Mesozoic–Cenozoic organic-walled dinoflagellate cyst (dinocyst)
biostratigraphy is a crucial tool for relative and numerical age control in
complex ancient sedimentary systems. However, stratigraphic ranges of
dinocysts are found to be strongly diachronous geographically. A global
compilation of state-of-the-art calibrated regional stratigraphic ranges
could assist in quantifying regional differences and evaluating underlying
causes. For this reason, DINOSTRAT is here introduced – an open-source,
iterative, community-fed database intended to house all regional
chronostratigraphic calibrations of dinocyst events (
Over 50 years of research efforts has established a framework to use organic-walled dinoflagellate cysts (dinocysts) as biostratigraphic and chronostratigraphic tools. Dinocyst biostratigraphy is particularly applied to sediments which are difficult to date otherwise, such as in restricted nearshore marine settings (e.g., Poulsen, 1994; Brinkhuis et al., 1998; Iakovleva et al., 2001; Śliwińska et al., 2012; Clyde et al., 2014) and polar regions (e.g., Sluijs et al., 2006; Bijl et al., 2013a; Houben et al., 2013; Radmacher et al., 2015; Śliwińska et al., 2020; Nøhr-Hansen et al., 2020). As with all biostratigraphy, the reliability of dinocyst biostratigraphy heavily depends on the accuracy, precision and regional consistency of the numerical ages of the first and last stratigraphic occurrences (FOs and LOs, hereafter jointly referred to as “events”) of easily recognized taxa. Through the past decades, numerical ages of dinocyst events have become increasingly better chronostratigraphically constrained, using independent age control from magnetostratigraphy (e.g., Brinkhuis et al., 1992; Powell et al., 1996), other biostratigraphic tools (e.g., Davey, 1979; Leereveld, 1997a, b; Oosting et al., 2006; Awad and Oboh-Ikuenobe, 2019) and astrochronology (Versteegh, 1997). However, efforts to compile a global chronostratigraphic calibration of dinocyst events have revealed strong diachroneity for many species between broad latitudinal bands and endemism of many species within latitudinal bands (e.g., Williams et al., 2004). Because this impacts the development of quasi-global dinocyst zonation schemes, as have been proposed for other microfossil groups (e.g., Martini, 1971; Gradstein et al., 2020), the question is, how should the research field of dinocyst biostratigraphy progress?
Two questions arise from the notion of the geographic diachroneity of dinocyst
events.
What kind of error or uncertainty should be applied to the numerical ages of
events? Now that diachroneity has been demonstrated, the next step is to
quantify the uncertainty in numerical ages of dinocyst events for each
species and to assess regional consistency. This is particularly important
when calibrated species ranges are geographically extrapolated over large
distances. And a related question is, what is the impact of regional
variability in the numerical ages of events on the regional consistency of the
stratigraphic order of events? What are the underlying causes for the observed diachroneity? Broadly, three
reasons could apply: (1) inaccurate or inadequate tie-in of dinocyst events
to the chronostratigraphic timescale can lead to apparent (but perhaps
false) diachroneity of species events between sites. (2) Complexities in
taxonomic concepts could obscure comparison of species ranges between sites.
