Karyological observations on plant taxa endemic to the Balearic Islands

Mitotic chromosome numbers are reported from 35 vascular plant taxa endemic to the Balearic Islands that are poorly known cytogenetically. The chromosome numbers of Chaenorhinum rodriguezii (2 n  = 14), Coristospermum huteri (2 n  = 22), Carduus ibicensis (2 n  = 26), Filago petro-ianii (2 n  = 26), Ononis crispa ssp. zschackei (2 n  = 30), and Ophrys bertolonii ssp. balearica (2 n  = 36) were determined for the first time. New chromosome numbers are reported in Pimpinella bicknellii (2 n  = 22), Calamintha rouyana (2 n  = 46), Vicia bifoliolata (2 n  = 14), and Teucrium asiaticum (2 n  = 30). A new diploid cytotype (2 n  = 16) was found in Ranunculus paludosus ssp. barceloi , which is restricted to populations from the western Balearics (Eivissa and Formentera), whereas tetraploid plants are known from the eastern Balearics (Mallorca). It is suggested that the tetraploid cytotype arose from the diploid cytotype through autopolyploidy. The idiograms of Coristospermum huteri , Pastinaca lucida , Pimpinella bicknellii , Carduus ibicensis , Helianthemum scopulicolum , Helichrysum crassifolium , Vicia bifoliolata , p aeonia cambessedesii , Helleborus lividus , Ranunculus paludosus ssp. barceloi , and Linaria aeruginea ssp. pruinosa are determined for the first time.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 463–476.     

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.     

Chromosome numbers in Hieracium and Pilosella species (Asteraceae) from the Iberian Peninsula and the Balearic Islands

Mitotic chromosome numbers are reported from 16 Hieracium and two Pilosella species from the Iberian Peninsula and the Balearic Islands that are little known cytogenetically. The chromosome numbers of Hieracium aragonense Scheele (2 n  = 27, 36), H. compositum Lapeyr. (2 n  = 27), H. murcandidum G. Mateo (2 n  = 27), H. spathulatum Scheele (2 n  = 27), H. segurae Mateo (2 n  = 27), H. teruelanum Mateo (2 n  = 27), H. valentinum Pau (2 n  = 27), Pilosella pseudovahlii (De Retz) Mateo (2 n  = 18), and P. tardans (Peter) Sójak (2 n  = 36) were determined for the first time. New cytotypes were detected in H. cordifolium Lapeyr. (2 n  = 27) and H. loscosianum Scheele (2 n  = 36). The karyotype of the studied species was similar in overall morphology and comprised metacentric and submetacentric chromosomes; secondary constrictions (up to two) were detected in the chromosome complements of some species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 311–320.     

A new annual species of Bellium (Asteraceae) from the Balearic Islands

A new species, Bellium artrutxensis (Asteraceae: Astereae), is described from the therophytic pastures of southern Minorca (Balearic Islands). The analysis of nuclear ribosomal internal transcribed spacer (ITS) sequences showed that the new taxon was closely related to Bellium bellidioides , but differed with regard to the annual lifespan, the absence of epigeal stolons, and the presence of a patent eglandular indumentum on the leaves. The new taxon shows morphological features that are well suited for the colonization of dry environments with a marked summer drought, and may be of adaptive significance. The new species is diploid (2 n  = 18) and shows up to two accessory chromosomes in root tissues. B chromosomes have not been reported previously in other species of Bellium .  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 65–77.     

The taxonomy of Galium crespianum J.J. Rodr. (Rubiaceae), a Balearic Islands endemic revisited

Galium crespianum J.J. Rodr. is a perennial plant inhabiting cliffs and crevices on the Balearic Islands. Although it was described from the mountains of Mallorca, several populations from the islands of Eivissa and Formentera have been traditionally considered as belonging to the same species. A careful comparison of specimens from the western and eastern Balearics revealed several discriminant features in gross morphology (robustness and habit of the stems, presence or absence of hairs at the internodes and leaves, waxiness of stems and leaves, shape of the inflorescence, flower colour and scent) and stem anatomy. In addition, several mutations in the ribosomal ITS sequences distinguished western and eastern plants. It is proposed that the plants from Eivissa and Formentera previously subsumed under G. crespianum be recognized at the specific level as G. friedrichii N. Torres, L. Saez, Mus & Rossello nom. nov. The new species is tetraploid (2n = 44) and shares morphological features with G. crespianum or with several taxa in the G. glaucum complex from the south east Iberian peninsula, to which it is most probably related.     

