See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/223786796

The Great Ordovician Biodiversification Event

(GOBE): The palaeoecological dimension

Article in Palaeogeography Palaeoclimatology Palaeoecology · August 2010

DOI: 10.1016/j.palaeo.2010.05.031

CITATIONS READS

114 552

5 authors, including:

Thomas Servais David A.T. Harper

French National Centre for Scientific Research Durham University
132 PUBLICATIONS 1,894 CITATIONS 323 PUBLICATIONS 14,727 CITATIONS

SEE PROFILE SEE PROFILE

Björn Kröger Axel Munnecke

University of Helsinki Friedrich-Alexander-University of Erlangen-N…
75 PUBLICATIONS 1,007 CITATIONS 117 PUBLICATIONS 2,580 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

RNames Stratigraphical Database View project

Ordovician Cephalopods of Spitsbergen View project

All content following this page was uploaded by Axel Munnecke on 17 December 2014.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document
and are linked to publications on ResearchGate, letting you access and read them immediately.
 Author's personal copy

Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119

Contents lists available at ScienceDirect

Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

The Great Ordovician Biodiversification Event (GOBE):

The palaeoecological dimension
Thomas Servais a,e,⁎, Alan W. Owen b, David A.T. Harper c, Björn Kröger a,d, Axel Munnecke e
a
FRE 3298 du CNRS (Géosystèmes), Université de Lille1, SN5, F-59655 Villeneuve-d'Ascq Cedex, France
b
Department of Geographical and Earth Sciences, University of Glasgow, Gregory Building, Lilybank Gardens, Glasgow G12 8QQ, Scotland, UK
c
Statens Naturhistoriske Museum (Geologisk Museum), Øster Voldgade 5-7, DK-1350 København K, Denmark
d
Museum für Naturkunde, Humboldt Universität zu Berlin, Invalidenstrasse 43, D-10115, Berlin, Germany
e
GeoZentrum Nordbayern, Fachgruppe Paläoumwelt, Loewenichstraße 28, 91054 Erlangen, Germany

a r t i c l e i n f o a b s t r a c t

Article history: The ‘Great Ordovician Biodiversification Event’ (GOBE) saw a spectacular increase in marine biodiversity at all
Received 15 January 2010 taxonomic levels largely within the phyla established much earlier during the so-called ‘Cambrian Explosion’. The
Received in revised form 5 May 2010 diversification was probably the result of a combination of several geological and biological processes and the
Accepted 20 May 2010
positive feedbacks resulting from them. The present paper reviews the palaeoecological dimension of the GOBE. It
Available online 1 June 2010
involved major increases in α, ß and γ biodiversity largely associated with the rise of the Paleozoic Evolutionary
Keywords:
Fauna dominated by suspension feeders and involving a greater occupation of ecospace and more complex
Ordovician ecological structures in the Ecological Evolutionary Units P1 and P2. In the benthos, these include more complex
Biodiversification food webs than those of the Cambrian, greater tiering, especially above the sediment–water interface, and the
Palaeoecology development of guild structures indicating increased competition between taxa for particular resources. The
Palaeoenvironment Ordovician is characterized by a profound change in reef composition, with a switch from microbial-dominated
reefs in the Early and Middle Ordovician to metazoan-dominated reefs in the Late Ordovician. Increases in
complexity of deep-water trace fossil assemblages began in the Early Ordovician and mark the increasing
exploitation in that environment and the development of hardgrounds permitted bioerosion and encrusting
strategies together with the appearance of cryptic communities.
Within the water column, the GOBE involved a significant increase in the diversity of the phytoplankton and the
development of a diverse zooplankton (including planktotrophic larvae from a range of invertebrate clades). This
revolution in the plankton enabled the establishment of a diverse fauna of pelagic vertebrates, molluscs and
arthropods and promoted the rise of suspension feeders in the benthos. An escalation amongst predators and thus
community evolution may also have been a major driver of biodiversification.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction successive rises to dominance of what he termed the ‘Cambrian’ and

1.1. The International Geoscience Programme (IGCP) n° 503 ‘Ordovician
Palaeogeography and Palaeoclimate’
Two sustained rises in the biodiversity of marine organisms took
place during Earth history. The first is recorded during the Early
Palaeozoic and the second started at the beginning of the Mesozoic (e.g.
Benton, 1995; Sheehan, 2001a; Crame and Owen, 2002; Stanley, 2007).
The Early Palaeozoic radiation of life in the oceans includes both the
‘Cambrian Explosion’ and the ‘Great Ordovician Biodiversification
Event’, or GOBE (Webby et al., 2004a; Harper, 2006). Based on diversity
counts, Sepkoski (1978, 1979, 1981, 1984) not only identified these
events, but also described their taxonomic content in terms of the
⁎ Corresponding author. FRE 3298 du CNRS (Géosystèmes), Université de Lille1, SN5,
F-59655 Villeneuve-d'Ascq Cedex, France.
E-mail address: thomas.servais@univ-lille1.fr (T. Servais).
‘Paleozoic’ Evolutionary Faunas. In addition, a major crash in diversity
took place close to the end of the Ordovician Period. This is the oldest
and second largest of the ‘Big Five’ Mass Extinctions and has been widely
linked to the Hirnantian glaciation, and thus to an episode of major
climate change (e.g., Brenchley et al., 1994; Sheehan, 2001b).
The highly successful International Geological Correlation Programme
(IGCP) project n° 410 ‘The Great Ordovician Biodiversification Event’
extended from 1997 to 2002, and resulted not only in the generation of
biodiversity curves for all Ordovician fossil groups (Webby et al., 2004a)
but also in the development of an improved globally-integrated
biozonation for graptolites, conodonts, chitinozoans and other fossil
groups (Webby et al., 2004b). Miller (1997a, 1998, 2000, 2004) argued
that global Ordovician biodiversity curves are a summation of patterns
that differ from palaeogeographical region to region and from clade to
clade. He emphasised the importance of dissecting global biodiversity
curves to reveal these underlying patterns and significant progress was
achieved in this respect in the individual chapters in Webby et al. (2004a),

0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2010.05.031
 Author's personal copy
100 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
compiled by teams of experts on each major clade. Moreover, while the
Ordovician diversification of some fossil groups was spectacularly rapid,
other groups evolved progressively over a longer period of time. It also
became clear that different methods of quantifying diversity (e.g. see
Cooper, 2004) or the time bins within which the data are grouped (e.g.
Alroy, 2000) strongly affected the diversity curves, and therefore their
possible interpretations (Servais et al., 2008, 2009a).
After the closing meeting of IGCP 410 in 2001 a successor project
was proposed to attempt to find the possible physical and/or chemical
causes of the Ordovician biodiversification, the end-Ordovician
extinction, and the Silurian radiation. This new project, IGCP n° 503
‘Ordovician Palaeogeography and Palaeoclimate,’ ran from 2004 to
2008, and continued ‘on extended term’ in 2009.
A wide range of factors has recently been postulated to have caused or
at least promoted the Great Ordovician Biodiversification Event. Trotter
et al. (2008), for example, considered global cooling as the main trigger of
the Ordovician biodiversification, while Achab and Paris (2007) and Paris
(2008) related the diversification to sea-level changes, volcanic events
and climate change. Miller and Mao (1995), for example, discussed the
relation between the Ordovician biodiversification and orogenic activities,
while Barnes (2004b) related it to a possible superplume event. In
contrast to all these terrestrial processes Schmitz et al. (2008) and Parnell
(2008) linked the biodiversification to an asteroid breakup event and its
consequent high meteorite flux. The present review paper is focused on
the palaeoecological dimension of the radiation and is part of a thematic
set of papers on Early Palaeozoic palaeoenvironments (Servais and Owen,
this issue) arising from what was originally intended to be the final
meeting of IGCP project 503 organised in August 2008 in Lille (France). It
draws on an extensive recent literature including earlier thematic sets of
papers arising from the project (Munnecke and Servais, 2007; Owen,
2008) and more broadly based reviews (Harper, 2006; Servais et al.,
2009a). A second special issue of the journal arising from the Lille meeting
is focused on climatic evolution and its impact during the Ordovician and
Silurian(Munnecke et al., this issue).
1.2. What is the ‘Great Ordovician Biodiversification Event?’
There is a long history of the production of diversity counts of fossil
organisms in order to understand the evolution of life through
geological time (e.g. Miller, 2009). As indicated in the review by
Harper (2006), one of the first attempts was that of Phillips (1860),
who, the year after Darwin's publication of the ‘Origin of Species,’
produced a semi-quantitative analysis of the biodiversity of Phaner-
ozoic life. The curve compiled by Phillips (Harper, 2006, Fig. 1) was
divided into three stratigraphically-based marine faunas, the first of
them corresponding to ‘Palaeozoic life.’ More than a century later, the
diversity counts of Sepkoski (1978, 1979, 1981, 1984, 1991, 2002)
confirmed the major radiations of marine diversity during the
Phanerozoic, and clearly pointed out that the most spectacular
increase of marine animal diversity at the family level took place
during the Ordovician, today known as ‘The Great Ordovician
Biodiversification Event’. Not only was the rapid increase of diversity
clear, but also the dramatic changes in palaeoecology, with the rise of
the ‘Paleozoic Evolutionary Fauna’ in the Ordovician progressively
replacing the ‘Cambrian Evolutionary Fauna’ (e.g. Sepkoski and
Sheehan, 1983; Sepkoski and Miller, 1985; Sepkoski, 1995; Sheehan,
1996; Droser and Sheehan, 1997).
The terms ‘Cambrian Explosion’ and ‘Great Ordovician Biodiversi-
fication Event’ are currently used to designate the two major events in
the evolution of marine life during the Early Palaeozoic. While many
authors consider these events to be separate, others believe that they
belong to a single, long-term radiation. The widely used (but
increasingly debated) term ‘Cambrian Explosion’ reflects the sudden
appearance of numerous organisms in the fossil record during the
Early and Middle Cambrian (e.g. Bottjer et al., 2000, 2001; Butterfield,
2001; Zhuravlev and Riding, 2001). Almost all animal phyla appeared
during the latest Proterozoic and Early Cambrian with only a few
making their appearance earlier or later in Earth history (e.g. Eble,
1998). Knowledge of the Lower and Middle Cambrian faunas is greatly
enhanced by several exceptionally preserved assemblages (e.g. the
Chengjiang, Sirius Passet and Burgess Shale biotas), which can be
considered as ‘taphonomic windows’ that allow a more complete view
of the body plans that appeared since the latest Precambrian. Such
‘Konservatfossillagerstätten’ are well known in the Cambrian, but are
rare in the Ordovician (e.g. Aldridge et al., 1994; Van Roy et al., 2010).
The extensively studied Cambrian ‘Fossillagerstätten’ are dated to
constrain the ‘Cambrian explosion’ to a time interval that had ended
by about 530 Ma. After this ‘explosion’ of animal phyla, over 40 myr
occurred before the start of the Early and Middle Ordovician
diversification (ca 485 to 460 Ma) that can be considered largely as
an ‘explosion’ of diversity at lower taxonomic levels. For several fossil
groups, this expansion of diversity at the order, family, genus and
species level occurred during a relatively short time span of only some
10 or 20 myr, and hence the Ordovician biodiversification is
considered by some authors as the most rapid diversity increase in
marine life during Earth history. Thus, for example, the brachiopods,
show a dramatic increase of diversity during a relatively short time
interval in the Floian–Darriwilian (Early and Middle Ordovician) (e.g.
Harper et al., 2004; Harper, 2006). Others such as corals diversified
later, but also very rapidly, during the Sandbian Stage, in the Late
Ordovician (Webby et al., 2004c). In contrast, yet other groups show a
continuous and long-term biodiversification that in some instances
started in the Cambrian. Such groups include some echinoderm
subphyla (e.g. Lefebvre and Fatka, 2003; Nardin and Lefebvre, this
issue) and the phytoplankton (e.g. Servais et al., 2008).
The Ordovician biodiversification is seen both in terms of
individual clades and in regional biodiversity curves (e.g. Miller,
1997b, 1997c, 2004). Hammer (2003) for example calculated the total
mean standing diversity of all fossil groups recorded from Baltica (the
most extensively studied palaeocontinent in terms of Ordovician
faunas) indicating a clear diversity increase with highest values
reached at the base of the Upper Ordovician. An obvious Ordovician
biodiversification is also recorded for all fossil groups from the South
Chinese Yangtze Platform (e.g. Chen et al., 2006).
On the other hand, the ‘Cambrian explosion’ and the ‘Ordovician
diversification’ can be viewed as belonging to a single ‘Cambrian–
Ordovician radiation’. Droser and Finnegan (2003), for example,
considered the Ordovician biodiversification to be rooted in the
Cambrian explosion and the Ordovician phytoplankton radiation was
part of a diversification that clearly started in the Cambrian (Servais et al.,
2008). The distinction between separate diversification steps in the
Cambrian and the Ordovician may in some instances be a sampling
artefact: Nardin and Lefebvre (this issue) consider that the relative lack
of extensive investigations in the Upper Cambrian may be responsible
for the apparently separate pulses in the Early Palaeozoic diversification
of blastozoans. The lack of Upper Cambrian studies on blastozoans is
possibly related to the absence of sedimentary rocks suitable for the
preservation of this fossil group, in particular in the ‘peri-Gondwanan’
areas that have been extensively studied for the lower and middle
Cambrian together with the Ordovician.
Similarly, the most recent diversity curve at the genus level by the
researchers working on the Paleobiology Database (Alroy et al., 2008)
suggested that the Cambrian and Ordovician events cannot easily be
distinguished, and that both the Cambrian explosion and the
Ordovician biodiversification belong to a single, large-scale radiation,
that started in the Precambrian and that was only halted at the base of
the Devonian (e.g. Alroy et al., 2008, Fig. 1).
1.3. Diversity counts: methods and results
Data through the Ordovician biodiversification event have been
assembled and analyzed in a variety of ways. The major global signal has
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
been recognized on the basis of fossil range data culled from literature,
principally the Treatise on Invertebrate Paleontology (e.g. Sepkoski,
1979, 1981, 1995, 2002) and the Fossil Record (e.g. Benton, 1993, 1995).
These compilations are, however, based on range-through data, a valid
assumption but one that fails to identify Lazarus taxa.
Interpretation of such curves, however, has been open to criticism.
For example Raup (1972) cited some nine important filters that could
bias the fossil record and later (Raup, 1975) noted that the Phanerozoic
diversity curve may reflect merely the amount of available rock volume
for any one time interval (see also McGowan and Smith, 2007) or the
efforts of taxonomists on selected intervals. Raup (1975) thus concluded
that the Phanerozoic biodiversity was probably saturated and in a state
of equilibrium for much of its duration. More recently Peters and Foote
(2002) have again emphasised the correlation between diversity and
numbers of available rock formations and Smith (2001) noted the
correspondence between high sea level and high diversity.
Nevertheless, Peters (2005) has suggested that there might be a
common cause for all three phenomena, rather than conclude that the
Phanerozoic record is a mere artefact of geological processes. Plate
tectonic processes and eustatic changes in sea level for example would
help accelerate diversity during intervals of high sea level, when platforms
were flooded also providing a much larger volume of sedimentary rocks,
generating a true biological signal. This is particularly relevant for the
GOBE, indicating that the accelerating diversity trends are real and not
simply a function of an increased area and volume of Ordovician rocks.
By contrast, the curves of Alroy et al. (2008) are based on locality-
based counts stored in the Paleobiology Database. Hammer (2003)
applied a similar strategy in the assembly of data from the Ordovician
of the Baltic province, demonstrating the unambiguous track of the
event, while deconstructing the curve into patterns and trends for
individual taxonomic groups. All these curves, however, are not free
from error. The global Phanerozoic curves developed by Alroy et al.
(2008) are, for example, subject to sampling bias. Localities in older
formations are possibly less numerous and thus the high levels of
diversity seen in the Modern Evolutionary Fauna may simply reflect
that localities within this unit are more accessible and numerous.
Nevertheless, when the curves are standardized against sample
size, the GOBE stands out as a massive hike in biodiversity rarely
equalled again throughout the rest of the Phanerozoic.
2. The Ordovician world
2.1. Ordovician stratigraphy
The objective of IGCP 503 was not to revise the stratigraphy of the
Ordovician System in detail, although the programme was based on a
strong interaction between the participants in IGCP 503 and the
Subcommission on Ordovician Stratigraphy (SOS). However, in order
to understand the diversity trends during the Ordovician and to
compare not only different trends from individual fossil groups, but
also to correlate trends from different palaeocontinents, a precise
stratigraphical correlation scheme is essential.
The marked faunal provincialism of the Ordovician has long hindered
international correlation. Although the British Ordovician series and
stages acted more or less as a sort of standard, especially in western,
southern and central Europe, the definition of these historical subdivisions
of the Ordovician System in the British Isles did not fulfil the
recommendations of the ICS (International Commission on Stratigraphy)
and of the IUGS (International Union of the Geological Sciences) for the
definition of global stratotypes. New Series and Stages have therefore
been defined in the last decade or so.
Based on the extensive work of numerous biostratigraphers, and
within the framework of the SOS, Webby et al. (2004b) published a
detailed Ordovician stratigraphical chart showing the correlations
between the main graptolite, conodont and chitinozoan zonal sequences
from the major palaeocontinents, including Laurentia, Gondwana, Baltica,
101
Avalonia and China. Most recently, Bergström et al. (2009a) have
published a chart correlating the main regional chronostratigraphical
schemes with the international Series and Stages (Fig. 1). They also
compiled a composite δ13C curve for the entire Ordovician. The new
international stratigraphical framework comprises three Global Series
(Lower, Middle, Upper) and seven Global Stages (successively: Trema-
docian, Floian, Dapingian, Darriwilian, Sandbian, Katian, Hirnantian)
(Bergström et al., 2006a). Finer scale correlation is facilitated by 19 Time-
Slices (TS) and 20 Stage-Slices (SS) defined by Webby et al. (2004b) and
Bergström et al. (2009a), respectively (Fig. 1). The latter units allow a
precise correlation in both carbonate and shale facies, and are tied to the
composite δ13C curve through the entire Ordovician. While this composite
global curve will doubtlessly be refined by further detailed regional
studies (e.g. Ainsaar et al., this issue), chemostratigraphy holds the
prospect of being a major correlation tool in the Early Palaeozoic as well as
a key to global environmental changes (Cramer et al., this issue).
The bases of the Global Stages are defined at the base of either a
graptolite or a conodont biozone of wide correlative value. Many of
the ‘time slices’ (Webby et al., 2004b) and ‘stage slices’ (Bergström et
al., 2009a), however, are of greater temporal extent than a single
biozone. The ‘stage slices’ (Bergström et al., 2009a) are intended to
replace the ‘time slices’ introduced by Webby et al. (2004b) and used
in the compilation of biodiversity curves for many groups of
organisms (Webby et al., 2004a).
Some biodiversity databases were established long before the
compilation of Webby et al. (2004a) and were based on Ordovician
subdivisions with boundaries that do not precisely match those of the
more recently defined units. Problems of correlation between
diversity trends are therefore possible and must be borne in mind
when biodiversity curves are compared and analyzed. Future
interpretations of biodiversity trends must thus be based on an
accurate stratigraphical correlation, and the work of Webby et al.
(2004b) and Bergström et al. (2009a) are important steps towards
this.
2.2. Ordovician palaeogeography
Progress made during the last 25 years in understanding Ordovi-
cian palaeogeography has resulted from the integration of tectonic,
stratigraphical, palaeoclimatological, palaeomagnetic and palaeonto-
logical evidence (e.g. Fortey and Cocks, 2003). After the benchmark
publication of Scotese and McKerrow (1990) numerous palaeogeo-
graphical reconstructions have been published including those of
Harper et al. (1996), Torsvik (1998), Cocks (2001) and Cocks and
Torsvik (2002, 2004, 2006). There is currently a general agreement on
the position of the major palaeocontinents: Laurentia (North America,
Greenland and slivers of crust in Norway and the British Isles) and
Siberia, both found in equatorial positions, Baltica, located at
intermediate latitudes in the southern hemisphere, the microconti-
nent of Avalonia that drifted from the high latitude margin of
Gondwana towards the tropics, and Gondwana (Fig. 2). This
supercontinent Gondwana was by far the largest continent. It
included South America, Africa, Antarctica, Australia, India and other
marginal (micro-) continents, amongst them North and South China.
Gondwana extended from the South Pole to the Equator and remained
separated from Laurentia throughout the Ordovician. Numerous other
microcontinents and terranes were present, and the precise positions
of many of them are widely discussed (e.g. Bassett, 2009).
The detailed analysis of the palaeobiogeographical distribution of
different fossil groups was a major research objective of IGCP 503 and will
be the subject of a special publication currently under preparation. Suffice
it to note here that at least to some extent the diversification of marine
organisms during the Ordovician was most probably related to the
palaeogeography of the Period. The Ordovician was a time of maximum
dispersal of the continents between episodes when they were amalgam-
ated in the supercontinents of Rodinia in the late Proterozoic and Pangaea
 Author's personal copy

