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Megisto cymela viola (Maynard, 1891)
(Little Wood-Satyr)

Pinned Specimens photo collection:


 

Live Adults photo collection:

Distribution and Larval Foodplants:

documented from the northern half of the Florida peninsula

Diagnosis

This is a M. cymela subspecies in Florida. It is characterized by brighter, more contrasty appearance; more pronounced marking, such as larger eye-spots and wider, more bowed brown bands; and larger size.

Synonymy

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

Genus Megisto Hübner, [1819]

Verz. bekannt. Schmett. (4): 54. Type-species: Papilio eurytus Fabricius, 1775, Syst. Entomol.: 487, no. 194 (= Papilio cymela, Cramer, 1777, Uitl. Kapellen 2(11): 55, pl. 132, figs. C, D; (16): 148 (index); synonym), by designation of Butler (1868), Cat. diurn. Lepid. Sat. Brit. Mus.: 14.

 

Megisto cymela (Cramer, 1777)

Uitl. Kapellen 2(11): 55, pl. 132, figs. C ♂ D, D ♂ V; (16): 148 (index).

Original Combination: Pap[ilio]. Nymph[alis]. Gemm[ata] Cymela

Type Locality: “Kaap de Goede Hoop” “Cap de bonne Esperance” in error; neotype from “Jacksonboro, Colleton County, South Carolina.”

Types: Type(s) probably lost; neotype in MGCL, designated by Gatrelle (2005), Tax. Rep. Int. Lepid. Surv. 6(5): 3, figs. 3 ♂ D, 4 ♂ V.

=    ‡eurytus (Fabricius, 1775)

Syst. Entomol.: 487, no. 194.

Original Combination: P[apilio]. D[anaus]. F[estivus]. Eurytus

Type Locality: “Iamaica” This locality was probably reversed with that of the taxon that preceded it (Papilio zangis, no. 193, from “Carolina”), since that taxon is only known from Jamaica. Defined as “Goose Creek, Berkeley County, South Carolina” by Gatrelle (2005), Tax. Rep. Int. Lepid. Surv. 6(5): 2.

Types: Lectotype (#127231) in HMUG, designated by Gatrelle (2005), Tax. Rep. Int. Lepid. Surv. 6(5): 2, figs. 1♂ D, 2 ♂ V.

Preoccupied by Papilio eurytus Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 487, no. 180.

=    eurytris (Fabricius, 1793)

Entomol. Syst. 3(1): 157, no. 485.

Original Combination: P[apilio]. N[ymphales]. Eurytris

Type Locality: “Jamaica”, but see comment above.

Types: Lectotype (#127231) in HMUG, designated by Gatrelle (2005), Tax. Rep. Int. Lepid. Surv. 6(5): 2, figs. 1♂ D, 2 ♂ V. This is the same specimen as the lectotype of eurytus.

This is probably an incorrect subsequent spelling of eurytus, supported by the spellings “eurithris” Fabricius, 1796, Index alph.: 122, and “eurythris” Fabricius, 1797, Epitome entomol: 135. Gatrelle (2005), Tax. Rep. Int. Lepid. Surv. 6(5): 2, proposed an entirely different idea, with novel application of this name as senior to viola below.

 

Megisto cymela viola (Maynard, 1891)

Butts. N. England (ed. 2): 71, pl. 10, fig. 11 ♂ V; figured by Maynard (1891), Manual N. Am. Butts.: fig. 36b.

Original Combination: Neonympha eurytris viola

Type Locality: “Enterprise, Florida” [Volusia County]

Types: Holotype in MCZ.

This taxon has been treated as distinct species by Oliver (1982), J. Lepid. Soc. 36(2): 153, Opler and Krizek (1984), Butts. East Great Plains: 185, and Opler (1992), Field Guide E. Butts.: 212. Catling and Calhoun (1997), Holarct. Lepid. 4(1): 27, determined this taxon to be a subspecies of M. cymela occurring in peninsular Florida.

