seasonal extent of sea-ice cover, and primary productivity
in Quaternary sediments (e.g., de Vernal and Marret, 2007;
de Vernal et al., 2007; Bonnet et al., 2012).
Paleoecology of extinct dinoflagellate cysts
Dinocysts are increasingly used for paleoenvironmental
reconstructions (Pross and Brinkhuis, 2005) but the definition of ecological preferences of extinct species remains
a challenge. Various combinations of actuopaleontological, empirical, and statistical approaches
including comparison with the morphology and ecology
of co-occurring extant genera and species, the identification of latitudinal and onshore-offshore gradients from
paleobiogeographic data, the interpretation of statistical
analyses (e.g., correspondence analysis) on dinocyst distribution in relation to independent paleoenvironmental
information, and the relation between dinocyst assemblages and geochemical proxies for water mass properties
may yield qualitative and quantitative ecological information on, e.g., temperature, salinity, onshore-offshore gradients, bottom water oxygenation, and productivity (e.g.,
Versteegh and Zonneveld, 1994; Pross and Brinkhuis,
2005; De Schepper et al., 2011; Bijl et al., 2011; Masure
et al., 2013; Schreck and Matthiessen, 2013). These ecological parameters are qualitatively known for some
extinct species, groups of taxa or complexes of genera
(e.g., Pross and Brinkhuis, 2005), and the correlation of
species abundance to geochemical proxies (e.g., Mg/Ca
temperatures on co-occurring planktonic foraminifera,
De Schepper et al., 2011) is promising for providing
quantitative data.
Biostratigraphy
Since the middle of the twentieth century, palynostratigraphy has emerged as a routine tool in both hydrocarbon exploration and academic research in Mesozoic and
Cenozoic sediments, and numerous biostratigraphic zonations have been erected for Triassic to Neogene sediments
(Stover et al., 1996). Dinocysts typically exhibit high
abundances in neritic settings; thus, the derived stratigraphic information is complementary to that obtained
from typically more offshore groups such as planktonic
foraminifers, coccolithophores, and radiolarians (Pross
and Brinkhuis, 2005). Significant progress has been made
during the past four decades of scientific ocean drilling
(DSDP, ODP, IODP) by assessing stratigraphic ranges
against independent chronostratigraphic information.
Recently, the focus is slowly moving from defining new
zonations toward calibrating bioevents to the geological
time scale on both regional (De Schepper and Head,
2008; Fensome et al., 2008a; Schreck et al., 2012) and
global scales (Williams et al., 2004). This avoids the inherent problem of zonations that zones named after the same
species may have different age ranges. However, few studies illustrate that dinocyst bioevents are rarely synchronous worldwide, and low-, mid-, and high-latitude
bioevents should be distinguished to account for the
observed latitudinal control on species ranges. Nonetheless, some bioevents are useful on regional and/or
supraregional scale and enable stratigraphic correlations
between different basins in the mid- and high latitudes
(Schreck et al., 2012).
Conclusions
In recent years, biological and paleontological studies
have provided a wealth of new information relevant for
the application of recent and fossil dinocysts in marine
geosciences. However, our knowledge of their ecology is
still biased to coastal and shelf environments, and many
open ocean regions such as the Pacific yet remain largely
unexplored. The phylogenetic relationship to the motile
form of many extant species is unknown, and molecular
genetic studies will be particularly useful to address the
long-standing question whether a single dinoflagellate
species forms different cyst species. In the geological
record, dinocysts are of eminent importance for
paleoenvironmental interpretation and biostratigraphy in
high latitudes where preservation of calcareous and
biosiliceous microfossils is poor, but data from lower latitudes are required to inevitably improve independent age
calibration of bioevents. This will also provide new data
on the temporal and spatial distribution of fossil dinocysts,
which together with calibration of species abundances to
geochemical proxies for, e.g., surface temperature, will
lead to a better understanding of cyst paleoecology.
