Variation patterns in the Phlomis × composita (Lamiaceae) hybrid complex in the Iberian Peninsula

Hybridization seems to be common in the genus Phlomis (Lamiaceae) in the Iberian Peninsula, especially in the P.  ×  composita complex. In order to detect patterns of morphological variation linked to eco- and geographical variation, ecological (cluster and canonical correspondence) and morphometric (principal component and discriminant function) analyses were performed. Character count procedure was applied to discern between divergence and reticulate events for the origin of the morphologically intermediate plants. Following these analyses clear patterns were detected suggesting the existence of four independent morphological groups also supported by the ecogeographical data. These are P. crinita ssp. crinita (Levante, eastern Spain), P. crinita ssp. malacitana (Andalusia, southern Spain) and P. lychnitis (widespread) being the extremes of the morphological variation, and P.  ×  composita including all the morphologically intermediate individual plants. Furthermore, at the population level significant differences in hybrid plant frequencies between areas were also found.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 97–108.     

Whole-genome duplications in South American desert rodents (Octodontidae)

The discovery of tetraploidy in the red viscacha rat, Tympanoctomys barrerae (4 n  = 102) has emphasized the evolutionary role of genome duplication in mammals. The tetraploid status of this species is corroborated here by in situ PCR and Southern blot analysis of a single-copy gene. The species meiotic configuration strongly suggests a hybrid derivation. To investigate the origin of T. barrerae further, the recently described Pipanacoctomys aureus was studied. This 92-chromosome species also has a duplicated genome size, redundant gene copy number and diploid-like meiotic pairing, consistent with an event of allotetraploidization. Phylogenetic analysis of mitochondrial sequences indicates sister-group relationships between these two tetraploid rodents. The new karyotypic data and the phylogenetic relationships suggest the participation of the ancestral lineages of Octomys mimax in the genesis of P. aureus . The high overall DNA similarity and shared band homology revealed by genomic Southern hybridization as well as matching chromosome numbers between O. mimax and the descendant tetraploid species support the notion of introgressive hybridization between these taxa.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 443–451.     

Two new species of Galium (Rubiaceae) from the Iberian Peninsula

Two new species of the genus Galium are described from the Iberian Peninsula: G. moralesianum Ortega-Olivencia & Devesa, and G. talaveranum Ortega-Olivencia & Devesa. The first grows on limestone or dolomitic-limestone outcrops in south-east Spain (the Segura and Gádor mountain ranges) and shows morphological similarities with G. boissieranum Ehrend. & Krendl, which is endemic to southern Spain (mountains of Málaga). The second inhabits the margins of water courses, alder stands and cork-oak dehesas (parkland-type systems) of the west of the Iberian Peninsula, and is morphologically reminiscent of G. mollugo L. The chromosome numbers of these two species and of G. boissieranum are given.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 177−187.     

Genome organization in diploid hybrid species of Argyranthemum (Asteraceae) in the Canary Islands

Two diploid species of hybrid origin, Argyranthemum lemsii and A. sundingii , have been described from different valleys in the Anaga peninsula, north-east Tenerife. They have previously been shown to originate from hybridization between the same parental species, the montane A. broussonetii and the coastal A. frutescens , A. broussonetii being the chloroplast donor in one valley and A. frutescens in the other. The specific status of the two hybrid species has been questioned. In this study we used karyotype analysis, FISH, and GISH to address the question of multiple diploid hybrid speciation. GISH did not discriminate clearly between the parental genomes, but differential labelling was observed in separate hybrid populations, indicating different chromosomal rearrangements in different valleys. Small karyotype differences and local loss of rDNA were also observed. Thus separate origins of the same hybrid combination in different valleys in Tenerife have been verified. Our results add some support to the recognition of two species of hybrid origin, but the case serves to illustrate some of the many problems connected with the species concept in plants.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 491–501.     

