Reproductive biology of mixed-species populations of Goodyera (orchidaceae) in northern Michigan

The reproductive biology ofGoodyera oblongifolia, G. repens var.ophioides andG. tesselata is discussed with emphasis on the reproductive isolating mechanisms operating in mixed-species populations. Cytology, phenology and pollination of the above three species were studied. Artificial hybridizations were made of these three species and ofG. pubescens. Results show that within mixed-species populations,G. oblongifolia,G. repens var.ophioides andG. tesselata are not completely isolated reproductively, and hybridization does occur. The deleterious effects of the loss of gametes to interspecific crosses are reduced by seasonal isolation, perennial growth and geitonogamous seed production which is encouraged by self-compatibility and clonal growth. Thus, hybridization in mixed-species populations is apparently kept to levels low enough to allow the species to maintain their identities.     

The phytogeography of leaf morphology in two species of North American Asters: Lasallea Novae-Angliae and L. Oblongifolia (Compositae)

Width-to-length ratios of upper stem leaves ofLasallea novaeangliae andL. oblongifolia were determined from herbarium specimens and field collections. No distinct subgroups based on suchratios were found in either species. InL. novaeangliae the curve of the ratio versus number of individuals was skewed toward the narrow-leaf range. This phenomenon was present but less pronounced inL. oblongifolia. The histogram of leaf ratios forL. novaeangliae was somewhat weakly bimodal, suggesting a possible broad-leaf race, which has not been recognized nomenclaturally. The absence of bimodality in the histogram of leaf ratios forL. oblongifolia indicates the need to place Shinners’ narrow-leaved varietyangustatus in synonymy. Other traits such as height, degree of branching and the nature of the pubescence were also found to be of no significance. Slight geographic trends in leaf width occur in both species. No real evidence was found for nomenclaturally recognizing any eastern variant in either species. Thus, Shinners’ var.orientis is not recognized inL. oblongifolia either.     

New chromosome numbers for plant taxa endemic to the Balearic Islands

Mitotic chromosome numbers are reported from 25 vascular plant taxa, endemic to the Balearic Islands that are poorly known cytogenetically. The chromosome numbers ofAnthyllis vulneraria subsp.balearica (2n=12),Cymbalaria fragilis (2n=56), andPolygonum romanum subsp.balearicum (2n=40) were determined for the first time. A new chromosome number was found in several populations ofAnthyllis hystrix (2n=70) suggesting that this species is decaploid, in contrast to an earlier work reporting a higher ploidy level (2n=12x=84). The new chromosome number 2n=32 was reported inHypericum hircinum subsp.cambessedesii. It is suggested that the previous count (2n=40) could be explained by the presence of anomalous pentaploid cells in some tissues, contrating with the presence of a regular tetraploid complement (2n=32). Cytogenetic observations suggest thatSibthorpia africana has a diploid chromosome complement of 2n=18, with 0–2 accessory chromosomes. Accessory chromosomes are also reported forPhlomis italica, being the first record of B chromosomes in this genus. Chromosomal instability was found inGalium crespianum andG. friedichii species, with three numbers 2n=44, 55 and 66. Two cytotypes differing in ploidy level were documented within single plants. It is suggested that both species share a regular complement of 2n=44 and that the past hybridization events and formation of regenerating roots from the typical rootstock ofG. crespianum andG. friedrichii could be involved in the genesis of chromosome variants through partial endopolyploidy and concomitant somatic segregation.     

Biochemical genetic studies on wild populations of three species of green leafhoppers,Nephotettix, from peninsular Malaysia

Nine populations of three species ofNephotettix (Insecta: Hemiptera) from Peninsular Malaysia were analysed for nine enzymes comprising 11 loci. Nei's (Genetics 89, 583, 1978) genetic distance,D, betweenN. virescens andN. malayanus was 0.181, that betweenN. virescens andN. nigropictus was 0.283, and that betweenN. malayanus andN. nigropictus was 0.203. The genetic distance betweenN. nigropictus from rice plant and from the weed-grassL. hexandra at Universiti Pertanian Malaysia was 0.004 and their genetic identity was 0.996, thus indicating that this insect species feeds on both host plants. The proportion of polymorphic loci and the observed heterozygosities were higher inN. nigropictus, with a wider range of host plants, than inN. virescens andN. malayanus, restricted to rice andL. hexandra, respectively.     

