Ocean basin and depth variability of oxygen isotopes in cenozoic benthic foraminifera

One of the longest, most detailed quantitative records of oceanographic change in the Cenozoic is that provided by oxygen isotope measurements made on the tests of foraminifera. As indicated by measurements on benthic foraminifera, the deep waters of the world ocean have undergone an overall cooling of about 10°C in the Cenozoic. This change has been neither monotonic nor gradual. Rather, it is evidenced by a few, relatively rapid increases in the ¹³O content of the benthic shells. These “steps” in the isotopic record have been associated with major evolutionary changes in the mean state of the deep ocean. The variance around this mean state has also changed through the Cenozoic. From relatively high variance in the Middle Eocene, the oceans showed low variance in the Late Eocene and Oligocene. In the Miocene the variance of the isotopic measurement again increased, reaching a maximum of short duration in the Middle Miocene. This maximum as well as that which occurred during the Late Pliocene and Quaternary, may be attributable to fluctuations in the isotopic composition of the oceans caused by growth and decay of large ice sheets.In the Late Miocene the benthic oxygen isotopes in Atlantic sites less than 3000 m deep have a higher variance than sites at similar depths in the Pacific and Indian Oceans. It is thought that this high variance results from long-term changes in the importance of the cool and salty North Atlantic Deep Water relative to that of the warmer and less saline Antarctic Intermediate Water at Atlantic sites between 1000 m and 3000 m water depth. Such significant differences in benthic isotopic variance between the ocean basins have been demonstrated only in post-Middle Miocene intervals.     

Ontogenetic effects on stable carbon and oxygen isotopes in tests of live (Rose Bengal stained) benthic foraminifera from the Pakistan continental margin

We determined the stable oxygen and carbon isotopic composition of live (Rose Bengal stained) tests belonging to different size classes of two benthic foraminiferal species from the Pakistan continental margin. Samples were taken at two sites, with water depths of about 135 and 275 m, corresponding to the upper boundary and upper part of the core region of the oxygen minimum zone (OMZ). For Uvigerina ex gr. Uvigerina semiornata and Bolivina aff. Bolivina dilatata, δ¹³C and δ¹⁸O values increased significantly with increasing test size. In the case of Uvigerina ex gr. U. semiornata, δ¹³C increased linearly by about 0.105‰ for each 100-μm increment in test size, whereas δ¹⁸O increased by 0.02 to 0.06‰ per 100 μm increment. For Bolivina aff. B. dilatata the relationship between test size and stable isotopic composition is better described by logarithmic equations. A strong positive linear correlation is observed between δ¹⁸O and δ¹³C values of both taxa, with a constant ratio of δ¹⁸O and δ¹³C values close to 2:1. This suggests that the strong ontogenetic effect is mainly caused by kinetic isotope fractionation during CO₂ uptake. Our data underline the necessity to base longer δ¹⁸O and δ¹³C isotope records derived from benthic foraminifera on size windows of 100 μm or less. This is already common practice in down-core isotopic studies of planktonic foraminifera.     

Late Quaternary deep and surface water mass evolution in the northeastern Indian Ocean inferred from carbon and oxygen isotopes of benthic and planktonic foraminifera

New planktonic and benthic foraminiferal stable isotope records from core YDY05 (northeastern Indian Ocean) provide new insights into paleoceanographic changes in the northeastern Indian Ocean since the last glacial period. The distinct δ¹⁸O decrease was observed since the beginning of the deglaciation to the mid-Holocene (∼8–6 kyr BP), possibly reflecting a reduction in surface salinity in the central Bay of Bengal (BoB) water, which probably resulted from strengthened precipitation, concurrent enhanced river discharge and rising sea-level, related to the intensification of Indian Summer Monsoon (ISM). Variations in benthic δ¹³C and δ¹³C_{Planktonic-Benthic} in our core site reflect significant variations in source water characteristics over the LGM-Holocene. The large δ¹³C_{Planktonic-Benthic} offset during the glacial period suggests a more sluggish deep water circulation, and lower δ¹³C_{Planktonic-Benthic} from the deglaciation to the Holocene suggests an enhanced deep water circulation in the central BoB. The drastic depletion in benthic δ¹³C during the glacial period suggests a significant reduction of North Atlantic Deep Water (NADW) intrusion and a progressive influx of Antarctic Bottom Water (AABW) and ¹²C-rich Circumpolar Deep Water (CDW) into the central BoB. In contrast, since the deglaciation, the central BoB experienced a drastically increased intrusion of better ventilated and ¹³C-rich NADW. The differences in benthic δ¹⁸O between the LGM section and the Holocene exceeds the ice volume effect by ∼0.5‰, providing further evidence that the deep water mass of the central BoB was influenced by the less dense NADW, instead of the AABW, since the last deglaciation.     

