Mimicry in Butterflies

d. Typical form

Chapter 124,746 wordsPublic domain

e. _Alcippina_ form f. _Daira_ form

_Hypolimnas misippus_ [F] g. Typical form h. _Alcippoides_ form i. _Inaria_ form

(After Aurivillius)

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PLATE X

SOUTH AMERICAN BUTTERFLIES

1. _Dismorphia cretacea_ [M] (Pieridae) 2. " _praxinoe_ [M] " 3. " " [F] " 4. _Perrhybris malenka_ [M] " 5. " " [M] (under surface) " 6. " " [F] " 7. _Mechanitis saturata_ (Ithomiinae) 8. _Papilio zagreus_ (Papilionidae) 9. _Protogonius tithoreides_ (Nymphalinae) 10. _Tithorea pseudonyma_ (Ithomiinae)

(NOTE. The figure of the _Mechanitis_ (Fig. 7) is taken from a rather worn specimen. The quality of the orange brown is better shewn by the specimen illustrated in Fig. 7 on Plate XV.)

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PLATE XI

SOUTH AMERICAN BUTTERFLIES

1. _Heliconius sulphurea_ (Heliconinae) 2. _Papilio pausanias_ (Papilionidae) 3. _Heliconius telesiphe_ (Heliconinae) 4. _Colaenis telesiphe_ (Nymphalinae) 5. _Heliconius melpomene_ (Heliconinae) 6. _Pereute charops_ [F] (Pieridae) 7. " " [M] " 8. _Papilio osyris_ [M] (Papilionidae) 9. " " [F] " 10. _Archonias critias_ [F] (Pieridae)

* * * * *

PLATE XII

SOUTH AMERICAN BUTTERFLIES

_Methona confusa_ (Ithomiinae) _Dismorphia orise_ (Pierinae) _Ituna phenarete_ (Danainae) _Castnia sp._ (Heterocera)

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PLATE XIII

MODELS 1. _Papilio nephalion_ 2. " _chamissonia_ 3. " _perrhebus_

MIMICS 4. _Papilio lysithous_ var. _lysithous_ 5. " " var. _rurik_ 6. " " var. _pomponius_

(For further details of this case see Jordan, _I^{er} Congr. Internat. d'Entomologie_, Bruxelles, 1911, p. 396.)

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PLATE XIV

1. _Methona confusa_, ×90 (Ithomiinae) 2. _Dismorphia orise_, ×150 (Pierinae) 3. _Thyridia themisto_, ×90 (Ithomiinae) 4. _Ituna ilione_, ×90 (Danainae) 5. _Castnia sp._, ×60 (Moth)

Microphotographs of the scales of various Lepidoptera in the S. American "Transparency group." For explanation see text, pp. 39-42.

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PLATE XV

CENTRAL AND SOUTH AMERICAN BUTTERFLIES

Illustrating the closely parallel series of patterns occurring in the two distinct groups Heliconinae and Ithomiinae.

1. _Heliconius mirus_ 2. " _telchinia_ 3. " _eucrate_ 4. " _pardalinus_ 5. " _splendens_ 6. _Mechanitis elisa_ 7. " _saturata_ 8. " _lysimnia_ 9. " _egaensis_ 10. " _methona_

* * * * *

PLATE XVI

NORTH AMERICAN BUTTERFLIES

1. _Papilio philenor_ (Papilionidae) 2. " _troilus_ " 3. _Argynnis diana_ [F] (Nymphalinae) 4. _Limenitis arthemis_ " 5. " _astyanax_ " 6. " _archippus_ " 7. " _floridensis_ (= _eros_) " 8. _Danais archippus_ (Danainae) 9. " _berenice_ "

