Phytogeography of the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica

Aim Central America is a biogeographically interesting area because of its location between the rich and very different biota of North and South America. We aim to assess phytogeographical patterns in the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica. Location Tropical America, in particular the montane area of Cordillera de Talamanca, Costa Rica. Methods The analysis is based on a new critical inventory of the montane bryophyte flora of Cordillera de Talamanca. All species were assigned to phytogeographical elements on the basis of their currently known distribution. Absolute and percentage similarities were employed to evaluate floristic affinities. Results A total of 401 species [191 hepatics (liverworts), one hornwort, 209 mosses] are recorded; of these, 251 species (128 hepatics, one hornwort, 122 mosses) occur in oak forests. Ninety‐three per cent of all oak forest species are tropical in distribution, the remaining 7% are temperate (4%) and cosmopolitan (3%) species. The neotropical element includes almost 74% of the species, the wide tropical element (pantropical, amphi‐atlantic, amphi‐pacific) only 19%. A significant part of the neotropical species from oak forests are species with tropical Andean‐centred ranges (27%). As compared with bryophyte species, vascular plant genera in the study region are represented by fewer neotropical, more temperate and more amphi‐pacific taxa. Bryophyte floras of different microhabitats within the oak forest and epiphytic bryophyte floras on Quercus copeyensis in primary, early secondary and late secondary oak forest show a similar phytogeographical make‐up to the total oak forest bryophyte flora. Comparison of oak forest and páramo reveals a greater affinity of the páramo bryophyte flora to temperate regions and the great importance of the páramo element in páramo. Surprisingly, oak forests have more Central American endemics than páramo. Main conclusions (1) Providing first insights into the phytogeographical patterns of the bryophyte flora of oak forests and páramo, we are able to confirm general phytogeographical trends recorded from vascular plant genera of the study area although the latter were more rich in temperate taxa. (2) Andean‐centred species are a conspicuous element in the bryophyte flora of Cordillera de Talamanca, reflecting the close historical connection between the montane bryophyte floras of Costa Rica and South America. (3) High percentages of Central American endemics in the bryophyte flora of the oak forests suggest the importance of climatic changes associated with Pleistocene glaciations for allopatric speciation.     

Brazilian Páramos II. Macro- and mesoclimate of the campos de altitude and affinities with high mountain climates of the tropical Andes and Costa Rica

Aim This paper presents a basic analysis of the macro- and mesoclimate of the Brazilian campos de altitude, a series of cool–humid, mountaintop grasslands in southeastern Brazil, and compares results with data from other tropical as well as temperate alpine sites. Location Beginning at altitudes of 1800–2000 m, the campos de altitude are found atop the highest summits of the main ranges of the southeastern Brazilian Highlands, between the states of Santa Catarina and Minas Gerais/Espírito Santo. Methods Macro- and mesoclimatic parameters for the campos de altitude are derived from both original data and previously reported results. Parameters include approximate radiation budgets, temperature lapse rates, seasonal and diurnal patterns in temperature, occurrence of frost, elevational gradients in precipitation, and interannual and seasonal patterns in precipitation. Using multivariate techniques and simple numerical contrasts, the climate of the campos de altitude is compared to climates of other tropical as well as temperate alpine sites. Results With respect to patterns of seasonality and the marked influence of polar frontal activity, the macroclimate of the campos de altitude is typically tropical-marginal. However, in reference to actual temperature and precipitation values, the length and profundity of the dry season, average humidities and cloudiness, the climate of the campus de altitude more closely corresponds to that of more inner-tropical systems. These commonalities are best developed with respect to páramo climates of the northern Andes and, especially, Costa Rica. Main conclusions Their very different latitudinal and geographic positions notwith- standing, the campos de altitude and high mountain formations of the N. Andean and Central American Cordillera show clear macroclimatic congruities. In these congruities reside both the environmental basis for strong Andean–southeast Brazilian biogeographic connections, and the context within which evolutionary and ecological parallelisms have developed in the biota of these two widely separate neotropical mountain systems.     

