Robert E. Silberglied, Annette Aiello, and Gerardo Lamas.
Neotropical Butterflies of the Genus Anartia: Systematics, Life Histories, and General Biology (Lepidoptera: Nymphalidae).
Psyche 86:219-260, 1979.

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NEOTROPICAL BUTTERFLIES OF THE GENUS ANARTIA: SYSTEMATICS, LIFE HISTORIES AND GENERAL
BIOLOGY (LEPIDOPTERA: NYMPHALIDAE)
BY ROBERT E. SILBERGLIED~
ANNETTE AIELLO~
AND
GERARDO LAMA$
INTRODUCTION
Butterflies of the genus Anemia Hiibner are among the most common and conspicuous diurnal Lepidoptera encountered in the New World tropics. While their abundance and ease of capture have made them popular subjects for research in various aspects of lepi- dopteran biology, the genus has never been thoroughly reviewed or revised. Two of the authors, (R.E.S. and A.A.) have been conduct- ing genetic, behavioral and ecological experiments on members of this genus for four years, and we feel it is both an opportunity and a necessity to condense the scattered published information with some of our own observations and results. Our experimental findings will be published separately.
As treated here, Anartia consists of five species (Figure 1) in three well-defined groups (Godman and Salvin, 1882). SYSTEMATICS
Genus Anartia Hiibner
Anartia Hubner, [1819]: 33.
Type species, Papilio jatrophae Linnaeus (Scudder, 1875: 11 1). Celaena Doubleday, [1849]: 214.
Type species, Papilio fatima Fabricius (Hemming, 1941: 425). Invalid and unavailable; published in synonymy (ICZN, Art. l id). Celoena Boisduval, 1870: 38.
Type species, Papilio fatima Godart (mon.). Junior subjective synonym. Anartia subgenus Anartiella Fruhstorfer, 1907: 112. Type species, Vanessa lytrea Godart (mon.). Junior subjective synonym. Smithsonian Tropical Research Institute, Box 2072, Balboa, Republics de Panama. Present address: Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138
Wuseo de Historia Natural "Javier Prado," Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Aptdo. 1109, Lima, Peru. Manuscript received by the editor February 26, 1980.



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220 Psyche [June-September
Medium-sized, lightly built butterflies. Head small; eyes hemi- spherical, prominent; labial palpi elongate, curved upwards and densely scaled, with first and third segments about 114 to 113 the length of the second (Reuter, 1896, fully describes the palpi); anten- nae slender, slightly shorter than the body in length, with slightly flattened, pointed, nine-segmented club; thorax strong, thinly scaled; forewing slightly angled apically, anterior margin curved at base, apex slightly truncate, outer margin sinuate, inner margin straight, Sc-R system variable among the species (Figure 2; Double- day, 1849; Schatz, [1887]; Godman and Salvin, 1882), costal cell open; hindwing somewhat quadrate, longer than wide, produced distally into a slight tail at vein M3, veins Rs, M 1 and M2 diverging nearly from the same point, costal cell open; prothoracic legs of male thin, tibia longer than the femur, a single tarsomere half the length of the tibia, clothed in fine white setae;prothoracic legs of female thicker than those of the male, tibia shorter than the femur, five tarsomeres, together nearly equalling the tibia in length, each tarsomere bearing stout spines, especially the apical one; meso- and metathoracic legs long, femora shorter than tibiae, tibiae and four basal tarsomeres spiny, claws moderately curved; abdomen equal in length to head and thorax combined; male genitalia (Figure 3) with a bifid, curved uncus and simple valves. Chromosome number, n=31 (A. amathea, A. fatima and A. jatrophae; Maeki and Reming- ton, 1961; Wesley and Emmel, 1975).
Scudder (1 893) suggests that the generic name is derived from the Greek for "incongruous; in allusion to its great difference in mark- ing from its fellows." Glaser (1 887) states that Anartia is a "prince of the caste of the children of the sun" (Indian mythology). The only common name used for the genus as a whole is "the American Peacocks" (Brown and Heineman, 1972).
Anartia amathea and A. fatima
Anartia amathea (Linnaeus)
amathea (Linnaeus), 1758: 478 (Papilio). Type locality: ["Indiis."]
[amalthea (Clerck), 1764: pi. 40, fig. 3. Emendation; see below.] amalthea (Cramer), 1780: 29, 173, pi. 209, fig. A, B. Unjustified emendation; see below.
roeselia (Eschscholtz), 1821: 207, pi. 5, fig. 9 (Cynthia). Type locality: bbBrasil."




