Evolution and speciation in Pellia,with special reference to the Pellia megaspora-endiviifolia complex (Metzgeriales), I. Taxonomy and distribution

Abstract

The genus Pellia, with three currently recognised species, includes a relatively primitive taxon, P. endiviifolia, in which the perichaetium is high and incised, with an enclosed calyptra. It is dioecious, produces anthocyanin wall pigments in exposed sites, reproduces asexually by repeatedly furcate, narrow fragmenting ‘autumnal innovations’, is freely branched and has relatively small spores. A species previously confused with this taxon, and here described as P. megaspora, is an endemic boreal American species, found from New York, Connecticut and Massachusetts north to Newfoundland and west to Minnesota. The plant has a tall perichaetium that is strikingly incised-ciliate to lacerate. It is dioecious but does not produce wall pigments and autumnal innovations, is simple or oncefurcate and has extremely large spores. Cytological and biochemical data support the specific recognition of P. megaspora. Biochemical data also suggest that Japanese populations currently assigned to P. endiviifolia are distinct and field observation suggests that these plants may represent an autonomous subspecies.     

Cytological diversity in Pellia endiviifolia (Dicks.) Dum

Abstract

Extensive cytological variation between British populations of Pellia endiviifolia (Dicks.) Dum. (n = 8+x/y) permits the recognition of six distinct karyotypes; one of them found also in Greece and another in the U.S.S.R. Although based on typically small amounts of constitutive heterochromatin, Giemsa C-banding patterns serve to identify each of three chromosomes, 1, 6 and 9, as dimorphic or polymorphic. Relative proportions of chromosomes differ between karyotypes and further variation relates toa nucleolar organiser region (NOR) on chromosome 6 of one population in addition to the NOR generally located in chromosome 7. These findings have repercussions intaxonomy and reproductive behaviour that are discussed in detail.     

Chromosomal distribution of rRNA genes in the karyotypes of two dioicous liverwort species from the genus Pellia Raddi

Over one hundred years have passed since the first cytogenetic studies were made on the liverwort genus Pellia Raddi. The karyotype of Pellia is characterised by large chromosomes, a varying heterochromatin content and the presence of sex chromosomes in the dioicous species. Most of the Pellia species are diploids with n?=?9, but one of them, Pellia borealis Lorb., has been described as an example of allopolyploidy in liverworts. Although the localisation of rRNA genes, which are essential components of the nuclear genome, remains a challenge in bryophytes, data on the number and chromosomal localisation of 35S and 5S rDNA in all of the Pellia species are now available. Previously, fluorescence in situ hybridisation using rDNA probes was performed on the mitotic chromosomes of 2 monoicous species. The aim of this study was to establish the number and chromosomal distribution of rRNA genes in 2 dioicous diploid species—Pellia endiviifolia (Dicks.) Dumort. and Pellia neesiana (Gottsche) Limpr. The relationships between the species within the genus Pellia can now be discussed in the context of the localisation of the rDNA sites and the range in the number of rDNA loci among bryophytes can also be verified.     

Distribution maps

Abstract

There are two groups of Pellia peroxidase isozyme phenotypes, one composed of P. epiphylla, P. neesiana and P. borealis, the other of P. endiviifolia from Europe, P. endiviifolia from Japan and P. megaspora. The phenotypes within the second group differ supporting the suggestion that there are three taxa present.     

Heterochromatin diversity in two species of Pellia (Hepaticae) as revealed by C-, Q-, N- and Hoechst 33258-banding

Giemsa C-banding of two liverwort species, Pellia epiphylla (L.) Corda (n=9) and P. neesiana (Gott.) Limpr. (n = X/Y + 8), identified each chromosome as unique. Almost the full complement of C-bands was defined by N-banding. Apart from minor variation in the prominence of certain bands, which were designated according to principles adopted in human cytology, the chief exceptions were the absence of bands 1p2, 1p4, 1q2, 1q4 and 7p6 from the N-banded karyotype of P. epiphylla and of band 7q25 from P. neesiana. Both quinacrine dihydrochloride and Hoechst 33258 showed reduced fluorescence in the constitutive heterochromatin of P. epiphylla. The effects of these two fluorochromes on P. neesiana, however, differed from each other and from their effect on P. epiphylla. Only one Q-band, the brightly fluorescing 1(X/Y)q24, occurred, whereas Hoechst 33258 resulted in considerable differentiation. The majority of C-bands were correlated with bright fluorescence with Hoechst 33258 but a few were dim. On the basis of these results, four major types of heterochromatin were identified in P. neesiana and two further types in P. epiphylla. They are discussed in the context of previously reported early replication of P. neesiana heterochromatin and point to considerable cytological evolution within these two species.     