Brazilian Páramos I. An introduction to the physical environment and vegetation of the campos de altitude

Aim The campos de altitude are a series of cool-humid, grass-dominated formations restricted to the highest summits of the southeastern Brazilian Highlands. Relatively little is known of the ecology, biogeography, or developmental history of this archipelago of mountaintop formations. This contribution presents an overview of our present state of knowledge with respect to the past and present physical environment and vegetation of the campos de altitude. The aim is (1) to introduce an international audience to the natural history of these diminutive yet important ecosystems, and (2) to provide the background for a series of forthcoming contributions that will treat the ecology of the campos de altitude and explore physical and biotic relationships between these ‘Brazilian páramos’ and similar formations in the high mountains of equatorial South and Central America. Location Beginning at altitudes of 1800–2000 m, the campos de altitude are found atop the highest summits of the main ranges of the southeastern Brazilian Highlands, between the states of Santa Catarina and Minas Gerais/Espírito Santo. Methods Drawing upon both original data and previously published results, this contribution reviews what we presently know about the physical environment and vegetation of the campos de altitude, in the context of geographic setting, geology, palaeoclimatology and palaeobotany, modern climate, modern vegetation, and conservation. Results and main conclusions Uplift of the southeastern Brazilian ranges to altitudes high enough to support orographic grasslands may have occurred as early as the middle- to late-Tertiary; pollen records show that campos de altitude have been extant on southeastern summits at least since the Late Pleistocene. The present-day climate of the campos de altitude is cool and (per)humid, but patterns of rainfall, temperature, and frost are distinctly seasonal. Although the flora of the campos de altitude is highly diverse and characterized by a high degree of endemism, the campos de altitude maintain strong floristic similarities to equatorial alpine formations of the Andean and Central American Cordillera; these similarities also extend to climate, soils, and vegetation physiognomy. Anthropogenic fires and grazing are widespread in the campos de altitude and probably contribute significantly to the modern structure of vegetation communities.     

云南哀牢山地区蕨类植物区系的垂直分布格局

根据大尺度的物种分布信息,利用聚类分析和数理统计的方法分析了云南哀牢山蕨类植物区系的垂直分布格局。研究结果表明,随着海拔的升高,热带区系成分所占比重呈显著递减趋势,而温带区系成分所占比重呈明显递增趋势。在整个海拔梯度上,东亚分布和热带亚洲分布成分始终占主导地位。区系过渡性随着海拔的升高呈先增强后减弱的单峰分布格局,平衡点出现于海拔1700 m左右。基于区系成分比重的聚类分析表明,在海拔1300 m,1900 m和2600 m左右,将海拔梯度上的蕨类植物分为4组,这与各海拔段内的物种组成和区系成分性质是一致的。     

罗霄山脉种子植物区系及其南北分化特征

本文基于野外调查和文献资料整理, 对罗霄山脉种子植物区系及其自北向南5条中型山脉之间的区系过渡性特征进行了系统分析。结果表明: (1)该区域内野生种子植物有179科1,107属4,314种; 区系表征科主要以亚热带山地为现代分布中心。科的区系组成为: 单型科25科, 寡种科61科, 单型属465属, 寡种属432属, 呈现出区系特征上的孑遗性, 也显示出本地区系成分与典型热带成分及典型温带成分间存在密切交流。(2)科的地理成分共分为10个类型, 以热带性成分占明显优势, 热带性科83科及温带性科40科分别占非世界性科总数的67.48%和32.52%; 属的地理成分包括15个分布区类型, 以温带成分占优势, 温带性属534属及热带性属484属分别占非世界性属总数的52.46%和47.54%。属分布区类型以泛热带分布、热带亚洲分布、北温带分布、东亚分布为主, 体现出本地区与古热带植物区系及温带第三纪植物区系的紧密联系。(3)罗霄山脉5条山脉间区系组成呈现出明显的南北分异特征, 自北向南热带属与温带属比值逐渐增大, 北段幕阜山脉温带性成分明显占优势, 与华中区系关联密切, 而南段万洋山脉、诸广山脉热带性属较占优势, 分布有丰富的华南区系成分。罗霄山脉地区是重要的区系交流通道, 历史时期的气候波动是本地区现代区系分布格局形成的主要原因。     

