Annette Aiello.
Adelpha (Nymphalidae): Deception on the wing.
Psyche 91(1-2):1-45, 1984.

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PSYCHE
Vol. 91 1984 NO. 1-2
ADELPHA (NYMPHALIDAE):
DECEPTION ON THE WING*
BY ANNETTE AIELLO
Smithsonian Tropical Research Institute
P.O. Box 2072, Balboa, Republic of Panama For the past century, lepidopterists have puzzled over the genus Adelpha Htibner, in an attempt to discover the secret character or combination of characters which might lead to a satisfying classification of the 100 or more butterfly species included in this large neotropical group. Several approaches have been tried (Godman & Salvin, 1884, 1901; Fruhstorfer, 1907; Forbes, un- published manuscript): wing pattern (both upperside and lower- side), wing venation, genitalia, and various combinations of these. If one attempts to coordinate all the information available, the result is a hopeless tangle. As a result, the most obvious set of characters (wing pattern) traditionally has been used in classification; other character groups (genitalia, larvae, pupae) which appear to confuse the situation have been largely ignored. A new strategy is needed, one which would both evaluate the reliability of the different groups of characters already surveyed, and search anew for overlooked sources of information. It was the purpose of my research to review what is known of the immature stages of Adelpha species and, based upon that information plus my own observations made in Panama between 1978 and the present, to make speculations regarding species relationships.
*Manuscript received by the editor January 12, 1984.



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The first attempt at classification within Adelpha was made by Godman & Salvin (1884, 1901) with their revision of the 32 species reported from Central America. At that time some 70 species were known for the genus. In their treatment, Godman & Salvin discuss Adelpha and its relationship to Limenitis and note that several species (bredowi, populi, carnilla, lorquini) are difficult to assign to either genus. Distribution of such characters as eye pubescence, venational differences, variation in proportions of male leg seg- ments, and peculiarities of male genitalia within Adelpha-Limenitis is surveyed. Classification of the 32 species begins with isolation of A. bredowi (eyes smooth in front) from all others (eyes hairy in front) and continues by arranging the other 31 species using gross features of wing pattern. The result is eleven groups, six of which are represented by single species.
With Fruhstorfer (1907) came the first and only published revision of the entire genus Adelpha. The 90 species treated are assigned to two main groups based upon the length of the forewing discal cell; short = Adelpha group, elongate = Heterochroa group. Upon this division, Fruhstorfer comments, "Anatomically there are also two series [male genitalia with or without clunicula] of species distinguishable. They, however, do not agree with those based upon the structure [discal cell length]." The linear arrangement of species within the largest group (Heterochroa with 82 species) reflects presumed relationships based upon wing pattern features. Fruhstorfer touches upon but does not pursue the possibility of a closer than realized alliance between New and Old World groups. In his introduction he notes that the male clasping organs of AdeZpha ". . .approach those of the [Old World] genus Pantoporia (At-) in such a way that. . .it would be quite impossible to ascertain where organs or photes [sic] of them belong to, which are not denorni- nated." Concerning the male valves, he further observes, ". . . there exist also nearly square ones with 2 or 3 small acicular teeth
(resembling a Limenitis [Moduza]procris from India and Ceylon)." Fruhstorfer's revision included a number of misidentifications, which presumably were corrected by Hall (1938) following exa min- ation of the original material.
At the time of his death (1968) W. T. M. Forbes had made a good deal of progress on a revision of Adelpha, and his manuscript is in



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19841 Aiello - Genus Adelpha 3
the Archives of the Museum of Comparative Zoology, Harvard University. About half of this manuscript is in a nearly illegible hand; half is typewritten. The typed portion includes part of an introduction, notes on each species, a key (based upon wing pattern) to species and species groups, and descriptions of four new species. Included also are more than 70 genitalic illustrations (inside of right valve) by Howarth. Forbes had analyzed these latter and was in the process of constructing a key to genitalia. His two approaches, of wing pattern and genitalia, yielded different species groupings, and it is not clear whether he favored one of these, or intended to use a combination of the two in his final classification. Of the immature stages of Adelpha, very little has been said, and what has been said has been largely ignored for the simple reason that this new information appears to confuse matters rather than clarify them.' Forbes (uncompleted manuscript) had read Moss's (1933) paper on Adelpha larvae and pupae, and had examined several pupal skins of Adelpha and Old World Limenitini in the collections of the British Museum (Natural History) when he commented, "The larvae and pupae are highly varied and unless a high percent are misdetermined, show characters wholly incongru- ent with adult structures and patterns." The fact that, within Adelpha, the study of adult characters results in species-alignments different from those obtained by consideration of the immature stages and/or genitalia, suggests that at least one set of characters is unreliable or perhaps even deliberately deceptive.
Due to taxonomic confusion within Adelpha, it is impossible to know how many species actually are represented by the 34 or so life history accounts published for this genus. My estimate is that at least 24 species are illustrated (including illustrations in this paper) as final instar larva, or pupa, or both, many of these by more than one author. Most accounts, whether illustrated or not, include a foodplant record.
'This author has located only four publications (Moss, 1933; Muller, 1886; Young, 1974; Cornstock & Vasquez, 1960) which figure any immature stages of Adelpha; several publications present descriptions only.



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4 Psyche [vo~. 91
In the present paper, all illustrated reports are reviewed to see whether it is possible to distinguish natural species groups within the genus Adelpha.
Judging from the few scattered reports (Table 1) of oviposition in the Limenitidini, eggs are laid singly and are usually placed at the tips of leaves. While many accounts do not specify whether placement is on the upper or lower surface, the majority that do specify, report upper surface oviposition. Adelpha iphicla females alight upon a leaf and walk backwards while searching for the desired oviposition site with the tip of the abdomen; the egg is then placed and the butterfly takes flight before her next oviposition, which may be on the same leaf. In the case of lower surface oviposition, the female merely bends her abdomen around the edge of the leaf and touches its tip to the leaf. Among the six species of Adelpha whose eggs were collected in Panama, four (Table 1) place their eggs along damaged portions of leaves, especially on jagged points, as well as at the leaf tip, and one of these (A. iphicla) is about as likely to lay eggs along an intact leaf margin as at the tip. As well, a female A. iphicla may return to the same leaf several times and place as many as four eggs on one leaf, often on its undersurface.
Eggs of the seven Adelpha species (basiloides, cocala, cytherea, marcia, iphicla, melanthe, phylaca aethalia, and salmoneus) exam- ined by this author, were all similarly sculptured (figure 1) with pits (hexagonal due to packing) and seta-like projections (one from each junction of three pits). This same sculpture type is figured for Limenitis by both 2Scudder (1889) and Eltringham (1923). Young's (1974) description of the egg of Adelpha leucophthalma as having the seta-like projections "aris[ing] from the facets," is doubtful.
Also doubtful is the account by Comstock and Vazquez (1960) which describes and illustrates the egg of A. celerio as having convex rather then concave hexagons. Such an error is easily made, as these *As Basilarchia.




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19841 Aiello - Genus Adelpha 5
Table 1. Placement of egg on leaf by various members of the Limenitidini (Nymphalidae)
EGG
PLACE-
MENT REFERENCE
-- -
Adelpha basiloides Bates TI, Dl Aiello
Adelpha celerio diademata Fruh. T Comstock & Vazquez (1960, pg. 407)
Adelpha cocala Cr. Ti, (Dl) Aiello
Adelpha cytherea (?) t Muller (1886, pg. 484) Adelpha iphicla L. TI(!), MQt) Aiello
Adelpha iphicla t Muller (1886, pg. 484) Adelpha isis Dru. Tl, Dl Miiller (1886, pg. 48 1) Adelpha leucophthalma tegeata Fruh. Dl Young (1974, fig. 2) Adelpha melanthe Bates Dl Aiello
Adelpha phylaca aethalia Feld. Tl Aiello Adelpha plesaure Tl Muller (1886, pg. 484) Adelpha salmoneus Butl. TI, (Dl) Aiello
Adelpha serpa Tl Muller (1886, pg. 484)
Adelpha syma Godt. T Hoffmann (1937, pg. 212) Athyma nefte Cr.
(as Parathyma nefte Cr.)
TI, (Mt )
Morrell (1954, pg. 160)
Afhyma opalina Kollar t Robson (1894, pg. 338) Lasippa tiga Moore
(as Neptis heliodore Fruh.) T Morrell (1954, pg. 160) Limenitis archippus
(as Basilarchia archippus) TI Scudder (1 889, fig. 16) Limenitis lorquini burrisonii Maynard T Dornfeld (1980, pg. 6 1) Moduza procris Cr. T Morrell (1954, pg. 157) Neptis nata Moore T Morrell (1954, pg. 162) T = leaf tip (tooth, in case of A syma), M = leaf margin (intact), D = damaged portion of leaf, I = upper surface of leaf, T = lower surface of leaf, () = less commonly Figure 1. Eggs of Adelpha (left) and Doxocopa (right).



