Upper Ordovician sequences of western Estonia

The Upper Ordovician (uppermost Caradoc-Ashgill) section of western Estonia consists of a series of seven open-shelf carbonate sequences. Depositional facies grade laterally through a series of shelf-to-basin facies belts: grain-supported facies (shallow shelf), mixed facies (middle shelf), mud-supported facies (deep shelf and slope) and black shale facies (basin). Locally, a stromatactis mud mound occurs in a middle-to-deep shelf position. Shallow-to-deep shelf facies occur widely across the Estonian Shelf and grade laterally through a transitional (slope) belt into the basinal deposits of the Livonian Basin.

Each sequence consists of a shallowing-upward, prograding facies succession. Sequences 1 (Upper Nabala Stage) and 2 (Vormsi Stage) record step-wise drowning of underlying shelf units (lower Nabala) that culminated in the deposition of the most basinal facies (Fjäcka Shale) in the Livonian Basin. Sequences 3–6 comprise the overlying Pirgu Stage and record the gradual expansion of shallow and middle-shelf facies across the Estonian Shelf. The Porkuni Stage (sequence 7) is bracketed by erosional surfaces and contains the shallowest-water facies of the preserved strata. The uppermost part of the section (Normalograptus persculptus biozone) is restricted to the Livonian Basin, and includes redeposited carbonate and siliciclastic grains; it is the lowstand systems tract of the lowest Silurian sequence 8. Sequence 7 and the overlying basinal redeposited material (i.e., the lowstand of sequence 8) correspond to the latest Ordovician (Hirnantian) glacial interval, and the bracketing unconformities are interpreted as the widely recognized early and late Hirnantian glacial maximums.

The sequences appear correlative to Upper Ordovician sequences in Laurentia. Graptolite biozones indicated that the Estonian sequences are equivalent to carbonate ramp sequences in the western United States (Great Basin) and mixed carbonate-siliciclastic sequences in the eastern United States (Appalachian Basin–Cincinnati Arch region). These correlations indicate a strong eustatic control over sequence development despite the contrasting tectonic settings of these basins.     

Late Ordovician (Turinian–Chatfieldian) carbon isotope excursions and their stratigraphic and paleoceanographic significance

Five positive carbon isotope excursions are reported from Platteville–Decorah strata in the Upper Mississippi Valley. All occur in subtidal carbonate strata, and are recognized in the Mifflin, Grand Detour, Quimbys Mill, Spechts Ferry, and Guttenberg intervals. The positive carbon isotope excursions are developed in a Platteville–Decorah succession in which background δ¹³C values increase upward from about −2‰ at the base to about 0‰ Vienna Pee Dee belemnite (VPDB) at the top. A regional north–south δ¹³C gradient, with lighter values to the north and heavier values to the south is also noted. Peak excursion δ¹³C values of up to +2.75 are reported from the Quimbys Mill excursion, and up to +2.6 from the Guttenberg excursion, although there are considerable local changes in the magnitudes of these events. The Quimbys Mill, Spechts Ferry, and Guttenberg carbon isotope excursions occur in units that are bounded by submarine disconformities, and completely starve out in deeper, more offshore areas. Closely spaced chemostratigraphic profiles of these sculpted, pyrite-impregnated hardground surfaces show that they are associated with very abrupt centimeter-scale negative δ¹³C shifts of up to several per mil, possibly resulting from the local diagenetic effects of incursions of euxinic bottom waters during marine flooding events.     