Bibliography

Original description from:

Maynard, Charles Johnson 1891. The butterflies of New England; with original descriptions of one hundred and six species, accompanied by an appendix containing descriptions of one hundred additional species. Illustrated with ten hand-colored plates, in which are given at least two hand-colored figures of each species (second edition). Newtonville, Massachusetts, C. J. Maynard: [4] + iv + 78 + [4] pp.; 10 pls. {1891}

p. 71

        NEONYMPHA  EURYTRIS  VIOLA, (Plate X,  Fig. 11) is larger in size than the typical northern N. eurytris, 
richer in color above,  where there is a violet iridescence and with more prominent markings above and be- 
low.     Habitat,  Middle  Florida.

Text from:

Maynard, Charles Johnson 1891. A manual of North American butterflies. Boston; De Wolfe, Fiske & Co.: iv + 226, 10 pls., 1 + 60 figs. {[May] 1891}

p. 109

       258a. NEONYMPHA  EURYTRIS  VIOLA Mayn., Fig. 36, b, differs from 
typical N. eurytris in being larger,  lighter in color, with spottings above 
and below very large and clear, while the small upper and lower spot on 
hind  wing  is minute,  and often missing.   Expands  l.60 to 1.75.   Hab- 
itat, Enterprise, Florida.  Specimens from which  I have made  the above 
description were obtained by my friend,  Mr. E. B. Clapp,  at Enterprise, 
Florida, in the spring of 1888,  and its distinctness from the more north- 
ern  form  appears  to entitle  it to the  sub-specific rank  given  it  in the 
Appendix of Butterflies of New England, page 71, December 1800. 

Text from:

Oliver, Charles G. 1982. Distinctiveness of Megisto c. cymela and M. c. viola (Satyridae). Journal of the Lepidopterists’ Society 36(2): 153. {24 Sep 1982}

p. 153



DISTINCTIVENESS OF MEGISTO C. CYMELA AND
M. C. VIOLA (SATYRIDAE)

Megisto cymela Cramer is remarkable for its lack of geographic variation over a range  
that extends from Manitoba and Quebec to the Gulf Coast. The subspecies M. c. viola  
Maynard, described from Florida, is highly distinctive, differing chiefly by its larger  
size and rich coloration on the ventral hindwing. Most references (e.g., A. B. Klots,  
1951, A Field Guide to the Butterflies, Houghton Mifflin, Boston, 349 pp.; W. H. Howe,  
1975, Butterflies of North America, Doubleday, New York, 633 pp.) consider that the  
name viola refers to all peninsular Florida populations and that M. c. cymela and M.  
c. viola blend phenotypically in southern Georgia and northern Florida.

An examination by me of series in The Florida State Collection of Arthropods,  
Gainesville, Florida, indicates that this is not the case. There is no sign of a phenotypic  
"blend zone," and populations of both typical M. c. cymela and M. c. viola exist in  
northern and central Florida. In addition, on the basis of data labels in The Florida  
State Collection, M. c. viola exists apparently sympatrically with M. c. cymela in south-  
ern Louisiana (Weyanoke, West Feliciana Parish) and Arkansas (Little Rock, Pulaski  
Co.). Moreover, in Florida M. c. viola appears strictly univoltine with a flight from late  
March to May. In the same general area M. c. cymela may have up to four broods with  
flights in April (Shalimar, Okaloosa Co.; Alachua Co.), July (Alachua Co.), October  
(Gainesvilla, Alachua Co.), and December (Sebring, Highlands Co.). The two entities  
thus may well be separate species.

Larvae of Pennsylvania M. c. cymela and Florida M. c. viola differ in life history in  
the laboratory. Although the pattern of larval markings is similar, larvae of M. c. cymela  
are a much darker shade of brown than M. c. viola. Broods of M. c. cymela derived  
from univoltine populations in Allegheny and Fayette Cos., Pennsylvania, develop  
without diapause and emerge as adults in 90 to 100 days when reared under conditions  
of 27°C days, 24°C nights and 16 hr light/24 hr. This correlates well with the three to  
four months between flights of M. c. cymela in Florida. Under the same conditions,  
however, growth of M. c. viola larvae from a Gainesville population differs greatly.  
Larvae hatched from eggs in early April 1979 grew at a very slow but steady rate and  
pupated and eclosed as adults in late February and early March 1980. This also cor-  
relates well with the flight time of M. c. viola in Florida if the cooler winter temper-  
atures in nature are taken into consideration.