Bibliography
Bijl, P. K., Pross, J., Warnaar, J., Stickley, C. E., Huber, M.,
Guerstein, R., Houben, A. J. P., Sluijs, A., Visscher, H., and
Brinkhuis, H., 2011. Environmental forcings of Paleogene
Southern Ocean dinoflagellate biogeography. Paleoceanography, 26, PA1202, doi:10.1029/2009PA001905.
Bonnet, S., de Vernal, A., Gersonde, R., and Lembke-Jene, L., 2012.
Modern distribution of dinocysts from the North Pacific Ocean
(37–64
N, 144
E–148
W) in relation to hydrographic conditions, sea-ice and productivity. Marine Micropaleontology,
84–85, 87–113.
Chen, B., Irwin, A. J., and Finkel, Z. V., 2011. Biogeographic distribution of diversity and size-structure of organic-walled dinoflagellate cysts. Marine Ecology Progress Series, 425, 35–45.
Dale, B., 1983. Dinoflagellate resting cysts: “benthic plankton”. In
Fryxell, G. A. (ed.), Survival Strategies of the Algae. Cambridge:
Cambridge University Press, pp. 69–144.
Dale, B., 1996. Dinoflagellate cyst ecology: modelling and geological applications. In Jansonius, J., and McGregor, D. C. (eds.),
Palynology: Principles and Applications. Dallas: American
Association of Stratigraphic Palynologists Foundation,
pp. 1249–1275.
Dale, B., 2009. Eutrophication signals in the sedimentary record of
dinoflagellate cysts in coastal waters. Journal of Sea Research,
61, 103–113.
De Schepper, S., and Head, M. J., 2008. Age calibration of dinoflagellate cyst and acritarch events in the Pliocene–Pleistocene of
the eastern North Atlantic (DSDP Hole 610A). Stratigraphy, 5,
137–161.
De Schepper, S., Fischer, E. I., Groeneveld, J., Head, M. J., and
Matthiessen, J., 2011. Deciphering the palaeoecology of Late
Pliocene and Early Pleistocene dinoflagellate cysts.
DINOFLAGELLATES
191
in Quaternary sediments (e.g., de Vernal and Marret, 2007;
de Vernal et al., 2007; Bonnet et al., 2012).
Paleoecology of extinct dinoflagellate cysts
Dinocysts are increasingly used for paleoenvironmental
reconstructions (Pross and Brinkhuis, 2005) but the definition of ecological preferences of extinct species remains
a challenge. Various combinations of actuopaleontological, empirical, and statistical approaches
including comparison with the morphology and ecology
of co-occurring extant genera and species, the identification of latitudinal and onshore-offshore gradients from
paleobiogeographic data, the interpretation of statistical
analyses (e.g., correspondence analysis) on dinocyst distribution in relation to independent paleoenvironmental
information, and the relation between dinocyst assemblages and geochemical proxies for water mass properties
may yield qualitative and quantitative ecological information on, e.g., temperature, salinity, onshore-offshore gradients, bottom water oxygenation, and productivity (e.g.,
Versteegh and Zonneveld, 1994; Pross and Brinkhuis,
2005; De Schepper et al., 2011; Bijl et al., 2011; Masure
et al., 2013; Schreck and Matthiessen, 2013). These ecological parameters are qualitatively known for some
extinct species, groups of taxa or complexes of genera
(e.g., Pross and Brinkhuis, 2005), and the correlation of
species abundance to geochemical proxies (e.g., Mg/Ca
temperatures on co-occurring planktonic foraminifera,
De Schepper et al., 2011) is promising for providing
quantitative data.
Biostratigraphy
Since the middle of the twentieth century, palynostratigraphy has emerged as a routine tool in both hydrocarbon exploration and academic research in Mesozoic and
Cenozoic sediments, and numerous biostratigraphic zonations have been erected for Triassic to Neogene sediments
(Stover et al., 1996). Dinocysts typically exhibit high
abundances in neritic settings; thus, the derived stratigraphic information is complementary to that obtained
from typically more offshore groups such as planktonic
foraminifers, coccolithophores, and radiolarians (Pross
and Brinkhuis, 2005). Significant progress has been made
during the past four decades of scientific ocean drilling
(DSDP, ODP, IODP) by assessing stratigraphic ranges
against independent chronostratigraphic information.
