关于底栖组合应用中若干问题的讨论

由于地质历史时期环境多变,同一深度带生境未必均一,加之生物自身对各环境因素的适应能力各不相同,因而各底栖组合的界线应有一定限度的变化范围。在正常情况下,每个底栖组合一般都有明确的岩性、岩相和生物群标志。研究者可以通过综合研究群落的组成、特征、埋藏特点和围岩特点等,确定每个群落所属的底栖生态位。生境的含氧性和构造环境等因素也可使底栖组合的正常分布受到干扰。     

General patterns of taxonomic and biomass partitioning in extant and fossil plant communities

A central goal of evolutionary ecology is to identify the general features maintaining the diversity of species assemblages. Understanding the taxonomic and ecological characteristics of ecological communities provides a means to develop and test theories about the processes that regulate species coexistence and diversity. Here, using data from woody plant communities from different biogeographic regions, continents and geologic time periods, we show that the number of higher taxa is a general power-function of species richness that is significantly different from randomized assemblages. In general, we find that local communities are characterized by fewer higher taxa than would be expected by chance. The degree of taxonomic diversity is influenced by modes of dispersal and potential biotic interactions. Further, changes in local diversity are accompanied by regular changes in the partitioning of community biomass between taxa that are also described by a power function. Our results indicate that local and regional processes have consistently regulated community diversity and biomass partitioning for millions of years.     

Major viral impact on the functioning of benthic deep-sea ecosystems

Viruses are the most abundant biological organisms of the world's oceans. Viral infections are a substantial source of mortality in a range of organisms-including autotrophic and heterotrophic plankton-but their impact on the deep ocean and benthic biosphere is completely unknown. Here we report that viral production in deep-sea benthic ecosystems worldwide is extremely high, and that viral infections are responsible for the abatement of 80% of prokaryotic heterotrophic production. Virus-induced prokaryotic mortality increases with increasing water depth, and beneath a depth of 1,000 m nearly all of the prokaryotic heterotrophic production is transformed into organic detritus. The viral shunt, releasing on a global scale approximately 0.37-0.63 gigatonnes of carbon per year, is an essential source of labile organic detritus in the deep-sea ecosystems. This process sustains a high prokaryotic biomass and provides an important contribution to prokaryotic metabolism, allowing the system to cope with the severe organic resource limitation of deep-sea ecosystems. Our results indicate that viruses have an important role in global biogeochemical cycles, in deep-sea metabolism and the overall functioning of the largest ecosystem of our biosphere.     

Ploughing the deep sea floor

Bottom trawling is a non-selective commercial fishing technique whereby heavy nets and gear are pulled along the sea floor. The direct impact of this technique on fish populations and benthic communities has received much attention, but trawling can also modify the physical properties of seafloor sediments, water–sediment chemical exchanges and sediment fluxes. Most of the studies addressing the physical disturbances of trawl gear on the seabed have been undertaken in coastal and shelf environments, however, where the capacity of trawling to modify the seafloor morphology coexists with high-energy natural processes driving sediment erosion, transport and deposition. Here we show that on upper continental slopes, the reworking of the deep sea floor by trawling gradually modifies the shape of the submarine landscape over large spatial scales. We found that trawling-induced sediment displacement and removal from fishing grounds causes the morphology of the deep sea floor to become smoother over time, reducing its original complexity as shown by high-resolution seafloor relief maps. Our results suggest that in recent decades, following the industrialization of fishing fleets, bottom trawling has become an important driver of deep seascape evolution. Given the global dimension of this type of fishery, we anticipate that the morphology of the upper continental slope in many parts of the world’s oceans could be altered by intensive bottom trawling, producing comparable effects on the deep sea floor to those generated by agricultural ploughing on land.     

