Floral development and evolution of capitulum structure in Anacyclus (Anthemideae,Asteraceae)

Background and Aims

Most of the diversity in the pseudanthia of Asteraceae is based on the differential symmetry and sexuality of its flowers. In Anacyclus, where there are (1) homogamous capitula, with bisexual, mainly actinomorphic and pentamerous flowers; and (2) heterogamous capitula, with peripheral zygomorphic, trimerous and long-/short-rayed female flowers, the floral ontogeny was investigated to infer their origin.

Methods

Floral morphology and ontogeny were studied using scanning electron microscope and light microscope techniques

Key Results

Disc flowers, subtended by paleae, initiate acropetally. Perianth and androecium initiation is unidirectional/simultaneous. Late zygomorphy occurs by enlargement of the adaxial perianth lobes. In contrast, ray flowers, subtended by involucral bracts, initiate after the proximal disc buds, breaking the inflorescence acropetal pattern. Early zygomorphy is manifested through the fusion of the lateral and abaxial perianth lobes and the arrest of the adaxials. We report atypical phenotypes with peripheral ‘trumpet’ flowers from natural populations. The peripheral ‘trumpet’ buds initiate after disc flowers, but maintain an actinomorphic perianth. All phenotypes are compared and interpreted in the context of alternative scenarios for the origin of the capitulum and the perianth identity.

Conclusions

Homogamous inflorescences display a uniform floral morphology and development, whereas the peripheral buds in heterogamous capitula display remarkable plasticity. Disc and ray flowers follow different floral developmental pathways. Peripheral zygomorphic flowers initiate after the proximal actinomorphic disc flowers, behaving as lateral independent units of the pseudanthial disc from inception. The perianth and the androecium are the most variable whorls across the different types of flowers, but their changes are not correlated. Lack of homology between hypanthial appendages and a calyx, and the perianth double-sided structure are discussed for Anacyclus together with potential causes of its ray flower plasticity.     

Generic interrelationships of theCardueae (Compositae): A cladistic analysis of morphological data

A cladistic analysis of 45 genera ofCardueae for 75 characters is presented, taking into account characters of bracts, corolla, stamens, styles and cypselas. Forty-six binary characters completeBremer's (1994) work. Some of the characters considered byBremer are criticised. The traditionalCarlininae is a paraphyletic group, the ten genera studied sharing no common derived character. A new classification of theCardueae is proposed and the distribution of the tribe is presented. Several new combinations are proposed. The nomenclature ofCentaurea is discussed and the status of the genusSerratula is analysed.     

Phylogenetic studies in the polyploid complex of the genus Leucanthemum Mill. (Compositae, Anthemideae) based on cpDNA sequence variation

To reconstruct the evolutionary history of the polyploid southern and central European genus Leucanthemum, comprising 41 species with ploidy levels ranging from 2x to 22x, we analysed chloroplast DNA sequence variation (psbA-trnH and trnL-trnF intergenic spacer regions) of 106 representatives belonging to 30 species and 41 taxa. In an unrooted haplotype network, which shows internal (ancestral) haplotypes that are mainly represented by diploid taxa (L. gallaecicum, L. gracilicaule, L. halleri, L. laciniatum, L. lithopolitanicum, L. rotundifolium, and L. tridactylites), we identified three major haplotype groups (HTGs) containing diploid and polyploid species. Whereas HTG?I contains most of the polyploid taxa of the genus, with a single diploid species from the SW Alps (L. virgatum), HTG?II consists of four diploid (L. burnatii, L. gaudinii, L. graminifolium, and L. vulgare) and six polyploid species, and HTG?III comprises one diploid (L. pluriflorum) and three polyploid species endemic to the NW part of the Iberian Peninsula. We also further found evidence for recurrent formation of at least three polyploid taxa (i.e., L. delabrei, L. ircutianum subsp. cantabricum, and L. pallens).     

Karyological studies in Tripleurospermum (Asteraceae, Anthemideae) from north-east Anatolia

Chromosome number and morphology in 14 taxa belonging to 19 populations of Tripleurospermum Sch. Bip. were studied using karyological and numerical taxonomical techniques. Data on chromosome measurements were analysed using cluster analysis. Chromosome numbers of these taxa are 2 n  = 2 x  = 18, 4 x  = 36 and 5 x  = 42–48. Seven records are new, two are not consistent with previous counts, and the remainder confirm the very limited previous data (one to three records). A new ploidy level (pentaploidy) is reported for the first time for the genus. Some correlations between ploidy levels and morphological characters are noted and several systematic and evolutionary aspects of the genus are discussed in the light of karyological data.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 146 , 427–438.     

