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.     

Thoughts on the ecology and evolution of the intertidal biota of the Azores and other Atlantic islands

The special features of the intertidal ecosystems of remote islands are reviewed briefly before focusing on the littoral zone of Macaronesia (Azores, Madeira and Canaries). Distribution patterns are briefly compared with other European shores. Species missing from the Azores compared with Madeira, the Canaries and continental European shores are listed. The degree of genetic differentiation within and amongst selected important species of gastropods (Patella spp.) and barnacles (Chthamalus spp.) in Macaronesia is reviewed. The role of isolated islands in allopatric speciation of these organisms is discussed in relation to dispersal. Possible threats to the populations and communities of rocky shores in Macaronesia are considered in the context of the endemic nature of certain species. The importance of studies of basic ecology and genetics to inform resource management and conservation is highlighted. Future research directions are indicated, emphasizing the usefulness of Macaronesia as a model system to explore speciation in intertidal organisms.     

Additive partitioning as a tool for investigating the flora diversity in oceanic archipelagos

This paper introduces the integration of additive partitioning with species—area relationships to island biogeography in order to address the question “How are the pteridophyte and spermatophyte native and endemic flora of different oceanic archipelagos partitioned across islands?”.Species richness data of all endemic species and all native species of pteridophytes and spermatophytes were obtained for the Azores, Canaries and Cape Verde in the Atlantic Ocean and Galápagos, Hawaii and Marquesas in the Pacific Ocean. Additive partitioning of species diversity was used to quantify how much of the total diversity of an oceanic archipelago flora (γ-diversity) is due to (i) the mean species richness of the flora of each island (α-diversity), (ii) the variability in species richness of the floras across islands (β_Nestedness) and (iii) the complementarity in species composition of the floras of different islands (β_Replacement). The analysis was separately performed for the native and endemic pteridophyte and spermatophyte floras.The diversity partitioning of the six archipelagos showed large differences in how the flora of each archipelago is partitioned among the α, β_Nestedness and β_Replacement components, for pteridophytes and spermatophytes and for all endemic species and all native species. The α-diversity was more important for all native species than for endemic species and more important for pteridophytes than for spermatophytes, with the Azores showing outstanding high values of α-diversity. The β_Nestedness was higher for pteridophytes than for spermatophytes and higher for endemic species than for all native species in both pteridophytes and spermatophytes. The values of β_Replacement suggested that: (i) the spermatophyte native flora is more differentiated across islands than the pteridophyte native flora and (ii) the pteridophyte endemic flora and, especially, the spermatophyte endemic flora are more differentiated across islands than the corresponding native flora. An outstanding value of β_Replacement for endemic and all native spermatophytes was found in Hawaii, confirming the biogeographical island differentiation in this archipelago.     

Molecular taxonomic identification in the absence of a ‘barcoding gap’: a test with the endemic flora of the Canarian oceanic hotspot

We use a comprehensive subset of Canarian angiosperms corresponding to 23 families, 35 genera and 60 Canarian endemic taxa to test whether this flora is suitable to taxonomic identification with the two proposed plant DNA barcode sequences and whether these sequences may reveal the existence of cryptic species overlooked by morphology. The rate of discrimination success between the insular congeneric samples using the rbcL+matK combination and a ‘character‐based’ approach (where we use only the combination of nucleotide positions in an alignment that allows unambiguous species identification) is higher (82.29%) than that obtained with the ‘distance‐based’ approach (80.20%) used by the CBOL Plant Working Group in 2009 and also when compared with tests conducted in other floras. This suggests that the molecular identification of the Canarian endemic flora can be achieved as successfully as in other floras where the incidence of radiation is not as relevant. The facts that (i) a distance‐based criterion was unable to discriminate between congeneric and conspecific comparisons and (ii) only the character‐based discrimination criterion resolved cases that the distance‐based criterion did not, further support the use of a character discrimination approach for a more efficient DNA barcoding of floras from oceanic islands like the Canaries. Thus, a barcoding gap seems not to be necessary for the correct molecular characterization of the Canarian flora. DNA barcodes also suggest the possible existence of cryptic taxa to be further investigated by morphology and that the current taxonomic status of some of the taxa analysed may need revision.     

