AMERICAN MUSEUM NOVITATES Published by Number 1229 THE AMERICAN MUSEUM OF NATURAL HISTORY May 5, 1943 New York City

THE ASCIA IN THE ANTILLES (, P1ERIDAE) BY WILLIAM P. COMSTOCK

The genus Ascia, in a broad sense, is from the Guianas, although it is not so considered to contain two occur- heavily marked as the specimen figured ring in the Antilles. The present status of by Kleemann. these species seems to be confused taxo- Gmelin (1790, p. 2262) concurred with nomically, and my present purpose is to Fabricius, giving additional references. present a more reasonable arrangement of Hutbner (1808, Sammlung, I, P]. cxxxvii, these species and their subspecies. figs. 1, 2, males; 3, 4, females) figured Mancipium Vorax monuste without lo- Ascia monuste monuste (Linnaeus) cality. These figures show heavily marked Surinam specimens with a strong ochre coloring on

Papilio monuste LINNAEUS, 1764, p. 237. the underside. There has been some discussion as to Latreille (1819, p. 141) gave a descrip- whether the name monuste should refer to tion which agrees with some of the male an American or an Asiatic species. specimens from the Guianas but said that Linnaeus described Papilio monuste in the species was found in China and asserted his division of the "Danai candidi" and that Fabricius was in error in selecting a made a second reference to it (1767, p. species from America with the underside 760), giving the locality, "Habitat in of the hindwing yellow. Barbaria." Boisduval (1836, p. 495) considered Muller (1774, p. 589) called monuste, monuste to be an American species. "Der barbarische Weissling" and said that Aurivillius (1882, p. 51) stated that the it came from "Barbarey," apparently type of monuste was not in the Museum following Linnaeus. Ludovicae Ulricae, and Jackson (1913) Fabricius (1775, p. 470) was the first did not list it in the collection of the reviser. He cited Kleemann (1761, p. 31, Linnean Society of London, nor in other P1. III, fig. 3) who gave an excellent but collections containing Linnean specimens. unnamed figure of the upperside of a male The type is presumably lost. Aurivillius of monuste such as occurs in the Guianas. gave his conception of monuste when he Fabricius gave "Habitat in America" and defined as "fig. typicae" those of Hiibner. remarked that the underside of the hind- Talbot (1929, p. 52) applied monuste to wing was yellow, which is a character more a Sumatran , described as cynis evident in the Antillean than in the conti- Hewitson (1866), and used the name nental populations of the species. Never- phileta Fabricius (1775) for the American theless his characterization is sufficient species. Later Talbot (1932, p. 207) to fix monuste as an American species. apparently reversed his opinion, for he However, various subsequent authors have accepted monuste as the genotype of had conflicting views. Ascia and as the stem name of various Cramer (1777, II, p. 71, P1. CXLI, fig. F) American subspecies and forms. called the species monusta, said that both Holland (1930, p. 133; 1931a, p. 278) surfaces were alike and that it was found considered monuste to be an American in China, gave the Linnean (1767) and the species. Kleemann references and figured the upper- The Linnean description, when care- side of a male which agrees with specimens fully read, defines a butterfly which is of 2 AMERICAN MUSEUM NOVITATES [No. 1229 common continental American occurrence. of the forewing with marginal spots de- The forewings are white with the apex, veloped on the hindwing at the veins. costa and outer margin fuscous; the hind- The coloration of the underside ranges wings are white with only the outer margin from pale cream-white to bright ochre, "denticulato-fuscus"; the underside is of sometimes with considerable brown mark- like color, but in place of the fuscous, only ing. The series in both sexes shows con- dusky at the margin. Males from Suri- nected intergradation between the ex- nam, or generally from northern South tremes, but these extremes, if considered America, are such as Linnaeus described, alone, are sufficiently different in both and he is known to have obtained speci- sexes to suggest two separate species. A mens of other species from Surinam. different picture is presented by a series The evidence provided by the original de- of thirty-four specimens from Dominica, scription, the opinion of Fabricius as first British West Indies, for the most part reviser, the figures of Kleemann and Cramer captured in November and December but and the likelihood that the type specimen showing no seasonal separation from others of monuste came from Surinam, all support in the series taken in January, April, Sep- the belief that the name monuste is properly tember and October. In this sample two applied to an American species and even extremes appear: nineteen males and more definitely to the particular form nine females of the light-bordered kind which occurs in Surinam and that this which, in a few specimens, show a slight locality may be fixed as the type locality increase in the width of the forewing of monuste. border; five males and one female of the The series of monuste in the collection broadly bordered kind; both kinds have of The American Museum of Natural the apex of the forewing and the entire History