Elemental composition of the sub-aquatic liverwort Pellia endiviifolia (Dicks.) Dum. in relation to heavy metal contamination

The concentrations of fifteen elements (B, Na, Mg, Al, P, K, Ca, Ti, Mn, Fe, Cu, Zn, Sr, Cd, Pb) in thalli of the sub-aquatic liverwort Pellia endiviifolia (Dicks.) Dum. collected from seven sites in Japan, were analyzed by inductively coupled plasma atomic emission spectrometer (ICP). The average concentration of major essential elements in the thalli was: 49 600 µg g^–1 K, 9 140 µg g^–1 Na, 6740 µg g^–1 Ca, 3 840 µg g^–1 P, and 3 060 µg g^–1 Mg. Thalli of P. endiviifolia from sites polluted by heavy metals (Cu, Zn, Pb) contained high concentrations of these elements (maxima 0.88% Cu, 0.55% Zn and 0.36% Pb in the older thallus), thus demonstrating the potentiality of this liverwort as an indicator for heavy metal contamination.     

Himalayan Moss Notes

Abstract

Studies of the relative and absolute lengths of mitotic chromosomes show that Pellia endiviifolia is cytologically distinct from the species P. megaspora, P. epiphylla and P. neesiana. This supports the recognition, on morphological grounds, of P. megaspora as a species distinct from P. endiviifolia.     

A cytological study of Pellia epiphylla (L.) Corda in Britain with reference to the status of Pellia borealis Lorbeer

Abstract

The history of Pellia borealis is outlined and its morphology compared with that of P. epiphylla. The techniques for preparing material for chromosome counts are described. Several gatherings of monoecious Pellia with large epidermal cells (P. borealis) were found to be haploid, n = 9. The size of the epidermal cells is shown to be valueless as a character for distinguishing the two species. It is concluded that only P. epiphylla should be retained as a species.

We wish to record our thanks to Dr S. Arnell for examining British material and for helpful correspondence; to Dr A. J. E. Smith for his valuable advice and criticism, and for the use of cytological facilities; and to all those who have contributed material of possible P. borealis.     

Response of bryophytes to exposure and water availability on some European travertines

Abstract

Seventeen travertine-bryophyte taxa were recorded in 212 quadrats from 17 sites, and the cover of the eight commonest species was analysed in terms of aspect, slope, exposure and water relations. Only Aneura pinguis was significantly affected by aspect, while Eucladium verticillatum and Hymenostylium recurvirostrum preferred steep slopes. The last two species, with Didymodon tophaceus, were also found to prefer sheltered sites, while Bryum pseudotriquetrum, Palustriella commutata var. commutata and Pellia endiviifolia were more common in exposed situations. Didymodon tophaceus, Hymenostylium recurvirostrum and Aneura pinguis were mostly confined to seepages and were kept moist by capillarity, while Palustriella commutata var. falcata and Pellia endiviifolia withstood high flows. Cratoneuron filicinum, Palustriella commutata var. commutata and Pellia endiviifolia had wide ecological amplitudes and were among the most frequently recorded species.     

Triploidy in Hochstetter's frog,Leiopelma hochstetteri,from New Zealand

Abstract

Autotriploidy is described in a female of the endemic New Zealand frog Leiopelma hochstetteri. This frog was found to have 3n=33 chromosomes plus 2 supernumerary chromosomes. All the chromosomes in the karyotype of this species contained C-band heterochromatin at the centromeres. A prominent C-band was found to be associated with a secondary constriction on chromosome no. 7. The supernumerary chromosomes in this species appear to be mitotically stable and contain C-band heterochromatin at the centromeres. From the limited data presently available, the triploid individual may have resulted from the fertilisation of a diploid egg produced when the second meiotic division had been suppressed.     

