The stratigraphic reinterpretation of the palynologically analysed Miocene succession of the Wijshagen borehole along the southern margin of the North Sea Basin allowed an age assessment – late Burdigalian to early Serravalian – for the Genk Sand Member of the Bolderberg Formation. The depositional environment varied during Burdigalian to Serravalian times from continental (peat formation) to open marine (glauconitic sands), respectively from south to north in the Roer Valley Rift System. The study area of the Wijshagen borehole is located in the central part of the Roer Valley Rift System between these extreme environments. During the Burdigalian, the glauconitic fine clayey sands of the Houthalen Sand Member were deposited in the study area. From the late Burdigalian onwards, the glauconite content decreased and lignite content increased as a result of high influx of clastic material in the Roer Valley Rift System, and marked the start of the deposition of the Genk Sand Member. The Genk Sand Member shows an overall coarsening-upwards trend, which is consistent with the gradual infill of the available accommodation space in the Roer Valley Rift System by northwest-prograding clastic delta sequences. Dinoflagellate cyst analyses indicate that the Genk Sand Member was largely deposited in a marginal marine environment with only short pulses of continental input. These pulses of continental input increase in a southerly or landward direction where they led to the development of thick lignite seams.

Both the deformation and rupture characteristics of rocks are related to geomechanics. In this paper, we identify the evaluation indexes related to coalbed methane (CBM) accumulation in strongly deformed strike-slip fault zones considering tectonics and fractures. We found that fault scale, the fault combination, the tectonic stress, the preservation conditions and fractures all have important effects on the CBM distribution. Areas near the large-scale opening faults are unfavourable to the preservation of coalbed methane. The distribution of gas wells with different capacities is influenced by tectonic extension and convergence. A 3D geomechanical method was used to analyse the influence of the ‘ribbon effect’ of strike-slip faults on the CBM distribution. Due to the influence of the ‘ribbon effect’, the tectonic stress presents a plane in situ stress heterogeneity, which in turn will affect the gas well productivity. We also calculated the integrated rupture rate (IF) to characterize the degree of tectonic fracture development in the target coal reservoir. The appropriate fracture development degree can improve the petrophysical properties of the coal reservoirs while maintaining good storage conditions, such that the gas wells can achieve a higher production capacity. This study is of great significance for the enrichment of the geomechanical theory of oil and gas exploration.

The terminal Ediacaran Dengying Formation (c. 551.1–538.8 Ma) in South China is one of two successions where Ediacara-type macrofossils are preserved in carbonate facies along with skeletal fossils and bilaterian animal traces. Given the remarkable thickness of carbonate-bearing strata deposited in less than 12.3 million years, the Dengying Formation holds the potential for construction of a relatively continuous chemostratigraphic profile for the terminal Ediacaran Period. In this study, a detailed sedimentological and chemostratigraphic (δ13Ccarb, δ18Ocarb, δ13Corg, δ34Spyrite, and 87Sr/86Sr) investigation was conducted on the Dengying Formation at the Gaojiashan section, Ningqiang County of southern Shaanxi Province, South China. Sedimentological results reveal an overall shallow-marine depositional environment. Carbonate breccia, void-filling botryoidal precipitates and aragonite crystal fans are common in the Algal Dolomite Member of the Dengying Formation, suggesting that peritidal facies were repeatedly karstified. The timing of karstification was likely early, probably soon after the deposition of the dolomite sediments. The presence of authigenic aragonite cements suggests high alkalinity in the terminal Ediacaran ocean. Geochemical analysis of micro-drilled samples shows that distinct compositions are registered in different carbonate phases, which should be considered when constructing chemostratigraphic profiles representative of true temporal variations in seawater chemistry. Integrated chemostratigraphic data suggest enhanced burial of organic carbon and pyrite, and the occurrence of extensive marine anoxia (at least in the Gaojiashan Member). Rapid basinal subsidence and carbonate accumulation during a time of elevated seawater alkalinity and increased rates of pyrite burial may have facilitated the evolutionary innovation of early biomineralizing metazoans.

