Tools for Non-Pollen Palynomorphs (NPPs) analysis: A list of Quaternary NPP types and reference literature in English language (1972–2011)

Abstract

A systematic study of Non-Pollen Palynomorphs (NPPs) from Quaternary sediments was begun in 1968 in the Hugo de Vries-Laboratory (Institute for Biodiversity and Ecosystem Dynamics — IBED) at the University of Amsterdam (The Netherlands). Since the publication of the famous paper by Bas van Geel (1972), more than 1300 NPPs have been described and many of them have been identified as the remains of diaspores, cyanobacteria, algae, rhizopods, fungi, invertebrates, and chormophytes. In order to assess the current state of the art in the description of NPP types, in this paper I reviewed the “classical NPP literature” and the most recent literature as of November 2011, and I present a list of the described NPP types, as long as published in the English language. Full list of the NPP types and the related literature are available online as supplementary materials (.xls file) in this article's online version. The NPP type list is a new tool for NPP analysts and aims to widen the NPP research community, increase the number of identified NPP, and reduce the inconsistencies in naming the same microfossils.

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Research paper
Tools for Non-Pollen Palynomorphs (NPPs) analysis: A list of Quaternary NPP types
and reference literature in English language (1972–2011)
Antonella Miola ⁎
Department of Biology, University of Padova, Via Ugo Bassi 58 B, 35131 Padova, Italy
abstractarticle info
Article history:
Received 19 December 2011
Received in revised form 25 June 2012
Accepted 27 June 2012
Keywords:
Non-Pollen Palynomorphs
Quaternary
fungi
algae
cyanobacteria
rhizopods
nomenclature
A systematic study of Non-Pollen Palynomorphs (NPPs) from Quaternary sediments was begun in 1968 in the
Hugo de Vries-Laboratory (Institute for Biodiversity and Ecosystem Dynamics —IBED) at the University of
Amsterdam (The Netherlands). Since the publication of the famous paper by Bas van Geel (1972), more
than 1300 NPPs have been described and many of them have been identified as the remains of diaspores,
cyanobacteria, algae, rhizopods, fungi, invertebrates, and chormophytes. In order to assess the current state
of the art in the description of NPP types, in this paper I reviewed the “classical NPP literature”and the
most recent literature as of November 2011, and I present a list of the described NPP types, as long as pub-
lished in the English language. Full list of the NPP types and the related literature are available online as sup-
plementary materials (.xls file) in this article's online version. The NPP type list is a new tool for NPP analysts
and aims to widen the NPP research community, increase the number of identified NPP, and reduce the in-
consistencies in naming the same microfossils.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
The Non-Pollen Palynomorphs (NPPs) are microfossils normally in-
cluded in Quaternary sediments that are processed for palynological
analysis. They are sub-fossil remains of a great variety of organisms
that can be sensitive to various ecological parameters or to human pres-
ence. Before the 1970s, it was a common practice for palynologists to
use only fossil pollen and fern spores, whereas many palynomorphs of
unknown origins were not considered. It is true that palynomorphs
other than pollen, mainly fungal and animal remains, had been consid-
ered in some earlier publications (e.g., Rudolph, 1917; Hoogenraad,
1935; Nilsson, 1952; Huikari, 1956; Frey, 1960; Graham, 1962;
Aartolahti, 1965). But it was not until 1968 that Bas van Geel, a master
student, started a systematic study of NPPs in palynological slides at
the Institute for Biodiversity and Ecosystem Dynamics (IBED) in the
University of Amsterdam (The Netherlands). His study aimed to im-
prove the quality of Quaternary palaeoenvironment reconstructions.
Bas van Geel introduced the practice to describe the NPP types and to
give them a name, even if they were of unknown origin (van Geel,
1972, 1978). This practice became the standard system for palynologists
that studied also Non-Pollen Palynomorphs in the pollen slides. Since
then, a variety of NPP types have been described and illustrated. Their
palaeo-environmental indicator values were mentioned and used in
later studies concerning Quaternary sediments and palaeoenvironment
reconstruction. Some NPPs have been identified as the remains of
diasporas, cyanobacteria, green algae, rhizopods, fungi, invertebrates,
and chormophytes. They have been used to provide additional and in-
dependent palaeoenvironmental information (see van Geel (2001) for
a summary). In one of the best known textbooks on pollen analysis,
Faegri and Iversen (1989) still discouraged the tendency to include mi-
crofossils of unknown origin under the term “palynology.”Neverthe-
less, they included a chapter titled “Some other microfossils”and cited
the articlesof Bas van Geel and co-authors. As compared to pollen anal-
ysis alone, the attention paid by Quaternary researchers to NPPs as bio-
logical proxies is steadily spreading, and their value has been
demonstrated in more than 200 scientific papers. Consequently, the
list of new NPPs is increasing, but many of them remain unidentified.
Obviously, the indicator-species approach (Birks and Birks, 1980)is
more difficult to apply to NPPs of unknown origin. The challenge for
NPP analysts is to identify at least the most common types of NPPs of
unknown origin, which seem to be indicative of environmental condi-
tions or human activities.
Since the certainty and speed of determinationrise with the number
of previously identified NPPs, the NPP researchers began to organize
systematic meetings, International workshops in Greifswald —March
2005; Innsbruck —August, 2006; Padova —June, 2008; Besançon —
June, 2010, and special symposia in congresses, International Palynolog-
ical Congresses Granada —July, 2004; Bonn —August, 2008; Bern —
August, 2011, in which information on identified and not-yet-identified
NPPs is shared. Two special issues of Review of Palaeobotany and Paly-
nology (141/1–2, 2006, edited by B. van Geel) and Vegetation History
and Archaeobotany (19/5–6, 2010, edited by J.N. Haas) published the
works presented on these occasions. A possible consequence of these
Review of Palaeobotany and Palynology 186 (2012) 142–161
⁎Tel.: +39 049 8276267; fax: + 39 049 8276260.
E-mail address: antonella.miola@unipd.it.
0034-6667/$ –see front matter © 2012 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2012.06.010
Contents lists available at SciVerse ScienceDirect
Review of Palaeobotany and Palynology
journal homepage: www.elsevier.com/locate/revpalbo

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events was that the number of scientific papers including NPP analysis
has increased in the last ten years (Fig. 1).
The diffusion of NPP studies and the necessity of the consistent com-
parison of the results require sharing common rules of nomenclature
and an updated list of NPPs. Common rules of nomenclature are quite
important for certainty of identification and for assessing the NPPs'
value as biological proxies. An updated list of NPPs should limit the
use of different terms/codes/taxon names to indicate the same object.
In contrast to pollen analysis, NPP analysis is not supported by a wide
availability of photographic reference materials, identification keys, or
regional and global databases (Hooghiemstra and van Geel, 1998).
The specialized literatures for algologists, mycologists or zoologists are
not always appropriate for palynologists, who can normally observe
only parts of the organisms, e.g., spores, cysts, eggs, and various kinds
of fragments. Recently, Gelorini et al. (2011) published an illustrated
guide to the identification of NPPs preserved in lake sediments from
Africa; Payne et al. (2012) provided a basic introduction to testate
amoeba taxonomy. Also, Komárek and Jankovská (2001) published a
useful guide for some genera of green algae,and Haas (1996) published
a useful description of a variety of species of Neorhabdocoela on the
basis of their oocytes. Normally, NPP analysts consult the papers of
van Geel and colleagues, the “classical NPP literature,”where they can
find descriptions and illustrations of NPP types that are both identified
and unidentified. A photocopy collection of illustrations and descrip-
tions of NPP types taken from the publications of van Geel and col-
leagues and from internal reports of the Hugo de Vries Laboratory at
the University of Amsterdam, from 1972 to 1997 was edited by van
Hoeve and Hendrikse in 1998 at Utrecht (A study of non-pollen objects
in pollen slides. The types as described by Dr. Bas van Geel and colleagues).
This work has been the “reference collection”for many of us. After over
15 years and more than 200 publications, an updated list of the pub-
lished NPP types should be a useful tool for both experienced palynolo-
gists and beginners.
In order to assess the current state of the art in the description of
NPP types, I first reviewed the “classical NPP literature.”Then, I
searched the most recent literature by means of Scopus (SciVerse
Scopus) and Google Scholar as of November 2011. Finally, on the
basis of the literature to which I could gain access, I drew up a list
of the NPP types that have been described and illustrated. A list of
the relative NPP literature is also given in this paper. Both lists are
available in the full version as supplementary materials in this
article's online version.
2. The NPP literature list
The list presented in Table 1 includes selected articles from scientific
journals and chapters in books, mainly in English language, concerning
Quaternary studies in whichpollen and NPP analyses have been applied
as the principal method of study and the results of NPP analysis have
been discussed. They have also been selected because they describe
and illustrate new or previously described NPP types or present discus-
sions about their indicator values for palaeoenvironment reconstruc-
tion. The list includes the papers from the “classical NPP literature”
and more recent papers (1990–2011) that have been searched by
means of Scopus (SciVerse Scopus) and Google Scholar for the term
“Non-Pollen Palynomorphs”anywhere in the journal papers and book
chapters. The automatic search also matches items with the terms
“non-pollen”or “palynomorphs.”These specialized search engines
offer a way to facilitate the searching of bibliographic materials for
academic purposes. Obviously they are limited to some selected sources
(please see the web sites of the search engines to view the database
sources), and they cannot find papers that are not included. This search
is therefore somewhat biased. The so called “grey literature,”for
example, cannot be retrieved in this way. Another bias concerns the
language: papers which arenot published inEnglish were not matched
if “Non-Pollen Palynomorphs,”“non-pollen,”and “palynomorphs”were
absent from the English abstract. For this reason, I added some further
studies known to me from other sources, but obviously, this list cannot
be complete.
The complete bibliographical information is provided in the full
list .xls file online. In addition, information concerning the presence
of NPP types' descriptions and/or illustrations in the articles is
given. In Table 1, each publication is described by its citation (first au-
thor, year of publication), the title of the journal/book, the presence of
descriptions and/or illustrations of new or previously published NPP
types (yes/no), the acronym used if NPP types have been described,
the age of the analyzed material (palaeo or modern material P/M),
and the continent and nation of the study materials.
It is obvious that a literature list is never complete. I would appre-
ciate very much receiving any relevant additional information.
3. The NPP type list
On the basis of the NPP selected literature listed in Table 1, a list of
NPP types has been compiled. It includes the NPP types that have
been published from 1972 to November, 2011. Since 1978, the defini-
tion of a new NPP type follows a procedure developed in the Univer-
sity of Amsterdam (van Geel, 1972, 1978). A new NPP type is defined
when the description of its morphologic and biometric characters,
good quality light microscope images, and information concerning
the chronological and stratigraphic contexts and its value as a biolog-
ical proxy are published in a scientific journal. “A Type can be consid-
ered as a provisionally, but not formally named form-species”(van
Geel, 1978, p. 47). It “has nothing to do with the taxonomical signifi-
cance of the world type (holotype). In most cases the types represent
not accurately identifiable, more or less homogeneous entities, as far
as a distinction of the particular structure is feasible”(van Geel et al.,
1981, p. 415). Initially the name of a type was formed by the term
“Type”followed by a progressive number (Type No. System). If iden-
tification could be suggested at various taxonomic levels, the number
was followed by the name of the taxon, e.g., Type 61 Mougeotia cf.
gracillima, or Type 62 Zygnemataceae (van Geel, 1978). This was a
good procedure until NPP analysis spread into many labs all over
the world and the description of new types and their publication
were proposed by labs apart from the Hugo de Vries Laboratory at
the University of Amsterdam. A new system, the “HdV-No. System,”
was introduced when the special issue on NPPs was published in
2010 (Vegetation History and Archaeobotany Special Issue 19/5–6,
2010) as a result of a consultation between the special issue editor,
Jean Nicolas Haas, and Bas van Geel. The word “Type”was thus re-
placed with an acronym that identifies the lab where the NPP type
has been described and where specimens preserved in palynological
slides should be stored. A list of the acronyms for the NPP types is
given in Table 2. The new practice is that each lab compiles its own
Fig. 1. Number of scientific papers including NPP analysis published between 1972 and
November, 2011. Data have been retrieved from a personal reference collection (1972–
1989) and by Scopus (Elsevier) and Google scholar for the term “Non-Pollen
Palynomorphs”(1990–2011).
