Leaf Phenology and Seasonal Carbon Gain in the Invasive Plant, Bunias orientalis L.

Abstract: In two potentially competing herbaceous plants, the invasive Bunias orientalis L. (Brassicaceae) and the native Picris hieracioides L. (Asteraceae), seasonal changes in leaf CO₂ gas exchange and plant growth were studied over an entire growing season from February 1998 to December 1998 in two experimental fields. The study was motivated by the hypothesis that pre-adaptive phenological displacement of alien species relative to the native flora may be an important reason for the observed expansion of B. orientalis in central Europe. We quantified the importance of phenological differences for annual carbon gain in both species by estimating total leaf carbon gain from the results of leaf CO₂ exchange and changes in plant leaf area. Bunias orientalis achieved almost half of its annual carbon gain in the time between early September and December, when competition for light by other species, like P. hieracioides , is low. Our quantitative approach corroborates the notion that the phenological shift of a relatively poor competitor, such as B. orientalis , could be of great importance for the success as an invasive species.     

Complex pattern of genome size variation in a polymorphic member of the Asteraceae

Aim  Although divergences in nuclear DNA content among different species within a genus are widely acknowledged, intraspecific variation is still a somewhat controversial issue. The aim of this study was to assess genome size variation in the polymorphic species Picris hieracioides L. (Asteraceae) and to search for potential interpretations of the size heterogeneity.
Location  Europe.
Methods  The genome sizes of 179 plants of P. hieracioides collected from 54 populations distributed across 10 European countries were determined by propidium iodide flow cytometry. Differences in nuclear DNA content were confirmed in simultaneous analyses.
Results  2C-values (population means) at the diploid level varied from 2.26 to 3.11 pg, spanning a 1.37-fold range. The variation persisted even after splitting the whole data set into two recently distinguished morphotypes (i.e. the 'Lower altitude' type and the 'Higher altitude' type) that possess significantly different nuclear DNA contents. Cluster analysis revealed the presence of three major groups according to genome size, which exhibited a particular geographical pattern. Generally, the genome size of both morphotypes increased significantly from south-west to north-east. A new cytotype, DNA triploid, was found for the first time.
Main conclusions  High intraspecific variation in the amount of nuclear DNA in P. hieracioides correlates with the extensive morphological variation found within the taxon. Despite the complex pattern that was presented, genome size variants were non-randomly distributed and reflected palaeovegetation history. We suggest that the complex evolutionary history of P. hieracioides (e.g. the existence of several cryptic lineages with different levels of cross-interactions) is the most plausible explanation for the observed heterogeneity in genome size.     

Karyotype of Four Species in Buddleja ( Loganiaceae) *

The chromosome numbers and karyotypes of the 4 species in genus Buddleja were reported. The karyotype formulas are 2n= 2x= 38= 22m+ 16sm ( B1 yunnanensis) , 2n= 2x= 38= 26m+ 10sm+ 2st ( B1 crispa) , 2n= 2x= 38= 20m+ 16sm + 2st ( B1 off icinalis ) , and 2n= 2x= 38= 20m+ 16sm+ 2st ( B1japonica) . Karyotype of B1japonica belongs to Stebbins. s 2B type and other 3 species belong to Stebbins. s 2A type. Based on the cytological data ( karyotypes and the recorded chromosome numbers) and the species morphologies, the evolution trend of the two series in Sect1Neemda was briefly discussed.     

Karyotypes of eight species of Euphorbia L. (Euphorbiaceae) from China

In this paper, eight species of the genus Euphorbia L. were cytologically studied. The three species of the subgenus Chamaesyce Raf., E. hirta, E. humifusa and E. hypericifolia, had chromosome numbers of 2n = 18, 22 and 32, with their basic chromosome numbers being x = 9, 11 and 8 respectively. The two species of the subgenus Poinsettia (Grah.) House. E. dentata, with 2n=28, a tetraploid, and E. cyathophora, with 2n= 56, a octoploid, had both the basic chromosome number of x= 7. The three species of the subgenus Esula Pers, E. lathyris, E. helioscopia and E. hylonoma, had chromosome number of 2n= 20, 42 and 20, with their basic numbers being x= 10, 7 and 10 respectively. The basic chromosome number of x = 8 is new for E. hypericifolia, in which x = 7 was previously reported. This indicates that this species had both ploidy(2n = 4x = 28, 8x = 56) and dysploidy(x = 7, 8) variations. In E. dentata, there occurred also ploidy variation (2n = 2x, 4x and 8x). A tetraploid cytotype of E. esula was found in China, its diploid cytotype and hexaploid cytotype being previously reported in North America, the Iberian Peninsula and some other European areas. Based on our results and those previously reported, we support the viewpoint that x＝10 may be the original basic chromosome number of Euphorbiaand discuss the role of polyploidy and dysploidy in the speciation and evolution of this genus     

