The late Kimmeridgian Nusplingen Plattenkalk (palaeolatitude ~30°N) is one of the well-known European Plattenkalk konservatlagerstätten. The laminated limestones of this lagerstätte have been deposited in a shallow lagoon, surrounded by sponge-microbial mounds, some of which formed small islands. Plattenkalk sediments are often thought to have been deposited below a halocline, which was induced by high evaporation rates. By measuring the stable isotope composition of belemnite rostra (n = 27) of the species Hibolithes semisulcatus, the depositional environment of the Nusplingen Plattenkalk has been investigated. Cathodoluminescence-microscopy and trace element analysis have been applied to check for diagenetic alteration of the studied rostra. The mean δ13C of the well-preserved rostra is +0.03 ± 0.27‰, the mean δ18O −0.68 ± 0.3‰. A narrow range of stable isotope data acquired from an accumulation of rostra, regurgitated by a fish or reptile, supports the notion that belemnite calcite reflects environmental conditions. The palaeontological and sedimentological findings suggest that H. semisulcatus was autochthonous to the Nusplingen Plattenkalk. Anoxic conditions prevailed in the sediments and temporarily in the water-column above the sediment/bottom water interface. A nektobenthic life style can be excluded for the Nusplingen belemnites; a demersal life style seems unlikely. Comparison with δ18Obel data from other latitudes indicates that a latitudinal gradient played a role in the δ18O composition of seawater. A pelagic, inner shelf setting is therefore suggested for the Plattenkalk, an interpretation that is supported by palaeontological evidence. It is here proposed that the formation of the Nusplingen Plattenkalk was not caused by salinity stratification. It is further suggested that the belemnites were not restricted to deep-water settings, but also occurred in shallow waters and higher in the water-column.


  1. Belemnites
  2. CL-microscopy
  3. Nusplingen Plattenkalk
  4. palaeotemperature
  5. salinity
  6. stable isotopes
The Late Jurassic Plattenkalke of Central Europe are well-known for their exceptional fossil preservation. Complete fossils showing soft-part preservation are common in the Nusplingen Plattenkalk (NP), exposed near the village of Nusplingen in south-western Germany (Fig. ). The NP was deposited during the late Kimmeridgian, thereby being slightly older than the early Tithonian Solnhofen Plattenkalk (Schweigert ). Current models suggest that the deposition of the NP took place in an enclosed lagoon, fringed by sponge-microbial mounds forming small islands (e.g. Dietl et al. ; Dietl & Schweigert ). For the Solnhofen Plattenkalk on the other hand, a widely accepted, but still controversial depositional model suggests deposition of the sediments below a halocline that formed due to high evaporation rates in the nearly enclosed basins (e.g. Barthel et al. ; Keupp et al. ). The salinity of this brine is suggested to have varied from 40–80‰ (Schwark et al. ). The distribution of rare earth elements in the Solnhofen Plattenkalk indicates that this brine was not anoxic (Kemp & Trueman ). While the Solnhofen Plattenkalk sensu stricto is the best studied deposit of its type in southern Germany, other lithologically similar levels reflect markedly different settings (e.g. Swinburne & Hemleben ; Fürsich et al. ).
Fig. 1. Location of the Nusplingen and Egesheim quarries in south-western Germany and palaeogeography during the late Kimmeridgian (modified from Klug et al. ).
The δ18O of the low-Mg calcite of belemnite rostra has been used as a palaeotemperature proxy for the Jurassic and Cretaceous since the studies of Urey et al., because it is believed to be relatively resistant to diagenetic overprint. The δ18O composition of marine biogenic low-Mg calcite skeletons is controlled by temperature and the δ18O of the ambient seawater. Evaporation and precipitation driven changes in the δ18O of the seawater are directly linked to seawater salinity, which results in the shell recording a combination of the two signals. The Mesozoic continents are considered to have had no extensive ice-sheets. The average seawater composition (δ18Osw) during such ice-free times can only be estimated by mass-balance calculations (Shackleton & Kennett ). Armendáriz et al. recently suggested that palaeotemperatures calculated from Mg/Ca ratios of belemnite calcite could be used to differentiate between temperature and salinity signals, which are both recorded in the δ18Obel values.
It remains, however, problematic that modern coleoid cephalopods have not been studied with respect to the applicability of this proxy. The results of other molluscs (bivalves, gastropods) either show a clear correlation between temperature and Mg/Ca (e.g. Klein et al. ) or the metabolism of the organisms largely obscures temperature effects (e.g. Purton et al. ; Izumida et al. ). Some studies of belemnite calcite show a correlation between δ18Obel and Mg/Ca (e.g. Rosales et al. ; McArthur et al. ; Armendáriz et al. ), while others found a better correlation between δ18Obel and Sr/Ca (McArthur et al. ; Li et al. ). Wierzbowski & Rogov did not observe any correlation between δ18Obel and Mg/Ca. Li et al. also did not observe any significant correlation between these two values, neither in belemnite populations nor in single specimens. Differences in the correlation of δ18Obel and Mg/Ca between different belemnite species indicate a presumably species-specific incorporation of trace elements, with species of Hibolithes showing no co-variation of the two values (McArthur et al. ; Wierzbowski & Joachimski ; Wierzbowski et al. ). These results cast doubt on the straightforward interpretation of elemental data in terms of absolute temperature. Absolute temperature determinations are complicate to interpret, due to changes of seawater Mg/Ca ratios through geological time. These are also reflected in fossil biogenic carbonates (e.g. Steuber & Rauch ).
In this study, the trace element and stable isotope data from belemnite rostra (Hibolithes semisulcatus) sampled bed-by-bed from the NP are presented. Included in the data set are four values derived from an accumulation of four regurgitated rostra (‘speiballen’, see Thies & Hauff ). Cathodoluminescence-microscopy and trace element analysis have been applied to check for diagenetic alteration of the rostra. Using these geochemical data, we want to investigate the environment during the formation of the NP. The data should also supply new information about the palaeoecology of H. semisulcatus.

Geological setting

The NP consists of laminated, fine-grained limestones, which are interbedded with coarser grained calciturbidites of varying thickness. The latter contain abundant fragments of siliceous sponges, and rare hermatypic coral fragments, from the reef bioherms surrounding the lagoon. Anoxic conditions prevailed in the sediment or in the bottom waters, but the surface layers supported rich planktonic and nektonic communities. A substantial part of the NP is made up of coccoliths or disintegrated coccoliths (Bantel et al. ).
The Nusplingen lagoon (palaeolatitude ~30°N) was surrounded by small islands, on which the well-preserved land plants, commonly found in the NP, and insects lived. The strata exposed in Nusplingen have a maximum thickness of 17 m (Dietl & Schweigert ). They were deposited in a time span of only a few thousand years (Bantel et al. ), and can be attributed to the late Kimmeridgian Hybonoticeras beckeri Zone (Schweigert ). The lagoonal system had an extent of ca. 1.5 km2 and is thought to have originally covered an area of less than 5 km2. Exact estimates of the expanse of the lagoon are largely impossible due to erosion of most of the NP sediments in this area. The deepest parts of the lagoon had a water-depth of about 80 m (Dietl & Schweigert ).