Volatile components vaxiation in the Teucrium marum complex (Lamiaceae) from the Balearic Islands

The volatile compounds of 18 samples from the western Balearic Islands belonging to three taxa of the Teucrium marum L. complex (T. marum subsp. marum, T. marum subsp. occidentale and T. marum subsp. drosocalyx) were fractionated by distillation-extraction and analysed by means of GC-MS. Seventeen components were characterized and the quantitative data were subjected to multivariate and clustering techniques. Two main groups appeared, one comprising T. marum subsp. occidentale samples and the other grouping T. marum subsp. marum and T. marum subsp. drosocalyx accessions. Chemical data and taxonomic groupings generally agreed, but quantitative and qualitative differences within and between taxa were found. Specimens from Minorca and Cabrera islands showing intermediate morphological features between T. marum subsp. marum and T. marum subsp. occidentale were included within the T. marum subsp. marum-T. marum subsp. drosocalyx group in the Principal Component Analysis. The origin of these intermediate specimens is discussed, and it is suggested that they are local variants of T. marum subsp. marum which had an independent origin throughout its area.     

Reproductive ecology of Cneorum tricoccon L. (Gneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Reproductive ecology of Cneorum tricoccon L. (Cneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Chromosome numbers in the tribe Anthemideae (Asteraceae) from north-east Anatolia

Twenty-two chromosome counts of 19 taxa in the tribe Anthemideae of the family Asteraceae are reported from north-east Anatolia, Turkey. The taxa belong to the subtribes Achilleinae (four Achillea taxa), Anthemidinae (five Anthemis taxa), Artemisiinae (two Artemisia species), Leucantheminae (one Lecanthemum species), Matricariinae (two Tripleurospermum species) and Tanacetinae (five Tanacetum taxa). Six counts are new reports, seven are not consistent with previous counts, and the remainder are confirmations of very limited previous data. Most of the populations of Anthemideae studied have the basic number x  = 9, with ploidy levels ranging from 2 x to 7 to 8 x , but dysploidy is also present, with one diploid species, Artemisia austriaca , having x  = 8.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 203–211.     

Contributions to the cytotaxonomy of the Commelinaceae. Chromosome evolution in Tradescantia section Cymbispatha

The five species of Tradescantia section Cymbispatha studied, including one species T. poelliae D. R. Hunt, have chromosome numbers of In = 12, 14, 16, 22, 28, 30 and 36 and karyotypes of acrocentric, metacentric or telocentric chromosomes, or mixtures of both acrocentric and metacentric chromosomes. The numbers of major chromosome arms of these cytotypes give a nombre fondamentaP series of 14, 28, 42 and 56 which, in combination with meiotic analyses, indicates plants which, in genetical terms at least, are diploid, tetraploid, hexaploid and octoploid. This series has evolved from a 2 n = 14 acrocentric or telocentric karyotype by a combination of Robertsonian fusion and polyploidy. Pseudo-iso-chromosomes are sometimes formed in this evolutionary development and can persist as stable members of normal complements.     

Contributions to the cytotaxonomy of the Commelinaceae. Chromosome evolution in Tradescantia section Cymbispatha

The five species of Tradescantia section Cymbispatha studied, including one species T. poelliae D. R. Hunt, have chromosome numbers of In = 12, 14, 16, 22, 28, 30 and 36 and karyotypes of acrocentric, metacentric or telocentric chromosomes, or mixtures of both acrocentric and metacentric chromosomes. The numbers of major chromosome arms of these cytotypes give a nombre fondamentaP series of 14, 28, 42 and 56 which, in combination with meiotic analyses, indicates plants which, in genetical terms at least, are diploid, tetraploid, hexaploid and octoploid. This series has evolved from a 2 n = 14 acrocentric or telocentric karyotype by a combination of Robertsonian fusion and polyploidy. Pseudo-iso-chromosomes are sometimes formed in this evolutionary development and can persist as stable members of normal complements.     

Chromosome numbers in the tribes Anthemideae and Inuleae (Asteraceae)

Twenty-two chromosome counts of 19 taxa (21 populations) in the tribe Anthemideae and one member (one population) of the tribe Inuleae of the family Asteraceae are reported. The Anthemideae studied belong to the subtribes Artemisiinae (14 Artemisia taxa, and one species each of the genera Dendranthema , Filifolium and Neopallasia ) and Tanacetinae (one species each of the genera Lepidolopha and Tanacetopsis ). From the Inuleae, we studied one Inula species. Five counts are new reports (including two at generic level), six are not consistent with previous counts and the remainder are confirmations of very limited (one to four records) previous data. Most of populations of Anthemideae studied have the basic chromosome number x  = 9, with ploidy levels ranging from 2 x to 10 x . Dysploidy is also present, with two x  = 8 diploid taxa. The species of Inuleae studied is a diploid with x  = 10, also indicating dysploidy, other members of the same genus Inula having basic numbers of x  = 9 or 8.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 77–85.     

