Karyology of theScorzonera (Compositae) species from the Iberian Peninsula

A karyological study of 15 taxa ofScorzonera L. from the Iberian Peninsula has been made. The chromosome numbers found inS. hispanica var.pinnatifida, S. baetica, S. reverchonii, S. angustifolia, S. laciniata var.calcitrapifolia and var.subulata (2n = 14) are new. Diploid cytotypes with 2n = 14 and 2n = 12 prevail, andS. hispanica var.crispatula is the only taxon which exhibits autopolyploidy (2n = 14, 28). x = 7 is considered to be the base chromosome number within the genus, with x = 6 being derived from it by translocation. This and detailed karyotype analyses allow to group the Iberian Peninsula species ofScorzonera into three groups.     

Reproductive biology ofFrangula alnus (Rhamnaceae) in southern Spain

Various aspects of the reproductive biology of the perennial hermaphroditeFrangula alnus Mill. were studied in two populations located in the province of Cádiz, southern Spain. Flowering extends from the second week of May to early July. The small, whitegreenish, entomophilous flowers are incompletely protandrous and last 8–10 days, but pollen transfer takes place only on days 1–3. A very weak, pleasant odour along with nectar and pollen attract over 45 insect species, of which 21 are probable pollinators. These are mainlyDiptera but also includeHymenoptera and the unusual (as flower visitors)Mecoptera. Reproduction is exclusively sexual and strictly xenogamous, with pollen transfer depending solely on insect vectors. Although flower morphology and individual flower phenology do not fully prevent self-pollination, and geitonogamy can easily take place, the level of autogamy was negligible. Therefore, some self-incompatibility mechanism is operative in this species. Only 2.8% of open-pollinated flowers set fruit. At the flower level, fruit initiation was apparently limited by availability of cross-pollen, which in turn seemed influenced by the structure of the population. The seed/ovule ratio in ripe fruits was ca. 50%. Predispersal (maternal) reproductive success (percent ovules becoming filled seeds) was 1.42%. For mature individuals this corresponded to ca. 430 to 1560 potential offspring per year.     

Sexual reproduction in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The reproductive biology of someScrophularia species in the Iberian Peninsula and Balearic Islands has been studied. Levels of fruit-set in bagged and unbagged inflorescences, the production of seeds/capsule, the effectiveness of seed-set following cross-pollination in the genus and the pollen/ovule ratios have been analyzed.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and Balearic Islands. II. For first part seeOrtega Olivencia & Devesa 1993.     

Different levels of morphometric variation in three heterogamous dogrose species (Rosa sect.Caninae, Rosaceae)

Difficulties in delimiting well-defined entities in the dogroses (Rosa sect.Caninae) has resulted in very variable taxonomic treatments. The present study was undertaken to provide a background for taxonomy as well as plant breeding. Morphometric diversity was analysed on seedlings obtained from field collections in South Sweden of three species,Rosa dumalis, R. rubiginosa andR. villosa. A canonical variates analysis showed that the three species are relatively distinct whereas two subspecies ofR. dumalis were less well discriminated. Analyses of variance demonstrated that intraspecific variation is pronounced inR. dumalis and, to a lesser extent, inR. villosa.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Reproductive biology in the primitive relic AngiospermDrimys brasiliensis (Winteraceae)

From field observations onDrimys brasiliensis, principally in the Botucatu region of São Paulo State, Brazil, new data on the reproductive biology, the rhythm of growth, and the development of lateral cymose inflorescences, flowers and fruits are presented. Pollination accelerates the rate of flower-development for about 4–6 days. Pollination experiments show thatD. brasiliensis is not self-sterile; because of mechanical devices the sticky pollen grains do not normally come into contact with the stigmata unless an animal pollen vector is involved. The pollinators are diurnalColeoptera, Diptera andThysanoptera which eat from the pollen, lick from the stigmatic exudates and (in case of the flies) probably also from the staminal glands. Fruit- and seedeaters are birds which seem to be the main dispersal agents. Establishment of new individuals normally is through seedlings, but also by vegetative propagation through plagiotropous branches which may root and separate from the mother plant. The morphological, developmental and reproductive aspects inD. brasiliensis are discussed in a wider context, compared with data from otherMagnoliidae, and related to aspects of early Angiosperm evolution.     