* * * * *

{183}

INDEX

_References to the plates are given in thicker type_

_Acraea_, taken by kestrel, 118; _A. axina_, 122; _A. caldarena_, 122; _A. egina_, 34, VI. 7; _A. encedon_, patterns of different forms in relation to those of _Danais chrysippus_, 29, 144; typical form of, IX. D; _alcippina_ form of, IX. E; _daira_ form of, IX. F; _A. halali_, 122; _A. violae_, 33 note; eaten by lizards, 108; attacked by birds, 110, 117 Acraeinae, as models for African butterflies, 33 Adaptation and Natural Selection, 61 _Adelpha_, 54 African butterflies, mimicry among, 28-36 _Alcidis agathyrsus_, 27, 145, III. 1 _Aletis helcita_, 36 _Amauris echeria_, 30, 148, VIII. 7; _A. hyalites_, 30, VI. 2; _A. niavius_, 30, VIII. 6 _Amphidasys betularia_, rapidity of increase in melanic sport of, 101 _Anosia plexippus_ (= _Danais archippus_), 113 _Anthomysa_, 41 _Aporia agathon_, 149 _Araschnia levana_, seasonal dimorphism in, 130; typical form, VI. 10; _prorsa_ form, VI. 9; _porima_ form, VI. 8 _Archonias_, 43, 56, 145; _A. critias_, XI. 10 _Argynnis diana_, 47, XVI. 3; _A. hyperbius_, 29; as mimic of _Danais plexippus_, 52; in mimicry ring, 66, IV. 3, 4 _Artamus fuscus_, 112 Asilid flies, as enemies of butterflies, 106 _Athyma punctata_, 53

Bates, G. L., on contents of birds' stomachs, 113 Bates, H. W., on mimicry, 9; on resemblances between unpalatable forms, 14; on initial variation in mimetic resemblance, 63; on S. American Pierines attacked by birds, 112 Bateson, 3 _Belenois_, 36 Bingham, on birds eating butterflies, 110 Birds, as enemies of butterflies, 109; stomach contents of, 113; feeding experiments with, 115; colour perception in, 119 Bowater, on _Amphidasys betularia_, 102, 137 note Breeding experiments, with _Hypolimnas dubius_, 30; with _Papilio polytes_, 84; with _Papilio memnon_, 89; with _Papilio dardanus_, 90; with _Pseudacraea eurytus_, 128 Bryant, on birds eating butterflies, 114 _Buchanga atra_, 111 _Byblia ilithyia_, 122 {184}

_Caduga tytia_, 24, 51, II. 2 _Callamesia pieridoides_, 56 _Calotes ophiomachus_, 107; _C. versicolor_, 107 Carpenter, on intermediates in _Pseudacraea eurytus_, 126; on breeding experiments with _Pseudacraea eurytus_, 128 _Castnia_, as mimic, 39, XII. 4; scales of, 41, XV. 5 _Catopsilia_, 121; _C. florella_, 111; _C. pyranthe_, 111 _Cerchneis rupicoloides_, 118; _C. naumanni_, 117 _Cercopithecus pygerythrus_, 121 _Charaxes athamas_, 110 _Citronophila similis_, 35 Classification of butterflies, 18-21 _Colaenis telesiphe_, 38, XI. 4 _Cyclosia hestinioides_, III. 3 _Cymatophora or_, establishment of melanic sport in, 102 note _Cyrestis thyodamas_, 110

Danainae, characteristics of, 22; as models for Oriental butterflies,23; as models for African butterflies, 28 _Danais_, 111, 145; _D. archippus_, 48; eaten by lizard, 108; rejected by bird, 113, XVI. 8; _D. berenice_, 48, XVI. 9; _D. chrysippus_, 23, 28; flight of, 55; in mimicry ring, 65; eaten by lizards, 108; eaten by Bee-eater, 111; eaten by Brown Shrike, 117; rejected by Kestrel, 118; rejected by baboon, 122; local variation in, 132; patterns overlapping with those of _Acraea encedon_, 144; _alcippus_ form, IX. B; _dorippus_ form, IX. C; typical form, IV. 1, VIII. 5; _D. plexippus_, as model for _Argynnis hyperbius_, 52; in mimicry ring, 65; eaten by Liothrix, 115 note, IV. 2; _D. petiverana_, 29, VI. 1; _D. septentrionis_, 23, 111, 112, I. 3; _D. vulgaris_, 150 Darwin, on natural selection, 1; on adaptation, 5; on initial variation in mimetic resemblance, 63; on a difficulty of the mimicry theory, 65 Defence in butterflies, 54 _Delias cathara_, 56; _D. eucharis_, 28, 115, 116, II. 1 de Meijere, on breeding _Papilio memnon_, 89 de Vries, 3 _Dismorphia_, as mimics of Ithomiinae, 38, 42; restricted range of many forms, 51; diversity of pattern in genus, 58; as Batesian mimics, 135; patterns parallel with those of Ithomiinae, 145; _D. astynome_, 151; _D. avonia_, 151; _D. cretacea_, 5, 8, 62, X. 1; _D. orise_, as mimic, 39, XII. 2; scales of, 40, XIV. 2; _D. praxinoe_, as mimic, 57, 62, X. 2, 3; as member of mimicry ring, 134 Distasteful groups, characteristics of, 55

