wunder · Library

Appendix Ii

Mimicry in Butterflies · Reginald Crundall Punnett — chapter 15 of 15 · ~6,438 words · public domain

The genus Papilio is a large and heterogeneous collection. It was pointed out by Haase that it falls into three distinct sections, of which one--the Pharmacophagus section--provides those members which serve as models in mimicry; while in the other two sections are found mimics, either of Pharmacophagus Swallow-tails, or of models belonging to other groups. Though Haase's terms have not yet come into general use with systematists, there is little doubt that the genus Papilio as it now stands must eventually be broken up on these lines. To say that one species of Papilio mimics another is therefore somewhat misleading; for the differences between the Pharmacophagus group and the other two are such as to constitute at any rate generic distinction in other groups. For convenience of reference a table has been added in which the various Papilios mentioned in the text have been assigned to their appropriate sections, and referred to their respective models. {158}

Pharmacophagus Papilio Cosmodesmus

(POISON-EATERS) (FLUTED SWALLOW-TAILS) (KITE SWALLOW-TAILS)

Antennae without Antennae without Antennae scaled scales. scales. on upper side.

Outer ventral row Outer ventral row As in Papilio. of spines of tarsi of spines of tarsi not separated separated from from the dorsal the dorsal spines spines by a spineless by a spineless longitudinal longitudinal depression. depression.

Larva covered with Larva either smooth Larva with third short hairs--with or with hard thoracic segment fleshy tubercles spiny tubercles. enlarged but no Third and fourth (known only in spines. thoracic segments a few species). enlarged.

Pupa with row of Pupa wrinkled--generally Pupa short with well-marked with long four-sided humps on each short dorsal thoracic horn. side of abdomen. horn. Humps if present very short.

Larva feeds on Larva does not feed As in Papilio. Aristolochia. on Aristolochia.

Abdominal margin Abdominal margin of hind wing of hind wing curved downwards bent over in , forming and with scent a kind of groove. organ in fold No scent organ. so formed.

{159}

LIST OF PAPILIONINE MIMICS

AMERICAN hahneli(mimic)| | |Methona confusa |zagreus | |} various Heliconinae |bachus | |} and Ithomiinae |euterpinus | |Heliconius melpomene | |pausanias | " sulphurea various species |hippason, etc. |lysithous | | | etc. | philenor |troilus | | |turnus | | |asterius | |

* * * * *

{160}

PLATE I

ORIENTAL BUTTERFLIES

1. Pareronia ceylonica } (Pieridae) 2. " " } 3. Danais septentrionis (Danainae) 4. Papilio xenocles (Papilionidae) 5. Hypolimnas bolina } (Nymphalinae) 6. " " } 7. Papilio clytia var. dissimilis } (Papilionidae) 8. " " var. lankeswara } 9. Elymnias singhala (Satyrinae) 10. Euploea core (Danainae)

* * * * *

PLATE II

ORIENTAL BUTTERFLIES

1. Delias eucharis (Pieridae) 2. Caduga tytia (Danainae) 3. Papilio agestor (Papilionidae) 4. Euploea mulciber } (Danainae) 5. " " } 6. Elymnias malelas } (Satyrinae) 7. " " } 8. Euploea rhadamanthus (Danainae) 9. Papilio mendax (Papilionidae)

* * * * *

PLATE III

ORIENTAL MOTHS AND BUTTERFLIES

The three upper figures are those of moths, and the three lower ones are those of butterflies.

1. Alcidis agathyrsus (New Guinea) 2. Papilio laglaizei " "

The moth is here supposed to serve as a model for the far rarer Papilio.

3. Cyclosia hestinioides 4. Ideopsis daos

The butterfly is very common and must be regarded as the model, the rarer moth as the mimic.

5. Epicopeia polydora (Assam) 6. Papilio bootes "

Both of these species are to be regarded as mimics of the abundant Pharmacophagus Papilio, P. polyxenus, which is very like P. bootes in appearance.

* * * * *

PLATE IV

ORIENTAL BUTTERFLIES

1. Danais chrysippus } (Danainae) 2. " plexippus } 3. Argynnis hyperbius } (Nymphalinae) 4. " " } 5. Elymnias undularis } (Satyrinae) 6. " " } 7. Hypolimnas misippus } (Nymphalinae) 8. " " }

The two Danaids together with the females of the other three species form a "mimicry ring." For explanation see text, pp. 65-69.

* * * * *

PLATE V

ORIENTAL BUTTERFLIES

1. Papilio polytes 2. " " , var. cyrus (M form) 3. " " , var. polytes (A form) 4. " " , var. romulus (H form) 5. " aristolochiae 6. " hector

The specimens figured on this plate were taken in Ceylon where they are all plentiful.