Late Devonian carbon isotope stratigraphy and sea level fluctuations, Canning Basin, Western Australia

The Upper Devonian reef complexes of the Canning Basin contain some of the world’s best exposed, continuous stratigraphic sections through the Frasnian-Famennian boundary. The facies distribution and composition of these reef complexes record interactions among sea level changes, sediment supply, ocean chemistry, and paleoecology. Changes in relative sea level produced spatial shifts in reef platform development and regional changes in sediment supply that can be correlated across facies boundaries using a combination of sequence stratigraphy, biostratigraphy, and carbon isotope stratigraphy. During the lowstand interval below the Frasnian-Famennian boundary, the reef margin advanced down the reef slope in shallow-water environments, and siliciclastics locally dominated in the marginal slope environment. Compilation of a broad late Frasnian to early Famennian sequence stratigraphic framework for the Canning Basin demonstrates that transgressive intervals correlate to positive carbon isotopic excursions within the basin. These isotopic shifts also can be correlated to time-equivalent positive carbon isotopic excursions reported from transgressive intervals in Europe. Thus, the late Frasnian transgressions in the Canning Basin were primarily eustatic rather than tectonic in origin, and positive carbon isotopic signatures of the Kellwasser horizons are globally correlative.     

The Middle Caradoc Facies and Faunal Turnover in the Late Ordovician Baltoscandian palaeobasin

A Late Ordovician episode of remarkable biotic, climatic, sea level and facies changes, named here as the Middle Caradoc Facies and Faunal Turnover, took place in the Baltoscandian area. This paper presents an integrated overview of these changes in the critical middle Caradoc interval. Data are given on carbonate rock composition, distribution and grain-size composition of the siliciclastic material and the carbon isotopic composition of whole-rock carbonates in cores of Estonia and Sweden.

The Middle Caradoc Facies and Faunal Turnover can be described as a succession of related environmental changes. The turnover began with a positive excursion in carbonate δ¹³C and continued with sea level changes that led to a sedimentary hiatus on the shelf and a change from carbonate-dominated to siliciclastic-dominated sedimentation in the basin. The turnover ended with an extinction event and associated microfaunal crisis.

The middle Caradoc turnover in Baltoscandia is comparable to a similar succession of changes in North America. The turnover affected two palaeocontinents, and reflects a widespread, possibly global environmental change. Onset of glaciation on Gondwana and/or increased orogenic activity might have initiated the changes in ocean circulation and led to the initial carbon isotope excursion. The following sea level rise and faunal changes affected several different continents.     

Aspects of recent advances in the Ordovician stratigraphy and palaeontology of China

The continuous and richly fossiliferous Ordovician succession of China (particularly South China) comprises a heterogeneous suite of litho- and bio-facies, which has been a main focus of stratigraphical and palaeontological research in recent years. Among the seven GSSPs established in China, three are within the Ordovician System, and the GSSP of the Darriwilian Stage at Huangnitang, Changshan County, western Zhejiang Province, was the first “golden spike” in China and the first for the Ordovician System. A series of case studies have revealed that: (1) the Ordovician radiation of some fossil groups on the Upper Yangtze Platform (e.g., brachiopods and graptolites) reached their first α-diversity acme in the Didymograptellus eobifidus Biozone, four zones earlier than the global trend; (2) the β-diversity peak was attained 3–4 zones later than the α-diversity peak; (3) many brachiopod communities or faunas first occurred in the central part of the Upper Yangtze Platform and subsequently expanded to both more offshore and near-shore facies; (4) diachroneity existed in many aspects of the radiation. The end-Ordovician mass extinction was a severe event in South China. Two pulses of the extinction are recognized for a number of major fossil groups, some being most strongly affected during the first pulse whereas the others suffered during the second pulse. Macroevolution during the Ordovician–Silurian transition has been investigated in detail, and the role of the Lazarus effect has been found to be less important than previously believed.     

Borings in trepostome bryozoans from the Ordovician of Estonia: two ichnogenera produced by a single maker, a case of host morphology control