143A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

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Table 1
The NPP literature list (1972–2011). Please note that the reference list at the end of the article incudes only references of citations in the text. The citations in the table not included
in the references can be found in the complete list published as “Supplementary material 1”on-line.
Citation Journal/proceed Des/Ill Acr. Age Continent Nation
Ahrens (1996) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Aichner et al. (2011) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Asia China
Almeida et al. (2005) Rev. Palaeobot. Palynol. No P America —north Mexico
Andreev et al. (2008) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Asia Russia
Anselmetti et al. (2010) Swiss J Geosci No P Europe Switzerland
Aptroot A Palyno-Bull No M Spain
Aptroot et al. (2006) Rev. Palaeobot. Palynol. Yes P Asia Russia
Argant et al. (2006) Rev. Palaeobot. Palynol. No P Europe France
Bakker et al. (1981) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Bakker et al. (1982) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
Barthelmes et al. (2006) Rev. Palaeobot. Palynol. Yes EMA P Europe Germany
Barthelmes et al. (2010) Folia Geobot. Phytotax. No P Europe Germany
Biserni et al. (2005) Rev. Palaeobot. Palynol. No P Europe Italy
Blackford (1998) In R. Preece (ed) P Europe United Kingdom
Blackford et al. (2006a) Rev. Palaeobot. Palynol. Yes QM M Europe United Kingdom
Blackford et al. (2006b) Rev. Palaeobot. Palynol. No P Europe United Kingdom
Bos et al. (2006) Veg. Hist. Archaeobot. Yes HdV P Europe The Netherlands
Bos et al. (2007) Quaternary Sci. Revi. No P Europe The Netherlands
Brinkkemper et al. (1987) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
Broerse (1991) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Buurman et al. (1995) Meded. Rijks Geol. Dienst Yes HdV P Europe The Netherlands
Carrion (2002a) Quaternary Sci. Revi. No P Europe Spain
Carrion (2002b) Rev. Palaeobot. Palynol. No M Europe Spain
Carrion et al. (1999) Rev. Palaeobot. Palynol. Yes HdV P Europe Spain
Carrion et al. (2000) Veg. Hist. Archaeobot. No P Africa Various
Carrion et al. (2001a) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Europe Spain
Carrion et al. (2001b) The Holocene No P Europe Spain
Carrion et al. (2001c) J. Ecol. No P EUrope Spain
Carrion et al. (2002) Ann. Bot. Fennici No P Europe Spain
Carrion et al. (2003) The Holocene No P Europe Spain
Carrion et al. (2004) Rev. Palaeobot. Palynol. No P Europe Spain
Carrion et al. (2005) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Europe Spain
Carrion et al. (2007) Quaternary Sci. Revi. No P Europe Spain
Carrion et al. (2010) Rev. Palaeobot. Palynol. No P Europe Spain
Carrión et al. (2010) The Holocene No P Europe Spain
Chambers et al. Mires and Peat Yes P
Chnura et al. (2006) Rev. Palaeobot. Palynol. Yes P America —north United States
Cook (2009) Rev. Palaeobot. Palynol. Yes A P Australia Victoria
Cook et al. (2011) Palynology Yes A P Australia Victoria
Costantini et al. (2009) Quatern. Int. No P Europe Italy
Court-Picon et al. (2005) Rev. Palaeobot. Palynol. No M Europe France
Court-Picon et al. (2006) Veg. Hist. Archaeobot. No M Europe France
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Davis (1987) Quaternary Res. Yes P America —north United States
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De Klerk et al. (2009) Boreas No M Asia Russia
de Vries (1992) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Demske et al. (2009) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Asia India
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Ejarque et al. (2011) Rev. Palaeobot. Palynol. No M Europe Andorra
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Feeser et al. (2010) Veg. Hist. Archaeobot. No P Europe Ireland
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Filipova-Marinova (2007) Compt. rend. de l'Acad. Bulg. Sci. No P Europe Bulgaria
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Garneau (1993) Academical Thesis Yes HdV P America —north Canada
Garneau (1998) Bull. Geol. Surv. Can. Yes Type P America —north Canada
Gauthier et al. (2010) Veg. Hist. Archaeobot. Yes IGA P America —north Greenland
Gelorini et al. (2011) Rev. Palaeobot. Palynol. Yes UG M Africa Uganda
Goris et al. (1981) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Goslar et al. (1999) J. Paleolimnol. No P Europe Poland
Grabandt (1990) In Rabassa J, Salemme M (eds) M America —north Canada
Graf et al. (2006) Rev. Palaeobot. Palynol. No M America —north Canada
Grenfell (1995) Rev. Palaeobot. Palynol. Yes P
Haas (1996) Rev. Palaeobot. Palynol. Yes P Europe Switzerland
Haas (2010) Veg. Hist. Archaeobot. Yes P
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144 A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

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Table 1 (continued)
Citation Journal/proceed Des/Ill Acr. Age Continent Nation
Hoaen et al. (2000) In Nicholson RA, O'Connor TP (eds) P
Hoshaw et al. (1988) Phycologia
Innes et al. (2003) J. Archaeol. Sci. No P Europe United Kingdom
Innes et al. (2004) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Europe United Kingdom
Innes et al. (2010) Veg. Hist. Archaeobot. No P Europe United Kingdom
Jankovska (1990) Folia Geobot. Phytotax. P Europe Czech Republic
Jankovska (2007) In Stursa J, Knapik R (eds) No P Europe Czech Republic
Jankovska et al. (1995) Folia Geobot. Phytotax. Yes P Europe Czech Republic
Jankovska et al. (2000) Folia Geobot. Phytotax. No P
Jankovska et al. (2002) Folia Geobot. Phytotax. No P Europe Czech Republic
Jankovska et al. (2006) Boreas No P Asia Russia
Kholeif et al. (2010) J. Afr. Earth Sci. No P Africa Egypt
Klaver (1981) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Koff et al. (2011) Est. J. Earth Sci. No P Europe Latvian
Komarek et al. (2001) Bibliotheca Phycologica Yes P
Kouli et al. (2009) Geobios No P Europe Greece
Kramer et al. (2010) Veg. Hist. Archaeobot. Yes NCP P Asia China
Kumaran et al. (2008) Curr Sci India No P Asia India
Kurhy (1985) Rev. Palaeobot. Palynol. Yes t/T P Europe Germany
Kurhy (1988) Rev. Palaeobot. Palynol. Yes t/T P America —south Colombia
Kurhy (1997) Rev. Palaeobot. Palynol. Yes t/T P America —north Canada
Kvavadze et al. (2008) Veg. Hist. Archaeobot. Yes P Asia Georgia
Kvavadze et al. (2010a) Veg. Hist. Archaeobot. Yes P Asia Georgia
Kvavadze et al. (2010b) Veg. Hist. Archaeobot. Yes P Asia Georgia
Laine et al. (2010) C. R. Biologies No P Europe France
Lamentowicz et al. (2010) J. Paleolimnol. No P Europe Poland
Ledru et al. (2006) Quat. Sci. Reviews No P America —south Brazil
Leroy et al. (2009) Quat. Sci. Reviews No M/P Europe Turkey
Leroy et al. (2010) Rev. Palaeobot. Palynol. No P Europe Turkey
Limaye et al. (2007) Curr Sci India No P Asia India
Looze (1989) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
López et al. (2003) Veg. Hist. Archaeobot. No P Asia Russia
López-Merino et al. (2006) Revista Españ. Micropal. No P Europe Spain
López-Merino et al. (2010) J. Archaeol. Sci. No P Europe Spain
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López-Merino et al. (2011) J. Palaeolimnol. No P Europe Spain
López-Sáez et al. (2006) Plant Ecol. No P Europe Spain
López-Sáez et al. (2007) Revista Españ. Micropal. No P Europe Spain
López-Sáez et al. (2009) Quatern. Int. No P Europe Spain
López-Sáez et al. (1998) Revue de Paleobiologie P
López-Sáez et al. (2002) Quatern. Int. No P Europe Spain
López-Sáez et al. (2009) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Europe Spain
Marinova et al. (2006) Rev. Palaeobot. Palynol. No P Europe Bulgaria
Mauquoy et al. (2004) Quaternary Res. Yes HdV P America —south Argentina
Mazier et al. (2006) The Holocene No M Europe France
Mazier et al. (2009) The Holocene M/P Europe Pays Basque
McAndrews et al. (2010) Veg. Hist. Archaeobot. Yes P America —north Canada
Medeanic (2004) Iheringia, Sér. Bot., Porto Alegre No M America —south Brazil
Medeanic (2006a) Iheringia, Sér. Bot., Porto Alegre No M America —south Brazil
Medeanic (2006b) Rev. Palaeobot. Palynol. Yes P America —south Brazil
Medeanic et al. (2005) J. Coastal Res. No M/P America —south Brazil
Medeanic et al. (2007) Gravel No M America —south Brazil
Medeanic et al. (2008) Rev. Geol. Am. Central Yes Type M America —south Costa Rica
Medeanic et al. (2010a) Int. J. Coal Geol. No P America —south Brazil
Medeanic et al. (2010b) J. Coastal Res. No M America —south Brazil
Menozzi et al. (2010) Veg. Hist. Archaeobot. Yes GE P Europe Italy
Merkt et al. (1999) Quatern. Int. No P Europe Germany
Miehe et al. (2009) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Asia China
Mighall et al. (2008) Veg. Hist. Archaeobot. No P Europe Ireland
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Pals et al. (1976) Ber. Rijksd. Oudheidk. Bodemonderz. Yes HdV P
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Pini (2004) J. Quaternary Sci. No P Europe Italy
(continued on next page)
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Table 1 (continued)
Citation Journal/proceed Des/Ill Acr. Age Continent Nation
Pini et al. (2009) Quat. Sci. Reviews No P Europe Italy
Pires et al. (2006) Revista Españ. Micropal. No P America —south Brazil
Prager et al. (2006) Rev. Palaeobot. Palynol. Yes EMA M Europe Germany
Ralska-Jasiewiczowa (1992a) Veg. Hist. Archaeobot. P Europe Poland
Ralska-Jasiewiczowa (1992b) Sveriges Geologiska Undrsokning, Ser. Ca P Europe Poland
Ran (1990) Meded. Rijks Geol. Dienst Yes HdV P Europe The Netherlands
Riera et al. (2006) Rev. Palaeobot. Palynol. P Europe Spain
Rull et al. (2008) J. Palaeolimnol. Yes LVA P America —south Venezuela
Rull et al. (2011) J. Palaeolimnol. No P Europe Spain
Ryan et al. (2010) Veg. Hist. Archaeobot. No P Europe United Kingdom
Schofield et al. (2011) Veg. Hist. Archaeobot. No P Europe Greenland
Slopsma (1997) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
Smith et al. (2009) Quat. Sci. Reviews No P Europe United Kingdom
Spaargaren (1988) Unpubb. Uni Amst. Yes HdV P Europe The Netherlands
Speranza et al. (2000) The Holocene Yes HdV P Europe Czech Republic
Starnberger et al. (2009) Quatern. Int. No P Europe Austria
Stolze et al. (2007) Rev. Palaeobot. Palynol. No P America —north Canada
Stutz et al. (2010) J. Paleolimnol. No P America —south Argentina
Texier et al. (2003) Prehistoria Alpina No P Europe France
Turton et al. (2006) Rev. Palaeobot. Palynol. Yes P America —north Canada
Urban et al. (2009) J. Arid Environ. No P Asia Oman
Valero-Garcés et al. (2006) unpublished No P Europe Spain
Valsecchi et al. (2010) J. Quaternary Sci. No P Europe Switzerland
van Bennekom (1992) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe The Netherlands
van Dam et al. (1988) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
van der Harst (1984) Int. Rap. H. de Vries Lab, Uni Amst. Yes HdV P Europe Ireland
van der Linden et al. (2006) Palaeogeogr. Palaeoclimatol. Palaeoecol. Yes HdV P Europe Sweden
van der Linden et al. (2008a) Palaeogeogr. Palaeoclimatol. Palaeoecol. Yes HdV P Europe Sweden
van der Linden et al. (2008b) Rev. Palaeobot. Palynol. Yes HdV P Europe Germany
van der Linden et al. (2008c) Palaeogeogr. Palaeoclimatol. Palaeoecol. No P Europe Sweden
van der Molen (1988) Proc. R. Ir. Acad. Yes Type P Europe Ireland
van der Plas et al. (2011) J. Quaternary Sci. No P Africa Mauritius
van der Wiel (1982) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
van Geel (1972) Acta Bot. Neerl. Yes HdV P Europe The Netherlands
van Geel (1976a) Unpubblished Uni Amst. Yes HdV P
van Geel (1976b) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
van Geel (1978) Rev. Palaeobot. Palynol. Yes HdV P Europe Various
van Geel (1986) In Berglund B.E. (eds) No P
van Geel (1998) Palynology P
van Geel (2001) In J.P. Smol, H.J.B. Birks and W.M. Last (eds) No P
van Geel (2006) Rev. Palaeobot. Palynol. No P
van Geel et al. (1978) Rev. Palaeobot. Palynol. Yes HdV P America —south Colombia
van Geel et al. (1981) Rev. Palaeobot. Palynol. Yes HdV P Europe The Netherlands
Van Geel et al. (1983a) Ber. Rijksd. Oudheidk. Bodemonderz. Yes HdV P Europe The Netherlands
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van Geel et al. (1988b) New. Phytol. Yes HdV P Europe Ireland
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146 A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

Author's personal copy
list of described NPPs, but lacking a system that controls the attribu-
tion of the names, the possibility of attributing the same name to dif-
ferent microfossils and vice versa exists and should be avoided. Each
microfossil should have its own name and precise literature refer-
ence. An updated NPP type list should contribute to correct name def-
initions. This is the aim of the list proposed below, which includes the
NPP types described in the reviewed literature listed in Table 1. In the
complete version proposed as a .xls file in the supplementary materi-
al, each type is described in terms of categories (hereafter) that allow
different queries and facilitate the researching of information
concerning published types.
