Scientific Contributions of the New York Zoological Society
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9 A Comparison of Eggs, Larvae and Pupae in Fourteen Species 1 2 of Heliconiine Butterflies from Trinidad, W. I . * William Beebe, Jocelyn Crane & Henry Fleming Department of Tropical Research, New York Zoological Society Zoological Park, New York 60, N. Y. (Plates I-XVI; Text-figures 1-14) [This paper is one of a series emanating from the 7. Heliconius aliphera 138 tropical Field Station of the New York Zoological 8. Heliconius melpomene ... 139 Society at Simla, Arima Valley, Trinidad, West 9. Heliconius numata 140 Indies. The Station was founded in 1950 by the 10. Heliconius erato 142 Zoological Society’s Department of Tropical Re- 11. Heliconius ricini 143 search, under the direction of Dr. William Beebe. 12. Heliconius sara 143 It comprises 200 acres in the middle of the North- 13. Heliconius wallacei 144 ern Range, which includes large stretches of undis- 14. Heliconius doris 144 turbed government forest reserves. The laboratory V. Pupae 145 of the Station is intended for research in tropical A. Characteristics of the Subfamily . 145 ecology and in animal behavior. The altitude of to 146 the research area is 500 to 1,800 feet, with an an- B. Key Pupae nual rainfall of more than 100 inches. C. Descriptions of the Species 147 [For further ecological details of meteorology 1. Agraulis vanillae 147 and biotic zones see “Introduction to the Ecology 2. Dione juno 147 of the Arima Valley, Trinidad, B.W.I.,” by William 3. Dryadula phaetusa 148 4. Dryas iulia 148 Beebe, Zoologica, 1952, Vol. 37, No. 13, pp. 157- 184]. 5. Philaethria dido 148 6. Heliconius aliphera 148 Contents page 7. Heliconius isabella 149 I. Introduction Ill 8. Heliconius melpomene .... 149 9. Heliconius numata 149 II. Materials and Methods 113 10. Heliconius wallacei 149 III. Eggs 115 11. Heliconius sara 149 IV. Larvae 115 12. Heliconius erato 149 13. Heliconius ricini 150 A. Characteristics of the Subfamily . 115 14. Heliconius doris 150 B. Key to Fifth Instar Larvae 127 VI. Discussion 151 C. Descriptions of the Species 129 1. Dione juno 129 VII. Summary 152 2. Agraulis vanillae 130 VIII. References 153 3. Dryadula phaetusa 131 4. Dryas iulia 132 5. Philaethria dido 135 I. Introduction 6. Heliconius isabella 137 HIS contribution is one of a series on the Contribution No. 1,000, Department of Tropical Re- biology and ecology of butterflies of the search New York Zoological Society. T nymphalid subfamily Heliconiinae. The 2This study has been completed with the aid of Nat- egg, larva and pupa of all of the 14 species re- ional Science Foundation Grant G 6376. corded from Trinidad are described. It is for- 111 112 Zoologica: New York Zoological Society [45: 9 tunate that all except one genus in the subfamily The data in the present study have accumu- are represented, as well as species from most lated slowly since the opening of the Zoological groups composing the large genus Heliconius. Society’s Trinidad Field Station in 1950, when The study has a triple purpose. First, it aims the first egg of Heliconius erato was accidentally to make available to taxonomic workers descrip- brought into the laboratory on a casually col- tions of the growth stages in a relatively homo- lected vine, and the larva reared by the senior geneous group of butterflies. Such descriptions author. Although most of the heliconiine butter- occur in the literature for only a few heli- flies known from Trinidad were soon discovered coniines; without exception these accounts lack near the Station, the identification and descrip- the detail necessary for comparative study, tion of the larvae and their respective food- either interspecifically or of different popula- plants proceeded erratically, through successive tions. The second purpose is to provide a foun- seasons, as a by-product of the work on imaginal dation for studies on the ethology and sensory behavior. Meanwhile, as our interest grew in the physiology of the immature stages. Thirdly, the ethology and phylogeny of the group, it became contribution is intended to serve as one of a essential to study the immature stages. Serious series of basic studies leading to an understand- attempts to complete the data for the present ing of the phylogeny of the group and of the contribution began in 1959, and it was not until evolution of its noteworthy adaptations. May, 1960, that the last unknown larva, that of Certain characteristics make the subfamily es- H. numata, was secured. pecially suitable for detailed investigation. The Since each of the 14 species has seven pre- group is widely distributed throughout the trop- imaginal stages (egg, five larval instars and ics and subtropics of the western hemisphere. It pupa), 98 different forms are concerned. Each includes numerous species, many having an ex- had to be described in life, studied in preserva- tensive range, and sympatry is common. It has tive, and variations established through com- furnished classic examples illustrating the the- parisons of series. Whenever