Review of Palaeobotany and Palynology 198 (2013) 2–13

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Review of Palaeobotany and Palynology

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Review papers

Reprint of ‘Eighty years of chitinozoan research: From Alfred Eisenack to

Florentin Paris’☆
Thomas Servais a,⁎, Aïcha Achab b, Esther Asselin c
a
Géosystèmes, UMR 8217 du CNRS, Université de Lille 1, SN5, F-59655 Villeneuve d'Ascq, France
b
Institut national de la recherche scientifique, INRS-ETE, 490, rue de la Couronne, Québec, QC G1K 9A9, Canada
c
Natural Resources Canada — Ressources Naturelles Canada, GSC-Quebec/CGC-Québec, 490, rue de la Couronne, Québec, QC G1K 9A9, Canada

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

Article history: In the early 1930s Alfred Eisenack first reported unknown, bottle-shaped, organic-walled microfossils that he had
Received 4 June 2012 discovered in erratic boulders from the south-eastern shores of the Baltic Sea. Eisenack erected the new group
Received in revised form 21 March 2013 Chitinozoa to classify these strange microfossils of unknown biological affinity. From the 1930s to the 1950s, a
Accepted 6 May 2013
few publications appeared reporting new findings and providing descriptions of these fossil organisms. It was
Available online 13 August 2013
only since the 1960s, with the development of the oil industry and the intensive biostratigraphical use of
Keywords:
organic-walled microfossils, that publications dealing with chitinozoans became more numerous and that the de-
Chitinozoa scription of new genera and species rapidly increased. The peak of description of new species was reached in the
zooplankton 1960s, but the number of publications remained high into the late 1990s. Since the 1990s the research activities
palynomorphs on chitinozoans are conducted by a much smaller number of scientists. One of the major driving forces of
Lower Palaeozoic chitinozoan research in the last forty years was Florentin Paris at the University of Rennes (Brittany, France).
He first established a high-resolution chitinozoan biostratigraphy of the Ordovician of southern Europe and
played an active role in bringing all scientists together for the development of global biostratigraphical schemes
and palaeobiogeographical scenarios of the Ordovician, Silurian and Devonian. It was also Florentin Paris, togeth-
er with his Estonian colleague Jaak Nõlvak, who suggested the now widely accepted biological interpretation that
Chitinozoa are most probably egg cases of a planktonic organism unknown from the fossil record. F. Paris was also
the first to collaborate w ith experts to use biogeochemical analyses and the C isotope signal of the chitinozoans
to better understand their biological affinity and detect biogeochemical changes in Palaeozoic oceans.
Crown Copyright © 2013 Published by Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Chitinozoan research since its beginnings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.1. The first discoveries: Alfred Eisenack . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.2. The first 40 years: from 1930 to the early 1970s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.3. The golden years of chitinozoan research and the contribution of Florentin Paris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Chitinozoa: a major palynomorph group in Palaeozoic rocks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.1. Taxonomical concepts and classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.2. Biological affinities of the chitinozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3.3. Biogeochemical analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4. Chitinozoans: a key group in Lower Palaeozoic biostratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4.1. Chitinozoan biostratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4.2. Impact of chitinozoan biostratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5. Biogeographical distribution patterns of the chitinozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6. Biodiversity studies and palaeoclimate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

☆ “This article is a reprint of a previously published article. A publishers' error resulted in this article appearing in the wrong issue. The article is reprinted here for the reader's convenience and
for the continuity of the special issue. For citation purposes, please use the original publication details; T. Servais et al./Review of Palaeobotany and Palynology 197 (2013) 143–151”.
DOI of original article: http://dx.doi.org/10.1016/j.revpalbo.2013.05.008.
⁎ Corresponding author. Tel.: +33 3 20 33 72 20; fax: +33 3 20 43 69 00.
E-mail address: thomas.servais@univ-lille1.fr (T. Servais).

0034-6667/$ – see front matter. Crown Copyright © 2013 Published by Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.revpalbo.2013.08.001
 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13 3

6.1. Chitinozoan biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6.2. Palaeoclimate: use of chitinozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7. Florentin Paris' impact on chitinozoan research and ‘northern Gondwanan’ studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

1. Introduction
Alongside the acritarchs (generally related to phytoplanktonic organ-
isms), the spores and pollen grains (related to land-plants) and the
scolecodonts (elements of the jaws of polychaetes), the chitinozoans
are considered today as one of the major groups of the Palaeozoic
organic-walled microfossils (palynomorphs). The group was first discov-
ered and described some 80 years ago by Eisenack (1930), who pro-
posed the name Chitinozoa for these bottle- or urn-shaped organic
microfossils. Since the 1960s the chitinozoans have become very impor-
tant in biostratigraphy, and today they are widely considered as one of
the major groups providing solid biostratigraphical correlations at local,
regional and global scales, with a similar or, in some intervals, even
more precise resolution than that of the graptolites and conodonts, con-
sidered as the two ‘classical’ biostratigraphic fossil groups of the Ordovi-
cian, Silurian and Devonian. In addition, chitinozoans have the advantage
that they can be easily found in both limestones and mudstones, unlike
graptolites and conodonts, respectively.
During the last 40 years, Florentin Paris, ‘Directeur de Recherches’ at
the Centre National de la Recherche Scientifique (CNRS) at the University
of Rennes (Brittany, France), was the driving force behind chitinozoan
research. He contributed to all aspects of chitinozoan investigations: tax-
onomy, biostratigraphy, palaeobiogeography, palaeoecology, palaeobio-
diversity and biogeochemistry. The stratigraphical information provided
by his work proved to be crucial for the understanding of regional geol-
ogy and palaeobiogeography, first of all of Brittany, but later of the entire
Armorican Terrane Assemblage, southern Europe and most areas of the
‘northern Gondwana domain’.
This paper attempts to outline the main trends in chitinozoan re-
search of the last 80 years, i.e. since the pioneering work by Alfred
Eisenack in the early 1930s to the standards reached at the beginning
of the 21st century. We do not attempt to provide an exhaustive list of
all the chitinozoan papers that have been published since 1930; readers
are invited to consult the listed bibliography to find the references not
cited herein. Because Florentin Paris was one of the major actors in
chitinozoan research during the last 40 years, this review will inevitably
highlight his contribution during that period. This review paper is the
first of a set of papers of a special issue on Palaeozoic marine
palynomorphs respectfully dedicated to Florentin Paris.
