Taxonomic significance of reflective patterns in the

JOURNAL
OF
THE LEPIDOPTERISTS'
Volume 27
SOCIETY
1973
Number 3
TAXONOMIC SIGNIFICANCE OF REFLECTIVE PATTERNS
IN THE COMPOUND EYE OF LIVE BUTTERFLIES: A
SYNTHESIS OF OBSERVATIONS MADE ON SPECIES
FROM JAPAN, TAIWAN, PAPUA NEW GUINEA
AND AUSTRALIA
ATUHIRO SIBATANI
30 Owen St., Lindfield, New South Wales 2070, Australia
During observations in the field of New South Wales, I came to notice
that some Australian Lycaenidae had unusual semi-transparent and sometimes brightly coloured eyes which I had not come across before in some
other parts of the world, including temperate and tropical Eurasia and
America. This character could be observed only in live or recently killed
butterflies. The regular occurrence of this type of eye in certain lycaenid
groups strongly suggested its taxonomic usefulness. Upon extending
my observation to other butterfly families, I soon realised that in such
semi-transparent eyes there were usually certain reflective spots which
changed their position according to the direction of observation, and
that these spots were observed almost invariably in Pieridae and Nymphalidae (s.str.), but not in Papilionidae and Hesperiidae, and variably
in Satyridae, Danaidae and Lycaenidae. Moreover, the pattem of these
spots also appeared to be of taxonomic significance. During the past
two years I have thus accumulated records of my own observations on
the superficial feature of the eye in butterfly species occurring in New
South Wales and Papua New Guinea.
Meanwhile, my attention was drawn to the extensive monograph, "The
Compound Eye of Lepidoptera," by Yagi and Koyama (1963). In this
work the authors not only recorded the pattem of reflective spots in
fresh eyes for the majority of butterfly species in Japan and many species
from Taiwan, but also correlated them to the histologic structure of the
ommatidium and thus clarified the optical basis of the appearance of
these spots. Although they gave a number of suggestions about the
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taxonomy and phylogeny of butterflies as well as other Lepidoptera
families based on their studies on the compound eye of Lepidoptera
at large, I feel that this extensive and important work treating the subject
in depth has so far failed to have an impact on the taxonomy of butterflies.
In order to pay due tribute to the work, and to fill the lacuna of
knowledge and interest in this aspect of butterfly morphology and
taxonomy among lepidopterists, I wish to briefly redescribe the superficial (but not histologic) features of the butterfly eye only to the extent
useful to careful field workers, on the basis of records compiled by
Yagi & Koyama (1963) as well as my own observations. The material
so collated covers species from Japan, Taiwan, Papua New Guinea and
Australia, thus representing some of the major butterfly groups in the
Palaearctic, Indo-Malayan and Australian Regions. The limitation of
this character as a taxonomic criterion is the fact that the eye must be
observed soon after an insect is captured, but this is certainly much less
demanding than is the knowledge of foodplants and early stages for
inclusion in taxonomic investigations. It is desired that, through the
cooperation of field workers in various countries, patterns of live butterfly eyes will soon be recorded for the groups not included in this
paper.
General Description and Terminology
The superficial appearances of butterfly eyes may be divided into two
major types. One is the eye looking totally dark; the other is the eye
looking semi-transparent, lightly or brightly coloured and bearing at
least one, and usually seven or more, dark spots which change their
position and sometimes their shape when observed from different directions.
The first type is observed in Papilionidae and Hesperiidae in which
the eye may reflect strong incident lights in the form of a hexagonal
bright central area. The images of such an eye are well recorded in
some photographs of butterflies taken in the field, especially those of
papilionid species: Papilio aegeus aegeus Donovan (Deger & Eden,
1970, p. 5, fig. 4); Papilio protenor demetrius Cramer ( Kohiyama,
Takase & Fujioka, 1971, fig. 5); Papilio demoleus sthenelus Macleay
(D'Abrera, 1971, p. 41). The hexagonal spot may be observed in the
eye of dead and dried specimens, suggesting that it is mainly due to
surface reflection, and thus is not the subject of this paper. In some
Satyridae and most Lycaenidae, the eye looks completely dark without
any hexagonal bright spot. An example may be seen in the photograph
of Jalmenus eichorni (sic) Staudinger given on p. 3 of Deger & Eden
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Fig. 1. A live eye of Pieris rapae Linnaeus, enlarged from a part of the original
photograph of Deger & Eden (1970), and reproduced through the courtesy of the
authors and publisher.
(1970), and of Taraka hamada Druce given in fig. 47 of Kohiyama et al.
(1971).
A good illustration of the second type of eye may be found in the
photograph of Pieris rapae (Linnaeus) shown in fig. 3 on p. 9 of Deger
& Eden (1970), of which I reproduce a part as Fig. 1 by courtesy of
these authors. As seen in the original photograph printed in colour, the
eye itself is pale greenish grey with a prominent dark central spot which
is surrounded by six conspicuous primary side spots arranged in a hexagonal array. Around these primary side spots, there are vague dark
shadows in the form of some discrete patches, which represent the
secondary side spots, which are rather ill-defined in this species.
These spots were called by Yagi & Koyama (1963) the "pseudopupils."
The name should have been phrased the "false pupils" to conform with
a uniform Latin derivation rather than a mixture of Greek and Latin
ones. However, the word "pupil" even with the adjective "false" sounds
inadequate because of its possible interpretational bias. The more
descriptive expression "reflective spots" is being used almost as a jargon
among Drosophia geneticists for a pair of apparently similar spots in
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the eye of the wild-type individuals of Drosophila melanogaster Meigen.