The evolution of borings has shown that the morphology of borings is a function of both the borer and its substrate. This study investigated the effect of bryozoan internal skeletal morphology on the dimensions and distribution of borings. One hundred and forty-three trepostome colonies from the Middle and Upper Ordovician strata of northern Estonia were examined. Of these, 80% were matrix entombed, longitudinally sectioned ramose and hemispherical colonies, and 20% were matrix-free hemispherical colonies that allowed examination of the colony surfaces. Seventy-one percent of the ramose colonies were bored, whereas 88% of the hemispherical colonies were bored. On average, only 8% of colony surface areas were bored out. Borings were more randomly oriented in the hemispherical colonies. In contrast in the ramose colonies, the borings tended to more restricted to the thin-walled endozone and thus parallel to the branch axis. This is interpreted to be a function of the thick-walled exozones controlling to some extent where the borer could bore. Based on morphology, the borings in the hemispherical colonies are referred to Trypanites and those in the ramose colonies to Sanctum . Sanctum is revised to include two possible openings and to recognize that boring shapes were inherently constrained by the thick-walled exozones of the host bryozoan colonies. Both trace fossils were probably produced by a boring polychaete that used the tubes as domiciles.     

Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

Lower Aptian carbon isotope stratigraphy from a distal carbonate shelf setting: the Cau section, Prebetic zone, SE Spain

A δ¹³C curve is reported for the latest Barremian to Early Aptian at a section located in the Prebetic zone (Cau section, SE Spain). The studied section records a hemipelagic succession of dark shales, deposited on a distal carbonate ramp with a high subsidence rate, adjacent to shallow carbonate environments. The integrated biostratigraphy of the section is based on ammonites, planktonic foraminifera and calcareous nannofossils, and it has allowed an accurate dating of the succession. The δ¹³C curve presented shows a distinctive evolution, leading to the recognition of three major excursions, as well as a subdivision into eight segments, which represents an improvement of the current biostratigraphic resolution. The correlation, both isotopic and biostratigraphical, with other well resolved sections is very accurate even at the higher resolution attained. Correlation with sections with low-resolution biostratigraphic characterisation from shallow platform limestones also gives good results, which supports the effectiveness of carbon isotope stratigraphy as a correlation tool.     

Huemacaspis (Trilobita, Kerfornellinae) from the Late Ordovician of the Argentine Cordillera Oriental

The trilobite genus Huemacaspis P?IBYL and VANěK, 1980 is redescribed from well-preserved material from the Santa Gertrudis Formation (early Caradoc), Argentine Cordillera Oriental. The new species H. gallinatoensis is described and figured. Huemacaspis is widely distributed in the southern part of the Central Andean basin during the early Caradoc and it is a key taxon for the correlation of different units from the Cordillera Oriental and Sierras Subandinas of Argentina and the Cordillera Oriental of Bolivia. This taxon is one of the few records of trilobites of early Caradoc age found in the shallow shelf environments of southwestern Gondwana. Huemacaspis is accommodated within the Kerfornellinae, a subfamily otherwise known from southern Central Europe and northern Africa. The trilobite fauna from the early Caradoc of the Central Andean basin suggests a faunal exchange with the latter regions indicating the persistence of a broadly similar biogeographic pattern to that documented in earlier Ordovician times.     

Ordovician meteoric carbon and oxygen isotopic values: implications for the latitudinal variations of ancient stable isotopic values

Columnar and clear blocky calcite cement from a Middle Ordovician carbonate succession in east Tennessee is interpreted as meteoric in origin. Columnar and clear blocky calcite from this succession does not show extremely large ¹³C depletions reported from meteoric phases of younger rocks. Meteoric fluid δ¹⁸O values calculated from clear blocky calcite are 2 to 3‰ more negative than approximately coeval sea water; a relationship typical of modern, low-latitude, coastal meteoric water. Comparison with meteoric δ¹⁸O values from Ordovician units elsewhere suggests that the geographic distribution of these values may be broadly similar to that observed today. Therefore, we tentatively suggest that geographic distribution of meteoric δ¹⁸O values during both icehouse and greenhouse eras are similar.     

