海洋微型底栖生物的多样性与地理分布

海洋微型底栖生物是指生活于海洋沉积物中及表面的所有单细胞原核和真核微型生物,包括原核微生物、真核微藻及原生动物等光合自养和异养的类群.与水体相比,海洋底栖生境孕育了形态和功能多样性更高的微型生物,在陆架浅海单位体积沉积物中其丰度较之水体中同类生物高一至几个数量级,而深海则孕育着特殊进化环境下新奇多样且数量庞大的微型底栖生物,是维持海洋生物多样性、海洋生态系统结构和功能不可或缺的部分.迄今对于微型生物是全球性还是限定性分布一直存在争议,对其解答受到分类研究欠缺及采样不足的制约.分子生物学从正反两方面提供了理论依据,但无法得出一个普遍接受的观点.海洋微型底栖生物的多样性研究侧重于物种多样性及群落结构与分子多样性,较为显著的进展体现在原核微生物的分子多样性及底栖真核微藻的物种多样性研究,对于海洋底栖原生动物的多样性研究则相较滞后.本文综述了国内外对海洋微型底栖生物各主要类群的分类学和多样性研究进展,探讨了各类群在全球的潜在物种多样性,并就我国未来加强海洋微型底栖生物多样性构成、分布与变动及驱动变化的因子以及底栖微食物网的研究提出了建议.     

Biodiversity and historical biogeography of stalked crinoids (Echinodermata) in the deep sea

About 95 species of stalked crinoids are now described from 60m to hadal depths, but our knowledge remains far from complete. Depending on which species concept is used, estimates of species richness can be dramatically different. It is necessary to have a homogeneous concept for taxonomic units. The abundance of the crinoid fossil record allows a discussion of the ancestry of deep sea crinoid fauna. Stalked crinoids have a horizontal diversity pattern with three regional centres of high diversity (i.e. western tropical Pacific, western tropical Atlantic and north-eastern Atlantic). Vertical patterns show two faunal strata which vary in importance among provinces. The epibathyal stratum has apparently remained relatively similar in intertropical areas since the Mesozoic. Despite environmental changes related to glaciation since the Middle Miocene, the deepest crinoid fauna (i.e. the deep sea fauna sensu stricto at depths more than 1000 ± 200 m) have a very ancient origin with a dispersion closely related to plate tectonics. The bathyal fauna on hard substrates includes a few living fossils and has a high historical interest.     

3. Biodiversity, biogeography and conservation of diatoms

Recent morphometric and breeding studies of diatoms show that the present species classification is too coarse and hides significant diversity. Many species are subdivided into phenodemes, which often differ in cell size, shape, stria density and pattern, but may also have different ultrastructural features. In raphid diatoms these can include the form of the raphe endings, details of the pore occlusions, and the structure of the girdle, while chloroplast structure can also vary. The phenodemes can be sympatric or allopatric. In Sellaphora pupula and other species, sympatric phenodemes are reproductively isolated. It is recommended that such demes are recognized as separate species; the total number of diatom species worldwide may thus be at least 2 × 10⁵. Use of a fine-grained classification reveals that many diatom species may be endemics, some restricted to a single lake or catchment, others to wider areas. Environmental impact assessments and conservation strategies must begin to take account of endemism and rarity among microscopic algae and protists.     

Biodiversity,biogeography, and connectivity of polychaetes in the world's largest marine minerals exploration frontier

Aim

The abyssal Clarion-Clipperton Zone (CCZ), Pacific Ocean, is an area of commercial importance owing to the growing interest in mining high-grade polymetallic nodules at the seafloor for battery metals. Research into the spatial patterns of faunal diversity, composition, and population connectivity is needed to better understand the ecological impacts of potential resource extraction. Here, a DNA taxonomy approach is used to investigate regional-scale patterns of taxonomic and phylogenetic alpha and beta diversity, and genetic connectivity, of the dominant macrofaunal group (annelids) across a 6 million km² region of the abyssal seafloor.

Location

The abyssal seafloor (3932–5055 m depth) of the Clarion-Clipperton Zone, equatorial Pacific Ocean.

Methods

We used a combination of new and published barcode data to study 1866 polychaete specimens using molecular species delimitation. Both phylogenetic and taxonomic alpha and beta diversity metrics were used to analyse spatial patterns of biodiversity. Connectivity analyses were based on haplotype distributions for a subset of the studied taxa.

