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Crane 1957 Zoologica.Pdf ~ Q' ~;:: tu ,J"' c 0 ·.;:::-, gu <n c ,_,Cl 12 (fJ j I~ rn I;;; Q Imaginal Behavior in Butterflies of the Family Heliconiidae: VJ 1 I~ c I if! I Changing Social Patterns and Irrelevant Actions 0 B(t, !, -' 0 0 h z Ill J ;,. JoCELYN CRANE w i u -;> i m :t [:; ,, D4!partme11t of Tropical Research, New York Zoological Society,New York 60, N. Y. ,_Dd 3 N t: ::J CD 'fi U) 1: h" ;1 (Plate I) ~~1••c- ..-- T(f) :1 CT) _J *=•~- ..-- :\ [This paper is one of a series emanating from the characteristic of one pattern by actions from g .j c\ tropical Field Station of the New York Zoological another behavioral field . "'1" Society at Simla, Arima Valley, Trinidad, British ~ The observations presented arose as unex­ .;·:. West Indies. The Station was founded in 1950 by \!._; {.>:__ ~ pected incidental results of a comparative etho­ ~.:. ( the Zoological Society's Department of Tropical Re­ 0 ) search, under the direction of Dr. William Beebe. It logical study of six species common on the island ~ I comprises 200 acres in the middle of the Northern of Trinidad, British West Indies. The publication ;j Range, which includes large stretches of undisturbed i~-- of these data appears to be a prerequisite to the [_'; government forest reserves; The laboratory of the adequate exposition of comparative specific iJ_ Station is intended for research in tropical ecology characteristics and to an illumination of the L and in animal behavior. The altitude of the research group's phylogeny. A first paper in the series, on 1- area is 500 to 1,800 feet, with an annual rainfall of H eliconius erato hydara Hewitson, bas been more than 100 inches. 1published in which the effect of age on behavior [For further ecological details of meteorology and -and the occurrence of displacement behavior biotic zones see "Introduction to the Ecology of the -~..::: were mentioned (Crani~, 1955). Five additional ~z,,_ Arima Valley, Trinidad, B.W.I.," William Beebe. (Zoologica, 1952, Vol. 37, No. 13, pp. 157-184.)] species, along with H. erato, are considered ...;:.. llJ CJ -, below. These consist of Dryas julia julia (Fab­ •'--- __c: f ~ (!) "'"- ricius), Heliconius melpomene euryades Rif­ cc ! <f CONTENTS Page <i. 0 0__ J r.: farth, H. sara rhea Cramer, H. ficini insulana n- u I ~- z I. Introduction . 135 Stiebel and H. isabella isabel/a Cramer. Details [\:) 0 >-- ,,, e? 0 (I) Il. Historical Review . 136 f- o: c ..c u.. (I) ill. Materials and Methods .............. 137 of their behavior patterns, accounts of releasing -~J ~'J w () if <f. 0 :::; n: IV. Survey of Social Behavior in Six Species mechanisms and discussion of phylogenetic as­ 0: "" 0 :r~~ U:J ~-' C' UJ 0 of Trinidad Heliconiidae . 137 pects are being reserved for more appropriate I< ·(;.__ _, __J l: 1-- V. Changes in Social Patterns with Age. 138 inclusion in forthcoming contributions. i ro ~) "- TI a:: ... , ((} :') y VI. Irrelevant Actions . 139 w .l:l .~ tj: cc . The risks of inaccuracy and incompleteness in 1-- ::- E u r-. vn. Discussion ........................ 141 the description of butterfly behavior became ap­ I • ---' > 143 f z ... m .- VIII. Summary . parent very early in the work, particularly in I $ .;;;. _U,i'""' L1.I.,, IX. References ........................ 143 ! !Z r- ~.: ,__, the study of courtships. The patterns seemed at once surprisingly variable within a species, un­ l. INTRODUCTION i:c expectedly similar among species and altogether HIS contribution is one of a series on the too unpredictable to be characteristic of arthro­ biology and ecology of butterflies of the pods, with their high degree of dependence on T neotropical family Heliconiidae. It con- stereotypic forms of behavior. cerns post-imaginal changes in social behavior Since the butterflies were being studied prin­ patterns, and various kinds of irrelevant be­ I cipally in outdoor insectaries, it was thought at , I I havior, including the displacement of actions first that the discrepancies might be the result of unnatural conditions-although even if that were icontribution No. 979, Department of Tropical Re­ so the interest of the variability of the be­ search, New York Zoological Society. havior would scarcely have been lessened. 135 -. 136 Zoologica: New York Zoological Society (42: 12 Crane: lmaginal Behavior in Butterflies of the Family Heliconiidae 137 1957] 1, I Enough corroborative observations have now I sequence of changes in colony functions have known, but they have reached their highest de­ by flowering weeds and by the pond. Branches been made in the field, however, to show that been studied in worker honeybees (Rosch, velopment in birds . So far as is at present with bromeliads attached stand upright at inter­ the apparent eccentricities are equally charac­ 1925), Polistes (Steiner, 1932) and in various known they have a comparatively insignificant vals against the netting, as a further aid in main­ teristic of free-flying butterflies. Further cor­ ants (e.g. Buckingham, 1910). Verlaine (1932) taining humidity