A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda)

Background The infraorder Anomura has long captivated the attention of evolutionary biologists due to its impressive morphological diversity and ecological adaptations. To date, 2500 extant species have been described but phylogenetic relationships at high taxonomic levels remain unresolved. Here, we reconstruct the evolutionary history—phylogeny, divergence times, character evolution and diversification—of this speciose clade. For this purpose, we sequenced two mitochondrial (16S and 12S) and three nuclear (H3, 18S and 28S) markers for 19 of the 20 extant families, using traditional Sanger and next-generation 454 sequencing methods. Molecular data were combined with 156 morphological characters in order to estimate the largest anomuran phylogeny to date. The anomuran fossil record allowed us to incorporate 31 fossils for divergence time analyses. Results Our best phylogenetic hypothesis (morphological + molecular data) supports most anomuran superfamilies and families as monophyletic. However, three families and eleven genera are recovered as para- and polyphyletic. Divergence time analysis dates the origin of Anomura to the Late Permian ~259 (224–296) MYA with many of the present day families radiating during the Jurassic and Early Cretaceous. Ancestral state reconstruction suggests that carcinization occurred independently 3 times within the group. The invasion of freshwater and terrestrial environments both occurred between the Late Cretaceous and Tertiary. Diversification analyses found the speciation rate to be low across Anomura, and we identify 2 major changes in the tempo of diversification; the most significant at the base of a clade that includes the squat-lobster family Chirostylidae. Conclusions Our findings are compared against current classifications and previous hypotheses of anomuran relationships. Many families and genera appear to be poly- or paraphyletic suggesting a need for further taxonomic revisions at these levels. A divergence time analysis provides key insights into the origins of major lineages and events and the timing of morphological (body form) and ecological (habitat) transitions. Living anomuran biodiversity is the product of 2 major changes in the tempo of diversification; our initial insights suggest that the acquisition of a crab-like form did not act as a key innovation.

Planktonic foraminiferal d18O and d13C values, derived sea-surface temperatures from d18O and selected planktonic foraminiferal species for core P69 of Hawke Bay off eastern North Island

Recent evidence suggests that the Subtropical Convergence (STC) zone east of New Zealand shifted little from its modern position along Chatham Rise during the last glaciation, and that offshore surface waters north of the STC zone cooled only slightly. However, at nearshore core site P69 (2195 m depth), 115 km off the east coast of North Island and ca 300 km north of the modern STC zone, planktonic foraminiferal species, transfer function data and stable oxygen and carbon isotope records suggest that surface waters were colder by up to 6°C during the late last glacial period compared to the Holocene, and included a strong upwelling signature. Presently site P69 is bathed by south-flowing subtropical waters in the East Cape Current. The nearshore western end of Chatham Rise supports a major bathymetric depression, the Mernoo Saddle, through which some exchange between northern subtropical and southern subantarctic water presently occurs. It is proposed that as a result of much intensified current flows south of the Rise during the last glaciation, a consequence of more compressed subantarctic water masses, lowered sea level, and an expanded and stronger Westerly Wind system, there was accelerated leakage northwards of both Australasian Subantarctic Water and upwelled Antarctic Intermediate Water over Mernoo Saddle in a modified and intensified Southland Current. The expanded cold water masses displaced the south-flowing warm East Cape Current off southeastern North Island, and offshore divergence was accompanied by wind-assisted upwelling of nutrient-rich waters in the vicinity of P69. A comparable kind of inshore cold water jetting possibly characterised most glacial periods since the latest Miocene, and may account for the occasional occurrence of subantarctic marine fossils in onland late Cenozoic deposits north of the STC zone, rather than invoking wholesale major oscillations of the oceanic STC itself.

Sedimentary record of the western Amundsen Sea Embayment after the last glacial maximum