Larvae of both entities were reared on potted Poa pratensis L., a grass native to the  
northern U.S. This could be a natural foodplant of M. c. cymela, but M. c. viola occurs  
south of its natural range. Nevertheless, M. c. viola larvae fed freely and produced  
adults of a size comparable to wild material. I consider it highly unlikely that differ-  
ences in development time were related to foodplant suitability.

Collectors in the Gulf Coast states should watch for localities where M. c. cymela  
and M. c. viola are sympatric and attempt to gather data on possible ecological differ-  
ences.
 

CHARLES G. OLIVER, RD. 1, Box 78, Scottdale, Pennsylvania 15683.

Authors' version of the manuscript published as:

Catling, Paul M. and John V. Calhoun 1997. Genus Megisto in Florida and the taxonomic status of Megisto viola (Lepidoptera: Nymphalidae: Satyrinae). Holarctic Lepidoptera 4(1): 27-33, 9 figs., 1 map., 2 tables {Mar, 25 Jun 1997}

p. 27-33, f. 1-10, t. 1-2, formatted by the authors, not as in the "Holarctic Lepidoptera", British spelling retained


GENUS MEGISTO IN FLORIDA AND THE TAXONOMIC STATUS OF
MEGISTO VIOLA (LEPIDOPTERA: NYMPHALIDAE: SATYRINAE)
 

PAUL M. CATLING1 AND JOHN V. CALHOUN2,3
 

1 2326 Scrivens Dr., R. R. #3 Metcalfe, Ontario K0A 2P0, Canada

2 977 Wicks Dr., Palm Harbor, Florida 34684, USA

 

ABSTRACT.– Megisto cymela (Cramer) and M. c. viola (Maynard) were studied to evaluate variation and subspeciation, using 274 specimens from Florida (239 from the peninsula and 65 from the panhandle) and 163 from the northern range limits in Ontario and Quebec. The few specimens suggesting multiple broods in Florida were either misidentified or probably mislabeled. The study sample and additional observations suggest a single flight period from late February to late April in the peninsula, late February to mid-June in the panhandle, and late May and June at the northern range limit in Canada. There are no constant morphological characteristics within the subspecific blend lone in northern Florida to suggest more than one species, but recognition of M. c. viola as a subspecies in peninsular Florida appears valid. Habitats and flight periods are also discussed.
 

KEY WORDS: Arkansas. Canada, cline, development, distribution, evolution, flight period, Georgia, Hermtuprychia, Louisiana, Nearctic, North America, Ontario, Pennsylvania, Quebec, symmpatry, USA, voltinism.

 

 

C. J. Maynard (1891a,b) briefly described Neonympha cymela viola Maynard, now in the genus Megisto, from specimens collected along the east coast of Florida, at Enterprise, Volusia Co. Subsequent literature (e.g., Klots, 1951; Emmel, 1975) and the most recent monograph of Megisto (Miller, 1976), indicate Megisto cymela (Cramer) to have two subspecies, the northern nominate M. c. cymela, blending into the southern subspecies, M. c. viola of Florida, southern Georgia and the Gulf coastal plain, west to southern Louisiana. The difficulty of assigning specimens from the blend zone to a subspecies caused Miller (1976) to doubt the validity of the two subspecies. However, some recent literature has elevated M. c. viola to species rank (Opler and Krizek, 1984; Opler and Malikul, 1992), based on the following observations by Oliver (1982):

(I) M. c. cymela and M. c. viola are sympatric in Florida, as well as parts of Louisiana and Arkansas, without a blend zone.

(2) The two subspecies have different flight periods in regions of symmpatry, M. c. cymela being multiple brooded while M. c. viola has a single brood.

(3) M. c. viola is highly distinctive, differing in its larger size and rich coloration on the ventral hindwing. To these differences, Opler and Malikul (1992) have added that M. viola has larger eyespots and a more strongly bowed postmedian line on both wings below.