Figures 1a--6a represent the under surfaces of the hind wings belonging to specimens 1--6.

* * * * *

PLATE VI

AFRICAN BUTTERFLIES

(except A. levana, Figs. 8--10, which is European)

1. Danais petiverana (Danainae) 2. Papilio leonidas (Papilionidae) 3. Amauris hyalites (Danainae) 4. Papilio leonidas var. brasidas (Papilionidae) 5. Pseudacraea boisduvali (Nymphalinae) 6. Papilio ridleyanus (Papilionidae) 7. Acraea egina (Acraeinae) 8. Araschnia levana var. porima 9. " " var. prorsa 10. " " 11. Precis octavia var. sesamus 12. " " var. natalensis

* * * * *

PLATE VII

TROPICAL AFRICAN BUTTERFLIES

1. Planema macarista (Acraeinae) 2. " " " 3. " tellus " 4. " paragea " 5. " epaea " 6. Pseudacraea hobleyi (Nymphalinae) 7. " " " 8. " terra " 9. Elymnias phegea (Satyrinae) 10. Papilio cynorta (Papilionidae)

(NOTE. Pseudacraea hobleyi and P. terra (Figs. 6--8) were at one time regarded as separate species. More recently they have been shewn to be forms of the polymorphic species, Pseudacraea eurytus.)

* * * * *

PLATE VIII

AFRICAN BUTTERFLIES

1. Papilio dardanus 2. " " , var. trophonius 3. " " , var. hippocoon 4. " " , var. cenea 5. Danais chrysippus (Danainae) 6. Amauris niavius " 7. " echeria " 8. Hypolimnas dubius var. mima (Nymphalinae) 9. " " var. wahlbergi "

* * * * *

Danais chrysippus a. Typical form b. Alcippus form c. Dorippus form

Acraea encedon d. Typical form e. Alcippina form f. Daira form

Hypolimnas misippus g. Typical form h. Alcippoides form i. Inaria form

(After Aurivillius)

* * * * *

PLATE X

SOUTH AMERICAN BUTTERFLIES

1. Dismorphia cretacea (Pieridae) 2. " praxinoe " 3. " " " 4. Perrhybris malenka " 5. " " (under surface) " 6. " " " 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.)

* * * * *

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 (Pieridae) 7. " " " 8. Papilio osyris (Papilionidae) 9. " " " 10. Archonias critias (Pieridae)

* * * * *

PLATE XII

SOUTH AMERICAN BUTTERFLIES

Methona confusa (Ithomiinae) Dismorphia orise (Pierinae) Ituna phenarete (Danainae) Castnia sp. (Heterocera)

* * * * *

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.)

* * * * *

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.

* * * * *

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 (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; cenea, 31, VIII. 4; dionysus, 31, 33; hippocoon, 31, VIII. 3; niavioides, 32, 33; planemoides, 31; ruspina, 33; trimeni, 31, 32, 33; {187} 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; lysithous, XIII. 4; rurik, XIII. 5; 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

CAMBRIDGE: PRINTED BY JOHN CLAY, M.A. AT THE UNIVERSITY PRESS

* * * * *

SELECTION FROM THE GENERAL CATALOGUE OF BOOKS PUBLISHED BY THE CAMBRIDGE UNIVERSITY PRESS

JOURNAL OF GENETICS. A periodical for the publication of records of original research in Heredity, Variation, and allied subjects. Edited by W. BATESON, M.A., F.R.S., and R. C. PUNNETT, M.A., F.R.S.

The Journal of Genetics is issued in parts as material accumulates, and a volume appearing, so far as possible, annually, consists of four such parts. The Subscription price for a volume is 30s net (post free) payable in advance; separate parts 10s net each.

Vols. I, II and III (1910-14) are now ready. Price in four parts, paper covers, 30s net each; bound in Buckram, 34s 6d net each.

MENDEL'S PRINCIPLES OF HEREDITY. By W. BATESON, M.A., F.R.S., V.M.H. Third impression with additions. Royal 8vo. With 3 portraits, 6 coloured plates and 38 figures. 12s net.