We document the occurrence of micro-diorite magmatic sills, with magmatic enclaves, in the Ceuta Peninsula within metapelites from the Lower Sebtides units (Internal Rif). All magmatic rocks show a primary magmatic mineralogy and geochemical signature diagnostic for high-K calc-alkaline to shoshonitic island arc magmatism. Moreover, these rocks are significantly affected by secondary metamorphic transformations under greenschist- to amphibolite-facies conditions, regionally dated at c. 21 Ma. Geometric relationships between the sills and the main regional foliation, developed under intermediate-pressure granulite-facies conditions at c. 28 Ma, demonstrate that the sills emplaced during the late stage of this main tectonic event. New U–Pb in situ analyses of monazite performed on the micro-diorite sills provide an age of 20.64 ± 0.19 Ma, coherent with this chronological framework and interpreted as the age of greenschist-facies re-equilibration. The discovery of pre-Miocene high-K calc-alkaline to shoshonitic arc-related magmatism is clearly consistent with the subduction context proposed for the Alboran Basin evolution, according to geophysical investigations. In this framework, the Lower Sebtides units could be considered as part of the upper plate of the subduction system, while the Upper Sebtides must be regarded as the lower subducted plate.

Low-pressure regional aureoles with steep metamorphic field gradients are critical to understanding progressive metamorphism in high-temperature metasedimentary rocks. Delicately layered pelitic and psammitic metasedimentary rocks at Mt Stafford, central Australia, record a greenschist- to granulite-facies Palaeoproterozoic regional aureole, associated with S-type granite plutons, reflecting metamorphism in the range 500–800 °C and at ∼3 kbar. The rocks experienced minimal deformation during metamorphism and partial melting. Partial melting textures evolve progressively along the steep metamorphic field gradient from the incipient stages of melting marked by cuspate grains with low dihedral angles, to melt proportions sufficient to form diatexite with schollen. Phase equilibria modelling in the NCKFMASHTO system for pelitic, semi-pelitic and high- and low-ferromagnesian psammitic samples quantitatively illustrates the dependence of partial melting on rock composition and water volume. Pelitic compositions are more fertile than psammitic compositions when the water content in the rocks is low, especially during the early stages of melting. The whole-rock ferromagnesian component additionally influences melt fertility, with ferromagnesian-rich psammite being more fertile than psammite with a lower ferromagnesian component. Subtle variations in free water content can result in obvious changes in melt volume but limited variation in melt composition. Distinct melting histories of pelitic and psammitic rocks inferred from field relationships may be partially attributed to potential differences in water volume retained to super-solidus conditions. Melt composition is more dependent on the rock composition than the variation in water content.

In this paper, zircon U–Pb geochronology, major and trace elements, and Sr–Nd isotope geochemistry of the Baiyanghe dolerites in northern West Junggar of NW China are presented. The U–Pb dating of zircons from the dolerites yielded ages of 272.2±4 Ma and 276.7±6.2 Ma, which indicate the emplacement times. The dolerites are characterized by minor variations in SiO2 (46.89 to 49.07 wt%), high contents of Al2O3 (13.60 to 13.92 wt%) and total Fe2O3 (11.14 to 11.70 wt%), and low contents of MgO (2.67 to 3.64 wt%) and total alkalis (Na2O+K2O, 5.1 to 5.97 wt%, K2O/Na2O = 0.37–0.94), which indicate affinities to metaluminous tholeiite basalt. The REE pattern ((La/Sm)N = 2.25–2.34, (La/Yb)N = 7.42–8.36), V–Ti/1000 and 50*Zr–Ti/50–Sm discrimination diagrams show that these rocks are OIB-type. The high contents of Zr and Ti indicate a within-plate tectonic setting, and samples plot in the ‘plume source’ field shown on the Dy/Yb(N) versus Ce/Yb(N) diagram. The positive εNd(t) values (+7.09 to +7.48), high initial 87Sr/86Sr ratios (0.70442 to 0.70682) and depletions of Nb and Ta elements in the samples can be explained by the involvement of subducted sediments. In summary, it is possible that the Baiyanghe dolerites were derived from an OIB-like mantle source and associated with a mantle plume tectonic setting. Therefore, our samples provide the youngest evidence for the existence of a mantle plume, which may provide new insights into the Late Palaeozoic tectonic setting of West Junggar.