2. Chitinozoan research since its beginnings
2.1. The first discoveries: Alfred Eisenack
It was Alfred Eisenack, a German amateur collector, trained as a
chemist and working as a school teacher, who first discovered and de-
fined the Chitinozoa in the 1930s. He continued research after World
War II as a scientific collaborator at the University of Tübingen. In
the first half of the 20th century Eisenack studied the numerous fossils
contained in the erratic boulders strewn by glaciers along the south-
eastern coast of the Baltic Sea near the town of Königsberg in Eastern
Prussia, the present-day Kaliningrad, Russia. Many of the erratic
Palaeozoic boulders yielded macrofossils (trilobites, graptolites, etc.)
that allowed a precise age determination and an interpretation of
their geographical origin. However, many boulders and erratic stones
did not yield any macrofossils. Alfred Eisenack dissolved these appar-
ently unfossiliferous samples in hydrochloric or hydrofluoric acids
for the search of microfossils. From the residues he observed different
organic-walled microfossils, not only acritarchs (named at that time
‘hystrichospheres’), but also melanosclerites and new organic-walled
microfossil groups that he described in a series of scientific publica-
tions (Eisenack, 1930, 1931, 1932, 1934, 1937, 1942, etc.). He named
these new bottle- and urn-shaped microfossils, never described be-
fore, Chitinozoa, a name he created by linking the Greek terms ‘chitin’
(organic) and ‘zoa’ (animal). Eisenack discussed the possible biological
affinities of the chitinozoans and concluded that they were the organic
remains of animals, although it was later discovered that the chemical
composition of the chitinozoan wall does not include any real ‘chitin’
(Dutta et al., 2007; Jacob et al., 2007; Voss-Foucart and Jeuniaux,
1972). The Chitinozoa still remain an enigmatic group in terms of
their biological affinities, but their probable relationship to soft-
bodied animals is now generally accepted.
2.2. The first 40 years: from 1930 to the early 1970s
In the first three decades (1930–1960), the chitinozoans were exclu-
sively studied by using the light microscope. The investigations focused
on the description (taxonomy) of new taxa and their stratigraphical dis-
tribution. The number of specimens investigated was generally rather
small and many taxa have often been described on the basis of a few
specimens only. The biometrical studies that are needed to understand
the full morphological variability were usually absent in earlier publica-
tions, and only simple measurements were provided. Concepts of bio-
stratigraphy and palaeobiogeography were in their infancy and the
potential role of organic microfossils in palaeoenvironmental recon-
struction not yet well perceived.
Following Eisenack's discovery, pioneering studies took place in
France (Deflandre, 1944–1949) and in South America where Lange
(1949, 1952, 1967a,b) documented chitinozoans from Brazil, while in
North America Collinson and Schwalb (1955) and Collinson and Scott
(1958) described chitinozoans from the Devonian and, slightly later,
Wilson and Clarke (1960) reported the first Ordovician chitinozoans.
As documented by several authors (Jenkins, 1970a; Miller, 1996; Paris,
1996; Servais and Paris, 2000; Servais and Wellman, 2004; Taugourdeau,
1966; Taugourdeau et al., 1967) a revolution in chitinozoan studies took
place in the late 1950s and the early 1960s in response to the significant
demand for stratigraphical investigations by the oil industry. The Commis-
sion Internationale de Microflore du Paléozoïque (CIMP) was created in
1958 by Carboniferous spore and pollen workers to discuss taxonomic
concepts. This new international society, that reached over 500 members
in the 1970s to 1990s, erected sub-commissions for the different organic
microfossil groups in the early 1960s, including the Chitinozoan
Subcommission, that is still active today.
The 1960s and the early 1970s were a key period in chitinozoan re-
search. Numerous positions in palynology were created both in the oil
industry and in academia. Chitinozoans were extensively studied from
various regions, such as the Saharan Platform (Taugourdeau, 1961;
Taugourdeau and de Jekhowsky, 1960), Spain (Cramer, 1964, 1967;
Cramer and Díez, 1978; Díez and Cramer, 1978), Canada (Jansonius,
1964; Legault, 1973), Sweden (Laufeld, 1967, 1974), the United
Kingdom (Downie and Ford, 1966; Jenkins, 1967), Russia (Umnova,
1969), Belgium (Martin, 1969, 1973), the United States (Taugourdeau,
1965; Jenkins, 1967, 1970b; Urban and Kline, 1970; Urban, 1972;
Urban and Newport, 1973; Wood, 1974; Wright, 1980; Wood and
4 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
Clendening, 1985), Brazil (see Costa, 1974 for more references) and
Australia (Combaz and Peniguel, 1972).
In France, A. Combaz, B. de Jekhowsky, L. Magloire, P. Millepied, C.
Poumot and P. Taugourdeau played an important role in chitinozoan re-
search. Their contributions dealt with chitinozoan extraction techniques,
morphology and classification (Taugourdeau, 1966) and chitinozoan
biostratigraphy of the Lower Palaeozoic oil-bearing successions
(Taugourdeau and de Jekhowsky, 1960; Taugourdeau, 1961). As active
members of the Chitinozoan Subcommission of the CIMP, they produced,
under the auspices of the French “Centre National de la Recherche
Scientifique” (CNRS) two important documents summarizing the up-
dated knowledge on the group. The first contribution (Taugourdeau
et al., 1967) documented and illustrated all published chitinozoan genera
and species with their known stratigraphic occurrences, the second
(Combaz et al., 1967) focused on chitinozoan structure and morphology.
These two papers were significant milestones in chitinozoan research,
because they established the common language and terminology that
were used by most chitinozoan palynologists in the following years. Be-
side this important French group of scientists, Alfred Eisenack remained
an active worker in Germany. He continued to publish important mono-
graphs, and also proposed a new chitinozoan classification scheme
(Eisenack, 1968, 1972a,b). His last contribution on chitinozoans
appeared when he was well in his eighties (Eisenack, 1976).
2.3. The golden years of chitinozoan research and the contribution of
Florentin Paris
A new generation of chitinozoan scientists came to the fore in the
1970s, they undertook detailed palynostratigraphic studies and devel-
oped regional biozonations. Among them Florentin Paris (e.g. Paris
and Deunff, 1970) started studying the Ordovician acritarchs of Brittany
with Jean Deunff, a palynologist working at the Université of Rennes
(France). In the course of his research he realized that he was more
attracted by the enigmatic group of microfossils that were the
chitinozoans. By that time a few of the scientists were studying
chitinozoans only, most of the palynologists, especially those from the
oil industry, were working on both acritarchs, chitinozoans and Late
Palaeozoic spores and pollen grains.
The first chitinozoan investigations carried out by Florentin Paris fo-
cused on the regional geology of the Armorican Peninsula in western
France. While completing his PhD (‘Doctorat de 3ème cycle’) he joined
the very productive Lower Palaeozoic team at Rennes University. Later
he joined the CNRS, as a research associate. His research on chitinozoans
was well in line with the 1960–70s pioneering work carried out, as
noted above, in other countries, especially in North America, England,
and Scandinavia. These studies highlighted the stratigraphic potential
of chitinozoans and paved the way to detailed biostratigraphic studies.
