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Chromosome Numbers in the Malvaceae. I

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Chromosome Numbers in the Malvaceae. I CHI%OMOSOME NUMBEgS IN THE MALVACEAE. I. BY A. SKOVSTED. (E,mpi~:e Cotton G~'owi,ng Corporation, Cotton Research Station, T~'inidad, B. W.I.) (With i[25 Figures in the Text.) CONTENTS. I'A (4 I'] I. Imbroducfion 2(]3 II. History 263 III. Technique a,nd ma,LeriM 264 IV. CybologicM observations 264 V. Disctission and generM summary 289 VI. Summary 292 I%eferences . 293 I. INTRODUCTION. THE present studies were undertaken as part of a programme for eluci- dating the cytology of the genus Gossypium. It was anticipated tha~ chromosome nmnbers and morphology might be useful in distinguishing the genus from closely related genera. Furthermore, it was thought that a cytological study of the entire family Malvaceae might contribute to a bettcr understanding of the evolution of this family and consequently to the origin of the genus Gossypium. The family Malvaceae contains at least 800 species. In the present investigation approximately 120 of these have been examined. As this represents a small percentage of the total number, and as further investi- g'ations are in progress, the present work is factual in nature, and the discussions and conclusions of a preliminary character. II. HIsTouY. Most of the relevant cytological work deals with the genus Gossypium only, the literature of which has recently been reviewed (Longley0a), Skovsted (19, 20)). A few species from ott~er genera have also been studied (Svensson-Stenar (23), Tischler (sh, 2a)). Only three comprehensive studies dealing with larger numbers of species have been published: two of these refer in particular to the flibiscus gro~ctp (Longley(*% Youngman(an); in the third, tile chromosome numbers of fourteen new species repre- 264 Chromosome Numbers in the Malvaceae. I. senting eight genera are recorded by Davie(a). These chromosome numbers are discussed in detail lager. III. TECHNIQUE AND ~ATEr~IAL. Both roo~4ips and pollen mother ceils were exami~led in the deter- minatiou of chromosome numbers. The majority of the root-tips were fixed in Navashin's solution, and the buds in @arnoy-Navashin. In the improved technique developed during the invesCigaCion later fixations of root-tips were made in LewiCsky's fixative (2J), and buds were treated according to the technique developed for cotton09). All slides were stained in Gentian violet. The taxonomic arrangement~ of genera is that of Engler and Prantl; nomenclature follows closely Index Kewe'nsis. Most of the species examined were obtained from seeds supplied through the Botanical Garden of the University of Copenhagen, or from seeds bought from Haage and Schmidt, Erfurt. Some wild species from different parts of the world eolleeted by Dr S. C. I-Iarland and by Mr J. B. Hutchinson in India were kindly placed at the writer's disposal. To Dr O. Hagerup the writer is indebted for fixed material of species grown a~ Timbuktu in the Sahara. Species native to Trinidad were collected from nature by the writer. IV. @YTOLOGICAL OBSEiZVATIONS. t YV~e tribe Malopeae. (1) The genus Malope. 2n Aut~hor M. trifida Car. (Fig. 1) ,t4 Davie (6), S. M. trifida (Fig. 2) 50 S. 2[. hispida Boiss. (Fig. 3) 44 S. In the present study the chromosome nmnber 2'n = 4,J: in M. t,r.ifida was confu'med from material obtained from Haage and Schmidt; another type of tile species obtained from Copenhagen had 2n = 50; morphologically the two types were very similar; chromosomes of sizes similar to those of M. tr'ifida were found in M. hislJida Boiss. (2) ~'he genus Kitaibelia. n 2n Aughor If. Li~dcmuthi Hort. (Fig. 4) -- 44 S. K. vitiJblia -- 42 D~vie (6) K. vitifolia WiHd, (2 types) (Figs. 5-6) 22 44 S. Numbm's de~ermined by the writer are m~rked wi~h "S". A. SKOVS~I'EO 265 In both K. Lindemuthi Herr. and K. vitifolia Willd. the chromosome number is here found to be 2n= 44, the chromosomes being of similar sizes; another type of K. vitiJblia showed n=22 in the first meiotic division. The t~'ibe Malveae. (1) The genus Abutilon. 'a 2n, Author A. striatum Thompso~d 8 -- Kuhn (26) A. striatum T/~om,pson.i 8 16 J)~vie (6) A. striatum :Dicks (Figs. 7-8) 8 16 S. A. 