Molecular research on the systematically challenging smoothhound shark genus Mustelus: a synthesis of the past 30 years

The species-rich genus Mustelus (smoothhounds) of the shark family Triakidae is one of the most bio-economically important groups of elasmobranchs in the world’s oceans. Despite the commercial value of Mustelus, the systematics of the group remains largely unresolved and there is no global review or synthesis of knowledge about the conservation status and conservation genetics of smoothhounds across all oceanic regions. Here, we analysed published studies as well as grey literature to gain insight into the biogeographic, ecological and behavioural factors that shape genetic diversity in smoothhounds, and we identify critical knowledge gaps. From a series of molecular phylogenetic studies it can be inferred that the genus Mustelus is paraphyletic and that the aplacental species evolved secondarily from the placental species of the genus. The increasing availability of genetic data aids in disentangling systematic issues, such that more meaningful morphological characters can be chosen for use in practical field-identification keys for co-occurring smoothhounds. An integrative taxonomic approach to the genus Mustelus may offer the best chance of recording and protecting the biodiversity of these sharks. Furthermore, it is evident that different smoothhound species exhibit unique gene-flow patterns, suggesting varying rates within species and hence that species-level conservation approaches would be most appropriate. Molecular studies have advanced our understanding of smoothhound biology (including reproductive traits), ecology and evolution. While many knowledge gaps remain, a crucial lesson from this review is that, when doing assessments on a molecular level, it is important to place genetic results in a broader context, by assimilating biological and ecological data, if definitive conclusions are to be drawn.     

Remarkably uniform oxygen isotope systematics for co-existing pairs of gem-spinel and calcite in marble,with special reference to Vietnamese deposits

Oxygen isotope systematics for co-existing pairs of gem-spinel and calcite in marble from Vietnam and other worldwide deposits have been determined in order to characterize the O-isotope fractionation between calcite and spinel. In Vietnam, the Δ¹⁸O_cc–sp (= 3.7 ± 0.1‰ for six samples from the An Phu and Cong Troi deposits) is remarkably constant. The combination of these data with those obtained on calcite–spinel pairs of Paigutan (Nepal, n = 2), Ipanko (Tanzania, n = 1), and Mogok (Myanmar, = 2) are also consistent with an overall Δ¹⁸O_cc–sp of 3.6 ± 0.3‰ for all the spinel samples (n = 11). The straight line correlation δ¹⁸O_cc = 0.96 δ¹⁸O_sp + 4.4 is excellent despite their worldwide geographic spread. The increment method of calculating oxygen isotope fractionation gave a geologically unreasonable temperature of formation for both minerals at 1374 °C when compared to temperatures obtained by mineral assemblage equilibrium of these marble type deposits, between 610 and 750 °C. The constant Δ¹⁸O_cc–sp reflects a constant temperature for this amphibolite facies assemblage, whose current best estimate is calculated at 620 ± 40 °C, but unquantified uncertainties remain.     

Palaeoproterozoic magnesite: lithological and isotopic evidence for playa/sabkha environments

Magnesite forms a series of 1‐ to 15‐m‐thick beds within the ≈2·0 Ga (Palaeoproterozoic) Tulomozerskaya Formation, NW Fennoscandian Shield, Russia. Drillcore material together with natural exposures reveal that the 680‐m‐thick formation is composed of a stromatolite–dolomite–‘red bed’ sequence formed in a complex combination of shallow‐marine and non‐marine, evaporitic environments. Dolomite‐collapse breccia, stromatolitic and micritic dolostones and sparry allochemical dolostones are the principal rocks hosting the magnesite beds. All dolomite lithologies are marked by δ¹³C values from +7·1‰ to +11·6‰ (V‐PDB) and δ¹⁸O ranging from 17·4‰ to 26·3‰ (V‐SMOW). Magnesite occurs in different forms: finely laminated micritic; stromatolitic magnesite; and structureless micritic, crystalline and coarsely crystalline magnesite. All varieties exhibit anomalously high δ¹³C values ranging from +9·0‰ to +11·6‰ and δ¹⁸O values of 20·0–25·7‰. Laminated and structureless micritic magnesite forms as a secondary phase replacing dolomite during early diagenesis, and replaced dolomite before the major phase of burial. Crystalline and coarsely crystalline magnesite replacing micritic magnesite formed late in the diagenetic/metamorphic history. Magnesite apparently precipitated from sea water‐derived brine, diluted by meteoric fluids. Magnesitization was accomplished under evaporitic conditions (sabkha to playa lake environment) proposed to be similar to the Coorong or Lake Walyungup coastal playa magnesite. Magnesite and host dolostones formed in evaporative and partly restricted environments; consequently, extremely high δ¹³C values reflect a combined contribution from both global and local carbon reservoirs. A ¹³C‐rich global carbon reservoir (δ¹³C at around +5‰) is related to the perturbation of the carbon cycle at 2·0 Ga, whereas the local enhancement in ¹³C (up to +12‰) is associated with evaporative and restricted environments with high bioproductivity.     