(4) Larvae of M. c. cymela are a darker brown and have a shorter development time (90-100 days instead of 300-360 days in the lab), correlating with the multiple broods. Oliver considered it unlikely that differences in development time were related to foodplant suitability.

Scott (1986), on the other hand, suggested that the species level distinction had not been proved, and he noted that the genitalia did not differ. He also noted that M. c. cymela generally has only one flight; this latter point in particular casting doubt on Oliver's (1982) contention. Recently, Calhoun (1996) noted the presence of M. cymela in four counties in the Florida panhandle and observed that these individuals appeared intermediate between M. c. cymela and M. c. viola, casting further doubt on the species status of the latter. These differences between recent authors have highlighted Oliver's earlier (1982) suggestion, that researchers in the Gulf Coast region should further assess the ecological differences between the two taxa. Here, we evaluate data relating to the recognition of M. c. viola as a possible separate species relative to Oliver's (1982) observations.

Although specimens attributed to M. c. viola have been recorded as far north and west as Arkansas, we have focused our study on Florida, where the type locality is located.

 

 

METHODS

Collection dates, locations, collectors, forewing lengths, band widths, eyespot widths and sexes were recorded from 274 specimens of Megisto from Florida. The sample included 239 from the peninsula and 65 from the panhandle (west of the Suwannee River) as well as 163 from the northern range limit in Ontario and Quebec. The Florida sample included 128 specimens in the Florida State Collection of Arthropods (FSCA), at Gainesville; 51 in the Allyn Museum of Entomology, Sarasota; 76 in the collection of J. Calhoun; 49 in the collection of P. Catling; and 23 in the collection of Marc C. Minno, Gainesville, Florida. Methods of measurement of eyespot width (lower spot on underside of forewing), forewing length and eyespot band width (distance between postmedial and submarginal brown lines on underside of forewing) are shown in Fig. 1. Each of the three data sets were separated into males or females for comparisons, so that differences between sexes could be separated from geographically based differences and potential infrataxa differences. Plots and histograms illustrating eyespot widths, forewing lengths, and band widths, were examined for bimodal patterns suggesting more than one taxon. A plot of frequency by Julian date was prepared to evaluate night periods in each of the three regions. The relationship between eyespot size and flight dale was evaluated separately in 29 females and 48 males, all from the Gainesville area of Alachua Co., using linear regression. Distribution in Florida is illustrated using different symbols for private and institutional records.

 

 

RESULTS AND DISCUSSION

Possible Errors and the Number or Broods

Oliver's (1982) conclusion about symmpatry and multiple broods was based largely on 8 of 128 Florida specimens in the Florida State Collection of Arthropods (FSCA) in Gainesville. Most of these specimens corresponded to M. c. cymela, and all were collected by H. V. Weems. Jr. Six were from Alachua Co., one was from Highlands Co. and one was from Dade Co.

With respect to the Alachua Co. material, one of six specimens is labeled 10 July 1947, giving the impression of a second brood. However, a second brood in the peninsula, or anywhere in the cast, is not supported by other material in the Gainesville collection, or by additional material examined in other collections (Allyn Museum, Calhoun, Catling, CNC, Minno: Fig. 2-3). Furthermore, it is not supported by the observations of Florida field biologists (Minno, Calhoun, etc.). Of particular importance regarding the concept of a second brood in Alachua Co. are the observations of Mr. Richard Worth, who kindly made notes on the occurrence of Megisto in Gainesville (Williston Rd. and SW 34th street) during his continuous 1996 study of another satyrine, Hermeuptychia sosybius (Fabricius). He observed the first specimens of Megisto (males) on 25 March and the last (female) on 7 May (R. Worth, pers. comm.). Thus, in a single year, the flight period at a single location lasted 6 weeks. With variation in the weather from year the year, records from a single location could easily extend over a period of 8 or possibly even 10 weeks. Mr. Worth was at the site continuously (including July) and could recognize Megisto easily in flight. The fact that he did not encounter a second brood of Megisto at the large population within his study area is further evidence that one does not occur there. We suspect that the specimen collected by Weems allegedly from Gainesville in July 1947 is mislabeled.

