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 RETURN TO 
 ENTOMOLOGY LIBRARY 
 
 Cornell University 
 Ithaca, N. Y. 
 
Bulletin of the Museum of Comparative Zoology 
 AT HARVARD COLLEGE. 
 
 Vol. XXX. No. 4. 
 
 ON THK COLOR AND COLOR-PATTERNS OF MOTllS 
 AND BUTTERFLIES. 
 
 I' (ioUDSBOROUGI! MAfGIt. 
 
 ca:\ieridge, mass., U. S.A. : 
 
 printed for the museum. 
 
 fubbuary, 1807. 
 
1 
 
ive Zobl^ /j J 
 
 Bulletin of the Mnseum of Comparat: 
 AT HARVARD COLLEGE, 
 
 Vol. XXX. No. 4 
 
 ON THE COLOR AND COLOR-PATTERNS OF MOTHS 
 AND BUTTERB^LIES. 
 
 By Alfred Goldsborough Mater. 
 
 With Ten Plates. 
 
 CAMBRIDGE, MASS., U. S. A. : 
 PRINTED FOR THE MUSEUM. 
 
 FEBRnART, 1897. 
 
> 
 
 
 '^^o^^']'JE7tt, 
 
 C,U..^H^ 
 
No. 4. — On the Color and Color-l^atterns of Moths and Butter- 
 flies} 
 
 Hv Aj.KKKI) (iol.DSliOKorCII Mayku. 
 
 This reseai-cli is an investigation of the general jilienoniena of Color 
 iti Lepidojitera, and also a special account of the Color-Pattei-ns of 
 the ])anaoid and Acraeoid Tlelieonidae, and of the I'apilios of 
 Tro])ical South America, and has been carried out under the direction 
 of my friend and instructor, Dr. Charles B. Davenport ; and the work 
 was done in connection with one of the coui'ses given liy him in 
 Harvard University in 1894-95. ■' I am indebted to Dr. Davenport 
 not only for suggesting the subject, but also for his kindness in devot- 
 ing much time to a criticism of the results. 
 
 The |)aper is divided into three parts. I'art A contains an ac- 
 count of the general phenomena of color in Lepidoptera; Part B 
 is devoted to a special discussion of the color-variations in the Ileli- 
 conidae, with special reference to the phenomena of mimicry; and 
 Part C consists of a summary of those results which are believed to 
 be new to science. A Table of Contents is given at the end of the 
 paper. 
 
 PART A. 
 
 GENERAL PHENOMENA OF COLOR IN LEPLDOJ^TERA. 
 L Classification ov Colous. 
 
 We follow Poulton ('90) in dividing L('])idoj)terous colors into (1) 
 pigmental and (ii) structural. 
 
 (1) Pigmental Colors are due to tlie presence of an actual ])ig- 
 ment within the scales, and although such colors are very common 
 in the Lepidojitera, it is frequently very difficult to say ol'f-hand 
 whether a given color is due to a ]>igrnent or to some structural effect. 
 Coste ('90-'91) and Urech ('9!!) have, however, given criteria for de- 
 termining whether a color is due to a pigment or to some other cause. 
 They succeeded, for example, in dissolving out the color in many 
 
 ' C(>iitril>iiti<ins from the Zoological Laboratory of the Mu.HCum of Comparative Zool- 
 ogy at Harvard College, E. L. Mark, Direutor, No. LXXIV. 
 
 »Thl8 iiaper was written in 18fl5 essentially as it now stands. 
 
170 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 cases, leaving the wing white or colorless. Coste used .is solvents 
 a number of strong acids and alkalis; while IJrech confined him- 
 self to the use of water, hydrochloric acid, and nitric acid. Their 
 results may be conveniently summarized as follows : — 
 
 Hkick according to Ureoh is a pigmental color, for it may be dis- 
 solved out of the wings by means of hydrochloric or nitric acid. 
 
 Urown is usually insoluble in water, but is soluble in hydrochloric 
 or nitric acid. 
 
 The red and orange pigments of the Pieridae, Lycaenidae, 
 Nymphalidae, Zygaenidae, and some Papilios are soluble in water. 
 They are insoluble in water in the Sphingidae, Arctidae, Bombycidae, 
 Saturnidae, and Geometridae. 
 
 Yelloio pigment is acted upon by reagents in almost the same way 
 as the red and orange, especially if both red and j'ellow appear upon 
 the same wing. It is soluble in the I'ieridae, Lycaenidae, Nyni- 
 j)halidae, Satyridae, and some Papilios, but insoluble in the Sphin- 
 gidae, Arctidae, Geometridae, and a few Noctuidae. 
 
 White is usually a structural color, but can be dissolved out 
 from the wings of the Pieridae by water, being in this case, of 
 course, due to a pigment. 
 
 Green pigment can be dissolved out by water in the cases of the 
 Pieridae, Lycaenidae, and Geometridae. In the vast majority of 
 cases, however, it is a structural color. 
 
 Violet and blue are almost always due to structural causes. In a 
 few cases, however, as in Smerinthus ocellatus, a blue pigment can 
 be dissolved out. 
 
 We see, then, that black, brown, red, orange, and yellow are 
 usually due to pigment, while white, green, violet, and blue are gen- 
 erally due to structural effects. 
 
 It is well known that the scales of Lepidoptera are essentially 
 hollow, flattened sacs often inclosing pigment, and Burmeister (78) 
 arrives at the conclusion, from a study of the scales in various spe- 
 cies of Castnia, that the pigment is for the most part attached to the 
 upper layer of the scale-sac, rendering it opaque, while the lower 
 layer receives less pigment and is, in consoiinence, a little more 
 translucent. 
 
 (2) Structural 6'oZor,s' owe their origin to the external structure of 
 the scales or wing-membranes and not to the presence of a pigment. 
 Thcv are often caused by diffraction, due to the scales being covered 
 with fine, parallel striae. Some of the most splendid colors in the 
 
MAYKU: COLOR AND COLOK-PATTERNS. 171 
 
 animal kingdom are duo to tliis cause ; sucli are the iridescent and 
 opalescent hues of many of the Morphos and Indo-Asiatic Papilios. 
 Very often the scales which display such brilliant colors contain no 
 pigment whatsoever ; for if one will merely soak them in alcohol, 
 ether, or water, all color disappears, and the scales become as trans- 
 parent as glass. This test was devised by Dimmock ('83), who 
 used it upon the brilliantl}' colored scales of many beetles. It 
 was first discovered by Burgess ('80), and has since been con- 
 firmed by Kellogg ('94), that the striae which produce these structural 
 colors are all upon the outer surface of the scale, i. e., the surface 
 which is away from the wing-membrane and exposed to the light. 
 Kellogg ('94) has determined the distance apart of the striae upon 
 the scales of many species of Lepidoptera. It appears, for example, 
 that the striae upon tlie scales of Danais plexippus are 2/* apart, 
 those upon the transparent scales of Morpho sp. l.^/x, upon the 
 pigment-bearing scales of Morpho 0.72/*, and upon Callidryas 
 eubule 0.9;u apart. It is very evident, then, that the brilliant color- 
 ation of the scales may be due to this fine striation, for the striae 
 upon Rowland's or Rutherfurd's finest gratings are a])proximately 
 1..5y«, apart, which is about the average distance between the ridges 
 of the scales. 
 
 Structural colors are, however, not always due to diffraction ; in 
 the case of white, for examjile, the color is almost invariably due to a 
 reflection of all, or nearly all, the light that impinges u])on the scales. 
 As long ago as 1855 Leydig pointed out that the silvery white color 
 seen in the scales of some spiders, such as Salticus and Tegenaria, 
 was due to air contained within them ; and more recently Dimmoi'k 
 ('83) has shown that silvery white and milk-white colorations are 
 due to optical effects produced by reflected light. In the silver}' white 
 scales, however, such as those of the under surface of the hind wings 
 of Argynnis, there must be a polished reflecting surface toward 
 the observer, for both silvery and milk-white colors appear simply 
 milk-white by reflected light. 
 
 (3) Combination Colors owe their richness and brilliancy to a 
 combination of structural and pigmental effects. The geranium-red 
 spots upon the hind wings of the Mexican Papilio zeunis Lucas owe 
 their red color to pigment, but over this red there plays, in certain 
 lights, a beautiful pearly iridescence, which, in combination with the 
 red, greatly enhances its charm. Urech ('92) has demonstrated that 
 in the Vanessas there are scales which have chemical coloring matter 
 
172 BULLETIN: MUSEUM OF COMPAUATIVE ZOOLOGY. 
 
 and interference colors also. In addition, he points out the interest- 
 ing case of certain Lycaenidae where the scales exhibit to the eye 
 only interference effects, and yet a pigment can be dissolved out of 
 them by the use of water. 
 
 (4) QuantUatioe DetermiiuUio/i of Pigmental Colors. I have 
 analyzed the colors of many butterflies by means of the spectroscope, 
 and also by Maxwell's discs. As is well known, Maxwell's discs are 
 colored circular discs of cardboard, perforated at the center and slit 
 along a radius so that two or more of them may be slid over each 
 other, thus exposing different proportions of each. Then by rapidly 
 rotating them the colors become blended, and thus it becomes 
 possible to match any color, and to discover its fundamental con- 
 stituents. Uy this means I have determined that the vast majority 
 of the colors found in Lepidoptera are impure ; that is to say, they 
 contain a large percentage of black. 
 
 P'or example the white of the upper surface of the wings of the 
 common Pieris rapae consists of: 17% black, 13% emerald -green, 
 10% lemon-yellow, and 60% white. 
 
 Also the so-called "blacks" found in butterflies are rarely jet-black, 
 but, almost always, only deep sliaiies of brown. For instance the 
 deep brown color of the under surface of the wings of lleliconius 
 mel])omene consists of 93% black, 3% lemon-yellow, 3.5% of 
 Maxwell's fundamental red (vermilion), and 0.5% of von Bezold's 
 fundamental blue-violet. 
 
 The purest color I have met with is the canary-yellow ground 
 color of the wings of Papilio turnus, which seems to consist of 
 wliito light with the addition of a little yellow. 
 
 Other colors all pos.sess considerable black. Thus the glaucous 
 green of Colaenis dido consists of black 29%, vermilion '24%, 
 emerald-green 37%, von Hezold's blue-violet 10%. 
 
 The sepia-brown ground color of Cercyonis alope consists of black 
 71%, vermilion 21.5%, emerald-green 7.5%. 
 
 The tawny rufous color of the wings of Mechanitis polyninia, etc., 
 is made up of black 46%, vermilion 40%, lemon-yellow 14%. 
 
 The rufous red patch on the upper surface of the fore wings of 
 lleliconius melpomene is made up of black 27%, vermilion 66.5%, 
 lemon-yellow 6.5%. 
 
 The yellow of the fore wings of Mechanitis polymnia consists of 
 lemon-yellow 67%, emerald-green 14%, and white 19%. 
 
MAYER: COLOR AND COLOR-l'ATTERNS. 173 
 
 (5) Spectrum Analysis of Colors of Lepidoptera. I have made 
 some spectrum analyses of the liglit reflected from tlie wings 
 of various butterflies, by means of a piece of apparatus most kindly 
 suggested for the purpose by Prof. Ogden N. Rood of Columbia 
 College. The arrangement is shown in Figs. 1, '2, Plate 1; Fig. 1 
 being a perspective view, and Fig. 2 a horizontal section of the 
 apparatus, which consists of a rectangular bo.\, blackened upon the 
 iii.side, and having a well-fitting cover. A rectangular slit (O) was 
 cut through one of the long sides of the box, near one end, and the 
 other end of the same side was perforated in order to allow the 
 admission of the direct-vision spectroscope (S). Imagine that we 
 wish to examine the yellow spots from a butterfly's wing. All of 
 the yellow spots from the wing are cut out, and pasted upon two 
 pieces of cardboard so as to make two large unbroken patches of 
 color. The pieces of cardboard are then blackened upon all those 
 places where the colored wing was not pasted. One of the card- 
 boards is then suitably mounted upon the back of the bo.x at B ; the 
 other is placed upon a vertical support (F'), the plane of which is 
 parallel to the back of the box. 
 
 The working of the apparatus is as follows : the sunlight enters 
 by the slit (O) and is reflected and diffused three or four times 
 between the pieces of colored wing mounted upon the back (H) of 
 the box, and the vertical support (F). The manner of this reflec- 
 tion and diffusion is shown by the dotted lines of Fig. 2. After 
 undergoing several reflections, the light enters the direct-vision 
 spectroscope (S). The slit of the spectroscope is wide open, and 
 thus the light which enters it may readily be examined. It was 
 fo\ind that it was necessary that the light be reflected more than 
 once from the wing before it enters the spectroscope, for the first 
 reflection shows so much white light that it is usually quite impossi- 
 ble to analyze the true color of the wing, the j)redominant colors 
 being obscured by a continuous spectrum. In general it was found 
 that the colors of the wings are not simple, but compound ; that is 
 to say, they are made up of a mixture of several different colors. 
 
 For example, the spectrum of the rufous ground color of the 
 upper surface of the wings of Danais plexippus consists of all of the 
 red and yellow of the spectrum and about 75% o^ tlic green. 
 
 The red spots upon the upper side of the fore wings of Ileliconius 
 Tuelpomene also consist of the red and yellow and a very faint, 
 hardly visible, trace of green. 
 
174 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 The glaucous green patches on the wings of Colaenis dido are 
 composed mainly of green and yellow, but there is also a faint develop- 
 ment of about half of the blue and a still fainter trace of red. 
 
 The iridescent blue-green ground color of the upper surface of the 
 wings of Morpho menelaiis, viewed in such a way that the light 
 makes an angle of about 20° with the normal to the surface of the 
 wing, gives a spectrum of green and blue about equally developed. 
 
 The yellow ground color found on the upper side of the wings of 
 Papilio turnus shows a continuous spectrum, in which the yellow 
 seems to be rather more brilliant than in the normal spectrum of 
 white light. 
 
 The sepia-brown ground color of the upper surface of the wings 
 of Cercyonis alope gives a spectrum which lacks only the blue-green 
 and blue. 
 
 (6) Smnmarij of Results. The researches of Coste ('90-'91) and 
 Urech ('93) have demonstrated that the colors of butterflies and 
 moths may be produced by two causes : by the presence of an actual 
 pigment, or by some structural effect. Some colors are due entirely 
 to pigment, others to structural causes, and still others to a combina- 
 tion of the two. 
 
 Black, brown, red, orange, and yellow are invariably due to 
 pigment. 
 
 Green is usually due to a structural effect, but in a few cases there 
 is a green pigment present. 
 
 White, blue, and violet are almost invariably due to structural 
 causes. 
 
 In addition to these facts I have found that most of the colors 
 which are displayed by Lepidoptera contain a surprisingly large 
 percentage of black. Also they are usually not simple colors, but 
 composed of a mixture of several different colors. It is remarkable 
 that Natural Selection, which is generally assumed to have been one 
 of the principal factors in bringing about the wonderful develop- 
 ment of colors in Lepidoptera, has not been potent enough to make 
 these colors purer than is the case in existing butterflies. 
 
 II. The kssentiai. Nature ok Pigmental Color in 
 Lepidoptera. 
 
 (1) Pigments of Laroae. Poulton ('85) showed that the jihy- 
 tophagous larvae of Lepidoptera " owe their colour and markings to 
 
MAYER: COLOR AND COLOR-PATTERNS. 175 
 
 two causes: (I) Pigments clerivecl from their food-plants, cliloro- 
 phyll and xantliophyll, and probably others ; (2) pigments proper 
 to the larvae, or larval tissues made use of because of some (merely 
 incidental) aid which they lend to the colouring, e. </. fat." Poulton 
 concludes that all green coloration is due to chlorophyll, and 
 that nearly all yellows are due to xanthophyll. All other colors, 
 including black and wliite and some yt'Hows, are due to pigments 
 proper to the larvae themselves. 
 
 Later, in 1893, Poultoii proved that the larvae of Tryphaena 
 pronuba could transform both etiolin and chlorophyll into a larval 
 coloring matter, which may be either green or brown. It thus 
 appears that some brown pigments are deprived from food, and are 
 merely modified plant pigments. Green larvae have green blood, 
 and this color is due to chlorophyll in solution. It is remarkable 
 that this clilorophyll solution is stable under the prolonged action of 
 light, and in this respect is different from any other known solution 
 of chloropliyll. It is worthy of note, further, that the spectrum 
 of this green blood shows a great resemblance to that of chlorophyll. 
 " In fact the two spectra are far nearer to one another than the 
 ordinary spectrum of chlorophyll in alcoholic solution, is to the 
 unaltered chlorophyll of leaves." 
 
 (2) Pigments of Imagines. In 1891, Urech showed that the 
 similarity between the color of the urine of butterflies and the 
 principal color of their scales is so close that it cannot be considered 
 as accidental, but rather must be regai-ded as physiological. Urech 
 compares in a table the color of the urine and that of the scales 
 of 29 species of Lepidoptera. In all but two species the resem- 
 blance is very close.' 
 
 Urech further shows that the color of the urine (and the corres- 
 ponding color of the scales) is not dependent upon the kind of food, 
 for one and the same food plant may be differently digested in 
 differentgroups of Lepidoptera. Thus he compares the behavior of a 
 Vanessa with that of one of the Microlepidoptera (leaf-rollers) . Both 
 of these feed upon the nettle (Urtica). In the larva of the Vanessa the 
 contents of the stomach are intensely green, but become red in the 
 pupa. In the case of the leaf-roller the contents of the stomach are 
 never markedly green and become insi])id in color during the pupal 
 stage. 
 
 1 Likewise, Hopkins ('04) has sliown that In the Pieridae the urine is tinged by a yellow 
 substance having exactly the color of the wings. 
 
176 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 Ponlton has shown that the reddish fluid voided by the Vanessas 
 immediately after emergence from the chrysalis contains uric acid, 
 and Hopkins ('94) says that when the yellow Pieridao emerge, they 
 often void from the rectum a large quantity of uric acid. It shonld 
 be borne in mind however, as Urech himself suggests, that the pig- 
 ment found within the wings may not be identical in chemical com- 
 position with the similarly colored fluid from the alimentary tract. 
 
 Hopkins ('89, '91, '94, '9(i) has discovered that the white pigment 
 found in the scales of Fieridae is uric acid, and that the red and 
 yellow pigments of the Pieridae are due to derivatives of uric 
 acid. He also says, " these uric acid derivatives used in ornamen- 
 tation, are apparently confined to the Pieridae alone among butter- 
 flies." Hence when a Pierid mimics an insect of another family, the 
 pigments in the two cases are chemically quite distinct. This is well 
 seen in the genera Leptalis (Pieridae) and Mechanitis (Danaidae). 
 
 In addition to this, Griffiths ('92) finds that the green pigment 
 found in Papilio, Parthenos, llesporia, Limenitis, Larentia, Ino, and 
 Ilalias is a derivative of uric acid, to which he gives the name of 
 "Lepidopteric acid " and as.signs tlie empyrical formula C^ H,;, Az^ 
 
 In a paper published in 1896 in the Bulletin of the Museum of 
 Comparative Zoology at Harvard College, Vol. 29, I have shown, 
 p. 226-230, that the pigments of the scales of Lepidoptera are 
 derived by various chemical processes from the lilood, or haemo- 
 lymph, of the pupa, and that the liaeTnoljMnph is a proteid substance 
 containing egg-albumen, globulin, tibrin, xantliophyll, orthophos- 
 phoric acid, iron, potassium, and sodium. 
 
 III. Dkvulopment of the various Colors in tiik Pupal 
 Wings. 
 
 A few researches have been carried out upon this interesting 
 topic, but as the literature is scattered and has never been brought 
 together, it will perhaps not be amiss to present a brief r6sum6 of 
 the principal facts which have been already ascertained. 
 
 (1) Jlistorical Account of preoious Researches. In 1889 
 Schiiffer ('89) discussed the question of the order and time of 
 appearance of the colors in the pupal wings of several of the 
 Vanessas. Unfortunately he apparently did not make his obser- 
 
MAYER: COLOR AND COLOR-PATTERNS. 177 
 
 vations at sufficiently close intervals of time, and was, therefore, 
 led into some misstatements, which have been corrected by van 
 Benimelen ('89) and Urech ('91). 
 
 Van Bemmclen carried out an elaborate research upon the 
 development of the various spots and colors upon the win<i;s of 
 Pyrameis cardui, Vanessa urticae, V. io, Pieris brassicae, and a few 
 other forms. He discusses in detail the time and manner of aj)pear- 
 ance of all of tlie different spots upon the wing. Into these details 
 we shall not follow him, but sliall merely present his general con- 
 clusions regarding the development of the various colors. In Pieris 
 brassicae it appears that during the first days of the pupal stage tlie 
 wings are colorless and transparent ; after a few days, however, the 
 fore wings become opaque, and white ; later the hind wing, also, 
 goes through the same changes. The wings then remain unaltered 
 until about two days before the butterfly issues. Then, very sud- 
 denly, the black spots and the yellow ground tone of the under 
 sides appear. "White is thus the primarj' color ; black and yellow 
 secondary. Tlie first color to make its appearance in the case of 
 Pyrameis cardui is a brown-yellow ground color, which may be 
 observed in pupae four days old. The hind wings are at this time 
 somewhat darker than the fore wings. The color then changes from 
 darker brown to cinnamon-brown. The black spots appear later upon 
 this delicate reddish brown ground color. The three fused sj)ots 
 which form the whitish band in the middle of the front edge of the 
 fore wing appear during the last days of development, just before 
 the completion of the final color-pattern. 
 
 Both van Bemmelen and Urech have shown that in Vanessa 
 urticae the order of a))pearance of the various colors is the same as 
 in Pyrameis cardui. The first color to appear in Vanessa urticae is 
 a faint reddish tinge ; this deepens and forms the ground color, and 
 later the black spots appear upon it. 
 
 Urech ('91) has made a careful study of the development of the 
 colors upon the pupal wings of Vanessa io. The wings are at lirst 
 wholly white. Then in a restricted area of this white is noticed the 
 appearance of a yellow, which forms the yellow of the matiu-e wings. 
 Almost contemporaneous with the development of the yellow comes 
 the red, which ap|)ears in another part of the primitively white field, 
 and gradually deepens in color until it forms the brownish red 
 ground color of the adult wings. Still later another portion of the 
 primitive white changes into the black of the mature wing. The 
 
17B BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 under side of the mature wings of Vanessa io is mainly uniform 
 black, and in this case also tliis color develops from the white at a 
 very rapid rate, near the end of the pupal stage. This development 
 of tlie black directly upon the white areas is quite remarkable in 
 Vanessa io, and very different from that of both Vanessa urticae 
 and Pyrameis cardui, where the black spots develop upon a field 
 already tinged with red. Urech j)oints out the fact, that some of 
 the white spots seen in the mature wings of the Vanessas represent 
 the " primitive white " of tlie pupal wings. 
 
 Finally, the latest paper upon the subject of the development 
 of color in the pupa is that of Haase ('93), who has examined 
 the pupae of a number of Papilios (e. r/., pliilenor, machaon, 
 asterias, turnus, and podalirius), and finds that during early pupal 
 life the wings are as transparent as glass ; after a time, however, 
 they ch.ange to an impure white, which soon becomes yellowish, and 
 then the various colors which are destined to adorn the mature 
 wings begin to appear. 
 
 If we are to learn much of fundamental import concerning the 
 phylogeny of color in Lepidoptera, the reseai-ches should be carried 
 out upon the lower moths, and not upon such highly specialized 
 forms of Rhopalocera as the Vanessae. 
 
 In my paper on Wing scales, etc. (Mayer, '96, p. 232), I have 
 come to the conclusion that dull ocher-j^ellow and drabs are, 
 phylogenetically speaking, the oldest pigmental colors in the Lepi- 
 doptera. The more brilliant colors, such as briglit yellows, reds, 
 and pigmental greens, are derived by complex chemical processes, 
 and are, phylogenetically speaking, of recent appearance. 
 
 I have made a study of the development of the colors and pattern 
 in the wings of Callosamia promethea Linn, and of Danais plexip- 
 pus Fab. 
 
 (2) Development of Color in the J'updl Winf/s of Cullosaniia 
 promethea. The cocoons of Callosamia promethea are very abundant 
 during the winter months, when they may be found hanging to the 
 stems of the food plants of tlie larvae. The pupal wings remain 
 pei-fectly trans|)arent all thi-ougli the winter, until about ten days 
 before the time when the moth is destined to issue ; they then become 
 opaque white. An examination of the wings at tliis period shows 
 that the scales are perfectly formed (Fig, 25, Plate 3), except for the 
 
MAYHR: COLOR AND COLOR-PATTERNS. 179 
 
 lack of pigment, which is developed later. If one treats the scales at 
 this stage with oil of cedar-wood or clove oil, they become practically 
 invisible under the microscope, thus demonstrating that there ia 
 no pigment within them. Fig. 2C, Plate 3, gives the appearance 
 presented by a scale taken from the light drab-colored margin of 
 the mature wing. This is about the lightest area upon the wing, 
 except the white spots ; but it will be seen that this scale is much 
 darker in ajjijcarance than the unpigmented one shown in Fig. 25. 
 The white or unpigmented condition of the wing lasts for about 
 four days. The wings then become uniformly tinged with an 
 impure yellow or light drab, and very soon after this the colors 
 begin to make their appearance. They first appear upon the lower 
 surface of the wings. Fig. 28, I'late 3, represents the under 
 surface of the fore wing of a female in a very eai'ly stage of color 
 development; in fact the upper surface shows, as yet, no trace of 
 the colors. It will be seen that a few dark red streaks have 
 appeared near the central portion of the wing, and it is worthy of 
 note that these occupy the iiitei-spaces between the nervitres. The 
 ocellus near the apex of the wing appears faintly outlined upon its 
 background of impure yellow. 
 
 Fig. 27, Plate 3, represents the under side of a hind wing of 
 a male in about the same stage as Fig. 28. Here, again, the red color 
 occupies the interspaces, and indeed it is only later that the nervures 
 become clouded over by it. 
 
 Figs. 29 and 30, Plate 3, represent, respectively, the under and 
 upper sides of the fore wing of a male about five hours after the 
 first appearance of the colors. Upon the ujjper side (Fig. 80) we 
 see two gray streaks near the base of the wing and a light cinnamon- 
 brown color extending from the lower edge toward the middle of 
 the wing. The ocellus near the apex is now quite apparent, but 
 still faint in color. On the under surface (Fig. 2!)) the red markings 
 have developed to a much greater extent than in Fig. 28. The 
 outermost of the two white spots which occupy the center of this red 
 area becomes the white central spot of tlie mature wing ; the inner- 
 most one is soon obliterated owing to its becoming clouded over 
 with red. 
 
 Figs. 87 and 30 represent respectively the upper surface of 
 the fore wing and the lower surface of the hind wing of a female, 
 slightly more advanced than in Fig. 30. Fig. 31 represents a male 
 and Fig. 38 a female about twelve hours after the tii-st appearance 
 
180 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 of the color. It is remarkable that in this stage the male and female 
 wings are quite similar in general appearance, except that the ground 
 color of the male is now a dusky gray, while that of the female is a 
 cinnamon-brown. 
 