A biogeographical study on tropical flora of southern China

The tropical climate in China exists in southeastern Xizang (Tibet), southwestern to southeastern Yunnan, southwestern Guangxi, southern Guangdon, southern Taiwan, and Hainan, and these southern Chinese areas contain tropical floras. I checked and synonymized native seed plants from these tropical areas in China and recognized 12,844 species of seed plants included in 2,181 genera and 227 families. In the tropical flora of southern China, the families are mainly distributed in tropical areas and extend into temperate zones and contribute to the majority of the taxa present. The genera with tropical distributions also make up the most of the total flora. In terms of geographical elements, the genera with tropical Asian distribution constitute the highest proportion, which implies tropical Asian or Indo‐Malaysia affinity. Floristic composition and geographical elements are conspicuous from region to region due to different geological history and ecological environments, although floristic similarities from these regions are more than 90% and 64% at the family and generic levels, respectively, but lower than 50% at specific level. These differences in the regional floras could be influenced by historical events associated with the uplift of the Himalayas, such as the southeastward extrusion of the Indochina geoblock, clockwise rotation and southeastward movement of Lanping–Simao geoblock, and southeastward movement of Hainan Island. The similarity coefficients between the flora of southern China and those of Indochina countries are more than 96% and 80% at family and generic levels, indicating their close floristic affinity and inclusion in the same biogeographically floristic unit.     

三峡大老岭植物区系的垂直梯度分析

为探讨山地植物区系构成特征及其垂直梯度的生态意义,根据对三峡大老岭地区植被垂直样带 调查获得的植物区系资料,分析了该地区植物区系成分构成的基本特征及其随海拔梯度的变化趋势,寻找了区系平衡点的位置;并利用聚类方法分析了山地气候垂直分异对区系成分构成的影响。结果表明：①大老岭植物区系具有温带性质,但仍反映了与热带区系的历史联系,有强烈的区域性;②属的分布区类型可归为热带分布、温带分布、地中海—中亚中心和东亚中心4组,各组区系成分的垂直梯度特征不同;热带、亚热带成分与温带成分的平衡点大致位于海拔650m;③区系成分构成和属的物种数量构成的聚类分析结果一致显示了植物区系构成与山地气候和植被垂直带相对应的格局。     

Phytogeographical relations of the Andean dry valleys of Bolivia

Aim The objective of this study is to examine the phytogeographical affinities of the Andean dry valleys of Bolivia in order to contribute to a better understanding of the Andean dry flora's distribution, origin and diversity. Particular emphasis is given to the analysis of the floristic connections of this flora with more austral parts of South America. Location The dry valleys of Bolivia are located in the Andes of the southern half of the country, at elevations between 1300 and 3200 m. Methods An extensive floristic list compiled by the author to evaluate plant diversity in these Andean regions was used as the base for this study. To accomplish this, all recorded genera and species were assigned, respectively, to 11 and 12 phytogeographical elements established previously by the author. Two phytogeographical spectra were thus obtained and analysed. Results At the genus level, the Andean dry valleys of Bolivia are clearly dominated by genera that have widespread distributions (cosmoplitan and subtropical genera). Many of these reached the Andes from the lowland region of the Chaco. At species level, Andean elements constitute more than 60% of the species total, most of which are restricted to the central‐southern Andes. This suggests that Chaco‐related and Andean genera had considerable levels of speciation in these valleys. Many genera and more than half the species have their northernmost distribution in the dry valleys of Bolivia, thereby underlining strong relationships with central‐southern South America (mainly Argentina, Paraguay and southern Brazil). The data supports the belief of the existence, in central‐southern Peru, of a floristic disjunction in dry to arid environments that separates a tropical dry flora north of this limit from a dry subtropical/warm temperate flora south of it. Main conclusions The Andean dry valleys of Bolivia are diverse plant communities with high levels of endemism (c. 18% of the species). The species of this region are more related to those present in central‐southern South America than to the flora of northern South America that ranges southwards to Peru. Many of the species have restricted distributions in the dry Andes of Bolivia and Argentina, and many genera of these dry valleys have their northernmost distribution in Bolivia/southern Peru, too. The data point to high levels of speciation also in the central Andes.     

