Systematics ofPrimula veris (Primulaceae)

Primarily, 50 characters taken from herbarium specimens, cultivated plants, and whole populations ofPrimula veris L. were critically evaluated. Only 24 characters were found to be suitable for taxonomic use. The geographical and multivariate analysis of about 1000 herbarium specimens from the whole area lead to a reduction of the number of subspecies. Only a less restricted subsp.veris (incl. subsp.canescens, subsp.inflata and probably including subsp.macrocalyx) and subsp.columnae (incl. subsp.suaveolens) can be distinguished considering the variability of characters within an individuum, under different culture conditions, and during the growth period. The status of the taxon macrocalyx cannot be cleared presently.     

Pollination ofNuphar (Nymphaeaceae) in Europe: Flies and bees rather thanDonacia beetles

Nuphar (Nymphaeaceae) comprises a small holarctic group of aquatic perennials whose flowers are pollinated by flies, bees, and beetles. We studied pollination in different populations of the two European speciesN. lutea andN. pumila in Norway and in Germany. Flowers are self-compatible and protogynous, preventing automatic selfing, and insect pollination is required for seed production. Sympatric populations were studied in Vest-Agder county in Norway to determine whetherN. lutea andN. pumila have the same or different pollinators. Allopatric populations ofN. lutea in Germany and Norway were then compared to determine whether their pollinator spectra differ as would be expected in an open flower with seemingly easily accessible pollen and nectar. Results of the present as well as previous studies of the pollination ofN. lutea andN. pumila show that both species are mainly pollinated by flies, including apparentNuphar specialists, such as the scatophagidHydromyza livens and the ephydridsHydrellia andNotiphila, the last also a long-known pollinator ofN. advena in Florida. Pollinator overlap between sympatric heterospecific populations was small, while allopatric conspecific populations had similar visitor and pollinator spectra. We found no evidence of pollination byDonacia beetles as reported from some North American populations ofNuphar. Dedicated to Univ.-Prof. DrF. Ehrendorfer on the occasion of his 70th birthday     

Daphnia Galeata × D. Longispina Hybrids in Western Norway

We describe the occurrence of D. galeata×longispina hybrids in two lakes of western Norway. Parental species and interspecific hybrids were characterised by both nuclear and mitochondrial molecular markers. In one of the populations, hybrids were shown to dominate the population over several years. A few individuals in both populations were probably not F1 hybrids, but possibly backcrosses or F2 hybrids. Most (possibly all) F1 hybrids were of D. galeata maternal origin. In addition, interspecific hybrids could be identified based on morphological characters, which were intermediate between the parental species. Interspecific hybridisation between these two species is remarkable, since they are distantly related.     

Estimates of hierarchical variation in flower morphology in natural populations ofScleranthus annuus (Caryophyllaceae), an inbreeding annual

The variation in two sets of morphological characters of the flowers of the highly inbreedingScleranthus annuus (Caryophyllaceae) was assessed using 15–20 plants from each of 20 natural populations from the southernmost region of Sweden. The stamen fertility data set consisted of 10 characters describing the degree of fertility of the ten stamens/staminoids, while the sepal/gynoecium data set comprised 10 sepal characters, together with style and stigma length. Substantial variation was found in both the degree of development and the fertility of the stamens, the degree of variability in stamen fertility being related to stamen position within the flower. Considerable variation was found in the characters of the sepal and the gynoecium. Hierarchical analyses of variance indicated that 29% of the variation in total male reproductive effort was distributed among populations, 28% among plants within populations and 43% represented within-plant variation. The corresponding averages for the characters from the sepal/gynoecium data set are 26, 38, and 35%: a greater proportion of the total variance in female reproductive characters is accounted for by among individual variation than is the case with the male reproductive characters. Significance tests of Mahalanobis distances derived by canonical variate analyses indicated that all populations were significantly separated using the sepal/gynoecium data set, while only 50% of the pairwise comparisons on the basis of the stamen fertility data set were significant. Cluster analysis did not reveal any aggregation of the populations. The incongruence of the two data sets and their ability to discriminate between the populations is discussed.     

