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~
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, : 33.
Type species, Papilio jatrophae Linnaeus (Scudder, 1875: 11 1). Celaena Doubleday, : 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.
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, ; 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."
19791 Silberglied, Aiello & Lamas - Genus Anartia 22 1 silvae Burmeister, 1861: 168.
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
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.
9791 Silberglied, Aiello & Lamas - Genus Anartia 223
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.
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
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
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.
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, : 34, pi. , fig. 547, 548. Type locality: Cuba, La Habana.
[litraea, Herrich-Schaffer, 1864: 163.
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.
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"],
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).
19791 Silberglied, Aiello & Lamas - Genus Anartia 23 1 corona Gosse, 1880: 199, pi. 8, fig. 1.
Type locality: Paraguay, "near Asuncion." Aberration. saturata Staudinger, : pi. 39, fig.  [as species]; : 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  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-
232 Psyche [June-September
ginal lunules, and edgings of the same color on the lines in the ventral hindwing. The center of the dark spot in hindwing cell M 1 is often suffused with blue scales.
Geographic and seasonal pattern variation in A. jatrophae have been the subjects of several papers (e.g., Munroe, 1942; Gillham, 1957; see also Longstaff, 1912). Consideration of large numbers of specimens from many localities and dates reveals an unusually large amount of phenotypic plasticity in pattern detail, both geographic and seasonal. The "several recognizable but not easily defined sub- species" (Riley, 1975) were the subject of a careful quantitative investigation by Gillham (1957), who concluded that they resulted from discordant variation in several characters. Although several modern authors (Brown and Heinemann, Howe, Klots, Riley, etc.) continue to recognize subspecies in jatrophae (especially for the West Indian populations), we see no reason to do so. They are better referred to as "jatrophae from . . ." than by taxonomic epithets that substitute for knowledge of the factors underlying their variation. Common names used for A. jatrophae include the "White Pea- cock" (e.g., Holland, 1898; Klots, 1951; Riley, 1975; Rawson, 1976) and the "Biscuit" (Barcant, 1971).
ADULT BEHAVIOR AND ECOLOGY
Habitats, seasonality and population structure Species of Anartia are found wherever their larval foodplants occur. Feeding as they do on herbaceous tropical weeds (Table I), they are restricted to well-watered, disturbed habitats. Under natu- ral conditions, these would include flood plains, landslide areas, treefall gaps, and similar sites, to 2,000 m or more, depending on latitude.
Human activities benefit Anartia. Their foodplants grow well along irrigation and drainage ditches, and large populations are found along roadsides, and in agricultural situations, throughout most of Latin America (see Young and Muyshondt, 1973; Young and Stein, 1976). Anartia are frequently found flying in the com- pany of Junonia spp., with whom they have several larval food- plants in common. A. jatrophae appears to succeed in drier sites, and those with lower vegetation (e.g., Leck, 1974), where foodplants not utilized by the other species grow. In seasonally dry areas, popu- lations usually diminish in size during the months of little rainfall
FOODPLANT SPECIES FAMILY USED BY LOCALITY REFERENCE Blechum brownei
Jaco binia magnifica
Miiller (1 886)
A.A. & R.E.S.
A.A. & R.E.S.
Young & Stein (1976)
Young & Muyshondt (1973)
Young & Stein (1976)
A.A. & R.E.S.
Young & Stein (1976)
cited in Lima (1967-1968)
cited in Howe (1975)
cited in Lima (1967-1968)
cited in Lima (1967-1968)
cited in Lima (1967-1968)
cited in Lima (1967-1968)
cited in Lima (1967-1968)
cited in Lima (1967-1968)
Scudder ( 1892)
A.A. & R.E.S.
Gundlach (1 891)
A.A. & R.E.S.
Jatropha manihot2 Eurphorbiaceae J Surinam Merian, Sepp, . . . - - -
Riley (1975) disputed Lippia as a larval foodplant. 2Undoubtedly a mistake, traceable to Merian (1705), who figured A. jatrophae on this species of plant; see text. A = A. amathea, C = A. chrysopelea, F = A. fatima, J = A. jatrophae.
234 Psyche [June-September
(cf. Emmel and Leck, 1970). Local extinctions are frequent. A. jatrophae usually persists for awhile after its local congener has disappeared. During these dry times, populations are restricted to moist refugia, and search for these otherwise common species may be frustrating (e.g., Hall, 1925). Although some individuals enter a nonreproductive physiological state during the dry season (0. R. Taylor, Jr., in ms.), there is no evidence for prolonged physiological diapause in Anartia. Among other things, their short adult longevity would seem to preclude survival through a long dry season. Groups of adults may seek shelter in the same location (Young, 1979), but they do not form structured aggregations characteristic of many other tropical butterflies.