Reliable dating of glaciomarine sediments deposited on the Antarctic shelf since the Last Glacial Maximum (LGM) is very challenging because of the general absence of calcareous (micro-) fossils and the recycling of fossil organic matter. As a consequence, radiocarbon (14C) ages of the acid-insoluble organic fraction (AIO) of the sediments bear uncertainties that are very difficult to quantify. In this paper we present the results of three different chronostratigraphic methods to date a sedimentary unit consisting of diatomaceous ooze and diatomaceous mud that was deposited following the last deglaciation at five core sites on the inner shelf in the western Amundsen Sea (West Antarctica). In three cores conventional 14C dating of the AIO in bulk sediment samples yielded age reversals down-core, but at all sites the AIO 14C ages obtained from diatomaceous ooze within the diatom-rich unit yielded similar uncorrected 14C ages ranging from 13,517±56 to 11,543±47 years before present (yr BP). Correction of these ages by subtracting the core-top ages, which are assumed to reflect present-day deposition (as indicated by 21044 Pb dating of the sediment surface at one core site), yielded ages between ca. 10,500 and 8,400 calibrated years before present (cal yr BP). Correction of the AIO ages of the diatomaceous ooze by only subtracting the marine reservoir effect (MRE) of 1,300 years indicated deposition of the diatom-rich sediments between 14,100 and 11,900 cal yr BP. Most of these ages are consistent with age constraints between 13.0 and 8.0 ka BP for the diatom-rich unit, which we obtained by correlating the relative palaeomagnetic intensity (RPI) records of three of the sediment cores with global and regional reference curves for palaeomagnetic intensity. As a third dating technique we applied conventional 53 radiocarbon dating of the AIO included in acid-cleaned diatom hard parts that were extracted from the diatomaceous ooze. This method yielded uncorrected 14C ages of only 5,111±38 and 5,106±38 yr BP, respectively. We reject these young ages, because they are likely to be overprinted by the adsorption of modern atmospheric carbon dioxide onto the surfaces of the extracted diatom hard parts prior to sample graphitisation and combustion for 14C dating. The deposition of the diatom-rich unit in the western Amundsen Sea suggests deglaciation of the inner shelf before ca. 13 ka BP. The deposition of diatomaceous oozes on other parts of the Antarctic shelf around the same time, however, seems to be coincidental rather than directly related.

Seawater carbonate chemistry, calcification and dissolution response, and skeletal mineralogy of benthic orhanisms during experiments, 2011

Increasing atmospheric pCO2 reduces the saturation state of seawater with respect to the aragonite, high-Mg calcite (Mg/Ca > 0.04), and low-Mg calcite (Mg/Ca < 0.04) minerals from which marine calcifiers build their shells and skeletons. Notably, these polymorphs of CaCO3 have different solubilities in seawater: aragonite is more soluble than pure calcite, and the solubility of calcite increases with its Mg-content. Although much recent progress has been made investigating the effects of CO2-induced ocean acidification on rates of biological calcification, considerable uncertainties remain regarding impacts on shell/skeletal polymorph mineralogy. To investigate this subject, eighteen species of marine calcifiers were reared for 60-days in seawater bubbled with air-CO2 mixtures of 409 ± 6, 606 ± 7, 903 ± 12, and 2856 ± 54 ppm pCO2, yielding aragonite saturation states of 2.5 ± 0.4, 2.0 ± 0.4, 1.5 ± 0.3, and 0.7 ± 0.2. Calcite/aragonite ratios within bimineralic calcifiers increased with increasing pCO2, but were invariant within monomineralic calcifiers. Calcite Mg/Ca ratios (Mg/CaC) also varied with atmospheric pCO2 for two of the five high-Mg-calcite-producing organisms, but not for the low-Mg-calcite-producing organisms. These results suggest that shell/skeletal mineralogy within some-but not all-marine calcifiers will change as atmospheric pCO2 continues rising as a result of fossil fuel combustion and deforestation. Paleoceanographic reconstructions of seawater Mg/Ca, temperature, and salinity from the Mg/CaC of well-preserved calcitic marine fossils may also be improved by accounting for the effects of paleo-atmospheric pCO2 on skeletal Mg-fractionation.

Stable isotope ratios of a oyster Hyotissa hyotis from the northern Red Sea

This study explores the giant oyster Hyotissa hyotis as a novel environmental archive in tropical reef environments of the Indo-Pacific. The species is a typical accessory component in coral reefs, can reach sizes of tens of centimetres, and dates back to the Late Pleistocene. Here, a 70.2-mm-long oxygen and carbon isotope transect through the shell of a specimen collected at Safaga Bay, northern Red Sea, in May 1996, is presented. The transect runs perpendicularly to the foliate and vesicular layers of the inner ostracum near the ligament area of the oyster. The measured d18O and d13C records show sinusoidal fluctuations, which are independent of shell microstructure. The d13C fluctuations exhibit the same wavelength as the d18O fluctuations but are phase shifted. The d18O record reflects the sea surface temperature variations from 1957 until 1996, possibly additionally influenced by the local evaporation. Due to locally enhanced evaporation in the semi-enclosed Safaga Bay, the d18Oseawater value is estimated at 2.17 per mil, i.e., 0.3-0.8 per mil higher than published open surface water d18O values (1.36-1.85 per mil) from the region. The mean water temperature deviates by only 0.4°C from the expected value, and the minimum and maximum values are 0.5°C lower and 2.9°C higher, respectively. When comparing the mean monthly values, however, the sea surface temperature discrepancy between reconstructed and global grid datasets is always <1.0°C. The d13C signal is weakly negatively correlated with regional chlorophyll a concentration and with the sunshine duration, which may reflect changes in the bivalve's respiration. The study emphasises the palaeogeographic context in isotope studies based on fossils, because coastal embayments might not reflect open-water oceanographic conditions.

