Taxonomic significance of reflective patterns in the compound eye of ...

JOURNAL OF

THE LEPIDOPTERISTS' SOCIETY

Volume 27 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

162 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

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

VOLUME 27, NUMBER 3 163

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


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.


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.


TABLE l. Distribution of eye types among various families of butterflies (0 , common, predominant types; and . , unusual types, in individual families).

Family II III

a b c d e g h

Hesperiidae 0 Papilionidae 0 Pieridae • 0 0 0 • Danaidae • • 0 0 Iithomiidae 0 Amathusiidae 0 Satyridae 0 • • • 0 Nymphalidae • • 0 0 0 0 • Acraeidae 0 Libytheidae 0 Lycaenidae 0 • • • • developmental 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

.'

o ·


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.


TABLE 2. Distribution of Type I and III eyes among some higher taxa of Satyridae.

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

Lethe europa, L. chandica, Ninguta, Neope Lopinga

Hypocysta

Argynnina, Oreixenica, Heteronympha penelope, H. paradelpha, H. banksii, H. cordace, Tisiphone

Erebia

Oeneis

Type III

Melanitis

Lethe marginalis, L. diana, L. sicelis, Harima Lasiommata, Kirinia Mycalesis, Orsotriaena

Harsiesis, Platypthima, Pieridopsis Geitoneura, Heteronympha merope, H. mirifica

Coenonympha

Minois

* 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.


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


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*.


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 Hewitson*, ]. 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


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.


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); Y pthima 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


(; (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.


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 clas

sification 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

Taxonomic significance of reflective patterns in the compound eye of ...

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