Material and methods

The belemnites used in this study were collected in the Nusplingen (48°7′26.76″N, 8°52′26.83″E) and Egesheim (48°7′23.20″N, 8°52′22.49″E) quarries (Fig. ). The lower ~2.5 m of the sediments in the Egesheim quarry are slightly older than the strata exposed in the Nusplingen quarry. The strata of both outcrops can be attributed to the Lithacoceras ulmense Subzone, and the Silicisphinctes hoelderi bio-horizon. The individual Plattenkalk packages are labelled Pk 1-6 in the Egesheim quarry and A–L in the Nusplingen quarry (Bantel et al. ). Taxonomically, the 27 belemnite rostra can be attributed to H. semisulcatus (Münster ), one of only two belemnite species recorded from the NP (Schweigert ; Schweigert & Dietl ). H. semisulcatus has been proposed as new type species for Hibolithes de Montfort to replace H. hastatus de Montfort and is considered to be conspecific with H. jurensis (Münster ) (see Riegraf et al. ; Schlegelmilch ). Four belemnites (no. 3–6 in Table ), which were lying only a few centimetres apart from each other on the same bedding plane, are here interpreted as material regurgitated by a predator (elasmobranch, holocephalan, marine reptile?). These four specimens are considered to have lived at the same time and are here being referred to as ‘speiballen’-specimens. Polished thin-sections of three of the ‘speiballen’-specimens have been prepared for CL-analysis to acquire detailed information about their diagenesis. After being covered with a thin layer of gold, the thin-sections were studied with a cathodoluminescence-microscope equipped with a hot-cathode.
Table 1. Stable isotope and elemental data from the Nusplingen and Egesheim quarries H. semisulcatus belemnites (late Kimmeridgian Hybonoticeras beckeri Zone, Lithacoceras ulmense Subzone, Silicisphinctes hoelderi bio-horizon). ‘Speiballen’-specimens are no. 3–6 (grey background).
[‰ V-PDB][‰ V-PDB][ppm][ppm][ppm][ppm][ppm]
*Considered diagenetically altered.
Excluding diagenetically altered samples.
120 cm above base B−0.45−0.6138881029151254161.3
2C 30–400.06−0.8938610030751265262.1
3C 70–78 (1)−0.10−0.95386390312712272576.4
4C 70–78 (2)0.08−0.9437570027301144141.2
5C 70–78 (3)0.27−0.6838368031171250123.2
6C 70–78 (4)−0.08−0.8038397030871263132.4
7D 0–100.25−1.063838902772123932910.1
8D 20–250.050.0038530029241247531.8
9F 5–10−0.21−0.4038631032961314441.0
10F 10–20−0.02−0.4438688030071197942.7
11F 20–250.23−0.5138662031031360231
12G 10–20−0.39−0.8138743026701194251.1
13G 20–30−0.29−0.7038605029481267171.3
14G 40–500.27−0.72382920332712901113.1
15G 60–700.38−0.8538635028791210181.2
16L 5–100.01−1.0738356028671252212.2
17L 40–50−0.130.0738491028661298451.9
18Pk3 0–100.16−0.8738548028251146331.6
19Pk4 10–20 (1)0.83−1.03386490298412693103.7
20Pk4 10–20 (2)0.00−0.8438242033771241313.7
21Pk5 0–5−0.19−0.7538733027911217151.6
22Pk5 0–10−0.08−0.6238433033571244152
23Pk5 35–400.12−0.4638479026321192756.7
24Pk6 0–5 (1)*−1.62−3.4139057015404946210
25Pk6 0–5 (2)−0.15−0.7238516027801132171.5
26Pk6 5–10*−0.05−1.5038458042831312347.7
27Pk6 15–250.58−0.3138362030781328261.4
 Average value0.03−0.6838498029811242662.6
 Standard deviation0.270.30248621156922.2
After preparation and cleaning, the belemnites were split in halves for sampling. The split-half was then shortly dipped in 10% HCl to remove any remaining contaminating carbonate dust still adjacent to the specimen and immediately afterwards rinsed in ultra-pure water. The samples were drilled by hand under a stereomicroscope with a 0.3 mm wide drill bit from clear calcite portions, avoiding the most likely altered regions at the outer margin, the apical line and the apex of the rostrum.
Elemental (Ca, Mg, Sr, Fe, Mn) and stable isotope data (δ13C, δ18O) of 27 samples have been obtained. Elemental analysis was performed by ICP-OES (iCap 6500 Thermo Electron Corporation) at the Ruhr-Universität Bochum. For analysis, samples (~1.5 mg) were dissolved in 3 M HNO3. Stable isotope composition of the samples was then determined at the GeoZentrum Nordbayern, Friedrich-Alexander Universität Erlangen-Nürnberg. Samples reacted with phosphoric acid, and were measured using a Gasbench II connected to a ThermoFinnigan Five Plus mass-spectrometer. Reproducibility and accuracy of measurements were monitored by analysis of international standards (NBS19 and LSVEC). Standard analytical errors have been determined as 0.08‰ or better for δ13C and 0.06‰ or better for δ18O. All isotope data are given in per mil (‰) relative to V-PDB (Vienna Pee Dee Belemnite). All specimens are housed in the Staatliches Museum für Naturkunde, Stuttgart (Germany).