This aspect relates to the ease by which subtle morphological differences
between species can be recognized (e.g., Hoyle et al., 2019). It also
relates to the question of whether the last occurrence of a fossil dinocyst
taxon reflects extinction of its producer, adjustment of cyst morphology by
its producer (e.g., Rochon et al., 2009), or a change in its life cycle
strategy (e.g., towards less preservable pellicle cysts; Bravo and Figueroa,
2014). (3) Finally, paleoenvironmental/paleoceanographic conditions can
impact species occurrence: ocean connectivity (Van Simaeys et al., 2005;
Bijl et al., 2013b; Van Helmond et al., 2016), leads and lags in the biotic
response to climate change (e.g., Sluijs et al., 2007), or the temperature
affinity of dinocyst taxa (Van Simaeys et al., 2005; Van Helmond et al.,
2016). For instance, in geologic time intervals of global climate cooling,
warm-loving plankton species have diachronous last occurrences which are
progressively later at lower latitudes. A good example is the modern
occurrence in the western Pacific warm pool of
A process that takes us towards answering these questions and improving the accuracy of dinocyst biostratigraphy requires a data compilation approach that incorporates data from as many sites as possible, with detailed metadata on the paleogeographic evolution of sites and the means of chronostratigraphic calibration. It further requires that such data compilations are constantly updated with new insights: updates to the geologic time scale and bio-magnetostratigraphic zonation schemes, altered taxonomic concepts, age models of sections, and stratigraphic sections. A complication on a logistical front is that dinocyst ranges are typically published in the closed-access peer-reviewed literature, which is not easily accessible to all, is inconsistent in its approach and is easily updated with new insights.
This paper introduces DINOSTRAT, an open-source, online platform intended to house, disseminate and iteratively update all published chronostratigraphic calibrations of dinocyst ranges: the way in which they are tied to the chronostratigraphic timescale and the (paleo)geographic position of the site from which they were calibrated. DINOSTRAT version 1.0 currently contains over 8500 entries of the first and last occurrences of over 1900 dinocyst taxa tied to the international timescale. These entries originate from 199 peer-reviewed papers presenting data from 188 sites. Including as many reports/sites as possible, with verifiable independent age control and their latitudinal evolution through time, allows for proper evaluation of error and uncertainty. DINOSTRAT will allow assessment and quantification of regional variability/consistency in event ages and provides the basic information to evaluate the paleoceanographic signal that diachroneity may hold. Open accessibility of the basic dinocyst stratigraphic data will further allow a proper evaluation and update of evolutionary patterns in dinocyst families (MacRae et al., 1996) with full disclosure of the underlying data. The approach to the selection of appropriate data and entry and calculations of ages and paleolatitudes is explained in Sect. 2. Section 3 presents examples of calibrated dinocyst events: the stratigraphic and paleolatitudinal distribution of selected modern dinocysts and that of extant and extinct dinocyst families, with selected taxa highlighted. Section 4 discusses the implications of the DINOSTRAT approach and future directions. This paper represents the start of a community-fed data assembly approach to iteratively improve regional constraints on dinocyst biostratigraphy.
DINOSTRAT version 1.0 represents a compilation of dinocyst events from peer-reviewed literature, with a publication date predating 1 January 2021 (see Table 1). The taxonomic nomenclature, supra-generic classification and synonymy cited in Williams et al. (2022) are followed. One inherent assumption in the initial setup of DINOSTRAT is that the authors of the reviewed literature have applied a consistent taxonomic framework. DINOSTRAT reports events of dinocyst species as they were presented in the papers but applying the synonymy index of Williams et al. (2022). Most dinocyst species are easily recognized, have a stable morphology (both regionally and through time) and clearly defined species concepts. However, some species (and subspecies) diagnoses are more subtle or represent endmembers in a continuum (e.g., Hoyle et al., 2019), in part imposed by environmental conditions (e.g., Ellegaard, 2000). Some authors tend to lump species in complexes, while others split them into subspecies. The international recognition of these lumps and splits may have evolved through time and may have restricted regional significance only. Therefore, subtle differences in species concept interpretation may exist between authors and regions, which the current approach was unable to account for, and identifying them is considered a next step for when individual studies or sites are revisited.
For the subfamily Wetzelielloideae, DINOSTRAT deviates from the taxonomic index of Williams et al. (2022). The fundamental redefinition of species concepts in the taxonomic revisions for the Wetzelielloideae (Williams et al., 2015) eliminates many stratigraphically useful Eocene dinocyst taxa (Bijl et al., 2016). Therefore, for this subfamily, the calibration of dinocyst species is presented in the taxonomic classification of Wetzelielloideae prior to Williams et al. (2015).