Seasonal carbon isotope discrimination in a grassland community

Summary Grassland communities of arid western North America are often characterized by a seasonal increase in ambient temperature and evaporative demand and a corresponding decline in soil moisture availability. As the environment changes, particular species could respond differently, which should be reflected in a number of physiological processes. Carbon isotope discrimination varies during photosynthetic activity as a function of both stomatal aperture and the biochemistry of the fixation process, and provides an integrated measure of plant response to seasonal changes in the environment. We measured the seasonal course of carbon isotope discrimination in 42 grassland species to evaluate changes in gas exchange processes in response to these varying environmental factors. The seasonal courses were then used to identify community-wide patterns associated with life form, with phenology and with differences between grasses and forbs. Significant differences were detected in the following comparisons: (1) Carbon isotope discrimination decreased throughout the growing season; (2) perennial species discriminated less than annual species; (3) grasses discriminated less than forbs; and (4) early flowering species discriminated more than the later flowering ones. These comparisons suggested that (1) species active only during the initial, less stressful months of the growing season used water less efficiently, and (2) that physiological responses increasing the ratio of carbon fixed to water lost were common in these grassland species, and were correlated with the increase in evaporative demand and the decrease in soil moisture.     

Magnesian calcite and chamositic ooids forming shoals peripheral to Late Ordovician (Ashgill) muddy siliciclastic shores: southern Ontario

In southern Ontario, ooids are associated with two distinct facies associations in the Queenston Formation, the final stage of Late Ordovician (Ashgill) Taconic basin fill. One facies consists of thin ooid and bioclastic grainstones interbedded with mudrock, and lies near the base of the formation, and, in southwestern Ontario, also forms a local NW-thickening wedge near the middle of the formation. Ooids have radial-fibrous and radial-concentric fabrics (Type A), with chamosite, illite, and Fe-oxide laths at intercrystalline sites. Vertical lithologic and ooid abundance patterns indicate that thresholds to carbonate production were sensitive to changes in terrigenous sediment supply, sea level, circulation, accommodation space, and tectonism.

Ooids in the second facies association are admixed with abraded fragments of open-marine biota, or occur burrow fills, within a <30-cm-thick interval of mudrock near the top of the preserved Queenston succession, a few metres below the Ordovician–Silurian unconformity. Ooids have radial concentric and crosscutting patchy microcrystalline fabrics (Type B). This unit may represent a transgressive or stillstand deposit modified by bioturbation.

The extent of preserved fabric suggests that both ooid types were originally magnesian calcite, but Type A ooids underwent greater burial alteration. This is shown by crystalline mosaics that cross-cut relict primary fabrics; δ¹³C values (−1.82‰ to +0.67‰) and δ¹⁸O values (−4.46‰ to −10.57‰) more negative than marine calcite of similar age; Mn and Fe concentrations more elevated above expected marine values; and a luminescence similar to that of intergranular cements. Burial meteoric diagenesis was likely promoted by excellent permeability of the host sand. We interpret authigenic chamosite and Fe-oxide to reflect diagenesis of iron-bearing and clay detritus trapped during ooid growth. Type B ooids suffered less alteration: δ¹³C (+1.1‰ to +6.64‰) and δ¹⁸O (−3.04‰ to −4.81‰) values overlap the expected marine range, including ¹³C enrichment that occurs within the Hirnantian (latest Ordovician) excursion. Although Mn and Fe values are still higher than those of modern calcitic ooids, negligible luminescence suggests that recrystallization occurred in the presence of marine-derived pore fluids. Further burial alteration was inhibited due to low permeability of the host mud.

Type A ooid facies in the Queenston Formation forms an ancient analogue for lesser known Quaternary ooid shoals peripheral to tropical deltaic systems. The facies of Type B ooids, while more enigmatic, may preserve a geochemical herald of latest Ordovician climate change. The presence of minor chamosite in Type A ooids defines a possible distal facies of the well-known oolitic ironstones of similar age in the mid-continental USA.     

