Table of Contents

18 May 1993, Volume 31 Issue 3

  

Research Articles
• Notulae de Ranunculaceis Sinensibus (XV)

  Wang Wen-tsai

  J Syst Evol. 1993, 31(3): 201-226.
  

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  (1) The confusion of the circumscription of Aconitum delavayi Franch., an alpine erect species endemic to the Mt. Maershan, province Yunnan, which was brought about by Finet & Gagnep. and Handel-Mazzetti with an inclusion of a lowland twinning species, Aconitum episcopale Lévl., is clarified. Three Chinese specimens cited by Handel-Mazzetti under Aconitum baicalense Turcz. ex Rapaics in fact represent three taxa of the genus Acontium other than A. baicalense. Examination of the type specimens shows that Clematis grata Wall. var. grandidentata Rehd. et Wils., with pinnate leaves and few-flowered axillary inflorescences, was erroneously referred to C. grate Wall. var. argentilucida (Lévl. et Van. ) Rehd., with ternate leaves and many-flowered axillary inflorescences, by Rehder. As a matter of fact, the former represents an independent species related to C. grata Wall. and should be raised to specific rank, and the latter should be a variety of C. apiifolia DC., being identical with C. apiifolia DC. var. obtusidentata Rehd. et Wils. (2) Aconitum delavayi Franch. var. leiocarpum Finet et Gagnep. (=A. contorum Finet et Gagnep. ), A. vilmorinianum Kom. var. altifidum W. T. Wang ( =A. episcopale Lévl.), Delphinium leptopogon Hand-Mazz. (=D. siwanense Franch. ), D. calleryi Franch. =D. anthriscifolium Hance var. calleryi (Franch.) Finet et Gagnep. ( =D. anthriscifolium Hance var. anthriscifolium), Thalictrum thibeticum Franch. (= T. delavayi Franch. ), and C. apiifolia DC. var. obtusidentata Rhed. et Wils. (=C. apiifolia DC. var. argentilucida (Lévl. et Van. ) W. T. Wang) are all reduced into synonymies. Owing to that Thalictrum delavayi Franch. var. decorum Franch. is a famous ornamental plant and according to Art. 14 of the International Code of Botanical Nomenclature, Thalictrum delavayi Franch. (1886) is suggested to be conserved against the earlier name Thalictrum thibeticum Franch. ( 1885). ( 3) Aconitum episcopale Lévl., Thalictrum lecoyeri Franch. T. macrostigma Finet et Gagnep. (= T. sino-rnacrostigma W. T. Wang), Clematis pterantha Dunn, C. tangutica (Maxin.) Korsh. var. obtusiuscula Rehd. et Wils., and Adonis davidii Franch., which were put into synonymies by various authors, are restored. (4) The infraspecific taxa of Thalictrum delavayi Franch. are reviewed and the revised diagnosis for each taxon is given. The affinities between the species of the Clematis acuminata group are discussed. A key to the species of ser. Acutangulae is given. (5) one series, three species, three varieties, and two forms are described as new. One new combination, and four new ranks are made, and one new name is given. (6) The foreign distribution of Aconitum hemsleyanum Pritz. var. circinatum W. T. Wang, Asteropyrum pehatum (Oliv.) Drumm. et Hutch., and Thalictrum alpinum L. var. elatum Ulbr., which were formerly known endemic to China, isreported.
• Micromorphology of Leaf Epidermis of Chimonobambusa (Bambusoideae)

  Chen Xiao-ya, Hu Cheng-hua, Yu Fu-geng, Ruan Ja-peng

  J Syst Evol. 1993, 31(3): 227-235.
  