Internal carotid arterial canal size and scaling in Euarchonta: Re-assessing implications for arterial patency and phylogenetic relationships in early fossil primates.

Primate species typically differ from other mammals in having bony canals that enclose the branches of the internal carotid artery (ICA) as they pass through the middle ear. The presence and relative size of these canals varies among major primate clades. As a result, differences in the anatomy of the canals for the promontorial and stapedial branches of the ICA have been cited as evidence of either haplorhine or strepsirrhine affinities among otherwise enigmatic early fossil euprimates. Here we use micro X-ray computed tomography to compile the largest quantitative dataset on ICA canal sizes. The data suggest greater variation of the ICA canals within some groups than has been previously appreciated. For example, Lepilemur and Avahi differ from most other lemuriforms in having a larger promontorial canal than stapedial canal. Furthermore, various lemurids are intraspecifically variable in relative canal size, with the promontorial canal being larger than the stapedial canal in some individuals but not others. In species where the promontorial artery supplies the brain with blood, the size of the promontorial canal is significantly correlated with endocranial volume (ECV). Among species with alternate routes of encephalic blood supply, the promontorial canal is highly reduced relative to ECV, and correlated with both ECV and cranium size. Ancestral state reconstructions incorporating data from fossils suggest that the last common ancestor of living primates had promontorial and stapedial canals that were similar to each other in size and large relative to ECV. We conclude that the plesiomorphic condition for crown primates is to have a patent promontorial artery supplying the brain and a patent stapedial artery for various non-encephalic structures. This inferred ancestral condition is exhibited by treeshrews and most early fossil euprimates, while extant primates exhibit reduction in one canal or another. The only early fossils deviating from this plesiomorphic condition are Adapis parisiensis with a reduced promontorial canal, and Rooneyia and Mahgarita with reduced stapedial canals.

Late Quaternary deep sea sedimentation off the Niger delta

The oxygen isotopes ratios of benthic foraminifera and detailed radiocarbon ages of the organic matter of an over 15 m long sediment core from the outer Niger delta allow us to date the oxygen isotope stage boundaries 1/2 to 11500 (+/- 650) years BP, 2/3 to approximately 23000 (+/- 2000) years BP. The composition of the predominantly terrigenous clays and accessory pelagic fossils reflects the evolution of the climate over the southwestern Sahel zone and the response of the Eastern Tropical Atlantic to these climatic fluctuations during the Late Quaternary. The dilution of the pelagic fossil concentrations by the terrigenous material and the oxygen isotopes ratios of planktonic foraminifera indicate large fluctuations in the freshwater discharge from the Niger, with high precipitations over the drainage area of this river from 4500 (+/- 300) to 11500 (+/- 650) years BP and from 11800 (+(- 600) to 13000 (+/- 600) years BP while the time intervals in between were as dry as today. Relative increase of kaolinite during wet phases and the association of smectite, chlorite and attapulgite during dry ones characterize the response of the weathering in the Niger drainage basins to the climatic fluctuations. The occurrence of 10-14 A mixed-layers prior to 26000 years BP is correlated with moderate alteration of the crystalline substratum outcrops from the middle-lower part of the Niger Basin. High quartz concentrations are particularly typical for the transition between oxygen isotope stages 1 and 2 at the inception of heavy precipitations in the southern Sahel zone. Sedimentation rates were quite constant, 30-35 cm/1000 years; they became unusually large at the beginning of the Holocene from 10900 (+/- 650) to 11500 (+/- 650) years BP where they reached more than 600 cm/1000 years. Bottom waters around 1100 m depth in the Gulf of Guinea responded to changes in paleo-oceanography of the entire Atlantic Ocean as well as to local influences. Abnormal carbon isotopes ratios and the drastic changes from a highly diversified fauna (during stages 2 and 3. and during the last part of stage 1 after approx. 7000 years BP) to a poorly diversified fauna in the intervenin time span point to the development of a local benthic environment which cannot easily be compared with the corresponding continental and slope environments of the entire Atlantic Ocean.

