZealandLycaena (Clark and Dickson,
1971, Gibbs, 1980), from Asian Heliophorus (Eliot, pers. comm.) and from
Papuan Melanolycaena (Sibatani, 1974). Thus, the Lycaeninae as a
whole seem to be a myrmecoxenous group with only few secondary
exceptions.
The Polyommatini show the reverse pattern: nearly all European
species are ant-associated (Table 1), the only exceptions being the
subgenera Agriades and Vacciniina. Both occur in arctic or alpine
tundra, or wet boreo-montane bogs with limiting nutritional resources
and few ant species present. The Polyommatini of Africa, Australia, and
North America also contain a large number of myrmecophilous species
(e.g. Clark and Dickson, 1971, Common and Waterhouse, 1981, Ballmer
and Pratt, 1988) with rather few exceptions (e.g. desert species, lycaenids
with endophytic larvae). Thus, the Polyommatini are basically ant-
associated and reductions of myrmecophily (and ant-organs) have oc-
curred in only a few species that occur where the ecological conditions did
no longer favor the symbiosis with ants.
The predominantly African tribe Aphnaeini is another strongly myrme-
cophilous group: the high proportion of obligately myrmecophilous spe-
cies in South Africa is mainly due to the Aphnaeini genera Aphnaeus,
Apharitis, Spindasis, Aloeides, Phasis, Poecilmitis, and the polyommatine
genus Lepidochrysops (Clark and Dickson, 1971; Claassens and Dickson,
1980) . The only representatives of the Aphnaeini in the Palaearctic
region (genus Cigaritis ) are specifically associated with ants of the
myrmicine genus Crematogaster (Rojo de la Paz, 1990).
The remaining and rather heterogeneous tribes Theclini and Eumaeini
show different pictures. The Theclini contain a large number of ant-
associated species in South Asia, Africa, and Australia (Clark and
Dickson, 1971, Corbet and Pendlebury, 1978, Common and Waterhouse,
1981) . The 3 European species as well as the 2 North American members
of this tribe, however, belong to the mainly Sino-Oriental sub tribe
Thecliti, and this whole subtribe has apparently reduced its ant-associa-
tions. Possibly the Thecliti (as the temperate-zone sister-group of the
Arhopaliti, a basically Oriental ant-associated lineage) reduced their
myrmecophily when adaptaing towards temperate regions. In South
East Asia and Australia, the Theclini subtribes Luciiti, Zesiiti, Ogyriti,
and Arhopaliti are predomionantly myrmecophilous, including a num-
ber of obligately ant-associated species (e.g. Common and Waterhouse,
1981, Fukuda et al., 1984, Fiedler, 1990a).
The tribe Eumaeini sensu Scott and Wright (1990) is the largest of the
whole family Lycaenidae. Myrmecophily and myrmecophilous organs
are known from its subtribes Amblypodiiti, Catapaecilmatiti, Loxuriti,
28(4):239-257, 1989(91)
245
lolaiti, Deudorigiti, and Eumaeiti (Fiedler, 1990a). In Europe there are
only a few representatives of the genus Tomares and the subtribe
Eumaeiti. The Tomares species possess a complete set of ant-organs and
are facultatively ant-associated. Tomares belongs to the Deudorigiti
(Eliot, pers. comm.) which subtribe contains a number of myrmecophi-
lous species in the tropics (e.g. Clark and Dickson, 1971).
The Eumaeiti are most diverse in America (Eliot, 1973) including the
species-rich genera Callophrys, Strymon, Satyrium. In the Palaearctic
region, comparatively few species (< 60) of the genera Satyrium s. 1. and
Callophrys s. 1. occur (Bridges, 1988). Within the genus Satyrium there
appears to be a marked tendency to reduce ant-associations, and this is
even more pronounced in Callophrys where only very few species have
been found to be tended by ants. Interestingly, both genera lack the
tentacle organs (Ballmer and Pratt, 1988), and the dorsal nectary organ
(“honey gland”) - though present - does not secrete nutritive liquids in
some species (e.g. Callophrys ruhi , Fiedler, 1990b). Hence, the Eumaeini
of the temperate regions seem to be a tribe with a basically low level of
myrmecophily and a high tendency to further reduce ant-associations
and the related organs. Unfortunately the current knowledge of the
biology of neotropical Eumaeiti is still too fragmentary to support further
interpretations. The ecological regimes selecting for the parallel reduc-
tion of myrmecophily in the Theclini and Eumaeini are not as clear as in
the Polyommatini. Possibly the preference for rather nutrient-poor food-
plants of the families Fagaceae, Betulaceae, Salicaceae and others may
play a role, together with the generally lower diversity and abundance of
ants in temperate woodlands (Jeanne, 1979), resulting in a lower chance
of maintaining stable trophobiotic associations there. Most of the
Palaearctic Polyommatini species, in contrast, occur in open habitats
which support a more diverse ant fauna (Seifert, 1986). Clearly further
studies are required on this interesting evolutionary feature.
It is now relevant to asses the importance of these results with respect
to the biogeography of lycaenid myrmecophily. Apparently the system-
atic composition of the lycaenid fauna of the biogeographical regions
largely influences the proportion and obligateness of myrmecophily. The
lycaenid fauna of Europe, for example, is predominated by species of the
Polyommatini (72 %), resulting in a high proportion of (at least faculta-
tively) ant-attended lycaenids. The Lycaenidae fauna of North America,
in contrast, contains a higher percentage of Eumaeiti and Lycaenini
species (67 % of the resident species). Not surprisingly, the proportion of
myrmecophily is lower. Among the African lycaenids, two thirds of the
fauna belong to the Aphnaeini, Old World sub tribes of the Eumaeini, and
Polyommatini: Most of these species are myrmecophilous. The remain-
ing third are species of the Poritiinae whose larvae in most cases do not
maintain close ant-associations (Clark and Dickson, 1971).
Thus, contradictory to the conclusions of Pierce (1987: “The distribu-
tion of ant association within the Lycaenidae is independent of phylog-
246
J. Res. Lepid.
eny”), this study suggests a high correlation between lycaenid phylogeny
and the evolution of myrmecophily. It also suggests that this systematic
effect may significantly influence the biogeography of lycaenid-ant-
associations. Notwithstanding the uncertainties of lycaenid systematics,
a substantial discussion of the biogeography and evolution of lycaenid
myrmecophily can hardly be realized without a phylogenetic approach.
Much more work needs to be done on the higher classification of the
Lycaenidae as well as in the description and analysis of larval life-
histories and ant-associations in all biogeographical regions to confirm
or reject the hypotheses given above. This paper is a first attempt to
understand a small region (Europe), and is intended to stimulate broader
and more thorough analyses.
Acknowledgements. My sincere thanks are addressed to all friends and colleagues
who contributed to this paper: to Dr. Gregory R. Ballmer, Dr. Phil J. DeVries, Lt.
Col. John N. Eliot, and Dr. David M. Wright for their discussions on lycaenid
phylogeny; to Zsolt Balint, Nicolas W. Elfferich, Andreas Hornemann, David
Jutzeler, Dr. Miguel L. Munguira, Dr. Alain Rojo de la Paz, Dr. Klaus G. Schurian,
Hans-J. Weidemann, and Martin Wiemers for reprints of their papers and much
useful information about lycaenid myrmecophily; Dr. Phil J. DeVries and Dr. Rudi
H. T. Mattoni for their substantial improvements of the manuscript; to Dr.
Bernhard Seifert for the determination of ants; and to my friend Wolfgang A.
Nassig for his permanent readiness to criticize and encourage that work. This
study was funded by a dissertation grant of the Studienstiftung des deutschen
Volkes, and by a postdoc fellowship from the Leibniz award of the Deutsche
Forschungsgemeinschaft to Prof. Dr. Bert Holldobler.
Literature Cited
Agenjo, R., 1963. Estudio de la “Moradilla del Fresno” Laeosopis roboris (Esp.) (Lep.
Lycaen.). Bol. Servicio Plagas Forest. 6:130-140.
Balint, Z., & A. Kertesz, 1990. A survey of the subgenus Plebejides (Sauter, 1986)
- preliminary revision. Linneana Belg. 12:190-224.
Ballmer, G. R., & G. F. Pratt, 1988. A survey of the last instar larvae of the
Lycaenidae of California. J. Res. Lepid. 27:1-81.
Baylis, M., & R. L. Kitching, 1988. The myrmecophilous organs of larvae of the
small blue, Cupido minimus (Lepidoptera; Lycaenidae). J. nat. Hist. 22:861-
864.
Baz, A., 1988. On the distribution of Tarucus theophrastus (Fabricius, 1793)
(Papilionoidea, Lycaenidae) in Morocco. Nota lepid. 11:177-181.
Bell, T. R., 1915. The common butterflies of the plains of India, part 17. J. Bombay
nat. Hist. Soc. 23:481-497.
Blab, J., & O. Kudrna, 1982. Hilfsprogramm fur Schmetterlinge. Okologie und
Schutz von Tagfaltern und Widderchen. Kilda Verlag, Greven. 135 pp.
Bridges, C. A., 1988. Catalogue of Lycaenidae and Riodinidae (Lepidoptera:
Rhopalocera). C. A. Bridges, Urbana/Illinois. vii + 816 pp.
Chapman, T. A., 1915a. On the early stages of Latiorina (Lycaena) pyrenaica, Boisd.
Trans, entomol. Soc. London 1915:397-410, 18 plates.
, 1915b. A contribution to the life history of Agriades escheri, Hb. Trans.
entomol. Soc. London 1915:411-423, 21 plates.
28(4):239-257» 1989(91)
247
— , 1915c. Notes on the early stages of Soditantides orion , Pall. Trans, entomol.
Soc. London 1915:424-427, 14 plates.
Chapman, T. A., & P. A. Buxton, 1919. Contributions to a life-history of Tarucus
mediterraneae Bethune-Baker. Entomol. mon. Mag. 55:163-173.
Clark, G. C., & C. G. C. Dickson, 1971. Life histories of the South African lycaenid
butterflies. Purnell, Cape Town, xvi + 272 pp.
Claassens, A. J. M., & C. G. C. Dickson, 1980. The butterflies of the Table Mountain
Range. C. Straik Publ., Cape Town. 160 pp.
Common, I. F. B., & D. F. Waterhouse, 1981. Butterflies of Australia. 2nd ed., Angus
& Robertson Publ., London/Sydney/Melbourne/ Singapore/Manila. 682 pp.
Corbet, A. S., & H. M. Pendlebury, 1978. The butterflies of the Malay Peninsula.
3rd ed. revised by J. N. Eliot. Malayan Nature Soc., Kuala Lumpur, xiv + 578
pp., 35 plates.
Cottrell, C. B., 1984. Aphytophagy in butterflies: its relationship to myrmecophily.
Zool. J. Linn. Soc. 79:1-57.
DeVries, P. J. , 1988. The larval ant-organs of Thisbe irenea (Lepidoptera: Riodinidae)
and their effects upon attending ants. Zool. J. Linn. Soc. 94:379-393.
, 1990. Evolutionary and ecological patterns in myrmecophilous riodinid
butterflies. In: Cutler, D. F., & C. Huxley (eds.): Interaction between ants and
plants. Oxford Univ. Press.
, 1991. Mutualism between Thisbe irenea butterflies and ants, and the role of
ant ecology in the evolution of larval-ant associations. Biol. J. Linn. Soc. (in
press).
Dujardin, F., 1972. Description de Callophrys avis barraguei subspecies nova et
observations biologiques sur la sous-espece nominative. Entomops 6:7-18.
Eliot, J. N., 1973. The higher classification of the Lycaenidae (Lepidoptera): a
tentative arrangement. - Bull. Brit. Mus. nat. Hist. (Entomol.) 28:371-505, 6
plates.
Emmet, A. M., & J. Heath (eds.), 1990. The moths and butterflies of Great Britain
and Ireland. Vol. 7/1: The butterflies. Harley Books, Colchester, ix + 370 pp.
Fellers, J. H.. 1987. Interference and exploitation in a guild of woodland ants.
Ecology 68:1466-1478.
— , 1989. Daily and seasonal activity in woodland ants. Oecologia 78:69-76.
Fiedler, K. 1987 Tetramorium caespitum (Linnaeus, 1758) (Hymenoptera:
Formicidae), eine weitere mit Polyommatus coridon (Poda, 1761) (Lepidoptera:
Lycaenidae) vergesellschaftete Ameisenart. Nadir, entomol. Ver. Apollo
Frankfurt, N. F. 8:60.
— , 1988. The preimaginal epidermal organs of Lycaena tityrus (Poda, 1761) and
Polyommatus coridon (Poda, 1761) (Lepidoptera: Lycaenidae) - a comparison.
Nota lepid. 11:100-116.
- • * , 1990a. Untersuchungen zur Funktion, Okologi© und Evolution der
Myrmekophilie der Blaulinge (Lepidoptepa: Lycaenidae). iv + 219 pp., Ph. D.
dissertation, University of Frankfurt.
, 1990b. Bemerkungen zur Larvalbiologie von Callophrys ruhi L. (Lepidoptera:
Lycaenidae). Nadu:, entomol. Ver. Apollo Frankfurt, N. F 11:121-141.
— , 1991. Zoogeographische Aspekte der myrmekophilen Beziehungen der
Blaulinge (Lepidoptera: Lycaenidae). Verb, dtsch. zool. Ges. 84 (in press).
Fiedler, K., & U. Maschwitz, 1988a. Functional analysis of the myrmecophilous
relationships between ants (Hymenoptera: Formicidae) and lycaenids
(Lepidoptera: Lycaenidae). II. Lycaenid larvae as trophobiotic partners of ants
- a quantitative approach. Oecologia 75:204-206.
248
J. Res. Lepid.
, 1987 [1988b]. Functional analysis ... III. New aspects of the function of the
retractile tentacular organs of lycaenid larvae. Zool. Beitr., N. F. 31:409-416.
, 1989a. Functional analysis ... I. Release of food recruitment in ants by
lycaenid larvae and pupae. Ethology 80:71-80.
, 1989b. The symbiosis between the weaver ant, Oecophylla smaragdina, and
Anthene emolus, an obligate myrmecophilous lycaenid butterfly. J. nat. Hist.
23:833-846.
Fiedler, K., & K. Rosciszewski, 1990. Myrmica sahuleti (Hym. Formicidae) - ein
weiterer Symbiosepartner von Polyommatus coridon (Lep., Lycaenidae). Nachr.
entomol. Ver. Apollo Frankfurt, N. F. 11: 141-142.
Fukuda, H., E. Hama, T. Kuzuya, A. Takahashi, M. Takahashi, B. Tanaka, H. Tanaka,
M. Wakabayashi & Y. Watanabe, 1984. The life histories of butterflies in Japan,
vol. 3. Hoikusha Publ., Osaka.
Gibbs, G. W., 1980. Reinstatement of a New Zealand copper butterfly, Lycaena
rauparaha (Fereday, 1877). New Zealand J. Zool. 7:105-114.
Harvey, D. J., 1987. The higher classification of the Riodinidae(Lepidoptera). Ph.D.
dissertation, University of Texas, Austin.
Harvey, D. J., & T. A. Webb, 1980. Ants associated with Harkenclenus titus ,
Glaucopsyche lygdamas, and Celastrina argiolus (Lycaenidae). J. Lepid. Soc.
34:371-372.
Hemmersbach, A., 1989. Zur Tagfalterfauna Kretas - Beobachtungen auf einer
Rundreise. Verb, westdtsch. Entomol. -Tag 1988:334-344.
Henning, S. F., 1983a. Biological groups within the Lycaenidae (Lepidoptera). J.
entomol. Soc. sth. Afr. 46:65-85.
, 1983b. Chemical communication between lycaenid larvae (Lepidoptera:
Lycaenidae) and ants (Hymenoptera: Formicidae). J. entomol. Soc. sth. Afr.
46:341-366.
Hesselbarth, G., &K. G. Schurian, 1984. Beitrag zur Taxonomie, Verbreitung und
Biologie von Tomares nogelii (Herrich-Schaffer, 1851) in der Turkei (Lepidoptera,
Lycaenidae). Entomofauna 5:243-250.
Higgins, L. G., & N. D. Riley, 1978. Die Tagfalter Europas und Nordwestafrikas.
2nd ed., Parey, Hamburg/Berlin. 377 pp.
Hinton, H. E., 1951. Myrmecophilous Lycaenidae and other Lepidoptera - a
summary. Proc. Trans. Sth. London entomol. nat. Hist. Soc. 1949-50:111-175.
Jarvis, F. V. L., 1958/59. Biological notes on Aricia agestis in Britain. - Entomol. Rec.
J. Var. 70:141-148, 169-187; 71:169-178, 2 plates.
Jeanne, R. L., 1979. A latitudinal gradient in rates of ant predation. Ecology
60:1211-1224.
Jordano, D., J. Fernandez Haeger & J. Rodriguez Gonzalez, 1990. The life-history
of Tomares ballus (Fabricius, 1787): phenology and host plant use in southern
Spain. J. Res. Lepid. 28:112-122.
Jutzeler, D. , 1989a. Einige weitere Eiablage- und Futterpflanzen von Polyommatus
eras Ochs, und Cupido minimus Fuessl. (Lepidoptera: Lycaenidae). Mitt,
entomol. Ges. Basel, N. F. 39:82-87.
, 1989b. Maculinea rebeli (Hirschke). Ein Raupenfund im Glarnerland
(Lepidoptera: Lycaenidae). Mitt, entomol. Ges. Basel, N. F. 39:92-93.
, 1989c. Weibchen der BlauhngsartLy caches idas L. riechen ihre Wirtsameisen
(Lepidoptera: Lycaenidae). Mitt, entomol. Ges. Basel, N. F. 39:95-118.
, 1989d. Kann das Weibchen von Plebejus argus (Linnaeus, 1761) Ameisen
28(4):239-257, 1989(91)
249
riechen? (Lepidoptera: Lycaenidae). Mitt, entomol. Ges. Basel, N. F. 39: ISO-
159.
, 1989e. Fine Besucherameise der Raupe von Lysandra bellargus Rott.
(Lepidoptera: Lycaenidae). Mitt, entomol. Ges. Basel, N. F. 39:92-93.
, 1990. Weibchen von Lycaeides idas (Linnaeus, 1761) riechen ihre
Wirtsameisen, Nachtrage aus den Jahren 1989 und 1990 (Lepidoptera:
Lycaenidae). Mitt, entomol. Ges. Basel, N. F. 40:66-71.
Kitching, R. L., & B. Luke, 1985. The myrmecophilous organs of the larvae of some
British Lycaenidae (Lepidoptera): a comparative study. J. nat. Hist. 19:259-
276.
Konig, F., 1988. Date morfologice, biologice si ecologice referitoare la Philotes
bavins hungarica Dioszeghy 1913 (Lepidoptera, Lycaenidae). 4th. Nation.
Conf. Entomol. Cluj-Napoca, pp. 175-182.
Kudrna, O., 1986. Butterflies of Europe. Vol. 8: Aspects of the conservation of
butterflies in Europe. Aula-Verlag, Wiesbaden. 323 pp.
Larsen, T. B., 1990. Butterflies of Egypt. Apollo Books, Svendborg, 112 pp.
Larsen, T. B., & K. Larsen, 1980. Butterflies of Oman. Bartholomew Books,
Edinburgh, 80 pp.
Larsen, T. B., & A. R. Pittaway, 1982. Notes on the ecology, biology and taxonomy
of Apharitis acamas (Klug) (Lepidoptera: Lycaenidae). Entomol. Gaz. 33:163-
168.
Leigheb, G., E. Riboni & V. Cameron-Curry, 1990. Kretania psylorita Freyer
(Lepidoptera, Lycaenidae). Discovery of a new locality in Crete. Nota lepid.
13:242-245.
Liebig, W. H., 1989. Bemerkungen zur Biologie von Maculinea alcon Schiff. (Lep.,
Lycaenidae). Entomol. Nachr. Ber. 33:117-121.
Malicky, H., 1969. Versuch einer Analyse der okologischen Beziehungen zwischen
Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera). Tijdschr. Entomol.
112:213-298.
Manino, Z., G. Leigheb, P. Cameron-Curry & V. Cameron-Curry, 1987. Descrizione
degli stadi preimmaginali di Agrodiaetus humedasae Toso & Balletto, 1976
(Lepidoptera, Lycaenidae). Boll. Mus. reg. Sci. nat. Torino 5:97-101.
Martin Cano, J., 1981. Similitudes biologicas y diferencias ecologicas entre
Glaucopsyche alexis (Poda) y Glaucopsyche melanops (Boisduval). (Lep.
Lycaenidae). Bol. Estacion centr. Ecol. 10:59-70.
-, 1982. La biologla de los licenidos espanoles (Lep. Rhopalocera). Misc.
Conmemorativa tJniv. autonoma Madrid 1982:1003-1020.
— — , 1984. Biologfa comparada de Lampides boeticus (L.), Syntarucus pirithous
(L.) y Polyommatus icarus (Rot.) (Lep., Lycaenidae). Graellsia 40:163-193.
Martin, J., & P. Gurrea, 1983. Relationship between Nordmannia esculi Huebner
(Lep.: Lycaenidae) and Camponotus cruentatus Latreille (Hym.: Formicidae).
Entomol. Rec. J. Var. 95:153-154.
Maschwitz,U.,M. Wust&K. Schurian, 1975. Blaulingsraupen als Zuckerlieferanten
fur Ameisen. Oecologia 18:17-21.
Mendel, H., & E. Parsons, 1987. Observations on the life-history of the silver-
studded blue, Plebejus argus L. Trans. Suffolk Nat. Soc. 23:2-8.
Munguira, M. L., & J. Martin, 1988. Variabilidad morfologica y biologica de Aricia
morronensis (Ribbe), especie endemica de la Peninsula Iberica (Lepidoptera:
Lycaenidae). Ecologfa 2:343-358.
, 1989a. Biology and conservation of the endangered lycaenid species of Sierra
Nevada, Spain. Nota lepid. 12, Suppl. 1:16-18.
250
J. Res. Lepid.
— — , 1989b, Paralelismo en la biologfa de tres especies taxonomicamente proximas
y ecologicamente diferenciadas del genero Ly sandra: L. dory las, L. nivescens y
L. golgus (Lepidoptera, Lycaenidae). Ecologfa 3:331-352.
Pierce, N. E., 1983. The ecology and evolution of symbioses between lycaenid
butterflies and ants. PhD dissertation, Harvard University, Cambridge/Mass.
, 1985. Lycaenid butterflies and ants: selection for nitrogen-fixing and other
protein-rich food plants. Am. Nat. 125:888-895.
