Nuclear DNA amounts in Macaronesian angiosperms

Nuclear DNA contents for 104 Macaronesian angiosperms, with particular attention on Canary Islands endemics, were analysed using propidium iodide flow cytometry. Prime estimates for more than one-sixth of the whole Canarian endemic flora (including representatives of 11 endemic genera) were obtained. The resulting 1C DNA values ranged from 0.19 to 7.21 pg for Descurainia bourgeauana and Argyranthemum frutescens, respectively (about 38-fold difference). The majority of species, however, possessed (very) small genomes, with C-values <1.6 pg. The tendency towards small nuclear DNA contents and genome sizes was confirmed by comparing average values for Macaronesian and non-Macaronesian representatives of individual families, genera and major phylogenetic lineages. Our data support the hypothesis that the insular selection pressures in Macaronesia favour small C-values and genome sizes. Both positive and negative correlations between infrageneric nuclear DNA amount variation and environmental conditions on Tenerife were also found in several genera.     

Evolutionary dynamics of satellite DNA family PIM357 in species of the genus Pimelia (Tenebrionidae,Coleoptera)

A large number of repeats of a satellite DNA (stDNA) family have been cloned and sequenced from species and populations of the genus Pimelia (Tenebrionidae, Coleoptera). The beetles were collected in the Canary Islands, Morocco, the Iberian Peninsula, and the Balearic Islands in order to analyze the evolutionary forces and processes acting on abundant stDNAs conserved at the genus level. This repetitive family is composed of an abundant A-T-rich stDNA, with basic units of 357 bp. All the sequences obtained showed similarity to the 22 repeat units of the PIM357 stDNA family described previously for six Iberian Pimelia species (Pons et al. 1997 ). An analysis based on similarity shows the presence of three different groups of sequences clearly in accordance with their geographical origin. One is composed of satellite sequences from Iberian and Balearic species, a second group from the Moroccan taxa, whereas the third one is from the Pimelia species endemic to the Canary Islands. The latter group shows higher nucleotide diversities for their stDNA sequences and a lack of relationship between transition stages to fixation and sequence divergence. Phylogeographic data of Canarian Pimelia show that the PIM357 stDNA family has persisted for more than 8 Myr and could probably be traced to the origin of the lineage. The data suggest that distinct demographic and phylogenetic patterns related to the colonization of the volcanic Canarian island chain account for particular evolutionary dynamics of the repeat DNA family in this group.     

Biogeographical and evolutionary patterns in the Macaronesian shield-backed katydid genus Calliphona Krauss, 1892 (Orthoptera: Tettigoniidae) and allies as inferred from phylogenetic analyses of multiple mitochondrial genes

Abstract.  We investigated the phylogenetic patterns, evolutionary processes, and their taxonomic implications, of two closely related shield-backed katydid genera endemic to the Macaronesian archipelagos: the monotypic Psalmatophanes Chopard, 1938 endemic to Madeira and Calliphona Krauss, 1892, which includes three species restricted to the Canary Islands. Two main hypotheses have been proposed to explain the origin and colonization pathways of these two genera: a single origin with subsequent sequential colonization of the islands, or three independent colonization waves from continental Africa. We used DNA sequence information from the mitochondrial genes cox1, tRNAleucine, rrnL and nad1 to infer phylogenetic relationships among Psalmatophanes and Calliphona species. Our results provide support for the independent colonization of Madeira and the Canary Islands, and suggest that Psalmatophanes is actually more closely related to the continental genus Tettigonia than to the Canarian representatives. Deep genetic divergence among Canarian species provides further support for the assignment of the Canarian species into two subgenera. Tree topology along with Bayesian-based estimates of lineage age suggest a pattern of colonization from Tenerife to La Palma, and from Tenerife to Gran Canaria with subsequent dispersal to La Gomera. We report the first collection of a Calliphona specimen in the island of El Hierro, which molecular data suggest is a recent immigrant from La Gomera. We hypothesize that the patterns of distribution and genetic divergence exhibited by Calliphona in the Canary Islands are compatible with a taxon cycle process. Our results have further implications for the higher level phylogeny of the subfamily Tettigoniinae and suggest that some of the tribes as currently delimited may not correspond to natural groups.     