Florentin Paris analyzed thousands of samples, and looked at tens of
thousands of chitinozoan specimens extracted from Ordovician, Silurian
and Devonian successions from southwestern Europe and Bohemia. He
classified them by using more adapted biometrical methods. Laufeld
(1974) was the first to use extensively the Scanning Electron Micro-
scope (SEM) for the observation of the chitinozoans from Gotland,
Sweden. He was followed by Florentin Paris who wisely exploited that
important change in microscopic observation techniques. Paris (1978)
published a laboratory procedure for the preparation and the storage,
after SEM observation, of chitinozoan specimens. This procedure is
still used by many chitinozoan workers. The advances in microscopy,
e.g. infrared and transmission electron microscopes, also appealed to
other workers to investigate the internal structural types and the ultra-
structure of the chitinozoan wall (e.g. Umnova, 1976; Grahn and
Afzelius, 1980; Mierzejewski, 1981).
The productive investigations of Florentin Paris during the 1970s led
him to publish his Doctor of Science dissertation entitled “Les
chitinozoaires dans le Paléozoïque du sud-ouest de l'Europe” in a special
issue of the “Société géologique et minéralogique de Bretagne” (Paris,
1981b). This monograph with over 400 pages and 40 photographic
plates has been up to now cited over 150 times in scientific journals
referenced in the current bibliographic databases, and it is still a key ref-
erence publication for all chitinozoan workers.
After the 1970s, Florentin Paris continued to contribute to the regional
geology of the Armorican Massif and other French areas (e.g. Paris and
Robardet, 1977; Paris, 1981a, Paris and Feist, 1983; Paris, 1986; Peucat
et al., 1986; Paris, 1988a; Paris and Robardet, 1990; Ballèvre et al., 1992;
Paris et al., 1996, 1999b; Trautmann and Paris, 2002) but he became
more and more involved in chitinozoan studies from various parts of
the ‘northern Gondwana Domain’ such as Portugal and Bohemia (e.g.,
Paris, 1979, 1981b; Paris et al., 1981; Paris and Kriz, 1984; Paris and
Mergl, 1984; Kriz et al., 1986), Morocco, Algeria and Libya (Elaouad-
Debbaj, 1984a,b; Boumendjel, 1985; Paris et al., 1985; Elaouad-Debbaj,
1986; Boumendjel, 1987; Elaouad-Debbaj, 1987; Elaouad-Debbaj et al.,
1987, Boumendjel et al., 1988; Elaouad-Debbaj, 1988; Paris, 1988b,c;
Streel et al., 1988; Oulebsir, 1992; Oulebsir and Paris, 1993, 1995;
Boumendjel and Paris, 1997; Paris et al., 2000a; Boumendjel, 2002; Jaglin
and Paris, 2002; Bourahrouh et al., 2004), Saudi Arabia (e.g. Paris and Al-
Hajri, 1995; Paris et al., 1995a; Al-Hajri and Paris, 1998; Paris et al.,
2000b), Turkey (e.g., Paris et al., 2007) and also from Podolia (e.g., Paris
and Grahn, 1996), Mauritania (Paris et al., 1998) and China (e.g., Tang
et al., 2007; Chen et al., 2008, 2009). These data contributed extensively
to the knowledge of the general geology, the geological history and the
palaeogeography of the investigated areas, and in particular the ‘northern
Gondwana Domain’.
During the same period numerous biostratigraphic studies based on
Palaeozoic chitinozoans were ongoing at several places around the
world. In France, Rauscher (1974a,b) investigated chitinozoans and
acritarchs from several areas. Martin (1969, 1973) published her first
chitinozoan studies of Belgium. The chitinozoan literature of this period
is very abundant, in particular from Baltoscandia and North America. In
Baltoscandia Y. Grahn and J. Nõlvak published on the chitinozoan asso-
ciations of the Ordovician, V. Nestor on those of the Silurian, while R.
Wrona documented those of the Upper Silurian–Lower Devonian in
southeast Poland (see detailed reference list in Wrona, 1980; Nõlvak
and Grahn, 1993; Nestor, 1994; Grahn and Nõlvak, 2010).
In North America, the most informative chitinozoan studies on the
Ordovician and the Ordovician–Silurian boundary were conducted in
the United States by Miller (1976), Grahn and Bergström (1984,
1985), Grahn (1985), Grahn and Miller (1986) and Hart (1986), and
in Canada by Neville (1974), Martin (1975, 1983), Achab (see Achab,
1989, for more references), Melchin (1982) and Melchin and Legault
(1985). Jenkins and Legault (1979) provided an up-date of the Lower
Palaeozoic chitinozoan species. Other studies dealt with Spitsbergen
(Bockelie, 1980), Russia and Siberia (Umnova, 1969, Zaslavskaya,
1980; Umnova, 1981, Zaslavskaya, 1983), Belgium (Verniers, 1982),
and China (Geng, 1984; Geng and Li, 1984; Geng et al., 1984; Geng,
1986; Geng et al., 1987; Fan and Hou, 1988; Geng and Cai, 1988;
Geng, 1990; Grahn and Geng, 1990; Wang and Chen, 1992, 1994;
Chen, 1994; Chen and Wang, 1996; Chen et al., 1996).
A common approach was behind most of these studies: chitinozoan
assemblages were carefully described and their stratigraphic occurrence
precisely documented. The biostratigraphic ranges of the chitinozoans
were related to those of the graptolites and conodonts and other fossils
in order to correlate precisely the chitinozoan biozones with the geologi-
cal time scale. These results have highlighted the high stratigraphic reso-
lution of chitinozoans and their stratigraphic value for local and long
distance correlations. On the basis of his studies Paris (1990) developed
a precise regional Ordovician chitinozoan biozonation for the northern
Gondwana margin, while Achab (1989) did the same for Laurentia and
Nõlvak and Grahn (1993) for Baltoscandia. These biozonations became
subsequently the regional standard chitinozoan zonation for these three
palaeocontinents (Paris et al., 2004; Webby et al., 2004b).
Communication between the members of chitinozoan community
was always harmonious, respectful and productive. It led to new
 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
developments in biostratigraphy, biodiversity and biological affinities of
chitinozoans. Florentin Paris was an active member of the community
collaborating with several colleagues such as Y. Grahn (Sweden and
South America), V. Nestor and J. Nõlvak (Estonia), J. Verniers
(Belgium), K. Boumendjel (Algeria), Z. Elaouad Debbaj (Morocco), T.
