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	1967	Journal of the Lepidopterists' Society	59
	VARIATION IN JUNONIA COENIA IN MISSISSIPPI (NYMPHALIDAE) Bryant Mather Box 2131, Jackson, Mississippi Concerning Junonia coenia, Klots (1951) wrote: aWe do not yet fully understand the relationships of the various forms of this butterfly. . . . A great deal of careful study and thorough analytical work is needed." The present paper discusses results of a study of some aspects of variation in Junonia coenia coenia (Hiibner) in Mississippi. The 105 Mississippi specimens which were studied were collected over a 10-year period, represented dates of capture in all 12 months and localities in 17 counties representing all sections of the state, and included 52 males and 53 females. Of the 105 specimens, 83 were taken in Hinds County and a total of 86 in the southwestern region of Mississippi as defined by Mather and Mather (1958), 12 were from the southeastern region, three from the northwestern, and two each from the northeastern and east central re­gions. No indication of intra-state geographical variation was detected. Features examined included: (a) Length of right forewing, measured with dividers and scaled to the nearest 1.0 mm; (b) Underside ground color (dark, intermediate, light); (c) Presence of subapical spot on forewing upperside as seen by the unaided eye (absent, faint, present); (d) Maximum diameter of each eyespot on the hindwing upperside, measured to the nearest 0.1 mm using an eyepiece micrometer and a magnification of 10 X; and (e) Ratio of diameters and areas of the eye-spots on the hindwing upperside. Data on each of these features are presented and discussed below. Forewing length The range of forewing lengths was 19 to 31 mm, distributed by month of capture as shown in Table 1; mean monthly values are indicated by open squares. The mean forewing length increases gradually from a minimum of about 21 mm in January to a maximum of about 27 mm in October, after which it decreases rapidly to about 24 mm in November and 22 mm in December. The peak in October and rapid decrease there­after may coincide with the beginning of cold weather; the earliest date for killing frost in central Mississippi is October 10 and the average date is November 5. In Virginia, Clark and Clark (1951) reported mean forewing lengths for the typical spring and fall form as 24 mm in males, 27 mm in females,

	60	Mather: Buckeye variation	Vol. 21, no. 1
	Table 1 Distribution of 105 Specimens of /. c. coenia from Mississippi  by Length of Forewing and Month in Which Taken, Showing Seasonal   Change in Mean Forewing Length number of specimens ForeWiNG LENGTH „ MM. iq 20 21 22 23 24 25 2& 2? 28 2^ 3o   31 ToTAL JAM I I H I I                                           [ 8 FEB I I B                                                     \ S MAR I I □ |                                                 3 APE I I El 3 I I                                M4- MAY I [¥] I                                            5 J UN 2 H] I                                      \ S JUL l[2]                                                            3 Aos I i 2 H] I                           7 SEP I 3 5 [7] I 2                          13 OCT I 2 3 [I] I I Z    I 13 NOV I I 3 D I ' '                      8 DEC 2 2 50112                                          21
	Total         S 6 12 14 15 IS 15 |o 4- 4 2 2 I Q - MEAM FoREWiN'6 LBN6TH oF SPECIMENS TAKEN IN INDICATED MON'TH			lOST
	and noted that summer specimens are a little larger and individuals ol another form confined to localized wet areas are still larger, about 28 mm for males and 30 mm for females. Field (1940) reported size values for the Kansas population, normal range 45 to 50 mm, 60 mm for large females, and 35 mm for "dwarfs." Assuming that these may be converted to forewing lengths by subtracting 2 mm from each value for thorax width and dividing the remainder by two, the normal Kansas forewing length is 21.5 to 24 mm and the extremes are 16.5 and 29 mm. It would

	1967	Journal of the Lepidopterists' Society	61
	Table 2 Distribution of 105 Specimens of /. c. coenia from Mississippi by Depth of Ground Color Below and Month in Which Taken, Showing Seasonal Change in Depth of Ground Color Underside
	JAnI Fee MAR Apr MAY JvjfJ JUL AU6 SEP Oct Nov £>£c total	| WRK IN TERMEPMH6 1 U6tfT i Wo % Mo % Nk> % ( 7 8fi I 12 1 £ loo - - 2 66 1 33 - I 7 3 Z| 10 72 -> - 5" loo - - S |0O - - 3 loo - 1 14- 6 86 8 6| l 8 4 31 6 4S S 4o 2 IS 3 37 4 So 1 13 1 18 8$ 2 IO I 5" 5o 18 37
	therefore appear that the individuals comprising the Mississippi popula­tion are somewhat smaller in size than those of the Virginia population and very slightly larger than those of the Kansas population; and that our sample includes three specimens that are larger than the largest Field found and none as small as the smallest he found. Ground color of underside The distribution of specimens by underside ground color and month of capture by number and percentage is shown in Table 2. No speci­mens with light ground color below were among those taken in January, February, or March; none with dark ground color below were among those taken in May, June, July, or August. The major seasonal shift from dark to light appears to take place between March and April, and

