The genus Calceolaria in NW South America VIII. The section Calceolaria and appendices to parts I–VIII

Sect. Calceolaria (= Aposecos) of Calceolaria (Scrophulariaceae) in NW South America is revised, and the other species of the section are briefly discussed. The section comprises annual herbs with somewhat succulent stems and leaf–blades that are usually pinnately dissected (pinnatifid). Four species are recognized in the investigated area, viz. C. mexicana, C. tenuis, C. tripartita , and C. chelidonioides , and their chromosome numbers are reported (2n = 32, 60, 64, and 60 respectively). The basic number in sect. Calceolaria is x = 8. Two new combinations are made, viz. C. mexicana ssp. prostrata (Kränzlin) Molau and C. mexicana ssp. perijensis (Pennell) Molau. The species of sect. Calceolaria are intersterile and facultatively autogamous, and each species comprises numerous pure lines, some of which may be morphologically discernible.
In the appendices, chromosome numbers are listed and the occurrence of elaiophores in all NW South American species of Calceolaria is indicated. The chromosome numbers of C. crenata ssp. australis, C. adenanthera, C. gossypina, C. semiconnata, C. stricta, C. sericea , and C. comosa (all 2n = 36) have not previously been reported. A revised key to the sections of Calceolaria in NW South America is also provided.     

朱砂根复合群形态性状的数量分析

朱砂根复合群—朱砂根(Ardisia crenata Sims)、红凉伞(A. crenata var. bicolor (E. Walker) C. Y. Wu C. Chen),和岭南紫金牛(A. linangensis C. M. Hu)之间存在一些过渡类型,很难依据少数几个关键性状对它们明确划分。本文运用E. Anderson的形象化散点图和柱状图的方法对全国14个省区135份标本的9个性状进行了分析,结果表明：岭南紫金牛应为独立的种,支持将红凉伞归并。     

Notes on Tragia, Dalechampia and Clutia (Euphorbiaceae) in Ethiopia and Somalia

Tragia mitis is lectotypified and T. cinerea is neotypified. T. abortiva, T. ashiae, T. crenata, T. doryodes, T. mixta, T. negeliensis, T. triumfettoides and T. uncinata are described as new. The new combination Dalechampia pavoniifolia is made. Clutia lanceolata is extended to include African material and the new combination Clutia lanceolata ssp. robusta made.     

Spatial arrangement of branches in relation to slope and neighbourhood competition

To gain a better understanding of the effects of spatial structure on patterns of neighbourhood competition among hardwood trees, the three-dimensional extension of primary branches was surveyed for ten community-grown Castanea crenata (Fagaceae) trees with respect to the positioning of neighbouring branches and the slope of the forest floor. There were significantly more branches extending towards the lower side of the slope than towards the upper side, but structural properties such as branch length and vertical angle were not affected by slope. When horizontal extension of a branch towards its neighbour was compared for a C. crenata branch and a neighbouring heterospecific, the former was significantly narrower than the latter when the inter-branch distance (horizontal distance between the base positions of two neighbouring branches) was short (< approx. 5 m). Castanea crenata branches tended to extend in a direction avoiding neighbouring branches of heterospecifics when the inter-branch distance was short. Furthermore, for an inter-branch distance <3 m, the horizontal extension of a C. crenata branch was less when it was neighbouring a heterospecific branch than when neighbouring a conspecific branch. These results suggest that horizontal extension of C. crenata branches is more prone to spatial invasion by nearby neighbouring branches of heterospecifics, and that the invasion can be lessened when C. crenata trees are spatially aggregated. The reason why such an arrangement occurs is discussed in relation to the later leaf-flush of C. crenata compared with that of other species in the forest.     

日本栗引种栽培试验研究

报道日本栗在桂林引种栽培研究结果。日本栗筑波、银寄品种种植表现出生长快、长势好、早实、早熟、丰产、粒大、质优等优良性状,是值得大力推广的优良品种。适当密植、幼树整形修剪、合理配植授粉树、加强肥水管理、人工辅助授粉及适时除雄是日本栗获得早实丰产的关健技术措施。     

Phylogeographical structure revealed by chloroplast DNA variation in Japanese beech (Fagus crenata Blume)