Winchester-Seeto (Australia) and A. Achab, E. Asselin and A. Soufiane
(Canada), among many others. These collaborations were productive
and resulted in many multi-co-authored publications.
Chitinozoan research continued to be productive during the next
two decades (1990–2010). The team at Ghent University (Belgium),
led by J. Verniers, documented Ordovician and Lower Silurian
chitinozoans from Avalonia in Belgium and the United Kingdom
(e.g. Van Grootel, 1990; Ancilletta, 1997; Verniers, 1999; Samuelsson
and Verniers, 2000; Samuelsson et al., 2000; Samuelsson and Servais,
2001, 2002; Verniers et al., 2002; Vanmeirhaeghe et al., 2005, 2006;
Vanmeirhaeghe, 2006; Verniers and Vandenbroucke, 2006;
Vandenbroucke, 2008; Vandenbroucke et al., 2009a). These studies
allowed to provide the basis of the global Silurian chitinozoan zonation
(e.g. Verniers et al., 1995, 2008), and also that of the Ordovician in
Avalonia (Vandenbroucke, 2008). In the United Kingdom, the
Llandovery and Wenlock Series were also documented by Swire
(1990), Verniers (1999), Mullins (2000), Mullins and Loydell (2001,
2002) and Mullins and Aldridge (2004), while Sutherland (1994) inves-
tigated the Ludlow Series. In the United States, Loydell et al. (2002,
2007), Butcher et al. (2010) and V. Nestor in Won et al. (2002) pub-
lished chitinozoan data on either Late Ordovician–Early Silurian, or Silu-
rian successions from the Illinois, New York and Alaska states. In the
meantime, Dufka (see, 1995 on his previous contributions) described
Early Silurian chitinozoans of the Prague basin, while Nõlvak (1999,
2002), Nõlvak and Grahn (1993), Nõlvak et al. (1999), Nestor (2005),
Loydell and Nestor (2005) and V. Nestor and J. Nõlvak in Rubel et al.
(2006) continued documenting them in the Baltoscandia area.
Chitinozoans from the Ordovician, Silurian and Devonian succes-
sions of the South American basins were also well documented by the
works of Grahn and his collaborations with other colleagues. For a
more exhaustive reference list about South American chitinozoan
data, we refer the readers to Grahn (1992, 2006), Grahn and Paris
(1992), Grahn and Gutiérrez (2001), Grahn and Melo (2002), Grahn
et al. (2000, 2003, 2010), Heuse et al.(1999), Wood (1994, 2004) and
Wood and Miller (1991) These studies resulted in regional chitinozoan
biozonations for the Ordovician, Silurian (Grahn, 2006) and Devonian
(Grahn, 2005; Grahn and Melo, 2004; Grahn et al., 2005, 2006) of
‘Northwestern Gondwana’.
In North America, Achab and Asselin (1995), Achab et al. (1997,
2003, 2011) and Soufiane and Achab (2000a,b) focused on Ordovician,
Silurian and Lower Devonian successions from eastern and northern
Canada (Anticosti, Gaspé peninsula, eastern-central Arctic) and western
U.S.A. (Nevada), whereas Albani et al. (2001) studied Middle Ordovician
chitinozoans from Newfoundland.
From Australia, Winchester-Seeto and Paris (1989), Winchester-
Seeto (1993a,b, 1996), and Winchester-Seeto et al. (2000) described
Ordovician and Devonian chitinozoans, while Quintavalle and
Playford (2006) reported Ordovician chitinozoans from the Canning
Basin. Several contributions also documented chitinozoan biostra-
tigraphy, diversification and palaeogeography from China (Wang
and Chen, 2004; Chen and Zhang, 2005; Tang et al., 2007; Chen
et al., 2008, 2009). Ghavidel-syooki and Vecoli (2007) reported on
chitinozoans from Iran, while Butcher (2009) reported on those
from Jordan.
3. Chitinozoa: a major palynomorph group in Palaeozoic rocks
3.1. Taxonomical concepts and classification
Of the different Palaeozoic marine organic-walled microfossil
groups the chitinozoans are very important, with over 1200 papers
5
describing chitinozoans published up to the 2000s. After the first
species defined by Eisenack in the 1930s, only a few species were de-
scribed between 1940 and 1955, but the number of new species ex-
ploded in the last part of the 1950s and in particular in the 1960s,
with over 100 new descriptions during this decade (Servais and
Paris, 2000). These numbers were never reached again in the
1980s and 1990s.
It is worth mentioning that the low number of chitinozoan workers
has not only permitted the close collaboration between scientists men-
tioned above, more importantly, it favoured widely accepted taxonomy
and biostratigraphic concepts. The more or less stable, well structured
and commonly used taxonomy of the chitinozoans (that are considered
by most authors as probably a monophyletic group) facilitates their use
for biostratigraphic purposes at the difference of other groups, such as
the acritarchs, whose taxonomy is still highly problematic and partly
chaotic. This is probably due to the fact that acritarchs, in contrast to
the chitinozoans, are clearly a polyphyletic group, and also because a
large number of specialists use different classification concepts
(Servais, 1996).
It is important to remember that the morphological classification of
chitinozoans is entirely artificial, being based on parataxonomy.
Chitinozoan classification is, as in most fossil groups, based solely on
morphological parameters that allow the creation of (morpho-) species,
(morpho-) genera and also suprageneric ranks that have actually no
true biological meaning. Nevertheless, chitinozoan classification has
benefitted from the stable principles and the logical hierarchy of mor-
phological features on which the classification is built. The fundaments
of chitinozoan classification were first established by Alfred Eisenack
(1931), reviewed later by Taugourdeau (1966), Jansonius (1970),
Eisenack (1972b) and Paris (1981b). We refer the readers to Miller
(1996) who discussed the evolutive change concerning the chitinozoan
classification between 1930 and 1981.
However, it was Florentin Paris (1981b) who proposed the more
elaborate and better structured classification framework on the basis
of a very high number of observations. He adapted the suprageneric
classification schemes of Eisenack (1931, 1972b) and Taugourdeau
(1966) and continued to follow, as suggested by Eisenack, the rules of
the International Code of Zoological Nomenclature (ICZN), in contrast
with acritarch taxonomy that follows the rules of the International
Code of Botanical Nomenclature (ICBN). Most chitinozoan workers
adopted and still follow the emended classification of Paris (1981b)
that became the standard presented in most palynology textbooks
(Miller, 1996). This classification scheme uses as highest taxonomic
subdivision the order. The order Operculatifera Eisenack, 1972b re-
groups all chitinozoans with an operculum whereas the order
Prosomatifera Eisenack, 1972b regroups all those with a prosome. The
lower ranks, commonly used by most chitinozoan workers, are the fam-
ilies (e.g., Desmochitinidae Eisenack, 1931, emend. Paris, 1981b), the
subfamilies (e.g., Desmochitininae Paris, 1981b), followed by the genera
and the species. Infraspecific ranks have also been commonly used in
the past, including subspecies and formas, but are currently avoided
and discouraged.