Temporal and palaeoclimatic context of the evolution of insular woodiness in the Canary Islands

Insular woodiness (IW), referring to the evolutionary transition from herbaceousness toward woodiness on islands, has arisen more than 30 times on the Canary Islands (Atlantic Ocean). One of the IW hypotheses suggests that drought has been a major driver of wood formation, but we do not know in which palaeoclimatic conditions the insular woody lineages originated. Therefore, we provided an updated review on the presence of IW on the Canaries, reviewed the palaeoclimate, and estimated the timing of origin of woodiness of 24 insular woody lineages that represent a large majority of the insular woody species diversity on the Canaries. Our single, broad‐scale dating analysis shows that woodiness in 60%–65% of the insular woody lineages studied originated within the last 3.2 Myr, during which Mediterranean seasonality (yearly summer droughts) became established on the Canaries. Consequently, our results are consistent with palaeoclimatic aridification as a potential driver of woodiness in a considerable proportion of the insular woody Canary Island lineages. However, the observed pattern between insular woodiness and palaeodrought during the last couple of million years could potentially have emerged as a result of the typically young age of the native insular flora, characterized by a high turnover.     

广东种子植物区系地理成分研究

广东种子植物区系共有２１９科１４３４属４９８６种。其中：①热带区系成分占有较大的比例,约占非世界属的６２．３％,表明广东区系受到热带区系的强烈渗透,次之为亚热带（至亚热带山地）成分约占３２．２％,及中国特有分布占５．６％;②以单型属（７２属）,单种属（５８５属）和寡种属（５６２属）占绝对优势,共占８５．０％,充分显示了广东区系的古老性和过渡性特点;③从区系组成和植被结构的特征成分来看,以华夏植物区系成分为主,包括：亚热带分布,亚热带山地分布,中国特有分布,东亚—北美间断分布,及一些亚洲热带分布中以亚热带为主的属：④在这些属中有东亚特有属１３２属,中国特有属７５属以及华南—西南—中南半岛特有属６８属等;这些属的存在毫无疑问地说明,广东区系是华夏植物区系的核心地区之一,同时也显示了热带区系与亚热带区系有着不可分隔的统一性。     

The endemic vascular flora of Peloritani Mountains (NE Sicily): Plant functional traits and phytogeographical relationships in the most isolated and fragmentary micro-plate of the Alpine orogeny

This study is aimed at (1) producing a complete and updated inventory of the endemic vascular flora of Peloritani Mountains, (2) defining the geographical limits of Peloritani, regarded here as a biogeographical district and (3) highlighting possible paleogeographic connections with other Mediterranean lands. The heterogeneity analysis of the endemic flora was performed by means of contingency tables, through the χ² test. The endemic flora of this area consists of 129 specific and infraspecific taxa, of which 15 are restricted to the Peloritani Mountains. The analysis of habitats revealed that endemic taxa are most abundant on cliffs, rangelands, woods and garrigues. A large number of surveyed taxa are endemic to central-southern Italy and Sicily, while the number of endemic taxa in common with Calabria, Etna and Aeolian Islands turned out to be rather low, in spite of the geographical proximity. The endemic flora of Peloritani allows to emphasize palaeogeographical relationships not only with the neighbouring Mediterranean territories, but also with currently remote ones, such as southern Spain, Sardinia and Corsica. The phytogeographical framework substantiates the hypothesis that the Peloritani floristic district coincides with the limit given to Peloritani Mountains by structural geologists.     

Timing and tempo of early and successive adaptive radiations in Macaronesia

The flora of Macaronesia, which encompasses five Atlantic archipelagos (Azores, Canaries, Madeira, Cape Verde, and Salvage), is exceptionally rich and diverse. Spectacular radiation of numerous endemic plant groups has made the Macaronesian islands an outstanding area for studies of evolution and speciation. Despite intensive investigation in the last 15 years, absolute age and rate of diversification are poorly known for the flora of Macaronesia. Here we report molecular divergence estimates and rates of diversification for five representative, putative rapid radiations of monophyletic endemic plant lineages across the core eudicot clade of flowering plants. Three discrete windows of colonization during the Miocene and early Pliocene are suggested for these lineages, all of which are inferred to have had a single colonization event followed by rapid radiation. Subsequent inter-archipelago dispersal events into Madeira and the Cape Verdes took place very recently during the late Pliocene and Pleistocene after initial diversification on the Canary Islands. The tempo of adaptive radiations differs among the groups, but is relatively rapid compared to continental and other island radiations. Our results demonstrate that opportunity for island colonization and successful radiation may have been constrained to discrete time periods of profound climatic and geological changes in northern African and the Mediterranean.     

