Growth rules based on the modularity of the Canarian Aeoniwm (Crassulaceae) and their phylogenetic value

Growth forms of 22 species of Aeonium (Crassulaceae) were quantified. Since all species are simple in their modular construction, models were developed to predict module length, branching mode and flowering probability using linear and logistic regression. When combined, the parameters of these models are species specific. A discriminant analysis generates a statistically significant separation of species at the level of phylogenetic sections. The results therefore demonstrate the phylogenetic value of growth rules in plants. This dynamic approach strongly contrasts with the traditional static view on forms in systematics and morphology. It also leaves scope for predicting the evolutionary pathways of morphological change which have caused the great diversity of growth forms in the genus Aeonium.     

The evolution of growth-forms in the Macaronesian genus Aeonium (Crassulaceae) inferred from chloroplast DNA RFLPs and morphology

Phylogenetic relationships among species of Aeonium were studied using morphological characters and chloroplast DNA RFLPs. Cladistic analysis of weighted morphological data indicates that the small, herbaceous and least woody species are basal in the genus. Chloroplast DNA data gave similar results, supporting the separation of the herbaceous or small, woody species from the large, hapaxanth rosettes, rosette trees, and branched subshrubs with yellow, white or red flowers as well as the only (herbaceous) species with axillary inflorescences. The relationships among the species descending from a polytomy that comprises all species of the genus as well as a polytomy which comprises 18 of the 26 species studied, are only very incompletely resolved, except for two monophyletic clades that contain the branched subshrubs with yellow flowers ( A . sect. Aeonium ) and the branched subshrubs and rosette trees with white or red flowers ( A . sect. Leuconium ), respectively. Cladistic analysis of the combined morphological and chloroplast DNA data improved resolution considerably. Four monophyletic clades are distinguished, each of which, except for three species, comprises only one of the five main growth-form types. Although Aeonium is generally regarded as an outstanding example of adaptive radiation, this mode of speciation seems to have been of minor significance in the evolution of the genus, because each growth-form apparently evolved only once. Instead, island speciation in the absence of major ecological shifts, is probably more important in the evolution of the genus.     

Macaronesian millipedes (Diplopoda) with emphasis on endemic species swarms on Madeira and the Canary Islands

Endemic species swarms constitute large fractions of the millipede faunas of Madeira (29 species of the Cylindroiulus madeirae group, plus six species of Acipes , out of a total of 60 species) and the Canary Islands (46 species of Dolichoiulus , plus four species of the Glomeris alluaudi-group , out of a total of about 79 species). The poorer faunas of the Azores (22 species) and the Cape Verde Islands (15 species) in contrast only include a few endemics. The Cylindroiulus madeirae group and Dolichoiulus show a high degree of diversity of structure (size, colour, leg length etc.) and habitat (laurisilva, xeric habitats, caves). The C. madeirae group, unlike Dolichoiulus , is strongly concentrated in the laurisilva. In this habitat, microhabitat differentiation is pronounced in both swarms.     

Mitochondrial DNA, ecology and morphology: interpreting the phylogeography of the Nesotes (Coleoptera: Tenebrionidae) of Gran Canaria (Canary Islands)

The genus Nesotes (Coleoptera: Tenebrionidae) is represented in the Canary Islands by 19 endemic species, the majority of which are single island endemics. Nesotes conformis and N. fusculus are described on four and three islands, respectively, but each forms a paraphyletic assemblage between Gran Canaria and the other islands. The other described species for Gran Canaria are N. quadratus, N. lindbergi and N. piliger. Thirty-six individuals representing the five species on Gran Canaria have been sequenced for 675 bp of the mitochondrial DNA (mtDNA) cytochrome oxidase II gene. Neighbour-joining analysis of maximum likelihood distances resulted in five distinct mtDNA lineages for N. quadratus, two of which also include mitotypes of N. conformis. Each of the other three species is found on only one mtDNA lineage. We propose from the molecular data that differentiation in a widespread N. quadratus-type ancestor was followed by morphological adaptation to coastal, pine and laurel forest habitats.     

