VOLUME 30, NUMBER 1 23

NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF BUTTERFLIES OF EL SALVADOR. VII. ARCHAEOPREPONA DEMOPHON CENTRALlS (NYMPHALIDAE)

ALBERTO M UYSHONDT 101 Avenida Norte #322, San Salvador, El Salvador

During August 1971, one of my sons found a strange-looking larva on a very small unknown shrub in a ravine near the city of San Salvador. The larva unfortunately died before pupating due to the lack of food. We could not identify the shrub because it was not in flower, and our efforts to substitute other similar plants for food were unsuccessful. Two years later, we found a female Archaeoprepona demophon centralis Fruhstor­ fer (Charaxinae) ovipositing on a larger flowering shrub and were able to rear the species to adult. The larvae were the same as the single specimen collected in 1971. Since that time we have reared the species several times. The rearing was done inside transparent plastic bags. The larvae were kept supplied with fresh leaves of the foodplant. The plastic bags were cleaned daily of excess moisture and excreta. Shortly before pupation, the larvae were transferred to a wooden box with mosquito-net windows. The adults emerged in the same box. Measurements of the different instars were recorded, and photographs were taken of all developmental stages. Some specimens of the early stages and exuvia were preserved in alcohol. These will be sent to the American Museum of Natural History, New York. The adults were determined by Dr. A. H. B. Rydon, the foodplant by Dr. D. Burch, University of South Florida.

LIFE CYCLE Egg (Figs. 1, 2). White, spherical but for slightly flattened base, smooth, no visible sculpturing at 15X magnification, 2.5 mm diameter. When ready to hatch in 6 days, head and body marks visible through eggshell. First instar larva (Fig. 4). General color pale brown. Head roundish, naked, with a dark brown M mark on front. Naked body thickens from head to 2nd abdominal segment, then narrows caudad, terminates in two short tails. Promi­ nence on dorsal meson of 3rd thoracic segment, another subdorsally on each side of 2nd abdominal segment directly above corresponding spiraculum. Thoracic segments with fine, dark brown lines. Abdominal segments darker brown dorsally. Anal prolegs smaller than others. Thoracic spiraculum larger than abdominals. Second abdominal spiraculum very high, completely out of line from others, except that 8th abdominal spiraculum also slightly higher. Grows from 4-10 mm in 13 days. Second instar larva (Fig. 5). Head with same aspect as in first instar except for one short horn on each epicranial apex. Body also as in first instar except dorsal hump more noticeable and tails longer and straight. Grows to 14 mm in 8 days. 24 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Figs. 1-6. Archaeoprepona demophon centralis. 1, Egg showing characteristic M on larval head, width about 2.5 mm; 2, egg showing dorsal markings of the larval body; 3, egg parasitied by larvae of an unknown Chalcididae; 4, first ins tar larva, about 6 mm long, on resting perch; 5, second instar larva, about 14 mm long, baring a vein; 6, third instar larva, about 27 mm long.

Third instar larva (Fig. 6). Same general colors as earlier. Head with longer horns; appearance of small, blunt, lateral projections below and behind horns. Mesal projection on 3rd thoracic segment and subdorsal ones on 2nd abdominal segment very pronounced. Dark brown line separating dorsal darker area from paler sub­ spiracular area which is criss-crossed by faint brown lines as is the thoracic zone. Caudal projections longer, still straight. Grows to 30 mm in 8 days. Fourth instar larva (Fig. 7). Head and horns covered by small tubercles, pro­ ducing a coarse aspect; lateral projections more noticeable. Body color darker, mostly over supraspiracular and dorsal areas, darker slanting lines. Appearance of tiny, bright blue spots along spiracular zone, around mesal prominence on 3rd thoracic segment and on longer, slightly crooked tails. Grows to 48 mm in 13 days. Fifth instar larva (Figs. 8, 9). Head and horns much rougher, lateral projections quite noticeable. Body color brown on thoracic segments and snbspiracularly on VOLUME 30, NUMBER 1 25