Genetic structure and morphological variation of British populations of the hybrid Potamogeton x salicifolius

Potamogeton x salicifolius Wolfg. is one of the three most frequently recorded Potamogeton hybrids in the British Isles and Europe. It is thought to be the hybrid between P. lucens and P. perfoliatus. Its scattered distribution suggests that it has arisen several times in Britain. Most British populations of P. x salicifolius can be identified by their morphological characteristics, which are intermediate between those of the putative parents P. lucens and P. perfoliatus . However, the population at the Ouse Washes, Cambridgeshire, differs from other populations in its greater similarity to P . lucens . A genetic study of eight British populations, using six isozyme systems, revealed that most populations consist of a single multi-enzyme phenotype. This suggests that they were the result of a single hybridization event and are therefore maintained through vegetative reproduction. By contrast, the Ouse Washes population consists of three multi-enzyme phenotypes. This variation is likely to have resulted from multiple hybridization events, although we cannot exclude the possibility that the plants are partially fertile. The isozyme systems studied were unable to identify P. lucens and P. perfoliatus unambiguously, and consequently did not provide evidence for their putative parentage of P. x salicifolius. However, at a local level the banding patterns of the hybrids were generally consistent with the local multi-enzyme phenotypes of these putative parents.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 99–111.     

Chromosome variation in the genus Pinellia (Araceae) in China and Japan

The chromosome numbers of 23 populations belonging to seven species of Pinellia Tenore were counted. The basic chromosome number of all the species studied was x  = 13, and the previously reported basic numbers x  = 9 and x  = 14 in P. ternata and P. cordata were not confirmed. Chromosome numbers of 2 n  = 26 in P. polyphylla and 2 n  = 78 in P. integrifolia are reported for the first time. Heptaploid (7 x ) and nonuploid (9 x ) are new ploidy levels reported for P. ternata , and hexaploid for P. cordata. Taxonomic, phylogenetic and phytogeographical inferences are made for the genus. Particular attention was given to P. ternata , the most widely distributed species in the genus, and considerable variation of chromosome number was found in its different populations. Based on chromosome studies of 11 populations of P. ternata , together with 12 populations reported in previous studies, the lower reaches of the Yangze River are identified as its centre of origin. From there it dispersed, with generation of hexaploid, heptaploid, octoploid and nonuploid forms.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 449–455.     

Hybrids between Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) do not show endogenous postzygotic isolation

Studies of prezygotic and postzygotic isolation in Drosophila have shown in general that species in sympatry tend to evolve prezygotic barriers earlier than do species in allopatry. However, postzygotic barriers tend to evolve at the same evolutionary rate in both sympatric and allopatric species. In contrast to these observations, the grasshoppers Chorthippus parallelus parallelus and C. p. erythropus show complete hybrid male sterility but only limited prezygotic isolation after an estimated 0.5 millions years of divergence. Like their congeners, C. brunneus and C. jacobsi form a hybrid zone where their ranges meet in northern Spain. However, the hybrid zone is mosaic and bimodal and, in contrast to the high levels of postzygotic isolation between C. parallelus subspecies, these two species showed no significant reduction in hybrid fitness in F₁ or backcross generations relative to the parental generations. The level of prezygotic isolation in laboratory tests was comparable to that between C. parallelus subspecies. These results suggest that endogenous postzygotic isolation does not play an important role in the reproductive isolation between C. brunneus and C. jacobsi , or in determining the structure of the hybrid zone. Exogenous postzygotic isolation may be present and should be tested in future studies.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 195–203.     

A new species of Valantia (Rubiaceae) from Spain

A new species of Valantia , V. lainzii Devesa & Ortega-Olivencia, endemic to the coastal zone of Granada (southern Spain) is described. The species recalls V. muralis L. in its general aspect, and V. deltoidea Brullo in the type of fructiferous body.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 331–335.     

Meiotic drive at the Om locus in wild-derived inbred mouse strains

Meiotic drive is an evolutionary force in which natural selection is uncoupled from organismal fitness. Recently, it has been proposed that meiotic drive and genetic drift represent major forces in the evolution of the mammalian karyotype. Meiotic drive involves two types of genetic elements, Responders and Distorters , the latter being required to induce transmission ratio distortion at the former. We have previously described the Om meiotic drive system in mouse chromosome 11. To investigate the natural history of this drive system we have characterized the alleles present at the distorter in wild-derived inbred strains. Our analysis of transmission of maternal alleles in both classical and wild-derived inbred strains indicated that driving alleles are found at high frequency in natural populations and that the existence of driving alleles predates the split between the Mus spicilegus and M. musculus lineages.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 487–492.     