Carbon and oxygen isotope geochemistry of live (stained) benthic foraminifera from the Aleutian Margin and the Southern Australian Margin

Comparisons of ambient bottom-water geochemistry and stable isotopic values of the tests of living (stained) calcareous benthic foraminifera from the North Pacific (on the Aleutian Margin, water depth 1988 m) and Murray Canyons group in the Southern Indian Ocean (Australian Margin, water depths 2476 m and 1634 m) provide modern environmental analogs to calibrate paleoenvironmental assessments. Consistent with the hypothesis that microhabitat preferences influence foraminiferal isotopic values, benthic foraminifera from both margins were depleted in ¹³C with respect to bottom-water dissolved inorganic carbon (DIC). The carbon isotope values of deep infaunal foraminifera (Chilostomella oolina, Globobulimina pacifica) showed greater differences from estimates of those of DIC than shallow benthic foraminifera (Bulimina mexicana, Bolivinita quadrilatera, Pullenia bulloides). This study provides new isotopic and ecological information for B. quadrilatera. The mean Δδ¹³C value, defined as foraminiferal δ¹³C values minus estimated ambient δ¹³C values from the Aleutian Margin, is 0.97‰ higher for G. pacifica than the mean from the Murray Canyon. This difference may result either from genetic or biological differences between the populations or from differences in environmental isotopic influences (such as pore water differences) that were not accounted for in the equilibrium calculations. These analyses provide calibration information for the evaluation of bottom water conditions and circulation patterns of ancient oceans based on fossil foraminiferal geochemistry.     

Factors controlling seasonal variability of benthic ammonium release and oxygen uptake in Alfacs Bay (Ebro Delta,NW Mediterranean)

The seasonal variability of sediment–water ammonium flux andoxygen uptake was studied in an estuarine bay (Alfacs Bay, Ebro Delta, NWMediterranean) influenced by temporal freshwater discharges. Three stationswith different organic loading were sampled. The relationships of benthicfluxes to bottom water (temperature, dissolved oxygen, ammonium, nitrateplus nitrite) and to sediment (porosity, chlorophyll a derivative pigments,organic carbon and nitrogen) variables were examined. Oxygen uptake rangedfrom 0.3 to 2.5 mmol m^–2 h^–1 and ammoniumrelease ranged from 6 to 230 µmol m^–2 h^–1.The lowest value was recorded at the station furthest from the freshwaterinputs, and the highest was at the littoral station nearest the freshwaterdischarge channels (for oxygen uptake) and at the deep station at the saltwedge front (for ammonium flux). Water temperature and the concentration ofchlorophyll a derivative pigments on the surface sediment were revealed asthe main variables to be taken into account to explain the variabilityfound. Changes in fluxes reflecting temperature changes were found at thestation furthest from the freshwater inputs, while at the other, fluxvariability was found to be related to the cycle of functioning offreshwater discharge channels. The different patterns of variability arediscussed in relation to the dynamics of the estuary and to the mainfeatures of benthic nitrogen cycling.     

Stable carbon isotope variability associated with taxonomic composition of lotic benthic algae

We examined the relationship between the δ¹³C and taxonomic composition of benthic algae collected from a riffle (fast current habitat) of a non‐shaded mountain stream, which is a tributary of the Kiso River, Japan. The benthic algal δ¹³C ranged from ?20.6 to ?14.2‰ and tended to be ¹³C‐depleted with increasing relative abundance of upright filamentous cyanobacteria and ¹³C‐enriched with increasing relative abundance of prostrate filamentous cyanobacteria. Using isotopic mass balance equations, the relative abundance of the dominant taxa, upright and prostrate filamentous cyanobacteria, small diatoms and others, explained 74% of δ¹³C variability. This study shows a case where the difference in taxonomic composition is a possible source of the isotopic variability of benthic algae, which is a mixture of taxa with distinct isotopic signatures.     