3.1. Acronym of the lab/university
The acronym followed by a hyphen and a progressive number in-
dicates that the NPP type could not be referred to a specifictaxon at
the moment of publication, e.g., HdV-303. The acronym identifies
the lab in which the NPP type was described and where specimens
preserved in palynological slides should be stored. Some authors
have used acronyms to indicate the study site, e.g., BRN —Burren, in
Feeser and O'Connell (2009), NCP —Naleng Cuo Palynomorphs, in
Kramer et al. (2010), or the types of fossils, micro (t) or macro (T),
in Kuhry (1988). The meaning of the acronyms and relative refer-
ences are shown in Table 2. In the NPP type list, the NPP types origi-
nally named as “Type No”by van Geel and colleagues have been
renamed by substituting “Type”for “HdV,”supposing that the speci-
mens are stored in the Hugo de Vries Laboratory (information is not
given in the papers). “Type”has been also maintained if the author
could not have access to the literature, e.g., Type-900, -901, -955
and -954 (Garneau, 1998 in Kuhry, 1997). The use of “Type”before
“HdV”or other acronyms should be avoided because these acronyms
are devoted only to types. By contrast, the acronym and number
should be retained, even when a taxonomical determination is pro-
posed, in order to maintain the link with the original publication
and especially with the lab specimens' collection. The online version
of the list also includes NPP types that have been found in palynolog-
ical slides and were soon identified, but were not referred to in terms
of NPP types at the time of their discovery, e.g., Closterium cf. kützingii
and Debarya aff. D. glyptosperma (van Geel and van der Hammen,
1978). In these cases the acronym and the related number are not
indicated.
3.2. Number
The number of the NPP type is not always indicated when the mi-
crofossil is directly defined with the taxonomical identification in the
original publication. Many HdV-types are not in the NPP type list, be-
cause they are no longer in use or have never been published or have
been skipped during the research (van Hoeve and Hendrikse, 1998),
e.g., HdV-153 to HdV-157, HdV-270 to HdV-302, HdV-428 to
HdV-460, etc. It is also possible that some of them have been de-
scribed in literature that the author could not review.
3.3. Letter
In some cases, a capital letter follows the type number without
space, e.g., HdV-128A, HdV-128B, and HdV-55A. It indicates a differ-
ent NPP type whose morphology may indicate a taxonomical rela-
tionship with the type having the same number. Small letters in the
captures of plates with illustrations of types have a different meaning.
A full stop and a small letter after the type number, e.g., HdV-231.a,
HdV-231.b or HdV-55A.a, HdV-55A.b, indicate different specimens
of the same type (van Geel et al., 1989).
3.4. Wide identification category
(Acritarchs, Algae, Protozoa, Fungi, Bryophyta, Pteridophyta,
Planta, Animalia, Unknown). The categories can be followed by a
question mark if the attribution of the NPP type to the category is in-
dicated as possible/probable in the original description.
3.5. Taxonomical identification
In this column, the taxonomical identification is indicated if it is
reported in the literature. The level of taxonomical identification is
quite various. It is reported as appears in the literature: i) NPP types
attributed to known species or genera or families, e.g., HdV-14 Meliola
elisii, HdV-32A Assulina muscorum, HdV-112 Cercophora sp./spec.,
HdV-58 Zygnemataceae, if the published literature allowed a high de-
gree of taxonomic precision. This means that the morpho-biometric
characters of the remains are certain and correspond to the characters
of such remains of the taxon; ii) genus-type, e.g., HdV-112
Cercophora-type, HdV-261 Arnium-type, when morphological resem-
blance was superficial or ambiguous. Such morphotypes mostly rep-
resent not accurately identifiable but more or less homogeneous
entities, based on conservative distinctions of general morphological
features of shape, size, and surface texture (van Geel, 1978; van
Geel et al., 1981); iii) “cf.”(confer,conferatur = Engl. compare with)
is generally used when a morphotype resembles a known species or
genus. In the extra-European NPP types, “cf.”is often used because a
positive association with European species or genera could not be
made due to fragmentary knowledge of the distribution of species,
e.g., UG-1250 Curvularia cf. comoriensis (Gelorini et al., 2011); iv) the
slash symbol separates different taxonomical identifications reported in
thereviewedliteratureforthesameNPPtype,e.g.,HdV-55CNeurospora
crassa/Neurospora sp.
3.6. Description
This column appears only in the online version. For the most com-
mon unknown NPP types the morpho-biometric description is
reported from literature. This column can be used to query the cata-
logue on the basis of morphological terminology (ascospore, conidia,
1-septate, 2-septate, oocyte, etc.).
Table 2
Acronyms used for the definition of the NPP types in the “HdV-No. System.”*In the full
version of Tab. 3 “micro”has been added to distinguish “t”from “T”, and to alphabeti-
cally order items according the column “Acr.”(= acronym).
Acron. Laboratories/study sites/other References
A Additional types to the hundreds of
Types already catalogued by van Geel
Cook (2009)
BRN BuRreN —Western Ireland Feeser and O'Connell
(2009)
EMA Ernst-Moritz-Arndt-University of
Greifswald —Germany
Prager et al. (2006)
GE Laboratory of Palynology, University
of Genova —Italy
Menozzi et al. (2010)
HdV Hugo de Vries Laboratory, University
of Amsterdam —The Netherlands
Van Geel et al. (2010)
IBB Palynology and Paleoecology Lab, Botanical
Institute of Barcelona (CSIC-ICUB) —Spain
Montoya et al. (2010)
IIB Innsbruck Institute of Botany —Austria Pers. com.
IGA Igaliku Lake —Southwestern Greenland Gauthier et al. (2010)
LVA Laguna Verde Alta, Mérida Andes —Venezuela Rull et al. (2008)
NCP Naleng Cuo Palynomorphs, Tibetan
Plateau —China
Kramer et al. (2010)
QM Department of Geography, Queen Mary,
University of London —Great Britain
Blackford and Innes
(2006)
t* Microfossil type Kuhry (1988)
T Macrofossil type Kuhry (1988)
TM University of Toulouse —le Mirail,
Toulouse —France
Cugny et al. (2010)
147A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

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Table 3
The list of NPP types described by Bas van Geel and colleagues and literature references. Acr.: acronym; N.: type number; L.: letter.
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 1 Fungi Gelasinospora sp. van Geel (1978) van Geel et al. (1981);
Kurhy (1985);
Kuhry (1997);
van Geel et al. (2006a);
Chambers et al. (2011);
Herrmann et al. (2010);
Medeanic et al. (2010a)
HdV 2 Fungi Gelasinospora cf. retispora van Geel (1978) van Geel et al. (1981);
van Geel et al. (2006a);
Chambers et al. (2011)
HdV 3 A Fungi van Geel (1978) van Geel et al. (1988b);
Kurhy (1985)
HdV 3 B Fungi Pleospora sp. van Geel (1978) Kurhy (1985);
van Geel et al. (2006a)
HdV 3 Fungi van Geel (1972) Kurhy (1985)
HdV 4 Fungi Anthostomella cf. fuegiana van Geel (1978) van Geel et al. (1981);
Kurhy (1985);
van Geel et al. (2006a)
HdV 5 Fungi van Geel (1978)
HdV 6 Fungi Coniochaeta xylariispora van Geel (1978) Kurhy (1985); van Geel et al. (2006a)
HdV 7 A Fungi Chaetomium sp. van Geel (1978) van Geel et al. (1981);
Kurhy (1985); van Geel et al. (2003a);
van Geel et al. (2006a);
Medeanic et al. (2010a)
HdV 7 B Fungi van Geel (1978)
HdV 7 Fungi van Geel (1972)
HdV 8 A Fungi van Geel (1978) van Geel et al. (1981)
HdV 8 B Fungi Microthyrium sp.? Mycrothyriaceae van Geel (1978) van Geel et al. (1981);Kuhry (1997)
HdV 8 C Fungi cf. Actinopeltis sp. van Geel (1978) van Geel et al. (2006a)
HdV 8 D Fungi van Geel (1978)
HdV 8 E Fungi van Geel (1978)
HdV 8 F Fungi Stomiopeltis sp. van Geel (1978) van Geel et al. (2006a)
HdV 8 G Fungi van Geel (1978)
HdV 8 Fungi ? Microthyrium spec. van Geel (1972) Herrmann et al. (2010)
HdV 9 Unknown van Geel (1978)
HdV 10 Fungi Trigocladium opacum van Geel (1978) Bakker et al. (1982); Kurhy (1985);
van Geel et al. (1988b);
Chambers et al. (2011)
HdV 11 Fungi van Geel (1978) van Geel et al. (1981)
HdV 12 Fungi van Geel (1978) van Geel et al. (1981);
Bakker et al. (1982); Kurhy (1985);
Chambers et al. (2011)
HdV 13 Fungi cf. Entophyctis lobata van Geel (1978) van Geel et al. (1981);
Kurhy (1985); Kuhry (1997)
HdV 14 Fungi Meliola ellisii (synonym
of M. niessleana)
van Geel (1978) Kurhy (1985); van Geel et al. (2006a)
HdV 14 Fungi Meliola ellisii (synonym
of M. niessleana)
van Geel (1978) van Geel et al. (2006a)
HdV 14 Fungi Meliola ellisii (synonym
of M. niessleana)
van Geel (1978) Kurhy (1985); van Geel et al. (2006a);