Pore water evolution in oilfield sandstones: constraints from oxygen isotope microanalyses of quartz cement

Oxygen isotope microanalyses of authigenic quartz, in combination with temperatures of quartz precipitation constrained by fluid inclusion microthermometry and burial history modelling, are employed to trace the origin and evolution of pore waters in three distinct reservoirs of the Brae Formation in the Miller and Kingfisher Fields (North Sea). Oxygen isotope ratios of quartz cements were measured in situ in nine sandstone thin sections with a Cameca ims-4f ion microprobe. In conjunction with quartz cement paragenesis in the reservoirs, constrained from textural and cathodoluminescence (CL) microscopy studies, pore water evolution was reconstructed from the time of deposition of the sandstones in the Upper Jurassic until the present.CL photomicrographs of quartz overgrowths in the Brae Formation sandstones show three cement zones (A, B and C) which can be related to different oxygen isotope compositions: (1) the earliest, and thinnest, zone A (homogeneous CL pattern with probable δ¹⁸O values between +23‰ and +26‰—direct measurements were not possible) precipitated in the sandstones at temperatures <60 °C; (2) the second zone B (complex CL pattern and directly measured δ¹⁸O values between +15‰ and +18‰) precipitated in the sandstones most likely between 70 and 90 °C; (3) the third zone C (homogeneous CL pattern and directly measured δ¹⁸O values between +16‰ and +22‰) precipitated in the sandstones most likely at temperatures >90 °C. Calculated oxygen isotope compositions of pore waters show that zone A quartz cements, and enclosing concretionary calcite, precipitated from a meteoric-type fluid (∼−7‰) during shallow burial (<1.5 km). Zone B quartz cements precipitated from fluids which evolved in composition from a meteoric-type fluid (δ¹⁸O −7‰) to a more ¹⁸O-enriched fluid (δ¹⁸O −4‰) as burial continued to ∼3.0 km. Data from zone C quartz cements are consistent with further fluid evolution from δ¹⁸O −4‰ to basinal-type fluids with δ¹⁸O similar to the present-day formation water oxygen isotope composition (+0.6‰ at 4.0 km burial). A similar pore water evolution can be derived for all three reservoirs studied, indicating that hydrogeologic evolution was similar across sandstones of the whole Brae Formation.The quartz cement zones observed in the Brae Formation sandstones, and the pore water history derived for the area studied, is analogous to published petrographic and pore water evolution data from the nearby Brent Group reservoirs and from reservoirs located in the Haltenbanken area on the Atlantic margin offshore Norway. Considering quartz cement is a major porosity-occluding phase in many reservoir sandstones, and because pore waters both dissolve quartz and carry the dissolved silica to cementation sites, the data presented are valuable for improving the understanding and prediction of reservoir quality development in sandstones globally.     

