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Junonia g. genoveva (Cramer, 1780)
(Mangrove Buckeye)

Type specimens photo collection:

Distribution and Larval Foodplants:

NE South America, possibly Central America

Diagnosis

Identity of J. genoveva and J. evarete has been utterly confused. Their males can be told apart by the color of the antennal club from below. In genoveva, ventral surface of the club is dark: brown or black (not pale), and contrasts with the lighter color of the antenna below. In female genoveva, the club color is similar to males, but in female evarate, color can be variable, however the most distal tip is typically pale.

Synonymy

(From: A Catalogue of the Butterflies of the United States and Canada, Jonathan P. Pelham)

Genus Junonia Hübner, [1819]

Verz. bekannt. Schmett. (3): 34. Type-species: Papilio lavinia Cramer, 1775, Uitl. Kapellen 1(2): 32, pl. 21, figs. C, D; (8): 153 (index) (= Papilio evarete Cramer, 1779, Uitl. Kapellen 3(17): 18, pl. 203, figs. C, D; (24): 174 (index); homonym), by designation of Scudder (1872), 4th Ann. Rept. Peabody Acad. Sci. (1871): 43. Wahlberg et al. (2005), Biol. J. Linn. Soc. 86(2): 236, 239, 250, determined that Junonia and Precis Hübner, [1819], are distinct genus-level taxa. The following arrangement of species-level taxa is preliminary, and largely follows Lamas et al. (2004), Atl. Neotrop. Lepid. Chcklst. Part 4A: 251, and Neild (2008), Butterflies of Venezuela. Part 2.

=    Alcyoneis Hübner, [1819]

Verz. bekannt. Schmett. (3): 34-35. Type-species: Alcyoneis almane Hübner, [1819], Verz. bekannt. Schmett. (3): 35, no. 293 (= Papilio almana Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 472, no. 89; synonym), by designation of Hemming (1933), Entomol. 66(844): 197.

=    Aresta Billberg, 1820

Enum. Ins. Mus. Billb.: 79. Type-species: Papilio laomedia Linnaeus, 1767, Syst. Nat. (ed. 12) 1(2): 772, no. 176 (= Papilio atlites Linnaeus, 1763, in: Johansson, Cent. Ins. Rar: 24-25, no. 72; synonym), by designation of Scudder (1875) Proc. Am. Acad. Arts Sci., Boston 10(2): 117, no. 113.

=    Kamilla Collins & Larsen, 1991

In: Larsen, Butts. Kenya, Nat. Hist.: 444. Type-species: Papilio cymodoce Cramer, 1777, Uitl. Kapellen 2(9): 5, pl. 99, figs. G, H; (16): 148 (index), by original designation.

 

Junonia genoveva (Cramer, 1780)

Uitl. Kapellen 4(25): 4, pl. 290, figs. E ♂ D, F ♂ V; (34): 249 (index).

Original Combination: Pap[ilio]. Nymph[alis]. Phaler[ata] Genoveva

Type Locality: “Surinamsche” “Suriname”; suggested to be “northeastern South America” by F. Brown and Heineman (1972), Jamaica Butts.: 179-180; considered to be “West Indian” by L. Miller and F. Brown (1981), Mem. Lepid. Soc. (2): 176.

Types: Possible Syntype(s) in BMNH.

Bibliography

Excerpt from page 247 in Appendix 4 in:

Neild, Andrew F. E. 2008. The Butterflies of Venezuela. Part 2: Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae, Ithomiinae, Morphinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publications, London. 276 pp., 84 pls., 31 figs., 2 tabs, 4 maps. http://www.thebutterfliesofvenezuela.com

p. 247

Junonia evarete and J. genoveva – a discussion of the early nomenclature.  The first  published name for  the species here called J. evarete was lavinia Cramer [1775], based on specimens from Surinam with iridescent blue on the DHW of males. However, this name is unavailable for use as it is a homonym of an earlier name (see the genus introduction). In order to establish which species the name lavinia applies to, I consulted Cramer’s original paintings (numbered plate 12, figs. B & B) of lavinia in the BMNH (London). These are more detailed than their published hand-painted counterparts (plate 21, figs. C & D). The finely painted antennae are creamy grey with creamy-yellow clubs (typical of blue forms of J. evarete), and the VHW is dark brown (also typical of this species in the Guianas). The sex of this specimen is not specifically mentioned, though in his description Cramer states that “green” (actually blue) is only present in males. However, blue is rarely found on some females of this species, and the width of the DHW orange submarginal bands strongly suggests that the specimen figured by Cramer is a female. In fact, with the sole exception of the blue iridescence, other characters such as the wing shape, general wing pattern, and markings are remarkably similar to Cramer’s later figure of the female of J. evarete.