辽西走廊白狼山保护区植物区系及其与邻近山地的关系

采用植物区系谱、排序分析、聚类模型和相关性等方法,研究了辽西白狼山植物区系属的地理组成以及与其它山地的相似关系,以揭示多个山地植物区系的相互关系及分布格局。结果表明:(1)白狼山自然保护区野生种子植物多样而丰富,共有101科394属790种;被子植物占总种数的98.35%,优势科现象明显,寡种属和单种属占优势,呈现一定残遗性;(2)属的地理成分以温带成分为主,共有273属,占总属数的81.3%,处于中国-日本森林植物亚区暖温带-温带过渡范围内,植物区系划归于华北植物区系平原山地亚地区。(3)相关分析表明,白狼山与努鲁儿虎山(r=0.999)、东灵山(r=0.998)之间的植物区系具有极显著相关关系;主坐标排序和聚类分析表明,白狼山种子植物属的分布区类型与努鲁儿虎山、东灵山、小五台山以及雾灵山等有着较大的相似性,不同纬度水热条件可能是影响植物区系地理分布格局的重要因素。     

云南省小黑山自然保护区种子植物区系研究

小黑山自然保护区有野生种子植物170科787属2195种。分析表明该区种子植物区系的地理成分复杂,联系广泛。热带性质的属有506属,占总属数的68.6%,温带性质的属有224属,占总属数的30.4%。热带性质的种890种,占总种数的40.9%,温带性质的种有1267种,占总种数的58.3%。表明该地区植物区系具有明显的亚热带性质,同时深受热带植物区系的影响。植物区系来源主要由东亚成分、热带亚洲成分(印度—马来西亚成分)、中国特有成分三部分融合而成。特有现象明显,有东亚特有科5科,中国特有属6属,中国特有种778种,占种总数的35.8%,而云南特有种有355种,占中国特有种的45.6%。保护区还拥有众多的珍稀濒危植物。     

福建梁野山国家级自然保护区种子植物区系研究

通过野外调查、采集标本和查阅相关文献资料,编制福建梁野山自然保护区野生种子植物名录,进行植物区系组成和分布区类型的分析,并将其与周边相邻2个保护区的植物区系进行比较。结果表明:(1)梁野山自然保护区共有野生种子植物154科662属1 498种,种类丰富。(2)区系地理成分以热带分布型为主,其中在科级水平上热带分布型所属科共有85科,占总科数的69.67%;在属级水平上热带性质分布型的属有371属,占总属数的60.82%。但相当数量温带成分的存在,体现了亚热带植物区系的特点。(3)对梁野山保护区与周边相邻的武夷山和南靖南亚热带雨林保护区的植物区系比较发现,梁野山种子植物区系属数的热带属数/温带属数(R/T值,1.66)高于武夷山的R/T值(1.10)而低于南靖南亚热带雨林保护区的R/T值(2.68),梁野山种子植物区系与该相邻2个保护区的相似性系数非常接近,但在区系成分上,梁野山既有与武夷山共有而南靖南亚热带雨林保护区没有分布的温带成分,也有与南靖南亚热带雨林保护区共有而武夷山没有的热带成分,说明梁野山植物区系与其地处武夷山脉南端的地理位置相一致,具有明显的从南亚热带向中亚热带过渡的特点。     