Genetic variation of ticks (Ixodes ricinus L.) in the Lithuanian and Norwegian populations

RAPD markers were used to measure the genetic diversity of 119 individuals of Ixodes ricinus collected from Lithuania and Norway. The samples were analysed within and also between the populations. We analysed 74 loci in each of 6 populations. Our results show high levels of diversity within the populations. The percentage of polymorphic loci of the six analysed populations: Birzai, Vilnius, Kretinga, Tjore, Kjosvik and Odderoya were 68.9%, 58.1%, 78.38%, 62.2%, 44.6% and 68.9%, respectively. The percentage of polymorphic loci in the Lithuanian populations was 93.2%, and in the Norwegian populations 81.08%. The genetic distance ranged from 0.019 to 0.079 within Norwegian populations and from 0.005 to 0.0967 within Lithuanian populations and between the countries from 0.022 to 0.146. The genetic variation of I. ricinus among Norwegian populations was lower than among Lithuanian populations. The highest part of genetic variation in I. ricinus ticks depends on variation within Kretinga (Lithuania) and Odderoya (Norway) populations situated in coastal areas where many migratory and sea birds are aggregated.     

Global Spread of the Eel Parasite Gyrodactylus Anguillae (Monogenea)

Gyrodactylus anguillae is known from four continents, and intraspecific variation in the morphology of specimens from each continent is negligible. However, morphological characters alone may not always be adequate for recognizing populations that have been isolated geographically over evolutionary time. We were unable to obtain specimens from Northeast Asia, but G. anguillae collected from North America, Europe and Australia had identical ribosomal DNA sequences (ITS1-5.8SrRNA-ITS2 region). This lack of even minor genetic variation in populations from three remote continents leads us to conclude that they share a contemporary common ancestor. This is one of at least five pests of eels to have invaded new continents as a result of the eel trade in the last four decades; these other known eel pests include Pseudodactylogyrus spp. (Monogenea) and Anguillicola spp. (Nematoda). Their dispersal has probably been secondarily augmented by longshore migration of infected eels, and perhaps also by transportation in ballast waters. The only other known gyrodactylid from eel gills, G. nipponensis from Japanese eel, is not closely related to G. anguillae, because the ITS1 region of the former is short (370 bases) and that of the latter long (685 bases). G. anguillae probably originated in Europe, because it was recorded there over 40 years ago, before the development of the international eel trade. We predict that the worm has a high potential to invade eel populations in East Africa, Southeast Asia, and New Zealand.     

A morphologically variant natural population of cyprinids without dorsal fin possibly derived from Mystacoleucus marginatus (Osteichthyes: Teleostes) from the Lancang Jiang River, Yunnan, China

In July 2001, 15 grotesque cyprinid specimens were collected in the Lancangjiang River (of the upper reaches of the Mekong River) in Menglun Town, Xishuanbanna Prefecture, Yunnan Province, PR China. These specimens are characterized by surprising characters that have not been observed in natural populations of cyprinids previously, such as an absence of a dorsal fin and of some supraneurals. Other characters, such as the number of branched rays of the anal fin, the formula of pharyngeal teeth, the condition of barbels, and the number of scales are not stable and vary greatly. The more stable characters of these variants, such as the formula of pharyngeal teeth, the pattern of striations on the scales, the number of vertebrae, the length of the gut, etc., have many similarities with those of Mystacoleucus marginatus (Cuvier & Valenciennes, 1842). These observations indicate that these variants may have originated from M. marginatus.     

