Palaeoceanography SOES 3015 – Palaeoclimate change
Tools and insights
LECTURE 1: Calcareous
Plankton and Benthic
Foraminifera Ecology and
Evolution
SeaWiFS Project, NASA/Goddard Space Flight Center and ORBIMAGE
Part I: Plankton biology and ecology
Phytoplankton
•Coccolithophores
•Diatoms
Zooplankton
•Foraminifera
•Radiolaria
Courtesy of UCL: http://www.ucl.ac.uk/GeolSci/micropal/
planktic foraminifer
~40 mm across
Courtesy of:
http://www.soes.soton.ac.uk/resources/collection/fossil
s/Forams/Eelco/Mediterranean/pages/sacculifer2.htm
Living coccolithophorids
Florisphaera
Gephyrocapsa
Coccolithus
Courtesy of UCL: http://www.ucl.ac.uk/GeolSci/micropal/images/calc/
Benthic foraminifera
Courtesy of Carlton university: http://hoopermuseum.earthsci.carleton.ca/2001_benthicforams_jk/tp.html
http://www.soton.ac.uk/~bam2/col-index/fossi-lindex/Forams/
Eelco/med-agean2/pages/pl-05.htm
Courtesy of Texas A/M univeristy: http://oceanworld.tamu.edu/students/forams/images/foramC-1.jpg
Feeding strategy:
• Foraminifera & radiolarians - Heterotrophs
• Diatoms & coccolithophorids - Autotrophs
Composition:
• Planktonic foraminifera & coccolithophorids - Calcareous
• Diatoms & radiolarians – Siliceous
• Benthic foraminifera - mixed
AGGLUTINATED
CALCAREOUS/ARAGONITIC
ORGANIC
Implications: evolutionary & palaeoceanographic
http://www.noc.soton.ac.uk/obe/PROJECTS/DEEPSEAS/pers_pages/gooday_pers_page.html
http://www.soes.soton.ac.uk/resources/collection/fossils/
Planktonic and Benthic foraminifera
Globigerina
bulloides
Vital statistics:
• Protozoan (single-celled)
Uvigerina
mediterranea
• Size: 50 mm - 5 cm
• Many produce a calcite shell
• Good fossilization potential
• Long fossil record
planktonics ~ 200 Ma Jurassic
benthics ~ 550 Ma Cambrian
• Way of living
planktonic foraminifera “0 - 2.5 km”
benthic foraminifera “sea-bottom”
• Both have a wide geographic distribution
from poles to the tropics
intertidal to deep ocean floor (benthics)
http://www.soes.soton.ac.uk/resources/collection/fossils/Forams/
Living specimen
planktonic foraminifera
Spinose
(heterotrophic)
• Free floating marine zooplankton
• Modern assemblages trace back ~1 Ma
(mid-Pleistocene)
• After death empty shells contribute
substantially to sediments
e.g., Globigerina ooze
Fossil specimen
• Abundance in sediments can reach
10s of thousands of individuals per cm3
http://www.soes.soton.ac.uk/resources/collection/fossils/, http://www.noc.soton.ac.uk/palaeo/index.php
Depth habitat, reproduction frequency, and temperature preference of planktic foraminifera
i
,
r
G. scitula
er
at
w
+
T. quinqueloba
re
u
t
ra
pe
m
te
G. menardii
D. anfracta
G. tenella
G. riedeli
G. ruber
G. theyeri
T. humilis
G. sacculifer
G. bulloides
N. pachyderma
G. inflata
G. uvula
O. universa
G. siphonifera I
G. conglobatus
N. dutertrei
G. glutinata
G. siphonifera II
H. pelagica
G. rubescens
sea surface
av.
pycnocline
R.