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  Micromorphological characters of leaf epidermis of 27 published species of Chimonobambusa Makino were examined by both light and scanning electron microscopes (Table 1). On the adaxial surface, bulliform cells are of irregular or tetragonal shape and in 2-- 3 rows between veins; stomata rare. On the abaxial surface, intercostal long cells are of thin and sinuous wall; short cells solitary, paired, or 3 to several in a row on veins; silica bodies narrow tall or saddle like; bicellular hairs common, with the basal cell slightly longer than or as long as the distal cell; hooks elliptic and apiculate, common in some species and rare in the others; papillae numerous, those overarching stomata showing different patterns (Plates 1, 2). The examined species exhibit three types of leaf epidermis (Table 2 ): 1, papillae widely spread, 4 rod- like papillae overarching the individual stomata, hooks common, including all the six examined species of Sect. Qiongzhuea and 9 species of Sect. Oreocalamus; 2, papillae around stomata only (with the exception of C. microfloscula), the 4 papillae branched or curved, plus several small ones, forming an arciform cover over the individual stomata, hooks sparse, including 7 species of Sect. Oreocalamus and 1 species of Sect. Chimonobambusa; 3, papillae overarching stomata intermediate between the other two types, with the other characters similar to type 1, including 2 species of Sect. Chimonobambusa and 2 species of Sect. Oreocalamus. The three types more or less correspond to the current classification of the genus based on gross morphology, but since Sect. Oreocalamus showed all the three types and type 3 was somewhat intermediate between the other two, present data support a broad sense of the genus Chimonobambusa, and the three sections proposed by Ohrnberger (1990) are to some extent acceptable. The results support the removing of C. luzhiensis from Sect. Qiongzhuea, but reassignment of C. metuoensis to Sect. Qiongzhuea cannot be confirmed. In addition, the leaf epidermis is of diagnostic value at species level. The data support the inclusion of C. linearifolia in C. angustifolia, but do not support that of C. rivularis in C. lactistriata, nor of C. yunnanensis and C. tuberculata in C. armata (Ohrnberger, 1990). In fact, C. yunnanensis, C. tuberculata and C. lactistriata, in addition to C. hirtinoda, are similar to C. qudrangularis in both leaf and gross morphological characters; so is C. pachystachys to C. utilis, C. neopurpurea to C. marmorea, and maybe C. rigidula to C. szechuanensis. They are possibly sister-species.or even conspecific, respectively. The various specific papillate patterns are characteristic of bamboo taxa, andinvestigation of wider samples will be of great interest in bamboo systematics.
• Chromosome Studies of Chinese Species of Malus Mill.

  Liang Guo-lu, Li Xiao-lin

  J Syst Evol. 1993, 31(3): 236-251.
  