 

The next available name for this species is that employed in this work, J. evarete Cramer [1779], while I consider J. genoveva Cramer [1780] to represent the second species under discussion. In both cases the text of the original description is of little help for species recognition. Hence in order to ascertain the most accurate characters for Cramer’s two taxa, I again consulted the original master paintings in the BMNH (J. evarete: plate numbered 172, figs. B & B, but published as plate 203, figs. C & D; J. genoveva plate numbered 267, figs. C & C, but published as plate 290, figs. E & F). These figures are more detailed than published versions, and were used to hand-paint the published plates. The figures were then compared with a series of specimens in the BMNH from Surinam (the type locality of both taxa), in order to establish characters for the two species in that country.

 

The Surinamese specimens which I examined in the BMNH collections (and about a dozen on loan from the RNHL) were separated into two species based upon the color of the antennae tips, and subsequently the two series were observed to agree with the other characters and generalizations noted on page 59. The first series, with light tipped antennae, little submarginal orange on the DHW, and a highly distinctive and dark VHW (with reduced eyespots which are rarely absent, and relatively monotonous coloration), clearly belong to the phenotype figured by Cramer as J. evarete (for figures of both sexes from French Guiana, see Brévignon, 2004, figs. 13–16). Cramer described both sexes of evarete but figured a female whose sex is confirmed by its forelegs, and by the complete orange submarginal band on the DHW. Cramer correctly noted that the male on the dorsal surface is not as brown as the female but is greyer. Unfortunately the figured antennae of evarete, though finely painted, are clearly not of a Junonia species (they are totally black on both surfaces), and we must assume that they were glued to the figured specimen. (NB: During a field trip to French Guiana (Surinam’s eastern neighbor) in late 2000 I had the opportunity to examine numerous individuals of nominate J. evarete and J. genoveva in the field. This demonstrated unequivocally that all specimens with dark brown coloration of the VHW do indeed possess pale-tipped antennae, and based on these characters alone were in all cases immediately distinguishable from individuals of J. genoveva. I also observed that males of J. evarete usually have blue on the DHW, while the opposite is the case for females).

 

The type of J. evarete has not been located, and appears to be lost (Gerardo Lamas, pers. comm.). Certainly neither I nor Phil Ackery (pers. comm.) have found any other potential type material in the BMNH, while Lamas has many years of experience searching for Neotropical type material in most of the major Western institutional collections. In order to objectively resolve the current confused taxonomic status of the name evarete, I am designating a neotype (see Figure A1). This male specimen, with forewing length 30 mm from base to apex, is in the RNHL collection (Nationaal Natuurhistorisch Museum, formerly the “Rijksmuseum van Natuurlijke Historie”) in Leiden, Netherlands (no suitable females exist in the BMNH or RNHL). Its data are as follows: “W. C. van Heurn, Paramaribo. Suriname. 1911” (printed on white card bordered on each side by a thin black line). Although the VHW markings and coloration of this neotype do not agree exactly with the Cramer figure, nevertheless the specimen is similar, and it certainly falls within the perceived variation of topotypic specimens. The dorsal surface of the neotype is very similar to the Cramer figure, as is the VFW. The VHW is a little paler, and further differs by the presence of a very narrow median line of lighter brown and whitish scales, and by a single small black ocellus in each of spaces 2 and 5, faintly and thinly ringed by black (diameter 2 mm). In fact the Cramer figure appears to be of a rather extreme form, for the VHW markings of “average” individuals tend to be more developed even than those of the neotype. The latter is further characterized by its torn left FW apex, and there is a section missing from each of the right FW tornus, HW tornus, and outer margin of space 2. Despite the relatively poor quality of the specimen, it has been selected as the only suitable topotypic (Surinamese) specimen. This neotype is simultaneously designated as the neotype for esra Fabricius (see below), a nomen dubium, for which clarification is necessary to ensure future stability of the nomenclature. In response to the qualifying conditions of Article 75.3 of the ICZN (1999), all relevant statements made in this revision of Junonia are to be treated as part of this neotype designation.