Epicuticular wax morphology and the taxonomy ofRosa (sectionCaninae, subsectionRubiginosae)

Classification within the genusRosa is problematic due to ± continuous variation of morphological characters. The dogroses, (Rosa, sectionCaninae) are classified only by their unique meiotic system not by directly visible characters. The intrasectional structure remains insufficient, as the characters (glands, hairs and prickles) are known to exhibit great morphological divergence affected by both genetic plasticity and environmental influences. Taxonomy of dogroses follows a highly artificial system which allows us to identify morphospecies, which are not necessarily evolutionary species. However, SEM-studies in theCaninae revealed that all taxa of the subsectionRubiginosae are characterised by a granule type of epicuticular waxes, whereas members from the other subsections (with three exceptions) form triangular rodlets. Comparison with specimens grown under the same conditions confirmed these findings on natural populations and additionally revealed that following artificial hybridism, offspring develop the maternal type of wax structure due to the matroclinal inheritance in this section.     

Quantitative morphological variation inTetramolopium (Asteraceae) in Hawaii and the Cook Islands

Multivariate analyses of quantitative characters were undertaken to examine the pattern of variation resulting from adaptive radiation ofTetramolopium in Hawaii and to investigate the systematics of an undescribed species in the Cook Islands and a group of populations of uncertain affinity on Kalaupapa Peninsula in Hawaii. Taxa differentiated on quantitative characters in the absence of qualitative characters. Differentiation was observed in multiple characters which could be organized into a few principal components. The analyses supported the species rank of the Cook Islands plants. The relationship of the Kalaupapa Peninsula populations to the rest of the taxa needs further study.     

New species in the rheophilous genus Nicsmirnovius Chiambeng & Dumont, 1999 (Branchiopoda: Anomopoda: Chydoridae) and reassignment of Alona eximia Kiser, 1948 and Alonella fitzpatricki Chien, 1970

A morphological comparison of specimens previously assigned to Alona eximia Kiser, 1948 from tropical Africa, Eastern Asia and the Americas shows that this species-group shares a number of morphological characters on the postabdomen, head pores, first antenna and second and fourth limb that separate them from Alona Baird, 1843 but unite them with Nicsmirnovius Chiambeng & Dumont, 1999. Alonella fitzpatricki Chien, 1970, formerly believed to be a junior synonym of A. eximia, is separated from the latter and assigned to the genus Nicsmirnovius. Two new taxa, from Africa and the Island of Socotra (Yemen) are added to the genus. The relationship between the specialised habitat of these chydorids and their morphology is discussed. The geographic range of all known populations is figured and a key to species is presented.     

Biosystematic study ofSedum L. SubgenusAizoon (Crassulaceae)

Variation in chromosome number and morphological characters ofSedum aizoon L. var.floribundum Nakai were investigated to analyze, correlations between them. Geographical variation in chromosome number was also examined. Chromosome numbers of 189 individuals from 55 localities were counted as 2n=32, 33, 34, 48, 61, 64, 78, 80, 84, 85, 88, 93, 94, 95, 96, 97 and 102. InSedum subgenusAizoon, which has the basic number of X=8, var.floribundum of the speciesaizoon showed a polyploid series from tetraploid to dodecaploid. Tetraploids were found most frequently in this variety. More than two chromosome numbers were found in all the populations with 2n=61 or more. Tetraploids were mainly distributed in the eastern part of Japan, and higher polyploids (higher than hexaploid level except octaploids) were distributed in the western part of Japan. Thirteen morphological characters were examined in 119 individuals belonging to 30 populations. The principal component analysis and the cluster analysis of these characters indicated that populations with similar chromosome numbers were not always morphologically similar. Ten morphotypes can be recognized based on the combination of chromosome number and morphological characters. ThusS. aizoon var.floribundum can be regarded as a polyploid-aneuploid complex.     