Florentin Paris was once more the leader of a research team that pro-
vided a revised classification of the chitinozoan group (Paris et al.,
1999a). The latter authors proposed a complete revision of all published
genera. They checked the definitions of the so far validly published 143
genera and subgenera and kept only 56 genera as valid. Many seemed to
have been junior synonyms or invalidly defined taxa. Paris et al. (1999a)
also proposed a suprageneric classification of the whole Chitinozoa
group based on diagnostic features whose hierarchy was established
on statistical and evolutionary grounds. This scheme has been followed
by the chitinozoan workers although since then, a few new genera have
been proposed. A few chitinozoan workers, however, preferred to use
an alphabetical listing of all taxa (as commonly used for the acritarchs)
because they consider a suprageneric classification of organisms of un-
known biological affinities to be meaningless.
6 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
3.2. Biological affinities of the chitinozoans
Even eighty years after the discovery of the first chitinozoans and sev-
eral decades of investigations to understand their biological affinities, it is
still unknown what the chitinozoans represent in terms of biological af-
finities. The reader can be referred to Paris (1981b), Taugourdeau
(1981), Miller (1996) and Paris and Nõlvak (1999) for a more detailed
history of the various interpretations of the biological affinities of the
chitinozoans, for which only a summary is provided below.
Eisenack (1930), in describing the first specimens, attributed the mi-
crofossils to a new group, convinced that they belonged to the Animal
Kingdom. Subsequently, Eisenack (1931) proposed that chitinozoans
are related to protozoans of the rhizopod Order Testacea, a hypothesis
later rejected by Eisenack (1932) himself because testaceans are
fresh-water organisms with a cellulosic test.
Chitinozoans have been subsequently compared with, and assigned
to, various groups. Their morphological characters, wall ultrastructure
and composition, and palaeoenvironmental and stratigraphic distribu-
tion were considered as evidence for various hypothetical affinities,
none of which has been conclusive. During the last eighty years,
chitinozoans have been successively assigned to various groups of pro-
tozoans, metazoans, protists (see Taugourdeau, 1981) and fungi
(Locquin, 1981). We refer the readers about this topic to Taugourdeau
(1981), Miller (1996) and Paris and Nõlvak (1999). The interpretations
by Cashman (1990, 1991) have been proven to have used falsified evi-
dence and were largely rejected by all other chitinozoan workers.
Paris (1981b) was the first to use not only the morphological simi-
larity to try to understand the biological affinity of the group. He also ex-
amined the contemporaneity of the time ranges of the known fossil
groups occurring between the Ordovician and Devonian with those of
the chitinozoans. Neither the graptolites nor another group matched,
however.
Chitinozoans are hollow and are often flattened; they may be excep-
tionally preserved in three dimensions. Paris et al. (1996) and
Munnecke and Servais (1996) described some very well preserved
specimens filled with calcite or sediments. That thus could suggest
that chitinozoan vesicles may have contained the ‘eggs’ of an unknown
organism. As such, they have been considered by some authors as ‘egg
cases’ or ‘egg capsules.’ The arrangement of chitinozoan tests in elongate
organic-walled sheets (‘cocoons’) was the reason why Kozlowski
(1963) considered them as analogous to eggs or cysts.
This ‘metazoan egg hypothesis’ was popular for many years (e.g.,
Laufeld, 1974; Grahn, 1981; Paris, 1981b), partly because some marine
invertebrates, such as some mollusks or some polychaetes, produce
long, linear or coiled chains of eggs. Grahn (1981) proposed the infor-
mal name ‘chitinozoophoran animal’ to designate the hypothetical or-
ganism that could have produced the chitinozoans, although this
‘chitinozoan animal’ has yet to be discovered (see also Grahn and
Paris, 2011). Several authors have been looking for the chitinozoan-
producing animal among fossil marine invertebrates. A recurrent inter-
pretation, some decades ago, was that chitinozoans might be related to
the planktonic graptolites (e.g., Jenkins, 1970a), because both groups
have a similar stratigraphical range. In detail, however, no direct corre-
lations are possible, as indicated above (Paris, 1981b; Paris and Nõlvak,
1999).
Although the chitinozoan-producing organism remains unknown
the ‘egg case hypothesis’ is still the most plausible interpretation, gener-
ally accepted by almost all specialists since the early 1980s. Only
Cashman (1990, 1991) proposed a different view, considering the
chitinozoans ‘juvenile’ Rhizopoda, an interpretation proposed first by
Eisenack (1931, see above), but not retained by most chitinozoan spe-
cialists (see Taugourdeau, 1981; Miller, 1996; Paris and Nõlvak, 1999).
The debate as to whether chitinozoans are a monophyletic or a poly-
phyletic group has also been animated, and will certainly go on for a
while as long as their biological affinities remain unresolved. Although
it is difficult to recognize phylogenetic relationships and evolutionary
lineages in organisms of uncertain biological affinities, and although it
appears difficult to propose phylogenetic trends for microfossils that
are considered as ‘eggs or ‘cysts,’ several authors have proposed evolu-
tionary lineages for the chitinozoans, based on the succession of mor-
phological features appearing in a clear stratigraphical order. Such
lineages (Paris, 1981b; Melchin and Legault, 1985; Miller, 1996, text-
fig. 10; Vandenbroucke et al., 2009a) have nevertheless the merit of
displaying the stratigraphic succession of the most significant
morphotypes, although the interpretation of evolutionary lineages
should be considered with care.
3.3. Biogeochemical analyses
The chemical composition of the chitinozoan vesicle wall is not fully
understood. Chitinozoans were considered to be composed of ‘chitin-
ous’ or ‘pseudochitinous’ organic matter, which also resulted in provid-
ing the name to the group. Voss-Foucart and Jeuniaux (1972) were
among the first to perform biogeochemical analyses on chitinozoan
walls and to show that, at least in the case of Cyathochitina
campanulaeformis, one of the most commonly distributed species, the
chitinozoan wall did not contain any chitin. Florentin Paris was also
highly interested by the biogeochemical analyses. During several at-
tempts he collected huge amounts of monospecific assemblages of
well preserved chitinozoans with nearly no thermal alteration and pro-
vided them to several types of chemical analyses (micro-FTIR, laser
micro-Raman spectroscopy and laser micropyrolysis–gas chromatogra-
phy–mass spectrometry). In these studies, Florentin Paris and his col-
leagues could prove that the composition of the chitinozoans was a
kerogen network dominated by aromatic groups with a low contribu-
tion of aliphatics and oxygen- or nitrogen-bearing compounds. No
chitin-related product was found, as was already postulated before. As
a result, Florentin Paris contributed to the very important paper with
Jacob et al. (2007) summarizing the chemical composition of the
chitinozoans. At about the same time, geochemical analysis by Dutta
et al. (2007) provided similar results and showed the highly aromatic
character of the chitinozoan vesicles. In addition, Florentin Paris also ini-
tiated a δ13C analysis on chitinozoan vesicles in order to determine the
isotopic fractionations among the different extinct palynomorph groups
(Lécuyer and Paris, 1997) and, on a larger extent, collaborated on pro-
jects dealing with biogenic phosphates (e.g., Lécuyer et al., 1996, 1998).