Figs. 7-11. Archaeoprepona demophon centralis. 7, Fourth ins tar larva crawling, about 45 mm long; 8, fifth instar larva recently moulted, about 50 mm long; 9 detail of head of fifth instal' larva; 10, prepupa in typical resting position, about 72 mm long; 11, prepupa ready to pupate, notice placement of tails. abdominal segments. Dark brown triangle dorsally covering area limited by 3 very prominent projections on 3rd thoracic and 2nd abdominal segments. Dorsa of remaining segments from spiracular line to meson, paler brown with darker brown slanting bands. Tails dark brown with yellow spots, very long and crooked. Profusion of tiny, bright blue spots on tails, along spiracular line and around thoracic prominence. Prolegs very thick. Grows to 70-80 mm in 18 days. Prepupa (Figs. 10, 11). Drastic color change. Head and body green, darker on head, thoracic prominence and along spiracular line from 6th-9th abdominal seg­ ments. Blue spots still visible. Pupa (Figs. 12-14). Bluish-green with scattered whitish, irregular spots; head bifid with points yellowish. Spiracula light brown surrounded by white ovals. Prior to adult emergence becomes very dark gray, some adult colors visible through cuticle. Lateral view: ventral profile only slightly convex; dorsal profile widens gradually from green cremaster to 5th abdominal segment, then the 4th forms a distinct hump, then narorws gradually to bifid head. Dorsal and ventral aspect: lateral profile widens smoothly from cremaster to wingcases, at thoracic level, then 26 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Figs. 12-18. Archaeoprepona demophon centralis. 12, Pupa, ventral aspect, about 42 mm long; 13, pupa, lateral aspect; 14, pupa, dorsal aspect; 15-18, sequence show­ ing emergence of adult male.

narrows abruptly to head. Measurements: 42 mm long, 18 mm laterally, and 20 mm dorso-ventrally at widest points. Adult (Figs. 19-22). Males and females shaped similarly. Forewing with faintly convex costal margin forming a rounded apex, concave outer margin to rounded tomus, straight inner margin. Hindwing with convex costal margin, strong humeral lobe, rounded outer angle continuing with rounded outer margin to VOLm-IE 30, NUMBER 1 27

Figs. 19-22. Archaeoprepona demophon centralis. 19, Adult male, dorsal view, wingspan about 82 mm; 20, adult female, dorsal view, wingspan about 103 mm; 21, adult male, ventral view; 22, adult female, ventral view. rounded anal angle, rounded inner margin with a fold. Dorsal ground color of forewing dull black in males, dark brown in females, with a greenish-blue iridescent isosceles triangle from median arca of wing with base on mid-inner margin; 2 small subapical, bluish spots. Hindwing with small basal, black triangle and broad, greenish-blue iridescent band from mid-costal margin ending in a point near inner margin, close to anal angle; this band with definite pale blue tinge along basal border. Inner fold and thin border along cestal margin, pale gray. Males with black brush of hairs near base of hindwing. Ventrally, dominant color pale brown with complicated pattern of darker brown areas and thin black lines; along outer margin of hindwing a row of small "eyes" with light blue pupils. Antennae and eyes black, proboscis orange. Dorsal and ventral surfaces of body concolorous with corresponding wing surfaces. Wing span 80 mm in males, up to 105 mm in females. Total developmental time for this species 85 days.

NATURAL HISTORY The females of Archaeoprepona demophon centralis deposit their eggs singly on the lower surfaces of the mature leaves of a shrub determined by Dr. D. Burch to be at least very close to Malpighia glabra L. (Mal­ pighiaceae). The pure white egg is quite visible against the dark green leaf. 28 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