Chambers et al. (2011)
HdV 15 Fungi HdV-20 van Geel (1978)
HdV 16 A Fungi Byssotecium circinnans van Geel (1978) van Geel et al. (1981);
van Geel et al. (2006a)
HdV 16 B Fungi van Geel (1978)
HdV 16 C Fungi van Geel (1978)
HdV 16 Fungi Curvularia tuberculata/
Byssothecium circinans
van Geel (1972) van Geel et al. (1988b);
van Geel et al. (2006a);
Medeanic et al. (2010a)
HdV 17 Fungi van Geel (1978)
HdV 18 A+ B Fungi van Geel (1978) van Geel et al. (1981)
HdV 19 Fungi van Geel (1978)
HdV 20 Fungi HdV-15 HdV-21 van Geel (1978) Kurhy (1985)
HdV 21 Fungi HdV-15 HdV-20 van Geel (1972)
HdV 22 Fungi Leptoshaeria/Herpotrichiella sp. van Geel (1978)
HdV 23 Fungi van Geel (1978) van Geel et al. (1981);
van Geel et al. (1988b)
HdV 24 Fungi van Geel (1978) Kurhy (1985)
HdV 25 Fungi cf. Clasterosporium caricinum van Geel (1978) Prager et al. (2006);
Medeanic et al. (2010a)
HdV 26 Fungi van Geel (1978)
HdV 27 Fungi Tilletia sphagni van Geel (1978) van der Wiel (1982); Kurhy (1985);
Kuhry (1997)
HdV 28 Animalia Copepoda Canthocamptus spermatophores van Geel (1978) van Geel et al. (1988b)
HdV 29 Unknown van der Hoven et al. eds (1998)
148 A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

Author's personal copy
Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 30 Fungi Helicosporium/Helicoon
pluriseptatum
van Geel (1978) van Geel et al. (1988b);
van Geel et al. (1981);
Kurhy (1985); Kurhy (1997);
Medeanic et al. (2010a)
HdV 31 A Protozoa Rhizopoda Amphitrema flavum HdV-186 van Geel (1978) van Geel et al. (1981);
van Geel et al. (1989);
Kurhy (1985); Kuhry (1997)
HdV 31 B Protozoa Rhizopoda Amphitrema
wrightianum
van Geel (1978)
HdV 32 A Protozoa Rhizopoda Assulina muscorum van Geel (1978) Kurhy (1985); Kuhry (1997);
Prager et al. (2006)
HdV 32 B Protozoa Rhizopoda Assulina seminulum van Geel (1978) Kurhy (1985)
HdV 32 Protozoa Rhizopoda Assulina sp. van Geel et al. (1988b) van Geel et al. (1989)
HdV 33 Unknown van Geel (1978) van Geel et al. (1981)
HdV 34 Unknown van Geel (1978)
HdV 35 Algae Dinoflagellates peridinoid van Geel (1978) Bakker et al. (1982);
van Geel et al. (1988b);
van Geel et al. (1989)
HdV 36 Animalia Acari Oribatida van Geel (1978)
HdV 37 Animalia Rotifera Habrotrocha (Callidina)
angusticollis
van Geel (1978) van Geel et al. (1981);
Kurhy (1985);
Kurhy (1997)
HdV 38 Unknown van Geel (1978)
HdV 39 Unknown van Geel (1972)
HdV 40 Unknown van Geel (1972)
HdV 41 Unknown van Geel (1978) Kuhry (1997)
HdV 42 Planta van der Hoven et al. eds (1998)
HdV 43 Planta van der Hoven et al. eds (1998)
HdV 44 Fungi cf. Ustulina deusta (=Kretzschmaria
deusta)/Xylariaceae e.g. Rossellinia —
van der Wiel (1982)
HdV-117 van Geel (1978) van Geel et al. (1981);
van der Wiel (1982);
van Geel et al. (1986);
van Geel et al. (2006a);
Prager et al. (2006)
HdV 46 Protozoa Rhizopoda Hyalosphaenia subflava van Geel (1978) Kuhry (1997)
HdV 47 Fungi van Geel (1978)
HdV 48 Fungi HdV-14 van der Hoven et al. eds (1998)
HdV 49 Fungi van Geel (1978)
HdV 51 Fungi cf. Sporidesmium spec. van Geel (1978)
HdV 53 Fungi van Geel (1978)
HdV 54 Fungi van Geel (1978) van Geel et al. (1988b)
HdV 55 A Fungi Sordaria‐type van Geel (1978) van Geel et al. (1981);
van Geel et al. (1983b);
van Geel et al. (1989);
Kuhry (1997);
van Geel et al. (2003a);
van Geel et al. (2006a);
Aptroot et al. (2006);
Prager et al. (2006);
Marinova et al. (2006);
van Geel et al. (2007);
Medeanic et al. (2010a)
HdV 55 A1 Fungi HdV-55A van Geel et al. (1981)
HdV 55 A2 Fungi van Geel et al. (1981) Bakker et al. (1982)
HdV 55 B Fungi Sordariaceae/Sordaria van Geel (1978) van Geel et al. (1983b);
Kurhy (1985);
Marinova et al. (2006);
Prager et al. (2006)
HdV 55 C Fungi Neurospora crassa/Neurosora sp. van Geel (1978) van Geel et al. (2006a);
Chambers et al. (2011);
Medeanic (2010a)
HdV 58 Algae Zygnemataceae van Geel (1978) van der Wiel (1982)
HdV 59 Unknown van Geel (1978) Bakker et al. (1982)
HdV 60 Algae Closterium idiosporum-type van Geel (1978) van Geel et al. (1981);
Kurhy (1985)
HdV 61 Algae Mougeotia cf. gracillima van Geel (1978) Kurhy (1985)
HdV 62 Algae Zygnemataceae van Geel (1978) Kurhy (1985)
HdV 63 A Fungi Lasiosphaeria cf. caudata/
L. caudata
van Geel (1978) van Geel et al. (2006a)
HdV 63 B Fungi Lasiosphaeria-like van Geel (1978)
HdV 63 C Fungi Lasiosphaeria spec. van Geel (1978)
HdV 63 Fungi Lasiosphaeria HdV-63A van Geel et al. (1981) Kuhry (1997)
HdV 64 Fungi van Geel (1978)
HdV 65 Fungi van Geel (1978) van Geel et al. (1981);
Bakker et al. (1982)
HdV 66 Algae Penium/Tetraedron cf. minimum van Geel (1978)
HdV 67 Fungi Cladosporium van Geel (1978)
(continued on next page)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 68 Planta Erica tetralix van Geel (1978)
HdV 69 Unknown van Geel (1978)
HdV 70 Animalia Rotifera Cephalodella exigua van Geel (1978)
HdV 71 Animalia Araneida van Geel (1978)
HdV 72 A Animalia Cladocera Alona rustica van Geel (1978)
HdV 72 B Animalia Cladocera Alona rustica van Geel (1978)
HdV 72 C Animalia Cladocera undiff. van Geel et al. (1983b)
HdV 72 D Animalia Cladocera Eurycercus cf. lamellatus van Geel et al. (1983b) van Geel et al. (1989)
HdV 72 Animalia Cladocera Alona rustica van Geel (1978) van Geel et al. (1983b)
HdV 73 Fungi? van Geel (1978)
HdV 74 Unknown van Geel (1978) Miola et al. (2006)
HdV 75 Animalia Invertebrata van Geel (1978)
HdV 76 Unknown van Geel (1978)
HdV 77 A Fungi Geoglossum sphagnophilum van Geel (1978) van Geel et al. (1981);
van der Wiel (1982);
van Geel et al. (2006a)
HdV 77 B Fungi Trichoglossum cf. hirsutum van Geel (1978) van der Wiel (1982);
van Geel et al. (2006a)
HdV 78 Fungi van Geel (1978)
HdV 79 Fungi van Geel (1978)
HdV 80 Planta Ericales van Geel (1978)
HdV 81 Animalia Insecta eyes? van Geel (1978)
HdV 82 Fungi van Geel (1978)
HdV 83 Fungi van Geel (1978) Kuhry (1997)
HdV 84 Animalia van Geel (1978)
HdV 85 Fungi Didymosphaeria massarioides van Geel (1978) van Geel et al. (2006a)
HdV 86 Unknown van Geel (1978)
HdV 87 Fungi van Geel (1978) van Geel et al. (1981)
HdV 88 A Animalia Invertebrata mandible van Geel (1978)
HdV 88 B Animalia Invertebrata mandible van Geel (1978)
HdV 89 Fungi Tetraploa aristata van Geel (1978) Marinova et al. (2006);
van Geel et al. (2011)
HdV 90 Fungi van Geel (1978) van Geel et al. (1981);
Kurhy (1985); Kuhry (1997)
HdV 91 Unknown van Geel (1978)
HdV 92 Fungi van Geel (1978)
HdV 93 Fungi Xylariaceae van Geel (1978)
HdV 95 Fungi van Geel (1978)
HdV 96 A Fungi Several species of dematiaceous genera van Geel (1978)
HdV 96 B Fungi van Geel (1978)
HdV 98 Fungi cf. Spadicoides bina van Geel (1978)
HdV 99 Fungi van Geel (1978)
HdV 100 Algae cf. Cylindrocystis brebissonii van Geel (1978) van Geel et al. (1981)
HdV 101 Animalia? van Geel (1978)
HdV 103 Animalia Rotifera Stephanoceros eichhornii van Geel (1978) Medeanic et al. (2010a)
HdV 104 Fungi van Geel (1978)
HdV 105 Fungi van Geel (1978)
HdV 106 A Animalia Rhysotritia ardua HdV-106 Bakker et al. (1982)
HdV 106 B Animalia Rhysotritia ardua Bakker et al. (1982)
HdV 106 Animalia Rhysotritia ardua Bakker et al. (1982)
HdV 106 Planta van Geel (1978)
HdV 107 Animalia Coleoptera Carabidae mandible van Geel (1978) Bakker et al. (1982)
HdV 108 Animalia Insecta Tupulidae Prionocera sp. van Geel (1978) van Geel et al. (1989)
HdV 109 Animalia Insecta Myrmicinae van Geel (1978) van Geel et al. (1989)
HdV 110 Animalia? van Geel (1978)
HdV 111 Animalia? van Geel (1978)
HdV 112 A Fungi Cercophora sp. van Geel et al. (1981)
HdV 112 B Fungi Cercophora sp. van Geel et al. (1981)
HdV 112 C Fungi Cercophora sp. van Geel et al. (1983b)
HdV 112 Fungi Cercophora-type/Cercophora sp. van Geel (1978) van Geel et al. (1981);
van der Wiel (1982);
van Geel et al. (1983b);
van Geel et al. (2003a);
van Geel et al. (2006a);
Menozzi et al. (2010)
HdV 113 Fungi Sporormiella-type/Sporormiella sp. van Geel et al. (2003a) van Geel et al. (2006a);
van Geel et al. (2007);
Medeanic et al. (2010a)
HdV 114 Planta Alnus/Corylus/Betula Pals et al. (1980) van der Wiel (1982);
van Geel et al. (1989);
Prager et al. (2006)
HdV 115 Acritarchs Mycrhystridium Pals et al. (1980) Bakker et al. (1982)
HdV 116 Acritarchs Cymatiosphaera Pals et al. (1980) Bakker et al. (1982)
HdV 117 Fungi cf. Ustulina deusta HdV-44 Pals et al. (1980) van Geel et al. (1981);
van Geel et al. (1988a)
HdV 118 Fungi Pals et al. (1980)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 119 Unknown Pals et al. (1980) Carrion et al. (2002)
HdV 120 Fungi Pals et al. (1980)
HdV 121 Fungi Pals et al. (1980) Kuhry (1997)
HdV 122 Fungi Pals et al. (1980)
HdV 123 Fungi Pals et al. (1980) van Geel et al. (1981);
Bakker et al. (1982)
HdV 124 Fungi cf. Persiciospora sp. Pals et al. (1980) Bakker et al. (1982);
van der Wiel (1982);
van Geel et al. (2006a)
HdV 125 Fungi Pals et al. (1980)
HdV 126 Fungi Gaeumannomyces/G.