102 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119

Fig. 1. The modern stratigraphical scheme for the Ordovician (Global Series, Global Stages, time slices, stage slices; Bergström et al., 2006a, 2009a; Webby et al., 2004b) plotted
against δ13C isotopes (Bergström et al., 2009a), sea level (Haq and Schutter, 2008), strontium isotopes (Shields et al., 2003; Shields and Veizer, 2004; Young et al., 2009), the reef
development (Webby, 2002), the climate signal (greenhouse–icehouse).

in the late Palaeozoic. Nearly four decades ago, Valentine and Moores
(1972) recognized a strong correlation between the position of the
palaeocontinents and biological diversity at a large (Phanerozoic)
temporal scale. They argued that during periods when continents were
fragmented there were more opportunities for allopatric speciation and
biological diversity was high. In contrast, when the continents were
grouped together with few geographical barriers and more limited
opportunities for allopatric speciation, diversity was low. The two major
radiations of marine life during the Phanerozoic can thus be related in
general terms to the breakup of the supercontinents of Rodinia during the
latest Proterozoic and early Palaeozoic and Pangaea in the Mesozoic
leading to the creation of more numerous and increasingly widely
separated continental platforms, new oceans and abundant new diversity
biodiversity foci. Miller and Mao (1995) and Owen and Crame (2002), for
example, also noted that the Ordovician biodiversification corresponded
to a period of time of major plate tectonic change. The same applies to the
Cambrian radiation, which Liebermann (2004) considered to have been
preceded by one of the most substantial plate reorganizations in Earth
history.
The Ordovician is not only considered to have been the Palaeozoic
period with greatest continental dispersal, but the distribution of the
continents together with high sea levels also led to the greatest
flooding of the continental shelves (Algeo and Seslavinsky, 1995) and
the greatest tropical shelf areas in the Phanerozoic (e.g. Walker et al.,
2002). Regions of high diversity are common in the tropics today and
it is reasonable to assume that such areas existed on the extensive
tropical shelves of the Ordovician oceans.
2.3. Ordovician geology: sea-floor spreading, volcanism, superplume
hypothesis
The very dynamic Ordovician palaeogeography with numerous
moving continents, terranes and island arcs, subduction zones and
 Author's personal copy

T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119 103

Fig. 2. Early Ordovician reconstruction with a view on the South Pole (based on Cocks and Torsvik, 2002), illustrating the debated palaeogeographical position of the Ordovician
palaeocontinents. The Gondwana supercontinent with its numerous marginal terranes extended from the South Pole to low latitudes in the northern hemisphere. The
microcontinent Avalonia drifted off Gondwana during the Early Ordovician to collide with Baltica during the latest Ordovician. Other major continents of the Ordovician include
Siberia and Laurentia at intermediate and low latitudes near the equator.

orogenic belts most probably had significant impacts on global
geochemical cycles and also on climate and palaeobiodiversity.
Together with the Cretaceous, the Ordovician Period had the greatest
amount of volcanism of the Phanerozoic (e.g. Stillman, 1984; Bergström
et al., 2004). Thick and very widespread volcanic ash falls, possibly the
largest in the Phanerozoic, have been recorded in the Ordovician
including the Millbrig and Kinekulle K-bentonites (Huff et al., 1992;
Kolata et al., 1996; Bergström et al., 1995, 2004). The intense volcanic
activity most probably had important consequences for climate change,
sea level and the creation of many, relatively short-lived oceanic and
marginal basins with volcanic islands and archipelagos. Moreover, in
addition to providing habitats for many groups of organisms, and
increasing γ biodiversity (see Section 3.4), volcanic environments could
also favour the exceptional preservation of biotas (e.g. Botting, 2002),
while generating local biodiversity peaks.
A postulated driver for the GOBE is a putative Middle Ordovician
superplume event (Barnes, 2004b) that may have induced major marine
transgression, high carbon dioxide levels together with a super
greenhouse effect, extensive (non-felsic) volcanism, widespread black
shales and a strontium isotope excursion indicative of increased mantle
input. The drastic change of the strontium isotope values (Shields et al.,
2003; Shields and Veizer, 2004), in particular, correlates roughly with
the Darriwilian–Sandbian boundary, an important interval in the
development of the Paleozoic Evolutionary Fauna, and in particular
the development of reefs and carbonate build-ups (Fig. 1). Recent work
by Young et al. (2009) has linked the major rapid drop in 87Sr/86Sr to
basalt weathering. Lefebvre et al. (in press) have argued that associated
with the postulated superplume event (Caldeira and Rampino, 1991;
Condie, 2001; Pavlov and Gallet, 2005; Courtillot and Olson, 2007), the
emplacement of a basaltic province at low latitudes, hypothetically
placed between Baltica and Laurentia, could have been responsible for
important climate changes. Such a basaltic province would have
impacted on the global atmospheric pCO2, with a temperature rise
during the Late Ordovician, followed by an important temperature fall as
the consequence of the erosion of basaltic rocks, changing dramatically
the global carbon cycle. The impact on the global carbon cycle of a
superplume event could thus have been a possible trigger for the short
global warming event (‘Boda Event’ of Fortey and Cocks, 2005) and the
Hirnantian glaciation (see below).
2.4. Ordovician climate, CO2, sea level, sea-water chemistry and oceanic
circulation patterns
It is beyond the scope of the present paper to discuss the climate
and sea-level changes of the Ordovician in detail. Recent studies
focused on these aspects are grouped elsewhere (Munnecke et al., this
 Author's personal copy
104 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
issue). Here, only a short summary of the palaeoclimatic conditions is
provided in order to place them within a palaeoecological context.
Understanding of the climate in the Early Palaeozoic has increased
significantly in the last decade or so. Munnecke et al. (2003), for
example, demonstrated that the Silurian was not a climatically
monotonous period, but an interval with intense climate changes
resulting in isotopic excursions, which are still not fully understood or
explained. The latest Ordovician (Hirnantian) has long been recognized
as one of the few intervals of the Phanerozoic when ice sheets covered
the poles of the Earth (e.g. Hambrey and Harland, 1981; Caputo, 1998).
The relationship between carbon isotopic excursions, sea-level fluctua-
tions and climate change is widely debated (e.g. Holmden et al., 1998;
Veizer et al., 2000; Joachimski et al., 2003; Tobin and Bergström, 2002;
Munnecke et al., 2003; Fanton and Holmden, 2007). However, extensive
documentation in recent years of carbon isotope changes has not only
thrown light on the details of the Hirnantian glacial event but has also
provided evidence for cooling beginning much earlier in the Ordovician
than was thought only 15 years ago (see Section 2.5) (e.g. Hatch et al.,
1987; Ainsaar et al., 1999, 2004; Kaljo et al., 2001, 2003; Saltzman and
Young, 2005; Young et al., 2005, 2008; Bergström et al., 2006b; Schmitz
and Bergström, 2007; Bergström et al., 2009b).
The Ordovician was a period of extremely high sea levels (Fig. 1),
considered by some scientists as the highest of the Palaeozoic, if not the
entire Phanerozoic (e.g. Haq et al., 1988; Hallam, 1992; Barnes, 2004a;
Miller et al., 2005; Haq and Schutter, 2008). Documentation and
understanding of Ordovician sea-level change is still at an early stage.
Regional sea-level curves have now been proposed for some palaeo-
continents, including Baltica (e.g. Dronov and Holmer, 1999, 2002;
Nielsen, 2004, 2007), Laurentia (e.g. Ross and Ross, 1992, 1995; Landing,
2002) and China (e.g. Su, 2001). However, international correlations of
sea-level fluctuations are still problematical and will continue as a major
research objective in the coming years (Munnecke et al., in press-b).
Similarly, models of oceanic circulation during the Ordovician are
also at best preliminary. Parrish (1982, 1987) and Parrish et al. (1983)
discussed Early Palaeozoic oceanic circulation patterns in the context of
upwelling zones and the research of petroleum source beds. In the early
1990s Jeppsson (1990) discussed an oceanic model for the Silurian,
whereas Wilde (1991) postulated climate zones and oceanic currents
for the Lower, Middle and Upper Ordovician, plotted on the palaeogeo-
graphical base maps of Scotese and McKerrow (1990). Subsequently,
Christiansen and Stouge (1999) proposed an oceanic current model to
explain faunal exchanges during the Ordovician. Most recently
Armstrong et al. (2009) have used oxygen isotope curves to track the
changing positions and influence of the Inter-tropical Convergence Zone
in the build up to the Hirnantian glaciation. Many palaeontologists have
also proposed scenarios to explain the evolution of the palaeogeo-
graphical distribution of their fossil groups, but rarely with fully justified
circulation patterns (see also Fortey and Cocks, 2003).
On the other hand, it has become increasingly evident that the sea-
water chemistry changed significantly during the Ordovician, and these
changes are currently discussed and debated (e.g. Stanley and Hardie,
1999; Saltzman, 2005). In particular, the precipitation of CaCO3 and sea-
water saturation states must be considered in the context of the
development of carbonate build-ups and reefs during the Ordovician
(e.g. Riding, 1993, 2000, 2002, 2006a, b; Riding and Liang, 2005; Palmer
and Wilson, 2004).
xMost importantly, in the last decade numerous papers have been
published regarding the modelling of atmospheric carbon dioxide.
Besides the GEOCARB models of Berner and his colleagues (e.g.
Berner, 1994; Berner and Kothavala, 2001), other authors have
proposed models for the Early Palaeozoic carbon cycle (e.g., Goddéris
et al., 2001). The strontium isotopic ratio of sea water has also been
discussed in several papers (e.g., François and Walker, 1992; Veizer et
al., 1999; Shields et al., 2003, Shields and Veizer, 2004; Young et al.,
2009), indicating a dramatic shift at the beginning of the Late
Ordovician (Fig. 1), that can probably be correlated with palaeoeco-
logical changes at this period including a shift in reef composition (see
below).
The physical and chemical changes in the Ordovician oceans, with
high sea levels and warm temperatures, and also with more abundant
nutrients available (as a consequence of high tectonic activity and
volcanism), were probably responsible for the increase in primary
production. This would have accompanied the increased diversity of
primary producers, although the relationship of biomass production and
diversity of the phytoplankton are difficult to quantify (see below).
2.5. Ordovician cooling and the Hirnantian glaciation
The impact of the Hirnantian glaciation on the ecosystem has been
discussed in many studies dealing with the Late Ordovician mass
extinction (for a summary, see Sheehan, 2001b). While the glaciation
was once thought to represent a very short cold interval in an otherwise
greenhouse climate (e.g. Brenchley et al., 1994) most recent work points
to cooling beginning much earlier in the Ordovician (Saltzman and
Young, 2005; Trotter et al., 2008; Vandenbroucke et al., 2009, this issue)
with the Hirnantian representing the most intense phase of a ‘long-lived
glaciation’ (Saltzman and Young, 2005) possibly starting in the Middle
Ordovician (Fig. 1) and lasting well into the Silurian (Grahn and Caputo,
1992; Díaz-Martínez and Grahn, 2007). The decrease of biodiversity in
many fossil groups following the early Late Ordovician (see Webby et al.,
2004a; Servais et al., 2008) may reflect a global cooling of the climate. A
short warming event in the Late Ordovician (the ‘Boda Event’ of Fortey
and Cocks, 2005, but see also Cherns and Wheeley, 2007) also had a clear
influence on the fossil record (e.g. Fortey and Cocks, 2005; Owen, 2007;
Jiménez-Sánchez and Villas, this issue).
Several causes of the Ordovician glaciation have been postulated
(see Armstrong, 2007). Changes of pCO2, palaeoceanography and sea
levels, palaeogeographical models including continental uplift and its
related weathering, movement of the Inter-tropical Convergence
Zone and the impact of a hypothetical superplume event have been
invoked (e.g. Kump et al., 1999; Gibbs et al., 1997; Herrmann et al.,
2004a,b; Lefebvre et al., in press; Armstrong et al., 2009). As with the
Ordovician biodiversification, global cooling and the glaciation in the
Late Ordovician were probably not triggered by a single abrupt factor,
but by the complex interplay of several drivers.
3. Palaeoecology of the GOBE
3.1. Historical overview
Interrogation of Phanerozoic diversity patterns and trends has
indicated that life is characterized by long intervals of stasis separated
by shorter intervals of change and reorganization (Sheehan, 1996).
Sepkoski's (1981) visionary studies divided Phanerozoic marine life into
three evolutionary faunas (EFs), the Ordovician biodiversification
forming the basis for his Paleozoic Evolutionary Fauna (Fig. 3). This
fauna was populated by communities of intermediate diversity,
dominated by suspension feeders and some detritivores and carnivores,
where tiering was common, and a plankton-based trophic structure, of
intermediate complexity, had already developed.
There is a clear and graphic distinction between sea-floor reconstruc-
tions of the Cambrian and Paleozoic evolutionary faunas (e.g. McKerrow,
1978), indicating a change in dominance from loosely-structured and
variable communities of trilobites, phosphatic brachiopods, early echino-
derms and molluscs to the more predictable community structures,
dominated by suspension feeders, in the Paleozoic Evolutionary Fauna.
However, Phanerozoic life can be further dissected into Ecologic
Evolutionary Units (EEUs) (Boucot, 1983; Sheehan, 1996; Fig. 3): the
Paleozoic Evolutionary Fauna containing four such intervals (P1–P4). Of
particular importance are the P1 and P2 intervals that cover the GOBE,
during which many new animal groups with new adaptive morphologies
generated increased complexity and diversity within communities during
 Author's personal copy

T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119 105

Fig. 3. The classical ‘Sepkoski’ diversity curve of marine invertebrate families through Phanerozoic time, documenting the Cambrian, Paleozoic and Modern Evolutionary Faunas, the
Great Ordovician Biodiversifiation Event (GOBE), and the ‘Big Five’ mass extinctions of marine invertebrates, plotted against the diversity curve of the Paleobiology Database (Alroy
et al., 2008). Note that in both datasets the GOBE stands out as a massive hike in biodiversity rarely equalled again throughout the rest of the Phanerozoic. Ecological-Evolutionary
Units (EEUs) after Sheehan (1996): C1–C2: Cambrian; P1–4: Palaeozoic; M1–3: Modern. Geological periods, from left to right: C: Cambrian, O: Ordovician, S: Silurian, D: Devonian,
C: Carboniferous, P: Permian, T: Triassic; J: Jurassic, C: Cretaceous, T: Tertiary.