Results

DNA taxonomy identified 291–314 polychaete species from the COI and 16S datasets respectively. Taxonomic and phylogenetic beta diversity between sites were relatively high and mostly explained by lineage turnover. Over half of pairwise comparisons were more phylogenetically distinct than expected based on their taxonomic diversity. Connectivity analyses in abundant, broadly distributed taxa suggest an absence of genetic structuring driven by geographical location.

Main Conclusions

Species diversity in abyssal Pacific polychaetes is high relative to other deep-sea regions. Results suggest that environmental filtering, where the environment selects against certain species, may play a significant role in regulating spatial patterns of biodiversity in the CCZ. A core group of widespread species have diverse haplotypes but are well connected over broad distances. Our data suggest that the high environmental and faunal heterogeneity of the CCZ should be considered in future policy decisions.     

低温细菌与古菌的生物多样性及其冷适应机制

低温细菌与古菌广泛分布于地球的低温环境,包括南极、北极及高山地带的冻土、低温土壤和荒漠、冰川、湖泊、海冰,以及深海、冰洞和大气平流层等.栖息在这些低温环境中的细菌与古菌具有丰富的多样性,主要为α,p和γ-Proteobacteria分支、CFB类群分支和革兰氏阳性细菌分支等.由于低温环境中的微生物流动性低,因而是研究微生物地理学理想的生态系统,有助于理解地球微生物的多样性、分布规律乃至形成机制.由于长期生活在冰冻环境中,低温细菌与古菌形成了多种适应低温环境的生理机制,如它们通过细胞膜脂类的组成来调节膜的流动性以维持正常的细胞生理功能;利用相容性溶质、抗冻蛋白、冰核蛋白及抗冰核形成蛋白等实现低温保护作用;产生冷激蛋白、冷适应蛋白和DEAD-box RNA解旋酶保持低温下RNA的正确折叠、蛋白质翻译等重要的生命活动;另外还产生低温酶,提高能量产生和储存效率等以适应低温环境.随着DNA序列分析技术的飞速发展,各类组学方法也用于揭示微生物全局性的冷适应机制.     

Biodiversity and biogeography of southern temperate and polar bryozoans

Aim To describe the distribution of biodiversity and endemism of bryozoans in southern temperate and polar waters. We hypothesized that we would find: (1) no strong latitudinal richness gradient; (2) striking contrasts in richness and endemism between clades and between regions; and (3) that faunal similarity of regions would cluster geographically around each southern continent. Location South Atlantic, Indian and Pacific Oceans and the Southern Ocean. Methods We constructed a data base from known literature, regional data bases and recent finds. We regionalized each southern continent, calculated levels of richness and endemism for each region and continent, and used primer 5 to perform multivariate statistical analysis. Results A third (1681) of global bryozoan species described occur south of 30° S, of which c. 87% were cheilostomes. In richness we found no latitudinal cline and change across longitude was stronger. New Zealand was richest and had the most (60%) endemic species, followed by Antarctica at 57%. There were striking contrasts in regional richness and endemism between clades but the highest levels of between‐region similarity were around Antarctica. The timing of past continent connectivity was reflected. Main conclusions Bryozoans show strong hemispherical asymmetry in richness and, like molluscs and corals, decrease away from Australasia rather than with latitude. Species endemism is much lower in Antarctic bryozoans than previously thought, and as this taxon is not particularly dispersive and is now amongst the best studied regionally, maybe Antarctic endemism in general is lower and Antarctica less cut‐off to species dispersal than previously thought. However, Antarctic generic endemism is double the level previously calculated and regional faunal similarities are much higher than around other continents – both reflecting long‐term isolation. Bryozoans, in contrast to the paradigm of Antarctic fauna, may be fairly robust to predicted climate change. Paradoxically, they may also be one of the best taxa to monitor to sensitively detect marine benthic responses.     

Biodiversity growth on the volcanic ocean islands and the roles of in situ cladogenesis and immigration: case with the reptiles

Models for biodiversity growth on the remote oceanic islands assume that in situ cladogenesis is a major contributor. To test this, we compiled occurrence data for 194 terrestrial reptile species on 53 volcanically‐constructed middle‐ to low‐latitude landmasses worldwide. Despite 273 native island‐species records, there are only 8–12 cases of the phenomenon, including just two radiations. Diversification frequencies are largely uncorrelated with island area, age, maximum altitude, and isolation. Furthermore, there is no indication that the presence of non‐sister congeners on an island stymies the process. Diversity on individual oceanic islands therefore results primarily from immigration and anageneis, but this is not a simple matter. Clusters that are difficult to reach (far or challenging to get to) or thrive upon (e.g. Canaries, Galápagos) have relatively few clades (3–8), some of which have many species (6–14), and all host at least one endemic genus. In these settings, diversity grows mainly by intra‐archipelago transfer followed by within‐island anagenetic speciation. In contrast, those island groups that are easier to disperse to (characterized by short distances and conducive transit conditions) and harbour more benign habitats (e.g. Comoros, Lesser Antilles) have been settled by many ancestor‐colonizers (≥ 14), but each clade has few derived species (≤ 4). These archipelagoes lack especially distinctive lineages. Models explaining the assembly and growth of terrestrial biotic suites on the volcanic ocean islands thus need to accommodate these new insights.     