八种国产大戟属植物的核型报道

8种大戟属Euphorbia L．植物的核型分析结果表明,大戟属不同亚属的染色体基数与其形态变 异的复杂性有一定关系。地锦草亚属subgen．Chamaesyce 3个种染色体基数分别为x=8,9,11;一品红 亚属subgen．Poinsettia两个种染色体基数均为x=7,分别为四倍体和八倍体;乳浆大戟亚属subgen． Esula 3个种,染色体基数分别为x=7,10,10。根据以前学者发表的资料分析,一品红亚属和大戟亚属 Subgen. Euphorbia的染色体基数是很稳定的,分别为x=7和x=10;通奶草E．hypericifolia为x=8 的四倍体,它不仅有染色体整倍性的变异,还有异基数性的变化。结合以前学者的研究,笔者支持x= 10为大戟属的最原始基数的观点。齿裂大戟E．dentata和通奶草具不同的染色体倍性,猫眼草E． esula的细胞染色体数目观察证实了我国存在四倍体的居群,与欧洲和北美的植物构成一个典型的多倍体复合体。     

山西关帝山部分植物的染色体研究

报道了山西关帝山庞泉沟自然保护区７科、１１属、１３种植物的染色体研究结果。其中８种作了核型分析,４种的核型和２种的染色体数目为首次报道。     

醉鱼草属四个种的核型分析

报道了醉鱼草属（Btuldleja）4个种的染色体核型。云南醉鱼草（B.yunnanensis）的核型公式为2n=2x=38=22m＋16sm,皱叶醉鱼草（B．crispa）为2n=2x=38=26m＋10sm＋2st,密蒙花（B．officinalis）为2n=2x=38=20m＋16sm＋2st,口本醉鱼草（B．japonica）为2n=2x=38=20m＋16sm＋2st。日本醉鱼草的核型为2B型,其它3个种的核型为2A型。根据核型分析结果,结合形态学特征和已有的细胞学资料,初步讨论了醉鱼草组（Sect．Neemda）两个系（Series）可能的演化关系。     

二倍体铁破锣的核型及四倍体细胞型的首次发现

本文重新检查了铁破锣的核型。来自湖南新宁的40个个体中,1个个体为四倍体,其核型公式为2n＝4x=32=16m+8sm+4st+4t;其余个体为二倍体,其核型公式为2n＝2x＝16=8m+4sm+2st+2t。来自云南大理的10个个体全为四倍体,其核型公式为2n=4x=32＝16m+8sm+4st+4t。据此认为商效民(1985)报道的该种的核型分析结果有误。他至少将第4对染色体的着丝点位置辨认错了。该对染色体不具有中部着丝点而实际上具有近端部着丝点。本文还比较了铁破锣和角叶铁破锣的核型差异,并详细讨论了铁破锣属的系统位置。     

中国水韭属植物的染色体数目及其分类学意义

利用细胞学方法观察统计了中国4种水韭属Isoestes植物的染色体数目。结果发现4个种的染色体基数均为x=11,其中高寒水韭I．hypsophila Hand.-Mazz．为2n=22,为首次报道;台湾水韭 I．taiwanensis DeVol为2n=22;中华水韭I．sinensis Palmer 2n=44为四倍体;而产于云南贵州一带并一直被当作是宽叶水韭I．japonica A．Br．的水韭属植物2n=22,与文献报道的宽叶水韭(2n=66,67,77,87,88,89)完全不同,应重新予以认识和评价。依据本文和其他相关的研究结果,对东亚水韭属植物进行了分类处理。     

Introgression mapping of genes for winter hardiness and frost tolerance transferred from Festuca arundinacea into Lolium multiflorum

