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Taxonomic Significance of Reflective Patterns in The 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 some­ times 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 Nym­ phalidae (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 but­ terflies. 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 super­ ficial (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 but­ terfly 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 di­ rections. 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 hexa­ gonal 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 164 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY 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 progres­ sively 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 pre­ dominant 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 modifica­ tions (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. VOLUME 27, NUMBER 3 165 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 fre­ quently encountered types in each of the butterfly families! occurring in the areas covered, except for Riodinidae for which no observation has been available.
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