	62	Mather: Buckeye variation	Vol. 21, no. 1
	lOO -r
	LIGHT
	r ■. •/•:			INTER­MEDIATE,
	DARlC
	w
	J F rAAhAJJ A S o N X> M oNTH Fig. 1. Percentage distribution by months of capture of Mississippi specimens of /. c. coenia having light, intermediate, and dark ground color on underside.
	from light to dark between August and September; the former being somewhat more abrupt than the latter. In central Mississippi the March-April shift coincides with a change in mean temperature from below 60° F to above 60° F (16° C), and the August-September shift with a change from above 80° F to below 80° F (27° C). The mean rainfall

	1967	Journal of the hepidopterists' Society	63
	for March is higher than for any other month (5.9 inches), and is lowest for October (2.5 inches). The August-September shift does not accom­pany an increase in average monthly rainfall; the mean values for August, September, and October being 4.0, 3.1, and 2.5 inches respectively. I have observed no association between depth of ground color below and moistness of habitat in Mississippi. The percentage relations of speci­mens of different depths of ground color below by months are shown in Figure 1. SUBAPICAL SPOT ON UPPERSIDE OF FOREWING Of the 105 specimens examined, 53 were classified as having the spot well developed; in 40 it was present but faint; and in 12 it was not visible to the unaided eye. Examination of the latter at a magnification of 30 X revealed that at least a few blue scales were present or that the specimen was so rubbed that it might be assumed that such scales could once have been present. The twelve specimens showing no visible spot had forewing lengths from 19 to 29 mm, six had forewing lengths of 19, 20, or 21 mm; thus suggesting a tendency for obsolescence of this spot to be associated with diminution in size. Relative size of spots of hindwing upperside The measured values for the maximum diameters of anterior and posterior spots for the 105 specimens are plotted in Figure 2. Solid triangles represent males, open circles represent females. For each speci­men represented by a spot in Figure 2 the ratio of the diameters of anterior to posterior spot was calculated and a frequency diagram of these ratios is given as Figure 3. Figure 3 indicates a skewed distribution. The most frequent ratio of diameters for the Mississippi population is about 1.65, with more specimens having values larger than this and fewer having smaller values. The diagram also suggests that the speci­men having the smallest ratio (1.26), reported previously (Mather and Mather, 1958) as suggestive of the phenotype of zonalis is more clearly a part of the Mississippi population as a whole than the specimen at the other end of the distribution which has a ratio of 2.59. In Figure 4 the points representing the four Mississippi specimens yielding the ex­treme data points in Figure 2 are replotted and values derived from one additional specimen in our collection, nine specimens figured in the literature (see Table 3), are indicated. The three specimens figured by Holland (1931) and Klots (1951) that are assigned to zonalis have ratios between 1.15 and 1.29. The two Mississippi specimens with the lowest diameter ratios are 1.26 and 1.36. The position in Figure 4 of the points for the six figured specimens of coenia suggest that the individuals com-

	64	Mather: Buckeye variation	Vol. 21, no. 1
	2.5                    3.o                  3,5"                   4,0                  4-S"                  5.o                  S.5 Dipy/v^Tf?/^ <of Posterior eye spot d miv) Fig. 2. Relation of maximum diameter of anterior and posterior eyespots on hind-wing upperside for 52 male and 53 female specimens of /. c. coenia from Mississippi, showing extreme ratios of diameters of 1.26 : 1 and 2.59 : 1.

	1967		Journal of the Lepidopterists' Society			65
	7 U Hi Q_ LL O
	Hi CD
	0,8		1.2 L4- 1-6 1-8 2-0 2-2 2-4- ?■& RATio of EYE SPoT PiRPOET^^S			Z8
	X			JL
	0.64 l.o 1.44 |.^6 2,sq 40 s.z<? RP,TiO OF EYE SPcT AR^S				676 784
	Fig. 3. Frequency distribution for ratios of diameters and ratios of areas of anterior and posterior eyespots on hindwing upperside for 105 specimens of /. c. coenia from Mississippi.
	prising the Mississippi population may have ratios that are slightly lower than average for a "typical" coenia population. Both Klots (1951) and Brown (1955) characterize coenia as having the anterior spot on the hindwing above "from two to three times as large as the posterior one" or "at least twice and often three times as large." This contrasts with zonalis in which the anterior spot is described as "little larger than the posterior one." Barnes and McDunnough (1916,

