Megathymus c. cofaqui (Strecker, 1876)
Distribution and Larval Foodplants:
[= harrisi], Georgia to NC
(From: A Catalogue of the Butterflies of the United States and Canada, Jonathan P. Pelham)
Genus Megathymus Scudder, 1872
4th Ann. Rep. Peabody Acad. Sci. (1871): 83. Type-species: Eudamus (?) yuccae Boisduval & Le Conte, , Hist. Lépid. Amér. sept. (23-26): pl. 70, figs. - ♂ [not ♀], larva and pupa, by original designation. I.C.Z.N. Opinion 483 placed this name on the Official List of Generic Names in Zoology as name no. 1222.
Megathymus cofaqui (Strecker, 1876)
Megathymus cofaqui cofaqui (Strecker, 1876)
Proc. Acad. Nat. Sci. Philad. 28(7): 148-150; figured by Strecker , Lepid. Rhop. Het. (15): pl. 15, fig. 2 ♀ D&V (holotype).
Original Combination: Ægiale Cofaqui
Type Locality: “Georgia”; defined as “somewhere in northern Florida near Georgia” by H. Freeman (1963), J. Res. Lepid. 2(2): 140; subsequently defined as “Burke County, Georgia” by Gatrelle (1999), Tax. Rep. Int. Lepid. Surv. 1(4): 3.
Types: Holotype in FMNH; figured by W. Barnes and McDunnough (1912), Contr. Nat. Hist. Lepid. N. Am. 1(3): 36, and Gatrelle (1999), Tax. Rep. Int. Lepid. Surv. 1(4): fig. 5 ♀ D.
= harrisi H. Freeman, 1955
Am. Mus. Novit. (1711): 2, figs. 3 ♂ D, 4 ♂ V (holotype); 7 ♀ D, 8 ♀ V, 27 ♂ genitalia, 28 ♀ genitalia.
Original Combination: Megathymus harrisi
Type Locality: “Stone Mountain, Georgia” [De Kalb County]
Types: Holotype in AMNH.
This synonymy determined by Gatrelle (1999), Tax. Rep. Int. Lepid. Surv. 1(4): 3.
Author's version of the text published as:
John V. Calhoun 2012. Notes on Megathymus yuccae as Illustrated by Boisduval & Le Conte (), with remarks about the holotype of M. cofaqui. News of the Lepidopterists' Society 54(1): 8-13, 15 figs.
p. 8-13, f. 1-15, formatted by the authors, not as in the "News of the Lepidopterists' Society", and with minor changes to the text
Notes on Megathymus yuccae as Illustrated by Boisduval & Le Conte (), with remarks about the holotype of M. cofaqui
John V. Calhoun
977 Wicks Dr., Palm Harbor, FL 34684, email@example.com
Plate 70 of the book Histoire générale et iconographie des Lépidoptères et des chenilles de l’Amérique septentrionale by Boisduval and Le Conte (1829-) portrays a butterfly that these authors named Eudamus (?) yuccae. This species is now recognized as the yucca giant-skipper, Megathymus yuccae. A small engraved inscription on the plate reads, "Abbot pinx.", meaning the illustration was derived from an original drawing by the artist-naturalist John Abbot ("pinx." is an abbreviation for the Latin "pinxit" = he/she painted it). The plate includes figures of the dorsal male, dorsal and ventral female, mature larva, pupa, and hostplant (Fig. 1). The engraver of this plate, known simply as Borromeé, rearranged the figures from Abbot’s original drawing—probably measuring about 25 x 34 cm—to accommodate the smaller format of the book (16 x 24 cm). Whereas the original drawing was probably rendered around the year 1820, the engraved plate was issued nearly two decades later (Boisduval & Le Conte ). No accompanying text was included, thus Plate 70 represents a description by indication; a named illustration with no written review of taxonomic characters.
Abbot’s original drawing is believed lost, but his accompanying notes are preserved in the Houghton Library, Harvard University. He called this species the "Great Georgia Skipper Butterfly" and wrote that it was "rare but most frequent in the lower parts of the Country." He credited John E. Le Conte, Jr. (co-author of Histoire générale) with discovering the caterpillar and its "manner of living." These naturalists first became acquainted in 1813, when Le Conte commissioned Abbot to complete a set of Lepidoptera drawings. In 1828, Le Conte traveled to Paris, France, to meet with the entomologist Jean B. A. D. de Boisduval. Le Conte took with him many drawings by Abbot, including those commissioned in 1813, as well as some that he had rendered himself. Boisduval decided to use the drawings for a book on North American Lepidoptera and he considered Le Conte, who lived in New York, as his co-author. Livraisons (parts) were issued beginning in 1829. Boisduval also intended to include moths, but he lost interest in the book and ceased its production in 1837.