Stable isotope composition of Late Cretaceous benthic foraminifera from the southern South Atlantic: Biological and environmental effects

The stable carbon and oxygen isotope composition of different benthic foraminiferal species of the latest Campanian and earliest Maastrichtian from Ocean Drilling Project Hole 690C (Weddell Sea, southern South Atlantic, ∼1800 m paleowater depth) have been investigated. The total range of measured isotope values of all samples exceeds ∼4‰ for δ¹³C and 1.1‰ for δ¹⁸O. Carbon isotope values of proposed deep infaunal species are generally similar or only slightly lower when compared to proposed epifaunal to shallow infaunal species. Interspecific differences vary between samples probably reflecting temporal changes in organic carbon fluxes to the sea floor. Constantly lower δ¹³C values for Pullenia marssoni and Pullenia reussi suggest the deepest habitat for these species. The strong depletion of δ¹³C values by up to 3‰ within lenticulinids may be attributed to a deep infaunal microhabitat, strong vital effects, or different feeding strategy when compared to other species or modern lenticulinids. The mean δ¹⁸O values reveal a strong separation of epifaunal to shallow infaunal and deep infaunal species. Epifaunal to shallow infaunal species are characterized by low δ¹⁸O values, deep infaunal species by higher values. This result possibly reflects lower metabolic rates and longer life cycles of deep infaunal species or the operating of a pore water [CO₃²⁻] effect on the benthic foraminiferal stable isotopes.Pyramidina szajnochae shows an enrichment of oxygen isotopes with test size comprising a total of 0.6‰ between 250 and 1250 μm shell size. Although δ¹³C lacks a corresponding trend these data likely represent the presence of changes in metabolic rates during ontogenesis. These results demonstrate the general applicability of multi-species stable isotope measurements of pristine Cretaceous benthic foraminifera to reconstruct past microhabitats and to evaluate biological and environmental effects on the stable isotope composition.     

Carbon and oxygen isotopic disequilibria of recent deep-sea benthic foraminifera

The carbon and oxygen isotopic compositions of 149 samples of benthic foraminifera from deep-sea core tops indicate that none of the nine species studied secrete calcium carbonate in isotopic equilibrium with ambient bottom water. Uvigerina, Pyrgo murrhina, and Oridorsalis tener are the closest to ¹⁸O equilibrium (with average deviations about −0.4‰), while Planulina wuellerstorfi and P. murrhina are the closest to ¹³C equilibrium (with average deviations about −1‰). P. wuellerstorfi shows the most systematic relationship between δ ¹³C and bottom water apparent oxygen utilization. The intraspecific variabilities in δ ¹⁸O and δ ¹³C suggest that estimates of bottom water paleotemperatures can be made to a precision of ± 0.7°C, while estimates of past apparent oxygen utilization (AOU) can be made to ± 35 μmol/kg. Based on intraspecific comparisons of the Recent samples with fossils, no temporal changes in the degree of either ¹⁸O or ¹³C disequilibrium have been detected for Planulina wuellerstorfi, Uvigerina, Oridorsalis tener and Globocassidulina subglobosa.     

Stable carbon and oxygen isotopes in human tooth enamel: Identifying breastfeeding and weaning in prehistory