Adults are also influenced by the availability of nectar sources, and may leave an otherwise suitable area if no flowers are in bloom. They take nectar from many species, especially Lantana camara (but not from L. trifolia; Shemske, 1976; Barrows, 1976; they feed only at the yellow flowers of L. camara), Hyptis mutabilis and Sida sp. (Fosdick, 1973). The seasonal fluctuation in quality, of larval and adult habitats, affects the biogeography (9.v.) of Anartia. Based on study of collecting localities and dates, we believe that much of the phenotypic variation seen in A. fatima and A. jatrophae is due in part to environmental conditions experienced during development.
The population biology of A. fatima has been studied in Costa Rica by Young (1972) and Young and Stein (1976), and in Panama by Silberglied, Aiello and Windsor (in prep.). A. amathea has been studied in Ecuador by Fosdick (1973; but cf. Sheppard and Bishop, 1973!). Population sizes differed considerably between the species and studies; in Panama, dramatic differences in population size were noted from one year to the next. During one year, striking cycles of recruitment from the immature stages occurred on a monthly basis (R. E. S., A. A. and D. M. Windsor, in prep.). In spite of a sex ratio of 1: 1 at eclosion in A. amathea (&q= 1.04, N= 1,957) and A. fatima, (($:$ = 0.96, N=2,281), samples from Anartia populations may be strongly skewed toward one sex or the other. The population of A. fatima on Barro Colorado Island, for example, always had a significant preponderance of males, due in part to greater emigration by females in search of oviposition sites (R. E. S., A. A. and D. M. Windsor, in prep.; Organization for Tropical Studies report, cited in Young and Stein, 1976). On the
19791 Silberglied, Aiello & Lamas - Genus Anartia 235 other hand, Fosdick's population of A. amathea in Ecuador was skewed toward females; it is likely that his site contained an abund- ant supply of larval foodplant on which females oviposited. Survivorship was low in all populations studied, and it appears that under natural conditions, adult life is short-averaging from one to two weeks (maximum 9 weeks) in the field (R. E. S. and A. A., unpubl.). Young (1972) reported a longevity of 45 days in the laboratory. Adults are subject to heavy predation during their adult lives (see below). There are no field studies of the immature stages of any species.
Palatability and natural enemies
Due to their wide geographic ranges and local abundance, the three mainland species of Anartia have frequently been used in experiments on butterfly palatability, mimicry and predator learn- ing. All three species were completely acceptable to the numerous insectivorous vertebrate and invertebrate predators to which they were offered (Table 2). Human subjects report that A. fatima have "no taste" or a "walnut flavor" (Emmel, et al., 1968). The predators of adult Anartia are those generalist insectivores common in dis- turbed habitats, especially spiders and insectivorous birds. Larvae probably suffer greatly from predation by social and solitary wasps. We have reared one (unidentified) tachinid parasitoid from a wild Anartia larva, but have never encountered viral or bacterial disease during the rearing of over 5,000 individuals. In spite of their palatability, Anartia are often the most common species in the habitats where they occur. The tremendous losses of adults, and probably greater losses of larvae, are more than com- pensated for by the great fecundity in this genus (see below). Function of coloration
Various functions have been suggested for the color patterns of Anartia species. Anartia orient to the sun and bask (Longstaff, 1912; Fosdick, 1973). There is no distal circulation in their wings, so only the colors of the body and wing bases play a role in thermoreg- ulation (see Wasserthal, 1975; Douglas, 1979). Brower, et al. (1 97 1) present convincing experimental evidence to support the idea that A. amathea is an "incipient" Batesian mimic of Heliconius erato. Caged predators that tasted, and learned to avoid, H. erato, also refused the similar-colored A. amathea, even though
Palatability tests using Anartia species: all three species tested were found to be palatable to all predators. PREDATOR PREY CONDITIONS LOCALITY
Argiope argentata A field obs. Panama (Dariin) R.E.S. & A.A. A rgiope argentata F field obs.
Panama R.E.S. & A.A.
HY MENOPTERA (Vespidae)
Polist es canadensis F, larva
Ameiva ameiva (Teiidae) F
"several lizard species" F
field obs. Panama
feeding exp. Costa Rica
R.E.S. & A.A.
R.E.S. & A.A.
R.E.S. & A.A.