I want to borrow this expression in place of the "pseudopupil," since
the latter has not yet been established among lepidopterists, and will
abbreviate it as RS in the rest of this paper.
Main Types and Subtypes of the Compound Eye
According to Yagi & Koyama (1963) the eye with RS may be divided
into two major types. One has a large central spot only. In the other the
central spot is smaller and usually accompanied by primary side spots
which however may become very faint, and frequently by the secondary
side spots which may surround strongly marked primary side spots.
The nature of the appearance of these RS has not been formulated
in exact physical or mathematical terms, but is correlated by Yagi &
Koyama (1963) with an at least partial absence of pigment around the
distal half of the ommatidium, allowing the oblique incident light to
pass through one ommatidium into another. The distribution and extent
of pigmentation seem to be related to the variation of the patterns of
RS, but I will not deal with this aspect here. When the butterfly dies,
the appearance of RS and the semi-transparency of the eye are progressively lost hand-in-hand, suggesting a change in the refractive index of
the substance filling the ommatidium. In the dark eye without RS, the
ommatidia appear to be optically isolated from one another by a predominant pigmentation along the septa of each ommatidium.
In the terminology of Yagi & Koyama (1963) the dark eye without RS,
the light eye having only one (large) central RS, and the light eye
having basically seven or more RS were called the non-pseudopupil
type, mono-pseudopupil type, and multi-pseudopupil type, respectively.
In this paper I will simply call them Type I, II and III eye, respectively.
If necessary, they may be called the eye with no RS, single RS and
multiple RS. I have never seen a Type II eye myself and its superficial
character is somewhat obscure to me. In Fig. 2 I have assembled
diagrammatic representations of these three basic types, and modifications (or subtypes) of the Type III.
In the subtype IlIa, the side spots are very obscure. The primary side
spots are conspicuous in IlIb; their position may be close to the central
spot or to the periphery of the eye. This last exampJe may be seen
in the photograph of Polyura pyrrhus sempronius (Fabricius) on p.
25 of D' Abrera (1971). In IIlc, the secondary side spots are apparent.
Sometimes the intensity of all the spots becomes more uniform, giving the
eye a mesh-like appearance. In IUd, radial striations appear, especially
around the central spot, and each RS may take a hexagonal shape. In
IIIe, individual RS's take a rectangular shape and tend to be confluent.
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I
II
rna
mb
me
md
me
IIIf
IIIg
IIIh
Fig. 2. Diagrammatic representation of types and subtypes of the reflective eye
pattern in butterflies. Modified from Yagi & Koyama (1963) except for HIe and h
which are original, through the courtesy of Prof. N. Koyama.
IIIf is characterised by dark areas intercalating the RS and tending to
connect one another to circumscribe the latter. IIIg represents a very
conspicuous type in which the RS's are longitudinally united to form a
striated pattern in the eye. IIlh is the most ill-defined pattern; here the
eye looks dark and the various modifications described in IIlc-g seem
to appear to various extents in combination.
Table 1 lists the distribution of the major types and some less frequently encountered types in each of the butterfly families! occurring
in the areas covered, except for Riodinidae for which no observation
has been available. Assignments of species to individual types and subtypes are compiled in the last section of this paper.
Intraspecific Variation
Repeated observations on different individuals of the same species
have given consistent results in most cases, but there does seem to be
some intraspecific variation in the appearance of the live eye. The
variation may fall into several categories: 1. individual variation; 2.
1 The higher classification of butterflies is in a state of confusion at the moment.
Almost
every author seems to have his or her own system of classification. The most logical approach
to the problem may be that of Ehrlich (1958) but his system has not been followed by the
majority of contemporary workers in the world. There is too little room for intel'mediate
classification in some of the Ehrlich's families which lumped together a number of distinct
taxonomic groups. Here I have followed the more conventional classification for that reason.
However, I admit that the standard of my recognising individual families is more arbitrary than
scientific. For definition of individual families I followed Miller (1968) for Satyridae, Fox
( 1956) for Ithomiidae, and Eliot (pers. comm.) for Lycaenidae. Also the recent trend of
splitting genera into smaller units is more (or too) conspicuous for European and Japanese
fauna than Australian and New Guinean ones, and this necessarily caused unevenness in
recognising individual genera.
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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY
TABLE l. Distribution of eye types among various families of butterflies (0 ,
common, predominant types; and . , unusual types, in individual families).
II
Family
III
a
Hesperiidae
Papilionidae
Pieridae
Danaidae
Iithomiidae
Amathusiidae
Satyridae
Nymphalidae
Acraeidae
Libytheidae
Lycaenidae
b
c
0
0
d
e
g
h
0
0
0
0
••
0
0
0
0
• •
0
• 0•
0
0
0
• • •
0
• •
• •
0
0
•
d evelopmental change taking place soon after emergence of the adult;
3. sexual dimorphism; 4. subspecific or seasonal difference; and 5.
genetic polymorphism in one population. At the present stage of investigation it is still difficult to make a distinction in this sense among
some of the observed examples of intraspecific variation. In the following
I give some examples worth noting.
1. Danaus chrysippus petilia Stoll (Danaidae) males caught at one
place (New South Wales) at the same time showed two subtypes of the
Type III eye: e and h. Slight variation between a and b, band c, or c
and g may be encountered among a number of species having the Type
III eye.
2. Yagi & Koyama (1963) noted that the eye of Aporia Hubner
(Pieridae) turns from Type IIIf to Type I soon after emergence, obviously
owing to a colour development around ommatidia during that period.