consists of over 600 specimens from hindwing on the underside ochre but many many localities in South, Central and narrow-bordered individuals are pale, while North America and the West Indies. This all broad-bordered specimens are strongly butterfly is a notable migrant, as stated ochre colored. If' the Dominican sample by Williams (1930, p. 126), and this habit was considered alone, the difference in the may account for a mingling of populations facies of the two forms is so marked that in the Antilles. In examining specimens it is certain that almost any taxonomist, from any particular insular region the without other information, would regard possibility of the influx of foreign elements them as two species. and of the interbreeding of various strains Several preparations made of the male must be considered, and it would not be genitalia of both kinds from Puerto Rico safe to reach positive conclusions as to the and from Dominica showed some very existence of stable geographical subspecies slight variation, but as this appeared even unless much larger series of specimens than between specimens having the same colora- are now available from reasonably segre- tion and pattern, it is considered to be gated populations could be examined. individual. The preparations agreed with Further, much more information is needed the figure of the monuste genitalia given about the nature of migrations and the by Klots (1933, P1. xii, fig. 95). I do not possible effects of immigrants upon in- consider that similarity of the genitalia is vaded populations. necessarily a proof that the two forms As an example, the variation observed examined are not specifically distinct. in a small sample from one population, a There are many cases known where the series of about 100 specimens from Puerto male genital armatures are similar in Rico, suggests extensive hybridization. several species which are distinct in pat- Great variation, which apparently is not tern. seasonal, is to be seen in both sexes. The There is a doubt in my mind as to marginal black-brown markings range in whether the narrow-bordered and broad- intensity from a narrow apical edging of bordered forms above referred to are dis- the forewing to a strongly dentate border tinct species or subspecies of one species. 1943] THE GENUS ASCIA IN THE ANTILLES 3

There is evidence in another migratory the intensity of this coloring is highly species that subspecies may occur to- variable, brown markings occur occa- gether under similar conditions. The sionally, but usually the surfaces are North American "Monarch," Danaus plainly yellow, and the veins are not out- plexippus, is known to migrate far to the lined in brown. south and has been captured in Puerto With an increased knowledge of monuste Rico where there exists the very distinct in the Antilles it might be possible to (probably largely sedentary) subspecific restrict the name eubotea to a definite population of Danaus plexippus portori- population. My use of the name for a censis Clark. composite group of populations which have In examining the various populations of principal characters in common is tentative monuste in the Antilles (including Florida) as a temporary aid in classification. there is a definite suggestion of underlying subspecific populations marked to a greater Ascia monuste phileta (Fabricius) or lesser degree, despite what appears to Florida me to be a general blending of the popula- Papilio phileta FABRICIUS, 1775, p. 471. tions suppositively caused by migrations. The population of monuste in Florida is Based on the material available and with a quite distinctive in that the males are full realization that my knowledge of it usually lightly bordered with black-brown is entirely morphological I now offer some on the forewing, lack marginal marks at the suggestions which I hope may be of aid in veins of the hindwing and are usually understanding the of monuste. palely colored on the underside. The I would first separate the Antillean females occur not infrequently with a dark populations of monuste as a whole from the smoky coloring on both sides of the wings, continental populations in a broad way but this kind intergrades to a normal whit- by the underside coloring. In addition I ish female. The name phileta has been would recognize two well-marked variant applied to the dark female as a dimorphic populations. These three I would classify form name. Fabricius described Papilio for the present as subspecies. phileta as related to monuste, giving the habitat as America. Although duskiness Ascia monuste eubotea (Latreille) of the females is not exclusively confined Antilles to the Floridian population, it certainly Pieris eubotea LATREILLE, 1819, IX, p. 144. appears more frequently in that population Pieris eubotea, BOISDUVAL, 1836, p. 500. than in others and to an extent to warrant Latreille described eubotea as a species its use as a subspecific character. Con- without locality, but Boisduval associated sidering the differentiation occurring in it with the female of monuste. The de- both sexes of Floridian specimens, I scription said that the upperside had suggest that this population should be dentate or crenulate borders on both wings recognized as a separate subspecies, Ascia and that the underside of the hindwing