Recently, the focus is slowly moving from defining new
zonations toward calibrating bioevents to the geological
time scale on both regional (De Schepper and Head,
2008; Fensome et al., 2008a; Schreck et al., 2012) and
global scales (Williams et al., 2004). This avoids the inherent problem of zonations that zones named after the same
species may have different age ranges. However, few studies illustrate that dinocyst bioevents are rarely synchronous worldwide, and low-, mid-, and high-latitude
bioevents should be distinguished to account for the
observed latitudinal control on species ranges. Nonetheless, some bioevents are useful on regional and/or
supraregional scale and enable stratigraphic correlations
between different basins in the mid- and high latitudes
(Schreck et al., 2012).
Conclusions
In recent years, biological and paleontological studies
have provided a wealth of new information relevant for
the application of recent and fossil dinocysts in marine
geosciences. However, our knowledge of their ecology is
still biased to coastal and shelf environments, and many
open ocean regions such as the Pacific yet remain largely
unexplored. The phylogenetic relationship to the motile
form of many extant species is unknown, and molecular
genetic studies will be particularly useful to address the
long-standing question whether a single dinoflagellate
species forms different cyst species. In the geological
record, dinocysts are of eminent importance for
paleoenvironmental interpretation and biostratigraphy in
high latitudes where preservation of calcareous and
biosiliceous microfossils is poor, but data from lower latitudes are required to inevitably improve independent age
calibration of bioevents. This will also provide new data
on the temporal and spatial distribution of fossil dinocysts,
which together with calibration of species abundances to
geochemical proxies for, e.g., surface temperature, will
lead to a better understanding of cyst paleoecology.
Bibliography
Bijl, P. K., Pross, J., Warnaar, J., Stickley, C. E., Huber, M.,
Guerstein, R., Houben, A. J. P., Sluijs, A., Visscher, H., and
Brinkhuis, H., 2011. Environmental forcings of Paleogene
Southern Ocean dinoflagellate biogeography. Paleoceanography, 26, PA1202, doi:10.1029/2009PA001905.
Bonnet, S., de Vernal, A., Gersonde, R., and Lembke-Jene, L., 2012.
Modern distribution of dinocysts from the North Pacific Ocean
(37–64
N, 144
E–148
W) in relation to hydrographic conditions, sea-ice and productivity. Marine Micropaleontology,
84–85, 87–113.
Chen, B., Irwin, A. J., and Finkel, Z. V., 2011. Biogeographic distribution of diversity and size-structure of organic-walled dinoflagellate cysts. Marine Ecology Progress Series, 425, 35–45.
Dale, B., 1983. Dinoflagellate resting cysts: “benthic plankton”. In
Fryxell, G. A. (ed.), Survival Strategies of the Algae. Cambridge:
Cambridge University Press, pp. 69–144.
Dale, B., 1996. Dinoflagellate cyst ecology: modelling and geological applications. In Jansonius, J., and McGregor, D. C. (eds.),
Palynology: Principles and Applications. Dallas: American
Association of Stratigraphic Palynologists Foundation,
pp. 1249–1275.
Dale, B., 2009. Eutrophication signals in the sedimentary record of
dinoflagellate cysts in coastal waters. Journal of Sea Research,
61, 103–113.
De Schepper, S., and Head, M. J., 2008. Age calibration of dinoflagellate cyst and acritarch events in the Pliocene–Pleistocene of
the eastern North Atlantic (DSDP Hole 610A). Stratigraphy, 5,
137–161.
De Schepper, S., Fischer, E. I., Groeneveld, J., Head, M. J., and
Matthiessen, J., 2011. Deciphering the palaeoecology of Late
Pliocene and Early Pleistocene dinoflagellate cysts.
DINOFLAGELLATES
191