3. A difference in the eye pattern between the two sexes was observed
in two lowland species of Heteronympha Wallengren (Satyridae) in
New South Wales: H. merope merope Fabricius and H. mirifica Butler,
in which males showed Illb or IIIb-g eyes and females typical IlIg
eyes. Another example was recognised in Ogyris amaryllis amaryllis
Hewitson (Lycaenidae) in which males showed IIlg eyes while a single
female specimen had IlIa eyes. Males of some other species of Ogyris
Westwood have IIlg eyes like the male of O. amaryllis, but I have not
yet examined females of these other species. My record includes a
somewhat doubtful case of I-Iypolimnas alimena eremita Butler (Nymphalidae): male, IIle; female, Illd-but this needs confirmation. In
most of the observations I have made, I have not recorded the sex of
the observed specimens explicitly, and it is possible that there are sexual
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dimorphisms among the species recorded without reference to sex in
this paper.
4. So far I have not come across any definite proof that subspecies
or seasonal forms of one and the same species can differ in the reflective
pattern of the eye. Nor do I know of any evidence for a genetic polymorphism related to this character among butterflies. However, in
Drosophila melanogaster (Diptera) the somewhat similar RS's are characteristic of wild-type eyes and are absent from all the known eye-colour
mutants.
A Brief Description of the Eye Pattern in Various Taxonomic
Groups and Evaluation of its Taxonomic Significance
Hesperiidae. All the examined species covering Coeliadinae, Pyrginae,
Trapezitinae and Hesperiinae have similar Type I eyes, so that this
eye pattern can be regarded as an invariable character of the family. The
pyrgine genus Chaetocneme Felder has red instead of dark brown or
black eyes whose colour is stable post mortem. It would be of interest
to know whether or not the eye of this genus follows the general pattern
of the family.
Papilionidae. All the genera so far examined and reported, covering
Parnassiinae (Zerynthiini and Parnassiini) and Papilioninae (Graphiini,
Papilionini and Troidini) show Type I eyes, so that this eye pattern
can be taken as a universal character of the family.
Pieridae. The family is characterised by the Type III eye. A remarkable exception is the genus Aporia Hubner having Type I eyes. Another
interesting point is the appearance of IIIf eyes in Gonepteryx Leach and
Ixias Hubner (Yagi & Koyama, 1963). The colour of the pierine eye
ranges from greenish yellow (Colias Fabricius and Eurema Hubner)
to bluish white (Prioneris Wallace), via the commonest pale greenish
or whitish grey of most of the genera including Catopsilia Hubner,
Pieris Schrank and Delias Hubner. The major subtypes of the eye are
IlIa, band c.
Danaidae. This family present certain difficult problems. Eyes of
many species are dark but still usually some dark and light patterns are
discernible. The diagram shown in Fig. 2 IIIh is an interpretation of
the complicated pattern encountered frequently in this family. Yagi
& Koyama (1963) noted that certain species of Euploea Fabricius in
Taiwan had eyes without RS. I also noted the same with several Euploea
species I collected in New Guinea. However, some specimens of certain
species in New Guinea or New South Wales had patterns like IlIg
eyes. I am not certain at the moment whether the variability of the
eye pattern in this group is at least partly due to the sexual dimorphism.
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TABLE 2.
Satyridae.
Distribution of Type I and III eyes among some higher taxa of
Taxa
BIINAE
ELYMNIINAE
Lethini
Lethe-Series
Pararge-Series
Mycalesini
SATYRINAE
Hypocystini
H ypocysta-Series
Tisiphone-Series*
Coenonymphini
Erebiini
Satyrini
Satyrus-Series
Oeneis-Series
Type I
Type III
Melanitis
Lethe europa, L. chandica,
Ninguta, Neope
Lopinga
Lethe marginalis, L. diana, L.
sicelis, Harima
Lasiommata, Kirinia
Mycalesis, Orsotriaena
Hypocysta
Harsiesis, Platypthima,
Pieridopsis
Geitoneura, Heteronympha
merope, H. mirifica
Argynnina, Oreixenica,
Heteronympha penelope,
H. paradelpha, H. banksii,
H. cordace, Tisiphone
Coenonympha
Erebia
Minois
Oeneis
* Miller (1968) called this the Xenica-Series, but Xenica W estwood, 1851, is an objective
synonym of Tisiphone Hubner, 1816-1826 (Hemming, 1967).
Working in the field it was not always possible to compare live males
and females of the same species. Also the specific identification of individual specimens of Euploea in the field posed some problems in New
Guinea. Obviously much more careful and extensive observations on
many individuals are needed before something more definite can be
stated about the eye of this difficult family.
Ithomiidae. The only known example (Tellervo Kirby) has subtype IIIb eyes with a vivid yellow ground colour. The eye of this family
therefore seems to be at variance with that of Danaidae or Satyridae.
Amathusiidae. The only known example (Taenaris Hi.ibner) showed
Type I eyes.
Satyridae. The majority of the species of this family have eyes of
Type IIIg with a grey or grey-brown, but sometimes yellow, ground
colour, but there occur a number of exceptions. A glance at Table 2
reveals that the difference is not apparently related to any taxonomic
groupings, because both Type I and III eyes are found side by side in
many recognised higher taxa, and even within a single genus, almost
throughout the family. However, in most cases the eye pattern may be
regarded as a good character of individual genera, especially in Hypocystini.
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Also noteworthy is a consistent difference between the two sexes as
exemplified by the two lowland species of Heteronympha (see above).
One theory might be that the difference in the eye pattern is more adaptive than phylogenetic, but nothing is known about the possible correlation of certain eye types with the habitat or some other ecological
or behavioral variables.