unusual periods of biotic change (Sheehan, 1996). Subsequent studies
involving the development of the palaeoecological architecture of
Phanerozoic life (Droser et al., 1997; Bottjer et al., 2001) have noted
important changes at virtually all levels of ecological structure. A series of
more focused papers has demonstrated that the taxic dominance of
communities and shell concentrations changed markedly from trilobite to
brachiopod-dominated assemblages within the early part of the Ordovi-
cian (Droser and Finnegan, 2003; Finnegan and Droser, 2008a). Trilobites
remained an important part of the Ordovician benthos, but there was a
spectacular increase in abundance, biomass, diversity and body size of the
Brachiopoda.
3.2. The rise of the Paleozoic Evolutionary Fauna
The Ordovician marks the rise to dominance of the Paleozoic
Evolutionary Fauna identified by Sepkoski (1981, 1984) in a multivariate
analysis of his marine biodiversity curves. It superseded the Cambrian
Evolutionary Fauna and established the broad pattern of marine life that
lasted until the end of the Permian. The Paleozoic Evolutionary Fauna was
dominated by a different set of phyla and classes than had typified the
Cambrian, including a major expansion in epifaunal suspension feeders. It
involved a significant increase in diversity at all levels (α, ß and γ) and
much greater ecological complexity including more complex food webs,
greater tiering both above and below the sediment–water interface, an
expansion of the plankton and greater occupation of the water column
(see Sheehan, 2001a for summary).
Jablonski et al. (1983) and Sepkoski and Sheehan (1983) documen-
ted a pattern of nearshore innovation and offshore expansion of
communities during the Ordovician (see also Sepkoski and Miller,
1985; Sepkoski, 1991; Harper, 2006). In gross terms this meant a
diachronous onshore–offshore change from trilobite and ‘inarticulate’
brachiopod-dominated palaeocommunities typical of the Cambrian
Evolutionary Fauna to the rhynchonelliformean brachiopod-domi-
nated Paleozoic Evolutionary Fauna and ultimately, in shallow-water
settings, the mollusc-dominated Modern Evolutionary Fauna. The
process may even have begun in the Cambrian. Bassett et al. (2002),
for example, traced the origins of the brachiopod-dominated faunas of
the Ordovician in Cambrian shallow-water carbonate environments of
Gondwana and its marginal terranes.
As with so many aspects of biodiversity change, the overall onshore–
offshore pattern of community change needs to be teased apart to be
fully understood. Thus, for example, Westrop et al. (1995) and Westrop
and Adrain (1998a) (see also Westrop and Adrain, 1998b; Miller et al.
(1998) have shown that trilobite alpha diversities remained remarkably
constant through the Ordovician across a range of shelf settings in
Laurentia. This, they argued, indicates that rather than trilobites being
displaced oceanwards as the changing onshore–offshore pattern might
superficially suggest, their contribution to the total α diversities of shelf
palaeocommunities was simply being ‘diluted’ by that of the more
rapidly radiating groups (but see Finnegan and Droser, 2008b). Those
trilobite families that diversified during the GOBE within the “Whiterock
Fauna” of Adrain et al. (1998, 2004) were part of the Paleozoic
Evolutionary Fauna and include the groups that survived the Late
Ordovician extinction. Adrain et al. (2000) also demonstrated that
Silurian trilobite α diversities across the shelf were comparable to those
of the Ordovician despite the marked drop in global clade diversity
during the end-Ordovician extinction event; emphasising the decou-
pling between different levels in the hierarchy of biodiversity metrics.
Ecological change through the GOBE cannot be numerically assessed in
the same way as taxonomic data. To date many ecological analyses have
 Author's personal copy
106 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
been both qualitative and subjective. A more structured and testable
approach to ecological change involves the analysis of changes in
palaeocological architecture during the event. The hierarchical ecological
framework, developed by Droser et al. (1997) and Bottjer et al. (2001),
helps to elucidate the magnitude of both diversifications and extinctions;
both can commonly be decoupled from taxonomic fluctuations and
provide more causal and fundamental models for diversity changes in
deep time. Droser et al. (1997) established four palaeoecological levels
together with their defining characteristics. Although it could be argued
that the possible colonization of the land by tracheophytes and the deep
sea by benthos are first level changes, the majority of ecological changes
took place at the second, third and fourth levels. At the second level, new
Bambachian megaguilds (Bambach, 1983) evolved. Bambach (1983)
introduced ‘adaptive strategies’ as a means of evaluating palaeoecological
changes through time and ‘megaguilds’ to decipher large-scale shifts in
the nature of ecospace utilisation. The Ordovician displays major changes
in its palaeoecological communities, from dominance by trilobites to that
by brachiopods, reflected also in changes in the composition of shell
concentrations, together with the establishment of Palaeozoic hard-
grounds permitting bioerosion and encrusting strategies and the
development of cryptic communities (see also Wilson and Palmer,
1992, 2002). Third level changes included the appearance of new
community types, particularly in deeper water and associated with
carbonate build-ups, an escalation in tiering complexity, both above and
below the substrate, and the further filling of already existing Bambachian
megaguilds to approach maxima for the Palaeozoic. Numerous, more
minor, changes took place at the fourth level with the appearance and loss
of communities.
3.3. Ecospace utilisation, competition and community saturation
An important aspect of understanding the palaeoecological context
of the GOBE is the analysis of ecospace utilisation (e.g. Bambach, 1983;
Bambach et al., 2002, 2007). Late Ordovician skeletal organisms show a
marked increase in the occupation of ecospace compared to those of the
Early and Middle Cambrian. They occupied 30 of the 216 theoretically
possible modes of life depicted in the 6 × 6 × 6 grid developed by
Bambach et al. (2007) and Bush et al. (2007) to show the basic
categories of topographical location relative to the sea floor, motility and
feeding mechanism. Early and middle Cambrian organisms with
mineralised skeletons occupied 19 modes of life on this scheme
(Bambach et al., 2007). The addition of information from Konservat-
Lagerstätten increased the Cambrian figure to 30 and it is reasonable to
assume that were such data available for the Late Ordovician, the
inclusion of soft-bodied organisms would significantly increase the
figure for that time interval. The main increases in ecospace utilisation
by skeletal organisms in the Late Ordovician are those identified in
earlier, qualitative, studies namely the range of surficial feeders, the
greater tiering both above and below the sediment–water interface and
the distribution of predators. Similar analyses of the occupation of
ecospace are needed for the Late Cambrian, Early and Middle Ordovician
to determine when the changes took place. Recent organisms with
regularly preserved hard parts occupy 62 fields in the ‘ecospace cube’
(Bambach et al., 2007, Fig 7B); a doubling of the late Ordovician figure.
As Bambach et al. (2007, p. 4) have stressed, the concept of ecospace
does not include any assessment of competition between organisms
within or between particular modes of life or of other limiting factors on
the distribution of species. The question then arises as to whether the
Great Ordovician Biodiversification Event was simply a passive filling
of ecospace or whether competition determined the structure of the
increasingly diverse communities. In demonstrating a considerable
constancy of trilobite α diversities in different shelf settings through the
Ordovician, Westrop and Adrain (1998a) argued that this indicates that
direct interactions between the members of the Evolutionary Faunas
were very limited and that the radiations were largely non-competitive.
For the brachiopods, however, various authors (e.g. Waisfeld et al., 2003)
recognized guild structures indicating competition between taxa for
particular resources (Bambach, 1983).
The expansion of predators both in terms of taxonomic groups and,
for skeletal groups at least, ecospace occupation (Bambach et al., 2007)
during the GOBE probably had a significant effect on the overall ecology
of the marine realm. Kröger and Landing (this issue) attribute major
changes in the disparity, body size and biomass of cephalopods in the
Early Ordovician of northeastern Laurentia to an escalation amongst
predators and thus community evolution which they consider a major
driver of the early phases of the Great Ordovician Biodiversification
Event. Bush et al. (2007) considered that the increase in predation from
the middle Palaeozoic to the late Cenozoic may have had a major effect
on suspension feeders by driving trends towards increased motility and
greater infaunalisation.
The effect of the GOBE was to create marine communities that were
markedly different from those of the Cambrian in terms of both their
ecological architecture and composition. Although the resultant overall
ecological patterns were very stable, Patzkowsky and Holland (2003)
have demonstrated that Late Ordovician communities in Laurentia were
not ‘saturated’ with species. They showed that local diversities were
strongly correlated with regional ones and thus there was a much stronger
influence of regional oceanographic controls on species numbers than
local inter-species competition. The latter should lead to some sort of
equilibrium, an upper limit on diversity. On a regional scale, Owen (2007)
documented a peak in the diversity of trilobites in Avalonia in the late
Katian, against the global trend. It was largely the result of immigration of
taxa from lower latitudes and not a burst of evolutionary originations.
Although the late Katian marked the first appearance, locally, of pure
carbonate depositional settings in Avalonia, much of this immigration was
into shelf environments that were long established and thus the
immigrants must have been able to insinuate themselves into existing
communities. Armstrong and Owen (2002a, b) recognized a similar
process in the conodont faunas. The process of insinuation rather than
displacement of existing taxa indicates that the ‘host’ communities were
not saturated. Patzkowsky and Holland (2003) raised the pertinent
question as to whether communities were ever saturated with species
during the remainder of the Palaeozoic. They considered the Late
Ordovician as the start of the Palaeozoic plateau of diversity in the
diversity curves of Sepkoski (2002 and references therein) and therefore
argued that unsaturated communities did not support Sepkoski's (1979)
coupled logistic model for global biodiversity change but favoured the
hierarchical niche model of Walker and Valentine (1984).
3.4. α, ß, γ diversity
The concepts of α (intra-community), ß (inter-community) and γ
(inter-provincial) diversity have been widely used in biological studies
since the early 1970s (see Whittaker, 1972; Owen and Crame, 2002). The
components of biodiversity can be deconstructed into increases within
individual communities, those apparent between adjacent communities
and those demonstrable between adjacent provinces. Sepkoski (1988)
applied these concepts to the GOBE, noting that only about 50% of the
increased diversity during the event could be accounted for by α and ß
diversity; the remaining increase was probably hidden in γ diversity and
possibly associated with hardground and reef communities, themselves
forming important aspects of ecological change during the event. Many
elements of the diversity increase can be modelled in terms of these three
overlapping components that can both elegantly explain some of the
increase and frame testable models (Harper, 2006). In many groups such
as the brachiopods (Harper, 2010) increases in diversity proceeded a pace
with the high levels of provincialism, generated by the dispersal of the
major continents and a high frequency of microcontinental fragments and
volcanic arcs, providing γ diversity (Harper and Mac Niocaill, 2002;
Harper et al., 2009). The migration of taxa into deep water and the
occupation of carbonate build-ups and reefs provided many possibilities
for new community types and ß diversity (e.g. Sepkoski and Sheehan,
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
1983; Bassett et al., 2002), whereas new guilds and the filling of
established guilds created new niches in existing community structures
and consequently increases in α diversity (Waisfeld et al., 2003; Harper,
2006).
4. A plankton revolution in the Ordovician?
4.1. Evolution of the phytoplankton
In modern oceans, the abundance of fish and marine mammals,
occupying the higher levels of the trophic chain, depends greatly on the
presence of bloom-forming phytoplankton. Food chains are very
complex, but the link between the phytoplankton at the base of the
food web, and sharks and whales, for example, on the top of the food
chain, are relatively short, with diverse metazoan zooplankton groups
being the intermediates.
Three major groups build up the modern-day marine phytoplankton.
All three groups diversified during the Early Mesozoic radiation: (1) the
organic-walled dinoflagellates first appeared during the Early Triassic; (2)
the calcareous nannoplankton (coccolithophores) originated in the Late
Triassic; and (3) the diatoms, representing the major part of the silicious
phytoplankton, emerged during the Early Cretaceous (e.g. Falkowski et al.,
2004). The presence of calcareous nannoplankton in the Palaeozoic is still
debated: some calcitarchs (Versteegh et al., 2009) may possibly be the
ancestors of calcareous microplankton (Munnecke and Servais, 2008),
including the calcareous dinoflagellates (Servais et al., 2009b). Diatoms
have no known Palaeozoic equivalent so far. The only elements in the
fossil record of Palaeozoic (and Precambrian) phytoplankton are therefore
the acritarchs, a group of organic-walled microfossils of unknown, but
most probably varied, biological affinities (Evitt, 1963).
Although acritarchs are, by definition, of unknown biological affinity,
many of the acritarchs from the Precambrian onwards, with size ranges
below 100 µm, are considered to be the cysts of diverse marine algae,
although other morphotypes may not be marine, nor part of the
microplankton (see Servais et al., 1997, for a review). Dinoflagellates
possibly first appeared in the Precambrian, as suggested by comparative
ultrastructure, molecular biology and biogeochemical studies, but
microfossils displaying all morphological parameters of modern
dinoflagellate cysts only appear in the Mesozoic (Fensome et al.,
1996). Nevertheless, many Palaeozoic palynologists now agree that
most acritarchs probably represent the cysts of marine phytoplankton,
including the ancestors of the dinoflagellates, because acritarchs not
only have a very similar size and morphology (Servais et al., 2004b), but
also have similar palaeoecologial and palaeogeographical distribution
patterns to those of modern dinoflagellates (e.g. Servais et al., 2003; Li
et al., 2004; Vecoli and Le Hérissé, 2004).
Diversification curves for the acritarchs have been presented since the
early 1970s, when Tappan and Loeblich (1973) reviewed the acritarch
literature and published a first ‘phytoplankton’ curve. This showed an
increase of acritarch species diversity in the Cambrian and a peak in the
Ordovician. The dataset of Downie (1984), also used by Strother (1996)
and Servais et al. (2004a), confirmed the increase of acritarch diversity
since the Late Precambrian with a peak in the Ordovician. Subsequently,
several authors (e.g. Katz et al., 2004, Fig. 3; Martin et al., 2008; Strother,
2008) used a curve that was based on unpublished data (and personal
communication by R.A. MacRae). This curve, based on the ‘Palynodata’
database (Fensome et al., 1996), used the information from the ‘acritarch
catalogue’ of Fensome et al. (1990a,b), building on a dataset of a
consortium of several oil companies.
A more recent re-evaluation of Palaeozoic acritarch diversity was
presented by Mullins et al. (2006) in the frame of the ‘PhytoPal’ project,
from which Servais et al. (2008) showed that the phytoplankton
diversity continuously increased from the Precambrian to reach the
highest levels of the Palaeozoic during the Middle Ordovician. Kröger
et al. (2009) have drawn a curve based on these data of Servais et al.
107
(2008), illustrating the evolution of the global diversity of the
Ordovician acritarchs (see Fig. 4 herein).
Katz et al. (2004) noted that there is an empirical relationship between
long-term sea-level change and diversities in the three modern-day
phytoplankton groups, in particular the calcareous nannoplankton and
the dinoflagellates. In addition, Katz et al. (2004) indicated that marine
invertebrate faunal diversity is highly correlated with phytoplankton
diversity and community composition. For example, diatom genera
richness is highly correlated with Cenozoic invertebrate richness, and the
combined diversity of calcareous nannoplankton and dinoflagellates is
strongly correlated with Mesozoic invertebrate diversity (see also Stover
et al., 1996). For the Palaeozoic, Katz et al. (2004, Fig. 6) noted only a weak
correlation between the diversity of marine invertebrates and the
diversity of the acritarchs. Servais et al. (2008), however, considered
that phytoplankton diversity should probably correlate strongly with that
of the marine invertebrates, but that complete datasets for Ordovician
acritarchs are still not available for most parts of the world. Servais et al.
(2008) tentatively correlated the diversity of acritarchs with that of
several groups of the zooplankton (see also Fig. 4). While some of these
correlations remain hypothetical, the complete dataset of Ordovician
acritarchs from the palaeocontinent Baltica by Hints et al. (this issue)
allows a more obvious correlation between the diversity of the acritarchs
and that of all marine invertebrates, as calculated by Hammer (2003). It
can therefore be assumed, that similar to the Mesozoic, the diversification
and abundance of phytoplankton was most probably linked to the
increase of the sea levels and the extension of flooded areas, and that the
presence of abundant phytoplankton possibly had a major impact on the
diversification of the zooplankton and the suspension feeders that feed on
the phytoplankton.
A major problem with the phytoplankton fossil record is the fact that
palaeopalynologists usually only analyze the larger fraction (N20 µm) of
the organic-walled microfossils. The fossil record thus only includes
relatively large cysts, although much smaller acritarchs are present in
most assemblages. Munnecke and Servais (1996), for example, recorded
numerous very small acritarchs from the Silurian of Gotland, often less
than 5 µm in diameter. This smaller fraction of a few µm is usually
attributed to the picoplankton. An even smaller fraction, attributed in
modern oceans to the bacterioplankton, is not present at all in the fossil
record, although it is highly probable that it was also present in the Early
Palaeozoic oceans. The picoplankton and the bacterioplankton repre-
sent a major part of the primary production in modern oceans and both
constitute a major part of the food of many invertebrate organisms,
including the zooplankton and the suspension feeders. Future studies,
both palaeontological (palaeopalynological) and biogeochemical, are
needed to analyze the fossil record of the picoplankton and the
bacterioplankton in the Palaeozoic oceans in order to understand their
impact on the marine trophic web and also the primary production and
the available biomass that also includes the cyanobacteria and other
organisms.
Although it is evident that the biodiversity of phytoplankton increased
significantly during the Ordovician, it is difficult to establish a relationship
between an increase in primary production and biodiversity. The
important palaeoecological changes require a large increase of primary
producers, in particular to trigger the abundance and size increases of
most organisms, such as the brachiopods, cephalopods and corals. Some
animal groups most probably specialized in feeding on specific groups of
primary producers, including the phyto-, pico- and bacterioplankton.
4.2. Evolution of the zooplankton
Although some planktonic, and even nektonic organisms are known
from the Cambrian (Butterfield, 1997, 2001) and although most animal
phyla were present since the ‘Cambrian explosion’, marine life was mostly
limited to that of marine invertebrates near the sea floor, in shallow-water
environments. Some of the major planktonic groups were absent in the
Cambrian and only originated in the Ordovician which may be considered
 108
Author's personal copy