 

The second series of Surinamese specimens (those with dark tipped antennae) are very distinct from J. evarete, and females of these clearly agree with Cramer’s figure of J. genoveva (Cramer does not say upon which sex his description is based, but he evidently figured a female: the meticulously painted forelegs are clearly articulated, and this sex is further suggested by the shape of the abdomen and wings, by the orange washed basal half of the upper surface of both wings, and by the width of the DHW orange submarginal band). The original painting clearly shows black tipped antennae, and this coupled with the orange-tinged basal half of the dorsal surface and the wide orange DHW band, permit identification of this taxon. There are only two female specimens of J. genoveva in the BMNH which could possibly be syntypic. Both come from the Felder collection (a source of many specimens which belonged to or were examined by Cramer). Though similar to the figures in Cramer, they clearly are not the figured specimen. Both antennae of one (from Surinam, “coll. Lenep [sic]”) have been glued on (one antenna appears to be from J. genoveva, the second from J. evarete), while the antennae of the second (without locality data, but with a handwritten label characteristic of specimens from Van Lennep’s collection) are old and discolored, and it is not easy to assign them to either species (however, my examination by microscope indicates these are the original antennae). The fact that Cramer’s description does not mention specimens in Van Lennep’s collection suggests neither specimen was part of the type series.

 

It is evidently desirable, given the obvious confusion in the literature and the similarity between this species and J. evarete, that the name-bearing type(s) be unequivocally identifiable. Granted that this is not the case, and since Gerardo Lamas (pers. comm.) is also of the opinion that their status as true syntypes is uncertain, it would be imprudent to designate either of these two specimens as a lectotype (following the same logic as for neotypes under Recommendation 75A of the ICZN, 1999). Lamas also informs me that he has been unable to locate any further type material in any Western institutional collection, nor have I nor Phil Ackery (pers. comm.) in the BMNH. In order to objectively resolve the current confused taxonomic status of the name genoveva, I am following Lamas’ advice and designating a neotype (I considered designating Cramer’s figure as the lectotype, following Article 74.4 of the ICZN, 1999, but this action seems wholly undesirable as it provides no specimen for physical examination). I have been unable to locate any topotypic (Surinamese) female in reasonable physical condition, and have therefore selected one from northern French Guiana, which is faunistically comparable (e.g. the nominate subspecies of J. evarete occurs in both localities). This neotype has a FW length of 26 mm, and data as follows (two rectangular white cards, with black print): Sinnamary (by bridge over estuary), NC French Guiana. 4.XII.00. Sea level. Leg. A. Neild. / Junonia genoveva genoveva ♀ (Cramer, [1780]). Det. A. Neild, 2000 (Neild collection, to be deposited in the BMNH upon publication). This specimen (see Plate 7, figures 199, 200), which is in excellent condition, agrees closely with that figured by Cramer, differing most significantly by its darker (less orange) basal half of the DHW, and by the presence of orange scaling in the distal (submarginal) half of the transverse white FW band. As noted above for the neotype of J. evarete, all relevant statements made in this revision of Junonia are to be treated as part of this neotype designation (Article 75.3 of the ICZN, 1999).

Figure A1. The male Neotype of Junonia evarete evarete designated in this work. (Photo: R. de Jong)
Image from: Neild, Andrew F. E. 2008. The Butterflies of Venezuela. Part 2: Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae, Ithomiinae, Morphinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publications, London. p. 247
http://www.thebutterfliesofvenezuela.com

Author's version of the text published as:

Calhoun, John V. 2010. The Identities of Papilio evarete Cramer and Papilio genoveva Cramer (Nymphalidae), with Notes on the Occurrence of Junonia evarete in Florida. News of the Lepidopterists' Society 52(2): 47-51, 18 figs.

p. 47-51, f. 1-18, formatted by the author, not as in the "News of the Lepidopterists' Society"

 

The Identities of Papilio evarete Cramer and Papilio genoveva Cramer (Nymphalidae),
with Notes on the Occurrence of Junonia evarete in Florida

 

John V. Calhoun

977 Wicks Dr.

Palm Harbor, FL  34684

bretcal1@verizon.net

 