 Fi'om this time onward, however, the wings of the two sexes begin 
 to differ more and more in ajjpearance, for the ground color of the 
 male becomes deep black, while tliat of the female remains cinnamon- 
 brown. This change is well exhibited by Figs. 32 and 39, Plate 3, 
 which give the appearance of the upper surfaces of the male and 
 female wings respectively at about twenty hours after the first appear- 
 ance of the colors. Fig. 33 represents the hind wing of the .same male 
 whose fore wing is shown in Fig. 3'2. Figs. 34, 35, 40, and 41 give 
 the appearance of the pupal wings just before emergence, when 
 the colors are completely formed. 
 
 To summarize ; Figs. 27, 29, 33, and 35 give successive stages in 
 the development of color in the male ; and Figs. 28, 36-41 give 
 similar stages for the female. It becomes evident, from a comparison 
 of these successive developmental stages, that the colors appear first 
 upon the central portions of the wings, and that the outer and costal 
 edges of the wings and the nervures are the last parts to acquire 
 the mature coloration. 
 
 It is worthy of remark that the color-pattern of the mature male 
 Callosamia jjromethea is quite a departure from the type of coloration 
 which is commonly found among the Saturnidae. The female, 
 however, conforms very well to the general pattern of the other 
 species of the family. It is quite evident that the deep black colora- 
 tion of the male is, phylogenctically speaking, a new acquisition, and 
 that the coloration of the female represents the less dilTerentiated 
 and therefore, more primitive type. 
 
 It is interesting in connection with these facts to observe that the 
 color-patterns of both male and female develop in almost identical 
 ways up to the twelfth hour after the first appearance of the color ; 
 that then, however, the grayish ground color of the male wings 
 begins to deepen into the characteristic jet black of the adult, while 
 the light cinnamon ground color of the female merely becomes 
 slightly darker a^ the wings mature. 
 
 (3) Development of Color in the Pupal Wi/iffs of Danais 
 plexippus. Figs. 42-45, Plate 3, are intended to illustrate four 
 stages in the development of color in the pupal fore wings of Danais 
 plexippus. The pupal stage of this species is of brief duration, last- 
 
MAYER: COLOR AND COLOR-rATTERNS. 181 
 
 iiig from one to two weeks only, according to the temperature to 
 which tiie chrysaHs is exposed. For the first few days the wings are 
 [)erfectly transparent, but about five days before the butterfly issues 
 they become ])ure white. An examination of the scales at this 
 period shows tliat they are completely formed and merely lack 
 pigment. In about 48 hours after this (see Fig. 42) the ground 
 color of the wings changes to a dirty yellow. It is interesting to 
 note that the white s])ots which adorn the mature wings remain 
 pure white. Fig. 43 illustrates the next stage, where the black has 
 begun to appear in the region beyond the cell. The nervures them- 
 selves, however, remain white. Fig. 44 shows a still later condi- 
 tion, where the dirty yellow ground color has deepened into rufou.s, 
 and the black has deepened and increased iii area and has also 
 begun to appear along the edges of the nervures. In Fig. 45 the 
 black has finally suffused the nervures, the base of the w'ing and 
 the submedian nervure being the only ])arts that still remain dull 
 yellow. It is apparent that in Danais plexippus, as in Oallosamia 
 promethea, the central arejis of the wings are the first to exhibit the 
 mature colors, and that the nervures and costal edges of the wings 
 are the last to be suffused. 
 
 IV. TiiK Laws which govekn thk O)L0K-PArrKKNs of IUittkr- 
 
 FI.IKS AND MoTIIS. 
 
 (1) Historical Account of precious Jiesearches. The earliest 
 paper upon this subject is by Iliggins ('08). He came to the 
 conclusion, that "the simplest type of color presents itself in the 
 plain uniform tint exhibited when the scales are all exactly alike." 
 He also thought it probable that "the scales growing on the mem- 
 brane upon or near the veins would be distinguished from the 
 scales growing on other parts of the membrane by a freer develop- 
 ment of pigmentary matter, and that in this manner would arise 
 a kind of primary or fundamental color-pattern, namely, a pale 
 ground with darker linear markings following the course of the veins, 
 e. ff. Fieris crataegi." He also attempted to explain the formation of 
 eye-spots by assuming that crescent-shaped markings migrate out- 
 wards from the sides of the nervures and meet so as to inclose a 
 space. 
 
182 
 
 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 It is, however, untrue that there is a freer development of pig- 
 ment within tiie scales Ij'ing upon the nervures; in fact, the reverse 
 is the case, as we have seen, in both Danais plexippus and Callo- 
 samia promethea. Iliggins's explanation of the formation of eye- 
 spots is also fallacious. 
 
 Darwin (71, Vol. 2, p. 133) published four excellent figures from 
 a drawing by Trimen, illustrating two simple ways m which eyc- 
 spots are actually formed, both diameti'ically opposed to Iliggins's 
 hypothesis. Darwin says that in the South African butterfly, Gyllo 
 leda, " in some specimens, large spaces on the upper surface of the 
 wings are coloured black, and include irregular white marks, and 
 from this state a complete gradation can be traced into a tolerably 
 perfect ocellus, and this results from the contraction of the irregular 
 blotches of colour. In another series of specimens a gradation can 
 be followed from excessively minute white dots, surrounded by a 
 scarcely visible black line, into perfectly .symmetrical and large 
 ocelli " with several rings. 
 
 Scudder ('88-'89) and, afterwards, Bateson ('94) have shown 
 that the ordinary eye-.spots, such as those found in Mori)ho and the 
 Satyridae, are invariably placed in the interspaces between the longi- 
 tudinal veins of the wings, and also that they are often found repeated 
 upon homologous |)laces of both pairs of wings. IJatesoii says that 
 ocelli are often seen upon both surfaces of the wing, the centei's of 
 the upper and lower ocelli coinciding. In the majority of cases, 
 liowever, the upper and lower ocelli, although coincident, have (juitc 
 different colors. The simpler sort of ocelli, such as those seen in the 
 Satyridae or in Morpho, have their centers on the line of the fold- 
 marks or creases of the wing. It sometimes happens that these 
 creases .seem to begin from the center of an ocellus. As the.se 
 creases commonly run midway between two nervures, it usually re- 
 sults that the center of the eye-spot is exactly half way between two 
 nervures. The large eye-spots of Parnassius apoUo are an exception 
 to this rule. In some Morphos, Satyridae, etc., in cell P of the hind 
 wing there are often two creases and two eye-spots, one for each 
 crease ; but if there be only one eye-spot present, its center does not 
 correspond with the middle of the cell, " but is exactly upon the 
 anterior of the two creases." I have observed the same law for the 
 white marginal spots in cell P in Ceratinia vallonia, C. fimbria, and 
 Mechanitis polyninia. 
 
 In 1889 Scudder, in his work upon the Butterflies of New 
 
MAYER: COLOR AND COLOK-l'ATTKRNS. 183 
 
 England, called attention to the following facts : the transverse 
 soi'ics of dark spots so often seen in the body of the wings of 
 Lcpidoptera arc invariably placed in the interspaces between the 
 longitudinal veins, never upon the veins themselves, excepting 
 only in rare instances, where the spots occur at the extreme margin, 
 lie also jjointed out that in many types of moths all differentiation 
 in coloring has been greatly retarded, so far as the hind wings are 
 concerned, by their almost universal concealment by day beneath 
 the overlapping front wings. In these cases " the simplest departure 
 from uniformity consists of a deepening of the tint next the outer 
 margin of the wing." It is but a step from this condition to a 
 band of dark color or a row of spots parallel with the margin. This 
 explains why the transverse style of markings, for the hind wings 
 at least, is so common. Scudder showed that "the number of 
 instances, in butterflies, in which similar markings appear in the 
 same areas of the two wings, and in the same relative position 
 in these areas, is far too common to be a mere coincidence. It is 
 most readily traced in the disposition of the ocelli, which are very 
 apt to be similar in size and perfection, and to be situated between 
 the same branches of homologous veins." 
 
 (2) Laws of Color-Patterns. As a result of my own study of 
 the wings of moths and butterflies, I am prepared to propose the 
 following additional laws of color-patterns, (a) Any spot found 
 upon the wings of a moth or biUterfly teiids to be bilaterally stpnmet- 
 rical both as regards form and color., the axis of symmetry being 
 a line passing through the center of the interspace in which the 
 spot is found, and parallel to the direction of the longitudinal 
 nervures. For example, in P^igs. 6 and 7, Plate 2, each spot is 
 bilaterally symmetrical about the axis HII. The same law holds 
 for the spots represented in Figs. 8-14 and 1(3. 
 
 (b) Spots tend to appear not in one interspace only, but as a row 
 occujjj'ing homologous places in successive interspaces. Indeed we 
 almost always And similar spots arranged in linear series, each sim- 
 ilar in shape and color to the others and occupying the center of its 
 interspace. The rows of spots represented in Figs. 8-14 and 16 
 will suffice to illustrate this law. 
 
 It is interesting to notice that bands of color are often made by 
 the fusion of a row of adjacent spots ; and, conversely, chains of 
 spots are often formed by the breaking up of bands, leaving 
 a row of spots occupying the interspaces. Many instances of this 
 
184 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 are to be seen in certain specimens of various species of the 
 Heliconidae. For example, in Heliconius eucrate (Fig. 58, Plate 
 4) I have observed that certain specimens show a row of distinct 
 spots in place of the, usually entire, band which crosses the middle 
 of the hind wing. In fact, the vast majority of bands can be 
 analyzed into a series of similar elements, each element occupying an 
 inters])ace. Thus, in Plate 2, Fig. 17, which represents a wing of 
 Saturnia spini, the band seen crossing the wing parallel with its 
 margin is made up of a scries of fused crescents, each crescent 
 occupying an interspace. 
 
 If, on the other hand, this band were to break away from the 
 ncrvures, the result would be a series of crescent-shaped spots each 
 occupying the center of an interspace. It is very interesting to 
 observe the manner in which bands degenerate and disajipear. 
 Numerous opportunities for doing this may be had among the Heli- 
 conidae. In some species, as in Melinaea {)arallelis, hardly any two 
 specimens are alike in the condition of the black band across the 
 middle of the hind wings. The most common m.ethod of disappear- 
 ance is a shrinkinff aioay of the band at one end. This is well illus- 
 trated in Figs. 84-87, Plate 7, which represent a sort of " Mercator's 
 Projection " of the wings of Mechanitis isthmia (for explanation of the 
 plan of projection see page 207.) Fig. 84 represents a male, showing 
 a well-marked band of hardly separated spots extending across the 
 middle of the hind wing. P'ig. 87 shows a female in which the 
 spots are thinner and more crescentic and the separations much 
 more marked. Fig. 85 is also drawn from a female, in which it will 
 be seen that tlie band has shrunk away leaving only a portion of it 
 at the right, and in Fig. 8(5, which represents another female speci- 
 men, only one faint spot is left. 
 
 It is very common to find bands shrinking away at one end. 
 Sometimes, however, they shrink away at both ends, and very often 
 they break up into a row of spots, which may then contract into the 
 centers of their interspaces and finally disapi)ear. It is worthy of 
 note that it is very rare to find a band breaking at tlie middle of its 
 length and each half receding from the other. Such a case is, how- 
 ever, shown by Melinaea parallelis (see Fig. 82, Plate 7), where one 
 sometimes finds specimens in which the black band across the middle 
 of the hind wings is complete and utibroken ; whereas in other 
 specimens, as in F'ig. 82, it is partially broken in the middle, and in 
 still otliers the bi-eak has become a wide gap by the drawing away of 
 the halves of the band from each other. 
 
MAYEH: COLOR AND COLOK-l'ATTKHNS. 185 
 
 We st'c, tlic'ii, tliat it is very common to find b;ui<l.s shrinking 
 aw.ay from either eml, but very rare to find them broken in the 
 middle region. This, however, is only a special case of the law 
 enunciated by IJateson ('94), that the ends of a linera series are more 
 variable than the middle. Almost any row of spots also e.\hibit.s 
 the same law, in that the spots occnpj'ing the middle portions of 
 the row are similar one to another, while those at the ends of the 
 series depart more or less from the type. (See Figs. 10-13, 
 IMiU-e 2.) 
 
 The position of si)Ots which are situated near the edge of the 
 wing is largely controlled by the wing-folds or creases. In Meli- 
 naea egiiia (Fig. 90, Plate 8) there is a row of white spots near the 
 outer edges of the wings, and each of these spots is cut in two by a 
 narrow black line which extends along the wing-fold. Also in Cera- 
 tinia vallonia (Fig. 81, i'late 7) and in many other forms of the 
 Danaoid Ileliconidae one often finds two creases in a cell, and in 
 this case tiiei'c are two marginal spots, one on each crease. In 
 many other cases, however, the marginal spots are double in each 
 cell, although there is but a single wing-fold ; the spots in these 
 cases are situated at some distance on either side of the fold. (See 
 Figs. 95, 96, Plate 8.) Another very common condition is e.\'em- 
 plified in Fig. 88, Plate 7, where there is a single marginal spot 
 situated upon the wing-fold in each cell. 
 
 (8) DeUiiled Discussion of the Z,aws of Color- Patterns. Figs. 
 ()-14 and Ki, Plate 2, are taken from special cases which serve 
 to illustrate the two chief laws of color-pattern, i. e., that spots tend 
 to be bilaterally symmetrical about an axis (HII, Figs. G, 7) passing 
 through the center of the cell parallel with the nervures ; and 
 also, that spots of similar shape and color tend to be repeated in 
 a I'ow of adjacent cells. 
 
 In Fig. 7 the spots are separated in the middle, but still incline 
 outward sj'mmetricaliy from the center ; indeed, instances of double 
 spots are very common. In such cases, however, each half spot is a 
 retleetioii of its mate on the other side of the axis passing through 
 the center of the cell. 
 
 Fig. 8 represents various eye-spots found in the Jlorphos, and 
 will serve to illu.strate the laws of eye-spots which have been enunci- 
 ated by Scudder ('89) and Bateson ('94) . These spots occupj' the 
 center of the cells in which they are found. In cell II, for example, 
 is a large eye-spot with a crescent in its center, and it will be 
 
18*5 BULLETIN: MUSFAJM OF COMl'ARATIVE ZOOLOGY. 
 
 observed that this cresoent follows the general law and is bilaterally 
 synunetrical abont the usual axis.' 
 
 Fig. 9 shows the law of repetition of some very eoniplex spots, each 
 being bilaterally symmetrical. It is found in Parthenos gambrisius. 
 
 Figs. 10 and 11 represent Ornithoptera urvilliana and (). prianius 
 respectively. In Fig. 10 we see an instance of a spot within a spot, 
 and in Fig. 11 an even more complex case, for here there are three 
 systems of spots one within another. 
 
 Fig. 1'2 represents the marginal markings found in Ilestia jasonia 
 and Fig. 13 Ilestia leuconoe var. clara. These two examples are 
 intended to illustrate the fact, that, although the markings are 
 situated iqyon the nervures, they are bilaterallj'^ symmetrical not 
 about the nervures as axes, but about the usual axis passing midway 
 between the nervures. In Fig. VI it will be seen that the two 
 curved markings situated upon nervures l"* and 2, and projecting 
 into cell P, are bilaterally symmetrical only in reference to the axis 
 through the middle of the cell. 
 
 In allied species the s]>ot situated upon nerviu'e 1'' is often absent. 
 The system of markings is therefore undergoing degeneration at thi.s 
 end (cf. Fig. 18, cell V) . The curved mark upon nervure 5 (Fig. 
 12) projecting into cell V is plainly symmetrical with respect to its 
 fellow in the opposite side of cell V, and not with its near companion 
 whidi projects into cell IV. The same is also true in the case of the 
 spots in cell VI. 
 
 In Fig. l-'} the spots appear at the first glance to be bilaterally 
 symmetrical about both nervures and centers of cells, but in cell IV 
 the marking situated on nervui-e 4 does not quite reach to the cen- 
 ter, and it is interesting to observe that its fellow on nervure 5 also 
 falls short of reaching the center and is therefore symmetrical with 
 respect to the other curved spot in cell IV. This case also furnishes 
 an instance of a break in the middle of a linear series. 
 
 Ficr. 14 is taken from the inider surface of the hind wing of 
 Papilio emalthion. It serves to illustrate the fusion of two orig- 
 inally separate rows of spots. In this case the crescent-shaped spots 
 above have fused with the rectang>ilar ones below, so as to inclose 
 a portion of the ground color of the wing. Sometimes two rows of 
 
 1 A veiy beaiitlfiU cxcei)tioii (Fig. ID, IMiite 2) to this rule for the crescents found in oye- 
 spots is seen in the under surface of tlie tore wint; of Missanga patinia Moore. It will lie 
 noticed that the large blaclc crescent found in this beautiful eye-spot is !I0° away from its 
 usual position. This is the only exception of the sort known to nie. 
 
MAYER: COLOR AND COLOR-PAT'PERTS'S. 187 
 
 spots of different colors fuse, giving a chain of spots wliicli arc of 
 one color above and anotlier below. 
 
 In Fig. 16 the spots composing the row 15M are blue (dark) 
 above, and red (light) below. It will be observed that the color is 
 bilatei'alljf symmetrical, as usual, about the axis tlirougli the middle 
 of the cell. Such bicolored spots are often due to a simple fusion, 
 as before stated ; but sometimes they may, perhaps, be intrinsically 
 bicolored. 
 
 Fig. 15 is a beautiful instance of an exception to the general rule 
 that spots are bilateral about the axis through the center of the cell. 
 It is taken from Ornithoptera trojana Staudinger.' The light spots 
 represented near the outer edge of the wing are of a brilliant irides- 
 cent green. It is evident that they are distinctly bilateral with 
 respect tothe tier v ur es ; especially is this true of the pair adjacent to 
 nervure 1 . Ornithoptera brookiana Wallace illustrates another 
 exception, though in a less marked degree.^ Otlier allied species of 
 Ornithoptera, however, would seem to show that these apparent 
 exceptions may have been derived from forms whicli exhibited two 
 spots in each cell and followed the usual rule. These are the only 
 instances of such exceptions known to me. I do not doubt, how- 
 ever, that fui-ther study would reveal others. 
 
 In Fig. 17 an exam|)Ie is given of the peculiar kind of eye-.spots 
 found in the Saturn idae. The sj)ecies from which the figure was 
 taken is Saturnia spini. It will be seen that this so-called eye-spot 
 is quite different in formation from the ocelli of butterflies. It is 
 simply a series of curved cross-bands between nervures, arranged 
 symmetrically on both sides of the cross vein CC. The "eye- 
 spots " upon the wings of Attacus luna and in the genus Telea are 
 also of this sort. True eye-spots, however, similar to those found 
 among the Morphos and Satyridae, occur in moths, as in the apex 
 of the fore wing of Samia cecropia, Callosamia promcthea, etc. 
 " False " eye-spots are also found on the wings of butterflies ; in 
 Vanessa io, for example, the so-called eye-spot of the fore wing has 
 been shown by Dixey ('90) to be made np of a series of fused 
 spots. It will be remembered that Merrifleld ('94, Plate 9, Fig. 4) 
 caused this " ocellus " to break up into its constituents by subjecting 
 the pupa to a tem]>erature of 1° C The ocellus upon the hind wing 
 of Vanessa io is no doubt a true eye-spot; the only evidence which 
 
 ' See Watkins, '91, Plate i. 
 'See Hewitaon, '5C-'7(i, Vol. 1. 
 
188 BULLETIN: MUSFAIM OF COMPARATIVE ZOOLOGY. 
 
 miglit lead one to infer that the occlhis of the fore wing was of the 
 same character is, that an aberrant form is sometimes found in 
 nature having the " eye-spots " on botli fore and hind wings 
 obliterated, thus indicating a possible connection between the two 
 (see South, '89). 
 
 Kig. 18 is intended to illustrate the process of degeneration occur- 
 ring in bands. Band BU is rej)resented as breaking down by the rare 
 method of parting in the middle. Example, Melinaea parallelis. 
 Band EE is degenerating at one end ; this is a very common 
 method. 
 
 Figs. 20-28 represent hypothetical conditions not found in nature ; 
 all being contrary to the conditions of the laws which have just been 
 stated. 
 
 In Fig. 20 row RR presents three spots for each cell. I believe 
 this has not been found in nature, but I should not be surprised if 
 it were discovered, for it is not contrary to any of the laws. 
 
 Row CC, on the other hand, is contrary to the law of bilaterality, 
 the crescents not being bilateral about axe.s passing thi-ough the 
 middle of the interspaces parallel with tlie longitudinal nervures. 
 
 Fig. 21 is intended to show a series of spots arranged side by 
 side in twos in each cell, and of different colors. This, I believe, is 
 impossible, for it is contrary to the law of bilaterality of color 
 arrangement about the usual axis (HH, Figs. 6, 7). 
 
 In Fig. 22 there are several conditions which are impossible ; e. </., 
 an eye-spot situated upon a nervure is never seen in nature, also 
 two spots originally side by side, as in cell III, never rotate around 
 each other so as to come to lie one above the other. Spots often 
 move, however, as shown by the arrows in cell IV, thus giving 
 rise to fusions ; or they may move away from each other, causing a 
 wider gap between the rows. In cell I'' are shown two looped 
 spots. One form (A) is quite usual, being found indeed in Cynio- 
 thoe caenis Drury.' The other form of spot (D) is an impossibility, 
 not being bilaterally symmetrical. 
 
 Fif. 23 illustrates other impossibilities in color-pattern, none of 
 them, of course, being found in nature. For example, one never 
 finds a row of slanting spots such as SS. Also one never sees a 
 row of similar spots in alternate interspaces, such as is shown in 
 DD, for this would be contrary to the law that similar spots are 
 repeated in a row of adjacent interspaces. These last four diagrams 
 
 1 See Cramer C17T!)-'82), Vol. 2, Plate 140. 
 
MAYER: COLOR AND COLOR-l'ATTERNS. 189 
 
 (Figs. 20-'2H) have been introduced merely to give an idea of the 
 curiously strict limitations wliich nature has imposed upon the differ- 
 entiation of the color-pattern. Many beautiful effects might have 
 been produced, such for example as that of alternate interspaces 
 showing (lifferent colors, but this is not seen in nature. 
 
 Ft is interesting to recall tlie fact, that the colors themselves are 
 impure and by no means so brilliant as the}-, perhaps, might have 
 been, had Natural Selection been more severe in regard to color. 
 
 There is doubtless some physiological reason whj' spots almost 
 invariably appear and disappear in {Xm middle of the inters}Kice.% and 
 when we know more of tlie anatomical and histological conditions 
 of tlie wing during the development of the colors, we may be able to 
 discover it. It will be remembered that in the developing pupal 
 wings of Callosamia promethea and Danais plexippus I found that 
 the colors first made their appearance in the interspaces, and finally 
 spread out so as to tinge over the nervures. 
 
 (4) Oriffin of Color- Variations. There is everj' reason to 
 believe that all kinds of spots and bands, which are essentially 
 only fused spots, may appear or disappear in any indivi<lual 
 specimen without going through a long course of Natural Selection 
 and slow phylogenetic differentiation. Darwin and Trimen ('71) 
 and Bateson ('94) have demonstrated that this is true for eye-spots. 
 In the Ileliconidae I have found that bands and rows of spots are 
 very variable in different specimens of the same species (see Plate 7, 
 Figs. 84-87). 
 
 There is a large and widely scattered literature recording the 
 appearance and disappearance of colors and markings upon the 
 wings of Lejjidoptera. Limits of time and .space prohibit my doing 
 justice to it here, but it may be well to call attention to a very few 
 of the more recent papers upon the subject. Many of the color- 
 aberrations recorded in tliis list of papers may be due to the direct 
 influence of environmental conditions upon the individual, but others 
 are no doubt true sports or, to speak crudely," congenital " variations, 
 and might under favorable conditions of life become the ancestors 
 of neio varieties or species. It seems highly probable that new 
 S])ecies often arise from just such sports in the manner so frecpiently 
 and ably expounded by Bateson. 
 
190 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 1'aktial BiDLioGRAi'iiv OF ukmaiikahle Color-Aherrations in 
 
 Llil'IDOPTERA. 
 
 Bairstow, S. D. 77. Ent. Mo. Mag., Vol. 14, p. 67. (Zygaena fllipendulae.) 
 Bkan, T. E. '95. Can. Ent., Vol. 27, p. 87-93, Plate 2. (Nemeophila petrosa 
 
 and varieties.) 
 Benson, E. F. '83. Entomologist, Vol. 16, p. 210. (Arge galathea.) 
 Bleuse, L. '89. Feuille jeiin. Natural., 19 Ann., p. 142-143. 
 Breionet, F. '90. Bull. Soc. Ent. France, (6), Tome 10, p. 29-30. (Theela 
 
 rubi ; Melithaea athalia.) 
 CARiiiN<iTON, J. '78. Entomologist, Vol. 11, p. 97, Fig. (Cidaria suffumata.) 
 Cakbington, J. '83. Entomologist, Vol. 16, p. 1, Fig. (Calliraorpliadominula.) 
 Carrington, J. '88. Entomologist, Vol. 21, p. 73, Fig. Editorial Note. (Arctia 
 
 caia.) And numerous other papers in the Entomologist. 
 Clark, J. A. '89. Entomologist, Vol. 22, p. 145-147, I'late 6. (Tripliaena 
 
 comes.) 
 CocKERELL, T. 1). A. '86. Eutomologist, Vol. 19, p. 230-231. (Epinephele 
 
 tithonus.) 
 CocKERELi,, T. n. A. '88. Entomologist, Vol. 21, p. 189. (Pieridae.) 
 Cockerei-l, T. D. a. "89. Entomologist. Vol. 22, p. 1-6, 13, 20-21, 26-29, 64- 
 
 56, 98-100, 125-130, 147-149, 185-186, 243-245. 
 Dewitz, H. '85. Berlin. Ent. Zeitschr., Bd. 29, p. 142, Taf. 2. (Precis amestris.) 
 Editors of Ento.molooist, '78. Entomologist, Vol. 11, p. 169-170, Plate 2. 
 
 (Vanessa atalanta and several Lepidoptera.) 
 Edwards, W. H. '68. Butterflies of North America. (Numerous plates.) 
 Fettig, F. J. '89. Feuille jeun. Natural., 19 Ann., p. 84. (Variations of 
 
 Lepidoptera in Alsace.) 
 Fitch, E. A. '78. Entomologist, Vol. 11, p. 50-61, Plate. (Colias edusa.) 
 Goss, H. '78. Entomologist, Vol. 11, p. 73-74, Fig. (Chelonia villica.) 
 ObbrtiiDr, C. '89. Bull. Soc. Ent. France, (6), Tome 9, p. 74-76. 
 OnERTiiiJR, C. '93. Feuille jeun. Natural., 24 Ann., p. 2-4. 
 PoujADE, G. A. '91. Ann. Soc. Ent. France, (6), Tome 11, p. 597-598, 
 
 PI. 16. (Thais rumina.) 
 Richardson, N. M. '89. Ent. Mo. Mag., Vol. 25, p. 289-291. (Zygaena filipen- 
 
 dulae.) 
 Scidder, S. H. '89. Butterflies of Nevf England, p. 1213. (Bibliography of 
 
 variations of Pieris rapae.) 
 South, R. '89. Entomologist, Vol. 22, p. 218-221, Plate 8. (Various Vanessidae.) 
 Speyer, a. '74. Stettiner Ent. Zeitung, Bd. 35, p. 98-103. 
 