Sources and fates of dissolved organic carbon in lakes as determined by whole-lake carbon isotope additions

Four whole-lake inorganic ¹³C addition experiments were conducted in lakes of differing trophic status. Inorganic ¹³C addition enriched algal carbon in ¹³C and changed the C-DOC by +1.5‰ to +9.5‰, depending on the specific lake. This change in C-DOC represented a significant input of algal DOC that was not completely consumed by bacteria. We modeled the dynamics in C-DOC to estimate the fluxes of algal and terrestrial carbon to and from the DOC pool, and determine the composition of the standing stock. Two experiments in lightly stained, oligotrophic lakes indicated that algal production was the source of about 20% of the DOC pool. In the following year, the experiment was repeated in one of these lakes under conditions of nutrient enrichment, and in a third, more humic lake. Algal contributions to the DOC pool were 40% in the nutrient enriched lake and 5% in the more humic lake. Spectroscopic and elemental analyses corroborated the presence of increased algal DOC in the nutrient enriched lake. Natural abundance measurements of the C of DOC in 32 lakes also revealed the dual contributions of both terrestrial and algal carbon to DOC. From these results, we suggest an approach for inferring the contribution of algal and terrestrial DOC using easily measurable parameters.     

Resource allocation to reproduction and soma in Drosophila: a stable isotope analysis of carbon from dietary sugar

Metabolic resources in adults of holometabolous insects may derive either from larval or adult feeding. In Drosophila melanogaster, reproduction and lifespan are differently affected by larval vs. adult resource availability, and it is unknown how larval vs. adult acquired nutrients are differentially allocated to somatic and reproductive function. Here we describe the allocation of carbon derived from dietary sugar in aging female D. melanogaster. Larval and adult flies were fed diets contrasting in sucrose (13)C/(12)C, from which we determined the extent to which carbon acquired at each stage contributed to adult somatic tissue and to egg manufacture. Dietary sugar is very important in egg provisioning; at every age, roughly one half of the carbon in eggs was derived from sugar, which turned over from predominantly larval to entirely adult dietary sources. Sucrose provided approximately 40% of total somatic carbon, of which adult dietary sucrose came to supply approximately 75%. Unlike in eggs, however, adult acquired sucrose did not entirely replace the somatic carbon from larvally acquired sucrose. Because carbon from larval sucrose appears to be fairly "replaceable", larval sucrose cannot be a limiting substrate in resource allocation between reproduction and lifespan.     

Discovery of the Ordovician Millbrig K-bentonite Bed in the Trenton Group of New York State: implications for regional correlation and sequence stratigraphy in eastern North America

The Ordovician (Chatfieldian) Millbrig K-bentonite Bed is a key stratigraphic marker horizon that is regionally synchronous over much of eastern and central North America. This prominent marker is an independent source of correlation among the major chronostratigraphic and sequence stratigraphic units in this region. The general stratigraphic position of the Millbrig K-bentonite has suggested to some authors that it is identical with the Hounsfield K-bentonite at Dexter, New York (the traditional type area of the Middle Ordovician, or Mohawkian Series in North America), but previously available geochemical and biostratigraphical evidence has been insufficient to confirm this correlation. Analyses of apatites and melt inclusions in quartz phenocrysts from the Millbrig K-bentonite at eight localities in Kentucky, Missouri, Virginia, and Pennsylvania and the Hounsfield K-bentonite at its type locality at Dexter, New York, using high-precision electron microprobe analysis techniques shows that the Millbrig and the Hounsfield have identical apatite and melt inclusion chemistry indicating a geochemical correlation. This correlation is supported by conodont biostratigraphy, and δ¹³C isotope chemostratigraphy. The new data demonstrate that the Millbrig K-bentonite, and therefore the base of the Chatfieldian Stage of the North American Mohawkian Series (by definition), lies very close to the base of the traditional Rocklandian Stage of New York. Furthermore, the Millbrig K-bentonite Bed lies in close proximity to the base of the Taconic supersequence over much of the Midcontinent region and in particular lies just below the M5 sequence boundary recognized in Kentucky and Tennessee. Our results permit extension of the Chatfieldian sequences into New York State and southern Ontario, and contributes to the resolution of the long-standing uncertainty about the position of the base of the Trenton Group in Ontario. Furthermore, viewed in a broad context of regional stratigraphic relations, we conclude that our results suggest that the persistent correlation difficulties reflect diachronous effects of widespread changes in oceanographic circulation patterns that emerged during Taconic Orogeny. Finally, regional differences in the timing and character of sequence bounding surfaces and facies similarities summarized here suggest that the causes of relative sea level change during the Chatfieldian may have been primarily tectonoeustatic mechanisms.     