"A new impression cannot fail to be welcomed by those who already have it as well as those who have it not.... Mendel's Principles of Heredity is already a classic. It marks a position of stability towards which previous work is now seen to have logically converged, and from which new and active research is to-day no less logically diverging. The various waves of biological thought are constantly intersecting, mingling, and passing on with altered rhythm, but it rarely happens that so many meet together at a nodal point as during the last decade.... As an analysis of that point, as a picture of how it has come into being, and as a foreshadowing of happenings in the near future, Mendel's Principles stands alone, and it is good to know that the generations of students now growing up cannot be cut off from the possession of a book so full of inspiration."--Gardeners' Chronicle

THE METHODS AND SCOPE OF GENETICS. An Inaugural Lecture delivered 23 October 1908 by W. BATESON, M.A., F.R.S. Crown 8vo. 1s 6d net.

ZOOLOGY. AN ELEMENTARY TEXT-BOOK. By A. E. SHIPLEY, Sc.D., F.R.S., and E. W. MACBRIDE, M.A., D.Sc. Third edition, revised and enlarged. Demy 8vo. With numerous illustrations. 12s 6d net.

EXPERIMENTAL ZOOLOGY. By HANS PRZIBRAM, Ph.D. Part I. Embryogeny, an account of the laws governing the development of the animal egg as ascertained by experiment. Royal 8vo. With 16 plates. 7s 6d net.

THE NATURAL HISTORY OF SOME COMMON ANIMALS. By OSWALD H. LATTER, M.A. Crown 8vo. With 54 illustrations. 5s net.

THE HOUSE-FLY (Musca Domestica Linn.): its structure, habits, development, relation to disease and control. By C. G. HEWITT, D.Sc., F.R.S.C. Demy 8vo. With 104 illustrations and a map. 15s net.

THE DETERMINATION OF SEX. By L. DONCASTER, Sc.D. Demy 8vo. With frontispiece in colour and 22 plates. 7s 6d net.

THE EVOLUTION OF SEX IN PLANTS. By JOHN MERLE COULTER, Head of the Department of Botany in the University of Chicago. Small 12mo, 4s net. (Published by the University of Chicago Press and sold in Great Britain and the British Empire (except Canada) by the Cambridge University Press.)

* * * * *

CAMBRIDGE MANUALS OF SCIENCE AND LITERATURE

Editors: P. GILES, Litt.D., and A. C. SEWARD, M.A., F.R.S.

Cloth, 1s net each; leather, 2s 6d net each.

"A very valuable series of books which combine in a very happy way a popular presentation of scientific truth along with the accuracy of treatment which in such subjects is essential.... In their general appearance, and in the quality of their binding, print and paper, these volumes are perhaps the most satisfactory of all those which offer to the inquiring layman the hardly earned products of technical and specialist research."--The Spectator

THE LIFE-STORY OF INSECTS. By Prof. G. H. CARPENTER. With 24 illustrations.

BEES AND WASPS. By O. H. LATTER, M.A., F.E.S. With 21 illustrations.

HOUSE-FLIES AND HOW THEY SPREAD DISEASE. By C. G. HEWITT, D.Sc. With frontispiece and 19 figures.

THE FLEA. By H. RUSSELL. With 9 illustrations.

SPIDERS. By C. WARBURTON, M.A. With 13 figures.

EARTHWORMS AND THEIR ALLIES. By FRANK E. BEDDARD, M.A. (Oxon.), F.R.S., F.R.S.E. With 13 figures.

PLANT-ANIMALS. By F. W. KEEBLE, Sc.D. With 23 figures.

PRIMITIVE ANIMALS. By G. SMITH, M.A. With 26 figures.

LIFE IN THE SEA. By JAMES JOHNSTONE, B.Sc. With frontispiece, 4 figures and 5 tailpieces.

THE INDIVIDUAL IN THE ANIMAL KINGDOM. By JULIAN S. HUXLEY, B.A. With frontispiece and 16 figures.

THE WANDERINGS OF ANIMALS. By HANS GADOW, F.R.S. With 17 maps.

THE MIGRATION OF BIRDS. By T. A. COWARD. With 4 maps.

HEREDITY IN THE LIGHT OF RECENT RESEARCH. By L. DONCASTER, Sc.D. With 12 figures.

THE COMING OF EVOLUTION. By JOHN W. JUDD, C.B., LL.D., F.R.S. With 4 plates.

A list of the eighty-six volumes now ready will be sent on application.

CAMBRIDGE UNIVERSITY PRESS C. F. Clay, Manager: Fetter Lane, London

* * * * *

NOTES

The Wisdom of God manifested in the Works of the Creation, London, 1691.

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."

Trans. Linn. Soc. vol. 23, 1862.

Trans. Linn. Soc. vol. 25, 1866.

Trans. Linn. Soc. vol. 26, 1870.

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.

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

Untersuchungen über die Mimikry, 1893.