Diplachne and leptochloa (Poaceae) in North America

Most North American botanists includeDiplachne inLeptochloa but good evidence for this treatment has never been provided. The diagnostic characters of these two closely related genera of the tribe Eragrosteae are presented and the seven species ofDiplachne and six ofLeptochloa occurring in America north of Mexico are distinguished. Three new combinations are made inDiplachne:D. sect. Ipnum, D. panicoides andD. monticola and the generic disposition of all the Central and North American species noted.     

New or rare data on chromosome numbers in several taxa of the genus<Emphasis Type="Italic">Artemisia (Asteraceae)</Emphasis> in Poland

Chromosome counts in 16 populations of fiveArtemisia species from Poland are presented in this paper. Those ofA. annua (2n=18) andA. dracunculus (2n=90) are reported for the first time in Polish populations. The decaploid level (2n=90) is described for the first time in non-cultivated populations ofA. dracunculus, and several cases of aneusomaty (intraindividual aneuploid variations in chromosome number: 2n=87, 88 and 89) have been detected in this species. In addition to the already known diploid number (2n=18), the tetraploid level (2n=36) has been detected inA. absinthium. The same two numbers have been recorded inA. abrotanum, which represents the first tetraploid count in populations of this taxon occurring outside botanical gardens. Finally, the chromosome number ofArtemisia campestris subsp.sericea (tetraploid, 2n=36) is reported for the first time. The relevance of polyploidy for the evolution of the genus and other cytotaxonomic or cytobiogeographical aspects are briefly discussed.     

Cytogeography of<Emphasis Type="Italic"> Ixeris nakazonei</Emphasis> (Asteraceae,Lactuceae) in the Ryukyu Archipelago of Japan and Taiwan

The cytogeographical structures of Ixeris nakazonei, a putative hybrid between I. debilis (6x) and I. repens (2x), were investigated in the Ryukyu Archipelago and Taiwan. In the Ryukyus, I. debilis occurs on Miyakojima Island of the southern Ryukyus and northward, while I. repens occurs on all islands except for Iriomotejima and Yonagunijima Islands. I. nakazonei, comprises six polyploid cytotypes, 3x, 4x, 5x, 6x, 7x and 8x, based on x=8. Four cytotypes from 3x to 6x occur in the central Ryukyus, while four cytotypes from 5x to 8x occur in the southern Ryukyus. The higher polyploids of I. nakazonei tend to be distributed in the more southerly area. Tetraploids of I. nakazonei always co-occur with I. debilis and I. repens, supporting the hybrid origin of this cytotype. Considering the chromosome number, octoploids, which predominate in the southern Ryukyus and Taiwan, may have derived directly from hybridization between I. debilis and I. repens. Odd-numbered polyploids of I. nakazonei, 3x, 5x and 7x, are relatively rare. Their chromosome numbers indicate that triploids and heptaploids are hybrids between the tetraploid of I. nakazonei and I. repens, and between the octoploid of I. nakazonei and I. debilis, respectively. Pentaploids of I. nakazonei in the central and southern Ryukyus are, respectively, hybrids between the tetraploid of I. nakazonei and I. debilis and between the octoploid of I. nakazonei and I. repens, indicating that pentaploids of I. nakazonei have at least two independent origins.     

A biosystematic study on polyploid populations of the genusSpiraea (Rosaceae) in Korea

A total of 61 chromosome counts from 11 taxa of Korean Spiraea was made. Our counts are the first report for eight taxa;S. blumei (2n=18, 36),S. chartacea (2n=36),S. chinensis (2n=36),S. fritschiana (2n=27, 36),S. microgyna (2n=18),S. prunifolia var.simpliciflora (2n=18),S. pseudocrenata (2n=36), andS. trichocarpa (2n=18). A new chromosome number of 2n=36 (tetraploid) is reported forS. pubescence. Populations of three species includingS. blumei, S. pubescence and .V.fritschiana, show different ploidy levels; diploid and tetraploid populations are found in the former two species and triploid and tetraploid ones, in latter species. Multiplication of chromosome numbers contributes to increase in size of pollen and stomata in the three species. Populations with different ploidy levels inS. blumei occupy different regions; diploid populations in inland Korea and tetraploid ones in Ullung Island. Island tetraploid population of 5.blumei might be originated from intra-island polyploidization through the introduction of diploid from inland Korea, considering the worldwide distribution of this species. Pollen fertilities of island populations of ,S′.blumei are relatively low, and sometimes no pollen grain is produced in anther sacs; it suggests that tetraploid population of the island is gynodioecious which may serve reduction of inbreeding depression.     