Eltringham, 17 note, 32 note, 36 note _Elymnias_, patterns in genus compared with those of Danaidae, 59, 144; _E. malelas_, 24, II. 6, 7; _E. phegea_, 35, VII. 9; _E. singhala_, 25, I. 9; _E. undularis_, in mimicry ring, 66, 115 note, 116, IV. 5, 6 _Epicopeia polydora_, 27, III. 5 Equilibrium, conditions of in mixed population, 93 _Eresia_, 134, 135 _Eugonia californica_, 114 _Euphaedra ruspina_, 36 _Euploea core_, 25, 108, 110, 112, I. 10; _E. mulciber_, 24, 51, II. 4, 5; _E. rhadamanthus_, 24, 51, II. 8; _E. rafflesii_, 110 Euploeinae, characteristics of, 22; as models for Oriental butterflies, 24; in relation to birds, 111, 112, 115 note {185} _Euripus halitherses_, 24

Feeding experiments, with Mantids, 105; with lizards, 107; with birds, 115; with mammals, 121 Finn, on feeding experiments with lizards, 108; on feeding experiments with Indian birds, 115; on feeding experiments with a Tree-shrew, 121 Flight, different in model and mimic, 55; difference of in _Papilio polytes_ and its models, 82 Fryer, on breeding _Papilio polytes_, 84; on relative abundance of females of_ Papilio polytes_ in Ceylon, 97; on birds eating "unpalatable" butterflies, 112

_Gerrhonotus infernalis_, 108

Haase, on mimicry, 16; on classification of Papilionidae, 25 Hahnel, on S. American Pierines attacked by birds, 112 Hardy, on conditions of equilibrium in a mixed population, 94 Hearsy, on birds eating butterflies, 111 _Hebomoia_, 110 Heliconinae, as models for S. American butterflies, 38 _Heliconius_, 145; _H. eucrate_, XV. 3; _H. melpomene_, as model, 42, 43, XI. 5; _H. mirus_, XV. 1; _H. pardalinus_, XV. 4; _H. splendens_, XV. 5; _H. sulphurea_, 43, XI. 1; _H. telchinia_, XV. 2; _H. telesiphe_, XI. 3 _Herpestes galera_, 121 Hess, on colour perception in birds, 119 Hopkins, on pigment of Pierids, 150 _Hypolimnas dubius_, polymorphism in, 30; as mimic of Danaines,30, VII. 8, 9; breeding experiments with, 30; var. _mima_ compared with model, 148; patterns of in relation to models, 149; _H. bolina_, 25, 117, I. 5, 6; _H. misippus_, 25, 29, as model, 53; flight of, 55; in mimicry ring, 66, 116; eaten by Brown Shrike, 117; _alcippoides_ form, IX. H; _inaria_ form, IX. I; typical form, IV. 7, 8, IX. G

_Ideopsis daos_, III. 4 Initial variation, difficulty of, 63 Insect enemies of butterflies, 105 Intermediates, between different forms of _Pseudacraea eurytus_, 128; in relation to mimicry,129, 140 Ithomiinae, characteristics of, 10; as models for S. American butterflies, 38 _Ituna_, 39; _I. ilione_, 40, XIV. 4; _I. phenarete_, XII. 3

Jacobsen, experiments with _Papilio memnon_, 89 Jordan, 40 note _Junonia_, 111

_Lanius cristatus_, 117 _Limenitis albomaculata_, 53; _L. archippus_, 49, 59, XVI. 6; _L. arthemis_, 47, 49, XVI. 4; _L. astyanax_, 47, XVI. 5; _L. floridensis (= eros)_, 49, XVI. 7; _L. proserpina_, 47 Lizards, as enemies of butterflies, 107 Local varieties, in connection with mimicry, 132 Lycaenidae, as mimics in Africa, 35 _Lycorea_, 145