Papers used in this review. Reference, geography, age base and age top (in Ma), tier (see Fig. 1), and means of calibration to the geologic time scale (GTS). The abbreviations of the microplankton zones indicated in the column “Calibrated to” (NJ, NC, NP, NN, CC, CP, CN, UC, N, P, E) are those commonly used in the literature and relate to the zones presented in the GTS2012 (Gradstein et al., 2012).
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A decision tree has been used to determine which papers to include in
DINOSTRAT (Fig. 1). This tree first discards studies in which dinocysts were
the only stratigraphic tool to date the sequence. Although these papers do
provide valuable information on the stratigraphic order of events, discarding
them from this review eliminates the risk of circular reasoning and
inherited chronostratigraphic ties. Only those dinocyst events are included
that could be calibrated against a stratigraphic tool that can be traced
back to the bio-, magneto- or chronozones in the geologic time scale 2012
(GTS2012; Gradstein et al., 2012). This decision tree can be used to define
five tiers of reliability in these papers (Fig. 1):
Tier 1 studies present dinocyst events along with magnetostratigraphic
constraints obtained from the same sedimentary section. The interpretation
of magnetochrons from the paleomagnetic signal was performed without the use of
dinocyst biostratigraphy. Since magnetic reversals are globally synchronous,
evaluating the synchroneity of dinocyst events with the use of
paleomagnetostratigraphy is most robust. Tier 2 studies present dinocyst events calibrated along with compromised or
problematic magnetostratigraphic constraints on the same sedimentary
section, for instance when the inclination signal suffers from a strong
overprint or when the magnetochron assignment is not clear. Studies in
which dinocyst events served as biostratigraphic tools for magnetochron
assignment are included in this tier as well. Tier 3 studies report dinocyst events together with biostratigraphic zones
(from nannoplankton, foraminifer or ammonite zones) identified in the same
sequence. These studies provide clear reports on the identification of these
zones in the sequence. Tier 4 studies report dinocyst events with biostratigraphy, of which either
the derivation is unclear or the tie to the geologic time scale (GTS; e.g., for outdated
ammonite zonations) or biostratigraphic data does not come from the same
sequence but, for example, has been interpreted from nearby outcrops. Tier 5 studies report dinocyst events with independent chronostratigraphy,
of which the derivation is unverifiable or represents a regional synthesis.
Decision tree for including studies in this review and categorization criteria for the five tiers.
The numerical age of each dinocyst event is not explicitly entered into
DINOSTRAT. Rather, its position within the zone it was calibrated to is
entered. Ages are subsequently calculated via linear interpolation between
these tie points, as follows:
Each event entry in DINOSTRAT (Dinoevents_Jan2021.csv in
Bijl, 2021) includes the (paleo)latitude of that event. This is
interpolated using the age of the event and its location, which has a
paleolatitude evolution through time (Paleolatitude.csv in Bijl, 2021; with
use of
DINOSTRAT currently contains dinocyst events from 199 publications and 188 sites. The wider North Atlantic–European area is strongly overrepresented (Fig. 2). Few sites are from the Pacific Ocean, the southern Atlantic and Indian Ocean, and the equatorial region. This probably reflects a genuine bias in the available information because of focus of the community towards economically interesting regions (e.g., for hydrocarbon industry). It may also in part reflect a bias towards research from developed nations and poor accessibility of publications from non-western societies.
Present-day geographic distribution of sedimentary sequences used
in DINOSTRAT (colors of the dots correspond to the to the tier status to
which the site has been allocated) and surface sediment stations (in grey
dots; Marret et al., 2020; Mertens et al., 2014).