The origin of clay minerals in active and relict hydrothermal deposits

Samples of Fe-oxide-rich hydrothermal sediments were collected from active and inactive portions of the TransAtlantic Geotraverse (TAG) hydrothermal field on the Mid-Atlantic Ridge. Clays separated from TAG metalliferous sediments in this study all consist of Al-poor nontronite. Oxygen isotope thermometry of the clays yields formation temperatures of 54-67°C for samples from the inactive Alvin mound compared with 81-96°C for samples from the active TAG site. The latter are the highest recorded temperatures for authigenic hydrothermal clays. Sr isotope analysis of the clays from the active mound suggests that they precipitated from seawater-dominated fluids, containing less than 15% hydrothermal end-member fluid. In contrast, nontronite from the inactive Alvin mound has ⁸⁷Sr/⁸⁶Sr values that closely resemble that of detrital North Atlantic clays, suggesting a dominantly continental source for the Sr. Rare earth element data are consistent with a significant detrital input to the inactive site but also demonstrate the extent of hydrothermal input to the low temperature fluid. Crystallographic fractionation of the trivalent REE is apparent in the heavy REE enrichments for all nontronite samples. The inferred formation-mechanism for nontronite-rich Fe-oxyhydroxide deposits at the surface of the active mound is by direct precipitation from low temperature fluids. At the inactive Alvin site, in contrast, the deposits form during alteration of pelagic sediments by diffuse fluids and replacement of biogenic carbonate with nontronite and Fe-oxyhydroxide. These two modes of formation are both important in seafloor hydrothermal settings where clay minerals are a significant component of the hydrothermal deposit.     

Petrological, geochemical, and stable isotope constraints on the genesis of the Miocene igneous rocks of Chetaibi and Cap de Fer (NE Algeria)

Miocene igneous rocks (diorites, andesites, dacites, rhyolites and microgranites) of Chetaibi and Cap de Fer massif, NE Algeria, are high-K calc-alkaline to shoshonitic rocks. Fresh diorites have δ³⁴S and δ¹⁸O values ranging between −2.5‰ and +5.9‰, +6.5‰ and +6.7‰ respectively, indicating a mantle origin. The relatively low δ³⁴S values (−5.4‰ to −12.2‰) and high δ¹⁸O (+8.3‰ to +9.0‰) of altered diorites indicate the input of a crustal component to the initial magma. The microgranites’ I-type signature is indicated by the geochemical data and the δ³⁴S and δ¹⁸O values of −1.2‰ and −3.6‰, and +7.8‰ to +10.4‰ respectively. The andesites show a large variation of δ³⁴S, between −33.2‰ and +25.7‰. Massive andesites with δ³⁴S between +6.8‰ and +7.6‰ preserve a ³⁴S-enriched mantle signature. The δ³⁴S of the lava flows between +25.7‰ and +25.8‰ are attributed to open system magma degassing, whereas the low δ³⁴S of two andesitic dyke samples (−13.7‰ and −33.2‰) strongly suggest a crustal sulphur input. High δ¹⁸O (+9.2‰ to +15.7‰) of andesites indicate post-magmatic alteration (mainly silicification); the flyschs with δ¹⁸O between of +13.3‰ and +21.7‰ are most likely the contaminant. Quartz veins within the andesites gave a δ¹⁸O value of +23.0‰ while silica-filling vesicles yielded a value of +13.8‰. Initial Sr-isotope data are rather high for all the rocks (diorites: 0.707–0.708, andesites: 0.707–0.710, and microgranites and rhyolites: 0.717–0.719), and because geochemical and stable isotope data do not indicate a substantial amount of crustal assimilation, an extensive enrichment of the mantle source by subducted sediments is called for. A metasomatized-mantle source, characterized by high radiogenic Sr and relatively high δ¹⁸O, has also been indicated for the genesis of similar Tertiary igneous rocks in the Western Mediterranean basin, e.g. the Volcanic Province of southeasten Spain [Benito, R., Lopez-Ruiz, J., Cebria, J.M., Hertogen, J., Doblas M., Oyarzun, R., Demaiffe, D., 1999. Sr and O isotope constraints on source and crustal contamination in the high-K calc-alkaline and shoshonitic neogene volcanic rocks of SE Spain. Lithos 46, 773–802] and some plutons of northeastern Algeria [Ouabadi, A., 1994. Pétrologie, géochimie et origine des granitoïdes peralumineux à cordiérite (Cap Bougaroun, Béni-Touffout et Filfila), Algérie nord-orientale. Thèse de Doctorat, Université de Rennes I, France, 257p; Fourcade, S., Capdevila, R., Ouabadi, A., Martineau, F., 2001. The origin and geodynamic significance of the Alpine cordierite-bearing granitoids of northern Algeria. A combined petrological, mineralogical, geochemical and isotopic (O, H, Sr, Nd) study. Lithos 57, 187–216].     