cf. caricis/Clasterosporium
caricinum hyphopodia
Pals et al. (1980) van der Wiel (1982);
van Geel et al. (1983b);
van Geel et al. (1989);
Kuhry (1997);
Carrion et al. (2002);
van Geel et al. (2006a);
Prager et al. (2006)
HdV 127 Planta Nymphaeaceae Pals et al. (1980)
HdV 128 A Algae? van der Wiel (1982) van Geel et al. (1983b);
van Geel et al. (1989);
Carrion et al. (2002);
Prager et al. (2006);
Kramer et al. (2010);
Miola et al. (2006)
HdV 128 B Algae? van der Wiel (1982) van Geel et al. (1983b);
van Geel et al. (1989);
Miola et al. (2006)
HdV 128 Unknown Pals et al. (1980) Carrion et al. (2000)
HdV 129 Planta Nymphaeaceae trichosclereids Pals et al. (1980) Kuhry (1997)
HdV 130 Algae Spirogyra 1 = type 315
= type C smooth
Pals et al. (1980) van der Wiel (1982);
van Geel et al. (1983b)
HdV 131 Algae Spirogyra 2Pals et al. (1980) van der Wiel (1982)
HdV 132 Algae Spirogyra reticulate Pals et al. (1980) van der Wiel (1982)
HdV 133 Algae Mougeotia Pals et al. (1980) van der Wiel (1982)
HdV 134 Algae Mougeotia Pals et al. (1980)
HdV 135 Algae Mougeotia Pals et al. (1980) van der Wiel (1982)
HdV 136 Algae Mougeotia Pals et al. (1980) van der Wiel (1982)
HdV 137 Planta Ceratophyllum sp. Pals et al. (1980) van Geel et al. (1981);
Kuhry (1985);
van Geel et al. (1989);
Kuhry (1997)
HdV 138 Animalia Porifera Ephydatia cf. fluviatilis Pals et al. (1980)
HdV 139 Animalia Hirudinea Piscicola geometra Pals et al. (1980)
HdV 140 Fungi Valsaria cf. variospora/V. variospora‐type van der Wiel (1982) van Geel et al. (1983b);
van Geel et al. (2003);
van Geel et al. (2006a)
HdV 141 Algae Mougeotia van der Wiel (1982)
HdV 142 Unknown van der Wiel (1982)
HdV 143 Fungi Diporotheca spec/D. rhizophila van der Wiel (1982) van Geel et al. (1986);
van Geel et al. (1989);
van Geel et al. (2003a);
van Geel et al. (2006a);
Prager et al. (2006)
HdV 144 Fungi van der Wiel (1982)
HdV 145 Fungi van der Wiel (1982)
HdV 146 Algae Gloeotrichia-type van der Wiel (1982) van Geel et al. (1983b);
van Geel et al. (1989);
van Geel et al. (1996b);
Kuhry (1997)
HdV 147 Unknown van der Wiel (1982)
HdV 148 Unknown van der Wiel (1982)
HdV 149 Unknown van der Wiel (1982)
HdV 150 Unknown van der Wiel (1982) van Geel et al. (1983b)
HdV 151 Fungi van der Wiel (1982)
HdV 152 Fungi van der Wiel (1982)
HdV 158 Fungi Klaver (1981)
HdV 159 Unknown Klaver (1981)
HdV 160 Unknown Dam et al. (1988)
HdV 161 Animalia Insecta Prionocera sp. van Geel et al. (1989)
HdV 165 Bryophyta cf. Riccia van Geel et al. (1983b)
HdV 166 A Algae Bulbochaete sp./Oedogoniaceae
1 sorocarpa
van Geel et al. (1983b)
HdV 166 B Algae Bulbochaete sp./Oedogoniaceae 2 van Geel et al. (1983b)
HdV 167 Algae? van Geel et al. (1983b)
HdV 168 Fungi van Geel et al. (1983b)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 169 Fungi Zopfiella type/Apiosordaria
verruculosa
van Geel et al. (1983b) van Geel et al. (2006a);
Cugny et al. (2010)
HdV 170 Algae Rivularia‐type van Geel et al. (1983b) van Geel et al. (1989);
van Geel et al. (1996b),
Menozzi et al. (2010)
HdV 171 Fungi Rhytidospora cf. tetraspora van Geel et al. (1983b) van Geel et al. (2006a)
HdV 172 Fungi Coniochaeta cf.
ligniaria/C. lignaria
van Geel et al. (1983b) Marinova et al. (2006);
van Geel et al. (2006a);
van Geel et al. (2011);
Medeanic et al. (2010a)
HdV 173 A Fungi van Geel et al. (1983b) van Geel et al. (1989)
HdV 173 B Fungi van Geel et al. (1989)
HdV 173 Fungi van Geel et al. (1983b)
HdV 174 Fungi van Geel et al. (1983b)
HdV 175 Animalia? van Geel et al. (1983b) Gelorini et al. (2011)
HdV 176 Animalia Copepoda Harpacticoidea van Geel et al. (1983b)
HdV 178 Animalia? Invertebrate? van Geel et al. (1983b)
HdV 179 Animalia Copepode or Protozoa van Geel et al. (1983b) van Geel et al. (1989);
Kuhry (1997);
Carrion et al. (1999);
Carrion et al. (2000);
Carrion et al. (2002)
HdV 180 Animalia Invertebrata HdV-179 van Geel et al. (1983b) van Geel et al. (1989)
HdV 181 Unknown van Geel et al. (1983b)
HdV 182 Unknown van Geel et al. (1983b) Carrion et al. (2002)
HdV 183 Unknown van Geel et al. (1983b)
HdV 184 Unknown van Geel et al. (1983b)
HdV 185 Unknown van Geel et al. (1983b)
HdV 186 Protozoa Rhizopoda Amphitrema flavum-like van Geel et al. (1983b)
HdV 187 A Animalia Invertebrata? van Geel et al. (1983b)
HdV 187 B Animalia Invertebrata? van Geel et al. (1989)
HdV 187 C Animalia Invertebrata? van Geel et al. (1989)
HdV 187 D Animalia Invertebrata? van Geel et al. (1989)
HdV 188 Unknown van Geel et al. (1983b)
HdV 189 A Animalia Cladocera Simocephalus sp. van Geel et al. (1983b)
HdV 189 B Animalia Cladocera Moina sp. van Geel et al. (1983b) van Geel et al. (1989);
Kuhry (1997)
HdV 189 C Animalia Cladocera Daphnia “pulex-type”van Geel et al. (1983b)
HdV 189 D Animalia Cladocera Leydigia-type van Geel et al. (1986)
HdV 189 E Animalia Cladocera Daphnia sp. van Geel et al. (1989)
HdV 190 Animalia cf. Herpobdella? van Geel et al. (1983b)
HdV 191 Animalia Turbellaria van Geel et al. (1983b)
HdV 192 Animalia Bryozoa Lophopus crystallinus van Geel et al. (1983b)
HdV 193 Planta van Geel et al. (1983b)
HdV 194 Animalia Oligochaeta van Geel et al. (1983b)
HdV 195 Animalia Conomelus cf. anceps van Geel et al. (1983b)
HdV 200 Fungi van Geel et al. (1989) Kuhry (1997);
Marinova et al. (2006);
Miola et al. (2006)
HdV 201 Fungi Xylomyces chlamydosporis/
aquaticus
van Geel et al. (1989) Kurhy (1985); Kuhry (1997);
Miola et al. (2010)
HdV 202 Fungi van Geel et al. (1989)
HdV 203 Fungi van Geel et al. (1989)
HdV 204 Fungi van Geel et al. (1989) Carrion et al. (2000)
HdV 205 Fungi Sordariaceae? van Geel et al. (1989) Kuhry (1997)
HdV 206 Fungi van Geel et al. (1989)
HdV 207 Fungi Glomus cf. fasciculatum van Geel et al. (1989) van Geel et al. (2003a);
Marinova et al. (2006)
HdV 208 Unknown van Geel et al. (1989)
HdV 209 Algae HdV-66 van Geel et al. (1989) Prager et al. (2006)
HdV 210 Algae Spirogyra species van Geel et al. (1989)
HdV 211 Algae Spirogyra species van Geel et al. (1989)
HdV 212 Algae Spirogyra undiff. van Geel et al. (1989)
HdV 213 Algae Zygnema type van Geel et al. (1989)
HdV 214 Algae Debarya sp. van Geel et al. (1989)
HdV 215 Bryophyta Meesia triquetra van Geel et al. (1989)
HdV 216 Planta Platanus (?) van Geel et al. (1989)
HdV 217 Planta Pteridophyta and
Spermatophyta
van Geel et al. (1989)
HdV 218 Planta Coniferae tracheids van Geel et al. (1989)
HdV 219 Animalia Invertebrata mandibles, labia van Geel et al. (1989)
HdV 220 Animalia Porifera sponges van Geel et al. (1989)
HdV 221 Animalia Invertebrata? van Geel et al. (1989) Kramer et al. (2010)
HdV 222 Unknown van Geel et al. (1989)
HdV 223 Unknown van Geel et al. (1989)
HdV 224 Unknown van Geel et al. (1989)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 225 Algae? van Geel et al. (1989) Kuhry (1997)
HdV 226 Algae? van Geel et al. (1989)
HdV 227 Unknown van Geel et al. (1989)
HdV 228 Algae? van Geel et al. (1989) Kuhry (1997);
Kramer et al. (2010)
HdV 229 Algae? HdV-74 van Geel et al. (1989)
HdV 230 Algae? Dinoflagellate marine? van Geel et al. (1989) Miola et al. (2010)
HdV 231 Unknown van Geel et al. (1989)
HdV 232 Unknown van Geel et al. (1989)
HdV 233 Unknown van Geel et al. (1989)
HdV 234 Unknown van Geel et al. (1989)
HdV 235 Unknown van Geel et al. (1989)
HdV 236 Unknown van Geel et al. (1989)
HdV 237 Planta Juniperus communis leaves van Geel et al. (1989)
HdV 238 Planta Salix van Geel et al. (1989)
HdV 239 Planta Agrostis-type fruit van Geel et al. (1989)
HdV 240 A Planta Phragmites van Geel et al. (1989) Kuhry (1997)
HdV 240 B Planta Phragmites australi s epidermis van Geel et al. (1989)
HdV 241 Planta Potamogeton sp. van Geel et al. (1989)
HdV 242 Unknown van Geel et al. (1989)
HdV 243 Planta van Geel et al. (1989)
HdV 244 Planta Equisetum fluviatile van Geel et al. (1989)
HdV 245 Planta van Geel et al. (1989)
HdV 246 Planta van Geel et al. (1989)
HdV 247 Planta van Geel et al. (1989)
HdV 248 Animalia Insecta Sialis cf. lutaria van Geel et al. (1989)
HdV 249 Animalia Insecta Sialis cf. lutaria van Geel et al. (1989)
HdV 250 Animalia van Geel et al. (1989)
HdV 251 Animalia Invertebrata among
which Trichoptera
van Geel et al. (1989)
HdV 252 Animalia Coleoptera van Geel et al. (1989)
HdV 253 Animalia Coleoptera van Geel et al. (1989)
HdV 254 Unknown van Geel et al. (1989)
HdV 255 Animalia Arthropoda? van Geel et al. (1989)
HdV 256 Fungi? Ustilaginales? Willemsen et al. (1996)
HdV 257 Unknown Willemsen et al. (1996)
HdV 258 Fungi Willemsen et al. (1996)
HdV 259 Fungi Mycogone sp. Willemsen et al. (1996)
HdV 260 Animalia Willemsen et al. (1996)
HdV 261 Fungi Arnium‐type/A. spp. van Geel et al. (2003) van Geel et al. (2006a)
HdV 262 Fungi Arnium imitans-type/A. spp. van Geel et al. (2003) van Geel et al. (2006a)
HdV 263 Fungi Valsaria-type van Geel et al. (2003) van Geel et al. (2006a);
Medeanic et al. (2010a)
HdV 264 Fungi van der Linden et al. (2006)
HdV 265 Fungi van der Linden et al. (2006)
HdV 266 Fungi van der Linden et al. (2008a)
HdV 267 Fungi van der Linden et al. (2008a)
HdV 268 Unknown van der Linden et al. (2008b)
HdV 269 Unknown van der Linden et al. (2008b)
HdV 303 Algae? van Geel et al. (1981) Miola et al. (2006)
HdV 304 Algae Staurastrum/Cosmarium/
Micrasterias
van Geel et al. (1981)
HdV 305 Fungi van Geel et al. (1981)
HdV 306 Fungi van Geel et al. (1981) van Geel et al. (1989)
HdV 307 A Unknown van Geel et al. (1981)
HdV 307 B Unknown van Geel et al. (1983b)
HdV 308 Unknown van Geel et al. (1981)
HdV 310 Fungi Amphisphaerella dispersella
(=A. amphisphaerioides)
van Geel et al. (1981) van Geel et al. (2006a)
HdV 312 Fungi van Geel et al. (1981)
HdV 313 A Algae Mougeotia sp. van Geel et al. (1981)
HdV 313 B Algae Mougeotia sp. van Geel et al. (1981)