Global biogeography of highly diverse protistan communities in soil

Protists are ubiquitous members of soil microbial communities, but the structure of these communities, and the factors that influence their diversity, are poorly understood. We used barcoded pyrosequencing to survey comprehensively the diversity of soil protists from 40 sites across a broad geographic range that represent a variety of biome types, from tropical forests to deserts. In addition to taxa known to be dominant in soil, including Cercozoa and Ciliophora, we found high relative abundances of groups such as Apicomplexa and Dinophyceae that have not previously been recognized as being important components of soil microbial communities. Soil protistan communities were highly diverse, approaching the extreme diversity of their bacterial counterparts across the same sites. Like bacterial taxa, protistan taxa were not globally distributed, and the composition of these communities diverged considerably across large geographic distances. However, soil protistan and bacterial communities exhibit very different global-scale biogeographical patterns, with protistan communities strongly structured by climatic conditions that regulate annual soil moisture availability.     

Atlantic reef fish biogeography and evolution

Aim To understand why and when areas of endemism (provinces) of the tropical Atlantic Ocean were formed, how they relate to each other, and what processes have contributed to faunal enrichment. Location Atlantic Ocean. Methods The distributions of 2605 species of reef fishes were compiled for 25 areas of the Atlantic and southern Africa. Maximum‐parsimony and distance analyses were employed to investigate biogeographical relationships among those areas. A collection of 26 phylogenies of various Atlantic reef fish taxa was used to assess patterns of origin and diversification relative to evolutionary scenarios based on spatio‐temporal sequences of species splitting produced by geological and palaeoceanographic events. We present data on faunal (species and genera) richness, endemism patterns, diversity buildup (i.e. speciation processes), and evaluate the operation of the main biogeographical barriers and/or filters. Results Phylogenetic (proportion of sister species) and distributional (number of shared species) patterns are generally concordant with recognized biogeographical provinces in the Atlantic. The highly uneven distribution of species in certain genera appears to be related to their origin, with highest species richness in areas with the greatest phylogenetic depth. Diversity buildup in Atlantic reef fishes involved (1) diversification within each province, (2) isolation as a result of biogeographical barriers, and (3) stochastic accretion by means of dispersal between provinces. The timing of divergence events is not concordant among taxonomic groups. The three soft (non‐terrestrial) inter‐regional barriers (mid‐Atlantic, Amazon, and Benguela) clearly act as ‘filters’ by restricting dispersal but at the same time allowing occasional crossings that apparently lead to the establishment of new populations and species. Fluctuations in the effectiveness of the filters, combined with ecological differences among provinces, apparently provide a mechanism for much of the recent diversification of reef fishes in the Atlantic. Main conclusions Our data set indicates that both historical events (e.g. Tethys closure) and relatively recent dispersal (with or without further speciation) have had a strong influence on Atlantic tropical marine biodiversity and have contributed to the biogeographical patterns we observe today; however, examples of the latter process outnumber those of the former.     

The role of ecological knowledge in explaining biogeography and biodiversity in Amazonia

Biogeographical studies in Amazonia have commonly taken a historical, rather than an ecological approach. General patterns have been sought in the distribution maps of different species, and these have been explained in terms of past or present distribution barriers, especially past climates and large rivers. Implicitly, and often also explicitly, it is assumed that Amazonia is ecologically so uniform that present-day ecological conditions are rather insignificant in determining species distribution patterns and speciation. However, this assumption is more based on the lack of relevant data than on actual observations of environmental uniformity or ecological unspecialization of the species. Recent studies have indeed documented ecological heterogeneity and floristic differences among sites that were previously thought similar. In the absence of direct knowledge of the past, more complete ecological and environmental understanding of the present-day Amazonia are needed for evaluating the relative roles of historical and ecological factors in Amazonian biogeography and biodiversity.     

Conservation biogeography of the Antarctic

Aim

To present a synthesis of past biogeographic analyses and a new approach based on spatially explicit biodiversity information for the Antarctic region to identify biologically distinct areas in need of representation in a protected area network.