 Thiele, H. '84.- Berlin. Ent. Zeitschr., Bd. 28, p. 161-162, Fig. (Apatura iris.) 
 TuTT, J. W. '89. Entomologist, Vol. 22, p. 15, 100-161. (Melanic Agrotis 
 corticea and pale variety of Lycaena bellargus.) 
 
 (5) Climate and Mela/iinni. I.oi'd Walsingham ('85), in liis 
 presidential address before the Yorksliire Naturalists' Union, brought 
 forward the idea, that, although Arctic insects might be perfectly 
 
MAYER: COLOR AND COLOR-l'ATTERNS. 191 
 
 able to withstand tlie most severe cold while in hibernation during 
 the winter, it is of great importance for them to absoi'b as much heat 
 as possible during the short summer. He placed several species of 
 lepidopterous larvae upon a snow surface exposed to bright sunshine. 
 The snow melted at different rates under the various larvae, and 
 in two hours the darkest insect had sunk by far the deepest into the 
 snow, proving that it was the best absorber of heat. This ingeni- 
 ous experiment of Lord Walsingham should be made the beginning 
 of an extensive and careful research. 
 
 Chapman ('88) has shown that it may be of advantage to moths 
 inhabiting wet regions to dis])lay dark colors, or become melanic. 
 His observations were made upon Diamea flagella, and he says that 
 upon one showery afternoon he observed that one side of the tree 
 trunks was wet and dai-k in color ; the other side being dry was 
 paler. " As a consequence, the dark specimens of flagella were very 
 conspicuous upon the dry portions, hardly visible on the wet, whilst 
 with the ordinary form the conditions were reversed, those on the 
 wet bark were conspicuous, those on the dry much less so." Per- 
 haps the dull coloration of Arctic moths may be partially due to the 
 effect of the somber backgroinid of rocks in the regions which they 
 iidiabit. 
 
 (6) Relation between Climate and Colors of Papilios. It is well 
 known that the Lepidoptera in the Tropics display the richest 
 variety and greatest number of colors. I have counted the colors 
 exhibited by the 22 species of Papilio enumerated by Edwards as 
 inhabiting North America north of Mexico, and also those which 
 are displayed by the 200 species of Papilio named in Schatz's list as 
 found in South America. The "colors" were determined by com- 
 parison with the colored plates in Kidgway ('8()). 
 
 In this manner it was determined that the North American 
 Papilios exhibit 17 colors, viz., black, brown, primrose-yellow, canary- 
 yellow, sulphur-yellow, orange, white, greenish white, apple-green, 
 cream-color, azure-blue, sage-green, rufous, ])earl-gray, indigo-blue, 
 iridescent blue, iridescent green. 
 
 On the other hand the South American Papilios exhibited 36 
 colors, viz., black, translucent black, brown, white, canary-yellow, 
 citron-yellow, olive-yellow, primrose-yellow, chrome-yellow, straw- 
 yellow, gamboge-yellow, cream-color, greenish white, apple-green, 
 malachite-green, emerald-green, sage-green, slaty green ii-idescence, 
 pea-green, azure-blue, iridescent Berlin-blue, indigo, pearl-blue, 
 
192 BULLETIN: MUSFAJM OF COMPARATIVE ZOOLOGY. 
 
 glaucous blue, salmon-buff, (f'cru-drab, flesh-color, coral-red, rose-red, 
 vermilion, rufous, geranium-red, geranimn-piiik, olive-buff, iridescent 
 geranium-pink (as in P. zeuxis), and transparent areas. 
 
 As "200 species in South America display but 3() colors, while 22 
 in North America show 17, it follows that, while the number of 
 species in South America is 9 times as great as in North America, 
 the number of colors dis])la3'ed is only a little more than twice as great. 
 The richer display of colors in the Ti'opics, therefore, may be due 
 simply to the far greater number of species, which gives abetter oppor- 
 tunity for color-sports to arise, and not to any direct influence of the 
 climate. The number of broods,' also, which occur in a year is much 
 greater in the Tropics than in the Temperate Zones, so that the Troj)- 
 ical species must possess a correspondingly greater o[)portunity to 
 vary. 
 
 V. The Causes which have led to the Development and 
 Preservation of the Scales of the liEi'iDoi-TEUA. 
 
 (1) Ea'jyeriments and Theory. It is well known that the scales 
 of Lepidoptera are morphologically identical with hairs. Indeed, a 
 graded series from simple hairs, such as arc found covering the 
 body-siH'face of most Arthropods, up to perfectly developed flat 
 scales bearing well differentiated striae may usually be found u])()n 
 one and the same insect. 
 
 It is also remarkable that the color-bearing scales of beetles have 
 been developed in the same manner as those of moths and butterflies, 
 and that in this case also hairs liave become differentiated into scales 
 which are precisely similar in a])pearance to tho.se of the Lepido]>tera 
 (see Dimmock, '88). 
 
 This is only another of the numerous instances met with in nature 
 where similar conditions of selection have developed complex organs 
 which are similar in appearance, though found in widely .sej)arated 
 groups. A list of papers I'clating to the development of scales has 
 been given by Dimmock ('88, p. 1-11). 
 
 Most of the haii-s which cover the body-surface in Arthropods arc 
 true sensory structures, the axis of each of which is a protoplasmic 
 process from a single cell of the hypodermis, which lies below the 
 cuticula. They liave probably been developed because the cuticula, 
 
MAVKU: COLOR AND COLOli-l'ATTKRNS. 193 
 
 bc'iiij^ liiiid, chitiiious, luid iiiHoxihlo, would seivo but poorly as a 
 ta(!tile or sensory surface. 
 
 Of course no one would venture to ascribe any sensory function to 
 the scales which cover the wing-nionibranes of the Le])idoptera. 
 We may, however, make severiil more or less reasonable hypotheses 
 concerning tlic ])robable uses of the scales, and by testing these sup- 
 ])ositions arrive ])erhaps at some plausible explanation of their reten- 
 tion and the complex development which they have undergone. 
 
 (1) They may have caused the wings of the ancestors of the 
 T.epidoptera to become more perfect as organs of flight, by causing 
 the frictional resistance between the air and the wing-surface to 
 become more nearly an optimum. 
 
 (2) The appearance and develo|)ment of the scales may have 
 served, as Kellogg ('94) has suggested, " to protect and to strengthen 
 the wing-membranes." 
 
 (3) The present development of the scales may be due to the 
 fact that they displayed colors which were in various ways advan- 
 tageous to the insects. 
 
 Concerning the first of these three hypotheses, the wing has, 
 broadly speaking, two chief functions to perform in flight. It must 
 beat more or less downward against the air, and must, in addition, 
 glide or cut through the air, supporting the insect in its flight. For 
 the mere beating against the air a relatively larc/c co-cflicient of 
 friction between the air and the wing might be advantageous ; but 
 for gliding and cutting through the air a «w?a// co-efficient of friction 
 would certainly be an advantage. There must therefore be an 
 optimum co-eflicient of friction, which lies somewhere between these 
 two. 
 
 In order to determine the co-eflicient of friction between the wing 
 and the air, use was made of a method which, in one form or 
 another, has long been known to engineers ; that is, of observing the 
 ratio of damping of the vibrations of a pendidum. 
 
 It is well known that when a pendulum is swinging free, and 
 uninfluenced by any fi-ictional resistances, the law of its motion is 
 expressed by the formula, 
 
 (i) d:= A sin ^ t 
 where d is the displacement of the pendulum from its middle 
 ])osition after the interval of time t, A is the maxinnim displace- 
 ment and T the time of a complete vibration, back and forth. If, 
 
194 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY, 
 liowever, frictioiial resistances interfere, the formula becomes, 
 
 (2) (1 = A c-K' sin — t 
 
 1 
 
 —log d T, 
 
 (5) Hence, K = -r~j --■ .,' ^. , , or if t = T, 
 
 ^ ' ' A log e sni '2ir t- ' ' 
 
 —log d 
 (^) K = ^^^ 
 
 where K is a constant dependent upon the friction, e is the base of 
 the Napierian system of logarithms and T, is the time of a complete 
 vibration, wliich may be different from the T, i-epresentiiig the time 
 of vibration when not under tlie influence of friction. 
 
 The plan was, then, to attach the wing of some large butterfly or 
 moth to the end of a short, light pendulum in such a way that it 
 would either fan against the air, or cut through it, and then to 
 observe the ratio of damjjing of the pendulum's vibrations. A 
 drawing of the pendulum with a wing attached is given in Plate 1, 
 Fig. 3. The wing is here shown in the position for "cutting or glid- 
 ing" through the air. It would be in the position for fanning against 
 the air, if it were rotated 90°. 'IMie pendulum was made of brass 
 and steel, the ends being of brass and the slender middle portion of 
 steel. Its vibrations were read off upon an arc graduated in milli- 
 meters. The readings were certainly accurate down to 0.5 mm. 
 The pendulum was hung upon a steel knife edge (n, n, P^ig. 3), 
 which rested upon Arm level glass bearings. The pendulum was 
 24.21 cm. long, and weighed 19.G1 grams. Its time of vibration 
 (Tj) was 0.877 seconds. This rate of vibration was practically 
 unaltered when a wing was fastened to the end of the pendulum, 
 the reason being that the wings were very light, the heaviest, that of 
 Samia cecropia, weighing only 0.038 grams. The wing to be experi- 
 mented upon was fitted into a deep, narrow slot at the fi-ee end of 
 the pendulum, and then cemented in by means of a little melted 
 beeswax. It thus became a perfectly rigid part of the pendulum 
 itself. 
 
 The pendiilmii with wing attaclied was deflected through a known 
 arc, read off upon the millimeter scale, and its reading at the end of 
 the first swing carefully observed. Then if A be the initial deflec- 
 tion, which we may call unity, and if d be the reading after the first 
 
 swing, the ratio of damping is given by the expression -j. In experi- 
 menting with a fore wing of Samia cecropia "fanning the air," it 
 
MAYKR: COLOR AND COLOR-l'ATrEUNS. 195 
 
 was found, as tlie mean of many trials, that this ratio of damping 
 was 0.919, that is to say, the amplitude of tlie 2d swing was 0.919 
 as great as the amplitude of the 1st, that of the 8d only 0.919 as 
 great as tliat of the '2d, and so on. The scales were tlien carefully 
 removed from the wing-membranes, by means of a camel's hair brush, 
 and by again testing the vibrations it was found that the new ratio of 
 damping was 0.917. This is so near the value of the ratio of damp- 
 ing with the scales on (0.919), that it may be considered identical, 
 the difference being due to errors of experimentation. 
 
 Hence we must conclude that the presence of the scales upon the 
 wing-membrane has not altered, appreciably, the co-eflicient of fric- 
 tion which would exist between scaleless wing-membranes and the 
 air. The results indicate rather, that when the scales appeared upon 
 the wings of the scaleless, clear-winged ancestors of the Lepidoptera, 
 the po-efKcient of friction remained unaltered. This tempts one to 
 the further conclusions, that the co-efficient of friction between the 
 air and the wings was already an optimum in these clear-winged an- 
 cestors before the appearance of the scales, and therefore that Natural 
 Selection would operate to keep it unaltered. 
 
 A wing of Samia cccropia cut so as to give it the same shape and 
 dimensions as one of Morpho menelaus, gave an identical damping 
 ratio. I conclude that the co-efficient of friction miiy be the same 
 for both moths and l)utterflies, at least for those which move theii- 
 wings at about the same rate in flight. 
 
 It was found in the case of the Samia cecropia wing, that when 
 it was vibrated in the position for " cutting through " the air, the ratio 
 of damping was 0.991. It will be remembered that, when the wing 
 "fanned" the air, this ratio was 0.917. We may find the ratio be- 
 tween the resistance encountered in " fanning" and that encountered 
 in "gliding" through the air by substituting these values in equa- 
 
 — log d 
 tion (4), K = ATTbi"^- 
 
 Thus for fanning, -r- = 0.917 and Tj = 0.877. Making A unity, 
 
 —log 0.917 
 
 K= ,, ,,,^, = 0.1. 
 
 0.877 log e 
 
 d 
 
 In cutting tln-ough the air, -r =0.991 and Tj as before = 0.87 * . 
 
 _losr 0.991 
 
 Hence in this case K =:jr~7r,,^-. = 0.01. 
 
 0.877 loii <J 
 
190 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 The wing, then, encounters at least 10 times the resistance in fanr 
 niiig that it does in gliding through the air. It should be said that 
 this last experiment is somewhat crude, for the wing necessarily 
 could not be made to cut the air with that delicate precision whicli 
 is probably realized by the insect in flight. I should not be 
 surprised, if in nature the insects encountered at least '20 times 
 the resistance in beating the air, that they do in merely gliding 
 through it. 
 
 Concerning Mr. Kellogg's supposition, that the scales were devel- 
 oped to "protect and to strengthen the wing-membranes," I will 
 admit that they may serve in some slight degree to protect the wing- 
 membranes from scratches, etc. ; but I am unable to accept his con- 
 clusion, that they strengthen the wing-membranes, any more than 
 that the shingles upon a roof serve to add strength to it. The 
 wing-membranes themselves are tough, elastic, and not easily torn o-r 
 scratched, and the scaleless wings of the Neuroptera and Ilj'me- 
 noptcra are very rarely found torn or scratched in nature. 
 
 In 1858 Mr. Alexander Agassiz called attention ('59) to the fact, 
 that " the nervures of the wings of butterflies are so arranged as to 
 give the greatest lightness and strength ; they are hollow, with theii- 
 greatest diameter at the base of the wing, the point of greatest 
 strain, their diameter gradually diminishing to the edge of the 
 membrane. If a section be made across such a wing parallel to the 
 axis of the body, we And very much the arrangement which has 
 been experimentally proved by P'airbain and vStephenson as giving 
 the greate.st strength of beams, as exemplifled in the tubular bridge. 
 We find the strongest nervure placed either on or near the anterior 
 edge of the upper wing ; there is no such nervure on the lower 
 wing, all being of nearly the same size, as such a one would have 
 prevented the elasticity of the wing from assisting the flight to 
 any considerable extent." Mr. Agassiz has informed me that he 
 carried out an extensive series of experiments upon the rigidity of 
 the wings of various species of Lepidoptera. lie placed little 
 platinum strips upon the wings and observed the extent of the 
 bending produced. His results demonstrated that the Sphinx moths 
 possess by far the strongest wings, and that the Danaoid and 
 Acraeoid Heliconidae have very weak wings. The reason for this 
 probably lies in the fact, that the Sphinx moths move their wings 
 with great rapidity, while, according to Bates {'(i2) and all sub- 
 sequent observers, the Heliconidae have a slow flight. 
 
MAYKK : COLOR AND COLOK-rATTKUKS. 197 
 
 As tlie scales liavc been developed not because they aided the 
 insects in flight or strengthened the wings, their retention must 
 have been due to some other cause, probably to their display- 
 ing colors which were advantageous to their possessors in various 
 ways. As Dimmock ('8H) says, " it is only in insects where certain 
 kinds of brilliant coloration have been developed that one finds 
 scales." Indeed, I believe that the vast majority of the scales 
 found in Lepi(lo])tera are merely color-bearing organs. Thej' prob- 
 ably first made their appearance upon small areas of the wings, 
 perhaps adjacent to the body, and were merely colored hairs, sim- 
 ilar to those of the surface of the body, which had grown out upon 
 the wings. In this position they displayed some color which was 
 of advantage to the insect ; perhaps serving to render it less con- 
 spicuous than formerly. Under these circumstances they would 
 naturally be preserved through the operation of selection until 
 finally they became modified into true scales; just as the hairs in 
 the Coleoptera have undergone a similar modification. If this 
 be true, it is easy to see how they might spread out over 
 the surfaces of the wings until the whole wing became covered 
 with scales. 
 
 (2) Summuri/ of Condtmons. The scales do not aid the insects 
 in flight, for the wings have precisely the same eiticiency as organs 
 of flight when the scales are removed. The phylogenetic appearance 
 and development of the scales upon the scaleless ancestors of the 
 Lepidoptera did not in the least alter the efficiency of their wings as 
 organs of flight. This efficiency of their wing surfaces was probably, 
 therefore, already an optimum liefore the scales appeared. The 
 .scales do not appreciably strengthen the wing-membranes, that 
 function being performed by the nervures. The majority of the 
 scales are merely color-bearing organs, which have been developed 
 under the influence of Natural Selection. 
 
 PART B. 
 
 COLOR-VARIATIONS IN THE IIELICONIDAE. 
 
 I. Genkuat. Causks which detkumine C'olokation i.n tub 
 
 IIeliconidae. 
 
 In 1801, after eleven years of study within the forests of South 
 America, Bates read his, now classic, paper upon the life and habits 
 
19S BULLETIN: MUSEUM OF COMPARATIVE Z()(')LOGY. 
 
 of the Helicoiiidae of tlic Amazon region. In it he first brought 
 forward his ingenious tlieory of Mimicry — a theory which, under 
 tlie able interpretations of Wallace and Fritz Muller, and in more 
 recent times, under the impetus of the zeal of their numerous disci- 
 ples, has yielded so much that is of interest to scientific men. 
 
 The rioliconidae are, above all, creatures of the forest, and ]}ates 
 found that the number of species increases as one travels inland 
 from the Lower iVmazons towards the eastern slopes of the Andes, 
 so that the liot Andean valleys near Bogota, or in Ecuador, contain 
 perhaps the greatest number. In their range they are resti-icted to 
 the Tropics of the New World. Only two species, Dircenna klugii 
 and lleliconins cliaritonius, extend so far north as the extreme South- 
 ern States of the United States, and none of them are found much 
 further south than iW S. Lat. 
 
 Bates and Felder first saw that the Heliconidae were naturally 
 divided into two distinct groups. One, the Danaoid Heliconidae, 
 consists of about twenty genera, all more or less closely related, and 
 evidently an offshoot from the great universal family, the Danaidae, 
 metnbers of which are found in both Hemispheres. The other grouj), 
 the true Heliconidae, is composed of two closely related genera, Jleli- 
 conius and Eueides. They are allied in structure to the Acrae- 
 idae and hence their name, Acraeoid Heliconidae. Schatz and llober 
 ('85-'92, p. 105) say of the Acraeoid Heliconidae: — They are an 
 offshoot of the great family Nymphalidae, which liave undergone a 
 remarkable <levelopment in the length of the fore wing, and in this 
 respect have been developed in a direction parallel with the Danaoid 
 Heliconidae. In their structure, however, they are quite distinct 
 from the Danaoid group. 
 
 Schatz has proposed a new classification for the Heliconidae. He 
 finds that the; genera Lycorea and Ituna, which Bates included among 
 the Danaoid Heliconidae, are very clo.sely allied to the Danaidae, he 
 therefore says that Dycorea should be placed among the Danaidae, 
 while Ituna is clearly midway between the Danaidae and the Dana- 
 oid Heliconidae. Schatz proi)o.ses tlie name " Neotropidae " for the 
 Danaoid Heliconidae. However, I think the name "Danaoid Heli- 
 conidae," being older and more descriptive of their relationship, 
 should by all means be retained. In this paper I shall follow Bates's 
 classification, and include among the Danaoid Helii^onidae the twenty 
 genera: Lycorea, Ituna, Athcsis, Thyridia, Athyrtis, Olyras, Kutre- 
 sis, Aprotopos, Dircenna, Callithomia, Kpithomia, Ceratinia, Sais, 
 
MAYER: COLOK AND COLOR-I'AT'l'KRNS. 199 
 
 Scada, Mechanitis, Na])eogones, Ithomia, Aeria, Meliiiaea, and 
 Tithorea. The Acraeoid Ileliooiiidae will tlioii consist of the two 
 remaining genera, llelieonius and Kueides. 
 
 Staudinger ('84-'88) records 458 species belonging to the Danaoid 
 grouj), and 150 belonging to the Acraeoid group. 
 
 Nearly all that we know concerning the early stages of the 
 Heliconidae is due to Wilhelm jNIUller ('8()), He gives figures and 
 more or less com|)lete descriptions of the early stages of Dircenna 
 xantho, Ceratinia eupompe, Ithomia neglecta, Thyridia theniisto, 
 Mechanitis lysimnia, and also of Ileliconius apseudes, If. eucrate, H. 
 doris, Eueides Isabella, K. aliphera, and E. pavana. Bates ('G'2, j). 
 596) says that he raised the larvae of Heliconius crato and Eueides 
 lybia. Schat/, and liober ('85-'92) figure the larva and pupa of 
 Ceratinia euryanassa. Edwards has given a detailed account of the 
 early stages of II. charitonius. 
 
 Miillcr found that the larvae of the Danaoid group feed on various 
 species of Solanum, while the genera Ileliconius and Eueides feed 
 upon the Passifloreae. The larvae are conspicuously colored, and 
 often gregarious ; they seem to take but little ]iains to liide them- 
 selves during the chrysalis stage, for Midler says that he has seen 
 the silver-spotted, white chrysalids of Ileliconius doris hanging in 
 great numbers in the near neighborhood of the larval food plant. 
 The mature insects also furnish a good example of what Wallace 
 ('67) designated as " warning coloration," for their tawny orange 
 and blac^k wings are very conspicuous as they sail slowly around in 
 circles, settling at frequent intervals in their lazy irregular flight. 
 
 Bates was the first to call attention to the circumstance that they 
 often possess a rather strong and disagreeable odor, and in 1878 
 Eritz Mtlller conlirmcd this observation for a number of the Heli- 
 conidae. He found, for e.xam])le, that the genera Ituna and Ilione 
 have a pair of finger-like processes near the end of the al)domen, 
 which can be protruded and then emit a rather disagreeable odor ; 
 and he also found that the Acraeoid Heliconidae, especially the 
 females, possess a disgusting odor. Seitz ('89), however, examined 
 about fifty species of Heliconidae and found that many of them 
 appear to have no odor. For example, he says that Ileliconius 
 eucrate and Eueide dianasa have no odor, but that some specimens 
 of Heliconius beskei, and Eueides aliphera have a horrid odor. 
 
 Whether they are odorous or not, it would seem that the Heli- 
 conidae have but few enemies to fear, for not one of the many 
 
200 BULLETIN: Ml'SKUM OF COMl'AIJATIVE ZOOLOGY. 
 
 skilled obsorvcrs wlio h;ive sUnlied them in their native haunts has 
 ever seen a bird attack them, and the onl_y ground for believing that 
 they are attacked rests upon the rather dubious evidence of a few 
 specimens found by Fritz Miiller having symmetrical pieces 
 apparently bitten out of the hind wings. Belt ('74) observed that a 
 pair of birds which were bringing large numbers of dragon-Hies and 
 butterflies to their j'oung never brought any of the Ilelicoiiidae, 
 although these were abundant in the neighborhood. In fact, Helt 
 was able to discover only one enemy of these butterflies, and that 
 was a yellow and black wasp, which caught them and stored them 
 up in its nest to feed its young. The Ileliconidae then, in spite of 
 their weak structure, conspicuous colors, and slow flight, enjoy a 
 peculiar immunity. 
 
 As is well known, Bates ('62) first called attention to the fact that 
 the Ileliconidae were "mimicked" or imitated both in color-pattern 
 and shape of wings by a number of other genera of butterflies and 
 even moths. Bates had no difliculty in showing that this mimicry 
 might easily be explained upon the ground that the Ileliconidae, on 
 account of their bad taste and .smell, were immune from the attacks 
 of birds and other in-sectivorous animals, and that therefore it gave a 
 peculiar advantage to a butterfly belonging to any other group not 
 thus protected, to assume the shape and coloration of the Ileliconidae ; 
 for then the birds could not perceive any difference between it and 
 the true Ileliconidae. Bates found that flftcen species of Pieridae 
 belonging to the genera Leptalis and Kuterpe, four Papilios, seven 
 Erycinidae, and among diurnal moths three Castnias and fourteen 
 Bombycidae imitate each some distinct species of the Ileliconidae 
 occupying the same district. He also found that all of these insects 
 were much rarer than the Ileliconidae which they imitated. In some 
 cases, indeed, he estimated the proportion to be less than one to a 
 thousand. Wallace ('89, p. 205), who has added so much to our 
 knowledge of this subject, aptly defines this kind of mimicry as an 
 "exceptional form of protective resemblance." 
 
 But by far the most remarkable tliscovery made by Bates was the 
 fact, that species belonging to different genera of the Ileliconidae 
 themselves mimic one another. Neither Bates nor Wallace was 
 able to give any satisfactory explanation of the cause of this latter 
 form of mimicry, for all of the genera of the Heliconidae are 
 immune. They therefore supposed it to be due to " unknown local 
 causes," or similarity of environment and conditions of life. 
 
MAYEH: COLOK AND COLOK-l'ATTKUNS. 201 
 
 Tims tlie iiiatter lestL-d until 1879, when Fritz Milller brouujht out 
 his well-known paper upon " Ituna and Thyridia, a remarkable 
 example of mimicry," in which he showed that both of these j^enera 
 arc i)rotected, yet they mimic each other. He also showed that this 
 mimicry miglit be due to Natural Selection brought about in the 
 following manner. It is possible that young birds, upon leaving the 
 nest, aie not furnished with an unalterable instinct which tells them 
 exactly what they should and should not eat; so they may try 
 experiments, and would then in all ])robability taste a few of the 
 Heliconidae before finding out that they were unfit to eat. Mtlller 
 then demonstrated that, if this supposition be true, it becomes a 
 decided advantage to the various species of Heliconidae to resemble 
 one another. Ilis reasoning was as follows : Let it be 8ui)po8ed 
 that the young and inexperienced birds of a region must destroy 
 1,200 specimens of any distasteful species of butterfly before it 
 becomes recognized as such, and let us assume further that there are 
 in existence 2,000 specimens of species A, and 10,000 of species B ; 
 then, if these species are different in appearance, each will lose 1,200 
 individuals, but if they resemble eacJi other so closely that they can- 
 not be distinguished apart, the loss will be divided pro rata between 
 them, and A will lose 200, and B 1,000; therefore A saves 1,000 or 
 50% and B saves only 200 or 2% of the total number of individuals 
 in the species; hence, while the relative numbers of the two species 
 are as 1 to 5, the relative advantage derived from the resemblance 
 is as 25 to 1. 
 
 Blackiston and Alexander ('84) have given a comj)lete mathe- 
 matical statement of Miiller's law, and have come to the conclusion 
 that, if tlie number of individuals destroyed is small compared with 
 the number constituting the species, the relative advantage is 
 inversely as the square of the original numbers ; but if the number 
 destroyed is large compared with the original number, the ratio of 
 advantage is much greater than the inverse squares of the original 
 numbers. Their deduction may be briefly stated as follows : — ■ 
 
202 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 Designation of Species 
 
 (1) Original number .... 
 