Evidence for deep-water production in the North Pacific Ocean during the early Cenozoic warm interval

The deep-ocean circulation is responsible for a significant component of global heat transport. In the present mode of circulation, deep waters form in the North Atlantic and Southern oceans where surface water becomes sufficiently cold and dense to sink. Polar temperatures during the warmest climatic interval of the Cenozoic era (approximately 65 to 40 million years (Myr) ago) were significantly warmer than today, and this may have been a consequence of enhanced oceanic heat transport. However, understanding the relationship between deep-ocean circulation and ancient climate is complicated by differences in oceanic gateways, which affect where deep waters form and how they circulate. Here I report records of neodymium isotopes from two cores in the Pacific Ocean that indicate a shift in deep-water production from the Southern Ocean to the North Pacific approximately 65 Myr ago. The source of deep waters reverted back to the Southern Ocean 40 Myr ago. The relative timing of changes in the neodymium and oxygen isotope records indicates that changes in Cenozoic deep-water circulation patterns were the consequence, not the cause, of extreme Cenozoic warmth.     

Rapid and early export of Phaeocystis antarctica blooms in the Ross Sea, Antarctica

The Southern Ocean is very important for the potential sequestration of carbon dioxide in the oceans and is expected to be vulnerable to changes in carbon export forced by anthropogenic climate warming. Annual phytoplankton blooms in seasonal ice zones are highly productive and are thought to contribute significantly to pCO2 drawdown in the Southern Ocean. Diatoms are assumed to be the most important phytoplankton class with respect to export production in the Southern Ocean; however, the colonial prymnesiophyte Phaeocystis antarctica regularly forms huge blooms in seasonal ice zones and coastal Antarctic waters. There is little evidence regarding the fate of carbon produced by P. antarctica in the Southern Ocean, although remineralization in the upper water column has been proposed to be the main pathway in polar waters. Here we present evidence for early and rapid carbon export from P. antarctica blooms to deep water and sediments in the Ross Sea. Carbon sequestration from P. antarctica blooms may influence the carbon cycle in the Southern Ocean, especially if projected climatic changes lead to an alteration in the structure of the phytoplankton community.     

Flushing submarine canyons

The continental slope is a steep, narrow fringe separating the coastal zone from the deep ocean. During low sea-level stands, slides and dense, sediment-laden flows erode the outer continental shelf and the continental slope, leading to the formation of submarine canyons that funnel large volumes of sediment and organic matter from shallow regions to the deep ocean(1). During high sea-level stands, such as at present, these canyons still experience occasional sediment gravity flows(2-5), which are usually thought to be triggered by sediment failure or river flooding. Here we present observations from a submarine canyon on the Gulf of Lions margin, in the northwest Mediterranean Sea, that demonstrate that these flows can also be triggered by dense shelf water cascading (DSWC)-a type of current that is driven solely by seawater density contrast. Our results show that DSWC can transport large amounts of water and sediment, reshape submarine canyon floors and rapidly affect the deep-sea environment. This cascading is seasonal, resulting from the formation of dense water by cooling and/or evaporation, and occurs on both high- and low-latitude continental margins(6-8). DSWC may therefore transport large amounts of sediment and organic matter to the deep ocean. Furthermore, changes in the frequency and intensity of DSWC driven by future climate change may have a significant impact on the supply of organic matter to deep-sea ecosystems and on the amount of carbon stored on continental margins and in ocean basins.     

Coral communities are regionally enriched along an oceanic biodiversity gradient

Ecological communities are influenced by processes operating at multiple scales. Thus, a better understanding of how broad- as well as local-scale processes affect species diversity and richness is increasingly becoming a central focus in modern community ecology. Here, in a study of unprecedented geographical scope, we show significant regional and local variation in the species richness of coral assemblages across an oceanic biodiversity gradient. The gradient that we sampled extends 10,000 km eastwards from the world's richest coral biodiversity hotspot in the central Indo-Pacific. Local richness and the size of regional species pools decline significantly across 15 islands spanning the gradient. In addition, richness declines across three adjacent habitats (reef slopes, crests and flats). In each habitat, a highly consistent linear relationship between local and regional species richness indicates strong regional enrichment. Thus, even on the most diverse coral reefs in the world, local coral assemblages are profoundly affected by regional-scale processes. Understanding these historical and biogeographical influences is essential for the effective management and preservation of these endangered communities.     