Location

Antarctica and the sub‐Antarctic.

Methods

We reviewed and summarized published biogeographic studies of the Antarctic. We then developed a biogeographic classification for terrestrial conservation planning in Antarctica by combining the most comprehensive source of Antarctic biodiversity data available with three spatial frameworks: (1) a 200‐km grid, (2) a set of areas based on physical parameters known as the environmental domains of Antarctica and (3) expert‐defined bioregions. We used these frameworks, or combinations thereof, together with multivariate techniques to identify biologically distinct areas.

Results

Early studies of continental Antarctica typically described broad bioregions, with the Antarctic Peninsula usually identified as biologically distinct from continental Antarctica; later studies suggested a more complex biogeography. Increasing complexity also characterizes the sub‐Antarctic and marine realms, with differences among studies often attributable to the focal taxa. Using the most comprehensive terrestrial data available and by combining the groups formed by the environmental domains and expert‐defined bioregions, we were able to identify 15 biologically distinct, ice‐free, Antarctic Conservation Biogeographic Regions (ACBRs), encompassing the continent and close lying islands.

Main conclusions

Ice‐free terrestrial Antarctica comprises several distinct bioregions that are not fully represented in the current Antarctic Specially Protected Area network. Biosecurity measures between these ACBRs should also be developed to prevent biotic homogenization in the region.     

Floristic biogeography of the Hawaiian Islands: influences of area, environment and paleogeography

Aim A detailed database of distributions and phylogenetic relationships of native Hawaiian flowering plant species is used to weigh the relative influences of environmental and historical factors on species numbers and endemism. Location The Hawaiian Islands are isolated in the North Pacific Ocean nearly 4000 km from the nearest continent and nearly as distant from the closest high islands, the Marquesas. The range of island sizes, environments, and geological histories within an extremely isolated archipelago make the Hawaiian Islands an ideal system in which to study spatial variation in species distributions and diversity. Because the biota is derived from colonization followed by extensive speciation, the role of evolution in shaping the regional species assemblage can be readily examined. Methods For whole islands and regions of each major habitat, species–area relationships were assessed. Residuals of species–area relationships were subjected to correlation analysis with measures of endemism, isolation, elevation and island age. Putative groups of descendents of each colonist from outside the Hawaiian Islands were considered phylogenetic lineages whose distributions were included in analyses. Results The species–area relationship is a prominent pattern among islands and among regions of each given habitat. Species number in each case correlates positively with number of endemics, number of lineages and number of species per lineage. For mesic and wet habitat regions, island age is more influential than area on species numbers, with older islands having more species, more single‐island endemics, and higher species : lineage ratios than their areas alone would predict. Main conclusions Because species numbers and endemism are closely tied to speciation in the Hawaiian flora, particularly in the most species‐rich phylogenetic lineages, individual islands’ histories are central in shaping their biota. The Maui Nui complex of islands (Maui, Moloka‘i, Lāna‘i and Kaho‘olawe), which formed a single large landmass during most of its history, is best viewed in terms of either the age or area of the complex as a whole, rather than the individual islands existing today.     

Biodiversity of Collembola and their functional role in the ecosystem

More than 6500 species of Collembola are known from throughout the world and these are only a small part of the still undescribed species. There are many checklists and catalogues of Collembola for smaller territories and entire continents. Biogeographical analyses have been made for some genera and smaller territories. The most serious problems for a global biogeographical analysis is the lack of enough records from immense territories of all continents. Local biodiversity of Collembola can be very high, reaching over 100 species in small mountain ranges. Sampling methods do not impede documenting biodiversity on a global scale. Collembola have well differentiated ecomorphological life-forms and feeding guilds which enable the functional role that Collembola play in ecosystems to be recognised in some degree. Collembola play an important role in plant litter decomposition processes and in forming soil microstructure. They are hosts of many parasitic Protozoa, Nematoda, Trematoda and pathogenic bacteria and in turn are attacked by different predators. They utilise as food Protozoa, Nematoda, Rotatoria, Enchytraeidae, invertebrate carrion, bacteria, fungi, algae, plant litter, live plant tissues, and some plant pathogens. Soil acidification, nitrogen supply, global climate change and intensive farming have greatly impacted collembolan diversity.     