Nymphalidae. Most species of this family have eyes of Type III,
but there is an odd example of Type I (Clossiana thore jezoensis Matsumura). I have a record of a live female of Hypolimnas deois divina
Fruhstorfer having Type I eyes, while another record of mine indicates
that the same species (male ?) had subtype IIIb eyes. Some species
(Dichorragia nesimachus nesiotes Fruhstorfer and H estina a8similis
formosana Moore) are assigned to Type II by Yagi & Koyama (1963).
Generally, the subtype IlIc predominates, but subtype IlId eyes are
widely seen among members of Nymphalinae (s.str.). The subtype IlIa
appears in Apaturinae. Many species have brightly coloured eyes:
yellow or orange predominates in Argynninae; grey or brown in Nymphalinae; bluish grey in Limenitinae; light orange or yellow-brown in
Apaturinae; green in some species of Cyrestis Boisduval, and bluish black
in Dichorragia Butler. In Charaxinae, Charaxes latona papuensis Butler
has bright orange eyes, while Polyura pyrrhus sempronius Fabricius
shows Type IIlb eyes which look almost as dark as Type I eyes.
Acraeidae. As far as known, the eye belongs to Type 1. Perhaps this
character may justify the family status of this group.
Libytheidae. The only known case is the subtype IlIf.
Lycaenidae. As noted by Yagi & Koyama (1963), the majority of the
species have Type I eyes. However, there are some genera with Type
III eyes, which are for some unknown reason frequently met with in the
Australian region. This latter type is found in various subfamilies, but
its occurrence generally serves as a criterion at least for a genus, sometimes for even a higher taxon.
Thus, in Polyommatinae, Zizula Chapman, Zizina Chapman and
Zizeeria Chapman (including the very anomalous Z. alsulus HerrichSchaeffer), which are somewhat remotely related, have yellowish grey
eyes of the subtype IlIa-b. Theclinesthes Rober and Catopyrops
Toxopeus stand out with similar yellowish grey eyes among the genera
related to Nacaduba Moore. Two species currently placed in Neolucia
Waterhouse and Turner: serpentata Herrich-Schaeffer and sulpitius
Miskin also have eyes similar to those of Theclinesthes, while Neolucia
agricola Westwood and N. mathewi Miskin have Type I eyes. I have
found that other features such as male genitalia and antennae of serpentata and sulpitius showed a very strong affinity to those of
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Theclinesthes and were unlike any other groups of Lycaenidae I know of,
let alone Neolucia agricola and N. mathewi. Obviously there is a need
of generic rearrangement for some species hitherto placed in Theclinesthes
and N eolucia, and the eye pattern is consistent with many other characters
with respect to this point. Among genera allied to Candalides Hubner
with Type I eyes, Erina Swainson is peculiar with Type III eyes, which
seems to point to a good generic status of Erina.
In Theclinae, Hypochrysops Felder & Felder and Philiris Rober have
yellow eyes belonging to subtype IlIa or b-g. Contrmy to previous
treatments by authors as recent as Common (1964), Tite (1963) or
D'Abrera (1971), the two genera are taxonomically closely related as
is seen from the wing shape, antennae and male genitalia. Eliot (pers.
comm. ) has reached the same conclusion independently. Paralucia
Waterhouse & Turner and Pseudodipsas Felder & Felder have grey subtype IlIa or b-c eyes and seem to stand fairly close to the preceding two
genera. More peculiar is Ogyris Westwood, males having characteristic
Type IIIg eyes with a grey-brown ground colour, like those of many
satyrid species, and the only female ever examined having subtype
IlIa eyes. Two species of Hypochlorosis Rober in New Guinea had
yellowish Type III eyes. According to a photograph shown by Kohiyama
et al. (1971), Spindasis takanonis Matsumura may have a brown Type
III eye, but this is not certain.
Curetinae seems to have disparate eye types in the single genus
Curetis Hubner. It would be of interest to see whether or not there
occur some clusters of genera having Type III eyes in areas other than
the Australian region.
Assignments of Various Species of Butterflies to Individual
Types and Subtypes of the Compound Eye
Assignments taken from Yagi & Koyama (1963) are marked with
an asterisk, and those inferred from the figures of Kohiyama et al. (1971)
with two asterisks.
SPECIES WITH TYPE I EYES
Hesperiidae. Chaospes benjamini japonica Murray*; Bibasis aquilina chrysaeglia
Butler*; Erynnis montanus Bremer*; Daimio tethys Mem\tries; Trapezites eliena
eliena Hewitson, T. iacchoides Waterhouse, T. phigalia phigalia Hewitson; Toxidia
peron Latreille; Signeta flammeata Butler; Mesodina halyzia halyzia Hewitson;
Leptalina unicoi or Bremer & Grey*; Aeromachus inachus Menetries*; Isoteinon
lamprospilus formosanus Fruhstorfer*; Notocrypta curvifascia C. & R. Felder*, N.