Stratified prokaryote network in the oxic-anoxic transition of a deep-sea halocline

The chemical composition of the Bannock basin has been studied in some detail. We recently showed that unusual microbial populations, including a new division of Archaea (MSBL1), inhabit the NaCl-rich hypersaline brine. High salinities tend to reduce biodiversity, but when brines come into contact with fresher water the natural haloclines formed frequently contain gradients of other chemicals, including permutations of electron donors and acceptors, that may enhance microbial diversity, activity and biogeochemical cycling. Here we report a 2.5-m-thick chemocline with a steep NaCl gradient at 3.3 km within the water column betweeen Bannock anoxic hypersaline brine and overlying sea water. The chemocline supports some of the most biomass-rich and active microbial communities in the deep sea, dominated by Bacteria rather than Archaea, and including four major new divisions of Bacteria. Significantly higher metabolic activities were measured in the chemocline than in the overlying sea water and underlying brine; functional analyses indicate that a range of biological processes is likely to occur in the chemocline. Many prokaryotic taxa, including the phylogenetically new groups, were confined to defined salinities, and collectively formed a diverse, sharply stratified, deep-sea ecosystem with sufficient biomass to potentially contribute to organic geological deposits.     

Ocean nutrient ratios governed by plankton biogeography

The major nutrients nitrate and phosphate have one of the strongest correlations in the sea, with a slope similar to the average nitrogen (N) to phosphorus (P) content of plankton biomass (N/P = 16:1). The processes through which this global relationship emerges despite the wide range of N/P ratios at the organism level are not known. Here we use an ocean circulation model and observed nutrient distributions to show that the N/P ratio of biological nutrient removal varies across latitude in Southern Ocean surface waters, from 12:1 in the polar ocean to 20:1 in the sub-Antarctic zone. These variations are governed by regional differences in the species composition of the plankton community. The covariation of dissolved nitrate and phosphate is maintained by ocean circulation, which mixes the shallow subsurface nutrients between distinct biogeographic provinces. Climate-driven shifts in these marine biomes may alter the mean N/P ratio and the associated carbon export by Southern Ocean ecosystems.     

Oceanic Cd/P ratio and nutrient utilization in the glacial Southern Ocean

During glacial periods, low atmospheric carbon dioxide concentration has been associated with increased oceanic carbon uptake, particularly in the southern oceans. The mechanism involved remains unclear. Because ocean productivity is strongly influenced by nutrient levels, palaeo-oceanographic proxies have been applied to investigate nutrient utilization in surface water across glacial transitions. Here we show that present-day cadmium and phosphorus concentrations in the global oceans can be explained by a chemical fractionation during particle formation, whereby uptake of cadmium occurs in preference to uptake of phosphorus. This allows the reconstruction of past surface water phosphate concentrations from the cadmium/calcium ratio of planktonic foraminifera. Results from the Last Glacial Maximum show similar phosphate utilization in the subantarctic to that of today, but much smaller utilization in the polar Southern Ocean, in a model that is consistent with the expansion of glacial sea ice and which can reconcile all proxy records of polar nutrient utilization. By restricting communication between the ocean and atmosphere, sea ice expansion also provides a mechanism for reduced CO2 release by the Southern Ocean and lower glacial atmospheric CO2.     