The CL-analysis of the three ‘speiballen’-specimens shows for most parts of the rostra intrinsic, blue CL-colours. Yellow–orange luminescence colours are found in the outer parts of the rostra (Fig. A), near the apical line and the apex (Fig. B, C), and along dark growth rings (laminae obscurae, sensu Müller-Stoll ). Microfractures filled with Yellow–orange luminescent calcite were observed in all parts of the studied rostra.
Fig. 2. CL-images and transmitted light images of belemnite rostra from the ‘speiballen’-specimens. A, transversal section through the middle part of a rostrum (C 70–78 (2)) combined from two images. The outer part of the rostrum shows diagenetic, Mn-rich yellow–orange calcite, mostly along growth rings (laminae obscurae, sensu Müller-Stoll ). The inner rostrum calcite is well-preserved, showing intrinsic, blue CL-colour. White arrows indicate microfractures infilled with diagenetic calcite. Black arrow indicates a scratch mark produced during preparation of the thin section. B, apical region of a rostrum (C 70–78 (4)). Apical line area shows possible enrichment in organic material and possibly former porous nature of the apex. White arrows indicate microfractures, red arrows indicate the heavily dissoluted and re-crystallised outer rostrum margin. C, CL-image and transmitted light image of heavily altered rostrum apex from a ‘speiballen’-specimen (C 70–78 (4)). Black arrows indicate probable former growth rings, originally containing high organic contents. The thickness of the rings increases nearer to the apex. White arrow indicates a well-defined boundary between two growth layers. All scale bars 500 μm.
The Fe contents of the studied rostra are between 12 and 329 ppm (Table ), but only four samples (no. 3, 7, 14, 19) show very high values of >100 ppm. The Mn contents vary between 1.0 and 10.1 ppm. The Sr contents for all samples are above 1100 ppm, except one (no. 24, 494 ppm). Sample no. 26 has a very high Mg content of 4283 ppm.
The carbon and oxygen isotope data are fairly uniform throughout the two sections (Table , Fig. ). The δ13C values range from −0.45 to +0.83‰, excluding negative outlier sample no. 24 with −1.62‰. The δ18O values range from −1.07 to +0.07‰, excluding the negative outlier samples no. 24 and 26 with −3.41 and −1.50‰, respectively. The average value for δ13C is +0.03 ± 0.27‰ and for δ18O −0.68 ± 0.3‰ (excluding samples no. 24 and 26). The data of the four ‘speiballen’-specimens show δ13C values of −0.10 to +0.27‰ and δ18O values of −0.95 to −0.68‰.
Fig. 3. Stable isotope data of H. semisulcatus belemnites from the Egesheim and Nusplingen quarries (late Kimmeridgian, Hybonoticeras beckeri Zone, Lithacoceras ulmense Subzone, Silicisphinctes hoelderi biohorizon). For details about the stratigraphy of the outcrops of the Nusplingen Plattenkalk, see Dietl et al. Black rectangles represent δ18O data from the Egesheim quarry and grey diamonds δ13C. Black crosses represent δ18O data from the Nusplingen quarry and grey triangles δ13C. Arrows indicate position of samples on the stratigraphic column.

Interpretation and discussion

Preservation of isotopic composition

The original mineralogical composition of the belemnite orthorostrum was of low-Mg calcite and a combination of low-Mg calcite and aragonite for the epirostrum (Bandel & Spaeth ). The low-Mg calcite fibres of belemnites show radiaxial-fibrous and fascicular-optic-fibrous fabrics (Richter et al. ). In some belemnite species also the front part of the orthorostrum containing the phragmocone (rostrum cavum) was made up of aragonite (e.g. Spaeth ; Košťák & Wiese ). Still controversial is the original porosity of the rostrum. Some authors (e.g. Veizer ; Spaeth ) suggest a primary porosity of up to 20%, while others argue for an extremely low porosity (e.g. Price & Sellwood ; Monks et al. ; Podlaha et al. ). Infilling of the original pores by early marine diagenetic calcite cement could result in apparently ‘cold’ palaeotemperatures, because the calcite has been precipitated from cold bottom or marine pore waters. This effect has been used to explain apparently too cold palaeotemperatures derived from foraminifera of the Cretaceous tropics (Pearson et al. ). There is, however, no indication, that this applies for the studied belemnites.
The CL-microscopy indicates that the studied rostra are for the most part composed of well-preserved calcite (Fig. ). The Yellow–orange CL-colours of the outer margin, the apical line and the apex indicate the presence of Mn-enriched, diagenetic calcite (e.g. Richter et al. ). Dark growth rings (laminae obscurae, sensu Müller-Stoll ), probably originally containing higher concentrations of organic material (e.g. Saelen ), are very prominent along the apical line (Fig. B). These growth rings increase in thickness closer to the apex, indicating an originally high organic content of the apex. The CL-images show, that the studied apex has been heavily altered, probably because of its originally high organic content (Fig. C). Microfractures filled with diagenetic, yellow–orange luminescent calcite, which have also been observed in the sampled rostrum portions, are here regarded to have had only a minor effect on the sample composition. They could, however, be the reason for at least part of the variation found in the isotope and trace element results.
High Fe (>200–250 ppm) and Mn (>100 ppm) as well as low Sr (<800 ppm) concentrations are proposed as indicators of diagenetic alteration in belemnite rostra (e.g. Veizer, ; Price et al. ; Rosales et al., ; Wierzbowski et al. ). All samples show low Mn contents (maximum 10.1 ppm), the ‘speiballen’-specimens have similar stable isotope values, with one sample having a very high Fe content (257 ppm). Because of this, the Fe content is considered not to be decisive for alteration in this study. Sample no. 24 is disregarded in the further considerations, because of its low Sr content of 494 ppm. Sample no. 26 is also neglected, because of a high Mg value of 4238 ppm. Both samples show aberrant low δ18O values, which is usually indicative of meteoric and burial diagenesis (e.g. Sharp ). This indicates that, if the other samples show slight effects from diagenesis, the original δ18O signals would be more positive than recorded.
Recent species of Sepia exhibit a strong metabolic influence on the δ13C values recorded from their internal skeletons (Rexfort & Mutterlose, ). A considerable variation of δ13C values has also been recorded from individual belemnite rostra (Dutton et al. ; Wierzbowski & Joachimski ). An in-depth interpretation of the acquired δ13Cbel values is therefore not intended, the metabolic effects on the incorporation of carbon in coleoid endoskeletons first need to be further clarified.
In some studies, belemnite δ18O values have been recorded giving palaeotemperatures lower than palaeotemperatures from co-occurring benthic fossils (e.g. Voigt et al. ). This suggests biofractionation during the precipitation of the belemnite rostrum calcite. While sepiids precipitate their aragonitic endoskeleton in oxygen isotope equilibrium with ambient waters (e.g. Rexfort & Mutterlose ), it is unclear, if the same applies for the calcitic belemnite rostrum. Such considerations, the unknown δ18Osw composition and the preservation cast doubt on calculated absolute palaeotemperature estimates from belemnite rostra.