The paleolatitudinal position of the sites through time confirms the strong overrepresentation of Northern Hemisphere mid-latitude sections (Fig. 3) and underrepresentation of the tropical regions, Pacific Ocean and southern mid-latitudes. The Paleogene has the largest latitudinal spread of records. The Mesozoic in particular has few entries from the Southern Hemisphere or equatorial regions. The Mesozoic records are predominantly calibrated to ammonite stratigraphy (tiers 3 and 4) and on some occasions to magnetostratigraphy (tiers 1 and 2; Fig. 3). Ammonite zones presented in the papers often had to be converted to those in GTS2012, which is not always straightforward, as the zone definitions have changed through time (Ogg and Hinnov, 2012a, b). The ammonite zonations are prone to regional diachroneity, which was demonstrated particularly for the Late Jurassic (Ogg and Hinnov, 2012b). This may create a level of circular reasoning when dinocyst events are calibrated against these zones because diachronous dinocyst events in DINOSTRAT may be the result of diachronous ammonite zones rather than actually being diachronous dinocyst events.
Data in DINOSTRAT.
DINOSTRAT version 1.0 includes over 8500 entries of calibrated dinocyst events (excluding the modern dinocyst database). On a species level, originations in DINOSTRAT peak in the Middle Jurassic (Bajocian–Callovian), the Early Cretaceous (late Valanginian–Barremian) and the Eocene (Ypresian; Fig. 3b). Extinctions peak in the Early Cretaceous (Berriasian–Barremian), Upper Cretaceous (Maastrichtian), Oligocene (Rupelian) and Miocene (Serravallian; Fig. 3b). This pattern is generally followed on a generic level, which likely has a stronger relation to the biologic diversity than dinocyst species diversity (Fensome et al., 1993).
The interpolated paleolatitudes for dinocyst events in DINOSTRAT allow detailed evaluation of the latitudinal synchroneity of dinocyst events. This paper presents a selection of the data in DINOSTRAT, focusing on the stratigraphic and geographic range of modern dinocyst species, of dinocyst families/subfamilies and of a selection of quasi-synchronous dinocyst events. Users can filter DINOSTRAT by locality (to present the stratigraphic order of events per site) and/or by taxon (to see the geographic variability in the range of any taxon), to serve their purposes.
Modern dinocysts from surface sediment samples (Marret et al., 2020,
Age and paleolatitude of first (in blue) and last (in red) occurrences of selected modern dinocyst species. Last occurrences come from both the surface sediment database (Marret et al., 2020, with Mertens et al., 2014) and entries in DINOSTRAT.
Range charts of the sites in DINOSTRAT are provided in the Supplement (see
“Sites” folder in Supplement File 2). The age-over-paleolatitude entry in
DINOSTRAT allows evaluation of the latitudinal difference in event ages for
each individual species in DINOSTRAT (
Order Gonyaulacales Family Areoligeraceae (Fig. 5)
Ages and paleolatitudes of first (solid line and triangles) and last (dashed line and circles) occurrences of dinocyst species of the family Areoligeraceae. Solid and dashed lines connect first and last occurrences, respectively, for each species, between sites. Colored lines represent quasi-synchronous species events.
Family Ceratiaceae (Fig. 6)
Same as Fig. 5 but for the family Ceratiaceae.
Family Cladopyxiaceae (Fig. 7)
Same as Fig. 5 but for the family Cladopyxiaceae.
Family Goniodomaceae (Fig. 8)
Same as Fig. 5 but for the family Goniodomaceae.
Family Gonyaulacaceae Subfamily Cribroperidinioideae (Fig. 9)
Same as Fig. 5 but for the family Gonyaulacaceae, subfamily Cribroperidinioideae.
Subfamily Gonyaulacoideae (Fig. 10)
Same as Fig. 5 but for the family Gonyaulacaceae, subfamily Gonyaulacoideae.
Subfamily Leptodinioideae (Fig. 11)
Same as Fig. 5 but for the family Gonyaulacaceae, subfamily Leptodinioideae.
Family Gonyaulacaceae, other (Fig. 12)
Same as Fig. 5 but for other genera in the family Gonyaulacaceae.
Family Mancodiniaceae (Fig. 13)
Same as Fig. 5 but for the family Mancodiniaceae.