Cenozoic stratigraphy of Taiwan: Window into rifting, stratigraphy and paleoceanography of South China Sea

Shallow marine sequences of the northern South China Sea (SCS) are uplifted and exposed by plate convergence in the Taiwan mountain belt. These deposits provide detailed geological information about the rifting event, stratigraphy, sedimentology, paleoclimate and paleoceanography of the shallow SCS to compare with what are recorded in the ODP 1148 deep-sea core. Seismic surveys and marine micropalentological studies show that Eocene sequences in the offshore Taiwan Strait and onland Taiwan mountain belt are all deposited in rifting basins and are covered unconformably by the Late Oligocene-Neogene post-rifting strata. Between syn-rifting and post-rifting sequences, there is a regional break-up unconformity throughout the island. Early Oligocene and Late Eocene strata are missing along the break-up unconformity equivalent to the T7 unconformity in the Pearl River Mouth Basin off south China. This may suggest that the SCS oceanic crust could have initiated between 33 and 39 Ma. Neither obvious stratigraphic gap nor slumping features are found in the Oligocene-Miocene transition interval of Taiwan. This observation highly contrasts with what has been documented from the ODP 1148 deep-sea core. This suggests that the stratigraphic gap and slumping features could only be recorded in the SCS deep sea region, but not in the shallow shelf near Taiwan. Compared to the Middle Miocene paleoceanographic re-organization events in the SCS deep sea, the geological history of the Taiwan shallow sequence shows changes of in sedimentation and faunal composition. Due to the Antarctic glacial expansion at~14 Ma, Middle to late Miocene strata of the Western Foothills show progressive regression sedimentations associated with a decrease of benthic foraminif-eral abundance and a sharp faunal turnover event. Many Early-Middle Miocene endemic benthic foraminifers were extinct in 14-13 Ma and new benthic foraminifers of the Kuroshio Current fauna appeared from 10.2 Ma, comparable with new occurrence of Modern benthic foraminifers at 9 Ma in the Java Sea area. This reveals that the Western Boundary Kuroshio Current in the North Pacific could initiate from 10-9 Ma due to closures of the Indo-Pacific seaways by convergent tectonics between the Australian Continent and the Indonesian Arc in 12-8 Ma. Subduction of the SCS oceanic lithosphere since the Middle Miocene resulted in formation of the Hengchun Ridge accretionary prism and the North Luzon Arc. Occurrence of these two bathymetric highs ( 2400 m) since the Middle Miocene and closures of the inter-arc passages in the North Luzon arc in the last 3.5 Ma would control the water exchanges between the West Pacific and the deep SCS. Accordingly, the tectonic evolution in the Central Range-Hengchun Peninsula accretionary prism and the arc-forearc Coastal Range not only control directly the route for water exchanges between the West Pacific and the SCS, but also indirectly shows a great influence on the geochemistry of deep SCS waters. The latter is best shown by much negative carbon isotope values of benthic foraminifers in the ODP 1148 deep-sea core than the West Pacific records in the last 14 Ma.     

Bacterial carbon processing by generalist species in the coastal ocean

The assimilation and mineralization of dissolved organic carbon (DOC) by marine bacterioplankton is a major process in the ocean carbon cycle. However, little information exists on the specific metabolic functions of participating bacteria and on whether individual taxa specialize on particular components of the marine DOC pool. Here we use experimental metagenomics to show that coastal communities are populated by taxa capable of metabolizing a wide variety of organic carbon compounds. Genomic DNA captured from bacterial community subsets metabolizing a single model component of the DOC pool (either dimethylsulphoniopropionate or vanillate) showed substantial overlap in gene composition as well as a diversity of carbon-processing capabilities beyond the selected phenotypes. Our direct measure of niche breadth for bacterial functional assemblages indicates that, in accordance with ecological theory, heterogeneity in the composition and supply of organic carbon to coastal oceans may favour generalist bacteria. In the important interplay between microbial community structure and biogeochemical cycling, coastal heterotrophic communities may be controlled less by transient changes in the carbon reservoir that they process and more by factors such as trophic interactions and physical conditions.     