- — -, 1987. The evolution and biogeography of associations between lycaenid
butterflies and ants. Oxvord Surv. evol. Biol. 4:89-116.
Pierce, N.E.,& S. Eastseal, 1986. The selective advantage of attendant ants for the
larvae of a lycaenid butterfly, Glaucopsyche lygdamus. J. Anim. Ecol. 55:451-
462.
Pierce, N. E., & P. S. Mead, 1981. Parasitoids as selective agents in the symbiosis
between lycaenid butterfly larvae and ants. Science 112:1185-1187.
Ravenscroft, N. O. M., 1990. The ecology and conservation of the silver-studded
blue butterfly Plebejus argus L. on the sandlings of East Anglia. Biol. Conserv.
53:21-36.
Rehfous, M., 1954. Fourmis et chenille de Lycenides. Mitt. Schweiz, entomol. Ges.
27:38-42.
Rojo de la Paz, A., 1990. Two new cases of myrmecophily in the Lycaenidae
(Lepidoptera): biology of Cigaritis zohra (Donzel, 1847) and Cigaritis allardi
(Oberthur, 1909) in Morocco. 7th Europ. Congr. Lepid., Lunz/Austria.
SBN, 1987. Tagfalter und ihre Lebensraume. Arten, Gefahrdung, Schutz.
Schweizerischer Bund fur Naturschutz, Basel, xii + 516 pp.
Scott, J. A., & D. M. Wright, 1990. Butterfly phylogeny and fossils. In: O. Kudrna
(ed.): Butterflies of Europe, vol. 2: Introduction to Lepidopterology. Aula-
Verlag, Wiesbaden, pp. 152-208.
Schurian, K. G., 1989. Revision der Ly sandra- Gruppe des Genus Polyommatus
Latr. (Lepidoptera: Lycaenidae). Neue entomol. Nachr. 24:1-181.
Schurian, K. G., & C. D. Thomas, 1985. Zur Biologie von Lysandra punctifera
Oberthur, mit Bemerkungen zu L. bellargus Rottemburg (Lep.: Lycaenidae).
Entomol. Z. 95:225-233.
Seifert, B., 1986. Vergleichende Untersuchungen zur Habitatwahl von Ameisen
im mittleren und sudlichen Teil der DDR. Abh. Ber. Naturkundemus. Gorlitz
59:1-124.
— — , 1988. A revision of the European species of the ant subgenus Chtonolasius
(Insecta, Hymenoptera, Formicidae). Entomol. Abh. staatl. Mus. Tierkunde
Dresden 51:143-180.
Sibatani, A., 1974. A new genus for two new species of Lycaeninae (s. str.)
(Lepidoptera: Lycaenidae) from Papua New Guinea. J. Austr. entomol. Soc.
13:95-110.
Thomas, C. D., 1985. Specializations and polyphagy of Plebejus argus (Lepidoptera:
Lycaenidae) in North Wales. Ecol. Entomol. 10:325-340.
Thomas, J. A., G. W. Elmes, J. C. Warblaw & M. Woyciechowski, 1989. Host
specificity among MacUlinea butterflies in Myrmica ant nests. Oecologia
79:452-457.
Warnecke, G., 1932/33. Ubersicht uber die bisher als myrmekophil bekannt
gewordenen palaarktischen Schmetterlingsraupen der Familie Lycaenidae.
Int. entomol. Z. Guben 26: 165-171, 215-219, 238-242, 285-291, 375-378, 431-
433, 460-462, 479-480, 514-516; 27: 44-46, 121-127, 145-150, 178-180, 305-309,
3 plates.
28(4):239-257, 1989(91)
251
Weidemann, H.-J., 1986. Tagfalter: Entwicklung - Lebensweise, vol. 1. Neumann-
Neudamm, Melsungen. 288 pp.
— — 1988. Tagfalter: Biologie - Okologie - Biotopschutz, vol. 2. Neumann-Neudamm.
Melsungen. 372 pp.
Wiltshire, E. P., 1944. Early stages of oriental palaearctic Lepidoptera. VII.
Entomol. Rec. J. Var. 56:111=114
, 1948. Middle East Lepidoptera, IX: Two new forms or species and thirty-five
new records from Cyprus. Entomol. Rec. J. Var. 60:79-89.
Zhdanko, A. B., 1983. Key to the lycaenid genera (Lepidoptera: Lycaenidae) of the
Soyjet Union, based on characters of the male genitalia. Entomol. Rev. 62:120-
140.
Table 1 : List of species, degree of myrmecophily, and associated ants of
European and North West African Lycaenidae and Riodinidae. Only field records
are included except where stated otherwise (lab.). Further explanations see text
and footnotes.
Species
Degree of
myrmecophily
Ant species
involved
Source of
information
Riodinidae
Hamearis lucina
Lvcaenidae
Aphnaeini:
0
Malicky 1969b (lab.)
Fiedler 1990a
Cigaritis zohra
4**
Crematogaster laestrygon Rojo de la Paz 1 990
C. allardi
3**
Crematogaster auberti
C. antaris
C. scutellaris
Rojo de la Paz 1 990
C. siphax
(3/4**)
?
no record
C. acamas
4**
Crematogaster sp.
Larsen & Pittaway 1 982
C. myrmecophila
4**
Crematogaster auberti
Cataglyphis bicolor?
Hinton 1951
Lvcaenini:
Lycaena phlaeas
0
—
Kitching & Luke 1985
L. helle
0
—
SBN 1987
L. dispar a
0/1?
Myrmica rubra
Hinton 1951
L vi rgaureae
0
—
SBN 1987
L. ottomanusb
(0?)
— .
Elfferich, pers. comm.
(lab.)
L. tityrus
0
__
SBN 1987
L. alciphron
0
—
SBN 1987
L. hippothoe
0
—
SBN 1987
L. candens
(0)
— -
no record
L. thersamon
0
—
Larsen 1 990
L. phoebus
0
—
Rojo de la Paz, pers. comm.
L. thetis
(0)
—
no record
L. ochimus
(0)
—
no record
252
J. Res. Lepid.
Theclini:
Theda betulae
larva: 0/1
pupa: 3
Lasius niger (pupa)
Maiicky 1969b, Kitching &
Luke 1985, Emmet & Heath
1990
Laeosopis roboris
(1 ?r
?
Agenjo 1963
Quercusia quercus
larva: 0/1
pupa: 2
Lasius sp.? (pupa)
Kitching & Luke 1985,
Emmet & Heath 1 990
Eumaeini:
Tomares ballus
2**
Plagiolepis pygmaea
+ indet.
Chapman & Buxton 1919,
Maiicky 1969b, Jordano
et al. 1990
T. mauretanicus
(2)**
?
Maiicky 1969b
T. nogelii
3D
indet.
Hesselbarth & Schurian 1984
Callophrys rubi c
larva: 0/1*
pupa: 2
??
Maiicky 1969b, Fiedler 1990d,
Emmet & Heath 1990
C. avis
(0)*
—
Dujardin 1972
Satyrium w-album
2*
indet.
Maiicky 1969b, Kitching &
Luke 1985, Schurian,
pers. comm.
S. spini
2*
indet.
Maiicky 1969b
S. ilids
2*
Camponotus aethiops
Maiicky 1969b, SBN 1987
S. esculi
2*
Camponotus cruentatus
Martin & Gurrea 1983
S. acadae
0*
—
Schurian, pers. comm.
S. (Fixsenia) pruni
Polvommatini:
Jamides section
0
Kitching & Luke 1985
Lampides boeticus
2**
Lasius niger Hinton 1 951 , Martin Cano
Camponotus compressus 1984; Schurian & Wiemers
C. cruentatus pers. comm.
C. sylvaticus
C. foreli
Prenolepis dandestina
Plagiolepis sp.
Tapinoma melanocephalum
Leptotes section
Leptotes pirithous d
2**
indet.
Hinton 1951, Clark &
Dickson 1971, Munguira,
pers. comm.
L. webbianus 8
0/1 **
?
Wiemers & Schurian, pers.
comm, (lab.)
Castalius section
Tarucus rosaceus
3**
Plagiolepis pygmaea
Camponotus sicheli
Monomorium salomonis
Chapman & Buxton 1919,
Rojo de la Paz,
pers. comm.
T. theophrastus
3**
indet.
Baz 1988
T. balkanicus
3 T)
indet.
Wiltshire 1945, 1948
Zizeeria section
Zizeeria knysna d
3**
Tapinoma
melanocephalum
Warnecke 1932/33, Clark
& Dickson 1971
28(4):239-257, 1989(91)
Cupido section
Eve res argiades d
2**
indet.
Warnecke 1932/33
E. decoloratus
(2*1
?
no record
E. alcetas
(2)**
?
Elfferich, pers. comm,
(lab.), Martin Cano 1982
Cupido minimus
3*
Lasius alienus Malicky 1969b, Baylis
Formica fusca & Kitching 1 988
F. rufibarbis
Plagiolepis vindobonensis
Myrmica rubra
C. lorquinii
3*
Plagiolepis pygmaea
Tapinoma nigerrimum
Munguira & Martin 1989,
Munguira, pers. comm.
C. osiris
3**
Lasius alienus
+ indet.
Malicky 1969b, SBN 1987
Azanus jesousd
3**
indet.
Bell 1915,
Clark & Dickson 1971
Lycaenopsis section
Celastrina argiolus
2**
Lasius niger
L. alienus
L. fuliginosus
Camponotus japonicus
C. nearcticus
Formica subsericea
F. truncorum
Myrmica sp.
Malicky 1969b, Harvey &
Webb 1980, Kitching
& Luke 1985, Emmet &
Heath 1990
Glaucopsyche section
Glaucopsyche alexis
3**
Lasius alienus
Formica pratensis
F. fusca
F. cinerea
F. nemoralis
F. subrufa
Camponotus aethiops
C. maxiliensis
Myrmica scabrinodis
Crematogaster auberti
Tapinoma erraticum
Malicky 1969b,
Martin Cano 1981,
SBN 1987
G. melanops
3**
Camponotus foreli
C. cruentatus
C. micans
C. sylvaticus
Malicky 1969b,
Martin Cano 1981
Maculinea arion
4*
Myrmica sabuleti
M. scabrinodis
Thomas et. al. 1989
M. teleius
4*
Myrmica scabrinodis
M. rubra
M. vandeli
M. sabuleti
Thomas et al. 1989
M. nausithous
4*
Myrmica rubra
M. scabrinodis
Thomas et al. 1989
M. alcon
4*
Myrmica ruginodis
M. rubra
M. scabrinodis
Thomas et al. 1989
Liebig 1989 (lab.)
M. rebeli
4*
Myrmica schencki
M. sabuleti
M. scabrinodis
M. sulcinodis
Thomas et al. 1989,
Jutzeler 1989b
253
254
J. Res. Lepid.
Jolana jolas
2*
Tapinoma erraticum
+ indet.
Warnecke 1932/33,
Malicky 1969b, Schurian,
pers. comm.
Turanana panagaea f
?
?
no record
Pseudophilotes
schiffermuelleri
(2)**
?
no record
Ps. baton
2**
Lasius alienus
Myrmica scabrinodis
Malicky 1969b, Blab &
Kudrna 1982
Ps. panoptes
(2**)
?
Nel 1982
Ps. barbagiae
(2**)
?
no record
Ps. abencerragus
(2**)
?
Martin Cano 1982
Ps. bavius
2*0
indet.
Konig 1988
Scolitantides orion
3**
Camponotus vagus
C. aethiops
Tapinoma erraticum
+ indet.
Chapman 1915c,
Malicky 1969b
Polyommatus section
Chilades trochylus d
3**
Prenolepis spp.
Pheidole quadrispinosa
iridomyrmex sp.
Malicky 1969b, Clark &
Dickson 1971 , Wasserthal,
pers. comm.
Plebejus argus
3/4**
Lasius niger
L. alienus
Formica cinerea ??
Kitching & Luke 1985,
C. Thomas 1985, Mendel &
Parsons 1987, Jutzeler
1989d, Ravenscroft 1990
P. vogelii
(3**)
?
no record
P. (Plebejides) martini
3**
Crematogaster sp.
Rojo de la Paz,
pers. comm.
P. (P.) trappi
3**
Formica lugubris
F. lemani
SBN 1987, Schurian &
Jutzeler, pers. comm.
P. (P.) hespericus
3**
Formica cinerea
F. subrufa
Plagiolepis pygmaea
P. schmitzi
Camponotus cruentatus
C. foreli
C. sylvaticus
Crematogaster auberti
Munguira & Martin 1989a,
Munguira, pers. comm.
P. (P.) sephirus
3**
Lasius near alienus 9 Balint & Kertesz 1 990,
Formica pratensis own observations
Camponotus aethiops
Tetramorium near caespitum 9
P.(Lycaeides) idas
3/4**
Formica cinerea
F. selysi
F. exsecta
F. lemani
F. pressiiabris
F. lugubris
F. lefrancoisi
F. fusca ?
Malicky 1969b, SBN 1987,
Jutzeler 1989c, 1990
P. (L.) argyrognomon
3**
Lasius alienus
L. niger
Myrmica scabrinodis
M. sabuleti
Malicky 1969b, Blab &
Kudrna 1982
P. (Kretania) eurypilus
f 9
?
no record
P. (K.) psylorita
?
?
Hemmersbach 1989,
Leigheb et al. 1990
Malicky 1969b
P. (Vacciniina) optilete 0
28(4):239"257, 1989(91) 255
Polyommatus (Aricia)
agestis
3**
Lasius alienus
L flavus
Myrmica sabuieti
Jarvis 1958/59, Kitching
& Luke 1 985, Emmet & Heath
1990, Schurian, pers. comm.
P. (A.) artaxerxes
3**
Lasius sp.
Maiicky 1969b, SBN 1987
P. (A.) cramera
(3*1
?
no record
P. (A.) morronensis
3**
Lasius niger
Crematogaster auberti
Tapinoma erraticum
T. nigerrimum
Munguira & Martin 1988
P. (A.) nicias
(3)**
?
Warneeke 1932/33
P. (A.) anteros
(3*1
?
no record
P. (A.) eumedon
3**
Myrmica sp.
Malicky 1969b, Weidemann
1986, SBN 1987, Schurian,
pers. comm.
P. (Albulina) orbitulus
(2)**
?
Warneeke 1932/33, SBN1987
P. (Agriades) glandon
0
—
Malicky 1969b, SBN 1987
P. (A.) zueilichi
0
Munguira & Martin 1989
P. (A.) pyrenaicus
0
—
Chapman 1915a
P. (A.) dardanus
(0)
no record
P. (A.) aquilo
(0)
no record
P. (Agrodiaetus) damonT*
Lasius niger
Formica pratensis
Warneeke 1932/33, SBN1987
Malicky 1969b
P. (A.) iphigenia
(3**)
?
no record
P. (A.) dolus
(3**)
?
Martin Cano 1982
P. (A.) ainsae
(3**)
?
Martin Cano 1982
P. (A.) admetus
(3)**
?
Warneeke 1932/33
P. (A.) fabressei
(3)**
?
Martin Cano 1982,
Munguira, pers. comm.
P. (A.) aroanensis
(3**)
?
no record
P. (A.) ripartii
3**
indet.
Munguira & Schurian,
pers. comm.
P. (A.) humedasae
(3)**
?
Manino et ai. 1987
P. (A.) thersites
3**
Lasius alienus
Myrmica scabrinodis
Tapinoma erraticum
Rehfous 1954,
Malicky 1969b,
Schurian, pers. comm.
P. (Cyaniris) semiargus 3**
Lasius sp.
Weidemann 1986
P. (C.) antbchena
(3**)
?
no record
P. (Lysandra) doryias
3**
Formica cinerea
Lasius alienus
Myrmica scabrinodis
Rehfous 1954, Weidemann
1986, SBN 1987
P. (L) golgus
3**
Tapinoma nigerrimum
Munguira & Martin 1989b
P. (L.) nivescens
3**
Tapinoma nigerrimum
Munguira & Martin 1989b
P. (L) atlantica
(3**)
?
no record
P. (L) amandus
3**
Lasius niger
Hornemann, pers. comm.
P. (L) escheri
3**
Formica cinerea
Myrmica specioides
Chapman 1915b, SBN 1987,
own observ.
P. (L) coelestina
(3**)
?
no record
P. (L) coridon
3**
Lasius niger Malicky 1969b, Kitching
L. alienus & Luke 1985, Fiedler 1987,
L flavus Fiedler & Roseiszewski 1990,
L. fuliginosus (??) own observ.
Plagiolepis vindobonensls
Formica rufa
Myrmica scabrinodis
M. sabuieti
M. schencki
Tetramorium caespitum
256
J. Res. Lepid.
P. (L.) hispanus 3’
Plagiolepis pygmaea
Maschwitz et al. 1975,
Schurian, pers. comm.
Schurian, pers. comm.
Malicky 1969b, Blab &
Kudrna 1982, Kitching &
P. (L.) albicans 3** indet.
P. (L.) bellargus 3** Lasius alienus
L. niger
Plagiolepis pygmaea
Myrmica sabuleti
M. scabrinodis
Tapinoma erraticum
Luke 1985, Jutzeler 1989e
P. (L.) punctiferus 3** Monomorium salomonis
Crematogaster scutellaris
P. (Meleageria) daphnis 3** Lasius alienus
Schurian & Thomas 1985
Schurian, pers. comm. &
own obs.
Formica pratensis
Tapinoma erraticum
P. (Polyommatus) icarus 2/3** Lasius alienus
Malicky 1969b,
Martin Cano 1984,
Kitching & Luke 1985,
SBN 1987, Jutzeler 1989d,
Emmet & Heath 1990
L. f lav us
L. niger
Formica subrufa
F. cinerea ?
Plagiolepis pygmaea
Myrmica sabuleti
P. (P.)eroides' (3**) ?
P. (P.) eros 3** Formica lemani
Myrmica gallienii
no record
Jutzeler 1989a
Footnotes to Table 1 :
a Only two (independent?) old records. In the extensive literature about this
locally endangered species and in recent textbooks, no mention of any ant-
associations is given. Like all other European Lycaena species for which
appropriate information is available, L. dispar is probably not truely
myrmecophilous.
b In laboratory experiments larvae of L. ottomanus were rather attractive to
Lasius niger ( Elfferich, pers. comm.).
c Only one very old and doubtful record of an ant-association; see Fiedler
(1990b) for detailed discussion.
d Information concerning myrmecophily only available from outside Europe.
e Wiemers observed no ants attending young larvae of L webbianus in the field.
Schurian, during his laboratory rearing, offered mature larvae to German Lasius
niger. First the larvae were rather unattractive, but after some time they were
palpated by the ants and the DNO was active. Thus the species may be at least
weakly myrmecophilous.
f Larval biology apparently unknown.
9 The Lasius workers collected in Hungary belong to a new species of the alienus
group with distinct pubescence on the clypeus (det. B. Seifert). The species will
be described by Seifert. Workers of the Tetramorium caespitum complex from
Hungary cannot be determined with certainty; sexuals would be required.
28(4):239-257, 1989(91)
257
Table 2: Numbers and percentages of facultatively (category 1=3) or obSigately
(category 4) myrmeoophilous, and of myrmecoxenous (category 0) lycaenid
species in Europe and North West Africa. The first table (a) is based exclusively
on certain field observations; species whose larval biology is insufficiently known
are excluded. The second table (b) is based on all assignments given in Table 1
(i.e. degrees of myrmecophily deduced from the situation in closely related
species or from the presence of larval ant-organs are included),
a)
Tribus
not
ant-associated
facultatively
ant-associated
obligateiy
ant-associated
Aphnaeini
..
1
3
Lycaenini
9
-
-
ThecSini
■ip
2
-
Eumaesni
4
0
-
Polyommatini
4
46
7
total: 82 (100%)
17(20.7%)
55 (67.1 %)
10 (12.2%)
b)
Tribus
not
ant-associated
facultatively
ant-associated
obligateiy
ant-associated
Aphnaeini
.
2
3
Lycaenini
13
-
-
Theclini
-
3
-
Eumaeini
4
7
-
Polyommatini
6
73
7
total: 118 (100%)
23 (19.5%)
85 (72.0 %)
10(8.5%)
Journal of Research on the Lepidoptera
28(4):258-262, 1989(91)
Detecting and recording the calls produced by
butterfly caterpillars and' ants
P.J. DeVries
Dept of Zoology, University of Texas, Austin, Texas 78712
and
Center for Conservation Biology, Stanford University, Stanford, California 94305
Introduction
Chirping crickets, shrilling cicadas, and buzzing flies are familiar
examples of how insects communicate with sound. Some sounds that are
fundamentally important to insect communication systems may, how-
ever, be inaudible to the human ear because they are produced at very low
amplitudes (Gogala 1985; M ark! 1983). For example, low amplitude air-
borne sounds produced by wing-flapping may provide vital cues for
species specific mate recognition in drosophilid flies (Hoy et ah 1.988), or
ants may use substrate-borne vibrations to recruit nestmates to a
resource (Baroni-Urbani et ah 1988). Although many insects may
produce low amplitude signals in their communication systems, investi-
gators require instruments to detect them before they can be studied.
Studies concerned with low-amplitude insect sounds are generally
conducted under laboratory conditions, and employ bulky and typically
expensive detection and recording instruments. However, a particle
velocity microphone and amplifier was recently designed by H. Bennet-
Clark (1984) that is inexpensive, portable, and shows great promise as a
tool for discovering and studying low amplitude insect sounds (e.g.,
Hunter 1987; Hoy et ah 1988). Using the Bennet-Clark particle velocity
microphone I have been able to investigate the low amplitude, substrate-
borne calls produced by riodinid and lycaenid butterfly caterpillars that
form symbioses with ants (DeVries 1.990; 1991). The purpose of this
paper is to briefly describe my methods and experience in detecting and
recording caterpillar and ant calls. My aim is to encourage a broader
interest in the documentation and study of these calls - an area of biology
where much remains to be explored.
The microphone and amplifier
Plans for the particle velocity microphone are found in Bennet-Clark
( 1984). My equipment was constructed by a friend, and modified from the
original design in three ways: 1) the microphone itself is simply wrapped
in flattened brass mesh (Fig la), 2) the monitor switch is spring loaded
to the off position to save battery power, 3) the amplifier was fitted into
an 140 x 75 x 32 mm aluminum box, and 4) the amplifier was fitted to
accept both sizes of headphone jacks (Fig. lb). To reduce bulk I use the
28(4):258-262, 1989(91)
259
smallest set of headphones I could find - not the finest, but easy to pack.