High level of genetic variation within clonal orchid Goodyera repens

Type of reproduction has an important effect on the maintenance of particular populations and species persistence in time and space. This trait significantly influences the ecological and genetic structure of populations, and in consequence the evolution of species. The primary objectives of this study were: to estimate genetic diversity within and among populations of clonal species Goodyera repens from different populations in northeastern Poland, and to amount factors shaping the genetic structure of this orchid. Based on 451 rosettes of G. repens from 11 localities in northeastern Poland, we conducted a genetic population analysis using allozymes. We included information on population size, flowering, fruit set and seed dispersal to elucidate their influences on genetic diversity of this species. Populations differed according to demographic properties. The majority of seeds (86.4–94.8 %) were found at a distance of 0.2 m. We observed a high level of genetic (P _PL = 50 %, A = 1.68, H _O = 0.210, H _E = 0.204) and genotypic diversity (G = 163, G/N _S = 0.66, G _U = 30.2 %), and low but statistically significant genetic differentiation among populations (F _ST = 0.060; P < 0.001). We suggest that the genetic diversity of G. repens is mainly an effect of the abundance of pine and spruce forest communities suitable for this species in NE Poland and the high level of sexual reproduction.     

Observations of meiotic chromosomes in Gaura (Onagraceae)

Observations of meiotic chromosomes are reported for all 21 species and 3 additional sub species ofGaura (Onagraceae), based upon a study of 647 individuals from 509 naturally occurring populations throughout the range of the genus. The basic chromosome number for the genus isx = 7, and 18 species are diploid withn = 7. Among these, the self-incompatible ones are often highly chromosomally heterozygous, with no homozygous individuals having been found in nature in the perenrennialsGaura lindheimeri andG. villosa, and two-thirds or more of the individuals apparently heterozygous in the following well-sampled species:G. calcicola, G. longiflora, andG. suffulta subsp.suffulta. In contrast, the autogamous species are entirely chromosomally homozygous or nearly so. Two species ofGaura are reported as chromosomal structural heterozygotes, with about 50% pollen abortion:G. biennis andG. triangulata; the translocation systems originated independently of one another. Two of the three polyploid species,G. sinuata andG. drummondii (G. odorata of many authors), are consistently tetraploid (n = 14) and, despite their cytological autotetraploidy, are thought to have originated following interspecific hybridization. They are the only rhizomatous species in the genus and may have had one ancestor in common. The remaining polyploid,G. coccinea, includes populations withn = 7, 14, 21, and 28, as well as evident interploid hybrids and, frequently, supernumerary chromosomes. The relationship among these populations is close and is maintained by frequent hybridization and exchange of genetic material. No other species seems to have participated in their origin, and the association of their chromosomes is consistently that characteristic of autopolyploidy in plants with tetraploid and higher chromosome numbers.     

Atlas of foliar surface features in woody plants,V. fraxinus (Oleaceae) of eastern North America

Scanning electron microscopy of foliar surfaces of the seven species ofFraxinus native in eastern North America confirms the importance of cuticular patterns (smooth, striated, striate-papillose, and coronulate) as useful taxonomic character states. There are there trichome types (acicular hairs ornamented with micro-striae or-papillae, peltate glands, and scales) which are found in nearly all species and which vary only in density. No differences in surface features are found by which determination can be made of either sex or ploidy level inF. americana. Hybridization betweenF. americana andF. pennsylvanica is considered to be rare in nature and is not a probable cause for the overlap of character states and concomitant difficulty of identification based upon gross morphological features. Likewise, the putative hybrid origins ofF. biltmoreana andF. profunda are not supported by micro-morphological features. Illustrations are given for all species.     