Intraspecific genetic variation in three non-coding chloroplast DNA (cpDNA) regions (trnT-L and trnL-F spacers, and trnL intron) of Japanese beech (Fagus crenata Blume) was investigated. This species is a major constituent of the typical cool-temperate deciduous forests in Japan. Twenty-one F. crenata populations from throughout Japan, and four F. japonica populations, a close relative of F. crenata, were examined. Seven haplotypes were distinguishable in F. crenata based on nucleotide substitutions and indels. Pairwise nucleotide diversities among haplotypes ranged from 0.0000 to 0.0042 for F. crenata, including F. japonica. The geographical distribution of cpDNA haplotypes was found to be highly structured in F. crenata. Four haplotypes predominated: haplotypes FC1 and FC4 are prevalent on the Pacific Ocean coast, haplotype FC6 is prevalent on the Japan sea coast from the San-in district to Hokkaido, whilst haplotype FC3 is restricted to northern Kyushu and the western-most part of Honshu. Two haplotypes (FC5 and FC7) are restricted to single populations and one haplotype (FC2) is a derivative of FC1. Each of these haplotypes, except FC2, are thought to be derived from different glacial refugia. Phylogenetic analysis showed that neither F. crenata nor F. japonica was monophyletic for the haplotypes, suggesting either ancestral polymorphism or ancient introgression between the lineages of these two Fagus species.     

Cytotaxonomical Studies of Tea Plants

Nine forms of Camellia sinensis (L) Kuntze from Guizhou, Yunnan, Sichuan and Fujian Provinces were cytotaxonomically studied in this work. The wall degradation hypotonic method was used for preparing chromosome samples which were stained in Giemsa. The micrographs of their somatic metaphase are shown in Plates 1-2, the ranges of chromosome relative lengths, arm ratios, karyotype formulae and classification of karyotypes according to Li and Chen (1985) are shown in Table 1 and the idiograms in Fig. l. The main points are mentioned as follows: (1) All these forms are found to be diploid with 2n = 30, and they are reported for the first time except Yunnan(Fengqing)cultivated form. (2) All the karyotypes of the forms examined are relatively uniform, which reveal the gradual decrease in size from the longest to the shortest chromosomes, similar chromosomal relative length, smaller chromosome types varying between 2- 5μm. The karyotype mainly consists of metacentric (m) and submetacentric (sm) chromosomes. Sat-chromosomes are difficult to be discovered on mid -metaphase chromosomes. The cytological information confirms the close relationship among these tea forms. (3) According to the asymmetry of karyotype, the nine tea forms are divided into two types: one consists of the arbor with macrophyll, i.e. Guizhou and Yunnan( Fengqing and Tengchong)cultivated forms, which have the”2A” karyotype possessing metacentric(m) and submetacentric(sm) chromosomes. The karyotypes might be of the most symmetrical or primitive type ;the other consists of the bush or microarbor with mostly medium- and microphyll, less frequently macrophyll, i.e. Qianmei 419, Qianmei 502, Yaan,Fuding, Sichuan and Qianmei 101 cultivated forms, which have the “2A” and “2B” karyotypes, consisting of metacentric (m), submetacentric (sm) and a pair of subtelocentric (st) chromosomes, and more asymmetrical than that of the first karyotype. They might be more advanced. (4) In recent years, two subspecies, i.e. ssp. “yunnan”and ssp. “bohea”, including seven varieties, in C. sinensis were recognized but not formally published by Zhuang et al. (1981). From the present work and the previous reports the authors agree with Zhuang’s classification, because recognition of two subspecies is remarkably consistent with the two kinds of karyotypes reported here. C.sinensis ssp. “yunnan” with arbor and macrophyll characters has more symmetrical karyotype than that of C. sinensis ssp. “bohea” with bush or microarbor and mostly mediumand microphyll characters. That is to say, ssp.”yunnan” is more primitive than ssp. “bohea”. (5) Based on the karyotype analysis of arbor and macrophyll teas reported before and here (see Table 3), the authors support the viewpoint that YunnanGui zhou Plateau is the original center of tea Plants, whereas Assam form in In-dia is derived.     

Search for the wild ancestor of buckwheat III. The wild ancestor of cultivated common buckwheat, and of tatary buckwheat

By surveying wild Fagopyrum species and their distribution in southern China and the Himalayan hills, I arrived at the conclusion that the newly discovered subspecies F. esculentum ssp. ancestralis Ohnishi is the wild ancestor of cultivated common buckwheat, while previously known wild tatary buckwheat,F. tataricum ssp. potanini Batalin is the wild ancestor of tatary buckwheat. Their original birthplace is revealed to be northwestern corner of Yunnan province for common buckwheat judging from the distribution of wild ancestor, and to be the northwest part of Sichuan province for tatary buckwheat judging from allozyme variability in wild tatary buckwheat. F. cymosum is not the ancestor of cultivated buckwheat; it is only distantly related to cultivated buckwheat, in morphology, isozymes and cpDNA. Several genetic, ecological and taxonomic categories which should be taken into consideration in examining the origin of buckwheat were discussed. Key Words: Fagopyrum esculentum ssp. ancestralis; Fagopyrum tataricum ssp. potanini; southern China theory of origin of buckwheat. Contribution from Plant Germ-Plasm Institute, Faculty of Agriculture, Kyoto University. No. 78.     