南海岛屿种子植物区系地理的研究

本文通过实地考察,广泛收集前人的研究资料,概述了南海岛屿地区的自然条件和植被,对南海岛屿种子植物的区系组成、特点、分布区类型、特有现象和替代现象等进行了较详细的分析,并与邻近植物区系进行了比较研究。同时,根据区内植物分布的特点和自然条件特征划分为５个植物区系小区,最后对南海岛屿地区植物区系的起源与演化进行了讨论。     

Spatial patterns of the Iberian and Balearic endemic vascular flora

The Iberian flora has a high degree of originality (1328 endemic species, 24% of endemism), comparable to other regions in the Mediterranean Basin. The richness of Iberian endemic species is unevenly distributed; the greatest diversity is found in the main mountain ranges although the southwestern Atlantic coast and specially the Balearic Islands are rich in range-restricted endemic species. The largest number of endemic genera is found in the northwestern mountains, which might have acted as a refugium area. The Baetic System, which includes nearly half (46%) of the total Iberian endemic species, is by far the richest region of the territory. Its endemic flora is characterized by the great richness of narrow endemics and the high species turnover rate. The k-means partitioning analysis enables us to identify 11 units, generally well defined by the natural geographic features. The clusters including the northwestern mountains, the Cantabrian Mountains, the southwestern coast and especially the Balearic Islands, the Pyrenees and the Baetic System are compact and consist of a high proportion of diagnostic species, and can therefore be considered areas of endemism on a large scale. The regionalization reflects a primary longitudinal division of Iberia between a basic eastern and an acidic western region, but also partly reveals a climatic division between Eurosiberian and Mediterranean regions. Southeastern Iberia seems to be an important center of differentiation for several typically Mediterranean genera (e.g. Centaurea, Linaria, Armeria, Teucrium and Thymus), but other large genera are also highly diversified.     

Origin and evolution of the endemic genera of Gonosperminae (Asteraceae: Anthemideae) from the Canary Islands: evidence from nucleotide sequences of the internal transcribed spacers of the nuclear ribosomal DNA

The Gonosperminae (Asteraceae) are composed of three genera endemic to the Canary Islands (GONOSPERMUM: Less., and LUGOA: DC.) and southern Africa (INULANTHERA: K?llersj?), and they are considered an example of a floristic link between these two regions. Phylogenetic analyses of ITS sequences reveal that the Canarian genera are not sister to INULANTHERA: and do not support the monophyly of the Gonosperminae. These results, coupled with previous phylogenetic studies of other groups, suggest that many of the putative biogeographic links between Macaronesia and southeast Africa need to be evaluated by rigorous phylogenetic analyses. INULANTHERA: forms part of the basal southern African radiation of the Anthemideae, and therefore it is closely related to other taxa from this region. Maximum likelihood and weighted parsimony analyses support a monophyletic group in the Canary Islands, that includes LUGOA:, Gonospermum, and three TANACETUM: species endemic to the island of Gran Canaria. Bootstrap support for the monophyly of this Canarian group is weak, and it collapses in the strict consensus tree based on unweighted parsimony. LUGOA: is nested within Gonospermum, and both interisland colonization among the western islands of La Gomera, El Hierro, La Palma and Tenerife, and radiation on the central island of Gran Canaria have been the major patterns of species diversification for these Canarian endemics.     

Opportunistic nectar-feeding birds are effective pollinators of bird-flowers from Canary Islands: experimental evidence from Isoplexis canariensis (Scrophulariaceae)