Genome size variation in Macaronesian angiosperms: forty percent of the Canarian endemic flora completed

Genome sizes for 127 Macaronesian endemic angiosperms from 69 genera and 32 families were estimated using propidium iodide flow cytometry. Only about 30-fold variation in 1C-values was found, ranging from 0.32 pg in Echium bonnetii to 9.52 pg in Scilla dasyantha. Taxa with very small DNA amounts (1C 1.4 pg) were the most dominant group (71.7%), whereas the frequency of other categories was much lower (18.9% and 9.4% in taxa with small (1.41–3.50 pg) and intermediate 1C-values (3.51–14.00 pg), respectively). Comparisons of average C- and Cx-values between Macaronesian endemics and non-Macaronesian representatives always revealed significantly smaller amounts in the former group at various taxonomic levels (genus, family, major phylogenetic lineage). Potential relationship between nuclear DNA content and insular burst of speciation is suggested owing to the marked prevalence of very small genomes among angiosperms that underwent rapid adaptive radiation. Merging all the genome size data on Macaronesian angiosperms available shows that this flora represents the best covered plant assemblage from the phytogeographic point of view.     

Spot variation in Maniola jurtina (L.) (Lepidoptera: Satyridae) in southern Portugal and a comparison with the Canary Islands

A sample of 2013 meadow brown butterflies, Maniola jurtina , was obtained in the Algarve, southern Portugal, on behalf of E.B. Ford, the founder of ecological genetics. Variation in the meadow brown occupied Ford's attention for more than 40 years. Ford died in 1988; most of his collection had earlier been destroyed, but the meadow browns remained and became available for analysis. The butterflies were scored for spotting characters on both fore- and hindwings. The population is homogeneous over the entire area of the Algarve and there is no conclusive evidence of a difference between the two years of collection (1986 and 1987). It is probably also homogeneous with southern Iberia in general, an area unique in the western Palaearctic range of the species as it is characterized by a high spot average in males and a low spot average in females. Algarve butterflies differ markedly from those of the Canary Islands, where there is considerable inter-island heterogeneity, supporting the view that the Canary Island populations did not originate from southern Iberia. In the Algarve, many females aestivate and by September there is an essentially all-female population. These are low-spotted and differ significantly from females active in May and June at the start of the annual (one generation) flight season.     

A new species of the wild dragon tree, Dracaena (Dracaenaceae) from Gran Canaria and its taxonomic and biogeographic implications

The arborescent taxa of Dracaena which form the dragon tree group comprise five species found in Macaronesia, Morocco (D. draco), East Africa (D. ombet, D. schizantha), Arabia (D. serrulata) and the island of Socotra (D. cinnabari). A new species of dragon tree, Dracaena tamaranae A. Marrero, R. S. Almeida & M. Gonzalez-Martin, is described from Gran Canaria, Canary Islands. This new species differs from D. draco, the only other Dracaena species currently known in Macaronesia, in having a growth form and inflorescence type and leaves more similar to the East African and Arabian species of Dracaena. In contrast, D. draco appears to be related to D. cinnabari. In this paper, we also present a study of the taxonomy, habitat and ecology of all the species of the dragon tree group. These are found in thermo-sclerophyllous plant communities of tropical-subtropical regions which are rather xerophilous and have a rainfall range of 200–500 mm. Our study indicates two independent colonization events for Dracaena in Macaronesia. In addition, we suggest that the dragon tree group provides an example of two major biogeographical disjunctions between East and West Africa. We postulate that this group has a Tethyan origin, a hypothesis supported by fossil and palaeoclimatic data, and thus parallels the distribution and dispersal pattern of other taxonomic groups.     

Microsatellite loci isolation in the endangered Gran Canarian blue chaffinch (Fringilla teydea polatzeki) and their utility in closely related taxa

The blue chaffinch, Fringilla teydea, is an endemic species of the Canary Islands. This species is formed by two subspecies: The Teneriffean blue chaffinch (F. t. teydea), and the endangered Gran Canarian blue chaffinch, (F. t. polatzeki). Here we report the isolation and characterization of nine tetranucleotide microsatellites (AAAG and AAAT) from the Gran Canarian subspecies, using an enrichment protocol. An average of 7.8 alleles per locus and an average observed heterozygosity of 0.773 were found (n = 28). The loci were tested for their ability to cross amplify in the Teneriffean subspecies and in the common chaffinch (Fringilla coelebs). These microsatellites will be used to manage a captive breeding programme for the endangered Gran Canarian subspecies.     