 (2) Number lost without imitation 
 
 (3) Remainders without imitation 
 
 (4) Number lo.st with imitation . 
 
 (5) Remainders with imitation 
 
 (6) Excess of remainders due 
 
 to imitation, or " abso- 
 lute advantage" (3) — (o) 
 
 (7) Ratio of excess to remainders 
 
 without imitation (()): (3), 
 ^proportional advantage. 
 
 (8) Ratio of proportional advan- 
 
 tage of B to proportional 
 advantajie of A. 
 
 (' 
 
 (a— e) 
 
 a 
 a+^b*^ 
 
 -'^] 
 a.+hj 
 
 be 
 
 a+b 
 
 a+b 
 
 B 
 
 b 
 
 = e 
 (b-e) 
 
 b 
 a + b ^ 
 
 b 1- 
 
 a+b 
 
 a e 
 a+b 
 
 a+b 
 
 a (a — e) 
 b (b— e)~ 
 
 (!zi) 
 
 It is evident, then, if e be small compared with a and b, that the 
 proportional advantage of R is to the proportional advantage of A 
 as a- is to b''. If, however, the loss (e) is great compared with a or 
 b, the relative gain for the weaker species becomes even greater than 
 the ratio of the squares of b and a. 
 
 If it be true, then, that young birds, when thej' leave the nest, do 
 not possess a directing instinct telling them what thej' should and 
 should not eat, but actually do experiment to some extent upon 
 various insects which they meet with, MUlIer's law is amply sufiicient 
 to account for the numerous cases of mimicry and remarkably close 
 resemblances Avhich are found among the .species of the Ileliconidae 
 themselves. 
 
 Unfortunately no direct experiments have ever been made upon 
 the feeding-habits of young South American birds, nor have the 
 contents of their stomachs been examined. There have been a few 
 exi)eriments, however, which seem to support the idea that some 
 animals do learn to associate an agreeable or disagreeable taste with 
 the coloration and appearance of their prey. It is well known that 
 Weismann ('82, p. 8BG-339) found that the black and yellow 
 larvae of Euchelia jacobaeae were refused by the green lizard of 
 Europe. He then introduced some young caterpillars of Lasiocampa 
 
MAYER; COLOR AND COLOR-PATTERNS. 203 
 
 rubi, which are very similar in appearance to those of Euchelia. 
 The lizards first cautiously examined tlie larvae, and finally ate them. 
 After this Weisniann reintroduced the E. jacobaeae larvae and the 
 lizards were seen to taste them, ap[)arently mistaking tliem for the 
 edible L. rubi caterpillars. 
 
 I'oulton ('87) carried out a most careful and well-conducted 
 research upon the protective value of color and markings in insects 
 in reference to their vertebrate enemies. lie experimented upon three 
 species of lizards and a tree-frog. Poulton combines his results with 
 those of other observers and presents them in the form of a table, 
 which certainly supports the suggestion of Wallace ('<37), that 
 brilliant and conspicuous larvae would be refused as food by some 
 at least of tlieir enemies. Poulton also shows that a limit to the 
 success of this method of defence (conspicuous larvae having 
 unpleasant taste or smell) would result from the hunger which the 
 success itself tends to ])roduce. In the Tropics, indeed, where 
 insectivorous birds and lizards are far more numerous than with us, 
 and where competition for food is great among them, " we may feel 
 sure that some at least would be sufficiently enterprising to make the 
 best of unpleasant food, which has at least the advantage of being 
 easily seen and caught." This last suggestion of Poulton certainly 
 seems reasonable ; moreover, it has occurred to me that young birds, 
 being but little skilled in the art of obtaining their food, might quite 
 often be forced by hunger to try various kinds of insects, and per- 
 haps even the Ileliconidae themselves. 
 
 Ueddard ('92, p. 153-167) reports the results of an extensive 
 series of experiments carried out by Mr. Finn and himself upon 
 marmosets, birds, lizards, and toads. lie arrives at conclusions which 
 are quite different from those of Poulton and others, but it appears 
 to me that his ex|)eriments were by no means so critically performed 
 as those of Poulton. He frequently threw larvae into a cage con- 
 taining many birds and observed them struggle for the })rey. It 
 may well be, however, that a bird would be quite willing to swallow 
 a very unsavory mouthful in order to prevent any of its companions 
 from, apparently, enjoying it. However, Beddard found that 
 toads will eat any insect without hesitation in spite of brilliant 
 coloration, strong odors, or stings. He also foutid that birds and 
 marmosets would often devour "conspicuously colored " larvae with- 
 out any hesitation, and that some "protectively colored" or incon- 
 spicuous larvae were refused. There can be no doubt that many 
 
204 BUI.LETIN: MUSEUM OF COMI'AKATIVE ZOOLOGY. 
 
 insectivorous animals pay but little attention to the colors of their 
 prey ; for example, it is well known to anglers that trout and salmon 
 will snap at the most gaudily colored "tiics," which may or niaj' not 
 have any counterpai't in nature. 
 
 The whole question of warning coloration will liave to be made 
 the subject of an extensive research upon both old and young 
 insectivorous animals before we can safely arrive at any certain con- 
 clusions respecting it. 
 
 II. Methods Puusued in Stuuyincj the Colou-Pattkrns ok 
 THK Heliconidak. 
 
 No comparative study of the color-patterns displayed by the 
 Ileliconidae has ever been made. In fact, very few such studies 
 have been carried out upon any Lepidoptera. The only works I 
 know of are those of Eimer ('89) and llaase ('9'2) upon the colora- 
 tion of the Papilios, and of Dixey ('90) upon the wing-markings of 
 certain genera of the Nyraphalidae and Pieridae. The family of the 
 Ileliconidae with its numerous species and comparatively simple 
 coloration affords an excellent opportunity for such a research. 
 
 In making this study of the Ileliconidae I was permitted through 
 the kindness of Mr. Samuel llenshaw to make free use of the collec- 
 tion in the Museum of Comparative Zoology at Harvard. I also 
 found the colored figures in the works of the following authors of 
 great service : Hewitson ('50-'76), 0. und R. Feldcr ('()4-'(J7), IlUb- 
 ner ('0G-"i5), Humboldt et Bonpland ('38), Cramer (1779-'82), Stau- 
 dinger ('84-'88), Godman and Salvin ('79-'8()), and M(f;nd-tri6s ('(i;!) ; 
 likewise the following shorter papers published in various serials : 
 Bates ('63, '65), Butler ('65, '69, '69-'74, '77), Druce ('76), (iodman 
 and Salvin ('80) , Ilewitson ('54), Snellenen van Leeuwen ('87), Srnka 
 ('84, '85), Staudinger ('8'2), and Weymer ('75, '84). I was thus 
 enabled to examine the color-patterns of 400 (89%) of the species 
 of the Danaoid group, and of 129 (86%) of the Acraeoid group, 
 either from the insects themselves or from figures given by the 
 authors named above. The remaining species were either inaccessi- 
 ble to mc, or were so vaguely described as to be unavailable. A 
 list of the species known to me is given in Table 28. 
 
 (1) The Two Types of Coloration in Ike Danaoid Ileliconidae. 
 It is very remarkable that the color-patterns of all of the Ileliconidae 
 
MAYER: COLOR AND COI.OR-PA'ITEUNS. 205 
 
 may be grouped into two very closely related types. To the one of 
 these I have given the name '■'■ Mclinaca type" for it is characteristic 
 of most of the species of the genus Melinaea. It is well represented 
 by Figs. 40, 48, 49, 51, and 55-57 (Plate 4). The insects which 
 belong to this type possess wings colored with rufous, black, and 
 yelloto. 
 
 The other type I designate as the " Ithoniia type," for it is very 
 characteristic of most of the species of the genus Ithomia. Figs. 47 
 and 52 (l^late 4) afford examples of it. This type differs from the 
 Melinaea in that the rufous and yellow areas upon the wings have 
 become traiispare/U. 
 
 There are, also, many species, found in numerous genera, which 
 fall between these two types of coloration, for the yellow and rufous 
 spots upon their wings have become translucent, so that one may 
 speak of them as "translucent yellow" and "translucent rufous." 
 These spots arc, -so to speak, in process of becoming transparent, but 
 a few yellow or rufous scales still remain dusted over the otherwise 
 flear spaces. Most of the Dircennas are good examples of this type ■ 
 (Fig. 54, Plate 4) . 
 
 Of the 400 species of the J)anaoid Ileliconidae, about 125 belong 
 to the " Melinaea type." It is well represented V)y most of the 
 species of the genera Lycorea, Athyrtis, Ceratinia, Mcchanitis, and 
 Melinaea. About 30 Ithomias and half a dozen Napeogenes also 
 belong to it. About KiO species belong to the " Ithomia type," and 
 of this number fully I'iO belong to the genus Ithomia. The others 
 are found in the genera Ceratinia, Napeogenes, Ituna, and Thyridia, 
 and many of them resemble the Ithomias so closely that they are 
 said to mimic them. About 100 species, some of which are found in 
 almost all of the genera, are intermediate in their color-patterns 
 between the Melinaea and the Ithomia types. The 15 remaining 
 species are represented by Melinaea gazoria (Fig. 53, Plate 4), 
 Ceratinia eupompe, and a few Ithomias, such as Ithomia hemixaiitho. 
 In these f(n'ms almost all color has disappeared, so that the whole 
 wing has become of a uniform dull translucent yellow, bordered on 
 the outer edges by a grayish black. 
 
 (2) Detailed Description of the Melinaea Type of Coloration. 
 Figs. 40, 48, 49, 51, and 55-57 (Plate 4) afford examples of this type 
 of coloration. In these insects we find the proximal half of the 
 central cell of the fore wing occupied by a rufous-colored area, which 
 I call the "inner rufous." It is marked I in all of the figures. 
 
206 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 JJeyond the " inner rufous " we find a black spot, mai'ked II in the 
 figures. It usually occupies the middle region of the cell of the fore 
 wing, and I have designated it as the " inner black." Beyond the 
 "inner black," and occupying most of the outer portion of the cell 
 of the fore wing, is a light-colored area, marked III in the figures. 
 This area is rufous in color in Fig. 49, but it is usually yellow, as in 
 Figs. 46, 48, 51, 54-57, and I have called it the " inner yellow." 
 Beyond the " inner yellow," and occupying the extreme outer 
 portion of the cell, lies the "middle black" (IV). In many species 
 it is fused, as in Figs. 4C-48, 56, 57, with the large black area, the 
 " outer black " (VII) , which occupies the greater portion of the outer 
 half of the fore wing. Just outside of the cell beyond the "-middle 
 black" one finds a well-developed yellow area (V), the " middle 
 yellow," and there is sometimes still another yellow patch beyond 
 this, which is marked VI and called the " outer yellow." Finally, 
 one often finds a row of white or yellow spots, the " marginal s])ots" 
 (IX), lying very near the outer margin of the fore wing (see Figs. 
 47-49, 51, 54, 56). These spots are very well developed in the 
 genera Ceratinia, Napeogenes, Ithomia, and Melir.aea. One more 
 very characteristic marking of the fore wing remains to be noticed ; 
 that is the longitudinal black stripe (VIII). It is also worthy of note 
 that the front costal edge of the fore wing is almost always tinged 
 with black. 
 
 The pattern of the hind wing is quite simple. The ground color 
 is usually rufous and a "middle black" band (XI) runs across the 
 middle of the wing. The outer edge is bordered by the "outer 
 black" (XIII). Above the "middle black" band lies the " inner 
 rufous " (X) of the hind wing, and below the "middle black " band 
 one finds the "outer rufous" (XII) of the hind wing. One often 
 finds a row of white or yellow dots within the outer black border of 
 the hind wing, and these I designate as the "marginal spots" of the 
 hind wing. 
 
 The Ithomia type of coloration, it will be remembered, may be 
 derived from the Melinaea, by simply imagining the rufous and 
 yellow areas to have become transparent. Also the outer black 
 usually suffers a reduction so as to become only a rather narrow 
 boi'der along the outer margin of the fore wing. Thyridia psidii 
 (Fig. 47) is a good example of this type. It will be seen that the 
 black areas remain about the same as in the Melinaea type, but that 
 
MAYER: COLOK AND COLOR-PATTERNS. 207 
 
 the rufous and yellow have become transparent. The middle and 
 outer yellow areas have also fused into a large transparent patch. 
 
 Itliomia .sao {V'lg. 52, Plate 4) is another good example of the 
 Ithoniia tyjje. In tliis particular s])ecies tlie " iinier black" of the 
 fore wing is absent, and the "middle black band" of the liind wing 
 has disai)peared. When we come to consider the other Ithomias, 
 we shall find that in this genus it has probably fused with the 
 marginal black of the hind wing. 
 
 I have made a record of the color-variations that affect the 
 various characteristic areas just considered, and have recorded them 
 for every one of tlie species of the Danaoid and Acraeoid llcliconidae 
 known to me. As these records are too extensive for convenient 
 inspection, I have condensed the results, and they will be found 
 in Tables 1-27 inclusive. Thus, Table 1 gives the variations in 
 color of the "inner rufous" area of the fore wing for each genus 
 of the Danaoid Heliconidae ; Table 2 records the variations of the 
 " inner black " ; Table 3 the " inner yellow " area, etc. In Table 1 
 we find, for example, opposite the genus Ituna, the number 2 in the 
 column labeled "transparent." This indicates that in two species of 
 Ituna the " inner rufous" area is transparent. 
 
 In order to facilitate the study of the color-patterns Dr. Daven- 
 port suggested that I make use of the ingenious projection method 
 invented by Keeler ('98). This method consists in " squaring the 
 wing" in the manner shown in Figs. 4 and 5 (Plate 1). In Fig. 4 
 the large rectangle (A, B. C, D) just at the right of the figure of the 
 hind wing represents a kind of Mcrcator's projection of the wing 
 itself. The nervures 1°, P, 2, 3, etc., are represented by the 
 vertical lines 1", P, 2, 3, etc., on the rectangle A, B, C, D. In cells 
 I», P, and P, (bounded by nervures 1", 1\ and 2,) one finds a 
 sinuous line winding across the middle of the cell. This line 
 appears in the same relative position upon the rectangle A, B, C, D. 
 The same is true of the eye-spot found in the cell bounded by 
 nervures 2 and 3, and of all the other markings of the wings. 
 The central cell of the wing itself is shown projected in the dotted 
 rectangle E, F, G, IT. 
 
 In the case of the /ore wing (Fig. .'i), the central cell of the wing 
 is dotted, and is shown projected upon the similarly dotted area 
 within the rectangle I, J, K, L. In other respects the method of 
 projection is the same as in the case of the hind wing. 
 
 In this manner the colors displayed by various species of Danaoid 
 
208 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 and Acraeoid Ileliconidae have been represented in color in Plates 
 5-8. Each largo rectangle upon the left hand side of the Plate 
 represents a hind wing, the small middle rectangles show the colors 
 of the cell of the hind wing, and the right hand rectangles give the 
 fore wings, all being projected in the manner illustrated in P'igs. 4 
 and 5, Plate 1. The chief advantage in Keeler's projection method 
 lies in the fact, that similar areas in the projection of the wings lie 
 vertically under or over one another, and thus by merely glancing 
 up or down the plates one may observe the color-variations which 
 occur in homologous cells of all the species represented. 
 
 III. General Discus.sion of the Color-Pattkrns and of 
 Mimicry in the Genera IIeliconius and Eueide.s. 
 
 Among the species of the genera Heliconius and Eueides we find 
 remarkably little variation in venation, but great diversity in color- 
 pattern of the wing.s, and in this respect they are very dilferent from 
 the Danaoid Ileliconidae, where, it will be remembered, we find fully 
 twenty different types of venation and only two types of color- 
 pattern . 
 
 (1) The Four Color Types in the Genus Heliconius. Schatz 
 and liober ('85-'9i2) divide the species of the genus Heliconius into 
 four groups based on color differences, as follows: — (1) the 
 "Antiochus group" (Plate 4, Fig. 50); (2) the "lirato group" 
 (Fig. 60); (3) the "Melpomene group" (Fig. 59); and (4) the 
 " Sylvanus group," a good example of which is Heliconius eucrate 
 (Fig. 58, Plate 4). 
 
 It will become apparent through an inspection of Figs. 50, CO, 59, 
 and 58, which represent respectively, Heliconius antiochus, H. erato, 
 H. melpomene, and H. eucrate, that the first three are quite closely 
 related in color-pattern, while the fourth (H. eucrate) approaches 
 very closely to the plan of coloration of the Melinaea type of the 
 Danaoid Heliconidae. In fact this resemblance is so close that it 
 may be safely said that the members of the " Sylvanus group," to 
 which H. eucrate belongs, mimic the Danaoid Heliconidae. 
 
 The "Antiochus group" is represented by Heliconius .anti- 
 ochus (Plate 4, Fig. 50, and Plato 5, Fig &1). II. sara, H. galanthus, 
 and H. charitonius (Plate 5, Figs. 01, 03, 04) are also members of 
 this group ; other e-vamplcs arc H. apseudes, II. cydno, II. chiones, 
 H. hahnesi, H. sappho, II. leuce, H. eleusinus, and H. clysonvnius. 
 
MAYER: COLOR AND COLOR-l'ATTICRNS. 209 
 
 These species are cliaracterizcd by their bhie iridcseonce, and tlie 
 narrow yellow or white bands upon the primaries ; the hind wings 
 are pointed at the outer apex, and the venation approaches the type 
 found in Euoides aliphera. H. ricini (Plate 5, Fig. G(j) is a good 
 example of a form intermediate in coloration between group 1 and 
 the " Erato group " (2). 
 
 The type of group 2 is Ileliconius erato (Plate 4, Pig. GO, and 
 Plate 5, Pigs. 67 and C8) . This group is closely allied to group 1 in 
 its characteristics. A good connecting link between groups 1 and.'i, 
 the "Melpomene group," is 11. phyllis (Fig. (55). 
 
 The third, or " Melpomene group," is represented by II. mel- 
 pomene, II. callicopis, II. cybele, II. thelxiope, and II. ve.sta (Plate 6, 
 Figs. 70-74, and Plate 4, Fig. 59). II. vulcanus, II. venus, II. 
 chestertonii, II. burneyi, and IT. pachinus are also examples of this 
 group. 
 
 (2^ Mimicvy betwuen. the Genus Jleliconius and the Damioid 
 Group. To Schatz's group 4, the "Sylvanus group," belong all those 
 species of Ileliconius which have departed widely from the colora- 
 tion pattern of the other three groups, and have come to resemble 
 various species of the genera Melinaea, Mechanitis, and Tithorea of 
 tlie Danaoid Ileliconidae. II. eucoma, H. eucrate, Il.dryalus, and H. 
 sylvana (Plate 8, Figs. 88, 89, 91, and 95) are good examples of 
 group 4. By glancing at the diagrams on Plate 8 it will be seen 
 that n. dryalus resembles Melinaea paraiya very closely ; in fact, the 
 likeness is so clo.sc tluit it is almost certain that no eye could distin- 
 guish between the two insects when they arc upon the wing. Another 
 startling resemblance is that between II. eucrate and Melinaea thera 
 (Plate 8, Figs. 91 and 92) ; moreover, there is but little difference 
 between the color-patterns of II. eucrate, Eueides dianasa, and 
 Mechanitis polymnia (Figs. 91, 98, and 94). IT. sylvana and 
 Melinaea egina (Figs. 95 and 90) are also said to mimic each other. 
 The resemblance certainly appears vei-y close at a casual glance, yet 
 when the colors are plotted, as in Figs. 95 and 90, the differences be- 
 come quite apparent. IT. claudia (Plate 5, Fig. 09) is a good con- 
 necting link between tlie Sylvanus group and the i\[elpomenc group. 
 In both the Melpomene and Sylvannsgroupsthe venation has departed 
 from the Eueides ali])hera type, and the contour of the hind wings is 
 much more rounded atul elliptical than is the case in tlie Antiochus 
 and Erato groups. (Compare Figs. 50 and 00 with Figs. 58 and 59, 
 I'latc 4.) There are rather less than twenty species which certainly 
 
210 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 belong to the Sylvanus group ; among them may be mentioned, 
 in addition to those already spoken of, Ileliconius nuniata, which 
 resembles Melinaea mneme and Tithorea harmonia ; II. zuleioa, which 
 resembles a Mechanitis and is a good copy of Melinaea hezia ; and 
 II. metalilis, which is said to mimic Melinaea lilis ; there are also 
 striking resemblances between 
 
 H. aurora and Melinaea luoiter ; H. messene and Melinaea inesenina ; 
 
 H. eucrate and Mechanitis lysimnia ; H. hecalesia and Tithorea liecalesina ; 
 
 H. hecuba and Tithorea bonplandii ; II. ethra and Mechanitis nesaea ; 
 
 H. formosiLs and Tithorea penthias ; H. pardalinus and Melinaea pardalis; 
 
 H. telcliina and Melinaea imitata ; II. isinenius aiui Melinaea mcssatis. 
 
 Most remarkable of all perhaps is the close resemblance between 
 Ileliconius aristiona, Mechanitis methonc, and Ithomia fallax of 
 Staudinger. In fact, Staudinger states in his " Exotische 8chmet- 
 terlinge " that he hesitated for some time to describe Ithomia fallax 
 on account of its close resemblance to Ilewitson's Mechanitis methone. 
 Good lists of the Heliconidae which are said to mimic one another 
 are given by Wallace ('89, p. 250, 251), and by Ilaase ('98", p. 
 146, 147). 
 
 (8) The Three Color- Types in the Germs Eueides. In the 
 genns Eueides we meet with three color-type.s represented by 
 E. aliphera, E. thales, and E. cleobaea. These insects are dis- 
 tinctly smaller than the species of the genus Ileliconius, and the 
 yellow spots upon their primaries are more ocherous in color than 
 in Ileliconius. E. aliphera (Plate 6, Fig. 77) represents the most 
 highly specialized color-type. Eueides mereaui (Fig. 70), however, 
 is a good connecting link between the color-patterns of E. aliphera 
 and E. thales (Fig. 75), and E. thales is almost identical in color- 
 pattern with Ileliconius vesta (Fig. 74). 
 
 The other type of Eueides is represented by E. cleobaea, E. 
 dianasa, E. Isabella, etc. (Plate 6, Fig. 78, and Plate 8, Fig. 93). 
 These resemble the Sylvanus group of Ileliconius or various Melinaeas 
 and Mechanitis. 
 
 (4) Detailed Discussion of Plates 5-8. Plate 5 is intended 
 to illustrate the types of coloration found in the Antiochus and 
 Erato groups of the genus Ileliconius. In IT. sara (Fig. 61) the 
 wings are suffused with a dark blue iridescence, and some narrow 
 yellow bands of color are found upon the primaries. In H. antiochus 
 (Fig. 62) we find similar bands of color upon the primaries, but 
 they are changed to white. H. antiochus may have descended 
 
MAYER: COLOR AND COLOU-rATTERNS. 211 
 
 from an all)inic s|)ort of H. sara. In IT. galanthns (Fig. 03) 
 the white areas liave greatly increased in size, and the iridescent 
 blue has become mucli hghter. In II. charitoniiis (Fig. 04) we 
 find the wings crossed Vjy yellow spots and bands, but in some speci- 
 mens tliis yellow color exhibits a decidedly reddish tinge. The figure 
 of II. charitoiiius in Staudinger's "Exotische Schmetterlinge " illus- 
 trates this ])eculiarit}' ; indeed, spots which are commoidy yellow are 
 often found red, and vice versa. In II. phyllis (Fig. (55) we find 
 along the upper part of the diagram of the hind wing a yellow mark- 
 ing, and a similarly shaped I'ed mark is found in its near ally, II. 
 thelxiope (Fig. 73, Plate 6). The same is also true of H. ricini 
 (Fig. (id, Plate 5). 
 
 II. erato (Figs. 07 and (iS, Plate 5, and Fig. GO, Plate 4) is very 
 remarkable, for there are no less than four distinct color-types 
 exhibited by different individuals of this species; one of them (Fig. 
 07) shows the basal half of the hind wing marked by six red tongues 
 of color edged with iridescent blue, and there is a dai-k rufous 
 suffusion upon some parts of the fore wing. In other si)ecimens 
 (Fig. 08) the red tongues of color which characterized the hind wing 
 of Fig. 07 are almost absent, and oidy the blue iridescence is left ; 
 also there is no rufous to be seen upon the fore wing. In another 
 type the blue iridescence of the hind wing has become green, and in 
 still other specimens the yellow stripes upon the fore wing have 
 become white. 
 
 As one looks over the diagrams upon Plates 5-8, it becomes evi- 
 dent that yellow frequently changes to white, for we often find one 
 or two species of a genus which exhibit white spots identical in shaj)e 
 and position with spots which are yellow in mo.stof the others. Good 
 examples of this are II. antiochus (Plate 5, Fig. 02), Melinaea 
 ])arallelis and Ceratinia leucania (Plate 7, Figs. 8'2and83) ; likewise 
 the white spot near the outer apex of the fore wing in II. eucrate 
 (Plate 8, Fig. 91), which is yellow in many individuals. Yellow 
 areas are also fre(iuently changed to rufous or red ; thus the yellow 
 basal half of the hind wing of II. eucrate (Plate 8, Fig. 91) is often 
 found of a rufous tinge in individual specimens of the species, and 
 among the specimens of this species in the jNIuseum of Comparative 
 Zoology one can trace a gradation of this ai-ea from bright yellow 
 to rufous. IT. Claudia (Plate 5, Fig. 09) is introduced in order to 
 exhibit some of the differences between the " Sylvanus " group, to 
 Avliich it belongs, and the " Antiochus " and " Erato " groups. 
 
21-2 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 Plate 6 is intended to exhibit the characteri.stic cokir-pattcrns 
 found in the Melpomene group and in the genus Eueides. Kig. 70 
 represents 11. inelpoinene, and Fig. 71 its near ally, H. callycopis, in 
 which the red area of the fore wing has become broken up, and some 
 red spots have made their ap])earance near the base of the liind wing. 
 In the ne.xt variety of H. melponiene, II. cybele (Fig. 72), it is 
 remarkable that the pattern of the fore wing has come to resemble 
 the Sylvanus type, and is identical in general plan of coloration with 
 the fore wings of the Melinaeas or Mechanitis (see Figs. 84 oi' 85, 
 Plate 7, or Figs. 92 or 94, Plate 8). In its close ally, II. thelxiope 
 (Fig. 73), a still nearer approach to the Melinaea type has come 
 about by the development of a black band across the middle of the 
 hind wing, and one has only to imagine a general fusion of the seven 
 club-shaped red stripes of the hind wing in Fig. 73, Plate 6, in order 
 to produce exactly the Melinaea type as exhibited, for example, by 
 Eucides cleobaea (Fig. 78). In this connection it is worthy of note 
 that Bates ('(52) showed that II. thelxiope was derived from II. 
 melpomene, there being between the two matiy intermediate forms. 
 