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19791 Silberglied, Aiello & Lamas - Genus Anartia 22 1 silvae Burmeister, 1861: 168.
New synonymy.
Type locality: Argentina, Tucuman, Manantial de Marlopa. amathea subspecies sticheli Fruhstorfer, 1907: 101. Type locality: Bolivia, "5 days north of Cochabamba." amathea subspecies thyamis Fruhstorfer, 1907: 102. Type locality: Brasil, S5o Paulo; Santa Catarina; Rio Grande [do Sul]; 'Paraguay."
[arnathea subspecies roeselia aberration "conjuncta" Zikan, 1937: 387. Type locality: Brasil, Minas Gerais, Passa Quatro, 900 m.] Linnaeus may have committed a 'lapsus calami,' or mistransliterated amathea from Greek to Latin. The etymologically correct spelling is amalthea, for the goat that nursed Jupiter. (The reddish, innermost satellite of the planet Jupiter is also named Amalthea.) Clerck's emendation was followed by Cramer, and used by many others since then, but A. amathea must stand as the nomenclaturally cor- rect name, since there is no "clear evidence of an inadvertent error" by Linnaeus (ICZN, 32 (a) (ii)), and no "demonstrably intentional change in the original spelling" by Clerck (ICZN, 33 (a)). Clerck's names have no standing in nomencla- ture, as he did not use the binomial system. The etymology of amathea is dis- cussed in greater detail by Fruhstorfer (1907). Anartia fatima (Godart)
fatima (Godart), [I 8241: 375 (Nymphalis). Type locality: "des Indes." Suggested replacement for fatima Fabricius. See below.
faiima Fabricius, 1793: 8 1 (Papilio).
Type locality: "Indiis." Junior homonym of Papilio fatima Cramer, 1780. Application for suppression of this name has been forwarded to the ICZN. See below.
fatima subspecies venusta Fruhstorfer, 1907: 1 1 1. Type locality: "Mexico," "Guatemala."
moreno Kruck, 1931: 234, fig. 1.
Type locality: Mexico, Oaxaca. Aberration. fatima form albifasciata Hoffman, 1940: 28 1. Type locality: "Mexico".
[fatirna aberration "albifusa" Hoffmann, 1940: 281, fig. 6, 7. Type locality: Mexico, Veracruz, Tierra Blanca.] [fatima subspecies venusta form 'bcolimensis" Hoffmann, 1940: 283, fig. 5b. Type locality: Mexico, Colima; [Michoacan], Rio Balsas.] [fatirna aberration "oscurata" [sic] Maza, 1976: 103, fig. 1. Type locality: Mexico, Veracruz, Cerro El Vigia.] [fatima mirus Martin, 1923: 54.
Type locality: Paraguay. Nomen nudum.]
If we were to follow strictly the rules of zoological nomenclature, A. fatima (Fabricius), as a junior primary homonym of the riodinid Emesis fatima (Cra- mer), would be invalid, since they were both described in the genus Papilio. However, considering the large amount of biological information published on this species, it would be in the best interest of a stable nomenclature if the specific epithet could be conserved. Accordingly, we have applied to the International



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222 Psyche [June-September
Commission on Zoological Nomenclature, for conservation of the name fatima, by recommending that the use of the name fatima Fabricius be suppressed until 1824, when Godart transferred the species to the genus Nymphalis. This would not affect the nomenclature of the riodinid, and would have the advantage of keeping the name fatima in use for what is certainly the most familiar Central American butterfly. The synonymy presented here reflects this recommendation. The name fatima was first used in Anartia in 1837, by Geyer, in Hiibner [1824-]1825[- 18371 (see Hemming, 1937, p. 479). Anartia amathea and A. fatima (Figure 1) are a pair of very closely-related species, restricted to the tropical and subtropical mainland of Latin America, including Trinidad and offshore islets. The ranges of these species abut in eastern Panama (Darih); hybrids between them have been collected in the field at the juncture of their distributions on several occasions (e.g., Brown, 1975). Inten- sive study of the mortality and development of Fl hybrids (Figure 4) and their offspring reveals strong hybrid breakdown, and behav- ioral research on courtship and mating preferences reveals a com- plex picture of assortative mating. These results and their evolution- ary consequences will be reported elsewhere; we here want to emphasize that we interpret amathea and fatima as biologically separate species.
The wing venation, male genitalia and larvae of amathea and fatima are, so far as we have been able to tell, identical. The wing venation (Figure 2) differs from that of other members of the genus by the two small veins that leave the Sc-R complex and branch towards the costa in the forewings. The valvae of the male genitalia (Figure 3) lack the basal swellings and sharp ventro-medial spines characteristic of chrysopelea and lytrea, and are similar to, but more lanceolate than, those of jatrophae.
A. amathea is easily distinguished from all other members of the genus by the extent of its vivid red coloration. On the dorsal surface, the red coloration extends into two spaces between the four postbas- all submedian lines of the anterior forewing, fills the median area of the posterior forewing, the submedian and median area of the hindwing (except for a dark line running through it from anterior to posterior), and the hind submarginal area of the hindwing. There are usually three to four subapical, five postmedial and four sub- marginal white spots on the forewing, and from one to four small submarginal white spots on the hindwing. The basal and postbasal regions of the wings are brown; all other markings are dark brown to black.