The Lower Cretaceous Wealden sideritic ironstones have a wide occurrence and great potential to aid the reconstruction of the depositional environments of the Weald Basin in SE England. However, mineralogical and geochemical datasets on the ironstones are scarce in the literature. Geochemical and mineralogical data on the sideritic ironstones are presented from the Wadhurst Clay Formation within the Weald Basin. The mineralogy of the ironstones was examined using a PANalytical X'Pert Pro X-ray diffractometer and PANalytical's HighScore Plus software. Elemental composition of the ironstones was measured using a PANalytical MiniPal2 ED-XRF (benchtop X-ray spectrometer). The examination of the mineralogy of the Wealden ironstones confirms the presence of early diagenetic siderites. The trace-element assemblage shows that the sideritic ironstones are chemically pure pointing to a freshwater origin. The sideritic ironstones reveal anoxic conditions and palaeo-salinity in the basin. More generally, it is suggested that the composition of the host rocks has significant controls on the composition of sideritic ironstones in sedimentary basins. This work reinforces the importance of the composition of sideritic ironstones as useful non-traditional data for understanding the depositional settings of sedimentary basins, especially when traditional datasets are not readily available or insufficient.

The Caledonides of Britain and Ireland include terranes attributed to both Laurentian and Gondwanan sources, separated along the Solway line. Gondwanan elements to the south have been variably assigned to the domains Ganderia and East Avalonia. The Midland Platform forms the core of East Avalonia but its provenance is poorly known. Laser ablation split-stream analysis yields information about detrital zircon provenance by providing simultaneous U–Pb and Lu–Hf data from the same ablated volume. A sample of Red Callavia Sandstone from uppermost Cambrian Stage 3 of the Midland Platform yields a U–Pb age spectrum dominated by Neoproterozoic and Palaeoproterozoic sources, resembling those in the Welsh Basin, the Meguma Terrane of Nova Scotia and NW Africa. Initial εHf values suggest that the Neoproterozoic zircon component was derived mainly from crustal sources < 2 Ga, and imply that the more evolved Palaeoproterozoic grains were transported into the basin from an older source terrane, probably the Eburnean Orogen of West Africa. A sample from Cambrian Stage 4 in the Bray Group of the Leinster–Lakesman Terrane shows, in contrast, a distribution of both U–Pb ages and εHf values closely similar to those of the Gander Terrane in Newfoundland and other terranes attributed to Ganderia, interpreted to be derived from the margin of Amazonia. East Avalonia is clearly distinct from Ganderia, but shows evidence for older crustal components not present in West Avalonia of Newfoundland. These three components of the Appalachian–Caledonide Orogen came from distinct sources on the margin of Cambrian Gondwana.

The recently uplifted and exposed Pliocene and Pleistocene sedimentary infill of the neotectonic Polis graben provides an excellent opportunity to understand extensional basin development in a marine setting. Fieldwork, facies analysis and dating using nannofossils and strontium isotopes reveal how the sedimentary conditions evolved during infill of the Polis graben during Pliocene and Pleistocene time, and allow a composite succession for the depocentre to be determined for the first time. Six lithofacies are recognized in the northern Polis graben, allowing evolving palaeoenvironments to be inferred. By the end of Miocene time (Messinian) a major c. N–S-trending graben was established; extensional faulting continued during the Pliocene–Pleistocene until recent time. Post-Messinian salinity crisis deposition began with deposition of hemipelagic muds (c. 5.08–2.76 Ma), equivalent to the Nicosia Formation. This was followed by upwards incoming of repeated normal-graded bioclastic carbonates (couplets) (c. 2.76–1.6 Ma), which are interpreted as age-equivalents of the Athalassa Formation elsewhere in Cyprus. The upwards sudden facies change is explained by tectonically controlled shallowing which enabled neritic carbonate production on the basin margins. The appearance of basement-derived material (e.g. ophiolitic extrusive detritus) in the highest stratigraphic levels of the basin fill in the north (c. 1.7–1.6 Ma) reflects onset of rapid surface uplift focused on the Troodos ophiolitic massif. Overall, the syntectonic basin infill appears to document a two-stage, pulsed uplift related to early-stage collision of the African and Eurasian plates in the easternmost Mediterranean region.