The true identities of Papilio evarete and Papilio genoveva have long been disputed.  Now placed in the genus Junonia Hübner, they were originally described by the Dutch naturalist Pieter Cramer (1721-1776) in his multivolume publication (completed by Casper Stoll) on the butterflies of Asia, Africa, and America.  Cramer’s type specimens are lost, but he provided hand-colored engraved illustrations of each species (Pl. 203, figs. C & D and Pl. 290, figs. E & F) (Cramer [1779], [1780]).  These names were subsequently used to recognize seasonal forms and subspecies.  They were even combined into the subspecies J. evarete genoveva (see Schwartz 1989 for a review of their usage).  Uncertainty also plagued other aspects of their status.  Miller & Brown (1981) mentioned that the type locality of P. genoveva was “not stated” and “probably West Indian,” yet Cramer indicated that both species were from “Suriname” (South America).  The identities of these taxa are of particular interest to those who study the butterflies of the southern United States and Latin America.     

    

A key investigation by Turner & Parnell (1985) confirmed that evarete and genoveva act as separate species in Jamaica, which corroborated the observations of Clench & Bjorndal (1980) in the Bahamas.  After consulting Cramer’s illustrations, Turner & Parnell concluded that J. evarete represented the species commonly known as the Mangrove Buckeye, while J. genoveva denoted the Tropical Buckeye.  Most subsequent authors followed this usage, but the application of these names remained irregular.  For his book on the butterflies of North America, Scott (1986) elected to follow the nomenclature of Clench & Bjorndal (1980), who applied these names to the opposite species (J. Scott pers comm.).  This enduring doubt caused some authors (e.g. Opler & Malikul 1992) to transpose facts about each species.  Based on an anticipated arrangement of Junonia by Lamas (2004), Opler and Warren (2002) also reversed the names of these species relative to Turner & Parnell (1985).  This nomenclature was adopted for other publications, including the popular field guide by Brock & Kaufman (2003).  Despite this trend, only anecdotal evidence supported its validity and online Lepidoptera talk groups continued to debate the issue.  These conflicting interpretations left lepidopterists without a clear concept on which to base identifications of evarete and genoveva.  This changed in 2008 with the publication of the second volume of the comprehensive series of guide books, The Butterflies of Venezuela, by Andrew Neild.  In fact, the nomenclature employed by Opler & Warren (2002) and Lamas (2004) was based on Neild’s unpublished research.   

 

To better understand the status of J. evarete and J. genoveva, Neild (2008, visit Andrew Neild "The Butterflies of Venezuela" book series web site at http://www.thebutterfliesofvenezuela.com) “dedicated a disproportionate amount of time to Venezuelan and continental Junonia in an attempt to unravel the perceived enigma.”  Because Cramer purportedly based his descriptions and figures on specimens from “Suriname,” Neild compared numerous specimens from that region of South America with the original drawings that served as the basis of Cramer’s published illustrations.  Rendered by the Dutch artist Gerrit Wartenaar Lambertz (1747-1803), these illustrations are generally more detailed than their engraved counterparts.  Although Turner & Parnell (1985) stated that they consulted the “original drawings of Cramer,” it does not appear that they examined the drawings by Lambertz, but rather used this phrase in reference to the published engravings.  Neild (2008) also argued that several characters used to separate these species in Jamaica are “of limited or no value for specific distinction of the continental populations.”  He found that some of the characters used to identify these species in Jamaica apply to the opposite species in the vicinity of Suriname.  Based on this evidence, Neild (2008) designated neotypes, which objectively defined these nominal species as Papilio evarete=Tropical Buckeye and Papilio genoveva=Mangrove Buckeye.  This action overturned the interpretation of Turner & Parnell (1985).  Although many of Cramer’s references to “Suriname” are erroneous, the similarity of his Junonia illustrations to the butterflies of that region strongly supports Neild’s conclusions.  To help familiarize other lepidopterists with his research, Andrew Neild kindly granted me permission to write this brief article and include relevant images in a comparative format (Figs. 1-12).  The original figures by Lambertz have not previously been published.     