Thus, all the components of the amplifier and microphone are compacted
for easy transport.
The recording stage
A serviceable recording stage can be made of two plastic Petri dishes
with a 75 mm diameter circle cut from their centers (I have used both
circular and rectangular types). The opposing bottoms of the Petri dishes
are fitted together and held in place with 4 nylon screws and nuts, with
a circular membrane .of paper or transparent mylar sandwiched between
the Petri dishes to provide the recording substrate (Fig la). The
interchangeable nature of the membrane will allow recording of caterpil-
lar calls as they are transmitted through different substrate materials
(e.g., plant material, metal, paper, wood).
The stage is supported above a table by an adjustable set of gator-jaw
clamps (lab hands') that are available from laboratory or electronic
supply houses. One of the gator jaws holds the stage, and the other jaw
is used to hold the microphone against the membrane from below (Fig
la&b). The user may want to make a more sophisticated recording stage
set-up, but the one described here is inexpensive, compact, and durable
in the field.
After connecting the microphone to the amplifier, detecting or record-
ing caterpillar calls is done simply by placing a caterpillar on the
membrane and allowing it to walk (be patient as it may take a few
minutes) and monitoring the activity with the headphones. A pair of
entomological forceps is useful for caterpillar manipulations. It is
advisable to occasionally check that the microphone is placed correctly
against the membrane (Fig. la). The cleanest signals are obtained from
lycaenid caterpillars that have been turned on their back - it eliminates
the scratching sound produced by their tarsi gripping the membrane
while walking. In the case of riodinid caterpillars, however, they quickly
right themselves, and typically produce a lot of high frequency back-
ground noise.
Recording
A caterpillar call can be recorded on any tape recorder, but those with
an adjustable gain yield the best results; the automatic gain on some tape
recorders tends to increase unwanted noise on the tape. F or my own work
I use a Marantz PMD - 420 portable cassette recorder / player and record
with high bias tape. While recording a call the tape recorder needs to be
isolated from the surface where the recordings are being made. Other-
wise the microphone will pick up the motor sounds of the tape recorder
transmitted through the table. I do this by cushioning the recorder on a
50mm thick foam pad placed on a chair or box isolated from the table
260
J. Res. Lepid.
'
Figure 1 : (A) Detail of the recording stage showing the gator-jaws, modified Petri
dishes, transparent mylar membrane, and particle velocity microphone set
up for recording caterpillar calls. (B) The amplifier, particle velocity
microphone and recording stage set-up. The tape recorder and head-
phones are not connected. An idea of scale can be gained from the 31cm
square floor tiles in background.
28(4):258-262, 1989(91)
261
surface holding the microphone, amplifier, and recording stage. Sec-
ondly, recording extraneous substrate-borne signals generated from
touching the table, the wires, or the amplifier (the microphone is
extremely sensitive) may be minimized by placing the recording stage on
a piece of foam rubber. Finally, a ground wire connected from the gator-
jaw stage support to the amplifier will further reduce or eliminate line
hum (Fig lb).
Suggestions
A major consideration in obtaining good recordings is the inherent
sensitivity of the equipment - ambient and incidental noise can be a
problem. In many instances the user will find that in addition to
caterpillar calls, the recordings will contain a seeming endless variety of
other sounds: wind, rain, bird, insect, and frog calls, vibrations of people
walking in the building, and perhaps most pestiferous, air conditioning
devices and 50-60 cycle electrical hum. Thus, it is an advantage to record
in a place where the investigator has some control over the environment.
Generally I record late at night in a building where the inhabitants have
left (or have been driven off) with the source of electricity shut off at the
mains, and work with a battery-operated headlamp for illumination.
Under conditions where the investigator cannot switch off the electrical
mains, and experiences severe electrical interference, a copper mesh
Faraday cage may be required. Field recordings are best made in a shed
or tent during the day to minimize picking up the calls of nocturnal
insects and amphibians on the recordings . However, at times rain, wind,
and animal calls can be an annoying problem. Finally, keeping the 9 volt
amplifier battery fresh will help reduce hum and flutter.
The silk normally laid down by walking caterpillars will build up on the
membrane after extended use and allow caterpillars a firm grip on the
substrate and generate unwanted noise as they walk. This source of
irritating high frequency noise can be minimized or avoided by replacing
or cleaning the stage membrane regularly . Using a mylar membrane
will result in cleaner recordings because it minimizes the ‘pops’ produced
by the caterpillar’s tarsi hooking into the substrate, it is easily cleaned,
and it has the added advantage of facilitating visual inspection of the
microphone position (Fig lb).
Ants are obviously important to the study of myrmecophilous caterpil-
lars. Recording ant stridulations must be done in such a way as to avoid
the excessive noise generated by the legs scrambling on the membrane.
Some species will happily walk on the membrane and produce substrate
stridulations or tapping. The industrious investigator may set up the
microphone such that it contacts the side of a container holding a captive
ant colony. However, the few times I tried this the typical frenzied
activity of an ant colony came through loud and clear, thus making the
recordings too cluttered for individual analysis. Holding ponerine or
262
J. Res. Lepid.
myrmecine ants with forceps such that the legs are completely restrained
(or removed), and touching the head or abdomen against the membrane
gives good recordings of ‘alarm’ stridulations.
The equipment described here, the heart of which is the Bennet-Clark
particle velocity microphone, has made it feasible for me to detect and
record caterpillar, pupae, ant and beetle sounds in Ecuador, Panama,
Costa Rica, Belize, the USA, Madagascar, England and Germany. As
simple as it is, my equipment has endured a lot of field time under what
may be termed ‘not exactly sterile laboratory conditions’, yet I have not
experienced any appreciable problems with it. I hope that these methods
described here will be expanded and improved upon through wider use
in the investigations of low amplitude insect sounds. Certainly they have
helped our understanding of the role of sound in caterpillar-ant symbio-
ses.
Acknowledgements . Thanks to R. Hoy for giving me the reference for the particle
velocity microphone, G. Raskin for building it, and a special thanks to H. Bennet-
Clark for originally inventing it , and many pleasant chats during my visit to Oxford
(Henry, I’m your greatest fan). This paper was encouraged by K. Fiedler, N. Greig,
D. Grimaldi, U. Maschwitz, and D. Nash. I gratefully acknowledge the support of
the MacArthur Foundation. This paper is dedicated to the memory of Dexter
Gordon and Gordon B. Small - both of whom were experts in sound.
Literature Cited
Baroni-Urbani, C., M.V. Buser,. & M. Schilliger 1988. Substrate vibration
during recruitment in ant social organization. Insectes Sociaux 35: 241-250.
Bennet-Clark, H.C. 1984. A particle velocity microphone for the song of small
insects and other measurements. J. exper. Biol. 108: 459-463.
DeVries, P. J. 1990. Enhancement of symbioses between butterfly caterpillars
and ants by vibrational communication. Science 248: 1104-1106.
DeVries, P. J. 1991. Call production by myrmecophilous riodinid and lycaenid
butterfly caterpillars (Lepidoptera): morphological, acoustical, functional,
and evolutionary patterns. American Museum Novitates (in press).
Gogara, M. 1985. Vibrational communication in insects (biophysical and
behavioural aspects). IN: Kalmring, K. & Eisner, N. (eds) Acoustic and
Vibrational Communication in Insects. Verlag Paul Parey, Berlin and
Hamburg, pp 117-126.
Hoy, R.R. , Hoikkala, A. & Kaneshiro, H. 1988. Hawaiian courtship songs:
evolutionary innovation in communication signals of Drosophila. Science
240: 217-219.
Hunter, M. 1987. Sound production in larvae of Diurnea fagella (Lepidoptera:
Oecophoridae). Ecol. Entomol. 12: 355-357.
Markl, H. 1983. Vibrational communication. IN: Huber, F. & Markl, H. (eds)
Neuroethology and Behavioral Physiology. Springer Verlag, Berlin Heidelberg,
pp 332-353.
Journal of Research on the Lepidoptera
28(4):263-276, 1989(91)
Genetics and Biogeography of the Oeneis chryxus
Complex (Satyrinae) in California
Adam H. Porter a
and
Arthur M. Shapirob
Department of Zoology, University of California, Davis CA 95616
Abstract. The nominal taxa Oeneis ivallda Mead and Oe. chryxus
Stanislaus Hovanitz are alpine butterflies endemic to the Sierra
Nevada of California. The range of Oe. c. Stanislaus is entirely con-
tained within the range of Oe. ivallda . The two intergrade gradually
in the north and abruptly in the south, and electrophoretic-genetic
analyses fail to demonstrate any interruption in gene flow between
them. This is consistent with the interpretation that ivallda and
Stanislaus are forms of a single species, and we recommend they be
classified as subspecies of Oe. chryxus Doubleday and Hewitson pend-
ing comparisons with Rocky Mountain Oe. chryxus chryxus. Hovanitz’s
(1940, Ecology 21:371) hypothesis that the color morphs are main-
tained by selection for crypsis breaks down in the northern Sierra,
where the pale ivallda morph is often found on dark substrates.
The peculiar distribution of these taxa suggests a double invasion of
the Sierra, with Stanislaus having arrived secondarily from the east
across the Great Basin. We discuss the plausibility of the easterly
colonization route, which remains controversial in the botanical
literature. Further genetic investigation of the chryxus complex may
provide a definitive test of this hypothesis.
Introduction
Oeneis ivallda Mead, a pale, sometimes nearly white Satyrine
butterfly ranging from Nevada to Inyo and Tulare Cos. in California,
is the only truly endemic butterfly to reach both the north and south
alpine limits of the Sierra Nevada. Its biogeographical relationship
with what is presently called Oeneis chryxus Stanislaus Hovanitz has
posed an evolutionary problem now recognized for over 50 yr. This re-
lationship bears in turn on the origin of the alpine biotic community in
this mountain range. Oe. chryxus Stanislaus was described by Hovanitz
(1937) from Sonora Pass, Alpine Co. It does not differ from Oe. ivallda
a Present address: Department of Zoology, University of Canterbury, Christchurch 1,
New Zealand.
6 To whom reprint requests should be directed.
264
J. Res. Lepid.
Fig. 1
121-
40* 120*
AVCASTLE PEAK
J DONNER PASS
. 1?^0 '
^ : . /'LAKE TAHOE
m//
CfiLl’S-//}'.
ECHO PASS n . (p *7FREEl PEAK
■ ) o
PARSON PASS /■/?
/eBBETTS PASS
SQNORA PAg§
WALKER
WASSUK MTS.
YOSEMITE
^?P, .. .....
Q. V-% TIOGA PASS
37°
SEQUOIA
PHENOTYPES
•
ivallda
■
Stanislaus
«■
intermediates
Distribution of alpine habitats in the Sierra Nevada of California with
known localities of Oeneis ivallda and Oe. chryxus Stanislaus, re-
drawn from Hovanitz (1940) with additional confirmed localities
added. Populations sampled in this study are in the larger sized font.
28(4):263-276, 1989(91)
265
in genital morphology, indeed, the only basis for diagnosis is color,
both sexes being a deep “butterscotch” brown. The geographic distri-
bution of Stanislaus was documented by Hovanitz (1940); it is entirely
contained in the central Sierra, from Carson Pass to Tioga Pass, with
pure ivallda distributed parapatrically to the N and S. His map is up-
dated and reproduced here (Fig. 1). The darkest Stanislaus occur
around Sonora Pass, with a gradual decrease in the frequency of dark
phenotypes N-ward to only —5% at Carson Pass (AMS, unpublished
data) and 0% near Donner Pass, but an abrupt transition zone to the S
“showing the entire range of variation from darkest to lightest” near
Tioga Pass (Hovanitz 1940, p. 371). This appears to be a unique geo-
graphical relationship, not reproduced in any pair or set of taxa in the
Sierran flora, for example (G. L. Stebbins, pers. comm.). Virtually the
same phenomenon has been reported in Andean birds, however, under
the rubric “leapfrog variation” (Remsen 1984). In the cases discussed
by Remsen, the N and S populations are treated as two geographic sub-
species, separated by a third, more distinct subspecies.
By 1940 Hovanitz had become convinced ivallda and Stanislaus were
in fact conspecific, although authorities subsequent to Hovanitz have
differed in their treatments. These authorities based their judgment on
inexplicit criteria, without any new biological data to justify them.
Garth and Tilden (1963) combined both names under ivallda as a single
subspecies of the widespread boreal and Rocky Mountain species
chryxus Doubleday and Hewitson. The same authors treated them as
separate species 23 yr later {Oe. ivallda and Oe. chryxus Stanislaus ;
Garth and Tilden 1986), as did Tilden and Smith (1986) and Miller and
Brown (1981). Emmel (1975) also kept them separate, but Scott (1986)
recognized both as subspecies of chryxus. The difference in wing-
pigment chemistry, the only diagnostic character reported to date, is
not in itself a basis for recognizing biological species, especially when
the color phenotypes intergrade. One way to approach the question of
species status independently is to look for biochemical-genetic evidence
indicating an interruption of gene flow (discussed below).
The ivallda/ Stanislaus problem is interesting from a biogeographic as
well as taxonomic standpoint because the localization of Stanislaus in
the central Sierra appears discordant with models of colonization from
the N but potentially concordant with immigration from the E, as
recognized by Hovanitz (1940):
“It may be postulated (1) That the Sierra Nevada was once entirely
populated by a white race and that the brown race has either originated
de novo in the central part or that it has come in via the high Basin
Ranges from other populations of the brown form; (2) that the Sierra
Nevada was once populated entirely by a brown race at either end of
which genes for whiteness developed greater concentrations, or (3) that
a uniform population never did exist in the Sierra Nevada (p. 373). ”
Hovanitz notes correctly that all the Rocky Mountain populations of
266
J. Res. Lepid.
chryxus are brown, and claims that “Individuals from these populations
could more easily reach the Sierra Nevada via the high Basin Ranges
which form a series of ‘stepping-stones’ across the uninhabitable desert
areas . . . than any other way.”
Because the Sierra Nevada is a young range, with most of the major
deformation leading to the modern fault-scarp topography occurring
only in the past 3 MY (Bateman and Wahrhaftig 1966) — corresponding
with the first evidence of glaciation (Curry 1966) — the origin and
evolution of the Sierra Nevada alpine biota represents a relatively
recent event. For this reason, it has received the attention of historical
biogeographers and community ecologists interested in the evolution
of new biotas. The conventional wisdom regarding the origins of the
Sierran alpine biota strongly favors a strictly northerly route of colon-
ization of alpine/boreal taxa (such as Oeneis) (Shar smith 1940, Axelrod
1957, 1977; Chabot and Billings 1972), although many endemic plant
species evolved in situ from dry-adapted continental taxa widespread
at lower altitudes (Stebbins and Major 1965).
Scenarios have been promoted involving easterly immigration of
some small proportion of the alpine biota from the Rocky Mountains
(Harshberger 1911; Major and Bamberg 1967; Major and Taylor 1977),
despite objections in the literature (Axelrod 1976). Although the
sequence of glacial events and the interglacial vegetation in the Sierra
Nevada remains very sketchy (Fullerton 1986), Wells (1983) published
data from wood rat ( Neotoma ) middens demonstrating the presence in
SE Oregon in full-Wisconsinian glacial time of a prostrate juniper
steppe with patterned ground, even below 1500 m. It is not difficult to
visualize Oeneis chryxus living in such a climate. Wells considers that
the low elevation desert trough E of the central Sierra has constituted
a major barrier to the W-ward dispersal of Rocky Mt. organisms. The
possibility that such organisms came in from the NE (N Great Basin)
along the shores of the pluvial lakes, spreading into the Sierra at lower
elevations initially on the E flank, must be taken seriously. At least
one “easterly” scenario has been borne out: Major and Bamberg’s
(1967) prediction that Pinus flexilis (Pinaceae) would be found to have
spread to the Sierra from the E in the Mojave sector has been amply
validated by Wells’ Neotoma data.
The biogeographical and evolutionary-ecological framework for inter-
pretation of the ivallda/ Stanislaus problem differs depending on
whether one is dealing with one species or two. As a single species,
both forms could have arrived in a polymorphic population colonizing
from the N or E; as separate waves of colonists from the same or dif-
ferent directions; or one form could have arisen and spread within the
Sierra. The relevant evolutionary questions would involve the biogeo-
graphy of diagnostic characters alone. This was the type of question
addressed by Hovanitz (1940) when he claimed that visual selection for
background matching (crypsis) by predators would favor the pale
28(4):263-276, 1989(91)
267
ivallda morph on granitic substrates and the darker Stanislaus morph
on andesite, and that the geography of the colors matched that of the
substrates. If the two forms belong to two species, one might interpret
the situation as one of competitive exclusion of ivallda by Stanislaus in
the central Sierra; the most interesting questions would address the
whole genomes of these taxa from the perspective of community
ecology — for example, “What ecological factors limit Stanislaus to the
central part of the high Sierra?” Here, we attempt to test the assump-
tion of conspecificity using electrophoretic characters, in an attempt to
refine evolutionary and biogeographical investigations to the proper
level of analysis.
Materials & Methods
Three samples were available for this study (Fig. 1): from the ridge
between Castle and Basin Peaks, Nevada Co., 2. VII. 1989 {leg. AMS)
(phenotypically pure ivallda , no Stanislaus or intermediates ever
recorded in 18 yr) (n = 20); Carson Spur along Hwy. 88, Alpine Co.,
4. VII. 1989 {leg. J. Mori) (mostly ivallda , approximately 5% inter-
mediate in long series, occasionally approaching full Stanislaus in color)
(n = 19); and Sonora Pass, Alpine-Tuolumne-Mono Cos., 5. VII. 1989
{leg. J. Mori) (phenotypically pure Stanislaus) in = 19). All individuals
were frozen alive at "80°C. The heads and thoraces were homogenized
for analysis; all wings and genitalia were retained as vouchers de-
posited in the Bohart Museum of Entomology at Davis. Every indivi-
dual was scored as ivallda , intermediate, or Stanislaus independently
by both of us; only two individuals (from Carson Spur) were ambiguous
(scored as intermediate by one of us, ivallda by the other).
Electrophoresis protocol followed Ayala et al. (1972) and Geiger and
Shapiro (1986), as modified by Porter and Matoon (1989). 16 loci were
studied: adenylate kinase ( AK-1 ; Enzyme Commission number: 2. 7. 4. 7),
aldolase (ALDO; 4.1.2.13), glucose-6-phosphate dehydrogenase (G6PD;
1.1.1.49), glutamic-oxaloacetic transaminase (two loci: GOT-1, GOT-
2; 2. 6. 1.1), glyceraldehyde-3-phosphate dehydrogenase (GAPDH;
1.2.1.12), a-glycerophosphate dehydrogenase (a-GPD; 1.1. 1.8), hexo-
kinase (HK-1; 2. 7. 1.1), isocitrate dehydrogenase (IDH-1; 1.1.1.42),
malate dehydrogenase (MDH-1, MDH-2; 1.1.1.37), malic enzyme (ME-
1, ME-2; 1.1.1.40), phosphoglucose isomerase (PGI; 5.3. 1.9), phospho-
glucomutase (PGM; 2. 7. 5.1), and the anodally migrating locus of
superoxide dismutase (SOD-2; 1.15.1.1). Zymograms were scored using
letter designations, with the fastest allele in the cathodal direction
given the letter A. Data were analyzed using the computer program
BIOSYS-1 (Swofford and Selander 1981). Formulae for the basic po-
pulation genetic parameters used here are given and discussed in most
introductory population genetics textbooks (e.g., Hedrick 1985) and
will not be repeated here. The banding patterns are entirely consistent
268
J. Res. Lepid .
Table 1.
Allelic frequencies at variable loci.
Population
Locus &
allele
( ivallda )
Castle Peak9
(. ivallda +
intermediates)
Carson Spur6
(. Stanislaus )
Sonora Pass*
AK-1
A
0.950
0.947
0.947
B
0.050
0.053
0.053
GAPDH
A
B
1.000
0.947
0.026
1.000
GOT-1
A
0.079
0.079
B
0.975
0.842
0.605
C
0.025
0.026
D
0.053
0.316
a-GPDH
A
B
1.000
0.974
0.026
1.000
HK
A
0.150
0.211
0.071
B
0.850
0.789
0.929
IDH-1
A
0.100
0.053
0.105
B
0.100
0.105
0.105
C
0.375
0.500
0.553
D
E
0.425
0.316
0.026
0.237
ME-1
A
0.026
0.179
B
C
1.000
0.947
0.026
0.821
PGI
A
0.375
B
0.550
0.868
1.000
C
0.075
0.132
PGM
A
0.625
0.447
0.263
B
0.375
0.553
0.737
3 n - 20. 6 n = 19. c n = 19, except n = 14 at HK & ME-1.
with those expected from segregating alleles of mono™, di-,and tetra-
meric enzyme systems reported for these loci in other organisms (Harris
and Hopkinson 1976; Kitching 1985). Since no breeding program has
been carried out, we made the usual assumptions in treating electro-
morphs as alleles for the purposes of genetic analysis.
28(4):263-276, 1989(91)
269
Table 2. Genetic variability in sample populations. A: mean alleles
per locus, Hobs : observed proportion of heterozygotes, Hexp :
heterozygote proportions expected from Hardy-Weinberg
ratios, P: percent of loci polymorphic, with more than one
allele detected. Standard errors in parentheses.
Population
A
P
Hoi&s
Hexp
Castle Peak
1.6 (0.2)
37.5
0.128(0.054)
0.133 (0.058)
Carson Spur
1.9 (0.3)
56.3
0.138 (0.048)
0.146(0.051)
Sonora Pass
1.6 (0.2)
37.5
0.146 (0.060)
0.132 (0.054)
Results
Allelic frequencies for the nine variable loci are given in Table 1; the
remaining loci (ALDO, GOT-2, G6PD, MDH-1, MDH-2, ME-2, and
SOD-2) were monomorphic. We found no deviations from Hardy-
Weinberg proportions. Genetic variability scores for all populations
are shown in Table 2. These values are in the normal range for most
invertebrates (Thorpe 1983), including butterflies (AHP, AMS, and HJ
Geiger, unpubl.), indicating that there is enough genetic variability
available to permit differentiation of these populations (and taxa) in
the absence of gene flow. Indeed, y2 contingency table analyses in-
dicate statistically significant differences in allelic frequencies among
populations at four of the nine variable loci (GOT-1: p < 0.0003; ME-1:
p < 0.02; PGI; p < 0.00001; PGM; p < 0.006). However, analysis using
Fst (Wright 1931) indicates that despite statistical significance, this
degree of differentiation is biologically minor (GOT-1: FST = 0.133;
ME-1: Fst = 0.083; PGI: FST = 0.212; PGM: FST = 0.088; other variable
loci: Fst < 0.03; mean FST = 0.081).
