Paleoflora of the Messel Formation

This is an overview of the {{Linktext|paleoflora}} of the Eocene Messel Formation as explored by the Messel Pit excavations in Germany. A former quarry and now UNESCO World Heritage Site, the Messel Formation preserves what lived in and around a meromictic lake surrounded by a paratropical rainforest during the latest early to earliest middle Eocene, approximately 47 Ma.{{cite journal |last1=Mertz |first1=D. |last2=Renne |first2=P. |year=2005 |title=A numerical age for the Messel fossil deposit (UNESCO World Heritage Site) derived from ⁴⁰Ar/³⁹Ar dating on a basaltic rock fragment |journal=Courier Forschungsinstitut Senckenberg |volume=255 |pages=7–75}} A complete list of plant taxa was published in 2024.{{cite journal |last1=Smith |first1=Krister T. |last2=Collinson |first2=Margaret |last3=Folie |first3=Annelise |last4=Haberesetzer |first4=Jörg |last5=Hennicke |first5=Florian |last6=Kothe |first6=Erika |last7=Lehmann |first7=Thomas |last8=Lenz |first8=Olaf K. |last9=Mayr |first9=Gerald |last10=Micklich |first10=Norbert |last11=Rabenstein |first11=Renate |last12=Racicot |first12=Rachel |last13=Schaal |first13=Stephan F. K. |last14=Smith |first14=Thierry |last15=Tosal |first15=Aixa |last16=Uhl |first16=Dieter |last17=Wappler |first17=Torsten |last18=Wedmann |first18=Sonja |last19=Wuttke |first19=Michael |title=The biodiversity of the Eocene Messel Pit |journal=Palaeobiodiversity and Palaeoenvironments |date=2024 |volume=104 |doi=10.1007/s12549-024-00633-2 |url=https://link.springer.com/article/10.1007/s12549-024-00633-2|doi-access=free }}

Several major monographs on the paleoflora have been published since the early 1900s, with {{ill|Hermann Engelhardt|de|Hermann Engelhardt (Paläontologe)}} producing the first leaf monograph in 1922, followed by Heidemarie Thiele-Pfeiffer in 1988 who provided the first in-depth palynological revision and expansion. Margaret Collinson, Steven Manchester, and Volker Wilde collaborated to study and redescribe the flowers, fruits, seeds, and other reproductive organs of the formation, with the monograph being published in 2011. Most recently, Johannes Bouchal, Christian Geier, Friđgeir Grímsson and colleagues have completed monographic studies of the palynomorphs, combining traditional light microscopy with scanning electron microscopy to achieve greater phylogenetic precision.{{cite journal |last1=Bouchal |first1=Johannes M. |last2=Geier |first2=Christian |last3=Ulrich |first3=Silvia |last4=Wilde |first4=Volker |last5=Lenz |first5=Olaf K. |last6=Zetter |first6=Richard |last7=Grímsson |first7=Friđgeir |title=Qualitative LM and SEM study of the Messel palynoflora: part I. Algae to Vitales |journal=Grana |date=2024 |volume=63 |issue=3 |pages=193-246 |doi=10.1080/00173134.2024.2375209 |url=https://www.tandfonline.com/doi/full/10.1080/00173134.2024.2375209|doi-access=free }}

Dinoflagellates

Only a single dinoflagellate taxon has been described formally from the Messel formation. Based on the predominance in layers where heavy nearshore runoff or slippage is present, its suggested Messelodinium thielepfeifferae was likely an inhabitant of warm nearshore waters in the adult stage or wind and wave action resulted in large accumulation of the cysts. M. thielepfeifferae populations are most prevalent in the older layers of the lake, when it was suggested to be holomictic and become absent in the youngest layers when green algae become dominant with the lake shifting to meromictic conditions and the water chemistry changed.{{cite journal |last1=Lenz . |first1=O. |last2=Wilde |first2=V. |last3=Riegel |first3=W. |last4=Heinrichs |first4=T. |title=Distribution and paleoecologic significance of the freshwater dinoflagellate cyst Messelodinium thielepfeifferae gen. et sp. nov from the Middle Eocene of Lake Messel, Germany |journal=Palynology |date=2007 |volume=31 |issue=1 |pages=119–134 |doi=10.2113/gspalynol.31.1.119}}

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Family ! Genus ! Species ! Authority ! Notes ! Image
Peridiniaceae \| †Messelodinium \| †Messelodinium thielepfeifferae \| Lenz et al., 2007 \| A peridinialean dinoflagellate. \|

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Family

! Genus

! Species

! Authority

! Notes

! Image

Peridiniaceae

†Messelodinium

†Messelodinium thielepfeifferae

Lenz et al., 2007

A peridinialean dinoflagellate.