皖西霍山县白马尖植物区系和植被

本文论述了北亚热带典型地段——大别山主峰白马尖的植被和植物区系的特点,提出白马尖地区是一“生物避难所”。与纬度相近地区的比较研究,结果表明,海拔1000米以上的山地,植被类型和植物区系成份与皖南黄山、浙北天目山较一致,而低海拔地区与江苏的宜溧山区相似,特别是常绿阔叶树的属、种极为一致。值得指出的是耐寒常绿阔叶林类型的发现,为北亚热带植被地带的概念增添了新内容。文中对原圈定佛子岭水库库区周围为水源涵养林自然保护区认为不妥,将白马尖主峰整个集水区包括在内,才是完善的。     

Biogeographical Divergence of the Flora of Yunnan,Southwestern China Initiated by the Uplift of Himalaya and Extrusion of Indochina Block

The floral composition of Yunnan is conspicuously linked to the biogeographical history of this extremely species-rich province in southwestern China. The floristic compositions of three representative regions in Yunnan were compared to reveal their variation with geography. From southern Yunnan, 4150 native species (including subspecies and varieties) from 1240 genera and 183 families of seed plants were recognized. From central Yunnan 3389 native species from 1095 genera and 167 families of seed plants were recognized. From northwestern Yunnan 6807 native species from 1296 genera and 166 families of seed plants were recognized. Although these three floras across Yunnan are similar in familial composition, similarities between the floras of southern and northwestern Yunnan are low at the generic and specific levels. The flora of northwestern Yunnan is dominated by families and genera with cosmopolitan and north temperate distributions, while the flora of southern Yunnan is dominated by tropical families and genera. Northwestern Yunnan is composed largely of temperate genera, of which the highest proportion has a north temperate distribution. In contrast, southern Yunnan has mainly tropical genera, of which most have a tropical Asian distribution. The flora of central Yunnan is a combination of southern and northwestern Yunnan. These three floras might be derived from a common Tertiary tropical or subtropical East Asian flora, but the geological history of each region has influenced its flora, and they have remained divergent since the late Tertiary. The flora of northwestern Yunnan has evolved with the uplift of the Himalayas and by gradual proliferation of mainly cosmopolitan and north temperate floristic elements, while the flora of southern Yunnan has evolved with extrusion of the Indochina block and the influence of mainly tropical Asian elements.     

Large-scale phytogeographical patterns in East Asia in relation to latitudinal and climatic gradients

Aim This paper aims at determining how different floristic elements (e.g. cosmopolitan, tropical, and temperate) change with latitude and major climate factors, and how latitude affects the floristic relationships between East Asia and the other parts of the world. Location East Asia from the Arctic to tropical regions, an area crossing over 50° of latitudes and covering the eastern part of China, Korea, Japan and the eastern part of Russia. Methods East Asia is divided into forty‐five geographical regions. Based on the similarity of their world‐wide distributional patterns, a total of 2808 indigenous genera of seed plants found in East Asia were grouped into fourteen geographical elements, belonging to three major categories (cosmopolitan, tropical and temperate). The 50°‐long latitudinal gradient of East Asia was divided into five latitudinal zones, each of c. 10°. Phytogeographical relationships of East Asia to latitude and climatic variables were examined based on the forty‐five regional floras. Results Among all geographical and climatic variables considered, latitude showed the strongest relationship to phytogeographical composition. Tropical genera (with pantropical, amphi‐Pacific tropical, palaeotropical, tropical Asia–tropical Australia, tropical Asia–tropical Africa and tropical Asia geographical elements combined) accounted for c. 80% of the total genera at latitude 20°N and for c. 0% at latitude 55–60°N. In contrast, temperate genera (including holarctic, eastern Asia–North America, temperate Eurasia, temperate Asia, Mediterranean, western Asia to central Asia, central Asia and eastern Asia geographical elements) accounted for 15.5% in the southernmost latitude and for 80% at 55–60°N, from where northward the percentage tended to level off. The proportion of cosmopolitan genera increased gradually with latitude from 5% at the southernmost latitude to 21% at 55–60°N, where it levelled off northward. In general, the genera present in a more northerly flora are a subset of the genera present in a more southerly flora. Main conclusions The large‐scale patterns of phytogeography in East Asia are strongly related to latitude, which covaries with several climatic variables such as temperature. Evolutionary processes such as the adaptation of plants to cold climates and current and past land connections are likely responsible for the observed latitudinal patterns.     