monuste phileta (Fabricius). In a series was yellow-ochre. Such females are not of seventy specimens from Florida there is uncommon in the Antilles, often lacking the no well-defined example of the generally blackish spot at the end of the cell on the distributed Antillean subspecies eubotea. forewings. Holland (1931a, p. 278, P1. LXVII, figs. For the present I suggest the use of the 15, 16) comments upon and figures a pair name eubotea for the most commonly taken in copula which are representative occurring manifestation of monuste in of monuste philela. the Antilles. The extent of the marginal Talbot (1932, p. 208) lists Ascia monuste niarkings of the upperside is variable, but cleomes (Boisduval and LeConte) from this margin is consistently dentate basad. southern United States. This is an The underside of the forewing usually has interesting form, about which there seems a distinct yellow apical area, and the hind- to be little information, but I believe that wing is entirely yellow on the underside; Talbot has properly recognized it. A 4 AMERICAN MUSEUM NOVITATES [No. 1229 single male from Virginia, No. 3935 of ward Islands there is a variant form which the Henry Edwards collection, deter- may be recognized as Ascia monuste vir- mined by Edwards as "cleomes Bdv. and ginia (Latreille). The evidence is that L.," agrees with the original description it occurs (perhaps as a migrant) both to and figures. This is a monuste of the conti- the north and south and that its region is nental t-ype with brown veins on the invaded from both directions (perhaps underside of the hindwing, but it is dis- through migrations) by the more wide- tinguishable from Mexican and Central spread forms, monuste eubotea and monuste American specimens. It is also distinct monuste. from phileta which is of the Antillean In one category or another, Talbot (1932, type. The name cleomes appears in North p. 207) lists eighteen names for monuste. American lists as a synonym of monuste. Some of these names appear to represent I suggest that it might be properly applied good continental subspecies such as raza to a subspecies with a more northern Klots from Lower California and automate range than phileta, but a study of more Burmeister from the Argentine. Talbot material is obviously needed. lists the remaining names variously as subspecies, forms and synonyms. Bois- Ascia monuste virginia (Latreille) duval (1836, pp. 493-495) described Antilles evonima, vallei and joppe from Cuba, all Pieris virginia LATREILLE, 1819, IX, p. 141. of which can be selected from a good Cuban Mylothris hemithea GEYER, 1832, Zutrage, IV, series of monuste eubotea, according to p. 24, Figs. 693, 694. Bates (1935, p. 116) who discussed monuste Latreille described virginia without a under the name phileta. At present, I locality, but Boisduval (1836, p. 494) gave include these Boisduval names as synonyms the locality as the Antilles. As described, of eubotea, but the first one might well be this is a form with a very narrow costal used to name a Cuban race when sufficient and outer-marginal bordering of dark knowledge of that population is available. brown in the forewing and otherwise It seems quite possible that with suffi- immaculate on the upperside. On the cient material and an increased knowledge underside, the apex of the forewing and the of the life histories, a series of insular sub- entire hindwing are plainly ochre-yellow. species of monuste might be shown to exist In a series of fifty specimens from the for which, incidentally, there is a suffi- Virgin Islands 40 per cent might be con- ciency of names available in the synonymy. sidered to qualify as virginia. The narrow- As previously indicated monuste monuste bordered females are particularly signifi- invades the Windward Islands from South cant. However, 60 per cent of the series America. Further, some males taken in I would consider to be monuste eubotea. Hispaniola are apparently monuste monuste, Males and females which would qualify being inseparable in appearance from as virginia occur in decreasing numbers specimens occurring in Central America. among populations of monuste eubotea Holland (1931, p. 256; 1931a, p. 278, in Puerto Rico and Jamaica. In a series P1. LXVII, fig. 17) described and figured of eighty-five specimens from Hispaniola a form from Florida, applying the "varietal none appears. or subspecific name crameri." As the form In St. Kitts, Antigua and Dominica, is described and figured this name is a virginia seems to be the prevalent form. synonym of monuste monuste. Its occur- In a series of thirty-four specimens from rence in Florida would be no more re- Dominica, twenty-eight specimens (over markable than in Hispaniola, but I have 80 per cent) qualify as virginia. This never seen specimens from Florida. form also appears in Guadeloupe and St. Life history information concernintg Lucia, but the material is insufficient to monuste is scant. Gundlach gave a de- draw any conclusions from these localities. scription of the and as occur- Summarizing the evidence on virginia, ring in Cuba. Cotton (1918, p. 281, it would seem that in the Virgin and Lee- Figs. 37, 38) presented further informa- 19431 THE GENUS ASCIA IN THE ANTILLES 5 tion. The half-dozen food plants men- which was described without