There is no correlation between δ18Obel and Mg/Ca in the current data set (Fig. ), indicating that temperature or salinity did not influence the Mg/Ca composition of the studied rostra. These interpretations exclude a further consideration of the Mg/Ca signal in terms of environmental information. The acquired data are in accordance with other studies showing higher incorporation of Mg in species of Hibolithes in comparison to other belemnite genera (e.g. McArthur et al.,b; Wierzbowski & Rogov ; Li et al. ; Wierzbowski et al. ). This suggests that metabolic effects on Mg incorporation largely obscured temperature effects on Mg/Ca in species of the genus Hibolithes.
Fig. 4. Scatter plot of δ18O and Mg/Ca data. There is no correlation between the two parameters in the data set. Rectangles represent samples from the Egesheim quarry and crosses samples from the Nusplingen quarry.

Belemnite habitat

Crucial for the interpretation of the stable isotope data is understanding whether the belemnites were autochthonous residents of the lagoon, or whether they have been transported into the lagoon by storms or currents. The H. semisulcatus specimens represent, together with ammonites, by far the most abundant macrofossils of the NP. Rostra and arm hooks (onychites) of all ontogenetic stages occur frequently in the NP deposits. An occurrence of larger numbers of juvenile belemnites with a rostrum length of ~10 mm is considered to be exceptional, since most belemnite bearing strata so far mainly supplied adult specimens (Rexfort & Mutterlose ). Some of the belemnite fossils of the NP are clearly victims of predation in the water column above their place of deposition (Schweigert ; Klug et al. ). Belemnites are almost exclusively found in the Plattenkalk beds, floating in the sediment. A transport by storms or currents into the Nusplingen lagoon can therefore be excluded. The rostra are often embedded vertically or subvertically together with the phragmocone. In other specimens the alveolus part of the rostrum is crushed, but the fragments are still embedded close to the rostrum. This indicates that the rostra were crushed by predators close to their place of deposition (Schweigert ). These taphonomic observations suggest that the belemnites inhabited the Nusplingen lagoon during their entire life, where they were devoured and embedded.
This is in contrast to the situation in Solnhofen, where belemnite rostra are extremely rare and are thought to be the remains of dead, floating specimens that have been washed into the Solnhofen lagoon (Schweigert & Dietl ). An autochthonous nature of the belemnites is therefore likely for Nusplingen. Considering the inner shelf position of the Nusplingen lagoon, fairly isolated from the landmasses to the north (Swinburne & Hemleben ; Dietl & Schweigert ), the abundance of belemnites is not surprising. Coccoliths and radiolarians are common in the NP and indicate a pelagic character of the Nusplingen flora and fauna (Bantel et al. ). The stable isotope data also support the autochthonous nature of the belemnites. The δ13Cbel and δ18Obel values show only a small variation for individual Plattenkalk beds; the ‘speiballen’-specimens in particular provided nearly identical stable isotope signatures (Table , Fig. ). These latter findings support the interpretation that the belemnite calcite reflects primarily environmental conditions. It is unlikely that these signals are a result of diagenesis, as the sampled rostrum portions are well-preserved, containing only slight amounts of diagenetic calcite in microfractures (Fig. ). It is of interest, that an intra-rostrum variation of ~0.75‰ in δ18Obel was found in a Bathonian H. semisulcatus (referred to as H. hastatus) by Wierzbowski & Joachimski. Two values from the NP (no. 8, 17 in Table ) show ~1‰ more positive δ18Obel values of 0.00 and +0.07 ‰ than the other rostra of their respective Plattenkalk bed. It seems possible that a variation of 0.75–1.00‰ in δ18Obel reflects the variation observed in the calcite of this belemnite species. Alternatively, the two rostra could be from belemnites, which migrated into the lagoon. This in turn indicates that by far the majority of the measured belemnite rostra are from autochthonous specimens.
Palaeotemperature estimates calculated from the data set, using the formula of Anderson & Arthur with a δ18Osw of −1‰ for a polar-ice free world (Shackleton & Kennett ), range from 12 to 16 °C (mean: 15 °C). Low palaeotemperatures reconstructed from belemnite calcite have also been found in various other studies. They were attributed to a deep-dwelling habitat of belemnites (e.g. Mutterlose et al., ) or a nektobenthic lifestyle (e.g. Wierzbowski & Joachimski ). A deep-dwelling, demersal lifestyle is a problematic assumption for the Nusplingen belemnites. The Nusplingen lagoon was shallow, with anoxic conditions prevailing in the sediment and temporarily also above the sediment/bottom water interface (Bantel et al. ). Fast-swimming cephalopods using jet-propulsion, are known to require high oxygen concentrations for their mode of locomotion (e.g. Seibel et al. ). If a fast-swimming, predatory lifestyle is accepted for the belemnites from Nusplingen, they were probably restricted to the oxygen rich layers higher in the water-column. This mode of life is consistent with the morphology of H. semisulcatus, as reconstructed by Klug et al. Voigt et al. and Alberti et al. explain palaeotemperatures of belemnites colder than those of the co-occurring benthic fauna by migration. Spawning induced migrations of adults to shallower waters can, however, be neglected for Nusplingen, since both juveniles and adults are common in the Plattenkalk. The Nusplingen H. semisulcatus belemnites therefore lived in the water column of the lagoon together with abundant ammonites. Vampyromorph coleoids (close relatives of the modern deep-sea cephalopod Vampyroteuthis infernalis) only occur as adults in the NP (Schweigert & Dietl ). Juvenile specimens of this group were presumably restricted to environments outside the lagoon, probably because of a nektobenthic lifestyle. The cephalopod fauna of the NP is thus composed of autochthonous (ammonites, belemnites) and allochthonous (vampyromorphs) elements (Fig. ).
Fig. 5. Sketch of the Nusplingen lagoon environment. The formation of anoxic sediments and temporarily anoxic bottom waters limited nektonic organisms to the water column. Hibolithes semisulcatus belemnites and different species of ammonites are abundant as fossils in the Nusplingen Plattenkalk, indicating a typical pelagic cephalopod fauna, with limited faunal exchange between the lagoon and the open ocean. Curiously, vampyromorph coleoids are only known from adult specimens (Schweigert & Dietl ).