Phenotypic plasticity inCrepis tectorum (Asteraceae)

Two experiments were carried out using two different approaches to compare populations ofCrepis tectorum (Asteraceae). One was based on a comparison of means of various vegetative and reproductive characters and another was based on a comparison of response patterns of the same characters in a series of environments. Population divergence within two earlier recognized form series, one from weed habitats and one from alvar habitats on Baltic islands, resulted in a partially overlapping pattern in cluster analyses based on character means. However, the pattern revealed by a comparison of the direction and amount of plastic response suggested that populations within the two form series had more similar response patterns than other combinations of populations. It was concluded that patterns of plasticity may provide useful additional information on the overall similarity among taxa. An hypothesis that plants in weed populations should exhibit a greater phenotypic response to the environments than plants in alvar populations was rejected.     

Molecular and morphological data suggest that Spinibarbus caldwelli (Nichols) (Teleostei: Cyprinidae) is a valid species

The taxonomic problem of the cyprinid species of genus Spinibarbus, occurring in southern China and northern Vietnam, was resolved on the basis of molecular and morphological analyses. Spinibarbus caldwelli and Spinibarbus hollandi have a smooth posterior edge of the last unbranched dorsal fin ray among species in the genus. Spinibarbus caldwelli is currently regarded as a junior synonym of S. hollandi because of ambiguities in diagnostic characters. In this article, 11 mtDNA cytochrome b sequences of Spinibarbus specimens were analyzed together with Barbodes gonionotus and Puntius conchonius as outgroups. Our results showed that specimens identified as S. hollandi from Taiwan were different from those from the Asian mainland at a high level of genetic divergence (0.097–0.112), which is higher than that between the two valid species, S. sinensis and S. yunnanensis (0.089), and suggested that Taiwan specimens should be considered as a different species from the Asian mainland one. In a molecular phylogenetic analysis, the sister-group relationship between Taiwan specimens and the Asian mainland specimens was supported strongly by a high confidence level (100% in bootstrap value). Further analysis of morphological characters showed that overlap of diagnostic characters is much weaker than previously suggested. Taiwan specimens had 8 branched rays in the dorsal fin, whereas those from the mainland had almost 9–10. The molecular and morphological differences suggest S. caldwelli to be valid. The molecular divergence shows the genetic speciation of S. hollandi and S. caldwelli might have occurred 5.6–4.9 million years ago; the former could be a relict species in Taiwan, and the latter dispersed in the Asian mainland.     

Morphometrics and character state recognition for cladistic analyses in theBletia reflexa complex (Orchidaceae)

Different criteria have been suggested for decisions on character state recognition for morphological characters showing continuous variation from taxon to taxon. A method proposed byMishler andDe Luna seeks to recognize as many character states as the groups of taxon means that are revealed by an analysis of variance (ANOVA) followed by multiple range tests. We apply this method as a guide in decisions of selecting taxonomic characters in species of theBletia reflexa complex. Seven species of the complex, represented by 156 individuals belonging to 54 populations, were the terminal taxa of the study. Character state codes were assigned to 20 characters. These characters together with 12 non-quantitative characters were incorporated into cladistic analyses. Results indicate that the complex is monophyletic only under one outgroup combination.     

Chlorophyll-deficient gene and morphological variations in Korean populations of maize (Zea mays)

We studied cultivated and naturalized Korean maize populations to determine the extent to which the chlorophylldeficient mutation and the phenotypic variations of two morphological characters (i.e., red coleoptiles and epicotyls, and the number of the first root hairs) are maintained. The frequency of the chlorophyll-deficient mutant gene (2.73% on average) was highly variable. Frequencies of red coleoptiles and epicotyls also were higher than expected from a mutation-selection balance. The average number of hairy phenotypes within populations was 1.8, ranging from 0.0 to 4.0. Naturalized populations were closely related to with cultivated communities. Most striking, however, was the more significant difference among populations than within populations with regard to both the frequency of chlorophyll-deficient mutant genes and the phenotypic variations of our two morphological characters. On a per-gene basis, the majority of the phenotypic variation (mean of 73.3%) resided among populations.     