waigensis waigensis Plotz; Thymelicus leoninus Butler*, T. sylvaticus Bremer*;
Ochlodes venata herculea Butler* , O. ochracea rikuchina Bulter*; Hesperia Florinda
Butler*; Potanthus flavus Murray*; Telicota eurotas eurychlora Lower; Cephrenes
augiades sperthias Felder; Polytremis pellucida Murray"; Pelopidas jansonis Butler*,
P. mathias oberthiiri Evans*; Parnara guttata Bremer & Grey*.
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Papilionidae. Pamassius eversmanni daisetsuzanus Matsumura*, P. stubbendorfii
hoenei Schweitzer*, P. glacialis Butler*; Luehdorfia puziloi inexpecta Sheljuzhko*,
L. japonica Leech*; Pachliopta aristolochiae interpositus Fruhstorfer*; Troides
aeacus kaguya Nakahara & Esaki*; Omithoptera priamus richmondius Gray; Byasa
febanus Fruhstorfer*, B. polyeuctes termessus Fruhstorfer* , B. alcinous Klug*;
Papilio machaon hippocrates C & R. Felder*, P. xuthus Linnaeus*, P. macilentus
Janson*, P. protenor amaura Jordan*, P. pl'otenor demetrius Cramer*, P. memnon
heromus Fruhstorfer*, P. castor formosanus Fruhstorfer", P. nephelus chaonulus
Fruhstorfer*, P. aegeus aegeus Donovan, P. aegeus ormenus Guerin, P. ambrax
ambrax BOisduval, P. euchenol' euchenor Guerin, P. helenus fortunius Fruhstorfer*,
P. thaiwanus Rothschild *, P. polytes pasikrates Fruhstorfer*, P. bianor takasago
Nakahara & Esaki*, P. bianor dehaani C. & R. Felder", P. ulysses autolycus Felder,
P. laglaizei Depuiset, P. anactus Macleay; Princips demoleus libanius Fruhstorfer*, P.
demoleus sthenelus Macleay; Chilasa agestor matsumurae Fruhstorfer*; Graphium
cloanthus kuge Fruhstorfer* , G. sarpedon nipponum Fruhstorfer*, G. sarpedon connectens Fruhstorfer*, G. sarpedon choredon Felder, G. doson positanus Fruhstorfer*, G.
eurypylus lycaonides Rothschild, G. agamemnon ligatus Rothschild, G. wallacei
wallacei Hewitson, G. macleayanus macleayanus Leach, G. weiskei Ribbe.
Pieridae. Aporia hippia japonica Matsumura*, A. crataegi adherbal Fruhstorfer*.
Danaidae. Idea leuconoe clara Butler"; Euploea leucostictos hobsoni Butler*,
E . sylvestor swinhoei Wallace", E. tulliolus koxinga Fruhstorfer*.
Amathusiidae. Taenaris myops kirschii Staudinger.
Satyridae. Lethe chandica rathnacri Fruhstorfer*, L. europa pavida Fruhstorfer*;
Ninguta schrenckii menalcas Fruhstorfer*; Neope goschkevitschii Menetries (or
N. niphonica Butler?) *; Lopinga achine achinoides Butler"; Hypocysta adiante
adiante Hi.ibner, H. pseudirius Butler, H. metirius Butler, H. aroa aroa BethuneBaker (?, from Madang, New Guinea), H. euphemia Westwood; Argynnina cyrila
Waterhouse & Lyell, A. tasmanica Lyell; Heteronympha paradelpha paradelpha
Lower, H. penelope penelope Waterhouse, H. banksii banksii Leach, H. cordace
cordace Geyer; Oreixenica lathoniella hel'ceus Waterhouse & Lyell; Tisiphone abeona
abeona Donovan; Erebia niphonica Janson*, E. ligea takanonis Matsumura*; Oeneis
daisetsuzana Matsumura *.
Nymphalidae. Clossiana thore jezoensis Matsumura"; Hypolimnas deois divina
Fruhstorfer 'i'.
Acraeidae. Acraea issoria formosana Fruhstorfer*; Miyana meyeri Kirsch.
Lycaenidae. Taraka hamada Druce"; Curetis brunnea Wileman *; Arthopoetes
pryeri Murray*; Ussuriana stygiana Butler*; Japnoica lutea H ewitson*, ]. saepestriata
Hewitson*; Shil'Ozua jonasi Janson*; Antigius attilia Bremer", A. butleri Fenton*;
Araragi enthea Janson*; Ravenna nivea Nire*; Wagimo signatus quercivorus
Staudinger*; Iratsume orsedice Butler*; Favonius saphirinus Staudinger*, F. orientalis
Murray", F. yuasai Shirozu*, F. cognatus Staudinger*, F. jezoensis Matsumura*,
F . ultramarinus hayashii Shirozu*; Neozephyrus taxila japonicus Murray*; Chrysozephyl'us smaragdinus Bremer", C. aUl'orinus Oberthur (Koyama, pel's. comm.),
C. ataxus kirishimaensis Okajima*; Narathura japonica Murray*, N. bazalus tUl'bata
Butler*, N. alkisthenes Fruhstorfer, N. micale centra Evans; Jalmenus ictinus Hewitson; Hypolycaena phorbas phorbas Fruhstorfer, H . danis del'ipha Hewitson; Deudorix
epijarbas diovis Hewitson; Rapala varuna simsoni Miskin, R. varuna formosana
Fruhstorfer*, R. arata Bremer*; Strymonidia w-album fentoni Butler*, S. mera
Janson*; Ahlbergia ferrea Butler*; Lycaena phlaeas daimio Seitz; Heliophorus ila
matsumurae Fruhstorfer*; Anthene lycaenoides Felder, subsp. (New Guinea);
Holochila neuropacuna Bethune-Baker, H. ardosiacea Tite, H. absimilis Felder, H.
consimilis Waterhouse; Cyprotides cyprotus Olliff; Candalides xanthospilos Hubner;
Microscene heathi heathi Cox; Niphanda fusca shijimia Fruhstorfer*; Prosotas dubiosa
dubiosa Semper, P. nora nora Felder, P. nora formosana Fruhstorfer*, P. papuana
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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY
Tite, P. felderi Murray; Nacaduba biocellata biocellata Felder, N. ruficirca Tite,
N. kurava parma Waterhouse & Lyell, N. berenice Herrich-Schaffer; Paraduba
owgarra Bethune-Baker, P. metriodes Bethune-Baker; Ionolyce helicon Felder, subsp.