Family Pareodiniaceae (Fig. 14)
Same as Fig. 5 but for the family Pareodiniaceae.
Family Scriniocassiaceae (Fig. 15)
Same as Fig. 5 but for the family Scriniocassiaceae.
Family Shublikodiniaceae (Fig. 16)
Same as Fig. 5 but for the family Shublikodiniaceae.
Family uncertain (Fig. 17)
Same as Fig. 5 but for the order Gonyaulacales, family uncertain.
Order uncertain Family Comparodiniaceae (Fig. 18)
Same as Fig. 5 but for the family Comparodiniaceae.
Family Stephanelytraceae (Fig. 19)
Same as Fig. 5 but for the family Stephanelytraceae.
Order Peridiniales Family Heterocapsaceae (Fig. 20)
Same as Fig. 5 but for the family Heterocapsaceae.
Family Peridiniaceae Subfamily Deflandroideae (Fig. 21)
Same as Fig. 5 but for the family Peridiniaceae, subfamily Deflandreoideae.
Subfamily Palaeoperidinioideae (Fig. 22)
Same as Fig. 5 but for the family Peridiniaceae, subfamily Palaeoperidinioideae.
Subfamily Wetzelielloideae (Fig. 23)
Same as Fig. 5 but for the family Peridiniaceae, subfamily Wetzelielloideae.
Family Peridiniaceae, other (Fig. 24)
Same as Fig. 5 but for other subfamilies in the Family Peridiniaceae.
Family Protoperidiniaceae (Fig. 25)
Same as Fig. 5 but for the family Protoperidiniaceae.
Order Nannoceratopsiales Family Nannoceratopsiaceae (Fig. 26)
Same as Fig. 5 but for the family Nannoceratopsiaceae.
Order Ptychodiscales Family Ptychodiscaceae (Fig. 27)
Same as Fig. 5 but for the family Ptychodiscaceae, subfamily Dinogymnioideae.
Order Suessiales Family Suessiaceae (Fig. 28)
Same as Fig. 5 but for the family Suessiaceae.
A suite of dinocyst events throughout the entire stratigraphic record have quasi-synchronous ages across all latitudes (Figs. 5–28). The uneven geographic spread of data, with voids in the equatorial region and the Pacific Ocean, makes global synchroneity of these events highly uncertain. Still, the synchronous events confirm the potential and value of dinocyst biostratigraphy to date complex sedimentary systems. They also imply that ocean connectivity did allow dinocyst species to migrate globally, as far as their environmental tolerances permitted.
Yet, the majority of dinocyst species have very diachronous ranges in DINOSTRAT, as well as latitudinally restricted geographic spreads, which confirms previous interpretations (Williams et al., 2004). By using DINOSTRAT the underlying causes of this diachroneity can now be further explored. The shortness of some of the records used in this review may lead to “false” events, i.e., those that represent re-appearance or temporal disappearance rather than “true” first or last occurrences (FOs and LOs, respectively). The obviously false FOs and LOs have been removed from DINOSTRAT by omitting events that occur at the base or the top of the sections. Particularly rare species or those occurring at the end of their preferred environmental niche come and go in stratigraphic sections, and these lead to false events in DINOSTRAT. Although such false FOs and LOs may obscure a uniform age of events over latitudes, they may still have important regional stratigraphic significance, which is why their entries are retained in DINOSTRAT. As a result, the age and region of the oldest FOs and youngest LOs have the most significance for the reconstruction of evolutionary patterns. Although caving of material typically falsely increases the age of the oldest FOs, this is unlikely to have a large influence on the entries in DINOSTRAT, as most studies come from core or outcrop material, and not from ditch cuttings, for which caving is much more likely. Reworking could falsely extend the age of the youngest LOs of species. Although species that were reported as reworked in the papers have been omitted from DINOSTRAT, some reworked dinocysts could have been falsely identified as in situ in the original papers. It cannot be excluded that this causes some level of diachroneity in LOs, although this is unlikely a large factor.