The African mimetic butterflies have been recently monographed by Eltringham in a large and beautifully illustrated work--African Mimetic Butterflies, Oxford, 1910.

Omitting the Hesperidae which hardly enter into questions of mimicry.

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

Cf. Shelford, Proc. Zool. Soc. 1902.

African Mimetic Butterflies, Oxford, 1910.

These African species of Hypolimnas are frequently referred to the genus Euralia.

Corresponding to the dorippus form of D. chrysippus (cf. Pl. IX) there is a rare form of trophonius known as dorippoides.

These two forms are figured on Plate 10 of Eltringham's African Mimetic Butterflies.

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.

Coloured figures of these and of the other African species referred to may be found in Eltringham's work on African Mimetic Butterflies.

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.

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.

Cf. Poulton, Essays on Evolution, 1908, pp. 264-6.

Cf. Poulton, Darwin and the 'Origin,' 1909, pp. 177-186.

The N. American members of this genus are often referred to as Basilarchia.

Darwinism, 1890 (1st Edition 1889), p. 264.

Macrolepidoptera of the World. Fauna Americana, p. 98.

Essays on Evolution, 1908, p. 381.

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

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

Macrolepidoptera of the World. Fauna Americana, pp. 98-104, Plates 28-30.

"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.

Darwinism, 1890, pp. 242-244.

Origin of Species, 6th Edition, 1891, p. 354.

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.

An English translation of Müller's paper is given by Meldola, Proc. Ent. Soc., 1879, p. xx.

Trans. Ent. Soc. Lond., 1908, p. 93.

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.

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.

Trans. Linn. Soc. vol. 24, 1866.

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

See Appendix II, p. 158.

Spolia Zeylanica, 1910.

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).

See later, p. 119.

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.

Philosophical Transactions of the Royal Society, vol. 204, 1913.

Tijdschr. voor Entomologie, vol. 53, 1909. A more accessible account is given by de Meijere, Zeit. f. indukt. Abstamm. u. Vererbungslehre, vol. 3, 1910.

For further information see Poulton, Trans. Ent. Soc. Lond. 1909, and various notes in Proc. Ent. Soc. Lond. subsequent to this date.

Science, July, 1908.

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.

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.

The Lepidoptera of Ceylon, 1880.

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

Intermediates may also occur in some strains (cf. Bowater, Journal of Genetics, vol. 3, no. 4, 1914).

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.

Trans. Ent. Soc. Lond. 1907.

Trans. Ent. Soc. Lond. 1902.

Proc. Zool. Soc. 1911.

Journ. Roy. Asiat. Soc. Bengal, vol. 65, 1897.

Spolia Zeylanica, 1910.

Biological Bulletin, vol. 5, 1903.

Trans. Ent. Soc. Lond. 1909.

Trans. Ent. Soc. Lond. 1902.

Trans. Ent. Soc. Lond. 1911.

Proc. Zool. Soc. 1913.

A Naturalist in Nicaragua, 1874, p. 316.

I^{er} Congr. Internat, d'Entomologie, Bruxelles, 1911.

Ibis, 1911.

Ibis, 1912.

The Condor, vol. 13, 1911, pp. 195-208.

Journ. Asiat. Soc. Bengal, vol. 64, 1895, and vol. 66, 1897.

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.

A form closely resembling P. ceylonica figured on Pl. I, fig. 1.

Proc. Zool. Soc. Lond. 1911.

Proc. Acad. Nat. Sci. Philadelphia, 1912.

C. Hess, Handbuch der vergleichenden Physiologie (herausgegeben von H. Winterstein), Bd. 4, 1912, p. 563.

Journ. As. Soc. Bengal, vol. 66^2, 1898.

Trans. Ent. Soc. Lond. 1902.

Marshall, loc. cit. p. 379.

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."

Cf. E. B. Poulton in Bedrock for Oct. 1913, p. 301.

Trans. Ent. Soc. London, 1914.

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.

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.

I^{er} Congr. Internat. d'Entom., Bruxelles 1911.

Cf. Poulton, Bedrock, Oct. 1913, p. 300.

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.

See Moulton, J. C., Trans. Ent. Soc. London, 1909.

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.

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.

Trans. Ent. Soc. Lond. 1909.

Cf. F. G. Hopkins, Phil. Trans. Roy. Soc. 1895.

Coloured representations of these two species will be found on Pl. 20 of Seitz, Macrolepidoptera of the World, Fauna Americana.

Untersuchungen über die Mimikry, 1893.

← Previous chapterAll chapters

Mimicry in Butterflies · The Wunder Library — complete classics, free to read, with narration.