Brower & Brower (1964)
Coppinger (1 970)
Table 2. continued
Coppinger (1 970)
Brower, et al. (1971)
Brower, et al. (1971)
Coppinger ( 1969)
Brower, et al. (1971)
Coppinger ( 1969)
Brower, et al. (1971)
Homo sapiens F taste test Costa Rica Emmel, et al. (1968) A = A. amathea, F = A. fatima, J = A. jatrophae.
238 Psyche [June-September
the patterns in which the colors are arranged are very different in the two species. Less convincing is Emmel's (1972) suggestion that A. fatima mimics other, striped, Heliconius species. A. fatima resem- bles far more closely various Adelpha, Doxocopa, and other pre- sumably palatable nymphalines.
The wing-bands of A. fatima are visible from both above and below, like those of a great many other banded butterflies (e.g., Graphium kirbyi, Cyrestis acilia, Limenitis arthemis, etc.). Such "disruptive" patterns presumably protect their bearers from preda- tors (Platt and Brower, 1968), but the only evidence available to date does not support this hypothesis (Silberglied, et al., 1980). The wings of Anartia are brittle and easily fractured; mutilated individu- als bearing evidence of unsuccessful attacks by predators, are com- mon (e.g., Longstaff, 1912; see Silberglied et al., 1980). The wing color patterns of Anartia spp. also play important intra- and interspecific communicatory roles between butterflies. These are discussed below under "courtship and mating." Flight and daily activity
A. amathea and A. fatima have a jaunty, somewhat erratic flight that enables them to move about beneath the foliage of low herba- ceous vegetation when seeking eclosing females (males) or oviposi- tion sites (females). However, much of their time is spent in more open spaces as they feed at flowers, bask, chase other butterflies, etc. A. jatrophae has a strikingly different flight, in which long glides are interrupted by abrupt, mid-air pauses (". . . spasmodic. . . alternate 'start' and 'glide,"' Walker in Brown and Heineman, 1972). Since less time is spent beating the wings, this type of flight requires less energy per unit distance travelled, than that of A. amathea and A. fatima; it may enable individuals to fly considerably greater distan- ces. When alarmed, A. jatrophae seems to use an ascending escape maneuver more often than A. amathea or A. fatima. Anartia species are active under sunny conditions, and during light rain. They avoid the dark interior of the forest, and rarely fly in strong winds (Young, 1979). Emmel (1 972) plotted morning court- ship activity curves for yellow- (young) and white-banded (older) male A. fatima, and Young (1972) reported daily oviposition activ- ity of A. fatima to be between 10:OO and 13:OO hours.
19791 Silberglied, Aiello & Lamas - Genus Anartia 239 Courtship and mating
Male A. fatima use both 'waiting' and 'seeking' behaviors (Mag- nus, 1963) to locate females. 'Waiting' males are found sitting on vegetation, often with wings slightly spread, from which they fly to inspect nearly any butterfly that passes. These chases may be quite prolonged, even when chasing other species or conspecific males. While such behavior has often been called 'aggressive' (e.g., Walker, in Brown and Heineman, 1972), we know of no way to differentiate it from simple inspection flights in which the responses of the pursued individual provides information to the pursuer (see Silber- glied, 1977). Under crowded conditions in flight cages, groups of males sometimes form 'strings,' each male courting the one ahead. Males often return to the same waiting site after an unsuccessful chase. A. jatrophae males seem to prefer lower waiting sites than males of A. amathea and A. fatima.
When chasing, the male of A fatima attempts to get above and slightly behind the female. If she does not avoid him, the male executes a 'bobbing' flight, during which he may be sending chemi- cal and/or visual signals. Such 'bobbing' pairs persist for up to several minutes, the female descending lower and lower until she alights upon vegetation. The male alights next to the female, and attempts to couple with her by bending his abdomen laterally as he walks forward (Emmel, 1972), but she may still refuse his advances by flying off, or by spreading her wings. A side-to-side motion of the sitting female has been reported as denial behavior in A. amathea (Fosdick, 1973).
'Seeking' males fly low into vegetation, where they are often suc- cessful in locating and mating with teneral females (Emmel, 1972). Females usually mate during their first two days of adult life, but males generally do not mate until the third day after eclosion (R. E. S. and A. A., in prep.). Males do not mate more than once per day, but may mate on several days in succession. We have known indi- vidual males to mate up to nine times and still be capable of produc- ing a spermatophore.