The newly hatched larva completely consumes the eggshell, leaving only the shiny base on the leaf, and stays for one day without further feeding. It then moves to the tip of the leaf and nibbles around the central vein, baring it and affixing to it small, frass pellets woven with silk until the vein is prolonged beyond the leaf limits. Pieces of leaf tissue hung from short lengths of silk are added to the resultant perch. This is used as a resting perch during the day where the small larva is very inconspicuous among the dried bits of leaf. It is abandoned by the larva only momentarily to feed at dawn or dusk. First, second and third instar larvae use a perch. If the leaf is consumed during this period, another perch is made on a second leaf, and in some cases, a third. From the fourth through fifth instars, the larva wanders about the plant, but usually stays motionless during the day resting on a twig, head and tail hanging at the sides, giving the dorsal anterior part of the body a rep­ tilian appearance (i.e., head of small snake or lizard) with the promi­ nences on the second abdominal segment resembling the eyes and the mesal prominence on the third thoracic segment, the snout. The effect is enhanced by the crawl of the larva which imitates an inquhing snake head. The mesal prominence on the third thoracic segment is retractile; when touched it almost disappears. The prepupal larva becomes green and very inconspicuous, blending with the green foliage. At this time, the larva wanders for two days searching for a suitable pupation site which may be on the same food­ plant. Once the site has been chosen, usually a twig, the larva weaves a silk pad girdling the twig, reinforcing the twig's attachment to the shrub. The anal prolegs are then applied to the pad, the tails are positioned on either side of the twig, and the larva hangs incurved ven­ trally with its head almost touching the anal prolegs. It hangs thus for two additional days. The pupa is also very cryptic due to its green color which matches the foliage. It is very passive, but when roughly handled it reacts by swinging laterally and then returning to the vertical position. The pupal color changes to very dark gray shortly before the adult emerges. Some of the adult colors are actually visible through the pupal cuticle before emergence occurs. The adult emerges rapidly and hangs from the smoky gray pupal cuticle while expanding its wings and ejecting the meconium (Figs. 15- 18). The adult is ready to fly in about 25 minutes. The adults of A. demophon centralis, like most Charaxinae, are strong, swift fliers, producing an audible rustling noise at short distance. They usually keep to tree-tops and come to the ground only to feed on fer- VOLUME 30, NUMBER 1 29

menting fruits (e.g., mangoes and guayabas) or vertebrate excrement. Both sexes are lured easily to baits of fem1ented banana. They also feed on sap oozing from tree wounds. The male exhibits a strong territorial behavior and perches on its chosen tree, chasing vigorously any tres­ passing butterfly. The female flies at lower levels, mostly when ready to oviposit. We have observed A. demophon centralis most frequently around wooded coffee plantations from 600-1200 m elevation. The foodplant, M alpighia glahra L. (?), is found most commonly in wooded ravines within the flight range of the adult. It is a small shrub with perennial, opposite, stipulate, ovate leaves and pinkish flowers with characteristic dentate petals arranged in cymes. The fruit is a red drupe when mature with up to 3 carpels. It was observed that the eggs of A. demophon centralis are very often parasitized by Chalcididae wasps (Fig. 3), and the very young larvae are preyed upon by spiders.