The complexity of taxonomic concepts in some dinocyst genera (species
definitions or morphological continua) hinders proper evaluation of
latitudinal synchroneity of events. The reviewed literature covers 50 years,
during which taxonomic concepts of dinocysts species have iteratively
evolved. The extensive synonymy database of Williams et al. (2022) does
deliver crucial organization of the taxonomic framework. Still, some of the
subtle morphological differences in species are limited to the expert eye of
individual researchers, and these may not have been recognized by others
(which occasionally has led to the presentation of taxa on a generic level,
instead of further specification to species level). Making the taxonomic
framework consistent for all studies now included in DINOSTRAT would be a
cardinal effort and will be part of the iterative setup of DINOSTRAT. For
example, reviews of dinocyst taxonomic frameworks on a per-family basis,
such as has been initiated for the
Events may also appear diachronous in DINOSTRAT because of inadequate or
inaccurate ties to the chronostratigraphic timescale. In such cases, minor
diachroniety (
Also, ecological reasons could cause geographically diachronous events. When
local environmental or depositional conditions change, assemblages adjust,
which leads to local and temporal (dis)appearances of species that may be
falsely interpreted as extinction or origination events. If so, dinocyst
taxa associated with the most dynamic environmental niches on the continental
shelf are expected to have the most diachronous events. Indeed, there are
particularly diachronous events in Goniodomaceae and Protoperidinioideae –
both families are associated with nearshore depositional settings
(Zonneveld et al., 2013; Sluijs et al., 2005; Frieling and Sluijs, 2018)
that are the most environmentally dynamic. Settings in which these species occur
offshore, such as in upwelling regions (Sangiorgi et al., 2018) or
hyperstratified waters (Reichart et al., 2004; Cramwinckel et al., 2019),
are environmentally equally dynamic. In contrast, families typically
associated with offshore conditions, such as the Wetzeliellioideae (Frieling
and Sluijs, 2018), reveal much more synchronous events. For regional
stratigraphy, the diachroneity is of less concern because these events can
still be used for regional stratigraphic correlation (e.g., as in Vieira and
Jolley, 2020). It does mean that for such species, dinocyst biostratigraphy
applies regionally, and caution should be taken to extrapolate event ages
far outside of these regions. There are also species that clearly show
regional inconsistency in origination or extinction ages because of climate
change – e.g.,
Diachroneity is usually larger between latitudinal bands than within
latitudinal bands. The sparsity of records from the SH high latitudes
complicates robust assessment of interhemispheric differences in dinocyst
event ages. In the Mesozoic, the diachroneity is likely related to the
inadequate calibration of events to the international timescale. DINOSTRAT
is short of Mesozoic records that are tied to stratigraphic tools other than
ammonites. For the Cenozoic, the diachroneity between hemispheres cannot be
explained by inadequate calibration since many events are calibrated against
magnetostratigraphy. For those, environmental reasons must be at play. While
in the early Paleogene many dinocyst events are quasi-synchronous (events
within the Wetzeliellioideae, of
Many dinocyst species and higher generic ranks have their oldest first occurrence and youngest last occurrence in NH mid-latitudes (for example Areoligeraceae, Cladopyxiaceae, Comparodiniaceae, Goniodomaceae, Nannoceratopsiaceae, Palaeoperidinioideae, Wetzeliellioideae; Figs. 5, 7, 18, 8, 26, 22, 23). This may be because of a much higher density of records at those latitudes. However, the vast continental shelf area in Europe throughout the Mesozoic and much of the Cenozoic did likely serve as the perfect habitat for taxa to find a new niche and to linger on. A higher record density in SH and equatorial regions should shed light on this idea.