Correlation of (Ordovician, Mohawkian) K-bentonites in the upper Mississippi valley using apatite chemistry: implications for stratigraphic interpretation of the mixed carbonate-siliciclastic Decorah Formation

Four Ordovician K-bentonites have been chemically fingerprinted using trace element content of apatite phenocrysts contained within the altered ash layers. Trace element analysis was performed using electron microprobe on individual crystals. The bentonites include the Deicke, Millbrig, Elkport, and Dickeyville beds. These beds lie within the Decorah Formation of the northern Mississippi Valley outcrop area. All four K-bentonite beds are believed to have a Taconic source whereas volcanic ash was transported by wind and deposited in the Ordovician North America epeiric sea.

The Decorah is a marine unit, shaly to the north and west and carbonate-rich to the south and east. Thirteen K-bentonite samples were collected from six localities. Only one locality (Dickeyville, WI) contains all four beds in succession. One hundred seventy-one apatite crystals were handpicked and analyzed for major, minor and trace elements using an electron microprobe. Each crystal was analyzed at three to six separate spots to ascertain compositional variation. The most discriminating and diagnostic elements were Mg and Mn; plotting Mg vs. Mn content produces data clusters characteristic for each K-bentonite. Clusters show only minimal change between localities, allowing individual K-bentonites to be identified and used for stratigraphic correlation.

Time slices provided by the K-bentonite horizons show that the Decorah Formation is composed of reciprocal wedges (stratigraphic sequences) of shale and carbonate. Shale was deposited in deeper water within the Hollandale Embayment but pinched out southeastward onto the flank of the Wisconsin Dome. Carbonates were formed largely in shoal water on the arch, prograding northwestward and shaling out downramp into the embayment.     

Oxygen and carbon isotope compositions of middle Cretaceous vertebrates from North Africa and Brazil: Ecological and environmental significance

In order to investigate mid-Cretaceous terrestrial climates of low paleolatitudes, Moroccan, Tunisian and Brazilian vertebrate apatites have been analyzed for their oxygen and carbon isotope compositions of phosphates (δ¹⁸O_p) and carbonates (δ¹⁸O_c, δ¹³C_c). At each site, coexisting theropod dinosaurs, titanosaurid sauropods, pterosaurs, crocodilians, turtles and fish have distinct δ¹⁸O_p and δ¹³C_c values reflecting their ecologies, diets and foraging environments. Oxygen isotope compositions of surface waters (δ¹⁸O_w) estimated from turtle and crocodile δ¹⁸O_p values range from − 5.0 ± 1.0‰ to − 2.4 ± 1.0‰, which do not differ from mean annual rainwater values occurring today under inter-tropical sub-arid to arid climates. High water temperatures ranging from 21 ± 6 °C to 34 ± 2 °C deduced from fish δ¹⁸O_p values are in agreement with those published for mid-Cretaceous low latitudes. Temporary or seasonal droughts are inferred from high δ¹⁸O_p values of lungfish teeth, even though lower reptile δ¹⁸O_p values suggest the use of distinct and most likely larger or regularly renewed bodies of water. Environmental conditions of the studied low latitude regions during the Aptian-Cenomanian interval were somewhat similar to those experienced today under semi-arid to arid tropical or equatorial climates, but with higher mean surface temperatures than present-day ones.     