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  The genus Malus Mill. of the Maloideae of the Rosaceae consists of about 35 recognized species, which are distributed in temperate region of the Northen Hemisphere including Asia, Europe and North America. Redher's classification divided the genus Malus into 6 series, 5 sections, of which 5 series, 3 section, including 27 species, occur in China. The chromosomal counts on 22 Chinese species were reported previously (Chen et al., 1986; Liang, 1987). However, no comprehensive karyotype analyses have been made for most species reported. Analyzed in this paper are karyotypes of 36 forms, 22 species, and synapsis configuration of 6 polyploid forms in Chinese Malus to explore the relationships among species and provide cytological evidence for delimitation of species in Malus. Chromosome classification follows Li and Chen (1985). The materials used are listed in Table 1 and the vouchers are deposited in our department. The cytological results are shown as follows (see the data in Table 1, Plates 1-- 8 and Fig. 1). 1 Sect. Malus (1) Ser. 1 Baccata The series consists of nine species endemic to China, 7 of which were cytologically observed: i.e. Malus baccata Borkh. 2n=34; Sooo1-3 of M. baccata Borkh. 2n= 34; M . mandshurica Kom. 2n= 34; M. rockii Rehd. 2n=34; Sooo 3-2 of M. rockii Rehd. 2n=51; M. sikkimensis Koehne. 2n=34; M. hupehensis Rehd. 2n=51; Sooo4-4 of M. hupehensis Rehd. 2n=51; M. halliana Koehne. 2n=34 and M. xiaojinensis Cheng et Jiang 2n=68. The karyotypes consist of sm (8- 9 homologous chromosomes) and m (6- 7 homologous chromosomes) as well as st (1-2 homologous chromosomes). The chromosomes range in relative length from 9.84 to 3.74, with the ratio of the longest to the shortest from 2.28 to 2.03 and the mean ratio arms from 2.08 to 1.86. All the karyotypes belong to Stebbins's 3B except 2B of M. Xiaojinensis. Pollen mother cell (PMC) of M. hupehensis contains 17I+17II, and the average synapsis configuration per cell is 2n=3x=51=17. 70I+ 15. 87II+ 0.52III, while PMC meiosis of M. xiaojinensis shows the synapsis configufations of 2n=4x= 68=4II+ 15IV. The latter has a remarkable and constant deleted chromosome, which is also the smallest in the karyotype. (2) Ser. 2 Pumilae. The series consists of six species endemic to China (Yu, 1979), all of which were cytologically examined, with the results: Sooo 9-8 of M. pumila Mill. 2n=34; Sooo 9-12 of M. pumila Mill. 2n=34; Sooo 9-7 of M. sp. 2n=34; M. asiatica Nakai. 2n=68; M. prunifolia Borkh. 2n=34; Soo 12-2 of M. prunifolia Borkh. 2n=34; M. spectabilis Borkh. 2n=51; M. micromalus Makino. 2n=34 and M. sieversii Roem. 2n=34. The karyotypes consist of sm(8-9 homologous chromosomes) and m (6- 7 homologous chromosomes) as well as st (1-2 homologous chromosomes). The chromosomes range in relative length from 10.77 to 3.42, with the ratios of the longest to the shortest from 2.95 to 2.04, and the mean arm ratios from 2.10 to 1.93. Therefore, all the karyotypes belong to 3B. Among more than 20 different forms of M. sieversii, there is only one ploidy level (2n=2x=34). PMC meiosis of M. spectabilis shows the synapsis configurations 2n=3x=51 =5I + 5II+ 12III. 2 Sect. II Sorbomalus (3) Ser. 3 Sieboldianae. This series has only one species endemic to China, M. sieboldii Rehd.: One triploid from Zunyi, Guizhou Province has 2n=3x=51=24m+24sm+3st. The chromosomes range in relative length 8.74-- 4.17, with the ratio of the longest to the shortest 2.08, and the mean arm ratio 1.80. The karyotype, therfore, belongs to 2B. PMC meiosis of this form shows the synapsis conriguration 2n = 3x = 51 = 6I+ 9II+ 9III. (4) Ser. 4 Kansuensis consisting of four species endemic to China, 2 of which were cytologically studied, with the results: M. toringoides Hughes. 2n=34, 51, 68 and M. transitoria Schneid. 2n=34, 51. The karyotypes consist of m (7—9 homologous chromosomes) and sm (7—9 homologous chromosomes) as well as a few st chromosomes. The chromosomes range in relative length from 9.44 to 4.01, with the ratios of the longest to the shortest from 2.26 to 1.83, and the mean arm ratios from 1.87 to 1.72. In M. toringoides four forms, 2x, 3x, 4x from Aba, and 3x from Barkang, in Sichuan Province, are of 2B karyotypes while 2x from Year zhu in Sichuan Province is of 2A. A mixploid with 3x, 4x in M. toringoides is found from Aba, with one smallest deleted chromosome at both levels much like that in M. xiaojinensis. PMC meiosis of triploid M. toringoides from Barkang showed the synapsis configurations of 10I+ 10II+ 7III. In M. transitoria 2x, 3x forms from Barkang, Sichuan Province and Lüda, Liaoning Province belong to 2A, while 2x, 3x forms from Aba, Sichuan Province both belong to 2B. PMC meiosis of triploid form from Lüda showed 9I+ 9II+ 8III synapsis configurations. (5) Ser. 5 Yunnanensis This series consists of four species endemic to China, 3 of which were cytologically studied with the results: M. ombrophila Hand-Mazz. 2n=34; M. honanensis Rehd. 2n=34, and M. yunnanensis Schneid. 2n= 34. The karyotypes consist of nine pairs of m, seven pairs of sm and one pair of st chromosomes. The chromosomes range in relative length from 8.43 to 4.37, with the ratios of the longest to the shortest from 1.88 to 1.86, and the mean arm ratios from 1.75 to 1.72. Therefore, the karyotypes belong to 2A. Two pairs of heterozygous chromosomes, 1st and 10th, are found carrying a small satellite. 