Cytotypes of Andropogon gerardii Vitman (Poaceae): fertility and reproduction of aneuploids

In many North American prairies, populations of Andropogon gerardii Vitman (Poaceae) are composed of hexaploid and enneaploid cytotypes (2 n  = 60, 90), with intermediates occurring occasionally. Under controlled pollination, the two common cytotypes can be crossed, producing progeny with a range of chromosome numbers. In an investigation of fertility and compatibilities of intermediate cytotypes, individuals with chromosome numbers between 60 and 90 were crossed with each other, with the 2 n  = 60 and 90 cytotypes, and with South American Andropogon species having 60 chromosomes. Regardless of cytotype, all A. gerardii plants had some fertility and virtually all crosses produced seeds. Cytotype is only partially predictive of fertility. Inter-specific hybrids between A. gerardii and South American hexaploid species were vigorous but sterile. Gene flow in natural A. gerardii populations of mixed cytotype probably involves plants of all cytotypes.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 95–103.     

Non-metric morphological divergence in the western house mouse, Mus musculus domesticus, from the Barcelona chromosomal hybrid zone

The effect of hybridization on morphological variation was investigated in 120 western house mice, Mus musculus domesticus , from the hybrid zone between the Barcelona and standard chromosomal races. The incidence of 37 non-metric cranial traits was calculated for standard mice (2 n  = 40) and Barcelona-standard hybrids (2 n  = 27–39). Subsequent analyses were conducted on several karyological subgroups, established by grouping the animals according to either their diploid number or their degree of chromosomal heterozygosity. Results revealed no significant difference by sex, asymmetry, or geographical distance. Significant phenetic divergences were found between the karyotypes studied in relation to several variants. Differences were especially substantial between the standard race and hybrid mice, even with respect to those hybrids with karyotypes close to that of the standard race. Within the hybrids, the maximum divergence corresponded to the 28-chromosome homozygotes, chromosomally close to the Barcelona race, and to the heterozygotes with more than two fusions. Since differences in non-metric trait frequencies are generally considered a measure of genetic divergence, the results suggest the occurrence of a barrier to gene flow in the Barcelona hybrid zone. The decrease of genetic exchange between the chromosomally differentiated mice might be due to reduced fertility in hybrids, associated with chromosomal heterozygosity.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 313–322.     

Morphology of Pinus uliginosa (Pinaceae) needles from populations exposed to and isolated from the direct influence of Pinus sylvestris

Eleven needle traits of two contrasting populations of Pinus uliginosa ( P. mugo complex), one located in a Pinus sylvestris forest, the other isolated from the influence of this species, were compared to quantify the degree of hybridization. Statistically significant differences between the studied populations were found, but both of them differed by a similar degree from P. sylvestris . Surprisingly, the P. uliginosa population surrounded by the P. sylvestris forest, which was expected to consist of individuals with intermediate traits, appeared to be more closely related to P. mugo than those isolated from the direct influence of P. sylvestris . Large differences in the distances between the analysed pair of populations of P. uliginosa and P. sylvestris have not influenced the morphological traits of the species. Gene flow from P. sylvestris to P. uliginosa is possible, but is likely to be small and does not differ compared with populations of P. uliginosa . The decline observed during the last three decades in both populations of the species has not influenced their needle morphological traits.  © 2003 The Linnean Society of London , Botanical Journal of the Linnean Society , 2003, 142, 83–91.     

Transitions in cuticular composition across a hybrid zone: historical accident or environmental adaptation?