A new genus Abyssogena from deep-water vents and seeps

A new genus Abyssogena is established for A. phaseoliformis (Métivier, Okutani & Ohta, 1986) and A. kaikoi (Okutani & Métivier, 1986), which were previously assigned to the genus Calyptogena Dall, 1891, and also for two new species, A. southwardae and A. novacula. The most characteristic features of Abyssogena are an elongate shell up to about 280 mm in length; a pallial line starting from the ventral margin of the anterior adductor scar; secondary pallial attachment scars developed dorsal to the pallial line; radially arranged hinge teeth with a reduced anterior cardinal tooth in the right valve; and presence of an inner ctenidial demibranch only. Abyssogena occurs in deep water from 2,985 to 6,400 m and is distributed in the Pacific and Atlantic Oceans at cold seeps along continental margins and hydrothermal vents at mid-oceanic ridges. Some species have a remarkably wide geographic distribution; A. southwardae is present throughout the Atlantic and A. phaseoliformis is present in Japan, Kuril-Kamchatka, as well as Aleutian Trenches. No fossils of Abyssogena are known.

Magnetic susceptibility and particle size for the Shilou red clay section on the eastern Chinese Loess Plateau

The Chinese Loess Plateau red clay sequences display a continuous alternation of sedimentary cycles that represent recurrent climatic fluctuations from 2.58 Ma to the Miocene. Deciphering such a record can provide us with vital information on global and Asian climatic variations. Lack of fossils and failure of absolute dating methods made magnetostratigraphy a leading method to build age models for the red clay sequences. Here we test the magnetostratigraphic age model against cyclostratigraphy. For this purpose we investigate the climate cyclicity recorded in magnetic susceptibility and sedimentary grain size in a red clay section previously dated 11Myr old with magnetostratigraphy alone. Magnetostratigraphy dating based on only visual correlation could potentially lead to erroneous age model. In this study the correlation is executed through the iteration procedure until it is supported by cyclostratigraphy; i.e., Milankovitch cycles are resolved in the best possible manner. Our new age model provides an age of 5.2Ma for the Shilou profile. Based on the new age model, wavelet analysis reveals the well-preserved 400 kyr and possible 100 kyr eccentricity cycles on the eastern Chinese Loess Plateau. Further, paleomonsoon evolution during 2.58-5.2Ma is reconstructed and divided into three intervals (2.58-3.6Ma, 3.6-4.5Ma, and 4.5-5.2Ma). The upper part, the youngest stage, is characterized by a relatively intensified summer monsoon, the middle stage reflects an intensification of the winter monsoon and aridification in Asia, and the earliest stage indicates that summer and winter monsoon cycles may have rapidly altered. The use of cyclostratigraphy along withmagnetostratigraphy gives us an effectivemethod of dating red clay sequences, and our results imply that many presently published age models for the red clay deposits should be perhaps re-evaluated.

Descriptions of 104 gravity and piston sediment cores from POLARSTERN expedition ARK-VIII/3 to the Arctic Ocean