3 Sect. III Docyniopsis. This section consists of three endemic species to China, 2 of which were studied with the results: M. formosana Kawak et Koidz. 2n= 34 and M. melliana Rehd. 2n= 34. Both karyotypes consist of nine pairs of m, seven pairs of sm and one pair of st chromosomes. The chromosomes range in relative length from 8.90 to 4.48, with the ratios of the longest to the shortest 1.85 and 1.83, and the mean arm ratios 1.67 and 1.66. Both karyotyoes brlong to 2A. As mentioned above, the karyotyoes of all the species examined are relatively uniform, i.e. chromosomes small; chromosomes gradually decreasing in size from the longest to the shortest; chromosomes mostly m and sm; and 1 —2 pairs of st chromosomes always present. The karyotypes vary in the ratio of the longeat to the shortest, the ratio of arms and the number of different chromosome types and satellites. From cytological investigations and previous reports (Chen et al., 1986; Liang, 1987), the cytology of the genus varies mainly in euploidy variation and chromosome structure. Intraspecific polyploids and polyploid species make up 58. 1% in 24 species endemic to China reported. The karyotype analyse has revealed the presence of three major types in Malus: 2A, 2B and 3B. The deletion in some polyploid forms shows another structural variation. It is clear that the three types are generally consistent with the separation of three sections of Chinese Malus. According to Stebbins's (1971), the evolutionary trend of karyotypes is from 2A to 2B and then 3B. The karyotypic data are in accordance with the evidence from morphology (Langenfeld, 1971; Huckins, 1972) and biochemistry (Williams, 1982). They both show that the section Docyniopsis and series Yunnanenses are the most preimitive, while the series Pumilae, Baccatae of the section Malus are the most advanced or derived. The most typical external characters in Docyniopsis and the series Yunnanenses e.g. grit-cells in fruit, leaves folded in bud, and the inflorescence according to Ponomarenko’s (1986) classification, might be very primitive. The 2A karyotypes in the genus correlated with many typically premitive characters. From the data mentioned above, we may suppose that the series Sieboldianae and Kansuenses in the section Sorbomalus are intermediate in evolution. In general, Southwest China is regarded as the probable area of the origin of Malus, and the section Docyniopsis and the series Yunnanenses as the relict primitive forms. This area is probably also the center of differentiation. A good deal of attention has been paid by horticulturists. A comparison of the karyotypes in the series Pumilae shows that the diploid M. sieversii has 14 m chromosomes, while the other diploids M. pumila, M. sp. and an alien species M. sylvestris Mill. from Europe ( Liang, unpub.) have 12 m chromosomes. Thus M. sieversii may be relatively primitive, although the ratio of the longest to the shortest and the ratio of mean arm ratio are similar. The present investigation confirms Ponomarenko’s view that the diploid M. sieversii might be an ancestor of cultivated apple (M. pumila). The wild polyploids of Malus were usually considered as allopolyploid (Brown, 1976). Studies on the meiotic pairing of five triploids and one tetraploid have revealed diverse types of polyploidy. M. hupehensis is a typical allotriploid species, M. spectabilis, M. sieboldii, M. toringoides and M. transitoria are segmental allotriploids, and M. xiaojinensis is almost an autotetraploid. Apomixis in all known polyploid species of Malus is facultative. The principal adaptive advantage of facultative apomixis is to make sterile individuals restore fertility, and also to maintain evolutionary potential. Therefore, it plays an important role in evolution of Malus, but taxonomy of apomictical groups remains highly controversial. Usually, the apomictical taxa are not treated as species level (Stace, 1980; Hong, 1990). In Rehder’s (1940) system, only M. hupehensis was recognized as species in apomictical taxa of Malus. We think this treatment to be reasonable because M. hupehensis is an allotripoid and so far its diplois form has not been found. New species M. xiaojinensis described by Cheng and Jiang (1985) is nearly an autotetraploid, and its karyotype constitution is extremely similar to the tetraploid form of M. toringoides from Aba , Sichuan Province, especially the remarkable similarities of the smallest deleted chromosome, morphology and biochemistry (Li, 1990). We think, therefor, that the triploid form of M. toringoides from Aba, Sichuan with 3x, 4x mixploidy might be the maternal plant of M. xiaojinensis. For this reason, M. xiaojinensis should be placed in apomictical polyploid complex of M. toringoides of the seriesKansuensis, and this is also in accordance with Rehder’s classfication. Decontaminated thianthrene disproportion. Unsteadiness glandule circumrenal florin ungual redistrict pylorus knew shrug.
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• A Numerical Taxonomic Study of Citrus and Its Close Relatives