Color and pattern appear to be important stimuli to males seek- ing females. Preference tests with dummies of A. fatima show that males prefer normally-colored females, and that obliteration of either the light bands or red markings reduces the number of approaches (Emmel, 1972; Taylor, 1973). Female A. fatima with
240 Psyche [June-September
yellow wing-bands are less attractive to males than those with white bands. The white band color reflects ultraviolet light more strongly (25% reflection) than does the yellow (14%)4; whether this compo- nent is important behaviorally remains to be determined. Males of A. amathea mate far less frequently with living females whose red color has been obliterated (R. E. S. and A. A., in prep.), than with red control females. Since A. amathea and A. fatima, like some other butterflies but unlike most other insects, see red (Bernard, 1979), it is not surprising that this color may be an important social signal. It has also been suggested that the black spots of A. jatro- phae may be important as a visual signal (Atsatt, 1968). Female mating behavior has been studied in A. amathea and A. fatima (R. E. S. and A. A., in prep.). Virgin females, isolated in flight cages from males, frequently approached other famales in what may be "solicitation" behavior, but of course this rarely would happen in nature.
The outcome of most courtships of non-teneral females is proba- bly determined primarily by female acceptance/rejection behavior (Taylor, 1972; R. E. S. and A. A,, in prep.). Females ofA. amathea mate assortatively, preferring conspecific males, but females of A. fatima do not discriminate between their own males and those of A. amathea. In A. amathea, females do not discriminate between con- specific males that had the red color obliterated, and normally- colored control males. The asymmetry of assortative mating, that results from these differences in female behavior of A. amathea and A. fatima, has evolutionary and ecological consequences at the junc- ture of the two species' ranges in eastern Panama. Copulation generally lasts from thirty minutes to one hour, but may be prolonged to as much as twelve hours. Young and Stein (1976) suggest that female A. fatima mate but once, Ehrlich and Ehrlich (1978) report a mean of 0.92 spermatophores per female in A. amathea (N=12, with no more than one per female) and Andersen (196?) reported a small number of female A. fatima with two spermatophores. Ehrlich and Ehrlich (1978) also suggest that Anartia might be a species capable of absorbing spermatophores; *Reflectance was measured densitometrically (Silberglied, 1976); on extreme yellow and white individuals. Emmel's (1972) description is misleading because, among other things, the television camera he used adjusts contrast automatically. Reflec- tance comparisons made with such a camera setup (Eisner, et al., 1969) are qualitative at best.
Silberglied, Aiello & Lamas - Genus Anartia Oviposition and Larval Foodplants
Females in search of oviposition sites fly within a few centimeters of low vegetation, and land frequently and briefly upon a variety of plants. Eggs are laid singly, usually, but not always, on the larval foodplant (Table 1). A. chrysopelea has been seen ovipositing on Tradescantia s p. (Det hier, 1 94 I), A. jatrophae on Cyperus diffusa, Oldenlandia corymbosa and Polygala verticillata, and A. fatima on "dead twigs, moss, rocks, walls, dry leaves, logs" (Young and Stein, 1976), "grasses, especially Oplysminus spp." (Young, 1972), Croton hirtus, Chaptalia nutans, garden hoses and cement walkways - none of which are acceptable larval foods. In the laboratory, A. fatima deposited more eggs on cage walls than on the Blechum brownei leaves provided. Apparently the only requirement for an oviposition site is that it be near the correct foodplant, but the stimuli important in eliciting oviposition behavior remain unknown (cf. Young and Stein, 1976). We have never seen any species of Anartia oviposit in an area that did not contain a real larval foodplant.
Anartia species have unusually high fecundity. A single female may lay several hundred eggs over the course of a few days (Young, 1972; Silberglied and Aiello, in prep.). Ehrlich and Ehrlich (1978) report that female A. amathea have approximately 100 eggs per ovariole, or a potential 800 eggs per female. Considering the sort of mortality for which such fecundity must compensate, the larva that survives to adulthood must be rare indeed. The larval foodplants of Anartia are listed in Table 1. While A. fatima and A. amathea accept Blechum brownei and B. costaricense as foodplants in Panama, in their natural habitat they would be less likely to encounter B. costaricense, a forest species. Neither A. ama- thea nor A. fatima will feed on Nelsonia brunellodes, another member of the same family, that often grows with B. brownei in Panama.
A. jatrophae has been reared upon numerous and diverse food- plants (see Table 1). Assuming that this pattern is real and not an artifact of limited data, we find it interesting that A. jatrophae, the most widespread of the three species, also has the broadest range of foodplants. This flexibility may enable it to coexist side by side with its congeners, with less direct competition for food. Furthermore, A.
242 Psyche [June-September
jatrophae's range extends to higher latitudes in both hemispheres than any other congener; its more polyphagous nature may be com- pared with the similar patterns found for temperate papilionids by Scriber (1973).