Climate and δ18Osw-distribution

The new δ18Obel values are similar to those described from the Kimmeridgian-Tithonian ‘ammonitico rosso’ facies of Mallorca, having had a slightly more southward latitudinal position on the northern Tethys shelf between 25–30°N (Price & Sellwood ). The Nusplingen data are on the same range as Oxfordian to lowermost Kimmeridgian values recorded from mainland Spain (Benito & Reolid ), as well as Oxfordian to Kimmeridgian belemnite data from the Swabian Alb and, to a lesser degree, those from Poland (Wierzbowski ) (Table ).
Table 2. Comparison of published Late Jurassic belemnite stable isotope data from the European Tethys Realm. The data from the western part are all similar, the Polish data are more positive. Mean δ18Obel values are given in parentheses.
LocalityPalaeolatitudeAgeδ18Obel [‰]Reference
Nusplingen, Swabian Alb~30°NLate Kimmeridgian−1.07 to +0.07 (−0.68)This study
Plettenberg, Swabian Alb~27.5°NLate Oxfordian−0.86 to −0.02 (−0.45)Wierzbowski
Prebetic Cordillera (southeastern Spain)20–30°NOxfordian – late Kimmeridgian−0.96 to +0.19 (−0.38)Benito & Reolid
Mallorca25–30°NLate Kimmeridgian – early Tithonian−1.00 to +0.04 (−0.48)Price & Sellwood
Kujawy area, Poland30–35°NEarly Middle Oxfordian−0.40 to +0.81 (+0.21)Wierzbowski
Western Carpathians, Poland30–35°N(latest Callovian?) Oxfordian – early Kimmeridgian−0.79 to +0.43 (−0.10)Wierzbowski
The Nusplingen lagoon (~30°N palaeolatitude) and other Plattenkalk lagerstätten of the Late Jurassic were situated in a winter-wet arid climate belt, comparable to the recent Mediterranean climatic situation, along the northern Tethys (e.g. Oost & De Boer ; Rees et al. ; Sellwood & Valdes ; Uhl et al. ). Sporomorph evidence from the southern North Sea indicates a long lasting phase of arid and warm climate in this area during the Kimmeridgian-Tithonian, probably triggered by the opening of the North Atlantic seaway (Abbink et al. ). General circulation models suggest that the Kimmeridgian climate was warmer than today, indicating aridity along the Tethys (Sellwood et al. ; Sellwood & Valdes ). Therefore, the apparent palaeotemperature signal of the Nusplingen belemnites of 12–16 °C, is too low.
Contemporaneous belemnite oxygen stable isotope records from higher palaeolatitudes (Scotland, ~40°N; Russian Platform, 40–50°N) are similar or often lower. They indicate ‘warmer’ conditions, than the data recorded from the Tethys (e.g. Gröcke et al. ; Nunn & Price ). This discrepancy could be the result of a δ18Osw latitudinal gradient (Lécuyer et al. ). Driven by precipitation, humid air, formed in the tropics, is enriched in 16O relative to the seawater. It evaporates and exports the 16O enrichment as precipitation to the high latitudes. This effect increases during warm climatic conditions, as assumed for the Late Jurassic. Palaeotemperature estimates at ~30°N would rise by 3–5 °C (Roche et al. ). Evidence for increased δ18Osw in latitudes between ~29–35°N comes from measurements of the phosphate oxygen isotope composition of turtles from various Plattenkalk deposits (Canjuers, Chassiron, Crayssac, Solnhofen, Cerin) of late Kimmeridgian-early Tithonian age. Known fractionation factors between ambient water and biophosphate from recent turtles allowed a reconstruction of δ18Osw to values between −0.7 and −0.2‰ (Billon-Bruyat et al. ). A latitudinally controlled gradient in the δ18Osw composition on δ18Obel values gives further support to the notion that belemnites inhabited the upper part of the water column, since latitudinal changes of δ18Osw are mainly affecting the surface waters (e.g. Roche et al. ).
In addition to this latitudinal effect, regional δ18Osw variations are also controlled by locally increased evaporation rates of restricted water-bodies or increased contribution of freshwater from rivers. The good fit of the acquired δ18Obel values along the northern Tethys (Table ) suggests that conditions of the Nusplingen site were similar to those of other inner-shelf settings. The high number of pelagic belemnites and ammonites as well as coccoliths in the NP supports this interpretation. The obtained data therefore do not indicate locally enhanced evaporation rates of the surface waters in the Nusplingen lagoon, salinity stratification was thus likely not the driving factor for the exceptional preservation of the Nusplingen fossils.


The stable isotope data acquired from H. semisulcatus belemnites of the Nusplingen Plattenkalk support the interpretation, that the belemnites are autochthonous. Anoxic conditions in the sediment and temporarily in the bottom waters of the lagoon restricted the belemnites to the well oxygenated upper parts of the water column. The active predatory lifestyle of the belemnites probably required high oxygen levels; hence, their δ18Obel values should reflect the environmental conditions of the upper water layers. The observed δ18Obel values are specific for individual Plattenkalk beds, they show the same range for four coeval, regurgitated belemnite specimens. The absence of a significant diagenetic overprint on these ‘speiballen’-specimens indicates that belemnite stable oxygen isotope data reflect the environmental conditions during the lifetime of the animal. This is supported by further δ18Obel records from other localities of Oxfordian to earliest Tithonian age. They reflect a latitudinal gradient of δ18Osw distribution. The increased δ18Osw values in the mid-latitudes (~30°N) explain at least in part the apparent ‘cold’ palaeotemperatures of 12–16 °C.
Our findings indicate: (1) A comparison with δ18Obel data from other localities of similar age along the northern Tethys margin indicates stable seawater temperature during the Oxfordian to earliest Tithonian in this region. This in turn suggests an open marine, inner shelf setting for the Nusplingen Plattenkalk; (2) the belemnite rostra are autochthonous to the shallow water setting of the Nusplingen lagoon; (3) an exclusively nektobenthic life style can be excluded for H. semisulcatus, a demersal life style seems unlikely; and (4) latitudinal as well as local salinity variations constrain absolute palaeotemperature determinations from belemnite calcite.


We thank the technicians and volunteers of the Staatliches Museum für Naturkunde Stuttgart for their efforts in exploiting the Nusplingen Plattenkalk site. Support from the GeoZentrum Nordbayern for stable isotope analysis and from the technical staff of the Ruhr-Universität Bochum for the preparation of belemnites, thin sections and trace element analyses is acknowledged. Furthermore, we thank Rolf Neuser for taking the CL-images. Two anonymous reviewers improved an earlier version of the manuscript by useful comments. Financial support is acknowledged by the DFG (Mu 667/43-1).