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6 Psyche [vo~. 91
eggs can present an optical illusion, however, an edge view of a partly eaten egg shell shows clearly that the hexagons are indeed pits.
Comstock and Vazquez (1960) also describe and figure a differently sculptured egg, which they call A. iphicia with the reservation that it might actually be an egg of 3Doxocopa. Because their larvae died soon after hatching, the authors never knew that they had indeed been fooled by the ovipositing Doxocopa female, a mimic of Adelpha. My own rearing of Doxocopa laure (LOT 83-6) was from two eggs with pattern (figure 1) identical to that in figure 33
of their paper. My reared Adelpha iphicla were from eggs patterned as those of all the other Adelpha species examined so far. Possibly this egg sculpture pattern will be found throughout the Limenitini and may prove useful in defining tribal limits. Oviposition was observed only for Adelpha iphicla, and each of the three fresh eggs collected required 5 days development before the first instars emerged. Of five other species collected as eggs, of unknown age, three had longer minimum egg-development times than did A. iphicla: A. basiloides (LOT 82-65), one egg hatched after 6 days; A. cytherea (LOT 83-3), one egg hatched after 7 days; and A. salmoneus (LOT 83-14), two eggs hatched after 6 days. Because individuals were collected at various stages of devel- opment, total time from hatching to eclosion could be determined for only a few individuals of ten species. In spite of the fact that the resulting data (Table 2) is not uniform, it is clear that development time is variable, and often more so within a species than between species. The extreme example of wide variation is in A. basiloides which has a longer development time than other species, and which sometimes passes through six instars instead of the usual five. Five 6-instar individuals of A. basiloides were encountered, and these were from four separate rearing lots, each of which included 5-instar individuals as well. Each of these lots was collected on Amaioua corymbosa; larvae collected on Alibertia edulis and Bertiera guianensis did not produce extra instars. In spite of the additional instar, 6-instar individuals did not require longer development times 3As Chlorripe.




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19841 Aiello - Genus Adelpha 7
than 5-instar individuals, however, larvae on Amaioua, regard less of number of instars, were slower to develop (41-48 days) than were larvae on Alibertia or Bertiera (32-44 days). Miles Moss reared twelve species of Adelpha from Para, Brasil, and illustrated (1933) the final instar larva and the pupa for each of them. In spite of the lack of detail in his illustrations and descriptions, and the fact that his scheme for numbering body segments omits abdominal segment-9, his little paper remains tile masterpiece on Adelpha immatures. It was Moss who realized (page 15) that:
". . .systematists,. . . by a careful examination of certain hither- to unsuspected points of likeness or dissimilarity between the species in their early stages, may perhaps be led to modify tile existing order and grouping of the butterflies of this difficult genus. It is just possible that a few unexpectedly close relationships may thus be established, while others at present confused, or regarded as near of kin, may be found to be more distantly related than was supposed."
He concluded that A. cytherea, and pseudococala are closely related, also delphicola and mesentina, and serpa and paraw. My analysis of published illustrations and live material lends support to Moss's groupings and adds several more.
FIRST INSTAR LARVAE
First instar larvae, of all species which I have seen, appear identical in form, and are some shade of brown or grey. The head bears setae, but none of the4chalazae of later instars; pale bumps on the body are found where the future ^coli will be, and the body is covered with tiny pale spots each centered with a minute seta. After their first meal, larvae take on the green color of their foodplant, although the head remains brown.
SECOND INSTAR LARVAE
From the second instar on, the head is ornamented with chalazae which give it a spiny appearance. The face has numerous round,



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[Vol. 91




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Table 2. (Continued) Duration (in days) of larval instars and pupal stage for twelve Adelpha species. leucophthalma
melanthe
nr. paraha -
phylaca aethalia m4
111
salmoneus (41, 5
PI
Unless otherwise noted, the above data was collected in Panama by the author. [l=n
() = commonest number of days in cases of strong bias towards one end of range; otherwise, ranges are normal curves. m = minimum number of days in cases where full time for an instar is not known.



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10 Psyche [vo~. 91
flatbottomed pits which first appear in small numbers in instar two, three, or four depending upon the species. These may be the same color as the rest of the head capsule or may be a contrasting color. Facial stripes appear in the fourth instar of some species as well. The body now bears stubby scoli (each with 3-5 radiating apical spines). These scoli are arranged in three rows (subdorsal, supra- spiracular, and subspiracular), thus, each body segment has three pairs of scoli. The subdorsal scoli on thoracic segments 2 & 3 and abdominal segments 2,7, & 8 are very slightly larger than the others, but all are similar in form.
Beginning with the second instar there are color and pattern changes in some species, but most are fairly uniformly colored (brown, green, or black) and have paler scoli and tiny spots, much as the first instar. In Adelpha basiloides, the larval color depends upon the foodplant; larvae are light brown or reddish brown on Amaioua, and dark brown on Alibertia and Bertiera. THIRD AND FOURTH INSTAR LARVAE
From instar three on, the face is framed by two distinct rings of chalazae, and body scoli show further development, especially those scoli which will be the largest or most distinctive in the final instar. By late fourth instar, body scoli have become swollen at the bases due to developing final instar scoli inside. In these two instars, the scolus spines are pale, and as before, the body is speckled with tiny pale spots.
A dorsal, paler patch ("saddle") appears in the third or fourth instar of many species. In A. salmoneus and cytherea the "saddle" is faint and extends from abdominal segment 2 or 3 through segment 6. A. cocala and leucophthalma may have a "saddle" on abdominal segments 5-6. A. justina is pale dorsally from thoracic segment 2 through abdominal segment 8. A. basiloides, nr. param and celerio have a triangular "saddle" with its base on the posterior portion of abdominal segment 4, and its apex at mid abdominal segment 6. In the latter two species the "saddle" is poorly defined; in A. basiloides it is sharply demarcated.
4Terminology of Peterson (1962). cHALAZAE(AE): a distinctly elevated cone-shaped area, bearing 1-3 simple setae; (SCOLUS(I): an elongated projection, bearing 4 or more setae or spines.




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Aiedo - Genus Adelpha
FINAL INSTAR LARVAE
Final nstar larvae (Figures 4 & 5) of all species studied have several tz mings in common.
The head (Figure 2) has a spiny
appearam e e owing to the numerous chalazae which frame the face. These s e c m to be fairly constant in number and position (Figure 3) and to v a Ty from species to species mainly in their relative size. A. iphicla f o example has a relatively smooth face because it lacks several D the chalazae found in other species. A, phylaca and especially mehihe are the spiniest in appearance due to numerous Wae. color and pattern also vary: A. celerio has a ~triped face, A. basiloides and A. cocala patterned ones. The pits of A. cytherea are darker than the rest of the face; the head of A. salmoneuS is reddish and constrasts strongly with its green body. Each b o d y segment bears three pairs of scoli (subdorsal, supra- ~~iracular- and subspiracular); there are no dorsal scoli. While the scoli are variable in form and length, in all species studied, those of the proth o Tax and abdomen-1 are either very short or are reduced to a few ~p5nes. Usually the longest are the subdorsal scoli of meso- and meta-zhorax and abdomen-2, -7 & -8, and the supraspiracular scoli of t h e meso-thorax. In many species the subdorsal scoli of abdomen-z are the most distinctive in form and often are curved backward S -
BO~Y scoli are diverse but the various forms can be grouped into two main types: those which are terete (round in cross-section), and those which are flattened.
The terete SCO~US, in its simplest form, is a short stalk with 3-5 spines radiating star-like from its apex (e.g., 5scoli A3-6 of A, celerio, cyzAerea, sa~moneus, and justina). More complex scoli are slender and longer with spines at intervals along their full length, and either with one to a few ascending spines towards the apex (thoracic scdi of many species, e.g., A. phylaca, melanthe, cocala, basiloide~, cytherea, justina, salmonem), or with 3-5 radiating apical spines (e.g., scoli of A3-6 in A. phylaca, melanthe). Scoli also may be short and thick with a dense covering of spines (e.g., A2 of A. cocala, Zeucophthalma, basiloides), or be club-shaped and more sparsely spined (e.g., A8 of basiloides). s~hroughout the remainder of this paper, "sco1us(i)*' refers to the subdorsal set unless otherwise noted; T = thorax, thoracic segment; A = abdomen, abdominal segment.



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Figure 2.
Faces of final instar larvae of Adelpha species reared in Panamh: B A S = basiloides, CEL = celerio, COC = cocala, CYT = cytherea marcia, IPH = iphicia, MEL = melanthe9 PHY = phylaca aethalia, SAL = salmoneus.