Insular floras, characterized by simple pollination networks, sometimes include novel mutualistic agents such as nonspecialist nectarivores. In this study we confirmed the effective pollination of Isoplexis canariensis by opportunistic nectar-feeding birds in Tenerife, Canary Islands, Spain. This plant is among the ornithophilous species of the Canarian flora that lack past and present specialist nectarivorous birds. Experimental hand pollinations revealed self-compatibility, but cross-pollinated flowers produced a greater percentage of viable seeds than self-pollinated ones. Flowers were visited by five species of birds (Phylloscopus canariensis, Parus caeruleus, Sylvia melanocephala, Serinus canarius, and Fringilla coelebs) and by the endemic lizard (Gallotia galloti, Lacertidae). Insect pollination was absent, and the few insect visitors acted as nectar thieves or secondary nectar robbers. Birds represented 93.1% of total visits, with the Canarian Chiffchaff, Ph. canariensis, being the most frequent visitor. Flowers visited by birds set more, larger, and heavier fruit than flowers from which birds were excluded. Bird visitation also enhanced seed viability. These results demonstrate the active role of these opportunistic birds as effective pollinators of this Canarian bird-flower species. Further, the results reveal the need to consider the effect of these birds on the evolution of ornithophilous floral traits in absence of specialist nectarivores.     

Using taxonomic and phylogenetic evenness to compare diversification in two Island Floras

This study compares the phylogenetic structure in the Canary Islands and Hawaii by means of the distributions of the species number for plant families (Taxonomic evenness) and lineages (Phylogenetic evenness) across archipelagos and across habitats in both archipelagos using the Gini coefficient. We then investigate phylogenies to identify particular habitats contributing to such differences using Taxonomic distinctness (AvTD) and its variation (VarTD).Our results show that the distribution of species number among Hawaiian lineages is much more uneven than the Canary Islands. In contrast, Hawaii produces a more even distribution of species number by family than the Canary Islands. This may be due to the Hawaiian Flora being derived from considerably fewer colonists than the Canarian Flora as a result of its much greater degree of isolation. At the same time, Hawaii is represented by the same number of families as the Canary Islands. This may stem from Hawaii's flora being derived from a greater range of source areas despite its isolation. Finally, there is much more diversification spread across a larger number of lineages in Hawaii. The higher degree of Hawaiian diversification may be due to a greater range of habitats, more diverse and phylogenetically distinct floristic sources, and low initial species diversity resulting from extreme isolation.Two Canarian habitats (Rock communities and Thermophilous habitats) and one Hawaiian habitat (Wet communities) contribute to the differences in phylogenetic structure between the two archipelagos. These habitats exhibit disproportionate levels of unevenness and may represent centres of diversification. We propose a combination of two habitat properties, high receptivity and low stability, to explain these results.     

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.     

The Azores diversity enigma: why are there so few Azorean endemic flowering plants and why are they so widespread?

Aim Endemism in the flora of the Azores is high (33%) but in other respects, notably the paucity of evolutionary radiations and the widespread distribution of most endemics, the flora differs markedly from the floras of the other Macaronesian archipelagos. We evaluate hypotheses to explain the distinctive patterns observed in the Azorean endemic flora, focusing particularly on comparisons with the Canary Islands. Location Azores archipelago. Methods Data on the distribution and ecology of Azorean endemic flowering plants are reviewed to ascertain the incidence of inter‐island allopatric speciation and adaptive, ecological speciation. These are contrasted with patterns for the Canary Islands. Patterns of endemism in the Azores and Canaries are further investigated in a phylogenetic context in relation to island age. beast was used to analyse a published molecular dataset for Pericallis (Asteraceae) and to investigate the relative ages of Azorean and Canarian lineages. Results There are few examples of inter‐island allopatric speciation in the Azorean flora, despite the considerable distances between islands and sub‐archipelagos. In contrast, inter‐island allopatric speciation has been an important process in the evolution of the Canary Islands flora. Phylogenetic data suggest that Azorean endemic lineages are not necessarily recent in origin. Furthermore, in Pericallis the divergence of the Azorean endemic lineage from its closest relative pre‐dates the radiation of a Canarian herbaceous clade by inter‐island allopatric speciation. Main conclusions The data presented do not support suggestions that hypotheses pertaining to island age, age of endemic lineages and ecological diversity considered individually explain the lack of radiations and the widespread distribution of Azorean endemics. We suggest that palaeoclimatic variation, a factor rarely considered in macroecological studies of island diversity patterns, may be an important factor. Palaeoclimatic data suggest frequent and abrupt transitions between humid and arid conditions in the Canaries during the late Quaternary, and such an unstable climate may have driven the recent diversification of the flora by inter‐island allopatric speciation, a process largely absent from the climatically more stable Azores. Further phylogenetic/phylogeographic analyses are necessary to determine the relative importance of palaeoclimate and other factors in generating the patterns observed.     