McAtee, on feeding experiments with birds, 118 Mammals, as enemies of butterflies, 121 Manders, on feeding experiments with lizards, 107; with birds, 117 {186} Mantids, as enemies of butterflies, 105 Marshall, on Müllerian mimicry, 72; on feeding experiments with Mantids, 105; on birds as enemies of butterflies, 107; on feeding experiments with S. African birds, 117; with monkeys, 121; on birds attacking Pierids, 150 _Mechanitis egaensis_, XV. 9; _M. elisa_, XV. 6; _M. lysimnia_, 151, XV. 8; _M. methona_, XV. 10; _M. saturata_, as model for _Dismorphia praxinoe_, 57, 62; as member of mimicry ring, 134, XV. 7 Melanic sports in moths, 101 _Melinaea_, 135 _Melinda formosa_, App. II _Melittophagus swinhoei_, 110 _Merops viridis_, 111 Merrifield, on seasonal dimorphism, 130 _Methona confusa_, XII. 1, XIV. 1 Migratory birds, suggested influence on mimicry of, 53 _Mimacraea_, 35 Mimetic resemblance, as induced through gradual slight changes, 64 Mimic, occupying same station as model, 51; occupying station apart from model, 53; scarcer than model, 56; pattern of in relation to allies, 57 Mimicry, Wallace's conditions of, 50; Batesian, 9; Müllerian, 14 Mimicry rings, 65; in S. American butterflies, 134; and natural selection, 136 Mimicry theory, difficulties of, 139 Monkeys, as enemies of butterflies, 121 Moths, mimicry in, 27, 36 Moulton, on S. American mimicry rings, 134 Müller, 14, 72 Müllerian mimicry, 53, 57, 66; difficulties of, 72 Mutation, see Sports _Mylothris_, 36

Natural selection and mimicry, 10-12, 61, 92, 152 Neal, on monkeys as enemies of butterflies, 123 _Nepheronia (= Pareronia) hippia_, 116 _Neptis imitans_, 24; _N. nemetes_, 54; _N. kamarupa_, 121 North American butterflies, mimicry among, 45 Norton, on rapidity of changes in mixed populations through natural selection, 94, App. I

Oriental butterflies, mimicry among, 23 Overlapping in patterns of different groups of butterflies, 144