Changes in chlorophyll a fluorescence, photosynthetic CO2 assimilation and xanthophyll cycle interconversions during dehydration in desiccation-tolerant and intolerant liverworts

The interactions among water content, chlorophyll a fluorescence emission, xanthophyll interconversions and net photosynthesis were analyzed during dehydration in desiccation-tolerant Frullania dilatata (L.) Dum. and desiccation-intolerant Pellia endiviifolia (Dicks) Dum. Water loss led to a progressive suppression of photosynthetic carbon assimilation in both species. Their chlorophyll fluorescence characteristics at low water content were: low photosynthetic quantum conversion efficiency, high excitation pressure on photosystem II and strong non-photochemical quenching. However, dissipation activity was lower in P. endiviifolia and was not accompanied by a rise in the concentration of de-epoxidised xanthophylls as F. dilatata. The photosynthetic apparatus of F. dilatata remained fully and speedily recuperable after desiccation in as indicated by the restoration of chlorophyll fluorescence parameters to pre-desiccation values upon rehydration. A lack of recovery upon remoistening of P. endiviifolia indicated permanent and irreversible damage to photosystem II. The results suggest that F. dilatata possesses a desiccation-induced zeaxanthin-mediated photoprotective mechanism which might aid photosynthesis recovery when favourable conditions are restored by alleviating photoinhibitory damage during desiccation. This avoidance mechanism might have evolved as an adaptative response to repeated cycles of desiccation and rehydration that represent a real threat to photosynthetic viability. Received: 12 January 1998 / Accepted: 14 July 1998     

Complete or almost complete analogy of chromosome banding between the baboon (Papio papio) and man

Summary The karyotype of the baboon (Papio papio) was studied using several staining techniques: Q bands, R bands, T bands, C bands, G 11.6 bands, and after incorporation of BrdU. When compared with the karyotypes of the Pongidae and of man, it appears that the quasi totality, if not the totality, of the chromosome bands are common, when the heterochromatin is exclude. Among the Pongidae, P. pygmaeus (orangutan) has the karyotype the closest to that of P. papio, differing by seven pericentric inversions, three complex rearrangements, and three fusions. By this intermediate stage, the origin of the human chromosomes may be retracted over several dozen million years.     

Satellite DNA probes of Alstroemeria longistaminea (Alstroemeriaceae) paint the heterochromatin and the B chromosome,reveal a G-like banding pattern,and point to a strong structural karyotype conservation

Alstroemeria species present a well-conserved and asymmetric karyotype. The genus is divided into a Chilean clade, rich in heterochromatin, and a Brazilian clade, poor in heterochromatin. We investigated the distribution of the main repetitive sequences in the chromosomes of the Brazilian species A. longistaminea (2n = 16 + 0-6B) aiming to evaluate the role played by these sequences on the structural organization of the karyotype. In situ hybridization of the three most abundant retrotransposons, corresponding to ~ 45% of the genome, was uniformly distributed. Three satellite DNA sequences, representing near half of the whole satellite fraction (1.93% of the genome), were mainly concentrated on the heterochromatin and one of them painted the whole B chromosome. Noteworthy, some satellites were located on euchromatin, either dispersed or concentrated in clusters along the chromosomes, revealing a G-band-like pattern. The two satellites that presented more C-band- and G-band-like labeling were also hybridized in situ in two other Alstroemeria species. They revealed astonishing similar patterns of distribution, indicating an unusually structural karyotype conservation among Brazilian species.

     

Karyosystematics of thePhleum alpinum polyploid complex (Poaceae)

The karyotypes of three taxa from thePhleum alpinum group of sect.Phleum (P. alpinum subsp.rhaeticum, 2n = 14,P. commutatum, 2n = 28, and informally namedP. commutatum, 2n = 14) were investigated by Giemsa C-banding. The overall similarity of diploid genomes suggests thatP. alpinum subsp.rhaeticum andP. commutatum are closely related — their karyotypes vary only with respect to their average amounts of telomeric heterochromatin. TheP. commutatum genome contains less telomeric heterochromatin than the genome ofP. alpinum subsp.rhaeticum, but in theP. alpinum group as a whole almost fluent transition between low (1.5%) and high (25.5%) amounts of telomeric heterochromatin was observed among populations. In the karyotype of tetraploidP. commutatum, seven distinguishable chromosome types were observed. Each of them is represented at somatic metaphase by four chromosomes. C-band structure of karyotype and average amount of telomeric heterochromatin suggest that this taxon has originated from hybridization between two related diploid forms of theP. commutatum — P. alpinum complex.     