Genes for winter hardiness and frost tolerance were introgressed from Festuca arundinacea into winter-sensitive Lolium multiflorum. Two partly fertile, pentaploid (2n = 5x = 35) F(1) hybrids F. arundinacea (2n = 6x = 42) x L. multiflorum (2n = 4x = 28) were generated and backcrossed twice onto L. multiflorum (2x). The backcross 1 (BC(1)) and backcross 2 (BC(2)) plants were preselected for high vigor and good fertility, and subsequently, a total of 83 BC(2) plants were selected for winter hardiness after 2 Polish winters and by simulated freezing tests. Genomic in situ hybridization (GISH) was performed on 6 winter-hardy plants selected after the first winter and shown to be significantly (P < 0.05) more frost tolerant than the L. multiflorum control. Among the analyzed BC(2) winter survivors, only diploid (2n = 2x = 14) plants were found. Five plants carried 13 intact L. multiflorum chromosomes and 1 L. multiflorum chromosome with a single introgressed F. arundinacea terminal chromosome segment. The sixth BC(2) winter survivor appeared to be Lolium without any Festuca introgression capable of detection by GISH. A combined GISH and fluorescence in situ hybridization analysis with rDNA probes of the most winter-hardy (after 2 winters) and frost-tolerant BC(2) plant revealed the location of an F. arundinacea introgression on the nonsatellite arm of L. multiflorum chromosome 2, the same chromosome location reported previously as a site for frost tolerance genes in the diploid and winter-hardy species Festuca pratensis.     

中国西南六种广义拂子茅属(禾本科)植物的染色体数目

广义拂子茅属（Calamagrostis）是一个世界温带广布的大属,有些作者又分为拂子茅属和野青茅属,但近期的研究表明处理为一个属较为合适。中国共有37种广义拂子茅属植物,但至今没有任何染色体的研究。本文报道了其中产于中国西南6种野青茅的染色体数目,其中Deyeuxia petelotii 4个居群,D．diffusa,D．moupinemis,D．nivicola和D．flvens各一个居群都是四倍体（2n=4x=28）,D．neglecta为六倍体（2n=6x=42）。根据广义拂子茅属植物染色体倍性特征,该属植物中至今未发现二倍体,四倍体是该属中倍性最低和最普遍的,广义拂子茅属的演化很可能是在四倍体的水平上进行的。由于以上几个四倍体种均是狭域分布的类群,所以可能是由四倍体的祖先隔离分化形成的。     

Correction of karyotype of diploid Beesia calthifolia and discovery of a tetraploid cytotype

In this paper, the chromosomes of Beesia calthifolia were re-examined. In the 40 plant individuals of the population from Xinning County, Hunan Province, central China, one was found to be tetraploid with the karyotype formula of 2n＝4x=32=16m+8sm＋4st + 4t, and the remaining were all found to be diploids with the karyotype formula of 2n＝2x ＝16=8m+4sm+2st+2t. All the 10 individuals of the population from Cangshan Mountain, Dali City, Yunnan Province, southwestern China, were unexpectedly found to be tetraploids with the karyotype formula of 2n＝4x＝32=16m+8sm+4st+4t. Tetraploid cytotype was reported in this species for the first time. Based on the results and those previously reported, it is considered that there may exist some errors in the results of the karyotype analysis of this species previously reported by Shang(1985). He might have at least mistakenly recognized the centromere position of the fourth pair of chromosomes. This pair of chromosomes should have subterminal rather than median centromeres. Furthermore, the karyotypic differences of B. calthifolia and B. deltophylla were analyzed and the systematicposition of the genus Beesia was discussed in detail.     

The Balkan endemic Picris hispidissima (Compositae): morphology, nuclear DNA content and relationship to the polymorphic P. hieracioides

The only Balkan endemic of the genus Picris, Picris hispidissima, was studied in detail using morphological and karyological methods. The species was shown to be morphologically distinct from the closest taxon, Picris hieracioides, by the pectinate–ciliate indumentum of involucral bracts, dilatation of the peduncle, length of the outer and inner bracts, and indumentum colour. No morphological variation that would require taxonomic classification was found within this species. Despite the diploid chromosome number (2n = 2x = 10) being confirmed for P. hispidissima, variation of up to 9.5% in genome size was found. The likely explanation for this variation is hybridization and introgression with closely related P. hieracioides. The most convincing evidence for this hypothesis is the detection of plants with two clearly different DNA contents arising from a single capitulum found in the location where P. hispidissima and P. hieracioides co-occur.     