The first reliable account of the immature stages of Anartia was that of Muller (1 886), who described five larval instars and the pupa of A. amathea, and correctly identified the foodplant family as Acanthaceae. Earlier authors (Merian, 1705, copied by Sepp, 1852- 1855 - see Muller, 1886; Seitz, 1914) erroneously reported the foodplant for A. jatrophae as "Manihot," and figured adult butterflies together with a pubescent moth-like larva lacking scoli, and a pupal exuvium of dubious affinity. Later accounts of the immature stages are given for A. jatrophae by Scudder (1893), Dethier (1941), Klots (1951), Riley (1975) and Rawson (1 976); for A. amathea by Riley (1975); for A. chrysopelea by Dethier (1941); and for A. fatima by Young and Stein (1976). Nothing is known concerning the immature stages of A. lytrea. Eggs
The eggs (Figure 6) of A. amathea and A. fatima are yellowish green, 0.65 to 0.70 mm in diameter, slightly taller than wide, and have eleven to thirteen longitudinal wax-crested ribs which extend to within 15 degrees of the upper pole, which is centered on the micropyle. The ventral surface is flat. The ribs are perpendicular to and rest upon 40-50 low ridges with which the egg is banded. The number of vertical ribs is variable within species and even among the eggs of a single female (Dethier, 1941). The eggs of A. jatrophae are similar (Dethier, 1941; Rawson, 1976), but those of chrysopelea are wider than they are high (Dethier, 1941). We were unable to distinguish the eggs of A. amathea and A. fatima from one another. Larvae
While Anartia larvae have been described by several authors, only Dethier (1941) used morphological terminology5 precise enough for 51n their accounts of larval armature, some authors refer to setae as "hairs" or "spines," and to scoli beset with numerous setae as "branched spines." Many other inaccuracies are found in several published larval descriptions. Our terminology follows that of Peterson (1962); bilaterally arranged thoracic and abdominal armature units (e.g., scoli, verrucae, chalazae, setae) are described in the singular.
19791 Silberglied, Aielh & Lamas - Genus Anartia 243
244 Psyche [June-September
comparative work. Since the known larvae differ little between spe- cies, the following account may be considered generic except as noted.
FIRST INSTAR (based primarily on A. fatima: Head (Figure 7): well- sclerotized, scoli lacking; labrum emarginate, bearing six pairs of setae; frons triangular, bearing three pairs of setae, ventral margin concave; two pairs of adfrontal setae, upper pair shorter than lower; epicraniurn rounded, each side bearing five long setae; three setae associated with the stemmata ("ocelli") and one with the antennae, as figured. Thorax: ventral eversible prothoracic gland between the legs and head; cervical shield with four pairs of setae, three dark pairs directed forwards, and one pale, thinner pair, directed back- wards; subdorsal chalaza on meso- and metathorax; supraspiracular chalaza on pro-, meso- and metathorax, that of prothorax bearing two setae; subspiracular chalaza on meso- and metathorax, situated slightly above plane of spiracles; prespiracular chalaza on pro- thorax, bearing two setae; a chalaza located between subspiracular and subventral chalazae on meso- and metathorax may be serially homologous with the prothoracic prespiracular chalaza; subventral chalaza on pro-, meso- and metathorax, that of prothorax bearing two setae. Abdomen: first segment darker than the others; subdorsal chalaza on segments 1-9, a small chalaza between, and posterior to, the subdorsal and subspiracular chalaza on segments 1-8 (situated posterior to subdorsal chalaza on segment 8); supraspiracular chal- aza on segments 1-9; subspiracular chalaza on segments 1-9, that of segment 9 bearing two setae; subventral chalaza on segments 1-10 (segment 10 with two, located posteriorly); suranal plate rounded; prolegs on segments 3-6 and 10, well developed; crochets uniserial, uniordinal, arranged in a circle; setae (one per chalaza except as noted) microscopically serrate, and curved anteriorly. SECOND INSTAR (based on A. amathea, A. fatima and A. jatrophae): Head (Figure 7): as in first instar except for addition of a pair of epicranial scoli, and secondary setae in epicranial, frontal, anterior, and ocellar areas. Thorax: ventral eversible prothoracic gland between legs and head; cervical shield with four pairs of setae, two dark pairs and two pale pairs; subdorsal scolus on meso- and meta- thorax; supraspiracular verruca on pro-, meso- and metathorax; subspiracular scolus on pro-, meso- and metathorax; prespiracular verruca on prothorax; a verruca located between subspiracular and
19791 Silberglied, Aiello & Lamas - Genus Anartia 245 Figure 7. Head capsules of Anartia spp. a = A. amathea, c = A. chrysopelea, f = A. fatima, j = A. jatrophae. Numbers on figure indicate instar number; numbers in brackets are head capsule widths in mm. f2 [width of head capsule = 0.6 mm], j2 [0.5], c2 [0.575, measurement and figure from Dethier, 19411, fl [O.4], j 1 [0.4], f3 [1.0], j3 [0.9], a-final [2.5], f-final [3.0], j-final [2.7]. subventral scoli on meso- and metathorax may be serially homolo- gous with the prothoracic prespiracular verruca; subventral scolus on pro-, meso- and metathorax. Abdomen: mediodorsal scolus on segments 1-8 (segment 8 with two, one anterior, one posterior); subdorsal scolus on segments 1-8; supraspiracular scolus on seg- ments 1- 10; subspiracular scolus on segments 1-8; subventral scolus (small) on segments 1, 2 and 7; pair of subventral verrucae, one anterior, one posterior, on segments 3-6; pair of posterior subven- tral verrucae, one above the other, on segment 10; suranal plate triangular; prolegs as in first instar; crochets uniserial, triordinal, arranged in a mesoseries; setae numerous on each scolus.