Cytogenetic studies in Pentatomidae (Heteroptera): A review

The suborder Heteroptera constitutes one of the most important insect groups because most species are plants feeders and cause damage on many plants of economic importance. One of the most important cytogenetic characteristics of Heteroptera is the holokinetic nature of the chromosomes. One particular feature of some species of Pentatomidae is the regular presence of an abnormal meiosis in one testicular lobe (harlequin lobe). From the 28 species cytogenetically analysed from Argentine material, 21 present the diploid number 2 n  = 14, four species present a reduced number (2 n  = 12) and another three species possess an increased diploid number (2 n  = 16); among all these only three present an harlequin lobe. In the present work, a bibliographic review of the chromosome number and sex determining system of 294 species and subspecies belonging to 121 genera within the subfamilies Asopinae, Discocephalinae, Edessinae, Pentatominae, Phyllocephalinae and Podopinae is presented. The male diploid numbers range from six to 27 with a mode in 14 chromosomes; this last diploid number is present in 85% of the species. The sex chromosome determining system is XY/XX except in three species: Macropygium reticulare (Fabricius, 1803), Rhytidolomia senilis (Say, 1832) and Thyanta calceata (Say, 1832) which present derived sex chromosome systems. Furthermore, the cytogenetic relationships with the other families of Pentatomoidea are discussed.     

Cytogenetic studies of Glomeris (Diplopoda: Glomeridae)

Karyotypes and meiosis of Glomeris hexasticha and G. connexa (Diplopoda: Glomeridae) from Poland were described using C-heterochromatin distribution and observations of the location of NORs. These species were characterized by 2n>=16 and the XY sex determination system. Differences were found in the amount of C-heterochromatin in X and Y chromosomes between the studied species. In G. hexasticha, supernumerary B chromosomes were described.     

Phylogeny, biogeography, and evolution of sex expression in the southern hemisphere genus Leptinella (Compositae, Anthemideae)

Leptinella is exceptional in the Anthemideae (Compositae) in its evolution of dimorphic sex expression. A molecular phylogeny including 40 of its 42 described taxa based on nucleotide sequences from two plastid regions (psbA-trnH and trnC-petN spacers) and one nuclear marker (nrDNA ITS) is presented. Phylogenetic reconstruction was hampered by inadequate phylogenetic signal indicating recent radiation of species during the last 5Ma and high level of reticulate evolution presumably caused by hybridisation and polyploidisation. Nevertheless, Leptinella is nested within a paraphyletic genus Cotula that also engulfs the South American genus Soliva. Within Leptinella, the highly polyploid and sexually polymorphic subgenus Leptinella is monophyletic, while subgenus Oligoleima as well as subgenus Radiata are polyphyletic. We found a basal split between a lineage of Australian and New Guinean taxa and one of largely New Zealand taxa. At least five long-distance dispersal events have to be assumed in order to explain the distribution pattern in Leptinella. Among those, one is from New Zealand to Australia, while the others are dispersals to South America and to several subantarctic islands. The phylogeny presented here indicates that the ancestral sex expression in Leptinella is monoecy and that dioecy and paradioecy are derived conditions. High ploidy is especially common in the dioica-group, where dioecy is also common. However, the occurrence of a dioecious sex expression in tetraploid representatives of this group and of polyploidy in other clades that only exhibit monoecious or paradioecious conditions indicate that there is no consistent correlation between these two characters.     

Cytogenetic studies of Poecilia (Pisces)

Natural populations of triploid females resembling the gynogenetic teleost, Poecilia formosa (Girard), occur in northeastern Mexico where they intermingle with diploid populations of this species and the members of congeneric bisexual species such as P. mexicana or P. latipinna. Mitotic configurations from gill epithelial cells show 46 chromosomes for the diploid fishes, but 69 chromosomes for members of the triploid clones associated with P. formosa. Triploid females have erythrocytes that are significantly larger than those from diploid specimens and also show a roughly 50% elevation in the average DNA content of their somatic nuclei. Similar analyses of two functionally incompetent males of P. formosa, of a number of bisexual F₁ and F₂ hybrid offpsring from P. latipinna x P. mexicana, and of females from several other poeciliid species consistently show only diploid DNA levels and somatic chromosome complements where 22N=46. Demonstration of cytogenetic criteria by which females from triploid clones may be clearly distinguished from sympatric diploid specimens of P. formosa or P. mexicana leaves unresolved, for the present, problems of an appropriate systematic designation for natural populations of triploid gynogenetic fishes. The role of sympatric speciation in the evolution of poeciliid genomes is discussed in terms of alternative mechanisms to account for the persistence in nature of a vertebrate triploid of hybrid origin.This work was supported by grants from the National Science Foundation (GB 7393) and from the U.S. Public Health Service (GM 14644).Recipient of a Research Career Development Award from the U.S. Public Health Service (1 K3 GM 3455).     