A comparative study on karyotypes of 28 taxa in Rubus sect. Idaeobatus and sect. Malachobatus (Rosaceae) from China

Rubus is a taxonomically difficult group and cytological data are expected hopefully to gain insight into the relationships of the genus. In this study the chromosome numbers and karyotypes of 18 taxa from sect. Idaeobatus and 10 taxa from sect. Malachobatus were investigated. Among them, the chromosome numbers of 10 taxa and karyotypes of 26 taxa were reported for the first time and mixoploidy was observed new in the genus. The chromosomes are small in size with a length of less than 3 µm and metacentric (about 90%) or submetacentric. All taxa have karyotypes of “1A” except R. cockburnianus, R. innominatus and R. ellipticus var. obcordatus, which have karyotypes of “2A”. No aneuploids were found in all the 28 taxa studied. Plants of sect. Idaeobatus have diploids with 2n=2x=14, except R. idaeopsis (2n=3x=21) and R. parvifolius (A mixoploid of 2n=2x=14 and 2n=4x=28). However, plants of sect. Malachobatus have tetraploids with 2n=4x=28, except for R. buergeri with 2n=8x=56. In addition, conspicuous karyotype differences existed within the 18 taxa belonging to 11 of 7 subsections in sect. Idaeobatus, and the differences between some species within subsections are greater than that between subsections, while uniform karyotypes within subsections and variable karyotypes between subsections were observed in six of 13 subsections in sect. Malachobatus represented by 10 taxa. Systematic values of the cytological data were discussed for some cases when applicable to the two sections.     

珍珠菜属3种植物的核型分析

采用常规压片法,对珍珠菜属(Lysimachia L.)3种植物的核型进行了研究。结果表明,长蕊珍珠菜的核型为2n=2x=24=12m 10sm 2st,显苞过路黄的核型为2n=2x=24=4m 6sm 6st 8t,均属首次报道。过路黄的核型为2n=2x=24=2m 2sm 4st 16t,与前人报道的有所不同。还对已报道的珍珠菜属的核型类型与不对称系数进行了比较。     

秦巴山区重楼属的核型研究

本文报道了陕西秦巴山区几种重楼属植物的核型研究结果。在秦岭(太白山)地区,北楼(Paris verticillata)具有一种核型,其核型公式为 2n=4 x=20=10m+2sm+4st+4t;狭叶重楼(P.polyphylla var.stenophylla)在一个居群内不同的个体间具有两种不同的核型,核型公式分别为2n=2x=10=5m+1sm+4t和2n=2x=10+2B=6m+4t+2B。在秦岭和巴山两地,宽叶重楼(P.polyphylla var.latifolia)和球药隔重楼(P.fargesii)分别具有两种不同的核型,前者的核型公式为2n=2x=10+3B=6m+3t+3B和2n=2x=10=5m+1sm+4t,后者的核型公式为2n=2x=10+2B=4m+2sm+4t+2B和2n=2x=10+2B=6m+4t+2B。这些材料均属2A核型,在核型结构及B染色体的有无和数目上与前人报道的结果不完全相同。     

Karyology and biogeography of Dugesia japonica and Dugesia ryukyuensis in Kyûshû, southern Japan

In southern Japan, two dugesiid species of freshwater planarians are known: Dugesia japonica Ichikawa et Kawakatsu, 1964, and Dugesia ryukyuensis Kawakatsu, 1976. D. japonica is a common and polymorphic species widely distributed in the Far East (karyotypes: n=8, 2x=16, 3x=24). D. ryukyuensis is a species recorded only from the Southwest Islands of Japan (Nansei Shotô) (karyotypes: n=7, 2x=14, 3x=21). Recently, small populations of D. ryukyuensis were found in the lowland areas in Kyûshû on the East China Sea (the Nishisonogi Peninsula, the Shimabara Peninsula, the Gotô Islands, the Satsuma Peninsula, and the Ôsumi Peninsula). The current geographical distribution of D. japonica and D. ryukyuensis in southern Japan can be explained from geological and faunal viewpoints, as follows: (1) two separate inversions by the ancestor of D. japonica, one in the Miocene and one after early Quaternary; (2) only one expansion of its domain by the ancestor of D. ryukyuensis in the Miocene.     