246 Psyche [June-September
MORPHOLOGICAL DIFFERENCES BETWEEN FIRST AND SECOND INSTARS: In the second instar6, scoli and additional setae appear on the head (Figure 7); the chalazae of the first instar are replaced by scoli and verrucae; the setae are no longer serrate; the central pair of setae on the cervical shield is pale; a prothoracic subspiracular scolus appears (no prothoracic subspiracular chalaza in the first instar); mediodorsal armature appears on abdominal segments 1-8; the fol- lowing armature is lost: subdorsal chalaza on segment 9, the small chalaza between and posterior to the subdorsal and supraspiracular chalazae on segments 1-8, the subspiracular chalaza on segment 9, and the subventral chalaza on segments 8 and 9; a supraspiracular scolus appears on segment 10; the subventral chalaza on segments 3-6 is now a pair of scoli; the suranal plate becomes triangular; the crochets become triordinal and are arranged in a mesoseries. THIRD AND SUBSEQUENT INSTARS: The head scoli are clubbed (slightly more so in A. amathea and A. fatima than in A. jatrophae) in the third through final instars (Figure 7). The head width increases by factors of 1.5 (A. fatima), and 1.6 (A. amathea and A. jatrophae) (see Figure 8). The adfrontal sutures become conspicuous in the final instar, by which time the body is black, the scoli are reddish brown, and there are often coarse longitudinal stripes composed of light dots. The prothoracic eversible gland is present in all instars. The interspecific differences in larval morphology are very subtle. A detailed, comparative larval study must await the discovery of the larva of A. lytrea, and the collection of new material of A. chrysopelea.
The number of instars is variable: A. amathea from Colombia had five instars (Muller, 1886; R. E. S. and A. A.); A. fatima from Panama had six (A. A. and R. E. S.); from Costa Rica five (Young and Stein, 1976); A. jatrophae7 from Panama had five (A. A. and R. ^ethierYs (1941) description, of a second instar A. jatrophae from Cuba, differs from ours in the number of scoli on abdominal segments 8- 10, and in the reported absence of a subspiracular scolus on the prothorax. From his account of the first four instars of A. chrysopelea, the larvae of that species are very similar in setal arrangement to the three described above. However, he reports that the setae of the first instar larva . . . do not arise from conspicuous sclerotized areas . . . ." ^awson (1976) reported three instars for A. jatrophae from Florida, but from his illustrations it is probable that he missed one or more instars; his "third" instar is probably a fifth or sixth. The sum of Rawson's development times is also unusually short. Further rearing in Florida should be done to corroborate his account.
Silberglied, AieZZo & Lamas - Genus Anartia jatrophae
width of head capsule (mm)
Figure 8. Dimensions of head capsules and scoli of the larvae of A. fatima and A. jatrophae, reared individually. Note that A. jatrophae has one less instar, but grows more per instar, than A. fatima. However, the final larval head capsule size of A. jatrophae is not quite as large as that of A. fatima. [Instar 1 lacks scoli.] E. S.). Development times for A. amathea, A. fatima and A. jatro- phae7 in Panama are given in Table 3 (reared individually) and Figure 9 (reared under crowded conditions). Similar times are given for partial life cycles of A. jatrophae by Dethier (1941) and Rawson (1976), and for A. chrysopelea by Dethier (1941). Young (1972) reported 28 days, and Young and Stein (1976) reported 46-49 days, both for A. fatima in Costa Rica. Under identical rearing condi- tions, A. jatrophae takes less time and fewer instars to develop to adult than does A. fatima (Table 3).