Herbivory and plant competition effects on the population dynamics of Naufraga balearica,a threatened narrow endemic species of the Balearic Islands

Naufraga balearica is a caespitose member of the Umbelliferae and is endemic to Majorca. We conducted a 12-year study in which permanent plots were monitored (each plot was divided into 100 cells) to assess (1) the natural population dynamics, (2) goat herbivory (1998–2003) and (3) plant competition. Using photographs and a G-test of independence, we analysed the frequency of cells in which a species was present and, for each annual transition, the frequency of cells where the species was stable, decreased or increased for each species and treatment (plots open to herbivores vs. closed plots). The natural dynamics of this threatened species showed a clear decline at the study site. Dactylis glomerata and Erodium reichardii were the species that appeared to have the greatest effect on the presence of N. balearica, especially in the closed plots. The competition effect of neighbouring species was not homogeneous throughout the 11 annual transitions analysed, suggesting that other factors affect the dynamics of the threatened species. Some herbivory pressure is beneficial to N. balearica because the species fluctuates more in the opened plots and the behaviour of the goats helps to control competing species, permitting the establishment of N. balearica.     

Spatial genetic structure of Aquilegia taxa endemic to the island of Sardinia

Background and Aims

The Mediterranean Basin is one of the most important regions for the Earth''s plant biodiversity; however, the scarcity of studies on fine scale patterns of genetic variation in this region is striking. Here, an assessment is made of the spatial genetic structure of all known locations of the three Sardinian endemic species of Aquilegia in order to determine the relative roles of gene flow and genetic drift as underlying evolutionary forces canalizing the divergence of Sardinian Aquilegia taxa, and to see if the spatial genetic structure found fits the current taxonomic differentiation of these taxa.

Methods

DNA from 89 individuals from all known locations of Aquilegia across Sardinia was analysed by means of amplified fragment length polymorphism (AFLP) markers. Both principal co-ordinates analysis (PCoA) and Bayesian clustering analyses were used to determine the spatial genetic structure irrespective of any taxonomic affiliation. Historical effects of gene flow and genetic drift were assessed by checking for the existence of isolation-by-distance patterns.

Key Results

STRUCTURE and PCoA analyses revealed a pattern of genetic variation geographically structured into four spatial genetic groups. No migration–drift equilibrium was detected for Aquilegia in Sardinia, when analysed either as a whole or in individual groups. The scenario approached a Case III pattern sensu Hutchinson and Templeton, which is associated with extreme isolation conditions where genetic drift has historically played a dominant role over gene flow.

Conclusions

The pattern of genetic variation of Sardinian taxa of Aquilegia indicates that genetic drift has been historically more influential than gene flow on population structure of Sardinian species of Aquilegia. Limited seed dispersal and divergent selection imposed by habitat conditions have been probably the main causes reinforcing post-Pleistocene geographical isolation of Aquilegia populations. The spatial genetic structure found here is not fully compatible with current taxonomic affiliations of Sardinian Aquilegia taxa. This is probably a consequence of the uncoupling between morphological and genetic patterns of differentiation frequently found in recently radiated taxa.     

Historical processes and environmental factors as determinants of inter-island differences in endemic faunas: the case of the Balearic Islands

Aim We describe current interisland similarities of endemic faunas, and elucidate the significance of historical factors and environmental ones in determining the pattern found. Location The six major islands of the Balearics (Western Mediterranean). Method An extensive review of all the endemic fauna ranging from platyhelminthes to mammals is made. From 568 presumed endemic species and subspecies, 230 full species with neither taxonomic nor distributional uncertainty are chosen. Inter-island similarities are determined using such a presence-absence matrix. Finally, relationships between the matrix of faunistic similarity and a number of matrices measuring environmental and historical factors are elucidated. Results Endemic fauna similarities depend clearly on historical factors. Dependence on environmental factors is unclear. Moreover, endemic fauna reveals two clear-cut clusters of islands within the Balearics: the Gymnesic Islands, in the NE, and the Pityusic Islands in the SW. Historical factors cluster the Balearic Islands in the same way. Contrasting, environmental variables show smoothed, no significant differences among the Gymnesics and the Pityusics. Main conclusions Pre-human flora (palynology) and fauna (bird and mammal fossil record) suggest that environmental differences among the Gymnesics and the Pityusics have now been reduced in comparison to the environmental differences at the Pleistocene and Holocene boundary. This environmental homogenization is likely related with human invasion. Historical effects of prehuman differences between Gymnesic and Pityusic Islands are still recognizable on endemic fauna. In contrast, there is no historical effects on interisland similarities using currently breeding birds (as an example of organisms well-dispersed and related to vegetation type). We explain the pattern of interisland similarities of endemic fauna as the result of the independent histories among the two islands groups. Contrasting, successive colonizations and extinctions would determine interisland similarities of breeding birds.     