This paper investigates the utility of stable carbon and oxygen isotopes in human dental enamel to reveal patterns of breastfeeding and weaning in prehistory. Enamel preserves a record of childhood diet that can be studied in adult skeletons. Comparing different teeth, we used δ¹³C to document the introduction of solid foods to infant diets and δ¹⁸O to monitor the decline of breastfeeding. We report enamel carbonate δ¹³C and δ¹⁸O of 33 first molars, 35 premolars, and 25 third molars from 35 burials from Kaminaljuyú, an early state in the valley of Guatemala. The skeletons span from Middle Preclassic through Late Postclassic occupations, ca. 700 B.C. to 1500 A.D. Sections of enamel were removed from each tooth spanning from the cusp to the cemento-enamel junction. Stable isotope ratios were measured on CO₂ liberated by reaction of enamel with H₃PO₄ in an automated carbonate system attached to a VG Optima mass spectrometer. Within a skeleton, teeth developing at older ages are more enriched in ¹³C and more depleted in ¹⁸O than teeth developing at younger ages. Premolars average 0.5% higher in δ¹³C than first molars from the same skeleton (P = 0.0001), but third molars are not significantly enriched over premolars. The shift from first molars to premolars may be due to the shift to solid foods from lipid-rich milk. After 2 years, when premolars begin to mineralize, the δ¹³C in childhood diets did not change systematically. First molars and premolars are similar in δ¹⁸O, but third molars average 0.7% lower than first molars (P = 0.0001) and 0.5% lower than premolars (P = 0.0003). First molar and premolar δ¹⁸O is heavier, because breast milk is more enriched in ¹⁸O than is drinking water. Hence, many children continued to nurse during the period of premolar formation. Together, these results indicate that Kaminaljuyú children had begun to eat solid maize foods before the age of 2 years but continued to drink breast milk until much later. Am J Phys Anthropol 106:1–18, 1998. © 1998 Wiley-Liss, Inc.     

Pliocene benthic foraminifera from the Blake Plateau: Faunal assemblages and paleocirculation

Relative abundance of benthic foraminifera have been analyzed from core V26-145 from the Blake Plateau. The investigated sequence represents the time interval between 1.8 and 4.6 Ma. In order to determine how different sieve sizes influence the relative abundance patterns, three sediment size fractions were studied separately. It becomes difficult to maintain consistent taxonomic concepts in the fraction 63–125 μm, partly because this fraction contains high abundances of juvenile forms. However, the 63–125 μm fraction holds high abundances of the important small speciesEpistominella exigua. Due to these reasons only the two larger fractions (125–250 μm and >250 μm) were considered meaningful to analyze for relative abundance patterns. An analysis of the two larger fractions (>125 μm; >250 μm) shows no consistency in relative abundance patterns.The relative abundance patterns for the 34 most common species in the size fraction >125 μm were analyzed by means of correspondence analysis. Three benthic foraminiferal assemblages (I, II, and III) were recognized and these can be associated with water masses. Assemblage I is associated with the Florida Current and consists of shallow water species (Amphistegina gibbosa, Compressigerina sp. A,Discorbinella biconcavus, Islandiella teretis, Reussella atlantica, andSiphonina pulchra). Assemblage II contains key species for North Atlantic Deep Water (NADW) (Cibicidoides kullenbergi, Epistominella exigua, Globocassidulina subglobosa, Lenticulina peregrina, Oridorsalis umbonatus, andPlanulina wuellerstorfi). The third assemblage (III) contains species associated with the Antilles Current (Bolivina rhomboidalis, Cassidulina obtusa, Cassidulina vortex, andNuttallides umbonifera). The correspondence analysis reveals an alternation in dominance between Assemblage I and Assemblage II prior to 3.3 Ma, suggesting lateral oscillations between the Florida Current and NADW. At about 3.3 Ma Assemblage I disappears and Assemblage III increases in importance, suggesting an increasing influence of the Antilles Current in the upper part of the record.     

Patterns and controls of seasonal variability of carbon stable isotopes of particulate organic matter in lakes

Carbon stable isotopes (δ¹³C) of particulate organic matter (POM) have been used as indicators for energy flow, primary productivity and carbon dioxide concentration in individual lakes. Here, we provide a synthesis of literature data from 32 freshwater lakes around the world to assess the variability of δ¹³C_POM along latitudinal, morphometric and biogeochemical gradients. Seasonal mean δ¹³C_POM, a temporally integrated measure of the δ¹³C_POM, displayed weak relationships with all trophic state indices [total phosphorus (TP), total nitrogen (TN), and chlorophyll a (Chl a)], but decreased significantly with the increase in latitude, presumably in response to the corresponding decrease in water temperature and increase in CO₂ concentration. The seasonal minimum δ¹³C_POM also correlated negatively with latitude while seasonal maximum δ¹³C_POM correlated positively with all trophic state indices, pH, and δ¹³C of dissolved inorganic carbon (DIC). Seasonal amplitude of δ¹³C_POM (the difference between seasonal maximum and minimum values) correlated significantly with pH, TP and Chl a concentrations and displayed small variations in oligotrophic, mesotrophic and low latitude eutrophic lakes, which is attributed to low primary productivity and abundant non-living POM in the low trophic state lakes and relatively stable environmental conditions in the subtropics. Seasonal amplitude of δ¹³C_POM was the greatest in high latitude eutrophic lakes. Greater seasonal changes in solar energy and light regime may be responsible for the large seasonal variability in high latitude productive lakes. This synthesis provides new insights on the factors controlling variations in stable carbon isotopes of POM among lakes on the global scale.     