the life history of ories of aposematic coloration and Mullerian another species became known to us, revisions mimicry. Polymorphism is so prevalent in some in our existing keys, descriptions, and phylo- localities that the systematic standing of many genetic hypotheses became necessary. Also forms remains in question. Patterns of social be- available for study were the growth stages, living havior in both larvae and imagos are proving to and preserved, of eight of the same species from be surprisingly complex, with involved sensory Surinam, which served for preliminary geo- bases. Finally, members of the family have graphic comparisons. shown themselves to be well adapted to life in The study presented here deals only with tropical insectaries and, as larvae, in the labora- major external characters and is still, therefore, tory; their behavior in all stages can, therefore, far from complete. The most notable gaps are in be studied by observation and experiment. the lack of work on setae and crochets in older Six contributions in the series have already larvae, although those of the first instar have been published (Beebe, 1955; Crane & Fleming, been treated in detail by Fleming (1960), on 1953; Crane, 1954, 1955, 1957; Fleming, 1960). detailed comparisons of spinules on the scoli In their most general characteristics, the life and in the omission of mouthpart studies. It is histories of all 14 species under consideration expected that future studies of behavior and are similar and may be informally summarized sensory physiology will contribute to the knowl- as follows: The egg, which is laid on Passiflora edge of these aspects. spp., is characterized externally by a network of For generic nomenclature, Michener’s (1942) strong horizontal and vertical ridges. The larva revision of the Heliconiinae is followed; the has numerous, paired, branching spines, includ- specific names are those of Kaye in “The Butter- ing one pair on the head and three pairs on most flies of Trinidad” (1921). The subspecies rec- body segments; the body is otherwise almost ognized by Kaye are as follows: smooth. The pupa varies from very rough, with Dione juno juno (Cramer) tubercles, flanges and spines, to almost smooth; Agraulis vanillae vanillae (Linnaeus) gold-colored spots and a pair of long cephalic Dryadula phaetusa phaetusa (Linnaeus) appendages may be present or absent. The dura- Dryas iulia iulia (Fabricius) tion of the egg stage is four days or less; the five Philaethria dido dido (Clerck) larval instars require a minimum total of 13 days Heliconius Isabella isabella (Cramer) for completion; the pupal stage lasts 9 or 10 Heliconius aliphera aliphera (Godart) days. Although the total minimum time of de- Heliconius melpomene euryades Riffarth velopment from oviposition to emergence of the Heliconius numata ethilla Godart butterfly is accordingly 26 days, development Heliconius erato hydara Hewitson often continues for three or four days longer. Heliconius ricini insulana Stichel 1960] Beebe, Crane & Fleming: Eggs, Larvae and Pupae in Heliconiine Butterflies 113 Heliconius sara thamar (Hiibner) four people, each of whom it is our pleasure to Heliconius wallacei wallacei Reakirt thank: Julie C. Emsley for all drawings of eggs Heliconius doris doris (Linnaeus) and larvae, except Text-figs. 5A, 5F, and 10-14; It will be noted that more than three-fifths Frances Waite Gibson for the latter text-figures of the species occurring on the island are re- and for all drawings of pupae (Pis. XIV and ferred to the mainland form originally de- XV); Russ Kinne for all the photographs in scribed. In the general geographical region Pis. I-XIII; and Sam Dunton for PI. XVI. We under consideration, infraspecific distinctions at wish also to express our appreciation to the present appear to be so much in need of revision following who, through their patience and skill that we have omitted subspecific names alto- in rearing the larvae during various seasons, gether from the body of the text. made this study possible: Susan Allan, Con- stance Carter, Frances Waite Gibson, To Fritz Muller we owe the beginning of a Rosemary comparative study of heliconiine development. Kenedy, Jane S. Kinne, Ellen Ordway and Bar- bara Working in Brazil, he described (1877, 1877- Young. 1878) morphological characters of eggs, larvae Finally, special appreciation goes to a group and pupae in seven species, in sufficient detail of organizations and individuals who have con- to be helpful today. Moreover, his attempt to tributed generously to the study: The National trace phylogenetic relationships in the group, Science Foundation, the National Geographic with reference both to ethological and morpho- Society, the Alcoa Steamship Company, Mrs. logical characters, is the only effort along these Mabel Ingalls and Mr. Kurt Reisinger. lines that has been made up to the present. II. Materials and Methods Muller’s work concerned the following species: Dione juno, Agraulis vanillae, Dryas iulia, Phil- Throughout the seasons at the field station, aethria dido, Heliconius eucrata (narcaea), H.