 II. vesta (Fig. 74) is evidently a close relative of II. thelxiope, 
 and what is still more worthy of note is, that it is almost identical in 
 the general effect of its color-pattern with Kueides thales (Fig. 75)! 
 The yellow spots upon the fore wing of E. thales are, however, duller 
 in hue than are those of H. vesta, and tlie insects are somewhat 
 different m size, IT. vesta spreaduig 78 mm., while E. thales spreads 
 only 6() mm. It will be noticed that the chief difference between 
 the color-patterns of these two species lies in the fact, that, while the 
 black stripes of the hind wings in II. vesta lie along the nervures, in 
 Eueides thales they occupy the middle of the cells themselves. The 
 general resemblance of the two color-i)atterns may of course be 
 merely accidental. An easy explanation, however, is afforded by 
 the theory of mimicry, for the two species look very much alike 
 until one subjects their color-patterns to close analysis, when 
 remarkable differences appear. E. thales (Fig. 75) may have been 
 derived from some such form as E. mereaui (Fig. 7G), for one has 
 merely to imagine a greater devel()j)ment of the black and a general 
 deepening of the rufous upon the hind wing of E. mereaui to make 
 it resemble E. thales quite closely. Finally, in E. aliphera (Fig. 77) 
 the l)lack serrated border of the hind wing is still more reduced, and 
 the black stripe which crosses the cell of the fore wing in E. mereaui 
 is not present. 
 
MAVKK : COLOK AND COLOH-I'ATTEKNS. 213 
 
 Pi.ATic 7 is iiitcniled to illustrate the peculiarities of color-])attern 
 found among the Danaoid Ileliconidae. Thyridia psidii (Fig. 79) 
 is an example of the transparent type of color-pattern found among 
 the Danaoid Ileliconidae, and especially prevalent among the 
 Ithomias. It will he seen hy comparing Fig. 79 with the other 
 figures upon J'lates 7 and 8, that the chief difference lies in the fact, 
 that in this type both the rufous and yellow areas have become 
 transparent. "J'he black area of the fore wing has also suffered a 
 reduction, especially along the outer margin of the wing. Inci- 
 dentally it should l)c mentioned, that in this particular species the 
 middle black band of the hind wing has become tilted up at a sharp 
 angle, instead of crossing the wing horizontally. A life-si/x' figure 
 of the wings of Thyridia psidii is given on Plate 4, Fig. 47. 
 
 In Napeogenes cyrianassa (Fig. 80) and Ceratinia vallonia (Fig. 
 81) j)ortions of the usually yellow and rufous areas have become 
 transparent. 
 
 The spots upon the fore wing of the Melinaeas are usually yellow, 
 but in Melinaea parallelis (Fig. 82) they are white. It would seem 
 that this form may have descended from some albinic sport. 
 Ceratinia leucania (Fig. 8.'i) resembles Melinaea parallelis so closely 
 in general plan of coloration, that it is very difficult to distinguish 
 between them, even when the two insects are seen side by side. 
 Ceratinia leucania, however, is somewhat smaller than Melinaea 
 parallelis. Hoth occupy the same region in Central America, and 
 the Ki)ecimeiis from which the diagrams were drawn came from 
 I'anania. 
 
 Figs. 84-87 are drawn from various specimens of Mechanitis 
 isthmia, all from Panama. They are intended to give .some idea of 
 the range of individual variation which is met with in this extremely 
 variable form. The contraction of the middle black band of the 
 hind wing in this form has already been noticed in the general 
 discussion of tlie laws of color-pattern (see page 184). In Fig 87 it 
 will be seen that the inner yellow stripe whicih usually crosses the cell 
 of the fore wing has become very narrow and changed to a rufoua 
 color. However, upon the under surface of the wing it still remains 
 as a yellow stripe. Indeed, in most color-changes the upper side of 
 the wing seems to take the initiative, the under surface being more 
 conservative. This is not true, however, in the Ithomias, where 
 the black areas of the under side of the wings often are found to be 
 rufous in color, while they still remain of the normal black upon the 
 
214 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 upper surface. The colors of the uiulor surface are, however, 
 usuallj' identical with those of the upper, though tliey are always 
 duller in hue. This may be due to the fact, that the colors of the 
 upper surface are more frequently seen than those of the lower, for 
 these insects often float lazily along with their wings horizontally 
 extended. The operation of Nat\u-al Selection would then be more 
 severe with the upper surfaces than with the lower. 
 
 Plate 8 gives an analysis of the color-patterns of some of the 
 Heliconinae and those Melinaeas, etc., which they resemble. H. 
 eucoma (Fig. 88) is a good e.xample of the Sylvanus type, and with 
 its rufous, 3'ellow, and black wings it is certainly a wonderfully close 
 copy of the color-pattern found so commonly among the species of 
 the genus Melinaea of the Danaoid Ileliconidae. 
 
 ITeliconius dryalus and Melinaea paraiya (Figs. 89, 90) resemble 
 each other so closely in size, shape, and coloration, that it must be 
 impossible to distinguish between them when the butterflies are in 
 flight; yet an analysis of their color-])atterns shows that there are 
 considerable differences between them. The shape of the yellow 
 bands upon the fore wings is quite different ; the inner black spot 
 within the cell is double in Melinaea paraiya, and there is also a row 
 of white spots along the margin of the fore wing. 
 
 A much closer resemblance is found between II. eucrate and 
 Melinaea thera (Figs. 91 and 92), where the Heliconius is almost a 
 true co))y of the Melinaea. 
 
 The color-patterns of Eueides dianasa (Fig. 93) and Mechanitis 
 polymnia (Fig. 94) are also very nearly the same. Both are 
 common species in Brazil. 
 
 Heliconius sylvana is said by Bates and by Wallace to mimic 
 Melinaea egina. It will be seen by reference to Figs. 9ri and 90 
 that their color-patterns are quite different in detail, yet the insects 
 look very much alike when placed side by side, and may easily be 
 mistaken for each other when upon the wing. Melinaea egina is 
 much more common than Heliconius sylvana. 
 
 IV. Gejjbb.vl Discussion ok the Colob-Patteuns a.vo op 
 
 MiMICBY AMONH THE DaN'AOID IIeUCONIDAE. 
 
 (1) The Origin of the Tloo Types of Coloration. The character 
 of the variation in the Danaoid Ileliconidae is very different from 
 that of the genera Heliconius and Eueides, for while there is great 
 
MAYEH: COLOR AND COLOll-rATTERNS. 215 
 
 diversity of color-pattern and very little variation in venation among 
 tlie species of the Acraeoid group, exactly the opposite condition is 
 met with in the Danaoid group, where we find at least twenty 
 dii^'erent types of venation and only two types of col()r-))attern. 
 One of these types of coloration is well exemplified by most of 
 the Mehnaeas (Fig. 48, I'late 4), and I liave therefore called it the 
 " 7l/e/iM«ea" type. The other type is exemplified by most of the 
 Ithomias (Figs. 47 and 52) and has been designated in this paper as 
 the "Ti/towta" type. In the Melinaeas, it will be remembered, we 
 find the rufous and black wings crossed by bands of yellow ; while 
 in the Ithomias, on the other hand, the rufous and yellow areas have 
 become transparent, often leaving the wing as clear as glas.s, and the 
 black, which is so characteristic of the outer lialf of the wing in the 
 Melinaea type, has shrunk away until it has come to lie along the 
 outer margin of tlie wing only. 
 
 By a study of all the genera of Danaoid Heliconidae we gain light 
 upon the question of the origin of the "Melinaea" and "Ithomia" 
 types of coloration. As we have seen (page 198), the Danaoid 
 Heliconidae are an offshoot from the great family Danaidae. Indeed, 
 two of the genera, Lycorea and Ituna, are so closely related to the 
 Danaidae that Schatz and IJober ('85-'S)'2) propose to include them 
 within that family. There can be but little doubt that Lycorea and 
 Ituna are remnants of the ancestral forms which long ago shot off from 
 the Danaidae to form the Danaoid Heliconidae ; and it is interesting to 
 note, that in these two patriarchal genera we find the two distinct 
 tyjies of color-pattern which are exhibited by tlie Danaoid Helico- 
 nidae, for all of the five known species of Lycorea are good examples 
 of the Melinaea ty])e (see Lycorea ceres, Fig. 46, Plate 4), while the 
 four known species of Ituna all exhibit the transparent, or Ithomia, 
 type of coloration. In fact, in their color-patterns the species of 
 Ituna remind one of gigantic Ithomias. The species of Lycorea, 
 however, are colored very much after the pattern of the Danaidae, 
 and indeed they have departed but little from the ty()e of the 
 member.s of the great family whence they sprang. On this account 
 I believe that the Melinaea type of coloration, which is so charac- 
 tei'istic of the species of Lycorea, is pliylogenetically older than 
 the Ithomia type. 
 
 In order to account for the origin of the Ithomia type, we may 
 assume that, shortly after the primeval forms of the Danaoid Heli- 
 conidae began to segregate out from the Danaidae, the species were 
 
21() BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 few and probably rare. Under these circumstances any given insect 
 would gain but little advantage by resembling merely the general 
 type of the coloration of its fellows. F'or the relative advantage 
 gained by such imitation, according to Fritz Miiller's law, increases 
 inversely as the scjuare of the fraction who.se numerator is the actual 
 number of the imitating form and whose denominator is tlie actual 
 number of the imitated. Therefore when the insects were still rare 
 there would be few to imitate and consequently but little advantage 
 would be gained by the imitation. Imaghie, for example, that a 
 single insect happens to imitate the color-pattern of a group of 100, 
 and that the advantage gained tliereliy is re[)resented by the number 
 1 ; it is evident from Fritz Miiller's law that, if it happened to 
 imitate the coloration of a group of 1,000, its relative advantage 
 would be 100 instead of 1. We see, then, that mimicry within the 
 grou]) of the Danaoid Helicouidae became an important factor only 
 after the group was well established and the insects became common. 
 During the early history of the race, then, there would be but little 
 tendency towards conservatism of color-patterns, and when the 
 "Ithomia" and "Melinaea" types of coloration made their aj)})ear- 
 ance, they both survived and now serve as tlie patterns for mimicry ; 
 and this accounts very well for the remarkable fact, that there 
 are no other types of coloration than these two to be found within 
 the whole group with its 450 species! 
 
 (2) Mimicry among the Danaoid Ileliconidae. The genus 
 Ithomia with its 230 sj)ecies is the dominant genus of the Danaoid 
 group, and in nearly all of the other genera individual species are 
 found which have departed widely from their generic type of 
 coloration and liave assumed the clear wings of the Ithomias. A 
 good idea of how far these interesting individuals may depart from 
 the coloration of their type may be gained by comparing Fig. 5;!, 
 Plate 4, which represents Melinaea gazoria, with Fig. 48, which 
 represents a typical Melinaea (M. paraiya). It is evident that 
 Melinaea gazoria is startlingly like an Ithomia both in size and 
 coloration, although it retains the venation and generic charac- 
 teristics of a Melinaea. 
 
 In Mechanitis, which is the most independent genus of the 
 Melinaea type of coloration, all of the species are fair examples of 
 the Melinaea type, except Mechanitis ortygia Druce, from Peru. 
 Druce ('70) in liis description of this curious little species states in 
 astonishment that it possesses the venation of a Mechanitis, but the 
 size and coloration of an Ithomia ! 
 
MAYER: COLOR AND COLOR-PATTERNS. 217 
 
 It is quite remarkable that altliougli tlie genera Meliiiaea and 
 Meclianitis serve as models of miiniery for the Acraeoid Ileliconidae, 
 they should themselves mimic Ithoiiiia. 
 
 The genus Ithomia is, however, the most independent of all the 
 genera of the Danaoid group, and I know of remarkably few good 
 instances in which an Ithotnia has apjjarently departed from the 
 coloration of its type to assume the guise of the Melinaeas. One good 
 example of such a change, liowcver, is afforded by Ithomia fallax of 
 Southern Peru, which resembles either Mechanitis methone or Ileli- 
 conius aristiona of Colombia (see page 210). There is apparently 
 a difficulty in ascribing this resemblance to mimicry, for the imitator 
 and imitated do not occupy the same geographical regions. 
 
 In direct contrast with the iiidei)endence of the Ithomias stands 
 the case of the genus Napeogenes ; for Godman and Salvin ('79-'86) 
 say of Napeogenes, that nearly every species mimics some Ithomia 
 which occupies the same district; and thus almo.st the very existence 
 of the genus would seem to depend upon its mimicry of Ithomia. 
 
 It is not the purpose of this paper to discuss, in detail, the numerous 
 interesting cases of mimicry which are believed to exist between 
 mendiers of the Danaoid Ileliconidae. An excellent discussion of 
 sudi cases, and of the relationships of the various genera, has been 
 given by IIaa.se ("93", p. 116-127). 
 
 V. Quantitative Detkumination ok the Variations of the 
 
 <;habactekistk; Wing-Markings in the Acraeoid and 
 
 Danaoid IIemconidae. 
 
 (1) Variatians of '■'■Inner linf'ous" Areas of the Fore and 
 Jlind Winf/s. Table 1 gives the color-variations which are exliibited 
 by the "inner rufous" area of the fore wings in the Danaoid Ileli- 
 conidae. This area is marked I in all of the figui-es upon Plate 4. 
 We learn from an inspection of Table 1 that this area is rufous in 
 color in 124 species of the Danaoid Ileliconidae, transparent in 152, 
 black in 24, and that in the remainder it is more or less translucent, 
 and of either a yellowish or rufous tinge. 
 
 Table 10 shows the variations which come over the "inner 
 rufous" area of the hind wings of the Danaoid Ileliconidae. This 
 area is marked X in tlie figures upon Plate 4. It is apparent at 
 a glance that the variations Avliich affect the inner rufous areas of 
 
218 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 both fore and hind wings arc vmy similar. In order to exhibit this 
 fact graphically, the color-variations have been laid off npon the 
 diagram, Fig. 97, Plate 9. The base line is marked at equal inter- 
 vals with the words " rufous," " translucent rufous," " translucent, 
 slightly rufous," " transparent," etc., and the ordinates sliow the 
 number of species which exhibit the various colors, rufous, trans- 
 lucent rufous, etc. For example, at the point " translucent rufous " 
 we find that the ordinate is 28 ; this indicates that in 23 species 
 the area is translucent rufous in color. The points thus foinid 
 upon the ordinates are successively joined by straight lines form- 
 ing a zig-zag figure. The full line represents the fore wing, and 
 the dotted line the hind wing, and it becomes clearly evident from 
 the closeness of these two zig-zag lines that the color of the inner 
 rufous area of the fore wing (area I, Plate 4) is almost always 
 sure to be identical with that of the inner i-ufous area of the hind 
 wing (area X, Plate 4). We see, therefore, that whatever color- 
 variation affects the inner rufous area of the fore wing, this area 
 in the hind wing is almost always affected in the same manner. 
 
 Fig. 99, Plate 9, is derived from Tables 15 and 24, which show 
 the color-variations in the fore and hind wings of the genera Ilelico- 
 iiius and Eueides. It is seen that here also the colors of these two 
 ai-eas in both the fore and hind wings are almost alwaj's identical. 
 We here meet with one of those interesitng physiological laws 
 M'hich are indej)endent of Natural Selection, and the meaning of 
 which remains a mystery, for surelj' we can see no reason on the 
 ground of adaptation why similar areas upon both fore and hind 
 wing should bear similar colors. 
 
 (2) The " Inner lilack" Spot. Table 2 shows the presence or 
 absence of the "inner black" spot in the Danaoid llelicouidac. 
 This spot is marked II in the figures upon Plate 4. When pres- 
 ent, it is always black in color and is usually found occupj'^ing the 
 middle region of the cell of the fore wing. The table shows that 
 it is about an even chance whether it be present oi- not, for it is 
 absent in 210 species and present in 190. In the genus Ithomia, 
 however, it is present in only one third of the species. What is 
 most worthy of note concerning it, is the fact that it almost always 
 appears, when present, as a single spot. Indeed, it appears as a 
 double spot in only 7 species, and 5 of these belong to the genus 
 Melinaea. A good example of its appearance as a double spot is 
 found in Melinaea paraiya (Fig. 48, Plate 4). It will be remem- 
 
MAYER: COLOR AND COLOR-PATTERNS. 219 
 
 bercd that there are 450 species in the Daiiaoid group ; 25 of 
 these belong to the genus Melinaea; yet among these 25 we find 
 5 exhil)iting this marking as a double spot. Assuming that the 
 doubling of this spot has arisen in each species as a sport, and that 
 such a sport is as likely to appear in one species as in any other of 
 the Danaoid group, then the chances against five such sports 
 appearing among the 25 Melinaeas is '^'Ij^^^xU^.^Jr^' "'" ^^°^^^ 
 2,830,000 to 1. Indeed, it is prol)able that all five of the species 
 of Melinaea which exhibit the doubling of this spot are descend- 
 ants of a sinyle ancestor in which it appeared for the first time 
 double, for the mathematical chance that one such ancestor should 
 appear among the Melinaeas, rather than in any other genus, is 
 evidently 1 in ^, or one chance in eighteen. The chance against 
 two such unrelated ancestors is, however, ■""'X'*^" or about 336 to 
 
 28x'i4 ' 
 
 1, and the chance against three is *^^^^~^, or 6,560 to 1, etc. 
 
 I}y reference to Table 16 we find that in the genera Ileliconius 
 and Eueides tlie inner black area is black or iridescent blue in all 
 of the species of Heliconius, but absent in 5 of the 18 species of 
 Eueides known to me. These 5 include Eueides alipliera and its 
 allies. Now there are 150 known species of the Acraeoid Helico- 
 nidae, and 24 of these belong to the genus Eueides ; so it is evident 
 that the mathematical chance against the supposition that five sports 
 arose independently in the genus Eueides, in which the inner black 
 was absent, is given by '"^^^j^^g^^^ or 13,900 to 1. It is there- 
 fore probable that the five Eueides lacking the inner black are 
 the descendants of a single ancestor. 
 
 (3) Variations of the '■'Inner Yellow'" and '■'■ 3{iddle Yellow" 
 Areas. Tables 3 and 5, and diagram Fig. 98, Plate 9, show the 
 color- variations of the "inner yellow" and "middle yellow" areas 
 in the fore wings of the Danaoid Ileliconidae. These areas are 
 marked III aiid y, respectively, in the figures upon Plate 4. The 
 " inner yellow " area, it will be remembered, occupies tlie outer por- 
 tion of the cell of the fore wing ; while the " middle yellow " is found 
 in the region just beyond tlie outer limits of the cell. The two areas 
 are often fused together as in Figs. 47, 48, 50, 51, and 55, Plate 4. 
 The inner yellow area is usually smaller than the middle yellow, 
 and a comparison of Tables 3 and 5 will show that it is much more 
 frequently obliterated by the encroachment of the rufous or black 
 
220 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 areas which suitouiuI it; for example, while the middle yellow is 
 rufous in color in only 14 species, the inner yellow is rufous in 56; 
 also the inner yellow area, being usually smaller and less conspicuous 
 than the middle yellow, is less important in cases of mimicry, and 
 the diagram Fig. 98, Plate 9, shows that it is much more variable 
 in color than the middle j'cUow. The full zig-zag line in this figure 
 represents the color-variations of the inner yellow, while the dotted 
 zig-zag line gives the color-variations of the middle yellow. As 
 there ai'e nine color-variations displayed by each of these two areas, 
 and as there are 400 species of the Danaoid Ileliconidae recorded 
 by me, it becomes evident that, if there tcere no color preferences 
 displayed by these areas, there would probably be about ^^, or 44.4, 
 species which would display it as rufous, 44.4 translucent, 44.4 yellow, 
 etc. The heavy, straight, dotted line (Fig. 98, Plate 9) represents 
 this ideal condition, which would be a])proximately realized were 
 one color as likely to occur as another in the respective areas. Now 
 it is evident from an inspection of the figure, that the f)dl zig-zag 
 line, which represents the color-variations of the " inner yellow," 
 approaches the sti-aight line condition more nearly than does the 
 dotted zig-zag line, which represents the middle yellow. i The 
 inner yellow is therefore more liable to color-variations than the 
 middle j-ellow ; and this is what we should expect on account of 
 its comparatively small size and its consequent inconspicuousness 
 as a characteristic marking in cases of mimicry. 
 
 A comparison of Figs. 97 and 98, Plate 9, is interesting, for it 
 shows that the color-variations of the inner rufous are quite similar 
 to those of the inner yellow and middle yellow. This serves to 
 illustrate the close phy.siological relationship which exists between 
 rufous and yellow. The two pigments are prol)ably closely related 
 chemically, for every ordinarily rufous area is sometimes found to be 
 yellow, and vice versa. Yellow areas also often change to white. 
 Rufous, yellow, and white are evidently closely related color-vari- 
 ations.''' 
 
 'This is not true for one color, white. 
 
 - It may be well to mention hero that the black areas upon the wings are subject to 
 very little color-variation. In some cases, however, especially upon the under surfaces 
 of the wings in Ithoinia, the black has changed to a rufous or russet color. For example, 
 Table 4 shows that the middle black area (IV in the figures upon Plate 4) is rufous in 
 only Vi species out of tlio 400 which are recordeil, and all of these 12 are Ithomias, Also 
 Tables 7 and 13 show that the outer black of the fore wing, and the outer black of the 
 hind wing are russet in 22 and 11 species, respectively. Evidently, black is a far more 
 conservative color than rufous, yellow, or white. Probably black is also quite different 
 from the other pigments chemically. 
 
MAYEK: COLOR ANT) COLOR-PATTERNS. 221 
 
 Tables 17 and 19 show the color-variations affecting the "inner 
 3'ellow " and " middle yellow " areas of the fore wing in Ileliconius 
 and Eueides. There is but little difference between the two tables, 
 except that in 15 species of Ileliconius the inner yellow is suffused 
 witli black or blue, while the middle yellow is never suffused hy the 
 outer black which surrounds it. P'ig. 100, Plate 10, exhibits 
 graphically the color-variation of these two areas. The "inner 
 yellow" is represented as a full line, and the "middle yellow" as 
 a dotted zig-zag. It is evident that here also the inner yellow is 
 more variable in color than the middle yellow, for not only does the 
 inner yellow area display two more colors, but its chart is a flatter 
 zig-zag. 
 
 (4) Variations of the " Middle Black " Mark of the Fore Wing. 
 Table 4 shows the color-variation of the middle black mark (area 
 IV in figures upon Plate 4) . This marking lies along the extreme 
 outer border of the central cell of the fore wing. It is small in area, 
 but is rendered very conspicuous from the fact that it is situated be- 
 tween the inner yellow and middle j'ellow markings. In spite of 
 its small size, however, it is a remarkably j)ermanent marking, for 
 Table 4 shows that it is absent in only 20 out of 400 Danaoid lleli- 
 conidae. In these 20 it has been obliterated by the fusion of the 
 inner and middle yellow areas. It is worthy of note that in 12 
 Ithomias it has become rufous in color. This change to rufous is 
 the only color-change wliicli the black areas of the wings ever 
 display. 
 
 Table 18 shows the variations of the middle black area for 
 Ileliconius and Eueides. 
 
 (5) Variations of the '■'■ Outer Yellow'''' Area of the Fore Winr/. 
 Table 6 shows the variations which affect the outer yellow ai-ea 
 of the fore Avings in the Danaoid Ileliconidae. This area is marked 
 VI in the figures upon Plate 4 ; it lies beyond the region of the 
 middle yellow, but is usually more or less fused* with it. Table G is 
 only approximately correct, owing to the difticultj- in many cases of 
 deciding whether the middle and outer yellow be really fused or not. 
 It will be seen that in the genus Ithomia the middle and outer 
 yellows are wliollj' fused in about 200 species. This is one of the 
 marked characteristics of this very independent genus. 
 
 Table 20 sliows the color-variations of the outer yellow area in 
 Ileliconius and Eueides. This marking is present in 81 and absent 
 in 48 of the Acraeoid group. It is much more widely separated 
 from the middle yellow than is the case in the Danaoid group. 
 
222 BULLETIN: MUSEUM OF COMl'ARATIVE ZOOLOGY. 
 
 (6) The relatiue Perrnaiieiicxj of the Black Areas upon the Fore 
 and Hind Wings. A study of the relative permanency of the 
 various characteristic black markings upon the wings is of interest, 
 for, if the generally accepted idea concerning the jirevalence of 
 mimicry within the group of the Danaoid Ileliconidae be true, we 
 should expect the most conspicuous markings to be the most perma- 
 nent, for they are evidently of the most importance for mimicry. 
 This is, however, not the case for the black markings. A good 
 example of this fact is afforded by a comparison of the relative 
 permanency of the black streak which extends along the extreme 
 costal edge of the fore wing with the inner black spot (II in figures 
 on Plate 4). The inner black spot is certainly a more conspicuous 
 marking than this narrow black streak along the costal edge ; yet it 
 is much more variable, for Table '1 shows that it is present in 210 
 and absent in 190 of the 400 Danaoid Ileliconidae. In other words, 
 it is about as likely to be present as absent. The black streak upon 
 the costal edge, on the other hand, is much more permanent, for it is 
 absent in only 52 species otit of the 400. 
 
 Another good example of the inaccurracy of the supposition that 
 large and conspicuously colored areas are always less variable than 
 small ones, is derived from a comparison of the relative variability 
 of the large outer black of the fore wing with the small outer 
 black of the hind wing. Although the outer black area of the fore 
 wing is usually much larger and more conspicuous than the outer 
 black margin of the hind wing, it is more variable in color, for it is 
 nifous in 22 species, while tlie outer black of the hind wing is 
 rufo)i8 in only 11, out of the 400. 
 
 In general, however, large colored areas are more permanent tlian 
 •small ones, as was found in the case of the inner and middle yellow 
 areas (see page 220). Indeed, a good in.stance of this greater vari- 
 ability of small color areas is afforded by the longitudinal black 
 stripe marked VIII in the figures of IMate 4, for this is more 
 variable than the larger outer black area of the fore wing. 
 
 (7) The " Middle Mack Htripe " of the Hind Wi/iff. In the 
 genus Ithomia the middle black stripe (XI, Plate 4) has migrated 
 downward, so that in many species it has become fused witli Ihe 
 outer black margin, as in Ithomia sao (Fig. 52, Plate 4). In other 
 cases there is still to be seen a narrow line of rufous color between 
 the middle black band and the outer black margin of the hind 
 wing. Such is the case in Ithomia nise (F'ig. 54, Plate 4). In 
 
MAYER: COLOR AND COLOR-l'A'ITERNS. 223 
 
 many other cases the outer black and middle black aie completely 
 fused, so far as the upper surface of the wings is concerned ; but, 
 if one examines the under surface of the hind wings, it will be 
 found that a narrow rufous streak still persists between the middle 
 black band and the outer black margin of the hind wing. 
 
 (8) Variations of the Marginal Spots of the Fore Win</. The 
 marginal spots are found very near the outer margin of the fore 
 wing; they are usually either yellow or white, but in some few 
 cases they are rufous. It ajipears from Table 9 that they are 
 present in 146 and absent in '254 species of the 400 Danaoid Ileli- 
 conidae known to me. Fig. 101, Plate 10, shows graphically 
 the manner in which these spots occur in those species which 
 possess them. It is evident from this curve that the number of 
 these spots is not determined merely by chance, for they show a 
 marked tendency to ap])ear either as 2 or 3, or as 6 or 7 sjiots. 
 It is due to this fact, that there are two maximum points upon 
 the diagram Fig. 101, Plate 10. In those species which exhibit the 
 "2- or 3-spot" condition, the spots are found near the front apex 
 of the fore wing. In the " 6- or 7-spot " condition they lie all along 
 the outer margin of the fore wing, one spot in each cell. In the 
 genera Ithomia, Napeogenes, and especially in Ceratinia these mar- 
 ginal spots have become large and conspicuous ornaments. (See 
 Fig. 49, Plate 4.) 
 