The specimen from Highlands Co., collected at Sebring on 24 Dec 1947, is also problematic. It is entirely alone among the peninsular specimens with regard to its collection date (Fig. 2-3). Specimens from the nearest sites in Hardee and Polk counties were collected from mid to late March.

Oliver (1982) did not mention a specimen in the FSCA collection (from Ross-Costello Hammock collected by Weems on 27 May 1967. Ross Hammock and Costello Hammock were once more or less connected hardwood hammocks in the pinelands of Dade Co. between Miami and Homestead. Not surprisingly, for someone who collected many thousands of insects and was not a Lepidoptera specialist, Weems did not recall this or his other records of Megisto (pers. comm.). The Ross-Costello record is potentially of great importance, being the southernmost record of Megisto, and referable to M. c. cymela rather than M. c. viola. Ross-Costello Hammock is a park that had an active interpretive center and continues to be much visited and used by biologists. Neither M. c. cymela nor M. c. viola was listed among the butterflies recorded from the park and it was never seen there by active biologists in the area (R. Hammer, W. P. Gould, pers. comm.). In fact, the species has not been found in the Miami area despite over a century of extensive field studies. Thus, the Ross-Costello record is likely the result of mislabeling.

The only other reference to Megisto in Florida that would suggest multiple broods is that of Grossbeck (1917), based on records of William T. Davis and Charles E. Sleight at Ortega (Volusia Co.) on 6 Sep 1911 and La Grange (Brevard Co.) on 9 and 11 Sep 1911. The voucher specimens for these reports, in the collection of the Staten Island Institute of Arts and Sciences, New York, were kindly examined by curator Ed Johnson who reported that they were referable to Hermeuptichia sosybius and classified in the collection with that species.

With the Weems records under suspicion and the Grossbeck reference discounted, the available data, including more than 70 locations and 300 records (Fig. 2-3), suggests that Megisto does not "have up to four broods" in Florida as suggested by Oliver (1982), and is in fact univoltine with a single flight beginning in March and ending in June depending on latitude.

The mid-June specimens from the panhandle in Fig. 1-2 are from the inland Appalachicola River area (Chattahoochee and Torreya State Park), where the earliest specimens were collected in mid-April. Consequently, they fall within the 6-10 week flight period to be expected at any site, and do not necessarily represent a second brood. Moreover, most of these specimens are worn, aluding to the end of a single spring brood.

 

Quantitative Characters

Maximum diameter of lower eyespot on underside of forewing (mm), maximum length of forewing along costal margin (mm), and maximum width of submarginal band on underside of forewing (mm), all decrease in value northwards (Table I, Fig. 4- 6). Females are consistently larger in these characteristics than males. There was no bimodal pattern in characters within a region to suggest more than one taxon (Fig. 4-6). The dispersion of variation is approximately the same for Canada, the Florida panhandle, and the Florida peninsula (Fig. 4), and in each of these plots of forewing length versus eyespot width, only one discrete cluster is evident. Band width was correlated with eyespot width, but demonstrated a little less geographic variability. Although the pattern of increasing forewing size, band width and eyespot size southward is very clear, the broad overlap of Florida peninsula and Florida panhandle records with records from the northern range limit in Canada (Fig. 4-9) strongly suggests that the recognition of M. c. viola as a separate species is unwarranted.

Eyespot width had the most significant relationship to both latitude and Julian day (increasing with decrease in either). Forewing lengths and band widths demonstrate a similar pattern but were less significant and were not always significant for both sexes (Table 2). The association of eyespot width with flight period is probably related to climate, the relatively greater significance of Julian day being a consequence of the fact that latitude does not allow compensation for inland effects. Some inland peninsula locations are cooler and subject to more frequent frosts than coastal areas on the same latitude.

