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Journal of the Lepidopterists' Society JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Volume 13 1959 Number 4 STUDIES OF THE CHROMOSOMES OF NORTH AMERICAN RHOPALOCERA. 1. PAPILlONID~~ by KODO MAEKI and CHARLES L. REMINGTON This is the first of a series of papers on the numbers and morphology of the chromosomes of butterflies north of the Isthmus of Panama, with discus­ sions of the phylogenetic significance of the cytological data. The first few papers will include, family by family, material collected in three lots during the 1959 season: 1) around the Rocky Mt. Biological Laboratory in western Colorado; 2) in northwestern Mexico, mainly at Ciudad Victoria (Tamauli­ pas) and El SaIto (San Luis Potosi); and 3) around New Haven, Connecti­ cut. MAEKI collected most of the material in Colorado, assisted by REMING­ TON and ROGER W. PEASE, JR. The Mexican collection was made by REM­ INGTON. The Connecticut specimens were taken by MAEKI, REMINGTON, and PEASE. ERIC E. REMINGTON brought in important living males in all three regions, and Dr. ]. R. TURNER took several of the Ciudad Victoria males. New studies will be reported in supplementary papers. Our findings with the cytology of interspecific hybrids will appear elsewhere. METHODS The techniques will be outlined only in this first paper, because they were used identically for all the families of butterflies which we studied. The tests provide by far the easiest material in which to investigate the chromosomes of butterflies. First, meiotic divisions exhibit the chromo­ some complement in hwploid condition, so that unequivocal counts of the rather large number of chromosomes are much easier to obtain than in somatic or pre-meiotic divisions. Second, several or many divid.ing cells can usually be found in the male gonads, whereas female meiosis is limited to the maturation divisions in the egg near the time of entry of the sperm. Third, with Rhopalocera, unlike most other Lepidoptera, male meiosis is usually in process in some cysts of the testes even in old, flown adult males; notable ex­ ceptions are the Parnassiime, Zerynthiinle, Asiatic Graphium, some Hesperi­ idle, and Megathymidle, where meiosis is nearly over by the time of edosion. 193 194 MAEKI & REMINGTON: Chromosomes Vo1.13: no.4 The collection of testes is a simple procedure. The testes, which are fused into a single mid-dorsal unit in most Rhopalocera, lie near the dorsal wall of the abdomen about one-half to two-thirds of the distance from the base to the anal end. Wild-caught males are held firmly but unharmed and a small incision is made with sharp, fine scissors in the abdominal wall, over the nor­ mal site of the testes. A gentle squeeze of the abdomen is usually sufficient to exsert the testes, with little other tissue emerging. The testes of many species differ strikingly in color from the other abdominal contents and are then readily recognized. They are pinched off with a fine forceps and placed in a vial of fixative. We used Allen's P. F. A. - 3, which has the following form­ ula: Picric acid - 75 cc Formalin - 15 cc Glacial acetic acid - 10 cc Urea- 1 gram A key number is placed in the vial with the testes. The same number is imme­ diately noted along with locality and date on the paper envelope in which the subsequently killed male is placed for permanent reference. The testes are left in the fixative overnight, for up to 24 hours, and then transferred for storage into a vial of 80% ethyl alcohol. A few preliminary examinations of freshly collected testes were made by squash preparations in aceto-orcein and aceto-dahlia stain-fixative, but only with species of which we had large samples. All the studies here re­ ported are based on permanent preparations, made at the Gibbs Research Laboratories at Yale University, as follows: the testes were imbedded in paraffin, sectioned at 10,u thickness, stained in Heidenhain's iron ha:matoxy­ lin, and counterstained with Light Green. The slides were studied with a Leitz LABOLUX microscope. Suitable nuclei were drawn with camera lucida, with the microscope having 100X objective and 20X oculars ·and the drawing set-up giving on the paper a total magnification of 5800X. The photographs were taken with a Leitz MIKAS camera through the above microscope, using the 100X objective and a lOX ocular; the actual magni­ fication on the negative is 333X. One series of counts had to be made primari­ ly from sections of the ovaries of Parnassius smintheus, a species in which we found only a single partially satisfactory division in adult males (see below). The slides and specimens from which the gonads were taken ,are be­ ing kept for permanent reference in the research series in the Peabody Museum of Natural History of Yale University. The key numbers on these slides and specimens are noted below in square brackets - [ ] and will allow future workers to examine our material. In addition to the eleven species of Papilionida: described below, we fixed and sectioned testes of 3 Papilio rutulus from Colorado, 2 P. glaucus L. from Connecticut, and 1 P. anchisiades Esper from EI Salto but found no dividing cells. The first two species were very old specimens, but the P. anchisiades was obviously a young, fresh male, and it may be that this species normally completes meiosis before eclosion. 