_Papilio aristolochiae_, as model for female of _P. polytes_, 13, 26, 52, 77; range of, 79; likeness to _P. polytes_, 80; characteristics of, 81; flight of, 82; eaten by lizards, 108; rejected by certain birds, 115, 116; disliked by Tree-shrew, 121, V. 5, 5A; _P. agestor_, 24, 51, II. 3; _P. asterius_, 46; _P. bachus_, App. II; _P. bootes_, 27, III. 6; _P. brasidas_, 29, VI. 4; _P. chamissonia_, 44, XIII. 2; _P. clytia_, 23, 25, 55, I. 7, 8; _P. coon_, 26, 89; _P. cynorta_, 35, 36, VII. 10; _P. dardanus_, investigated by Trimen, 14; mimicry in, 30; breeding experiments with, 90; polymorphic forms of in relation to models, 149 note; var. _humbloti_, 32; var. _meriones_, 32; [F] _cenea_, 31, VIII. 4; [F] _dionysus_, 31, 33; [F] _hippocoon_, 31, VIII. 3; [F] _niavioides_, 32, 33; [F] _planemoides_, 31; [F] _ruspina_, 33; [F] _trimeni_, 31, 32, 33; {187} [F] _trophonius_, 31, 122, VIII. 2; _P. delesserti_, App. II; _P. demoleus_, 111, 121; _P. echerioides_, App. II; _P. erithonius_, 110; _P. euterpinus_, 42, 43; _P. glaucus_, 45; var. _turnus_, 46; _P. hahneli_, 39; _P. hector_, model for female of _P. polytes_, 13, 52, 78; range of, 79; characteristics of, 81; flight of, 82; eaten by lizards, 108; eaten by birds, 110, 117, V. 6, 6A; _P. hippason_, App. II; _P. laglaizei_, 27, 124, III. 2; _P. leonidas_, 29, VI. 3; _P. lysithous_, polymorphism in, 44; [F] _lysithous_, XIII. 4; [F] _rurik_, XIII. 5; [F] _pomponius_, XIII. 6; _P. macareus_, 23, 111; _P. memnon_, 26, 89; _P. mendax_, 24, 51, II. 9; _P. nephalion_, 44, XIII. 1; _P. osyris_, XI. 8, 9; _P. paradoxus_, 25; _P. pausanias_, 43, XI. 2; _P. perrhebus_, 44, XIII. 3; _P. philenor_, as model, 45; taken by lizard, 108, XVI. 1; _P. polytes_, polymorphism in females of, 13, 75; mimic of Pharmacophagus Papilio, 26; habits of, 52, 124; often more abundant than models, 56; description of, 76-78; relative abundance of models in Ceylon, 79; breeding experiments with, 84; equilibrium among females of in Ceylon, 96; relative abundance of three forms of female of in Ceylon, 97; historical notes on abundance of forms of female in Ceylon, 98; origin of forms of female in, 125, 141; relation of polymorphic forms to models in, 149 note; preyed on by Wood-Swallow, 112; feeding experiments with, 116, V. 1-4, 1A-4A; _P. polyxenus_, 27; _P. rex_, App. II; _P. ridleyanus_, 34, 36, VI. 6; _P. sarpedon_, 110; _P. troilus_, 45, XVI. 2; _P. xenocles_, 23, 111, I. 4; _P. zagreus_, 43, X. 8 Papilionidae, as mimics of Oriental models, 23-25; of African models, 29, 30, 35; of S. American models, 43; of N. American models, 45 Parallel patterns, in different butterfly groups, 144 _Pareronia_, 145, 149; _P. ceylanica_, 23, 59, 116 note, I. 1, 2 Pattern and physiological properties, possible connection between, 137 Patterns, overlapping series of in different groups of butterflies, 145 _Pedaliodes_, 135 _Pereute charops_, 42, XI. 6, 7 _Pericopis_, 39 _Perrhybris_, as mimics of Ithomiines, coloration of male in _P. malenka_, 62; as members of mimicry rings, 134, 135; _P. demophile_, 151; _P. lorena_, 151; _P. malenka_, X. 4, 5, 6 Pharmacophagus Swallow-tails, characteristics of, 22, App. II; as models for Oriental butterflies, 25; absence of in Africa, 35; as models in S. America, 43; as models in N. America, 45 _Phrissura_, 36 _Phyciodes_, 38, 54 Physiological properties, possible connection of with pattern, 137 Pieridae, as models for Oriental butterflies, 28; mimicry in African, 36; mimicry in S. American, 43; frequency of bird attacks on, 150 _Planema epaea_, 35, VII. 5; _P. macarista_, sexual difference in, 34, VII. 1, 2; mimicked by _Elymnias phegea_, 35; by _Pseudacraea eurytus_, 126; _P. poggei_, as model for _planemoides_ female of _Papilio dardanus_, 31; _P. paragea_, 126, VII. 4; _P. tellus_, 126, VII. 3 {188} Poison-eaters, see Pharmacophagus Swallow-tails Polymorphism, in females of mimicking species, 13; among females of _P. dardanus_, 30; among females of _P. polytes_, 75 Population, conditions of equilibrium in mixed, 93 Poulton, 17; on N. American mimetic butterflies, 45; on the "Transparency group," 41; on mimicry through agency of migratory birds, 53; on _Hypolimnas misippus_, 66 note; on the relation between mimetic forms of _P. polytes_, 90; on predaceous insects, 105; on relative proportion of different forms of _Pseudacraea eurytus_, 127; on local variation in _D. chrysippus_, 132 _Precis_, 111, 122, 131; _P. octavia_, seasonal dimorphism in, 131, VI. 11, 12 _Prioneris_, 110; _P. sita_, 28 Pritchett, feeding experiments with lizards, 108 Protective resemblance, 8 _Protogonius_, as mimics of Ithomiines, 38; as members of mimicry rings, 134, 135, 138; _P. tithoreides_, X. 9 _Pseudacraea_, 59, 144; _P. boisduvali_, 34, VI. 5; _P. eurytus_, relative proportion of different forms in, 127; polymorphism of in relation to model, 149 note; var. _hobleyi_ as mimic of _Planema macarista_, 35, 127, VII. 6, 7; var. _terra_, as mimic of _Planema tellus_, 126, VII. 8; var. _obscura_ as mimic of _Planema paragea_, 126

Ray, on adaptation, 4, 6 Rodents, bearing on mimicry of recent genetic work with, 147