locality. tioned in the literature include various Latreille stated, however, that the speci- species of Brassica, plants of the Chicory mens belonged to M. Dufresne, and and Caper families and Tropaeolum (Nas- Grimshaw (1900, p. 6) discovered types turtium). Breeding in quantity might (male and female) in the Dufresne collec- repay the investigator. tion in the Edinburgh Museum of Science The second species known from the and Art. Grimshaw said of josephina, Antilles is classified in the subgenus "This species, which comes from St. . It is also polytypic and has a Domingo and Mexico, is quite distinct from varied distribution. P. amaryllis, Fab., which is a native of The combination Papilio amaryllis was Jamaica." I recognize josephina as the first used by Cramer (1784, IV, p. 210, PI. subspecies occurring in Hispaniola. CCCXCI, figs. A, B) for a Palearctic species in the Satyridae. Fourcroy (1785, II, p. Ascia (Ganyra) josephina paramaryllis, 240) and Borkhausen (1788, I, p. 80), at new name later dates, separately used the same combsi- Jamaica nation to rename another satyrid but both The identity of Papilio amaryllis Fab- of these christenings fall as synonyms as ricius is established by the original de- well as homonyms. Still later Fabricius scription. Donovan (1800, P1. xxviii, used the combin,tion, creating' another fig. 1) figured it, possibly from the type. homonym, but this time the name was used Fabricius states that his specimen was in for a pierid, which is recognized as Ascia the collection of Dr. Hunter, and the type amaryllis (Fabricius) (1793, p. 189) or may still exist in the collection of the placed in a subgenus as Ascia (Ganyra) University of Glasgow. Grimshaw's de- amaryllis. The condition is unfortunate, termination of this subspecies as that one for according to the code a homonym is which occurs in Jamaica is now generally permanently defunct and amaryllis cannot accepted. be used for the stem name of the species, nor as the genotype of Ganyra. Ascia (Ganyra) josephina josepha There are three Antillean forms occur- Salvin and Godman ring, respectively, in Jamaica, Hispaniola Guatemala, Mexico, Nicaragua and Puerto Rico, and another in Central Pieris josepha SALVIN AND GODMAN, 1868, p. America, which are now listed as subspecies 150. Guatemala. under the stem name of Ascia amaryllis Salvin and Godman differentiated the (Fabricius) by Talbot (1932, p. 211). Central American subspecies under the These forms are sufficiently distinct in name josepha. The two names proposed facies to be readily separated, but for three by Fruhstorfer (1907, p. 139), gervasia and of the subspecies examined there appears protasia, are synonyms. Such individual to be no genitalic difference in the males. variants as he most briefly described are Therefore it seems correct to consider this to be found in any good series of specimens. butterfly as one species divided into four There is a connected intergradation be- geographical subspecies. Thus the prob- tween the light and datk females. lem becomes taxonomic, that is, to name correctly these subspecies. Ascia (Ganyra) josephina krugii (Dewitz) Ascia (Ganyra) josephina josephina Puerto Rico (Latreille) Pieris josephina var. krugii DEWITZ, 1877, p. Hispaniola 235, P1. i, fig. 3. Pieris josephina LATREILLE, 1819, p. 158. Although I have not seen specimens, Asca josephina, HEMMING, 1934, p. 194. krugii appears, from the description and Succeeding the homonym amaryllis, figure, to be a distinct subspecies. the next available name which may be used At present there is no subspecies of as a stem name is josephina Latreille, josephina recognized from Cuba. There 6 AMERICAN MUSEUM NOVITATES [No. 1229 is, however, Pieris menciae Ramsden have the principal veins of the forewing (1915, p. 15) which appears from the de- and to some extent the veins of the hind- scription to be very closely related to wing overlaid with chalk-white scales. josephina paramnaryllis, differing notably Not having examined specimens of menciae, only in the absence of the black spot at the I cannot make a definite statement, but end of the forewing cell. Like other sub- it seems quite possible that this is the species of josephina, the males of menciae Cuban subspecies of josephina.

KEY TO SUBSPECIES OF Ascia (Ganyra) josephina 1.-Length of forewing usually less than 35 mm.; black spot at distal end of forewing cell narrow, not more than 1 mm. wide...... 2. Length of forewing usually more than 35 mm.; black spot at distal end of forewing cell broad, at least 1.75 mm. wide...... 3. 2.-Male and female immaculate white except for a spot at distal end of forewing cell. (Jamaica) ...... paramarylli8. Female with dark spots along veins M3 and Cul of forewing on upperside; male and female with traces of a black bar beyond distal end of hindwing cell. (Puerto Rico) ...... krugii. 3.-Forewing falcate, outer margin concave from Ml to Cu2; hindwing margin angulate at Ms; vein M3 of hindwing longer from base to apex than distance from its base to base of R.. (His- paniola) ...... josephina. Forewing not falcate, outer margin scarcely concave; vein M3 of hindwing equal or shorter from base to apex than distance from its base to base of RB. (Central America) ...... josepha.

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