Banded karyotype of spined loach Cobitis taenia and triploid Cobitis from Poland

The present work provides new data on the banding pattern of diploid Cobitis taenia and its triploid hybrid females, which belong to the diploid–polyploid complex in the Vistula River tributary. C-banding, silver-staining (Ag), and fluorescent staining with chromomycin A₃ techniques were used to describe the diploid and triploid karyotype. The karyotype of Cobitis taenia of 2n=48 was characterised by one pair of NOR-bearing subtelocentric chromosomes and at least four chromosomes with CMA₃-positive sites. The C-positive heterochromatin was present in the centromeres of almost all chromosomes and the pericentromeric regions of several metacentric and submetacentric chromosomes. The triploid females of 3n=74 had two pairs of chromosomes with active NORs. The NORs-sites were located terminally on two biarmed and two uniarmed chromosomes. The CMA₃-staining revealed at least six A₃-positive sites. The C-banded and A₃-stained triploid karyotype was composed of haploid set of Cobitis taenia and diploid set of unidentified species, so heterochromatin pattern confirmed the possibility of their hybrid origin. The characteristics of banded diploid and triploid karyotype, and the hypothetical karyotype of an unknown species of 2n=50 is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chromosomal evolution inCallithrix emiliae

We studied the karyotype of specimens ofCallithrix emiliae (Callithricidae, Primates) from Rondonia, Brazil. Comparison with the karyotype ofCallithrix jacchus showed that, even though these two species show many karyotypic similarities, they differ by a Robertsonian translocation, a paracentric inversion and large-scale addition of heterochromatin. TheC. emiliae species appears to be in an active phase of chromosome evolution by the addition of constitutive heterochromatin.     

A new karyotype for <Emphasis Type="Italic">Cavia magna</Emphasis> (Rodentia: Caviidae) from an estuarine island and <Emphasis Type="Italic">C. aperea</Emphasis> from adjacent mainland

We describe a new karyotype for Cavia magna Ximenez, 1980 from an estuarine island and the karyotype of Cavia aperea Erxleben, 1777 from an adjacent mainland. The species have differences in diploid number (2n), autosomal fundamental number, quantity, and distribution of heterochromatin as dissimilar distributions of the nucleolus-organizing regions (Ag-NORs). The C. aperea karyotype has a diploid number of 64 as previously reported for C. aperea and most other Cavia species. In contrast, this new C. magna karyotype exhibits a variant diploid number of 2n = 62, considering that previous work reported a karyotype of 2n = 64 for C. magna. The discovery of a distinct diploid number within C. magna represents the first record of intra-specific chromosomal variation in a species of Caviidae. The diploid number of 2n = 62, heterochromatin quantity, Ag-NOR distributions, and inversed X chromosome from this population of C. magna are as seen in the geographically proximate (Cavia intermedia Cherem Olimpio and Ximenez; intermediate Cavy). These data provide further evidence supporting C. magna as the sister species of C. intermedia.     

Polymorphism of the X-chromosome,Y-chromosome and autosomes in the Australian hopping mice,Notomys alexis,N. cervinus and N. fuscus (Rodentia,Muridae)

All three species of Notomys so far studied possess a diploid number of 48. Many elements in the karyotype of N. alexis are polymorphic due to variation in heterochromatin, but the variation is most marked in autosomal pair 1, which occurs in at least four forms, the X-chromosome, which occurs in three forms, and the Y-chromosome which occurs in many forms. N. cervinus is unique in the genus in possessing an entirely biarmed karyotype due mainly to the addition of heterochromatic short arms. The X-chromosome of N. cervinus occurs in three forms and the Y-chromosome in two forms. The karyotype of N. fuscus is mainly telocentric although two autosomal pairs are polymorphic due to pericentric inversions. The X- and Y-chromosomes both occur in two forms in N. fuscus. Chromosome measurements and C-banding show that most of the variation in the size and morphology of the sex-pair both within and between species is due to variation in constitutive heterochromatin.     

Heterochromatin in mitotic chromosomes of theVirilis species group ofDrosophila

The amount of heterochromatin in the genome of ten members of thevirilis species group was determined as the length of C-band chromosome material relative to the total karyotype length. Thevirilis phylad (Drosophila virilis, D. novamexicana, D. americana americana, andD. americana texana) has significantly greater amounts of heterochromatin in the genome than do members of the montana phylad (D. montana, D. lacicola, D. flavomontana, D. borealis, D. ezoana, D. littoralis). Thus, the significant karyotypic change accompanying diversification of these species has involved reduction in their total constitutive heterochromatin. These changes have apparently involved reductions in the amount of centromeric heterochromatin in the autosomes.