100 m
= DCM
P. obliquiloculata
Reproduction
on a half
G. crassaformis
synodic
200 m
lunar
cycle
G. tumida
G. hirsuta
400 m
Reproduction on a
G. truncatulinoides
synodic lunar cycle
2000 m
Reproduction on
an annual cycle
After Schiebel & Hemleben, 2005, Palaeontologische Zeitschrift
Benthic foraminifera
(heterotrophic)
• living in or on the sediment (infaunal and
epifaunal respectively)
• Modern assemblages trace back ~23
Ma (Neogene)
Cibicidoides spp. epifaunal
• density of living benthic foraminifera can
exceed 106/m2, extremely diverse to
wide variety of habitats
Uvigerina spp. infaunal
http://www.soes.soton.ac.uk/resources/collection/fossils/index.html
TROX Model
Benthic foraminifera
Figure redrawn and modified from: Jorissen, F.J., de Stigter, H.C., Widmark, J.G.V., (1995) A conceptual model
explaining benthic foraminiferal microhabitats. Marine Micropaleontology, v. 26, no. 1-4, p. 3-15.
TROX = Trophic condition and Oxygen concentration
Calcareous nannoplankton & nannofossils
Scanning electron microscope
modern
modern
Paleog.
Light microscope
modern
Plio.
Pleisto.
Pleisto.
Eoc. & modern
Plio.
Scale varies
• Free floating marine phytoplankton
• Modern assemblages trace back ~4 Ma
(Pliocene)
• After death coccospheres collapse,
the cocoliths contribute substantially to
sediments, e.g., Upper Cretaceous chalks
• Abundance in sediments can reach
billions of individuals per gram
Courtesy of UCL: http://www.ucl.ac.uk/GeolSci/micropal/calcnanno.html
Additional photos courtesy of Samantha Gibbs
SeaWiFS Project, NASA/Goddard Space Flight Center and ORBIMAGE
Calcareous nannoplankton
Vital Statistics:
• marine algae (single-celled)
• Size: 50-100 µm (liths – 1-20 mm)
COCCOSPHERE
• secrete tiny platelets of calcite - liths
• Good fossilization potential
• Long fossil record
~ 230 Ma Upper Triassic
Coccolithus pelagicus
• Way of living
surface waters 0 - 200 m
• Wide geographic distribution
from subpolar to the tropics
COCCOLITH
Courtesy of UCL: http://www.ucl.ac.uk/GeolSci/micropal/calcnanno.html
Depth stratification in nannofossil ecologies
To view figure please follow the link provided
in the reference below the figure caption.
Emiliania huxleyi
http://www.noc.soton.ac.uk/soes/staff/tt/eh/index.html
Florisphaera profunda
- thermocline dweller
Courtesy of UCL: http://www.ucl.ac.uk/GeolSci/micropal/calcnanno.html
Original Figure caption: Cell density distribution of
coccolithophores and dominant species off Bermuda
through time (January 1991–January 1994) and water
depth (0–200 m): (a) filter samples analysed, (b) total
coccolithophore cell densities, (c) Emiliania huxleyi;
(d) Florisphaera profunda. Densities shaded in units of
20×103 cells l−1. Fig. 4 (cont.) (e) sample distribution;
(f)Umbellosphaera tenuis; (g) Umbellosphaera
irregularis. Densities shaded in units of
5×103 cells l−1.
Haidar, A.T., Thierstein, H.R., (2001), Coccolithophore dynamics off Bermuda (N. Atlantic). Deep-Sea
Research Part II: Topical Studies in Oceanography, v. 48, no. 8-9 p. 1925-1956.
Calcareous plankton distributions:
Planktonic
foraminiferal
provinces
planktonic foraminifera
• Distinct ‘assemblages’ within
different oceanic regimes:
eg. tropical, subtropical,
polar water masses
• Largely absent from continental
shelves
planktonic foraminiferal
‘bio provinces’
Reproduced by permission from Macmillan Publishers Ltd: Molecular evidence for genetic mixing of Artic
and Antarctic subpolar populations of planktonic foramanifers. Darling, K.F., Wade, C.M., Stewart, I.A., Kroon, D., Dingle, R.,
Leigh Brown, J.J., Nature, v. 405, p. 43-47. Copyright (2000), Not under CC License.