  Zhong Guang-yan, Ye Yin-min

  J Syst Evol. 1993, 31(3): 252-260.
  

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  Fifty nine taxa of Citrus, Fortunella and Poncirus were studied by hierarchical agglomerative clustering analysis and quantas type 3 analysis using 86 morphological characters. Five affinity groups were obtained in Citrus. The first group includes C. medica, C. limon, C. limonia, C. aurantifolia and C. iambhiri; the second includes C. grandis, C. aurantium, C. sinensis and C. paradisi; the third is mandarin (C. reticulata); the forth is C. ichangensis; The last is C. hongheensis. Mandarin oranges show two subgroups: one comprise satsuma, King, Shagan, Tankan and Bendiguangju, and the other consists of the remaining typical mandrins. The two papeda oranges, honghe papeda and ichang papeda, could not be clustered into the same group, indicating that they might have evolved from two compeletly different ancestors. Pumelos(C. grandis) show some relationships to honghe papeda orange. C. limonia is assumed to be a hybrid between rough lemon and mandarin. The sweet orange(C. sinensis) is considered to be the offspring of pomelo and mandarin. Zhekiang jinju a small mandarin generally considered similar to calamondin in China, should be a true mandarin orange based on our results. Poncirus was well distinguished from the other two genera, Citrus and Fortunella, by the numerical methods used in this study. Our results show that citron (C. medica), mandarin (C. reticulata) and pumelo (C. grandis) are the original species of the subgenus Citrus, which was congruent with the re-sults obtained by Barrett and Soost (1976), Potvin (1983) and Handa (1985).
• Surface Ultrastructure of Seeds of Chinese Cuscuta

  Huang Jian-zhong, Li Yang-han, Zhang Yu-ying

  J Syst Evol. 1993, 31(3): 261-265.
  

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  The surface ultrastructure of seeds of chinese Cuscuta was observed under SEM. The main results are as follows: the seed surface could be divided into three parts: hilum, hilum-rim and spermoderm. The hilum is collar-shaped or higher or lower than level of hilum-tim. Hilum-rim cells are mostly radiately arranged and less frequently in reticulate or other types of arrangement. The spermoderm shows at least three patterns of ornamentation, cerebelloid, striate or irregular, on which some tuberculate or small clumps of granules could be found sometimes According to the surface ultrastructure of the hilum, hilum-rim and spermoderm, the observed dodder seeds could be divided into three types. The surface characters of dodder seeds are also important both for diagnosis ofspecies and for quarantine of dodders.
• New Taxa of Labiatae from China

  Guo Xin-hu

  J Syst Evol. 1993, 31(3): 266-272.
  

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  One new genus, one new species and one new subspecies of Labiatae from Anhui and Zhejiang province of China are described and one new combination is made in this paper. They are Pogonanthera H. W. Li et X. H. Guo, P. caulopteris H. W. Li et X. H. Guo, P. intermedia (C. Y. Wu et H. W. Li) H. W. Li et X. H. Guo and Paraphlomis foliata (Dunn) C. Y. Wu et H. W. Lissp. montigena X. H. Guo et S. B. Zhou.
• A Revision of the Genus Leptomischus Drake

  Lo Hsien-shui

  J Syst Evol. 1993, 31(3): 273-276.
  