Origin(s), evolution,and systematics ofCucurbita pepo (Cucurbitaceae)

Numerical studies of morphological data and starch gel electrophoresis have provided a new perspective on important issues pertinent to the origin(s) and subsequent evolution of domesticatedCucurbita pepo. Wild C.texana and/orC. fraterna appear to be the most likely candidates for progenitor(s) of the domesticate. Populations of texana-like plants from beyond Texas share various attributes with Texas populations, suggesting that C. texana once had a more widespread distribution to the northeast. The possibility exists thatC. pepo was domesticated independently in eastern U.S., as well as in Mexico, which is in line with recent archeological findings. Multiple domestications are also supported by allozyme data indicating a substantial divergence within the species. A new classification consisting of C.pepo ssp.pepo (origins in Mexico),C. pepo ssp.ovifera var. ovifera (origins in eastern U.S.), and C.pepo ssp.ovifera var. texana (spontaneous populations in eastern U.S.) is proposed.     

Polymorphic chloroplast simple sequence repeat primers for systematic and population studies in the genus Hordeum

In this study we report the development of primers to amplify polymorphic chloroplast simple sequence repeats in the genus Hordeum , which includes cultivated barley ( H. vulgare ssp. vulgare ) and its wild progenitor H. vulgare ssp. spontaneum . Polymorphic products were amplified in a wide range of Hordeum spp. and intraspecific variation was detected in both cultivated and wild barley. A decrease in cytoplasmic diversity was observed between sspp. spontaneum and vulgare as well as between ssp. vulgare landraces and cultivars, which is characteristic of domestication processes in many crop species. We also observed possible evidence for reticulate evolution of H. brachyantherum polyploids, with apparent multiple cytoplasmic introgressions during successive polyploidization events.     

A taxonomic revision of Secale (Triticeae, Poaceae)

Several taxa have previously been recognized within Secale , but most of them are difficult or even impossible to distinguish morphologically. We recognize only three species: S. sylvestre, S. strictum , and S. cereale. Secale strictum has priority over S. montanum and has two subspecies, ssp. strictum and ssp. africanum , and two varieties within ssp. strictum , van strictum and var. ciliatoglume comb. nov. Secale cereale is also treated as having two subspecies. The cultivated taxa, marked by their tough rachises, are placed in ssp. cereale and the wild or weedy taxa that have more or less fragile rachis, in ssp. ancestrale. A complete synonymy is given for S. cereale , but typification has been omitted because, in many instances, type material does not exist or has been impossible to trace.     

High segregation distortion in maize B73 x teosinte crosses

Two genetic linkage maps of cultivated maize inbred lines and teosinte species were constructed. One population comprised 81 F(2) individuals derived from a cross between maize inbred line B73 and Zea mays ssp parviglumis, while the second consisted of 63 backcross individuals from a cross of maize inbred line B73 with Z. mays ssp diploperennis. In the B73 x Z. mays ssp parviglumis F(2) population, 172 simple sequence repeat (SSR) markers were mapped to 10 chromosomes, which covered 2210.8 cM. In the B73 x Z. mays ssp diploperennis backcross population, 258 SSR markers were mapped to 10 chromosomes, covering 1357.7 cM. Comparison of the two maps revealed that the total map length of Z. mays ssp diploperennis covers 1357.7 cM, which is about 61.4% of that of Z. mays ssp parviglumis (2210.8 cM). Extensive segregation distortion regions were found on chromosomes 1, 2, 3, 5, 6, 7, and 10 in the B73 x Z. mays ssp parviglumis F(2) population and on chromosomes 1-5 and 8-10 in the B73 x Z. mays ssp parviglumis backcross population. Segregation distortion analysis confirmed that the segregation distortion ratio in the interspecific population B73 x Z. mays ssp diploperennis was higher than in B73 x Z. mays ssp parviglumis. We found that the recombination distances are highly variable in these genetic crosses between cultivated and wild species of maize.     