The phylogenetic relationships and evolution of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae): Evidence frommatK and ITS sequences, and floral morphology and anatomy

The phylogenetic relationships of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae) are investigated through a cladistic analysis of sequence variation of parts of the chloroplast genematK and the ITS region of nuclear ribosomal DNA. The floral anatomical and morphological characteristics ofIxanthus based on paraffin sections and SEM studies are also discussed. In the molecular analysesIxanthus is part of a clade of mostly temperate Erythraeinae and Chironiinae sensu Gilg, in which it is sister to the mostly Mediterranean annualBlackstonia. This relationship is supported by farreaching similarities in flower morphology and anatomy, and the shared possession of at least some perfoliate leaves. This finding contradicts most former classifications in whichIxanthus was placed either in subtribe Gentianinae which never includedBlackstonia, or in a subtribe of its own which was suggested to connect woody tropical with herbaceous temperate taxa. The phylogenetic relationships found here show that 1) the sister group ofIxanthus is distributed in the Mediterranean area, and that 2) its habit as a basally woody suffrutescent herb is best interpreted as an advanced character state.     

Pollen morphology of endemic species of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex (Asteraceae: Anthemideae) in the Canary Islands (Spain), and its taxonomical implications

The pollen of nine endemic taxa of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex and of T. parthenium L. (introduced species from the Canary Islands) was studied using light and scanning electron microscopy with the aim of extending the knowledge of their morphology and to investigate whether pollen morphology may help clarifying the controversial taxonomy of these taxa. Twenty six populations in the Canary Islands (Spain) were studied and the following parameters were measured: polar axis (P), equatorial diameter (E), polar exine thickness (Ex1), length (L) and thickness (A) of the spines in the meridional optical section, exine thickness at the centre of the mesocolpium in the interapertural area (Ex2) and distance between spines (De) in the equatorial optical section. The studies show that the pollen of the three genera is very similar regarding shape, apertures, and ornamentation, but may be distinguished by size and exine thickness.     

Spatial and temporal variation in distribution of Gelidium canariensis (Rhodophyta) from natural populations of the Canary Islands

This study was designed to investigate spatial and temporal variation in Gelidium canariensis populations at two shores in northern Gran Canaria during two years. Spatial scales ranged from some hundred meters (distance between shores), 10 to 30 m (distance between plots) to less than 3 m (distance between quadrats). Gelidium individuals were defined as distinct Gelidium clumps. The results show a significant difference in size of clumps between shores, but not on the smaller spatial scales. No significant temporal variation was found. There was no significant temporal or spatial variation in standing crop or density (counts made in quadrats where Gelidium was present, rather than counts for the total shore). Sporophytic and gametophytic clumps were also distinguished by identifying reproductive structures in the field. The total proportion of sporophytes was larger than the proportion of gametophytes, but at a smaller scale there could be a shift in dominance. The survival rate of clumps was similar between shores with a mean survival rate of 85%, but there was a significant difference in recruitment between shores. The results indicate a stable population structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Patterns of genetic variation in the adaptive radiation of New World crossbills (Aves: Loxia)