(New Guinea); Erysichton lineata meiranganus Rober; Neolucia agricola agricola
Westwood, N. mathewi Miskin; "Castalius" mindams mindams Felder; Pistoria
nigropunctatus nigropunctatus Bethune-Baker; Callictita cyara Bethune-Baker; Danis
hymetus taygetus Felder, D. hymetus hymetus Felder, D. albula Grose-Smith, D.
hebes Druce; Pepliophoms inops piluma Druce; Jamides purpurata purpurata GroseSmith, J. aratus Cramer, subsp. (?), ]. nemophila electus Grose-Smith, ]. croUus
pseudeuchylas Strand, ]. aleuas nitidus Tite, J. celeno sun dana Fruhstorfer; Catochrysops panormus papuana Tite, C. amasea Waterhouse & Lyell; Lampides boeticus
Linnaeus; Syntamcus plinius pseudocassius Murray; Everes argiades hellotia
Menetries*, E. lactumus Hubner, subsp. (New Guinea); Tongeia fischeri Eversmann*; Pithecops nih ana urai Bethune-Baker*; Celastrina argiolus ladonides de
l'Orza *, C. sugitanii Matsumura *, C. limbata himlicon Fruhstorfer*, C. albocaerulea
sauteri Fruhstorfer*, C. philippina nedda Grose-Smith, C. argioloides Rothschild,
C. drucei drucei Bethune-Baker, C. leucothalia Jordan, C. meeki meeki BethuneBaker; C. owgarra Bethune-Baker; Udara rona Grose-Smith, U. dardia owgarra
Bethune-Baker; Eupsychellus dionisius Boisduval; Maculinea teleius kazamoto Druce*;
Sinia divina barine Leech*; Euchrysops cneius cnidus Waterhouse & Lyell; Plebeius
argus micrargus Butler*; Lycaeides argyrognomon praeterinsularis Verity *, L.
subsolana yagina Strand*, L. subsolana yarigatakeana Matsumura*.
SPECIES WITH TYPE II EYES
Nymphalidae. Dichorragia nesimachus nesiotes Fruhstorfer*; Restina assimilis
formosana Moore*.
SPECIES WITH TYPE III EYES
Subtypes are shown in parentheses after each names wherever adequate records
are available.
Pieridae. Colias erate poliographus Motschulsky (c) *, C. palaeno aias Fruhstorfer
(c)*, C. palaeno sugitanii Esaki (c)*; Eurema hecabe mandarina de I'Orza (b)*, E.
hecabe hobsoni Butler (b)*, E. laeta bethesba Janson (b)*, E. esakii Shirozu (b)*,
E. blanda arsakia Fruhstorfer (b) *, E. smilax Donovan (b), E. candida puella
Boisduval (b); Catopsilia pomona Fruhstorfer (c) *, C. crocale Cramer (c), C.
pyranthe pyranthe Linnaeus (c) *, C. pyranthe crokera Macleay (b); Gonepteryx
mahaguru niphonica Verity (f) *; Appias indra aristoxenus Fruhstorfer (c) *; Delias
nysa nysa Fabricius (c); Cepora nandina eunama Fruhstorfer (c) *, C. perimele
latilimbata Butler (c), C. euryxantha Homath (c), C. abnormis Wallace (c);
Elodina egnatia angulipennis Lucas (b), E. parthia Hewitson (b), E. padusa
Hewitson (a); Prioneris thestylis formosan a Fruhstorfer (c) *; Ixias pyrene insignis
Butler (f) *; Anthocharis scolymus Butler (c) *, A. cardamines isshikii Matsumura
(b) *; Pieris rapae rapae Linnaeus ( c), P. rapae curcivora Boisduval ( c) *, P.
canidia canidia Linnaeus (c) *, P. melete Menetries (c) *, P. napi nesis Fruhstorfer
(c) *, P. napi iaponica Shirozu (c) *; Leptosia nina niobe Wallace (c) *; Leptidea
amurensis Menetries (a) *.
Danaidae. Danaus plexippus Linnaeus (1- IIIh ); Limnas chrysippus petilia Stoll 3
(g-h, e), t;l (d or e-h); Tirumala hamata septentrionis Butler (g) *; Radena similis
similis Linnaeus (g) *; Parantica aglea maghaba Fruhstorfer (g) *, P. melaneus swinhoei
Moore (g) *, P. sUa niphonica Moore (g) *, (h?) **; Euploea althaea iuvia Fruhstorfer
(h?)*, E. mulciber barsine Fruhstorfer (h?), E. core corinna Macleay t;l (g).
Ithomiidae. Tellervo zoiltts hempsal Fruhstorfer (b).
Satyridae. Penthema formosan us Rothschild (b?); M elanitis leda leda Linnaeus
(f)*, M. leda destitans Fruhstorfer (g), M. phedima oitensis Matsumura (f)*, M.