Development time (days) for two species of Anartia, reared as isolated individuals under identical conditions in Panama. The difference between total mean development times for the two species is significant (t = 5.599, df = 21, p< .0005). A. jatrophae A. fatima
(N=7) (N= 1 6)
STAGE mean s.d. mean s.d.
TOTAL 31.72 0.76 34.38 1.54
A larva nearing pupation wanders for about one day, then pre- pares a silk pupation platform several centimeters above the ground on the underside of a leaf or twig. Platform making usually begins in the late afternoon or early evening, and is quickly followed by spin- ning of the silk stalk from which the pupa will be suspended. During platform making and stalk spinning, larvae evert the whitish gland located ventrally on the prothorax between the legs and head. The function of this gland is not known. Once the silk stalk is completed, the larva walks forward until its tenth segment prolegs are posi- tioned over it. These prolegs are then used to pull and shape the stalk before they finally clamp onto it, and support the larva during its final molt. By midnight most larvae have let go with all but the tenth segment prolegs, and now hang in a "J" position until 8 or 9 AM, when ecdysis takes place.
In the laboratory, larvae hang from the cage cover to pupate. Under crowded conditions, freshly-formed pupae may be cannibal- ized by hungry final instar larvae.
Pupae of A. amathea (Muller, 1886), A. fatima (Young and Stein, l976), and A. jatrophae (Scudder, 1893; Wolcott, 195 1; Rawson, 1976) are 15-22 mm long, smooth, spindle-shaped and without pro- tuberances. They are usually translucent jade green in color, with dark spots (Figure 6; see also Young and Stein, 1976) in the same positions occupied by scoli in the final larval instar, plus a few additional dark marks on the wings. Occasional individuals of all three species are black (Scudder, 1893; A. A. and R. E. S.). days since oviposition
Figure 9. Development times of male and female (stippled) A. amathea (N= 1,764) and A. fatima (N= 1,579), reared under crowded conditions (up to 20 larvae per container).
19791 Silberglied, Aiello & Lamas - Genus Anartia 249 The pupal period usually lasts six to eight days in the three species we have studied (see also Rawson, 1976; Young, 1972; Young and Stein, 1976). A day or two before eclosion, color changes can be seen through the pupal skin. The eyes turn yellow and finally brown, and the wings become pale brown (in A. jatrophae) or black (A. amathea and A. fatima). In A. fatima the wing-bands show clearly by the morning of eclosion. Adults emerge during the morning and are ready to fly within one to two hours. BIOGEOGRAPHY
The three distinctive evolutionary lines of the genus Anartia have well-defined and interesting geographic distributions (Figure 10). A. lytrea and A. chrysopelea are West Indian endemics on Hispaniola and Cuba, respectively. (A. chrysopelea is also known from the Isle of Pines, Swan Island, and southern Florida.) A. fatima and A. amathea are widely distributed in tropical Central and South Amer- ica, respectively. A. jatrophae is ubiquitous throughout all the warm regions of the Western Hemisphere, including the southern United States, the Bahamas and West Indies, all of Central and most of South America, to about 30 degrees north and south latitude. It has been difficult to determine the limits in some areas because of the tendency of collectors not to collect common species once a series has been obtained. Since all species can be collected around human habitations, many collectors do not bother with them soon after arrival in the tropics. Another problem has been the profusion of mislabelled specimens and erroneous reports, such as A. fatima from "Brasil," and A. amathea from "Mexico" (da Silva, 1907) and "Havane" (Lucas, 1857). The southernmost limits of A. jatrophae and A. amathea are poorly documented; neither species occurs in Chile.
Within these broad distributions, Anartia spp. are restricted to moist, or at least not very dry, disturbed habitats where their larval foodplants grow. Distributions may change markedly during the year in areas having pronounced dry seasons. Local extinctions of many populations occur through the dry season, with recoloniza- tion following the start of the rains. For example, during 1977, A. fatima went extinct throughout most of central and southern Panama, with occasional individuals remaining at isolated refugia (including dripping air conditioners and lawn sprinklers), but with a
Geographical distributions of Anartia species. See "Biogeography."