Genetic diversity of wild grapevine populations in Spain and their genetic relationships with cultivated grapevines

The wild grapevine, Vitis vinifera L. ssp. sylvestris (Gmelin) Hegi, considered as the ancestor of the cultivated grapevine, is native from Eurasia. In Spain, natural populations of V. vinifera ssp. sylvestris can still be found along river banks. In this work, we have performed a wide search of wild grapevine populations in Spain and characterized the amount and distribution of their genetic diversity using 25 nuclear SSR loci. We have also analysed the possible coexistence in the natural habitat of wild grapevines with naturalized grapevine cultivars and rootstocks. In this way, phenotypic and genetic analyses identified 19% of the collected samples as derived from cultivated genotypes, being either naturalized cultivars or hybrid genotypes derived from spontaneous crosses between wild and cultivated grapevines. The genetic diversity of wild grapevine populations was similar than that observed in the cultivated group. The molecular analysis showed that cultivated germplasm and wild germplasm are genetically divergent with low level of introgression. Using a model‐based approach implemented in the software structure , we identified four genetic groups, with two of them fundamentally represented among cultivated genotypes and two among wild accessions. The analyses of genetic relationships between wild and cultivated grapevines could suggest a genetic contribution of wild accessions from Spain to current Western cultivars.     

Lentil domestication: On the quality of evidence and arguments

The current views on lentil domestication are based on biological attributes of the wild progenitorLens culinaris ssp. orientalis and on assumptions which have never been tested. Seed dormancy, a major factor in the adaptation of ssp.orientalis to its natural habitat, makes it inappropriate for cultivation, because poor germination causes seed yield following cultivation to be equal to the amount of sown seeds. Higher yield, resulting from the evolution of a non-dormant type can be obtained only after five or six cycles of unprofitable cultivation. It is doubtful that incipient farmers would have undertaken such an endeavor without preexisting knowledge that non-dormant types could eventually be obtained. Experiments involving the sowing of wild lentil would have been much more successful if the non-dormant types were present in appreciable quantities in the seed stock. Establishment of that type in the natural population would have required a period of seven to eight years with favorable growing conditions allowing the non-dormant type to become widespread in the population, followed by massive predation by man reducing the hazard of a population explosion. The close similarity between isozyme profiles of the cultivated lentil and its wild progenitor indicates that lentil cultivation was attempted with seeds derived from different populations and in different areas.     

Experimental studies of adaptation in Clarkia xantiana. II. Fitness variation across a subspecies border

Because the range boundary is the locale beyond which a taxon fails to persist, it provides a unique opportunity for studying the limits on adaptive evolution. Adaptive constraints on range expansion are perplexing in view of widespread ecotypic differentiation by habitat and region within a species' range (regional adaptation) and rapid evolutionary response to novel environments. In this study of two parapatric subspecies, Clarkia xantiana ssp. xantiana and C. x. ssp. parviflora, we compared the fitness of population transplants within their native region, in a non-native region within the native range, and in the non-native range to assess whether range expansion might be limited by a greater intensity of selection on colonists of a new range versus a new region within the range. The combined range of the two subspecies spans a west-to-east gradient of declining precipitation in the Sierra Nevada of California, with ssp. xantiana in the west being replaced by ssp. parviflora in the east. Both subspecies had significantly higher fitness in the native range (range adaptation), whereas regional adaptation was weak and was found only in the predominantly outcrossing ssp. xantiana but was absent in the inbreeding ssp. parvifilora. Because selection intensity on transplants was much stronger in the non-native range relative to non-native regions, there is a larger adaptive barrier to range versus regional expansion. Three of five sequential fitness components accounted for regional and range adaptation, but only one of them, survivorship from germination to flowering, contributed to both. Flower number contributed to regional adaptation in ssp. xantiana and fruit set (number of fruits per flower) to range adaptation. Differential survivorship of the two taxa or regional populations of ssp. xantiana in non-native environments was attributable, in part, to biotic interactions, including competition, herbivory, and pollination. For example, low fruit set in ssp. xantiana in the east was likely due to the absence of its principal specialist bee pollinators in ssp. parviflora's range. Thus, convergence on self-fertilization may be necessary for ssp. xantiana to invade ssp. parviflora's range, but the evolution of outcrossing would not be required for ssp. parviflora to invade ssp. xantiana's range.     