Intensified deep Pacific inflow and ventilation in Pleistocene glacial times

The production of cold, deep waters in the Southern Ocean is an important factor in the Earth's heat budget. The supply of deep water to the Pacific Ocean is presently dominated by a single source, the deep western boundary current east of New Zealand. Here we use sediment records deposited under the influence of this deep western boundary current to reconstruct deep-water properties and speed changes during the Pleistocene epoch. In physical and isotope proxies we find evidence for intensified deep Pacific Ocean inflow and ventilation during the glacial periods of the past 1.2 million years. The changes in throughflow may be directly related to an increased production of Antarctic Bottom Water during glacial times. Possible causes for such an increased bottom-water production include increasing wind strengths in the Southern Ocean or an increase in annual sea-ice formation, leaving dense water after brine rejection and thereby enhancing deep convection. We infer also that the global thermohaline circulation was perturbed significantly during the mid-Pleistocene climate transition between 0.86 and 0.45 million years ago.     

Episodic fresh surface waters in the Eocene Arctic Ocean

It has been suggested, on the basis of modern hydrology and fully coupled palaeoclimate simulations, that the warm greenhouse conditions that characterized the early Palaeogene period (55-45 Myr ago) probably induced an intensified hydrological cycle with precipitation exceeding evaporation at high latitudes. Little field evidence, however, has been available to constrain oceanic conditions in the Arctic during this period. Here we analyse Palaeogene sediments obtained during the Arctic Coring Expedition, showing that large quantities of the free-floating fern Azolla grew and reproduced in the Arctic Ocean by the onset of the middle Eocene epoch (approximately 50 Myr ago). The Azolla and accompanying abundant freshwater organic and siliceous microfossils indicate an episodic freshening of Arctic surface waters during an approximately 800,000-year interval. The abundant remains of Azolla that characterize basal middle Eocene marine deposits of all Nordic seas probably represent transported assemblages resulting from freshwater spills from the Arctic Ocean that reached as far south as the North Sea. The termination of the Azolla phase in the Arctic coincides with a local sea surface temperature rise from approximately 10 degrees C to 13 degrees C, pointing to simultaneous increases in salt and heat supply owing to the influx of waters from adjacent oceans. We suggest that onset and termination of the Azolla phase depended on the degree of oceanic exchange between Arctic Ocean and adjacent seas.     

Reduced North Atlantic Deep Water flux to the glacial Southern Ocean inferred from neodymium isotope ratios

The global circulation of the oceans and the atmosphere transports heat around the Earth. Broecker and Denton suggested that changes in the global ocean circulation might have triggered or enhanced the glacial-interglacial cycles. But proxy data for past circulation taken from sediment cores in the South Atlantic Ocean have yielded conflicting interpretations of ocean circulation in glacial times--delta13C variations in benthic foraminifera support the idea of a glacial weakening or shutdown of North Atlantic Deep Water production, whereas other proxies, such as Cd/Ca, Ba/Ca and 231Pa/230Th ratios, show little change from the Last Glacial Maximum to the Holocene epoch. Here we report neodymium isotope ratios from the dispersed Fe-Mn oxide component of two southeast Atlantic sediment cores. Both cores show variations that tend towards North Atlantic signatures during the warm marine isotope stages 1 and 3, whereas for the full glacial stages 2 and 4 they are closer to Pacific Ocean signatures. We conclude that the export of North Atlantic Deep Water to the Southern Ocean has resembled present-day conditions during the warm climate intervals, but was reduced during the cold stages. An increase in biological productivity may explain the various proxy data during the times of reduced North Atlantic Deep Water export.     