Late Quaternary paleoproductivity changes off the Congo deduced from stable carbon isotopes of planktonic foraminifera

A detailed record (≈2-kyr intervals) of the difference in stable carbon isotopes (Δδ¹³C) between Globigerina bulloides and Globigerinoides ruber (pink) is used to reconstruct changes in upwelling intensity off the Congo River for the last 190,000 yr. Comparisons of the oxygen and carbon isotope data from this core with records from the Niger Fan and from pelagic cores in the eastern equatorial South Atlantic indicate that the Congo Fan isotope records do not contain a strong freshwater signal as is described off other major rivers. The temporal pattern of the Δδ¹³C signal correlates with the marine organic carbon record from the Congo Fan. Thus the planktonic Δδ¹³C record, reflecting past changes in upwelling intensity and nutrient content, corroborates the signal provided by sedimentary organic carbon, which is presumed to indicate changes in the amount of biological productivity and export flux to the seafloor.

The planktonic Δδ¹³C signal is characterized by a dominant 23-kyr periodicity which provides evidence for a strong response of upwelling fluctuations off the Congo to precessional forcing. Minima in the Δδ¹³C record are aligned with periods of minimum boreal summer insolation over Central Africa reflecting an increase of upwelling and biological productivity off the Congo at periods of enhanced zonal intensity of southeast trades and corresponding weak southeast monsoon over the eastern South Atlantic. A strong response to changes in river discharge probably did not occur, indicating that fertilization by river-derived nutrients has played only a minor role with respect to Late Quaternary changes in the total amount of primary productivity off the Congo.     

Magnesium and strontium compositions of Recent benthic foraminifera from the Coral Sea, Australia and Prydz Bay, Antarctica

Analyses of the Mg/Ca and Sr/Ca ratios of the modern benthic foraminifera, Cibicides wuellerstorfi (epifaunal) and Uvigerina species (infaunal) from the Coral Sea, and Cibicides refulgens (epifaunal) and Trifarina angulosa (infaunal) from Prydz Bay, Antarctica revealed relationships with temperature that have possible applications for reconstructions of bottom-water paleotemperatures. A positive relationship exists between the Mg/Ca and Sr/Ca ratios of Cibicides wuellerstorfi and Cibicides refulgens and ambient temperatures, at least within the range of 2 and 6°C. For the correlation between Mg/Ca compositions and temperature the r² values range from 0.78 (C. wuellerstorfi alone) to 0.88 (C. wuellerstorfi and C. refulgens together). At present, the Mg/Ca-temperature relationship must be regarded as tentative because of significant overlap of standard error values. The relationship between the Sr/Ca compositions of C. wuellerstorfi and bottom-water temperature yields an r² value of 0.95. These results indicate that Sr/Ca and possibly Mg/Ca ratios of Cibicides wuellerstorfi may provide useful information for the assessment of paleotemperature. Single-species data are presently insufficient to assess the influence of ambient temperature on trace-element compositions of Uvigerina species. Trifarina angulosa may have Mg/Ca compositions which are positively related to temperature, but Sr/Ca values seem unaffected by temperature. This may be due to pore-water influences on infaunal tests or to vital effects. Although more modern data are needed, our present results suggest that Sr/Ca ratios and possibly Mg/Ca ratios of some benthic foraminifera have the potential to be useful paleothermometers, at least within a temperature range of −2 to 6°C.     