HdV 313 C Algae Mougeotia sp. van Geel et al. (1981) van der Wiel (1982)
HdV 313 D Algae Mougeotia sp. van Geel et al. (1981)
HdV 313 E Algae Mougeotia sp. van Geel et al. (1981)
HdV 313 F Algae Mougeotia cf. punctata HdV-A van Geel et al. (1983b) van Geel et al. (1989)
HdV 313 Algae Mougeotia sp. div. van Geel et al. (1981)
HdV 314 Algae Zygnema type van Geel et al. (1981) van Geel et al. (1989)
HdV 315 Algae Spirogyra sp. HdV-C, HdV-130 van Geel et al. (1981)
HdV 317 Algae van Geel et al. (1981)
HdV 320 Unknown van Geel et al. (1981)
HdV 322 Fungi van Geel et al. (1981)
HdV 324 Fungi van Geel et al. (1981)
HdV 327 Algae cf. Hypoxylon argillaceum van Geel et al. (1981)
HdV 332 A Algae Cosmarium protractum van Geel et al. (1981)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 332 B Algae Cosmarium turpinii var. eximium van Geel et al. (1981) van Geel et al. (1989)
HdV 332 C Algae Euastrum insulare var. lacustre van Geel et al. (1981) van Geel et al. (1989)
HdV 332 D Algae Cosmarium cf. sportella van Geel et al. (1981)
HdV 332 E Algae Cosmarium cf. controversum van Geel et al. (1981)
HdV 332 F Algae Cosmarium cf. botrytis van Geel et al. (1989)
HdV 332 Algae Desmidiaceae van Geel et al. (1981)
HdV 333 Algae cf. Penium van Geel et al. (1981)
HdV 334 Fungi van Geel et al. (1981)
HdV 336 Fungi cf. Apiospora montagnei van Geel et al. (1981)
HdV 339 Fungi van Geel et al. (1981)
HdV 340 Bryophyta van Geel et al. (1981) van Geel et al. (1989)
HdV 341 A Algae? van Geel et al. (1981)
HdV 341 B Algae? van Geel et al. (1981)
HdV 342 Algae Spirogyra cf. scrobiculata van Geel et al. (1981) Kurhy (1985); van Geel et al. (1989);
Miola et al. (2010)
HdV 343 Fungi cf. Scopinella barbata van Geel et al. (1981) van Geel et al. (2006a)
HdV 350 Fungi van Geel et al. (1981)
HdV 351 Fungi van Geel et al. (1981) Miola et al. (2010)
HdV 352 Protozoa Rhizopoda Arcella sp. van Geel et al. (1981) Kuhry (1997); Herrman et al. (2010)
HdV 353 A Animalia Rhabdocoela–Turbellaria van Geel et al. (1981) Miola et al. (2010)
HdV 353 B Animalia Rhabdocoela–Turbellaria van Geel et al. (1981) Miola et al. (2010)
HdV 353 C Animalia Rhabdocoela–Turbellaria van Geel et al. (1981) van Geel et al. (1989)
HdV 353 D Animalia Rhabdocoela–Turbellaria van Geel et al. (1981) van Geel et al. (1989)
HdV 354 Bryophyta van Geel et al. (1981)
HdV 356 Algae? van Geel et al. (1981)
HdV 357 Fungi Puccinia sp. van Geel et al. (1981)
HdV 358 Fungi van Geel et al. (1981)
HdV 359 Fungi Brachysporium obovatum/Brachysporium
bloxami/Bactrodesmium betulicola
van Geel et al. (1981) Carrion et al. (2002)
HdV 360 Fungi cf. Brachysporium sp. van Geel et al. (1981) Medeanic et al. (2010a)
HdV 361 Fungi van Geel et al. (1981) van der Wiel (1982)
HdV 364 Fungi Thecaphora sp. van Geel et al. (1981)
HdV 365 Fungi van Geel et al. (1981)
HdV 366 Unknown van Geel et al. (1981)
HdV 367 Fungi Polyadosporites/Nectria peziza/
Pleosporaceae (e.g. Herpotrichia
lignicola and Parodiella perisporioides)/
Amphisphaeriaceae (e.g. Ceriophora
palustris)
van Geel et al. (1981) van Geel et al. (1983b);
Carrion et al. (1999);
Carrion et al. (2002)
HdV 368 Fungi Podospora-type/P. sp. van Geel et al. (1981) van Geel et al. (2003a);
van Geel et al. (2006a);
van Geel et al. (2007)
HdV 369 Planta cf. Lemna van Geel et al. (1981)
HdV 370 Fungi van Geel et al. (1981)
HdV 371 Algae Tetraedron cf. minimum Bakker et al. (1982)
HdV 372 Algae Closterium cf. rostratum van Geel et al. (1981) Kurhy (1985)
HdV 373 Algae Mougeotia cf. laetevirens van Geel et al. (1981) van der Wiel (1982);
van Geel et al. (1989);
Miola et al. (2010)
HdV 375 A Planta Betula cf. fruticosa van Geel et al. (1981)
HdV 375 B Planta Betula cf. nana van Geel et al. (1981) van Geel et al. (1989)
HdV 375 C Planta Betula cf. pubescens van Geel et al. (1981)
HdV 376 Planta Betula van Geel et al. (1981)
HdV 377 Planta Pinus van Geel et al. (1981)
HdV 378 Fungi Ophiostomataceae van Geel et al. (1981)
HdV 379 Planta Populus tremula van Geel et al. (1981)
HdV 380 A Planta Pinus sylvestris van Geel et al. (1981)
HdV 380 B Planta Pinus sylvestris van Geel et al. (1981)
HdV 381 Planta Pinus sylvestris van Geel et al. (1981)
HdV 382 Planta Populus tremula van Geel et al. (1981)
HdV 383 Unknown van Geel et al. (1981)
HdV 384 Algae Chara van Geel et al. (1981)
HdV 385 Bryophyta Sphagnum van Geel et al. (1981)
HdV 386 Bryophyta Sphagnum van Geel et al. (1981)
HdV 387 Planta Pinus sylvestris van Geel et al. (1981)
HdV 388 Unknown van Geel et al. (1981)
HdV 390 A Animalia Bryozoa Plumatella repens van Geel et al. (1981)
HdV 390 B Animalia Bryozoa Cristatella mucedo van Geel et al. (1981)
HdV 390 C Animalia Bryozoa Hyalinella punctata van Geel et al. (1981)
HdV 391 Planta Pinus sylvestris van Geel et al. (1981)
HdV 392 Planta van Geel et al. (1981)
HdV 393 Planta van Geel et al. (1981)
HdV 394 Planta van Geel et al. (1981)
HdV 395 Planta van Geel et al. (1981)
HdV 396 Animalia Acari Oribatida HdV-36 van Geel et al. (1981)
HdV 397 Animalia Invertebrata van Geel et al. (1981)
154 A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 398 A Planta Betula dwarf van Geel et al. (1989)
HdV 398 B Planta Betula tree van Geel et al. (1989)
HdV 399 Planta Juncus van Geel et al. (1989)
HdV 400 Unknown Hofwegen (1983)
HdV 401 Unknown HdV-146 Hofwegen (1983)
HdV 402 Unknown Hofwegen (1983)
HdV 403 Unknown Hofwegen (1983)
HdV 404 Fungi Hofwegen (1983)
HdV 405 Fungi Hofwegen (1983)
HdV 406 Fungi Hofwegen (1983)
HdV 407 Unknown Hofwegen (1983)
HdV 408 Fungi Hofwegen (1983)
HdV 409 Unknown Hofwegen (1983)
HdV 410 Unknown Hofwegen (1983)
HdV 411 Fungi Hofwegen (1983)
HdV 412 Unknown Hofwegen (1983)
HdV 413 Unknown Hofwegen (1983)
HdV 425 Unknown Ahrens (1996)
HdV 426 Unknown Ahrens (1996)
HdV 427 Fungi Ahrens (1996)
HdV 461 Fungi Kuhry (1985)
HdV 462 Fungi cf. Brachysporium sp. Kuhry (1985)
HdV 463 Fungi Stuartella sp.? Kuhry (1985)
HdV 464 Unknown Kuhry (1985)
HdV 465 Fungi cf. Trichocladium opacum Kuhry (1985)
HdV 466 Fungi Podospora sp./Zopfiella sp. Kuhry (1985)
HdV 470 Unknown Haaster (1984)
HdV 471 Unknown Haaster (1984)
HdV 473 Unknown Haaster (1984)
HdV 480 Unknown Brinkkemper et al. (1987)
HdV 481 Animalia Arthropoda Brinkkemper et al. (1987)
HdV 483 Unknown Brinkkemper et al. (1987)
HdV 484 Animalia Insecta Trichoptera mandible Brinkkemper et al. (1987)
HdV 486 Unknown Brinkkemper et al. (1987)
HdV 487 Animalia Brinkkemper et al. (1987)
HdV 488 Unknown Brinkkemper et al. (1987)
HdV 489 Animalia Brinkkemper et al. (1987)
HdV 490 Animalia Brinkkemper et al. (1987)
HdV 491 Animalia Insecta Brinkkemper et al. (1987)
HdV 492 Unknown van Smeerdijk (1989)
HdV 493 Unknown van Smeerdijk (1989)
HdV 494 Fungi van Smeerdijk (1989)
HdV 495 Fungi van Smeerdijk (1989) Cugny et al. (2010)
HdV 496 Fungi van Smeerdijk (1989)
HdV 497 Algae HdV-356 van Smeerdijk (1989)
HdV 498 Fungi Dyctyosporium van Smeerdijk (1989)
HdV 501 Fungi Zopfiella lundqvistii van Geel et al. (1986) van Geel et al. (2006a)
HdV 502 Fungi Bactrodesmium type van Geel et al. (1986)
HdV 503 Fungi Drechselera cf. state of
Cochliobolus specifer
van Geel et al. (1986)
HdV 504 Planta van Geel et al. (1986)
HdV 505 Planta Coniferae van Geel et al. (1986)
HdV 506 Unknown van Geel et al. (1986)
HdV 507 Unknown van Geel et al. (1986)
HdV 508 Planta van Geel et al. (1986)
HdV 509 Animalia Chironomids
Glyptotendipes gr. pallens
van Geel et al. (1986)
HdV 510 Planta Dicotyledons van Geel et al. (1986)
HdV 511 Fungi Ran (1990)
HdV 512 Fungi Ran (1990)
HdV 513 Unknown Ran (1990)
HdV 514 Unknown Ran (1990)
HdV 515 Unknown Ran (1990)
HdV 516 Unknown Ran (1990)
HdV 517 Unknown Ran (1990)
HdV 518 Unknown Ran (1990)
HdV 519 Unknown Ran (1990)
HdV 520 Unknown Ran (1990)
HdV 521 Unknown Ran (1990)
HdV 522 Unknown Ran (1990)
HdV 523 Unknown Ran (1990)
HdV 524 Unknown Ran (1990)
HdV 525 Unknown Ran (1990)
HdV 526 Unknown van Hoeve et al. eds (1998)
HdV 527 Fungi Enthorhiza sp. van Geel et al. (1983a)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 528 Fungi Gelasinospora cf. calospora van Geel et al. (1983a)
HdV 529 Fungi Puccinia cf. magnusiana van Geel et al. (1983a)
HdV 530 Protozoa Rhizopoda Centropyxis ecornis van Geel et al. (1983a)
HdV 531 Animalia Nematodes Trichuris spec. van Geel et al. (1983a)
HdV 532 Animalia Insecta Spaheroceridae van Geel et al. (1983a)
HdV 545 Unknown van der Harst (1984)
HdV 546 Fungi Trichodelitschia van Hofwegen (1983) Marinova et al. (2006);
Cugny et al. (2010)
HdV 547 Fungi van Hofwegen (1983)
HdV 548 Unknown van Hofwegen (1983)
HdV 549 Fungi Looze (1989)
HdV 550 Fungi van der Molen (1988)
HdV 551 Fungi van der Molen (1988)
HdV 552 Fungi van der Molen (1988) Willemsen et al. (1996)
HdV 553 Unknown van der Molen (1988)
HdV 554 Unknown Dam et al. (1988)
HdV 555 Animalia Insecta Trichoptera Dam et al. (1988)
HdV 560 Animalia? Spaargaren (1988)
HdV 561 Unknown Spaargaren (1988)
HdV 562 Animalia? Spaargaren (1988)
HdV 563 Unknown Spaargaren (1988)
HdV 564 Fungi Broerse (1991)
HdV 565 Unknown Broerse (1991)
HdV 566 Fungi van Bennekom (1992)
HdV 567 Unknown de Vries (1992)
HdV 568 Fungi de Vries (1992)
HdV 569 Fungi TM 11 de Vries (1992) Cugny et al. (2010)
HdV 570 Unknown de Vries (1992)
HdV 571 Fungi Speranza et al. (2000)
HdV 572 Fungi cf. Endophragmia Speranza et al. (2000) Menozzi et al. (2010)
HdV 573 Fungi HdV-527 Slopsma (1997)
HdV 574 Fungi HdV-206 Slopsma (1997)
HdV 575 Fungi Bombardioidea type Bos et al. (2005)
HdV 580 Fungi Isthmospora spinosa van Geel et al. (2006b) Yeloff et al. (2007a)
HdV 581 Fungi Yeloff et al. (2007a)
HdV 582 Fungi Yeloff et al. (2007a)
HdV 600 Algae Aphanizomenon cf. gracile Ralska-Jasiewiczova et al.