Satyrinae, transparency in S. American, 42 _Sceleporus floridanus_, 108 Schaus, on birds as enemies of butterflies, 112 Seasonal dimorphism, 130 Seitz, 44, 52, 58 Shelford, 56 note S. American butterflies, mimicry among, 38 Sports, as foundation of mimetic resemblances, 70, 91, 143 Sweet-peas, experiments on, 91 Swynnerton, on contents of stomachs of birds, 114

_Telipna sanguinea_, 36 _Terias brigitta_, 35; _T. hecabe_, 110 _Thyridia_, 40, XIV. 3 _Tithorea pseudonyma_, X. 10 "Transparency group," in S. America, 39 Trimen, on mimicry in African butterflies, 13 _Tupaia ferruginea_, 121

Variation, difficulty of initial, 63

Wade, on relative abundance of the three forms of _P. polytes_ in Ceylon, 99 Wallace, on mimicry in Oriental butterflies, 12; on the conditions of mimicry, 50; on the females of _P. polytes_, 76; on initial variation, 64 Warning colours, 10, 11 Weismann, 1, 2

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NOTES

[1] _The Wisdom of God manifested in the Works of the Creation_, London, 1691.

[2] Ray gives the case of an elephant "that was observed always when he slept to keep his trunk so close to the ground, that nothing but Air could get in between them," and explains it as an adaptation in habit to prevent the mice from crawling into its lungs--"a strange sagacity and Providence in this Animal, or else an admirable instinct."

[3] _Trans. Linn. Soc._ vol. 23, 1862.

[4] _Trans. Linn. Soc._ vol. 25, 1866.

[5] _Trans. Linn. Soc._ vol. 26, 1870.

[6] In attributing this quality to the butterflies in question I am merely stating what is held by the supporters of the mimicry theory. I know of scarcely any evidence either for or against the supposition.

[7] It is assumed that the intelligence of the birds is such that they can learn a pattern after a single disagreeable experience of it.

[8] _Untersuchungen über die Mimikry_, 1893.

[9] The African mimetic butterflies have been recently monographed by Eltringham in a large and beautifully illustrated work--_African Mimetic Butterflies_, Oxford, 1910.

[10] Omitting the Hesperidae which hardly enter into questions of mimicry.

[11] The classification adopted is that used by Dr Sharp in the "Cambridge Natural History," _Insects_, vol. 2, 1901.

[12] Cf. Shelford, _Proc. Zool. Soc._ 1902.

[13] _African Mimetic Butterflies_, Oxford, 1910.

[14] These African species of _Hypolimnas_ are frequently referred to the genus _Euralia_.

[15] Corresponding to the _dorippus_ form of _D. chrysippus_ (cf. Pl. IX) there is a rare form of _trophonius_ known as _dorippoides_.

[16] These two forms are figured on Plate 10 of Eltringham's _African Mimetic Butterflies_.

[17] _Acraea violae_, the only representative of the group in S. India and Ceylon, is nevertheless a very abundant insect. It cannot, however, be said that it is definitely mimicked by any other species in this region.

[18] Coloured figures of these and of the other African species referred to may be found in Eltringham's work on _African Mimetic Butterflies_.

[19] These descriptions are taken from preserved specimens which I owe for the most part to the kindness of Dr Jordan. I have not had an opportunity of examining fresh ones.

[20] This is more marked in _Castnia_ than in _Anthomysa_. It appears to be a peculiarity of many members of the genus _Castnia_ that the scales do not lie so tight as generally in moths. Owing to this, some of the large whole-coloured species have a somewhat fluffy look.

[21] Cf. Poulton, _Essays on Evolution_, 1908, pp. 264-6.

[22] Cf. Poulton, _Darwin and the 'Origin,'_ 1909, pp. 177-186.

[23] The N. American members of this genus are often referred to as _Basilarchia_.

[24] _Darwinism_, 1890 (1st Edition 1889), p. 264.

[25] _Macrolepidoptera of the World._ _Fauna Americana_, p. 98.

[26] _Essays on Evolution_, 1908, p. 381.

[27] These "unpalatable" butterflies are sometimes extensively preyed upon by insectivorous birds, when they fall an easier prey owing to their slowness (cf. p. 112).

[28] See Shelford, PROC. ZOOL. SOC. 1902, p. 260. A coloured figure of both species is given in the paper.