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Aiello - Genus Adelpha
Figure 3.
Location of head chalazae 1-5, referred to in text. Flattened scoli have a plumose or leaf-like appearance (e.g., A. celerio, serpa, paraena) which is due not only to the arrangement of spines in two opposite rows along the scolus, but also to the flattening and widening of those spines which may take on spatulate, elliptical, or lanceolate forms, and may be so crowded that they overlap one another. In the case of extremely condensed scoli, the spines appear to fuse with one another (see A. serpa in Muller, 1886).
Upon hatching, a larva eats some or all of its egg shell and then feeds on the leaf tip leaving the midrib intact. Larvae rest out on the midribs which they have exposed, and eventually extend them by addition of fecal pellets held in place with silk. Most larvae fashion several such supports during each instar, often using lateral veins instead of the midrib. When not feeding, Adelpha larvae rest out on their supports, facing away from the leac molting takes place on the support as well. After they molt to the final instar, larvae abandon their supports and thereafter rest on the upper surface of a leaf.



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14 Psyche [vo~. 91
In addition to the above behavior, typical of many nymphalid butterfly species, first through fourth instar Adelpha larvae engage in an odd practice not known for other New World butterflies; they accumulate their fecal pellets, fastening them in place with silk, to form a mass which either surrounds the base of the support or is suspended beneath it. This mass may include bits of leaf in A. iphicla, or consist of more leaf than feces in A. phylaca and melanthe. The work of A. basiloides is the most distinctive because, in addition to the mass just described, this species consistently constructs a small, usually curved, larva-form mass on the leaf surface, several mm away from either the leaf edge or the usual mass. Upon viewing this artistry, one cannot help imagining that it serves as a decoy larva to discourage would-be predators. A single reared individual of A. iphicla also engaged in this behavior, but excepting that, I have observed it only in A. basiloides. Resting larvae position their bodies in one of several ways: (1) Straight Position (figures 4 & 5). Larvae in any instar may use this position, and typically when out on their supports they rest this way.
(2) Front-Curved Position, as shown in Young (1974, fig. 2. B, C). In this position, the larva grasps its support with the prolegs only, and raises and curves its anterior portion (head through A2) so that the head is somewhat inclined, and the toracic scoli are directed forwards. Larvae about to molt use this position; the second instar in Young's (1974, fig. 2C) photograph has a swollen pro-thorax and is probably preparing to molt to third instar. Final instar larvae of A. justina use a raised but uncurved version of this position instead of the usual final instar stance, described next, (3) Front-Arched-Rear-Up Position.
This position is typical of
final instar larvae but occurs in earlier instars as well. Involved are the raising and arching of the anterior pdrtion of the body (head through A2), ,plus elevation of the posterior portion (A7-10). In addition, the thoracic scoli are directed forwards, and those of A2 are held backwards. In this position, the face and the area at the top of the arch (T3 through A2) are parallel to each other and to the substrate. This and the next position are assumed when the larva is disturbed.
(4) Curled Position. The larva curls to one side into a "C" or "J" shape on the upper surface of the leac the rear portion (A7-10) may or may not be elevated. A. celerio often rests in this position,



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19841 Aiello - Genus Adelpha 15
and when it does, the scoli on the outside of the curve stick out all around and the animal resembles a bit of moss. Moss (1933) reported the same behavior and appearance for A. thesprotia. At first inspection, it would appear that larvae of Adelpha butterflies are not very particular about their foodplant selection. Indeed, some 56 plant species, representing 42 genera and 16 families have been reported as larval foodplants of Adelpha. Moreover, many Adelpha species are known to be polyphagous: A. melanthe, delphicola, and isis have each been reared on three plant genera, A. celerio on four, and A. cocala and iphicla each on eight. Cecropia (Cecropiaceae), Sabicea (Rubiaceae), and Vitex (Verben- aceae) are each attacked by five different species of Adelpha. However, when larval foodplants are grouped by the butterfly species which feed upon them (Table 3), a pattern does emerge: Adelpha species fall nicely into two main feeding groups: (1) Rubiaceous feeders, and (2) Non-rubiaceous feeders. With the following three exceptions, butterfly species that feed upon members of the Rubiaceae have not been reported on plants outside that family. A. boreas tizona was reared on both Rubiaceae and Ericaceae (Marquis and DeVries, unpublished); A. cocalay reared on seven members of the Rubiaceae, also has been reported on Emm~tum of the Icacinaceae (Moss, 1933); A. syma, on Rubiaceae and Rosaceae. The record of A. iphicla iphicleola on Celtis (Ulmaceae) (Comstock and Vazquez, 1960) is in error; their butterfly was actually Doxocopa, misidentified as Adelpha. It is interesting to note that the rubiaceous genera, utilized as foodplants by Adelpha, belong to at least seven of the 18 tribes outlined for the Rubiaceae by Kirkbride (1982). Several non-rubiaceous feeders do show wide foodplant pref- erences, the extreme examples being: A. melanthe which feeds on three plant genera representing three families, and A. celerio which feeds on five genera in four families. The only common bond among the foodplants of these two butterflies seems to be that all are scabrous- or pubescent-leaved, second-growth trees. Some plants attract a more specialized group of feeders: Vitex (Verbenaceae) is attacked by A. abia, calliphaney epizygis, and jordani, which have not been reported on any other plants. A. naxia ipiphilca, also



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16 Psyche [vo~. 91
Figure 4.
Final instar larvae of Adelpha species reared in Panama: abbreviations are the same as in Figure 2 except that CEL = nr. CEL; in addition nr. PAR = near para2na.




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Aiello - Genus Adelpha
Figure 4. (Continued)




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18 Psyche [VOI. 91
Table 3. Larval foodplants reported for Adelpha species, arranged alphabetically by butterfly species.
A DELPHA SPECIES/ PLANT
FOODPLANT FAMILY LOCALITY REFERENCE
ABIA
Vitex sp.
BASILEA
Calycophyllum sp.
BASILOIDES
Alibertia edulis
Amaioua corymbosa
Bertiera guianensis
Ixora nicaraguensis
BOETIA OBERTHURI
Luehea seemannii
BOREAS TIZONA
Satyria sp.
Chomelia bispinosa
BREDOWI CALIFORNICA
Quercus chrysolepis
Quercus spp. (especially
chrysolepis
Qyercus spp.
BREDOWI EULALIA
Quercus spp.
CALLIPHANE
Vitex montevidensis
CALLIPHICLEA
Ilex paraguariensis
CELERIO
Cecropia peltata
Miconia argentea
Ochroma pyramidale
Ochroma pyramidale
Urera sp.
Verbenaceae Brasil
Rubiaceae
Rubiaceae
Rubiaceae
Ru biaceae
Ru biaceae
Tiliaceae
Ericaceae
Ru biaceae
Fagaceae
Fagaceae
Fagaceae
Fagaceae
Cuba
Panama
Panama
Panama
Panama
Costa Rica
Costa Rica
Costa Rica
California
California
California
Mexico, US
Verbenaceae Brasil
Aquifoliaceae Paraguay
Cecropiaceae Panama
Melasto- Costa Rica
mataceae
Bombacaceae Panama
Bombacaceae Panama
Urticaceae Costa Rica
Dewitz (1879)
Aiello LOTS 79-12],
80-38, 81-17, 81-42,
81-49, 83-16, 83-42
Aiello LOTS 82-59,
82-65, 82-70, 82-72,
83-42
Aiello LOTS 82-64,
82-73, 83-87
Mallet in DeVries
(unpub)
Mallet in DeVries
(unpub)
Marquis in DeVries
(unpub)
DeVries (unpub)
Orsak (1977)
Howe (1975)
Dornfeld (1 980)
Ferris & Brown (1 980)
cited in Lima (1968)
Jorgensen (1921)
Coley LOT 15
Mallet in DeVries
(u~PW
Aiello LOT 82-41
Coley LOT 22
DeVries (unpub)




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19841 Aiello - Genus Adelpha 19
Table 3. (continued)
A DELPHA SPECIES/ PLANT
FOODPLANT FAMILY LOCALITY REFERENCE
CELERIO DIADEMATA
Conostegia xalapensis
Miconia sp.
nr. CELERIO
Heliocarpus
popayanensis
COCALA
Calycophyllum
candidissimum
Calycophyllum
candidissimum
Chomelia psilocarpa
Emmotum nitens?
Malanea sp.
Melasto-
mataceae
Melasto-
mataceae
Tiliaceae
Rubiaceae
Rubiaceae
Rubiaceae
Icacinaceae
Rubiaceae
Pentagonia macrophylla Rubiaceae
Psychotria sp.
Uncaria tomentosa
Warscewiczia coccinea
CRETON
Quercus?
CYTHEREA
Sabicea aspera
CYTHEREA MARCIA
Sabicea panamensis
Sabicea villosa
Sabicea villosa
DELPHICOLA
Bombax munguba
Cecropia sp.
Pourouma sp.
DEMIALBA
Rondeletia sp.
DONYSA
Quercus?
Rubiaceae
Rubiaceae
Rubiaceae
Fagaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Mexico
Mexico
Panama
Panama
Costa Rica
Panama
Brasil
Brasil
Costa Rica
Costa Rica
Panama
Panama
Mexico
Brasil
Panama
Panama
Costa Rica
Bombacaceae Brasil
Cecropiaceae Brasil
Cecropiaceae Brasil
Rubiaceae Costa Rica
Fagaceae Mexico
Comstock & Vazquez
( 1 960)
Comstock & Vazquez
( 1 960)
Aiello LOTS 82-75,
83-68
Aiello LOT 81-44
Janzen 8 1 -SRNP-740
in DeVries (unpub)
Aiello LOT 81-14
Moss (1933)
Moss (1933)
DeVries (unpub)
Marquis in DeVries
(unpub)
Aiello LOT 81-54
Aiello LOT 82-66,
82-71, 82-74
Miller & Miller (1970)
Moss (1933)
Aiello LOT 83-129
Aiello LOTS 82-26
83-3
DeVries (unpub)
Moss (1933)
Moss (1933)
Moss (1933)
Haber in DeVries
(unpub)
Miller & Miller (1970)




