Research paperCoccoliths from recent sediments of the central Portuguese margin: Taphonomical and ecological inferences
Introduction
Calcareous nannoplankton, predominantly represented by coccolithophores, has the best fossil record of all marine phytoplankton, except in polar regions (Ziveri et al., 2004). Their calcitic cell-covers (coccoliths sensu lato) are produced by biomineralization of tiny calcite crystals arranged with preferential orientations, and represent morphological elements which are relevant for taxonomy and phylogeny (Pienaar, 1994, Young, 1994). The preservation of coccoliths in the geological archives provides valuable information on paleoenvironmental conditions for the photic zone (McIntyre and Bé, 1967, Roth, 1994, Andruleit et al., 2000, Baumann et al., 2000, Andruleit and Rogalla, 2002, Boeckel and Baumann, 2008). As such, they can be used as indicators of paleoceanographic processes, sea surface water masses, productivity and climate change (e.g. Flores et al., 2000, Beaufort et al., 2001, Ziveri et al., 2004, Silva et al., 2008, Beaufort et al., 2011).
The correspondence between the coccolith species assemblages preserved in the seabed and the living communities thriving in the overlying photic layer is, however, complex, particularly on the continental shelf (e.g. Andruleit et al., 2000, Ziveri et al., 2000a, Ziveri et al., 2000b, Andruleit and Rogalla, 2002, Andruleit et al., 2003, Baumann et al., 2005). Complexity increases even more in heterogeneous and dynamic areas such as submarine canyons, which often act as traps of particulate matter derived from the continent and shelf, and also as preferential conduits for transport of sediment from the coastal area to the deep sea (e.g. dense-water cascading and sediment gravity flows) (Schmidt et al., 2001, Van Weering et al., 2002, Canals et al., 2006, De Stigter et al., 2007, Oliveira et al., 2007, De Stigter et al., 2011).
In modern ecosystems, submarine canyons may show increased phytoplankton density, resulting from upwelling and enhanced vertical mixing in the upper canyon reaches (e.g. Hickey, 1995, Ryan et al., 2005, Kampf, 2006, Skliris and Djenidi, 2006, Ryan et al., 2010, Mendes et al., 2011). Concentration of marine fauna in and around the canyons, related to local enhancement of primary production, has been reported in several studies (Macquart-Moulin and Patriti, 1996, Allen et al., 2001, Bosley et al., 2004, Palanques et al., 2005, David and Di-Méglio, 2012). Whether any of this enhanced biological productivity is reflected in fossil assemblages in canyon sediments is problematic, since high terrigenous sediment input and strong bottom dynamics typical of submarine canyons are expected to dilute and disturb the paleoecological signal.
Here we report the results of a study of recent coccolith assemblages obtained from 92 surface sediment samples collected from the central Portuguese margin, where two major submarine canyons are located: the Nazaré Canyon and Lisbon–Setúbal Canyon. Although information on the effect of physical processes operating in these canyons on phytoplankton ecology is scarce, there is some evidence suggesting that these canyons promote phytoplankton production near the coast. Mendes et al. (2011), investigating the distribution and composition of phytoplankton assemblages in Nazaré Canyon during an upwelling event, reported the highest Chl-a concentrations to occur in the canyon head. According to these authors, persistently high concentrations of diatoms south of the canyon reflect intensified upwelling along the southern rim of the canyon, a phenomenon persisting even during the relaxation of winds causing upwelling. In a study of late-winter coccolithophore assemblages from the Nazaré Canyon region, Guerreiro et al. (2014) hypothesized the presence of local diversity and productivity “hotspots” related to the canyon topography. While Emiliania huxleyi, Gephyrocapsa ericsonii, Coronosphaera mediterranea and Gephyrocapsa muellerae were the dominant species overall, Gephyrocapsa oceanica was relatively more abundant at the canyon head and adjacent shelf south of the canyon, whereas Syracosphaera spp. and Ophiaster spp. were consistently more abundant further offshore above the middle Nazaré Canyon. Persistently high Chl-a concentrations in the Nazaré Canyon head observed from satellite data were interpreted by the same authors as indicative of persistent high phytoplankton production.