 

Neild (2008) asserted that males of J. evarete and J. genoveva can generally be separated by the color of the ventral surface of the antennal club.  In evarete it is usually pale and similar in color to the ventral shaft, while that of genoveva tends to be dark brown or brownish-black, contrasting with the color of the shaft.  In female evarete the ventral club is variable in color, yet the extreme distal tip is usually pale.  The ventral club of female genoveva is usually like that of the male.  Turner & Parnell (1985) did not discuss the genders of Cramer’s figured specimens.  Pelham (2008) identified both as males, but Neild (2008) concluded that they were likely females, though the figures of evarete possess some male characteristics.  Despite this assessment, a male specimen was selected to serve as the neotype of evarete, as Cramer’s written description was based on both sexes and the ventral antennal club of female evarete is occasionally darker, resembling that of genoveva (A. Neild, pers. comm.).  Antennal coloration is seemingly reliable in most areas, but this and other diagnostic features reportedly break down in parts of Mexico (A. Warren pers. comm.) and possibly elsewhere.  Hafernik (1982) suspected that these species are involved in a complex pattern of interrelationships that may not easily be reconciled through conventional taxonomic categories.  Phylogenetic studies of the genus Junonia by Kodandaramaiah & Wahlberg (2007) support the separation of evarete and genoveva (at least between some West Indian and Brazilian populations), but evidence suggests that additional subspecies and/or species await description within this group (Brévignon 2004, Lamas 2004, Neild 2008).  Images of these species from various geographic locations are available on the valuable website, Butterflies of America (Warren et al. 2010).    

 

Junonia evarete (Tropical Buckeye) ranges throughout much of the Neotropics northward to the southwestern United States and Florida.  Populations in southwestern North America are extremely variable and include the melanistic subspecies J. e. nigrosuffusa Barnes & McDunnough, whose status remains unclear (it may involve multiple species).  Florida populations are considered to represent the subspecies J. e. zonalis (C. Felder & R. Felder).  Neild (2008) designated a male lectotype of Junonia zonalis from among three syntypes that were collected in Cuba during the mid-19th century by Johannes (Juan) Gundlach (1810-1896).  These specimens were mentioned in the original description of zonalis by Felder and Felder (1867).  The first author to document this butterfly was Sloane (1725), who figured a specimen from Jamaica (Figs. 13, 14).  Sloane used no name, but described the species as “A small dark brown colour’d Butterfly, with black spots like Eyes and some rusty marks.”  He also remarked, “’Tis to be met with plentifully in the Savannas where it frisks up and down taking no long Flight.”     

 

Butterflies recognized as J. genoveva (Mangrove Buckeye) are found over a large portion of the Neotropics northward to Florida and Texas.  It occurs locally in Florida along the coast of the central and southern peninsula in association with its hostplant, black mangrove (Avicennia germinans (L.)L., Avicenniaceae).  Populations in Texas are confined to the southern coast near tracts of black mangroves, but these butterflies are poorly understood and often confused with phenotypes of J. evarete.  Hybridization in this region between these taxa and J. coenia complicates their identification.  Northern populations of J. genoveva are not taxonomically defined. 

 

Andrew Neild has contributed much to our basic understanding of J. evarete and J. genoveva.  Additional studies of Neotropical Junonia will undoubtedly reveal more surprises.  Visit www.thebutterfliesofvenezuela.com for more information about The Butterflies of Venezuela.  A glowing review of the second volume of this book was published recently by Penz (2010). 

 

Junonia evarete in Florida. During November and December of 1981, J. evarete was found at several locations in the Florida Keys and in the vicinity of Homestead on the Florida mainland (Baggett 1982a, 1982b, pers. comm.).  This was thought to be first documented occurrence of this species in Florida.  However, I discovered in the University of Michigan Museum of Zoology a single male of this species from Key Largo, collected on 16 August 1961 by Thomas E. Pliske (Fig. 15).  It is possible that J. evarete has long occurred in Florida as an irregular colonist, most likely from Cuba, but overlooked because of its similarity to J. genoveva and especially the abundant J. coenia.  The 1961 specimen was found among a series of J. genoveva, thus other Florida specimens of J. evarete may be misidentified in collections.  It is also conceivable that purported Florida hybrids between J. coenia and J. genoveva (Remington 1968, Scott 1986) include specimens of J. evarete.  The individual of J. evarete found in 1961 was thought to be such a hybrid (T. Pliske, pers. comm.) and I initially mistook individuals of this species to be hybrids when I encountered them on Plantation Key in 1981.  Rutkowski (1971) observed on Big Pine Key what he believed were “copulating pairs representing various intergradations” between J. coenia and J. genoveva.  Despite this potential confusion, Marcus (2007) confirmed that the DNA of J. genoveva and J. evarete from Florida show evidence of hybridization with J. coenia