Spatial and environmental determinants of vascular plant species richness distribution in the Iberian Peninsula and Balearic Islands

Using an exhaustive data compilation, Iberian vascular plant species richness in 50 times 50 UTM grid cells was regressed against 24 explanatory variables (spatial, geographical, topographical, geological, climatic, land use and environmental diversity variables) using Generalized Linear Models and partial regression analysis in order to ascertain the relative contribution of primary, heterogeneous and spatially structured variables. The species richness variation accounted for by these variables is reasonably high (65% of total deviance). Little less than half of this variation is accounted for spatially structured variables. A purely spatial component of variation is hardly significant. The most significant variables are those related to altitude, and particularly maximum altitude, whose cubic response reflects the occurrence of the maximum number of species at the highest altitudes. This result highlighted the importance of Iberian mountains as hotspots of diversity and the relevance of large and small scale historical factors in contemporary plant distribution patterns. Climatic or energy-related variables contributed little, whereas geological (calcareous and acid rocks) and, to a lesser extent, environmental heterogeneity variables (land use diversity and altitude range) seem to be more important.     

A review on the effects of alien rodents in the Balearic (Western Mediterranean Sea) and Canary Islands (Eastern Atlantic Ocean)

Invasions of alien rodents have shown to have devastating effects on insular ecosystems. Here we review the ecological impacts of these species on the biodiversity of the Balearic and the Canary Islands. A total of seven species of introduced rodents (two rats, three mice, one dormouse, and one squirrel) have been recorded (six in the Balearics and four in the Canaries). Some of them can occasionally be important predators of nesting seabirds, contributing to the decline of endangered populations in both archipelagos. Rats are also known to prey upon terrestrial birds, such as the two endemic Canarian pigeons. Furthermore, rats actively consume both vegetative and reproductive tissues of a high number of plants, with potential relevant indirect effects on vegetation by increasing erosion and favoring the establishment of alien plants. In the Balearics, rats and mice are important seed predators of endemic species and of some plants with a restricted distribution. In the Canaries, rats intensively prey upon about half of the fleshy-fruited tree species of the laurel forest, including some endemics. In both archipelagos, alien rodents disrupt native plant–seed dispersal mutualisms, potentially reducing the chances of plant recruitment at the same time that they modify the structure of plant communities. We further suggest that alien rodents played (and play) a key role in the past and present transformation of Balearic and Canarian native ecosystems.     

Chromosome counts in Asian Artemisia L. (Asteraceae) species: from diploids to the first report of the highest polyploid in the genus

This work comprises 24 reports of chromosome numbers in 24 Artemisia L. species from Asia. Ten are included in the subgenus Dracunculus Besser and the rest belong to other subgenera. Seven counts are new reports, 14 are consistent with scarce previous ones, and three contribute new ploidy levels. That carried out in A. medioxima reports the highest ploidy level ever counted for the genus (16 x ). There is only one species with x  = 8 as the basic chromosome number. In the remaining x  = 9-based species, ploidy levels range from 2 x to 16 x , illustrating the great role played by polyploidy in the evolution of the genus.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 301–310.     

Genetic variation in Femeniasia (Compositae, Cardueae), an endemic and endangered monotypic genus from the Balearic Islands (Spain)

Femeniasia balearica is the only representative of its genus and is endemic to a small area on the northern coast of Minorca (Balearic Islands, Spain). The entire range of the species covers only 18 km of coastline. It is extremely rare and classified as endangered in the National Catalogue of Threatened Species and as a priority species in the EU Habitats Directive. We carried out a census of current population size and estimated genetic diversity based on AFLP markers to facilitate conservation of this unique species. In the 66 individuals analysed (∼10% of population), 225 bands were scored and the level of diversity was relatively high. Three divergent population groups corresponding to geographical areas (Western, Central and Eastern) were identified. In AMOVA and Bayesian analyses, most of the diversity was found within populations but there was strong differentiation between the three population groups. Genetic and geographical distances between the populations were strongly correlated. Our results show that the populations of F. balearica are not genetically depauperate, in spite of their small sizes. Unexpectedly, our results suggest that as many as three MUs (management units) should be recognized in this very small area, because the low levels of gene flow among them indicate contemporary demographic independence.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 97–107.