DINOSTRAT presents for the first time a quantitative overview of the
stratigraphic and paleolatitudinal distribution of fossil and modern
dinocyst taxa. Through that, it refines with coherent, independent,
open-access data the evolutionary patterns presented previously (e.g.,
Fensome et al., 1993; MacRae et al., 1996) and adds their latitudinal
distribution through time. Following up on 60 million years of
experimentation in cyst formation among a wide group of dinoflagellates
(Figs. 13, 15, 16, 18–20, 26, 28), gonyaulacoid dinocysts developed their
most fundamental taxonomic features in a rapid diversity phase in the
Bajocian (
Once downloaded, DINOSTRAT can be filtered by location, allowing users to compare newly generated dinocyst chronologies to calibrated regional dinocyst events nearby. DINOSTRAT can also be filtered by species, genus or higher taxonomic rank for further evaluation of the latitudinal spread of any species of interest. The data in DINOSTRAT are readily visualized in Supplement File 2, and these plots can be adjusted and reproduced using the R markdown file plot creator.Rmd in Bijl (2021). The community is invited to contact the author either via email or through GitHub, with suggestions, error reports, and/or additional papers or data to be entered so that the data content of DINOSTRAT is iteratively improved.
DINOSTRAT will be regularly updated. Annual minor updates include the addition of sites, adjustments in the current entries (e.g., through the feedback process) or minor revisions in taxonomy/stratigraphy. Major updates will occur in a 3-year cycle and will be the result of updates to the geologic time scale or profound revisions in dinocyst taxonomic concepts. Major updates will be accompanied by a short communication in this journal; minor updates will be communicated through the GitHub repository. Updates of the geologic time scale (e.g., to GTS2020; Gradstein et al., 2020) will be implemented once the metadata of that geologic time scale have become available. All versions of DINOSTRAT will remain archived on GitHub.
The database is available under a CC BY 4.0 license on GitHub (Bijl, 2021;
This paper presents the database DINOSTRAT version 1.0 (Bijl, 2021), a
database containing
Many dinocyst taxa show quasi-synchronous events latitudinally, which can be widely used to stratigraphically date complex sedimentary sequences. Latitudinal diachroneity in events can be the result of inadequate calibration to the chronostratigraphic timescale, false interpretations of true events, complicated species concepts or paleoceanographic reasons. In any case, this dictates caution when extrapolating ages of dinocyst events to far distances and demonstrates the need for regionally calibrated dinocyst zonations, which DINOSTRAT here provides. It further provides a solid foundation to review spatio-temporal patterns in dinocyst evolution, dispersal and extinction. DINOSTRAT is freely available under a CC BY 4.0 license. It allows the user to filter by region or by species, genus or higher taxonomic rank.
Supplement File 1 is a table of conversions of published zones to those in GTS2012. Supplement File 2 is a zip file containing ages and latitudes of events in individual
dinocyst species (1914 plots), grouped by genus (459 plots) and by family (27
plots), including modern cyst species (92 plots) and the range charts for all
sites (188 plots). The supplement related to this article is available online at:
The contact author has declared that there are no competing interests.
Publisher's note: Copernicus Publications remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
I thank Henk Brinkhuis, Bas vd Schootbrugge, Francesca Sangiorgi and Appy Sluijs for useful discussions. The “Advanced course in organic-walled dinoflagellate cyst taxonomy, stratigraphy and paleoecology” has been a great “playground” to discuss progress in the field, and for that I have Martin Head, Martin Pearce, Jörg Pross, Jim Riding and Poul Schiøler to thank. I acknowledge the then research assistants who helped in building predecessors of DINOSTRAT: Tjerk Veenstra, Keechy Akkerman and Caroline van der Weijst. Thanks to Martin Schobben and Ilja Kocken for help with the data analysis and visualization in R and to Douwe van Hinsbergen for help reconstructing the paleolatitudes of the sites. James Ogg is thanked for providing the data from GTS2012. The constructive and detailed comments from Henrik Nøhr-Hansen and Ian Harding greatly improved the final paper.
The LPP Foundation has financially supported the development of DINOSTRAT.
This paper was edited by Thomas Blunier and reviewed by Henrik Nøhr-Hansen and Ian Harding.