T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119

Fig. 4. Changes in the diversity of the major fossil groups in the water column (plankton, mobile benthos and predators) during the GOBE. Phytoplankton: acritarchs (redrawn from Kröger et al., 2009, based on Servais et al., 2008);
zooplankton: chitinozoans (redrawn from Paris et al., 2004), graptolites (redrawn from Cooper et al., 2004), radiolarians (redrawn from Noble and Danelian, 2004); mobile benthos: ostracodes (redrawn from Braddy et al., 2004), trilobites
(redrawn from Adrain et al., 2004); pelagic predators: orthoceritid and lithuitid cephalopods (redrawn from Kröger et al., 2009).
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
as a ‘revolution in the oceanic trophic chain’ or an ‘Ordovician plankton
revolution’ (Servais et al., 2008). Interpretations of the patterns of
diversity and the proportional abundance of demersal organisms,
plankton and nekton in the Palaeozoic oceans clearly indicate the
‘plankton revolution’ in the Ordovician and the ‘nekton revolution’ in
the Devonian (Klug et al., in press, Fig. 1).
One of the major planktonic groups, the graptoloids, evolved in the
Early Ordovician. Although graptolites appear in the Middle Cambrian,
planktonic forms only radiated in the Ordovician (Chen et al., 2006) and
occupied different pelagic zones. The graptolites show increasing
diversity in the Early and Middle Ordovician (Cooper et al., 2004),
with a peak in the Darriwilian, similar to that of the acritarchs (Servais
et al., 2008, Fig. 2; Fig. 4).
Another group with a clear planktonic distribution is the chitinozoans.
They are interpreted to be possibly the eggs of a planktonic organism, but
the ‘chitinozoan animal’ still remains unknown. Their distribution and
diversification are well known. The group first appears in the Tremadocian
and diversified rapidly on the three major palaeocontinents, Gondwana,
Baltica and Laurentia (Paris et al., 2004; Achab and Paris, 2007). The global
Ordovician diversity trends of the chitinozoans partly mirror those of the
acritarchs (Fig. 4), but clearer correlations can be seen at the regional level,
in particular in the dataset from ‘northern Gondwana’ (southern Europe
and North Africa), as documented by Vecoli and Le Hérissé (2004), but a
correlation is also observed on Baltica (Hints et al., this issue). As with the
graptolites, the food and the feeding behaviour of the ‘chitinozoan animal’
are not clearly understood. However, it can be suggested that these
organisms fed on the smaller parts of the phytoplankton, possibly also on
the picoplankton (b5 µm).
Two of the major microfossil groups occupying the pelagic realm of
modern oceans are the radiolarians and the foraminifers. In modern-
day marine environments, radiolarians represent a major part of the
smaller zooplankton. Some of them have dinoflagellate (and/or
foraminifer) symbionts, while others feed on the phytoplankton
(and/or the foraminifers). Records of foraminiferans from the Lower
Palaeozoic are lacking or problematical but the radiolarian fossil
record indicates that few taxa are present in the Cambrian and a
diversification took place during the Ordovician (Noble and Danelian,
2004). The diversity curves, although preliminary, show similar
trends compared to the graptolites and also reflect the peak of the
phytoplankton in the Darriwilian (Fig. 4).
A major part of the modern-day zooplankton is constituted by
small crustaceans, which form the major food of fishes. In modern
oceans, one of the major groups of smaller crustaceans is the
copepods, which appeared in the Miocene. The caryocaridid arthro-
pods (phyllocarids) may constitute a possible Early Palaeozoic
equivalent, representing a major zooplankton group and an important
zooplanktonic component in Ordovician ecosystems, where they
developed and radiated (Vannier et al., 2003; Whittle et al., 2007).
Possibly some of the ostracodes, that rapidly developed during the
Ordovician (e.g. Braddy et al., 2004) also constituted a part of the
zooplankton. The diversity of this group also increased continuously
during the Ordovician (Fig. 4). Most of the Ordovician ostracodes can
probably be attributed to the mobile benthos, the earliest known
pelagic ostracodes are only recorded in the Silurian (Perrier et al.,
2008).
The information from various fossil groups thus clearly indicates that
a major change in the trophic structure of the Ordovician oceans took
place with the rise of the zooplankton, confirming the observation of
Sepkoski in the 1980s.
4.3. The development of planktotrophy
A census of the larval stages of marine invertebrates was not directly
present in the diversity curves of Sepkoski's (2002) compendium or in
the dataset of the Paleobiology Database (e.g. Alroy et al., 2008).
However, the larvae of many benthic organisms today represent an
109
important component of the plankton. The evolution of larval
development probably dates back to the late Precambrian. Signor and
Vermeij (1994) had already assumed that planktonic feeding larvae
arose during the Late Cambrian–Early Ordovician and that plankto-
trophy was an escape strategy from an increasing predation pressure
resulting from the appearance of benthic suspension feeders. With an
increasing diversity and abundance of phytoplankton in the water
column, not only the zooplankton developed, but also the suspension
feeders that filtered the phytoplankton (or picoplankton) out of the
water column at increasing heights above the sea substrate (develop-
ment of tiering, see below). The development of larval stages and
planktotrophy would not only allow the escape from shallow-water
levels where predation was high, but also enable a wider distribution of
taxa, with larval stages being able to be transported over longer
distances, possibly even crossing oceans to settle on other continents.
Nützel and Frýda (2003) and Nützel et al. (2006) found the first
evidence for the origin of planktotrophy in gastropods at the Cambrian–
Ordovician transition, confirming previous assumptions by Chaffee and
Lindberg (1986) and Signor and Vermeij (1994). Nützel et al. (2006) also
considered that planktotrophy may have played a role as an escape
strategy from the benthos, and they indicated that the expansion of the
suspension feeders and the development of planktotrophy were
most probably based on a higher nutrient supply and an increased
palaeoproductivity.
Direct evidence for planktotrophy seems to be absent from the
Cambrian (e.g. Runnegar, 2007) and recent molecular clock data seem
to corroborate the assumption that it developed only during the late
Cambrian. Peterson (2005) plotted the different types of plankto-
trophic metazoan larvae on a phylogenetic tree and noted that
planktotrophy has evolved several times between the latest Cambrian
and the Middle Ordovician.
Several major developments thus took place during the interval
between the late Cambrian and the Middle Ordovician. Based on the
available phytoplankton, planktotrophy developed in several inver-
tebrate clades and an increasing number of marine invertebrates
probably developed planktonic larval stages. The development in the
Late Cambrian and Early Ordovician of the new feeding strategies
would thus have had a major impact on the trophic chains.
5. The invasion of the pelagic realm
5.1. Development of motility
The successively increasing complexity of the pelagic food chain is
indicated by the development of a diverse fauna of pelagic Ordovician
vertebrates, molluscs and arthropods. Based on the increase of the
available phytoplankton, it seems reasonable to assume that zoo-
plankton developed during the Ordovician and constituted the
foundation for freely swimming animal groups that became predators
in the open seas of the pelagic realm.
Very rare in pre-Ordovician assemblages (Schoenemann et al.,
2008), pelagic trilobites developed during the Ordovician and fairly
diverse mesopelagic and epipelagic trilobite communities were
established by the Tremadocian (e.g. Adrain et al., 2004; Racheboeuf
et al., 2009). As is noted above, another group of arthropods, the
caryocaridids, is also considered to have occupied the pelagic realm
from the earliest Ordovician and to have diversified significantly
during the Middle Ordovician (Vannier et al., 2003; Whittle et al.,
2007).
With cephalopods as likely predators of these pelagic arthropods the
Ordovician can be considered as the time of the initial establishment and
diversification of widespread pelagic ecosystems with large macro-
predators. However, little or no evidence for the occurrence of fish in the
open seas is known today and cephalopods appear to have been the
largest animals in the open seas of the Ordovician.
 Author's personal copy
110 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
5.2. First fishes
Apart from the feeding apparatuses of conodonts, some of which
belonged to taxa with distributions indicating a pelagic mode of life
(Rasmussen 1998; Armstrong and Owen, 2002a), the fossil record of
Ordovician vertebrates is scattered, and its incompleteness leaves
many open questions. Nonetheless, large-scale patterns of Ordovician
vertebrate evolution and palaeobiogeography are becoming clearer
for the earliest skeletal vertebrates (e.g. Blieck and Turner, 2003;
Turner et al., 2004). The earliest dermoskeletal vertebrate remains are
known from arandaspids in Floian siliciclastic sediments of Australia
(see review in Young, 2009; Davies and Sansom, 2009). Because Early
Ordovician occurrences are restricted to Gondwana, an ‘out of
Gondwana’ scenario for the evolution of these early vertebrates was
suggested by Elliott et al. (1991). Smith et al. (2002) proposed an
alternative ‘out of Laurentia’ model. Conversely, Young (2009)
contended that the poorly understood fossil record of the earliest
vertebrates means that such scenarios are very provisional. He argued
that a primitive cosmopolitanism at higher taxonomic levels contrasts
with a marked endemism at genus, and species level. The endemism,
which is one of the prominent features of the Ordovician vertebrate
record (Blieck and Turner, 2003), can be explained by the very
shallow marine habitats of Ordovician vertebrates (Sansom and
Smith, 2001; Allulee and Holland, 2005; Davies and Sansom, 2009).
The marked endemism contrasts with global patterns of Ordovician
vertebrate evolutionary and diversity trends. Arandaspids and an array
of other primitive vertebrates diversified strongly worldwide during the
Darriwilian–Katian and Turner et al. (2004, p. 334) considered that all
major vertebrate clades seem to be present by the Caradoc. The
tendency of a strongly increasing number and diversity of armoured
vertebrates indicates that such armour against predators played a major
role in the diversification process.
5.3. Swimming eurypterids
The appearance and diversification of swimming eurypterids early in
the Late Ordovician roughly corresponds to the first radiation of armoured
vertebrates (Braddy et al., 2004; Tetlie, 2007; Tetlie and Cuggy, 2007;
Tollerton, 2003). Ordovician eurypterids occur most commonly in near
shore marine deposits (Tetlie, 2007) and thus their habitats overlap with
those of early vertebrates. Trophic interactions between eurypterids and
Ordovician fishes may therefore have taken place. Additionally, co-
occurrences of cephalopods in Ordovician fish localities (e.g. Sansom et al.,
2009) may indicate trophic interactions between fish and nautiloids.
5.4. Evolution of the cephalopods
Based on actualistic comparisons, nautiloids seem to have been
carnivores from the beginning, but direct evidence of Ordovician
buccal apparatuses is lacking. Large slit-like shell injuries in
Ordovician cephalopod shells (Kröger, 2004) and fossil assemblages
(Frey, 1989) suggest a durophagous habit for at least some nautiloids.
Changes in the fossil assemblages from the Beekmantown Group, New
York, USA, show evidence of increasing competition and levels of
predation in the mollusc faunas since the Early Ordovician (Kröger
and Landing, 2007, 2008, 2009, this issue). Kröger and Landing (this
issue) argued that an increasing competition and habitat differenti-
ation may not have been restricted to Lower Ordovician tropical
shallow-water environments, but potentially were an important
aspect of the Ordovician diversification in general, including the
successive invasion of the open water by larger swimming animals.
Nautiloids were restricted to the palaeotropical carbonate plat-
forms during the earliest Ordovician, but soon expanded their habitats
and occur in distal, deep-water sediments in the middle Tremadocian
(Kröger et al., 2009). The pelagic cephalopod diversification peaked
during the Darriwilian (Frey et al., 2004; Kröger and Zhang, 2009;
Kröger et al., 2009). Kröger et al. (2009) argued that the timing and
regional pattern of the occupation of the pelagic realm by cephalo-
pods partially contradicts the idea of an invasion by simply an escape
from benthic competition and predation (Signor and Vermeij, 1994).
They suggested that possibly an increasing complexity and sustain-
ability of the Ordovician pelagic food chain itself was a major factor in
driving the diversification.
The foci of the Ordovician cephalopod diversification, however, were
in the shallow-water carbonate platforms of the palaeotropics. There,
cephalopod faunas changed drastically over the epoch and reached a
diversity climax during the Katian (Fig. 4). Additionally, Kröger et al.
(2009) demonstrated that the diversification of some of the most
diverse Ordovician cephalopod groups paralleled the diversification of
reef builders and their associated fauna.
5.5. Radiation of trilobites into the water column
Trilobites are one of the key components of the Cambrian
Evolutionary Fauna and thus one of the main elements of the mobile
benthos prior to the Ordovician. Setting aside the agnostids, whose
mode of life has been greatly debated (see review by Fortey and
Owens, 1999, pp. 455–458) but which might have been planktic in at
least some taxa, few trilobites lived in the water column during the
Cambrian (see Fortey and Rushton, 2007; Schoenemann et al., 2008).
A substantial number of trilobite clades diversified during the Great
Ordovician Biodiversification Event along similar trajectories to those
of other invertebrate groups that typify the Paleozoic Evolutionary
Fauna (Adrain et al., 2004). As a result of this diversification, trilobites
reached a peak of morphological disparity during the Ordovician
(Foote, 1991; Fortey and Owens, 1999). Fortey and Owens (1990,
1999) identified sets of recurring morphotypes that they linked to the
autecology of the trilobites concerned (see also Adrain et al., 2004).
The vast majority of species were members of the benthos but wide
palaeogeographical distributions (e.g. Servais et al., 2005), the
development of extremely large eyes and, in some instances, a highly
streamlined shape, indicate a pelagic mode of life in some Ordovician
clades (Fortey and Owens, 1987, 1999). These comprise both
epipelagic taxa, typified by the Telephinidae, and mesopelagic groups
especially the Cyclopygidae and related families (see Adrain et al.,
2004; Owen and Bruton, 2008). Such groups appeared in the
Tremadocian and reached a peak of generic diversity in the
Darriwilian. The epipelagic trilobites included the most widespread
of all trilobite species (McCormick and Fortey, 1999) and many of the
cyclopyid trilobites were both widespread and morphologically very
conservative over long periods (Adrain et al., 2004). Despite this
success, none of the pelagic clades survived into the Silurian and the
water column was never occupied by post-larval trilobites again.
Chatterton and Speyer (1989) showed that not only did trilobite
groups with pelagic modes of life in the holaspis stage become extinct
by the end of the Ordovician but so did the majority of those benthic
taxa that had entirely planktic protaspides. These too began to decline
after the Darriwilian and Chatterton and Speyer (1989) suggested that
there was a link between susceptibility to extinction in the Late
Ordovician and dependence on planktic food. They considered that
the start in the decline in diversity of pelagic trilobites and taxa with
wholly planktic larval stages coincided with the onset of the Late
Ordovician glaciation. While evidence for the latter was at the time
limited to reports of glacigenic rocks older than the unequivocal
Hirnantian deposits (Hambrey and Harland, 1981), their comments
are remarkably prescient both in the light of the more recent isotopic
evidence for a much earlier initiation of global cooling (e.g. Trotter
et al., 2008; Young et al., 2008) and also in the light of the new
diversity curves for the phytoplankton and zooplankton.
Fortey and Owens (1999, p. 453) suggested that not only did
microplankton form a food source for pelagic trilobites but some taxa
may have fed on smaller plankton feeders, such as phyllocarid
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
crustaceans. The occupation of pelagic environments by trilobites
undoubtedly can be seen as part of the ‘revolution in the oceanic
trophic chain’ (Servais et al., 2008).
6. “The booming of suspension feeders”—transition in the benthos
6.1. Evolution of tiering
Tiering strategies were essential to develop the complex community
structures of the Paleozoic Evolutionary Fauna (Ausich and Bottjer, 1982;
Bottjer and Ausich 1986). Tiering took place below the sediment–water
interface but during the GOBE changes above the substrate were most
marked. The basal levels (0–5 cm) were occupied by sponges, corals,
bryozoans, rhynchonelliformean brachiopods, bivalves and both diplo-
poran and rhombiferan echinoderms. The higher levels (5–20 cm) were
characterized by sponges, corals, bryozoans, and diploporan echino-
derms, together with crinoids. Whereas the levels at 20–50 cm were
populated by diploporan echinoderms and crinoids, the highest tiers at
50–100 cm only hosted crinoids. Secondary tiering was also important
with for example brachiopods attaching to bryozoans (Harper and
Pickerill, 1997) and crinoids (Sandy, 1996). This tiering structure
remained more or less stable until the end of the Cretaceous (Ausich
and Bottjer, 1982). By contrast there was a much more limited escalation
in the depth and complexity of infaunal tiering during the GOBE perhaps
reflecting the lack of intense predation pressure (Bottjer and Ausich,
1986). However, as indicated by Sheehan and Schiefelbein (1984), the
appearance of Thalassinoides galleries during the GOBE was a major
event. Late Ordovician Thalassinoides galleries penetrated 2 m into the
sediment, while prior to the GOBE Thalassinoides-like burrows were
confined horizontally. Trace fossil data indicate bioturbators had reached
the deep sea during the event (see below). An obvious interpretation of
the development of epifaunal tiering could be the increasing presence of
highly diverse phytoplankton (but also bacterioplankton and picoplank-
ton) in the water column, that allowed the different suspension feeding
organisms to filter their food out of the water in increasing heights.
6.2. Evolution of benthic suspension feeders
A key characteristic of the GOBE was the marked diversification of
the suspension feeding benthos, extensively documented across a
range of animal phyla. During the GOBE a complex, tiered network of
suspension feeders was established for the first time and dramatically
altered not only the composition and structure of many sea floors but
also trophic feeding webs and possibly hydrographic conditions at the
substrate–water interface. Nevertheless, different fossil groups show
different diversity patterns and trends, indicating the absence of a
single external driver. For some groups, explosive radiations at several
intervals were the norm, and for others, a more progressive and
steady increase in diversity was more typical.
The evolution of benthic suspension feeders is particularly well
documented by the brachiopod fossil record. Brachiopod diversifica-
tion was stepwise but sustained during the GOBE. The global curve for
the rhynchonelliformeans (Harper et al., 2004; Fig. 5) indicates four
phases of radiation, during the late Tremadocian, the Darriwilian, the
Sandbian and the Katian. The initial radiation was associated with
holdovers from the Cambrian and early Tremadocian, commonly
occupying inshore carbonate environments, whereas the second,
Darriwilian, pulse was associated with the establishment of the
Ordovician brachiopod fauna dominated by orthides and strophome-
nides. Deeper-water taxa, associated with offshore migrations,
contributed to the third phase while the fourth phase is linked to
diversifications in warm, carbonate environments. During the GOBE
brachiopods generally increased in size, developed a complex set of
onshore–offshore community spectra and established a range of
morphotypes that, with the exception of a few aberrant Permian
forms, typified the subsequent history of the phylum. They dominated
111
benthic communities both above and partly within the sea bed. They
also provided a substrate for attaching and encrusting epifauna,
providing support, protection and a source of food through inhalant
and exhalent currents. Few detailed bed-by-bed studies of brachiopod
diversity through the event are available, although Rasmussen et al.
(2007) and Rasmussen and Harper (2008) have provided data for the
Darriwilian phase of the diversification in the East Baltic and Zhan
et al. (2006) have provided data from South China.
Another major group of lophophorate suspension feeders is the
Bryozoa, the fossil record of which is now known to extend back to the
Upper Cambrian (Landing et al., 2010). These colonial suspension feeders
are a major component of some Ordovician sea-bed communities, in
particular in the Upper Ordovician of the American mid-continent, where
they are present in rock-forming abundance (Taylor and Ernst, 2004),
commonly also forming mound-like structures. A major radiation of the
phylum occurred during the Ordovician, with the rapid appearance of
many Phanerozoic orders of marine bryozoans, principally within the
stenolaemates, rapidly occupying and in some cases dominating the low-
level filter-feeding benthos. The increase was exponential from the Floian
to the Katian, followed by a diversity decline in the Late Ordovician
(Taylor and Ernst, 2004; Fig. 5). During this interval a number of ecological
changes associated with the diversification of adaptive colony types,
together with the taphonomically favourable conditions of the Calcite
Seas enhanced the spectacular pattern of radiation through the middle
part of the Ordovician.
Although already well established in the Cambrian, sponges
diversified markedly during the Ordovician with continuous diversity
increases from the earliest part of the Period and a more pronounced
diversification in the Late Ordovician (Carrera and Rigby, 2004; Fig. 5). In
contrast to the Cambrian sponge faunas dominated by archaeocyathans
and thin-walled weakly fused spiculate demosponges and hexactinel-
lids, the Ordovician sponge fauna is more typically dominated by the
heavier, thick-walled demosponges in carbonate environments where-
as the siliceous hexactinellids were more prominent in siliclastic facies.
During the period, however, demosponges became less important as
rugose and tabulate corals and especially stromatoporoids began occupy
these types of benthic niches.
The stromatoporoids diversified only from the middle Darriwilian
onwards, forming some of the first extensive metazoan reef structures
prior to becoming much more diverse and widespread (cosmopolitan)
in the Silurian (Webby, 2004; Fig. 5).
The first appearance of the corals is still debated, although current
evidence suggests that the tabulates appeared first in the Early
Ordovician, with simple cerioid growth modes, lacking septa and mural
pores and possessing only occasional tabulae. The rugose corals
apparently appeared later during the Middle Ordovician developing
heavy, robust skeletons. These two major Palaeozoic groups, however, the
Rugosa and Tabulata, diversified rapidly during the Ordovician with an
‘explosion’ of diversity early in the Late Ordovician (Webby et al., 2004c).
The diversity patterns of the echinoderms differ markedly across
the different groups (classes). Following an important radiation in the
Cambrian (when the early members of the phylum were an important
component of the Cambrian Evolutionary Fauna) and a decline in the
late Cambrian, echinoderms diversified again significantly during the
Ordovician, to become one of the dominant faunal elements in
Palaeozoic benthic marine environments for the next 200 Ma (e.g.
Sprinkle and Guensburg, 2004; Lefebvre and Fatka, 2003; Rozhnov,
2007; Nardin et al., 2009; Nardin and Lefebvre, this issue). In
particular stalked crinoids dominated the Ordovician echinoderm
fauna, generating widespread, crinoidal limestones commonly in
shallow-water environments. Blastozoans, including the cystoids,
diploporites, and rhombiferans often formed higher tiers but were
generally less abundant than the crinoids. Their biodiversification has
recently been analyzed in detail (Nardin and Lefebvre, this issue;
Fig. 5) and shows a continuous increase of diversity to a peak in the
Sandbian.
 112
Author's personal copy

