Chromosomes of North American Rhopalocera. Part 3
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1960 Journal 0/ the Lepidoptl'rists' Society 127 STUDIES OF THE CHROMOSOMES OF NORTH AMERICAN RHOPALOCERA. 3. LYClENIDlE, DANATNlE, SATYRINlE, MORPHINlE by KODO lVIAEKI and CHARLES L. REMINGTON This paper is a continuation of our first two, which gave the sources of most of our material, our techniques, and an account of the chromosomes of Papilionidce, Pieridce, Hesperiidce. and Megathymidce (Maeki & Reming ton, 1960a, 1960b). The subject of the fourth, concluding, paper of this series will be the remaining groups of the family Nymphalidce. In general, EHRLICH'S reclassification (1958a, 1958b) of the Rhopalocera is accepted, and we follow him in placing the Riodinids in the Lyccenidce and the Danaids, Satyrids, and Morphids as subfamilies of the huge family Nymphalidce. The inclusion of the Lyccenidce in the present paper is not m~ant to suggest close phylogenetic affinity with the other groups but is merely for convenience in fitting the size requirements of this J ourna!. Similarly, the Pieridce were paired with the Hesperioidea in our preceding paper only for convenience. Most of the specimens for which the chromosomes are reported here were taken in 1959. We now are processing many additional species from North America, for which counts will be published in a supplementary paper, and there are 4 Danaince and 1 Satyrince in our forthcoming report on S. A. Hessel 's 1960 African material. As before, the numbers in square brackets are the designations of indi viduals studied and can be found on the specimens and slides preserved for permanent reference in the Peabody "Museum of Yale University. In the fol lowing lists "n" represents the haploid chromosome number, "I" refers to the primary ,permatocyte division and "TI" to the secondary spermatocyte di vision. No female meiosis was investigated in these groups. Fixed testes of several species not listed below were sectioned and ex amined but did not show any meiotic divisions. For some of these species we have studied enough testes to conclude that divisions are rare or absent after eclosion of the imago (or barely possibly that divisions are limitf,d to a brief time of day when we would not normally be dissecting and fixing material, such as 5 :00 a.m.). The numbers of specimens of these species are as fol lows: several I ncisalia augustinlls (Westwood) , I. polios Cook & Watson, and 1. irus (God'Ht) from Connecticllt and New Jersey, 6 Florida Satvrill m favonius (J. E. Smith) , 7 Colorado CE neis Chr-YXllS Doubledav, 4 Colorado Erebia raflias Edwards, 3 Colorado E. epipsodea Butler, 11 Florida Eupty rhia hermes (Fabricius) , and 3 Florida E. areolata (J E. Smith). Fewer testes were examined but all gave negative results for the following: 1 Colo rado Plebejus scc pio/us (Boisduval), 1 Colorado Chrvsophanus titw (Fab ricius), 2 New Mexico Sandia macfarlandi Ehrlich & Clench, 1 Colorado CEneis uhleri Reakirt, 1 Colorado Ca:nonympha ochracea Edwards. 128 MAEKI & REMINGTON: Chromosomes Vo1.14: no.2 A. L YClENIDlE: 1. Calephelis virginiensis (Guerin). N = 45. Counts were made in 3 nuclei (I) and 4 nuclei (II) in testes of one male [F81 J taken in the Cork screw Swamp, near Immokalee, Collier Co., Florida, 19 April 1960, leg. K. Maeki. There are 9 chromosomes distinctly larger than the remainder. This is the first Riodinid species to be studied cytologically. 2. Lycccna helloides (Boisduval). N = 24. Counts were made in 28 nuclei (I) and 18 nuclei (II) in testes of one male [63] taken at Gothic, el. 9500', Gunnison Co., Colorado, 20 July 1959. About 7 chromosomes are large, about 12 are medium, and about 5 are smaller in size. There is some uncertainty about the placement of this Colorado form in L. hello ides or L. dorcas (Kirby) (see Clench, 1958: p.59). 3. Lycccna nivalis (Boisduval). N = 24. Counts were made in 106 nuclei (I) and 68 nuclei (II) in testes of 11 males [9, 29-1, 29-2, 29-3, 34-1, 34-2, 34-3, 34-5, 99, 101, 102] all taken at Gothic in July 1959. Six other males showed no meiotic meta phases. The size relations of the chromosomes are similar or identical to those of L. helloides. One anomalous metaphase plate at the second division in male No. 99 shows 25 elements, instead of 24; the other nuclei have 24 elements in this individual. There is a possibility that this male is a hybrid. 4. Lyccena snowi (Edwards). N = 24. Counts were made in 12 nuclei (I) in the testes of one male [183] taken at Copper Lake, el. 11,000', Gunnison Co., Colorado, 1 Aug. 1959. No suitable secondary spermatocyte divisions were found. As with L. helloides, there are 7 large chromosomes. Probably L. snowi is a race of L. cupreus (Edwards), but for the present we list it as distinct. 