 Table 22 shows the manner of ai)pearanee of these spots in the 
 genera Heliconius and Eueides. They are found in only 26 species 
 of the 129 known to me ; and this number is far too small to war- 
 rant general conclusions concerning the order of their appearance. 
 
 (9) The Marginal Spots of the Hind Win//. Table 14 illus- 
 trates the manner in which the marginal sjjots of the hind wings 
 make their appearance. They are absent in 279 and present in 
 121 of the 400 species of the Danaoid group. Thus they occm- 
 rather less frequently than the marginal spots of the fore wing. In 
 the 121 species in which these spots are found they show a decided 
 tendency to appear either as 4 or as 5 spots. Fig. 102, Plate 10, 
 is a graphic representation of the disti-ibution of these spots, derived 
 from Table 14. It appears that the outline of the figure approaches 
 a probability curve, and is approximjitely symmetrical about the 
 mean ordinate (A, B), sitnated at 4.54. 
 
224 lU'LLETlN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 VI. CoMPARISOX OF THE CoLOK-V.\KI.\T10XS OF THE PaI'ILIOS OF 
 
 South Amkuica witii thosh of thk Heliconidae. 
 
 In order to cmpliasize the peculiarities of the coloration of the 
 Heliconidae, I will conclude by instituting a conii)arison between 
 their variations and those of the South American Papilios. There 
 are about '200 species of Papilio in South America, and these display 
 in all 36 distinct colors. The colors have been determined bj' 
 reference to the plates in liidgwa^f's " Nomenclature of color for 
 naturalists." A list of the colors which are displayed by these Pa])i- 
 lios has already been given upon page 191. 
 
 By exercising a very fine discrimination in distinguishing color we 
 may count 15 distinct colors whicii are displayed by the 450 mem- 
 bers of the Danaoid Heliconidae, as follows : black, brown, translu- 
 cent black, sulphur-yellow, canary-yellow, citro!i-yellow, jjrimrose- 
 yellow, yellow-rufous, reddish rufous, rufous, white, translucent yel- 
 low, translucent rufous, transparent areas upon the wings, transpar- 
 ent areas which display iridescence. We see, then, that while the 
 200 species of Papilio display SO different colors, the 450 Danaoid 
 Heliconidae exhibit only 15. In other Avords, the numbers of the 
 species and of the colors are almost in inverse ratio in the two 
 groups; for while the Papilios are only ^ as n\iinerous as the 
 Danaoid Heliconidae, they display almost 'j"- times a's many colors ; 
 and this is all the more remarkable when we remember that the gen- 
 eral class of coloration in the Papilios and Danaoid Heliconidae is 
 apparently the same. That is to saj', in both gr()U])s we find all of 
 the sijecies displaying decidedly conspicuous colors, the coloration 
 of the upper surfaces of the wings being in both rather more bril- 
 liant than that of the lower surfaces, but without essential differences 
 in color-jjattern. Nor is there an attempt in either case at protective 
 resemblances, such as the imitation of the coloration of bark, leaves, 
 etc. The color-patterns of the I'apilios are, moreover, extremely 
 complex, and upon comparing the different species, there are seen 
 to be frequent fusions and obliterations of the characteristic mark- 
 ings, so that Haase ('93), who has made an extensive study of their 
 color-patterns, is forced to divide them into many small groups 
 of a few species each. The variation in the form of the wings is 
 also very great among the Papilios, for while P. protesilaus ])os- 
 sesses upon its hind wings, long tail-like appendages, the hind 
 wings of P. hahneli are rounded off and without marked appendages. 
 
MAYEK: COLOK AND COLOR-l'ATTERNS. 225 
 
 There is, apparently, but one important respect in whicli the 
 Danaoid Heliconidae are more variable than the Papilios, and that 
 is size. For example, Lycorea ceres, which is probably the largest 
 of the Danaoid group, has 2.2 times the spread of wing of Ithomia 
 nise, which is one of the smallest (see Plate 4, Figs. 4G and 54). 
 The largest Papilio, P. androgea, on the other hand, spreads only 
 ■J. 10 times as much as the smallest, P. triopas. 
 
 There is another minor respect in which the color-patterns of the 
 Papilios are different from those of the Heliconidae. In the Heli- 
 conidae the fore wing slightly overlaps the hind wing, and that por- 
 tion of the hind wing whicli is hidden from view is always dull in 
 color (see Plates 5-8) . In the Papilios, however, the fore wing 
 does not overlap the hind wing to such an extent as in tlie Helico- 
 nidae, .and it is worthy of note that the costal edges of the hind 
 wings in the Papilios are as brilliantly colored as are any other por- 
 tions of the wings. 
 
 It is difficult to account for the remarkable conservatism in respect 
 to coloi'-variations among the Heliconidae, unless we resort to the 
 explanation afforded by the theory of mimicry ; for, while there is 
 such remarkable simplicity and uniformity of color-pattern through- 
 out the whole group of the Heliconidae, individual variations 
 are very common. In the collection at the Museum of Compara- 
 tive Zo6logj% for example, one finds a regularly graded series of 
 specimens of Heliconius eucrate ; at one end of this series the 
 " inner rufous " area of the hind wing is bright yellow, and at 
 the other end it is rufous ; intermediate specimens are found in 
 which this area is yellow, but dusted over with rufous scales. Also 
 the " middle black band " of the hind wings in Melinaea parallelis 
 is very variable, some specimens showing it broken in the middle 
 (Plate 7, Fig. S2), and others having it as an entire band. I have 
 also seen one specimen of H. burneyi in which the commonly 
 yellow spots upon the under surface of the wings were changed to 
 white. Another good instance of individual variability is afforded 
 by H. phyllis (Plate 5, Fig. 65), for in this species the series of 
 small red spots sometimes found just below the yellow band upon 
 the hind wing is very variable, and more often absent than j)rescnt. 
 Still other instances of individual variability are seen in the yellow 
 stripes upon the wings of II. charitonius (Plate 5, Fig. 64), which 
 are often found tinged with rufous. Also the remarkable diversity 
 in Mechanitis polymnia, and M. isthmia (Plate 7, Figs. 84-87) are 
 
226 BULLETIN: MUSEUM OF COMl'AKATIVE ZOOLOGY. 
 
 Other examples which sliow that there is no lack of individual vari- 
 ability among the Heliconidae. Yet the Daiiaoid species as a whole 
 vary but little from the two great tyj)es of coloration represented 
 by Ithomia and Melinaea, and in this respect tliey are very differ- 
 ent from the Papilios, where we fiiul a great many color-types and 
 great diversity in shape of wings. Surely there must be some cause 
 for the remarkable fact that the Danaoid Heliconidae with their 
 453 species should display but two types of color-pattern. I can 
 think of but one explanation, which is afforded by Fritz Miiller's 
 theory of mimicry. 
 
 In conclusion it gives me pleasure to thank those friends whose 
 generous aid and kindness have done so much to render the prose- 
 cution of this research a pleasure to me. I wish to express my 
 gratitude to Dr. Charles B. Davenport, who is the real instigator 
 and promoter of this research ; to Mr. Samuel Ilenshavv, to wliom 
 I am indebted for numerous kindnesses, and Avho placed at my dis- 
 posal the extensive entomological collections and library of the 
 Museum of Comparative Zoology at Harvard ; to Prof. Edward L. 
 Mark for his kindness in revising the manuscript of this paper, and 
 for the numerous valuable suggestions which he has made ; to Dr. 
 Samuel H. Scudder, to whom I am gratefid for much kind advice 
 and for the use of rare works in his library ; to Prof. Ogden N. 
 Rood for his valuable suggestions in i-egard to the spectroscoj)ic 
 apparatus; to Dr. Alpheus Hyatt for his valued and kind advice, 
 and to my father, Prof. Alfred M. Mayer, for the use of Maxwell's 
 discs and the direct-vision spectroscope. 
 
 PART C. 
 
 GENERAL SUMMARY OF RESULTS BELIEVED TO 
 BE NEW TO SCIENCE. 
 
 (1) The great majority of the colors of Lepidoptera contain a 
 surprisingly large percentage of black (p. 172). 
 
 (2) The colors displayed by the scales are not simple, but com- 
 pounded of several different colors (p. 173). 
 
 (3) The jjigments of the scales of Lepidoptera are derived by 
 various chemical processes from the blood, or haemolympli, of the 
 
MAYKR: COLOR AND COLOR-l'ATTERNS. 227 
 
 pupa. Tlie puj)al blood of tlic (Satuniidae is a proteid substance 
 containing egg albumen, globulin, fibrin, xanthopliyll, orthophos- 
 phoric acid, iron, potassium, and sodium (p. 17G). 
 
 (4) In Callosamia prometliea and Danais plexippus the pupal 
 wings are at first perfectly transparent, then white, then impure 
 yellow, excepting upon those portions wliich are destined to remain 
 white in the mature wing. The mature colors then begin to appear 
 near the central areas of tlie wings and bettoeen the nervures. Last 
 of all, the nervures themselves become tinged with the mature 
 colors. The central portions of the wings acquire their mature 
 colors before the outer and costal edges, or the root of the wing 
 adjacent to the body (p. 178, Plate 3). 
 
 (5) The white stage in the development of color in the pupal 
 wings represents the condition in which the scales are perfectly 
 formed but lack the pigment which is destined to be introduced 
 later (p. 178). (See, also, Mayer, '9G, p. 230.) 
 
 (0) Dull ocher-yellows and drabs are, phylogenetically speaking, 
 the oldest pigmental colors in the Lepidoptera. The more brilliant 
 colors, such as bright yellows, reds, and pigmental greens, are 
 derived by complex chemical processes and are, phylogenetically 
 speaking, of recent appearance (p. 178). (See, also, Mayer, '96, p. 
 23'2.) 
 
 (7) While the number of species of PapiUo in South America 
 is 9 times as great as in North America, the number of colors which 
 they display is only twice as great. Hence the greater number of 
 colors displayed by the tropical forms may be due simply to the 
 far greater number of the species, and not to any direct influence 
 of the climate (p. 191). 
 
 (8) The following laws control the color-patterns of butterfiies 
 and moths : (a) 7\ny spot found upon the wing of a butterfly or 
 moth tends to be bilaterally symmetrical, both as regards form and 
 color ; and tlie axis of symmetry is a line passing through the center 
 of the interspace in which the spot is found, parallel to the longi- 
 tudinal nervures (p. 18.3). (b) Spots tend to appear not in one 
 interspace only, but in homologous places in a row of adjacent 
 interspaces (p. 183). (c) Bands of color are often made by the 
 fusion of a row of adjacent spots, and, conversely, chains of spots 
 are often formed by the breaking up of bands (p. 183). (d) AVhen 
 in process of disappearance, bands of color usually shrink away at 
 one end (p. 184). (e) The ends of a series of spots are more 
 
228 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 variable than the middle. Thi,s i.s only a special case of Bateson's 
 ('94) law (p. 185). (£) The position of spots situated near the 
 outer edges of the wing is largely controlled by the wing-folds or 
 creases (p. 185). 
 
 (9) The scales in Lepidoptera do not strengthen the wings or 
 aid the insects in flight. The vast majoritj' of the scales are merely 
 color-bearing organs, which have been developed under the influence 
 of Natural Selection. The ])hylogeiietic appearance and develop- 
 ment of scales upon the originally scaleless ancestors of the Lepi- 
 doptera did not alter the efficienci/ of their wings as orr/cmtt of fli<jht. 
 It is probable, therefore, that this etticiency was an optimum before 
 the scales appeared (p. 197). 
 
 (10) A systematic study of the Danaoid Helioonidae demon- 
 strates that their color-patterns can be jjlaced in two types. Type 1, 
 the more complex, is closely related to the coloration of the Danaidae 
 from which the Danaoid Ileliconidae sprang, and is therefore, 
 phylogenetically speaking, the older type of coloration. This type 
 is characteristic of the genera Lycorea, Melinaea, and Mechanitis, 
 and I have called it the " Melinaea" type. It is characterized by 
 the fact that the wings are rufous and black in color, and crossed by 
 a definite system of yellow bands. Type '2, the " Ithomia " type, is 
 characteristic of the genera Ithomia, Ituna, and Thyridia. The 
 "Ithomia" type has been derived from the "Melinaea" by the 
 originally rufous and yellow areas upon the wings having become 
 transparent (p. 204). 
 
 (11) The phylogenetic origin of the "Melinaea" and "Ithomia" 
 types of coloration can be accounted for upon the supposition, that 
 when the species of the Danaoid Ileliconidae began to segregate 
 out from the Danaidae they were for a time rare (p. 215). • 
 
 (12) A record of the characteristic markings upon the wings of 
 the Danaoid and Acraeoid Heliconidae shows that, physiologically 
 speaking, the colors red, rufous, j'ellow, and white are closely 
 related, and that black is quite disthict from these, being the least 
 variable color of all (p. 220). 
 
 (13) In both the Danaoid and Acraeoid Heliconidae, whatever 
 color-variation affects that part of the fore wing which is adjacent 
 to the body of the insect, almost always the same color-variation 
 affects the homologous area of the hind wing in a similar mannei- 
 (p. 218, and Fig. 99). 
 
 (14) The smaller yellow spots upon the wings of the Ileliconi- 
 
MAYEK: COLOR AND COLOU-PATTEKNS. 229 
 
 (lae are move liable to color-variations than are the larger ones. 
 This is what we should exi)ect, if the theory of mimicry be true ; 
 for large spots are more conspicuous, and therefore their ])reservation 
 is more important (p. '220). This rule, however, docs not hold for 
 the black markings of the wing (p. 222). 
 
 (15) The mathematical chance against five similar and inde- 
 pendent color-sports arising in the genus Melinaea is as 2,830,000 
 to 1. Hence, the five Melinaeas which display the "inner black" 
 as a double spot are probably descended from a single ancestor (p. 
 219). 
 
 (16) The marginal spots of the fore wing in the Danaoid Heli- 
 conidae show a marked tendency to appear either as 2 or 3, or else 
 as 6 or 7 spots (p. 228, Fig. 101). The marginal spots of the hind 
 wing show a marked tendency to apj^ear either as 4 or 5 spots 
 (p. 223, and Fig. 102). 
 
 (17) The 200 species of PapiUo in South America display 36 
 distinct colors, while the 450 species of Danaoid Ileliconidae exhibit 
 only 15. Ilenee the numbers of the species and of the colors are 
 almost in inverse ratio in the two groups. This may be explained 
 by the fact, that the Danaoid Heliconidae mimic one another, while 
 the Papilios do not (p. 224). 
 
 (18) The colors are dull upon those portions of the hind wing 
 which are hidden from view by the ovcrlaj)ping fore wing (p. 225). 
 
 (19) There is no lack of individual variability among the species 
 of the Danaoid Ileliconidae ; yet the species as a whole vary but 
 little from the two great types of color-pattern represented by 
 Melinaea and Ithomia. In order to account for this remarkable 
 fact I am forced to resort to Fi-itz Mtiller's theory of mimiciy (p. 
 225). 
 
230 
 
 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 1. 
 
 Showing the Variations in Color of the " Inner Rufous " (Area 
 I in Figures on Plate 4) of the fore icing in the Danaoid Heli- 
 conidae. 
 
 (Jf.xkua. 
 
 2 
 1 
 
 o 
 II 
 
 is 
 
 s s 
 
 lis 
 
 1 
 
 1 
 
 H 
 
 2 
 1 
 5 
 
 1 
 4 
 
 7 
 
 12 
 120 
 
 ft 
 
 lis' 
 go 
 
 > 
 
 
 is 
 
 Lycorea 
 Ituna . . 
 Athesis 
 Thyridia 
 Athyrtis . 
 Olyras . . 
 Eutresis 
 Aprotojjos 
 Dircenua . 
 Callitlioiiiia 
 Bpithomia . 
 Ceratinia . 
 Sais . . . 
 Scada . . 
 Mechaiiitis 
 Napeogenes 
 Ithomia 
 Aeria . . 
 Melinaea . 
 Tithorea . . 
 
 
 
 5 
 
 2 
 
 1 
 
 1 
 1 
 1 
 28 
 5 
 
 18 
 
 7 
 
 29 
 
 22 
 4 
 
 2 
 
 3 
 
 1 
 1 
 
 3 
 13 
 
 2 
 
 1 
 1 
 
 2 
 
 2 
 15 
 
 2 
 
 1 
 
 1 
 26 
 
 1 
 
 7 
 
 3 
 5 
 4 
 2 
 
 1 
 1 
 
 1 
 
 1 
 
 1 
 1 
 
 2 
 2 
 
 4 
 3 
 3 
 
 6 
 
 1 
 
 Total 
 
 124 
 
 23 
 
 23 
 
 152 
 
 30 
 
 22 
 
 24 
 
 1 
 
 Excluding Ithomia 
 
 95 
 
 10 
 
 8 
 
 32 
 
 4 
 
 17 
 
 1 
 
 21 
 
 
 
 Note: The costal edge of the fore wing is usually black; it is rufous or brown, how- 
 ever, in 47 Ithomias and dull yellow in one; it is rufous In two species of Sais, in one 
 species of Ceratinia, and in one species of Athesis. Hence it is black in 34k species and 
 llgiit <:oIored in 52. 
 
MAYER; COLOR AND COLOR-l'ATTKRNS. 
 
 231 
 
 TABLE 2. 
 
 Showing the Variation (presence or absence) of the "Inner 
 Black" (Area II) of the /ore wing in the Danaoid Heliconidae. 
 
 
 «3 
 
 *i 
 
 
 Genera. 
 
 S 
 
 1 
 
 RemarkH. 
 
 
 
 £• 
 
 
 
 I 
 
 < 
 
 
 Lycorea 
 
 5 
 
 
 
 Ituna . . 
 
 
 
 
 3 
 
 
 
 Athesis 
 
 
 
 
 2 
 
 1 
 
 
 Thyridia 
 
 
 
 
 6 
 
 
 
 Athyrtis 
 
 
 
 
 2 
 
 
 
 Olyras . . 
 
 
 
 
 4 
 
 
 
 Eutresis 
 
 
 
 
 2 
 
 
 
 Aprotopos 
 
 
 
 
 2 
 
 
 
 Dircenna . 
 
 
 
 
 8 
 
 4 
 
 
 Callithomia 
 
 
 
 
 3 
 
 
 
 Epithomia 
 
 
 
 
 2 
 
 
 
 Ceratinla . 
 
 
 
 
 29 
 
 12 
 
 
 Sais . . . 
 
 
 
 
 4 
 
 1 
 
 
 Scada . . 
 
 
 
 
 
 7 
 
 
 Mechanitis 
 
 
 
 
 23 
 
 1 
 
 
 Napeogenes 
 
 
 
 
 13 
 
 17 
 
 Appears as 2 spots in 1 species. 
 
 Ithoinia 
 
 
 
 
 72 
 
 140 
 
 Appears as 2 spots in 1 species. 
 
 Aeria . . 
 
 
 
 
 
 4 
 
 
 Melinaea . 
 
 
 
 
 21 
 
 3 
 
 Appears as 2 spots in 5 species. 
 
 Tithorea 
 
 10 
 
 
 
 Total 
 
 210 
 
 190 
 
 
 Excluding Ith 
 
 om 
 
 ia 
 
 
 138 
 
 60 
 
 
232 
 
 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 3. 
 
 Showing the Variations in Color of the "Inner Yellow" (Area 
 III) of the /"ore wing in the Danaoid Heliconidae. 
 
 ^i 
 
 £■38 
 m2 
 
 Lycorea . . . . 
 
 Itiiiia 
 
 Athesis . . . . 
 Thyridia . . . . 
 Athyrtis . . . . 
 
 Olyras 
 
 Eutresis . . . . 
 Aprotopos . . . 
 Oircenna . . . . 
 Callithoinia . . . 
 Epitlioinia . . . 
 Ceratiuia . . . . 
 
 Sais 
 
 Scada 
 
 Meclianitis . . . 
 Napeogenes . . . 
 Ithoniia . . . . 
 
 Aeria 
 
 Melinaea . . . . 
 Tithorea . . . . 
 
 Total 
 
 Excluding Ithomia 
 
 66 
 
 158 
 
 38 
 
 31 
 
 04 
 
MAYER: COLOR AN]) COLOR-l'ATTERNS. 
 
 233 
 
 TABLE 4. 
 
 Sliowiiig till' presence or ab.scncc of the " Mitldle Black " Mark 
 (Area IV) of tlH'/c>?'t' whu/ in the Danaoid Ileliconidae. 
 
 (tCNEUA. 
 
 Present. 
 
 Absent. 
 
 Present, but olianKed to 
 some color other tlian black. 
 
 Lycorea 
 
 Ituna 
 
 Atliesis 
 
 Thyridia 
 
 Athyrtis 
 
 Olyras 
 
 Kutresis 
 
 Aprotopos .... 
 
 Dircenna 
 
 Callithoniia .... 
 Kpithomia .... 
 
 Ceratinia 
 
 Sais 
 
 Scada 
 
 Meclianitis .... 
 Napeogenes ... 
 
 Ithoniia 
 
 Aeria 
 
 Melinaea 
 
 'Pithorea 
 
 6 
 
 3 
 
 3 
 
 S 
 
 2 
 
 2 
 
 2 
 
 2 
 
 11 
 
 3 
 
 2 
 
 34 
 
 6 
 
 7 
 
 24 
 
 25 
 
 194 
 
 4 
 
 23 
 
 10 
 
 1 
 
 7 
 
 r, 
 
 
 1 
 
 12 rufous or brown. 
 
 'I'otal 
 
 366 
 
 20 
 
 12 
 
 E.xchiding Ithomia ". 
 
 172 
 
 14 
 
 
234 
 
 RULLETIN: MUSEUM OF COMl'AHATIVE ZOOLOGY. 
 
 tai5i:k fi. 
 
 Showing the Variation in Color of the " Middle Yellow " Hand 
 (Area V) of the /'ore winff in the Danaoid Heliconidae. 
 
 Lycorea . . . . 
 
 Ituua 
 
 Athesis . . . . 
 Thyridia . . . . 
 Atliyrtis . . . . 
 
 Olyras 
 
 Eutresis . . . . 
 Aprotopos . . . 
 Dircenna . . . . 
 Callithomia . . . 
 Epithoinia . . . 
 Ceratiiiia . . . 
 
 Sais 
 
 Scada 
 
 Mechanitis . . . 
 NapeogencK . . . 
 Ithoniia . . . . 
 
 Aeria 
 
 Melinaea . . . . 
 Tithorea . . . . 
 
 Total 
 
 Excluding Itliomia 
 
 169 
 36 
 
MAYKR: COLOR AND COLOR-PATTERNS. 
 
 235 
 
 TAULK G. 
 
 Sliowliiir approximately tlic; Nimiber of Species in which the 
 " Outer Yellow " (Area VI) of the fore, loinf/ in the Danaoid Heli- 
 conidae appears as a separated Marking. It is usually fused with 
 the " Middle Yellow " Area. 
 
 
 Wholly fused 
 
 Partially fused 
 
 
 
 (iKiVUltA. 
 
 with miiiaic 
 
 with middle 
 
 .Separate. 
 
 .\l)Nent. 
 
 
 .VBllow. 
 
 yellow. 
 
 
 
 Lycorea 
 
 
 1 
 
 4 
 
 
 Ituna . . 
 
 
 
 
 
 2 
 
 1 
 
 
 
 Athesis 
 
 
 
 
 
 3 
 
 
 
 
 Thyridia . 
 
 
 
 
 
 5 
 
 
 
 
 AthyrtLs 
 
 
 
 
 
 
 
 2 
 
 
 Olyras 
 
 
 
 
 
 1 
 
 S 
 
 
 
 Kutresis . 
 
 
 
 
 
 
 2 
 
 
 
 Aprotopos 
 
 
 
 
 
 
 2 
 
 
 
 Dircenna 
 
 
 
 
 
 7.> 
 
 5 .? 
 
 
 
 Callithoiiiia 
 
 
 
 
 
 
 1 
 
 2 
 
 
 Epitlioiiiia 
 
 
 
 
 
 
 1 
 
 1 
 
 
 Ceratiiiia . 
 
 
 
 
 
 22 
 
 10 about 
 
 ;! about 
 
 
 Sais . , 
 
 
 
 
 
 
 
 2 
 
 8 
 
 Scada . . 
 
 
 
 
 
 
 
 4 
 
 8 
 
 Meclianitis 
 
 
 
 
 
 
 1 
 
 20 
 
 
 Napeogenes 
 
 
 
 
 
 
 
 
 24 about 
 
 
 Ithomia 
 
 
 
 
 
 200 about 
 
 
 
 
 Aeria . . 
 
 
 
 
 
 4 
 
 
 
 
 Melinaea . 
 
 
 
 
 
 1 
 
 
 17 
 
 (i 
 
 Tithorea . 
 
 
 
 
 
 
 
 10 
 
 
 Total . . . 
 
 
 
 
 
 about 250 
 
 
 
 about 30 
 
 '"abourSd" 
 
 perhaps 20 
 
236 BULLETIN: MUSEUM OF COMPAUATIVE ZOOLOGY. 
 
 TAKLE 7. 
 
 Showing the Degree of Dcvclo])iiient of the " Outer Black " (Area 
 VII) of the fore loing in the Danaoid llelicoiiidae. 
 
 
 Well devel- 
 
 Heduced 
 
 
 
 oped over a 
 
 to the outer 
 
 Present, but changed to 
 
 6ENEBA. 
 
 large area of 
 
 margin of the 
 
 another color. 
 
 
 - 
 
 the fore wing. 
 
 fore wing. 
 
 
 Lycorea .... 
 
 5 
 
 
 
 I tuna 
 
 
 
 
 
 
 
 ■> 
 
 
 A thesis 
 
 
 
 
 
 
 2 
 
 I 
 
 
 Thyridia 
 
 
 
 
 
 
 
 ■> 
 
 
 Athyrtis 
 
 
 
 
 
 
 2 
 
 
 
 Olyras . 
 
 
 
 
 
 
 3 
 
 1 
 
 
 Eutresis 
 
 
 
 
 
 
 2 
 
 
 
 Aprotopos 
 
 
 
 
 
 1 
 
 1 
 
 
 T)ircenna . 
 
 
 
 
 
 5 
 
 7 
 
 
 Callithomia 
 
 
 
 
 
 :? 
 
 
 
 Epitliomia 
 
 
 
 
 
 2 
 
 
 
 Ceratinia . 
 
 
 
 
 
 28 
 
 l:! 
 
 2 partly nifoiis. 
 
 Sais . . . 
 
 
 
 
 
 2 
 
 
 3 partly nifovis. 
 
 Scada . . 
 