T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119

Fig. 5. Changes in the diversity of the major benthic fossil groups during the GOBE. Sessile benthic suspension feeders: stromatoporoids (redrawn from Webby, 2004), bryozoans (redrawn from Taylor and Ernst, 2004), blastozoan
echinoderms (Nardin and Lefebvre, this issue), sponges (redrawn from Carrera and Rigby, 2004), brachiopods (redrawn from Harper et al., 2004); detritus and suspension feeders: bivalves (redrawn from Cope, 2004) and gastropods
(redrawn from Frýda and Rohr, 2004).
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
A number of molluscan groups, for example the Pteriomorphia,
contain suspension feeding taxa, but to date their trends have not
been isolated from those documented in these molluscan groups as a
whole. The radiation of the bivalve and rostroconch molluscs is not
presently understood in the same detail as for example the
brachiopods, with peaks in the diversity curves resulting mostly
from important monographic bursts. Nevertheless, despite this
potential bias, the diversification of the bivalves clearly also follows
the general increasing trend of the Ordovician biodiversification
(Cope and Babin, 1999; Cope, 2004) but with peaks in the Floian,
Darriwilian and Katian (Fig. 5). In general terms the Pteriomorphia
together with the Anomalodesmata (Cope, 2004, Fig. 20.6) markedly
increased in diversity through the Sandbian and Katian interval,
marking their migration to low-latitude carbonate platforms.
Frýda and Rohr (2004) summarized the Ordovician diversification
of the gastropods, which is as complex as that of the brachiopods with
different diversity trends for the different gastropod groups. However,
a clear increase from the basal part of the Ordovician up to the middle
Late Ordovician is also observed in this clade (Novack-Gottshall and
Miller, 2003a, 2003b; Frýda and Rohr, 2004), punctuated by
origination events in the Floian, early Darriwilian and early Sandbian
(Fig. 5). The group is dominated by detritus feeders and statistically
forms a part of Sepkoski's Modern Evolutionary Fauna although it was
an intimate part of many shallow-water, suspension-dominated
communities during the Ordovician.
6.3. Colonization of the deep sea floor
In his review of Cambrian to Carboniferous deep marine trace fossils,
Orr (2001) identified the Late Cambrian to Early Ordovician as a crucial
interval in the evolution of communities in that environment. He
attributed the major restructuring of deep marine communities to the
offshore displacement of ichnotaxa resulting from increased competi-
tion in shallow marine environments (see also Droser and Bottjer, 1993;
Mángano and Droser, 2004). Most recently, Buatois et al. (2009) have
documented an upper Tremadocian deep marine ichnofauna from
Argentina that is intermediate between the Ediacaran–Cambrian low
diversity matground faunas and the more diverse and morphologically
much more varied deep-water trace fossils that have been widely
documented from the subsequent Ordovician. The Argentinian fauna
includes traces indicating the trapping of microorganisms and bacterial
farming and thus indicates that the Agronomic Revolution (see Seilacher
and Pflüger, 1994) had started in the deep sea by or during the late
Tremadocian. Graphoglyptid traces, however, are much rarer and
geometrically simpler than later in the Ordovician.
7. Development of carbonate build-ups and reefs
The Ordovician is characterized by a profound change in reef
composition, with a shift from microbial-dominated reefs in the Early
and Middle Ordovician to metazoan-dominated reefs in the Late
Ordovician. The most comprehensive summary of the reef development
is provided by Webby (2002). In this review the Ordovician reefs are
subdivided in five constructional types: (a) true reefs, built in situ by
metazoans that produce a rigid framework, (b) biostromes, similar to
(a) but without significant topographic relief, (c) reef mounds, con-
structed by skeletal organisms but with very limited framework rigidity,
(d) carbonate mud mounds, composed mainly of micrites with sparse
skeletal organisms, and (e) microbial reefs, stromatolites and thrombo-
lites. Because of the high sea level at least 40% (possibly 60%) of the
continents were flooded (Walker et al., 2002), and these large epeiric seas
provided in many areas suitable habitats for reef growth. Similar to
modern reefs, most of the Ordovician reefs occur in palaeolatitudes
between 30°N and 30°S in shallow-water settings (Webby, 2002, 2004). It
is important to note that the exact stratigraphical position is somewhat
113
uncertain for many reefs because graptolites, the main stratigraphical
index fossils, are rare in reefal deposits.
The early Tremadocian reefs were small (mostly b1 m) and
consisted of stromatolites and thrombolites with abundant calcifying
cyanobacteria of the Epiphyton and Renalcis groups (Wood, 1999).
Associated metazoans are rare and consist of lithistid sponges and the
small tabulate coral Lichenaria (Webby, 2002; Carrera and Rigby,
2004; Webby et al., 2004c). Reefs of this type are reported from North
America and Argentina (Kiessling, 2002). In the late Tremadocian
reefs became larger, they occupied a greater range of environmental
settings, and the diversity especially of microbial reef-building biotas
increased. For the first time both Calathium (receptaculitalean) and
lithistid sponges became associated with the microbial reefs (Nitecki
et al., 2004). This trend continued into the Floian but with addition of
the first metazoan-dominated reefs consisting of up to metre-sized
skeletons of the probable stromatoporoid Pulchrilamina (Webby,
2002). In the Dapingian and early Darriwilian no significant changes
are observed in the composition of the microbial reefs although it
seems that the sponge–Calathium associations became more impor-
tant at the expanse of the microbial associations (Webby, 2002).
However, few new reef types were developed: large stromatactis-
bearing mud mounds are reported from North America, and mud
mounds as well as small bryozoan reefs from Russia (Kiessling, 2002).
A major turnover in reef composition is observed in the late
Darriwilian, and the newly developed reef types became the basis not
only for the Late Ordovician reef ecosystems but also for those of the
entire Silurian and Devonian. During the Middle and Late Ordovician,
thrombolites as well as calcifying cyanobacteria such as Epiphyton and
Renalcis disappeared, because these forms obviously lost their ability to
calcify (Webby, 2002; Riding, 2006b). Labechiid stromatoporoids,
solenoporacean algae, tabulate corals, and bryozoans start to play a
major role as frame-builders. Sponges and bryozoans developed a variety
of growth forms (Taylor and Ernst, 2004) and many reefs contain a
diverse epifauna of trilobites, brachiopods, bivalves, gastropods, and
ostracodes (Webby, 2002). Whereas stromatoporoid-dominated reefs
occupied shallow subtidal environments, bryozoan-dominated reefs
seem to favour somewhat deeper-water conditions. In the Sandbian,
reefs become even more diverse, and depending on the palaeo-water
depth different reef types are observed with stromatolitic and/or tube
worm-dominated intertidal reefs, patch reefs of different size in subtidal
environments consisting of corals, stromatoporoids, bryozoans, and
sponges (in varying amounts), and large bryozoan-dominated reefs in
shelf-edge environments (Webby, 2002). In the Katian, the metazoan
reefs became more widespread around the globe, with stromatoporoid-,
coral- and Solenopora-dominated reefs occurring for the first time on
Baltica, and pelmatozoan–bryozoan mud mounds on the peri-Gondwana
platform (Vennin et al., 1998). As indicated by Sheehan (1985) there was
a long interval between the Middle Cambrian (decrease in archaeocyathid
reefs) and the start of the GOBE lacking colonial animals with massive
skeletons, and thus an absence of colonial animals in reefs. The
development of colonial animals with massive skeletons during the
GOBE was therefore a major new lifestyle shared by several phyla.
As a result of the glacio-eustatic sea-level drop in the Hirnantian, gaps
are observed in shallow-water successions, and consequently reports of
reefs are rare. In the Hirnantian a major positive δ13C and δ18O excursion
is reported from several palaeocontinents (see review in Bergström et al.,
2009a). It is interesting to note, however, that in favourable settings reefs
occur even during intervals of maximum δ18O values (Long, 1993; Kaljo
et al., 2001, 2004; Ernst and Munnecke, 2009), which commonly are
interpreted as proxies for maximum cooling and/or volume of ice
shields. The latest Ordovician extinction event terminated reef growth
globally, and it took several million years before the first reefs
reappeared in the Silurian (Yue and Kershaw, 2004; Copper, 2001a,b).
The controlling factors of the Ordovician reef development,
especially for the major faunal turnover in the Darriwilian, are still a
matter of debate. Most likely a combination of several factors must be
 Author's personal copy
114 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
considered, e.g., tectonic setting and sea-bed topography, temperature,
salinity, climate conditions and weathering on land, CO2 and O2 content
of the atmosphere, ocean circulation, and sea-level development
(Munnecke et al., in press-b). Oolites, precipitating directly from sea
water, are more abundant in the Lower Ordovician (Wilkinson et al.,
1985) compared to the Middle and Upper Ordovician, which indicates
elevated calcium carbonate saturation states of the sea water. The high
abundance of microbial carbonates in the Lower and Middle Ordovician
might be the result of elevated water temperatures (see discussion in
Webby, 2002), which seems to be supported by new δ18O data (Trotter
et al., 2008). In contrast to metazoans which secrete calcium carbonate
intracellulary (biologically controlled), calcifying cyanobacteria secrete
calcium carbonate outside their cells (biologically induced by photo-
synthesis; Riding 2006b) in more or less direct contact to the ambient
sea water, and thus higher water temperatures and the resulting lower
CO2 content in sea water would favour cyanobacteria calcification (and
oolite precipitation). However, the reconstructed atmospheric CO2,
which was 14 to 18 times present day level (e.g., Berner, 1994), is
somewhat problematic because even the comparatively low increase of
pCO2 observed in modern oceans has strong effects on calcium
carbonate precipitating organisms especially those secreting aragonite
(Kleypas et al., 1999; Guinotte et al., 2006). So why didn't the Ordovician
carbonates dissolve? Ridgwell (2005) suggested that due to the lack of
pelagic calcareous plankton in the pre-Mid-Mesozoic oceans the sea
water in general reached much higher states of saturation with respect
to calcium carbonate.
8. Food and biomass
There has been relatively little exploration of the importance of the
impact of bioproduction on the radiations and extinctions of marine life,
although it appears evident that the diversification of the marine
biosphere is not only linked to the biogeochemical cycles (carbon,
phosphorus, etc.), but also to the availability of nutrients and the levels
of bioproductivity. Valentine (1971) attempted to understand the
diversity patterns in relation to the nutrient and food supply, while
Bambach (1993, 1999) started to analyze the changes in biomass and
productivity in the marine ecosystems. More recent papers on this topic
include those by Martin (1996, 2003) and Martin et al. (2008). It can be
assumed that increasing nutrient availability in the Early Palaeozoic
accelerated primary production (bacterio-, pico- and phytoplankton
biomass), and thus food for marine organisms. The observed increasing
diversity of the acritarchs (e.g. Servais et al., 2008) is not in itself a
measure of the abundance of phytoplankton in the ancient oceans,
however. It is very difficult to speculate about the quantity of
phytoplankton and of fresh dead organic matter in the oceans.
Geochemical proxies can be indirect evidence for the changes in the
sea-water chemistry and nutrient fluctuations (e.g. Saltzman, 2005).
In order to understand radiations and extinctions, and in particular
the Ordovician biodiversification and its possible links to the ‘Cambrian
explosion,’ future studies are needed to understand the relationship
between the nutrients in the water column and the development of the
marine trophic chains. Work within IGCP 410 and IGCP 503 has mostly
focused on the geological parameters (e.g. sea level, climate change,
sedimentology, geochemistry, etc.) of the Ordovician biodiversification.
To understand how diversity at the species and genus level ‘exploded’,
the palaeoecology of each fossil group must be better understood.
Feeding strategies must be more widely characterized and trophic
structures more wisely explored.
9. Future perspectives
The most major diversification of marine life, the Ordovician Radiation
or Great Ordovician Biodiversification Event (GOBE) warranted little
mention in either Darwin's ‘Origin of Species’ (in 1859) or more
importantly a year later in John Phillip's ‘Life on Earth: Its origin and
succession’, nor did the end-Ordovician extinction or Silurian recovery.
Rather, in the latter publication, the diversification and extinction is
disguised in a gradual taxonomic increase of Early Palaeozoic life that
continued until the end of the Era. Nevertheless we recognize now that
the GOBE is one of the turning points in the history of life (Sepkoski, 1981;
Sheehan; 1996; Droser and Sheehan, 1997) signalling the most intense
phase of species radiation of the Palaeozoic Era (Webby et al., 2004a,b,c),
prompting irreversible changes in the biological composition and
structure of the planet's oceans (Harper, 2006). The impact of this event
was immense. The sea floors were populated as never before, together
with tiers above and within the substrate, the water column was filled
with diverse phyto- and zooplankton (Servais et al., 2008) and heavily
skeletalized benthos contributed to a major carbonate factory that would
have consequences for long-term climate change on the planet (Pruss et
al., 2010). This hike in diversity was, however, punctuated by a major
extinction and a relatively rapid recovery during the Late Ordovician and
Early Silurian and diversifications and extinctions continued through the
subsequent part of the Silurian.
There is currently no single explanation that can account for the GOBE
(Servais et al., 2009a), but rather a coincidence of biological and geological
factors combined to help drive and promote the biodiversification.
Irrespective of its causes, the diversification changed the oceans forever
and set a new agenda for marine life (Harper, 2006). Similarly there are a
number of hypotheses for the cause of the end-Ordovician glaciation and
the subsequent warming event and a strong research focus on the precise
relationship of the glaciation and its effects, on extinction. Nevertheless,
the Paleozoic Evolutionary Fauna that became established during the
GOBE was relatively stable, surviving the end-Ordovician and late
Devonian extinctions and continuing for some 200Ma. The end Permian
extinction event destroyed its suspension feeding structures and a new
marine ecosystem, based on the detritus feeding ecosystem of the
Modern Evolutionary Fauna, diversified during the Triassic.
During the last decade, IGCP 410 and 503 have established without
doubt the reality and broad importance of the Ordovician biodiversi-
fication but we are only beginning to understand its significance.
Global biodiversity curves are only an aggregation of local and
regional signals (Miller, 1998). The GOBE was diachronous across
different environmental and geographic zones whereas diversifica-
tions and extinctions in biotic groups were rarely coincident. Many
more careful local and regional studies, based on sound taxonomy and
high precision stratigraphy, are required to accurately document
biotic change through these key intervals. These systematic studies
are being aided by better descriptive and statistical techniques based
on a new generation of equipment and software. The increasing use of
GIS software to enable the present locations of samples to be
converted into their positions on palaeogeographical maps is enabling
the spatial distribution patterns of Ordovician organisms to be readily
assessed. Major advances are also expected in our detailed under-
standing of changes in climate and environment during the
Ordovician and Silurian signalled by sea level and geochemical
changes (Munnecke et al., in press-b); the latter detected by a new
generation of analytical machines and techniques. The next challenge
we face is to assemble all our local and regional biotic and proxy data,
construct better global databases and achieve much better syntheses
of our biological, chemical and physical data. In this way we may yet
come closer to identifying the more primary causes of the Early
Palaeozoic diversification and extinction events.
Acknowledgments
This paper is a summary of many of the activities of IGCP n° 503 that
ran from 2004 to 2009. We acknowledge all the participants of the project
who discussed aspects of the present work with us, and in particular the
two co-leaders LI Jun (Nanjing, China) and Peter Sheehan (Milwaukee,
Wisconsin, USA). We are particularly grateful to the two referees who
provided very valuable comments to improve the manuscript (Arnie
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
Miller, Cincinnati, Ohio, USA, and Peter Sheehan, Milwaukee, Wisconsin,
USA). BK acknowledges the University of Lille1 (ATER position) and the
Alexander von Humboldt-Foundation (Bonn, Germany, Feodor-Lynen
programme) for a post-doctoral position at Lille. TS acknowledges the
Alexander von Humboldt-Foundation for a Nachkontakt-Programm at
the Universität Erlangen-Nürnberg and the Naturkundemuseum Berlin
(Germany). DATH acknowledges support from the Danish Council for
Independent Research (FNU). This is a contribution to IGCP 503 and to the
SYSTER project (INSU-CNRS) ‘Climat de l'Ordovicien.’
References
Achab, A., Paris, F., 2007. The Ordovician chitinozoan biodiversification and its leading
factors. Palaeogeography, Palaeoclimatology, Palaeoecology 245, 5–19.
Adrain, J.M., Fortey, R.A., Westrop, S.R., 1998. Post-Cambrian trilobite diversity and
evolutionary faunas. Science 280, 1922–1924.
Adrain, J.M., Westrop, S.R., Chatterton, B.D.E., Ramsköld, L., 2000. Silurian trilobite alpha
diversity and the end-Ordovician mass extinction. Paleobiology 26, 625–646.
Adrain, J.M., Edgecombe, G.D., Fortey, R.A., Hammer, Ø., Laurie, J.R., McCormick, T.,
Owen, A.W., Waisfeld, B.G., Webby, B.D., Westrop, S.R., Zhou, Z.-Y., 2004. Trilobites.
In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 231–254.
Ainsaar, L., Meidla, T., Martma, T., 1999. Evidence for a widespread carbon isotopic
event associated with late Middle Ordovician sedimentological and faunal changes
in Estonia. Geological Magazine 136, 49–62.
Ainsaar, L., Meidla, T., Martma, T., 2004. The Middle Caradoc Facies and Faunal Turnover