Plant genetic diversity in the Canary Islands: a conservation perspective

The Canary Islands are an Atlantic volcanic archipelago with a rich flora of ～570 endemic species. The endemics represent ～40% of the native flora of the islands, and ～20% of the endemics are in the E (endangered) category of the International Union for Conservation of Nature. A review of allozyme variation in 69 endemic species belonging to 18 genera and eight families is presented. The average species-level genetic diversity (H(T)) at allozyme loci is 0.186, which is twice as high as the mean reported for endemics of Pacific archipelagos. Possible factors contributing to this higher diversity are discussed, but the reasons remain obscure. An average of 28% of the allozyme diversity within species resides among populations, indicating a high level of interpopulational differentiation. Studies of reproductive biology indicate that many of the endemic species are outcrossers. The high total diversity within species, the relatively high differentiation among populations, and the outcrossing breeding systems have implications for species conservation. Decreased population sizes in outcrossing species would promote biparental inbreeding and increase inbreeding depression. The relatively high proportion of allozyme diversity among populations indicates that the most effective strategy for preserving genetic variation in species is to conserve as many populations as possible. The genetic diversity in many Canary Island endemics is endangered by: (1) overgrazing by introduced animals, such as barbary sheep, goats, mouflons, rabbits, and sheep; (2) interspecific hybridization following habitat disturbance or planting of endemics along roadsides or in public gardens; (3) competition with alien plant species; and (4) decline of population size because of urban development and farming.     

南岳衡山蕨类植物区系研究

南岳衡山蕨类植物共计235种(含种下等级),隶属于36科75属。区系分析表明:该地蕨类植物具有较强的热带性质,同时也显示出该区蕨类植物区系向温带渗透和过渡的性质,科、属的分布区类型均以热带、亚热带分布型为主,而种以温带分布型为主。热带、亚热带分布型在该区有18科38属69种,分别占科、属、种数的50.00%、50.67%和29.36%;其中泛热带分布型有14科21属5种,分别占科、属、种数的38.89%、28.00%和2.13%;温带性分布型有3科18属158种,分别占科、属、种数的8.33%、24.00%和67.23%。表明泛热带分布型是该区科、属最重要的分布类型;在种的区系上,温带性成分占明显优势,反映了该区热带成分与温带成分的交汇与兼容。该区虽没有自己的特有属,但具1种特有种,而且中国特有种丰富,共计46种,占总种数的19.57%。该区系与周邻联系广泛,地理成分以华东成分为主。在区系关系上,该区与武夷山、庐山的关系密切,而与万佛山的关系较为疏远。     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Are island plant communities more invaded than their mainland counterparts?

Questions: Are island vegetation communities more invaded than their mainland counterparts? Is this pattern consistent among community types? Location: The coastal provinces of Catalonia and the para‐oceanic Balearic Islands, both in NE Spain. These islands were connected to the continent more than 5.35 million years ago and are now located <200 km from the coast. Methods: We compiled a database of almost 3000 phytosociological relevés from the Balearic Islands and Catalonia and compared the level of invasion by alien plants in island versus mainland communities. Twenty distinct plant community types were compared between island and mainland counterparts. Results: The percentage of plots with alien species, number, percentage and cover percentage of alien species per plot was greater in Catalonia than in the Balearic Islands in most communities. Overall, across communities, more alien species were found in the mainland (53) compared to the islands (only nine). Despite these differences, patterns of the level of invasion in communities were highly consistent between the islands and mainland. The most invaded communities were ruderal and riparian. Main conclusion: Our results indicate that para‐oceanic island communities such as the Balearic Islands are less invaded than their mainland counterparts. This difference reflects a smaller regional alien species pool in the Balearic Islands than in the adjacent mainland, probably due to differences in landscape heterogeneity and propagule pressure.     