[29] _Macrolepidoptera of the World._ _Fauna Americana_, pp. 98-104, Plates 28-30.

[30] "In what way our _Leptalis_ (= _Dismorphia_) originally acquired the general form and colour of Ithomiae I must leave undiscovered." _Trans. Linn. Soc._ vol. 23, 1862, p. 513.

[31] _Darwinism_, 1890, pp. 242-244.

[32] _Origin of Species_, 6th Edition, 1891, p. 354.

[33] _H. misippus_ was at one time regarded as a clear case of Batesian mimicry. But in view of its plentifulness, of the fact that it may be abundant outside the area inhabited by its model, and of the ease with which it can establish itself in parts remote from its original habitat, _e.g._ S. America, it has come to be regarded by certain supporters of the mimicry theory as a Müllerian mimic. Cf. Poulton, _Essays on Evolution_, 1908, pp. 215-217.

[34] An English translation of Müller's paper is given by Meldola, _Proc. Ent. Soc._, 1879, p. xx.

[35] _Trans. Ent. Soc. Lond._, 1908, p. 93.

[36] Provided of course that the type form remains in the majority. If the variation occurred simultaneously in more than 50% of A the advantage would naturally be with the variation.

[37] It is possible to imagine an exceptional case though most unlikely that it would occur. Suppose for example that there were a number of distasteful species, say 20, all of different patterns, and suppose that in all of them a particular variation occurred simultaneously; then if the total shewing that variation from among the 20 species were greater than the number of any one of the species, all of the 20 species would come to take on the form of the new variation. In this way it is imaginable that the new pattern would gradually engulf all the old ones.

[38] _Trans. Linn. Soc._ vol. 24, 1866.

[39] These darker ribs are also present in the male and M female but are obscured owing to the generally deeper colour.

[40] See Appendix II, p. 158.

[41] _Spolia Zeylanica_, 1910.

[42] We shall take it for the present that, from the point of view of mimicry, birds are the main enemies of butterflies (cf. Chap. IX).

[43] See later, p. 119.

[44] The specimens figured on Pl. V were dried in papers when taken. The body is consequently much compressed and the characteristic scarlet of _P. hector_ and _P. aristolochiae_ is largely hidden.

[45] _Philosophical Transactions of the Royal Society_, vol. 204, 1913.

[46] _Tijdschr. voor Entomologie_, vol. 53, 1909. A more accessible account is given by de Meijere, _Zeit. f. indukt. Abstamm. u. Vererbungslehre_, vol. 3, 1910.

[47] For further information see Poulton, _Trans. Ent. Soc. Lond._ 1909, and various notes in _Proc. Ent. Soc. Lond._ subsequent to this date.

[48] _Science_, July, 1908.

[49] If for example there were 5000 dominants and 4000 recessives, and if only half of the population survives to mate, then we should be left with 2500 dominants and 2000 recessives as parents of the next generation. But if there were also a 10% selective disadvantage working against the recessives, their numbers would be further reduced from 2000 to 1800 and the proportion of dominants to recessives would be changed from 5:4 to 25:18.

[50] As these larvae were for the most part found simply over a considerable time it follows that they are the offspring of different females and represent the relative proportions of the three forms in the general population.

[51] _The Lepidoptera of Ceylon_, 1880.

[52] From the experience of breeders it would appear that the melanic form is somewhat hardier, at any rate in captivity.

[53] Intermediates may also occur in some strains (cf. Bowater, _Journal of Genetics_, vol. 3, no. 4, 1914).

[54] An interesting case of a similar nature has recently been published by Hasebroek (_Die Umschau_, 1913, p. 1020). A melanic form of the moth, _Cymatophora or_, suddenly appeared near Hamburg in 1904. This new form, to which the name _albingensis_ was given, rapidly became the predominant one. In 1911-1912 over 90% of the moths reared from caterpillars taken in the open were of the _albingensis_ form; nor were any intermediates found between it and the typical form. Some experiments were also made which shew that the _albingensis_ form behaves as a dominant to the original type form.

[55] _Trans. Ent. Soc. Lond. 1907._

[56] _Trans. Ent. Soc. Lond. 1902._

[57] _Proc. Zool. Soc._ 1911.

[58] _Journ. Roy. Asiat. Soc. Bengal_, vol. 65, 1897.

[59] _Spolia Zeylanica_, 1910.