Reproductive biology of the Andean-disjunct genusRetanilla (Rhamnaceae)

Various aspects of the reproductive biology of the Andean-disjunct genusRetanilla (DC.) Brongn. were studied in populations ofR. trinervia andR. ephedra located in central Chile and ofR. patagonica in southern Argentina. Flowering extends from -austral- winter (early spring inR. patagonica) to early summer. The small, white, entomophilous flowers are incompletely protandrous and last four to five days. A weak pleasant odor along with nectar and pollen attract thirty-seven diurnal insect species, ten of which are probable pollinators: these include the honey bee, eight species of solitary bees pertaining toAnthophoridae (1 sp.),Colletidae (3 spp.) andHalictidae (4 spp.), and a nemestrinid fly. Although flower morphology and individual flower phenology do not fully prevent self-pollination (and geitonogamy can easily take place), the level of autogamy is low. Therefore, some self-incompatibility mechanism seems operative inRetanilla. Fruit set of open-pollinated flowers is extremely low, with a maximal value of c. 3% inR. patagonica. In this species, ripe fruits contain on average 1.37 viable seeds. Predispersal (maternal) reproductive success (percent ovules becoming viable seeds) is 2.3%. For medium to full-sized individuals this corresponds to c. 4600 potential offspring per year. As presently known,Retanilla is a reproductively uniform group, in which the Andean disjunction seems to have exerted no particular impact, which is consistent with the view that pollination generalization exerts some stabilizing influence on floral morphology and other reproductive traits.Retanilla expresses a basic rhamnacean set of traits (including protandry and self-incompatibility) showing also high pollen production and secondary pollen presentation. The latter two traits seem characteristic of theRetanilla-Trevoa clade, suggesting that a trend to increased male effort and, perhaps, dependence upon polleneating has evolved within the tribe Colletieae.     

DNA fingerprint variation in some blackberry species (Rubus subg.Rubus,Rosaceae)

We have analysed samples from Sweden, Denmark, and Germany of six facultatively apomictic blackberry species to investigate the accordance between a taxonomy based on morphological characters on the one hand, and distribution of genetic variation estimated by DNA fingerprinting on the other hand. DNA fingerprint variation was found to be quite restricted in all species investigated. The first taxonomic group included three species related toRubus nessensis, two being characterized by one very widespread DNA fingerprint in all three countries and a few rare ones, whereas the third species differed between Sweden and Germany. The second taxonomic group included species related toR. gracilis. Two of these species exhibited very similar DNA fingerprints, whereas the third species deviated clearly. The utilization of DNA fingerprinting as a tool in taxonomy is discussed; most likely this method could become a useful complement to morphology, especially in plant groups with reduced levels of genetic recombination.     

Reproductive biology ofAsphodelus aestivus (Asphodelaceae)

InAsphodelus aestivus the large number of showy flowers opened per inflorescence, high nectar and pollen production, and absence of automatic self-pollination, so that insect visitation is necessary for pollination, indicate that cross-fertilization is favoured. LargeHymenoptera are the main pollinators. Hand self-pollinations result in some degree of fruit-set, but cross-pollination also yield low fruit- and seed-set. The perennial habit and polyploid status are in agreement with the breeding system of this species, with sexual reproduction being rather inefficient and active vegetative propagation.     

Multivariate and cladistic analyses of the purple-flowered species ofErysimum (Cruciferae) from the Iberian Peninsula

A taxonomically difficult purple-flowered group within the genusErysimum, restricted to the Iberian Peninsula, is analyzed by multivariate and cladistic analyses. 51 specimens have been scored for 14 characters. Both principal components and discriminant analyses provide support to the recognition of the five species considered by the author, namelyE. linifolium, E. lagascae, E. baeticum, E. popovii, andE. cazorlense. Cladistic analysis, using 7 characters resulted in a single most parsimonious cladogram containing no homoplasies. The pattern of morphologic divergence follows a clear NW.-SE. trend, which is congruent with the topology of the cladogram. This trend significantly affects growth-form as well as fruit characters, both providing the main grounds for species recognition. The different behavior and significance of several characters in both kinds of analysis is discussed. The co-occurrence of morphologically similar individuals differing in the flower color is discussed, too. Possible explanations for this phenomenon involve hybridization in a wide sense or, alternatively, rejecting the assumption of monophyly for the group.     