Interspecific Hybridization of Pennisetum glaucum (L) R. Br. with Wild Relatives

To widen the germplasm base for the introgression of economically important traits such as resistance to biotic and abiotic stresses from related species, crosses of cultivated pearl millet were made with pollen from four related species differing in the basic chromosome number (x=5,7,8 and 9). Embryo rescue technique was used to obtain viable progeny. Pollinations of pearl millet with Pennisetum ramosum (2n=2x=10) did not give any viable progeny. Pearl millet interspecific hybrids with P. schwelnfurthii (2n-2x-14), P. mezianum (2n=4x-32) and P. orientale (2n=2x=18) were obtained. The hybrid between P. glaucum and P. mezianum (2n=23) is the first successful report. Interspecific hybrid plants resembled their corresponding pollen parents. Southern blots of Psfl digested DNAs from interspecific hybrids and the parental species were hybridized to a full length rDNA to further confirm their hybridity. This further revealed differential amplification of two rDNA repeats among the F₁ hybrids from the same cross (P. glaucum X P. orientale).     

Variability of the 5S and 45S rDNA sites in Passiflora L. species with distinct base chromosome numbers

Cytologically, the species of Passiflora with known chromosome number can be divided into four groups: (1) 2n = 12, 24, 36; (2) 2n = 24; (3) 2n = 18, 72; and (4) 2n = 20. The base chromosome number proposed for the genus is x = 6, with x = 9, x = 10 and x = 12 being considered secondary base numbers. In the present study, variability of 5S and 45S rDNA sites was investigated in 20 species of these four groups to check the reliability of this hypothesis. In the group with x = 6, five diploid species (2n = 12) exhibit two 5S rDNA sites and two (P. capsularis, P. morifolia and P. rubra) or four (P. misera 2x and P. tricuspis) 45S rDNA sites. The hexaploid cytotype of P. misera had 12 45S rDNA sites and six 5S rDNA. A tetraploid species, P. suberosa, had ten 45S rDNA sites and four 5S rDNA sites, both in the same chromosomes as the 45S rDNA sites. In the group with x = 9, P. actinia, P. amethystina, P. edmundoi, P. elegans, P. galbana, P. glandulosa and P. mucronata displayed six 45S rDNA sites, whereas P. alata, P. cincinnata, P. edulis f. flavicarpa, P. edulis var. roxo and P. laurifolia had four sites. In this group, all species were diploid (2n = 18) and had only two 5S rDNA sites. Passiflora foetida, the only species with 2n = 20, had six 45S rDNA sites and four 5S rDNA sites. The species with x = 12 (2n = 24), P. haematostigma and P. pentagona, showed four 45S rDNA sites and two 5S rDNA. In general, the number and location of 5S and 45S rDNA sites were consistent with the hypothesis of x = 6 as the probable ancestral genome for the genus, while the groups of species with x = 9, x = 10 and x = 12 were considered to be of tetraploid origin with descending dysploidy and gene silencing of some redundant gene sites, mainly those of 5S rDNA.     

New Chromosome Counts of Some Dicots in the Sino-Japanese Region and Their Systematics and Evolutionary Significance

