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Hybrids Between Papilio Xuthus and the P. Polyxenes-Machaon Group

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Hybrids Between Papilio Xuthus and the P. Polyxenes-Machaon Group 1960 Journal of the Lepidopterists' Society A STUDY OF HYBRIDS BETWEEN PAPILlO XUTHUS AND THE P. POLYXENES-MACHAON GROUP by SHIGERU ALBERT AE Papilio xuthus Linne (pU: fig.2) is found commonly but only in East Asia. It is distributed in Japan, Korea, Manchuria, Amur, China, North Burma, Formosa, Luzon, Guam, etc. It is very common in most parts of Japan, but becomes rare toward the northern part of Hokkaido. It seems also uncommon in hot regions, as in Formosa. In Japan, it is usually not found in the high mountains, unlike P. hippocrates Felder & Felder (Fig. 1), which is found in the high mountains as well as the lower places. The adult of P. xuthus resembles the adult of the P. machaon group, and P. xuthus is usually placed in the P. polyxenes-machaon group. However, the pupa and larva of P . xuthus do not resemble the pup~ and larv~ of species of that group. They rather resemble the pup~ and larv~ of P. protenor demetrius Cramer, P . memnon thunbergii von Siebold, P. helenus nieconicolens Butler, and P. macilentus Janson in Japan. The pup~ and larv~ of these four Japanese black swallowtails resemble each other closely. The larval and pupal resemblances between P. xu thus and P. helenus Linne were pointed out by REMINGTON (1960). These four black swallowtails are Rutace~ feeders as is P. xuthus. Therefore, the phylogenetic position of P. xuthus in the genus Papilio is in­ teresting and a study of it may help in analyzing the evolutionary process of species in the genus. Dr. C. L. REMINGTON obtained very important data in this problem by interspecific crossings between P. xuthus and the polyxenes-machaon group ( 1(60). Namely, he succeeded in obtaining one male hybrid between P. hip­ pocrates and P. xuthus and two male hybrids between P. polyxenes Linne and P. xut hus. Since P. hippocrates is very common in Japan as well as P. xuthus. it is very easy to work on hybridization of these two species in Japan. Therefore, the writer obtained many hybrids of this kind in 1958 and 1959 at Nanzan University, Nagoya, Japan, following Dr. REMINGTON'S work in 1957. The writer also obtained some hybrids between P. xuthus and P. poly­ xent'S (Fig.3 ) and between P. xuthus and P. "brucei Edwards" (FigA). The data are not sufficient to give a final conclusion on the relation between P. xuthus and P. machaon relatives, and crossing experiments are still in pro­ gress in the writer's laboratory, along with hybridization tests between P. xuthus and the four above-mentioned Japanese black swallowtails. However, it takes years to complete this kind of work. Therefore, the writer presents here the data mainly in genetics of interspecific characters on the hybrids be­ tween P . xuthus and P. machaon relatives obtained in 1958 and 1959, so that they are available for the workers in the same field. AE: Papilio xu/hus hybrids Vol.1+: no.1 Pup.:e of P. polyxenes and P. "brucei" were reared by the writer in 1957 at Yale University and the Rocky lVlountain Biological Laboratory re­ spectively and sent to the writer at N anzan University in winter by the kind­ ness of Dr. REMINGTON. The so-called" brucei" is the high mountain Papilio of Colorado and is considered by R EMINGTON not to be true brucei (Maeki & Remington 1960: p.196). A permit to import these pup.:e to Japan was given to N anzan University by the Agricultural Ministry of Japan. P. hippo­ crates pup.:e were obtained by the rearings of local wild larv.:e and eggs from local wild females in the fall of 1957 and 1958, and wild P. xulhl/s pup.:e were collected from Karatachi (Poncitrus trifoliata Rafin., Rutace.:e) fences in Nagoya and Osaka in the winters of 1957-1958 and 1958-1959. Wild adults of P. hippocrates and P. xuthus in the vicinity were also used in 1958 and 1959, especially males. The prefix of individual designation used for P. "brucei", P. hippocrates, P. p'olyxenrs, and P. xuthus are respectively B, H, P, and X. The individual clesignation of a female was also used as the designation of its brood. \Vhen a female mated twice or more, this brood was separated; e.g. H-3a and H-3b. All the matings were obtainecl by hand-pairing (Clarke & Sheppard, 1956). Eggs were obtained from females in glass cylinders on potted food plants. The containers were placed under incandescent and fluorescent lights. For the rearing of larv.:e, insect cages about 30 cm. cubed were used. These experiments were carried on exclusively in