Discussion Species belonging to the genus Archaeoprepona, prior to its establish­ ment by Fruhstorfer in 1916, were placed in the genus Prepona Boisduval (Hemming, 1967). The type species for Archaeoprepona was designated as demophon Linneaus. Le Moult (1932) chose to ignore Fruhstorfer's genus and established the invalid synonym Pseudoprepona with demo­ phon again as the type species. Some modern authors (e.g., Descimon, et aI., 1973) consider Archaeoprepona to be a subgenus of Prepona. Basic differences in the adults: black hair tufts on the hindwing of Archaeoprepona, against the honey-colored tufts of Prepona and the male genitalic differences and antennal colorations as noted by Fruhstorfer (1916) consistently correspond to morphological differences during the larval and pupal stages. The data pertaining to the immature stages are found in a number of sources. Included are the descriptions of the early stages made by Muller (1886) of Archaeoprepona amphimachus Fabri­ cius, A. catachlora Staudinger, A. demophon extincta Staudinger (all un­ der the generic name Prepona) and Prepona laeries Hubner. Lichy's (1933) description of the early stages of Prepona omphale guatemalensis Le Moult and that of P. omphale octavia Fruhstorfer by Muyshondt ( 1973a) also are important sources. In addition to the above life history of Archaeoprepona demophon centralis, another study that will soon be published on a yet, undetermined species of Archaeoprepona is taken into consideration. All the descriptions of the mentioned Prepona species show that the two horns on the head are fused and appear as a single epicranial pro- 30 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY jection. The Archaeoprepona species have two distinct horns, one on each epicranial apex plus a shorter horn below and behind each large epi­ cranial horn. The general habitus of the larvae of both groups is similar except Prepona species lack the median dorsal prominence on the third thoracic segment which is present in all known Archaeoprepona larvae. Prepona larvae do not show the drastic color change when entering the prepupal stage which we noticed in rearing Archaeoprepona demophon centralis and the undetermined A. sp. Muller's (1886) descriptions make no mention of prepupal color changes. The pupae of Prepona gradually are humped dorsally while Archaeoprepona species show a very pro­ nounced hump on the fourth abdominal segment. Also the pupae of Prepona have orange, irregular spots; these are white in Archaeoprepona pupae. All these differences favor the concept of two genera, but the many similarities, including behavioral patterns, indicate that both genera belong under the same immediate higher classification. On the same basis it also is very evident that these genera are closely related to the Zaretis-Siderone complex (Rydon, 1971; Muyshondt, 1973b and ms. in prep.). These facts seem to be in agreement with Rydon's treatment of the Charaxidae (with family status) which, in addition to other sub­ families that include Old World genera, separates the American genera into three subfamilies: Preponinae (Prepona, Archaeoprepona, Agrias, Anaeomorplw and N oreppa ), Zaretidinae (C oenophle bia, Zm'etis and Siderone) and Anaeinae (Fountainea, Hypna, Anaea, Polygrapha, Con­ sul, Cymatogramma and Memphis) with Zaretidinae being the link be­ tween Preponinae and Anaeinae. Some authors have tried to prove close phylogenetic relations between the Charaxinae and other groups, e.g., Limenitinae (Reuter, 1896; Muller, 1886). According to Fruhstorfer (1924), Hahnel thought there was an affinity between Prepona and the Apaturinae. If the consensus is fol­ lowed that the more reliable morphological characters to determine phylogenetic relationships are those which resist, to a greater degree, changes induced by selection, and these are the characters of the early stages (Brower, et aI., 1963), it appears that neither the Limenitinae nor the Apaturinae show much in common with the Charaxinae, nor with any other group of the Nymphalidae. Rydon (1971) may be right in assign­ ing family status, as other authors have also done, to the Charaxinae. The smooth, spherical eggs (flattened base and micropylar area or not), the spineless (Dornenlossen of Muller [1886]) larvae and the peculiar pupae of this group have nothing in common with either the very sculp­ tured (pineapple-like) eggs, the odd-looking, spine.·covered larvae or the VOLUME 30, NUMBER 1 31 pupae with characteristic projections of the Limenitinae. In a series of future artieles observations on the local Adelpha (Limenitinae) will cmphasize this point. The spherically ribbed eggs and the flattcned pupae of the Apaturinae do not have much in common with the Charaxinae either. It is true that the Charaxinae show some behavioral similarities with other nymphalids (e.g., the making of resting perches during the larval stage, adult feeding of fermenting fruit, etc.), but this might very well be the result of selective forces producing independently convergent successful strategies in species otherwise distantly related as is the case in many color similarities between species forming Miillerian mimicry complexes. It frequently happens that during the larval stage of a given species (e.g., Morpho peleides Kollar, Dione moneta Hiibner, various Apatura spp. and Smyrna hlomfildia [Fabricius] in particular), or during the pupal stage (e.g., Opsiphanes tamarindi Felder, O. cassina Felder, Zaretis itys [Cramer], and Anaea eurypyle [C. & R. Felder]), there are very noticeable differences in coloration even though during the early stages there should be less chance for diversification. There are also dif­ ferences of this sort in the adults of the same species, even coming from the same brood. We once obtained three adult males of Morpho peleides hyacinthus Butler, with four "eyes" on the ventral side of the forewing while another 18 specimens, males and females, only had three as is common. All 21 specimens were from eggs deposited by the same female on the same date in our insectary. They wcre fed their normal foodplants and wcre kept under identical conditions until adult emergcnce. Le Moult's (1932) opinion was that species of Prepona and Arclweoprepona are less variable than their near relatives, the Agrias, and have stable external characteristics. According to Dr. Descimon (pers. comm.), thc numerous new species and subspecics that Le Moult named on the basis of external charactcristics has caused grcat confusion. V ane-Wright ( 1974) thought many of Le Moult's taxonomic conclusions were unsound because he "split" many previously accepted species on little evidence. Much "lumping" and "splitting" has bccn done in the past without having enough data to form sound conclusions. It is important that more investigations on the early stages of the neotropical Rhopalocera be conducted so that systematists can use the results to establish a more accurate scheme of classification.

