VOLUME 24, NUMBER 4 297 This is not the first time that material from the highlands west of Jalapa in the Schaus collection has been described as from thc city. The scarcity of this species, and for that matter of the Central American diocles, may be caused by collectors seeking these butterflies in tropical habitats, more typical for Adelpha, rather than in the high, moist montane valley where thcse specics occur. Acknowledgments We would like to thank Mr. R. 1. Vane-Wright of the British Museum (Natural History) and Mr. William D. Field of the U. S. National Mu­ seum for their help in comparing our material with the types of species in the respective museums. Literature Cited CLE:\fCH, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47: 1021-1034. FHUHSTORFER, H. 1915. Adelpha. In A. Seitz, Die Gross-schmetterlinge der Erde 5: 51O~533. GODMAN, F. D. AND O. SALVIN. 1870-1901. Biologia Centrali-Americana. Insecta. L epidoptera-Rhopalocera. London, 2 vols. THE AFFINITIES OF THE ITHOMIINAE AND THE SATYRINAE ( NYMPHALIDAE ) LAWRENCE E. Gn.BERT AN)) PAUL R. EHRLICH Department of Biological Sciences, Stanford University, Stanford, California Fox (1956) called into qucstion the classical association of the nymph­ alid subfamilies Danainae and Ithomiinae, arguing that the Ithomiinae are actually more closely related to the Satyrinae, especially the glossy-winged genera such as Pierella. Miller (1968) in his excellent revision of the Satyrinae quoted Fox's view and remarked on the presence of a vestige of vein 3V (.3A) as "structural evidence" supporting "the obvious similar tendency of the ithomiids and the satyrines to have hyaline wings" (p. 15). As Miller noted, however, this characteristic is also shared by the danaines. There is, of course, massive morphological evidence clearly showing that ithomiine adults are more similar to danaines than to satyrines (Ehrlich, 1958, Ehrlich and Ehrlich, 1967). In this paper we present further evi­ dence that Fox's judgement was in error. 298 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY In December, 1969, in the course of a field study of butterfly popula­ tions at the Beebe Tropical Research Station, Simla, Trinidad, W.I., we used "fedegoso" (dried plants of H eliotropum indicum L. (Boraginaceae) as an attractant to samplc populations of ithomiines for mark-release­ recapture experiments. We were impressed that this plant attracted not only very large numbers of male ithomiines (Ithomia drymo, Hymenitis andromica, Tithorea harmonia, Melinaea lilis, Hypothyris euclea, Mecha­ nitis isthmia, and others), but also largc numbers of males of the danaine, Lycorea ceres. No other butterflies were seen to pay the slightcst attcn­ tion to the fedegoso. A check of the literature reveals that Beebe himself ( 1955) had recorded ithomiines and both Lycorea and Danais plexippus visiting the lure. It is interesting to note here, that on Samoa and other South Pacific islands, danaines such as Euploea spp. and Danais melissa (again all males) are attracted by the dead twigs of Tournefortia argentia, another borage (Buxton, 1926, Poulton, 1936). An additional similarity among these isolated observations in the Neotropics and in Oceania is that amabd moths are attracted to these lures in both regions (sce also :\1oss 1947). Other behavioral traits link the Ithomiinae and Danainae. Both fly in the sedate manner which is typical of distasteful butterflies, and put on displays of agility only when attacked. Pierella, on the othcr hand, is one of the most rapidly flying satyrines, P. hyalinus displaying behavior char­ acteristic of butterflies with "flash and dazzle" coloration. We havc scen both danaines and ithomiines commonly taking nectar at flowers in the tropics, behavior rarely encountered in tropical satyrines. On the other hand, Pierella, Euptychia, and Taygetis all assemble on decaying fruit and are easily baited with it, but in our experience fruit baited traps havc never attracted ithomiincs or danaincs. Similar obscrvations on the absence of danaincs on fruit baits havc becn made in Africa (Hydon, 1964) where danaines wcre much more prominent in the butterfly fauna. In Euploea, at least, among the danaines, males are reputed to "assemble" females (Latter, 1935). It is also strongly suspected that male ithomiines "assemble" females (Gilbert, 1969). However, this point will remain uncertain until more is known about the courtship behavior of forest danaines, ithomiines, and satyrines. Finally, both Lycorea ceres and all of the ithomiincs we have investigated will feign death on capture, often remaining inert for up to five minutes. Wc have never observed this behavior in satyrines. One might argue that these behavioral similarities are all due to the involvement of the ithomiines and danaines in mimetic complexes, and indeed they may be. It should be noted, however, that Heliconills numata VOLUME 24, NUMBEH 4 289 and H. isabella (Nymphalinac), and Dismorphia amp hi one (Pieridae, Dismorphiinae) are involved in a mimicry complex with Lycorea ceres and Tithorea, Mechanitis, Meiinaea, and Hypothyris around Simla. Al­ though these share flight characteristics with Lycorea, Mechanitis, Meli­ naea, Hypothyris, and Tithorea, they do not visit fedegoso, they do not feign death, and, at least in the H eliconiini, males do not assemble females. We consider the morphological and behavioral evidence of the closeness of adult ithomiines to danaines significant when compared to the super­ ficial similarity in wing transparency of some ithomiines with a few satyrines. Similarly, evidence from immature stages indicates close affinity of ithomiines with danaines, not satyrines. The larvae of the first two always feed on dicotyledons, the third always on monocotyledons. Both Itho­ miinae and Danainae feed on plants rich in alkaloids, the former almost exclusively on Solanaceae and the latter primarily on Asclepiadaceae and Apocynaceae. Furthermore, Tithorea harmonia megara, one of the threc New World ithomiines reported to feed on non-solanaceous plants, fceds in Trinidad on Echites ( = Mesechites trifida Jacq., Apocynaceae; Guppy 1894, 1904). In Costa Rica, Fountaine (1911 ) reared Tithorea "pinthias" (= 1'. tal'ricina duenna Bates; see Fox, 1968) on a plant which from its description is probably Echites or another Apocynaceous vine. Guppy ( 1894) also reports finding Sais (= Ael'ia) eUl'ymedia on Echites in Trinidad. Morphologically, larvae of the danaine Lycorea and the ithomiine Tithorea are strikingly similar (descriptions in Guppy, 1894, 1904 and Fountaine, 1911 and illustrations in Guppy, 1904), and when disturbed have the same behavior of nervously twitching the long flexible protuber­ ances which arise from the first segment behind the head. Guppy (1894) describes A. eUl'ymedia larvae as almost identical to those of Tithorea except for their much smaller size at pupation. The larvae of such itho­ miine genera as Dil'cenna, Hypothyris, Ithomia, Goclyris, and Mechanitis do not superficially bear such strong resemblance to the danaines, nor to one another for that matter (Gilbert, personal observation). In any case, ithomiine larvae do not seem suggestive of the satyrines. Muller (1884) in his classic work "Sudamerikanische Nymphaliden Raupen" pointed out two characteristics in common among danaines and ithomiines. One is primary bristle development around larval bristle la. The second major characteristic suggesting close relationship is the rigidity of the pupae. The only two major differences between these groups according to Muller are, firstly, that danaines possess and itho­ miines lack the "Scherndornen" (long flexible protuberances), and sec- 300 JOUHNAL OF THE LEPlDOPTEHISTS' SOCIETY ondly that danaines feed on Asclepiadaceae whereas ithomiines feed on Solanaceae. Unfortunately, Millier apparently never worked on the life history of Tithorea, which represents a major exception to both of his criteria for separating thcse two groups. Acknow ledgments We would like to thank Michael C. Singer for advice in translating the Muller paper. Literature Cited BEEBE, w. 1955. Two little-known selective insect attractants. Zoologica 40: 27-32. BUXTO:'>!, P. A. 1926. Ellploea spp. frequenting dead twigs of TOtlrnefortia Argentea in Samoa and Tonga. Proc. Ent. Soc. Lond. 1: 35-37. EHRLICH, P. R. 1958. The comparative morphology, phylogeny and higher classifi­ cation of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull. 39: 305-.370. EHHLICH, P. R. AND ANNE H. EHRLICII. 1967. The phenetic rclationships of the butterflies. 1. Adult taxonomy and the nonspecificity hypothesis. Systematic Zoology 16: 301-317. FOUNTAINE, M. E. 1911. Five months butterfly collecting in Costa Rica in the summer of 1911. Entomologist 46: 189-194. Fox, R. M. 1956. A Monograph of the Ithomiidae (Lepidoptera) Part 1. Bull. Amer. Mus. Nat. Hist. 111(1). Fox, R. M. 1968. Ithomiidae (Lepidoptera: Nymphaloidea) of Central America. Trans. Amer. Ent. Soc. 94: 155-208. GILBEHT, L. E. 1969. Some aspects of the ecology and community structure of itho­ mid butterflies in Costa Rica-Mimeo Res. Rep. Adv. Pop. Bio. Organization for Tropical Studies, Costa Rica, pp. 69-93. LATTER, O. H. AND ELTRINGHAM, H. 1935. The epigamic behavior of Ettploea (Crastia) core asela (Moore) (Lepidoptera Danainae). Proe. Roy. Soc. (B) 117: 470- 482. MILLER, LEE D. 1968. The higher classification, phylogeny, and zoogeography of the Satyridae (Lepidoptera). Mem. Amer. Ent. Soc. 24. Moss, A. M. 1947. Notes on the Syntomidae of Para with special refcrence to wasp mimicry and fedegoso, Heliotropillm indicttm (Boraginaceae) as an attractant. Entomologist 80: 30-35. MULLER, W. 1886. Sudamerikanisehe Nymphaliden Raupen. Zool. Jahrb. 1. POULTON, E. B. 1936. Euploeine butterfly feeding at the same broken surface of a Toumefortia branch and some days later on the withered leaves of the same branch: Tulagi, Solomon Islands 1936. Proc. R. Ent. Soc. Lond. (A) 11: 94-96. .