Global planktonic foraminiferal diversity
Reprinted by permission from Macmillan Publishers Ltd: Rutherford, S., D’Hondt, S., Prell, W., Environmental controls on the geographic distribution
of zooplankton diversity. Nature, v. 400, p. 749-753. Copyright (1999). Not under CC licence.
•
•
•
•
•
•
Pole to equator gradient with peak diversity in middle latitudes
SST explains 90% geographic variation
Direct control by thermal properties of surface waters – control vertical
niche availability
High lats – virt. absent thermocline & little vertical partitioning of niches
Mid lats – higher SST; thick, permanent thermocline
Tropics – even higher SST (lower diversity), shallow thermocline
planktonic foraminifera species distributions
• Species possess endemic
distributions (their ‘homeland’) but
can periodically range beyond this.
• Evidence suggests endemic
distributions are determined by an
inability to maintain populations in
certain regions, not by an inability
to disperse beyond home range.
• Oceanographic parameters
important in delimiting distribution
of various species:
- temperature
- thermocline and halocline
structure (depth of mixed layer)
- sea-ice extent
- seasonality, upwelling,
phytoplankton blooms
An example of planktonic
Foraminifera species distribution
for G. miocenica, G. menardii.
Pliocene Distribution is shown
with Atlantic Endemics and indopacific Menardelids
The figure can be found in the
following text: (see link in
reference)
Norris, R.D., (1999) Hydrographic and tectonic
control of plankton distribution and evolution in;
Abrantes, F., Mix, A.C.,(eds) Reconstructing
ocean history; a window into the future monogr.
Diversity - ‘Cryptic species’
species with similar morphologies but genetically distinct
= >additional diversity
May have subtly different
ecologies
Reprinted by permission from Macmillan Publishers Ltd:
Darling, K.F., Wade, C.M., Stewart, I.A., Kroon, R.,
Dingle, R., Leigh Brown, J., Molecular evidence for genetic
mixing of Arctic and Antarctic subpolarpopulations of
planktonic foraminifers, Nature, v. 405, p. 43-47 (
Copyright, 2000). Not under CC licence.
Darling et al., (2000), Nature
Images: http://www.soes.soton.ac.uk/resources/collection/fossils/index.html
Hydrographic control on ‘cryptic species’
E.g., Orbulina universa
Courtesy of the National Academy of Sciences: de Vargas, C., Norris, R., Zaninetti, L., Gibb, S.W., Pawlowski, J., (1999) Molecular evidence of cryptic
speciation in planktonic foraminifers and their relation to oceanic provinces. Proceedings of the National Academy of Sciences of the United States of
America, v. 96, no. 6, p. 2864-8.
Hydrographic control on ‘cryptic species’
E.g., Globigerinella siphonifera
The example for Globigerinella siphonifera,
can be found within the following reference:
de Vargas, C., Bonzon, M., Rees, N.W., Pawlowski, J.,
Zaninetti, L., (2002) A molecular approach to
biodiversity and biogeography in the planktonic foraminifer Globigerinella siphonifera (d'Orbigny). Marine
Micropaleontology, v. 45, no. 2, p.101-116.
Hydrographic control on ‘cryptic species’
E.g., Globigerinella siphonifera
The example for Globigerinella siphonifera,
can be found within the following reference:
de Vargas, C., Bonzon, M., Rees, N.W., Pawlowski, J.,
Zaninetti, L., (2002) A molecular approach to
biodiversity and biogeography in the planktonic foraminifer Globigerinella siphonifera (d'Orbigny). Marine
Micropaleontology, v. 45, no. 2, p.101-116.