(1992a)
van Geel et al. (1994);
van Geel et al. (1996b)
HdV 601 Algae Anabaena sp. van Geel et al. (1994)
HdV 700 Protozoa Protozoa Foraminiferae Bakker et al. (1982) van Geel et al. (1983a);
Kholeif (2010)
HdV 701 Unknown Bakker et al. (1982)
HdV 702 Unknown Bakker et al. (1982)
HdV 703 A Algae Dinophyceae Bakker et al. (1982)
HdV 703 B Algae Dinophyceae Bakker et al. (1982)
HdV 703 C Algae Dinophyceae Bakker et al. (1982)
HdV 704 A Algae Operculodinum cf. centrocarpum Bakker et al. (1982)
HdV 704 B Algae Bakker et al. (1982)
HdV 704 C Algae cf. Hystrichosphaera Bakker et al. (1982)
HdV 704 Algae “Histrix”Bakker et al. (1982)
HdV 705 Unknown Bakker et al. (1982)
HdV 706 Fungi Bakker et al. (1982)
HdV 707 Fungi Culcitalna achraspora Bakker et al. (1982)
HdV 708 Unknown Bakker et al. (1982)
HdV 709 Unknown Bakker et al. (1982)
HdV 710 Algae Bakker et al. (1982)
HdV 711 Algae Mougeotia sp. Bakker et al. (1982)
HdV 712 Unknown HdV-38 Bakker et al. (1982)
HdV 713 Unknown Bakker et al. (1982)
HdV 714 Unknown Bakker et al. (1982)
HdV 715 Unknown Bakker et al. (1982)
HdV 716 Unknown Bakker et al. (1982)
HdV 717 Unknown Bakker et al. (1982)
HdV 718 Unknown Bakker et al. (1982)
HdV 719 A Unknown Bakker et al. (1982)
HdV 719 B Unknown Bakker et al. (1982)
HdV 720 Unknown Bakker et al. (1982)
HdV 721 Unknown Bakker et al. (1982)
HdV 722 Algae Bakker et al. (1982)
HdV 723 Unknown Bakker et al. (1982)
HdV 724 Unknown Bakker et al. (1982)
HdV 725 Unknown Bakker et al. (1982)
HdV 726 Unknown Bakker et al. (1982)
HdV 727 Unknown Bakker et al. (1982)
HdV 728 Unknown Bakker et al. (1982)
HdV 729 Fungi Bakker et al. (1982) van Geel et al. (1983b)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 730 Fungi Zopfiaspec. Bakker et al. (1982)
HdV 731 Unknown Bakker et al. (1982) van Smeerdijk (1989)
HdV 732 Unknown Bakker et al. (1982)
HdV 733 Unknown Bakker et al. (1982)
HdV 734 Unknown Bakker et al. (1982)
HdV 735 Unknown Bakker et al. (1982)
HdV 736 Animalia Bryozoa Bakker et al. (1982)
HdV 737 Animalia? Plumatella cf. fungosa Bakker et al. (1982)
HdV 740 Animalia Nanhermannia nanus Bakker et al. (1982)
HdV 741 Unknown Bakker et al. (1981)
HdV 742 Animalia Bakker et al. (1981)
HdV 743 Unknown Bakker et al. (1981)
HdV 744 Animalia Bakker et al. (1981)
HdV 745 Unknown Bakker et al. (1981)
HdV 748 Animalia? Bakker et al. (1981)
HdV 749 Fungi cf. Zopfia nicotia Bakker et al. (1981)
HdV 750 Unknown Bakker et al. (1981)
HdV 751 Unknown Bakker et al. (1981)
HdV 752 Unknown Bakker et al. (1981)
HdV 753 Unknown Bakker et al. (1981)
HdV 754 Unknown Bakker et al. (1981)
HdV 755 Unknown Bakker et al. (1981)
HdV 756 Unknown Bakker et al. (1981)
HdV 757 Unknown Bakker et al. (1981)
HdV 758 Unknown Bakker et al. (1981)
HdV 759 Unknown Bakker et al. (1981)
HdV 760 Algae Pediastrum Bakker et al. (1981)
HdV 764 Algae Chara spec. glomerus Bakker et al. (1981)
HdV 765 Animalia Daphnia sp. ephippium Bakker et al. (1981)
HdV 766 Algae Botryococcus spec. Bakker et al. (1981) Medeanic et al. (2008);
Herrmann et al. (2010);
Miola et al. (2010)
HdV 767 Planta Lemna spec. seed Bakker et al. (1981)
HdV 768 Planta Bakker et al. (1981)
HdV 770 Algae Scenedesmus cf. quadricauda Bakker et al. (1981)
HdV 771 Algae Bakker et al. (1981)
HdV 772 Algae Mougeotia Bakker et al. (1981)
HdV 773 Algae Spirogyra spec. Bakker et al. (1981)
HdV 774 Planta Myrica gale Bakker et al. (1981)
HdV 775 Planta Bakker et al. (1981)
HdV 776 B Planta Betula Bakker et al. (1981)
HdV 776 C Planta Betula Bakker et al. (1981)
HdV 776 Planta Betula Bakker et al. (1981)
HdV 777 Planta Betula Bakker et al. (1981)
HdV 778 Fungi? Bakker et al. (1981)
HdV 779 Planta Betula periderm Bakker et al. (1981)
HdV 780 Planta Bakker et al. (1981)
HdV 781 Planta Betula? Bakker et al. (1981)
HdV 782 Planta Bakker et al. (1981)
HdV 783 Planta Bakker et al. (1981)
HdV 784 Planta Bakker et al. (1981)
HdV 785 Planta Bakker et al. (1981)
HdV 786 Planta Bakker et al. (1981)
HdV 787 Planta Bakker et al. (1981)
HdV 788 Planta Empetrum nigrum Bakker et al. (1981)
HdV 789 Planta Ericaceae? Bakker et al. (1981)
HdV 790 Planta Bakker et al. (1981)
HdV 791 Planta Bakker et al. (1981)
HdV 792 Planta Bakker et al. (1981)
HdV 793 Unknown Bakker et al. (1981)
HdV 794 Unknown Bakker et al. (1981)
HdV 795 Planta Bakker et al. (1981)
HdV 796 Planta Bakker et al. (1981)
HdV 797 Planta Bakker et al. (1981)
HdV 798 Planta Bakker et al. (1981)
HdV 799 Unknown Bakker et al. (1981)
HdV 800 Planta Triglochin maritima Bakker et al. (1981)
HdV 804 Planta Campanula spec. seed Bakker et al. (1981)
HdV 805 Planta Campanula spec. seed Bakker et al. (1981)
HdV 806 Bryophyta Bakker et al. (1981)
HdV 807 Planta Bakker et al. (1981)
HdV 808 Pteridophyta Lycopodium inundatum Bakker et al. (1981)
HdV 809 Planta Nymphaea Bakker et al. (1981)
HdV 810 Fungi Mauquoy et al. (2004)
HdV 811 Fungi? Mauquoy et al. (2004)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 812 Fungi Mauquoy et al. (2004)
HdV 813 Fungi Mauquoy et al. (2004)
HdV 814 Fungi Mauquoy et al. (2004)
HdV 815 Fungi Yeloff et al. (2007b)
HdV 816 Fungi Yeloff et al. (2007b)
HdV 817 Fungi Yeloff et al. (2007b)
HdV 818 Planta Yeloff et al. (2007b)
HdV 819 Unknown Yeloff et al. (2007b)
HdV 820 Animalia Tardigrada? Yeloff et al. (2007b)
HdV 821 Unknown Yeloff et al. (2007b)
HdV 823 Fungi Yeloff et al. (2007b)
HdV 900 Algae Pediastrum Kuhry (1997)
HdV 901 Algae Botryococcus Kuhry (1997)
HdV 902 Animalia Tardigrada Garneau (1993)
HdV 903 Animalia Scolecodonte Garneau (1993)
HdV 904 Animalia Hydrozoa Garneau (1993)
HdV 905 Animalia Insecta Trichoptera Garneau (1993)
HdV 906 Animalia Cladocera Garneau (1993)
HdV 907 A Animalia Insecta Chironomidae larvae Garneau (1993)
HdV 907 B Animalia Insecta Chironomidae pupae Garneau (1993)
HdV 908 Animalia Coleoptera 3 Garneau (1993)
HdV 909 Animalia Insecta Hemiptera Garneau (1993)
HdV 910 Animalia Insecta Diptera 1 Garneau (1993)
HdV 911 Animalia Insecta Ephemeroptera Garneau (1993)
HdV 912 Animalia Insecta Lepidoptera Garneau (1993)
HdV 913 Animalia Insecta Hymenoptera Garneau (1993)
HdV 914 Animalia Insecta Homoptera Garneau (1993)
HdV 915 Animalia Insecta Pseudoscorpiones Garneau (1993)
HdV 916 Fungi Garneau (1993)
HdV 917 Fungi Garneau (1993)
HdV 918 Fungi Garneau (1993)
HdV 919 Fungi Garneau (1993)
HdV 920 Fungi Garneau (1993)
HdV 921 Fungi Garneau (1993)
HdV 922 Unknown Garneau (1993)
HdV 923 Fungi? Garneau (1993)
HdV 924 Fungi Garneau (1993)
HdV 925 Fungi Garneau (1993)
HdV 926 Fungi Garneau (1993)
HdV 927 Fungi Garneau (1993)
HdV 928 Fungi Garneau (1993)
HdV 929 Fungi Garneau (1993)
HdV 930 Fungi Garneau (1993)
HdV 931 Fungi Garneau (1993)
HdV 932 Fungi Garneau (1993)
HdV 933 Fungi Garneau (1993)
HdV 934 Fungi Garneau (1993)
HdV 935 Fungi Garneau (1993)
HdV 936 Unknown Garneau (1993)
HdV 937 Fungi Garneau (1993)
HdV 938 Fungi Sordariaceae Garneau (1993)
HdV 939 Fungi Garneau (1993)
HdV 940 Fungi Garneau (1993)
HdV 941 Unknown Garneau (1993)
HdV 942 A Unknown Garneau (1993)
HdV 942 B Unknown Garneau (1993)
HdV 942 C Unknown Garneau (1993)
HdV 942 D Unknown Garneau (1993)
HdV 943 Fungi Garneau (1993)
HdV 944 Fungi Garneau (1993)
HdV 945 Fungi Garneau (1993)
HdV 946 Fungi Garneau (1993)
HdV 947 Fungi Sordariaceae Garneau (1993)
HdV 948 Fungi Garneau (1993)
HdV 949 Fungi Garneau (1993)
HdV 950 Unknown Garneau (1993)
HdV 951 Unknown Garneau (1993)
HdV 953 Fungi Sordariaceae Garneau (1993)
HdV 954 Unknown Garneau (1993) Kuhry (1997)
HdV 955 Fungi Garneau (1993) Kuhry (1997)
HdV 956 Unknown Garneau (1993)
HdV 957 Fungi Garneau (1993)
HdV 958 Unknown Garneau (1993)
HdV 959 Fungi Garneau (1993)
HdV 960 Fungi Garneau (1993)
HdV 961 Fungi Garneau (1993)
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Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 962 Fungi Garneau (1993)
HdV 963 Unknown Garneau (1993)
HdV 964 Algae Rivulariaceae Garneau (1993)
HdV 965 Unknown Garneau (1993)
HdV 966 Acritarchs Garneau (1993)
HdV 967 Unknown Garneau (1993)
HdV 968 Unknown HdV-726 Garneau (1993)
HdV 969 Fungi Garneau (1993)
HdV 970 Fungi Garneau (1993)
HdV 971 Fungi Garneau (1993)
HdV 972 Fungi Garneau (1993)
HdV 973 Fungi Garneau (1993)
HdV 974 Fungi Garneau (1993)
HdV 975 Fungi Sordariaceae HdV-112B Garneau (1993)
HdV 976 Unknown HdV-184 Garneau (1993)
HdV 977 Fungi Garneau (1993)
HdV 978 Fungi Garneau (1993)
HdV 979 Fungi Garneau (1993)
HdV 980 Unknown HdV-128B Garneau (1993)
HdV 981 Fungi Diaporthaceae–Gnomonia
dryadis/Herpotrichia diffusa/
Leptosphaeria discors similar
Carrion et al. (1999)
HdV 982 Algae Oedogonium (O. pseudo‐cleveanum/
O. sphaerocephalum/O. ibadanense/
O. subspirale/O. henriquesii similar)
Carrion et al. (1999)
HdV 983 Algae Desmidiaceae (Staurastrum, e.g. S.
