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Karyotype Alteration and Phylogeny, VI Karyotype Analysis in Palmae by Dyuhei Sato Receivednovember 12, 1946

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Karyotype Alteration and Phylogeny, VI Karyotype Analysis in Palmae by Dyuhei Sato Receivednovember 12, 1946 174 Cvtologia 14 Karyotype Alteration and Phylogeny, VI Karyotype analysis in Palmae By Dyuhei Sato ReceivedNovember 12, 1946 I Introduction The karyotype, namely, chromosome morphology seems to be more essential and less changeable characteristic than the other external mor phology or phenotype. The so-called karyotype analysis has been believed to be one of the important methods to discuss the phylogeny. Accordingly the system in monocotyledons has to be reinvestigated from the stand point of the karyotype alteration and phylogeny. O. DRUDE mentions that Pallmae forms a very particular group free from other families, but HUTCHINSON (1932) suggests the origin of this family which is derived from the Liliaceous stock, to be perhaps through part of the Agavales (cf. Dracaena, Yucca, Cordyline, etc.). The writer (SATO 1935, 1938, 1942) clearly showed the phylogenetic significance of the Yucoa-Agave karyotype which supported the HUTCHINSON's opinion and here undertakes to pursue the origin and further development of this and similar karyoypes, namely, Liliales-Agavales-Palmales. The chromosome numbers in Palmae were insufficiently reported by several workers, NEMEC 1910, SODERBERG1919, SUSSENGUTH1920, RADER MACHER1925, SANTOS 1928, SINOTO 1928, PATEN and NARAYAMA1937 and BEAL 1937, because the cytology of tropical plants is confronted by the difficulty of obtaining the favourable materials, more properly speaking, difficulty of cytological observation based upon bad condition of fixation, abundance of small chromosomes, agglutination of chromosomes, etc. In the present case the writer reports on the materials which were certainly observed cytologically. II Material and Methods Some species reported here were obtained in the Ponape Island and the other species were pot plants most of which were raised in the Ko isikawa Botanic Garden of the Tokyo Imperial University and also in the Kosobe Green House of the Kyoto Imperial University. For the species names the label names on the seed bags imported were adopted in most 1946 Karyotype Alteration and Phylogeny, VI. Karyotype analysis in Fa mae 175 cases. The root-tips were fixed with NAVASHIN'S solution, and the para ffin sections were cut 15 micra in thickness and NEWTON'S gentian violet method was used for staining. III Observation Various symbols for the description of karyotypes have been independently employed by many investigators. Accordingly the general reprt sentation of karyotypes was proposed by SINOTO (1944) in details for the prevention of such confusion and then this method of abbreviation may serve for the comparison of various results of different authors in the future. In the description of karyotypes the letters A, B and C denote respectively long, medium and short chromosomes, for which the letters, L, M and S have been used in previous papers of the present -writer (1935-1944). When only long and short chromosomes are present, the letters A and C instead of A and B are employed for the distinct differ ence between them and also for the comparison of similar karyotypes. To distinguish the Sat-chromosomes from other chromosomes the suffix cs or t is added on the left shoulder (or foot) of the abbreviated letters above mentioned, and then the former means the Sat-chromosome with a second ary constriction at the short arm (or long arm) and latter the,Sat-chromo some with a trabant. 1 Corypheae Trithrinax brasiliensis 2n=36(2b)=8A+8B+2esB+18C (Fig. 1). This species has thirty-six somatic chromosomes of which three pairs of long chromosomes have median constrictions and one pair of long chromosomes and five pairs of medium ones have subterminal conric tions and nine remaining pairs of small chromosomes have median or submedian constrictions. One pair of medium chromosomes is a Sat chromosome with a secondary constriction. The chromosome complement is larger than the other typical tropical species. Livistona subglobosa 2n=36(2b)=4A+32C (Fig. 2). This species has thirty-six chromosomes of which two pairs of long chromosomes have subterminal constrictions. This karyotype has smaller chromosome complement and seems to be the typical one of tropical Palmae. Trachycarpus excelsus 2n=36(2b)=6A+12B+18C (Fig. 3). This species has thirty-six somatic chromosomes of which two pairs of long chromosomes have submedian constrictions and one pair of long chromosomes have subterminal ones and six pairs of medium chromosomes and also nine pairs of short chromosomes have subterminal or submedian constrictions. According to SINOTO (1928) haploid number of Trachycarpus excelsus 176 D. SATO Cytologia 14 1946 Karyotype Alteration and Fhylogeny, VI. Karyotype analysis in Palmae 177 and variety Fortune is eighteen. Trinax argen.tea 2n=36(2b)=16A+20C (Fig. 4). This species has small chromosome complement consisting of thirty six somatic chromosomes and gradual difference between long and short chromosomes can not certainly be detected by their continuous sizes. Rhapis humilis 2n=36(2b)=8A+10B+18C (Fig. 5). R. fabelliformis 2n=36(Zb)=8A+10B+18C (Fig. 6). These species have comparatively larger chromosome complements as well as Trithrinax and Thrachycarpus. Sabal adansoni 2n=36(2b)=2A+34C (Fig. 7). S. Blackbuirtianum. 