1959 Journal of the Lepidopterists' SOI-iety 195 RESULTS 1. Parnassius smintheus Doubleday. The haploid chromosome comple­ ment is n = 30. This was determined in diploid condition by 6 countable nuclei undergoing oogonial division in the ovaries of a young female with no sphragis (presumably unmated) collected at Copper Lake, e1. 10,500', Gun­ nison Co., Colo., on 7 August [199J. In each of these nuclei the 2n comple­ ment of 60 was visible. The apparent differences in size shown in the draw­ ing are probably due to the orientation of the chromosomes and not to actual differences. The base number was verified in haploid condition from one cell in diakinesis in which 30 chromosomes were seen; this was in a male from Cumberland Pass, 11 August [224]. Testes were also collected from 11 males from Copper Lake (27 July), Cumberland Pass (11 August), and Treasure Mt. (21 August), but no other suitable dividing cells were present. There were manv maturation divisions with abnormal meiosis leading to the forma­ tion of apyrene sperm. It is usual for Parnassius males to have little or no further normal meiosis after eclosion from the pupal shell. In order to study spermatocyte divisions we will need to collect testes from larvx and pupx. This species has recently been listed under the Palearctic phcebus (Fab.), but we consider this a premature step not justified without biological evidence. 2. Battus philenor (Linne). The haploid chromosome number is 30. Counts were made in 41 nuclei in 2 males, taken at Ciudad Victoria, Tamps., Mexico, on 1 August [MI2] and at EI Salto, S. L. P., Mexico, on 4 August [M53]. All suitable nuclei were primary spermatocyte divisions. No normal nuclei at the secondary spermatocyte division were found. The chromosomes of B. philenor are all similar in size. 3. Papilio ornythion Boisduval. The haploid chromosome number is 30. Counts were made in 15 nuclei in the primary spermatocyte division and 13 nuclei in the secondary spermatocyte division, from 4 males taken at Ciudad Victoria, 2 August [M40-A3, M40-A4, M40-A6, M40-A7]. The size of all the chromosomes is similar. Four other males taken with the above 4 showed no meiotic divisions. 4. Papilio cresphontes Cramer. The haploid chromosome number is 30. Counts were made in 8 nuclei in the primary spermatocyte division and 3 nuclei in the secondary spermatocyte division, from 2 males taken at Ciudad Victoria, 1 August rMII-2, MII-3], and 15 nuclei in the primary spermat­ ocyte division from a male taken at EI Saito, 4 August [M51]. A third male collected at Ciudad Victoria with the first two showed no normal divisions. There appear to be 4 chromosomes distinctly smaller than the other 26; the latter are similar in size. The identifications for P. cresphontes and P. thoas were verified from the male genitalia, which differ widely. 5. Papilio thoas autocles Rothschild & Jordan. The haploid chromosome number is 27. Counts were made in 36 nuclei, all in the primary spermatocyte division, from a single male taken at Ciudad Victoria, 2 August [M40-B]. Estimating chromosome size can be imprecise due to effects of variation in 196 MAEKI & REMINGTON: Chromosomes VoLl3: no.4 fixation and staining processes and the angle of the sections. Nevertheless, it is clear that only 2 very small chromosomes are present, and we estimate that there are 10 medium-sized, 13 large, and 2 very large chromosomes. Assum­ ing that the ancestral condition for thnas w as n = 30 with a caryotype simi­ lar to that of cresphontes, it is possible that the evolution of the present cary­ otype of thoas involved a) fusion of two of the four smallest chromosomes and b) fusions of fou r of the medium- or large-sized chromosomes to produce the two very large elements. As remarkable as the reduced chromosome number in thoas is the very large size of virtually all of its chromosomes, as compared to primary spermatocyte chromosomes in other Papilio. (See discussion below.) 6. Papilio polyxenes Fabricius. The apparent haploid chromosome num­ bers are 30 and 31. This is a remarkable situation and resembles the previous­ ly known case of Pieris rapce in Japan, discussed below. Four males of P. polyxenes taken in New Haven Co., Connecticut, 23 August to 7 September, gave counts as follows for 32 nuclei: Specimen 1st Division 2nd Division symbol n=30 n=31 n =30 n=31 [389J 2 3 0 0 [393] 9 5 0 0 [312] 4 0 3 0 [345J 1 1 4 0 Total nuclei: 16 9 7 0 The thirty-first apparent chromosome is a smaller body than the other 30 and different in its behavior.
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