Genesis of sulfide-rich chromite ores by the interaction between chromitite and pegmatitic olivine–norite dikes in the Potosí Mine (Moa-Baracoa ophiolitic massif, eastern Cuba)

The Potosí Mine is located in the Moa-Baracoa massif in the easternmost part of the Cuban Ophiolitic Belt. Chromite mineralization occurs within the mantle-crust transition zone. Two events of magma intrusion overprint the chromitite bodies: one gave rise to the crystallization of pegmatitic olivine-norite dikes, and the other produced pegmatitic gabbro dikes. Sulfide-poor chromite ores, brecciated chromite ores, and sulfide-rich chromite ores can be distinguished in the different chromitite bodies. Sulfide-poor ores represent more than 80 vol% of the chromitites. This type occurs far from the zones intruded by pegmatitic gabbro dikes and shows petrographic and chemical features similar to other chromitite bodies described in the Moa-Baracoa massif. Brecciated chromite ores occur within pegmatitic gabbro dikes. In this type, chromite crystals occur included within chromian diopside and plagioclase. These silicates often contain droplet-like sulfide aggregates. Sulfide-rich ores are spatially associated to the contacts between sulfide-poor chromite and pegmatitic olivine-norite dikes. These ores mainly consist of recrystallized (coarse) chromite with interstitial pyrrhotite, pentlandite, cubanite, and chalcopyrite. Chromite from sulfide-rich ores exhibits TiO₂, FeO, V₂O₃, MnO, and especially, Fe₂O₃ contents, higher than those of chromite from brecciated ores and much higher than those of chromite from sulfide-poor ores. The sulfide-rich ores are PGE-rich (up to 1,113 ppb of total PGE), and show nearly flat chondrite-normalized PGE patterns, slightly above 0.1 times chondritic values. Mineralogical and chemical data indicate that the chromite ores of the Potosí Mine were modified by the intrusions of olivine-norite and gabbro dikes. The interaction between pre-existing sulfide-poor chromite ores and the intruding volatile-rich silicate melts produced strong brecciation, partial dissolution, and recrystallization (coarsening) of chromite. The sulfide assemblage formed by fractionation of the immiscible sulfide melt segregated from the volatile-rich silicate melt that generated the pegmatitic olivine-norite. The segregation of the sulfide melt can be interpreted as the consequence of chemical interaction between intruding melts and the host chromite. The variable extent of this interaction produced chromite ores with variable sulfide ratios. The magmatic nature of the sulfide mineralization is supported by sulfur isotope data, which range from -0.4 to +0.9‰. Sulfide melt collected incompatible PGE (Rh, Pt, Pd) to produce the typical flat chondrite-normalized pattern of sulfide-rich chromite ores.     