The weak differentiation among populations is perhaps reflected in a
more familiar way by the low genetic distances (Fig. 2). Using UPGMA
as a clustering algorithm (Sneath and Sokal 1973), eight of ten inde-
pendent analyses grouped Carson Pass (mostly ivallda by wing color)
with Sonora Pass ( Stanislaus ), white the other two grouped Carson
with Castle Peak {ivallda). Notably, the distance between nodes in
these analyses was small, and the greatest genetic distance shown was
always within the range shown by subspecies or consubspecific popul-
ations in other animal groups (Thorpe 1983). There was nothing to sug-
gest the interruption of gene exchange expected across a species bound-
ary. Taken together, the genetic analyses indicate that (i) the Carson
Pass population is more intermediate than wing color data would sug-
gest, and (ii) these populations are only weakly differentiated.
Discussion
Genetic population structure and species -level taxonomy. — There is
nothing in our data which would suggest that the ivallda and Stanislaus
270
J. Res. Lepid.
NEI'S UNBIASED D
0.02 0.01
1 I
0
J
Castle Peakivallda
Carson Pass intermediate
Sonora Pass Stanislaus
0.08
|
0.99
NEI'S UNBIASED I
RODGERS D
0.06 0.04
__L_
0.02
_JL_
Castle Peakivallda
Carson Pass intermediate
Sonora Pass Stanislaus
I II I I
0.92 0.94 0.96 0-98 1.0
RODGERS I
Fig. 2. Genetic distance (Nei 1978, Rodgers 1972) phenograms clustered
using UPGMA (Sneath and Sokal 1973). The distances shown are
well within the ranges exhibited by subspecies in most butterflies
studied. Their lack of concordance indicates the close relationship
between these populations.
populations belong to separate species. We found intermediate wing
color phenotypes, no diagnostic enzyme loci, and no consistent clinal
patterns from locus to locus. At the same time, it must be stressed that
analyses of electrophoretic data are logically designed to detect inter-
ruption of gene flow. Failure to detect such interruption is not equi-
valent to demonstrating that gene flow is occurring, because very
similar allelic distributions could conceivably be generated by parallel
stabilizing selection even in the absence of gene exchange. However,
alternative scenarios which do not invoke gene flow become less prob-
able as more loci are examined and found with high similarities: each
enzyme locus is likely to respond differently to selection pressures
(Carter and Watt 1988), but all loci respond similarly to gene flow.
By assuming that selection on enzyme alleles is weak when averaged
over loci, and with the knowledge that genetic drift inescapably differ-
entiates populations, we can use population genetic theory to generate
a simplest-case scenario to explain the observed genetic similarity in
terms of gene flow. This scenario represents a most-parsimonious
hypothesis which can potentially be falsified with data from additional
28(4):263-276, 1989(91)
271
enzyme loci, with knowledge of the responses to selection by the alleles
at sampled loci, or with evidence of geographic variation in previously
undetected alleles. If the gene flow estimate we generate is high, then
the populations are conspecific under the most parsimonious inter-
pretation of the data.
Under an infinite island model with no selection, the FST scores we
report are explainable by genetic drift counteracted by an average
gene exchange rate of approximately 2.8 individuals migrating be-
tween these populations each generation (using Wright’s [1931] for-
mulation Nm « (1/Fst - l)/4, where Nm is the rate of gene exchange
among populations). This gene flow rate is sufficiently strong to unite
the gene pools of these populations (Wright 1931). However, the in-
finite island model of population structure seems unrealistic for Oeneis.
We have observed “hilltopping” behavior (Shields 1967) in both ivallda
and Stanislaus : males aggregate on mountain peaks and ridges in
search of receptive females. This mating system and the relative con-
tinuity of the alpine habitat (Fig. 1) imply that these butterflies are
distributed among geographically adjacent or semi-connected popul-
ations, and are better described using an isolation-by-distance model of
population structure. When sampled populations are separated by
intervening populations in an isolation-by-distance model (as here),
then the gene flow estimated from FST represents the average rate of
genes diffusing between the sampled populations — the rate of indivi-
dual animals actually exchanged is a function of the distance between
samples, and can be much higher between adjacent local populations
(Slatkin and Barton, 1989). Thus, under a model of genetic population
structure which seems realistic for these butterflies, gene flow between
these taxa is likely to be only weakly interrupted at best.
Although Oeneis seem ideal for mark-recapture experiments, no
estimates of individual dispersal are presently available to compare
against our estimates from genetic data. Garth and Tilden (1963, p. 77)
document the ability of ivallda to colonize an unusual habitat 300 m
lower in elevation than others. AMS visits Donner Pass (2100 m), 2 km
from Castle-Basin Peak, regularly during Oeneis flight season and has
observed two individual ivallda there in 18 yr (1 male, 1 female), a
high enough incidence to suggest fairly frequent dispersal beyond the
alpine zone. The distribution of Oeneis in W Nevada (discussed below)
also suggests the ability to move among habitat patches.
Taxonomists often assume that characters diagnostic of parapatric or
allopatric taxa are indicative of more fundamental genetic differen-
tiation. If Hovanitz’s (1940) scenario is correct, the two Oeneis color
morphs are maintained by selection for crypsis on different substrates.
Such a selection differential will erect a partial barrier to the flow of
genes between them (Barton 1979, 1983). The barrier and its resulting
cline arise because neutral genes are linked on the chromosomes to
the color genes experiencing selection, and can only cross the barrier
272
J. Res. Lepid.
after recombination links them to the favored color alleles. The pre-
sence of this barrier might seem to indicate that the genetic similarity
shown between these taxa is an artifact of history rather than a result
of contemporary population processes. However, Barton and Bengtsson
(1986; see also Barton 1986) have shown that such a barrier will only
slow neutral genes, but cannot stop them for long unless (a) there is
very strong selection against intermediate genotypes and (b) there are
so many genes involved in the characters under selection that recom-
bination will not provide an escape from linked deleterious alleles.
Thus, current theory supports the notion that the diagnostic character
(color) distinguishing ivallda from Stanislaus should be treated as
genetically independent of other potential taxonomic characters — an
unreliable indicator in itself of species status or gene flow. A similar
lack of congruence between wing characters and electrophoretic data
has been found by Porter and Geiger (1988) and Porter and Mattoon
(1989) in the Satyrine genus Coenonympha. There is no biological in-
consistency between the distribution of wing color morphs and our
electrophoretic data.
The taxonomy most consistent with the available data recognizes
ivallda and Stanislaus as members of the same biological species. The
genetic relationship between these taxa and the polytypic Oeneis
chryxus remains unclear, but we recommend that ivallda and Stanislaus
remain classified as subspecies of Oe. chryxus pending further study.
In the meantime, the evolutionary interpretation of the ivallda/ Stanislaus
distribution problem is best addressed from a population biology, rather
than community ecology, perspective — that is, the evolutionary ecology
and biogeography of individual ivallda and Stanislaus traits should be
considered separately.
How convincing is the crypsis scenario ? — In the S, which Hovanitz
knew best, the geography of substrate color matches Oeneis color
morph distributions quite well. Slemmons (1966, p. 206) maps the
central Sierran andesites; the S limit of the andesite belt is in fact at
Tioga Pass, and to the S the alpine is nearly pure granite with some
darker volcanic rock in the vicinity of Mammoth Mt. The zone of rapid
transition from Stanislaus to ivallda morphs in the S corresponds well
to this conspicuous feature of Sierran geology, although detailed map-
ping of color frequencies remains to be done.
In the N, the alpine zone is more fragmented, and probably not all
Oeneis populations are known. Most of the northern alpine is, however,
on andesite — not granite. There is no corresponding geological feature
to account for the reappearance of the ivallda morph and the gradual
N ward disappearance of the dark Stanislaus phenotype. At Carson
Pass some 75% of the Oeneis habitat is on andesitic mudflows (lahars)
of the same character and color as those illustrated by Hovanitz, but
the frequency of the ivallda morph is high (95%). The rarity of dark
and intermediate morphs there strongly suggests that something other
than background matching is limiting their N-ward spread: a selection
28(4):2S3-278s 1989(91)
273
regime strong enough to produce the sharp cline at Tioga Pass should
also favor the Stanislaus morph at Carson, (It remains possible that the
hypothetical predator drops out or switches prey just S of Carson Pass.)
There is granitic alpine in the Crystal Range WSW of Lake Tahoe —
although it is unlikely that the high frequency of the ivallda color
morph at Carson Pass could be maintained by massive gene flow from
there, this may be the historical source of the three N- most ivallda
populations (Mt. Lola, Castle-Basin Peaks, Anderson Peak). However,
given the inconsistencies in the crypsis scenario for ivallda populations
in the N (Hovanitz [1940] discussed the tenuousness of crypsis and
mimicry hypotheses when experimental data were unavailable), we
recommend that it be treated cautiously pending experimental con-
firmation with a known predator.
Plausibility of an Easterly Invasion by the Stanislaus Color Genes —
Austin and Murphy (1987) recorded the ivallda morph in Nevada only
in the Carson Range (Carson City and Washoe Cos.), just a few km E of
the Sierra Nevada, and nominate Rocky Mountain univoltine chryxus
in extreme E Nevada (Elko, Lincoln, White Pine Cos.). Since then
Stanislaus (indistinguishable from Sierran) has been found in the
Sweetwater Mts. in Lyon Co. on the California border, again just a few
km E of that morplTs range in the Sierra Nevada (G. T. Austin, pers .
comm.). These occurrences seem to be due to W-to-E dispersal from the
Sierra, and are uninformative about colonization routes into the Sierra.
An absence of relictual Oeneis in central Nevada is mirrored in other
alpine butterflies, and in other alpine organisms generally (Billings
1978, Harper and Reveal 1978).
The genetic analyses provide no additional characters associated with
either ivallda or Stanislaus morphs for comparison to other chryxus
populations to the N and E, leaving wing color as the only reliable
character. All other chryxus , and most other Oeneis , are colored like
Stanislaus , or even darker (Ferris and Brown 1980). In the absence of a
phytogeny for Oeneis , we assume that the original invaders of the
Sierra were this color and that the ivallda color is a uniquely derived
autapomorphy. But if the Stanislaus morph came from the N, why are
the relict plesiomorphous color genes in the central Sierra and not also
in the N — especially if andesitic substrates are relevant? As Hovanitz
noted, the distribution would make more sense if the invasion of the
Stanislaus color genes had come directly across the Great Basin. Oeneis
chryxus apparently does not occur in Oregon (Dornfeld 1980) or in
the Klamath Mts. of N California (Shapiro et al. 1981), reaching its
southern limit (W of the Rockies) in Washington. Its range as mapped
by Scott (1986, p. 248) raises a very serious question of where an in-
vasion from the N might have come from; no relicts of the hypothetical
N route have been found.
At least one other butterfly taxon is distributed along the hypotheti-
cal NE invasion route. Limenitis lorquini weidemeyerii (Edwards), a
Rocky Mountain middle-elevation nymph alid butterfly, is restricted to
274
J. Res. Lepid.
montane riparian canyon habitats in the Great Basin (Austin and
Murphy 1987; Porter 1989). Its distribution closely follows the colon-
ization route in the Humboldt drainage proposed by Major and Bamberg
(1967): it crosses the low-elevation desert into the Wassuk Mts. and
reaches its W distribution limits on the N shore of Mono Lake (see
Fig. 1), where it hybridizes with the Si err an L. lorquini lorquini
(Boisduval) (Porter 1989). “Pure” L. 1. weidemeyerii and hybrid wing
pattern morphs are sympatric with Oe c. Stanislaus near Sonora Pass
where montane and subalpine habitats interdigitate. The distribution
of the weidemeyerii morph is entirely consistent with an invasion of
Stanislaus color genes from the E into the central Sierra. The presence
of L. lorquini burrisoni Maynard (a weakly defined taxon quite similar
in phenotype to nominate lorquini ) in Oregon, Washington and into
western Montana makes a northerly route of Sierran invasion by the
weidemeyerii form seem highly unlikely. Although Limenitis is found
in montane habitats and Oeneis rarely strays from the alpine, similar
immigration corridors may have been used at different times by both.
The simplest scenario consistent with the Oeneis distribution data
has chryxus invading first (from the N or E) and evolving the distinc-
tive ivallda color, with a second chryxus invasion from the E injecting
the distinctive Stanislaus color into the central Sierran populations.
However, this is by no means the only available scenario, and it is even
possible that the butterscotch brown of the Stanislaus morph represents
a character reversal, and not evidence of two Sierran invasions. Genetic
studies of the entire Oeneis chryxus complex in western North Ameri-
ca, with an eye towards alleles linking Sierran and potential source
populations, may expose characters which will help resolve the bio-
geographical problem. The answer will be of considerable value in the
interpretation of the origins of the entire Sierran alpine biotic
community.
Acknowledgements. This study would not have been possible without the
help of Jim Mori, who collected two samples for us. We thank Brad Shaffer for
the loan of his facilities and G. L. Stebbins, J. Major, and D. Elliott Fisk for
discussions of Sierran historical biogeography, and George Austin for com-
municating unpublished Nevada data. Beth Jakob suggested several im-
provements in the manuscript. This research was supported by California
Agricultural Experiment Station project CA-D*-AZO-3394-H, Climatic Range
Limitation of Phytophagous Lepidopterans (AMS, Principal Investigator).
Literature Cited
AUSTIN, G. T., AND D. D. MURPHY., 1987. Zoogeography of Great Basin butterflies:
patterns of distribution and differentiation. Great Basin Nat 47:186-201.
AXELROD, D. L, 1957. Late Tertiary floras of the Sierra Nevada uplift (California-
Nevada). Geol. Soc. Amer. Bull. 68:19-45.
28(4):263-276, 1989(91)
275
AXELROD, D. I., 1976. History of the coniferous forests, California and Nevada.
Univ. Calif. Publ. Bot. 70:1-62.
AXELROD, D. I., 1977. Outline history of California vegetation, pp. 139-220. In
M. Barbour and J. Major, eds., Terrestrial Vegetation of California. Wiley -
Interscience, New York.
AYALA, F. J., J. R. POWELL, M. L. TRACEY, C. A. MOURAO, AND S. PEREZ-SALAS., 1972.
Enzyme variability in the Drosophila willistoni group. IV. Genetic vari-
ation in natural populations of Drosophila willistoni. Genetics 70:113-139.
BARTON, N. H., 1979. Gene flow past a cline. Heredity 43:333-339.
BARTON, N. H., 1983. Multilocus dines. Evolution 37:454-471.
BARTON, N. H., 1986. The effects of linkage and density-dependent regulation on
gene flow. Heredity 57:415-426.
BARTON, N. H., AND B. O. BENGTSSON., 1986. The barrier to genetic exchange be-
tween hybridizing populations. Heredity 56:357-376.
BATEMAN, P. C., AND C. WAHRHAFTIG., 1966. Geology of the Sierra Nevada, pp.
105-172. In E. H. Bailey (ed.), Geology of Northern California. Calif. Div.
Mines and Geology, Bull. 190.
BILLINGS, D. W., 1978. Alpine phytogeography across the Great Basin. Great
Basin Nat. Mem. 2:105-117.
CARTER, P. A., AND W. B. WATT., 1988. Adaptation at specific loci. V. Metabolically
adjacent enzyme loci may have very different experiences of selective
pressures. Genetics 119:913-924.
CHABOT, B. F„ AND W. D. BILLINGS., 1972. Origins and ecology of the Sierran
alpine flora and vegetation. Ecol. Monog. 42:163-199.
CURRY, R. T., 1966. Glaciation about 3,000,000 years ago in the Sierra Nevada.
Science 154:770-771.
DORNFELD, E. J., 1980. The Butterflies of Oregon. Timber Press, Forest Grove,
Oregon. 276 pp.
EMMEL, T. C., 1975. Satyridae, pp. 79-111. In W. J. Howe (ed.), The Butterflies of
North America. Doubleday, New York. 633 pp.
FERRIS, C. D., AND F. M. BROWN., 1981. Butterflies of the Rocky Mountain States.
University of Oklahoma Press, Norman. 442 pp.
FULLERTON, D. S., 1986. Chronology and correlation of glacial deposits in the
Sierra Nevada, California. Quart. Sci. Rev. 5:161-170.
GARTH, J. S„ AND J. W. TILDEN., 1963. Yosemite butterflies. J. Res. Lepid. 2:1-96.
GARTH, J. S., AND J. W. TILDEN., 1986. California Butterflies. University of Cali-
fornia Press, Berkeley. 246 pp.
GEIGER, H. J., AND A. M. SHAPIRO., 1986. Electrophoretic evidence for speciation
within the nominal species Anthocharis sara Lucas (Pieridae). J. Res.
Lepid. 25:15-24.
HARPER, K. T., AND J. L. REVEAL., 1978. Intermountain biogeography: a sym-
posium. Great Basin Nat. Mem. 2:1-268.
HARRIS, H., AND D. A. HOPKINSON., 1976. Handbook of Enzyme Electrophoresis in
Human Genetics. North Holland, Amsterdam.
HARSHBERGER, J. W., 1911. Phytogeographic Survey of North America. Verlag W.
Engelmann, Leipzig. 790 pp.
HEDRICK, P. W., 1985. Genetics of Populations. Jones & Bartlett, Boston. 629 pp.
HOVANITZ, W., 1937. A new race of Oeneis chryxus (Dbld. & Hew.) (Lepidoptera:
Satyridae). Ent. News 48:228.
HOVANITZ, W„ 1940. Ecological color variation in a butterfly and the problem of
“protective coloration.” Ecology 21:371-380.
270
J. Res . Lepid.
KITCHING, I. J., 1985. Allozyme variation in the milkweed butterflies (Lepidop-
tera: Danainae). Zool. J. Linn. Soc. 86:367-389.
MAJOR, J., AND s. A. BAMBERG., 1967. Some cordilleran plants disjunct in the
Sierra Nevada of California, and their bearing on Pleistocene ecological
conditions, pp. 171-188. In H. E. Wright, Jr., and W. H. Osburn (eds.),
Arctic and Alpine Environments. Indiana Univ. Press, Bloomington.
MAJOR, J., AND D. w. TAYLOR., 1977. Alpine, pp. 601-675. In M. G Barbour and J.
Major (eds.), Terrestrial Vegetation of California. Wiley and Sons, New
York.
MILLER, L. D„ AND F. M. BROWN., 1981. A catalogue/checklist of the butterflies of
America north of Mexico. Lepid. Soc. Mem. 2:1-280.
NEI, M\, 1978. Estimation of average heterozygosity and genetic distance from a
small number of individuals. Genetics 89:583-590.
PROTER, A. H., 1989. Gene Flow Statistics and Species-Level Systematics in
Butterflies. Ph. D. Thesis. University of California, Davis.
PROTER, A. H., AND H. J. GEIGER., 1988. Genetic and phenotypic population struc-
ture of the Coenonympha tullia complex (Lepidoptera: Nymphalidae:
Satyrinae) in California: no evidence for species boundaries. Can. J. Zool.
66:2751-2765.
PORTER, A. H., AND S. O. MATTOON., 1989. A new subspecies of Coenonympha tullia
(Muller) (Nymphalidae: Satyrinae) confined to the coastal dunes of
northern California. J. Lepid. Soc. 43:229-238.
REMSEN, J. V., JR., 1984. High incidence of “leapfrog” pattern of geographic vari-
ation in Andean birds: implications for the speciatioe process. Science
224:171-173.
RODGERS, J. s., 1984. Measures of genetic similarity and distance, pp. 145-153.
In Studies in Genetics VII. Univ. Texas Pub. No. 7213.
SCOTT, J. A., 1986. The Butterflies of North America; A Natural History and
Field Guide. Standford University Press. Stanford, CA. 436 pp.
SHAPIRO, A. M., C. A. PALM, AND K. L. WCISLO., 1979(81). The ecology and biogeo-
graphy of the butterflies of the Trinity Alps and Mount Eddy, northern
California. J. Res, Lep. 18:69-152.
SHARSMITH, C. W., 1940. A contribution to the history of the alpine flora of the
Sierra Nevada. Ph. D. Thesis. Univ. of California, Berkeley.
SLATKIN, M. AND N. H. BARTON., 1989. A comparison of three indirect methods for
estimating average levels of gene flow. Evolution 43:1349-1369.
SNEATH, P H. A., & R R. SOKAL., 1973. Numerical taxonomy. W. B. Freeman, San
Francisco. 573 pp.
STEBBINS, G. L., AND J. MAJOR., 1965. Endemism and speciation in the California
flora. Ecol. Monogr. 35:1-35.
SWOFFORD, D. L., AND R. B. SELAMDER., 1931. A computer program for the analysis
of allelic variation in genetics. J. Hered. 72:281-283.
THORPE, J. P., 1933. Enzyme variation, genetic distance and evolutionary
divergence in relation to levels of taxonomic separation, pp. 131-153. In:
G. S. Oxford and D. Robinson (eds.), Protein Polymorphisms: Adaptive and
Taxonomic Significance. Academic Press. New York.
TILDEN, J. W., AND A. SMITH., 1986. Field Guide to Western Butterflies. Houghton
Mifflin, Boston. 370 pp.
WELLS, P. V., 1983. Paleobiogeography of montane islands in the Great Basin
since the last glaciopluvial. Ecol. Mon. 53:341-382.
WRIGHT, s., 1931. Evolution in Mendelian populations. Genetics 16:97-159.
Journal ofReasearch on the Lepidoptera
28(4):277-282, 1989(91)
A New Polythrix From Central America
(Lepidoptera: Hesperiidae)
John A. Shuey*
Battelle Great Lakes Environmental Center, 739 Hastings, Traverse City, MI 49684
Abstract. Polythrix kanshul is described as a new species. It differs
from its nearest relative, P. metallescens in several details of wing
pattern and in many genitalic characters. These two species, along
with P. eudoxus , form a monophyletic lineage defined by the morphol-
ogy of the uncus. Polythrix kanshul in known from Palenque, Chiapas,
Mexico and Bayano, Panama.
The genus Polythrix is distributed from the southern border of the
United States south to Argentina with the bulk of the 15 recognized
species found in Central America and northern South America (Evans
1952). Freeman ( 1979) recorded six species in Mexico and provided a key
to their identification while Llorente, et al, (1990) listed seven species.