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phedima polishana Fruhstorfer (f)*, M. amabilis Boisduval subsp. (New Guinea)
(g); Mycalesis gotama fulginia Fruhstorfer (g)*, M. francisca formosana Fruhstorfer
( g) *, M. terminus kyllenion Fruhstorfer (g); Lethe marginalis Motschulsky (g) *,
L. diana Butler (g) *, L. siceUs Hewitson (g) *; H arima callipteris Butler (b-c) *;
Kirinia epaminondas Staudinger (g) *; Lasiommata deidamia interrupta Fruhstorfer
(g) *; Harsiesis hygea hygea Hewitson (g); Pieridopsis virgo Rothschild & Jordan
(g); Platypthima homochroa Rothschild & Jordan (g), P. decolor Rothschild &
Jordan (g); Geitoneura klugi klugi Guerin (b), G. acantha acantha Donovan <f'
(b-g); H eteronympha merope merope Fabricius it; (b), <f' (g) , H . mirifica Butler
it; (b-g), <f' ( g); Ypthima arctous papuana Fruhstorfer (g), Y. arctous arctous
Fabricius (g), Y. argus Butler (g) *, Y. yamanakai Sonan (g) *, Y. multistriata
Butler (g) *, Y. esakii Shirozu (g) *; Coenonympha oedippus annulifer Butler (g) *;
Minois dryas bipunctatus Motschulsky (g) *.
Nymphalidae. Cupha prosope oderca Fruhstorfer (b); Mellicta ambigua niphona
Butler (b)*; Melitaea scotosia Butler*; Clossiana freija asahidakeana Matsumura
(b-c) *; Brenthis daphne rabdia Butler (c) *, B. ina tigroides Fruhstorfer (c) *;
Argynnis paphia geisha Hemming (c) *, A. anadyomene midas Butler (c) *; Damora
sagana liane Fruhstorfer (c) *; F abriciana adippe pallescens Butler (c) *, F. nerippe
C. & R. Felder (c)*; Mesoacidalia charlotta fortuna Janson (c)*; Argyronome laodice
japonica Menetries (c) *, A. ruslana lysippe Janson (c); Argyreus hyperbius hyperbius
Linnaeus (c) *, A. hyperbius inconstans Butler (b); Vindula arsinoe rebeli Fruhstorfer
( c); Vagrans egista offaka Fruhstorfer (c-d); Araschnia burejana strigosa Butler
(b)*, A.levana obscura Fenton (b)*; Polygonia c-aureum Linnaeus (d)*; P. c-album
hamigera Butler (d) *, P. vau-album samurai Fruhstorfer (d) *; Kaniska canace nojaponicum von Siebold (d) *, K. canace drillon Fruhstorfer (d) *; Nymphalis antiopa
asopos Fruhstorfer (d) *, N. xanthomelas japonica Stichel (d) *; Inachis io geisha
Stichel (d) *; Aglais connexa Butler (d) *; Cynthia cardui Linnaeus (c) *; Vanessa
indica Herbst (e)*; Precis almana almana Linnaeus* , P. lemonias lemonias Linnaeus*,
P. iphita iphita Cramer*, P. erigone tristis Miskin (b) ; Yoma algina etonia Fruhstorfer
(c); Symbrenthia hippocle formosana Fruhstorfer*, S. hippocle hippocrates Staudinger; Hypolimnas misippus Linnaeus (RS round and H-shaped) *, H . holina Linnaeus*, H. bolina nerina Fabricius (e), H. alimena eremita Butler it; (e), <f' (d),
H. deois divina Fruhstorfer (b); Doleschallia bisaltide guralca Grose-Smith (c);
Kallima inachus formosana Fruhstorfer*; Mynes geoffroyi ogulina Fruhstorfer (b);
Cyrestis achates achates Butler (b), C. thyodamas mabella Fruhstorfer*; Phaedyma
shepherdi damia Fruhstorfer (b); Neptis praslini meridionalis Talbot (b), N.
sappho intermedia Pryer (c) *, N. hylas luculenta Fruhstorfer*, N. nata lutatia
Fruhstorfer*, N. philyra excellens Butler*, N. rivularis insularum Fruhstorfer*, N.
pryeri jucundita Fruhstorfer*, N. alwina kaempferi de l'Orza*; Athyma selenophora
laela Fruhstorfer (b-d) *, A. perius perius Linnaeus *; Ladoga camilla japonica
Menetries (b-d)*, L. glorifica Fruhstorfer (b-d)*; Helcyra chionippe thyiada Fruhstorfer (c); Apatura ilia substituta Butler (a-b)*; Hestina japonica C. & R. Felder
(a-b) *; Sasakia charonda Hewitson (a-b) *; Charaxes latona papuensis Butler (c);
Polyura pyrrhus sempronius Fabricius (I-IIIb).
Libytheidae. Libythea celtis celtoides Fruhstorfer ( f ) *, L. celtis formosana
Fruhstorfer (f) *.
Lycaenidae. Curetis acuta paracuta de Niceville (a) *; Para lucia aurifer Blanchard
it; (b-c), P. aenea aenea Miskin it; (b-c); Pseudodipsas brisbanensis brisbanensis
Miskin it; (b), P. cuprea Sands it; (a); Hypochrysops delicia delicia Hewitson it;
(b), H. ignita ignita Leach it; (a), H. cyane Waterhouse & Lyell it; (b-g), H.