in the Late Ordovician Baltoscandian palaeobasin. Palaeogeography, Palaeoclima-
tology, Palaeoecology 210, 119–133.
Ainsaar, L., Kaljo, D., Martma, T., Meidla, T., Männik, P., Nõlvak, J., Tinn, O., 2010. Middle
and Upper Ordovician carbon isotope chemostratigraphy in Baltoscandia: a
correlation standard and clues to environmental history. Palaeogeography,
Palaeoclimatology, Palaeoecology. 294, 189–201.
Aldridge, R.J., Theron, J.N., Gabbott, S.E., 1994. The Soom Shale: a unique Ordovician
fossil horizon in South Africa. Geology Today 10, 218–221.
Algeo, T.J., Seslavinsky, K.B., 1995. The Paleozoic world: continental flooding,
hypsometry, and sea-level. American Journal of Science 295, 787–822.
Allulee, J.L., Holland, S.M., 2005. The sequence stratigraphic and environmental context of
primitive vertebrates: Harding Sandstone, Upper Ordovician, Colorado, USA. Palaios
20, 518–533.
Alroy, J., 2000. New methods for quantifying macroevolutionary patterns and
processes. Paleobiology 26, 707–733.
Alroy, J., et al., 2008. Phanerozoic trends in global diversity of marine invertebrates.
Science 321, 97–100.
Armstrong, H.A., 2007. On the cause of the Ordovician glaciation. In: Williams, M.,
Haywood, A.M., Gregory, F.J., Schmidt, D.N. (Eds.), Deep-Time Perspectives on
Climate Change: Marrying the Signal from Computer Models and Biological Proxies.
Special Publications of the Micropalaeontological Society: The Geological Society,
London, pp. 101–121.
Armstrong, H.A., Owen, A.W., 2002a. Euconodont diversity changes in a cooling and
closing Iapetus Ocean. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeography and
biodiversity change: the Ordovician and Mesozoic-Cenozoic Radiations: Geological
Society, London, Special Publications 194, pp. 85–98.
Armstrong, H.A., Owen, A.W., 2002b. Euconodont biogeography and the closure of the
Iapetus Ocean. Geology 30, 1091–1094.
Armstrong, H.A., Baldini, J., Challands, T.J., Grocke, D.R., Owen, A.W., 2009. Response of
the Inter-tropical Convergence Zone to Southern Hemisphere cooling during Upper
Ordovician glaciation. Palaeogeography, Palaeoclimatology, Palaeoecology 284,
227–236.
Ausich, W.I., Bottjer, D.J., 1982. Tiering in suspension-feeding communities on soft
substrata throughout the Phanerozoic. Science 216, 173–174.
Bambach, R.K., 1983. Ecospace utilization and guilds in marine communities through
the Phanerozoic. In: Tevesz, J.S., McCall, P.L. (Eds.), Biotic Interactions and Fossil
Communities. Plenum Press, New York, London. 719-146.
Bambach, R.K., 1993. Seafood through time: changes in biomass, energetics, and
productivity in the marine ecosystem. Paleobiology 19, 372–397.
Bambach, R.K., 1999. Energetics in the global marine fauna: a connection between
terrestrial diversification and change in the marine biosphere. Geobios 32, 131–144.
Bambach, R.K., Knoll, A.H., Sepkoski, J.J., 2002. Anatomical and ecological constraints on
Phanerozoic animal diversity in the marine realm. Proceedings of the National
Academy of Sciences 99, 6854–6859.
Bambach, R.K., Bush, A.M., Erwin, D.H., 2007. Autecology and the filling of ecospace: key
metazoan radiations. Palaeontology 50, 1–22.
Bassett, M.G. (Ed.), 2009. Early Palaeozoic peri-Gondwana terranes: new insights from
tectonics and biogeography: Geological Society, London, Special Publications, 325.
288 pp.
Bassett, M.G., Popov, L., Holmer, L.E., 2002. Brachiopods: Cambrian-Tremadoc precursors
to Ordovician radiation events. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeography
and biodiversity change: the Ordovician and Mesozoic-Cenozoic Radiations:
Geological Society, London, Special Publications, 194, pp. 13–23.
Barnes, C.R., 2004a. Ordovician Oceans and Climates. In: Webby, B.D., Droser, M.L., Paris,
F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event. Columbia
University Press, New York, pp. 72–76.
115
Barnes, C.R., 2004b. Was there an Ordovician superplume event. In: Webby, B.D.,
Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification
Event. Columbia University Press, New York, pp. 77–80.
Benton, M.J. (Ed.), 1993. Fossil Record 2. Chapman and Hall and the Palaeontological
Association, London.
Benton, M.J., 1995. Diversification and extinction in the history of life. Science 268,
52–58.
Bergström, S.M., Huff, W., Kolata, D., Bauert, H., 1995. Nomenclature, stratigraphy,
chemical fingerprinting, and real distribution of some middle Ordovician -bentonites
in Baltoscandia. GFF 117, 1–13.
Bergström, S.M., Huff, W.D., Saltzman, M.R., Kolata, D.R., Leslie, S.A., 2004. The greatest
volcanic ash falls in the Phanerozoic: Millbrig and Kinekulle K-bentonites. The
Sedimentary Record 2 (4), 4–7.
Bergström, S.M., Finney, S.C., Chen, Xu., Goldman, D., Leslie, S.A., 2006a. Three new
Ordovician global stage names. Lethaia 39, 287–288.
Bergström, S.M., Saltzman, M.R., Schmitz, B., 2006b. First record of the Hirnantian
(Upper Ordovician) δ13C excursion in the North American Midcontinent and its
regional implications. Geological Magazine 143, 657–678.
Bergström, S.M., Chen, X., Gutierrez-Marco, J.C., Dronov, A., 2009a. The new chronostrati-
graphic classification of the Ordovician System and its relations to major regional
series and stages and to δ13C chemostratigraphy. Lethaia 42, 97–107.
Bergström, S.M., Chen, X., Schmitz, B., Young, S., Jia-Yu, R., Saltzman, M.R., 2009b. First
documentation of the Ordovician Guttenberg δ13C excursion (GICE) in Asia:
chemostratigraphy of the Pagoda and Yanwashan formations in southeastern
China. Geological Magazine 146, 1–11.
Berner, R.A., 1994. Geocarb II: a revised model of atmospheric CO2 over Phanerozoic
time. American Journal of Science 294, 56–91.
Berner, R.A., Kothavala, Z., 2001. GEOCARB III: a revised model of atmospheric CO2 over
Phanerozoic time. American Journal of Science 301 (2), 182–204.
Blieck, A.R.M., Turner, S., 2003. Global Ordovician vertebrate biogeography. Palaeogeo-
graphy, Palaeoclimatology, Palaeoecology 195, 37–54.
Botting, J., 2002. The relationship between pyroclastic volcanism and Ordovician
diversification. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeography and
Biodiversity Change: The Ordovician and Mesozoic–Cenozoic Radiations: Geolog-
ical Society, London, Special Publications, 194, pp. 99–113.
Bottjer, D.J., Ausich, W.I., 1986. Phanerozoic development of tiering in soft substrata
suspension-feeding communities. Paleobiology 12, 400–420.
Bottjer, D.J., Hagadorn, J.W., Dornbos, S.Q., 2000. The Cambrian substrate revolution.
GSA Today 10 (9), 1–7.
Bottjer, D.J., Droser, M.L., Sheehan, P.M., McGhee Jr., G.R., 2001. The ecological
architecture of major events in the Phanerozoic history of marine life. In: Allmon,
W.D., Bottjer, D.J. (Eds.), Evolutionary Paleoecology. The Ecological Context of
Macroevolutionary Change. Columbia University Press, New York, pp. 35–61.
Boucot, A.J., 1983. Does evolution take place in an ecological vacuum? II. Journal of
Paleontolology 57, 1–30.
Braddy, S., Tollerton, V.P., Racheboeuf, P.R., Schallreuter, R., 2004. In: Webby, B.D.,
Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification
Event. Columbia University Press, New York, pp. 255–265.
Brenchley, P.J., Marshall, J.D., Carden, G.A.F., Robertson, D.B.R., Long, D.G.F., Meidla, T.,
Hints, L., Anderson, T.F., 1994. Bathymetric and isotopic evidence for a short-lived
Late Ordovician glaciation in a greenhouse period. Geology 22, 295–298.
Buatois, L.A., Mángano, M.G., Brussa, E.D., Benedetto, J.L., Pompei, J.F., 2009. The
changing face of the deep: colonization of the Early Ordovician deep-sea floor,
Puna, northwest Argentina. Palaeogeography, Palaeoclimatology, Palaeoecology
280, 291–299.
Bush, A.M., Bambach, R.K., Daley, G.M., 2007. Changes in theoretical ecospace utilization in
marine fossil assemblages between the mid-Paleozoic and late Cenozoic. Paleobiology
33, 76–97.
Butterfield, N.J., 1997. Plankton ecology and the Proterozoic–Phanerozoic transition.
Paleobiology 23, 247–262.
Butterfield, N.J., 2001. Ecology and evolution of the Cambrian plankton. In: Zhuravlev, A.Y.,
Riding, R. (Eds.), The Ecology of the Cambrian Radiation. Columbia University Press,
New York, pp. 200–216.
Caldeira, K., Rampino, M., 1991. The mid-cretaceous super plume. Geophysical Research
Letters 18, 987–990.
Caputo, M.V., 1998. Ordovician – Silurian glaciations and global sea-level changes. In:
Landing, L., Johnson, M.E. (Eds.), Silurian Cycles — Linkages of Dynamic Stratigraphy
with Atmospheric, Oceanic, and Tectonic Changes: New York State Museum Bulletin,
491, pp. 15–25.
Carrera, M., Rigby, K., 2004. Sponges. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G.
(Eds.), The Great Ordovician Biodiversification Event. Columbia University Press,
New York, pp. 102–111.
Chaffee, C., Lindberg, D.L., 1986. Larval biology of Early Cambrian molluscs: the
implication of small body size. Bulletin of Marine Science 39, 536–549.
Chatterton, B.D.E., Speyer, S.E., 1989. Larval ecology, life history strategies, and patterns of
extinction and survivorship among Ordovician trilobites. Paleobiology 15, 118–132.
Chen, X., Zhang, Y.D., Fan, J.-X., 2006. Ordovician graptolite evolutionary radiation: a
review. Geological Journal 41, 289–301.
Cherns, L., Wheeley, J.R., 2007. A pre-Hirnantian (Late Ordovician) interval of global
cooling — the Boda event re-assessed. Palaeogeography, Palaeoclimatology,
Palaeoecology 251, 449–460.
Christiansen, J.L., Stouge, S., 1999. Oceanic circulation as an element in palaeogeo-
graphical reconstructions: the Arenig (early Ordovician) as an example. Terra Nova
11, 73–78.
Cocks, L.R.M., 2001. Ordovician and Silurian global geography. Journal of the Geological
Society, London 158, 197–210.
 Author's personal copy
116 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
Cocks, L.R.M., Torsvik, T.H., 2002. Earth geography from 500 to 400 million years ago: a
faunal and palaeomagnetic review. Journal of the Geological Society, London 159,
631–644.
Cocks, L.R.M., Torsvik, T.H., 2004. Major terranes in the Ordovician. In: Webby, B.D.,
Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification
Event. Columbia University Press, New York, pp. 61–67.
Cocks, L.R.M., Torsvik, T.H., 2006. European geography in a global context from the Vendian
to the end of the Palaeozoic. In: Gee, D.G., Stephenson, R.A. (Eds.), European
Lithosphere Dynamics, 32. Geological Society, London, Memoirs, pp. 83–95.
Condie, K.C., 2001. Mantle Plumes and Their Record in Earth History. Cambridge
University Press, Cambridge. 315 pp.
Cooper, R.A., 2004. Measures of biodiversity. In: Webby, B.D., Droser, M.L., Paris, F.,
Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event. Columbia
University Press, New York, pp. 52–57.
Cooper, R.A., Maletz, J., Talor, L., Zalasiewicz, J.A., 2004. Graptolites: patterns of diversity across
paleolatitudes. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great
Ordovician Biodiversification Event. Columbia University Press, New York, pp. 281–293.
Cope, J.W.C., 2004. Bivalves and rostroconchs. In: Webby, B.D., Droser, M.L., Paris, F.,
Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event. Columbia
University Press, New York, pp. 196–208.
Cope, J.W.C., Babin, C., 1999. Diversification of bivalves in the Ordovician. Geobios 32,
175–185.
Copper, P., 2001a. Evolution, radiations, and extinctions in Proterozoic to Mid-Paleozoic
reefs. In: Stanley, G. (Ed.), The History of Sedimentology of Ancient Reef Systems
Topics in Geobiology. Plenum Press, New York, pp. 89–119.
Copper, P., 2001b. Reefs during the multiple crisis towards the Ordovician–Silurian
boundary: Anticosti Island, eastern Canada, and worldwide. Canadian Journal of
Earth Sciences 38, 153–171.
Courtillot, V., Olson, P., 2007. Mantle plumes link magnetic superchrons to Phanerozoic
mass depletion events. Earth and Planetary Science Letters 260, 495–504.
Crame, J.A., Owen, A.W. (Eds.), 2002. Palaeobiogeography and biodiversity change: the
Ordovician and Mesozoic–Cenozoic Radiations: Geological Society, London, Special
Publications, 194. 206 pp.
Cramer, B.D., Kleffner, M.A., Brett, C.E., McLaughlin, P.I, Jeppsson, L, Munnecke, A.,
Samtleben, C., 2010. Paleobiogeography, high-resolution stratigraphy, and the future
of Paleozoic biostratigraphy: Fine-scale diachroneity of the Wenlock (Silurian)
conodont Kockelella walliseri. Palaeogeography, Palaeoclimatology, Palaeoecology
294, 232–241.
Davies, N.S., Sansom, I.J., 2009. Ordovician vertebrate habitats: a Gondwanan perspective.
Palaios 24, 717–722.
Díaz-Martínez, E., Grahn, Y., 2007. Early Silurian glaciation along the western margin of
Gondwana (Peru, Bolivia and northern Argentina). Palaeogeographic and geody-
namic setting. Palaeogeography, Palaeoclimatology, Palaeoecology 245, 62–81.
Downie, C., 1984. Acritarchs in British stratigraphy. Geological Society, London, Special
Report 17, 1–26.
Dronov, A., Holmer, E., 1999. Depositional sequences in the Ordovician of Baltoscandia.
Acta Universitatis Carolinae, Geologica 43, 133–136.
Dronov, A., Holmer, L., 2002. Ordovician sea-level curve: Baltoscandian view. In: Satkunas,
J., Lazauskiene, J. (Eds.), Basin Stratigraphy — Modern Methods and Problems, 5th
Baltic Stratigraphical Congress, Vilnius, pp. 33–35. Extended Abstracts.
Droser, M.L., Bottjer, D.J., 1993. Trends and patterns of Phanerozoic ichnofabrics. Annual
Reviews of Earth and Planetary Sciences 21, 205–225.
Droser, M.L., Sheehan, P.M., 1997. Palaeoecology of the Ordovician Radiation: resolution
of large-scale patterns with individual clade histories, palaeogeography and
environments. Geobios 20, 221–229.
Droser, M.L., Finnegan, S., 2003. The Ordovician radiation: a follow-up to the Cambrian
explosion. Integrative Comparative Biology 43, 178–184.
Droser, M.L., Bottjer, D.J., Sheehan, P.M., 1997. Evaluating the ecological architecture of
major events in the Phanerozoic history of marine invertebrate life. Geology 25,
167–170.
Eble, G.J., 1998. The role of development in evolutionary radiations. In: McKinney, M.,
Drake, L., James, A. (Eds.), Biodiversity Dynamics: Turnover of Populations, Taxa,
and Communities. Columbia University Press, New York, pp. 132–162.
Elliott, D.K., Blieck, A.R.M., Gagnier, P.Y., 1991. Ordovician vertebrates. In: Barnes, C.R.,
Williams, S.H. (Eds.), Advances in Ordovician Geology. Geological Survey of Canada,
Paper, 90-9, pp. 93–106.
Ernst, A., Munnecke, A., 2009. A Hirnantian (latest Ordovician) reefal bryozoan fauna
from Anticosti Island, eastern Canada: taxonomy and chemostratigraphy. Canadian
Journal of Earth Sciences 46, 207–229.
Evitt, W.R., 1963. A discussion and proposals concerning fossil dinoflagellates,
hystrichospheres and acritarchs. II. Proceedings of the National Academy of
Sciences 49, 298–302.
Falkowski, P.G., Katz, M.E., Knoll, A.H., Quigg, A., Raven, J.A., Schofield, O., Taylor, F.J.R.,
2004. The evolution of modern eukaryotic phytoplankton. Science 305, 354–360.
Fanton, K.C., Holmden, C., 2007. Sea-level forcing of carbon isotope excursions in epeiric seas:
implications for chemostratigraphy. Canadian Journal of Earth Sciences 44, 807–818.
Fensome, R.A., Williams, G.L., Barss, M.S., Freeman, J.M., Hill, J.M., 1990a. Acritarchs and
fossil prasinophytes: an index to genera, species and intraspecific taxa. American
Association of Stratigraphic Palynologists, Contribution Series 25 771 pp.
Fensome, R.A., Williams, G.L., Barss, M.S., Freeman, J.M., Hill, J.M., 1990b. Acritarchs and
fossil prasinophytes: an index to genera, species and intraspecific taxa. American
Association of Stratigraphic Palynologists Contribution Series 25, 1–771.
Fensome, R.A., MacRae, R.A., Moldowan, J.M., Taylor, F.R.J., Williams, G.L., 1996. The
early Mesozoic radiation of dinoflagellates. Paleobiology 22, 329–338.
Finnegan, S., Droser, M.L., 2008a. Body size, energetics, and the Ordovician restructuring
of marine ecosystems. Paleobiology 34, 342–359.
Finnegan, S., Droser, M.L., 2008b. Reworking diversity: effects of storm deposition on
evenness and sampled richness, Ordovician of the Basin and Range (UT and NV).
Palaios 23, 87–96.
Foote, M., 1991. Morphological patterns of diversification: examples from trilobites.
Palaeontology 34, 461–485.
Fortey, R.A., Cocks, L.R.M., 2003. Palaeontological evidence bearing on global Ordovician–
Silurian continental reconstructions. Earth-Science Reviews 61, 245–307.
Fortey, R.A., Cocks, L.R.M., 2005. Late Ordovician global warming—the Boda event.
Geology 33, 405–408.
Fortey, R.A., Owens, R.M., 1987. The Arenig Series in South Wales. Bulletin of the British
Museum Natural History (Geology) 41, 69–307.
Fortey, R.A., Owens, R.M., 1990. Evolutionary radiations in the Trilobita. In: Taylor, P.D.,
Larwood, G.P. (Eds.), Major Evolutionary Radiations. Systematics Association,
Special Volume, 42. Clarendon Press, Oxford, pp. 139–164.
Fortey, R.A., Owens, R.M., 1999. Feeding habits in trilobites. Palaeontology 42, 427–465.
Fortey, R.A., Rushton, A.W.A., 2007. The mid-Cambrian trilobite Beishanella, and the limits
of convergent evolution. Memoirs of the Association of Australasian Palaeontologists
34, 391–399.
François, L.M., Walker, J.C.G., 1992. Modelling the Phanerozoic carbon cycle and
climate: constraints from the 87Sr/86Sr isotopic ratio of seawater. American Journal
of Science 292, 81–135.
Frey, R.C., 1989. Paleoecology of a well-preserved nautiloid assemblage from a Late
Ordovician shale unit, Southwestern Ohio. Journal of Paleontology 63,
604–620.
Frey, R.C., Beresi, M.S., Evans, D.H., King, A.H., Percival, I.G., 2004. Nautiloid cephalopods.
In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 209–213.
Frýda, J., Rohr, D.M., 2004. Gastropods. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G.
(Eds.), The Great Ordovician Biodiversification Event. Columbia University Press, New
York, pp. 185–195.
Gibbs, M., Barron, E., Kump, L.R., 1997. An atmospheric pCO2 threshold for glaciation in
the Late Ordovician. Geology 25, 447–450.
Goddéris, Y., Francois, L.M., Veizer, J., 2001. The early Paleozoic carbon cycle. Earth and
Planetary Science Letters 190, 181–196.
Grahn, Y., Caputo, M.V., 1992. Early Silurian glaciations in Brazil. Palaeogeography,
Palaeoclimatology, Palaeoecology 99, 9–15.
Guinotte, J.M., Orr, J., Cairns, S., Freiwald, A., Morgan, L., George, R., 2006. Will human-
induced changes in seawater chemistry alter the distribution of deep-sea
scleractinian corals? Frontiers in Ecology and the Environment 4 (3), 141–146.
Hallam, A., 1992. Phanerozoic Sea-level Changes. Columbia University Press, New York.
266 pp.
Hambrey, M.J., Harland, W.B., 1981. Earth's Pre-Pleistocene glacial record. Cambridge
University Press, Cambridge.
Hammer, O., 2003. Biodiversity curves for the Ordovician of Baltoscandia. Lethaia 36,
305–314.
Haq, B.U., Schutter, S.R., 2008. A chronology of Paleozoic sea-level changes. Science 322,
64–68.
Haq, B.U., Hardenbol, J., Vail, P.R., 1988. Mesozoic and Cenozoic chronostratigraphy and
eustatic cycles. Sea-level changes: an integrated approach. SEPM Special Publica-
tion 42, 71–108.
Harper, D.A.T., 2006. The Ordovician biodiversification: setting an agenda for marine
life. Palaeogeography, Palaeoclimatology, Palaeoecology 232, 148–166.
Harper, D.A.T., 2010. The Ordovician brachiopod radiation: roles of alpha, beta and
gamma diversity. Special Paper, Geological Society of America 466, 69–83.