polytrichum,S. teliferum,S. asperum,
S. hirsutum,S. pilosum)
HdV-181 Carrion et al. (1999) Carrion et al. (2002)
HdV 984 Algae Euastrum ansatum or oblongum similar Carrion et al. (1999) Carrion et al. (2002)
HdV 985 Algae Euastrum bidentatum? Carrion et al. (1999) Carrion et al. (2002);
Menozzi et al. (2010)
HdV 986 Unknown Carrion et al. (1999)
HdV 987 Algae? Carrion et al. (1999)
HdV 988 Unknown Carrion et al. (1999)
HdV 989 Unknown HdV-984 Carrion et al. (1999)
HdV 990 Algae cf. Euastrum Carrion et al. (1999)
HdV 1001 Fungi Caryospora sp. van Geel et al. (2011)
HdV 1002 Fungi Sporoschisma spp. Gelorini et al. (2011)
HdV 1004 Fungi van Geel et al. (2011)
HdV 1005 Fungi Brachydesmiella sp. van Geel et al. (2011)
HdV 1006 Fungi van Geel et al. (2011)
HdV 1007 Fungi van Geel et al. (2011)
HdV 1008 Fungi cf. Valsaria sp. van Geel et al. (2011)
HdV 1009 Fungi Chaetomium sp. van Geel et al. (2011)
HdV 1010 Fungi van Geel et al. (2011)
HdV 1011 Fungi Epicoccum purpurascens van Geel et al. (2011)
HdV 1012 Fungi cf. Sordaria sp. van Geel et al. (2011)
HdV 1013 Fungi Cercophora-type van Geel et al. (2011)
HdV 1014 Fungi Rosellinia type van Geel et al. (2011)
HdV 1015 A Fungi Dictyoarthrinium cf. sacchari van Geel et al. (2011)
HdV 1015 B Fungi Dictyoarthrinium cf. sacchari van Geel et al. (2011)
HdV 1015 C Fungi Dictyoarthrinium cf. sacchari van Geel et al. (2011)
HdV 1018 A Fungi Spegazzinia tessarthra van Geel et al. (2011)
HdV 1018 B Fungi Spegazzinia tessarthra van Geel et al. (2011)
HdV 1020 Fungi van Geel et al. (2011)
HdV 1022 Fungi Clasterosporium sp. Gelorini et al. (2011)
HdV 1023 Fungi Spegazzinia intermedia van Geel et al. (2011)
HdV 1024 Fungi Brachysporium cf. pulchrum van Geel et al. (2011)
HdV 1025 Fungi Diplocladiella cf. scalaroides van Geel et al. (2011)
HdV 1026 Fungi Peziza/Scutellinia van Geel et al. (2011)
HdV 1027 Fungi Munkovalsaria donacina van Geel et al. (2011)
HdV 1028 Fungi van Geel et al. (2011)
HdV 1029 A Fungi Curvularia cf. intermedia Gelorini et al. (2011)
HdV 1029 A Fungi Curvularia spp. van Geel et al. (2011)
HdV 1029 B Fungi Curvularia spp. van Geel et al. (2011)
HdV 1030 Fungi cf. Byssothecium sp. van Geel et al. (2011)
HdV 1031 Fungi Acrodictys sp. van Geel et al. (2011)
HdV 1032 Fungi van Geel et al. (2011)
HdV 1033 Fungi van Geel et al. (2011)
HdV 1034 Fungi cf. Alternaria sp. van Geel et al. (2011)
HdV 1035 A Fungi Paraphaeosphaeria cf. michotii van Geel et al. (2011)
HdV 1035 B Fungi Paraphaeosphaeria cf. michotii van Geel et al. (2011)
HdV 1036 Fungi cf. Brachysporium sp. van Geel et al. (2011)
HdV 1037 Fungi cf. Helminthosporium sp. van Geel et al. (2011)
HdV 1038 Fungi Arecophila sp. van Geel et al. (2011)
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3.7. Similar
In this column, similar NPP types are indicated as reported in the
literature.
3.8. First publication/other publications
In the first column, the citation of the paper in which the NPP type
has been described is indicated. Further descriptions and/or illustra-
tions of the NPP types can be found in the literature cited in the follow-
ing column.
A partial NPP type list is presented in Table 3, where the NPP types
described by van Geel and colleagues have been selected. They have
been chosen among all NPPs because they are the most frequently cited.
4. Final remarks
On the basis of the literature that the author could review, the num-
ber of NPP types is more than 1300 (1314). They have been described in
about 100 papers. More than 600 types (653) have been identified on
some taxonomical level, e.g., EMA-56 Alysidium resinae, HdV-990 cf.
Euastrum,HdV-1031Acrodictys sp., A-12 Invertebrata?. Four hundred
forty-six have only been identified at the level of the wide categories
used in this review, e.g., HdV-120 Fungi, some of them dubiously,
e.g., EMA-52 Algae?. The collaboration of specialists could increase the
knowledge regarding these types. Two hundred fifteen remain of
completely unknown origin. Nevertheless, at least 16 of them have
been described and illustrated two or three times and recorded many
times after their first publication, e.g., HdV-128. It is possible that
many of the unidentified NPP types are peculiar to the site of the
study and/or are rare, and probably for this reason, they are not yet
identified by the NPP research community. They probably do not
deserve our attention, and their description and publication should be
discouraged in the future.
A certain number of NPPs (121 included in the 653 identified)
have been identified at a certain taxonomical level when they were
described and published, but a number (Type No. or or other-No.)
were not assigned to them. Some of them probably have a
corresponding NPP type among those named using the “HdV-No. Sys-
tem,”for instance, zygospores of Mougeotia (van Geel, 1976; van Geel
and van der Hammen, 1978; van Geel and Grenfell, 1996; Medeanic
et al., 2008; Chambers et al., 2011) have also been identified as
HdV-B, -133,-134, -135, -136, -141, -313, -313A-F, -418, -711, -772
and as Type 414 (van Geel, 1976; Pals et al., 1980; van Geel et al., 1981;
Bakker and van Smeerdijk, 1982; van der Wiel, 1982; Kuhry, 1997). A
possible correspondence exists also between the HdV-353A-D cocoons
(vanGeeletal.,1981) and the oocytes of Neorhabdocoela described by
Haas (1996).
The information concerning taxonomical identification and type
description has been reported from the literature. Few attempts
have been made to harmonize vocabulary, e.g., “one septum”,
“uni-septate”,“mono-septate”become 1-septate, in order to avoid
the introduction of misunderstandings or subjective interpretations.
No attempts have been made to harmonize NPP nomenclature be-
cause that was not the aim of this paper.
An effort should be made to coordinate the description and publi-
cation of new NPPs. A central NPP recording place could be useful in
selecting the publication of new types of NPPs, and avoiding the pro-
liferation of rare types and the attribution of different names to the
same type of palynomorphs. The harmonization of the NPP nomen-
clature, the definition of common standards for the description, mea-
surement, and digital images of NPPs by a central organization, could
improve the quality of NPP analysis, and as a consequence, it could in-
crease the number of identified NPP types. A good collection of im-
ages linked to the reference literature, for example, could avoid the
replication of names in the future. All the above-mentioned activities
begin with the knowledge of an update list of the NPP types, and this
paper is to be a first step in this.
Table 3 (continued)
Acr. N. L. Category Taxonomical identification Similar First publication Other publications
HdV 1039 Unknown van Geel et al. (2011)
HdV 1040 Fungi Isthmospora spinosa van Geel et al. (2011)
HdV 1041 Fungi Spegazzinia deightonii van Geel et al. (2011)
HdV 1042 Fungi Montagnula sp. van Geel et al. (2011)
HdV 1043 Fungi cf. Lasiodiplodia theobromae Gelorini et al. (2011)
HdV 1044 Fungi van Geel et al. (2011)
HdV 1045 Fungi van Geel et al. (2011)
HdV 1046 Fungi van Geel et al. (2011)
HdV 1047 Fungi Rhytidospora cf. tetraspora van Geel et al. (2011)
HdV 1048 Fungi van Geel et al. (2011)
HdV 1049 Fungi cf. Mitteriella ziziphina van Geel et al. (2011)
HdV 1051 Fungi van Geel et al. (2011)
HdV 1052 Fungi Xylariaceae van Geel et al. (2011) Gelorini et al. (2011)
HdV 1053 Fungi Dictyosporium cf. heptasporum van Geel et al. (2011)
HdV 1054 Fungi Rosellinia cf. valdiviensis van Geel et al. (2011)
HdV 1055 Fungi cf. Ustilago sp. van Geel et al. (2011)
HdV 1056 Fungi van Geel et al. (2011)
HdV 1057 Fungi cf. Anthostomella vestita van Geel et al. (2011)
HdV 1058 A Fungi Gelorini et al. (2011)
HdV 1058 Fungi van Geel et al. (2011)
HdV 1059 Fungi van Geel et al. (2011)
HdV 1061 Fungi van Geel et al. (2011)
HdV 1062 Animalia van Geel et al. (2011)
HdV 1064 Unknown van Geel et al. (2011)
HdV 1093 Fungi Gelasinospora cf. cratophora van Geel et al. (2011)
HdV 1103 Fungi Glomus sp. van Geel et al. (2011)
HdV 1245 Fungi Diporotheca sp. van Geel et al. (2011)
HdV 1351 Fungi Gelasinospora cf. dictyophora van Geel et al. (2011)
HdV 1400 Unknown van Geel et al. (2010)
HdV 1401 Unknown van Geel et al. (2010)
HdV 1402 Unknown HdV-91
HdV-819
van Geel et al. (2010)
160 A. Miola / Review of Palaeobotany and Palynology 186 (2012) 142–161

Author's personal copy
Supplementary materials related to this article can be found on-
line at http://dx.doi.org/10.1016/j.revpalbo.2012.06.010.
Acknowledgments
This work was supported by the PICAR project (PRIN 2008FJCEF4,
Scient. coord. A. M. Mercuri). The author would like to thank the
anonymousreviewers, L. Kooistra and J.N. Haas for their useful remarks
and suggestions, and Barbara Gaudioso and Bruna I. Menozzi for their
valuable support.
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