Abbot’s drawings were the basis of 61 of the 78 hand-colored engraved illustrations in Histoire générale (Calhoun 2004). Many of these original watercolors are now preserved in the Irvin Department of Rare Books and Special Collections, University of South Carolina (Calhoun 2004). Among them is a life-sized drawing of a mature Megathymus larva, undoubtedly rendered by Le Conte (Fig. 12). Across the top of the drawing in Le Conte’s hand is written "Larva de le grande Hesperia [Larva of the large Hesperia]. Beneath the drawing, Le Conte wrote a brief Latin account of the life history of the species. Manuscripts at Harvard University reveal that the American entomologist Samuel H. Scudder sketched Le Conte’s Megathymus drawing during a visit with Boisduval in 1871. Scudder later sent his sketch to the entomologist Charles V. Riley, asking if it represented M. yuccae, to which Riley enthusiastically replied, "Without doubt!" Riley (1876) published a detailed treatise on the life history of M. yuccae.
A second species. Nearly 120 years after the publication of Plate 70 in Histoire générale, Evans (1955) realized that the figure of the dorsal female actually portrays the cofaqui giant-skipper, Megathymus cofaqui (Strecker). Unaware of this earlier determination, I reached the same conclusion (Calhoun 2004). The wing markings of this figure are very similar to specimens of cofaqui from Georgia, including those described by Freeman (1955) as Megathymus harrisi (Figs. 2, 3). The ground color of yuccae is dark brown, while that of cofaqui tends to be a warmer brown. This is reflected by the dorsal figures on Plate 70; the male is seal brown and the female is more russet. The forewing discal spots in nominotypical cofaqui are usually connected to the large spot in cell M3, forming an unbroken orange band across the wing. These pattern elements are separated in yuccae. A small submedian forewing spot in cell CuA2 (Cu2) is generally present in females of cofaqui, but is virtually always absent in females of yuccae. The female on Plate 70 bears a sizable spot at this location. The figure in Histoire générale is the earliest known depiction of cofaqui.
Megathymus skippers were a great oddity during the early 19th century and specimens were extremely rare. For many years Abbot and Le Conte were probably the only naturalists who knew of their existence. The first species to be described was yuccae, followed four decades later by cofaqui. They were so misunderstood that some early authors believed they belonged in the moth family Castniidae. Western taxa of Megathymus, and those of the related genus Agathymus, were mostly described during the 20th century. Only a handful of specimens of cofaqui are known from Georgia prior to 1950 (Skinner 1911; Harris 1931, , 1954, 1972). This species was first recorded in Florida around 1885 when it was collected by Charles W. Johnson in the vicinity of St. Augustine (Skinner 1911). Although the hostplant of cofaqui was suspected for many years (e.g. Riley 1882), it was not confirmed until Bonniwell (1916) found larvae and pupae in Florida on Yucca aloifolia L. Only after the biology of cofaqui was fully revealed by Bonniwell (1918) and Harris (1954) were numerous additional populations discovered, as adults are rarely encountered in nature. Many records are based solely on the presence of larval tents. The species was first recorded in South Carolina in 1976 (Mather 1977), in North Carolina in 1992 (K. Roever pers. comm.), and in Alabama in 1994 (Scholtens 1996; Howell & Charny 2010).
Source of the figured cofaqui. There are two plausible scenarios to account for the presence of M. cofaqui on Plate 70 of Histoire générale. The simplist explanation is that John Abbot collected at least one female cofaqui before 1820 and believed it represented yuccae. This would not be surprising, as cofaqui remained unrecognized until 1876. In addition, Abbot mentioned in his notes for his life history drawing of the "Great Georgia Skipper Butterfly" that one larva pupated on 17 May and the adult emerged on 20 June. This is very late for yuccae, which flies in Georgia from mid-February to early May (Harris 1972). Even in Florida, yuccae is not known to fly during June. This emergence date, however, may be applicable to cofaqui. In Georgia, adults of cofaqui have been recorded from early July to early September, but most records are from the piedmont and mountain regions. The species may begin to emerge earlier within the southern coastal plain. The entire flight period of cofaqui in southern coastal Georgia may be similar to that of the Florida panhandle, which is located within a similar physiographic region. Adults in the Florida panhandle fly from mid-June through early October, with occasional emergences in April (Zeiger 1979; M. Friedman pers. comm.; N. Grishin pers. comm.; K. Roever pers. comm.). Abbot perhaps intended to illustrate the life history of yuccae, but mistakenly inserted a female of an undescribed species. He did this on several other occasions, such as portraying two different Erynnis skippers as the male and female of the same species. The relationships between species were poorly understood and numerous taxa remained undescribed.