Harper, D.A.T., Pickerill, R.K., 1997. Mid Ordovician commensal relationships between
articulate brachiopods and a trepostome bryozoan from eastern Canada. Atlantic
Geology 32, 181–187.
Harper, D.A.T., Mac Niocaill, C., 2002. Early Ordovician rhynchonelliformean brachiopod
diversity: Comparing some platforms, margins and intra-oceanic sites around the
Iapetus Ocean. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeography and Biodiversity
Change: The Ordovician and Mesozoic–Cenozoic Radiations: Geological Society,
London, Special Publications, 194, pp. 25–34.
Harper, D.A.T., Mac Niocaill, C., Williams, S.H., 1996. Palaeogeography of early
Ordovician Iapetus terranes: integration of faunal and palaeomagnetic data.
Palaeogeography, Palaeoclimatology, Palaeoecology 121, 297–312.
Harper, D.A.T., Cocks, L.R.M., Popov, L.E., Sheehan, P.M., Bassett, M.G., Copper, P.,
Holmer, L.E., Jin, J., Rong, J.-Y., 2004. Brachiopods. In: Webby, B.D., Droser, M.L.,
Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event.
Columbia University Press, New York, pp. 157–178.
Harper, D.A.T., Owen, A.W., Bruton, D.L., 2009. Ordovician life around the Celtic fringes:
Diversifications, Extinctions and Migrations of Brachiopod and Trilobite Faunas at
Middle Latitudes. In: Bassett, M.G. (Ed.), Early Palaeozoic Peri-Gondwanan
Terranes: New insights from tectonics and biogeography: Geologocal Society,
London, Special Publications, 325, pp. 157–170.
Hatch, J.R., Jacobson, S.R., Witzke, B.J., Risatti, J.B., Anders, D.E., Watney, W.L., Newell, K.D.,
Vuletich, A.K., 1987. Possible late Middle Ordovician organic-carbon isotope
excursion — evidence from Ordovician oils and hydrocarbon source rocks,
Midcontinent and East-Central United-States. AAPG Bulletin-American Association
of Petroleum Geologists 71, 1342–1354.
Herrmann, A., Haupt, B., Patzkowsky, M., Seidov, D., Slingerland, R., 2004a. Response of
Late Ordovician paleoceanography to changes in sea level, continental drift, and
atmospheric pCO2: potential causes for long-term cooling and glaciation.
Palaeogeography, Palaeoclimatology, Palaeoecology 210, 385–401.
Herrmann, A., Patzkowsky, M., Pollard, D., 2004b. The impact of paleogeography, pCO2,
poleward ocean heat transport and sea level change on global cooling during the
Late Ordovician. Palaeogeography, Palaeoclimatology, Palaeoecology 206, 59–74.
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
Hints, O., Delabroye, A., Nõlvak, J., Servais, T., Uutela, A., Wallin, A., 2010. Biodiversity
patterns of Ordovician marine microphytoplankton from Baltica: Comparison with
other fossil groups and sea-level changes. Palaeogeography, Palaeoclimatology,
Palaeoecology 294, 161–173.
Holmden, C., Creaser, R.A., Muehlenbachs, K., Leslie, S.A., Bergström, S.M., 1998. Isotopic
evidence for geochemical decoupling between ancient epeiric seas and bordering
oceans: implications for secular curves. Geology 26, 567–570.
Huff, W., Bergström, S.M., Kolata, D., 1992. Gigantic Ordovician ash fall in North America
and Europe: biological, tectonomagmatic, and event stratigraphic significance.
Geology 20, 875–878.
Jablonski, D., Sepkoski Jr, J.J., Bottjer, D.J., Sheehan, P.M., 1983. Onshore–offshore
patterns in the evolution of Phanerozoic shelf communities. Science 222,
1123–1125.
Jeppsson, L., 1990. An oceanic model or lithological and faunal changes tested on the
Silurian record. Journal of the Geological Society, London 147, 663–674.
Jiménez-Sánchez, A., Villas, E., 2010. The bryozoan dispersion into the Mediterranean
margin of Gondwana during the pre-glacial Late Ordovician. Palaeogeography,
Palaeoclimatology, Palaeoecology 294, 220–231.
Joachimski, M.M., Horacek, S., Breisig, S., Buggisch, W., 2003. The oxygen isotopic
composition of biogenic apatite — no evidence for a secular change in seawater
δ18O. European Geophysical Society, Geophysical Research Abstracts 5, 10792.
Kaljo, D., Hints, L., Martma, T., Nolvak, J., 2001. Carbon isotope stratigraphy in the latest
Ordovician of Estonia. Chemical Geology 175, 49–59.
Kaljo, D., Martma, T., Männik, P., Viira, V., 2003. Implications of Gondwana glaciations in
the Baltic Later Ordovician and Silurian and a carbon isotopic test of environmental
cyclicity. Bulletin de la Société géologique de France 174, 59–66.
Kaljo, D., Hints, L., Martma, T., Nolvak, J., Oraspold, A., 2004. Late Ordovician carbon
isotope trend in Estonia, its significance in stratigraphy and environmental
analysis. Palaeogeography, Palaeoclimatology, Palaeoecology 210, 165–185.
Katz, M.E., Finkel, Z.V., Grzebyk, D., Knoll, A.H., Falkowski, P.G., 2004. Evolutionary
trajectories and biogeochemical impacts of marine eukaryotic phytoplankton.
Annual Review Ecology Evolution S. 35, 523–556.
Kiessling, W., 2002. Secular variation in the Phanerozoic reef system. In: Kiessling, W.,
Flügel, E., Golonka, J. (Eds.), Phanerozoic Reef Patterns: SEPM Special Publication,
72, pp. 625–690.
Kleypas, J.A., Buddemeier, R.W., Archer, D., Gattuso, J.-P., Langdon, C., Opdyke, B.N.,
1999. Geochemical consequences of increased atmospheric carbon dioxide on coral
reefs. Science 284, 118–120.
Klug, C., Kröger, B., Kiessling, W., Mullins, G.L, Servais, T., Fryda, J., Korn, D., Turner, S., in
press. The Devonian nekton revolution. Lethaia (Electronic publication ahead of
print). doi:10.1111/j.1502-3931.2009.00203.x.
Kolata, D.R., Huff, W.D., Bergström, S.M., 1996. Ordovician K-bentonites of eastern
North America. GSA Special Paper 313, 1–84.
Kröger, B., 2004. Large shell injuries in Middle Ordovician Orthocerida (Nautiloidea,
Cephalopoda). GFF 126, 311–316.
Kröger, B., Landing, E., 2007. The earliest Ordovician cephalopods of eastern Laurentia -
Ellesmerocerids of the Tribes Hill Formation, eastern New York. Journal of
Paleontology 81, 841–857.
Kröger, B., Landing, E., 2008. Onset of the Ordovician cephalopod radiation—evidence
from the Rochdale Formation (middle Early Ordovician, Stairsian) in eastern New
York. Geological Magazine 145, 490–520.
Kröger, B., Landing, E., 2009. Cephalopods and paleoenvironments of the Fort Cassin
Formation (Upper Lower Ordovician), eastern New York and adjacent Vermont.
Journal of Paleontology 83, 664–693.
Kröger, B., Landing, E., in press. Early Ordovician community evolution with eustatic
change through the Beekmantown Group, northeast Laurentia. Palaeogeography,
Palaeoclimatology, Palaeoecology.
Kröger, B., Zhang, Y.B., 2009. Pulsed cephalopod diversification during the Ordovician.
Palaeogeography, Palaeoclimatology, Palaeoecology 273, 174–183.
Kröger, B., Servais, T., Zhang, Y.B., 2009. The origin and initial rise of pelagic cephalopods
in the Ordovician. PLoS ONE 4 (9), e7262. doi:10.1371/journal.pone.0007262.
Kump, L.R., Arthur, M.A., Patzkowsky, M., Gibbs, M., Pinkus, D., Sheehan, P., 1999. A
weathering hypothesis for glaciation at high atmospheric pCO2 during the Late
Ordovician. Palaeogeography, Palaeoclimatology, Palaeoecology 152, 173–187.
Landing, E., 2002. Early Paleozoic sea levels and climates: New evidence from the east
Laurentian shelf and slope. In: McLelland, J., Karabinos, P. (Eds.), Guidebook for
Fieldtrips in New York and Vermont, New England Intercollegiate Geological
Conference 94th Annual Meeting and New York State Geological Association 74th
Annual Meeting. Lake George, New York. C6-1-C6-22.
Landing, E., English, A., Keppie, J.D., 2010. Cambrian origin of all skeletalized metazoan
phyla – Discovery of Earth’s oldest bryozoans (Upper Cambrian, southern Mexico).
Geology 6, 547–550.
Lefebvre, B., Fatka, O., 2003. Palaeogeographical and palaeoecological aspects of the
Cambro-Ordovician radiation of echinoderms in Gondwanan Africa and peri-
Gondwanan Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 195, 73–97.
Lefebvre, V., Servais, T., François, L., Averbuch, O., in press. Did a Katian large igneous
province trigger the Late Ordovician glaciation? A hypothesis tested with a carbon
cycle model. Palaeogeography, Palaeoclimatology, Palaeoecology
Li, J., Servais, T., Yan, K., Zhu, H., 2004. A nearshore-offshore trend in the acritarch
distribution of the Early-Middle Ordovician of the Yangtze Platform, South China.
Review of Palaeobotany and Palynology 130, 141–161.
Liebermann, B.S., 2004. Geobiology and paleobiogeography: tracking the coevolution of the
Earth and its biota. Palaeogeography, Palaeoclimatology, Palaeoecology 219, 23–33.
Long, D.G.F., 1993. Oxygen and carbon isotopes and event stratigraphy near the
Ordovician–Silurian boundary, Anticosti Island Quebec. Palaeogeography, Palaeo-
climatology, Palaeoecology 104, 49–59.
117
Mángano, M.G., Droser, M.L., 2004. The ichnologic record of the Ordovician Radiation.
In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 369–379.
Martin, R.E., 1996. Secular increase in nutrient levels through the Phanerozoic: implications
for productivity, biomass, and diversity of the marine biosphere. Palaios 11, 209–219.
Martin, R.E., 2003. The fossil record of biodiversity: nutrients, productivity, habitat area
and different preservation. Lethaia 36, 179–193.
Martin, R.E., Quigg, A., Podkovyrov, V., 2008. Marine biodiversification in response to
evolving phytoplankton stoichiometry. Palaeogeography, Palaeoclimatology,
Palaeoecology 258, 277–291.
McCormick, T., Fortey, R.A., 1999. The most widely distributed trilobite species:
Ordovician Carolinites genacinaca. Journal of Paleontology 73, 201–218.
McGowan, A.J., Smith, A.B., 2007. Are global Phanerozoic marine diversity curves truly
global? A study of the relationship between regional rock records and global
Phanerozoic marine diversity. Paleobiology 34, 80–103.
McKerrow, W.S. (Ed.), 1978. Ecology of Fossils. Duckworth Press, London. 383 pp.
Miller, A.I., 1997a. Dissecting global diversity patterns. Annual Review of Ecology and
Systematics 28, 85–104.
Miller, A.I., 1997b. Comparative diversification dynamics among palaeocontinents
during the Ordovician Radiation. Geobios, Mémoire Spécial 20, 397–406.
Miller, A.I., 1997c. A new look at age and area: the geographic and environmental
expansion of genera during the Ordovician radiation. Paleobiology 23,
410–419.
Miller, A.I., 1998. Biotic transitions in global marine diversity. Science 281, 1157–1160.
Miller, A.I., 2000. Conversations about Phanerozoic global diversity. In: Erwin, D.H.,
Wing, S.L. (Eds.), Deep Time: Paleobiology's Perspective: Paleobiology, Supple-
ment, 26, pp. 53–73.
Miller, A.I., 2004. The Ordovician Radiation: towards a new global synthesis. In: Webby,
B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversifica-
tion Event. Columbia University Press, New York, pp. 380–388.
Miller, A.I., 2009. The consensus that changed the paleobiological world. In: Sepkoski,
D., Ruse, M. (Eds.), The Paleobiological Revolution. University of Chicago Press,
Chicago, pp. 364–382.
Miller, A.I., Mao, S., 1995. Association of orogenic activity with the Ordovician radiation
of marine life. Geology 23, 305–308.
Miller, A.I., Holland, S.M., Droser, M.L., Patzkowsky, M.E., 1998. Dynamics of the Ordovician
Radiation: a comment on Westrop and Adrain. Paleobiology 24, 524–528.
Miller, K.G., Kominz, M.A., Browning, J.V., Wright, J.D., Mountain, G.S., Katz, M.E.,
Sugarman, P.J., Cramer, B.S., Christie-Blick, N., Pekar, S.F., 2005. The Phanerozoic
record of global sea-level change. Science 310, 1293–1298.
Mullins, G., Servais, T., Vecoli, M., 2006. The Ordovician phytoplankton database:
progress report. In: Yang, Q., Wang, Y.-D., Weldon, E.A. (Eds.), Ancient Life and
Modern Approaches. Abstracts of the Second International Palaeontological
Congress. University of Science and Technology of China Press, pp. 317–318.
Munnecke, A., Servais, T., 1996. Scanning Electron Microscopy of polished, slightly etched
rock surfaces: a method to observe palynomorphs in situ. Palynology 20, 163–176.
Munnecke, A., Servais, T., 2007. What caused the Ordovician biodiversification?
Palaeogeography, Palaeoclimatology, Palaeoecology 245, 1–4.
Munnecke, A., Servais, T., 2008. Palaeozoic calcareous plankton: evidence from the
Silurian of Gotland. Lethaia 41, 185–194.
Munnecke, A., Samtleben, C., Bickert, T., 2003. The Ireviken Event in the lower Silurian
of Gotland, Sweden — relation to similar Palaeozoic and Proterozoic events.
Palaeogeography, Palaeoclimatology, Palaeoecology 195, 99–124.
Munnecke, A., Calner, M., Harper, D.A.T. (Eds.), this issue. Lower Palaeozoic sea-levels
and palaeoclimate. Palaeogeography, Palaeoclimatology, Palaeoecology.
Nardin, E., Lefebvre, B., 2010. Unravelling extrinsic and intrinsic factors of the Early
Palaeozoic diversification of blastozoan Echinoderms. Palaeogeography, Palaeocli-
matology, Palaeoecology 294, 142–160.
Nardin, E., Lefebvre, B., David, B., Mooi, R., 2009. La diversification des échinodermes
primitifs au Paléozoïque inférieur: l'exemple des blastozoaires. CR Palevol 8,
179–188.
Nielsen, A.T., 2004. Ordovician sea level changes: a Baltoscandian perspective. In:
Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 84–93.
Nielsen, A.T., 2007. Sea-level changes, glaciations and oxygen level during the Early
Palaeozoic: controlling factors for the Cambrian explosion and the Ordovician
radiation? In: Ebbestad, J.O.R., Wickström, L.M., Högström, A.E.S. (Eds.), Wogogob
2007, 9th meeting of the Working Group on Ordovician Geology of Baltoscandia —
Field guide and Abstracts: Sveriges geologiska undersökning, Rapporter och
Meddelanden, 128, pp. 106–107.
Nitecki, M.H., Webby, B.D., Spjeldnæs, N., Zhen, Y.Y., 2004. Receptaculitids and algae. In:
Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 336–347.
Noble, P.J., Danelian, T., 2004. Radiolarians. In: Webby, B.D., Droser, M.L., Paris, F.,
Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event. Columbia
University Press, New York, pp. 97–101.
Novack-Gottshall, P.M., Miller, A.I., 2003a. Comparative geographic and environmental
diversity dynamics of gastropods and bivalves during the Ordovician Radiation.
Paleobiology 29, 576–604.
Novack-Gottshall, P.M., Miller, A.I., 2003b. Comparative taxonomic richness and
abundances of the Late Ordovician gastropods and bivalves in the mollusc-rich
strata of the Cincinnati Arch. Palaios 18, 559–571.
Nützel, A., Frýda, J., 2003. Paleozoic plankton revolution: evidence from early gastropod
ontogeny. Geology 31, 829–831.
Nützel, A., Lehnert, O., Fryda, J., 2006. Origin of planktotrophy — evidence from early
molluscs. Evolution and Development 8, 325–330.
 Author's personal copy
118 T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
Orr, P.J., 2001. Colonization of the deep-marine environment during the early
Phanerozoic: the ichnofaunal record. Geological Journal 36, 265–278.
Owen, A.W., 2007. Trilobite diversity in Avalonia prior to the end Ordovician extinction —
the peak before the trough. Palaeogeography, Palaeoclimatology, Palaeoecology 245,
264–271.
Owen, A.W., 2008. Ordovician and Silurian environments, biogeography and biodiver-
sity change: an introduction. Lethaia 41, 97.
Owen, A.W., Bruton, D.L., 2008. The environmental significance and fate of the
trilobite fauna of the Pyle Mountain Argillite (Upper Ordovician), Maine. In:
Rábano, I., Gozalo, R., García-Bellido, D. (Eds.), Advances in Trilobite Research.
Cuadernos del Museo Geominero, Instituto Geológico y Minero de España,
Madrid, 9, pp. 283–287.
Owen, A.W., Crame, J.A., 2002. Palaeobiogeography and the Ordovician and Mesozoic-
Cenozoic radiations. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeography and
Biodiversity Change: The Ordovician and Mesozoic–Cenozoic Radiations: Geolog-
ical Society, London, Special Publications, 194, pp. 1–11.
Palmer, T.J., Wilson, M.A., 2004. Calcite precipitation and dissolution of biogenic
aragonite in shallow Ordovician calcite seas. Lethaia 37, 417–427.
Paris, F., 2008. Meteoritic spur to life? Nature Geoscience 1, 18–19.
Paris, F., Achab, A., Chen, X., Grahn, Y., Nolvak, J., Obut, O., Samuelsson, J., Sennikov, N.,
Vecoli, M., Verniers, J., Wang, X., Winchester-Seeto, T., 2004. Chitinozoans. In:
Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Biodiversification Event. Columbia University Press, New York, pp. 294–311.
Parnell, J., 2008. Global mass wasting at continental margins during Ordovician high
meteorite influx. Nature Geoscience 2, 57–61.
Parrish, J.T., 1982. Upwelling and petroleum source beds, with reference to the
Paleozoic. AAPG Bulletin 66, 750–774.
Parrish, J.T., 1987. Palaeo-upwelling and the distribution of organic-rich rocks. In:
Brooks, J., Fleet, A.J. (Eds.), Marine Petroleum Source Rocks: Geological Society
Special Publication, 26, pp. 199–205.
Parrish, J.T., Ziegler, A.M., Humphreville, R.G., 1983. Upwelling in the Paleozoic Era. In: Seuss,
E., Thiede, J. (Eds.), Coastal Upwelling, Its Sedimentary Record: Part B. Sedimentary
Records of Ancient Coastal Upwelling. Plenum Press, New York, pp. 553–578.
Patzkowsky, M.E., Holland, S.M., 2003. Lack of community saturation at the beginning of
the Paleozoic plateau: the dominance of regional over local processes. Paleobiology
29, 545–560.
Pavlov, V., Gallet, Y., 2005. A third superchron during the Early Paleozoic. Journal of
International Geoscience 28 (2), 78–84.
Perrier, V., Vannier, J., Siveter, D.J., 2008. The Silurian pelagic myodocope ostracod
Richteria migrans. Earth and Environmental Science, Transactions of the Royal
Society of Edinburgh 98, 151–163.
Peters, S.E., 2005. Geologic constraints on the macroevolutionary history of marine
animals. Proceedings of the National Academy of Sciences 102, 12326–12331.
Peters, S.E., Foote, M., 2002. Determinants of extinction in the fossil record. Nature 416,
420–424.
Peterson, K.J., 2005. Macroevolutionary interplay between planktic larvae and benthic
predators. Geology 33, 929–932.
Phillips, J., 1860. Life on Earth: Its Origin and Succession, Cambridge and London.
Pruss, S.B., Finnegan, S., Fischer, W.W., Knoll, A.H., 2010. Carbonates in skeleton-poor seas:
new insights from Cambrian and Ordovician strata of Laurentia. Palaios 25, 73–84.
Racheboeuf, P.R., Crasquin, S., Brussa, E., 2009. South American Ordovician phyllocarids
(Crustacea, Malacostraca). Bulletin of Geosciences 84, 377–408.
Rasmussen, C.M.Ø., Harper, D.A.T., 2008. Resolving early Mid-Ordovician (Kundan)
bioevents in the East Baltic based on brachiopods. Geobios 41, 533–542.
Rasmussen, C.M.Ø., Hansen, J., Harper, D.A.T., 2007. Baltica: a mid Ordovician diversity
hotspot. Historical Biology 19, 255–261.
Rasmussen, J.A., 1998. A reinterpretation of the conodont Atlantic Realm in the late Early
Ordovician (early Llanvirn). In: Szaniawski, H. (Ed.), Proceedings of the Sixth
European Conodont Symposium (ECOS VI): Palaeontologica Polonica, 58, pp. 67–77.
Raup, D.M., 1972. Taxonomic diversity during the Phanerozoic. Science 177, 1065–1071.
Raup, D.M., 1975. Taxonomic diversity estimation using rarefaction. Paleobiology 1, 333–342.
Ridgwell, A., 2005. A Mid Mesozoic Revolution in the regulation of ocean chemistry.
Marine Geology 217, 339–357.
Riding, R., 1993. Phanerozoic patterns of marine CaCO3 precipitation. Naturwis-
senschaften 80, 513–516.
Riding, R., 2000. Microbial carbonates: the geological record of calcified bacterial algal
mats and biofilms. Sedimentology 47 (Suppl. 1), 179–214.
Riding, R., 2002. Structure and composition of organic reefs and carbonate mud
mounds: concepts and categories. Earth Science Reviews 58, 163–231.