 

Junonia evarete remains restricted in Florida to the extreme southern peninsula and Keys, where it inhabits weedy disturbed habitats in association with its hostplant, blue porterweed (Stachytarpheta jamaicensis (L.)Vahl; Verbenaceae).  Several locations in the Keys where J. evarete occurred have been lost to development.   This species is now most frequent along the grassy margins of drainage canals in western and southern Miami-Dade County.  It is multivoltine and adults can be found throughout the year.  They are mainly active during mid-afternoon, when they perch and bask beside low levees that parallel the larger canals (Figs. 16-18).  Adults routinely settle on the ground, but are extremely wary and take flight at the slightest provocation.  The butterflies briefly visit flowers, especially beggerticks (Bidens alba (L.)DC).  In late afternoon I have observed both sexes retreating to brushier areas, presumably to rest for the evening.  The origin of Florida populations remains under investigation (J. Marcus pers comm.).     

 

Acknowledgements: Thanks are extended to Andrew Neild for his meticulous research, as well as for generously sharing his evidence and supporting this article.  He also offered helpful comments on an early draft of the manuscript.  Rienk de Jong granted permission to reproduce images of the P. evarete neotype.  Mark O’Brien photographed the 1961 Florida specimen of J. evarete and Thomas Pliske recalled its capture.  Linda Cooper provided the photos of living J. evarete.  Nick Grishin, Jeffrey Marcus, Charlie Sassine, James Scott, John Shuey, and Andy Warren discussed their experiences with Junonia.  Beverly Pope and John Heppner assisted in obtaining literature.  Thanks also to Mark Salvato for directing me to areas in southern Florida where J. evarete was recently observed.  

 

 

Literature Cited

 

Baggett, H. D.  1982a.  Report on the 1981 Homestead meeting. So. Lepid. News 3:19-23.

_____.  1982b.  1981 season summary: Florida. News Lepid. Soc. No. 2 (Mar/Apr):26-27.

Brévignon, C.  2004.  Description de deux nouvelles sous-espèces guadeloupéennes du genre Junonia Hübner, 1819 (Lepidoptera, Nymphalidae, Nymphalinae) Lambillionea 104 (1): 72-80.

Brock, J. P. & K. Kaufman.  2003.  Butterflies of North America. Houghton Mifflin, New York, New York. 383 pp.     

Clench, H. K. & K. A. Bjorndal.  1980.  Butterflies of Great and Little Inagua, Bahamas. Ann. Carnegie Mus. 49:1-30.

Cramer, P.  [1779].  De Uitlandsche kapellen voorkomende in de drei waereld-deelen Asia, Africa en America [Papillons exotique des trois parties du monde l’Asia, l’Afrique et l’Amerique]. Vol. III. S. J. Baalde; Utrecht, Barthelemy Wild., Amsterdam. 1-128, pls. 193-264.

_______.  [1780].  De Uitlandsche kapellen voorkomende in de drei waereld-deelen Asia, Africa en America [Papillons exotique des trois parties du monde l’Asia, l’Afrique et l’Amerique]. Vol. IV. S. J. Baalde; Utrecht, Barthelemy Wild., Amsterdam. 1-28, pls. 289-304.

Felder, C. & R. Felder.  1867.  Reise der Österreichishischen Fregatte Novara um die erde in den jahren 1857, 1858, 1859 unter den befehlen des Commodore B. Von Wüllerstorf-Urbair. Zoologischer theil. Zweiter band. Zweite abtheilung: Lepidoptera. Rhopalocera. 3:379-536, pls. 48-74.      

Hafernik, J. E., Jr.  1982.  Phenetics and ecology of hybridization in buckeye butterflies (Lepidoptera: Nymphalidae). Univ. Calif. Publ. Entomol. 96:i-vii, 1-109.

Kodandaramaiah, U. & N. Wahlberg.  2007.  Out-of-Africa origin and dispersal-mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae). J. Evol. Biol. 20:2181-2191.

Lamas, G. (ed.).  2004).  Checklist: Part 4A: Hesperioidea – Papilionoidea. In Heppner, J. B. (ed.), Atlas of Neotropical Lepidoptera. Vol. 5A. Assoc. Tropical Lepid.; Sci. Publ., Gainesville, Florida.    