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9791 Silberglied, Aiello & Lamas - Genus Anartia 223



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224 Psyche [June-September
The pattern is similar but much paler on the ventral surface. Most of the markings that are black dorsally are brown ventrally. There are, in addition, two dark postmedian spots, an elongate one in forewing cell Cu2 and a small, round one in hindwing cell M 1. The saturation of the red color, and the overall contrast of the pattern, is more pronounced among males than among females, especially on the dorsal wing surfaces. The red color is noticeably faded on older individuals and on old museum specimens. Figure 2.
Wing venation of the five species of Anartia. a = A. amathea, f = A. fatima, c = A. chrysopelea, I = A. lytrea, j = A. jatrophae. Scales = 1 cm. See text.



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I 9791 Silberglied, Aiello & Lamas - Genus Anartia 225 Specimens of A. amathea from the southeastern part of its range are characterized by fusion of the five postmedian forewing spots into a broad, white band. Examination of large numbers of speci- mens reveals that this spot fusion exhibits much variation both within and between populations, and forms a cline running from northwest to southeast (Figure 5). Accordingly, we recognize Esch- scholz' "roeselia" simply as that end of the cline showing the most distinctive forewing banding, not as a subspecies. Burmeister's "sil- vae," and Fruhstorfer's "thyamis" and "sticheli," are poorly- characterized variants that fall well within the ordinary range of variation.
Several common names have been coined for amathea, including the "Coolie" (Barcant, 1971), the "Tomato" (Kaye, 1921), and the "Red Anartia" (Riley, 1975).
The wing pattern of A. fatima is built around elements similar to those of A. amathea, but modified and colored in such a manner as to produce quite a different appearance. The wings are dominated by the distinctive bands, composed in the forewings of seven, and in the hindwings of five, enlarged postmedian spots, fused with one another. When A. fatima is at rest, the forewing and hindwing bands are joined in a continuous line. A. fatima also has three to four subapical and one to four submarginal spots on the forewing, of the same color as the band. The red coloration is restricted to a narrow median band on the hindwing (composed of four spots, distal to the position of the dark median line of A. amathea), and along the hind margin in some specimens. The remainder of the wings is largely dark brown to black, including the spaces between the four black postbasal/submedian lines on the anterior forewing. The ventral surface is similar in pattern to, but much lighter than, that of the dorsum; the bands are occasionally infuscated with darker scales beneath, and there is usually a well-developed, black, postmedian c-shaped mark in hindwing cell Ml just basal to the band. Males and females have similar patterns, but that of the male is generally more saturated and of higher contrast than that of the female.
The color, nature and function of the distinctive bands of A. fatima have been subjects of much research. In all populations, individuals can be found with yellow bands, white bands, or any shade from yellow to white. Fruhstorfer (1907) considered the white-banded form to be a distinct subspecies, venusta. Emmel



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Figure 3.
Male genitalia of the three species-groups of Anartia. Left: ventral view; Right: lateral view with left valve removed. Scale = 1 mm. [Specimen data: A. fatima: Panama, Zona del Canal, Barro Colorado Island; A. chrysopelea: "Cuba"; A. jatrophae: Colombia, Cali.]
(1972, 1973), assuming that the band color differences had a genetic basis, reported "phenotype" frequencies, as well as behavioral experiments designed to learn more about the maintenance of such a polymorphism. Taylor (1973) disputed Emmel's assumptions, demonstrating that the band color was age- and sex-related, and correlated with physical and physiological measures of age. Young and Stein (1976) showed that the band colors, of marked individuals in a population, fade with time; they also reported some equivocal data on the colors of individuals at eclosion. Our own (R.E.S. and A.A., in prep.) studies, which include rear- ings of over a thousand individuals, and the following of over a thousand marked individuals in a natural population, will be reported in detail elsewhere. But our clear-cut results are relevant to a consideration of the nature of Fruhstorfer's venusta and can be summarized as follows: Males always eclose with clear yellow wing- bands (N= 11 19). The color of the female wing-bands at the time of eclosion is variable, and may be anywhere in the continuum from



