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Dendritic and/or rosetted microborings in calcareous and osteic skeletal substrates have a diverse trace fossil record, spanning most of the Phanerozoic, whereas the ichnodiversity of comparable bioerosion traces produced in modern seas is rather limited. The most prominent occurrences are known from Devonian brachiopods and from Upper Cretaceous belemnite rostra. Ichnotaxonomically, they are comprised within one of the few ichnofamilies established to date, the Dendrinidae Bromley et al., 2007. As an outcome of the present revision of this ichnofamily, the plethora of 84 ichnospecies established within 25 ichnogenera since the erection of the type ichnogenus Dendrina Quenstedt, 1849 was considerably condensed to 22 ichnospecies included in 7 ichnogenera, based on a coherent morphological categorisation and ichnotaxobasis assessment. The suite of ichnogenera now subsumed within the Dendrinidae includes Dendrina Quenstedt, 1849; Clionolithes Clarke, 1908; Calcideletrix Mägdefrau, 1937; Dictyoporus Mägdefrau, 1937; Abeliella Mägdefrau, 1937; Nododendrina Vogel et al., 1987; and Pyrodendrina Tapanila, 2008. New combinations thereby concern Dendrina dendrina (Morris, 1851) comb. nov., Clionolithes pannosus (Solle, 1938) comb. nov., C. alcicornis (Vogel et al., 1987) comb. nov., C. convexus (Hofmann, 1996) comb. nov., Calcideletrix anomala (Mägdefrau, 1937) comb. nov., C. fastigata (Radtke, 1991) comb. nov., Dictyoporus balani (Tavernier et al., 1992) comb. nov., Nododendrina europaea (Fischer, 1875) comb. nov., N. incomposita (Mägdefrau, 1937) comb. nov. and N. paleodendrica (Elias, 1957) comb. nov. Investigation of new material and a reassessment of 63 dendrinid microborings previously addressed in informal nomenclature allowed the establishment of two complementing ichnogenera, Rhopalondendrina igen. nov. and Antodendrina igen. nov., and eight new ichnospecies, comprising Pyrodendrina arctica isp. nov., P. belua isp. nov., P. villosa isp. nov., Rhopalondendrina avis igen. et isp. nov., R. acanthina igen. et isp. nov., R. contra igen. et isp. nov., R. tigris igen. et isp. nov. and Antodendrina ligula igen. et isp. nov. In densely bioeroded calcareous substrates, different dendrinids and other bioerosion traces may be found in direct contact with each other, forming composite trace fossils, but some of these associations appear rather systematic in nature and could be the work of the same tracemaker under different behavioural modes, thus forming compound trace fossils. In these cases, however, the distinction between the two concepts remains largely equivocal. Dendrinid microborings are primarily found in living and dead calcareous skeletal substrates of bivalves, brachiopods, belemnites and corals, with complementing records from six other substrate types. Facing considerable sampling artefacts, evidence for true substrate specificity or symbiotic relationships is inconclusive as yet, whereas there is direct evidence for post-mortem infestation in several cases, such as the diverse dendrinid associations in Upper Cretaceous belemnite guards. Despite a wealth of available interpretations, the actual biological identity of the dendrinids’ tracemakers remains largely speculative. The most convincing evidence has been put forward in support of foraminiferans as the producers of Nododendrina, and excavating micro-sponges producing Clionolithes and some Calcideletrix. Since most of the dendrinids are found in aphotic (palaeo-)environments, these two principal types of organotrophic tracemakers are also potential candidates for the other ichnogenera. With regards to evolutionary patterns through geologic time, strong adaptive radiations are evident from the ichnodiversity of dendrinid ichnospecies in the Early to Mid-Palaeozoic, reflecting the “Ordovician Bioerosion Revolution” (sensu Wilson & Palmer 2006) and the “Mid-Palaeozoic Precursor of the Mesozoic Marine Revolution” (sensu Signor & Brett 1984), respectively, and in the Mesozoic, coinciding with the prominent “Marine Mesozoic Revolution” (sensu Vermeij 1977). This pattern mimics that of
other micro- and macro-bioerosion trace fossils and is interpreted as a reflection of increased predation pressure and consequent infaunalisation. For extinction events, in turn, a differential effect is recorded in that the first four of the “Big Five” mass extinctions appear not to have had any noticeable effect on dendrinid ichnodiversity, whereas the end-Cretaceous mass-extinction resulted in a 77% drop following the Cretaceous peak ichnodiversity of 13 dendrinid ichnospecies.