Although Abbot possibly encountered cofaqui in Georgia, I do not think that he mixed two species of Megathymus in his drawing used for Plate 70 of Histoire générale. Abbot routinely duplicated his compositions to more easily fulfill orders for his drawings. He maintained a book of template drawings which he used for many years (Calhoun 2007). Despite Abbot’s penchant for duplication, I have found no representations of cofaqui among his thousands of surviving Lepidoptera drawings. In fact, his only other illustrations known to portray Megathymus are two watercolors of yuccae at The Natural History Museum, London. Those drawings, probably rendered during the 1790s, accurately depict the dorsal and ventral aspects of two specimens, male and female, respectively. They were completed before the life history of yuccae was known. Abbot’s notes for these drawings, as transcribed by his London agent, John Francillon, read, "Taken 13th April in Oak Woods, sometimes in Fields near Swamps, very Rare." This date is consistent with the flight period of yuccae in Georgia. Abbot was then residing in Burke County, Georgia, not far from the town of Waynesboro (Fig. 15). He collected insects in Georgia primarily within the counties bordering the Savannah River, between Burke and Chatham Counties, westward to Bullock County. It was previously thought that Abbot lived and collected almost exclusively in Screven County. In reality, he resided there only for about three years, from 1813 to 1816.
The figures on Plate 70 of Histoire générale were credited to John Abbot, but it was probably Boisduval who was responsible for the inclusion of the female cofaqui. Issued in the tenth livraison of Histoire générale is a page entitled "Avis de l’un des Auteurs" [Opinion of one of the Authors], in which Boisduval announced, "I will touch on the nature of the drawings of Abbot, and I will completely redo them when they offer some inaccuracies" (translated from French). Plate 64, based on an Abbot drawing and issued with the 22nd livraison, apparently was altered because Boisduval disagreed with Abbot’s representation of the butterfly now recognized as Libytheana carinenta bachmanii (Kirtland) (Calhoun 2004). Boisduval probably was not familiar with this species and believed that Abbot’s figures were poor representations of the closely related Libytheana motya (Hübner), which was represented in his collection. It is conceivable that a female specimen of cofaqui likewise persuaded Boisduval to "correct" Abbot’s female figure of yuccae. Such a specimen most likely came from Abbot, or even Le Conte, whose family owned a rice plantation in southeastern Georgia, called Woodmanston, located near Riceboro in present-day Liberty County. In a letter to the American entomologist Thaddeus W. Harris, Le Conte remarked that he had sent Boisduval "complete collections" of Lepidoptera from New York and Georgia (18 July 1840; Ernst Mayr Library, Harvard University). Although Boisduval referred to Le Conte as his "collaborateur", Le Conte was not involved in the actual publication of Histoire générale. Le Conte provided drawings, specimens, and life history notes, but the plates and text were supposedly prepared entirely under Boisduval’s supervision (LeConte 1874). In fact, Boisduval did not provide Le Conte with all the parts of the book as published and he kept all of Le Conte’s materials (drawings and specimens) after production of the work had ceased (LeConte 1874; letter from J. E. Le Conte to T. W. Harris, ibid.).
Evidence suggests that the dorsal female on Plate 70 was only partially copied from Abbot’s original figure. The shape of the wings does not agree with the shorter and broader dimensions of cofaqui (Figs. 2, 3). However, the wing shape is virtually identical to Abbot’s earlier female of yuccae (Fig. 5), and the body is essentially copied from that of the male figure on the same plate. In keeping with his earlier compositions of yuccae, the original drawing for Plate 70 almost certainly included a female of yuccae. After learning the species’ life history from Le Conte, Abbot probably rendered figures of the early stages and combined them with adult figures of yuccae which he derived from his template drawings. The engraver of Plate 70 seemingly retained the wing shape of Abbot’s original figure, but changed the wing pattern and other elements at the behest of Boisduval.