Fluid inclusions,stable isotopes and gold deposition at Björkdal,northern Sweden

The Björkdal gold deposit is located in the eastern part of the Early Proterozoic Skellefte district in northern Sweden. The ore zone is hosted by a granitoid which intrudes a 1.9 Ga old supracrustal sequence and consists of a network of quartz veins between two shear zones. The ore mineralogy, alteration assemblages, ore fluid characteristics and general setting of the Björkdal deposit reveal many similarities with mesothermal Archean systems. Three types of fluids are represented by fluid inclusions observed in quartz, scheelite and calcite. The first type consists of a CO₂-rich fluid which is syngenetic with the formation of the quartz veins. These inclusions occur in quartz and scheelite. Isotopic equilibrium temperatures derived from quartz-scheelite pairs reflect depositional temperatures around 375 °C. Molar volumes of the carbonic fluid inclusions, ranging down to 55 cm³mole, indicate a maximum lithostatic trapping pressure of 1.8 kbar. These fluids were generated at depth in conjunction with early orogenic magma-forming processes. The gold was introduced to the vein system by the carbonic fluid but the gold was deposited after reactions between this fluid and the wall-rock, producing a slightly alkaline, more CH₄-rich aqueous type 2 fluid. Fluid inclusions of this chemically modified fluid indicate that the precipitation of the gold, together with pyrrhotite, pyrite and chalcopyrite, occurred under heterogenous conditions at a temperature of 220 °C and a hydrostatic pressure of 0.5 kbar. The gold deposition occurred from fluids with a δ ¹⁸O signature of around +8‰ and δD values close to zero per mil. Any metamorphic influence on the stable isotopic signatures is regarded as minimal. The isotope data suggest rather that a surface-derived fluid component had access to the vein system during this process. At a post-vein forming stage (metamorphic stage ?) a secondary episode of gold mobilization occurred as suggested by the aqueous type 3 inclusions trapped in cross-cutting microfractures in quartz and randomly in calcite, and with homogenization temperatures between 145–220 °C and a salinity up to 11eq. wt.% NaCl. The Skellefte district is a major ore province, which forms a 200 by 50 km area in northern Sweden (Fig. 1), comprising numerous stratabound massive sulfide ore deposits. During the last decade epigenetic gold deposits have received increasing interest from a prospecting point of view. The Björkdal deposit is one of several epigenetic gold discoveries made recently in the Skellefte district. In 1985 a geochemical survey, designed on a grid-pattern basis, revealed a gold anomaly about 12 km north-east of the Boliden community and three years later the Björkdal gold mine was in operation. The annual production is about 960 000 metric tons of ore (1992) and the total reserves are estimated at a minimum of 7 Mton of ore with a gold grade of 2.9 ppm. This paper reports on the geological features of the Björkdal deposit and discusses the genesis of the deposit on basis of fluid inclusions and distribution of oxygen and hydrogen isotopes.     

Fluid–rock interaction at a carbonatite-gneiss contact, Alnö, Sweden

We evaluate balanced metasomatic reactions and model coupled reactive and isotopic transport at a carbonatite-gneiss contact at Alnö, Sweden. We interpret structurally channelled fluid flow along the carbonatite-gneiss contact at ～640°C. This caused (1) metasomatism of the gneiss, by the reaction: ${\hbox{biotite} + \hbox{quartz} + \hbox{oligoclase} + \hbox{K}_{2} \hbox{O} +\,\hbox{Na}_{2}\hbox{O} \pm \hbox{CaO} \pm \hbox{MgO} \pm \hbox{FeO} = \hbox{albite} + \hbox{K-feldspar} + \hbox{arfvedsonite} + \hbox{aegirene-}\hbox{augite} + \hbox{H}_{2} \hbox{O} + \hbox{SiO}_{2}}We evaluate balanced metasomatic reactions and model coupled reactive and isotopic transport at a carbonatite-gneiss contact at Aln?, Sweden. We interpret structurally channelled fluid flow along the carbonatite-gneiss contact at ∼640°C. This caused (1) metasomatism of the gneiss, by the reaction: , (2) metasomatism of carbonatite by the reaction: calcite + SiO₂ = wollastonite + CO₂, and (3) isotopic homogenization of the metasomatised region. We suggest that reactive weakening caused the metasomatised region to widen and that the metasomatic reactions are chemically (and possibly mechanically) coupled. Spatial separation of reaction and isotope fronts in the carbonatite conforms to a chromatographic model which assumes local calcite–fluid equilibrium, yields a timescale of 10²–10⁴ years for fluid–rock interaction and confirms that chemical transport towards the carbonatite interior was mainly by diffusion. We conclude that most silicate phases present in the studied carbonatite were acquired by corrosion and assimilation of ijolite, as a reactive by-product of this process and by metasomatism. The carbonatite was thus a relatively pure calcite–H₂O−CO₂–salt melt or fluid.