Evolutionary history, biogeography and eco-climatological differentiation of the genus Anthemis L. (Compositae, Anthemideae) in the circum-Mediterranean area

Aim To reconstruct the temporal, geographical and eco‐climatological differentiation of the genus Anthemis (Compositae, Anthemideae) in the circum‐Mediterranean region, in order to evaluate the relative importance of geographical vs. climatological differentiation processes in influencing the actual distribution patterns in this plant group. Location The circum‐Mediterranean region, including the Iberian Peninsula, northern Africa, the Italian and Balkan peninsulas, the Aegean region and Anatolia, the Caucasus, the Arabian Peninsula and western Asia. Methods The phylogeny of the genus Anthemis was obtained from a maximum likelihood analysis based on nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) sequence data, and the chronology of diversification was derived using a penalized likelihood approach. The reconstruction of the spatial diversification of the genus was based on a dispersal/vicariance (DIVA) analysis. Eco‐climatological niche differentiation was inferred by optimizing 19 bioclimatic variables onto the phylogeny. A multi‐dimensional hypervolume, proposed as a representation of the eco‐climatological niche and defined by the combination of ranges for all bioclimatic variables, was calculated for each taxon and each internal node. To identify ‘eco‐climatological vicariance’ events in the phylogeny, the pairwise overlap among hypervolumes of sister groups was calculated. Finally, the temporal and clade‐wise relative importance of geographical vs. eco‐climatological vicariance events was estimated. Results The temporal reconstruction shows a constant increase of lineages through the last 12 Myr. The geographical reconstruction suggests that Anthemis diverged from the rest of the Compositae–Anthemideae in the eastern Mediterranean region, and from there radiated into the whole circum‐Mediterranean region through successive dispersal and vicariance events. The reconstruction of the eco‐climatological niches suggests a progressive adaptation from a montane‐humid climate towards arid environments and the typical mediterranean climate. Main conclusions The results presented here involved phylogenetic, geographical and eco‐climatological reconstructions; joint analyses of all of these aspects have assessed the relative importance of geological vs. climatic forces that have affected the distributional history of the genus Anthemis. Large‐scale differentiation patterns triggered by geological forces appear to have influenced the evolutionary history of the genus in a rather constant manner over the last 12 Myr, whereas climatic forces seem to have played an important role in two phases of the radiation process: at around 9 Ma, when the area experienced the onset of a trend towards aridification, and during the last 3.5 Myr, with the establishment of the typical mediterranean climate and the influence of Pleistocene climate oscillations.     

Cytogenetic studies of cultivatedCrocus sativus (Iridaceae)

Meiosis and mitosis are described in cultivatedCrocus sativus of Iran. This indicates that this species is an autotriploid and sterile. Karyotype analysis, rare inversions, laggard chromosomes and distribution of chromosomes in the first anaphase are described, and the reasons for its sterility are given.     

Karyological studies in someHypochoeris spp. (Compositae) from Sicily

FiveHypochoeris spp. from Sicily have been investigated:H. glabra L. (2n=10),H. radicata L. (2n=8),H. cretensis L. (2n=6),H. laevigata L. (2n=12),H. robertia Fiori (2n=8). Basic chromosome numbers are very variable, x = 3, 4, 5, 6. The karyotype of each species is presented. Geographical origin (S. America or Mediterranean region) of the genusHypochoeris and the taxonomic position ofH. robertia are discussed.     

Chromosome studies: Mexican Compositae

Chromosome observations reported here from 70 collections representing 65 species from 40 genera of Compositae (mostly Heliantheae) provide taxonomically useful information. First counts for 28 species, including the first count for Psacaliopsis, may prove to be especially interesting in relation to phyletic and taxonomic alignments of the taxa. Of special interest among first reports is 2n = 5 II for Stevia lita; other stevias have x = 11, 12, or 17. Thirteen of our counts differ from prior reports and may also be of particular interest in phyletic and taxonomic contexts.     

AFLP fingerprinting suggests an allopolyploid origin of two members of the Leucanthemum vulgare aggregate (Compositae,Anthemideae) in central Europe

In order to clarify the origin of the wide‐spread tetraploid Leucanthemum ircutianum and the hexaploid L. adustum, both found in the alpine and dealpine parts of central Europe together with the diploid L. vulgare, we carried out a genetic analysis based on the distribution of amplified fragment length polymorphism (AFLP) fragments in populations of these three taxa, both in the northwestern Alps (Allgäu region, south Germany and adjacent parts of Austria) and the Franconian Jura (vicinity of Regensburg, Bavaria, Germany). Statistical analyses based on AFLP fragments (principal co‐ordinate analysis, PCoA; analysis of molecular variance, AMOVA; Bayesian analysis of population structure, BAPS) suggest that neither of the polyploids is of autopolyploid origin based on L. vulgare, but rather are allopolyploids with the second (maternal) diploid parent still unknown.