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20 Psyche [VOI. 91
Table 3. (continued)
A DELPHA SPECIES1 PLANT
FOODPLANT FAMILY LOCALITY REFERENCE
EPIZYGIS
Vitex montevidensis
EROTIA
Tetrapteris sp.
FESSONIA
Randia echinocarpa
Randia learstenii
IPHICLA
A ntirrhoea trichantha
Alseis blackiana
Bathysa sp.
nr. barbinervis
Calycophyllum
candidissim um
Ca/ycophy/Zum
candidissim urn
Gonzalea spicata
Gonzalea spicata
Isertia haenkeana
Rondeletia panamensis
Verbenaceae Brasil
Malpighi-
aceae
Rubiaceae
Rubiaceae
R u biaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Uncaria (as Ourouparia)
guianensis Rubiaceae
Uncaria tomentosa Rubiaceae
IPHICLA EPHESA (as ephicla ephesa)
Bathysa sp. Rubiaceae
IPHICLA IPHICLEOLA
Calycoph yllurn
candidissimum Rubiaceae
Brasil
Costa Rica
Costa Rica
Panama
Panama
Brasil
Panama
Cuba
W.I.
W.I.
Panama
Panama
Brad
Panama
Brad
Costa Rica
cited in Lima (1968)
Janzen 79-SRNP-2 16
in DeVries (unpub)
Janzen 79-SRNP-725
in DeVries (unpub)
Aiello LOTS 81-25,
82-27, 83-41
Aiello LOTS 82-36,
82-56, 82-63, 82-69
Mtiller (1886)
Aiello LOTS 81-34,
81-78
Riley (1975)
Barcant (1 970)
Riley (1975)
Aiello LOT 8 1-16
Aiello LOTS 83-101,
83-1 1 1
Moss (1933)
Aiello LOTS 81-46,
81-51
cited in Lima (1968)
Janzen 79-SRNP-135
in DeVries (unpub)
IPHICLA IPHICLEOLA (Doxocopa misidentified as Adelpha) Celtis sp. Ulmaceae Mexico Comstock & Vazquez ( 1 960)




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19841 Aiello - Genus Adelpha 21
Table 3. (continued)
A DELPHA SPECIES/ PLANT
FOODPLANT FAMILY LOCALITY REFERENCE
ISIS
Cecropia pachystachia
Cecropia pachystachia
Coussapoa schottii
Coussapoa schottii
Pourouma acutijlora
Pourouma acutiflora
JORDAN1
Vitex trzjlora
JUSTINA JUSTINA
Alternate-leaved plant
LARA
LEUCOPHTHALMA
Pentagonia wendlandia
Undetermined
Undetermined
MELANTHE
Cecropia sp.
Trema micrantha
Trema micrantha
Urera sp.
MELONA
Malanea sp.
MESENTINA
Pourouma sp.
MYTHRA
Bathysa sp.
nr. barbinervis
NAXIA IPIPHILCA
Piper areanum
Vitex cooperi
PARAENA
Isert ia longzjlora
Remijia amazonica
Cecropiaceae Brasil
cited in Lima (1968)
Cecropiaceae Brasil Mtiller (1886)
Cecropiaceae Brasil
cited in Lima (1968)
Cecropiaceae Brasil Mtiller (1886)
Cecropiaceae Brasil
cited in Lima (1968)
Cecropiaceae Brasil Muller (1886)
Verbenaceae Brasil Moss (1933)
9 Panama Aiello 83-23,
83-67
Cecropiaceae Trinidad
M. J. W. Cock
(pers. comm.)
Rubiaceae
Costa Rica Young (1974)
Rubiaceae Panama Aiello LOTS 83-22,
83-24, 83-25
Rubiaceae Costa Rica DeVries (unpub)
Cecropiaceae Costa Rica DeVries (unpub)
Ulmaceae Panama Aiello LOT 83-8
Ulmaceae Costa Rica DeVries (unpub)
Urticaceae Costa Rica DeVries (unpub)
Rubiaceae Brasil Moss (1933)
Cecropiaceae Brasil Moss (1933)
Rubiaceae Brad Mtiller (1886)
Piperaceae Costa Rica Marquis in DeVries (unpub)
Verbenaceae Costa Rica DeVries (unpub)
Rubiaceae Brasil Moss (1933)
Rubiaceae Brasil Moss (1933)




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22 Psyche [VOI. 91
Table 3. (continued)
A DELPHA SPECIES/ PLANT
FOODPLANT FAMILY LOCALITY REFERENCE
nr. PARAENA
Combrefum decandrum Combreta- Panama
ceae
PHLIASSA
Aliberfia edulis Rubiaceae Brasil
Berfiera guianensis Rubiaceae Brasil
PHYLACA AETHALIA
Cecropia obtusifolia Cecropiaceae Panamti Cecropia peltata Cecropiaceae Panama
Cecropia peltata Cecropiaceae Panama
Cecropia sp. Cecropiaceae Panama
Aiello LOT 82-55
Moss (1933)
Moss (1933)
Aiello LOTS 82-39,
82-68, 82-76, 83-78
Aiello LOT 8 1-70
Coley LOT 5
Aiello LOT 82-40
PLESAURE
Bathysa? sp. Rubiaceae
PSEUDOCOCALA
Sabicea aspera Rubiaceae
PSEUDOCOCALA? (as nr. cocala)
Sabicea sp. Rubiaceae
SALMONEUS
Sabicea panamensis Rubiaceae
SERPA
Miconia minutiflora Melasto-
mataceae
Misc. spp. Melasto-
mataceae
SERPA HYAS
Brasil Muller (1886)
Brasil Moss (1933)
Brasil Muller (1886)
Panama Aiello LOTS 82-37,
83- 14
Brad Moss (1933)
Brasil Mtiller (1886)
Ilex paraguariensis Aquifoliaceae Barsil SOPHAX (= zalmona zalmona)
SYMA
Rubus fructicosus Rosaceae Brasil
Rubus?sp. ("bramble") Rosaceae Brasil
Cephalanfhus sarandi Rubiaceae Brad
THESPROTIA?
Alternate-leaved climber ? Brad
TRACTA
Undetermined Rubiaceae Costa Rica
ZALMONA ZALMONA (as sophax)
Sabiceae aspera Rubiaceae Costa Rica
cited in Lima (1968)
Mtiller (1886)
Jones & Moore (1882-3)
cited in Lima (1968)
Moss (1933)
Haber, Chacon in
DeVries (unpub)
DeVries (unpub)




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19841 Aiello - Genus Adelpha 23
reared on Vitex, has been found on only one other plant species (Piper, Piperaceae),
Adelpha pupae (Figures 5 & 6) are diverse in form, yet have several features in common:
(1) The body is slender and tapers towards the cremaster, so that the posterior wing margins protrude and appear as keels. (2) Segments T2 and A2 are expanded dorsally to form two projections of varying shape and size.
The thoracic projection is the smaller of the two and may be pointed and directed posteriorly (A. salmoneus, cytherea, iphicla, justina), or pointed and oriented almost perpendicularly to the body (A. melanthe, phylaca aethalia), or it may be reduced to a slight hump (A. celerio), or smooth curve (A. basiloides). The abdominal projection is the larger and more variable of the two. It may be short, pointed, and directed anteriorly (A. basiloides, justina), a bit larger and curved anteriorly (A. iphicla, celerio, cytherea, cocala), even larger and directed posteriorly (A. sal- moneus), or it may be a huge laterally flattened hook (A. melanthe, phylaca aethalia).
(3) The head usually has a pair of apical projections ("horns") of varying shape and size. These may be long, slender, and slightly curved (A. celerio), sickle-shaped and recurved to the sides (A. Figure 5. Final instar larva and pupa of Adelpha justina justina, (JUS) reared in Panama.




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24 Psyche [vo~. 91
SAL P H
2SG H HASBROUCK
Figure 6.
Pupae of Adelpha species reared in Panama: abbreviations are the same as in Figures 2 & 4; CEL = celario.