5. LYC<Ena hypophleas (Boisduval). N = 24. Count·s were made in 31 nuclei (I) and 28 nuclei (II) in the testes of 3 males [331, 333, 381] taken in New Haven, Connecticut, September 1959. Two chromosomes are dis tinctly smaller than the remainder; the others are uniform in size and are rather large. We classify hypophleas as a full species until its genetic relations with Palearctic forms are better known. 6. Lycccna rubidus (Edwards). N = 38. Counts were made in 87 nuclei (I) and 52 nuclei (II) in testes of 5 males [175, 190-1, 190-2, 190-3, 190-4] taken at Gothic, 31 July and 2 Aug. 1959. There are 6 or 7 large and 5 or 6 small chromosomes. r 7. LycrCna heteronea (Boisduval). N = 68. Counts were made in \ 106 nuclei (1) and 93 nuclei (II) in the testes of 9 males [64-1, 64-3, 64-4, 64-5,77-1,77-2,77-3,71-1, 92-1] taken at Gothic on 20 and 22 July 1959. Two chromosomes are very large,S or 6 medium, and the others small. 8. J caricia icarioides (Boisduval). N = 24. Counts were made in 23 nuclei (I) in testes of one male [106-2] taken at Gothic, 23 July 1959. There are 7 large, 13 medium, and 4 small chromosomes. Two other males showed no countable nuclei. CHROMOSOMES PLATE 1 .- .-. ~ -.- :.-. •••••.. .. • •••• ••-.. •••••• .. :::.. • ••• •• • • • ••••• ••••••.... .. :• • •••••• .....• ••• : . ..... - ••• • • ••• ••• •• .: .. •••••• .11 • ••• •• • Ib 2a 2b la • •• • •• • •••• • •••• • • •••••• • • ••••• • •• • ••••• • •••• • ••• • • •••• •• •• •••• ••• ••••• •• • 3b 3c 3a ••••• • • • : • •••• •••••• •• • ·• •••.... ••• •• • • •••••• • •••• •• • • ••• • • ••• ••••• • • 5b 4 .- 5a • • • •• • • ••••••••• • • • .- -..... •••• •••• • •••••• -•..... ••••• . ..- .-....... ,.... :.. .... - -.. -......• •• . • -.... 7a • ••• • 6a •• • . .....•••• . • • •••••••• . ... •• •••••• -.: : ..... • ••• e •• : .....• ~:.::.: • •• • -. ... • • •• .•• ..... •• . .. , • • 7b . 6b Fig. 1a - ealephelis 'Virginiensis (I); fig. 1b - same (II); fig. 2a - Lycama hello ides ( I ) ; fig. 2b - same (II); fig. 3a-L. ni'Valis (I); fig. 3b - same (II; n= 24); fig. 3c - same (II; n = 25); fig. 4 - L. sllowi (I); fig. Sa - L. hypophleas (I); fig. 5b - same (II); fig. 6a - L. rubidus (I); fig. 6b - same (II); fig. 7a L. heterollea (I); fig. 7b - same (II). 129 130 MAEKI & REMINGTON: Chromosomes Vo1.14: no.2 9. Lycceides melissa (Edwards). N = 24. Count~ ; were made in 18 nuclei (1) in testes of one male [114J taken at Gothic, 24 July 1959. ~o suitable m::taphase nuclei in the secondary division were found. The size relations of the chromosomes appear to be exactly like those of I. icarioides and A. glandon. 10. A griades glandorz (de Prunner). N = 24. Counts were made in 13 nuclei (II) in testes of one male [93J taken at Gothic, 22 July 1959. No countable nuclei in the primary division were found. A second male showed no meiotic divisions. This is rusticus (Edwards), presumed to be a race of the holarctic A. glandon. II. F:veres amyntula (Boisduval). N = 24. Counts were made in 26 nuclei (J) and 17 nuclei (II) in testes of 2 males [109, 110J taken at Gothic, 2\ J ulv 1959. There are 5 chromosomes distinctly larger than the other 19; the latter are rather uniform in size. A third male showed no meiotic divisions. 12. Everes romyntas (Godart). N = 2+. Counts were made in II nuclei (I1) in testes of 2 males taken in New Haven, Connecticut, 3 Sept. [319-2] and 19 Sept. [395 J 1959. No suitable primary divisions were found. No countable nuclei were found in testes of 4 other males taken with No. 319-2. There appear to be 4 large, 18 medium. and 2 small chromosomes, and this seems to be a significant distance from the 5-19-0 size relations of E. amyntula. The species distinctness of these two Everes has been questioned by several authors (e.g., Clench, 1958: pp. 59-60). 13. Eumceus debora (Hubner). N = 24. Counts were made in 66 nuclei (1) and 22 nuclei (II) in the testes of one male [MI6] taken at Ciudad Victoria, Tamaulipas, :Mexico, 1 Aug. 1959. The caryotype includes 8 or 9 larger, 11 or 12 medium, and 4 smaller chromosomes. 14. Satyrium sylvinus (Boisduval). N = 41. Counts were made in 10 nuclei (I) and 14 nuclei (II) in the testes of one male [282 J taken near Somerset, Gunnison Co .. Colorado, 15 Aug. 1959. There appear to be about 10 chromosomes distinctly larger than the others, but all are large. IS. Calycopi.f cecroj>s (Fabricius). N = 24. Counts were made in 15 nuclei (I) and 7 nuclei (TT) in testes of 2 males [FI54, F192J taken at the Archbold Biological Station, Highlands Co., Florida, 21 and 22 April 1960, leg. K. Maeki. 'Many additional suitable nuclei were present and could have been counted. There are 20 large and 4 small chromosomes. A third male taken at the same time had no meiotic divisions, only maturing sperms. 16. Strymo71 melinus Hubner. N = 24. Counts were made in 16 nuclei (I) and 34 nuclei (II) in testes of 3 males taken on West Rock, New Haven, Connecticut, 23 Aug.