The theory that Boisduval modified the dorsal female in Plate 70 is further supported by the gender of the ventral figure. During production of the plates for Histoire générale, Boisduval often altered the abdomens of Abbot’s figures to appear leaner (Calhoun 2004). This would explain why the abdomen of the ventral figure of Plate 70 is consistent with a male, but the wing shape and pattern are reminiscent of a female. This figure is comparable to Abbot’s earlier drawings of a female yuccae (Figs. 6, 7). If Abbot originally figured a female of cofaqui, logic dictates that he would have portrayed the ventral surface of that specimen, not that of another (i.e. a female yuccae).
Some lepidopterists have questioned the presence of the pale postmedian spot on the hindwing of the ventral figure on Plate 70 (Fig. 7). This spot is evocative of cofaqui, suggesting that Abbot rendered a composite or "hybrid" figure. However, this well-defined spot is undoubtedly an artifact of the reproduction process and does not accurately reflect a discrete character found on Abbot’s original drawing. This opinion is based upon Abbot’s earlier figure of the ventral female of yuccae, which includes a faithful representation of a small patch of pale scales (a "phantom spot") that is very distinct in some individuals of yuccae (Fig. 8). Because Abbot frequently duplicated his drawings, this feature was likely present on the ventral figure consulted for Plate 70. When reproducing this figure for the engraving, the colorists simply overemphasized this character. The engraved plates for Histoire générale include many such erroneous enhancements.
Finally, the larva on Plate 70 (Fig. 13) bears little resemblance to the actual larva of yuccae. Although it is difficult to determine the accuracy of Abbot’s original figure, the engraved version is consistent with his artistic style. He typically tapered the thorax of skipper larvae to create an exaggerated "neck". The prothorax is generally smaller than the head in most hesperiids, but the opposite is true in Megathymus. The shape of the larva on Plate 70 is basically an enlarged version of Abbot’s other hesperiid larvae (Fig. 14). The pupa is likewise similar in form to Abbot’s other hesperiid pupae. It is possible that Abbot did not personally rear this species, but based his figures and notes on information from Le Conte. The colorists possibly contributed to the inaccuracy of the larva on the plate, such as omitting the dark head and adding longitudinal bands.
It is very important to remember that embellishments were a common component of hand-colored engraved illustrations. The abilities of the original artist, as well as those of the engraver and colorists, must be considered. We are so accustomed to photographic reproduction that we routinely accept such images as accurate portrayals. Historical illustrations require that we refer back to the original drawings, or at least other drawings by the same artist, to help reveal the true nature of the subjects depicted.
The holotype of cofaqui. Strecker (1876) based his description of Aegiale cofaqui on a single female from Georgia. In his original description, Strecker perceived the similarity between the holotype and the female figured in Histoire générale. He wrote, "In markings of upper surface this species resembles somewhat closely the lowermost of Boisduval’s figures on plate 70…but the outline of the wings is entirely different." Strecker also cited the presence of the forewing spot in cell CuA2 of the figure, noting that Charles V. Riley, who had studied yuccae in detail, never found this spot on specimens of that species.
The holotype of A. cofaqui, now deposited in the Field Museum of Natural History (Chicago, Illinois; FMNH), was first figured by Strecker () (Fig. 4). Photographs of this specimen were subsequently published, but they were not in color (Barnes and McDunnough 1912; Harris 1954) or were unclear and did not include all the associated labels (Gatrelle 1999). With the help of James H. Boone of FMNH I obtained new high resolution images for review (Figs. 9-11). Strecker created large drawer labels which he placed at the head of each series of specimens. He also used smaller labels to denote type specimens. Both kinds of labels are associated with the holotype of A. cofaqui (Fig. 11). These labels show that the specimen was received from the ardent collector, Herbert K. Morrison. In a list of the type specimens contained in his collection, Strecker (1900) also attributed the specimen to Morrison, but offered no additional locality data. Strecker likely based the name cofaqui on the Muskogean settlement of Cofaqui, located near today’s town of Louisville in Jefferson County, Georgia. The explorer Hernando De Soto traveled through this area in 1540 (Hodge 1907), and Strecker may have become aware of Cofaqui from Irving (1857) who translated De Soto’s travels from contemporary accounts.