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19841 Aiello - Genus Adelpha 25
basiloides), shaped like tiny asymmetrical leaves (A. cocala, leucophthalma), small and triangular, like cat ears (A. iphicla, cytherea, salmoneus), smaller triangles which are bent to the sides (A. justina, phylaca aethalia), or they may be reduced to two tiny rounded projections (A. melanthe).
(4) Pupal color varies from pearly white, through straw-color, brown, green, copper, or shimmering gold or silver. Whatever its color, an Adelpha pupa gives the impressing that it is empty or diseased. The pearly white pupa of A. basiloides is especially deceptive; it appears more to be an abandoned skin than a solid object. The shimmering silver pupae of A. celerio and nr. pamna have black sutures and both give the impression that they are transparent with black lines showing through from the other side. As far as I know, the only other instance of a totally silver pupa occurs in Mechanitis of the Ithomiidae. The pupa of A. salmoneus is coppery brown and resembles a dead or diseased individual. In exploring for possible correlations between pupal and larval types, I began with species whose pupae have a large flat hook-like projection from the dorsum of A2. As it turned out, all have similar larvae, foodplants, and adult male genitalia, yet, the adult wing patterns are diverse. Further analysis revealed several other species groups that show similar correlations. Based upon this work, I have come to the conclusion that the genus Adelpha comprises five to seven natural groups, and possibly more. The only set of characters that doesn't hold together within these groups is adult wing pattern. Some twenty species, for which adequate illustrations are available, are included in the classification which follows, and several others whose immatures were described only, are discussed. The outline and discussion of Adelpha species groups includes brief final instar descriptions for species reared by me in Panama, as well as comments upon previously published accounts of other species belonging to these groups. The MAJOR CHARACTERISTICS listed at the beginning of each group do not function as a key, rather, they represent defining characters for those groups, except as noted in parentheses. FOODPLANTS may host more than one butterfly group: members of GROUPS I and I1 have been reared on plants from



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26 Psyche [vo~. 91
a number of families, GROUP I11 on Verbenaceae, and the remaining GROUPS (IV-VII) on Rubiaceae.
At this stage of our knowledge, an attempt to work out a phylogeny within Adelpha would be premature indeed; more information on life histories of Adelpha and its relatives (especially Old World genera) would be necessary to such an undertaking. *KEY TO ADELPHA SPECIES GROUPS BASED UPON FINAL INSTAR LARVAE, PUPAE. AND FOODPLANTS A. LARVA: body scoli, especially scolus of A2, flattened and feather-like; face with dark stripes. PUPA: shimmering silver with slender, very slightly twisted horns. FOODPLANTS: non-rubiaceous. (A. celerio, sp. nr. celerio, serpa, paraena) A.
LARVA: body scoli not flattened or feather-like; face patterned or plain, but not striped. PUPA: perhaps with some metallic portions, but not totally metallic; head horns, various ... B B. LARVA: scoli of A2-8 of similar form, although those of A2, and/or 7 & 8 may be longests. FOODPLANTS: non- ......................................
rubiaceous C
C. LARVA: pale grey or pale brown with sides of thorax through A2 much darker than rest of body; scolus of A2 long. PUPA: with huge, hook-like dorsal pro- jection from A2; head horns various,. ........... ............... (A. phylaca aethalia, melanthe, isis, mesentina, delphicola, (?)call@ hiclea, (?)abyla) C.
LARVA: dark, or not patterned as above; scolus of A2 short. PUPA: hangs tilted with ventral side superior; head horns short points bent sharply to sides. .... ........................... (A. justina, jordani) B.
LARVA: scolus of A2 different in form from scoli of A3-6 (in A. iphicla the difference is merely the swelling at base of A2 scolus) FOODPLANTS: rubiaceous.. ............ D D. LARVA: scoli of T2 or 3, and A2 & 7 swollen at base; (in A. iphicla, T2 & A2 only); face not patterned; head chalaza-1 not darker than others (don't known for A. pseudococala). PUPA: horns small, triangular "cat- ears" ....................................... E *The reader should bear in mind that in all Adelpha species, the scoli of Tl and A2 are tiny.




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Aiello - Genus Adelpha 27
LARVA: scoli of A2 conical, close together, and with fewer spines at base; scoli of A3-6 very short ..................................... . (A. cytherea, salmoneus, pseudococala) LARVA: scolus of A2 not conical, similar to other scoli except for its swollen base; scoli of A3-6 of moderate length; face smoother than other spe- cies due to reduction or loss of most chalazae (A. iphicla)
..............................
D. LARVA: body scoli about same thickness at middle as at base; scolus of A2 strongly arched towards the posterior, thick and densely spined; face patterned; head chalaza-1 dark, and chalazae-3 & 4 pale com- pared to rest of face. PUPA: head horns curved or recurved to sides ............................ F F, LARVA: scoli of T3 and A7 long, slender, and tapered to sharp point; scoli of A7 & 8 differ in form from each other (that of A8 club-shaped). ............
PUPA: head horns sickle-shaped
(A. basiloides, phliassa)
....................
F.
LARVA: body scoli not tapered to sharp point; scoli of A7 & 8 similar in form. PUPA: head horns like tiny asymmetrical leaves .......... (A. cocala, leucophthalma)
.................
MAJOR CHARACTERISTICS:
(1) Larval body scoli flattened;
(2) Larva with face striped;
(3) Larval head chalaza-1 dark (also true of GROUPS VI & VII);
(4) Larvae assume a curled resting position; (5) Pupa shimmering silver;
(6) Valves of male genitalia without clunicula; FOODPLANTS: Aquifoliaceae, Bombacaceae, Cecropiaceae, Com- bretaceae, Melastomataceae, Rubiaceae, Urticaceae. Two larval types have been observed within this group: A. Final instar larvae with A2 scolus broad, curved posteriorly, and with median spines spreading ....... nr. celerio, serpa.



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28 Psyche [VOI. 91
A. Final instar larvae with A2 scolus longer, slender, straight (although directed posteriorly), and with median spines ascend- ing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . celerio, paraena Pupae vary only slightly, the more apparent differences being in the length, and degree of curvature of the head horns, and in the shape of the dorsal projections on T2 and A2. Head horn shape and size do not appear to correlate with the two larval types seen so far, and perhaps varies even within a species. celerio, Panama (Aiello, Coley)
Larvae are dark, mostly black above and green laterally, with pinkish lateral spots on A2-4, 7 & 8; scoli on thorax and A7 & 8 are pale, while that of A2 is black; subspiracular scoli A2-4 are mixed pale lime and bright green. A black subspiracular stripe passes along the thorax and turns sharply upward across A1 and fuses with the dark scolus of A2. A few days before pupation the pinkish lateral marks change to green, as do the scoli of T3 and A7 & 8; the scolus of A2 remains dark. Just prior to pupation larvae fade to a dirty yellow. Larvae rest in the curled position with the posterior end elevated and the white undersurface of A8 & 9 exposed, and in this configuration resemble a bit of fallen moss and lichen mingled,
Pupal head horns are long, and slightly curved near their tips. Adults resemble iphicla in wing pattern. nr, celerio, Panama (Aiello)
A larva resembling celerio produced, but with the A2 scolus short and broad, a pupa that had short head horns and appeared identical to my nr. paragna.
Unfortunately the pupa died and its identity remains a mystery.
celerio diademata, Mexico (Comstock & Velazquez, 1960) The illustrations of this larva and its pupa indicate member- ship in GROUP I; the larva is clearly of the celerio type having long narrow scoli, even on A2. The pupa closely resembles that of the celerio reared by me in Panama; the head horns are long and slightly curved just before their tips.
The authors report that their larva faded to yellow just before pupation.




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19841 Aiello - Genus Adelpha 29
paraha, Brasil (Moss, 1933)
From its illustration, the larva of this species is of the slender scolus type. Moss noted that it was brighter green than serpa and had none of the orange spines. The record of Rubiaceae for paraena is unique for this group.
nr. paraena, Panama (Aiello)
In pattern, this larva is very similar to A. celerio, but is paler above, being mottled brown and black, with a white dorsal patch joining A6 & 7. The scoli are brown, that of A2 being the darkest; subspiracular scoli A2-4 are pink. The larva turned yellowish just prior to pupation.
The pupal head horns are shorter than those of celerio, and curve outward at about their midpoint.
Adults resemble celerio so closely that I did not realize they were different species until rearing them, and doubtless they are mingled in many collections.
serpa, Brasil (Moss, 1933)
The larva of serpa has the scolus of A2 shorter and denser than the other major scoli, and I take it to be of the same general type as celerio. Moss described the scolus of A2 (segment 6 in his system) as being bushy and dark green, and the other major scoli as ochreous (orange). His larva rested with the anterior portion turned, and resembled a bird dropping.
serpa, Brasil (Muller, 1886)
Muller's illustrations of serpa show the larval face, and a scolus from A2. The face is clearly striped, and the scolus is short, thick, and has flattened spines overlapping each other; each of these conditions of the scolus is more extreme than in nr. celerio. MAJOR CHARACTERISTICS:
(1) Larva pale, with sides of thorax through A2 darker than rest of body;
(2) Pupa with huge, laterally flattened, hook-like projection from dorsum of A2;
(3) Pupal head horns diverse: leaf-shaped, bent triangular, rounded;