Late Holocene deep-sea benthic foraminifera from the Sea of Marmara

The benthic foraminiferal assemblages of two cores from the late-Holocene, organic-carbon-rich and carbonate-poor, deep-sea sediments of the eastern depression of the Sea of Marmara have been studied. They were deposited under high level of primary productivity and poorly oxygenated bottom-water conditions; they show low diversity and are dominated by a group of species adapted to an infaunal life style with wide bathymetric distribution (ca. 70–2000 m) in the Mediterranean Sea. Oxygen deficiency down to about 0.5 ml/l does not seem adversely to affect the rate of reproduction of the dominant species belonging to Melonis, Chilostomella, buliminids, and bolivinitids. Their distribution is primarily controlled by substrate conditions.Faunal similarities with fossil assemblages in association with some late-Quaternary sapropels and related facies from the eastern Mediterranean basins suggest that they were deposited under palaeo-oceanographic conditions closely similar to those of the modern Sea of Marmara.     

Distribution of live benthic foraminifera in the Ría de Vigo (NW Spain)

The spatial and vertical distributions of live benthic foraminifera from the muddy central axis of the Ría de Vigo were examined under downwelling and upwelling conditions. The spatial distribution of the main benthic foraminiferal species is controlled by salinity variations (San Simón Assemblage), food availability (Outer Ría Assemblage) and oxygen concentrations and/or reducing microenvironmental conditions in the sediment (Inner Ría Assemblage).The microhabitat distribution of benthic foraminifera is mainly dependant on the oxygenation/redox conditions in the sediment and the vertical segregation of food. In the inner areas of the ría, low oxygen and/or reducing microenvironmental conditions prevent the vertical segregation of microhabitats. In outer areas oxygen penetration is deep but microhabitat partitioning only occurs in the presence of additional food resources (mainly fresh organic carbon from phytoplankton blooms) which triggers the colonisation of the surface layer by new species and the migration upwards and reproduction of opportunistic species.     

Morphology and microhabitat preferences of benthic foraminifera from the northwest Atlantic Ocean

The distribution of Rose Bengal stained calcareous benthic foraminifera was determined in six ☐ cores raised from water depths between 200 and 3000 m on the Nova Scotian continental margin and Gulf of Maine. The taxa can be separated into four microhabitats within the surficial sediments. Epifaunal taxa are generally found in the top cm, intermediate infaunal taxa are found from about 1 to 4 cm and deep infaunal taxa are found at > 4 cm sediment depth in at least one ☐ core. A fourth group, shallow infaunal taxa, is found in the top 2 cm and is inferred to be infaunal based on wall porosity characteristics and test shapes similar to infaunal taxa. The epifaunal, shallow infaunal and intermediate infaunal taxa maintain their positions within the sediments from core to core, whereas the deep infaunal taxa are found at progressively shallower sediment depths in cores within increasing organic carbon contents from shallower water depths.Each microhabitat category has distinct morphological characteristics. Epifaunal taxa have plano-covex or biconvex cross sections, trochospiral coiling and large pores absent or found on only one side. Shallow infaunal taxa have uniserial, triserial, or planispiral coiling, with surface ornamentation present on a number of taxa. The intermediate infaunal taxa have rounded peripheries, pores over the entire test and planispiral coiling, with the exception ofCibicidoides bradyi which has trochospiral coiling. The deep infaunal taxa have, in general, planispiral or triserial coiling with cylindrical or ovate shaped tests.     

Baseline study of soft bottom benthic assemblages in the Bay of Santander (Gulf of Biscay)

The Bay of Santander constitutes one of the most important estuarine areas in northern Spain (Gulf of Biscay) which has been significantly stressed by sewage discharges. However, an improvement in the environmental quality of the bay is expected once the new sewer system comes into operation. The evaluation of such an evolution needs to be contrasted with the current situation, established in specific baseline studies. For this purpose, a field study was carried out on the soft bottom assemblages of the estuary in spring of 1998. At each station, the benthic fauna abundance, grain size, organic matter, carbon, nitrogen, phosphorus and heavy metals concentrations were analysed. The community structure and the physical and chemical characteristics of the sediment reflected the higher impacts both near the points of discharges and in the low water exchange zones, whereas those areas showing higher tidal flows presented an acceptable quality. High heavy metal concentrations for Cd, Hg, Pb and Zn indicated contamination of industrial sources in several areas of the Bay. This information allows to establish a reference situation in order to improve the design of a specific monitoring program for this area.     