Many of Morrison’s letters to Strecker are preserved among the extensive Strecker correspondence at FMNH. The late F. Martin Brown organized these manuscripts during the early 1960s and his photocopies are now deposited in the archives of the McGuire Center for Lepidoptera and Biodiversity (Gainesville, Florida; MGCL). According to this correspondence, Morrison collected in Georgia and North Carolina from early February through most of September 1876. Only five of Morrison’s letters from this trip have survived, dated 3 March-13 April. All are postmarked from Parramore Hill, Screven County, Georgia (often spelled "Scriven" on older maps). Parramore Hill was situated along the Central of Georgia Railway, about 60 miles south of Augusta (Fig. 15). The site of this community is now located within Jenkins County, which was created in 1905. Morrison is not known to have visited Georgia prior to 1876; his letters from 1872-1875 were postmarked from Boston or Cambridge, Massachusetts, where he lived.
Morrison traveled to Georgia specifically to look for insects where John Abbot had explored many years earlier (Anonymous 1876a; Mann 1885). Morrison made a business of collecting insects and he offered Strecker many specimens from this trip, with the expectation that Strecker would advertize his collecting activities to other potential customers. On 23 March, Morrison advised Strecker, "I will not promise definitely to give you the rare things you may want, but if I hear from any that give me orders that it was from you they got their information, it would make a good deal of difference." A letter to Strecker from the entomologist Henry Edwards, dated 19 December 1876, reveals that Morrison offered as many as 250 specimens from Georgia for $25. During that same month, Morrison told Strecker that he had collected additional specimens of cofaqui, but had already sold them. Strecker doubtless never knew exactly where these specimens were found, as Morrison was notoriously secretive about his collecting localities.
Citing information from F. Martin Brown, Harris (1972) implied that the holotype of cofaqui most likely came from the vicinity of Parramore Hill. However, Morrison did not confine himself to that portion of Georgia. On 13 April 1876, he told Strecker, "I am going down the Ogeechee [River] this next week, I will not be in Parramore for a month to come – I think I shall get good things down there." The Ogeechee River flows from east-central Georgia to the Atlantic Ocean, just south of Savannah. In an unpublished circular sent to his entomological customers, dated, 1 September 1876, Morrison announced, "During the past summer I have been engaged in collecting the Insects of Southern and middle Georgia and of the Black Mountains of North Carolina." Although Gatrelle (1999) argued that there is no evidence that the holotype of cofaqui came from anywhere other than the vicinity of Burke/Screven Counties of Georgia, Morrison presumably explored a sizeable portion of Georgia, perhaps reaching deep into the southern counties. Gatrelle (1999) noted a resemblance between the holotype of cofaqui and specimens of the species from peninsular Florida, which he described as the subspecies M. c. slotteni. He downplayed this similarity under the assumption that the holotype was collected in east-central Georgia. Although Gatrelle (1999) defined the type locality as Burke County, Georgia, we should consider the possibility that the holotype was collected in southern Georgia.
The holotype’s date of capture also hints at a more southern origin. According to letters from Edward J. Nolan of the Academy of Natural Sciences of Philadelphia, Strecker presented his manuscript describing A. cofaqui on 29 June 1876. It was approved by the publications committee on 13 July 1876 (it was officially entered as a new manuscript on 4 July (Anonymous 1876b) and it was published the following September). Assuming that Strecker had previously completed the bulk of the manuscript, it is reasonable to assume that Strecker required several days to add the section on cofaqui and submit it for publication. Morrison’s letters indicate that he sent specimens to Strecker as he collected them in Georgia and it took five or six days for them to reach their destination in Reading, Pennsylvania. Consequently, the holotype was probably collected no later than mid-June 1876. This is consistent with the June emergence date in the notes for Abbot’s drawing of yuccae, which may refer to an individual of cofaqui.