Abbink, O., Targarona, J., Brinkhuis, H. & Visscher, H. 2001: Late Jurassic to earliest Cretaceous palaeoclimatic evolution of the southern North Sea. Global and Planetary Change 30, 231–256.
Alberti, M., Fürsich, F.T. & Pandey, D.K. 2012: The Oxfordian stable isotope record (δ18O, δ13C) of belemnites, brachiopods, and oysters from the Kachchh Basin (western India) and its potential for palaeoecologic, palaeoclimatic, and palaeogeographic reconstructions. Palaeogeography, Palaeoclimatology, Palaeoecology 344–345, 49–68.
Anderson, T.F. & Arthur, M.A. 1983: Stable isotopes of oxygen and carbon and their applications to sedimentological and paleoenvironmental problems. In Arthur, M.A., Anderson, T.F., Kaplan, I.R., Veizer, J. & Land, L.S. (eds): Stable Isotopes in Sedimentary Geology: SEPM Short Course 10, 1.1–1.151. Society of Economic Paleontologists and Mineralogists, Dallas.
Armendáriz, M., Rosales, I., Bádenas, B., Aurell, M., García-Ramos, J.C. & Piñuela, L. 2012: High-resolution chemostratigraphic records from Lower Pliensbachian belemnites: Palaeoclimatic perturbations, organic facies and water mass exchange (Asturian basin, northern Spain). Palaeogeography, Palaeoclimatology, Palaeoecology 333–334, 178–191.
Armendáriz, M., Rosales, I., Bádenas, B., Piñuela, L., Aurell, M. & Garcia-Ramos, J. 2013: An approach to estimate Lower Jurassic seawater oxygen-isotope composition using δ18O and Mg/Ca ratios of belemnite calcites (Early Pliensbachian, northern Spain), Terra Nova 25.6, 439–445.
Bandel, K. & Spaeth, C. 1988. Structural differences in the ontogeny of some belemnite rostra. In Wiedmann, J. & Kullmann, J. (eds): Cephalopods Present and Past, 247–271. SchweitbartSche, Stuttgart.
Bantel, G., Schweigert, G., Nose, M. & Schulz, H.-M. 1999: Mikrofazies, Mikro- und Nannofossilien aus dem Nusplinger Plattenkalk (Ober-Kimmeridgium, Schwäbische Alb). Stuttgarter Beiträge zur Naturkunde, Serie B 279, 55.
Barthel, K.W., Swinburne, N.H. & Conway Morris, S. 1990. Solnhofen – A Study in Mesozoic Palaeontology, 236 pp. Cambridge University Press, Cambridge.
Benito, M.I. & Reolid, M. 2012: Belemnite taphonomy (Upper Jurassic, Western Tethys) part II: fossil–diagenetic analysis including combined petrographic and geochemical techniques. Palaeogeography, Palaeoclimatology, Palaeoecology 358–360, 89–108.
Billon-Bruyat, J.-P., Lécuyer, C., Martineau, F. & Mazin, J.-M. 2005: Oxygen isotope compositions of Late Jurassic vertebrate remains from lithographic limestones of western Europe: implications for the ecology of fish, turtles, and crocodilians. Palaeogeography, Palaeoclimatology, Palaeoecology 216, 359–375.
Dietl, G. & Schweigert, G. 2004: The Nusplingen lithographic limestone – a ‘fossil lagerstaette’ of Late Kimmeridgian age from the Swabian Alb (Germany). Rivista Italiana di Paleontologia e Stratigrafia 110, 303–309.
Dietl, G., Schweigert, G., Franz, M. & Geyer, M. 1998: Profile des Nusplinger Plattenkalks (Oberjura, Ober-Kimmeridgium, Südwestdeutschland), Stuttgarter Beiträge zur Naturkunde, Serie B 265, 37 pp.
Dutton, A., Huber, B.T., Lohmann, K.C. & Zinsmeister, W.J. 2007: High-resolution stable isotope profiles of a dimitobelid belemnite: implications for paleodepth habitat and late Maastrichtian climate seasonality. Palaios 22, 642–650.
Fürsich, F.T., Werner, W., Schneider, S. & Mäuser, M. 2007: Sedimentology, taphonomy, and palaeoecology of a laminated Plattenkalk from the Kimmeridgian of the northern Franconian Alb (southern Germany). Palaeogeography, Palaeoclimatology, Palaeoecology 243, 92–117.
Gröcke, D.R., Price, G.D., Ruffell, A.H., Mutterlose, J. & Baraboshkin, E. 2003: Isotopic evidence for Late Jurassic-Early Cretaceous climate change. Palaeogeography, Palaeoclimatology, Palaeoecology 202, 97–118.
Izumida, H., Yoshimura, T., Suzuki, A., Nakashima, R., Ishimura, T., Yasuhara, M., Inamura, A., Shikazono, N. & Kawahata, H. 2011: Biological and water chemistry controls on Sr/Ca, Ba/Ca, Mg/Ca and δ18O profiles in freshwater pearl mussel Hyriopsis sp. Palaeogeography, Palaeoclimatology, Palaeoecology 309, 298–308.
Kemp, R.A. & Trueman, C.N. 2003: Rare earth elements in Solnhofen biogenic apatite: geochemical clues to the palaeoenvironment. Sedimentary Geology 155, 109–127.
Keupp, H., Koch, R., Schweigert, G. & Viohl, G. 2007: Geological history of the Southern Franconian Alb – the area of the Solnhofen Lithographic Limestone. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 245, 3–21.
Klein, R., Lohmann, K.C. & Thayer, C.W. 1996: Bivalve skeletons record sea-surface temperature and δ18O via Mg/Ca and 18O/16O ratios. Geology 24, 415–418.
Klug, C., Schweigert, G., Dietl, G. & Fuchs, D. 2005: Coleoid beaks from the Nusplingen Lithographic Limestone (Upper Kimmeridgian, SW Germany). Lethaia 38, 173–192.
Klug, C., Schweigert, G., Fuchs, D. & Dietl, G. 2009: First record of a belemnite preserved with beaks, arms and ink sac from the Nusplingen Lithographic Limestone (Kimmeridgian, SW Germany). Lethaia 43, 445–456.
Košťák, M. & Wiese, F. 2008: Lower Turonian record of belemnite Praeactinocamax from NW Siberia and its palaeogeographic significance. Acta Palaeontologica Polonica 53, 669–678.
Lécuyer, C., Picard, S., Garcia, J.P., Sheppard, S.M.F., Grandjean, P. & Dormart, G. 2003: Thermal evolution of Tethyan surface waters during the Middle-Late Jurassic: evidence from δ18O values of marine fish teeth. Paleoceanography 18, 1076.
Li, Q., McArthur, J.M. & Atkinson, T.C. 2012: Lower Jurassic belemnites as indicators of palaeo-temperature. Paleogeogeography, Paleoclimatology, Palaeoecology 315–316, 38–45.