While identifying material from southern Mexico, I realized that a new
species of Polythrix was present in the sample. My purpose here is to
describe it and to document its relationship with other species in the
genus.
Polythrix kanshul , Shuey New species
Diagnosis of male: The wing pattern (Figures 1-4) ofP. kanshul is very
similar to P. metallescens (Mabille) (Figures 5-8) with the following
exceptions: the ground color of P. kanshul is darker brown and the dorsal
metallic blue-green body and hindwing scaling is brighter than in P.
metallescens ; P. kanshul has four apical spots, P. metallescens has three;
and ventrally, the narrow white discal band on the hindwing is shorter
in P. kanshul , extending only between veins 2 A and Cu2 while in P.
metallescens this band extends between veins 2A and Sc+Rr A cblor
photograph of P. metallescens can be found in Lewis (1973).
Description of male: Figures 1-4. Forewing: ground color brown;
fringes brown; metallic green hairs cover the basal one-third dorsally —
one-sixth ventrally; four apical spots present; three hyaline spots - one
located in the distal portion of the discal cell and one each in the mid-
points of cells M3 and Cu2; costal fold present; ventral hair- tuft covers the
origin of vein Cu2; ventral cells Cu2 and 2 A gray. Hindwing: ground color
brown; fringes white; vein 2 A extended, forming a blunt tail; metallic
green hairs and scales cover the inner two-thirds dorsally — one-third
ventrally extending downward at the tails; ventral surfaces with a
narrow white discal band between veins 2A and Cu2. Head, palpi, and
*Research Associate, Carnegie Museum of Natural History, Pittsburgh PA
278
J. Res. Lepid.
Figures 1 -4. Polythrix kanshui; 1 , dorsal view, holotype male, Palenque, Mexico; 2, ventral view of previous specimen; 3, paratype
male, Bayano, Panama; 4, ventral view of previous specimen.
Figures 5-8. Polythrix metailescens; 5, dorsal view; male, Madden Forest, Panama, 3 Aug. 1969; 6, ventral view of previous
specimen; 7, female, Cayuga, Guatemala, May; 8, ventral view of previous specimen.
28(4):277-282, 1989(91)
279
thorax ground color brown but densely covered with metallic green hairs
and scales.
Male (Figure 9) valvae elongated into a curved, heavily toothed, blunt
projection; upper edge of sacculus (sensu Klots, 1970) heavily sclerotized
and wrinkled. Uncus (Figures 9-10) fused into a single posterior projec-
tion with lateral socii originating from tegumen on both sides. Aedeagus
(Figure 11) with anterior extension beyond membranous ejaculatory
duct; posterior cornuti plate-like and hinged ventrally.
Female: Unknown. If the pattern of sexual dimorphism is similar to
that ofP. metallescens (Figures 7-8), the female ofP. kanshul should be
similar to the male, but with longer tails and duller green iridescence.
Types: Holotype - Mexico, Chiapas, Ruinas Palenque, approx 17°30' X
92°05', 21-VIII-1987, J.A. Shuey, collector (Carnegie Museum of Natural
History). Paratypes - one specimen with the same locality data as the
holotype, collected 20-VIIX-1987 (J. A. Shuey collection): One specimen,
Panama, Panama, Bayano, 12-X-1974, G.B. Small, collector (United
States National Museum of Natural History).
Etymology: The specific name reflects the long and splendid history of
the type locality, Palenque, and is a latinization of Kan-Xul (kan-shool).
Kan-Xul was the second son of Pacal to assume the rulership of Palenque,
and along with his father and brother, was responsible for much of the
magnificent architecture of this site. Kan-Xul ruled Palenque at its
zenith, but was captured in warfare with neighboring Tonina, and
presumably sacrificed there (Scheie and Miller, 1986). An accession
portrait of Kan-Xul, in stucco relief, still survives within the palace at
Palenque. Kan xul is Mayan for “magnificent animal”. My name for this
insect is meant as a double tribute; first to the Maya, past and present,
whose world view and beliefs continue to shape much of Central America;
second, to the insect, which is truly ‘kan xul’.
Habitat and Distribution: The habitat at Palenque is mature to young
“selva alta perennifolia” (perennial high rainforest) (Miranda and Gyves,
1979). The entire forest in the vicinity of the ruins has presumably
regrown since approximately 1000 BP. The classic Maya developed the
entire area around the core of the ruins, and little or no forest probably
occurred in the immediate vicinity at the time of active occupation
(Andrews, 1975). Portions of the present day forest represent the original
old-growth forest that blanketed the site after Palenque was abandoned,
but much of the surrounding forest represents more recent regrowth that
followed the clearing of the ruins during the late 1800’s to the present.
The holotype was captured along a well worn trail through a part of the
rainforest that may represent part of the older growth. It was perched
on the underside of a leaf when captured. The other Palenque specimen
was collected at a nearby motel which is surrounded by young rainforest
regrowth. This specimen was collected at night on a white sheet
illuminated by ultraviolet light. It was probably dislodged from its
280
J. Res. Lepid.
Figures 9-11. Polythrix kanshul, holotype male genitalia; 9, lateral view; 10, dorsal
view of uncus and tegumen; 1 1 , lateral view of aedeagus.
Figures 12 - 14. Polythrix metallescens, male genitalia; 12, lateral view; 13, dorsal
view of uncus and tegumen; 14, lateral view of aedeagus.
Figures 15-16. Polythrix genitalia, uncus and tegumen, dorsal view; 1 5, P. caunus,
R. Yanacani, E. Bolivia, alt. 600m., March 1915; 16, P. auginus, Cayuga,
Guatemala, August.
28(4):277-282, 1989(91)
281
nocturnal perch during an intense evening rainstorm, and subsequently
attracted to the light.
The perennial high rainforest habitat is widespread throughout the
lowlands of Central America, and P. kanshul is probably found in all of
the intervening countries between Chiapas and Panama. Perennial high
rainforest in Mexico extends northward into Tabasco and in fragmented
form into Veracruz, and of course, southward throughout much of South
America; P. kanshul should occupy a more extensive area than is
presently known. Like all rainforest life, this species is certainly
extirpated from the portions of its original range which have been
converted for agricultural uses.
Discussion: Polythrix kanshul and P. metallescens are each other’s
closest known relatives. They differ from all other Polythrix species in
the distinctive configuration of the hyaline forewing spots and in the
relatively large amount of metallic over-scaling on the wings and body.
In other species of Polythrix , the fore wing spots are fairly broad and
overlap. These overlapping spots, in conjunction with the prominent
apical spots and hindwing tails, give the genus Polythrix its distinctive
appearance. InP. kanshul andP. metallescens these spots are taller than
they are wide, and generally do not overlap.
Despite their similarities, I would argue that the divergence of P.
kanshul andP. metallescens from a common ancestor is not recent. While
the genitalia of these two species differ most conspicuously in the
configuration of the valvae, the details of almost every other structure
differ also (Figures 9 - 14). The accumulation of so many structural
differences indicates that these taxa have followed different evolutionary
paths for a long time. The broad geographic overlap of these two taxa also
supports this contention. Polythrix metallescens occurs from Belize
south through Central America and into at least amazonian Brazil.
Polythrix kanshul is known from Panama and Chiapas Mexico, indicat-
ing a broad overlap in the known range of these two species. Recent
differentiates are generally allopatric.
The fused uncus is apparently an apomorphy which defines a lineage
composed of three species, P. metallescens , P. kanshul , and P. eudoxus
(Stoll). My inclusion ofP. eudoxus in this lineage is somewhat tentative,
but Evans’ (1952) caricature of the genitalia of this species suggests that
it too has the fused uncus and lateral socii. All other species of Polythrix
have a more typical Pyrginae uncus composed of two lateral prongs.
However, some of these species may form a transition series to this
apomorphic character state; P. caunus (Herrich-Schaffer) andP. auginus
(Hewitson) for example, have the basal portion of the uncus elongated,
with the two prongs reduced to small hooks on the distal end (Figures 15
and 16), and the beginnings of enlarged lateral socii. The homology of this
transitional state is tentative, but its configuration is certainly sugges-
tive.
The addition of P. kanshul to Mexico’s fauna raises the number of
282
J. Res. Lepid.
Polythrix species known from that country to eight. It seems likely that
P. metallescens also will be found to occur in southern Mexico. This
skipper is known from Belize and Guatemala, and may eventually be
found in the dense rainforests of the Lacadon Forest and Montes Azulies
Biosphere Reserve of eastern Chiapas.
Acknowledgements. Foremost, I thank Judith A. Cox-Shuey for accompanying
me in Central America on so many occasions, and for her tolerance of my
entomological preoccupations. Dr. John W. Peacock was also present on the trip
which produced the specimens described here, and it was at his UV light that the
first Mexican specimen of P. kanshul was taken; he and Dr. John Rawlins
reviewed an early draft of this manuscript. Drs. John Burns, National Museum
of Natural History, and John Rawlins, Carnegie Museum of Natural History,
kindly lent comparative material used in this study.
Literature Cited
Andrews, G. F. 1975. Maya cities: Placemaking and urbanization. Univ. of
Oklahoma Press, Norman.
Evans, W. H. 1952. A catalogue of the American Hesperiidae indicating the
classification and nomenclature adopted in the British Museum (Natural
History): Part II, Pyrginae, Section 1. British Museum, London. 178 p.
Freeman, H. A. 1979. Review of Mexican Polythrix Watson 1893 (Hesperiidae). J.
Lepid. Soc. 33: 124-128.
Klots, A. B. 1970. Lepidoptera. in S. L. Tuxen ed., Taxonomists glossary of
genitalia in insects. Munksgaard, Copenhagen. 359 p.
Lewis, H. L. 1973. Butterflies of the World. Follett Publ. Co., Chicago. 312 p.
Llorente-Bousquets, J. , A. Luis-Martinez and I. Vargas-FernAndez. 1990. Catalogo
sistematico de los Hesperioidea de Mexico. Publicaciones Especiales del
Museo de Zoologfa No. 1. Univ. Nacional Autonoma de Mexico. Mexico, D.F.
70 p.
Miranda, E. G. and Z. F. Gyves. 1979. Nueva atlas porrua de la Republica
Mexicana. Editorial Porrua, S. A. Mexico D. F. 197 p.
Schele, L. and M. E. Miller. 1986. The blood of kings: Dynasty and ritual in Maya
art. Kimbell Art Museum, Fort Worth, Texas. 335 p.
Journal of Research on the Lepidoptera
28(4):283-288, 1989(91)
Three unusual species of Parades from South
America (Lepidoptera: Arctiidae)
Vitor 0. Becker
Centro de Pesquisa Agropeeuaria dos Gerrados, Caixa postal 700023, 73300-Planaltina,
DF, Brasil
and
Scott E. Miller
Bishop Museum, Box 19000-A, Honolulu, Hawaii 96817, USA.
Abstract. Two new species of Parades Walker are described from the
northern Andes, which differ from congeners by their small size and
uniform brown coloration: Parades minuta n. sp. (Colombia) and P.
diminuta n. sp. (Venezuela). Thagona medinata (Dognin) is recognized
as a Parades, and transferred from Lymantriidae; it differs from other
Parades in its immaculate white coloration.
Introduction
In the course of research on zygaenoid moths, we encountered two
undescribed species of the arctiid genus Parades Walker which superfi-
cially resemble megalopygids, especially Podalia bolivari (Heylaerts)
(Miller and Becker, in press). These two species are very similar to the
few uniform brown species of Parades , especially P. obscurior (Schaus)
(see Watson, 1973: 33, pis. 3 Id, 89a, b), but differ from these in their much
smaller size (fore wing lengths of about 9 mm versus 18 mm). Both the
new species are known only from males. It is possible that the females
are brachypterous, as are some others in the genus, e.g., Parades
deserticola (Berg, 1875: 212) andP. imitatrix (Rothschild, 1922: 493).
We also take the opportunity to transfer Thagona medinata (Dognin)
from the Lymantriidae to its proper place among Parades , and give
illustrations to permit its identification.
The proper generic name of this group has been confused in the past.
We follow Watson (1980) and Watson and Goodger (1988: 32) in using
Parades . Before Watson and Goodger (1986), most of the species were
placed in Palustra , Antarctia , or Maenas .
Holotypes are deposited in the National Museum of Natural History
(USNM). Other collection acronyms follow Heppner and Lamas (1982).
284
Figs. 1-4. Parades male genitalia, ventral view, aedeagus removed (paratypes);
Figs. 1-2: P. diminuta ; Figs. 3-4: P. minuta .
28(4):283-288, 1989(91)
285
Figs. 5-7. Parades male left wings;
Fig. 5: P. minuta (holotype);
Fig. 6: P. diminuta {holo-
type); Fig. 7 P. medinata
(lectotype).
Figs. 8-9. Parades medinata , male genitalia, ventral view, aedeagus removed
(paralectotype).
286
J. Res. Lepid.
Taxonomy
Parades minuta Becker & Miller, new species
Figs. 3-5
DIAGNOSIS. -Very similar to Parades obscurior , but much smaller
and with darker ground color. Similar toP. diminuta, but antennae with
longer ciliation and forewings lacking maculation at end of discal cell
(Fig. 5); base of valva lacking setose costal lobe (Fig. 3).
ADULT MALE (Fig. 5). -Forewing length 8.5 mm.
Head densely hairy, dark brown. Antennae strongly bipectinate,
pectinations three flagellum diameters long. Thorax and abdomen
densely hairy, dark brown dorsally, pale brown ventrally. Legs pale
brown, tarsi with light and dark brown banding. Forewings dark brown,
costal margin slightly concave, thinly scaled with elongate scales yield-
ing translucent appearance. Hindwings similar, slightly lighter. Ven-
tral wings lighter, except costal margins which are dark brown with some
patches of lighter scales.
MALE GENITALIA (Figs. 3, 4). -Uncus tapered, slightly expanded at
middle, apex rounded; tegumen long, bent ventrad; valvae short, simple,
covered with short setae distally; juxta an inverted, broad trapezium; !
saccus triangular, slightly rounded anteriorly. Aedeagus short, bent
ventrad at middle; vesica smooth (one of two preparations has a very
long, thin cornutus; presumably lost in preparation of the second speci-
men).
ADULT FE MALE . -U nkno wn .
TYPE LOCALITY. -Colombia, Cundinamarca, Bogota, “Pueblo Guasca”.
IMMATURE STAGE S . -U nkno wn .
FLIGHT PERIOD. -Unknown.
DISTRIBUTION. -Known only from the vicinity of Bogota, Colombia.
MATERIAL EXAMINED. -Holotype (USNM) and 33 male paratypes:
COLOMBIA: Cundinamarca : Bogota, 2800-3200 m [no date], A.H. Fassl
(USNM); “Pueblo Guasca, Bogota”, [no date], “F. Johnson/donor” (BMNH,
BPBM, CMNH, LACM, USNM, VOB, ZSBS).
Parades diminuta Becker & Miller, new species
Figs. 1, 2, 6
DIAGNOSIS. -Similar to Parades minuta , but fore wings more rounded
and with pale mark across end of discal cell (Fig. 6), and antennae with
shorter ciliation; base of valva with setose costal lobe (Fig. 1).
ADULT MALE (Fig. 6). -Forewing length 9 mm.
Entirely brown, except vertical tan line at end of discal cell. Hindwing
slightly lighter than forewing. Antennae narrow, without pectination;
ciliation as long as flagellum diameter.
MALE GENITALIA (Figs. 1, 2). -Uncus tapered, slightly constricted at
28(4):283-288, 1989(91)
287
middle; apex pointed; tegumen rounded; valvae simple, narrow, base of
costa expanded into a short, irregular lobe covered with short setae; juxta
weak, nearly rectangular, slightly constricted laterally; saccus broadly
rounded. Aedeagus nearly straight; vesica expanded, with a broad
scobinate area; a small area with short, triangular spines at edge of
scobinate area.
ADULT FE MALE . -U nknown .
TYPE LOCALITY.-Venezuela, Merida, Mucuy Fish Hatchery, 7 km E
of Tabay, 6600 feet [2000 m].
IMMATURE STAGES. -Unknown.
FLIGHT PERIOD. -February.
DISTRIBUTION. -Known only from the type locality.
MATERIAL EXAMINED. -10 males from the type locality (holotype
[USNM] and 9 paratypes), all collected 10-13-11-1978 by J.B. Heppner at
blacklight (BMNH, LACM, UCV, USNM, VOB).
Parades medinata (Dognin), new combination
Figs. 7-9
?Trochuda medinata Dognin, 1920: 4.
Thagona medinata : Schaus, 1927: 549, pi. 74c.
This species was described from an unspecified number of males and
one female from “Medina, est de la Colombie, 500 metres (Fassl)”. Three
males and one female from the Dognin Collection are present in the
USNM; we hereby designate the male which bears Dognin’s “type” label,
as well as USNM type number 29743, as lectotype. We hereby transfer
this species to Arctiidae. The characters of wing venation, antennae, and
male genitalia are typical of Parades , and very similar to others in the
genus (as illustrated by Watson, 1971, 1973).
A series of specimens has been collected at Planaltina, DF, Brazil, by
the first author.
Acknowledgements. Most of this work was carried out at the Smithsonian
Institution. The photographs were taken by Victor Krantz of the Smithsonian
Institution. D.C. Ferguson, Systematic Entomology Laboratory, U.S. Depart-
ment of Agriculture, assisted in placing the new species. J.P. Donahue, D.C.
Ferguson, and J.E. Rawlins reviewed the manuscript.
Literature Cited
Berg, C. 1875. Patagonische Lepidopteren beobachtet auf einer Reise im Jahre
1874. Bulletin de la Societe Imperiale des Naturalistes de Moscou 49: 191-
245.
j Dognin, P. 1920. Heteroceres nouveaux de 1’Amerique du Sud. Fascicule XVIII.
Imprimerie Oberthtir, Rennes. 13 pp.
! Heppner, J.B. and G. Lamas. 1982. Acronyms for world museum collections of
insects, with an emphasis on Neotropical Lepidoptera. Bulletin of the
Entomological Society of America, 28: 305-315.
288
J. Res. Lepid.
Miller, S.E. and V.O. Becker. 1991. Podalia bolivari: a highly sexually dimorphic
neotropical megalopygid pest (Lepidoptera).
Rothschild, L.W. 1922. A preliminary list of the Arctiinae of Para, Brazil, and a
few from other localities. Annals and Magazine of Natural History (ser. 9) 9:
457-494.
Schaus, W. 1927. Lymantriidae. pp. 535-564 in A. Seitz (ed.), Macrolepidoptera
of the World, Volume 6. Alfred Kernen, Stuttgart.
Watson, A. 1971. An Illustrated Catalog of the Neotropic Arctiinae Types in the
United States National Museum (Lepidoptera: Arctiidae). Part I. Smithsonian
Contributions to Zoology, 50: iii + 361 pp.
Watson, A. 1973. An Illustrated Catalog of the Neotropic Arctiinae Types in the
United States National Museum (Lepidoptera: Arctiidae). Part II. Smithsonian
Contributions to Zoology, 128: iii + 160 pp.
Watson, A., D.S. Fletcher and I.W.B. Nye, 1980. In I.W.B. Nye (ed.), The generic
names of moths of the world, v. 2. British Museum (Natural History), London,
xiv + 228 pp.
Watson, A. and D.T. Goodger. 1986. Catalogue of the Neotropical Tiger-moths.
Occasional Papers on Systematic Entomology, British Museum (Natural
History), 1: 1-71
Journal of Research on the Lepidoptera
28(4):289-296, 1989(91)
Selection of Lepidopterologically Interesting Areas
in Central Spain Using UTM Distribution Maps
J.L. Viejo, C. de Silva, C. Ibero and J. Martin.
Dep. de Biologfa, C-XV, Universidad Autonoma de Madrid. 28049 Madrid. Spain.
Abstract. This paper deals with species richness and biogeographic
interest of the butterfly fauna ( Papilionoidea & Hesperioidea ) of the
Madrid province, using its one hundred and nine 100 sq. km UTM
squares data. Richest species squares (80-102 species) are on the north
(Sierra de Guadarrama) and the poorest ones on the centre and south.
There is a slightly rich area on the southeast. Fauna’s biogeographic
interest (chorological index sensu Kudrna) shows a different pattern,
being maximum on the southeast squares, lightly high on the south and
centre, and low on the north. In conclusion, richest species squares are
not necessarily those of maximum average chorological index. This is
explained by the environmental similarity (climate, vegetation, etc)
between Sierra and the European generality , while plant formations
on the south (typically xerophytic) are peculiar in comparison with the
rest of the continent, which have been used as a biogeographic refer-
ence. Consequently, species richness appears as a limited criterion
when focussing the selection of areas lepidopterologically interesting.
Qualitative criteria must be also considered to establish possible zones
to protect their butterfly communities, such as the biogeographic
interest of the fauna, provided by UTM species distribution maps.
Resumen. Este trabajo estudia el numero de especies y el valor
biogeografico de la fauna de mariposas (. Papilionoidea & Hesperioidea )
presente en cada una de las 109 cuadrfculas UTM de 100 kilometres
cuadrados de la provincia de Madrid. Las cuadrfculas con mayor
numero de especies (entre 80 y 102) se situan al norte del territorio
(Sierra de Guadarrama) y las mas pobres en el centro y sur. Al sureste
hay una zona moderadamente rica. El valor biogeografico de la fauna
(fndice corologico de Kudrna) tiene un reparto bien distinto, ya que es
inaximo en las cuadrfculas del sureste, moderadamente alto en el sur
y centro, y bajo en el norte. Se inhere, por tanto, que las cuadrfculas mas
ricas en especies no son necesariamente las de mayor fndice corologico
medio, lo que atribuimos a la similitud ambiental (clima, vegetacion,
etc) de la Sierra con la mayor parte de Europa, mientras que las
formaciones vegetales del sur (encinares, coscojares y quejigares sobre
todo) son mas singulares (mas xerofilas) con respecto al continente,
ambito de referenda biogeografica utilizado. Concluimos con que el
criteria del numero de especies es de utilidad limitada en la seleccion
de areas de in teres lepidopterologico, y que son necesarios tambien
criterios cualitativos, como el valor biogeografico de la fauna, para
establecer zonas susceptibles de proteccion por su fauna de mariposas,
a partir de mapas UTM de distribucion de las especies.
290
J. Res. Lepid.
Introduction
Decreasing numbers in many butterfly and skipper populations are
awakening, among numerous naturalists, the interest for their conser-
vation.