Reproductive biology and pollination ecology of Triplaris gardneriana (Polygonaceae): a case of ambophily in the Brazilian Chaco

• Triplaris gardneriana (Polygonaceae) is a dioecious pioneer tree reported as insect‐pollinated, despite possessing traits related to anemophily. Here, we analyse the possible roles of insects and wind on the pollination of this species to establish whether the species is ambophilous.
• We carried out observations of floral biology, as well as on the frequency and behaviour of pollinators visiting flowers in a population of T. gardneriana in the Chaco vegetation of Brazil. We conducted experimental pollinations to determine the maternal fertility of female plants and whether they were pollen‐limited, and we also conducted aerobiological experiments to provide evidence of how environmental factors influence atmospheric pollen dispersal.
• The population comprised an area of approximately 152.000 m² and was composed of 603 female and 426 male plants (sex ratio = 0.59:0.41). We observed 48 species of insects visiting flowers of T. gardneriana, of which the bees Scaptotrigona depilis and Apis mellifera scutellata were the most effective pollinators. We recorded pollen grains dispersed by wind on 74% of the glass slides placed on females, located at different distances (1–10 m) from male plants.
• Airborne pollen concentration was negatively correlated with relative humidity and positively correlated with temperature. Our observations and experimental results provide the first evidence that T. gardneriana is an ambophilous species, with pollen dispersal resulting from both animal and wind pollination. This mixed pollination strategy may be adaptive in T. gardneriana providing reproductive assurance during colonisation of sites with different biotic and abiotic conditions.

     

Reproductive biology ofPolyalthia littoralis (Annonaceae)

Flowering phenology of some annonaceous trees and reproductive biology ofPolyalthia littoralis (Annonaceae) were studied. The trees showed various types of flowering phenology within the family. Among them,P. littoralis had hermaphroditic and protogynous flowers, and exhibited continuous flowering throughout the year. Bagged flowers set fruits and seeds comparable to the control. The observations of meiotic stages and the results of castrated tests indicated no possibility of apomictic reproduction. The abscised anthers attached to torus by the extended spiral thickenings, and accomplished self-pollination. Outbreeding possibly occurs, but the following self-pollination guarantees seed set.     

Chromosome numbers of European blackberries (Rubus subg.Rubus,Rosaceae)

Chromosome numbers are presented for 32 collections of 29 European blackberry species (Rubus subg.Rubus) from Germany. One species is triploid (2n = 21), 27 species are tetraploid, (2n = 28), and one species is pentaploid (2n = 35). Chromosome numbers are reported for the first time ofR. adspersus, R. amisiensis, R. calvus, R. conothyrsoides, R. contractipes, R. demissus, R. elegantispinosus, R. ferocior, R. foliosus, R. hypomalacus, R. leucandrus, R. nemorosus, R. platyacanthus, R. praecox, R. rhombifolius, andR. rhytidophyllus. Chromosome numbers forR. dasyphyllus, R. gelertii, R. glandithyrsos, R. lamprocaulos, R. lindebergii, R. macrophyllus, R. montanus, R. muenteri, R. pedemontanus, R. polyanthemus, R. senticosus, R. silvaticus, andR. vigorosus are confirmed.     

Honeyeater (Meliphagidae) pollination and the floral biology of PolynesianHeliconia (Heliconiaceae)

The primary pollinator of Polynesian heliconias,Heliconia laufao andH. paka, is the Wattled Honeyeater,Foulehaio carunculata. This report is the first documentation of pollination by honeyeaters in the genusHeliconia and the first record ofF. carunculata as a pollinator of a plant species. The Polynesian heliconias bear inflorescences that produce 2–4 hermaphroditic flowers per day for a period of 2–3 months. Each flower secretes abundant nectar (125–184 l) with low sugar concentration (15–18% sucrose-equivalents, weight per weight basis) which is available at anthesis just before dawn. Ninety percent of flower visits occur between anthesis and mid-morning. The honeyeaters perch on inflorescence bracts, and probing of the flower results in pollen deposition on the head and bill from where pollen is transferred between flowers. No statements on compatibility can be made forHeliconia paka; however,Heliconia laufao appears to be self-compatible. Calculations of energetic values of nectar of the Polynesian heliconias and Daily Energy Expenditures ofF. carunculata suggest that populations ofH. laufao andH. paka serve as rich energy resources for their pollinators.     

濒危植物金花猕猴桃繁殖生物学初步研究

繁殖生物学是目前濒危植物保护生物学研究的重点领域之一,金花猕猴桃(Actinidia chrysantha)是猕猴桃属濒危物种之一,目前未见其繁殖生物学相关报道。因此,该文以分布于花坪国家级自然保护区的野生金花猕猴桃为研究对象,用游标卡尺测量了花器官及果实形态,通过野外观察记录了其物候、访花昆虫及开花结果习性,用人工授粉和套袋法确定其传粉媒介,开展田间播种试验确定种子繁殖力,对其繁殖生物学开展了较为系统的研究。结果表明：金花猕猴桃物候因海拔高度不同而不同,较低海拔地区5月中下旬开花,高海拔地区5月下旬至6月上旬开花,花期持续7~10 d,果实每年9月下旬至10月上旬成熟;雄株花枝率76.5％,雌株果枝率61.9％,果实长圆柱形、短圆柱形或椭圆形,平均单果重7.34~27.53 g,最大果重35.0 g;金花猕猴桃为虫媒和风媒共同授粉,主要访花昆虫有蜜蜂科、细蜂科、鼻蝇亚科、食蚜蝇科、蜡蝉科、大蚊科长脚蚊属昆虫等;金花猕猴桃种子发芽率低,参试的3个居群的种子发芽率存在差异,分别为花坪17.5％,资源车田15.36％,贺州姑婆山0;4种不同种子处理方式中,低温＋GA3处理的种子发芽率(22.67％)最高。综上所述,金花猕猴桃不存在传粉障碍,种子萌发率低可能是致其濒危的重要原因。该研究结果为保护金花猕猴桃种质资源提供了科学依据。     