Nannoplankton biogeographic zones/bioprovinces
Surface water circulation
– compare with bioprovinces
Redrawn by university of Southampton from McIntyre
& Be, 1967, Deep Sea Research
Atlantic biogeography of calcareous nannoplankton
(core top samples)
Follow link to see figure showing Atlantic biogeography for
calcarous nanno plankton Umbilicosphaera and Calcidiscus
Boeckel, B., Baumann, K.H., Henrich, R., and Kinkel, H., (2006) Coccolith distribution patterns in South Atlantic
and Southern Ocean surface sediments in relation to environmental gradients Deep Sea Research I, p. 1073-1099
Umbilicosphaera
oligotrophic, warm-water
favouring
Calcidiscus
More mesotrophic favouring
All species found
everywhere but flourish
where conditions favorable
Boeckel et al. (2006) Deep Sea Research I
Also nannofossil evidence for genetic cryptic species
comes from research by Quinn et al (2004) and
Knappertsbusch et al., (1997)
E.g., Calcidiscus leptoporus
Follow links below for full articles:
Quinn, P.S., Saez, A.G., Baumann, K.H., Steel, B.A., Sprengel, C., Medlin, L.K., (2004) Coccolithophorid
biodiversity: evidence from the cosmopolitan species Calcidiscus leptoporus IN: Thierstein, H.R., Young, J.R., [Eds].
Coccolithophores: from molecular processes to global impact. Springer, i-xiii, p.299-326.
Knappertsbusch, M., Cortes, M.Y., Thierstein, H.R., (1997) Morphologic variability of the coccolithophorid
Calcidiscus leptoporus in the plankton, surface sediments and from the early Pleistocene. Marine
Micropaleontology, v. 30, no.4, p. 293-317.
Biogenic sedimentation
How does surface water production translate into sediment
accumulation?
• Species surface water abundance/distribution patterns correlated to
oceanographic parameters, e.g. SST, salinity.
• Do seafloor assemblages reflect environmental conditions in the
overlying water column where organisms lived?
Consider:
•
•
•
lateral transport
dissolution
On a broad scale flora/fauna in sediment 
surface flora
But:
• On a smaller scale, flora/fauna in sediment =
surface flora
Calcareous plankton and carbonate sedimentation
Pelagic species diversity, biogeography, and evolution
Norris, Richard D Paleobiology. Vol. 26, no. sp4, pp. 236-258.
Dec 2000.
As a generalization:
planktonic foraminifera carbonate
production in surface waters is dominant in
the “eutrophic” upwelling area.
Calcareous nannoplankton carbonate
production in surface waters is dominant in
the “oligotrophic” central gyres.
After Antoine et al., 1996, redrawn by Norris, Palaeobiology 2000
Superimpose water column factors: Carbonate
sedimentation
•
carbonate begins to dissolve at lysocline
•
Carbonate Compensation Depth (CCD)
•
cold water: more CO2, carbonic acid, more dissolution
•
in theory no calcareous sediment below CCD
CCD – ‘snowline’
Map of seafloor sediments
Image sourced from: Open University Course Team (eds) Ocean Chemistry and deep sea
sediments, (1989), Open University.
•
Pattern of sediments is controlled by a combination of water depth, water
saturation state, export production.
Part 2: Calcareous Plankton Evolution
Planktonic foraminifera
Foraminifera evolution
Recognize cycles of evolutionary
change - useful for biostratigraphy
Cretaceous-Paleogene Boundary
Richard D. Norris. Biased extinction and evolutionary trends
Paleobiology 1991 17: 388-399.
http://rcp.missouri.edu/geosci_macleod/research/ktstory1.html(images from Brian Huber)
Speciation models for pelagic environments
In theory dispersal not
limited so how do we get
speciation events?
•
Perhaps there actually is
dispersal limitation
•
Isolation by distance
•
Isolation by depth
•
Isolation by season
•
Changes in reproductive
timing
Surprisingly high rates of
origination and
extinction?
Norris, R.D., Pelagic species diversity, biogeography, and evolution
Paleobiology. Vol. 26, no. sp4, pp. 236-258. Dec 2000.