Acesta excavata (Fabricius, 1779) is a slow growing bivalve from the Limidae family and is often found associated with cold-water coral reefs along the European continental margin. Here we present the compositional variability of frequently used proxy elemental ratios (Mg/ Ca, Sr/Ca, Na/Ca) measured by laser-ablation mass spectrometry (LA-ICP-MS) and com- pare it to in-situ recorded instrumental seawater parameters such as temperature and salin- ity. Shell Mg/Ca measured in the fibrous calcitic shell section was overall not correlated with seawater temperature or salinity; however, some samples show significant correlations with temperature with a sensitivity that was found to be unusually high in comparison to other marine organisms. Mg/Ca and Sr/Ca measured in the fibrous calcitic shell section display significant negative correlations with the linear extension rate of the shell, which indicates strong vital effects in these bivalves. Multiple linear regression analysis indicates that up to 79% of elemental variability is explicable with temperature and salinity as independent pre- dictor values. Yet, the overall results clearly show that the application of Element/Ca (E/Ca) ratios in these bivalves to reconstruct past changes in temperature and salinity is likely to be complicated due to strong vital effects and the effects of organic material embedded in the shell. Therefore, we suggest to apply additional techniques, such as clumped isotopes, in order to exactly determine and quantify the underlying vital effects and possibly account for these. We found differences in the chemical composition between the two calcitic shell lay- ers that are possibly explainable through differences of the crystal morphology. Sr/Ca ratios also appear to be partly controlled by the amount of magnesium, because the small magne- sium ions bend the crystal lattice which increases the space for strontium incorporation. Oxi- dative cleaning with H2O2 did not significantly change the Mg/Ca and Sr/Ca composition of the shell. Na/Ca ratios decreased after the oxidative cleaning, which is most likely a leaching effect and not caused by the removal of organic matter.
The Arctic Svalbard Archipelago hosts the world’s northernmost cold-water ‘carbonate factories’ thriving here despite of presumably unfavourable environmental conditions and extreme seasonality. Two contrasting sites of intense biogenic carbonate production, the rhodolith beds in Mosselbukta in the north of the archipelago and the barnacle-mollusc dominated carbonate sediments accumulating in the strong hydrodynamic regime of the Bjørnøy-Banken south of Spitsbergen, were the targets of the RV Maria S. Merian cruise 55 in June 2016. By integrating data from physical oceanography, marine biology, and marine geology, the present contribution characterises the environmental setting and biosedimentary dynamics of these two polar carbonate factories. Repetitive CTD profiling in concert with autonomous temperature/salinity loggers on a long-term settlement platform identified spatiotemporal patterns in the involved Atlantic and Polar water masses, whereas short-term deployments of a lander revealed fluctuations of environmental variables in the rhodolith beds in Mosselbukta and at same depth (46 m) at Bjørnøy-Banken. At both sites, dissolved inorganic nutrients in the water column were found depleted (except for elevated ammonium concentrations) and show an overall increase in concentration and N:P ratios toward deeper waters. This indicates that a recycling system was fuelling primary production after the phytoplankton spring bloom at the time of sampling in June 2016. Accordingly, oxygen levels were found elevated and carbon dioxide concentrations (pCO2) markedly reduced, on average only half the expected equilibrium values. Backed up by seawater stable carbon and oxygen isotope signatures, this is interpreted as an effect of limited air-sea gas exchange during seasonal ice cover in combination with a boost in community photosynthesis during the spring phytoplankton bloom. The observed trends are enhanced by the onset of rhodophyte photosynthesis in the rhodolith beds during the polar day upon retreat of sea-ice. Potential adverse effects of ocean acidification on the local calcifier community are thus predicted to be seasonally buffered by the marked drop in pCO2 during the phase of sea-ice cover and spring phyto-plankton bloom, but this effect will diminish should the seasonal sea-ice formation continue to decline. Among the 25 macrobenthos taxa identified from images captured by the lander’s camera system, all but three species were calcifiers contributing to the carbonate production. Biodiversity was found to be much higher in Mosselbukta (21 taxa) compared to Bjørnøy-Banken (8 taxa), which is considered as a result of enhanced habitat diversity provided in the rhodolith beds by the bioengineering crustose alga Lithothamnion glaciale. Filter-feeding activity of selected key species did reveal group-specific but no common activity patterns. Biotic disturbance of the filtering activity was common, in contrast to abiotic factors, with hermit crabs representing the primary trigger. Motion tracking of rhodoliths revealed a high frequency of dislocation, triggered not by abiotic factors but by the activity of benthic invertebrates, in particular echinoids ploughing below or moving over the rhodoliths. The echinoid Strongylocentrotus sp. is the most abundant component of the associated fauna, thereby considerably contributing both to carbonate production and to grazing bioerosion. Together, these results portray a high degree of seasonal as well as short-term dynamics in environmental conditions that despite many similarities support distinctly different communities and biodiversity patterns in the calcifying macrobenthos at the two studied polar carbonate factories.