Marcus, J. M.  2007.  A history of invasion and hybridization in the buckeye butterflies (genus Junonia) of Florida. Presentation abstract. Pp. 15, In Program of the 58th Annual Meeting of the Lepidopterists’ Society. Bakersfield, California.  

Miller, L. D. & F. M. Brown.  1981.  A catalogue/checklist of the butterflies of America north of Mexico. Lepid. Soc. Mem. No. 2. vii+280 pp.         

Neild, A. F. E.  2008.  The butterflies of Venezuela. Part 2: Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae, Ithomiinae, Morphinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publ., London. 276 pp, 84 pl. http://www.thebutterfliesofvenezuela.com

Opler , P. A. & V. Malikul.  1992.  A field guide to eastern butterflies. Houghton Mifflin, Boston, Massachusetts. xvii+396 pp.     

Opler, P. A. & A. D. Warren.  2002.  Butterflies of North America. 2. Scientific names list for butterfly species of North America, north of Mexico. Contrib. C. P. Gillette Mus. Arth. Biodiv., Colorado State Univ., Ft. Collins, Colorado. 79 pp.  

Pelham, J. P.  2008.  A  catalogue of the butterflies of the United States and Canada, with a complete bibliography of the descriptive and systematic literature. J. Res. Lepid. 40:i-xiv, 1-652.

Penz, C.  2010.  Book review: the butterflies of Venezuela, Part 2: Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae, Ithomiinae, Morphinae), by Andrew F. E. Neild. J. Lepid. Soc. 64:51-51.   

Remington, C. L.  1968.  Suture-zones of hybrid interaction between recently joined biotas. Evol. Biol. 2:321-428.

Rutkowski, F.  1971.  Notes on some south Florida Lepidoptera. J. Lepid. Soc. 25:137-139.

Scott, J. A.  1986.  Butterflies of North America: a natural history and field guide. Stanford Univ. Pr., Stanford, California. xv+583 pp., 64 pl.

Sloane, 1725.  A voyage to the islands Madera, Barbadoes, Nieves, St. Christophers, and Jamaica; with the natural history of the herbs and trees, four-footed beasts, fishes, birds, insects, reptiles, &c. of the last of those islands. To which is prefix’d, an introduction, wherein is an account of the inhabitants, air, waters, diseases, trade, &c. of that place; with some relations concerning the neighbouring continent, and islands of America. Illustrated with the figures of the things described, which have not been heretofore engraved. In large copper-plates as big as the life. Vol. II. Author, London, England. xviii+499 pp., pls. 157-274.

Schwartz, A.  1989.  The butterflies of Hispaniola. Univ. Florida Pr., Gainesville, Florida. xii+580 pp.

Turner, T. W. & J. R. Parnell.  1985.  The identification of two species of Junonia Hübner (Lepidoptera: Nymphalidae): J. evarete and J. genoveva in Jamaica. J. Res. Lepid. 24:142-153.

Warren, A. D., K. J. Davis, N. V. Grishin, J. P. Pelham, E. M. Stangeland.  2010.  Butterflies of America.  Interactive listing of American butterflies.  Website: http://www.butterfliesofamerica.com. 

 

 

 

Figures

 

 

 

Explanation of Figures

 

1) dorsal engraving of Papilio evarete from Cramer ([1779]). 2) original dorsal drawing of P. evarete by G. W. Lambertz*. 3) male neotype of P. evarete (dorsal) from Suriname. 4) ventral engraving of P. evarete from Cramer ([1779]). 5) original ventral drawing of P. evarete by Lambertz*. 6) male neotype of P. evarete (ventral) from Suriname. 7) dorsal engraving of Papilio genoveva from Cramer ([1780]). 8) original dorsal drawing of P. genoveva by Lambertz*. 9) female neotype of P. genoveva (dorsal) from French Guiana. 10) ventral engraving of P. genoveva from Cramer ([1780]). 11) original ventral drawing of P. genoveva by Lambertz*. 12) female neotype of P. genoveva (ventral) from French Guiana. 13) dorsal engraving of J. evarete zonalis from Sloane (1725). 14) ventral engraving of J. e. zonalis from Sloane (1725). 15) earliest known specimen of J. evarete (dorsal) from Florida. (*© The Natural History Museum, London).

 

Junonia evarete zonalis, Miami-Dade Co., Florida: 16) dorsal male. 17) dorsal female. 18) ventral male.

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