Two subspecies of the grasshopper Chorthippus parallelus form a hybrid zone in the Pyrenees. Transitions across the zone, including changes in mating signals and reproductive isolation have been intensively studied. Cuticular pheromones have been identified as likely mate recognition signals. Since the major role of the cuticle is in waterproofing, environmental adaptation of cuticular composition has the interesting potential to generate assortative mating as an incidental by-product. We describe the pattern of variation in cuticular hydrocarbon blend in four transects through the hybrid zone. We find no evidence for a previously observed displaced cline in one blend component. There were differences between subspecies but these varied among transects and were small compared with variation between transects. We examined environmental variation within one transect and found a correlation between vegetation and cuticular composition, suggesting that environment influences the constitution of the cuticle, and hence natural selection may interact with mating signals in this species.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 193−201.     

Karyotype evolution in the genus Boronia (Rutaceae)

New chromosome counts are reported for Boronia clavata 2 n  = 14, B. heterophylla 'Near White' 2 n  = 15, B.  'Carousel' 2 n  = 16, B. deanei 2 n  = 22, B. chartacea 2 n  = 32, B. keysii 2 n  = 32, B. pilosa 2 n  = 44, B. anethifolia 2 n  = 36 and B. citriodora 2 n  = 108. Studies in 20 genotypes of 18 species and one interspecific hybrid revealed that they are highly complex in terms of chromosome number, ploidy level, chromosomal length, karyotype constitution and asymmetry. Karyotype analysis indicated that Boronia taxa with high chromosome numbers are primitive and those with lower numbers are derived. The basic chromosome number for this genus is suggested to be x = 18. Analysis of chromosome number, variations of total chromosome length (TCL) and average chromosome length (ACL), Nombre Fondamental (NF) and karyotype asymmetry suggest that dysploid reduction is the major mechanism in Boronia karyotype evolution. Chromosomal rearrangements might also have been involved. Origin, chromosome number changes and spread of Boronia are discussed in relation to the species divergence and the geological and climatic changes of the Australian continent.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 309–320.     

Determination of ploidy and nuclear DNA content in populations of Atriplex halimus (Chenopodiaceae)

Nuclear DNA contents were determined by flow cytometry for 20 populations of the perennial C₄ shrub Atriplex halimus L. (Chenopodiaceae) originating from the Mediterranean basin and Fuerteventura (Canary Islands). Two populations were also analysed for chromosome number: one (from Ibiza, Spain), with a 2C nuclear DNA content of 2.40 pg, was shown to be diploid (2 n  = 2 x  = 18), whilst the other (from Sicily, Italy), with 5.11 pg, was tetraploid (2 n  = 2 x  = 36). With respect to nuclear DNA content, two groups of populations were detected, diploids with 2.40–2.44 pg and tetraploids with 4.77–5.13 pg. The diploid populations were mainly from the western Mediterranean (Spain and France) and Fuerteventura, whereas tetraploids were generally, but not exclusively, from more arid areas in North Africa and the eastern Mediterranean. In general, the diploid and tetraploid populations corresponded to the subspecies halimus and schweinfurthii , respectively. For certain populations having morphologies intermediate between those considered typical of these two subspecies, nuclear DNA contents showed them to be tetraploid. There was significant variation in nuclear DNA content among the tetraploid populations, with greater values in the more easterly populations.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 147 , 441−448.     

Convergent evolution of the cavefish Astyanax (Characidae, Teleostei): genetic evidence from reduced eye-size and pigmentation

More than 20 populations of the cave-dwelling characid Astyanax occur within a restricted karst area in Mexico. The fish possess reduced eyes without lenses and visual cells. It is still an open question as to whether this condition evolved convergently after multiple entries of the surface ancestor into the different caves or whether a single cave ancestor, already characterized by reduced eyes, spread secondarily within them. In the crosses between specific populations, specimens appear that deviate considerably from those of the parents. They possess larger and better-developed eyes with histologically intact lenses and visual cells; they thus have the structural potential for vision. This indicates that in different cave populations, different mutations in the eye gene system have occurred. In cases where these non-functional rudimentary genes are recombined in hybrid specimens, gene expression may be restored. This is the result of separate evolution of several Astyanax cave populations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 545−554.     