The Upper Cretaceous Kanguk Formation of the Sverdrup Basin, Canadian Arctic Islands, contains numerous diagenetically altered volcanic ash layers (bentonites). Eleven bentonites were sampled from an outcrop section on Ellesmere Island for U–Pb zircon secondary ion mass spectrometry dating and whole-rock geochemical analysis. Two distinct types of bentonite are identified from the geochemical data. Relatively thick (0.1 to 5 m) peralkaline rhyolitic to trachytic bentonites erupted in an intraplate tectonic setting. These occur throughout the upper Turonian to lower Campanian (c. 92–83 Ma) outcrop section and are likely associated with the alkaline phase of the High Arctic Large Igneous Province. Two thinner (<5 cm) subalkaline dacitic to rhyolitic bentonites of late Turonian to early Coniacian age (c. 90–88 Ma) are also identified. The geochemistry of these bentonites is consistent with derivation from volcanoes within an active continental margin tectonic setting. The lack of nearby potential sources of subalkaline magmatism, together with the thinner bed thickness of the subalkaline bentonites and the small size of zircon phenocrysts therein (typically 50–80 μm in length) are consistent with a more distal source area. The zircon U–Pb age and whole-rock geochemistry of these two subalkaline bentonites correlate with an interval of intense volcanism in the Okhotsk–Chukotka Volcanic Belt, Russia. It is proposed that during late Turonian to early Coniacian times intense volcanism within the Okhotsk–Chukotka Volcanic Belt resulted in widespread volcanic ash dispersal across Arctic Alaska and Canada, reaching as far east as the Sverdrup Basin, more than 3000 km away.

An Ordovician–Silurian boundary section marked by an uninterrupted, relatively high rate of black shale sedimentation and abundant, diverse graptolites is described from the south-central Pyrenees. The structurally simple Estana section comprises the uppermost part of the quartzite-dominated Bar Formation and overlying black shales of late Hirnantian and early Rhuddanian age, which have been dated by graptolites to the upper Metabolograptus persculptus and lower–middle Akidograptus ascensus–Parakidograptus acuminatus biozones. Due to the absence of M. persculptus, a Metabolograptus parvulus Biozone correlative with the upper part of the persculptus Biozone is recognized below the lowest occurrence of akidograptids, which indicate the base of the Silurian System. The graptolite fauna comprise 27 species including Normalograptus minor, N. lubricus, N. rhizinus, Hirsutograptus, Korenograptus bifurcus, K. bicaudatus, K. lanpherei and Nd. shanchongensis, most of which were formerly considered to be endemic to the low-latitude palaeobiogeographical province of China, Siberia and northern North America. Two new species, N. baridaensis and N. ednae, are described. The succession of graptolite assemblages in the Estana section, and occurrence of several cosmopolitan taxa in its parvulus and lower ascensus–acuminatus biozones that are unknown elsewhere in peri-Gondwanan Europe, suggest that strata immediately surrounding the Ordovician–Silurian boundary may be absent, highly condensed or oxic and barren of graptolites in other sections of northwestern peri-Gondwana. Common graptolite synrhabdosomes and abnormal rhabdosomes may indicate some environmental stress in the parvulus Biozone, although the rather uniform black shale lithology, total organic carbon content and δ13Corg values suggest uninterrupted sedimentation under stable, anoxic conditions.

We describe here a new turtle from the early Eocene of Wutu, Shandong Province, China. This turtle with a full row of well-developed inframarginal scutes is assigned to the basalmost testudinoids while stem testudinoids were believed to disappear by the Palaeocene–Eocene boundary. This account shows that stem testudinoids crossed this boundary in their original range. The first comprehensive phylogenetic analysis of stem and modern testudinoids performed here demonstrates that the stem testudinoids, previously placed in the family ‘Lindholmemydidae’, do not form a monophyletic group, and the two major clades of testudinoids (Emydidae and Geoemydidae+Testudinidae) split one from another well before the Palaeocene/Eocene boundary, prior to the Late Cretaceous.

Palaeogeographic reconstructions have been proposed for years. The technique employed, however, is more or less always the same: it consists of determining the palaeoenvironment at the local scale and extending it to the regional scale. Such work is carried out in a maximum number of locations all over the planet and the global palaeogeography is the result of interpolation of those reconstructions. Advances in palaeogeography can be made via an alternative way, which consists of integrating and then coupling various global models. It results in the proposal of synthetic palaeogeographies that can be compared a posteriori to local or regional data. The advantage is twofold: (1) the view is really global and it avoids gaps (in particular in the oceanic realm) in the reconstructions, and it is very much less focused on the coastline; (2) it takes advantages from almost all the fields of geosciences, so that reconstructions can be constrained from a large variety of data. The two techniques – the ‘classic’ and the ‘alternative’ – are not contradictory but complementary, and it is desirable that one feeds the other and the study of palaeogeography be revived.