[60] _Biological Bulletin_, vol. 5, 1903.

[61] _Trans. Ent. Soc. Lond._ 1909.

[62] _Trans. Ent. Soc. Lond._ 1902.

[63] _Trans. Ent. Soc. Lond._ 1911.

[64] _Proc. Zool. Soc. 1913._

[65] _A Naturalist in Nicaragua_, 1874, p. 316.

[66] _I^{er} Congr. Internat, d'Entomologie_, Bruxelles, 1911.

[67] _Ibis_, 1911.

[68] _Ibis_, 1912.

[69] _The Condor_, vol. 13, 1911, pp. 195-208.

[70] _Journ. Asiat. Soc. Bengal_, vol. 64, 1895, and vol. 66, 1897.

[71] Nevertheless a Liothrix is recorded as eating _Danais plexippus_ and a _Euploea_ even though two male specimens of the palatable _Elymnias undularis_ were in the cage.

[72] A form closely resembling _P. ceylonica_ figured on Pl. I, fig. 1.

[73] _Proc. Zool. Soc. Lond. 1911._

[74] _Proc. Acad. Nat. Sci. Philadelphia_, 1912.

[75] C. Hess, _Handbuch der vergleichenden Physiologie_ (herausgegeben von H. Winterstein), Bd. 4, 1912, p. 563.

[76] _Journ. As. Soc. Bengal_, vol. 66^2, 1898.

[77] _Trans. Ent. Soc. Lond._ 1902.

[78] Marshall, _loc. cit._ p. 379.

[79] In this connection may be quoted a letter from Capt. N. V. Neal near Lagos to Mr W. A. Lamborn which was recently published in the _Proceedings of the Entomological Society_.

"You have asked me about monkeys eating butterflies. This is very common, as every native will tell you. I have seen it myself. The monkey runs along a path, sees some butterflies fluttering round some filth, goes very quietly, and seizes one by the wings, puts the solid part (body) into his mouth, then pulls the wings off. The poor butterfly goes down like any oyster.... The dog-faced baboon and the large brown monkey with a very long tail, which seems to be the most common species in this colony, are great butterfly-eaters. The little spider-monkey also considers a butterfly a treat, and prefers one to a spider."

[80] Cf. E. B. Poulton in _Bedrock_ for Oct. 1913, p. 301.

[81] _Trans. Ent. Soc. London, 1914._

[82] In the female _hobleyi_, with rare exceptions, the orange of the male is replaced by white, and it has received the name _tirikensis_. The female of _P. macarista_ also shews white in place of the orange of the male.

[83] Cf. Poulton, E. B., _I^{er} Congr. Internat. d'Entomol._, Bruxelles 1911. This proportion is founded on several hundreds caught at random. Observers are agreed that _Pseudacraea_ is both a warier insect and a stronger flyer than the various Planemas which it resembles.

[84] _I^{er} Congr. Internat. d'Entom._, Bruxelles 1911.

[85] Cf. Poulton, _Bedrock_, Oct. 1913, p. 300.

[86] The size of the white spot may shew much variation in specimens from the same region. I have seen African specimens in which it is large, while in the Ceylon specimen figured on Plate IV it is as small as in the typical African specimen shewn on Plate VIII.

[87] See Moulton, J. C., _Trans. Ent. Soc. London_, 1909.

[88] In this connection it is of interest that a recent observer with considerable breeding experience finds that the dark _doubledayaria_ variety of the Peppered Moth is more hardy than the typical form (cf. p. 101). The swift success of the dark variety led some to regard it as better protected against bird enemies. It is, however, not unlikely that the deeper pigmentation is associated with some physiological difference which makes for greater hardiness. See Bowater, _Journal of Genetics_, vol. 3, 1914.

[89] As examples may be mentioned _P. polytes_, _Hypolimnas misippus_, _H. dubius_, and _Pseudacraea hobleyi_. With the exception of the _planemoides_ form it is true also for _P. dardanus_, the most polymorphic of them all.

[90] _Trans. Ent. Soc. Lond. 1909._

[91] Cf. F. G. Hopkins, _Phil. Trans. Roy. Soc. 1895._

[92] Coloured representations of these two species will be found on Pl. 20 of Seitz, _Macrolepidoptera of the World, Fauna Americana_.

[93] _Untersuchungen über die Mimikry_, 1893.