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19791 Silbergfied, AieIIo & Lamas - Genus Anartia 227 yellow to white; it appears to be independent of the length of the period of larval and pupal development. The bands of males, and of females that are not already white, always fade to white over a period of approximately two weeks under natural conditions. Fad- ing appears to be related to light exposure; it can be induced in dead specimens exposed to sunlight (Taylor, 1973), but does not occur in museum specimens protected from light.
A. fatima shows marked variation in the extent of red coloration on the hindwings, with an increasing expression of red on the hind margin of the hindwing, in the northwestern part of its range. Hoff- mann's "colimensis" (Figure 5) represents the extreme expression of red in A. fatima. There is an intriguing resemblance between this variant of A. fatima, and the banded "roeselia" of A. arnathea, at the northern and southern extremes of their respective ranges. Figure 4. Fl hybrids of A. amaihea and A. fatima; male above, female below. Reared on Blechum brownei at Barro Colorado Is., Panama. Left: A. amathea female x A. fdmu male: male AF-17, 1977; female AF-160, 1976. Right: A. fatma female x A. amaha mate: male FA-151, 1976; female FA-274, 1976.



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228 Psyche [June-September
Several other variants of A. fatima have also been reported or described, most of which are well within the normal phenotypic range. The more unusual forms include Kruck's "moreno" (a melanic lacking the characteristic band), and Hoffmann's striking "albifusa" (with the forewing band extending toward and fusing with the subapical white spots). Aiello and Silberglied (1978) reported, but did not describe taxonomically, an aberration with orange instead of red hindwing markings3, apparently due to the homozygous condition of a recessive allele at a single locus. A sim- ilar, probably homologous aberration apparently exists in amathea (A. Shapiro, pers. comm.).
The only common name we know for A. fatima is simply "Fatima" (Klots, 195 1).
Fl hybrids between amathea and fatima are illustrated in Figure 4. These reared specimens closely resemble those captured by G. B. Small, K. S. Brown (1975), and ourselves at several localities in eastern Panama where the two distributions are contiguous. The two reciprocal hybrids are intermediate between the parental spe- cies, and similar to one another, in color and pattern, and there is relatively little variation among the offspring of either cross. A paper illustrating and describing the Fl, backcross and F2 genera- tions, and discussing the interspecific genetics of pattern characters, is in preparation.
Anartia chrysopelea and A. lytrea
Anartia chrysopelea Hiibner
chrysopelea Hiibner, [1831]: 34, pi. [95], fig. 547, 548. Type locality: Cuba, La Habana.
[litraea, Herrich-Schaffer, 1864: 163.
Misspelling.]
lytrea subspecies eurytis Fruhstorfer, 1907: 112 (Anartia (Anartiella)). Type locality: "Haiti (?), Puerto Rico (?)." Anartia lytrea (Godart)
lytrea (Godart), 18 19: 299 (Vanessa).
Type locality: unknown; "de l'expedition du capitaine Baudin." dorninica Skinner, 1889: 86.
Type locality: Haiti, [Artibonite], Samana Bay. 3We have since found that the red color of normal fatima (and amathea) can be changed to orange, identical to that of this aberration, by immersing the wings in dilute hydrochloric acid.




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I 9791 Silberglied, Aieilo & Lamas - Genus Anartia 229 These two species, endemic to the largest islands of the West Indies, are very similar to one another and evidently closely related. They are identical in wing venation and genitalic structure, and differ only slightly in size, wing shape and wing pattern. Seitz (1 924) and Bates (1935) considered them to be subspecies ('choromorphs' of Bates). However, since there is little variation within each of these entities, since the differences between them are very consistent, and since they are welt-isolated geographically, they are evidently bio- logically separate species and are so treated here. The venational features that distinguish these species are the com- bination of a single vein crossing from R to Sc, and three veins Figure 5,
Above: variation among specimens of A. amathen in the expression of postmedial forewing banding. Specimen data, from left: Colombia, Viliavicencio, Dcpt. Meta, 588 m, 28 Sep 1942, M, Bates; Peru, La Merced; Brasil, Rio del Janeiro; Brad, Pelotas, C. Biezanko ["roeselia"]. Below: A western Mexican specimen of A. fatima illustrating extreme expression of dorsal hindwing red coloration, and ventral infuscation of the band. Specimen data: Mfexico, Colirna, Jacob Doll coil, ["coiimen- sis"],