Obviously , the bigger the information about species the more efficient
will be the measures to propose towards its conservation. Therefore, it is
necessary to deep in the knowledge of some aspects such as the precise
geographic distribution, environmental preferences, life cycles, interac-
tions with foodplants, parasites and any other biological aspect affecting
different species. But, in view of the fast butterfly and skipper commu-
nities impoverishment process, generally caused by different human
activities, it is fairly evident that we can not wait the results of the
aforementioned autoecological studies to adopt protection criteria.
Nevertheless, we believe suitable to begin applying protection mea-
sures based only on geographic range data, given that, as repeatedly has
been said, ecosystem conservation, as opposed to species approach to
butterfly protection, would seem to be the most effective policy to be
followed (Thomas & Mallorie, 1985; Munguira, 1987; Viejo, Viedma &
Martinez, 1989). And Lepidoptera atlases are very useful for those
preliminary studies.
In some European countries, such as Great Britain (Heath & Skelton,
1983) or Switzerland (Gonseth, 1987), their butterfly distribution maps
are already concluded, at a national scale and following UTM 100 km2 !
squares system. On the opposite, in Spain we are still well behind to
complete our butterfly distribution national maps, although a valuable
effort on the elaboration of regional atlases within the last ten years has j
been made, and some of them, both from the north (Gomez de Aizpurua,
1977; 1979; 1988) as well as from central Spain (Viejo, 1983; Gomez de
Aizpurua, 1987) have been already published.
Methods
The Atlas of the Lepidoptera of Madrid (Gomez de Aizpurua, 1987) provided
data for this study, which compiles 153 distribution maps of species of Zygaenoidea,
Papilionoidea & Hesperioidea in the Madrid province. We have excluded the 13
species of Zygaenoidea, and from the lasting 140 we have eliminated 4 because
of uncertain data, as well as the records prior to 1950 with no later confirmation.
A presence-absence matrix (1-0) with the faunistic data from the one hundred '
and nine 100 km2 squares of Madrid was made. From this matrix we could obtain
the species number and the Average Chorological Index (Kudrna, 1986) of each (
square, which have been used as criteria to establish the conservation interest of
the study area, given the linking relationship between butterflies and specific j
vegetation communities (Uherkovich, 1983; Viejo & Templado, 1986).
Species number is a variable frequently used in conservation studies (Margules
& Usher 1981; Galiano, Sterling & Viejo, 1985; Usher, 1986), because of its |
convenient obtention and handling, although it offers, by itself, just a limited ,
information.
The chorological index proposed by Kudrna (op. cit.) is used here, having been
28(4):289-296, 1989(91)
291
n Holm oak woods (on basic soils)
K / ^ » » >. (on acid soils)
” ” ” (wetter climate)
tX-jvJ Lusitanian oak woods
39MI Pyrenean
no .. (atlantic climate)
f I Beech woods
Pine
ecSM Alpine meadows
Fig. 1 . Map of the climax vegetation of Madrid province (modified from Rivas
Martinez, 1982).
used before by the authors in butterfly conservation studies (Sanchez & Viejo,
1988; Viejo & Viedma, 1988; Viejo, Viedma & Martinez, op. cit .) and it is the sum
of three variables related to species range: size, composition and affinity. This
index ranges from 4 to 14; high values mean biogeographically peculiar species
(European endemic species with a very small range), while low values correspond
to widely distributed species. The mean of the chorological index of the species
occuring within a square is the square’s Average Chorological Index. The higher
this value, the more peculiar fauna, biogeographically speaking, in the consid-
ered square.
Data were processed with the BMDP ID program at the Computer Center of
the Universidad Autonoma de Madrid.
292
J. Res. Lepid.
Fig. 2. Map of species richness in each 100 km2 UTM grid in Madrid province.
Area of study
This paper is based on faunistic data of the Madrid province, located in
the center of the Iberian Peninsula, between the 40° and 41° N parallels
and the 3° and 4° W meridians. It is approximately triangle-shaped and
has a surface of 8,000 km2.
Geomorphologically, Madrid can be divided into two parts: the Sierra
de Guadarrama (North) and the Llanos del Sur (southern Plains),
according to Hernandez Pacheco (1941).
The Sierra de Guadarrama.- These mountains are included in the
Sistema Central, that goes across Madrid province following the main
direction of this range, that is from east-northeast to west-southwest,
and runs along the north border of the province for 100 km, ranging from
1,000 m (altitude at the surrounding plain) to 2,430 m a.s.l. It is
essentially constituted by archaic siliceous rocks (mainly granites and
28(4):289-296, 1989(91)
293
7. 0 - 7. 5
6. 5-6. 9
I 6.0-6. 4
X § 5-5.9
Fig. 3. Map of the Average Chorological Index values in each 1 00 km2 LJTM grid in
Madrid province.
gneiss), although marly and cretaceous limy lands, miocenic arkosic
sands and quaternary alluvial soils are also present. Its climate even
inside the general continentality, is more humid and colder than that of
the Llanos del Sur, and it is classified as Humid Mediterranean type,
following to Emberger (Viejo, 1982). From a botanic point of view, the
Sierra de Guadarrama belongs to the Mediterranean Region, Carpetano-
Iberico-Leonesa province (Rivas Martinez, 1982; Izco, 1984) and three
bioclimatic levels can be distinguished: Supramediterranean,
Oromediterranean and Crioromediterranean.
The climax vegetation of each level is respectively: Holm oak ( Quercus
ilex) and Pyrenean oak ( Q . pyrenaica ) woods, Scottish Pine ( Pinus
sylvestris ) woods scattered with J uniper J uniperus communis) trees and
high mountain alpine meadows. Cattle raising and forestry are wide-
spread land uses in the Sierra.
294
J. Res. Lepid .
Llanos del Sur.- Located at the south of the Sierra, it is a wide and flat
region. Its altitude ranges from 500 to 1,000 m a.s.l. This region is lightly
south-exposed, and the Tagus river traverses it along its southern
border. Siliceous alluvial soils (arkosic sands) are dominant on the north
and west, as well as evaporitic rocks (loams, gypsums, and pontiensic
limestone)are on the south and east. Its climate can be classified between
Temperate Mediterranean and Semiarid Mediterranean (Viejo, 1982), 1
being much dryer and warmer than that of the Sierra, increasing in this
tendency while going further south. This climate, the substratum and
the vegetation establish a transition zone in the meeting region with the
Sierra, sharing at this point some features with it. Llanos del sur are
included in the Mediterranean Region, Carpetano-Iberico-Leonesa and
Castellano-Maestrazgo-Manchega botanical provinces, and only the
Mesomediterranean bioclimatic level is present. The climax vegetation
consists of Holm oak and Lusitanian oak (Quercus faginea) woods, as well
as Mediterranean shrubs, although it is very disturbed by land uses,
mainly agriculture and urbanism.
Results and discussion
Figure 2 shows species richness in each square, that ranges from 10 to
102 species. The north of the province has the highest species richness
per square. In this area two very rich zones can be distinguished: One on
the center and the other on the northeast end (Viejo, Martin & de Silva
1988). Another relatively rich region appears on the southeast, with GO-
75 species squares. The mid-province region is rather poor, coinciding
with the metropolitan area of Madrid. The highly cultivated Tagus
Valley, at the south end of the province, is the poorest region. Comparison
between the species richness and climax vegetation maps (fig. 1) shows
that highest species numbers correspond, to a large extent, with the
Pyrenean Oak ( Q.pyrenaica ) and Lusitanian Oak (Q. faginea ) climax
areas, at the north and at the southeast regions respectively.
Figure 3 shows Average Chorological Index of every square, which
varies from 5.5 to 7.5. The distribution of this variable is different than
that of the former (Species Richness). The highest values correspond to
the Mesas del Sureste (southeast Plateaux), climax domain of the
Lusitanian Oak, although there are also some high ones on the central
and southern areas of the province. Lowest values appear on the most
altered areas (furthest south end) and on the Sierra de Guadarrama.
There is an interesting point to comment: Richest squares are not
necessarily those with highest Average Chorological Index (correlation
between both variables, r= 0.06). This is because of the own landscape
nature and, consequently, because of the different lepidopteran species
that occur in them. The north of the province is mainly cool and humid,
and its vegetation corresponds to the phytosociological series of the
Pyrenean Oak and Scottish Pine, and these vegetal formations are much
closer (as a floristic whole) to those mideuropean-atlantic, than the Holm
28(4):289-296, 1989(91)
295
Oak, Lusitanian Oak and Kermes Oak woods of the south of the province
are, which is a highly Mediterranean area. In other words, there are more
species with low chorological index in the northern Mountains than in
the southern Plains, given that the environmental conditions on the
Sierra de Guadarrama (mainly climate and vegetation) are very close to
those on west and Central Europe. On the other hand, endemic species
and biogeographically “rare” species (high chorological index) occur in
typically Mediterranean biotopes (Baz, 1991).
Note that even farming lands, at least those of non irrigated croplands
(olive groves, vineyards or cereal fields), present high Average Chorological
Index; that means, many biogeographically interesting species can be
found here, even if species richness is not high at all (Viejo, 1985).
Conclusions
Obviously the lepidopterologically interesting areas selection, pointing
towards their protection, must be based on deeper studies than just the
analysis of the species range UTM maps. But it is also evident that in the
meanwhile these maps are the only useful argument to establish possible
protected zones. Nevertheless, we consider that species richness is a
limited criterion, because if we apply no other criterion, no attention will
be paid to areas with a low species richness, but may be sheltering a
biogeographically interesting fauna; that means the south of Madrid in
the present case. By these reasons, we believe absolutely necessary to
deep in the analysis, and applying other criteria as well, such as the
biogeographic interest of the fauna (Idle, 1986), easily provided by the
UTM maps.
Finally, we want to point out the interest that a rather mideuropean
fauna has, inside a tipically Mediterranean environment, feature that
increases the peculiarity of Sierra de Guadarrama fauna, at least from
an Iberian perspective.
Literature cited
Baz, A. 1991. Ranking species and sites for butterfly conservation using presence-
absence data in Central Spain. Nota Lepid. Suppl. 2:4-12.
Gallano, E.F., Sterling, A. & Viejo, J.L., 1985.The role of riparian forests in the
conservation ofbutterflies in a Mediterranean area. Environmental Conservation
12:361-362.
Gomez de Aizpurua, C., 1977. Atlas provisional de los lepidopteros del norte de
Espana. Diputacion Foral de Alava. Vitoria.
1979. Atlas provisional de los lepidopteros del norte de Espana. Anexo 1.
AEPNA. Vitoria.
1980. Atlas provisional de los lepidopteros de Madrid (Papilionoidea,
Hesperioidea & Zygaenoidea]. Comunidad de Madrid.
1988. Atlas provisional de los lepidopteros de la zona norte. Gobierno vasco.
Vitoria.
Gonseth, Y., 1987. Atlas de distribution des Papillons diumes de Suisse (Lepidoptera,
Rhopalocera). Documenta Faunistica Helvetiae 5: 242 pp. CSCF Neuchatel.
296
J. Res. Lepid.
Heath, J. & Skelton, H. J., 1973. Provisional Atlas of the Insects of the British Isles.
Part 2: Lepidoptera. BRC, Nature Conservancy.
Hernandez Pacheco, F., 1941. Caracterfsticae fisiograficas del territorio de Madrid.
Anal. Cienc. Naturales I. Jose Acosta pp. 36-46.
Idle, E.T., 1986. Evaluation at the local scale: A region in Scotland. In Usher (ed.)
Wildlife Conservation Evaluation, pp. 181-198. Chapman & Hall. London.
Izco, J., 1984. Madrid verde. Ministerio de Agricultura, Pesca y Alimentation y
Comunidad de Madrid. 517 pp. Madrid.
Kudrna, O., 1986. Butterflies of Europe. Vol. 8. Aspects of the conservation of
butterflies in Europe. Aula Verlag. 323 pp. Wiesbaden.
Margules,C.R. & Usher, M.B., 1981. Criteria used in assesing wildlife conservation
potential: A review. Biol. Conserv. 21: 79-109.
Munguira,M.L., 1987. Biologla y biogeograffa de los licenidos ibericos en peligro de
extincion (Lepidoptera, Lycaenidae}. Tesis doctoral inedita. 462 pp. Madrid.
Rivas Martinez, S., 1982. Mapa de las series de vegetation de Madrid .Diputacion
de Madrid.
Sanchez, B.L. & Viejo, J.L., 1988. Estudio preliminar de las mariposas de la parte
oriental de los Montes de Toledo (Lepidoptera: Papilionoidea & Hesperioidea).
SHILAP Revta. lepid. 16 (61): 31-37.
Thomas, C.D. & Mallorie, H.C., 1985. Rarity, species richness and conservation
butterflies of the Atlas Mountains in Morocco. Biol. Conserv. 33: 95-117.
Uherkovich, A., 1983. A zselic nagylepkefaunaj a IV. Kelet-Zselic: Pale komyeke
(Lepidoptera). A Janus Pannoius Muzeum Evkonyve 27: 37-50. Pecs.
Usher, M.B., 1986. Wildlife Conservation Evaluation : Attributes, Criteria and
Values. In Usher (ed.) Wildlife Conservation Evaluation pp. 3-44. Chapman &
Hall. London.
Viejo, J.L., 1982. Fenologfa y preferencias climaticas de los Pieridae (Lepidoptera)
en la provincia de Madrid. Miscelanea Commemorativa . Universidad Autonoma
de Madrid, pp. 973-1000.
1983. Las mariposas de la Depresidn del tajo . Universidad Complutense.
Madrid.
1985. Diversity and Species Richness of Butterflies and Skippers in Central
Spain Habitats. J. Res. Lepid. 24 (4): 364-371.
Viejo, J.L., Martin, J. & de Silva, C., 1988. Patrones de distribution de las
mariposas de Madrid (Insecta, Lepidoptera, Papilionoidea & Hesperioidea)
Ecologfa 2:359-368.
Viejo, J.L. & Templado,J., 1986. Los pieridos,satfridos y ninfalidos (Lep.) de la
region de Madrid en relation con las formaciones vegetales. Graellsia 42: 237-
265.
Viejo, J.L., & Viedma, M.G., 1988. Los bosques y la conservation de las mariposas
en el centro de la Peninsula Iberica (Lep. Papilionoidea & Hesperioidea). Bol.
R. Soc. Esp. Hist. Nat. 84 (1-2): 153- 164.
Viejo, J.L., Viedma, M. G. &• Martinez FalEro, E. , 1989.The importance of Woodlands
in the Conservation of Butterflies (Lep. Papilionoidea & Hesperioidea) in the i
Centre of the Iberian Eeninsula. Biol. Conserv. 48: 101-114.
Journal of Research on the Lepidoptera
28(4):297-309, 1989(91)
An unrecognized, now extinct, Los Angeles area
butterfly (Lycaenidae)
Rudolf H. T. Mattoni
9620 Heather Road, Beverly Hills, CA 90210, USA
Abstract. Philotes sonorensis has been regarded as a geographically
invariant species. This historic viewpoint is corrected and several
geographic variants and patterns of variation are described. A new
subspecies, P. sonorensis extinctis, is named. The subspecies became
extinct in 1967 consequent to an engineering program for water
diversion. The relationship of P. s. extinctis and parapatric P. s.
sonorensis are discussed.
Introduction
California not only leads the nation as the trendsetter of fashion, the
capitol of entertainment, and the model of buoyant lifestyle, but also as
the exterminator of species, including butterflies. “Species” is used here
in the context of the federal Endangered Species Act which for inverte-
brates includes subspecies. Extinction means globally lost, versus extir-
pation, which refers to extinction in only part of the range.
The first recorded North American butterfly extinction was Cercyonis
sthenele sthenele, last collected in 1880, followed by Glaucopsyche
lygdamus xerces in 1943-44. Both were victims of land conversion of the
San Francisco sand dunes, dunes which now underlie about half the area
of the city and which today are scarcely recognizable. Loss of the Xerces
blue was especially unfortunate as its populations were a highly poly-
morphic complex ranging from the spectacular xerces phenotype to that
of the surrounding parapatric and widespread subspecies incognita. The
pattern of variation may have been an ecologic/genetic parallel to the
situation described by this paper. In 1958 Parnassius clodius strohbeeni
was last seen in the Santa Cruz mountains, a possible victim of
overcollecting. The next known extinction was the unexplained disap-
pearance of Argynnis (Speyeria) adiaste atossa around 1960 (Emmel and
Emmel, 1973). This fritillary was formerly abundant in the Tejon
Mountains near Los Angeles. After 1983 Glaucopsyche lygdamus
palosverdesensis of suburban Los Angeles was no longer seen, in spite of
intensive attempts by a squad of experienced collectors under the able
leadership of Jess Morton (Mattoni, unpublished). The species was lost
to a combination of overcollecting, poor weather and habitat fragmenta-
tion. The time of the last flight of an undescribed subspecies of Plebejus
saepiolus in the Big Pine area of the San Gabriel mountains was 1985
(Emmel, pers. comm.). At least two additional species are in imminent
danger of extinction: Argynnis ( Speyeria ) adiaste clemencei and
Euphydryas editha quino (= wrightii of authors)( Allen, Brown, Ballmer
298
J. Res. Lepid.
& Mattoni, unpublished data). Although neither were seen for several
years after 1986, the fritillary was widespread with only a single
population of the checkerspot reported in 1990. These last observations
are hardly encouraging. Several other species are probably not too far
behind. These events were so rapid that no timely help was provided by
the listing process under the Endangered Species Act. Between wide-
spread political attacks to weaken the Act and serious understaffing of
agencies, the future for biodiversity is indeed bleak.
The list can now be expanded by a previously unreported subspecies
which became extinct in 1967. The event passed unnoticed because of an
unrecognized systematic situation I will in part rectify with this paper.
Failure to formally notice significant geographic variation in Philotes
sonorensis was perhaps a function of later authors assuming authority
of earlier authors who did not notice consistent patterns of variation
other than naming one form and one aberration. The species clearly
stands apart from all Scolitantidine blues, without apparent sister
species, in the monotypic genus Philotes . The entire species is almost
completely confined to the California Floristic Province (described by
Raven, 1988), a trait shared with only eight other butterflies. This
isolation, combined with a striking appearance, may have biased observ-
ers into overlooking complex variations. However, Langston ( 1963) broke
with tradition and cited a substantial and consistently different appear-
ance of specimens from central coastal California when compared with
those from the south, figuring females of each. Langston later (1972)
referred to macule and aurora variation in northern California colonies.
From his thesis on Philotes Shields (1973) noted that Los Angeles County
specimens are larger with the females more boldly marked. He found no
geographic variation in valve teeth number in males, cited the Mattoni
and Seiger ( 1963) report of intrapopulation variation of UFW postmedian
macule number in populations of the San Gabriel Canyon wash, and let
the matter rest. During the same time period Fred Thorne (pers. comm.)
provided specimens and advised that San Diego County populations
from the desert (Sentenac canyon area) and coast (Pt. Loma/La Jolla)
were sufficiently distinct to warrant subspecific status. Coastal San
Diego County populations no longer exist, although there may be rem-
nants along the northern Baja California coast (Brown and Faulkner,
pers. comm.). The species distribution is shown in figure 1.
Inspection of series of specimens from throughout the range shows
several distinct sets of wing pattern types which beg further systematic
study. With escalating destruction of natural habitat such study should
be undertaken soon. While preparing a guide for identification and
conservation biology of butterflies of the Los Angeles area (Mattoni,
1990), it was necessary to formally name the unique population described
below:
28(4):297-309, 1989(91)
299
Fig. 1 . Distribution map of Philotes sonorensis. Data after Shields 1973 with a few
recent records.
300
J. Res. Lepid.
Philotes sonorensis extine tis Mattoni new subspecies
Males. Upperside, Cyanic overlay as in nominotypical species. Forew-
ing:: postmedian macule number vary in number from none to li ve, with
frequency distribution given in table 1; holotype with four. Hindwing as
in nominotypical species. Underside. Ground medium grey-brown.
Fringes well distinguished at all veins. Forewing. Macule pattern as in
nominotypical species. Hindwing. Median space between sub-basal and
postmedian macule usually lightened against ground by whitish suffu-
sion, postmedian space darker grey than ground with submarginal
macule absent and with submarginal space light grey usually most
strongly marked in M3, Cu. and Cu2. Marginal macules faint.
Females, Upperside, Cyanic overlay similar to nominotypical species,
but slightly and uniformly darker due to a higher proportion of melanic
scales. Marginal band wider and macules 10 to 30% larger on average
than other populations presenting a darker overall aspect. Forewing.
Postmedian macule number in interspaces Ou.;! and Oil, vary from 0 to 3.
Hindwing. Macules and orange aurora larger than other populations,
entire spaces anterior to Rs with melanin suffusion, again presenting a
darker overall appearance than nominotypical species. Underside, In
all aspects similar to males.
Types: All specimens taken in the upper San Gabriel wash from
February through April over a period of 1922-1987, after which they were
extinct. Older specimens are variously labelled San Gabriel Canyon, San
Gabriel Canyon wash, Fish Canyon, and Azusa. Holotype male and
allotype female III 24 1963, R. H. T. Mattoni leg.
Type disposition: The holotype and allotype will be placed in the
Smithsonian Institution. 115 paratypes will remain in the author's
collection until further systematic issues are resolved and will then be
placed in an appropriate institution. The Los Angeles County Museum
of Natural History has 255 paratypes. All specimens figured will be
deposited in the Los Angeles County Museum of Natural History.
Etymology: The subspecies name calls attention to the fate of the taxon, j
I suggest the common name Human Folly Blue because the extinction
was due to a short term engineering fix without recognition of long range
environmental impacts. The U. S. Army Corps of Engineers destroyed i
the habitat to provide a spreading basin for ground water recharge. Two
consequent ironies of the action are that the Corps of Engineers would
today be prevented from such action by its own mandate to preserve
riparian habitat and that the groundwater basin being recharged is now
contaminated with chlorinated organic chemicals. This historical lesson
of environmental tinkering appears forever condemned to repetition.
Nomenclature and Synonymy: C, and R. Felder (1865. Reise Novara )
2:281 & plate 35 figs. 3,4) named Lycaena sonorensis with the habitat I
designation of Sonora (Lorquin). The Felder (Lorquin) “Sonora” type j
locality issue was discussed by Brown (1967). Both G. Shields and J.
28(4):297-309, 1989(91)
301
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described in text and shown in figure 2. The samples are grouped into the San Gabriel Mountains, which had two distinct phenotypic class
populations; the San Diego coast and desert, each with a distinctpopuSation; the central California coast; and Chili Bar in the northern Sierra
Nevada Mountains. When total sample of female set is less than 6, the class frequency is not scored, but the class size is given (East Fork
only). For sample site 4, 1963, only data for classes A and D are available.