Stable isotopes, ecological integration and environmental change: wolves record atmospheric carbon isotope trend better than tree rings

Large-scale patterns of isotope ratios are detectable in the tissues of organisms, but the variability in these patterns often obscures detection of environmental trends. We show that plants and animals at lower trophic levels are relatively poor indicators of the temporal trend in atmospheric carbon isotope ratios (delta13C) when compared with animals at higher trophic levels. First, we tested how differences in atmospheric delta13C values were transferred across three trophic levels. Second, we compared contemporary delta13C trends (1961-2004) in atmospheric CO2 to delta13C patterns in a tree species (jack pine, Pinus banksiana), large herbivore (moose, Alces alces) and large carnivore (grey wolf, Canis lupus) from North America. Third, we compared palaeontological (approx. 30000 to 12000 14C years before present) atmospheric CO2 trends to delta13C patterns in a tree species (Pinus flexilis, Juniperus sp.), a megaherbivore (bison, Bison antiquus) and a large carnivore (dire wolf, Canis dirus) from the La Brea tar pits (southern California, USA) and Great Basin (western USA). Contrary to previous expectations, we found that the environmental isotope pattern is better represented with increasing trophic level. Our results indicate that museum specimens of large carnivores would best reflect large-scale spatial and temporal patterns of carbon isotopes in the palaeontological record because top predators can act as ecological integrators of environmental change.     

Response of Late Ordovician paleoceanography to changes in sea level, continental drift, and atmospheric pCO2: potential causes for long-term cooling and glaciation

We performed sensitivity experiments using an ocean general circulation model at two stages of the Late Ordovician (Caradoc, 454 Ma; Ashgill, 446 Ma) under a range of atmospheric pCO₂ values (8–18× PAL; pre-industrial atmospheric level) at high and low sea level.

The model results indicate that the long-term cooling trend during the Late Ordovician can be explained by progressive cooling of the global ocean in response to falling levels of atmospheric pCO₂, sea level change, and paleogeographic change. These results also explain the occurrence of low latitude cool-water carbonates in North America.

In all simulations, a drop in sea level led to a reduction in poleward ocean heat transport. This indicates a possible positive feedback that could have enhanced global cooling in response to sea level drop during the Late Ordovician. Alterations in poleward ocean heat transport linked to changes of atmospheric pCO₂ also indicate that there is a threshold of 10× PAL, above which ocean current change cannot be responsible for glaciation in the Late Ordovician. Continental drift could explain the observed global cooling trend in the Late Ordovician through a combined poleward ocean heat transport feedback and increased ice-albedo effect if atmospheric pCO₂ was low during the entire Late Ordovician.

The model results further indicate that the response of meridional overturning to changes in paleogeography, atmospheric pCO₂, and sea level is stronger than the response of surface circulation to these perturbations. Because the overturning circulation is so strong, meridional overturning was the dominant mechanism for described changes in heat transport in the Late Ordovician.     

Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state

Carbon stable isotope analysis was carried out on zooplankton from 24 United Kingdom lakes to examine the hypothesis that zooplankton dependence on allochthonous sources of organic carbon declines with increasing lake trophy. Stable isotope analysis was also carried out on particulate and dissolved organic matter (POM and DOM) and, in 11 of the lakes, of phytoplankton isolates. In 21 of the 24 lakes, the zooplankton were depleted in ¹³C relative to bulk POM, consistent with previous reports. δ¹³C for POM showed relatively little variation between lakes compared to high variation in values for DOM and phytoplankton. δ¹³C values for phytoplankton and POM converged with increasing lake trophy, consistent with the expected greater contribution of autochthonous production to the total organic matter pool in eutrophic lakes. The difference between δ¹³C for zooplankton and that for POM was also greatest in oligotrophic lakes and reduced in mesotrophic lakes, in accordance with the hypothesis that increasing lake trophic state leads to greater dependence of zooplankton on phytoplankton production. However, the difference increased again in hypertrophic lakes, where higher δ¹³C values for POM may have been due to greater inputs of ¹³C-enriched organic matter from the littoral zone. The very wide variation in phytoplankton δ¹³C between lakes of all trophic categories made it difficult to detect robust patterns in the variation in δ¹³C for zooplankton. Received: 2 November 1998 / Accepted: 3 December 1999