Li, Q., McArthur, J.M., Doyle, P., Janssen, N., Leng, M.J., Müller, W. & Reboulet, S. 2013: Evaluating Mg/Ca in belemnite calcite as a palaeo-proxy. Palaeogeography, Palaeoclimatology, Palaeoecology 388, 98–108.
McArthur, J., Mutterlose, J., Price, G., Rawson, P., Ruffell, A. & Thirlwall, M. 2004: Belemnites of Valanginian, Hauterivian and Barremian age: Sr-isotope stratigraphy, composition (87Sr/86Sr, δ13C, δ18O, Na, Sr, Mg), and palaeo-oceanography. Palaeogeography, Palaeoclimatology, Palaeoecology 202, 253–272.
McArthur, J.M., Janssen, N.M.M., Reboulet, S., Leng, M.J., Thirlwall, M.F. & van den Schootbrugge, B. 2007a: Palaeotemperatures, polar ice-volume, and isotope stratigraphy (Mg/Ca, δ18O, δ13C, 78Sr/86Sr): the Early Cretaceous (Berriasian, Valanginian, Hauterivian). Palaeogeogeography, Palaeoclimatology, Palaeoecology 248, 391–430.
McArthur, J.M., Janssen, N.M.M., Reboulet, S., Leng, M.J., Thirlwall, M.F. & van de Schootbrugge, B. 2007b: Testing palaeo-environmental proxies in Jurassic belemnites: Mg/Ca, Sr/Ca, Na/Ca, δ18O and δ13C. Palaeogeography, Palaeoclimatology, Palaeoecology 252, 464–480.
Monks, N., Hardwick, J.D. & Gale, A.S. 1996: The function of the belemnite guard. Paläontologische Zeitschrift 70, 425–431.
de Montfort, D. 1808: Conchyliology systématique et classification méthodique des coquilles 1, 495 pp. Schoell, Paris.
Müller-Stoll, H. 1936: Beiträge zur Anatomie der Belemnoidea. Nova Acta Leopoldina, N. S. 4, 159–226.
Münster, G. Graf zu., 1828: Über die Versteinerungen von Solnhofen. In Keferstein C. (ed.): Teutschland, Geognostisch-Geologisch Dargestellt und mit Charten und Durchschnittszeichnungen Erläutert 5, 578–581. Verlag des Landes-Industrie-Comptoirs, Weimar.
Münster, G. Graf zu 1830: Bemerkungen zur näheren Kenntnis der Belemniten, 18 pp. Birner, Bayreuth.
Mutterlose, J., Malkoc, M., Schouten, S., Sinninghe Damsté, J.S. & Forster, A. 2010: TEX86 and stable δ18O paleothermometry of early Cretaceous sediments: implications for belemnite ecology and paleotemperature proxy application. Earth and Planetary Science Letters 298, 286–298.
Mutterlose, J., Malkoc, M., Schouten, S. & Sinninghe Damsté, J.S. 2012: Reconstruction of vertical temperature gradients in past oceans – proxy data from the Hauterivian-early Barremian (Early Cretaceous) of the Boreal Realm. Paleogeogeography, Paleoclimatology, Palaeoecology 363–364, 135–143.
Nunn, E.V. & Price, G.D. 2010: Late Jurassic (Kimmeridgian-Tithonian) stable isotopes (δ18O, δ13C) and Mg/Ca ratios: new paleoclimate data from Helmsdale, northeast Scotland. Paleogeogeography, Paleoclimatology, Palaeoecology 292, 325–335.
Oost, A.P. & De Boer, P.L. 1994: Tectonic and climatic setting of lithographic limestone basins. Geobios 27, 321–330.
Pearson, P.N., Ditchfield, P.W., Singano, J., Harcourt-Brown, K.G., Nicholas, C.J., Olsson, R.K., Shackleton, N.J. & Hall, M.A. 2001: Warm tropical sea surface temperatures in the Late Cretaceous and Eocene epochs. Nature 413, 481–487.
Podlaha, O., Mutterlose, J. & Veizer, J. 1998: Preservation of and in belemnite rostra from the Jurassic/Early Cretaceous successions. American Journal of Science 298, 324–347.
Price, G.D. & Sellwood, B.W. 1994: Palaeotemperatures indicated by Upper Jurassic (Kimmeridgian-Tithonian) fossils from Mallorca determined by oxygen isotope composition. Palaeogeography, Palaeoclimatology, Palaeoecology 110, 1–10.
Price, G.D., Ruffell, A.H., Jones, C.E., Kalin, R.M. & Mutterlose, J. 2000: Isotopic evidence for temperature variation during the Early Cretaceous (late Ryazanian – mid Hauterivian). Journal of the Geological Society, London 157, 335–343.
Purton, L.M., Shields, G.A., Brasier, M.D. & Grime, G.W. 1999: Metabolism controls Sr/Ca ratios in fossil aragonitic mollusks. Geology 27, 1083–1086.
Rees, P.M., Ziegler, A.M. & Valdes, P.J. 2000: Jurassic phytogeography and climates: new data and model comparisons. In Huber, B.T., McLeod, K.G. & Wing, S.L. (eds): Warm Climates in Earth History, 297–318. Cambridge University Press, Cambridge.
Rexfort, A. & Mutterlose, J. 2006: Stable isotope records from Sepia officinalis—a key to understanding the ecology of belemnites? Earth and Planetary Science Letters 247, 212–221.
Rexfort, A. & Mutterlose, J. 2009: The role of biogeography and ecology on the isotope signature of cuttlefishes (Cephalopoda, Sepiidae) and the impact on belemnite studies. Palaeogeography, Palaeoclimatology, Palaeoecology 284, 153–163.
Richter, D.K., Götte, T., Götze, J. & Neuser, R.D. 2003: Progress in application of cathodoluminescence (CL) in sedimentary petrology. Mineralogy and Petrology 79, 127–166.
Richter, D.K., Neuser, R.D., Schreuer, J., Gies, H. & Immenhauser, A. 2011: Radiaxial-fibrous calcites: a new look at an old problem. Sedimentary Geology 239, 23–36.
Riegraf, W., Janssen, N.M.M. & Schmitt-Riegraf, C. 1998: Cephalopoda dibranchiata fossiles (Coleoidea) II. In Westphal, F. (ed.): Fossilium Catalogus: I, Animalia 135, 512 pp. Backhuys Publishers, Leiden.
Roche, D.M., Donnadieu, Y., Pucéat, E. & Paillard, D. 2006: Effect of changes in δ18O content of the surface ocean on estimated sea surface temperatures in past warm climate. Paleoceanography 21, PA2023.
Rosales, I., Quesada, S. & Robles, S. 2001: Primary and diagenetic isotopic signals in fossils and hemipelagic carbonates: the Lower Jurassic of northern Spain. Sedimentology 48, 1149–1169.
Rosales, I., Robles, S. & Quesada, S. 2004: Elemental and oxygen isotope composition of Early Jurassic belemnites: salinity vs. temperature signals. Journal of Sedimentary Research 74, 342–354.