302
J. Res. Lepid ,
Emmel provided additional information (in lilt.). A population from
‘'environs de Los Angeles" was named L. regia by Boiduval (1889. Ann.
Ent. Soc. Belgique 12: 46) but was subsequently synonymized with L.
sonorensis by Reakirt (1878. Butterflies and Moths of North America).
Comparison of published figures indicate the Boisduval specimens differ
phe i.:to typically from those of the Felders. Both Felder and Boisduval
material clearly was collected by Lorquin, but the exact origin of any of
the specimens remains obscure. Two pairs of Boisduval specimens
labelled “type" are in the USNM (Oberthur collection). The two well worn
Felders syntypes are males in the BM(NH). Photographs of a pair of
Boisduval syntypes appear similar to extinctis , with the dark postmedian
space. The other pair is marked as the widespread montane populations.
The specimen figured by the Felders is not extinctis , but appears similar
the nearby populations and lighter Boisduval syntypes.
Reconstructing the Lorquin type localities revealed the specimens
were likely taken in 1852 when Lorquin travelled around Los Angeles
and also in San Diego. During travel near Los Angeles he took
Glaucopsyche piasus sagittigera , most likely near the Verdugo Hills and
may well have collected L. sonorensis at same time. Neither named taxa
conforms to extinctis, A type locality must be designated when the species
is thoroughly studied.
Diagnosis
The San Gabriel wash population was distinct and deserves special
recognition for its combination of three characteristics: 1) postmedian
macule pattern frequency and dimorphism of a unique form, 2) complex
difference in the underside ground and maculation pattern in almost all
individuals, 3) very high population densities.
1). POSTMEDIAN MACULE PATTERN FREQUENCY AND DIMOR-
PHISM OF THE UNIQUE FORM “COMSTOCKT
The postmedian macule patterns were arbitrarily designated by letter
for males and number for females and are illustrated in figure 2. The
male classes X, A, B, C, D, G, and I represent a decreasing macule number
in interspaces Rs to Cu 1 ranging from 5 to 0 macules. Classes B and C
both have 3 macules but different positions. The female classes range
from 3 to 0 macules within interspaces Cu 1 and Cu 2.
The form “comstocki” (CM) is illustrated in figure 2, second specimen
in row 3 and Mattoni ( 1964: specimen 15). On the up per side, male CM are
indistinguishable from “normal” specimens that are without macules
(pattern class I), but the underside is obviously distinct. The ground is
entirely the darker grey that is restricted to the postmedian interspace
in the normal. The hindwing macules are absent and the forewing
postmedian macules are aggregated into a single discoid al macule.
Female CM undersides are as the male, but the upperside fore wing
macules are distributed as on the underside. The CM character state was
28(4):297-309, 1989(91)
303
Fig. 2. Specimens of F. sonorensis extinctis from San Gabriel Canyon wash showing
classes of upperside pattern and underside variation. Left to right. Row 1 ,
males: X, A, B, C; row 2, males: D, G, I (or CM, difference in underside),
asymmetric, D on left, B on right; Row 3, undersides: wild type, CM
(“comstocki”), D/CM, upper San Gabriel canyon; row 4, females: 0, 1 , 2, 3.
See text for further explanation. Unless otherwise stated all specimens in
figures leg. R. Mattoni.
probably controlled by a recessive gene that modified melanin deposition
at a critical stage during pigment formation in the pupa. The hypothesis
that the CM variant was environmentally induced cannot be discarded,
yet failure to observe CM in any other populations and its relatively high
frequency at San Gabriel strongly supports a genetic explanation. Reces-
siveness is inferred from a report of all wild type progeny from a CM
female by an early collector, but both report and undocumented data are
hearsay.
Following the conclusion of their fieldwork, Mattoni and Seiger (1963)
noted an additional distinct variant class: rare males with state D
upperside macule pattern and females with a 0 macule pattern, but with
a underside primary postmedian macule series less than half the dis-
tance from the discoidal macule to the distal wing margin. This variant
(D/CM) is illustrated by specimen 11 in figure 2, 13 in figure 3, and
specimens 13, 14, and 20 in Mattoni ( 1964). Our hypothesis was that this
variant represented the heterozygote of CM, as its frequency approxi-
304
J. Res. Lepid.
Fig. 3. Variation in upperside patterns in females and underside patterns comparing
specimens from lower San Gabriel canyon wash (extinctis) and upper San
Gabriel canyon (sonorensis). Row 1 , females, wash. Row 2, females, upper
canyon at Coldbrook ranger station. Row 3, undersides, wash. Row 4,
undersides, Coldbrook ranger station. Row 5, undersides, fire road or site
7, intermediates, see text.
mated the Hardy-Weinberg equilibrium in the small sample we made in
1963. Mattoni (1964) published a color plate illustrating these forms as
well as samples from other populations. The legend for this plate is given
below, as this information is not elsewhere available.
The frequency of all the postmedian macule classes and CM are given
for populations from which more than 20 specimens were available. It
should be noted that asymmetry is exceptional. The 14-specimen East
Fork sample was included to increase the upper San Gabriel canyon I
population. Three conclusions can be drawn from these data: wash
population ( extinctis ) males had a significantly greater frequency of class
D (except Atascadero) and a significantly lower frequency of class X than ,
any other populations, wash females had a higher frequency of class 3
i
28(4):297-309, 1989(91)
305
Fig. 4. Specimens representative of different geographic areas. Row 1 , females, El
Dorado county, Chili Bar, leg. O. Shields. Row 2, undersides, same data as
row 1 . Row 3, undersides, Santa Clara county, Alum Rock Park, leg. R.
Langston. Row 4, female and 3 undersides, San Diego county, Sentenac
Canyon, leg. F. Thorne. Row 5, female and 3 undersides, Santa Barbara
county, Santa Barbara, leg. R. Denno.
than any population (except San Diego coast, possibly representing a
sampling error, but see below), no valid specimen of CM has ever been
observed from any but the San Gabriel canyon wash population. Since
j the extinction of extinctis , local collectors mostly take their specimens
from other parts of the San Gabriel mountains, usually in Brown’s Gulch,
located 3 miles north of what was the wash. Form CM has never been seen
in spite of a thousand or more takes in the vicinity. A specimen of CM
reported by Shields (1973) from Ventura County (Henne, leg.) was
i apparently a class I specimen in which the underside was not inspected.
| The Henne collection in the LACM has a Ventura I male with a normal
underside.
306
J. Res. Lepid.
Fig. 5. Map of distribution of Philotes sonorensis in the lower San Gabriel wash prior
to 1 967. Areas in black remained in 1 968, although no butterflies remained.
These last remnants destroyed in 1 980’s.
2). THE UNDERSIDE PATTERN COMPLEX
Virtually every specimen from the wash population can be separated
from the populations in upper San Gabriel canyon and most other
localities by the underside pattern. The character is illustrated in figures
2,3, and 4 as well as Mattoni ( 1964). The difference between the wash and
upper canyon populations is most striking in figure 3, comparing rows 3
and 4. The border between these character states is abrupt, the limit
apparently having been the edge between the wash and the steep slopes
marking the beginning of the canyon walls. This border is shown on the
map, figure 5. Butterflies taken at this interface are shown in figure 3,
row 5. The specimens taken here, which were rare, indicate a zone of
intergrades and segregates. Since early 1980 access to the area has been
blocked, so status of the species is unknown at the site.
The distinct dark grey postmedian space on the secondaries occurs in
coastal San Diego county populations (extirpated, see Mattoni 1964
figure 21) and some other alluvial washes from the south slopes of the
San Gabriel mountains. The latter have not been well sampled and today
few, if any, remnants of these wash populations are extant.
Samples of underside patterns from other populations are illustrated
for comparative purposes. These include desert San Diego county figure
28(4):297-309, 1989(91)
307
4, row 4 and Mattoni 1984, 29-32), Santa Barbara (figure 4, row 5 and
Mattoni 1964, 25-28), Alum Rock Park, Santa Clara county (figure 4, row
3), and Chili Bar, El Dorado county (figure 4, row 2). The Chili Bar
population is also singular in that 90% of the sample lacked checkered
fringes.
3). HIGH POPULATION DENSITIES
The reason the San Gabriel wash was the long favored locality of
collectors of the Human Folly Blue was the extremely high population
numbers of the butterfly in the small circumscribed area where it
occurred (figure 5). The 1955 and 1956 study of Mattoni and Seiger ( 1963,
and unpublished) indicated total standing populations in those years on
the order of tens of thousands in the 8 square kilometer area the
population inhabited. During the period beginning with the discovery of
the population until its destruction in 1967, collectors could easily take
several hundred specimens in a day. No other known population of the
species had or has the potential of such yields. Abundance of individuals
of the extinctis population was unique in terms of high densities in every
year for which records are available. The density characteristic was not
a function of foodplant density, as many other populations (i.e. Baja
California, central California coast) are found in regions where Dudleya
lanceolata and D. cymosa are among the dominants in their plant
communities yet the butterfly remains rare.
Rarity has only recently been viewed from the standpoint of relating
the characteristics of species that define rareness (Rabinowitz, 1981).
Through most of its range P. sonorensis is rare in the sense of being
constantly sparse yet occurring across several limited habitats. Under all
conditions it is distribution limited by the occurrence of its foodplant,
usually a colonial and local plant. Because butterflies are all r- strate-
gists, excepting possibly the giant Ornithoptera , rarity must have an
ecological and/or genetic bases. All populations have the potential of
rapidly achieving high density, but do so only on occasion. The very dense
population of extinctis occurring adjacent to, and probably interbreeding
with, low density sonorensis implies a gap in adaptive characteristics.
Populations of the species from the nearby Big and Little Dalton, Santa
Anita and Eaton washes occur(red) only in low density and without the
diversity of forms found at San Gabriel.
Mierogeographic distribution and systematic implications
The wash population distribution as known in 1963 is mapped in figure
5. At that time there was undisturbed wash habitat to the south of the
extant population, but no butterflies could be found although foodplant
was present. The areas to the east of 3 and west of 1 had been denatured
by residential construction. It is unknown if the butterfly ever occurred
in these sites. The black overlay denotes undisturbed sites remaining in
1968. Both were scouted in that year without finding specimens, al-
808
J. Res. Lepid.
though a few were taken on the fire road, where they must still occur, but
is now inaccessible. These last sites were denatured by construction and
clearing in about 1980.
Site 7 (fire road) referred to all the steep slope east and north of site 6
and the bridge. The flat wash immediately north of 6 is an orchard. At this
point the road was located within a few feet of the river and it was possible
to walk about a mile upriver. Although foodplant was present over this
entire area, the butterfly was uncommon. Specimens sampled in site 6
were all of the extinctis pattern. The sedentary nature of the species
(Mattoni and Seiger, 1963 and unpublished; Keller, Mattoni and Seiger,
1966) probably limited movement across the river between sites 6 and 7.
To what extent the distinct patterns and characteristics of sonorensis
and extinctis were maintained by selection as opposed to loss by hybrid-
ization remains unknown.
Coda
The Philotes sonorensis sonorensis / extinctis relationship had the
potential of providing a fascinating case for investigating evolutionary
biology in sedentary butterflies. The contrast of two adjacent populations
with different complex wing patterns, clear-cut polymorphisms, and
ecologies presented a singular situation. Destruction of the wash habitat
and attendant extinction of extinctis co-opted further investigation. Yet
at some future time, when human species density is of necessity reduced
and constrained by resource limitation, and the San Gabriel river dams
no longer function due to siltation, the wash habitat may be revegetated
and a population of the butterfly could re-invade the current biological
desert. Should curiosity of biological matters survive for future humans,
this note may be useful.
Acknowledgements. The original manuscript was substantially rewritten and
vastly improved following input and sometimes pungent comment from Mike
Collins, John Emmel, Tom Emmel, Merrill Peterson, Barry Prigge, and Oakley
Shields. W. D. Field of the Smithsonian Institution generously provided photo-
graphs of one pair of the regia types. Chris Henne translated the Boisduval
description from the French.
Legend for Mattoni (1964). Specimens read left to right. Top five rows, all
San Gabriel Canyon wash 1963, Males wing pattern class: 1, C. 2, B. 3,
A. 4, X. 5, Female “comstocki” (CM). Males: 6, 1. 7, G. 8, D. Females: 9,
3 10, 2. 11, 1. 12, 0. Undersides: 13, D/CM?. 14, D/CM?. 15, CM. 16,
Female, darkly marked. Undersides: 17-19 variants of wild type. 20, D/
CM?. 21, San Diego, Paradise Valley (Fred Thorne, leg.). 22-24, Upper
San Gabriel Canyon, East Fork (Mattoni, leg.) Underside, 22. Female,
23. Male, 24. 25-28, Santa Barbara, (R. F. Denno, leg.). 29-32, San Diego
Co., Sentenac Canyon (Fred Thorne, leg.).
28(4):297-309, 1989(91)
309
Literature cited
Brown, F. M. 1967. Lorquin’s localities “Sonora” and “Utah”. Jr. Lepid. Soc. 21:271-
274.
Emmel, T. C. and J. F. Emmel. 1973. The butterflies of southern California. Sci. Ser.
Nat. Hist. Mus. L. A. No. 26
Hovanitz, W. 1967. Natural Habitats. Jr. Res. Lepid. 6: 199-202
Keller, E. C., R. H. T. Mattoni and M.S.B. Seiger. 1966. Preferential return of
artificially displaced butterflies. Anim. Behav. 14:197-200.
Langston, R. 1963. Philotes of central coastal California. Jr. Lepid. Soc. 17:201-223.
1972. The Sonora blue in 1971 the earliest season for the north. Pan Pac. Ent.
48:67).
Mattoni, R. H. T. 1964. Jf. Res. Lepid. 3 (1) cover illustration.
— — 1990. Butterflies of greater Los Angeles. Lepidoptera Research Foundation,
Beverly Hills, CA
Mattoni, R.H.T. and M.S.B. Seiger. 1963. Techniques in the study of population
structure in Philotes sonorensis. Jr. Res. Lepid. 1:237-244.
Rabinowitz, D. 1981. Seven forms of rarity, in Singe, H. The biological aspects of
rare plant conservation. Wiley. London.
Raven, P. H. 1988. The California flora, in Barbour, M. G. and J. Major, eds.
Terrestrial vegetation of California. California Native Plant Soc. Sacramento.
Shields, 0. 1973. Studies on North American Philotes II. The biology, distribution,
and taxonomy of Philotes sonorensis (F. & F.) Bull. Allyn Mus. 15:1-16.
— — - 1973a. Studies on North American Philotes I. Roosting behavior, tending
ants, parasites, and predators. Bull. Allyn Mus. 10:1-5.
28(4):310-315, 1989(91)
Notes
Gesneriaceae as a larval hostplant of Hyposcada virginiana
(Nymphalidae: Ithomiinae)
Knowledge of the hostplant interactions within the subfamily Ithomiinae have
been important to our understanding of neotropical butterfly evolution and
ecology (e.g., Mielke & Brown 1979; Gilbert 1983; Ackery & Vane-Wright 1984;
Brower 1984; Boppre 1984; Brown 1987; DeVries & Stiles 1990; Vasconcellos
Neto 1991). Broad patterns of hostplant use in the family Nymphalidae are well
known, and it is clear that the Ithomiinae is largely composed of specialists on the
Solanaceae, with a few species that use the Apocynaceae (Drummond & Brown
1987; DeVries 1986, 1987; Ackery 1988). There is, however, one frequently cited
exception to the overall pattern of ithomiine hostplant interactions : Haber (1978)
reported that Costa Rican Hyposcada virginiana evanides Haensch may oviposit
on the genera Columnea and Drymonia (Gesneriaceae).
The potential use of Gesneriaceae as a larval hostplant by any member of the
Nymphalidae is singular. In fact, the only records of Gesneriaceae in the
extensive, world-wide review of nymphalid hostplants by Ackery ( 1988) are those
of Haber ( 1978). In conflict with Haber’s records (but in line with other ithomiine
host records) are observations by Drummond & Brown (1987) that Brazilian
Hyposcada egra (Hewitson) oviposited on a plant thought to be Markea
(Solanaceae), and that the larvae were reared in the laboratory on Juanulloa
(Solanaceae). Thus, with our understanding of nymphalid hostplant relation-
ships in general, and those of the Ithomiinae in particular, there is need for
verification or rejection of Gesneriaceae as a hostplant of Hyposcada (DeVries
1986, 1987; Drummond & Brown 1987; Ackery 1988). Here I provide the first
direct field corroboration of Haber’s (1978) suggestion that Hyposcada uses
Gesneriaceae as a larval hostplant.
On 12 April 1990 at 0810 hrs I observed a female Hyposcada virginiana oviposit
3 eggs on the underside of an intermediate age leaf of Drymonia sp. (Gesneriaceae)
at JardinBotanico Wilson, San Vito de Java, Costa Rica. The woody, hemiepiphytic
plant with glabrous, semi-succulent leaves, was attached to the side of a palm
tree approximately 5 m above the ground, and was in shade at the time of
oviposition. Each oviposition act was separated by about a 30 second interval,
and the eggs were deposited near the middle of the leaf. The white eggs were
large for an ithomiine (« 2 mm diameter), bore a sculpturing that could be
detected without the aid of a lens, and were slightly wider towards the micropylar
region than the base. First instar larvae (body entirely pale grey with shiny black
head, and no tubercles) hatched 5 days later, ate the egg shell, rested for 24 hours,
and then began eating small, round holes in the leaf. Second, third, and fourth
instars bore no projections or papillae, all were shiny, semi-transparent grey with
a dull yellow band at the interface of venter and lateral areas that extended from
segment A-8 to T-l, continuing across the anterior margin of T-l. In all instars
the head was shiny black, and without patterns or relief. Although all three
larvae were healthy and growing vigorously, a necessary move to a different field
site where no acceptable hostplant occurred prohibited rearing them beyond
28(4):310-315, 1989(91)
311
fourth instar. The larvae were preserved in ETOH and specimens are in both the
author’s voucher collection and that of the Museo Nacional de Costa Rica.
None of the larvae fed at the leaf margin, but as is typical of many ithomiines
that feed on Solanaceae, they ate round holes in the interior of the leaf blade, then
moved to another undamaged section to eat another hole. The larval feeding
behavior left the once entire Drymonia leaf with a large number of irregular
holes. The larvae were cryptic while on the plant and fell into what Ackery ( 1988)
pointed to as the typical Solanaceae feeding type of ithomiine. A casual
inspection of 10 other Drymonia sp. plants in the area showed that most of their
leaves had feeding damage similar to that caused by H. virginiana larvae.
However, I found no other H. virginiana larvae at this or subsequent inspection
over an intermittent three month period.
The observations here raise three points. First, although it is unknown
whether the H. virginiana larvae would have produced adults, the present
observations support Haber’s (1978) records that Gesneriaceae is a hostplant for
Costa Rican Hyposcada. Secondly, the general vegetative similarity between
some hemiepiphytic Solanaceae (i.e. , Markea , J uanulloa ) and some hemiepiphytic
Gesneriaceae (rounded, glabrous, semi-succulent leaves) allows for the possibil-
ity that the H. egra oviposition record (Drummond & Brown 1987) was actually
on a Gesneriaceae. Finally, when the observations of Drummond & Brown ( 1987)
and those here are considered together, they suggest the possibility of some
chemical similarity between Solanaceae and certain Gesneriaceae.
Acknowledgements . Thanks to L. D. Gomez for identifying the hostplant, and J.
Clark, B. Hawkins, and J. Longino for field assistance. Supported by a fellowship
from the MacArthur Foundation, and dedicated to Chano Pozo.
Literature Cited
Ackery, P. R. 1988. Hostplants and classification: a review of nymphalid
butterflies. Biol. J. Linn. Soc. 33: 95-203.
Ackery, P. R. & R. I. Vane- Wright. 1984. Milkweed butterflies: their cladistics and
biology. London: British Museum (Nat. Hist.), Entomology.
Boppre, M. 1984. Chemically mediated interactions between butterflies. Symp.
Roy. Ent. Soc. 11: 259-275.
Brower, L. P. 1984. Chemical defenses in butterflies. Symp. Roy. Ent Soc. Lond.
11: 110-134.
Brown, K. S. Jr. 1987. Chemistry at the Solanaceae/Ithomiinae interface. Ann.
Missouri Bot. Garden 74: 359-397.
DeVries, P. J. 1986. Hostplant records and natural history notes on Costa Rican
butterflies (Papilionidae, Pieridae & Nymphalidae). J. Res. Lep. 24: 290-333.
— — . 1987. The butterflies of Costa Rica and their natural History. Princeton
University Press, Princeton, New Jersey.
DeVries, P. J. & F. G. Stiles. 1990. Attraction of pyrrolizidine alkaloid seeking
Lepidoptera to Epidendrum paniculatum orchids. Biotropica 22: 290-297.
Drummond, B. A. & K. S. Brown Jr. 1987. Ithomiinae (Lepidoptera: Nymphalidae):
summary of known larval food plants. Ann. Missouri Bot. Gard. 74: 341-358.
Gilbert, L.E. 1983. Coevolution and mimicry. IN: D. Futuyma & M. Slatkin (eds.),
Coevolution. Sinaur, Sunderland, pp. 263-281.
Haber, W. A. 1978. Evolutionary ecology of tropical mimetic butterflies. Ph. D.
thesis, Univ. of Minn.
312
J. Res. Lepid.
Vasconcellos Neto, J. 1991. Interactions between ithomiine butterflies and
Solanaceae: feeding strategies and reproductive strategies. IN: P. W. Price, T.
M. Lewinsohn, G. W. Fernandes, & W. W. Benson (eds.), Plant-animal
interactions: evolutionary ecology in tropical and temperate regions. John
Wiley & Sons, Inc. pp. 291-313.
P.J. DeVries. Dept, of Zoology, University of Texas, Austin, Texas 78712 and
Center for Conservation Biology, Stanford University, Stanford, California
94305
Manifesto on Conservation
Editor’s note:
The Lepidopterological Society of Japan held its first seminar on the conservation
of butterflies in June 1 990. The conveners drafted, and the members present passed,
a resolution on conservation. This resolution was provided by Prof. Atuhiro Sibatani
and is published in its entirety below.
This action by the Japanese Society is a highly significant event. It makes a strong
statement on behalf of a large grassroots body of both professional and amateur
biologist members in a leading industrialized nation. It is a particularly remarkable
document given its origins in a country heretofore notorious for neglecting environ-
mental concerns in favor of immediate economic benefit. The thrust of the resolution
correctly emphasizes the need to protect total biodiversity while minimizing the
importance of individual species. We publish this as a model statement for many
circumstances.