罗霄山脉种子植物区系的特有现象与残遗现象

罗霄山脉种子植物区系非常丰富, 是连接华东、华南、华中植物区系的重要通道, 区系组成具有明显的古老性和过渡性特征。本文针对该区域种子植物的特有现象和残遗现象, 分析了其组成特点及形成原因。结果表明: (1)罗霄山脉共有中国特有科3科, 特有属55属, 特有种1,624种, 罗霄山脉区域特有种43种7变种; 特有属中以古特有属为主, 并以温带成分占优势(达55.91%), 体现出明显的古老性和孑遗性。(2)罗霄山脉共有孑遗属165属, 以热带亚洲分布、东亚-北美间断分布、东亚特有及中国特有成分为主, 包括木本属132属。(3)受中新世气候波动以及第四纪冰期的影响, 许多北方热带植物群的常绿成分、北极-第三纪的落叶成分, 在罗霄山脉得以保存下来成为古特有和孑遗成分。特有现象、残遗现象的分析结果表明罗霄山脉是一个重要的生物避难所, 其对中国东部植物区系的保存和重新扩散具有重要意义。     

Marine macroalgal biodiversity hotspots: why is there high species richness and endemism in southern Australian marine benthic flora?

The southern Australian marine macroalgal flora has the highest levels of species richness and endemism of any regional macroalgal flora in the world. Analyses of species composition and distributions for the southern Australian flora have identified four different floristic elements, namely the southern Australian endemic element, the widely distributed temperate element, the tropical element and a cold water element. Within the southern Australian endemic element, four species distribution patterns are apparent, thought to largely result from the Jurassic to Oligocene fragmentation of East Gondwana, the subsequent migration of Tethyan ancestors from the west Australian coast and the later invasion of high latitude Pacific species. Climatic deterioration from the late Eocene to the present is thought responsible for the replacement of the previous tropical south coast flora by an endemic temperate flora which has subsequently diversified in response to fluctuating environmental conditions, abundant rocky substrata and substantial habitat heterogeneity. High levels of endemism are attributed to Australia's long isolation and maintained, as is the high species richness, by the lack of recent mass extinction events. The warm water Leeuwin Current has had profound influence in the region since the Eocene, flowing to disperse macroalgal species onto the south coast as well as ameliorating the local environment. It is now evident that the high species richness and endemism we now observe in the southern Australian marine macroalgal flora can be attributed to a complex interaction of biogeographical, ecological and phylogenetic processes over the last 160 million years.     

中国东北地区植物特有性研究

为了更好地理解植物区系的起源、种系分化及演化进程,推进特有植物保护工作,本文对东北地区内分布的中国特有植物、东北地区特有植物和植物区系地区特有植物的物种组成、科属结构、生活型组成、区系特征和地理分布情况进行了系统研究。结果表明：①研究区域内共有中国特有植物289种、东北地区特有植物109种、长白植物区系地区特有植物154种、大兴安岭植物区系地区特有植物16种、蒙古草原植物区系地区特有植物10种、华北植物区系地区特有植物20种。②各类特有植物中均以草本植物种数最多,其中多年生草本植物占绝对优势,乔木、灌木和藤本相对较少。③中国特有植物集中分布在东北地区大兴安岭山脉和长白山山脉。④东北地区特有植物集中分布地区与东北地区主要山脉走向一致,说明东北地区特有植物以山地起源植物为主。⑤长白植物区系地区植物特有性较强,表现出区系较为成熟的性质。大兴安岭植物区系地区、蒙古草原植物区系地区和华北植物区系地区特有性较弱,表现出区系较年轻的性质。⑥长白植物区系地区特有植物集中分布在吉林长白山山脉和长白山山脉向南延伸的余脉上。大兴安岭植物区系地区特有植物在区内分布较均匀,华北植物区系地区和蒙古草原植物区系地区特有植物在区内分布较零散。     

安徽珍稀濒危植物区系探讨

在深入调查的基础上,对安徽珍稀濒危植物区系组成和地理成分进行了详细的统计分析,探讨了该区系的组成特点及地理分布特征。结果表明,该区系属温带亚热带性植物区系,具有珍稀濒危物种丰富,古老、孑遗植物众多;地理成分复杂,区系联系广泛;过渡性质明显,温带亚热带成分占优;特有现象显著,华东区系特征典型等基本特征。在植物区系亲缘上,该区系隶属东亚植物区、中国-日本森林植物亚区、华东地区,与华中植物区系关系密切,受西南植物区系影响深刻。     

河南种子植物区系地理研究

河南省地处中原 ,属于我国南北过渡和东西过渡的重要区域 ,植物区系成分复杂多样并于周围地区联系广泛 ,对其植物区系研究对深入认识本省的自然环境特征及其在我国多种自然区划中的位置等有着重要意义。该文在最新资料的基础上 ,运用区系学原理对河南省种子植物区系的种类组成、地理成分 (属、种 2个层次上 )等进行了系统的分析 ,在此基础上概括出河南植物区系的基本特征为 :(1 )植物种类比较丰富 ,多样性较高 ;(2 )起源古老 ;(3 )地理成分复杂 ,温带成分略占优势 ,过渡性突出 ;(4 )中国特有植物比较丰富。     