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  In this revision, three species are recognised. A discussion is made on merging the genus Polysolenia into Leptomischus. Indopolysolenia burmanica Deb et Rout is reduced to a synonum of Leptomischus primuloides Drake. A key to species is given below: l. Flowers larger, with corollas more than 2cm long. 2.Stipules 3-fid or torn; Leaves narrowly lanceolate; stigmas with 2 short obtuse lobes ...................................................... 1.L. wallichii 2.Stipules entire; leaves obovate to elliptic; stigmas with 2 slender elongate lobes ......................................................... 2. L. primuloides 1.Flowers smaller, with corollas 6-6.5mm long; stigmas with 2 lanceolate lobes .......................................................... 3. L. parviflorus
• A New Series of Parnassia L. (Saxifragaceae)

  Ku Tsue-chih

  J Syst Evol. 1993, 31(3): 276-276.
  

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• New Taxa of Salix L. from West China

  Ding Tuo-ya, Fang Chen-fu

  J Syst Evol. 1993, 31(3): 277-280.
  

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  Two new species and one new variety of Salix L. from western China, namcl. S. ludingensis T. Y. Ding et C. F. Fang, S. neoamnematchinensis T. Y. Ding et C. F. Fang, and S. taoensis Gorz var. leiocarpa T. Y. Ding et C. F.Fang, are reported in this paper.
• The Origin of Chinese Traditional Drug “Shuigaoben” from Hubei

  Zhang Hai-dao, Fang Shu-min

  J Syst Evol. 1993, 31(3): 281-285.
  

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  A new variety of Ligusticum L. (Umbelliferae), L. sinense var. hupehense collected from Hubei province, is described in this paper.. Cytological investigation shows that the new plant is a diploid, with the karyotypic formula 2n=22=14m+ 8sm. The parameters of chromosomes and the idiogram are given. The comparison of ARC (arm ratio curve) diagram between the new variety andL. sinense is also given.
• The Identification and Revision on the Type of Cochleariopsis Y. E. Zhang (Cruciferae)

  Lu Lian-li

  J Syst Evol. 1993, 31(3): 286-287.
  

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  The genus Cochleariopsis Y.H. Zhang was described in 1985, with C. zhejiangensis Y. H. Zhang, the only member of the genus, as its type. However, this species had been published by O.E. Schulz earlier in 1923, named Cochlearia warburgii O.E. Schulz. Hence, this species is not new one, and the type of the ge-nus should be Cochleariopsis warburgii (O. E. Schulz)L. L. Lu .
• One New Species of Pedicularis L. from Xizang

  Yang Han-bi

  J Syst Evol. 1993, 31(3): 288-290.
  

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  Pedicularis nyingchiensis H. P. Yang et Y. Tateishi is described as newfrom Xizang, China.
• One New Species of Mecodium (Hymenophyllaceae) from Guangxi

  Zhou Hou-gao, Chu Wei-ming

  J Syst Evol. 1993, 31(3): 291-293.
  

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  One new species of the genus Mecodium, M. paramnioides H. G.Zhou et W. M. Chu, is described from Guangxi, China.
• A New Morphological Interpretation of the Female Reproductive Organs in Ginkgo biloba L., with a Phylogenetic Consideration on Gymnosperms

  Fu De-zhi, Yang Qin-er

  J Syst Evol. 1993, 31(3): 294-296.
  