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30 Psyche [vo~. 91
(4) Valves of male genitalia armed with row of teeth beginning at apex and extending along lower edge; clunicula broad; FOODPLANTS: Aquifoliaceae, Bombacaceae, Cecropiaceae, Ulrna- ceae, Urticaceae.
abyla, Jamaica (Swainson, 190 1)
There are several things in Swainson's description that lead me to place abyla tentatively with GROUP 11. "On first segments are two brown horns with sharp points bending over the head; from this to the end of the third segment is silvery grey; then two tiny horns bending backward." "The [pupa] shape is very curious, resembling the pictures of 'Punch,' long nose and all." It appears that larval segments Tl and A1 were overlooked, probably because of their small size and tiny scoli. The silver area would then be bounded by scoli of T2 and A2, suggesting the two-toned larva of GROUP 11. The pupal description may be in reference to the long abdominal hook of GROUP 11, or it may be an over- reaction to the shorter dorsal projection of a species in some other group (iphicla perhaps). Unfortunately, no foodplant is mentioned.
calliphiclea, Paraguay (Jorgenson, 192 1) Jorgensen's description of the pupa as having the abdomen prolonged toward the thorax, forming a large hook, induces me to place calliphiclea in GROUP 11: "El torax arriba con una carena alta como el abdomen en el dorso este ultimo hacia el torax prolongado, formando un gran gancho." In addition, the non-rubiaceous foodplant (Ilex) is consistent with this group. Each of the five species remaining, possess characteristics 1 & 2 above, and there is no doubt in my mind that they are all close relatives. Pupal head horn shape may have taxonomic value within the group but that cannot be determined until a larger number of species has been reared.
delphicola, Brasil (Moss, 1933)
The pupal head horns curve to the sides and resemble tiny asymmetrical leaves; the hook on dorsal A2 is rounded at the tip. isis, Brasil (Muller, 1886).
Muller's illustration of the pupa is a lateral view so the head horn shape does not show, however, it is stated in the text that



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19841 Aiello - Genus Adelpha 31
the horns are small points bent to the sides. The hook on A2 is enlarged slightly at the tip much as in melanthe. melanthe, Panama (Aiello)
The larva is grey (darker above) excepting the dorsal thorax through A2, which is dirty yellow, and the sides of that same area, which are dark brown. The scoli are grey with black spine bases and apices. The larva has a frosted appearance partly due to its patterned scoli, but mostly because the entire body is densely clothed with short thick grey setae. It is the only Adelpha species so far that has rounded pupal head horns. The hook on A2 is slightly expanded at the tip.
Adults of melanthe look like large salmoneus but the undersurface is quite different.
mesentina, Brasil (Moss, 1933)
From Moss's illustrations, mesentina and delphicola larvae are similar, and no doubt these two species are close relatives. As a result of their close resemblance, Moss did not realize that he had two species until the adults emerged. If his illustrations are accurate, then mesentina has a less rounded pupal hook than delphicola, but the head horns are nearly the same. phylaca aethalia, Panama (Aiello, Coley) The larva is greenish or pinkish grey, and speckled with darker grey (especially on the dorsum); the sides of the thorax through A2 are dark brown, and some individuals have a dark brown oval on each side of A5, just below the scolus. Pupal head horns are short triangles bent to the sides.
Adults are very close look-alikes of cocala. MAJOR CHARACTERISTICS:
(1) Larval scoli A2-6 all similar and short; (2) Pupa tilted so that ventral side is superior (at least in just ina);
(3) Pupal head horns small triangles bent to sides (also occurs in GROUP 11);
(4) Valves of male genitalia unarmed (also true of GROUP IV);




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32 Psyche
[Vol. 91
FOODPLANTS: Verbenaceae.
The most interesting thing about this group is the fact that the scolus of A2 is just the same as those of A3-6. Both species, thus far included, rest in the front-curved position, as shown in Moss's illustration of jordani. Also in both, the pupal dorsal A2 projection is a short point, directed anteriorly, and the T2 projection is smaller.
This and GROUP IV have similar genitalia, but feed on different foodplant groups. I see them as close but distinct relatives. jordani, Brasil (Moss, 1933)
The larva is dull green with black-spined, orange scoli; the pupa is white with black spots.
justina, Panama (Aiello & Small)
Early final instar larvae are dark brown (lighter dorsally) with paler scoli (black at tips of those on T2 and A7 & 8), but later change to black and white checkered with transverse orange bands joining members of scolus pairs (A2-8). The straw-colored pupa has a dorsal metallic sheen, from head through Al, and lacks the prominent spotting of jordani. Adults of justina closely resemble leucophthalma and the two can be found flying together.
MAJOR CHARACTERISTICS:
(1) Larva with scoli of A2 conical, close together, and with fewer spines at base;
(2) Larval face with chalazae reduced in size; (3) Pupal head horns of this and Group V, are small, triangular "cat-ears;"
(4) Valves of male genitalia unarmed (also true of GROUP 111);
FOODPLANTS: Sabicea (Rubiaceae).
nr. cocala, Brasil (Mliller, 1886)
The larva illustrated does not at all resemble that of cocala, so I guess Muller's label "bei cocala" to have been based upon adult appearance. Like cytherea, this larva shows the conical scolus of A2, the thick based scoli of T2, and A7, and the very short scoli of A3-6. As well, Mliller's larva fed on Sabicea.



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19841 Aiello - Genus Adelpha
cytherea, Brasil (Moss, 1933)
Of this larva, Moss commented, "Now in two shades of brown, on the stem or near flower-head easily mistaken for a part of the plant. . . . "
cytherea marcia, Panama (Aiello)
The larva is patterned dark and light brown; with dark brown, oblique lateral stripes, which cross segments A3-8. A thin white subspiracular line expands into a lime-green mark on A7-9. The pupa is straw-color but takes on a silver or gold sheen at certain angles.
pseudococala, Brasil (Moss, 1933)
Moss states merely that the larva of this species is identical to cytherea, although larger and greener at the ends. The pupal abdominal projection is directed towards the posterior as in salmoneus.
salmoneus, Panama (Aiello)
The dark brown second through fourth instar larvae, with their grey, oblique, lateral stripes, are well camouflaged amongst the dark, pale-veined, young leaves of their foodplant. Fourth and fifth instars have red-brown heads.
Early final instar larvae are brownish green, and become greener with age. By late final instar, larvae are bright yellow- green; the dorsal area of A4 & 5 is pale purple-brown; A4 & 5 each bear a dark lateral purple mark; the head is red-brown; scoli of T2 & 3, and A2 & 7 are brown-purple, while that of A8 is still green.
The pupa is bronze with dorsal gold, and has an A2 projection that is prolonged more towards the posterior end than the anterior.
MAJOR CHARACTERISTICS:
(1) Larval scoli of T2 and A2 swollen at base, otherwise, all body scoli usually similar in form;
(2) Pupal head horns of this and GROUP IV, are small, triangular "cat-ears";
(3) Larval head uniformly dark, and face smooth except for usual pits, and two pairs of reduced chalazae (nos. 2 & 4);



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34 Psyche [vo~. 91
(4) Valves of male genitalia armed with apical row of teeth which extend to outside of valve;
FOODPLANTS: Antirrhoea, Alseis, Bathysa, Calycophyllum, Gon- zalea, Zsertia, Rondeletia, Uncaria (Rubiaceae). iphicla, Panama (Aiello)
The color of iphicla larvae varies from dark grey, through golden brown, and red-brown, to almost black, regardless of the foodplant eaten. The head is always smooth (by Adelpha standards) and also varies, from yellow-brown with black punctations to uniform dark brown or black. Final instar larvae show the most pattern (especially light-colored individuals): dark, oblique lateral stripes on A2-5, which terminate part way up the bases of the scoli, and often but not always, white lateral marks on A2, 7 & 8.
The pupa varies from waxy white to straw-color or pale brown, and often is partly or entirely burnished gold or silver, especially on dorsal Tl & 3, and Al. Almost always there is a small silver diamond located on the base of each mesothoracic leg.
iphicla, Brasil (Moss, 1933)
Moss described iphicla larvae as "very similar to cytherea, the two species evidently being closely related, as is also shown by the pupae."
GROUPS IV and V may be related; the "cat-eared" pupae, and oblique-striped larvae of both, seem to hint at that. As well, both groups of larvae display varying degrees of reduction of head chalaze, and swelling of certain scoli bases. However, the genitalia of the two groups are different, and, in iphicla the scoli on A3-6 are longer and the face smoother than in Group IV. And so, for the moment they will be kept separate, but near to each other.
GROUP VI & VII
MAJOR CHARACTERISTICS:
(1) Larva with scolus of A2 arched towards the posterior, thick, and densely spined;
(2) Larval face patterned;
(3) Larval head chalza-1 dark (also true of GROUP I);