Genetic insight into Mediterranean chukar (Alectoris chukar, Galliformes) populations inferred from mitochondrial DNA and RAPD markers

The chukar (Alectoris chukar, Galliformes) is one of the most important game birds as it is widely distributed and hunted over the whole of its range. The aim of this work was to assess the genetic differentiation as well as the possible presence of hybrid specimens in A. chukar populations from Italy, Greece and Cyprus. To provide phylogenetic context, conspecific, allopatric specimens from Israel, Georgia, Armenia, Kazakhstan, Afghanistan, Pakistan, Mongolia, China and USA were compared. Sequencing of the mitochondrial DNA (mtDNA) Control Region supplied information on the ancestry of A. chukar populations, whereas Random Amplified Polymorphic DNA (RAPD) fingerprinting was used to assess whether hybridization had occurred. The Italian population was found to be an inter-specific mixture of A. chukar and A. rufa (i.e., the red-legged partridge) mtDNA lineages, whereas the representatives from Greece and Cyprus showed only the A. chukar maternal line. RAPD markers revealed introgression with A. rufa genes in the Italian population, whereas no A. chukar × A. rufa hybrid specimens were detected in the eastern Mediterranean populations. The genetic data obtained from the Italian A. chukar population as well as from a few Greek specimens pointed against their Mediterranean kinship, suggesting relationships with A. chukar subspecies from the easternmost part of the Asian continent.     

Identification of immature tubificids (Oligochaeta) of Lake Constance and its influence on the evaluation of species distribution

In the course of an extensive taxonomical treatment of Oligochaetes collected from Lake Constance the immature specimens of all species were identified, mainly by setal characters. The method is presented, several hints are given and some characters of additional taxonomic value are described in a comparative way.The large extent to which evaluation of species distribution may be misleading if based only on the mature specimens is demonstrated.     

Morphometric variation inLomelosia crenata (Dipsacaceae)

Morphometric characters on 118 herbarium specimens ofLomelosia crenata subsp.crenata (regarded as including subsp.pseudisetensis) were statistically evaluated to detect discontinuities in the variation pattern. Three morphotypes were detected: specimens with short stems and long calyx bristles, specimens with short stems and short bristles, and specimens with long stems and short bristles. The first morphotypic group is separated from the others, which are partly overlapping. These groups broadly correspond toL. crenata subsp.crenata, toScabiosa crenata var.glabriuscula, and toL. crenata subsp.pseudisetensis, respectively.     

Floral development and systematic position of <Emphasis Type="Italic">Arillastrum</Emphasis>, <Emphasis Type="Italic">Allosyncarpia</Emphasis>, <Emphasis Type="Italic">Stockwellia</Emphasis> and <Emphasis Type="Italic">Eucalyptopsis</Emphasis> (Myrtaceae)

We have investigated the floral ontogeny of Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis (of the eucalypt group, Myrtaceae) using scanning electron microscopy and light microscopy. Several critical characters for establishing relationships between these genera and to the eucalypts have been determined. The absence of compound petaline primordia in Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis excludes these taxa from the eucalypt clade. Post-anthesis circumscissile abscission of the hypanthium above the ovary in Stockwellia, Eucalyptopsis and Allosyncarpia is evidence that these three taxa form a monophyletic group; undifferentiated perianth parts and elongated fusiform buds are characters that unite Stockwellia and Eucalyptopsis as sister taxa. No floral characters clearly associate Arillastrum with either the eucalypt clade or the clade of Stockwellia, Eucalyptopsis and Allosyncarpia.We gratefully acknowledge Clyde Dunlop and Bob Harwood (Northern Territory Herbarium) for collecting specimens of Allosyncarpia, and Bruce Gray (Atherton) for collecting specimens of Stockwellia. The Australian National Herbarium (CANB) kindly lent herbarium specimens of Eucalyptopsis for examination. This research was supported by a University of Melbourne Research Development Grant to Andrew Drinnan.