ACKNOWLEDGMENTS We wish to express our gratitudc to Dr. A. B. Klots for encouraging us to publish the results of our studies and to Dr. G. L. Godfrey for read- 32 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY ing, criticising and suggesting improvements for our article. We espe­ cially thank Dr. A. H. B. Rydon and Dr. H. Descimon, Ecole Normale Superieure, Paris, for supplying us with a wealth of reference material and Dr. L. D. Miller for sending Vane-Wright's publication on Prepona. Special mention is due to my children: Orlando, who first found the larva of the species reported in this study; Marilynn, who first saw the ovi­ positing female; and the youngest member of the team, Pierre, who obtained many other eggs and carefully fed the larvae during their long developmental time.

LITERATURE CrI'ED BROWER, L. P., J. Vz. BROWER, & C. T. COLLINS. 1963. Experimental studies of mimicry. 7. Relative palatability and Mi.illerian mimicry among neotropical butterflies of the subfamily Heliconiinae. Zoologica (New York) 48: 65-84. DESCIMON, H., J. M. DE MAEGHT, & J. R. STOFFEL. 1973. Contribution a ['etude des Nymphalides neotropicales. Description de trois nouveaux Prepona mexicains. Alexanor 8: 155-159. FRUHSTORFER, H. 1924. Genus Prepona Bvd., p. 550-566. In A. Seitz (ed.), The macrolepidoptera of the world. Vo!' 5. Stuttgart. HEMMING, F. 1967. The generic names of the butterflies and their type-species. (Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology. Supp!. 9. 509 p. LE MOULT, E. 1932. Etudes sur les Prepona. Nov. Entomo!', fasc. 2. LICHY, R. 1933. Observations biologiques sur les differents etats de Prepona omphale s/sp. guatemalensis Le Moult. (Lep., Nymph.). Nov. Entomo!', fasc. 3. MULLER, W. 1886. Si.idamerikanische Nymphalidenraupen. Versuch eines natiir­ lichen Systems der Nymphliden. Zoo!. Jahrb. Zeitschr. Syst., Geogr., BioI. der Thiere 1: 417-678. MUYSHONDT, A. 1973a. Notes on the life cycle and natural history of butterflies of EI Salvador. T. Prepona omphale octavia (Nymphalidae) . J. Lepid. Soc. 27: 210-219. ---. 1973b. Notes on the life cycle and natural history of butterflies of El Salvador. II. Anaea (Zaretis) itys. (Nymphalidae). J. Lepid. Soc. 27: 294-302. REUTER, E. 1896. Dber die Palpen der Rhopalocera. Helsingfors. RYDON, A. H. B. 1971. The systematics of the Charaxidae (Lepidoptera: Nympha­ loidea). Ent. Rec. J. Var. 83: 219-233, 283-287, 310-316,336-341, 384-388. VANE-WRIGHT, R. I. 1974. Eugene Le Moult's Prepona types (Lepidoptera: Nymphalidea, Charaxinae). Bull. Allyn Mus. 21: 1-10.