4. Chitinozoans: a key group in Lower Palaeozoic biostratigraphy
4.1. Chitinozoan biostratigraphy
Some fossil groups, such as the ammonites of the Jurassic and Creta-
ceous, are considered stratigraphic index fossils. In the Lower
Palaeozoic, the classical stratigraphic fossils for the Cambrian are the tri-
lobites, while the planktonic graptolites were considered for many years
the most important index fossils for the Ordovician, Silurian, and Lower
Devonian (Lapworth, 1879; Webby, 1998). With the development of
micropalaeontology in the second half of the 20th century, the
conodonts also became a very important biostratigraphic tool for the
entire Palaeozoic and were used extensively for international correla-
tions. For most Ordovician stratotypes, including the Global Boundary
Sections and Points (GSSP), the first occurrence of either a graptolite
or a conodont species is used (Webby, 1998; Webby et al., 2004b;
Bergström et al., 2009).
In order to serve as an index fossil in global correlations, a fossil
taxon must not only be abundant and easy to recognize, but must also
display a wide geographic distribution. Planktonic microfossils display
these characteristics, and may be considered as stratigraphic index
guides, if their palaeogeographic distribution is well understood. With
the development of the oil industry, acritarchs and chitinozoans have
been intensively used in biostratigraphic studies in the Lower
Palaeozoic. It should be however noted that today, while the use of
 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
acritarchs is still limited, the chitinozoans are playing a significant role,
in particular in Ordovician, Silurian and Devonian successions (Verniers
et al., 1995; Paris, 1996; Paris et al., 2000c; Servais and Paris, 2000).
Chitinozoans are considered today as one of the three major
biostratigraphically useful fossil groups for the Ordovician and Silurian,
being important not only in recognising the GSSPs (e.g., Paris et al.,
1981; Verniers and Vandenbroucke, 2006), but also in defining time-
slices and stage boundaries (Webby et al., 2004b; Bergström et al.,
2009). This is clearly the result of the detailed biostratigraphic studies
carried out on well-dated geological sections during the last decades.
These studies have shown that chitinozoans, by their occurrence in al-
most all marine sediments and the short stratigraphic ranges of some
species, are a high-resolution stratigraphic tool that can easily compete
with graptolites and conodonts. Among these studies, those conducted
by Florentin Paris on Ordovician, Silurian and Devonian chitinozoan
faunas from various areas of the ‘northern Gondwana palaeocontinent’
are of great importance; they contributed extensively to the knowledge
of the general geology and the geological history of this domain.
Florentin Paris' knowledge of Ordovician successions allowed him to
be for many years a titular member of the International Subcommission
of Ordovician Stratigraphy (ISOS) and of the International Subcommission
of Silurian Stratigraphy (ISSS). He was also one of the co-leaders of the In-
ternational Geoscience Programme (IGCP) No. 410 ‘The Great Ordovician
Biodiversification Event’. Animated by Barry Webby (Sydney, Australia),
Mary Droser (Riverside, USA) and Florentin Paris, this programme
brought together scientists from all over the world during the years
1997–2002 (Webby et al., 2004a). To fulfil the objectives of this project
that attempted to understand the diversification trends of all fossil groups
from all palaeocontinents, the need for an integrated, well-calibrated Or-
dovician time-scale was crucial. The regional chitinozoan biozones were
used at the same level as those based on graptolites and conodonts to es-
tablish the stratigraphic framework and to subdivide the Ordovician into
19 times-slices, that constituted the reference stratigraphic scale for
documenting faunal diversities (Webby et al., 2004b).
As co-leader of IGCP 410 and leader of the “Chitinozoan Clade”,
Florentin Paris convened almost all chitinozoan specialists (Paris et al.,
1999c) to participate in this IGCP project and to build regional
chitinozoan databases for Gondwana, Baltica, Laurentia and Avalonia.
This was a prerequisite for correlating the Ordovician regional
biozonations and documenting chitinozoan diversity through the entire
Ordovician (Paris et al., 2004).
Although Florentin Paris' main interest was in Ordovician
chitinozoan stratigraphy, he was also active in the development of Silu-
rian and Devonian biostratigraphic schemes, as for example, his collab-
orative work on the Siluro-Devonian boundary in Bohemia (Paris et al.,
1981). He was also an important member of the team that developed
the global chitinozoan biozonation for the Silurian (Kriz et al., 1986;
Verniers et al., 1995), and took a leading role in the establishment of
the global chitinozoan biozonation of the Devonian (e.g., Paris et al.,
2000c).
4.2. Impact of chitinozoan biostratigraphy
The impact of a fossil group can be estimated by the biostratigraphic
information it provided on sediments or stratigraphic units of unknown
age, in particular in areas where the geological history is complex.
Chitinozoan data allowed Montenari et al. (2000) to establish a Silurian
age for metasediments located in the Black Forest in Germany, whose
age had long been totally unknown. This age assignment permitted a re-
consideration of the geodynamic evolution of this important part of the
European Variscan chain. Chitinozoans were also used for precisely dat-
ing the Ordovician impact craters in Baltoscandia (Grahn et al., 1996),
and in providing a precise age of the Ordovician Fossil-Lagerstätte of
the Soom Shale (Vandenbroucke et al., 2009b). Chitinozoans also
helped to provide a precise age of the first occurrence of trilete spores
in the Katian of the Arabian Peninsula (Steemans et al., 2009).