Incipient species groups or young adaptive radiations such as crossbills (Aves: Loxia) present the opportunity to investigate directly the processes occurring during speciation. New World crossbills include white-winged crossbills (Loxia leucoptera), Hispaniolan crossbills (Loxia megaplaga), and red crossbills (Loxia curvirostra complex), the last of which is comprised of at least nine morphologically and vocally differentiated forms ('call types') where divergent natural selection for specialization on different conifer resources has been strongly implicated as driving diversification. Here we use amplified fragment length polymorphism (AFLP) markers to investigate patterns of genetic variation across populations, call types, and species of New World crossbills. Tree-based analyses using 440 AFLP loci reveal strongly supported clustering of the formally recognized species, but did not separate individuals from the eight call types in the red crossbill complex, consistent with recent divergence and ongoing gene flow. Analyses of genetic differentiation based on inferred allele frequency variation however, reveal subtle but significant levels of genetic differentiation among the different call types of the complex and indicate that between call-type differentiation is greater than that found among different geographic locations within call types. Interpreted in light of evidence of divergent natural selection and strong premating reproductive isolation, the observed genetic differentiation suggests restricted gene flow among sympatric call types consistent with the early stages of ecological speciation.     

Comparative chromosomal analysis and phylogeny in four Ctenomys species (Rodentia, Octodontidae)

A cytogenetic chromosome study was carried out on specimens of four species of Ctenomys – C. talarum, C. rionegrensis, C. pearsoni and C. dorbignyi –from 10 different populations. The analysis of chromosomes was performed through sequential uniform stain, G and C-banding, and with restriction enzymes. The results obtained are discussed in relation to phylogeny. Chromosome evolution in the species studied suggests that the chromosome number has increased due to fissions, and that a reduction of the amount of constitutive heterochromatin has occurred. Different types of heterochromatin, with different patterns, have been added in parallel during evolution. Some taxonomic suggestions can be deduced from this cytogenetic study.     

Characterization of ten polymorphic microsatellite loci for the threatened species Grimmia curviseta Bouman (Grimmiaceae,Musci)

Grimmia curviseta Bouman is a moss species endemic to the Canary Islands in the North Atlantic Ocean, which has a very restricted distribution across the summit areas of Tenerife and La Palma islands. Using massive sequencing, we developed ten polymorphic microsatellite markers for this species. The pattern of microsatellite alleles per locus provides preliminary evidence that G. curviseta is allodiploid. The average number of alleles per locus ranged from 2 to 7, and observed heterozygosities varied from 0.022 to 1.000. A significant genetic differentiation was observed between the Tenerife and La Palma populations, suggesting that there are some limitations to dispersal. This set of microsatellites constitutes a valuable tool to investigate patterns of genetic diversity across the distribution range of G. curviseta, information that may eventually be used to establish conservation strategies for the species. Owing to the cross-amplification with the closely related species Grimmia montana Bruch & Schimp., our study also demonstrates the utility of these markers for population-level genetic analyses in the genus Grimmia.     

The long-term ecology of the lost forests of La Laguna, Tenerife (Canary Islands)

Aim We report the first analysis of the long‐term ecology of Tenerife, in order to establish a pre‐colonization base‐line and to assess the effect of human activity and the role of climatic variation on vegetation during the Late Holocene. Location A former lake bed in the city of La Laguna (Tenerife, Canary Islands, Spain). Methods A sedimentary sequence of over 2 m was obtained from the former lake bed. Fossil pollen and microfossil charcoal concentrations were analysed. Radiocarbon dating of the sequence indicates that it spans approximately the last 4700 years. The pollen diagram was zoned using optimal splitting within psimpoll 4.25. Results Three pollen zones were differentiated: (1) in Zone L1 (c. 4700–2900 cal. yr bp ) a mixed forest was dominated by Quercus, Carpinus, Myrica and Pinus; (2) in Zone L2 (c. 2900–2000 cal. yr bp ) the laurel forest taxa increased, while Pinus, Juniperus and Phoenix declined; and (3) Zone L3 (c. 2000–400 cal. yr bp ) was characterized by the decline of Carpinus and Quercus and the abundance of laurel forest taxa (e.g. Myrica). Neither Carpinus nor Quercus was hitherto considered to be native to the Canary Islands. Their decline started c. 2000 years ago, coinciding with microfossil charcoal evidence of increased burning and with archaeological evidence for the first human settlement on Tenerife. Main conclusions Between c. 4700 and 2000 cal. yr bp , the composition of the forest in the valley of La Laguna was very different from what it is at present. In particular, Quercus and Carpinus appear to have been significant components, alongside components of the present‐day laurel forest, and the native pine (Pinus canariensis) forest and thermophilous woodland were also more prevalent in the region (but probably not within the lake basin itself) until 3000 cal. yr bp . The subsequent decline of Quercus and Carpinus led to the establishment of the present laurel forest in the region and a shift to more open vegetation types. These changes indicate that the aboriginal inhabitants of the islands, the Guanches, had a far more profound impact on the vegetation of Tenerife than hitherto realized.     