Saelen, G. 1989: Diagenesis and construction of the belemnite rostrum. Palaeontology 32, 765–798.
Schlegelmilch, R. 1998. Die Belemniten des Süddeutschen Jura. Ein Bestimmungsbuch für Geowissenschaftler und Fossiliensammler, 151 pp. Gustav Fischer Verlag, Stuttgart.
Schwark, L., Vliex, M. & Schaeffer, P. 1998: Geochemical characterization of Malm Zeta laminated carbonates from the Franconian Alb, SW-Germany (II). Organic Geochemistry 29, 1921–1952.
Schweigert, G. 1999: Erhaltung und Einbettung von Belemniten im Nusplinger Plattenkalk (Ober-Kimmeridgium, Beckeri-Zone, Schwäbische Alb. Stuttgarter Beiträge zur Naturkunde, Series B 273, 35.
Schweigert, G. 2007: Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen 245, 117–125.
Schweigert, G. & Dietl, G. 2010: The Coleoidea of the Upper Kimmeridgian Nusplingen Lithographic Limestone – diversity, preservation and palaeoecology. In Fuchs, D. (ed.): Proceedings of the 3rd International Symosium, Coleoid Cephalopods Through Time, Ferrantia, volume 59, 43–45.
Seibel, B.A., Thuesen, E.V., Childress, J.J. & Gorodezky, L.A. 1997: Decline in pelagic cephalopod metabolism with habitat depth reflects differences in locomotory efficiency. Biological Bulletin 192, 262–278.
Sellwood, B.W. & Valdes, P.J. 2008: Jurassic climates. Proceedings of the Geologists’ Association 119, 5–17.
Sellwood, B.W., Valdes, P.J. & Price, G.D. 2000: Geological evaluation of multiple general circulation model simulations of Late Jurassic palaeoclimate. Palaeogeography, Palaeoclimatology, Palaeoecology 156, 147–160.
Shackleton, N.J. & Kennett, J.P. 1975: Paleotemperature history of the Cenozoic and initiation of Antarctic glaciation: oxygen and carbon isotope analyses in DSDP sites 277, 279 and 281. Initial Reports of the Deep Sea Drilling Projects 29, 743–756.
Sharp, Z. 2007: Principles of Stable Isotope Geochemistry, 344 pp. Pearson Prentice Hall, Upper Saddle River.
Spaeth, C. 1971: Aragonitische und calcitische Primärstrukturen im Schalenbau eines Belemniten aus der englischen Unterkreide. Paläontologische Zeitschrift 45, 33–40.
Spaeth, C. 1975: Zur Frage der Schwimmverhältnisse bei Belemniten in Abhängigkeit vom Primärgefüge der Hartteile. Paläontologische Zeitschrift 49, 321–331.
Steuber, T. & Rauch, M. 2005: Evolution of the Mg/Ca ratio of Cretaceous seawater: implications from the composition of biological low-Mg calcite. Marine Geology 217, 199–213.
Swinburne, N.H. & Hemleben, C. 1994: The plattenkalk facies: a deposit of several environments. Geobios 27, 313–320.
Thies, D. & Hauff, R.B. 2012: A Speiballen from the Lower Jurassic Posidonia Shale of South Germany. Neues Jahrbuch für Geologie und Paläontologie 267, 117–124.
Uhl, D., Jasper, A. & Schweigert, G. 2012: Charcoal in the Late Jurassic (Kimmeridgian) of Western and Central Europe—palaeoclimatic and palaeoenvironmental significance. Palaeobiodiversity and Palaeoenvironments 92, 329–341.
Urey, H.C., Lowenstam, H.A., Epstein, S. & McKinney, C.R. 1951: Measurement of paleotemperature and temperature of the Upper Cretaceous of England, Denmark and the southern United States. Bulletins of the Geological Society of America 62, 399–416.
Veizer, J. 1974: Chemical diagenesis of belemnite shells and possible consequences for paleotemperature determinations. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 147, 91–111.
Veizer, J. 1983: Chemical diagenesis of carbonates: Theory and trace element technique. In Arthur, M.A., Anderson, T.F., Kaplan, I.R., Veizer, J., Land L.S. (eds): Stable Isotopes in Sedimentary Geology: SEPM Short Course 10, 3-1-3-100. Society of Economic Paleontologists and Mineralogists, Dallas.
Voigt, S., Wilmsen, M., Mortimore, R.N. & Voigt, T. 2003: Cenomanian palaeotemperatures derived from the oxygen isotopic composition of brachiopods and belemnites: evaluation of Cretaceous palaeotemperature proxies. International Journal of Earth Sciences (Geologische Rundschau) 92, 285–299.
Wierzbowski, H. 2004: Carbon and oxygen isotope composition of Oxfordian-Early Kimmeridgian belemnite rostra: palaeoenvironmental implications for Late Jurassic seas. Palaeogeography, Palaeoclimatology, Palaeoecology 203, 153–168.
Wierzbowski, H. & Joachimski, M.M. 2009: Stable isotopes, elemental distribution, and growth rings of belemnopsid belemnite rostra: proxies for belemnite life habitat. Palaios 24, 377–386.
Wierzbowski, H. & Rogov, M. 2011: Reconstructing the palaeoenvironment of the Middle Russian Sea during the Middle-Late Jurassic transition using stable isotope ratios of cephalopod shells and variations in faunal assemblages. Palaeogeography, Palaeoclimatology, Palaeoecology 299, 250–264.
Wierzbowski, H., Rogov, M.A., Matyja, B.A., Kiselev, D. & Ippolitov, A. 2013: Middle-Upper Jurassic (Upper Callovian-Lower Kimmeridgian) stable isotope and elemental records of the Russian Platform: indices of oceanographic and climatic changes. Global and Planetary Change 107, 196–212.

Information & Authors


Published In

Volume 47Number 41 October 2014
Pages: 512523


Received: 10 October 2013
Accepted: 6 March 2014
Published online: 7 July 2014
Issue date: 1 October 2014



Kevin Stevens [email protected]
Institut für Geologie, Mineralogie und Geophysik, Ruhr-Universität Bochum, Universitätsstr. 150, 44801, Bochum, Germany;
Jörg Mutterlose [email protected]
Institut für Geologie, Mineralogie und Geophysik, Ruhr-Universität Bochum, Universitätsstr. 150, 44801, Bochum, Germany;
Günter Schweigert [email protected]
Staatliches Museum für Naturkunde, Rosenstein 1, 70191, Stuttgart, Germany;

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