7
Another example is the contribution of chitinozoans, together with
sequence stratigraphy, to the regional correlations of the Upper Ordovi-
cian Ellis Bay Formation of Anticosti Island, Canada, or the Upper Ordo-
vician sequences of western Estonia, in the context of the understanding
of the global Late Ordovician extinction (Kaljo et al., 2008; Achab et al.,
2011, Kaljo et al., 2011). The biozonation established by Florentin Paris
for the Late Ordovician was instrumental in deciphering the timing
and tempo of the Hirnantian glaciation by showing that most of the
‘North Gondwanan’ glacial deposits belong to the same chitinozoan
biozone (Paris et al., 1995b, 1998; Bourahrouh et al., 2004). It also led
to the exact location and the dating of the glaciation in Turkey
(Monod et al., 2003). The collaboration between palynologists and sed-
imentologists specialized in sequence stratigraphy (e.g., Ghienne et al.,
2007; Le Héron et al., 2008; Desrochers et al., 2010; Loi et al., 2010)
also led to a better understanding of the glacial dynamics not only
from the Gondwanan area, but also from those regions at lower lati-
tudes without glacial deposits (Harris et al., 2004; Achab et al., 2011).
Such integrated biostratigraphy studies now also include the Silurian
(Davies et al., 2013). Chitinozoan data also contributed to the under-
standing of the sedimentologic and the stratigraphic frameworks of
Palaeozoic sequences, as illustrated by the collaborative work of
Florentin Paris on the northern Gondwanan domain from Brittany and
North Africa (e.g. Dabard et al., 2007; Videt et al., 2010).
5. Biogeographical distribution patterns of the chitinozoans
As a fossil group commonly interpreted as belonging to the plank-
ton, the chitinozoans have for a long time been considered to be of lim-
ited use in biogeographic reconstructions. Like the graptolites and the
acritarchs, they have been considered (e.g., Fortey and Mellish, 1992)
to be geographically widespread and often of cosmopolitan distribution,
which is an advantage for global correlations, but a disadvantage for de-
fining biogeographic provinces or establishing palaeogeographic
boundaries. However, owing to the accumulation of data through
time, it was possible to demonstrate that the chitinozoans also display
palaeogeographic and palaeolatitudinal distribution patterns. Achab
(1988, 1989) and Achab et al. (1992), for example, were the first to doc-
ument the latitudinal distribution of chitinozoans and the presence of
chitinozoan provinces, which were confirmed by Paris and others
(Paris, 1991, 1993; Oulebsir and Paris, 1995; Paris et al., 1995a; Paris,
1996; Paris et al., 1998; Paris, 2008b) and Miller (1996). Today, it is
well understood that geographical distribution patterns do exist, and
that chitinozoans can be used as palaeobiogeographic tools, as other
planktonic and even nektonic organisms (Servais et al., 2005). In the Or-
dovician, up to now, three major distinct chitinozoan provinces have
been recognized; they correspond to the margins of the three major Or-
dovician palaeocontinents: Gondwana, Baltica and Laurentia (Achab
and Paris, 2007). Again, Florentin Paris has been an active contributor
in these developments of chitinozoan research.
6. Biodiversity studies and palaeoclimate
6.1. Chitinozoan biodiversity
Research in micropalaeontology and palynology has undergone a cy-
clic evolution through time. Strongly supported by the oil industry in
the second half of the 20th century, micropalaeontological studies
were mostly focused on stratigraphy, and, since most palynologists
were stratigraphers, this became, as indicated above, the main trend
in chitinozoan research. At the beginning of the 1980s an additional
focus became important for Palaeozoic palaeontologists: the study of
palaeogeography and the attempt to produce palaeogeographical re-
constructions, which considered a particular interesting task for pre-
Pangaean times.
In the two last decades a further major research question animated
the scientific debate among geologists and palaeontologists: the
8 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
question of global climate change. With the debate on the impact on our
society on the global climate, questions about the drop of biodiversity
are constantly being raised in the media. To better understand the mod-
ern changes of biodiversity, scientists seek to learn from the past by an-
alyzing fossil sequences that might have worked in an analogous way.
For these reasons, most palaeontological and micropalaeontological
studies today are focused on Quaternary and sub-Recent fossils, as
well as on the problem of the impact of Late Quaternary climate changes
on global diversity. The Ordovician mass extinction (that is usually cor-
related with the Hirnantian glaciation) can also be compared to the
Quaternary (although it is not a direct analogue), and geologists and
palaeontologists working in the Ordovician might be able to contribute
to this issue. Can the first of the ‘Big Five’ extinctions during Earth Histo-
ry be compared with the modern world?
In the late 1990s, the International Geological Correlation Pro-
gramme (IGCP) 410, was proposed precisely to understand the ‘Great
Ordovician Biodiversification Event’ (GOBE). Beside a major contribu-
tion to the Ordovician stratigraphic framework and the definition of
time slices (Webby et al., 2004b), the chitinozoan team (Paris, 1999;
Paris et al., 1999c, 2004) depicted the chitinozoan biodiversification pat-
terns for the three major palaeocontinents that were Gondwana,
Laurentia and Baltica and proposed a global curve integrating all region-
al data.
Following this very successful IGCP 410, a new IGCP project was
launched in 2004 to understand not only the geological and biological
factors that triggered the Ordovician biodiversification, but also the Late
Ordovician extinction and the succeeding Silurian radiation. This IGCP
project no. 503 named ‘Ordovician Palaeogeography and Palaeoclimate’
run from 2004 to 2009 (Harper et al., 2011). It attempted to answer the
question of the impact of changing palaeogeography and changing
palaeoclimate on the biodiversity trends, resulting in several review pa-
pers that summarize the geochemical background (Munnecke et al.,
2010) and the palaeoecological context (Servais et al., 2010). A major
contribution on the chitinozoans to IGCP 503 was the paper published
by Achab and Paris (2007), which, on the basis of the data acquired by
the IGCP 503 chitinozoan team (Paris et al., 2004), summarized the diver-
sity trends of the chitinozoans and the triggering factors of the diversity
increases and drops, between the first occurrence of the chitinozoans in
the earliest Ordovician to the Hirnantian extinction. The authors
questioned all possible causes and factors that could have influenced
the evolution of the group, including palaeogeography, sea-level changes,
climatic trends, volcanism, etc. In a more recent paper, Grahn and Paris
(2011) discussed in detail the emergence, biodiversifaction and the ex-
tinction of the chitinozoans and the chitinozoophorans.
This and similar contributions on other fossil groups made it possible
to better understand the evolution of the fossil food chains. The
chitinozoans, together with the planktonic graptolites and the radiolar-
ians, evolved very rapidly during the Ordovician, probably because of
the increasing presence of the phytoplankton, which was probably an
important component at the base of the food chain (Servais et al.,
2008, 2009, 2010). Grahn and Paris (2011) noted that the marine inver-
tebrates, probably producing the chitinozoans, called chitinozoophorans,
were most likely to have been small, pelagic or necto-pelagic, soft-
bodied, probably wormlike animals, measuring from a few millimetres
to a few centimetres in length. Most probably there must have been a re-
lation between the zooplanktonic chitinozoophorans and the phyto-
plankton, which most probably served as food to the chitinozoan
producing invertebrate organism. Grahn and Paris (2011) also pointed
out that there is no evidence of a large colonization of the pelagic niche
in the Cambrian, but from the Early Ordovician onward, this niche was
exploited chiefly by graptolites and chitinozoophorans. Both groups
inhabited nearshore and offshore habitats, but in contrast to the grapto-
lites, the chitinozoans displayed their highest diversity and abundance at
high latitudes, in less distal environments (Grahn and Paris, 2011).