Systematics, ecology and phylogeographic significance of Olea europaea L. ssp. maroccana (Greuter & Burdet) P. Vargas et al., a relictual olive tree in south-west Morocco

Within the genus Olea , the olive ( Olea europaea L.) possesses the largest geographic distribution, and several subspecies have been described. South-west Morocco harbours an endemic olive, O. europaea ssp. maroccana (Greuter & Burdet) P. Vargas et al. , located in the western part of the High Atlas (mainly on the southern slope of the Ida-ou-Tanane massif) and in the western Anti Atlas. The existence and the taxonomic position of this tree have long remained uncertain, but the combination of several morphological traits is unique. Nuclear and cytoplasmic DNA polymorphism also demonstrate that this tree is a well-differentiated and relictual taxon, probably originating from an ancestral unit within tropical Africa. Our findings refute the opinions of several authors who consider ssp. maroccana to be intermediate between the cultivated olive ( O. europaea ssp. europaea ) and the Saharan olive ( O. europaea ssp. laperrinei (Batt. & Trab.) Cif.). According to cpDNA and mtDNA analysis, the Moroccan olive and the Canarian olive ( O. europaea ssp. guanchica P. Vargas et al. ) are related and belong to the same sub-taxon. This biogeographical affinity is strongly supported by other recent phylogenetic studies. The long-term viability of ssp. maroccana is threatened by several factors: (1) limited distribution and reduced numbers of individuals within isolated populations, (2) fragmentation processes related to human and livestock pressures, and (3) putative gene flow between this taxon and the cultivated olive. In order to ensure the long-term survival of this endemic olive, a specific program of conservation should be implemented in the near future.     

A comparative analysis of genetic polymorphism in wild and cultivated barley from Tibet using isozyme and ribosomal DNA markers.

This study was conducted to address some of the issues concerning the possible significance of Tibet in the origin and evolution of cultivated barley. A total of 1757 barley accessions from Tibet, including 1496 entries of Hordeum vulgare ssp. vulgare (HV), 229 entries of the six-rowed wild barley H. vulgare ssp. agriocrithon (HA), and 32 entries of the two-rowed wild barley H. vulgare ssp. spontaneum (HS), were assayed for allozymes at four esterase loci. A subsample of 491 accessions was surveyed for spacer-length polymorphism at two ribosomal DNA loci. Genetic variation is extensive in these barley groups, and the amount of genetic diversity in cultivated barley of this region is comparable with that of cultivated barley worldwide. The level of genetic variation of HA is significantly lower than the other two barley groups, and there is also substantial heterogeneity in the level of polymorphism among different agrigeographical subregions. However, little genetic differentiation was detected among the three barley groups (HV, HA, and HS), as well as among different agrigeographical subregions. Comparison of the results from this and previous studies indicated a strong differentiation between Oriental and Occidental barley, thus favoring the hypothesis of a diphyletic origin of cultivated barley.     

Structural diversity and endotoxic activity of the lipopolysaccharide of Yersinia pestis

The endotoxic activity of the lipopolysaccharides (LPS) with defined chemical structure from Yersinia pestis strains of various subspecies differing in their epidemic potential was studied. The LPS of two strains of Y. pestis ssp. caucasica and ssp. altaica, whose structures have not been studied earlier, were analyzed by high-resolution mass spectrometry. In addition to reported structural changes, an increase in the degree of LPS phosphorylation was observed when strain I-2377 (ssp. altaica) was cultivated at an elevated temperature. A high tumor necrosis factor alpha(TNF-alpha)-inducing activity observed for LPS samples from Y. pestis cultures grown at 25 degrees C correlated with an increased degree of lipid A acylation, particularly, with the presence of the hexaacyl form of lipid A, which was absent from the LPS when bacteria were cultivated at 37 degrees C. No correlation was found between the lethal toxicity of the LPS in vivo and its ability to induce TNF-alpha production in vitro.     

Genetic relationships among subspecies of Vigna unguiculata (L.) Walp. based on allozyme variation

 The cowpea [Vigna unguiculata (L.) Walp.] is a morphologically and genetically variable species composed of wild perennial, wild annual, and cultivated forms that are mainly used for edible seeds and pods. In this study, genetic variation in 199 germplasm accessions of wild and cultivated cowpea was evaluated using an allozyme analysis. The results from this survey showed that wild cowpea exhibits genetic variation perfectly fitted with the existing morphological classification. The cowpea gene-pool is characterized by its unusually large size. It encompasses taxa (ranked as subspecies) that could be considered as different species considering the high genetic distances observed between accessions belonging to different taxa. These subspecies can be classified into three groups characterized by their breeding systems: perennial outcrossers, perennial out-inbreds, and inbred annuals. Allozyme data confirm this grouping. Perennial outcrossers look primitive and are more remote from each other and from perennial out-inbreds. Within this large gene-pool, mainly made of perennial taxa, cultivated cowpeas (ssp. unguiculata var. unguiculata) form a genetically coherent group and are closely related to annual cowpeas (ssp. unguiculata var. spontanea) which may include the most likely progenitor of cultivated cowpeas. Received: 15 June 1998 / Accepted: 29 September 1998