Different fates of island brooms: contrasting evolution in Adenocarpus, Genista, and Teline (Genisteae, Fabaceae) in the Canary Islands and Madeira

Analysis of sequence data from the internal transcribed spacers (ITS) and 5.8S region of nuclear ribosomal DNA show that Canarian and Madeiran brooms (Genisteae) of the genera Teline, Adenocarpus, and Genista are related to Mediterranean species and not to species from adjacent parts of Morocco. Each separate colonization of the islands has resulted in contrasting patterns of adaptation and radiation. The genus Teline is polyphyletic, with both groups (the "T. monspessulana group" and the "T. linifolia group") separately nested within Genista. Genista benehoavensis (La Palma) and G. tenera (Madeira) form, with G. tinctoria of Europe, a single clade characterized by vestigially arillate seeds. The Canarian species of Adenocarpus have almost identical sequence to the Mediterranean A. complicatus and are likely to be the result of island speciation after a very recent colonization event. This Canarian/Mediterranean A. complicatus group is sister to the afrotropical montane A. mannii which is probably derived from an earlier colonization from the Mediterranean, possibly via the Red Sea hills. The independent colonization and subsequent radiation of the two Teline groups in the Canary Islands make an interesting comparison: the phylogenies both show geographical structuring, each with a central and western island division of taxa. Within the "T. monspessulana group" there is some evidence that both continental and Madeiran taxa could be derived from the Canary Islands, although it is likely that near contemporaneous speciation occurred via rapid colonization of the mainland and islands. The finding of two groups within Teline also has implications for patterns of hybridization in those parts of the world where Teline species are invasive; in California members of the T. monspessulana group hybridize readily, but no hybrids have been recorded with T. linifolia which has been introduced in the same areas.     

我国南沙群岛的植物与植被概况

南沙群岛是我国最南端的一个植物地理区域,但目前我国植物学家对这里的植物与植被还了解得不多。本文通过实地考察个别人工岛,并广泛收集国内外的有关文献资料,概述了南沙群岛的植物与植被的组成、特点及其与邻近地区植物区系的关系,为我国南海岛屿植物区系的研究提供基本资料．     

华中地区藤本种子植物区系的研究

以湖南和湖北两省为代表对华中地区藤本种子植物的区系进行了研究。本区共有藤本种子植物62科175属838种,其中土著种类有60科159属784种。在科、属、种的水平上对藤本种子植物区系特性进行了较深入的统计和分析,并对本区藤本种子植物与邻近地区的关系、本区内的藤本种子植物的地带性分异进行了探讨。统计表明,本区藤本种子植物种数占该区种子植物总种数的11%,且61.7%的种类主要集中在30种以上的大科中;热带分布型明显多于温带分布型;泛热带、热带亚洲、东亚（中国）特有和东亚—北美间断分布科属比较集中,其中许多属为古老成分;本区藤本种子植物特有成分丰富,计有东亚特有科1科、中国特有属6属、中国特有种507种（其中华中地区特有种35种）。通过研究认为：本区藤本种子植物具有种类丰富、分布类型多样、特有成分较多、区系较古老的特点;与本区的全部种子植物区系相比,藤本种子植物区系的热带性更强,具有明显的热带亲缘性,并与喜马拉雅及中国西南有较强的地理渊源。     

中国森林种子植物区系的特征

本文主要根据中国种子植物区系研究的最新资料,简要介绍中国现代森林种子植物区系的多样性、地理成分、区系性质及地区差异等特征,供森林研究和中国生物多样性研究参考。     

四川被子植物区系特征的初步研究

四川地区被子植物区系188科1493属8711种及1252变种,全球50％以上的科都有代表,与毗邻地区及世界植物区系有着广泛而密切的联系。区系古老而原始,珍稀孑遗类群丰富;单型或少种的科属众多,含100种以上的大科和大属也十分繁荣;含5种以下的属占76.89%,但占总属数约22.64%的多种属和大属是区系的主体,包含区系总种数的63.28%。科的分布类型有明显热带性质,属、种的分布类型反映出典型的亚热带属性。优势现象明显,特有类群和表征类群丰富。该地区被子植物区系对中国乃至东亚都有代表性。