 
 
 
 
 3 
 
 4 
 
 
 Mechanitis 
 
 
 
 
 
 22 
 
 ^ 
 
 
 Napeogenes 
 
 
 
 
 
 26 
 
 4 
 
 
 Ithomia 
 
 
 
 
 
 101 
 
 54 
 
 10 nifous or brown. 
 
 Aeria . . 
 
 
 
 
 
 4 
 
 
 
 Melinaea . 
 
 
 
 
 
 24 
 
 
 1 partly rufous. 
 
 Tithorea . 
 
 
 
 
 
 10 
 
 
 
 Total 
 
 305 
 
 05 
 
 22 partly rufous. 
 
MAYER: COLOK AND COLOR-PATTERNS. 
 
 237 
 
 TABLE 8. 
 
 Showing the presence or absence of the " Longitudinal Black 
 Stripe" (Area VIII) whicli runs parallel with the lower Edge of 
 the fore wing in the Danaoid Heliconidae. 
 
 Genera. 
 
 Present and 
 
 wen developed 
 
 as a stripe. 
 
 Much 
 reduced. 
 
 Absent. 
 
 Whole area 
 
 suffused with 
 
 black. 
 
 Lyeorea 
 
 Ituna 
 
 Athesis 
 
 Thyridia . . . . 
 
 Athyrtis 
 
 Olyras 
 
 Eutresis 
 
 Aprotopos .... 
 
 Uircenua 
 
 Callithomia . . . . 
 
 Epitliomia 
 
 Ceratiuia 
 
 Sais 
 
 Scada 
 
 Mechanitis .... 
 Napeogeues .... 
 
 Ithomia 
 
 Aeria 
 
 Melinaea 
 
 Tithorea 
 
 5 
 3 
 3 
 5 
 2 
 3 
 2 
 1 
 7 
 
 1 
 37 
 
 3 
 
 (> 
 
 17 
 
 20 
 
 200 
 
 4 
 14 
 
 6 
 
 3 
 
 1 
 2 
 
 1 
 
 
 
 
 6 
 
 1 
 
 1 
 1 
 2 
 
 4 
 
 6 
 
 1 
 
 1 
 2 
 2 
 
 2 
 
 5 
 4 
 2 
 
 5 
 
 Total 
 
 388 
 
 24 
 
 14 
 
 24 
 
238 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 9. 
 
 Showing the Manner of Occurrence of tlio Marginal Spots (Area 
 IX) of the fore wing in the Danaoid Heliconidae. 
 
 
 Wltll- 
 
 1 
 
 2 
 
 s 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 e 
 
 
 spets 
 
 spot. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 Lycorea . 
 Ituna . . 
 
 
 
 5 
 3 
 
 
 
 
 
 
 
 
 
 Athesis 
 
 
 
 :! 
 
 
 
 
 
 
 
 
 
 
 Tliyridia . 
 Athyrtis . 
 Olyras 
 Eutresis . 
 
 
 
 4 
 1 
 3 
 2 
 
 
 
 1 
 
 
 
 1 
 
 1 
 
 
 
 Aprotopos 
 Dircenna . 
 
 
 
 1 
 11 
 
 1 
 
 1 
 
 
 
 
 
 
 
 
 (Jallithomia 
 
 
 
 1 
 
 
 
 
 2 
 
 
 
 
 
 
 Epithomia 
 Oeratinia 
 
 
 
 2 
 1!) 
 
 1 
 
 3 
 
 1 
 
 
 
 4 
 
 12 
 
 1 
 
 
 Sais . . 
 
 
 
 6 
 
 
 
 
 
 
 
 
 
 
 Scada . . 
 
 
 
 4 
 
 
 
 1 
 
 
 1 
 
 
 1 
 
 
 
 Mechaiiitis 
 
 
 
 17 
 
 
 1 
 
 
 2 
 
 
 1 
 
 3 
 
 
 
 Napeogenes 
 Ithoinia 
 
 
 
 14 
 1.37 
 
 2 
 
 1 
 14 
 
 3 
 14 
 
 1 
 
 7 
 
 16 
 
 6 
 
 14 
 
 3 
 8 
 
 3 
 
 
 Aeria . . 
 
 
 
 4 
 
 
 
 
 
 
 
 
 
 
 Melinaea . 
 
 
 
 13 
 
 
 1 
 
 
 
 2 
 
 4 
 
 3 
 
 1 
 
 
 Tithorea . 
 
 
 
 5 
 
 
 1 
 
 
 1 
 
 
 
 2 
 
 
 1 
 
 Total .... 
 
 264 
 
 4 
 
 22 
 
 20 
 
 13 
 
 19 
 
 29 
 
 33 
 
 6 
 
 I 
 
MAYKR : COLOR AND COLOR-l'ATTERNS. 
 
 239 
 
 TABLE 10. 
 
 Showing the Color-Variations affecting tlie " Inner Rufous" (Area 
 X) of the hind wing in the Danaoid Ileliconidae. 
 
 (iHNKUA. 
 
 
 §2 
 
 3 
 
 eg 
 
 1 
 
 p.. 
 Si 
 
 1 
 
 1 
 
 S>;.| 
 
 Is. 
 
 1 
 
 it 
 S 
 
 7>ycorea 
 
 I tuna 
 
 Atliesis . 
 
 Thyridia 
 
 Athyrtis 
 
 Olyras . 
 
 Kutresis 
 
 Aprotopos 
 
 Diroenna 
 
 Callithomia 
 
 Kpithoinia 
 
 Ceratinia 
 
 Saia . . 
 
 Scada 
 
 Mecliauitis 
 
 Napeogenes 
 
 Ithoinia 
 
 Aeria 
 
 Meliiiaea 
 
 Tithorea 
 
 
 
 
 6 
 
 2 
 3 
 
 1 
 
 3 
 
 2 
 
 23 
 
 6 
 
 16 
 
 7 
 
 31 
 
 19 
 6 
 
 1 
 
 2 
 
 6 
 14 
 
 2 
 
 8 
 3 
 
 12 
 
 2 
 
 1 
 4 
 
 1 
 
 2 
 
 6 
 
 6 
 
 133 
 
 1 
 
 2 
 6 
 
 6 
 14 
 
 1 
 1 
 
 7 
 
 6 
 5 
 
 4 
 
 2 
 
 4 
 
 1 
 4 
 
 1 
 
 4 
 2 
 
 4 
 
 Total .... 
 
 123 
 
 23 
 
 26 
 
 166 
 
 29 
 
 23 
 
 12 
 
 10 
 
240 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 11. 
 
 Showing the Variations of the " Middle Black Stripe" (Area XI) 
 of the kind tohig in the Danaoid Heliconidae. 
 
 
 
 
 Fused with 
 
 
 Geneba. 
 
 Present. 
 
 Absent. 
 
 the marginal 
 black. 
 
 Changed color. 
 
 Lycorea 
 
 5 
 
 
 
 
 Ituna 
 
 2 
 
 1 
 
 
 
 Athesis 
 
 2 
 
 1 
 
 
 
 Thyridia 
 
 2 
 
 2 
 
 1 partially 
 
 
 Atliyrtis . . . 
 
 2 
 
 
 
 C 1 changed 
 
 Olyras 
 
 1 
 
 2 
 
 
 } to translu- 
 ( cent yellow .' 
 
 Eutresis 
 
 
 2 
 
 
 
 Aprotopos .... 
 
 2 
 
 
 
 
 Dirceiina 
 
 4 
 
 7 
 
 1 
 
 
 Oallithoinia .... 
 
 
 
 3 
 
 
 Epithoinia .... 
 
 
 2 
 
 
 
 Ceratinia 
 
 21 
 
 
 20 
 
 
 Sais 
 
 
 1 
 
 4 
 
 
 Scada 
 
 
 7 
 
 
 
 Mechaiiitis .... 
 
 22 
 
 2 
 
 
 
 Napeogenes .... 
 
 15 
 
 
 15 
 
 
 Ithomia 
 
 45 
 
 1 
 
 168 
 
 
 Aeria 
 
 
 4 
 
 
 
 Melinaea 
 
 15 
 
 9 
 
 
 
 Tithorea 
 
 8 
 
 2 
 
 
 
 Total 
 
 146 
 
 43 
 
 212 
 
 1? 
 
MAYEH: COLOK AND COLOK-l'ATTERNS. 
 
 241 
 
 TABLE 12. 
 
 Showing tlie Color- Variations of the "Outer Hufou.s" (Area XII) 
 of the kind iring in the Danaoid Ileliconidae. 
 
 Gk 
 
 SKUA. 
 
 j 
 
 || 
 
 in 
 
 a 
 t 
 
 Hi 
 
 c 
 s o 
 
 |-s. 
 
 1 ' 
 
 i 
 
 1 
 
 ^11 
 
 Lycorea 
 
 Ituna 
 
 Athesis 
 
 Thyridia 
 
 Athyrtis 
 
 Olyras . 
 
 Eutresis 
 
 Aprotopos 
 
 Dircenna 
 
 Callithoinis 
 
 Epithomia 
 
 Ceratinia 
 
 Sais . . 
 
 Scada 
 
 Mechanitis 
 
 Napeogene 
 
 Ithomia 
 
 Aeria . 
 
 Melinaea 
 
 Tithorea 
 
 
 
 
 
 
 5 
 
 2 
 3 
 
 1 
 1 
 2 
 1 
 2 
 29 
 6 
 
 21 
 15 
 44 
 
 17 
 6 
 
 1 
 
 1 
 
 4 
 1 
 
 2 
 
 3 
 
 1 
 
 2 
 
 1 
 4 
 
 1 
 3 
 
 7 
 2 
 
 1 
 
 1 
 1 
 
 1 
 7 
 
 1 
 
 2 
 
 2 
 2 
 
 3 
 16 
 165 
 4 
 6 
 2 
 
 1 
 
 Total 
 
 
 163 
 
 7 
 
 fl 
 
 20 
 
 3 
 
 8 
 
 3 
 
 199 
 
 1 
 
 Excluding Ithomia 
 
 . 109 
 
 7 
 
 * 6 
 
 18 
 
 2 
 
 8 
 
 3 
 
 34 
 
 1 
 
242 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 18. 
 
 Showing the presence or absence, and Color-Changes of tlie 
 "Outer Black" (Area XIII) of the hind wing in the Danaoid 
 Helicon idae. 
 
 Gbnbra. 
 
 Present. 
 
 .Vbsent. 
 
 Changed color. 
 
 Lycorea 
 
 Ituna 
 
 Athesis 
 
 Thyridia 
 
 Athyrtis 
 
 Olyras 
 
 Eutresis 
 
 Aprotopos .... 
 
 Dircenna 
 
 Callithomia .... 
 Epithomia .... 
 
 Ceratinia 
 
 Sais 
 
 Scada 
 
 Mechanitis .... 
 Napeogenes .... 
 
 Ithoniia 
 
 Aeria 
 
 Melinaea 
 
 Tithorea 
 
 5 
 
 3 
 
 3 
 
 5 
 
 2 
 
 4 
 
 2 
 
 2 
 
 12 
 
 3 
 
 2 
 
 41 
 
 5 
 
 7 
 
 24 
 
 30 
 
 210 
 
 4 
 
 24 
 
 10 
 
 1 
 
 1 1 changed to rufous or brown. 
 
 Total 
 
 398 
 
 1 u 
 
MAY15U: COLOK AND COLOR-PAITERNS. 
 
 243 
 
 TABLE 14. 
 
 Showing tlu' Number of tlic Marginal Spots of the hind wing in 
 the Danaoid Ileliconidae. 
 
 Genera. 
 
 With- 
 
 l' 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 
 
 out 
 spots. 
 
 spot. 
 
 1 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 spots. 
 
 1 
 
 spots. 
 
 spots. 
 
 Lycorea . . . 
 
 
 
 
 
 3 
 
 
 Ituna . . 
 
 
 
 2 
 
 1 
 
 
 
 
 
 
 
 
 
 Athesis 
 
 
 
 
 
 
 1 
 
 1 
 
 
 1 
 
 
 
 
 Thyridia . 
 
 
 
 4 
 
 1 
 
 
 
 
 
 
 
 
 
 Athyrtis . 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 1 
 
 Olyras . . 
 
 
 
 :! 
 
 
 1 
 
 
 
 
 
 
 
 
 Eutresis . 
 
 
 
 1 
 
 
 
 
 
 
 1 
 
 
 
 
 Aprotopos 
 
 
 
 
 
 
 2 
 
 
 
 
 
 
 
 Dirceniia . 
 
 
 
 10 
 
 
 1 
 
 
 
 
 
 
 1 
 
 
 Callithoinia 
 
 
 
 2 
 
 
 1 
 
 
 
 
 
 
 
 
 Epitliomia 
 
 
 
 1 
 
 
 
 
 
 1 
 
 
 
 
 
 Ceratinia . 
 
 
 
 18 
 
 8 
 
 1 
 
 2 
 
 5 
 
 6 
 
 2 
 
 4 
 
 1 
 
 
 Sais . . . 
 
 
 
 4 
 
 
 
 
 1 
 
 
 
 
 
 
 Scada . . 
 
 
 
 ;! 
 
 
 
 
 1 
 
 1 
 
 2 
 
 
 
 
 Median itis 
 
 
 
 18 
 
 
 1 
 
 1 
 
 2 
 
 2 
 
 
 
 
 
 Napeogenes 
 
 
 
 21 
 
 
 1 
 
 1 
 
 1 
 
 5 
 
 
 1 
 
 
 
 Ithoniia . 
 
 
 
 164 
 
 
 8 
 
 6 
 
 12 
 
 9 
 
 (5 
 
 6 
 
 1 
 
 
 Aeria . . 
 
 
 
 4 
 
 
 
 
 
 
 
 
 
 
 Melinaea . 
 
 
 
 19 
 
 
 
 
 1 
 
 
 
 3 
 
 1 
 
 
 Tithorea . 
 
 
 
 4 
 
 
 
 
 2 
 
 2 
 
 2 
 17 
 
 
 4 
 
 
 Total .... 
 
 279 
 
 
 
 14 
 
 13 
 
 26 
 
 25 
 
 16 
 
 1 
 
244 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE 15. 
 
 Showing the Color- Variations of the "Inner Kufoiis" Area of the 
 fore wing in Helioonius and Eueides. 
 
 
 Rufous. 
 
 KedcUsh 
 rufous. 
 
 Yellow. 
 
 Ocher. 
 
 White. 
 
 Black. 
 
 Irides- 
 cent 
 blue. 
 
 Heliconius . . 
 Eueides . . . 
 
 35 
 U 
 
 8 
 8 
 
 11 
 
 1 
 
 2 
 
 20 
 3 
 
 20 
 
 Total . . . 
 
 49 
 
 U 
 
 : 
 
 2 
 
 32 
 
 26 
 
 Note: The costal edge of the fore wing is always black. 
 
 TAIJLE 16. 
 
 Showing the Variations affecting tlie " Inner Hlack " Area of the 
 fore wing in Heliconius and Eiieides. 
 
 
 Black. 
 
 Iridescent blue. 
 
 Rufous. 
 
 Helioonius 
 
 Eueides 
 
 85 
 13 
 
 26 
 
 5 
 
 Total 
 
 98 
 
 26 
 
 5 
 
 Note: In M species of Heliconius the inner rufous is entirely suffused with black. 
 
 TABLE 17. 
 
 Showing the Color- Variatioii.s of the " Inner Yellow " Area of the 
 fore wing in Heliconius and Eueides. 
 
 
 Rufous. 
 
 Red. 
 
 Yellow. 
 
 Ocher. 
 
 White. 
 
 Black. 
 
 Irides- 
 cent 
 blue. 
 
 Heliconius . . 
 Eueides . . . 
 
 11 
 
 
 12 
 
 54 
 
 12 
 
 20 
 
 12 
 
 3 
 
 Total . . . 
 
 17 
 
 12 
 
 54 
 
 12 
 
 20 
 
 12 
 
 3 
 
MAY UK: COLOR AND COLOU-l'ATTRRNS. 
 
 245 
 
 TAHLE 18. 
 
 The " Middli' IJlack " Area in the fore winy in Ileliconiiis is 
 present as a l}lack or I^hie Marking in 99 Species and absent in 12. 
 It is present as a lilack Mark in all 1 8 Species of Eueides. 
 
 TAHLK 19. 
 
 Showing the Color- Vai'iation of the " Middle Yellow " Area of the 
 fore loin;/ in Helieonius and Eueides. 
 
 
 RufDUS. 
 
 lletUlish 
 rufous. 
 
 Yellow. 
 66 
 
 Ocher. 
 
 White. 
 
 Black. 
 
 Heliconius 
 Kueides . . 
 
 11 
 5 
 
 12 
 
 12 
 
 23 
 1 
 
 
 Total . ■ . 
 
 16 
 
 12 
 
 65 
 
 12 
 
 24 
 
 
 TABLE 20. 
 
 Showing the (Jolor- Variations of the "Outer Yellow " Area of the 
 fore wing in Heliconius and Eueides. 
 
 
 Kufous. 
 
 RviUlish 
 rufous. 
 
 Yellow. 
 
 Oclier. 
 
 White. 
 
 Blacl(. 
 
 Irides- 
 cent 
 blue. 
 
 Heliconius . . 
 Eueides . . . 
 
 2 
 
 1 
 
 47 
 
 6 
 
 24 
 
 1 
 
 3S 
 11 
 
 4 
 
 Total .... 
 
 2 
 
 1 
 
 47 
 
 6 
 
 26 
 
 44 
 
 4 
 
 TAHLE 21. 
 
 The " Outer Black " Area of the /ore wing in all the 111 species 
 of Heliconius known to me is Black or L-idescent Blue. 
 It is Black in all 18 Eueides. 
 
24(3 
 
 BULLETIN: MUSEUM OF COMl'ARATIVE ZOOLOGY. 
 
 TABLE '22. 
 
 Sliowiiig tlu' .Manner of Occurrence of tlie IMargiTial Spots of tlio 
 /o)-e wiiiff in Heliconins and Eueides. 
 
 
 With- 
 out 
 spots. 
 
 1 
 spot. 
 
 2 
 spots. 
 
 3 
 spots. 
 
 4 
 spots. 
 
 5 
 spots. 
 
 u 
 spots. 
 
 7 
 spots. 
 
 8 
 spots. 
 
 il 
 spots. 
 
 Heliconius . . 
 Eueides . . . 
 
 89 
 14 
 
 
 6 
 3 
 
 8 
 
 2 
 
 5 
 
 3 
 3 
 
 4 
 
 1 
 
 2 
 2 
 
 — 
 
 1 
 
 Total .... 
 
 103 
 
 2 
 
 5 
 
 5 
 
 1 
 
 TAI5LK 23. 
 
 Showing the Variations affecting tlie " Longitudinal Black Stripe " 
 of the fore loing in Heliconins and Eueide.s. 
 
 
 Whole area suf- 
 fused with black. 
 
 Well developed as 
 a black stripe. 
 
 .■Vbsent ( area suf- 
 fused with rufous). 
 
 Heliconius .... 
 Eueides 
 
 •2. 
 
 23 
 
 1(1 
 
 13 
 
 Total 
 
 77 
 
 30 
 
 18 
 
 TABLE 24. 
 
 Showing the Color- Variations- of the " Liner Rufous" Area of the 
 hind wing in Heliconius and Eueides. 
 
 
 1 
 
 % 
 
 i 
 
 >< 
 
 i 
 1 
 
 n 
 u a 
 
 ii 
 
 1 
 
 
 
 
 
 Heliconius . . 
 
 42 
 
 7 
 
 3 
 
 
 1 
 
 10 
 
 28 
 
 10 
 
 1 
 
 5 
 
 Eueides . . . 
 
 15 
 
 
 
 2 
 
 
 1 
 
 
 
 
 
 Total .... 
 
 57 
 
 7 
 
 3 
 
 2 
 
 1 
 
 17 
 
 20 
 
 10 
 
 1 
 
 - 
 
MAYER: COLOR AND COLOR-PATTKRNS. 
 
 247 
 
 TABLE 25. 
 
 Showing the Variations of the "Midrllo Black Stri|)i' " of the Itiml 
 wing in Ileliconius and Eueides. 
 
 
 Well (level- 
 
 o|ie(l as a more 
 
 or less distinct 
 
 stripe. 
 
 Absent (suf- 
 fused with 
 black). 
 
 59 
 
 Absent 
 (l)lace taken 
 by red or ru- 
 fous). 
 
 5 
 
 10 
 \ 
 
 Absent 
 
 (]>lace taken 
 
 by oolier 
 
 color). 
 
 Helioonius .... 
 Eueides 
 
 47 
 C 
 
 1 
 
 Total 
 
 53 
 
 59 
 
 15 
 
 1 
 
 TAMLE 20. 
 
 Showing the Color-Variations of the "Outer I?ufous" Area of the 
 hind roiiu/ in Fleliconius and Eueides. 
 
 Ileliconius 
 Eueides . 
 
 Total . . 
 
 Rufous. 
 
 Reddish 
 rufoufi. 
 
 Yellow. 
 
 White. 
 3 
 
 Ocher. 
 
 Black. ; 
 
 30 
 
 17 
 
 47 
 
 4 
 
 19 
 19 
 
 
 49 
 1 
 
 50 
 
 4 
 
 3 
 
 
 Iride.i- 
 cent 
 blue. 
 
 TABLE 27. 
 
 Tlie " Outer Black " Area of tlie hind whiff is Black in 106 species 
 of Ileliconius, White in 4, and Yellow in 1. It is Black in all the 18 
 species of Eueides known to me. 
 
248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY. 
 
 TABLE -28. 
 
 Showing tlic Number of Species in each (Tenus of the IleUooiiidae 
 examined, and also tlie Number known according to tlie Enumera- 
 tion of Staiidinger ('84-88). 
 
 (flCMIillA. 
 
 Number of species ex- 
 iiinined by me. 
 
 Number of species 
 known to .Staiidinger 
 
 (•84-'88). 
 
 Lycorea 
 
 Ituna 
 
 Athesis 
 
 Tliyridia 
 
 Athyrtis 
 
 Olyras 
 
 Eutresis 
 
 Aprotopos 
 
 Dircemia 
 
 Callithomia 
 
 Epilhoinia 
 
 Ceratinia 
 
 4 species and 1 var. 
 3 
 3 
 5 
 2 
 4 
 2 
 2 
 12 
 3 
 
 41 
 
 7 
 
 10 spoH'ies, 14 var. 
 
 30 
 
 212 
 
 4 
 
 24 
 
 10 
 
 4 species and 1 var. 
 4 
 4 
 4 
 2 
 5 
 2 
 4 
 20+ 
 8 
 2 
 
 60+ 
 4 
 
 .Scada 
 
 Mechanitis 
 
 Napeogenes 
 
 Ithomia 
 
 !) 
 
 10 species, 13 var. 
 
 30+ 
 
 230+ 
 
 4 
 
 Melinaea 
 
 Titliorea 
 
 25 
 18 
 
 Total of Danaoid Heliconidae. 
 
 400 
 
 453+ 
 
 Helicoiiius 
 
 111 
 18 
 
 130 
 24 
 
 
 
 
 529 
 
 607+ 
 
 
JIAYKH: COLOU AND COLOR-l'ATTKUNS. 249 
 
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 Zeit.sclir., Bd. 28, p. 163-165. 
 Srnka, A. 
 '85. Neue Siidamerikanisohe Danaidae und Heliconidae. Berlin. Ent. 
 Zeitschr., Bd. 29, p. 121-1.30, Tiif. 1. 
 Staudinger, (). 
 '82. Oil three new and interesting Species of Rhopalocera. I'roc. Zool. Soc. 
 London, 1882, p. 890-^98, pi. 24. [New Heliconidae.] 
 Staudinger, (). 
 '84-'88. Exotische Tagfalter. (Exotische Schnietterlingc von Staudinger 
 und Schatz.) Fiirth, 1. Theil, 8:^:! pp. 100 Taf. [New Heliconidae.] 
 Urech, F. 
 
 '91. Beobachtungen iiber die verschiedenen Schuppenfarben und die zeit- 
 liche Succession ilires Auftretens (Farbenfelderung) auf den I'uppen- 
 fliigelchen von Vane.ssa urticae und io. Zool. Aiizeiger, .lalirg. 14, p. 4(i(i- 
 473. 
 Urech, F. 
 '92. Uber Eigenschaften der Schuppenpigmenle einiger I.cpidopteren- 
 Species. Zool. Anzeiger, Jahrg. 15, p. 299-;!0(i. 
 Urech, F. 
 '93. Beitrilge zur Kenntniss der rarb(i von Insektousehnppen. Zeitschr. f. 
 wiss. Zool., Bd. 57, p. 300-384. 
 Van Bemmelen. See Bemmelen, .1. F. van. 
 Wallace, A. R. 
 '07. [Theory of Warning Coloration.] 'Praiis. Kiit. Soc. Lonilou, Ser. 3, 
 Vol. 5, p. 80-81 , I'roo. 
 Wallace, A. R. 
 
 '89. Darwinism. London juid New York, 1(1 + 49-1 pp. [p. 2:!:'.--J(i7.1 
 
254 BULLETIN: MUSEUM OF COMPAHATIVE ZOOLOGY. 
 
 Walsinghaiii. 
 '«:"). Oil some probable Causes of a Tendency to melanic Variation in 
 Lepidoptera of liigli Latitudes. Trans. Yorkshire Union, pt. 5, p. 113-140. 
 Cf. Nature, Vol. ;!1, p. QOr). 
 Watkins, W. 
 '91. Ornithoptera trojana, Staudinger. Entomologist, Vol. 24, p. 178-17!), 
 pi. 4. 
 Weismaim, A. 
 '82. Studies in the Theory of Descent. Translated and edited by Haphael 
 Meldola. London, 2 Vols., 28 + 72!) pp., 8 pis. [p. 1-1.18.] 
 Weyiuer, (1. 
 '"■'). Exotische Lepidopteren. Stettiner ICnt. Zeit., .lahrg. ."!(!, j). ;i()8-385, 
 Taf. 1-2. 
 Weymer, G. 
 '84. Exotische Lepidopteren. II. Stettiner Ent. Zeit., .lalng. 4.'>, \i. 7-28, 
 Taf. 1-2. 
 
MAYK,H: roi.OI! A\l) COT,()H-l'ATTKRNS. 
 
 TABM'; OK roXTRNTS. 
 PART A. 
 
 (lIvXKIIAI. PlIKNOMKNA 1)1' C'oi.DII IN l,i: |-| DOI'TI.II ^ . 
 