The female with the largest eyespots (Fig. 7) was found at Kelly Park, Orange Co., on 25 March 1996 and was worn at a time when males were prominent and fresh, suggesting that eyespots of exceptional size may be correlated with abnormally early emergence. However, we were unable to detect significant differences within a sex in the samples from Alachua Co. (P=O.18 for males, P=O.28 for females). The possibility of a significant relationship between eyespot diameter and flight time at a single location within the relatively long flight period needs to be assessed with more data from single sites, and could perhaps be achieved as part of mark-recapture studies so as to limit impact. Nevertheless the variation in eyespot diameter within a site on a single day also makes the formal taxonomic recognition of a species with large eyespots inappropriate. For example, on 2 April 1996, in San Felasco Hammock, Alachua Co., a male with unf lower eyespot 4.6mm in diameter (corresponding to some from Kelly Park, Orange Co., near the southern range limit), was flying with another that had an eyespot diameter of 3.2mm, which is consistent with some from the northern range limit in Ontario (Fig. 8-9).

 

Development Times

Oliver (1982) believed that the 90·100 day development time of M. cymela from Pennsylvania correlated with the alleged 3-4 months between flights in Florida, but it also correlates approximately with the length of frost-free period or growing season in Pennsylvania after June and before June the following year. The development times of M. cymela from Pennsylvania (Aowertown, Montgomery Co.) noted by Mr. Richard W. Boscoe were variable extending from 108-162 days (R. W. Boscoe, pers. comm.). Oliver (1982) recorded development times of approximately twice as long for Megisto from Gainesville, and Boscoe (pers. comm.) recorded development times of 192-291 days in eggs from a population in Withlacoochee State Forest, in Hernando Co., Florida. The longer development time of the Florida individuals correlates with the longer growing season there. Through either a shorter development time in the north or a longer one in the south, young larvae are able to take advantage of the relatively moist early spring period of fresh graminoid growth in deciduous forests. Adaptation of races to different lengths of growing season does not, however, require that the races be recognized as distinct species.
 

Evolutionary Considerations

For both male and female Megisto in both the Florida peninsula and panhandle there was a significant decrease in eyespot size in relation to increasing latitude and later collection dates, but a significant relationship was not found within the Canadian sample. This and the dispersion in Fig. 4 suggests that the cline exists mostly at the southern limit of the range of Megisto in eastern North America, and particularly within the Florida panhandle. Florida populations occupy more or less isolated southern hardwood forests, but interestingly the most extreme variants do not occur in the most well established area of relict flora and fauna in the Appalachicola drainage , but much further south in the peninsula. This suggests the possibility that the larger size and eyespots in the Florida populations may be a selected trait conferring advantage in predator evasion in a situation where the insects are more vulnerable to predators due to lack of leaves on most woody vegetation during the relatively early night period. There is much more leaf cover available during the later night periods northward, even in the Florida panhandle. The development time may also play a role. 

 

Distribution in Florida

Within Florida, Megisto occurs in areas of more or less isolated hardwood forests south to near the middle of the peninsula in northern Hardee Co., near the west coast, and 10 Orange Co., near the east coast (Fig. 10). It was not found in the region around Archbold Biological Station, Highlands Co., despite extensive field surveys in that area (M. C. Minno, pers. comm.).

 

 

CONCLUSIONS

The Florida specimens referred to M. c. viola are extremes of clinal variation and Florida populations overlap in variation in eyespot size, forewing band width and forewing length with similar amounts of variation further north, and even to the northern range limit in Canada. There is no evidence for two separate species in scatter plots of putatively distinctive characters in Florida and there is only one brood, although this brood may peak at different times over short distances in the Florida panhandle. Shorter development times in more northern populations may be explained in terms of adaptation to spring flush of growth in woodland graminoids and latitudinal differences in length of growing season. A significant decrease in eyespot size in relation to increasing latitude and later collection dates within both the Florida peninsula and the panhandle suggests that the cline exists mostly at the southern limit of the range of Megisto in eastern North America.

The pattern of geographic variation is of great interest in terms of evolution, and although taxonomic recognition of M. c. viola as a separate species appears inappropriate, subspecies rank seems valid. With a narrow cline in the Florida panhandle, it appears that M. c. viola may be limited to the northern half of the Florida peninsula. Additional research is needed on the relationships of Florida populations of M. cymela and those elsewhere along the Gulf Coast and lower Mississippi Valley.