epicurus Miskin, H. byzos Boisduval, H. pythias pythias Felder it; (b), H. rufinus
Grose-Smith <f' (b), H. polycletus 1'CX Boisduval (b-g); H. argyriorufa van E ecke it;
(a), H. chrysargyra Grose-Smith it; (b-g); Philiris innotatus Miskin it; , P. albihumerata Tite it; (a), P. unipunctata Bethune-Baker it; (a), P. griseldis griseldis
Staudinger (b), P. moira moira Grose-Smith (b); Ogyris genoveva gela Waterhouse
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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY
(; (g), O. ianthis Waterhouse (; (g), O. abrota Westwood (; (g), O. amaryllis
amaryllis Hewitson (; (g), <;1 (a); Hypochlorosis lorquini metilia Fruhstorfer (a),
H. humboldti Druce (a); Spindasis takanonis Matsumura ?**; Erina acasta Cox
(; (a), E. hyacinthina hyacinthina Semper (; (a); Catopyrops ancyra mysia Waterhouse & Lyell (; (a-b), C. florinda estrella Waterhouse & Lyell (; (a-b); Theclinesthes miskini Lucas (; (a), T. onycha onycha Hewitson (; (a-b); "Neolucia"
sulpitius sulpitius Miskin (a), "N." serpentata serpentata Herrich-Schaffer (a);
Zizeeria maha argia Menetries (b) *, Z. maha okinawana Matsumura (a) *, Z. knysna
karsandra Moore (a), "Z." alsulus alsulus Herrich-Schaffer (a); Zizina otis aruensis
Swinhoe (a); Zizula hylax dampierensis Rothschild (a).
SUMMARY
The observations of Yagi & Koyama (1963) on the superficial reflective
pattern of fresh compound eyes based on butterflies from Japan and
Taiwan are extended to those from Papua New Guinea and Australia.
Three basic patterns of the eye are recognised: Type I, with no reflective
spots (RS); Type II, with single RS; and Type III, with multiple RS.
The last one is divided into several subtypes according to the number,
shape, distribution and modification of RS. A number of higher taxa
may be well characterised by the eye patterns. Records of assignments
of individual species to various eye types are compiled.
ACKNOWLEDGMENTS
Thanks are due to Prof. N. Koyama, Shinshu University, Ueda, Japan,
for his permission to utilise some of the original figures and valuable
suggestions; to Mrs. Dorothy Deger and Mr. Anthony Healy, Sydney, for
their supply of a photograph and permission to reproduce a part of a
figure in the book of which Mrs. Deger is a co-author; to Mr. D. P.
Sands, DASF, Lee, Papua New Guinea, for his critical reading of the
manuscript; and to Prof. T. Shirozu, Kyushu University, Fukuoka, Japan,
for drawing my attention to the work by Yagi & Koyama (1963). I am
also indebted to Lt. Col. J. Eliot, Taunton, England, for information on
his recently completed work on the higher classification of Lycaenidae
which is now in press. I also acknowledge the permissions granted by
the National Parks and Wildlife Service, Government of New South
Wales, and the Department of Agriculture, Stock and Fisheries, Papua
New Guinea, to collect specimens used in this work in Ku-ring-gai Chase
National Park, New South Wales, and in Papua New Guinea. I also
express my deep appreciation to Messers T. L. Fenner, DASF, Konedobu;
A. Hutton, DASF, Garaina; R. Straatman, Sogeri; and Dr. J. L. Glessitt
and his staff at the Bernice P. Bishop Museum, New Guinea Field Station,
Wau (now Wau Ecology Institute), for their generous help to my collecting activities in Papua New Guinea.
VOLUME
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LITERATURE CITED
COMMON, I. F. B. 1964. Australian Butterflies. Jacaranda, Bristbane. 132 p.
D'ABRERA, B. 1971. Butterflies of the Australian Region. Landsdowne, Melbourne.
415 p.
DEGER, D. & R. EDEN. 1970. Collecting Australian Butterflies. Horwitz, North
Sydney. 52 p.
EHRLICH, P. R. 1958. The comparative morphology, phylogeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci.
Bull. 39: 305-370.
Fox, R. M. 1956. A monograph of the Ithomiidae (Lepidoptera), Part 1. Bul!.
Amer. Mus. Nat. Hist. lll: 1-76.
HEMMING, F. 1967. The generic names of the butterflies and their type-species
(Lepidoptera: Rhopalocera). Bull. Brit. Museum (Nat. Hist.): Entomo!.
Supp!. 9: 1-510.
KOHIYAMA, K., T. TAKASE & T. FUJIOKA. 1971. Butterflies of Japan. Yama-toKeikoku, Tokyo. 204 p. (in Japanese).
MILLER, L. D. 1968. The higher classification, phylogeny and zoogeography of
the Satyridae (Lepidoptera). Mem. Amer. Entom. Soc. 24: 1-174.
TITE, G. E. 1963. A revision of the genus Candalides and allied genera (Lepidoptera: Lycaenidae). Bull. Brit. Museum (Nat. Hist.): Entomol. 14: 199-260.
YAGI, N. & N. KOYAMA. 1963. The Compound Eye of Lepidoptera: Approach
from Organic Evolution. Shinkyo, Tokyo. 320 p.
STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN
NEW ENGLAND. IV. A PRELIMINARY ANALYSIS OF
BEAK-DAMAGED SPECIMENS, WITH DISCUSSION OF
ANOMALY AS A POTENTIAL ANTI-PREDATOR FUNCTION
OF HINDWING DIVERSITY
THEODORE
D.
SARGENT
Department of Zoology, University of Massachusetts,
Amherst, Massachusetts 01002
An intensive study of the biology of the Catocal,a moths is presently
being conducted in southern New England (Sargent & Hessel, 1970;
Kellogg & Sargent, 1972; Sargent, 1972a). During the course of these
investigations, substantial numbers of beak-damaged specimens have
been collected. The present study was undertaken in hopes that an
analysis of such specimens might shed some light on various aspects of
the predator-prey relationships involving birds and these moths.
Prior Studies
Beak-damaged Lepidoptera have attracted considerable attention in
the literature, though most prior studies have been concerned almost