2n=36(21))=2A+34C (Fig. 8). These species have smaller chromosome complement of which one pair of long chromosomes has submedian constriction and most of remain ing short chromosomes indicate no distinct constrictions on account of extreme contraction. The presence of such dot or contracted chromosomes is one of the characteristic features of karyotypes in the tropical Palmae. Chameerops humilis 2n=36(2b)=6A+12B+18C. Seven genera, nine species in Corypheae have similarly thirty-six somatic chromosomes, but their karyotypes are different from each other, though they are of diploid nature having two nucleoli in the telophase. The report of NEMEC (1910) in Pritchardia filementosa (2n=24) is doubtful judging from these karyotypes in the other genera. The karyotypes of Livistona (b=18=2A+16C) and Sabal(b=18A+17C) are similar to those of Cordyline (b=19=2A+17C), Dracaena (b=19=2A+17C) and and Sansevieria (b=20=2A+18C) in Yucceae of Agavaceae and the other karyotypes in Corypheae are similar to those of Norina (b=18=6A+12C) and Dasylirion (b=19=4A+15C) in Nolineae of Agavaceae. 2 Borasseae Hyphaene wildbrandi 2n=36(2b)=2A+6B+2'C-r26C (Fig. 9). Figs. 1-32. Karyotypes in Corypheae (1-8), Borasseae (9), Calameae (10). Areceae (11-20), Cocoineae (21-31) and Phoeniceae (32). 1, Trithrinax brasiliensis 2n=86(2b). 2, Livistona subglobosa 2n=36(2b). 3, Traehcarpus excelsus 2n=36(2b). 4, Trinax argentea 2n=36 (2b). 5, Rhapis humilis 2n=36(2b). 6, H. flabellifornzis 2n=36(2b). 7, Sabal adansoni 2n=36(2b) (trop.). 8, S. Blackburnianum 2n=36(2b) (trop.). 9, Raphia Rut is 2n=32(2b) (trop.). 10, Hyphaene wildbrandi 2n=36(2b) (trop). 11, 12, Exorrhiza Savoryana 2n=32 (2b). 12, its somatic doubling of chromos_??_mes 2n=64(4b). 13, Ptychosperma Maearthurii 2n=32 (2b). 14, 15. Oreodoxa regia 2n=38(2b) (trop). 14, plant in the Ponape Island and 15, pot plant in Koishikawa Botanic Garden. 16, Howea Belmoreana 2n=36(2b). 17, Hyophorbe Verschatielti 2n=32(2b). 18, Caryota wrens 2n=32(2b). 14, A,enga saceharifera 2n=32(2b) (trop.). 20,A. Engleri 2n=32(2b) (Trop.). 21-23, Martinezia caryotaelolia 2n=32 (2b) (trop.). 22, its somatic doubling of chromosomes 2n=64(4b). 23. its mixoploid cells especially shown their nucleoli respectively. 24, Elaeis gusneensis 2n=32(2b) (trop.). 25, Cocos odorata 2n=32(2b) (trop.). 26, C. australis 2n=32(2b) (trop.). 27, C. capitata 2n=32 (2b)(trop.). 28. Attalea Cehuni 2n=32(2b). 29, Jubaea spectabilis 2n=32(2b). 30, Eutia bonneti 2n=32(2b), 31, Ar,ca.strum romanzofinum 2n=32(2b). 32, Phoenix canariensts 2n=36 (2b). •~2200 except 23 (•~1100. 178 D. SATO Cytologia 14 This species has thirty-six somatic chromosomes of which one pair of long chromosomes and three pairs of medium ones have subterminal constrictions and one pair of short chromosomes has satellites. This karyotype has a smaller chromosome complement and similar to those of Corclyline, Dracaena and Sansevieria in Yucceae of Agavaceae. 3 Calameae Raph.ia Ruffle 2n=32(2b)=6A+8B+18C (Fig. 10). This species has thirty-two somatic chromosomes of which three pairs of long chromosomes have subterminal constrictions and four pairs of medium chromosomes and nine remaining pairs of short ones have sub erminal or submedian constrictions. This karyotype is clearly different from those in Corypheae and Borasseae (b=18) by having another basic number (b=16) and similar to the karyotypes in the following tribes, Areaceae and Cocoineae (b=16). This karyotype is similar to that of Phormium (b=16=4A+12C) especially in basic num'b,ar and also similar to the Dracaena type (b=19=2A+17C). 4 Areceae (1) Euareceae Exorrhiza Savoryana 2n=32(2b)=14A+18C (Figs. 11, 12).•V •V (somatic doubling) 2n=64(4b)=28A+36C (Fig. 12). This species has thirty two somatic chromosomes of which seven pairs of long chromosomes have subterminal or submedian constrictions and nine pairs of short chromosmmes have submedian or subterminal constric tions. This karyotype has a larger chromosome complement than those of tropical Palmae and somatic doubling of chromosomes is also found in the mixoploid tissue of the root-tips which showed two or four corre sponding nucleoli at telophase. Area Catechu 2n=32(2b)=6A+8B+18C. (2) Ptychospermeae Ptychosperma Macarthurii 2n=32(2b)=6A+8B+18C (Fig. 13). This species has thrity-two somatic chromosomes of which three pairs of long chromosomes have subterminal constrictions, four pairs of medium chromosomes and remaining nine pairs of short chromosomes have sub median or subterminal constrictions. (3) Oncospermeae Oreodoxa regia 2n=38=(2b)=2A+6B+30C (Figs.
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