Eight latest Eocene to earliest Miocene stratigraphic surfaces have been identified in petroleum well data from the Taranaki Basin, New Zealand. These surfaces define seven regional sedimentary packages, of variable thickness and lithofacies, forming a mixed siliciclastic–carbonate system. The evolving tectonic setting, particularly the initial development of the Australian–Pacific convergent margin, controlled geographic, stratigraphic and facies variability. This tectonic signal overprinted a regional transgressive trend that culminated in latest Oligocene times. The earliest influence of active compressional tectonics is reflected in the preservation of latest Eocene – Early Oligocene deepwater sediments in the northern Taranaki Basin. Thickness patterns for all mid Oligocene units onwards show a shift in sedimentation to the eastern Taranaki Basin, controlled by reverse movement on the Taranaki Fault System. This resulted in the deposition of a thick sedimentary wedge, initially of coarse clastic sediments, later carbonate dominated, in the foredeep close to the fault. In contrast, Oligocene active normal faulting in a small sub-basin in the south may represent the most northerly evidence for rifting in southern Zealandia, related to Emerald Basin formation. The Early Miocene period saw a return to clastic-dominated deposition, the onset of regional regression and the southward propagation of compressional tectonics.

A pycnodont premaxilla bearing a large incisiform tooth with an unusual crown morphology is described from the Palaeocene (Thanetian) phosphate deposits of Morocco. This peculiar tooth shows a broad, fan-shaped multicuspid crown with nine cusps, well adapted for benthic macroalgal scraping. This morph, assigned to a new species of Pycnodus (P. multicuspidatus sp. nov.), emphasizes the phenotypic plasticity of the group and documents an additional trophic specialization among Palaeogene pycnodontiform fishes. In the post-K/Pg boundary marine ecosystem of the Ouled Abdoun Basin, P. multicuspidatus sp. nov. may have opportunistically replaced Maastrichtian fish taxa with a similar front dentition and feeding behaviour, such as the putative specialized pycnodontiforms Stephanodus and Hadrodus.

The first find of microdiamond in the Nordøyane ultra-high-pressure (UHP) domain of the Western Gneiss Region (WGR) of the Scandinavian Caledonides reshaped tectonic models for the region. Nevertheless, in spite of much progress regarding the meaning and significance of this find, the history of rock that the diamonds were found in is complex and still largely ambiguous. To investigate this, we report U–Pb zircon ages obtained from the exact crushed sample material in which metamorphic diamond was first found. The grains exhibit complicated internal zoning with distinct detrital cores overgrown by metamorphic rims. The cores yielded a range of ages from the Archaean to the late Neoproterozoic / early Cambrian. This detrital zircon age spectrum is broadly similar to detrital signatures recorded by metasedimentary rocks of the Lower and Middle allochthons elsewhere within the orogen. Thus, our dating results support the previously proposed affinity of the studied gneiss to the Seve–Blåhø Nappe of the Middle Allochthon. Metamorphic rims yielded a well-defined peak at 447 ± 2 Ma and a broad population that ranges between c. 437 and 423 Ma. The data reveal a prolonged metamorphic history of the Fjørtoft gneiss that is far more complex then would be expected for a UHP rock that has seen a single burial and exhumation cycle. The data are consistent with a model involving multiple such cycles, which would provide renewed support for the dunk tectonics model that has been postulated for the region.

The abundance of specimens of Ichthyosaurus provides an opportunity to assess morphological variation without the limits of a small sample size. This research evaluates the variation and taxonomic utility of hindfin morphology. Two seemingly distinct morphotypes of the mesopodium occur in the genus. Morphotype 1 has three elements in the third row: metatarsal two, distal tarsal three and distal tarsal four. This is the common morphology in Ichthyosaurus breviceps, I. conybeari and I. somersetensis. Morphotype 2 has four elements in the third row, owing to a bifurcation. This morphotype occurs in at least some specimens of each species, but it has several variations distinguished by the extent of contact of elements in the third row with the astragalus. Two specimens display a different morphotype in each fin, suggesting that the difference reflects individual variation. In Ichthyosaurus, the hindfin is taxonomically useful at the genus level, but species cannot be identified unequivocally from a well-preserved hindfin, although certain morphologies are more common in certain species than others. The large sample size filled in morphological gaps between what initially appeared to be taxonomically distinct characters. The full picture of variation would have been obscured with a small sample size. Furthermore, we have found several unusual morphologies which, in isolation, could have been mistaken for new taxa. Thus, one must be cautious when describing new species or genera on the basis of limited material, such as isolated fins and fragmentary specimens.