19791 Silberglied, Aiello & Lamas - Genus Anartia 25 1 substantial population remaining active on the moister Caribbean side of the isthmus. Within two months after the rains began, the species had reestablished itself in most of its former habitats. All species of Anartia are highly vagile. Based on our study of the Barro Colorado Island population, it appears that females of A. fatima emigrate more than males do. The ecological adaptations of A. fatima as a colonizing species have been summarized by Young (1972) and Young and Stein (1976). A. jatrophae, with its gliding flight similar to that of many migratory species, appears to be the most vagile, for it usually recolonizes former habitats long before A. fatima arrives. It also has a wider geographic range, and its popula- tions, while variable, are not strongly differentiated from one another, suggesting considerable gene flow. Vagrants of all species fly considerable distances, and occasionally establish local, tempo- rary populations beyond the normal range. For example, A. fatima reaches Kansas (Howe, 1975), A. jatrophae reaches Kansas and southern New England (Ehrlich and Ehrlich, 196 l), A. chrysopelea has turned up in southern Florida on at least two occasions (Ander- son, 1974; Bennett and Knudson, 1976) and possibly once on Anti- gua (Fruhstorfer, 1907), Godman and Salvin's (1882) record of A. lytrea on Jamaica, while unconfirmed (Brown and Heineman, 1972), is certainly within the realm of possibility, and A. amathea ". . . occurs sporadically. . . on Antigua, Grenada and Barbados. . . no doubt a vagrant . . . sometimes established for short periods (God- man and Salvin, 1896; Riley, 1975).
The distributions of all Anartia species, except A. jatrophae, are strictly allopatric of one another. A. jatrophae coexists with all; it is a better colonizer, utilizes a wider array of larval foodplants (Table l), takes less time to develop (Table 3), and withstands drier condi- tions. Such correlation of ecological distinctness with coexistence illustrates well the concept of limiting similarity of sympatric con- geners, and their comparative ecology would be worth a more detailed study (see also Young and Stein, 1976). EVOLUTION
Relationships to other genera
Young and Stein (1976) reported the "outstanding discovery" that the immature stages of A. fatima are similar to those of Siproeta, and suggested a close relationship between the two genera. This similarity, which involves the larval foodplants, egg and larval mor-
252 Psyche [June-September
phology, and open discal cell in both fore- and hindwing, had been discovered and published ninety years earlier by Miiller (1886; see also Brown and Heineman, 1972). While we agree with such an assessment, we want to take this opportunity to point out that this section of the subfamily is replete with genera of uncertain affinity, and we feel it is unwise to speculate further on phylogenetic posi- tions until a broad, modern generic revision of the Nymphalinae, employing larval and biological as well as adult characters, is undertaken.
Evolution within the genus
Anartia clearly contains three distinct phylogenetic lines: (1) A. amathea and A. fatima probably represent a pair of sister- species, derived from a widespread neotropical ancestor. We envi- sion a scenario in which the populations of this ancestor were isolated from one another during the Tertiary subsidences of the Panamanian isthmus, after which time the distinctive colors and patterns of the two species evolved. The secondary contact and occasional hybridization between A. amathea and A. fatima in the Darien represents a recent event in geologic time, the consequences of which are of considerable interest.
(2) A. chrysopelea and A. lytrea probably represent another pair of sister-species, derived from a common ancestor (Bates, 1 935). Their physical isolation on separate islands probably fostered their dif- ferentiation.
(3) A. jatrophae is a widely-distributed species of great geographic variation. Tendencies toward the formation of distinct, geographi- cally isolated populations are thwarted by the high vagility of individuals.
These three species-groups differ from one another in only a few morphological characters. It is not possible at present to decide which character states are plesiomorphic, and which derived, for these features. For this reason we do not feel it would serve a useful purpose to present speculations on the branching sequence within the genus.
DEPOSITION OF SPECIMENS
Voucher specimens of the immature stages of A. amathea, A. fatima and A. jatrophae have been deposited in the Museum of Comparative Zoology (MCZ).
All adult specimens illustrated,
19791 Silberglied, Aiello & Lamas - Genus Anartia 253 except for A. lytrea (American Museum of Natural History [AMNH]), are in the MCZ collection. Fl hybrids have also been deposited with G. B. Small, K. S. Brown, the Museo de Historia Natural "Javier Prado," the Peabody Museum (Yale University), the AMNH, and the National Museum of Natural History (Smith- sonian Institution), as well as the MCZ. ACKNOWLEDGEMENTS
We thank the numerous individuals and institutions who made specimens and data available to us. In particular, R. E. S. and A. A. thank G. B. Small for introducing them to hybridization between A. amathea and A. fatima, K. Radcliffe Dressier for help with rearing, J. M. Sigda for statistical assistance, E. Mayr for suggestions on conservation of the same fatima, and the Smithsonian Tropical Research Institute for financial support and the use of facilities. AIELLO, A., AND SILBERGLIED, R. E.
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