	Table 3 Relation of Maximum Diameter of Anterior and Posterior Eyespots on Hindwing Upperside in Mississippi J. coenia and Specimens Figured in the Literature as J. zonalis. Data Plotted in Figure 4
	DiArv\ET* R. OF SPOT? ETER5 MM DATE -taken* LOCALITY RjTFE-ReNiCEL A,MT- 1 ERioR. Post­erior A 4.3 23 1-87 2o 7«>vjM iq^ CUNTonI, (VMS5. 1 6 4.8 3.8 \n(o 25 |0 SEP |<H<? CLINTOK, tV\^S. MATHE-i? AND MftTMER. 0^S&"> " L\KtT ZOKADS " c q.z 5.3 174 3o 2o OCT 1^57 6R0WrtSV/lLLF,WS.S. \d q.3 4.s 2.o7 2q 22 SEP 1^57 PETERSB0R6, VA- E 5". 7 2-2 2.5<? 11 8 Ao£ 1^54- Clinton,, iv\\ss. F (oA 2* 2.2o - — CHAPEL HILL, N.C KLOTS (|95l) , PL 14 , ^\<k IZ "cpetnia " ^ G 4.5" 3.5 I-2t - — — HOLLAND (iqsO,?1--^.^ 9 "6ENOVEVA" CT* (<*) H 44 3.4 l-2i - — T\T05V)LLC( FLA- KLOTS O55-0-5 ?L- 14- , R|^ >3 "Zonalis " ^ I 4.o 3.5 l-is- - - — HOLLAND cIR3D;PL-XX, FiG S "LAx/jNiJl" cr^ fc) J 7-o 3.£> |.<?4 - iq SePiqz5 CASiM ODhlN, /V\J). CLARK (1932.) ^ PL. 7 , Fl<^ "2> 11 CoENfi_g " D«/ FoRrA ^ K 75 AS 1-^7 - — — Bro^JM C&5S) ? p loA " Lftv/imiQ f CoErJiQ " L 8.5 A.£> 1-85 - 255&P iq25" CABiK JOMN, •**>. CLARK AMO CLARfc. (i9SD 5 PL-5, Frf<3- 3 "coisniQ " wet form ?■ M <f.O s.o 1-80 - — — Holland (\wf)? Plx*, fi<s 7 "CQENift " ?- r 4.9 1.(o l-fc<? - — — Mftcy AMD 5BEPARD0940, pL • I " ©OCKEYE " CCQgNift^
		< o
	(?) BOTH REFERABLE TO ZONALIS (KLqTS , 19 Si)

	1967	Journal of the Lepidopterists Society	67
	X -----O-----------------------------7 1 D / ] M / / / - / Dj / J / °F / ▲ Mississippi cPc? "j A E O Ml S5I5S IPPI $> X VIRGINIA Q FIGURED IN LITERATURE! / dn <4 / <v HI 2 3 4 ^ <£ 7 <9
	3                       4                       5"                       <£ J)lAfNAPTER OP POSTERIOR EY5 sPoT ,		M M
	Fig. 4. Relation of maximum diameter of anterior and posterior eyespots on hind-wing upperside for four Mississippi specimens of /. c. coenia, one from Virginia, six figured in the literature, and three specimens figured in the literature representing /. zonalis. Data for these fourteen points are given in Table 1.
	Plate IX, Figs. 6, 7) figure a female coenia from Long Island, N. Y. and a female genoveva from Miami, Florida. The latter is similar to that figured by Klots (1951) from Titusville, Florida as zonalis. Barnes and McDunnough emphasize that the feature selected to differentiate geno­veva from coenia was the replacement by orange suffusion in genoveva of the brownish encirclement bounded outwardly by the black of the large eyespot on the dorsal surface of the forewing in coenia. This dif­ference is clearly shown in Klots' figure but not emphasized by him in the text. The Mississippi specimen (Fig. 5 B) having hindwing eyespots in the size relation of figured specimens assignable to zonalis does not

	68	Mather: Buckeye variation	Vol. 21, no. 1
	Fig. 5. Five Mississippi specimens of /. c. coenia. Specimens A, B, C, E yielded data shown in Figure 4; A has the smallest anterior spot of the sample; B has the