Radiodonta, apex Cambrian predators such as Anomalocaris have been known from the Kinzers Formation (Cambrian Series 2, Stage 4 – Pennsylvania, USA) for nearly 100 years. Work over the last ten years, mainly on radiodont material from the Chengjiang (Cambrian Series 2, Stage 3 – Yunnan, China) and Burgess Shale (Miaolingian, Wuliuan – British Columbia, Canada), has greatly improved our knowledge of the diversity and disparity of radiodonts and their frontal appendages, including the description of new species, genera and families. Previous work identified two species of radiodonts from the Kinzers Formation: Anomalocaris pennsylvanica Resser, 1929 and Anomalocaris? cf. pennsylvanica based on isolated frontal appendage material (Briggs, 1979). A restudy of Kinzers Formation material shows that only some of the specimens can be confirmed as Anomalocaris pennsylvanica, and a number of specimens previously attributed to Anomalocaris in fact belong to other more recently discovered radiodont genera Amplectobelua and Tamisiocaris. This reinterpretation makes the Kinzers Formation the most diverse Cambrian Stage 4 Burgess Shale Type Lagerstätten in terms of number of radiodont species. This assemblage includes the youngest known Tamisiocaris and the first from outside Greenland, the only Amplectobelua from Stage 4 and the oldest from Laurentia, two specimens tentatively assigned to the recently described Chengjiang genus Laminacaris, and the endemic Anomalocaris pennsylvanica. The identification of these new radiodont taxa increases the total known diversity of the Kinzers Formation to more than ten species, and so it should now be considered a Tier 2 Lagerstätte.

Whether the latitudinal distribution of climate-sensitive lithologies is stable through greenhouse and icehouse regimes remains unclear. Previous studies suggest that the palaeolatitudinal distribution of palaeoclimate indicators, including coals, evaporites, reefs and carbonates, has remained broadly similar since the Permian period, leading to the conclusion that atmospheric and oceanic circulation control their distribution rather than the latitudinal temperature gradient. Here we revisit a global-scale compilation of lithologic indicators of climate, including coals, evaporites and glacial deposits, back to the Devonian period. We test the sensitivity of their latitudinal distributions to the uneven distribution of continental areas through time and to global tectonic models, correct the latitudinal distributions of lithologies for sampling- and continental area-bias, and use statistical methods to fit these distributions with probability density functions and estimate their high-density latitudinal ranges with 50% and 95% confidence intervals. The results suggest that the palaeolatitudinal distributions of lithologies have changed through deep geological time, notably a pronounced poleward shift in the distribution of coals at the beginning of the Permian. The distribution of evaporites indicates a clearly bimodal distribution over the past ~400 Ma, except for Early Devonian, Early Carboniferous, the earliest Permian and Middle and Late Jurassic times. We discuss how the patterns indicated by these lithologies change through time in response to plate motion, orography, evolution and greenhouse/icehouse conditions. This study highlights that combining tectonic reconstructions with a comprehensive lithologic database and novel data analysis approaches provide insights into the nature and causes of shifting climatic zones through deep time.

The transition between the seemingly disparate Ediacaran and Cambrian faunas is both enigmatic and body-fossil poor. The Chapel Island Formation on the Burin Peninsula, Newfoundland, Canada, contains a rich diversity of ichnofossils, providing new insight into the nature of the Ediacaran–Cambrian transition and early Fortunian ecosystems. Five ichnoguilds are recognized within the Treptichnus pedum zone. Ichnologic data are analysed from an ecospace perspective, revealing a more protracted transition between Ediacaran and Cambrian ecosystems. Our analysis documents the appearance of limbs, vertical burrows and uncontroversial equilibrium structures, as well as the retention of ‘other’ feeding styles, such as microbial grazing and chemosynthesis.