Chloroplast DNA diversity in the genusRubus (Rosaceae) revealed by Southern hybridization

The variability in chloroplast DNA type of 20Rubus genotypes was examined by Southern hybridization. DNA extracted from theRubus accessions was digested with two restriction enzymes (EcoRI and EcoRV) and heterologous chloroplast DNA sequences from barley and pea were used as probes to detectRubus chloroplast DNA sequences on Southern blots ofRubus total DNA. Restriction fragment length polymorphism was detected and a total of 92 restriction fragments were generated by the probe/enzyme combinations examined. Cladistic principles based on the parsimony assumption were used to assemble a phylogenetic tree based on chloroplast restriction fragment length data. The phylogenetic tree grouped the taxonomically defined species and is in general agreement with information based on morphological criteria. However, the Japanese red raspberryR. illecebrosus was shown to have diverged considerably in terms of evolutionary time from other species in subg.Idaeobatus. Furthermore, the molecular approach provides a quantitative estimate of the relationship between species that is difficult to obtain from morphological data. In order to complement the chloroplast DNA information a ribosomal DNA probe was also included in the analysis and provided further information on the phylogenetic relationships withinRubus.     

Genetic diversity inChaenomeles (Rosaceae) revealed by RAPD analysis

Genetic relatedness inChaenomeles was studied by RAPD analysis in 42 plants representing accessions of three wild species and one hybrid taxon. Amplification with 17 primers yielded a total of 156 polymorphic RAPD bands. Estimates of genetic relatedness suggest thatC. cathayensis andC. japonica are the most distantly related species, and that the former is comparatively homogeneous.Chaenomeles speciosa, which may have arisen through hybridization betweenC. cathayensis andC. japonica, takes an intermediate position between these two species. Analysis of diagnostic bands demonstrate that neitherC. speciosa norC. ×superba has any bands that do not occur in at least one ofC. cathayensis orC. japonica. Moreover,C. speciosa and the partly overlapping taxonC. ×superba are comparatively heterogeneous, which is also in accordance with a hybrid origin. Intraspecific variation was studied mainly inC. japonica; plants obtained from different sources of material formed well separated groups in the cluster analysis.     

Comparisons of within-population genetic variation in sexual and agamospermousAmelanchier (Rosaceae) using RAPD markers

We compared genetic variation of sexualAmelanchier bartramiana and facultatively agamospermous (asexually seed-producing)A. laevis at one site where the two species are sympatric. We analyzed 77 random amplified polymorphic DNA (RAPD) markers in 29A. bartramiana individuals and 76 RAPD markers in 31A. laevis individuals. The two species do not differ significantly in mean genetic variation. However, 22.4% of genetic similarity values betweenA. laevis individuals exceed the highest value ofA. bartramiana and may represent the effect of agamospermy. Variation withinA. laevis may be the result of sexuality, hybridization, polyploidy, and other factors.     

Reproductive biology of the genusDamasonium (Alismataceae)

The reproductive biology of the genusDamasonium was studied in cultivation and in nature. A correlation was shown between flowering time and latitude. All species are self-compatible but the N. American member of the genus (D. californicum) is strongly protandrous and probably experiences high levels of outcrossing in nature. In the European taxa the stamens and the stigmas mature simultaneously but the levels of inbreeding appear to be greater in the southern species (D. bourgeai, D. polyspermum) than in the northern (tetraploid) speciesD. alisma. The Australian memberD. minus resembles the southern European species in being strongly autogamous and bud-pollination was shown to occur. Floral morphology, stigma exertion, pollen/ovule (P/O) ratio and seed weight each show a strong correlation with the degree of autogamy. Vegetative reproduction occurs only inD. californicum andD. alisma through the production of corm-buds. This study suggests that autogamy inDamasonium is accompanied by a shift in resource allocation from male (pollen) toward female (seed) function. However, further studies are required to assess the quality of seeds and their survivorship in nature.