 I. Cl.AS.SIKir.VTlDN OF (\>I,()ICS. 
 
 I'AdK 
 
 (1) Pigmental Colors 18» 
 
 (2) Structural Colors 170 
 
 (3) Combination Colore 171 
 
 (4) Quantitative Determination of Pigmental Colors .... 172 
 
 (5) Spectrum Analysis of Colors of IiC])i(lopt.pra 173 
 
 ((!) Summary of Hesults 174 
 
 II. TUE ESSKNTIAI, NaTIUIK OF I'Ki.MKXTA I. Coi.Oll rx LkiTDOI'TK It A . 
 
 ( 1 ) Pigments of Larvae 1 74 
 
 (2) Pigments of Imagines 17') 
 
 III. DkVKUH'MKNT ok TMIi VAlll<)l!S Cor.OllS IN TIIK PlI'Al. WlXCS. 
 
 (1) Historical Account of previous Researches 17(i 
 
 (2) Development of Color in the Pupal Wings of Callosaniia promethea 178 
 
 (3) Development of Color in the Pupal Wings of Danais plexijjpus 
 
 (archippus) 180 
 
 IV. TnK Laws wnicn oovkrn tiik Coi.or-Pattkkns or Br-rrKid-LiEs and 
 Morns. 
 
 (1) Historical Account of previous Researches 181 
 
 (2) Laws of Color- Patterns 183 
 
 (3) Detailed Discussion of the Laws of Color- Patterns . 
 
 (4) Origin of (^olor-Variations 
 
 Bibliography of Color-Aberrations 
 
 (5) Climate and Melanism 
 
 (»i) Relation between Climate and Colors of Papilios 
 
 18S 
 189 
 190 
 190 
 191 
 
 V. 'I'liK Cacshs wnicn have leu to the Develoi'.ment and Prksehvation 
 OK the Scales ok the Lepidoi'Teiia. 
 
 (1) ICxperiments and Theory 192 
 
 (2) Summary of (Conclusions 197 
 
 PART B. 
 
 Coi.oh-Vaiiiations in the Heliconidae. 
 
 I. (iENKiiAi. (Causes which nETEWMiNB Coloration in the Heliconidak. 
 
 IT. Methods itiisied in studying the Coi.or-Patterns ok the 
 Heliconidak. 
 
 (1) The Two Types of Coloration in the Danaoid Ileliconidae 204 
 
 (2) Detailed Description of the Melinaea 'I'ype of Coloration 206 
 
256 lU'LLETIN: MUSEUM OF COMPARATIVE 7A)OUH',Y. 
 
 III. General Disciissiox op the Coi.or-Patteiins and oi- Mijik'HV is 
 THE Genera Heliconius and Eukides. 
 
 (1) The Four Color-Types in the Genus Heliconius . . . 208 
 
 (2) Mimicry between the genus Heliconius and the Danaoid 
 
 Group 20!) 
 
 (3) The Three Color-Types in the Genus Eueides . 210 
 
 (4) Detailed Discussion of Plates 5-8 210 
 
 IV. General Dlscussion ok the Color-Patterns and of Mimicry 
 
 AMONO THE DaNAOID HbLICONIDAE. 
 
 (1) The Origin of the two Types of Coloration . . . 214 
 
 (2) Mimicry among the Danaoid Heliconidae .... 2l(i 
 
 V. Quantitative Determination ok the Variations of the char- 
 acteristic Wing-Markinos in the Acraeoid and Danaoid 
 Heliconidae. 
 
 (1) Variations of the "Inner Rufous" Areas of the Fore and 
 
 Hind Wings 217 
 
 (2) The "Inner Black "Spot 218 
 
 (3) Variations of the " Inner Yellow " and "Middle Yellow " 
 
 Areas 210 
 
 (4) Variations of the " Middle Black " Mark of the Fore Wing 221 
 
 (5) Variations of the " Outer Yellow " Area of the Fore Wing 221 
 
 (6) The Relative Perraaneiioy of the Black Areas upon the Fore 
 
 and Hind Wings 222 
 
 (7) The "Middle Black Stripe "of the Hind Wing 222 
 
 (8) Variations of the Marginal Spots of the Fore Wing . 223 
 
 (9) The Marginal Spots of the Hind Wing .... 223 
 
 VI. Comparison ok the Color-Variations ok the Papilios of 
 South America with those of the Heliconidae. 
 
 PART C. 
 
 General Sum.mauv ok Results iikmeved to he new to Soiknck. 
 
 Tables 230 
 
 Bibliography 240 
 
 Explanation of Plates 
 
Ma YBB. — Color and (^olnr-l'iittPrn*. 
 
 Abbreviations. 
 
 B. Back surface covered with wings. (). Orifice for admission of light. 
 F. Front surface covered witli wings. S. Spectroscope. 
 
 Arrow indicates directions of rays of light. 
 
 Fig. 1. Perapeotive view of spectroscopic apparatus used in determining the 
 
 compcxsition of the colore of Lepidoptera. 
 Fig. 2. Horizontal section of same. See p. 175. 
 Fig. .3. Pendulum used in determination of the frictional resistance between 
 
 the air and the wings of Lepidoptera. See p. 103. 
 Figs. 4, 6. Diagrams to illustrate Keeler's method of projection, as applied to 
 
 Lepidoptera. See p. 207. 
 
Mayer- Coj,or and Color Patterns. 
 
 Plate. 1. 
 
 M 
 
 /> \ ^\ 
 
 
 ^ 
 
 
 
 
 
 
 =: 
 
 ® 
 
 
 n 
 
 ^ 
 
 — 
 
 M 
 
 lUliP 
 
 I 
 
 ®m 
 
 Z' !• / l' / a' 3 45 6 7 8D0H 
 I' l' i» 
 
 BullMus Comp. ZoolVol XXX. 
 
Maykh, — Color and Color ratteiiis. 
 
 Diagrama to illustrate the laios which govern the Color-Patterns of Lepidnpterti. 
 
 Fig. 6. Euthalia bellata (W. L. Distant, '82-'80, Plate 4:i, Fig. 12). Illustrates 
 
 the law of bilaterality of spots. See p. 183. 
 Fig. 7. Zethera inusa (G. Semper, •8(i-'92, Taf. 7, Fig. 10). Bilaterality of 
 
 double spots. See p. 188. 
 Fig. 8. Eye-spots in Morpho. See p. 182, 18;{. 
 Fig. {). Parthenos gambrisius (W. L. Distant, ■82-'8(i, Plate 1 1 , Fig. 7 ). A series 
 
 of complex spots, each one being similar to the rest, and bilaterally 
 
 symmetrical. 
 Figs. 10, 11. Ornithoptera urvillana and <). priamus (R. II. F. Ripon, '89-'93). 
 
 Spots within spots, all being bilaterally symmetrical. 
 Figs. 12, 13. Hestia jasonia and H. leuconoe. Axis of lateral symmetry 
 
 (H, H),for spots passes through center of interspace. H. jasonia 
 
 (F. Moore, '90-'96, Plate 3, Fig. 1). H. leuconoe (G. Semper, 
 
 '8(i-'92, Taf. 1, Fig. 3). 
 Fig. 14. Papilio emalthion, to illiustrate fusion of two rows of spots. 
 Fig. IT). Ornithoptera trojana, an apparent exception to the law of bilaterality. 
 
 Sec p. 187. 
 Fig. l(i. Limenitis proserpina (S. H. Scudder, '88-'89, Plate 2, Fig. 9), showing 
 
 fusion of two rows of diiferently colored spots. See p. 187. 
 Fig. 17. Saturnia spini, false eye-spot. See p. 187. 
 Fig. 18. Cases of degeneration of bands of color. See p. 184. 
 Fig. 19. Missanga patina (F. Moore, '90-'9(i, Plate 72, Fig. 2"'). Exceptional 
 
 form of eye-spot. See foot note p. 186. 
 Figs. 20-23. Hypothetical conditions of coloration, not found in nature, being 
 
 contrary to the laws of color-pattern. See p. 188. 
 
Mayer- Color and Color Paherns. 
 
 Plate. 2. 
 
 BULL.MUS COMP. ZOOL.VOL. XXX, 
 
MwEii. — Color and Color-Patterns. 
 
 To illustrate color-development in Callosainia prometkea and Danais plexippiiit. 
 
 Fig. U4. Enlarged view of pupal wing of C. promethea in the "white stage." 
 
 See p. 178. 
 Fig. 25. Scale from wing of C. promethea in white stage of color-development, 
 
 sliowiug the total absence of pigment in the scale. See p. 178. 
 Fig. 2(i. Scale from light drab-colored area of matnre wing of C. promethea. 
 Figs. 27, 3(), and 33. Successive stages in the formation of color in pupal hind 
 
 wing of C. promethea. 
 Figs. 28, 37-40. Successive stages in the formation of color in the \n\yA\ furc 
 
 wing of ? C. promethea. See p. 17i), 180. 
 Figs. 20, 30-3!). Successive stages in the formation of color in the pupal wings 
 
 of (J ('. promethea. (Fig.s. 20, ;!0-33, 3r) fore wing; Fig. 34 hind 
 
 wing.) See p. 170, 180. 
 Figs. 34 and 41. Pupal hind wings of C. promethea, respectively mature $ 
 
 and ?. 
 Figs. 42-45. Successive stages in the color-development of I), plexippus. See 
 
 p. 180-181. 
 
Mayer.- Color and Color Pattern : ■ 
 
 Plate. 3. 
 
 :.!. Mus. CoMP. ZouL Vui. KXiX.. 
 
1 
 
Maver. — Color ami Color-Patterns. 
 
 I'LATK 4. 
 
 Systematic aiiali/sis of the chariv.teriatic markings upon the xuinys of the Ileli- 
 r.nnidae. Homologous markings are designated by the same numerals, I, II, 
 III, etc. 
 
 Fig. 46. Lyoorea ceres; an example of the '■ Melinaea type" of coloration. 
 
 See p. 205. 
 
 Fig. 47. Thyridia p.sidii ; an example of the " Ithomia type " of coloration. 
 
 See p. 200 ami Plate 7, Fig. 7!). 
 
 Fig. 48. Melinaea paraiya. 
 
 Fig. 49. Ceratinia ninonia. 
 
 Fig. 50. Heliconius aiitiochus. 
 
 Fig. 51. Niujeogenes due.ssa. 
 
 Fig. 52. Itlwmia sao. 
 
 Fig. 6:i. Melinaea gazoria. 
 
 Fig. 54. Ithomia nise. 
 
 Fig. 55. Mechanitispolyiiinia. 
 
 Fig. 50. Eueides cleobaea. 
 
 Fig. 57. Tithorea furia var. 
 
 Fig. 58. Heliconius eucrate. 
 
 Fig. 59. Heliconius melpomene. 
 
 Fig. 00. Heliconius erato. 
 
Mayer - Color and Color Patterns, 
 
 Plate. 4. 
 
 BullMus Comp. Zool Vol XXX. 
 
Mavek. — Color and C:olor-Patteriis. 
 
 I'LATK T). 
 
 Color-patterns of tlie Anliochiin anil Krnto ijronpn of the IleUc.onklac projccleii 
 by Keeler's method. See p. 207. 
 Figs. 01, ()2. Heliconius saraaiul II. aiitioclius, to .sliow variation of yellow to 
 
 white. See p. 2tO, and Fig. 51), I'late 4. 
 Fig. (!.'!. M. galanthus, showing devclopiuent of wliite. 
 Fig. ()4. 11. charitonia, rows of double spots. 
 Fig. ()o. H. phyllis, close relation between yellow and red. 
 Fig. (i(). II. ricini. 
 
 Figs. (>7, 08. II. erato; two color-types. 
 Fig. (ii). IT. Claudia; an example of the Sylvanus group. 
 
Mayer - Color and Color PAnERNS. 
 
 Plate. 5. 
 
 HIND WING. 
 I" l'' 1^ U III W V VI VII VIII 
 
 FORE WING, 
 n 111 IV V v: 
 
 BuLL.Mus CoMR ZoolVol, XXX. 
 
Mayek. — Color :iii(l Color-l'atterns. 
 
 Color-patterns of the Melpomene group of lleliconiun and of the genus 
 Eueides. 
 
 Fig. 70. H. melpomene, the type of the Melpomene group. See p. 212, and 
 Fig. oi), Plate 4. 
 
 Fig. 71. H. melpomene var. callicopi.s, showing the breaking up of the red 
 area of the primaries. 
 
 Fig. 72. H. melpomene var. cybele ; the fore wing has assumed a color-pattern 
 which recalls the " Melinaea type" of coloration found in the 
 Danaoid lleliconidae. 
 
 Fig. 73. II. thelxiope, derived phylogenetically from H. melpomene, and 
 showing a rather close approach to the " Melinaea type" of color- 
 ation. See p. 212. 
 
 Fig. 74. H. vesta. 
 
 Fig. 7"). Eueides thales $ ; represented to show the close resemblance of il.s 
 color-pattern to H. vesta. See p. 212. 
 
 Fig.s. 70, 77. E. mereaui and E. aliphera. E. mereaui is intermediate in color- 
 pattern between E. thales and E. aliphera. 
 
 Fig. 78. Eueides cleobaea, to show the close approach of this insect to the 
 "Melinaea type" of coloration. 
 
Mayer- Color and Color PAnERNs. 
 
 HIND WING 
 I" i'' i"^ II III IV V VI VII VIII 
 
 Plate. 6. 
 
 FORE WING 
 I' l'' Jl III IV V VI 
 
 Helkviiius 
 nielpomene'linn.. 
 
 '/ irif/pomen^ var. 
 ntlfycopis Gum. 
 
 /Imelpom^e. var. 
 (yheie Cram. 
 
 UIBBttga 
 
 f! vesl/v Gxuiv. 
 
 /■Meidfs Uiales 
 
 IiLAJu 
 
 mk 
 
 (ram. J' 
 
 I'.niereaiii 
 
 Hiibn. 
 
 E. /i/iphem 
 (iodi. 
 
 /■' ilfoba-ea 
 /Kb,i. 
 
 ACM del H M.lsti lilh IdsIh 
 
 Bull Mus Comp. ZoolVol XXX. 
 
M A VKR. — Color anil C'olor-I'attcrns. 
 
 Intended to ahow sDtnc types of coloration which are found in the Vanaoid 
 Ileliconidae, and also the remarkable individual variation in Meehanitis isthmia. 
 
 Fig. 7f>. Thyridia psidii, an example of the "Ithomia" type of coloration. 
 
 See p. 213, and Plate 4, Fig. 47. 
 Fig. 80. Napeogene.s cyrianassa, showing senii-translncent condition of wings. 
 
 See p. 21:!. 
 Fig. 81 . Ceratinia vallonia. 
 Fig. 82. Melinaea parallelis; albinism of spots on primarie.s ; black band of hind 
 
 wing broken in the middle. See p. 188, 213. 
 Fig. 83. Ceratinia leucania, which probably mimics M. pai'allelis. 
 Figs. 84-87. Meehanitis isthmia, showing remarkable individual variation in 
 
 the black stripe of the hind wings, and also in the " inner yellow " 
 
 spot of the fore wings. See p. 184, 213. 
 
Mayer -CoioR AND Color Patterjis 
 
 Plate. 7. 
 
 HIND WING^ FORE WING 
 
 i'' I'- I! Ill N V VI VII VIII I" i'' i: 111 IV ■.' 
 
 [MID^ 
 
 7'li.vnduv 
 p.ii'du f.utji . 
 
 BliU. 
 
 ( tniiuualmni/iM 
 
 Mi'rhaiiUts 
 u-il/irtini Ratfg. 
 
 ^ i; j( jfl 6 Meisftl lilfi Boslen 
 
 BullMus Comp. ZoolVol XXX. 
 
Mavku. — Color and Coloi-l'atterns. 
 
 Illustrates the mimicry between members of the " St/lvanus " group of the tjenus 
 Heliconius and various Meiinaeas, etc. 
 
 Fig. 88. Heliconius eucoma, an example of the " Sylvanus " type of coloration 
 
 in genus Heliconius. See p. 214. 
 Figs. 89, !)0. Heliconius dryalus and Melinaea paraiya ; close resemblance of 
 
 their color-patterns. See p. 214. 
 Figs. 91, 92, 93,94. Respectively Heliconius eucrate, Melinaea thera, Eueides 
 
 dianasa, and Meclianitis polynmia; showing close resemblance 
 
 between color-patterns. See p. 214. 
 Figs. 95, 90. Heliconius sylvana and Melinaea egina; tliese two forms are said 
 
 by Bates to mimic each other. See p. 214. 
 
Mayer- Color and Color Pattems 
 
 Plate. 8. 
 
 H IN n WIN (I 
 
 FORE WING 
 
 1" 1° r li !:■ " ' /I VII VIII !■> l'' II III iv V VI 
 
 //I'/uv/i/'us 
 •h'n/iii (mill. 
 
 . ^Icliiiiira 
 I//II/I (iiirn 
 
 Bull Mus Comp. Zool Vol, XXX, 
 
Mavkk— Color ami ("olor-FatleriiB. 
 
 PLATP; !). 
 
 lliagrams to illustrate color-variations. 
 
 The various colore are laid off at definite intervals along the axis of abscissae, 
 and the ordinates represent the number of species which exhibit the various 
 colors. 
 
 Fig. 07. Hepresents the color-variations of the " inner rufous " area of tlii^ fore 
 and hind wings in the Danaoid Heliconidae. The full line re))resents 
 the variations of the fore wing. The dotted line those of the hind 
 wing. The closeness of these two lines shows the intimate relations 
 between the color-variations of the " inner rufous " areas upon fore 
 and hind wing. See p. 218. 
 
 Fig. 08. 'I'he full line represents color-variations of " inner yellow " spot of fore 
 wings in Danaoid Heliconidae. The dotted line represents same for 
 "middle yellow." It is apparent that the "inner yellow" is more 
 variable than the -'outer yellow." and also that the variations of 
 both are quite similar to thr)se of the "inner rufous." See p. 210. 
 
 Fig. 00. Color-variations of "inner rufous" areas of Acraeold Heliconidae. 
 The full line represents the fore wing and the dotted line the hind 
 wing. See p. 218. 
 
Mayer- Coior and Color Patterns 
 
 Plate. 9. 
 
 T 
 
 ', '■ 
 — 1 
 
 v~~ 
 
 ■ : i 1 
 i i. . 
 
 
 ---- 
 
 
 1 
 
 \ 
 
 ^ 
 
 1 
 
 i 
 i 
 
 — f 
 
 :| : 
 \ ."!"■•< i:^__ 
 
 • 
 
 :: J. 
 
 ; j - i 4"j 
 \':i rr 
 
 ■ • ■■ '] 
 . .J 
 
 3|S I 
 
 SO r 
 
 li^ 
 
 
 
 
 
 i 
 
 . 
 
 
 
 
 
 
 
 
 
 
 r-. L 
 
 j rj 
 
 ^[!- 
 
 i::^ 
 
 \\l[ 
 
 Ml! 
 
 ;■'■- 
 
 1 
 
 V 
 
 
 1 
 
 
 
 i 
 
 t ^"^ 
 
 1 
 
 )0 - 
 
 1 1 
 
 
 ! i i 
 1 ' 
 
 : ; I i 
 
 -' 
 
 I 
 
 h 
 
 
 -;::h 
 
 llij 
 
 
 li ■ : : 
 
 tjlT 
 
 ' 
 
 -U-LL 
 
 -i- 
 1. 
 
 5 - 
 
 - 
 5 - 
 
 1- 
 
 It!" 
 
 1 : 1 
 
 it' 
 j ' 1 
 
 Hi' 
 
 
 !:i 
 
 ^■^ 
 
 ^ 
 
 1 ■•' 
 
 — - 
 
 ■iil' 
 
 -J— 
 
 
 L L 
 
 ■V- 
 
 'j1 
 
 in 
 
 ■fit* 
 
 r 
 
 i 
 
 'tl 
 
 BullMus Comr Zool.Vol. XXX. 
 
Mayer. —Color and Color-Patterns. 
 
 PLATE 10. 
 
 Fig. 100. Coloi-variatious of " inner yellow " spots on fore wings of Aoraeoid 
 
 Hcliconidae. Tlie full line represents the " inner yellow," the dotted 
 
 line the " middle yellow." See p. 221 . 
 Fi';. 101. Variations of marginal spots upon fore wing in Danaoid Heliconidae. 
 
 These spots tend to appear either as 2 or :i, or as (> or 7 spots. See 
 
 p. 22:!. 
 Fig. 102, Variations of marginal spots of hind wings in Danaoid Heliconidae. 
 
 These spots tend to appear either as 4 or as 5 spots. See p. 22:5. 
 
Maykk - CninR and Color Pait 
 
 Platk, 10. 
 
 BullMus Co MR ZooLVm. XXX. 
 
\- 
 
I 
 
The (oliowing Publications of tiie Museum of Comparative Zo51o§y 
 are in preparation: — 
 
 Ueport* on the Kes.iItB of Dredging Operations In 1877, 1878, 1879, and 1880, 1„ charge of Alex- 
 A.Ni-icit A<iA»8./„ by the U. S. Coast Survey Steamer "Blake,- a. follows:- 
 A. .MILNE-KinVAKDS. Crustacea of the " Blake." 
 !■:. lOHLKRS. 'I'lie Annelids of the " Blake." 
 C. HAKl'LAUU. The Coniatula) of the " Blake," with 15 Plates. 
 11. LUDWIU. The tienus I'entacrlnus. 
 A. K. VKKKILL. Tlio Alcyonaria of the " Blake." 
 Illustrations of North American JIAKINE lNVKKl'KBB.vrKS, fr,,,,, DrawiuKs by BuiiK 
 iiAKDr, SdNiiBi,, and A. Agassi/,, prei.are.l under the Direction of L. Aoassiz. ' 
 
 A. AUASSIZ. A Visit to the Great Barrier Keef of Australia. 
 LOUIS CABOT. Immature State of the 0<lunata, Part IV. 
 K. L. MARK. Studies on Lepiilosteua, continued. 
 
 " On Arachintctis. 
 
 R. T. HILL. On the Geology of the Istlimus of Panama. 
 
 On the Geology of Jamaica. 
 CHAKIES W.VCIISAILJTH and FRANK SPRINGER. The North American Fossil Crinoi- 
 dea Uunerata. With an Atlas of 83 Plates. Will be is«ucd as Vols. X.X. a«d XXI. of 
 tiifl Aloinoii*s> 
 
 ^""T^nZl^l"' '"^ ''°'^'^«l'''^^'' I.ABOUATOKV, in charge of Professor E. L. Mahk, 
 
 W WHITNEY. The Histology of Tliyone. 
 T. (i LICE. The Suprareiials In Amphibia. 
 
 Contribution, from the GEOLOGICAL LABOllATOKV, in charge of Professor N. S. SHVLEn 
 
 "'"t^^^lZ::':^"^' ^-^"'^"^ .AUOUATORV, commumcatoa by A.kxa.o... 
 
 A. AGASSlZandA. G. MAYEIl. The Acalephe of the East Coast of the Uni....i «.„„. 
 »n.oo,., i™ " Oil DactylometraqalnquocirraAgass. 
 
 AGASSIZ and WHITMAN. Pelagic Fishes. Part II., with 14 Plates 
 
 Reports on the Resnlts of the Expedition of 1891 of the U. S. Fish Comn.ls.ion steuncr 
 •Albatross," Lieutenant Commander Z. L Ta^nk.,, f. S. N., Commanding incbareeof 
 Albxa.ni>i:kAoassi/., as follows:- " "manning, in charge of 
 
 A.AGASSIZ. TMie Pelagic Fauna. C. F. LIJTKEN and TH. MORTKNSEN. 
 I he IK blnl. Tiig Opiilurldaj 
 
 I F BFNI.-M,.r:ir"f "?"■''"' ^"""*- 0'"*'A'^«- The Acalephs. 
 
 C HU VNM T nie V!".""'- ''■ '" '^''^"X- The Aotinarians. 
 
 IV. HU.VM.I. I osagitue. joHN MUKUAY. The Bottom Spedmons, 
 
 C OllUV TheVinbo r'""""- ROBERT U11.GW.AY. 'Ilie Alcoholic Birds. 
 
 C CHUN ,,^eS!P^™o..ores^ P. SCHIEJIKNZ. The Pleropods and llete- 
 
 J ho fcyos of DceiKSea Crustacea. ropods 
 
 W. H. DALL. TheMollusks. W PFUCY si aiww -ii c. „, 
 
 S. GARM AN. The Fishes. , STF wfrR T^; ., ''f.f "^'""'*»- 
 
 H .1 llANSI-v n.^ /-i_ • . ,. '-• •■'ii-.iM'.t.l'.R. The Heptilos. 
 
 w. A ™^AN"";:;:rz "•"™"^- rr "'h^^^pstf;" v;""'"'-- 
 
 w-'e'^ivIT- ...•''"-^"•"«""-"'- J.oitiid.:.''"'"'"^- '••"'«'"'■'■•—'' 
 
 K. Von r.KNDFNI.-l-i t> -ii n. , «■> «ILSON. The Sponges. 
 
 res-vrntofgnnsofF!;!,:.. '"""""•" "^ '"''' WOODWORTH. The Planarians. 
 
PUBLICATIONS 
 MUSEUM OF COMPARATIVE ZOOLOGY 
 
 AT IIAUVAUU COLLIXiE. 
 
 TlicTo liiive been published of the Bi-LLtriNs Vols. 1. to XXIX. ; 
 of the Memoirs, Vols. I. to XXII. 
 
 Vols. XXVIII. and XXX. ol' the Bili.ktin, and Vols. XIX. to 
 XXI. and XXIII. of the Mkmoiks, are now in conv.se ol" imblieatiou. 
 Tiie liri.i.KTiN and Memoiks are devoted to the pnblii-alion of 
 original work by the Professors and Assistants of the Musenni, of 
 investigations carried on by students and othcr.s in the ditfcrent 
 Laboratorie.s of Natural History, and of work by specialists based 
 upon the Museum Collections. 
 
 The following pnl)lieations are in prciiaralion : — 
 Kqiorls on thf Uesiills of Dredging Opemtiima from 1877 to 1880, in cliarg« of 
 
 Alo.XiUiilcr Agassi/, by ttie U. S. Coast Survey Steamer " Bluke," Lielit. 
 
 C'oTnman.kr ('. D. Sigsbce, U. S. N.,aml Commander J. H. Bartlett, U. S. N., 
 
 Conimaniling. 
 lU-porls on the Uesults of the Kxpe-lition of 18!ll of thv U. S. Kish Commission 
 
 Steamer " All)atross," Lieut. Commiimler Z. I.. Tanner, U. S. N., Com- 
 
 iiianilinj^, in cliarge of Alexander Agassiii. 
 Cuntiibutiotis from the Zoiilogital Laboratory, in charge of I'rof.-' I 1^ 
 
 Mark. 
 Contributions from the Geological Laboratory, in charge of Professor M. S. 
 
 Slialer. 
 Similes from the Newport Miirine T/il....-'i"i-v •■„ umic.'ilea bv AlcNander 
 
 AgHHbi/. 
 
 Sid)scriptions for the pnliiicatii' .iiMimu ■• i 
 
 on the following terms: — 
 
 For the I3i;i,i.i:tin, $4.00 per volume, payable in advance. 
 For the MiofoiKS, $8.00 " 
 These publications are issued in numbers at irregnlar inter- 
 vals ; one volume of the Bulletin (Svo) and half a volimie of the 
 Memoirs (4to) usually appear annually. Each number of tlie Bul- 
 letin and of the IMonioirs is also sold separately. A price list of 
 the publications of the Museum will be sent on application to tlie 
 Director of the Museum of Comparative Zoology, Cambridge, Mass. 
 ALEXANDER AGASSIZ, Direcim: 
 
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 JUL - - 1987