 

 

ACKNOWLEDGEMENTS

We would not have been able to complete this work without the substantial help of several entomologists. Lee D. Miller and Jacqueline Y. Miller kindly provided data and measurements from specimens in the Allyn Museum of Entomology, Sarasota, Florida. Marc C. Minno (Gainesville, Florida) provided data from his personal collection, as well as a very useful review of the manuscript. Richard W. Boscoe (Lafayette Hill, Pennsylvania) provided data on reared specimens. Richard Worth (formerly Gainesville, Florida; now Belmont, California) made continuous observations over a period of several months in the Gainesville area and also collected specimens. Ed Johnson (Staten Island, New York) provided information concerning specimens in the Stalen Island Institute of Arts and Sciences. Information on butterfly occurrences at Ross-Costello Hammock was supplied by Roger Hammer and Waller P. Gould (Miami. Florida). John B. Heppner (FSCA, Gainesville, Florida) and Thomas C. Emmel (University of Florida, Gainesville) provided useful advice.

 

 

LITERATURE CITED

Calhoun, J. V.

1996. Florida - season summary. News Lepid. Soc. (Los Angeles). 38([3]):52.

Emmel, T. C.

1975. Family Satyridae. In W. H. Howe (ed.), Butterflies of North America, 79-111. New York: Doubleday. 633pp, 97 pI.

Grossbeck, J. A.

1917. In F. E. Watson (ed.), Insects of Florida. 4. Lepidoptera. Bull. Amer. Mus. Nat. Hist. (New York), 31:1·141.

Klots, A. B.

1951. A Field Guide to the Butterflies of North America East of the Great Plains. Boston: Houghton Mifflin. 349pp, 40 pI.

Maynard, C. J.

1891a. The Butterflies of New England (2nd ed.). Newtonville. 78pp, 10 pI.

1891b. A Manual of the North American Butterflies. Boston: De Wolfe. Fiske & Co. 226pp. 10 pI.

Miller, L. D.

1976. Revision of the Euptychiini (Satyridae). Bull. Allyn Mus. (Sarasota), 33:1-23.

Oliver, C. G.

1982. Distinctiveness of Megisto c. cymela and M. c. viola (Satyridae). J. Lepid. Soc. (Los Angeles), 36: 153.

Opler, P. A., and G. O. Krizek

1984. Butterflies East of the Great Plains. Baltimore: J. Hopkins Univ. Pr. 294pp, 54 pl.

Opler, P. A., and V. Malikul

1992. A Field Guide to Eastern Butterflies. Boston: Houghton Mifflin. 396pp, 48 pI.

Scott, J. A.

1986. The Butterflies of North America. Stanford: Stanford Univ. Pr. 583pp, 64 pI.




References:

Maynard, Charles Johnson 1891. The butterflies of New England; with original descriptions of one hundred and six species, accompanied by an appendix containing descriptions of one hundred additional species. Illustrated with ten hand-colored plates, in which are given at least two hand-colored figures of each species (second edition). Newtonville, Massachusetts, C. J. Maynard: [4] + iv + 78 + [4] pp.; 10 pls. {1891}. p. 71, pl. 10, f. 11

Maynard, Charles Johnson 1891. A manual of North American butterflies. Boston; De Wolfe, Fiske & Co.: iv + 226, 10 pls., 1 + 60 figs. {[May] 1891}. p. 109, f. 36 b

Oliver, Charles G. 1982. Distinctiveness of Megisto c. cymela and M. c. viola (Satyridae). Journal of the Lepidopterists’ Society 36(2): 153. {24 Sep 1982}

Catling, Paul M. and John V. Calhoun 1997. Genus Megisto in Florida and the taxonomic status of Megisto viola (Lepidoptera: Nymphalidae: Satyrinae). Holarctic Lepidoptera 4(1): 27-33, 9 figs., 1 map., 2 tables {Mar, 25 Jun 1997}.

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