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19841 Aiello - Genus Adelpha 35
(4) Valves of male genitalia armed with apical row of 2-6 fairly uniform teeth;
(5) Larval head chalazae 3 & 4 pale, and contrasting with face. FOODPLANTS: Ru biaceae.
These two groups are very closely related and perhaps will be merged when more is known of their larval and pupal diversity. For now they are placed apart on account of the specialized scoli in basiloides, and differences in the form of the pupal head horns. ADDITIONAL CHARACTERISTICS:
(5) Larval scoli of T3 and A7 long, slender, and tapered to sharp point;
(6) Larval scolus of A8 club-shaped;
(7) Larval face pattern, independent of pits; (8) Pupal head horns sickle-shaped;
FOODPLANTS: Alibertia, Amaioua, Bertiera, Ixora (all Rubiaceae). basiloides, Panama (Aiello)
Larvae are mottled black and brown, and are paler towards the posterior end. The dark and pale portions intersect obliquely along a line beginning at the subspiracular scolus of A4, and terminating dorsally at the beginning of segment A7. The dorsal dark portion thus comes to a point at the beginning of A7. As larvae approach pupation, they may become tinged with rose or green, especially at scoli bases, and laterally on A4-6. The pupa is pearly white with black tiped head horns, and appears empty. Head horns vary somewhat in length and degree of curvature, even among larvae collected on the same plant. Both extremes are illustrated (Figure 6). phliassa, Brasil (Moss, 1933)
Comparing my observations of basiloides with Moss's descrip- tion and illustrations of phliassa, it appears that the two are extremely close relatives.
plesaure, Brasil (Muller, 1886)
The pupa illustrated appears identical to those of phliassa and basiloides. Hall (1 938) feels that plesaure and phliassa are one and the same species.




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GROUP VII
ADDITIONAL CHARACTERISTICS:
(4) Larval face pattern due to dark pits against a paler face; (5) Pupal head horns shaped like tiny asymmetrical leaves (also occurs in GROUP 11);
FOODPLANTS: Calycophyllum, Chomelia, Malanea, Pentagonia, Psychotria, Uncaria, Warscewiczia (all Rubiaceae). cocala, Panama (Aiello)
Early fifth instars are crypticaly patterned with golden brown and black; later the pattern remains but the colors become moss- green, black, and cream, and a pinkish grey and black area appears on dorsal A3-6, plus a broad, oblique, lateral pinkish or yellowish stripe across A4 and 5 together. Some individuals have a subspiracular lime-green mark on each of A7 & 8. The pupa is dark green.
cocala, Brasil (Moss, 1933)
The curved, thick scoli of A2 show clearly in Moss's illustration, but his larvae had more than one lateral oblique stripe. The pupae appear nearly identical. His record of Emmotum (Icacinaceae) as a foodplant, in addition to Malanea (Rubiaceae) is odd, and is the only reported deviation, by cocala, from Rubiaceae.
leucophthalma leucophthalma, Panama (Aiello & Small) The larva of leucophthalma is apparently indistinguishable from that of cocala (G. Small, personal communication) and the two species must be closely related.
The pupa also is very similar to cocala, but differs in being brown or copper-color, and in having the abdominal projection rather square, the thoracic projection more pointed, and the head horns farther appart at their bases.
leucophthalma leucophthalma (as I. tegeata), Costa Rica (Young, 1974)
The observations by Young are consistent with those of Gordon Small and myself.
erotia, Brasil (MUller, 1886)
An isolated scolus, figured by Muller and labelled erotia, would



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19841 Aiello - Genus Adelpha 37
place that species in GROUP I, but I would like to see more evidence. In addition, the erotia wing-undersurface pattern does not have the sharply stamped appearance typical of GROUP I, and it is possible that the butterly was misidentified. iphicla iphicleola, Mexico (Comstock & Vazquez, 1960) This record was in error; the authors misidentified Doxocopa as Adelpha.
rnelona, Brasil (Moss, 1933)
The larval description and illustrations in Moss do not give sufficient detail to place this species, although from the pupa it belongs in GROUP VII.
thesprotia, Brasil (Moss, 1933)
The larval description and illustration in Moss do not give sufficient detail to place this species, although I suspect that it may represent an eighth group. As well, there is a good deal of disagreement concerning the identity of Moss's thesprotia; Forbes (unpublished) believed that Moss actually had nea, a species which on the basis of genitalia probably belongs to GROUP I. Moss's statement that the larva rested in a curled position (typical of GROUP I), lends support to Forbe's suspicion. However, from what can be seen of the scoli in Moss's illustration, they are all similar in size and form, are not flat, and the pupa looks more like those of GROUP VIII. When adult specimens of Adelpha are sorted into the species groups just outlined, the result is a jumble of wing pattern types that would make any biologist wince, this author included. However, it would seem even less natural to split groups that are based upon the presumably more conservative characters of the immatures. I prefer to think that in Adelpha, wing pattern not only does not reflect species relationships, but instead may be intended to deceive. One might expect adult wing characters to be the most specialized and difficult to interpret; adults move about with ease and interact with each other and with other butterfly species, as well as with potential predators, while in most cases the immature stages must contend, in



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38 Psyche [vo~. 91
a less active way, with predators and parasites alone. An explana- tion of Adelpha's wing pattern madness, might be that this genus is comprised of several groups, of various affinities, which together form a large mimicry complex, perhaps based upon members whose larvae feed on alkaloid-bearing plants (e.g., Rubiaceae). Possible examples are found among the unrealted look-alikes reared by me in Panama (Figure 7): phylaca aethalia (non-rubiaceous feeder (NR) ) closely resembles cocala (rubiaceous feeder (R) ); justina (NR) resembles leucophthalma (R); melanthe (NR) is larger than, but very similar in pattern to salmoneous (R); and celerio and nr. paraena (both NR) are very similar in pattern to iphicia and basiloides (both R). The idea is intriguing, but must be labelled "pure speculation" until data on palatability of Adelpha butterflies is available. In its defense, however, I would like to point out that there are several examples of Adelpha look-alikes in uncontestably unrelated butterflies. Nymula velabrum (Riodinidae), whose wing pattern closely resembles that of A. iphicla, and several other Nymula species that are look-alikes of other Adelpha types, all depart from the basic wing pattern of their genus. Female Doxocopa laure and pavon (Nymphalidae) resemble A. iphicla,
while female D. clothilda are even more convincing look-alikes of A. salmoneus. Male Doxocopa clothilda andpavon differ from the females of their species, and do not resemble Adelpha, while the male of D. laure has an Adelpha-like wing pattern with overlying purple iridescence. Pyrrhogyra hypsenor (Nymphalidae) is similar in appearance to Adelpha species that have broad white bands, and has fooled at least one lepidopterist (Muyshondt, 1974). The few published accounts of life histories for Old World Limenitidinae provide intriguing glimpses into the trove of informa- tion awaiting the attention of lepidopterists. Many larval and pupal forms, similar to those of Adelpha, occur among Old World genera, and in addition, many utilize the same foodplant genera as Adelpha and display similar larval behavior. Life history studies, that compare Old and New World groups, are essential to our understanding of generic limits and relationships within this subfamily.




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19841 Aiello - Genus Adelpha 39
6Morrell (1954) gives a fascinating account of larval behavior in several Malayan genera of the tribes Limenitidini (Athyma, Moduza, Pandita), and 'Neptini (Lasippa, Neptis, Phaedyma). Except for Neptis leucoporos, the larvae, of the butterflies he observed, expose the midrib, or a secondary leaf vein, by feeding around it (just as Adelpha does); young larvae rest out on these slender supports. Like Adelpha also, Athyma, Moduza, and Pandita accumulate their feces to form a mass at the base of the support vein; Neptini do not.
Judging from Morrell's illustrations, the larva of Moduza resembles a combination of Adelpha cocala and basiloides, es- pecially in having short thick scoli on A2; its pupa is remarkably like A. cocala or leucophthalma, but with slightly stalked head horns; the host plants are mainly Rubiaceae (Mussaenda, Nauclea, Timonias, Uncaria, Wendlandia), with one record on Oleaceae (Olea).
The larvae of Athyma kanwa feeds on Uncaria, and is described by Morrell (1960) as "brown with delicate green branched spines, and resembles a growth of moss on a decaying patch of leaf;" the pupa (illustrated in Morrell, 1954) is silver with long head horns and to me looks for all the world like a pupa of Adelpha celerio. In form, Athyma nefte larvae (Morrell, 1954) are reminiscent of a very plump Adelpha iphicla larva, and have been reported on Glochidion (Euphorbiaceae) and Mussaenda (Rubiaceae); the pupa is "golden brown in colour, with two dorsal plates of brilliant metallic gold," and from what can be determined from its picture, it looks like the pupa of Adelpha iphicia or cytherea, but with an exagerated A2 projection, and like those species it appears to have short pointed head horns.
Pandita sinope is described by Morrell (1954) as having a larva similar to that of Athyma nefte in color and general appearance; the illustration of the pupa is also similar to A. nefte, but has a less according to Corbet and Pendlebury (1978), five of the seven names used by Morrell (1954) should be amended: his Neptis columella = Phaedyma columella; Neptis heliodore = Lasippa tiga; Neptis nata = Neptis leucoporos; Parathyma kanawa = Athyma kanwa; Parathyma nefte = Athyma nefte. His Moduza procris and Pandita sinope remain unchanged.
'Neptini is defined by Eliot (1969) to include Aldania, Lusippa, Neptis, Pantoporia, and Phaedyma.