	1967	Journal of the Lepidopterists' Society	69
	have the forewing eyespots surrounded by orange, but it also lacks a clear development of the blacker outer boundary of the brownish zone sur­rounding this spot. The eyespots in the hindwing upperside in coenia are not exactly circular but were considered to be so for the calculation of areas from diameters. If two spots have the same diameter they were assumed to have the same area. In the absence of qualifying comment to the con­trary I would assume that authors who describe a spot as "twice or three times as large" as another spot must refer to the area relations of the spots in question. For this reason, Figure 3 has been provided with two scales, one showing ratios of diameters, the other showing ratios of areas computed from the diameters. The range in ratios of diameters of spots of specimens yielding data plotted in Figure 4 is from 1.15 to 2.59, hence the range in areas ("sizes") is from about 1.3 to 6.9. The ratio of spot diameters of the figures of specimens of coenia given in Klots (1951) and Brown (1955) is respectively 2.20 and 1.67 which correspond to 4.8 and 2.8 in ratios of area. The most frequent diameter ratio for the specimens of the Mississippi sample is about 1.65 which corresponds to a ratio of areas ("sizes") of about 2.7. No reference has been found in the literature to a specimen of coenia having an anterior spot that has an area approximately seven times that of the posterior spot (Fig. 5 E). Based on the data and the relations presented in Figures 2, 3, and 4, it is suggested that zonalis may be characterized in part as having hind-wing eyespots above in which the diameter of the anterior spot is less than 1.4 times (or the area less than twice) that of the posterior spot; while coenia may be characterized, in part, by spots in which the diam­eter of the anterior spot is more than 1.4 times (or the area more than twice) that of the posterior spot and including individuals in which these ratios go up at least to 2.6 and 7 respectively. Figure 5 is a photograph of five Mississippi specimens including those representing the observed extremes of variation in the dimensional rela­tions of the eyespots in the hindwing upperside. Those designated A, B, C, and E define the limits of the observed data for the Mississippi sample as these limits are shown in Figure 4. In specimen B the larger spot is about one and one-half times as large as the smaller spot; in speci­men E the larger spot is nearly seven times as large as the smaller; these two specimens account for the two bars in the graph (Fig. 3) that are separated from the mass of the data for Mississippi.
	<- smallest ratio of diameters (1.26 : 1); C has the largest anterior and posterior spots; E has the largest ratio of diameters (2.59 : 1) and the smallest posterior spot; X has the smallest space between the anterior and posterior spots.

	70	Mather: Buckeye variation	Vol. 21, no. 1
	Summary The Mississippi population of Junonia represents a single variable population that may be called Junonia coenia coenia (Hiibner). J. c. coenia is a population, the individuals of which characteristically have hindwing eyespots of such proportionate sizes that the diameter of the anterior spot is more than 1.4 times, and its area is more than twice, that of the posterior spot. All but two of 105 Mississippi specimens studied had hindwing spots with ratios of diameters of 1.4 or more. The Mis­sissippi specimens have the hindwing spots of upperside somewhat more nearly equal in size than appears to be the case for coenia in the rest of its range. The Mississippi individuals range in size from those having a forewing as short as 19 mm to others having a forewing as long as 31 mm. The size increases from a January average minimum of about 21 mm to an October average maximum of 27 mm and then decreases rather rapidly. The ground color of underside is predominantly dark from September through March, and light from April through August. More individuals having light ground color persist after August than do individuals with dark ground color after March. No association of ground color of underside with moistness of habitat or variation of aver­age rainfall was found. The change in depth of ground color of under­side with season appears more abrupt than does the change of size with season. A majority of the Mississippi specimens showed a well-developed subapical spot in the forewing above; in most of the remaining specimens such a spot was visible but faint; in 12 of 105 specimens it was not visible to the unaided eye. An association of tendency for obsolescence of this spot with diminution of wing length was noted. Literature Cited Barnes, W., & J. H. McDunnough. 1916. Notes on North American diurnal Lepi- doptera. Contrib. to the Nat. Hist, of the Lepid. of No. Amer., 3(2): 53-152 (p. 101, PL IX, Fig. 5, 6). Brown, F. M. 1955. Colorado butterflies, part II, Danaidae, Heliconidae, Nym- phalidae. Denver Mus. Nat. Hist. Proc., 4: 33-112. Clark, A. H. 1932. The butterflies of the District of Columbia and vicinity. U. S. Natl. Mus. Bull. 157, 337 pp. Clark, A. H., & L. F. Clark. 1951. The butterflies of Virginia. Smithson. Misc. Coll., 116 (7), 239 pp. Field, W. D. 1940. A manual of the butterflies and skippers of Kansas (Lepidop- tera, Rhopalocera). Bull. Univ. Kans., 39, 328 pp. Holland, W. J. 1931. The butterfly book (revised edition). Doubleday & Co., Garden City, N. Y., 424 pp. Klots, A. B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Boston, 349 pp. Macy, R. W., & H. H. Shepard. 1941. Butterflies. Univ. of Minn. Press, Minne­apolis, 247 pp. Mather, B., & K. Mather. 1958. The butterflies of Mississippi. Tulane Univ. Stud. Zool., vol. 6, pp. 63-109.