In the present analysis of coccoliths from surface sediments of the central Portuguese margin, we aim to assess whether the ecological differentiation as observed in living coccolithophore assemblages from surface waters can still be discerned in coccolith assemblages preserved in the underlying seabed sediments, or whether this differentiation is obliterated by the dynamic sedimentary processes typical for this heterogeneous continental margin area. We use the term “(paleo)ecology” to highlight that we are dealing with sediment thanatocoenoses representing very recent geological timescales, which are expected to present reduced diagenetic effects. With our study we aim to contribute to the knowledge of this phytoplankton group, in particular its potential as paleoecological and paleoceanographic proxies in continental margin areas.
Section snippets
Central Portuguese margin
The central Portuguese margin has a relatively narrow shelf with a variable width of 20–50 km and a gradient of < 1°, at the shelf break at 160–200 m depth passes into a steep irregular slope with a gradient of 6–7° (Fig. 1). The shelf is underlain by a thick accumulation of Cenozoic detritical formations, filling structural basins formed during earlier rifting phases. The margin is dissected by a number of long submarine canyons, of which the Nazaré and Lisbon–Setúbal canyons are the most
Cruises and surface sediment sampling
Surface sediment box- and multicores used in this study were recovered during several cruises with RV Pelagia of Royal NIOZ, in November 2002, October 2003, April/May 2004, May 2005, September 2006 and March 2011 on the central Portuguese margin (cruises 64PE204, 64PE218, 64PE225, 64PE236, 64PE252 and 64PE332, respectively). Coring equipment and methodology are described in De Stigter et al., 2007, De Stigter et al., 2011. Sediment coring was concentrated on six transects crossing the central
Calcareous nannoplankton in surface sediments
A total of 15 distinct coccolith taxa were recognized using polarizing light microscopy. The complete list of the observed taxa is presented in Appendix A.
Total coccolith concentrations of species ≥ 3 μm varied between a minimum of 3 × 106 nanno/g at the shelf of Estremadura Spur and a maximum of 3 × 109 nanno/g reached at the lower slope off Cape Mondego. G. muellerae had the highest concentrations, followed by C. leptoporus, G. oceanica, the taxonomic group composed by U. sibogae, U. irregularis and
(Paleo)ecology versus taphonomy
Linking living coccolithophore communities to the corresponding coccolith assemblages preserved in seabed sediments is complex (Andruleit et al., 2000, Ziveri et al., 2000a, Ziveri et al., 2000b, Baumann et al., 2005), particularly in dynamic coastal areas and in places where the continental margin is dissected by submarine canyons promoting the interplay of continental and marine factors (e.g. Roth, 1994, Steinmetz, 1994). Sediments in such areas may be expected to contain an averaged signal
Conclusions
In sediments of the central Portuguese margin, concentrations of total coccoliths as well as of individual species generally do not reflect coccolithophore productivity in surface waters, but appear governed by taphonomical processes favoring their preservation in fine-grained hemipelagic sediments of the continental slope and fine-grained lithogenic-rich sediments of the Nazaré and Lisbon–Setúbal canyons. Neither redistribution of coccoliths by bottom currents nor dilution of coccoliths by
Acknowledgments
Data for this study were collected in the framework of the EU-funded EUROSTRATAFORM (contract EVK3-CT-2002-00079) and HERMES EU-funded projects (contract GOCE-CT-2005-511234), “Lead in Canyons” and “Pacemaker” projects funded by the Netherlands Organization for Scientific Research, and by the Portuguese Science Foundation (FCT) (FCT-PTDC/MAR/102800/2008). Multicores were collected during RV Pelagia cruises funded by the Netherlands Organisation for Scientific Research. Coccolith samples were
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