Hyrrokkin sarcophaga is a parasitic foraminifera that is commonly found in cold-water coral reefs where it infests the file clam Acesta excavata and the scleractinian coral Desmophyllum pertusum (formerly known as Lophelia pertusa). Here, we present measurements of the trace element and isotopic composition of these parasitic foraminifera, analyzed by inductively coupled optical emission spectrometry (ICP-OES), electron probe microanalysis (EPMA) and mass spectrometry (gas-source MS and inductively-coupled-plasma MS). Our results reveal that the geochemical signature of H. sarcophaga depends on the host organism it infests. Sr / Ca ratios are 1.1 mmol mol−1 higher in H. sarcophaga that infest D. pertusum, which could be an indication that dissolved host carbonate material is utilized in shell calcification, given that the aragonite of D. pertusum has a naturally higher Sr concentration compared to the calcite of A. excavata. Similarly, we measure 3.1 ‰ lower δ13C and 0.25 ‰ lower δ18O values in H. sarcophaga that lived on D. pertusum, which might be caused by the direct uptake of the host's carbonate material with a more negative isotopic composition or different pH regimes in these foraminifera (pH can exert a control on the extent of CO2 hydration/hydroxylation) due to the uptake of body fluids of the host. We also observe higher Mn / Ca ratios in foraminifera that lived on A. excavata but did not penetrate the host shell compared to specimen that penetrated the shell, which could be interpreted as a change in food source, changes in the calcification rate, Rayleigh fractionation or changing oxygen conditions. While our measurements provide an interesting insight into the calcification process of this unusual foraminifera, these data also indicate that the geochemistry of this parasitic foraminifera is unlikely to be a reliable indicator of paleoenvironmental conditions using Sr / Ca, Mn / Ca, δ18O or δ13C unless the host organism is known and its geochemical composition can be accounted for.
Hyrrokkin sarcophaga is a parasitic foraminifer that is commonly found in cold-water coral reefs where it infests the file clam Acesta excavata and the scleractinian coral Lophelia pertusa. Here, we present measurements of the elemental and isotopic composition of this parasitic foraminifer for the first time, analyzed by inductively coupled optical emission spectrometry (ICP-OES), electron probe micro analysis (EPMA) and mass spectrometry (MS). Our results reveal that the geochemical signature of H. sarcophaga depends on the host organism it infests. Sr/Ca ratios are 1.1 mmol mol-1 higher in H. sarcophaga that infest L. pertusa, which could be an indication that dissolved host carbonate material is utilised in shell calcification, given that the aragonite of L. pertusa has a naturally higher Sr concentration compared to the calcite of A. excavata.Similarly, we measure 3.1 ‰ lower δ13C and 0.25 ‰ lower δ18O values in H. sarcophaga that lived on20 L. pertusa, which might be caused by the direct uptake of the host’s carbonate material with a more negative isotopic composition or different pH regimes in these foraminifera (pH can exert a control on the extent of CO2 hydration/hydroxylation) due to the uptake of body fluids of the host. We also observe higher Mn/Ca ratios in foraminifers that lived on A. excavata but did not penetrate the host shell compared to specimen that penetrated the shell, which could be interpreted as a change in food source, changes in the calcification rate, Rayleigh fractionation or changing oxygen conditions. While our measurements provide an interesting insight into the calcification process of this unusual foraminifer, these data also indicate that the geochemistry of this parasitic foraminifer is unlikely to be a reliable indicator of paleoenvironmental conditions using Sr/Ca, Mn/Ca, δ18O or δ13C unless the host organism is known and its geochemical composition can be accounted for.