Genetic characterization of hybridization between the New Zealand everlastings Helichrysum lanceolatum and Anaphalioides bellidioides (Asteraceae: Gnaphalieae)

Intergeneric hybridization between the endemic New Zealand species Helichrysum lanceolatum and Anaphalioides bellidioides (Asteraceae: Gnaphalieae) at two Banks Peninsula populations was investigated by DNA sequencing and amplified fragment length polymorphism fingerprinting. Crosses in both directions have contributed to the generation of the F1 hybrids, but in the study season hybridization was far more common with H. lanceolatum as the maternal parent. F1 hybrids have reduced, but considerable, fertility and seed collected from them includes back-crosses. Two hybrid plants sampled from the wild appear to be back-crosses to H. lanceolatum. Despite back-crossing, introgression was not demonstrated and is at most limited in scope. These results also corroborate important roles for small population size and rarity in the formation of hybrid lineages in New Zealand's everlastings.  © 2007 Landcare Research New Zealand Ltd. Journal compilation © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 89–98.     

Karyological and taxonomic notes on Ophrys (Orchidoideae, Orchidaceae) from the Iberian Peninsula

Karyological information on Iberian Ophrys species is very limited. This paper provides the haploid and diploid chromosome numbers of 11 taxa of sect. Pseudophrys and sect. Ophrys , both of which are well represented in the Iberian Peninsula, and two taxa from Tunisia. The first data on chromosome numbers for O. vasconica (2 n  = 72, 74), O. ficalhoana (2 n  = 36), O. picta (2 n  = 36), O. sphegifera ( n  = 18, 2 n  = 36, 38) and O. passionis (2 n  = 36) are also presented, confirming the stability of the chromosome number in Ophrys . In addition, populations of the group O. omegaifera ( O. dyris and O. vasconica ), together with tetraploidy, pentaploidy and the existence of aneuploid phenomena, are reported for the first time in Iberia. The basic diploid number is always 2 n  = 36. The karyotypes of several species were analysed. Evolutionary trends in Ophrys chromosomes are discussed. Taxonomic and phytogeographical details are provided on several species or groups of species from Iberia.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 142 , 395−406.     

Phylogeography of five Polytrichum species within Europe

Using allozymes and microsatellites we have analysed the genetic structure among European populations for several Polytrichum species to infer relevant factors, such as historical events or gene flow, that have shaped their genetic structure. As we observed low levels of genetic differentiation among populations, and no decreasing levels of genetic variation with increasing latitude within most of the examined species, no genetic evidence was obtained for a step-wise recolonization of Europe from southern refugia after the latest glacial period for P. commune , P. uliginosum , P. formosum and P. piliferum . The near absence of population substructuring within these species does indicate that extensive spore dispersal is the most important factor determining the genetic structure among European Polytrichum populations. Gene flow levels have apparently been sufficient to prevent genetic differentiation among populations caused by genetic drift, and to wipe out any genetic structure caused by the postglacial recolonization process. On the other hand, increased genetic differentiation of alpine P. formosum populations suggests that mountain ranges might restrict gene flow significantly among Polytrichum populations. In contrast to most examined Polytrichum species, P. juniperinum showed high levels of genetic differentiation and a profound genetic structure. Assuming that gene flow is not more restricted in P. juniperinum , these findings suggest that this species has recolonized Europe after the latest glacial period from two different refugia, one possibly being the British Isles.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society 2003, 78, 203–213.     

High levels of genetic diversity in populations of Iris aphylla L. (Iridaceae), an endangered species in Poland

We used RAPDs (Random Amplified Polymorphic DNAs) to investigate genetic diversity and its partition within and between three populations of Iris aphylla in Poland. Analysis of Molecular Variance (AMOVA) of 84 distinct RAPD multiband genotypes revealed higher variation within populations (77.2%) than genetic differentiation between them (22.8%, P  < 0.002). Values of genetic diversity indices ( H ) were similar in all three sites (0.21–0.24). The differentiation of the populations corresponded to low average gene flow ( Nm  = 0.81). Our results indicated that genetic diversity was independent of population size. We concluded that although sexual reproduction and gene flow between populations of I. aphylla were very limited, they preserved high levels of genetic diversity. Relatively large number of seeds, which migrated in the past to populations, as well as patterns of reproduction and life history of I. aphylla may explain this situation.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 65–72.