Hints et al. (2010) were able to document a very close correlation
between the diversity trends of all marine invertebrates of the
palaeocontinent Baltica and that of the acritarchs, chitinozoans
(chitinozoophorans) and scolecodonts. The chitinozoan-producing or-
ganism did probably feed on (bacterio-, pico- and) phytoplankton and
was also probably part of the food chain that allowed larger animals,
such as fishes and cephalopods, to develop and diversify rapidly during
the Ordovician (Kröger et al., 2009). The chitinozoans are, thus, together
with other groups of zooplanktonic animals, such as the graptolites and
radiolarians, a key element of the ‘Ordovician plankton revolution’
(Servais et al., 2008) that completely changed the life in the oceans
and probably was a major trigger of Great Ordovician Biodiversification
Event, GOBE (Paris, 2008a; Servais et al., 2009, 2010). It is thus today
considered as evident that the chitinozoans, so far absent in the Cambri-
an seas, filled the Ordovician oceans rapidly and became an important
part of the plankton.
6.2. Palaeoclimate: use of chitinozoans
Many microorganisms and microfossil groups have been used in the
last decades to depict climatical and palaeoclimatical changes, in partic-
ular in the Cenozoic. Achab and Paris (2007) considered a series of geo-
logical factors to have had a major impact on the climate changes. The
chitinozoan palaeobiodiversity fluctuations, illustrated by radiation
and extinction phases, could be correlated with these geological and
palaeoclimatical changes. It was suggested that acritarch and
chitinozoan distribution is controlled by palaeolatitudes (Achab, 1991;
Paris, 1991; Achab et al., 1992; Paris, 1996). However, more recently
Florentin Paris contributed to the mapping of the chitinozoan distribu-
tion in the Late Ordovician which enabled the detection of a latitudinal
temperature gradient (Vandenbroucke et al., 2010a) or the shifting of
the polar front during the Late Ordovician (Vandenbroucke et al.,
2010b). In addition to their use in palaeobiogeography, chitinozoans
may thus be able to play a more important role than previously thought
in Lower Palaeozoic climate studies.
7. Florentin Paris' impact on chitinozoan research and ‘northern
Gondwanan’ studies
This short review on chitinozoan research, from the first discovery in
1930 to the present day, clearly shows that Florentin Paris was one of
the major contributors of chitinozoan studies in the last part of the
20th and the beginning of the 21st centuries. At the beginning of his ca-
reer he focused his research on the detailed stratigraphic documenta-
tion of chitinozoan faunas, establishing a high-resolution chitinozoan
biostratigraphy of the Ordovician of southwestern of Europe. On the
basis of an uncountable number of observations and descriptions, he
was able to refine and structure in a logical way the classification
schemes previously proposed by A. Eisenack and P. Taugourdeau. He
then played an active role in bringing all scientists together for the de-
velopment of global biostratigraphical schemes of the Ordovician, Silu-
rian and Devonian, in promoting the use of chitinozoans for the
definition of international boundary stratotypes, or at least for the rec-
ognition of the chronostratigraphic boundaries elsewhere. His work
was instrumental for the refinement of the ‘Northern Gondwanan’
palaeobiogeography. As the biological affinities of chitinozoan remained
unknown, he also collaborated with J. Nõlvak to propose the now wide-
ly accepted biological interpretation that Chitinozoa are most probably
egg cases of an unknown planktonic organism. During the last decade
he was actively involved in the new focus of chitinozoan research di-
rected toward chitinozoan biodiversity and its relationship with envi-
ronmental changes. He was also the first to use biogeochemical
analyses and the C isotope signal of the chitinozoan wall to better un-
derstand their biological affinity and detect biogeochemical changes in
Palaeozoic oceans. In all these aspects of research, Florentin Paris was
the driving force.
However, notwithstanding his significant contribution to palaeon-
tology, Florentin Paris always remained at heart a field geologist
 T. Servais et al. / Review of Palaeobotany and Palynology 198 (2013) 2–13
Fig. 1. Florentin Paris in the field at Touijinet, Mauritania.
(Fig. 1), contributing, for example, his share to the geological mapping
of Brittany (e.g., Bonjour et al., 1988; Paris and Le Hérissé, 1992). As
member of the very productive “Lower Palaeozoic team” at the Univer-
sity of Rennes, he published several significant papers on the geology
and palaeogeography of the Armorican Peninsula and the ‘North Gond-
wana’ domain making him the expert in the Lower Palaeozoic geology
and stratigraphy of the ‘Armorican terrane assemblage’ and North
Africa.
The leading role of Florentin Paris was probably favoured by his per-
sonal and human qualities, his open mind, his willingness to help other
scientists, to invite them to Rennes, and his ability to bring people to-
gether. Many of his colleagues remember his hospitality, not only in
his research department, but also at his home, where he and his wife
Jacqueline welcomed many colleagues and visitors.
Florentin Paris was in close contact with most chitinozoan workers.
Many of the recent developments involved research students and it is
not surprising that Florentin Paris was directly implicated in the training
of many of them. Among the students that now constitute the youngest
generation of chitinozoan workers are some of the young scientists that
have recently been recruited by several universities or research institu-
tions: A. Butcher (Portsmouth, UK), S. de la Puente (Mendoza,
Argentina), Tang Peng (Nanjing, China), and T. Vandenbroucke (Lille,
France), to name just a few. These colleagues and the entire chitinozoan
community will continue to benefit from the standards that Florentin
Paris set for the future generations of palynologists.
Acknowledgments
We are particularly grateful to Jaak Nõlvak, Thijs Vandenbroucke
and Jacques Verniers for reviewing the paper and for adding useful
‘background information’. This study is a contribution to the IGCP 591
“The Early and Middle Palaeozoic Revolution” project. It was supported
by the Natural Science and Engineering Research Council of Canada
(Grant RGPIN/4226-2006 to Aicha Achab). It is a Natural Resources
Canada — Earth Sciences Sector Contribution 20120108. We wish to
thank Kheira Boumendjel, Yngve Grahn, Jean-François Ghienne and
Jaak Nõlvak for valuable information and John Riva for reviewing the
English of an earlier version of the manuscript. On behalf of all our col-
leagues, we dedicate this review paper to our colleague and eminent
scientist, Florentin Paris.
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