Manifesto
A statement developed at the first seminar on The Conservation of Butterflies
as a Part of Nature, 2-3 June 1990, Osaka, co-sponsored by the Lepidopterological
Society of Japan and the Osaka Museum of Natural History.
Participants at the first seminar of the Lepidopterological Society of Japan
(LSJ) have confirmed the following statements.
• The first cause of nature conservation shall not be the protection from
extinction of individual species, but preservation of diversity in biological
communities and ecosystems.
• This means to maintain, along with persistent diversity of. individual gene
pools and species, the integrity of systems which may well entail local
alterations of these systems.
® Butterflies are pertinent bioindicators of the terrestrial ecosystem. Hence, the
conservation of butterflies implies conservation of the entire ecosystem.
• In order to fulfill the objective of nature conservation, we are responsible and
willing to work responsibly, not only to maintain nature’s biodiversity, but also
to restore this diversity in Japan as well as the world at large, under
cooperation with other bodies and using butterflies as indicators.
28(4):310-315, 1989(91)
313
Implications and further explanation
I. The Position of the LSJ
A. Since 1965, LSJ has engaged in activities for conservation of the Japanese
Lepidoptera (mainly butterflies) by setting up the Committee for Study of Nature
Conservation Issues (since 1975 the Committee for Nature Conservation) and by
issuing occasional announcements and appeals. The society also edited and
published Decline and Conservation of Butterflies in Japan I ( 1989/90). With this
publication, LSJ demonstrated in advance of other national bodies concerned
with insects, its interests in and responsibilities for the conservation of butter-
flies and shed light on some of the problems of butterfly conservation. We must
admit, however, that LSJ has lagged behind related organizations in Europe and
America. Our failure to establish reliable principles about the possible relation-
ship between butterfly conservation and collecting as well as scientific studies
thereof, may account for this lag.
B. Fortunately, along with the growth of scientists’ concerns in problems of
both domestic and foreign nature conservation, we have witnessed significant
progress in the theory of butterfly conservation which has profoundly extended
our understanding of the larger issues. We now wish to present, on the basis of
our accumulated experience and the recent theoretical advances, the following
plans for concrete action:
1. To hold, for several years, and every year if feasible, seminars,
international symposia, etc. on conservation of butterflies in order to examine
general theories of conservation as well as concrete means to counteract decline
and extinction in individual cases, and also to work out guidelines for actions and
practice.
2. To compose guidelines for butterfly collecting and investigation.
Until its completion the relevant clauses in the corresponding codes of the Royal
Entomological Society (U.K., 1968) and the Lepidopterists’ Society (U.S.A., 1982)
should be consulted for ethical norms.
3. To compile, as soon as possible, estimations of the danger and threat
of extinction for all the butterfly species of Japan and suggestions for concrete
actions to be taken for their protection and, where applicable, eventual local
reestablishments.
C. Although opinions expressed herein were supported at the first seminar of
LSJ, they do not necessarily represent opinions of the Society as a whole.
Endeavors will be made to have this manifesto endorsed by the Society.
II. Scientific Understanding of Butterfly Conservation.
A. Butterflies have a high reproductive capacity, generally undergoing large
fluctuations of population size in nature, but being capable of recovery from
serious declines of population density. Ordinary, disciplined, modest collecting
does not threaten sustained survival of any butterfly populations unless their
habitats are destroyed or disturbed for other reasons.
B. The concept that butterflies can be used as significant indicator organisms
for terrestrial ecosystems has been adopted in many countries. We also agree
with this attitude. Today butterfly conservation does not simply imply butterfly
preservation alone, but has become an indispensable means to maintain the
persisting diversity of terrestrial ecosystems at large.
C. The decline and extinction of many butterfly populations, as witnessed in
recent years in Japan, has largely been the consequence of the loss of habitat
314
J. Res. Lepid.
caused by the recent rapid and profound structural alteration of industry and the
extensive development which occurred in parallel with it. Fortunately, there has
been no record of extinction of butterfly taxa (species and subspecies) in Japan
yet. However, danger of complete extinction may be imminent for some taxa.
D. At present, the most endangered species are those which inhabit rural
modified environments closely situated to human residences.
E. Because of these circumstances as well as for the reasons described in C, we
wish to point out that the policy of butterfly conservation as adopted by the state
and local governments in Japan and many other countries, comprising prohibi-
tion of collecting at either specific or population level, with occasional inclusion
of protection of the habitat, has not properly served the newly defined purpose of
conserving the persistent diversity of butterflies or ecosystems.
F. Butterflies are components of diverse ecosystems. Reasons for their decline
and extinction are complex and far from uniform, varying from species to species
and even from population to population within one species. Protecting butterflies
from extinction accordingly requires scientific analyses at various levels, for
which continued training of young butterfly workers, capable of undertaking
scientific surveys in the field is needed, as well as education of the general public.
III. Nature Conservation in General
A. In natural ecosystems species diversity is generated and maintained by
interactions among numerous and complex components. While individual
ecosystems give rise to persistent diversity, they undergo perpetual alterations
to their component species and the size of their populations. Moreover, it is these
alterations which provide the mechanism enabling the maintenance of diversity.
For this reason, the temporary conservation of a particular species within a small
area is sometimes incompatible with securing species diversity in an ecosystem.
Since ecosystems are usually undergoing a process of perpetual change and
transition, human interference is necessary to keep them “stable” or constant.
B. Traditional agri- and silviculture and natural disasters are two main
external causes of “sound” changes in ecosystems. They do this by interrupting
vegetational succession at various stages resulting in rejuvenation, therein
guaranteeing species diversity. Thus, before the advent of modern civilization,
traditional society and culture were integral parts of the mechanism by which
diversity of the ecosystem was perpetuated. Today the situation is completely
changed. New industrial structures have either destroyed the natural environ-
ment or affected its simplification with the calamitous loss of factors which
generated the previously extant diversity. The recent impoverishment of the
butterfly fauna reflects this very well.
C. A highly desirable conservation policy would not consist of promoting a list
of species whose collection is formally prohibited, but would aim at providing
safety for all the butterfly species occurring in individual regions and localities.
However, since natural environment is in constant transition, such a policy
should not imply an unnatural fossilization of the current situation, but should
aim at instituting both natural and cultural mechanisms to enhance biodiversity
through modest man-made (or human-sized) interference with the system. The
usual practice of environment impact assessments which refer to the status quo
as a criterion for preservation, does not serve the objective of nature conservation.
Official bodies and grass-roots movements active in nature conservation are at
a turning point where they should reappraise the nature of the problems they are
faced with in order to achieve the goals to which they aspire.
28(4):310-315, 1989(91)
315
An Attractant for Zerene eurydice (Pieridae)?
During the summer of 1981, while living in an apartment near the campus of
California Polytechnic State University, San Luis Obispo, I observed unusual
behavior of Zerene eurydice (Boisduval) toward a hedge of the cultivated shrub
called xylosma, Xylosma congestum (Lour) Merrill (Flacourtiaceae). Through-
out the summer, especially during August, males and females of Z. eurydice were
commonly seen in westward flight which took them over or around the hedge of
xylosma. One day the hedge was trimmed to shape by gardeners and the
clippings were left on the ground next to it. During the next two weeks or so after
the trimming, individuals of both sexes of Z. eurydice would approach the hedge
in their normal manner, but once within about one meter from the shrub they
would drop down to alight on the clippings. None were noticed to extend their
proboscis or move any part of their body; they remained motionless with wings
folded as if basking in some welcome scent. If allowed to remain undisturbed, the
butterflies would return to flight after about 3 minutes with no apparent effects.
Attempts to approach the butterflies startled them and they took flight. No less
than eight individuals displaying the behavior were counted and recorded; others
were casually noticed, but were neither recorded nor captured. This behavior
ended about two weeks after the hedge was trimmed, possibly because of
evaporation or decomposition of compounds within the xylosma clippings.
The cause of such behavior is a mystery, but perhaps involves attractance to
chemical compounds within the plant; xylosma gives off a characteristic odor
especially after it has been trimmed. Two other reported attractants for Z.
eurydice are purple flowers (Emmel, T.C. & J.F. Emmel, 1973, The butterflies of
southern California, Natural History Museum of Los Angeles County, Science
Series 26: 1-148. See page 20) and fresh horse manure (Garth, J.S. & J.W. Tilden,
1986, California butterflies, California Natural History Guides: 51, U.C. Press,
Berkeley. Seepage 110). Once isolated, the chemical(s) in Xylosma might prove
to be a worthwhile attractant for Z. eurydice; more reliable than finding purple
flowers and less offensive than horse manure!
Robert L. Allen, Museum of Systematic Biology, University of California,
Irvine, CA 92717.
28(4):316-318, 1989(91)
Book Reviews
—
BUTTERFLIES OF EUROPE. Vol. 2. INTRODUCTION TO LEPID-
OPTEROLOGY. 1990. Otakar Kudrna (ed.). AULA-Verlag, Wiesbaden, 557 pp.,
93 figs., 4 col. pis., 25 tables, 2 diagrams. ISBN 3-89104-033-4. Available from:
AULA-Verlag GmbH, Postfach 1366, Luisenplatz 2, 6200 Wiesbaden 1, Ger-
many, Price: 248 DM.
This is the third volume to appear in the eight volume series Butterflies of
Europe and is intended as a text and reference book for advanced students.
Ch. 1 is a short introduction by O. Kudrna. Lepidopterology is the study of
Lepidoptera from different viewpoints such as taxonomy, ecology, physiology,
cytology, behavior, etc.
Ch. 2, Lepidopterology in Europe, by Kudrna and M. Wiemers, is an address
guide to major European museums, institutes, and societies possessing impor-
tant collections and libraries. BMNH has the largest butterfly collection and the
best entomological library in the world. Also included are current European
Lepidoptera journals and a “Who’s Who” of over 300 “great” European lepidop-
terists, complete with obituary references and collection depositions.
Ch. 3. early stages, by J. P. Brock gives general characteristics, keys to families
for larvae and pupae, detailed line drawings of morphology, and a brief account
on preserving early stages.
Ch. 4, adult structure and function, by J. A. Scott, discusses internal and
external morphology with the aid of numerous labelled drawings. It also covers
breathing and blood circulation; feeding, digestion and excretion; and nervous,
sensory, and endocrine systems.
Ch. 5. butterfly phylogeny and fossils, by Scott and D. M. Wright, reviews
chemical-genetic, intuitive, phenetic, biological species, and cladistic methods of
phylogeny study. There is a good discussion of cladistic methods which they apply
in the remainder of the chapter. Sections list synapomorphies for Pyraloidea-
Macrolepidoptera, Macrolepidoptera, Hedyloidea-Hesperioidea-Papilionoidea,
and each Rhopalocera group at the family, subfamily, and sometimes tribal
levels. The outgroup for Hedyl.-Hesp.-Pap. is not specified but is probably most
Ditrysian moths. They accept Hedylidae as the sister-group to all butterflies. A
cladogram of subfamilies and some tribes is presented, calculated by hand from
the character data, though it is difficult to see how the most parsimonious choice
was made given the large number of taxa (cf. Felsenstein, 1978, Syst. Zool. 27 : 27-
33). Showing the distribution of the specific characters on the cladogram would
have clarified the analysis for the reader (cf. de Jong, 1983, Zoologische
Mededelingen 57: 243-270; Nielsen, 1987, Invertebr. Taxon. 1: 201-229). Their
data also should be run on a computer for more objective results. In their view,
the frenulum of Euschemon is the expression of a suppressed gene rather than
an indicator of primitiveness. Also, they show Lycaenidae evolving from the
ancestor of Nymphalidae via ancestral Riodinidae, implying foreleg reduction
reverted to a normal foreleg condition in Lycaenidae. It is unclear how they
determined that the listed synapomorphies only apply to each group in question
and are absent in all the remaining groups. This is the most comprehensive
cladistic analysis of butterflies yet attempted, but butterfly fossils are all too
briefly discussed in less than a page and are not updated since the late 1970’s.
28(4):316-318, 1989(91)
317
Ch. 6. origins and phylogeny of butterflies, by Brock, considers phenetics and
cladistics as “bogus” and heavily criticizes Scott’s work. Cladistics is not without
controversy, but personal vendettas have no business being aired in an introduc-
tory text. He discusses character correlations and notes that characters used in
higher level classification are largely stabilized by canalization and that paral-
lelism has introduced confusion into Lepidoptera phylogeny schemes. He dis-
cusses butterfly characters in relation to those of moths and from this limited set
of selected characters arrives at conflicting conclusions: that butterflies arose
from Microlepidoptera and Hedylidae by monophyly! However, butterflies may
be diphyletic (Shields, 1989, Tyo to Ga 40: 197-228) or even triphyletic if
Hedylidae is another entry-point. There is some discussion of only two other
proposed theories: ancestry from Pyraloidea and cossid-castniid. I believe he is
correct in questioning Scott’s papilionid-pierid eleven “shared” traits. Numerous
differences between papilionids and pierids (e.g. egg, pterin array, osmaterium,
aorta, Baronia immatures unlike pierids, etc.) should not be ignored even though
they are autapomorphies since they add up to major differences.
Ch. 7, genetics of European butterflies, by R. Robinson, discusses inheritance,
epistasis and hypostasis, genes, multiple alleles, inviability and impenetrance,
gene linkage, polymorphism, electrophoretic and quantitative variation, cytoge-
netics, chromosome number variation, chiasma, centromeres, sex chromosomes,
sex chromatin, and supernumerary chromosomes, with a section on genetics of
selected species. A table lists haploid chromosome numbers that are known for
ca. 60% of European butterfly species with references. The text, however, is
difficult to follow for the non-devotee (such as myself).
Ch. 8, case studies in ecological genetics, by P. M. Brakefleld, defines ecological
genetics as “an integrated study of ecology and genetics concerned with under-
standing evolutionary processes,” e.g. industrial melanism. Case studies include
eyespots in M aniola jurtina and Coenonympha tullia. He also describes methods,
field surveys, and genitalia variation.
Ch. 9, butterfly chromosomes and their application in systematics and phylog-
eny, by Z. Lorkovic, is based mainly on the classical paraffin-cut method which
requires little lab assistance. Special attention is devoted to illustrations, mainly
drawings, of the numbers, sizes, etc. of chromosomes and their significance in
phylogeny. Discussions include spermatogenesis, oogenesis, fission and fusion
of chromosomes, supernumerary chromosomes, individual variation of chromo-
somes, behavior of chromosomes in hybrids, and significance of karyotypes for
taxonomy and phylogeny, along with a section on procedures, methods, and
techniques for chromosome number frequency for European butterflies. This
chapter is clearly and concisely written — a model summary. He suggests that
chromosome numbers do not support an intimate relationship of Satyridae with
Nymphalidae and Riodinidae with Lycaenidae. There are two color plates of
hybrid pierids.
Ch. 10, enzyme electrophoretic methods, by H. Geiger, summarizes electro-
phoresis methods as applied to butterflies. Electrophoresis is not for amateurs
since a well-equipped lab and special training are mandatory. Electrophoresis
is a useful tool for analyzing problems in systematics and evolutionary biology.
He also discusses analysis of enzyme electrophoretic data, steps in drawing
dendrograms, relevant mathematics, and the method’s disadvantages.
Ch. 11, experimental breeding of butterflies, by S. R. Bowden, notes that
experiments can determine if varieties are due to environment or are genetic. He
covers hybridization, fertility, pairing cages, growing nectar flowers, larval
318
<7. Res. Lepid .
housekeeping, breeding synchronization, sex-ratio, and record keeping. Fresh
material is often needed in electrophoresis and chromatography. There are two
color plates of genetic forms of the P ieris napi group. This chapter is quite brief
and is highly critical of cladistics.
Ch. 12, parasitoids, by M. R. Shaw, covers Hymenoptera and Diptera that
attack the early stages of butterflies. He notes they are little studied and that
misidentifications abound. He also discusses parasitoid biology, difficulties
encountered in their study, an outline of the principle groups, collecting and
rearing techniques, labelling, and transport. Their host associations can be
phylogenetic or ecological in nature.
Ch. 13, butterfly behavior, by T. G. Shreeve, discusses thermoregulation, mate-
location, mate-recognition, egg-laying, and feeding and their methods of study,
though no mention is made of such topics as roosting, fear flight, etc.
Ch. 14, butterfly movements, by Shreeve, is divided into movement, dispersal,
and migration and is often theoretical. Books devoted to migration are not
mentioned.
Chapters 3, 4, 7, and 12 have glossaries of terms and there are many useful
references cited in most chapters. The book begins with a comprehensive table
of contents, ends with an index to scientific names, general index, and author
addresses, and is well-organized. By-and-large, its multi-disciplined approach is
an excellent introduction to the study of butterflies and fills a previous void with
its publication.
Oakley Shields, 6506 Jerseydale Road, Mariposa, CA 95338, USA.
THE BUTTERFLIES OF EGYPT. Torben B. Larsen. 1990. Apollo Books,
Lundbyvej 36, DK-5700 Svendborg, Denmark, 112 pp. (Available from the
publisher for 240 Danish kroner + postage.)
Torben Larsen has systematically documented the butterfly fauna of much of
the Near East, beginning with his classic The Butterflies of Lebanon ( 1974). This
latest contribution presents the small fauna (58 species, fewer than recorded in
most California counties) of a large, but very arid, country. Although the total
documentation is much less than that available for some other regional faunas
published by Larsen, he does a thorough and painstaking job as usual. The
habitats and the butterflies themselves are illustrated in handsome and well-
produced color plates. There are the usual Larsen treatments of biogeography,
faunistics, migration and life-history strategies. The format is meant to match
the Lebanon book.
At this writing, there are about 6.4 kroner to the dollar. This comes out to about
$37.50, or 33e/page. In other words, this is not a cheap book. It will be
indispensable for regional specialists, but something of a luxury for the average
collector. For those interested in life-history evolution, adaptation to seasonality,
or community composition, all of Larsen’s work provides raw material for the
study of functional convergence. This is no small contribution.
Arthur M. Shapiro, Department of Zoology, University of California, Davis,
CA 95616, USA.
INSTRUCTIONS TO AUTHORS
Manuscript format: Two copies must be submitted, double-spaced, typed, with wide
margins. Number all pages consecutively. If possible italicize rather than underline scientific
names and emphasized words. Footnotes are discouraged. Do not hyphenate words at the right
margin. All measurements must be metric. Time must be cited on a 24-hour basis, standard
time. Abbreviations must follow common usage. Dates should be cited as: day-Arabic numeral;
month-Roman numeral; year-Arabic numeral (ex. 6. IV. 1984). Numerals must be used for ten
and greater e.g. nine butterflies, 12 moths.
Electronic submission: The Journal is now being produced via desktop publishing, allowing
much shorter publication times. Although typewritten manuscripts are acceptable, those
submitted on computer disc are highly preferred. After being notified of your paper's
acceptance, submit either a Macintosh or IBM disc version. Include on your disc both the fully
formatted copy from your word processing program and a text-only (ASCII) copy. The two most
preferred formats are Microsoft Word for the Macintosh and either Microsoft Word or Word
Perfect for the IBM, although translation utilities will allow conversion from most formats. Put
returns only at the ends of paragraphs, not at the end of each line. Use one tab to indent each
paragraph. Even if your printer is incapable of outputting italics, please specify italics rather
than underline in your disc copy. Please note any special characters that are used in either the
body of the text or the tables (e.g. e, u, °, §, p., <J , 9 ).
Title material: All papers must include the title, author's name, author's address, and any
titular reference and institutional approval reference. A family citation must be given in
parenthesis (Lepidoptera: Hesperiidae) for referencing.
Abstracts and Short Papers: All papers exceeding three typed pages must be accompanied
by an abstract of no more than 300 words. Neither an additional summary nor key words are
required.
Name citations and Systematic Works: The first mention of any organism should include
the full scientific name with unabbreviated author and year of description. There must be
conformity to the current International Code of Zoological Nomenclature. We strongly urge
depositing of types in major museums, all type depositories must be cited.
References: All citations in the text must be alphabetically listed under Literature Cited in
the format given in recent issues. Abbreviations must conform to the World List of Scientific
Periodicals. Do not underline or italicize periodicals. If four or less references are cited, please
cite in body of text not in Literature Cited. For multiple citations by the same author(s), use
six hyphens rather than repeating the author’s name.
Tables: When formulating tables, keep in mind that the final table will fill a maximum space
of 11.5 by 19 cm either horizontally or vertically oriented. Number tables with Arabic
numerals. When submitting tables on disc, use tabs between columns rather than multiple
spaces.
Illustrations: Color can be submitted as either a transparency or print, the quality of which
is critical. Black and white photographs should be submitted on glossy paper, and, as with line
drawings, must be mounted on stiff white cardboard. Authors must plan on illustrations for
reduction to page size. Allowance should be made for legends beneath, unless many consecutive
pages are used. Drawings should be in India ink. Include a metric scale. Each figure should
be cited and explained as such. Each illustration must be identified by author and title on the
back. Indicate whether you want the illustration returned at your expense. Retain original
illustrations until paper is accepted. Legends should be separately typed on pages entitled
"Explanation of Figures." Number legends consecutively with separate paragraph for each
page of illustration.
Review: All papers will be read by the editor(s) & submitted for formal review to two referees.
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
Volume 28 Number 4 Winter 1989(1991)
IN THIS ISSUE
Date of Publication: September 15, 1991
European and North West African Lycaenidae 239
(Lepidoptera) and their associations with ants
Konrad Fiedler
Detecting and recording the calls produced by 258
butterfly caterpillars and ants
P, J. DeVries
Genetics and Biogeography of the Oeneis chryxus 263
Complex in California
Adam H. Porter and Arthur M. Shapiro
A new Polythrix from Central America 277
(Lepidoptera: Hesperiidae)
John A. Shuey
Three unusual species of Parades from South America 283
(Lepidoptera: Arctiidae)
Vitor O. Becker and Scott E. Miller
Selection of Lepidopterologically Interesting Areas 289
in Central Spain Using UTM Distribution Maps
J. L. Viejo, C. de Silva, C. Ibero and J. Martin
An unrecognized, now extinct, Los Angeles area 297
butterfly (Lycaenidae)
Rudolf H. T. Mattoni
Notes
310
Book Reviews
316
Cover Illustration: Collage of Philotes sonorensis sonorensis in flight.
From cover illustration of R. Mattoni, Butterflies of Greater Los Angeles , 1990.
Original photograph by Greg Ballmer. ml
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