中国帕米尔高原种子植物的区系地理研究

中国帕米尔高原约有种子植物59科303属963种(包括变种)。分析表明:该区种子植物区系的地理成分较为复杂,联系广泛。温带性质的属有254属,占总属数的83.8%,热带性质的属有13属,占总属数的4.3%;温带性质的种有723种,占总种数的75.1%,热带性质的种有17种,占总种数的1.8%。表明该地区植物区系具有明显的温带性质,同时受到热带植物区系的影响。另外在区系科属分级水平上,寡属科、单属科占绝对优势,而超过20属的大科在该区仅有3科,却占总属数、总种数的35.9%和38.6%,区系优势现象十分明显;单种或少种属也较多,属种比值偏高,表明区系组成上的复杂性。     

贵州太平山自然保护区种子植物区系

经野外调查和资料整理,贵州省黎平县太平山自然保护区共有野生种子植物147科492属1 178种(含种下等级)。区系成分分析表明:在科级水平上,热带分布科占保护区非世界分布科的65. 25%,温带分布科占34. 75%,热带成分明显占优;在属级水平上,热带分布属占保护区非世界分布属的51. 92%,温带分布属占46. 05%,热带成分有微弱优势,说明该区域的植物区系具有亚热带性质。区系相似性比较分析表明,该区域的种子植物区系与黄桑自然保护区、佛顶山自然保护区、花坪自然保护区的关系较为亲近。     

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。     

贡嘎山蕨类植物区系的特点

贡嘎山蕨类植物区系共含４０科,９３属,３９９种,最主要的是耳蕨属Ｐｏｌｙｓｔｉｃｈｕｍ,鳞毛蕨属Ｄｒｙｏｐｔｅｒｉｓ,蹄盖蕨属Ａｔｈｙｒｉｕｍ以及水龙骨科Ｐｏｌｙｐｏｄｉａｃｅａｅ等系统演化上较高级的类群,所含的种子是喜马拉雅和中国西南当地分化的种;其特有成分多为新特有属,可认为它是随青藏高原隆起而形成的较年青的蕨类区系,热带属在本区数量多而种类少,在山上河可与北温带种类并存。贡嘎山处于蕨类物种东     

Phytogeographical patterns of genera of endemic flora in relation to latitudinal and climatic gradients in China

This paper aims at detecting the relationships between phytogeographical patterns of genera of Chinese endemic seed plants and latitude or climatic factors. The landmass of China was divided into four latitudinal zones, each of c. 8°. Based on a total of 1664 indigenous genera of Chinese endemic seed plants which were grouped into fifteen geographical elements, belonging to three major categories (cosmopolitan, tropical and temperate) and which were absent or present in 28 provinces in China, we analyzed the phytogeographical patterns of genera of Chinese endemic seed plants and detected the relationships between them and main climatic factors. Our results showed that the proportion of tropical genera decreases with the increase in latitude; the proportion of temperate genera increases with the increase in latitude; and the proportion of cosmopolitan genera increased gradually increased gradually with latitude. There are a slow decrease in the proportion of tropical genera and slow increase in the proportion of temperate genera at latitudes 35°–40°. Alternatively, the tropical genera and the temperate genera have the same proportion at latitude c. 25°. These changes and issues about the different genera also appeared in main climate factors. In general, the genera present in a more northerly flora are a subset of the genera present in a more southerly flora. In summary, the large-scale patterns of phytogeography of endemic flora in China are strongly related to latitude, which covary with several climatic variables such as temperature.