Riding, R., 2006a. Cyanobacterial calcifcation, carbon dioxide concentrating mechan-
isms, and Proterozoic–Cambrian changes in atmospheric composition. Geobiology
4, 299–316.
Riding, R., 2006b. Microbial carbonate abundance compared with fluctuations in
metazoan diversity over geological time. Sedimentary Geology 185, 229–238.
Riding, R., Liang, L., 2005. Geobiology of microbial carbonates: metazoan and seawater
saturation state influences on secular trends during the Phanerozoic. Palaeogeo-
graphy, Palaeoclimatology, Palaeoecology 219, 101–115.
Ross, J.R.P., Ross, C.A., 1992. Ordovician sea-level fluctuation. In: Webby, B.D., Laurie, J.R.
(Eds.), Proceedings of the Sixth International Symposium on the Ordovician
System, University of Sydney, Australia, 15 – 19 July 1991: Global Perspectives on
Ordovician Geology. A. Balkema, Rotterdam, pp. 327–335.
Ross, C.A., Ross, J.R.P., 1995. North American depositional sequences and correlations.
In: Cooper, C., Droser, M., Finney, S. (Eds.), 7th International Symposium on the
Ordovician System, Las Vegas, SEPM Publication, 77, pp. 309–314.
Rozhnov, S.V., 2007. Changes in the Early Palaeozoic geography as a possible factor
of echinoderm higher taxa formation: delayed larval development to cross
the Iapetus Ocean. Palaeogeography, Palaeoclimatology, Palaeoecology 245,
306–316.
Runnegar, B., 2007. No evidence for planktotrophy in Cambrian molluscs. Evolution and
Development 9 (4), 311–312.
Saltzman, M.R., 2005. Phosphorus, nitrogen, and the redox evolution of the Paleozoic
oceans. Geology 33, 573–576.
Saltzman, M.R., Young, S.A., 2005. Long-lived glaciation in the Late Ordovician? Isotopic
and sequence-stratigraphic evidence from western Laurentia. Geology 33, 109–112.
Sandy, M.R., 1996. The oldest record of pedunculate attachment of brachiopods to
crinoid stems, upper Ordovician, Ohio, U.S.A. (Brachiopoda; Atrypida: Echinoder-
mata; Crinoidea). Journal of Paleontology 70, 532–534.
Sansom, I.J., Smith, M.M., 2001. The Ordovician radiation of vertebrates. In: Ahlberg, E.
(Ed.), Major Events in Vertebrate Evolution. Taylor & Francis, London, pp. 156–171.
Sansom, I.J., Miller, C.G., Heward, A., Davies, N.S., Booth, G.A., Fortey, R.A., Paris, F., 2009.
Ordovician fish from the Arabian Peninsula. Palaeontology 52, 337–342.
Schmitz, B., Bergström, S.M., 2007. Chemostratigraphy in the Swedish Upper
Ordovician: Regional significance of the Hirnantian δ13C excursion (HICE) in the
Boda Limestone of the Siljan region. GFF 129, 133–140.
Schmitz, B., Harper, D.A.T., Peucker-Ehrenbrink, B., Stouge, S., Alwmark, C., Cronholm,
A., Bergström, S.M., Tassinari, M., Wang, X.F., 2008. Asteroid breakup linked to the
Great Ordovician Biodiversification Event. Nature Geoscience 1, 49–53.
Schoenemann, B., Clarkson, E.N.K., Ahlberg, P., Dies Àlvarez, M.E., 2008. A Furongian
polymerid planktonic trilobite. In: Rábano, I., Gozalo, R., García-Bellido, D. (Eds.),
Advances in trilobite research. Cuadernos del Museo Geominero: Instituto
Geológico y Minero de España, Madrid, 9, pp. 361–364.
Scotese, C.R., McKerrow, W.S., 1990. Revised world maps and introduction. In:
McKerrow, W.S., Scotese, C.R. (Eds.), Palaeozoic Palaeogeography and Biogeogra-
phy: Memoir of the Geological Society, London, 12, pp. 1–21.
Seilacher, A., Pflüger, F., 1994. From biomats to benthic agriculture: a biohistoric
revolution. In: Krumbien, W.E., Paterson, D.M., Stal, L.J. (Eds.), Biostabilization of
sediments. Bibliotheks und Informationssystem der Carl von Ossietzky Universität
Oldenburg, pp. 97–105.
Sepkoski Jr., J.J., 1978. A kinetic model for Phanerozoic taxonomic diversity: I. Analysis
of marine orders. Paleobiology 4, 223–251.
Sepkoski Jr., J.J., 1979. A kinetic model for Phanerozoic taxonomic diversity: II. Early
Phanerozoic families and multiple equilibria. Paleobiology 5, 222–251.
Sepkoski Jr., J.J., 1981. A factor analytical description of the Phanerozoic marine fossil
record. Paleobiology 7, 36–53.
Sepkoski Jr., J.J., 1984. A kinetic model for Phanerozoic taxonomic diversity: part 3. Post-
Paleozoic families and multiple equilibria. Paleobiology 10, 246–267.
Sepkoski Jr., J.J., 1988. Alpha, beta, or gamma: where does all the diversity go? Palaeobiology
14, 221–234.
Sepkoski Jr., J.J., 1991. A model for onshore–offshore change in faunal diversity.
Paleobiology 17, 157–176.
Sepkoski Jr, J.J., 1995. The Ordovician Radiations: diversification and extinction shown by
global genus-level taxonomic data. In: Cooper, C., Droser, M., Finney, S. (Eds.), 7th
International Symposium on the Ordovician System, Las Vegas, SEPM Publication, 77,
pp. 75–80.
Sepkoski Jr., J.J., 2002. The compendium. In: Jablonski, D., Foote, M. (Eds.), A
compendium of Fossil Marine Animal Genera: Bulletins of American Paleontology,
363, pp. 10–560.
Sepkoski Jr., J.J., Sheehan, P.M., 1983. Diversification, faunal change, and community
replacement during the Ordovician radiation. In: Tevesz, M.J.S., McCall, P.M. (Eds.),
Biotic Interactions in Recent Fossil Benthic Communities. Plenum Press, New York.
Sepkoski Jr., J.J., Miller, A.I., 1985. Evolutionary faunas and the distribution of benthic
communities in space and time. In: Valentine, J.W. (Ed.), Phanerozoic Diversity
Patterns: Profiles in Macroevolution. Princeton University Press and Pacific
Division, American Association for the Advancement of Science, Princeton, New
Jersey.
Servais, T., Owen, A.W., this issue. Preface — Early Palaeozoic environments: an
‘explosion’ of diversity of marine species, genera and families. Palaeogeography,
Palaeoclimatology, Palaeoecology.
Servais, T., Brocke, R., Fatka, O., Le Hérissé, A., Molyneux, S.G., 1997. Value and meaning
of the term acritarch. In: Fatka, O., Servais, T. (Eds.), Acritarcha in Praha 1996: Acta
Universitatis Carolinae, Geologica, 40, pp. 631–643.
Servais, T., Li, J., Molyneux, S.G., Raevskaya, E., 2003. Ordovician organic-walled
microphytoplankton (acritarch) distribution: the global scenario. Palaeogeogra-
phy, Palaeoclimatology, Palaeoecology 195, 149–172.
Servais, T., Li, J., Stricanne, L., Vecoli, M., Wicander, R., 2004a. Acritarchs. In: Webby, B.D.,
Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversification
Event. Columbia University Press, New York, pp. 348–360.
Servais, T., Stricanne, L., Montenari, M., Pross, J., 2004b. Population dynamics of galeate
acritarchs at the Cambrian–Ordovician transition in the Algerian Sahara. Palaeon-
tology 47, 395–414.
Servais, T., Blieck, A., Caridroit, M., Chen, X., Paris, F., Tortello, F., 2005. Use and utility of
plankton and nekton for Ordovician palaeogeographical reconstructions. Bulletin
de la Société géologique de France 176, 531–543.
Servais, T., Lehnert, O., Li, J., Mullins, G.L., Munnecke, A., Nützel, A., Vecoli, M., 2008. The
Ordovician Biodiversification: revolution in the oceanic trophic chain. Lethaia 41, 99–109.
Servais, T., Harper, D.A.T., Li, J., Munnecke, A., Owen, A.W., Sheehan, P.M., 2009a.
Understanding the Great Ordovician Biodiversification Event (GOBE): influences of
paleogeography, paleoclimate, or paleoecology? GSA Today 19 (4/5), 4–7.
Servais, T., Munnecke, A., Versteegh, G., 2009b. Silurian calcispheres (Calcitarcha) of Gotland
(Sweden): comparisons with calcareous dinoflagellates. CR Palevol 8, 527–534.
Sheehan, P.M., 1985. Reefs are not so different — they follow the evolutionary pattern of
level-bottom communities. Geology 13, 46–49.
 Author's personal copy
T. Servais et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 294 (2010) 99–119
Sheehan, P.M., 1996. A new look at Ecological Evolutionary Units (EEUs). Palaeogeo-
graphy, Palaeoclimatology, Palaeoecology 127, 21–32.
Sheehan, P.M., 2001a. History of marine diversity. Geological Journal 36, 231–249.
Sheehan, P.M., 2001b. The Late Ordovician mass extinction. Annual Reviews of Earth
and Planetary Sciences 29, 331–364.
Sheehan, P.M., Schiefelbein, J.D.R., 1984. The trace fossil Thalassinoides from the Upper
Ordovician of the eastern Great Basin: deep burrowing in the early Paleozoic.
Journal of Paleontology 58, 440–447.
Shields, G.A., Veizer, J., 2004. Isotopic signatures. In: Webby, B.D., Droser, M.L., Paris, F.,
Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event. Columbia
University Press, New York, pp. 68–71.
Shields, G.A., Carden, G.A.F., Veizer, J., Meidla, T., Rong, J.Y., Rong, Y.L., 2003. Sr, C, and O
isotope geochemistry of Ordovician brachiopods: a major isotopic event around the
Middle–Late Ordovician transition. Geochimica Cosmochimica Acta 67, 2005–2025.
Signor, P.W., Vermeij, G., 1994. The plankton and the benthos: origins and early history
of an evolving relationship. Paleobiology 20, 297–319.
Smith, A.B., 2001. Large-scale heterogeneity of the fossil record, implications for Phanerozoic
biodiversity studies. Philosophical Transactions of the Royal Society B 356, 1–17.
Smith, M.P., Donoghue, P.C.J., Sansom, I.J., 2002. The spatial and temporal diversification
of Early Palaeozoic vertebrates. In: Crame, J.A., Owen, A.W. (Eds.), Palaeobiogeo-
graphy and biodiversity change: the Ordovician and Mesozoic–Cenozoic Radia-
tions: Geological Society, London, Special Publications, 194, pp. 69–83.
Sprinkle, J., Guensburg, T.E., 2004. Crinozoan, blastozoan, echinozoan, asterozoan, and
homalozoan echinoderms. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G.
(Eds.), The Great Ordovician Biodiversification Event. Columbia University Press,
New York, pp. 266–280.
Stanley, S.M., 2007. An analysis of the history of marine animal diversity. Paleobiology
33, 1–55.
Stanley, S.M., Hardie, L.A., 1999. Hypercalcification; palaeontology links plate tectonics
and geochemistry to sedimentology. GSA Today 9, 1–7.
Stillman, C.V., 1984. Ordovician volcanicity. In: Bruton, D.L. (Ed.), Aspects of the Ordovician
System. Palaeontological Contributions from the University of Oslo, 295, pp. 183–194.
Stover, L.E., Brinkhuis, H., Damassa, S.P., Verteuil, L., De Helby, R.J., Monteil, E., Patridge,
A.D., Powell, A.J., Riding, J.B., Smelror, M., Williams, G.L., 1996. Mesozoic–Tertiary
dinoflagellates, acritarchs and prasinophytes. In: Jansonius, J., McGregor, D.G. (Eds.),
Palynology: Principles and Application. American Association of Stratigraphic
Palynologists Foundation. Publishers Press, Salt Lake City, Utah, USA, pp. 641–750.
Strother, P.K., 1996. Acritarchs. In: Jansonius, J., McGregor, D.G. (Eds.), Palynology:
Principles and Application. American Association of Stratigraphic Palynologists
Foundation. Publishers Press, Salt Lake City, Utah, USA, pp. 81–106.
Strother, P.K., 1973. A speculative review of factors controlling the evolution of
phytoplankton during Paleozoic time. Revue de Micropaléontologie 51, 9–21.
Su, W.B., 2001. On the Ordovician Sequence Stratigraphy and Sea-level Changes at the
Southeast Margin of the Upper Yangtze Platform. Geological Publishing House,
Beijing. 106 pp (in Chinese).
Tappan, H., Loeblich Jr., A.R., 1973. Evolution of the oceanic plankton. Earth-Science
Reviews 9, 207–240.
Taylor, P.D., Ernst, A., 2004. Bryozoans. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G.
(Eds.), The Great Ordovician Biodiversification Event. Columbia University Press,
New York, pp. 147–156.
Tetlie, E.O., 2007. Distribution and dispersal history of Eurypterida (Chelicerata).
Palaeogeography, Palaeoclimatology, Palaeoecology 252, 557–574.
Tetlie, E.O., Cuggy, M.B., 2007. Phylogeny of the basal swimming eurypterids (Chelicerata;
Eurypterida; Eurypterina). Journal of Systematic Palaeontology 5, 345–356.
Tobin, K.J., Bergström, S.M., 2002. Implications of Ordovician (= 460 Myr) marine
cement for constraining seawater temperature and atmospheric pCO2. Palaeogeo-
graphy, Palaeoclimatology, Palaeoecology 181, 399–417.
Tollerton, V.P., 2003. Summary of a revision of New York State Ordovician eurypterids:
implications for eurypterid palaeoecology, diversity and evolution. Transactions of
the Royal Society of Edinburgh, Earth Sciences 94, 235–242.
Torsvik, T.H., 1998. Palaeozoic palaeogeography: a North Atlantic viewpoint. GFF 120,
109–118.
Trotter, J.A., Williams, I.S., Barnes, C.R., Lécuyer, C., Nicoll, R.S., 2008. Did cooling oceans
trigger Ordovician biodiversification? Evidence from conodont thermometry. Science
321, 550–554.
Turner, S., Blieck, A.R.M., Nowlan, G.S., 2004. Cambrian–Ordovician vertebrates. In:
Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician
Bioiversification Event. Columbia University Press, New York, pp. 327–335.
Valentine, J.W., 1971. Resource supply and species diversity patterns. Lethaia 4, 51–61.
Valentine, J.W., Moores, E.M., 1972. Global tectonics and the fossil record. Journal of
Geology 80, 167–184.
Vandenbroucke, T.R.A., Armstrong, H.A., Williams, M., Zalasiewicz, J., Sabbe, K., 2000.
Ground-truthing Late Ordovician climate models using the paleobiogeography of
graptolites. Palaeoceanography 24 (PA4202), 1–19. doi:10.1029/2008PA001720.
Vandenbroucker, T.R.A., Armstrong, H.A., Williams, M., Sabbe, K., Zalasiewicz, J.A.,
Nõlvak, J., Verniers, J., 2010. Epipelagic chitinozoan biotopes map a steep latitudinal
temperature gradient for earliest Late Ordovician seas: implications for a cooling
Late Ordovician climate. Palaeogeography, Palaeoclimatology, Palaeoecology 294,
202–219.
Vannier, J., Racheboeuf, P.R., Brussa, E.D., Williams, M., Rushton, A.W.A., Servais, T.,
Siveter, D.J., 2003. Cosmopolitan arthropod zooplankton in Ordovician seas.
Palaeogeography, Palaeoclimatology, Palaeoecology 195, 173–191.
View publication stats
119
Van Roy, P., Orr, P.J., Botting, J.P., Muir, L.A., Vinther, J., Lefebvre, B., el Hariri, K., Briggs, D.E.G.,
2002. Ordovician faunas of Burgess Shale type. Nature 465, 215–218.
Vecoli, M., Le Hérissé, A., 2004. Biostratigraphy, taxonomic diversity and patterns of
morphological evolution of Ordovician acritarchs (organic-walled microphyto-
plankton) from the northern Gondwana margin in relation to palaeoclimatic and
palaeogeographic changes. Earth-Science Reviews 67, 267–311.
Veizer, J., Goddéris, Y., François, L.M., 2000. Evidence for decoupling of atmospheric CO2
and global climate during the Phanerozoic eon. Nature 408, 698–701.
Veizer, J., Ala, D., Azmy, K., Bruckschen, P., Buhl, D., Bruhn, F., Carden, G.A.F., Diener, A., Ebneth,
S., Goddéris, Y., Jasper, T., Korte, C., Pawellek, F., Podlaha, O.G., Strauss, H., 1999. 87Sr/86Sr,
δ13C and δ18O evolution of Phanerozoic seawater. Chemical Geology 161, 59–88.
Vennin, E., Alvaro, J.J., Villas, E., 1998. High-latitude pelmatozoan–bryozoan mud-mounds
from the Ordovician northern Gondwana platform. Geological Journal 33, 121–140.
Versteegh, G.J.M., Servais, T., Streng, M., Munnecke, A., Vachard, D., 2009. A discussion and
proposal concerning the use of the term calcispheres. Palaeontology 52, 343–348.
Waisfeld, B.G., Sánchez, T.M., Benedetto, J.L., Carrera, M.G., 2003. Early Ordovician
(Arenig) faunal assemblages from western Argentina; biodiversification trends in
different geodynamic and palaeogeographic settings. Palaeogeography, Palaeocli-
matology, Palaeoecology 196, 343–373.
Walker, L.J., Wilkinson, B.H., Ivany, L.C., 2002. Continental drift and Phanerozoic
carbonate accumulation in shallow-shelf and deep-marine settings. The Journal of
Geology 110, 75–87.
Walker, T.D., Valentine, J.W., 1984. Equilibrium models of evolutionary species diversity
and the number of empty niches. American Naturalist 124, 887–889.
Webby, B.D., 2002. Patterns of Ordovician reef development. In: Kiessling, W., Flügel,
E., Golonka, J. (Eds.), Phanerozoic Reef Patterns: SEPM Special Publication, 72,
pp. 129–179.
Webby, B.D., 2004. Stromatoporoids. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G.
(Eds.), The Great Ordovician Biodiversification Event. Columbia University Press,
New York, pp. 112–118.
Webby, B.D., Droser, M.L., Paris, F., Percival, I.G., 2004a. The Great Ordovician
Biodiversification Event. Columbia University Press, New York. 484 pp.
Webby, B.D., Cooper, R.A., Bergström, S.M., Paris, F., 2004b. Stratigraphic framework
and time slices. In: Webby, B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The
Great Ordovician Biodiversification Event. Columbia University Press, New York, pp.
41–47.
Webby, B.D., Elias, R.J., Young, G.A., Neumann, B.E.E., Kaljo, D., 2004c. Corals. In: Webby,
B.D., Droser, M.L., Paris, F., Percival, I.G. (Eds.), The Great Ordovician Biodiversifica-
tion Event. Columbia University Press, New York, pp. 124–146.
Westrop, S.R., Adrain, J.M., 1998a. Trilobite alpha diversity and the reorganization of
Ordovician benthic marine communities. Paleobiology 24, 1–16.
Westrop, S.R., Adrain, J.M., 1998b. Trilobite diversity and the Ordovician Radiation: a
reply to Miller et al. Paleobiology 24, 529–533.
Westrop, S.R., Trembley, J.V., Landing, E., 1995. Declining importance of trilobites in
Ordovician nearshore communities: dilution or displacement? Palaios 10, 75–79.
Whittaker, R.H., 1972. Evolution and measurement of species diversity. Taxon 21,
213–251.
Whittle, R.J., Gabbott, S.E., Aldridge, R.J., Theron, J.N., 2007. Taphonomy and
palaeocology of a Late Ordovician caryocaridid (Crustacea, Phyllocarida) from the
Soom Shale Lagerstätte, South Africa. Palaeogeography, Palaeoclimatology,
Palaeoecology 251, 383–397.
Wilde, P., 1991. Oceanography in the Ordovician. In: Barnes, C.R., Williams, S.H.
(Eds.), Advances in Ordovician Geology. Geological Survey of Canada, Paper, 90-9,
pp. 283–298.
Wilkinson, B.H., Owen, R.M., Carroll, A.R., 1985. Submarine hydrothermal weathering,
global eustasy, and carbonate polymorphism in Phanerozoic marine oolites. Journal
of Sedimentary Petrology 55, 171–183.
Wilson, M.A., Palmer, T.J., 1992. Hardgrounds and hardground faunas. Institute of Earth
Studies Publications, vol. 9. University of Wales, Aberystwyth. 131 pp.
Wilson, M.A., Palmer, T.J., 2002. The Ordovician bioerosion revolution. Geological
Abstracts with Programs 33, 248.
Wood, R., 1999. Reef Evolution. Oxford Univ. Press, Oxford. 414 p.
Young, G.C., 2009. An Ordovician vertebrate from western New South Wales, with
comments on Cambro-Ordovician vertebrate pattern. Alcheringa 33, 79–89.
Young, S.A., Saltzman, M.R., Bergström, S.M., 2005. Upper Ordovician (Mohawkian)
carbon isotope (δ13C) stratigraphy in eastern and central North America: regional
expression of a perturbation of the global carbon cycle. Palaeogeography,
Palaeoclimatology, Palaeoecology 222, 53–76.
Young, S.A., Saltzman, M.R., Bergström, S.M., Leslie, S.A., Chen, X., 2008. Paired δ13Ccarb
and δ13Corg records of Upper Ordovician (Sandbian-Katian) carbonates in North
America and China: implications for paleoceanographic change. Palaeogeography,
Palaeoclimatology, Palaeoecology 270, 166–178.
Young, S.A., Saltzman, M.R., Foland, K.A., Linder, J.S., Kump, L.R., 2009. A major drop in
seawater 87Sr/86Sr during the Middle Ordovician (Darriwilian): links to volcanism
and climate. Geology 37, 951–954.
Yue, L., Kershaw, S., 2004. Reef reconstruction after extinction events of the latest
Ordovician in the Yangtze platform, South China. Facies 48, 269–284.
Zhan, R.B., Jin, J., Rong, J.Y., 2006. β-diversity fluctuations in Early–Mid Ordovician
brachiopod communities of South China. Geological Journal 41, 271–288.
Zhuravlev, A.Y., Riding, R., 2001. The Ecology of the Cambrian Radiation. Columbia
University Press. 525 pp.