Tectonic-palaeoceanographic control on evolution
Closing of Central American seaway
Onset of Northern Hemisphere Glaciation
Both affected the evolution of Menardellids, For figure see link below:
Norris, R.D.,(1999) Hydrographic and tectonic control of plankton distribution and
evolution in monograph: Reconstructing ocean history; a window into the future
eds: Abrantes, F; Mix, A. C., ISBN 0-306-46293-1
Benthic foraminifera evolution
•
Much longer geological record than planktonics
•
Molecular studies suggest pre-skeletal record
•
Similar test morphology and wall types developed several times
The evolution of early Foraminifera. Pawlowski, J; Holzmann, M; Berney, C; Fahrni, J; Gooday, AJ; Cedhagen, T; Habura, A; Bowser, SS Proceedings of the National
Academy of Sciences, USA, v. 100, no. 20, p. 11494-11498. 30 Sep 2003. Copyright (2003) National Academy of Sciences, U.S.A.
Calcareous nannoplankton evolution
Coccolithophore and nannolith family-level phylogeny
Redrawn and modified from Bown et al. (2004) Coccolithophores
Nannofossil biodiversity and evolution
•
Biases – much lost from surface waters.
•
Broad correlation with sea-level, ocean cycling, temperature
•
Punctuated by mass extinctions
•
Evolution in late Triassic
Evolution of calcareous nannoplankton
NEOGENE
120
100
Species richness/Rs/Re
80
60
40
Pliocene
20
0
My
Species richness/Speciation/Extinction
0
Miocene
20
PALEOGENE
Oligocene
40
Eocene
PETM
Paleocene
60
K/T BOUNDARY
Maastrichtian
Nannoplankton
diversity
Campanian
CRETACEOUS
Coniacian
Turonian
Cenomanian
JURASSIC
TRIASSIC
OAE2
100
OAE1d
OAE1c
OAE1b
120
OAE1a
Albian
Aptian
Barremian
Hauterivian
Valanginian
Berriasian
Redrawn and modified from
Bown et al. (2004) Coccolithophores
80
Santonian
140
Tithonian
Kimmeridgian
Oxfordian
160
Callovian
Bathonian
Bajocian
Aalenian
180
Toarcian
TOARCIAN OAE
EXTINCTIONRATE (Re)
Pliensbachian
Sinemurian
Hettangian
Rhaetian
200
T/J BOUNDARY
Norian
Carnian
220
Extinction rates
100
Rate of extinction
66.5 Ma K/T boundary
90
Paleogene
Neogene
Jurassic
Cretaceous
80
70
204.5 Ma Triassic/Jurassic boundary
60
54.5 Ma Paleocene/Eocene boundary
Rate
of 50
extinction
60.5 Ma mid-Paleocene
6.5 Ma latest Miocene
45.5 Ma middle Eocene
33.5 Ma Eocene/Oligocene boundary
144.5 Ma Jurassic/Cretaceous boundary
40
30
3.5 Ma midPliocene
background
20
10
0
0
10
20
30
40
Rank Order
Redrawn and modified from: Bown 2005, Micropaleontology
50
60
70
80
Implications:
- Biostratigraphy
- palaeoceanographic reconstructions –
extant and extinct species
Biostratigraphic applications of calcareous plankton
e.g., Discoasters are important biostratigraphic markers for the Cenozoic
MIOCENE
http://paleopolis.rediris.es/cg/CG2005_M01/images/TN_CG2005_M01_Fig_27.gif
Reproduced by permission from Cambridge University Press: Perch Nielson, K., Saunders, J.B., Bolli, H.M., (eds) Plankton
Stratigraphy, vol 1., Planktic Foraminifera, Calcareous Nannofossils and Calpionellids. Cambridge Earth Science Series, 608 pp. (1985)
Biostratigraphic applications of calcareous plankton
IODP Research Vessel Joides
Resolution, Courtesy of IODP
sediment cores
Courtesy of IODP
Planktonic foraminifera
biostratigraphy
ODP Site 762
Exmouth Plateau, NW.
Australian margin
Courtesy of the Micropalaeontological Society: Hancock, C., Dickens, Henderson., (2002) Early Palaeogene planktic foraminiferal
and carbon isotope stratigraphy, Hole 762C, Exmouth Plateau, Northwest Australian margin, v. 21, p. 29-42.
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