a double screened rearing room to prevent any accidental release of American butterflies in 1958. This room was windowless and lighted by fluorescent and incandescent lights usually during the day time but with no definite period. The rearing of the larv.:e of the hybrids between P. hippocrates and P. xuthus in 1959 was carried on at the windows ide of the laboratory. There was no control of temperature or humidity, and no heater was used in either room. REMINGTON (1960) pointed out that P. hippocrates should be placed as a species separate from P. machaon, and the writer has no objection on it. HYBRIDS BETWEEN Papilio xuthus AND P. hippocrates Nine P. hippocrates females were hand-paired with P. xuthus males in 1958. Only two of the females, H-3 and H-Il, laid fertile eggs. Broods H-3a and H-l1 were obtained from them. One more female laid a few eggs but they were all infertile. H-3 was remated with another P. xu/flus male and later again with a P. hippocrates male, and Broods H-3b and H-3c were obtained. Seven P. Xl/thus females were hand-paired with P. hippocrates males in 1958. Four females laid eggs, but only one, X-26, laid fertile eggs. The fertility of matings is shown in Table 1. An egg is yellowish white when laid, and if it starts a development, a brown mottling or ring appears (Rem­ ington, 1960). This phenomenon was used as an indication of fertilization. When a larva is well formed in an egg shell, the egg color turns black. All larv.:e which hatched in Brood X-26 were put into plastic containers with Seri (ill nanthe stolonifera DC., U mbellifer.:e) and Karatachi. The 1%0 Journal 0/ the Lepidopterists' Society 7 larv~ which chose Karatachi were reared on Karatachi. The number of larv~ which started to feed on Seri was larger than the number which started to feed on Karatachi. Fifteen larv~ of Brood H-3a were reared on Seri, 14 on Natsumikan (Citrus natsudaidai Hayata) and 4 on Karatachi, respectively. The larv~ which were reared on N atsumikan were moved to Seri at their 4th instar, because of the shortage of fresh N atsumikan leave'S. The larv~ of H-3b were reared on Mitsuba (Crytotcl'nia japonica Hassk., U mbellifer~), and the larv~ of H-3c were reared on Serio The number of pup~ which were obtained from these rearings and the pupal lengths are shown in Table 3. Although the eggs of Brood H -3c were laid after the female mated with a wild P. hippocrates male for more than one hour, the larv~ from them were identified as hybrids. Broods H-3a, X-26, and X-27 (from eggs laid by a wild P. xu thus ) were reared at the same time (April to :YIay) , and the pup~ of X-27 were larger than the pup~ of the hybrids. Therefore, the small size of the hybrid pup~ are not seasonal. Since three pup~ of P. xuthus, which were obtained from the larv::c reared on Seri, were as large as the other P. xut hus reared on Karatachi, effects of food plant on hybrid larv~ do not seem to be the cause of the small size. [Po hippocrates larv~ feed on Umbellifer~ and occasionally on Rutace~, and P. xuthus feeds on Rutace~ in nature in Japan (Esaki & Shirozu, 1951) 1. No difference in developmental rates was observed in these rearings. The P. hipp·ocrates larv~ were not reared at the same time, but their summer form, which comes from spring larv~, is usually larger than the summer form of P. xuthus. The hybrid butterflies which emerged from these small pup~ were consequently very small. Loss during the larval stage in hybrids was noticeable. Some definitely died of diseases, but some deaths could have been due to hereditary causes from hybridization. Some pup~ were malformed to some extent. The larv~ of Brood H-11 were reared outdoors under a net and fed on Natsumikan branches. Some of them re'ached the 4th instar, but all died without having another molt. Somc- additional data were obtained in 1959. Ten P. hippocrates females were hand-paired with P. xul fLus males. Seven of the females laid eggs, and the eggs laid by five of them contained some fertile ones. Twelve P. xuthw females were hand-paired with P. hippocrates males. Nine of the females laid eggs, and the eggs laid bv four of them contained some fertile eggs. Table 2 shows the fertility of these matings. Only one larva, which hatched from the eggs laid by female H-29-13, was reared on Seri and reached the pupal stage. This was sent to Mr. K. lV1AEKI for chromosomal study. The two larv~ which hatched from the eggs laid by female H -29-16 were killed accidentally. Fifteen larv~ which hatched from the eggs laid by female H-34-5 were reared on Seri, and 6 pup~ were obtained; four of them emerged.
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