Endemic plants of the arid succulent karoo in Namibia: towards hypotheses for their evolution

This study investigated endemic plants in the Sperrgebiet in the succulent karoo – a global biodiversity hotspot in southern Africa. In order to develop hypotheses regarding the evolution of this endemic flora, the following questions were posed: 1) Are taxonomic patterns similar at different levels of taxonomic order? 2) Are adaptations of Sperrgebiet endemic taxa different from the overall flora? 3) Are adaptations different in related endemic and non‐endemic taxa? There is an over‐representation of species of Mesembryanthemaceae in the endemic flora, which is in accordance with patterns in the succulent karoo biome overall. The nearly 37% contribution of Mesembryanthemaceae to the endemic flora is, however, remarkably higher than elsewhere in the biome. The over‐representation of this family is not maintained at a higher level of taxonomic order, supporting the notion of recent speciation in this family. Regarding plant functional attributes, major differences between the endemic and non‐endemic flora exist. Compact leaf‐succulents with canopy stored, water‐dispersed seeds, as well as bulbs and shrubby leaf‐succulents predominate the endemic flora. Differences in reproductive (seed storage and dispersal distance) and growth form attributes between the endemic and non‐endemic flora were largely influenced by phylogenetic history. Minutism (plant height <10 cm) was favoured by many endemic plants, but this was not only influenced by phylogenetic factors. Two hypotheses emerged from these observations: 1) the evolution of minutism is largely driven by external factors in the Sperrgebiet, and 2) external factors and phylogenetic history are likely of equal importance in the development of the Sperrgebiet endemic flora.     

江西木本植物区系成分及其特征的研究

江西位于我国中亚热带东部,地史悠久,自然地理条件优越,植物种类相当丰富。本文根据笔者历年的调查研究,在前人的基础上探讨了江西木本植物区系的起源和演化过程,该区系主要起源于中生代的古热带植物区系和泛北极植物区系。并对该区系各类地理成分进行了统计和分析,江西的野生木本植物计有102科430属约2000种(含种下单位),木本植物以热带亚洲、泛热带、东亚和北温带分布型成分为主体。将该区系的特征初步归结为如下几点:(1)地理成分复杂,区系联系广泛;(2)区系起源古老,子遗树种众多;(3)热带成分占优势,特有现象较显著;(4)南北区系交汇过渡,具明显的中亚热带华东区系特点;(5)珍稀濒危树种繁多,森林区系急待保护。     

广东种子植物区系科的组成及其特点

广东共有种子植物２１９科１４３４属４９８６种．本文主要对其科的组成进行了统计和分析,结果表明广东植物区系具有明显的过渡性特点,但在北回归线两侧地区的植物区系仍是一整体．且由于中国特有科、东亚特有科、亚洲特有科、东亚和北美间断分布科,以及各种区系和植被表征科的存在表明广东植物区系是东亚植物区的一个组成部分。     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

海南竹类的地理分布及区系特征

通过对海南岛竹类区系地理成分进行统计、分析和对其分布、与邻近地区竹类区系关系的探讨,其区系特征归纳如下:1.拥有丰富的特有成分。2.区系的热带性质明显。3.分布上差异显著。4.区系原始、古老。5.与毗邻的广东竹类区系联系较多。     

秦岭的珍稀特有植物及其区系特征

本文论迹了秦岭地区珍稀、特有植物的种类组成,历史成因,及其区系特征和与邻近地区植物的关系,并分析了它们的分布特点。1.秦岭是我国南北植物的分界线,有珍稀植物66种,特有植物169种。2.秦岭珍稀植物具有相对的古老性,它们与华中亚热带区系的历史渊源较深。而秦岭特有植物却具有相对的年青性,表现出明显的温带性特点和剧烈的分化作用,大部分为分化较晚的新特有种。3.秦岭地区地史上和现在的南北地质、气候条件的分异,决定了其珍稀植物的特点,也使其本身成为我国大多数亚热带珍稀植物分布的北界。而复杂的地形造成不同气候条件的交错和变化,及明显的新构造运动引起垂直气候带谱的变化,导致了秦岭植物的强烈变异和分化,以及大量新特有种的形成。4.秦岭珍稀植物具有多种区系成分,与邻近地区有着广泛的联系,尤其与川东-鄂西地区联系最为密切。我们认为从这里经大巴山东端到秦岭中部,再向西扩展,是秦岭与华中亚热带植物区系相互渗透的一条主要路线。