Recent benthic foraminifera from the continental margin of northwest Africa: Community structure and distribution

Sediment grab samples were collected at 107 locations along the continental margin of northwest Africa. These samples form a series of depth transects between the Straits of Gibraltar and Dakar, Senegal. The greater than 250 μm size fraction was retained for a census of both the live and dead foraminifera. After deleting trace occurrences, Jaccard and correlation coefficient-based cluster analyses were performed to decipher the community structure for this margin.The geographic distribution of the four major faunal provinces recognized is conventional in that for both the live and dead assemblages three are related to depth: upper slope and shelf, middle slope, and lower slope and continental rise. The biotopes and thanatotopes within these provinces are strongly restricted along vertical or latitudinal boundaries when the data are analysed using Jaccard coefficients. Dendrograms constructed from correlation coefficients emphasize depth-related faunal communities. Dominant species such asCibicides lobatulus, Trifarina fornasinii, Planulina ariminensis, Uvigerina finisterrensis, andC. wuellerstorfi are generally distributed within a definite depth range along this margin and strongly influence the correlation coefficient-based dendrograms. Other distributions are clearly not depth-related but correspond to various environmental variables, for example:Cancris auriculus — coarse substrate;Bolivina subaenariensis — oxygen minimum;Cibicidoides kullenbergi andHoeglundina elegans — low organic carbon.Uvigerina peregrina is also a notable exception to depth-dependence in that this dominant species and the province it represents are generally confined to the continental slope south of Cape Blanc. Its preference for that region may arise from the high organic carbon and fine grain-size of the sediment there, from low salinity of the bottom water, or from other unknown variables. The most important vertical faunal boundary occurs between Cape Barbas and Cape Blanc (between 22° and 21°N latitude). In contrast, the Canary Island Ridge intersects the continental margin at a saddle depth of about 1500 m, but that physiographic barrier has little effect on provincialism of the deep-water benthic foraminifera.     

Stable carbon isotopes and the metabolism of the terrestrial Devonian organism Spongiophyton

Devonian fossils of Spongiophyton have been identified as a terrestrial evolutionary intermediate between algae and vascular land plants on the basis of their dichotomously branched, tubular morphology, thick cuticles, and the scattered distribution of surface pores. Our understanding of their physiology is, however, severely limited, but may be increased through the use of stable carbon isotope measurements. One such study led to the hypothesis that Spongiophyton carried a carbon isotope (δ¹³C) signature characteristic of lichens ( Jahren et al., 2003 Geology 31 , 99–102). Here, we outline three difficulties with accepting this idea and report independent isotopic measurements of Spongiophyton fossils from Canada and Ghana. Our results show that the isotopic discrimination of analogous tissues of Spongiophyton, extant lichens, liverworts and mosses are statistically indistinguishable. We suggest therefore that claims to have definitively identified lichen metabolism are premature, and cannot be sustained.     

Climate-related variability in carbon and oxygen stable isotopes among populations of Aleppo pine grown in common-garden tests

The Aleppo pine (Pinus halepensis Mill.) is found in the Mediterranean under a broad range of moisture and thermal conditions. Differences in severity and duration of water stress among native habitats may act as selective forces shaping the populations’ genetic make-up in terms of contrasting drought strategies. We hypothesised that these strategies should translate into intraspecific variation in carbon isotope composition (δ¹³C, surrogate of intrinsic water-use efficiency, WUE_i) of wood holocellulose, and such variation might be linked to changes in oxygen isotope composition (δ¹⁸O, proxy of stomatal conductance) and to some climatic features at origin. Thus, we evaluated δ¹³C, δ¹⁸O, growth and survival for 25 Aleppo pine populations covering its geographic range and grown in two common-garden tests. We found intraspecific variability for δ¹³C and growth, with high-WUE_i populations (which showed ¹⁸O-enriched holocellulose) having low growth. These results suggest stomatal regulation as common control for δ¹³C and productivity. We also detected sizeable relationships between δ¹³C and climate factors related to the magnitude and timing of drought such as the ratio of summer to annual rainfall. The main climate variable associated with δ¹⁸O was minimum temperature, but only in the coldest trial, suggesting differences in growth rhythms among sources. Overall, slow growing populations from highly-seasonal dry areas of the western Mediterranean exhibited a conservative water-use, as opposed to fast growing sources from the northernmost distribution range. The particular behaviour of the Mediterranean Aleppo pine as compared with other conifers demonstrates different selective roles of climate variables in determining intraspecific fitness.