Biodiversity and biogeographic affiliation of Bryozoa from King George Island (Antarctica)

King George Island (KGI), which is located between the Antarctic and South American continents, may play a crucial role in the exchange of Bryozoa amongst the various Antarctic sectors and across the Polar Front. Knowledge regarding the biological diversity of this area could help us understand the evolution of the Antarctic ecosystem and its connectivity to the South American continent as well as the colonization ability of particular species. Here, we investigate the patterns of diversity and biogeographic affiliation of the cheilostome Bryozoa from KGI and the surrounding areas. Of 114 identified taxa from a depth range of 6–492 m, 26 species were reported for the first time in KGI. The most speciose genera were Camptoplites, Osthimosia, Smittina, and Cellarinella. Species richness at KGI consisted of 70% of the total bryozoans at the South Shetland Islands (SSI). Fifty-nine per cent of the bryozoans from KGI are endemic to Antarctica, which closely reflects the previously estimated endemism rate for bryozoans and other Antarctic taxa. Cluster analysis indicated that the strongest faunal links of SSI bryozoans were with Antarctic Peninsula assemblages, corresponding to the physical distance between both locations. The biogeographic similarities between SSI and South America confirm the broad trend of existing Antarctic–South American faunal links previously observed in bryozoans and many other taxa and indicate that SSI might be an important transitional zone between Antarctica and South America.     

Biodiversity issues for the forthcoming tropical Mediterranean Sea

Present-day Mediterranean marine biodiversity is undergoing rapid alteration. Because of the increased occurrence of warm-water biota, it has been said that the Mediterranean is under a process of ‘tropicalization’. This paper analyses the main patterns of the Mediterranean Sea tropicalization and considers briefly its extent and consequences. As happened during previous interglacial phases of the Quaternary, Atlantic water, entering via the Straits of Gibraltar, carries into the Mediterranean species that are prevalently of (sub)tropical affinity. On the other side of the basin, Red Sea species penetrate through the Suez Canal, a phenomenon called lessepsian migration from the name of F. de Lesseps, the French engineer who promoted the cutting of the Canal. Also the many exotic species introduced by humans voluntarily or involuntarily are nearly always typical of warm waters. Climate change combines with Atlantic influx, lessepsian migration and the introduction of exotic species by humans to the establishment of tropical marine biota in the Mediterranean Sea. Present-day warming ultimately favours the spread of warm-water species through direct and indirect effects, and especially by changing water circulation. It is impossible at present to foresee to what extent the exuberance of warm-water species will affect the trophic web and the functioning of marine ecosystems in the Mediterranean Sea of tomorrow. While Mediterranean Sea communities are modifying their pattern of species composition, they do not seem to be acquiring a more marked tropical physiognomy: Mediterranean coastal marine ecosystems are still dominated by frondose algae (even if the species that are gaining ascendancy are of tropical origin) and not by corals as is normal in tropical seas.     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Taxonomy and Biogeography of Apomixis in Angiosperms and Associated Biodiversity Characteristics

Apomixis in angiosperms is asexual reproduction from seed. Its importance to angiospermous evolution and biodiversity has been difficult to assess mainly because of insufficient taxonomic documentation. Thus, we assembled literature reporting apomixis occurrences among angiosperms and transferred the information to an internet database (http://www.apomixis.uni-goettingen.de). We then searched for correlations between apomixis occurrences and well-established measures of taxonomic diversity and biogeography. Apomixis was found to be taxonomically widespread with no clear tendency to specific groups and to occur with sexuality at all taxonomic levels. Adventitious embryony was the most frequent form (148 genera) followed by apospory (110) and diplospory (68). All three forms are phylogenetically scattered, but this scattering is strongly associated with measures of biodiversity. Across apomictic-containing orders and families, numbers of apomict-containing genera were positively correlated with total numbers of genera. In general, apomict-containing orders, families, and subfamilies of Asteraceae, Poaceae, and Orchidaceae were larger, i.e., they possessed more families or genera, than non-apomict-containing orders, families or subfamilies. Furthermore, many apomict-containing genera were found to be highly cosmopolitan. In this respect, 62% occupy multiple geographic zones. Numbers of genera containing sporophytic or gametophytic apomicts decreased from the tropics to the arctic, a trend that parallels general biodiversity. While angiosperms appear to be predisposed to shift from sex to apomixis, there is also evidence of reversions to sexuality. Such reversions may result from genetic or epigenetic destabilization events accompanying hybridization, polyploidy, or other cytogenetic alterations. Because of increased within-plant genetic and genomic heterogeneity, range expansions and diversifications at the species and genus levels may occur more rapidly upon reversion to sexuality. The significantly-enriched representations of apomicts among highly diverse and geographically-extensive taxa, from genera to orders, support this conclusion.