and natural conditions. The roboration of the natural prevalence of the types reported differences between young and old role among insects. A thwarted solitary wasp, Bembex rostrata, when forced to remove pebbles wild banana group (Heliconia) beside the bog of behavior under consideration comes from the mason wasps in nest-repairing and provision be­ forms the coolest, most shady corner of the cage. fact that, now that the causes are better under­ havior, that of old bees late in the season being repeatedly from the mouth of its burrow merely buzzed loudly and ran around in a wide arc H. erato, melpomene and other shade-lovers in­ stood, the patterns shown in the insectaries are incomplete; this observation invalidated a con­ variably seek it out during the heat of the day. highly predictable. clusion drawn by Fabre (1879; ed. 1920) from (Nielsen, 1945). When experimenting with but­ terflies attracted to pieces of coloured paper In contrast, other species, such as H. isabella, Some of the behavioral discrepancies have experiments which he performed, using aged which is most active around noon, frequent the individuals. Pardi (1947) found that age was Dr. D. Ilse noticed movements which might proved to be due simply to age differences in the open center of the cage around a group of Lan­ one of the factors determining the status of in­ have been displacement activities (personal individuals observed. The full courtship pattern tana, B.idens and Asclepias. In this cage all of dividual Polistes females in a dominance hier­ communication) . Possibly one of the factors re­ characteristic of the species, for example, is the latter, favorite food blossoms of the heli­ elicited only in and by individuals between cer­ archy. Nielsen & Nielsen (1952) reported that sponsible for the apparently slower speciation of the migratory period of a pierid butterfly insects than birds (Mayr, 1942) is the greater coniids, thrive and are allowed to grow freely tain ages, although successful reproduction can over most of the cage. This scattering of a take place throughout a much longer portion of (Astia) was confined to a single day of the five­ displacement-proneness of the latter." day life-span. As an example of intrageneric A few recent observations and comments, natural food supply encourages a normal amount imaginal life. of flight by the butterflies. Other irregularities appear when, as Tin­ variation, the parasitic wasp genus (Opius) may however, suggest that displacement activities The success of the new design is attested by bergen (1952, p. 26) defines the conditions for be cited: males of certain species cannot mate may prove, after all, to be widespread among the fact that all of the six species (p. 135) of displacement activities, "a strongly activated for five days or more after emergence, although · higher arthropods. References to date appear heliconiids discussed in the present paper feed, drive is denied discharge through its own con­ in other species they do so early in the, imaginal to be confined to the following: salticid spiders court, mate and lay eggs. Many individuals re­ summatory act(s)." However, in the present period (Hagen, 1953). Finally, entomologists (Crane, 1949), mantids (idem, 1952), the but­ paper the term "irrelevant behavior" (suggested would probably agree that female insects that terfly Heliconius erato (idem, 1955), Drosophila main alive, barring accidents, for one to three and one-half months, although differences in by Rand, 1943) will be employed as a more have recently molted into the final instar are in (Bastock & Manning, 1955) and fiddler crabs, general term than "displacement behavior." The general more attractive to males than are older genus Vea (Gordon, 1955, and Crane, 1957). viability are shown. H. isabella is the most dif­ latter, it seems, may be usefully restricted to the individuals. It will be noted, however, that use ficult to maintain, and observations on this III. MATERIAL AND METHODS definition given by Bastock, Morris & Moynihan of the inexact term "recently" is necessary. species are still somewhat deficient. Two locally rare heliconiids, Philaetraea dido (Linnaeus) (1~5~, p. 25) :. "A displacement activity is an Rockstein (1956) discussed the unreality of The studies were all conducted in out-of-door activity belongmg to the executive motor pat­ a sharp boundary between the pupal and imag­ wire mesh insectaries in Trinidad (Crane & Flem­ and Heliconius wallacei Reakirt, have not been tern of an instinct other than the instincts inal stages in insects, citing recent reasearcb on ing, 1953; Crane, 1955) between 1954 and successfully maintained. They live a few days activated." biochemical changes occurring after emergence 1957. During the past two seasons a new in­ and feed, but do not "settle in" and spend most This distinction between terms seems desir­ in the .worker honeybee, house fly, Drosophila, sectary, designed as were the earlier ones by of their active periods batting against the roof.
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