Systematic relationships in the white-rayed species of Melampodium (Compositae)

The white-rayed species complex ofMelampodium has been treated by various authors as containing one, two, or three species. A re-evaluation of the taxa in this complex by use of Chromatographic profiles of methanolic leaf extracts from numerous populations over the range of the group has revealed three distinct patterns which, along with morphological and ecological data, provide support for the recognition of three species within the group. Distributional analysis of previously and newly reported chromosome counts of 157 plants in 99 populations shows tetraploid races ofn = 20 in bothM. leucanthum var.leucanthum andM. cinereum var.cinereum to be located primarily in central and south central Texas but locally intermixed with diploid races. Interpretation of morphology, meiotic configurations, and chemical profiles confirms that these cytotypes are autopolyploid in origin and should not be given formal recognition. All data are used to interpret evolutionary relationships, and it is believed thatM. leucanthum var.argopkyllum may closely resemble the progenitor of the entire complex.     

Restriction fragment length polymorphism of chloroplast DNAs in some species of fir (Abies sp.)

Restriction fragment analysis of the chloroplast gene psbA was carried out in twelveAbies species native to Asia, Europe and North America using the four restriction enzymes. The pair of Asiatic representatives ofA. koreana andA. veitchii differed profoundly not only from each other but also with respect to the European and Northamerican species. The variation observed within the last mentioned groups of firs was mainly due to the different restriction profiles of the gene inA. alba andA. nordmanniana species of the former region as well as owing to its heterogenous nature in all the four species from North America. The RFLP data presented to far only partially correlate with the taxonomic status and hybridological relationships of the species concerned.     

Systematics and gene geography of Juniperus communis L. inferred from isoenzyme data

Using isoenzyme analysis, 35 populations of Juniperus communis L. from various parts of the Russian species range and by one population from Sweden and Alaska were studied. The total sample size was 1200 plants. As a result, the existence of J. communis var. oblonga in North Caucasus and J. communis var. depressa in North America was confirmed, but genetic differences between J. communis var. communis and J. communis var. saxatilis were not detected in the main part of the Russian species range (European part of Russia, Ural, Siberia). These populations proved to be genetically uniform with the same predominant allelic frequencies, which may evidence recent settling of this species from one of Central or East European refugium. J. communis var. saxatilis from northeastern Russia inhabiting the region behind Verkhoyansk mountain and Russian Far East showed considerable differentiation in frequencies of alleles at three loci and geographical subdivision. These populations also exhibit high intrapopulation variation. This can be connected with the refugium in this territory. The origin of this group is probably connected with migrations from Central Asia (Tibet) in the direction to northeastern Russia along mountains connecting Central and North Asia. It is also assumed that migrations of this species previously proceeded across the Beringian land bridge.     

Chromosome numbers,polyploidy, and hybridization in castilleja (scrophulariaceae ) of the Great Basin and Rocky Mountains

Chromosome numbers are reported for 190 collections ofCastilleja of the Great Basin-Rocky Mountains and adjacent regions in western North America. Polyploidy has played a major role in evolu tion of this genus. Of the 35 taxa represented, 14 are exclusively diploid (n =12). The remaining 21 taxa exhibit polyploidy: eight are tetraploid only, two are octopIoid only, while 11 of the more wide-ranging species have two or more levels of polyploidy. BothC. rhexifolia andC. sulphured have diploid, tetraploid, and octoploid races while the widespreadC. miniata has four (2x, 4x, 8x, and 10x). Where appropriate, the role of polyploidy and hybridization inCastilleja evolution and their relevance to taxonomy are discussed for individual species.     

New species and a new combination in Myrtaceae from northeastern South America

Three new species of Myrtaceae (Calyptranthes bracteata, Eugenia gonglycocarpa, andMyrcia rupta) from northeastern South America are described and illustrated, and a new combination (Eugenia tetramera) is proposed. The closed-calyx and the completely or partially fused cotyledons ofMyrcia rupta, unusual features for the genus, are discussed and compared with related species inMyrcia andMarlierea.     

Chromosome numbers in callirhoe (Malvaceae)

Forty-four chromosome counts from the North American genusCallirhoe are reported, including first-recorded counts ofC. alcaeoides, C. digitata, C. bushii, C. triangulata, andC. involucrata var.tenuissima. Information concerning chromosome numbers from present and previous reports is summarized and evaluated. Observations are presented on meiosis for the genus in general, for male sterile individuals of gynodioecious species, and for selected F₁ hybrids. Pollen fertility data are recorded for the latter. Chromosomally three species groups are recognized: an exinvolucellate, diploid species group withn=14; an involucellate, tetraploid and octoploid species pair,n=28 and 56; and an involucellate, diploid and tetraploid species group withn=15 and 30. Intrageneric relationships and those within the tribe Malveae subtribe Malvinae are discussed.     