Morrison also collected yuccae during his trip to Georgia. He mailed a postcard to Strecker from Parramore Hill, postmarked 1 April 1876, in which he announced, "Caught a perfect ♀ Castnia yuccae Bdv. Lec. today." Morrison offered the specimen to Strecker for two dollars—equivalent to about $40 today. Although Morrison mailed this and five other specimens to Strecker on 12 April, a search of the Strecker collection revealed no yuccae from Morrison. Instead, Strecker’s collection contains only three yuccae from Stewart County, Georgia, which were collected by A. W. Latimer, a druggist and publisher who lived in Lumpkin, Georgia and nearby Eufaula, Alabama. Many of Latimer’s letters to Strecker, dated 1875-1900, are preserved at FMNH (copies at MGCL). During the 1880s, Latimer sometimes wrote letters with the letterhead "A. W. Latimer, Collector and Dealer in Insects…Lepidoptera a Speciality." These letters reveal that Latimer sent many specimens to Strecker. In a letter dated 7 April 1879, Latimer included a sketch of a female yuccae that he had reared from a pupa. This excited Strecker, who repeatedly asked Latimer to send specimens. After many failed attempts to rear more adults (most were parasitized), Latimer sent several yuccae to Strecker in 1882. The absence of Morrison’s "Castnia yuccae" in Strecker’s collection implies that this was the specimen that Strecker described as A. cofaqui. Although populations of cofaqui in coastal Georgia may present a rare spring emergence, a capture date as early as 1 April is improbable. Strecker sold and exchanged many specimens from his collection, thus it is more likely that Morrison’s yuccae was given to another entomologist. Strecker’s correspondence also reveals that in 1876 he received several Florida specimens of yuccae which were collected by the Ohio naturalists Charles Dury and Edward Mills. These specimens were possibly discarded, as they were not in perfect condition.
Morrison’s relationship with Strecker was tumultuous. After providing Strecker countless specimens over the course of a decade, Morrison died of dysentery on 15 June 1885 at the young age of 31 years. Two weeks later, Strecker received a letter from Morrison’s wife, Ada, which included a bill for $35.50 toward outstanding debts. On the envelope, Strecker curtly scrawled, "no answer…I don’t owe the wretches a cent."
Acknowledgements. Thanks are extended to Nick Grishin and Andrew D. Warren for permission to reproduce an image from the Butterflies of America website (http://www.butterfliesofamerica.com/). Elizabeth Sudduth (Irwin Dept. of Rare Books and Special Collections, Univ. of South Carolina) granted permission to publish J. E. Le Conte’s drawing. James H. Boone (Dept. of Zoology, FMNH) arranged for photography of the holotype of M. cofaqui and searched for specimens in the Strecker collection. C. Howard Grisham provided data from specimens collected by Ronald R. Gatrelle and others. James K. Adams, MaryAnn Friedman, Nick Grishin, Harry E. LeGrand, Jr., and Killian Roever kindly shared their observations of M. cofaqui. Finally, Jacqueline Y. Miller and Andrew D. Warren permitted access to documents and specimens at MGCL.
Anonymous. 1876a. [H. K. Morrison collecting insects in Georgia and North Carolina]. Can. Entomol. 8:239.
_____. 1876b. [Proceedings of] July 4. Proc. Acad. Nat. Sci. Phil. 28:140.
Barnes, W. & J. H. McDunnough. 1912. Revision of the Megathymidae. Contrib. Nat. Hist. Lepid. No. Amer. 1(3):1-43, pls. I-VI.
Boisduval, J. B. A. D. de & J. E. Le Conte. 1829-. Histoire générale et iconographie des Lépidoptères et des chenilles de l’Amérique septentrionale. Librairie Encyclopédique de Roret, Paris. 228 pp., 78 pl.
_____. . Livraison 24. Pp. 205-212, pls. 70-72. In Histoire générale et iconographie des Lépidoptères et des chenilles de l’Amérique septentrionale. Librairie Encyclopédique de Roret, Paris.
Bonniwell, J. G. 1916. Location of pupae of Megathymus cofaqui (Lep.). Entomol. News 27:372.
_____. 1918. Collecting on the Gulf coast of Florida in the spring of 1917. Lepid. (Boston) 2:108-109.
Calhoun, J. V. 2004. Histoire générale et iconographie des Lépidoptéres et des chenilles de l’Amérique septentrionale by Boisduval & Le Conte (1829-): original drawings for the engraved plates and the true identities of four figured taxa. J. Lepid. Soc. 58:143-168.
_____. 2007. John Abbot’s butterfly drawings for William Swainson, including general comments about Abbot’s artistic methods and written observations. J. Lepid. Soc. 61:1-20.
Evans, W. H. 1955. A catalogue of the Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and Megathyminae. Brit. Mus. (Nat. Hist.), London. vi+499 pp., pls. 54-88.
Finley, A. 1823. [Map of Georgia]. The author, Philadelphia.