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230 Psyche [June-September
branching from Sc toward the costa in the forewing (Figure 2). The male genitalia (Figure 3) are very distinctive, with a pronounced basal swelling and ventro-medial spine on each valve. A. chrysopelea is the smallest member of the genus, and has the most strongly developed "tail" at hindwing vein M3. The dorsal ground color of the wings is very dark brown, the males being darker than the females. A 2-3 mm wide, white postmedian band crosses the forewing, similar to that of A. fatima but composed of only five fused postmedian spots, and extending only to Cu2. Each hindwing bears an oval to rhomboid median white macula. A round 'ocellus' (eye-spot), consisting of black ringed with dull orange, is located in the anal angle of both fore- and hindwing, that of the forewing being slightly larger than that of the hindwing. The fine, dark, postbasal and submedian lines are present but obscured. Both wings have a series of dull orange submarginal lunules. The ventral ground color is lighter; the hindwing macula is infus- cated and crossed basally by a narrow stripe that extends from the costal margin to, and nearly surrounding, the 'ocellus.' The ventral forewing 'ocelli' are of the same relative size as they are dorsally. Occasional specimens have a suffusion of lavender scales postme- dially in the ventral hindwing.
A. lytrea is somewhat larger than A. chrysopelea, with lighter brown ground color and less distinct markings. Dorsally, the white bands are slightly infuscated, with less sharply defined edges. Those of the hindwings are more elongate, and not as wide in the middle. The orange ring surrounding the 'ocellus' in the anal angle of the forewing is much wider in A. lytrea, and the hindwing 'ocelli' are far smaller than the forewing 'ocelli.' As in A. chrysopelea, there is a narrow row of submarginal orange lunules, more strongly curved in A. lytrea. The underside pattern is modified in a way similar to that of A. chrysopelea, but in the hindwing the orange ring does not quite surround the 'ocellus.' The "tail" at hindwing vein M3 is not as pronounced in A. lytrea as in A. chrysopelea. A. chrysopelea and A. lytrea have been called "Huebner's Anar- tia" and "Godart's Anartia," respectively (Riley, 1975). Anartia jatrophae
Anartia jatrophae (Linnaeus)
jatrophae ([Linnaeus] in Johansson), 1763: 25 (Papilio). Type locality: "America"; Surinam (Munroe, 1942: 2).



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19791 Silberglied, Aiello & Lamas - Genus Anartia 23 1 corona Gosse, 1880: 199, pi. 8, fig. 1.
Type locality: Paraguay, "near Asuncion." Aberration. saturata Staudinger, [1885]: pi. 39, fig. [6] [as species]; [1866]: 104 [as variety]. Type locality: Haiti, Port-au-Prince; "Puerto Rico." jatrophae variety jamaicensis Moschler, 1888: 27. Type locality: "Jamaica."
[fatrophae aberration "margarita" Oberthur, 1896: 30, pi. 9, fig. 18. Type locality: Brasil, Bahia.]
jatrophae subspecies luteipicta Fruhstorfer, 1907: 1 12. Type locality: "Honduras."
jatrophae variety pallida Kohler, 1923: 24, pi. 2, fig. 12. Type locality: Argentina, Misiones.
jatrophae subspecies luteopicta Munroe, 1942: 2. Type locality: Honduras. Incorrect spelling, not available. jatrophae subspecies guantanamo Munroe, 1942: 2. Type locality: Cuba, Oriente, Guantanamo, San Carlos Estate. jatrophae subspecies semifusca Munroe, 1942: 3. Type locality: Puerto Rico, San Juan.
jatrophae subspecies intermedia Munroe, 1942: 4. Type locality: "St. Croix."
The name Jatrophae (which may be a misnomer based on Merian's [1705] erroneous larval foodplant association) has been attributed to Johansson, but we agree with Hodges (1971, p. 29-30) that authorship should properly be ascribed to Linnaeus.
A. jatrophae has the widest geographic range of any species in the genus. Morphologically, it is distinctive in the combination of two veins crossing separately from R to Sc, and three veins branching from there to the costa (Figure 2). The male genitalia (Figure 3) are most similar to those of A. amathea and A. fatima, but have blunter and slightly asymmetrical valves.
A. jatrophae has a distinctive appearance that sets it apart from the other species. The ground color of most of the wings is dirty white or light gray, with marginal and submarginal dull, rusty orange in some populations. The wings often have a pearly lustre, especially beneath. The pattern is quite complex and highly variable in the tone of pigmentation, distal ground color and expression of certain details. Besides the intricate series of dark lines, chevrons and lunules in the lighter field (better studied in the photographs then described), there are three characteristic postmedian dark spots: a large one in forewing cell Cul and others in hindwing cells Ml and Cul.
The ventral surface is much lighter in ground color, and even more variable than the dorsum. There are often red-orange submar-



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