'wmbellatum Sweet (Figs. 9-10) -- 16 S. A. crispum Sweet (Figs. 1 [-12) 7 14 S. A. gla,ttcnm G, ~on (Fig. 13) 14 -- S. A. Avicennae -- 42 :Bresl,~vetz (3) A. Avicennae Gaertn. (2 types) (Figs. 1,1-] 5) 21 42 S. A. giganteum ]?resl (Fig. 16) -- 42 S. A. indicu,n+ Sweet (Figs. 17-18) 2i 42 S. A. molle Sweet (2 types) (Pigs. 19-20) 2I 4:2 S. A..Regncllii Niq. (3 types) (Figs. 21-22) 21 42 S. Abutilon sp. (Fig. 23) -- 42 S. A burliest, sp. (fl'om Sudan) (Fig. 24) -- 42 S. Following the cytological observations reported here the genus ha, s been divided into two groups, one with 8 and the other with 7 as the basic chromosome number. 1 The first group contains: A. st,riatum with n=8 and 2n=16 and A. umbel,k~,tum Sweet. The latter species pos,~essed %=16 (Fig. 9) in five of the six roots examined, while one root was tetraploid with 2n = 32 (Fig. 10). The second group (basic number 7) comprises nine species, only one of which has been studied by previous workers : cytologically, A. crisl)um Sweet is the more primitive type with n = 7 and 2n = 14, while A. glaucum G. Don has n -- 1~; the remaining seven species all have ~ = 21 or 2n = ~2. Chromosome sizes differ somewhat from one species to another, but, since considerable variation in size is also found among the chromosomes in a single cell, no particular inference can be drawn. Most species possess one or two pairs of chromosomes which are easily distinguishable by their satellites. Only in A. crisl)Um was it possible to distinguish the wl{ole set of chromosomes on minute morphological differences. 1 It is interesting to note that the cytological division appears to be associated with morphological characters : the species witch n = 8 having five-lobed capsules and the species of the seven series poly-lobed capsules. I-[erbarium studies indicate that this morphological division is m~tural throughout the genus. Most species from all parts of the world have poly-lobed e~psules, while only a few--all confined to the New World--have five-lobed capsules. 266 Chromosome Numbers in the Malvaceae. I. (2) The genus Wissadula. 2~ Author W. contracta ]%. E. Fries (Fig, 25) 14 S. The t4: somatic chromosomes k~re smMler than those of Abutilon crispum Sweet (Fig. 11). (3) The genus Sphaeralcea. n 2)~ Author S. miniata Spaeh. (Figs. 26-27) 5 10 S. S, ,u~nbellata G. Don (Fig. 28) -- 34 S. The chromosomes are much larger than those of the above-mentioned genera. (~t) The genus )JIodiola. 2n Author M. "m'ullli[ida Moeneh (2 ~ypes) (Fig. 29) 18 S. The 18 somatic chromosomes are similar iu size to those of S'phae,ralcea. (5) The genus Lavatera. n 2v, Author L. trimestris 7 14 Davie (6) L. lrimesO'is L. (3 types) -- 14 S. L. arborca L. -- 36 D~vie (6) L. arborea (2 gypes) (Fig. 30) -- 42 S. L. cclche,m,iria~a 22 -- Davie (6) L. cachemiriana Cambess (Figs. 31-32) -- ,12 S. L. ,mauritanica Dueh. (Fig. 33) -- c, 84 S. L. cretica L. (Fig. 34) -- c, 112 S. L. Olbia L. 20 40 D~vie (6) L. th,u,ri'~lgiuca 20 -- Svensson-Sten~r (23) L. thuringiaca (2 types) (Fig. 35) 22 4~ S. L. ~)lcbcia Sims (Fig. 36) -- 43 S. The chromosome numbers of five species have already been recorded. Four of these were examined dnrhlg the presegt investigations, but only the chromosome number of L. t~'i,mest~'is was confirmed. Two types of L. a,rborea L. and L. eachem@iana Cambess had 2n=~2; in the latter species, tetraploid tissue with 2n=8L1 was observed. L. mauritanica Duch. had % = e. 84 and L. eretica L. approximately 112 somatic chromo- somes. These species are apparently poiyploids of seven compared with L. thuringiaee L. which has n=22 and 2n=44; L. plebeia Sims affords an intermediate number with 2n = 43; the meiotic divisions of this species have not yet been studied. It may be anticipated that cytological studies of other species in this genus will provide useful results as different chromosome numbers are apparently represented. n. ~KOVSTE]) 267 (6) l'he genus Althaea. n 2~ Author A. can~;mbina L. (Fig. 37) -- 28 S. A, a,ptcrocarl)r,~ Fcnzt (Fig. 38) -- 42 S, A . .fic~blia -- 44 Davie (6) A. fieifolia (Jay. (Fig. 39) -- '.42 S, A. Heldreichii Boiss. (Fig. 40) -- 42 S, qOicinalis -- c. 40~t4 D,~vie (6) q[#ci*~ali,s L. (2 typos) (Fig. 41) -- 42 S, pcdlida W~dcl. et [{it. (Fig. 42) -- 42 S. TOSEC(, 21 -- Kcssetcr (12) ~'08~Cb ] 3 -- Burkcbt (4) rosen C~v. (4 types) (Fig. 43) -- 42 S, h ybrida Heft. (Fig, 44.) -- 42 S. ,si'nc~sis C~v, (]fig. 45) -- 70 S, ar,meniaca Tcnoro -- c. 84 S, Hoke;~ackcri ]3oiss. ot }Iuot. -- c. 84 S. ]e,;'agujevace~mis :P~ncic (Fig. 46) -- c. 84 S. In the presen~ study twelve species proved to be simple polyploids Of seven. Slight variations were observed in the chromosome sizes of the different species. 0nty A. eannabina has chromosomes significantly larger than the other species. (7) The genus Malva. n 2n Author M. oxyloba ]3oiss.
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