Freeman, H. A. 1955. Four new species of Megathymus (Lepidoptera, Rhopalocera, Megathymidae). Amer. Mus. Nov. 1711:1-20.
Gatrelle, R. R. 1999. Subspecific status of southeastern U.S. Megathyus cofaqui and M. yuccae: renaming of the Florida subspecies of M. cofaqui. Taxon. Rpt. 1(4):1-6.
Harris, L., Jr. 1931. A list of the butterflies of Georgia. Trans. Georgia Nat. Club 1:1-27.
_____. . The butterflies of Georgia. Revised. Bull. Georgia Soc. Nat. 5:1-33.
_____. 1954. An account of the unusual life history of a rare yucca skipper (Megathymidae). Lepid. News 8:153-162.
_____. 1972. Butterflies of Georgia. Univ. Oklahoma Pr., Norman, Oklahoma. xvi+326 pp., 23 pl.
Hodge, F. W. (ed.). 1907. Handbook of American Indians north of Mexico. Part 1. Govt. Printing Off., Washington, D.C. ix+972 pp.
Howell, W. M. & V. Charney. 2010. Butterflies of Alabama. Pearson Learning Solutions, Boston, Massachusetts. 509 pp.
Irving, T. 1857. The conquest of Florida, under Hernando De Soto (revised edition). G. P. Putnam & Co., New York, New York. xxii+457 pp.
LeConte, J. L. 1874. Hints for the promotion of economic entomology. Proc. Amer. Assoc. Adv. Sci. 23(II [B]):10-22.
Mann, B. P. 1885. Herbert Knowles Morrison. Psyche 4:287.
Mather, B. (ed.). 1977. Zone 6. South Carolina. The annual field summary for 1977. News Lepid Soc. No. 2 (Mar/Apr):14.
Riley, C. V. 1876. Notes on the yucca borer, Megathymus yuccae (Walk.). Trans. Acad. Sci. St. Louis 3:323-344.
_____. 1882. Further notes on the pollination of Yucca and on Pronuba and Prodoxus. Proc. Amer. Assoc. Adv. Sci. 29:617-639.
Scholtens, B. G. (ed.). 1996. Season summary. Zone 9. Alabama. News Lepid. Soc. (Apr.):51.
Skinner, H. 1911. The larger boreal American Hesperiidae, including Eudamus, Erycides, Pyrrhopyge and Megathymus. Trans. Amer. Entomol. Soc. 37:169-209.
Strecker, F. H. H. 1876. Description of a new species of Aegiale and notes on some other species of North American Lepidoptera. Proc. Acad. Nat. Sci. Philadelphia 28:148-153.
_____. . Pp. 135-143, pl. XV. In Lepidoptera, Rhopaloceres and heteroceres, indigenous and exotic; with descriptions and colored illustrations. Author, Reading, Pennsylvania.
_____. 1900. List of types in the collection of Lepidoptera of Dr. Herman Strecker. Lepidoptera, rhopaloceres and heteroceres, indigenous and exotic. Suppl. 3. 37 pp.
Zeiger, C. (ed.) 1979. Current zone reports and planned activities. Zone III. Florida. So. Lepid. News. 1(2):4.
1) Plate 70 of Histoire générale. 2) Dorsal female M. cofaqui from Plate 70 of Histoire générale. 3) Female (dorsal) M. cofaqui, Fulton County, GA (MGCL) (© Nick Grishin). 4) Female holotype of Aegiale cofaqui, as figured by Strecker (). 5) Dorsal female M. yuccae by J. Abbot, ca. 1792*. 6) Detail of ventral female M. yuccae by J. Abbot, ca. 1792* (arrow indicates "phantom spot"). 7) Detail of ventral female M. yuccae from Plate 70 of Histoire générale (arrow indicates overemphasized spot). 8) Ventral female M. yuccae, Charleston Co., SC (MGCL) (arrow indicates well-defined "phantom spot"). 9) Holotype (dorsal) of A. cofaqui (FMNH). 10) Holotype (ventral) of A. cofaqui (FMNH). 11) Labels associated with the holotype of A. cofaqui (FMNH). 12) Megathymus larva by J. E. Le Conte. 13) Larva from Plate 70 of Histoire générale. 14) Examples of hesperiid larvae by J. Abbot. (*© The Natural History Museum, London).
Fig. 15. Early map of Georgia (Finley 1823) with locations relevant to this study (inset shows area covered).