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42 Psyche [vo~. 91
exagerated A2 projection and thus appears almost identical to pupae of Adelpha iphicla and cytherea. Larvae of P. sinope feed on Uncaria (Rubiaceae).
The valves of the male genitalia in Pandita sinope and Athyma nefte are unarmed, have cluniculae, and look very similar to those of Adelpha GROUPS 111 & IV.
A number of other life history accounts for Old World Limenitidini have been published, but it is not the intent of this paper to report them in detail. Instead, I merely wish to point out a few of the intriguing similarities that exist between Old and New World groups, in hope of prompting other workers to investigate this promising group.
Many people contributed to this project, and I am indebted to each of them for their enthusiastic help and support. I would especially like to thank Gordon Small for his encouragement during this project, as well as for generous logistical help, identification of butterflies, and for the many valuable larvae which he has brought to me. Richard Vane-Wright of the British Museum provided photographs of a number of Adelpha type specimens, as well as helpful comments about them; Adelpha celerio andphylaca aethalia were identified by comparison with these photographs. Robert Diaz, James Mallet, and Henry Stockwell also located caterpillars for this project, and Bob Diaz and Dagmar Werner contributed their time and effort to translate several German papers. Lissy Coley made specimens, from her own project, available for study. I would like to thank Ricardo Cortez for logistical and technical help, and Robert Robbins and Gordon Small for reading the manuscript. This paper would be deficient without the carefully executed drawings of Marshall Hasbrouck, which were made using a combination of live and preserved material plus photographs, or without the foodplant list compiled by Philip DeVries. Phil has made this list available for use by other lepidopterists, in spite of the fact that it is as yet unpublished. Robert Silberglied gave encourage- ment in the earliest stages of this project. I would like to thank the Image Systems Branch, Tropic Test Center, Corozal, Panama, for the photographs (Figure 7) of adult specimens. Without the facilities and library of the Smithsonian Tropical Research Institute,



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19841 Aiello - Genus Adelpha 43
Panama, and the support of National Geographic Society Grant 2444-82, this work would not have been possible. AIELLO, A.
(Unpub)
Insect Rearing Records (mainly for Lepidoptera). Each year is in a separate notebook; lot numbers used in the present paper refer to pages in these notebooks.
BARCANT, M.
1970. Butterflies of Trinidad and Tobago. 314 pp.; Collins. London. COLEY, L.
(Unpub) L. Coley has reared a number of Lepidoptera on Ochroma and Cecropia. Numbers given are her identification numbers. COMSTOCK, J. A. AND L. VAZQUEZ G.
1960.
(1961)
Estudios de 10s ciclos biol6gicos en Lepid6pteros Mexicanos. AN INST BIOL MEXICO 31: 349-488.
CORBET, A. S., AND H. M. PENDLEBURY
1978. The Butterflies of the Malay Peninsula. 3rd edition revised by J. N. Eliot. xiv + 578 pp., 35 pi., 438 figs. Malayan Nature Society. DEVRIES, P. J.
(Unpub)
List of host plants of butterflies of Costa Rica, compiled from his own data plus that of a dozen other workers. The last compilation was prepared during early 1982.
DEWITZ, H.
1879. Naturgeschichte Cubanischer Schmetterlinge. In: Zeitschrift fur die gesammten Naturwissenschaften. Bd. 52: 166- 167. DORNFELD, E. J.
1980. The Butterflies of Oregon. xiv + 276 pp.; Timber Press. Forest Grove, Oregon.
ELIOT, J. N.
1969. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). 155 pp., 3 pi., 101 text-figs. Supplement 15 to BULL BRIT MUS.
ELTRINGHAM, H.
1923.
Butterfly Lore. 180 pp.; Oxford, at the Claredon Press. FERRIS, C. D., AND F. MARTIN BROWN (editors) 1980.
Butterflies of the Rocky Mountain States. xviii + 442 pp.; University of Oklahoma Press. Norman.
FRUHSTORFER, H.
1907. Adelpha. pp. 510-533 + pis. 106-1lOa. In: SEITZ, A. The Macro- lepidoptera of the World. Div. 11, Vol. 5, viii + 1139 pp. GODMAN, F. D., AND 0. SALVIN
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190 1.
Biologia Centrali-Americana. Insecta. Lepidoptera-Rhopalcera. Vol. I, xlv + 487 pp. (1879-1901); Vol. 11, 782 pp. (1887-1901).



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HALL, A.
1938.
On the types of Adelpha (Lep., Nymphalidae) in the Collection of the British Museum. ENTOMOLOGIST 71: 184-187,208-211,232-235.257-259, 284-285.
HAYWARD, K. J.
1950. Las especies y formas Argentinas del gfenero Adelpha. ACTA ZOOL LILLOANA 9: 375-393.
HOFFMANN, F.
1937.
Beitriige zur Naturgeschichte brasilianischer Schmetterlinge. 11. ENTO- MOLOGISCH ZEITSCRIFT 51: 21 2-213, 23 1-232. HOWE, W. H.
1975. The Butterflies of North America. xiii + 633 pp., 97 color plates containing 2093 illus. and 32 text figs. Doubleday & Company, Inc. Garden City, New York.
JONES, E. D., AND F. MOORE
1883. Metamorphoses of Brazilian Lepidoptera from San Paulo, Brazil. Second Series. PROC LIT PHILOS soc LIVERPOOL 37: 229-259. JORGENSEN, P.
1921.
Sobre algunos nuevos enemigos de la yerba-mate, Ilex paraguariensis. REV soc CIENTIFICA PARAGUAY l(1): 27-30. KIRKBRIDE, A. C. G.
1982.
A preliminary phylogeny for the neotropical Rubiaceae. PL SYST EVOL 141: 115-122.
MILLER, L. D., AND J. Y. MILLER
1970.
Notes on two rare Mexican Adelpha and related Central American species (Nymphalidae). J LEP soc 24(4): 292-297. MORRELL, R.
1954. Notes on the larval habits of a group of nymphalid butterflies. MALAYAN NAT J 8: 157-164.
1960. Common Malayan Butterflies. Malayan Nature Handbooks. xii + 64 pp., 20 pis. Longman. London,
Moss, A. M.
1933. Some generalizations on Adelpha, a neotropical genus of nymphalid butterflies of the group Limenitidi. NOV ZOOL 39: 12-20 + pis. 1 & 2. M~LLER, W.
1886. Sudamerikanische Nymphalidenraupen, versuch eines naturlichen Sys- tems der Nymphaliden. ZOOL JAHRB, ZEIT SYST, GEOGR BIOL THIERE 1: 417-678 + pi. 12-15.
MUYSHONDT, A.
1974.
Notes on the life cycle and natural history of butterflies of El Salvador. V. A. Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae). J NY ENT soc 82(3): 163-172.
ORSAK, L. J.
1977. The Butterflies of Orange County, California. xii + 349 pp.; University of California, Irvine. Center for Pathobiology MISC PUBL No. 3; Museum of Systematic Biology RES SER No. 4.




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PETERSON, A.
1962. Larvae of Insects: An Introduction to Nearctic Species. Part I. Lepidoptera and Plant Infesting Hymenoptera. Columbus, Ohio. RILEY, N. D.
1975.
A Field Guide to the Butterflies of the West Indies. 224 pp. + 338 color illus.; Collins. London.
SCUDDER, S. H.
1889. Butterflies their Structure, Changes and Life-histories with special reference to American Forms. viii + 322 pp.; Henry Holt & Co. New York.
SILVA, A. G. D'A. E., C. R. GONCALVES, D. M. GALVAO, A. J. L. GONCALVES, J. GOMES, M. DO N. SILVA, AND L. DE. SIMONI. 1967-
1968.
Quartro Catalogo dos insetos que vivem nas plantas do Brasil, sues parasites e predadores. parte I (2 vol., 906 pp.) & parte I1 (2 vol., 887 pp.). Ministerio da Agricultura. Rio de Janeiro, GB, Brasil. SMALL, G.
(Unpub)
List of Panamanian Butterflies. This list is based upon twenty years of observations in Panama.
SWAINSON, E. M.
1901.
Notes on lepidopterous larvae from Jamaica, B.W.1. J NY ENT soc 9: 77-82.
YOUNG, A. M.
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Notes on the natural history of a rare Adelpha butterfly (Lepidoptera: Nymphalidae) in Costa Rica high country. J NY ENT soc 82(4): 235-244.



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