Reconciling gene trees with organism history: the mtDNA phylogeography of three Nesotes species (Coleoptera: Tenebrionidae) on the western Canary Islands

The processes of island colonization and speciation are investigated through mtDNA studies on Canary Island beetles. The genus Nesotes (Coleoptera: Tenebrionidae) is represented by 19 endemic species on the Canary Islands, the majority of which are single island endemics. Nesotes conformis is the most widespread, occurring on Gran Canaria, Tenerife, La Palma and El Hierro. Nesotes conformis forms a paraphyletic assemblage, with a split between Gran Canaria and the other three islands. Nesotes conformis of the western Canary Islands cluster with Nesotes altivagans and Nesotes elliptipennis from Tenerife. Fifty‐two individuals from this western islands species complex have been sequenced for 675 base pairs of the mtDNA cytochrome oxidase II gene, representing Tenerife, La Palma and El Hierro. A neighbour joining analysis of maximum likelihood distances resulted in three distinct mtDNA lineages for N. conformis, two of which also include mitotypes of N. altivagans and N. elliptipennis. Through application of parametric bootstrap tests, we are able to reject hypotheses of monophyly for both N. conformis and N. altivagans. Nesotes altivagans and N. elliptipennis are poorly separated morphologically and mtDNA sequence data adds support to this being one species with a highly variable morphology. We propose that N. altivagans/N. elliptipennis is recently derived from two ancestral mtDNA lineages within N. conformis from the Teno region of Tenerife. We further propose colonization of the younger islands of La Palma and El Hierro by N. conformis from a mitochondrial lineage within the Teno massif (colonization; diversification; mitochondrial DNA; Canary Islands; Coleoptera).     

Phylogeography of the endangered darkling beetle species of Pimelia endemic to Gran Canaria (Canary Islands)

Phylogenetic and geographical nested clade analysis (NCA) methods were applied to mitochondrial DNA sequences of Pimelia darkling beetles (Coleoptera, Tenebrionidae) endemic to Gran Canaria, an island in the Canary archipelago. The three species P. granulicollis, P. estevezi and P. sparsa occur on the island, the latter with three recognized subspecies. Another species, P. fernandezlopezi (endemic to the island of La Gomera) is a close relative of P. granulicollis based on partial Cytochrome Oxidase I mtDNA sequences obtained in a previous study. Some of these beetles are endangered, so phylogeographical structure within species and populations can help to define conservation priorities. A total of about 700 bp of Cytochrome Oxidase II were examined in 18 populations and up to 75 individuals excluding outgroups. Among them, 22 haplotypes were exclusive to P. granulicollis and P. estevezi and 31 were from P. sparsa. Phylogenetic analysis points to the paraphyly of Gran Canarian Pimelia, as the La Gomera P. fernandezlopezi haplotypes are included in them, and reciprocal monophyly of two species groups: one constituted by P. granulicollis, P. estevezi and P. fernandezlopezi (subgenus Aphanaspis), and the other by P. sparsa'sensu lato'. The two species groups show a remarkably high mtDNA divergence. Within P. sparsa, different analyses all reveal a common result, i.e. conflict between current subspecific taxonomic designations and evolutionary units, while P. estevezi and P. fernandezlopezi are very close to P. granulicollis measured at the mtDNA level. Geographical NCA identifies several cases of nonrandom associations between haplotypes and geography that may be caused by allopatric fragmentation of populations with some cases of restriction of gene flow or range expansion. Analyses of molecular variance and geographical NCA allow definition of evolutionary units for conservation purposes in both species-groups and suggest scenarios in which vicariance caused by geological history of the island may have shaped the pattern of the mitochondrial genetic diversity of these beetles.