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  The ovuliferous structure of Ginkgo biloba L. has been variously interpreted morphologically. As a result the systematic position and the relationship with other gymnosperms of this ancestral gymnosperm have long been under dispute. In the present paper, a brief survey of the main views as to the nature of the ovuliferous structure is givcn. Based on morphological and teratological data previously reported, a new interpretation is proposed. The essential points are summarized as follows: 1. In morphological essence, a fertile dwarf shoot with some ovuliferous structures in Ginkgo biloba L. might as a whole be nothing but a megasporophyll strobilus (female cone), which is shared actually by all the conifers in the gymnosperms. The fertile dwarf shoot has appearance extremely similar to that of the vegetative dwarf shoot, suggesting that in Ginkgo biloba L. the vegetative organs and the reproductive organs have not been yet well differentiated, and thus its megasporophyll strobilus might represent one of the most primitive compound strobilus types. 2. In Ginkgo biloba, the ovuliferous structure borne in the axil of a scale leaf (sometimes a normal leaf) on the dwarf shoot, together with the scale leaf itself, might be the homogenous organ corresponding to the bract-scale and seed-scale complex of the compound female strobilus of the typical conifers. The complex is a relatively isolated reproductive unit on the strobilus. The normal leaves and the scale leaves on the dwarf shoot might be equivalent to the bract-scales in the typical cones, though the normal leaves still retain the vegetative nature as the foliage leaves on the vegetative shoot. The stalk hearing ovules at its top might be equivalent to a seed-scale of the typical cones. 3. The megasporophyll strobilus in Ginkgo biloba, namely a whole fertile dwarf shoot as mentioned above, seems to show much more primitive characteristics than those of typical conifers. In this plant it is very difficult to distinguish the fertile dwarf shoot from the common vegetative dwarf shoot before reproduction time. Moreover, its megasporophyll strobilus often exhibits more atavistic abnormalities than those of other conifers. All the evidence indicates that the primitive ancestor of conifers might have had the fertile organs which might be of basically identical morphology as vegetative shoots, except that in the fertile organs there might exist numerous fertile leaves bearing one or many ovules. 4. The longer stalk of the ovuliferous structure in Ginkgo biloba might have come from mainly a secondary elongation growth of the seed scale, and only a little part of it might be the remains of the original shoot. The fork structure bearing ovules at the top of the stalk might be the rudementary part of the petioles of the only two extremely reduced megasporophylls. The collar around the base of the ovule might be a secondary protective structure. 5. A correct morphological interpretation of the female strobilus in Ginkgo biloba is doubtless of important significance for our better understanding of the evolution of the female reproductive organs in conifers. According to our interpretation mentioned above, together with the concept of the bractscale and seed-scale complex proposed in the present paper, which is mainly based on the concept of the seed-scale complex propose by Florin, here we put forward an evolutionary theory of the bract-scale and seed-scale complex. According to this theory, the female reproductive organs of the ancestral conifers should be very similar, as mentioned above, to the sterile foliage shoot except that the former might have some fertile leaves which could produce ovules at reproduction time. This ancestral female reproductive organ type might have had evolved towards two directions and thus formed two main evolutionary lines. One is represented by the genus Cycas and we may call it the Cycas Evolutionary Line (C-line), in which the megasporophyll strobilus is monopodial, with the fertile leaves and sterile bracts occurring directly on the main axis. The Cycadaceae is the only living gymnosperm member along this evolutionary line. The second line is represented by all the conifers including Ginkgo, which all have the structure of the bract-scale and seed-scale complex, and thus we called it the "Bract-scale and Seed-scale Complex Evolutionary Line" (BS-line). The members along this line have multipodial female strobilus, i.e. compound strobilus. On the main axis occur some sterile vegetative bracts. In the axils of some or most of the bracts occur the seed-scales. The seed-scales are actually the remains of the extremely, or smetimes completely reduced fertile shoots. Each part of the bractt-scale and seed-scale complex and the main axis of the strobilus could have undergone independent or correlated changes, and thus have had formed various types of strobilus which are found in the living conifers. 6. Our theory on the evolution of the bract-scale and seed-scale complex seems to support the division of all the gymnosperms into two major groups as proposed by Chamberlain, and is also in favour of the placement of Ginkgo biloba into the conifers as the most primitive member along BS-line. 7. Based on their similar morphological characters, it can be considered that Ginkgo biloba might have close relationships with the Nageiaceae, Ephedraceae, Welwitschiaceae and Araucariaceae. All these groups have multinerved leaves without costa. These living gymnosperms might have a more direct relationship withthe ancestral cordaites.