New somatic chromosome numbers for nine species eight families and eight gen era in the Sino-Japanese Region are reported here as shown in Table 1. Data of six genera are previously unknown cytologically. The bearings of these new data on the systematics and evolution of the related species, genera or families are discussed as follows: (1) Platycarya strobilacea Sieb. et Zucc. (Juglandaceae). The chromosome number of this species is 2n=24, with a basic number of x=12, which deviates from 2n=32 occurred in Juglans, Carya, Pterocarya and Engelhardtia with the basic number x= 16. The Juglandaceae appears to be fundamentally paleotetraploid, with an original basic number of x = 6 in Platycarya and x-8 in the other four genera, although secondary polyploidy occurs in Carya. Based on the remarkable morphological differences between Platycarya and the rest seven genera of the family, Manning (1978) established two subfamilies: Platycaryoideae for Platycarya and Juglandoideae for the other genera. Iljinskaya (1990), however, recently established a new subfamily: Engelhardioideae for Engelhardtia. Lu (1982) points out that because of a great number of primitive characters occurring in Platycarya, the genus could not be derived from any other extant juglandaceous taxa but probably originated with the other groups from a common extinct ancestor. The present cytological data gives support to Manning′s treatment. We are also in favor of Lu′s supposition and suggest that basic aneuploid changes, both ascending and descending, from a common ancestor with the original basic number x=7, took place during the course of early evolution of the Juglandaceae and led to the origin of taxa with x=6 and 8. Subsequent polyploidy based on these diploids occurred and brought forth polyploids of relic nature today, whereas their diploid progenitors apparently have become extinct. (2) Nanocnide pilosa Migo (Urticaceae). The chromosome number of this Chinese endemic is 2n-24, with a basic number of x=12. An aneuploid series occurs in the Urticaceae, with x--13, 12, I1, 10, 9, 8, 7, etc. According to Ehrendorfer (1976), x = 14, itself being of tetraploid origin, is the original basic number of the whole Urticales, and descending aneuploid changes took place in the early stage of evolution of the Urticaceae and Cannabinaceae. In addition to Nanocnide, x= 12 also occurs in Australina, Hesperonide and Lecanthus, and partly in Chamabainia, Elatostema, Girardinia, Pouzolzia and Urtica. (3--4) Sedum sarmentosum Bunge and S. angustifolium Z. B. Hu et X. L. Huang (Crassulaceae). The former is a member of the Sino-Japanese Region, while the latter is only confined to eastern China. The chromosome number of Sedum is remarkably complex with n=4-12, 14-16…74, etc. S. angustifolium with 2n=72 of the present report is evidently a polyploid with a basic number of x =18 (9^?) Previous and present counts of S. sarmentosum show infraspecific aneupolyploidy: n = c. 36 (Uhl at al. 1972) and 2n=58 (the present report). These two species are sympatric in eastern China and are morphologically very similar, yet distinguishable from each other (Hsu et al. 1983) S. sarmentosum escaped from cultivation in the United States gardens exhibited high irregularity in meiosis (Uhl et al. 1972). Uhl (pets. comm. ) suspected strongly that it is a highly sterile hybrid. R. T. Clausen (pets. comm.) found that plants of S. sarmentosum naturalized in the American Gardens propagated by means of their long stolons and broken stem tips, and could not yield viable seeds. Hsu et al. (1983) found that some of the plants of S. sarmentosum and S. angustifolium did yield a few seeds, but other did not. These species are, therefore, by the large vegetatively apomictic. (5) Glochidion puberum (L. ) Hutch. (Euphorbiaceae). The genus Glochidion includes about 300 species, but only eigth species from the Himalayas have been studied cytologically, with n= 36 and 2n= 52, having a basic number of x= 13. The present count for the Chinese endemic G. puberum establishes the tetraploid chromosome number 2n= 64, and adds a new basic number x= 16 to the genus. (6) Orixa japonica Thunb. (Rutaceae). Orixa is a disjunct Sino-Japanese monotypic genus. Out of the 158 genera of the Rutaceae, chromosome numbers of 65 genera have hitherto been investigated, of which 42 genera are with x=9 (66.61%), some with x=7, 8 and 10, and rarely with x=13, 15, 17 and 19. The present count of 2n=34 for O. japonica may have resulted from a dibasic tetraploidy of n=8+9. (7) Rhamnella franguloides (Maxim.) Weberb. (Rhamnaceae). The chromosome number of this member of the Sino-Japanese Region is 2n= 24. with a basic number of x= 12. The basic number x= 12 also occurs in Hovenia, Paliurus, Sageretia, Ceanothus and Berchemia. Hong (1990) suggested that x= 12 in Rhamnaceae may be derived from descending aneuploidy of a paleotetraploid ancestor. (8) Sinojackia xylocarpa Hu (Styracaceae). The chromosome number of this rare Chinese endemic is 2n= 24, with a basic number of x =12, which is identical with that in Halesia and Pterostyrax, but deviates from that in Styrax (x=8). The basic number x=8 in the Styracaceae may be derived from the original basic number x=7 by ascending aneuploidy in the early stage of evolution of the family, and x=12 may be derived from polyploidy. (9) Thyrocarpus glochidiatus Maxim. (Boraginaceae). The chromosome number of this Chinese endemic species is 2n=24, with a basic number of x=12. An extensive aneuploid sequence of x = 4-12 occurs in the Boraginaceae, of which x = 8, 7 and 6 are the most common. The basic number x=12 also occurs in Cynoglossum and Mertensia. It is evident that aneuploid changes, both descending and ascending, from an ancestor with x = 7, have taken place in the primary phase of evolutionary diversification of the Boraginaceae, and subsequent polyploidy has given rise to x=15, 17 and 19 in a few genera (e. g. Amsinskia and Heliotropium). The origin of x=12 is not certain. Either it be a result of ascending aneuploidy, or a product of polyploidy on the basis of x = 6. The present authors are in favorof the latter.