Polar ocean stratification in a cold climate

The low-latitude ocean is strongly stratified by the warmth of its surface water. As a result, the great volume of the deep ocean has easiest access to the atmosphere through the polar surface ocean. In the modern polar ocean during the winter, the vertical distribution of temperature promotes overturning, with colder water over warmer, while the salinity distribution typically promotes stratification, with fresher water over saltier. However, the sensitivity of seawater density to temperature is reduced as temperature approaches the freezing point, with potential consequences for global ocean circulation under cold climates. Here we present deep-sea records of biogenic opal accumulation and sedimentary nitrogen isotopic composition from the Subarctic North Pacific Ocean and the Southern Ocean. These records indicate that vertical stratification increased in both northern and southern high latitudes 2.7 million years ago, when Northern Hemisphere glaciation intensified in association with global cooling during the late Pliocene epoch. We propose that the cooling caused this increased stratification by weakening the role of temperature in polar ocean density structure so as to reduce its opposition to the stratifying effect of the vertical salinity distribution. The shift towards stratification in the polar ocean 2.7 million years ago may have increased the quantity of carbon dioxide trapped in the abyss, amplifying the global cooling.     

Chameleon radiation by oceanic dispersal

Historical biogeography is dominated by vicariance methods that search for a congruent pattern of fragmentation of ancestral distributions produced by shared Earth history. A focus of vicariant studies has been austral area relationships and the break-up of the supercontinent Gondwana. Chameleons are one of the few extant terrestrial vertebrates thought to have biogeographic patterns that are congruent with the Gondwanan break-up of Madagascar and Africa. Here we show, using molecular and morphological evidence for 52 chameleon taxa, support for a phylogeny and area cladogram that does not fit a simple vicariant history. Oceanic dispersal--not Gondwanan break-up--facilitated species radiation, and the most parsimonious biogeographic hypothesis supports a Madagascan origin for chameleons, with multiple 'out-of-Madagascar' dispersal events to Africa, the Seychelles, the Comoros archipelago, and possibly Reunion Island. Although dispersal is evident in other Indian Ocean terrestrial animal groups, our study finds substantial out-of-Madagascar species radiation, and further highlights the importance of oceanic dispersal as a potential precursor for speciation.     

Archaeal dominance in the mesopelagic zone of the Pacific Ocean

The ocean's interior is Earth's largest biome. Recently, cultivation-independent ribosomal RNA gene surveys have indicated a potential importance for archaea in the subsurface ocean. But quantitative data on the abundance of specific microbial groups in the deep sea are lacking. Here we report a year-long study of the abundance of two specific archaeal groups (pelagic euryarchaeota and pelagic crenarchaeota) in one of the ocean's largest habitats. Monthly sampling was conducted throughout the water column (surface to 4,750 m) at the Hawai'i Ocean Time-series station. Below the euphotic zone (> 150 m), pelagic crenarchaeota comprised a large fraction of total marine picoplankton, equivalent in cell numbers to bacteria at depths greater than 1,000 m. The fraction of crenarchaeota increased with depth, reaching 39% of total DNA-containing picoplankton detected. The average sum of archaea plus bacteria detected by rRNA-targeted fluorescent probes ranged from 63 to 90% of total cell numbers at all depths throughout our survey. The high proportion of cells containing significant amounts of rRNA suggests that most pelagic deep-sea microorganisms are metabolically active. Furthermore, our results suggest that the global oceans harbour approximately 1.3 x 10(28) archaeal cells, and 3.1 x 10(28) bacterial cells. Our data suggest that pelagic crenarchaeota represent one of the ocean's single most abundant cell types.     

Patterns and processes in reef fish diversity

A central aim of ecology is to explain the heterogeneous distribution of biodiversity on earth. As expectations of diversity loss grow, this understanding is also critical for effective management and conservation. Although explanations for biodiversity patterns are still a matter for intense debate, they have often been considered to be scale-dependent. At large geographical scales, biogeographers have suggested that variation in species richness results from factors such as area, temperature, environmental stability, and geological processes, among many others. From the species pools generated by these large-scale processes, community ecologists have suggested that local-scale assembly of communities is achieved through processes such as competition, predation, recruitment, disturbances and immigration. Here we analyse hypotheses on speciation and dispersal for reef fish from the Indian and Pacific oceans and show how dispersal from a major centre of origination can simultaneously account for both large-scale gradients in species richness and the structure of local communities.