<Emphasis Type="Italic">Quinua</Emphasis> and Relatives (<Emphasis Type="Italic">Chenopodium</Emphasis> sect.<Emphasis Type="Italic">Chenopodium</Emphasis> subsect.<Emphasis Type="Italic">Celluloid</Emphasis>)

Traditionally viewed as an Andean grain crop,Chenopodium quinoa Willd. includes domesticated populations that are not Andean, and Andean populations that are not domesticated. Comparative analysis of leaf morphology and allozyme frequencies have demonstrated that Andean populations, both domesticated(quinua) and free-living(ajara), represent an exceptionally homogeneous unit that is well differentiated from allied domesticates of coastal Chile(quingua) and freeliving populations of the Argentine lowlands(C. hircinum). This pattern of relationships indicates that Andean populations represent a monophyletic crop/weed system that has possibly developed through cyclic differentiation (natural vs. human selection) and introgressive hybridization. Relative levels of variation suggest that this complex originated in the southern Andes, possibly from wild types allied withC. hircinum, with subsequent dispersal north to Colombia and south to the Chilean coast. Coastal populations were apparently isolated from post-dispersal differentiation and homogenization that occurred in the Andes. Other data point toward a center of origin in the northern Andes with secondary centers of genetic diversity subsequently developing in the southern Andes and the plains of Argentina. Comparative linkage of South American taxa, all tetraploid, with North American tetraploids of the subsection will eventually clarify this problem. While the possibility of a direct phyletic connection betweenC. quinoa and the Mexican domesticate(C. berlandieri subsp. nuttalliae,) cannot be excluded, available evidence indicates that the latter represents an autonomous lineage that is associated with the basal tetraploid, C. b. subsp.berlandieri, through var.sinuatum, whereas South American taxa show possible affinities to either var. zschackei or var.berlandieri. An extinct domesticate of eastern North America,C. b. subsp.jonesianum, represents either another instance of independent domestication, possibly from subsp. b. var.zschackei, or a northeastern outlier of subsp.nuttalliae.     

Natural hybridization in JapaneseCalamagrostis

Relationships amongCalamagrostis longiseta var.longiseta, C.l. var.longe-aristata andC. fauriei, which are principally at the 2n=28 chromosome level, were studied mainly on the basis of population samples. The populations comprised (1) “pure” populations of each ofC.l. longiseta, C.l. longe-aristata andC. fauriei and (2) populations showing a gradation fromC.l. longiseta toC. fauriei via the intermediateC.l. longearistata and fromC.l. longiseta toC.l. longe-aristata. Hybridization betweenC.l. longiseta andC. fauriei was estimated to be ultimately related to the geneses of the population structures detected, and it was concluded thatC.l. longe-aristata must be of hybrid origin betweenC.l. longiseta andC. fauriei. Populations showing a gradation fromC.l. longiseta toC.l. longe-aristata were regarded to have been brought about by the extinction ofC. fauriei from the hybridizing population coupled with the change of environments and also by the secondary contact betweenC.l. longiseta and plants ofC.l. longe-aristata migrating from the area that produced it. No significant reduction of pollen fertility and seed productivity was found inC.l. longe-aristata. Based on the results of population analyses, the delimitation of the taxa concerned was made clearer. The examination of herbarium specimens showed that the range ofC.l. longe-aristata considerably exceeds the overlapping area of the present ranges ofC.l. longiseta andC. fauriei. The three taxa were deviated ecologically, and “pure” populations ofC.l. longe-aristata were found on several damp subalpine fields in central and northern Honshu that seemed to represent the most suitable niche for this taxon. It was estimated thatC.l. longe-aristata would evolve into a new good species if the future organism-environment interactions are favorable for it.     

The taxonomic status of physalis lanceolata (Solanaceae) in the Carolina Sandhills

In the most recent monograph of the genus in North America,Physalis lanceolata was considered to be a hybrid betweenP. heterophylla andP. virginiana. Comparative morphological and Chromatographic studies indicate, however, thatP. lanceolata is not intermediate between its supposed parents. This evidence plus its high pollen viability and seed set indicate that the species is not a hybrid. Natural hybrids betweenP. heterophylla andP. virginiana are sterile and are intermediate between these species in morphology and Chromatographic pattern.