Sediment descriptions and lithostratigraphy (chapter 6.4) NANSEN BASIN The upperrnost 20-50 cm of sedirnents in the Nansen Basin norrnally cornprise soft dark brown, brown-grayish and brown clay. Except for the toprnost clay, the four piston cores retrieved, contained quite different lithologies: a rnuddy diarnicton with outsized clasts (PS2157-6), sandy-silt beds alternating with clay beds (PS2159-6), and silty clay beds of brownish and grayish colours (PS2161-3). Core PS2208-3 was retrieved frorn a plateau on a searnount. The plateau was serni-encircled by hills. The upper 250 cm of this core cornprise brown and olive brown clays. Below these are several sandlayers and a 74 cm thick unit of a sandy mud with rnud-clasts up to 20 cm in diameter. GAKKEL RIDGE The uppermost 20-50 cm of sediments on the Gakkel Ridge comprise soft dark brown, brown, grayish brown clay. In most of the cores there are two horizons of brown clay separated by olive brown clay. The upper horizon is darker. The older stratigraphy is rather varied. Core PS2165-1 contains several thin gray sandlsilt layers, probably distal turbidites. The sarne is found in Core PS2167-1. This core also has a thick (approx. 2 rn) coarse grained turbidite containing large rnud clasts and basaltic rock fragrnents. The color of the turbiditic layers is dark gray. There are several horizons of hernipelagic sandylsilty clays with quite a variety in colours; black, gray, olive, brown, yellowish brown and reddish. The colour variation rnay be due to hydrotherrnal activity or provenance or a shift in redox potential. Cores PS2168-2 and PS2169-1 have typical sequences of very dark gray sandy mud with sharp lower boundaries grading upwards into olive brown clay. Below the lower boundary is often a thin (1-2 cm) gray clay layer. AMUNDSEN BASIN The giant box cores (GKG) provided in most cases excellently preserved sedirnent surfaces which consisted in the entire Amundsen Basin of dark brown to dark grayish brown silty clay with few dropstones and common calcareous microfossils (foraminifers and calcareous nannofossils). The brown and grayish brown color of the sediment surface is a result of the oxidizing conditions at the seafloor due to the rapid renewal of the bottom water rnasses. Planktic forarninifers and calcareous nannofossils are relatively frequent and well preserved despite the rernote location of the basin and its water depths of >4000 rn. Srnear slide descriptions have shown that the surface sedirnents consist dorninantly of clays to silty rnuds with clay rninerals and quartz as the rnost important constituents. The coarse fractions contained besides planktic and benthic forarninifers and coarse clastic rnaterials, rare bivalves, dropstones and mud clasts. The Station PS2190 at the North Pole is a particular good exarnple of the type of sedirnents deposited at the sea floor surface of the Arnundsen Basin, with hornogenous dark brown soft clay covering a sedirnent sequence of highly variable cornposition. Nurnerous giant box cores also provide insight into the detailed lithostratigraphy of the upperrnost sedirnent layers. Twelve box cores have been collected frorn the Arnundsen Basin. Below the youngest unit of 5-20 crn thick silty clays deposits of variable stratigraphies have been found, rnostly consisting of clays or silty clays. In a few instances turbidites have been observed. Benthic forarninifers have not been found in the surface sedirnents. Other fossils were extrernely rare. Bioturbation is weakly developed on all stations. Benthic anirnals seern to live only in and on the upperrnost 2 cm of the uppermost sediment layer. They cornprise amphipods (on all stations) and holothurians, bryozoans, polychaetes, and porifers at one station each. LOMONOSOV RIDGE Sediments from the Lomonosov Ridge show a variety of colors and textures. Following smear slide analyses they are composed mostly of clay minerals and quartz with mica and feldspars, especially in the siltier and sandier parts. Volcanic glass, microcrystalline carbonate, opaque minerals and green amphibole are occasional accessories. The sediments from the Lomonosov Ridge show a noticeable difference from sediments collected from the surrounding basins. Lomonosov Ridge sediments are richer in silt and sand than basin sediments. Occasional turbidites occur in ridge sediments but these must be of entirely local origin. The ridge sediments include frequent layers of "cottage cheese" texture made up of what appear to be small, angular mud clasts of a variety of colors.

(Table 1) Types of giant magnetofossils in ODP holes

Magnetotactic bacteria biomineralize magnetic minerals with precisely controlled size, morphology, and stoichiometry. These cosmopolitan bacteria are widely observed in aquatic environments. If preserved after burial, the inorganic remains of magnetotactic bacteria act as magnetofossils that record ancient geomagnetic field variations. They also have potential to provide paleoenvironmental information. In contrast to conventional magnetofossils, giant magnetofossils (most likely produced by eukaryotic organisms) have only been reported once before from Paleocene-Eocene Thermal Maximum (PETM; 55.8 Ma) sediments on the New Jersey coastal plain. Here, using transmission electron microscopic observations, we present evidence for abundant giant magnetofossils, including previously reported elongated prisms and spindles, and new giant bullet-shaped magnetite crystals, in the Southern Ocean near Antarctica, not only during the PETM, but also shortly before and after the PETM. Moreover, we have discovered giant bullet-shaped magnetite crystals from the equatorial Indian Ocean during the Mid-Eocene Climatic Optimum (~40 Ma). Our results indicate a more widespread geographic, environmental, and temporal distribution of giant magnetofossils in the geological record with a link to "hyperthermal" events. Enhanced global weathering during hyperthermals, and expanded suboxic diagenetic environments, probably provided more bioavailable iron that enabled biomineralization of giant magnetofossils. Our micromagnetic modelling indicates the presence of magnetic multi-domain (i.e., not ideal for navigation) and single domain (i.e., ideal for navigation) structures in the giant magnetite particles depending on their size, morphology and spatial arrangement. Different giant magnetite crystal morphologies appear to have had different biological functions, including magnetotaxis and other non-navigational purposes. Our observations suggest that hyperthermals provided ideal conditions for giant magnetofossils, and that these organisms were globally distributed. Much more work is needed to understand the interplay between magnetofossil morphology, climate, nutrient availability, and environmental variability.