Charophyta

Olaf Lenz et al (2011) briefly noted the presence of the Zygnemataceae fossil genus Ovoidites, but only gave general population densities with no enumeration on species information.

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Family ! Genus ! Species ! Authority ! Notes ! Image
Zygnemataceae \| †Ovoidites \| Undescribed{{cite journal \|last1=Lenz \|first1=O. \|last2=Wilde \|first2=V. \|last3=Riegel \|first3=W. \|year=2011 \|title=Short-term fluctuations in vegetation and phytoplankton during the Middle Eocene greenhouse climate: a 640-kyr record from the Messel oil shale (Germany) \|journal=International Journal of Earth Sciences \|volume=100 \|issue=8 \|pages=1851–1874\|doi=10.1007/s00531-010-0609-z \|bibcode=2011IJEaS.100.1851L \|s2cid=52968593 \|doi-access=free \|url=https://e-docs.geo-leo.de/bitstream/11858/6785/2/531_2010_Article_609.pdf }} \| \| A minor Charophyte green algae component of the Messel Formation. \|

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Family

! Genus

! Species

! Authority

! Notes

! Image

Zygnemataceae

†Ovoidites

Undescribed{{cite journal |last1=Lenz |first1=O. |last2=Wilde |first2=V. |last3=Riegel |first3=W. |year=2011 |title=Short-term fluctuations in vegetation and phytoplankton during the Middle Eocene greenhouse climate: a 640-kyr record from the Messel oil shale (Germany) |journal=International Journal of Earth Sciences |volume=100 |issue=8 |pages=1851–1874|doi=10.1007/s00531-010-0609-z |bibcode=2011IJEaS.100.1851L |s2cid=52968593 |doi-access=free |url=https://e-docs.geo-leo.de/bitstream/11858/6785/2/531_2010_Article_609.pdf }}

A minor Charophyte green algae component of the Messel Formation.

Chlorophyta

Several chlorophyte green algae are identified from the Messel Formation, with the greatest concentrations being found in the youngest layers. In the older layers Tetraedron minimum is a distinct component of the yearly varves in the lake, with large blooms occurring to form the lighter summer portions of the varve couplets. In the younger layers, Botryococcus species become the most prolific, having taken over as a major phytoplankton taxon from T. minimum and the dinoflagellate Messelodinium thielepfeifferae.

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Family ! Genus ! Species ! Authority ! Notes ! Image
Botryococcaceae \| Botryococcus \| Botryococcus braunii{{cite journal \|last1=Adam \|first1=P. \|last2=Schaeffer \|first2=P. \|last3=Albrecht \|first3=P. \|year=2006 \|title=C₄₀ monoaromatic lycopane derivatives as indicators of the contribution of the alga Botryococcus braunii race L to the organic matter of Messel oil shale (Eocene, Germany) \|journal=Organic Geochemistry \|volume=37 \|issue=5 \|pages=584–596\|doi=10.1016/j.orggeochem.2006.01.001 }} \| Kützing \| A chlorophyte green algae. \|
Peridiniaceae \| Tetraedron \| Tetraedron minimum \| (A. Braun) Hansgirg \| A chlorophyte green algae The main component in the spring/summer layers of the Messel varves. \|

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Family

! Genus

! Species

! Authority

! Notes

! Image

Botryococcaceae

Botryococcus

Botryococcus braunii{{cite journal |last1=Adam |first1=P. |last2=Schaeffer |first2=P. |last3=Albrecht |first3=P. |year=2006 |title=C₄₀ monoaromatic lycopane derivatives as indicators of the contribution of the alga Botryococcus braunii race L to the organic matter of Messel oil shale (Eocene, Germany) |journal=Organic Geochemistry |volume=37 |issue=5 |pages=584–596|doi=10.1016/j.orggeochem.2006.01.001 }}

Kützing

A chlorophyte green algae.

Peridiniaceae

Tetraedron

Tetraedron minimum

(A. Braun) Hansgirg

A chlorophyte green algae
The main component in the spring/summer layers of the Messel varves.

Pteridophytes

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Family ! Genus ! Species ! Authority ! Notes ! Image
Aspleniaceae \| Asplenium \| Asplenium sp.{{cite journal \|last1=Engelhardt \|first1=H. \|year=1922 \|title=Die alttertiäre Flora von Messel bei Darmstadt \|journal=Abhandlungen der Hessischen Geologischen Landesanstalt zu Darmstadt \|volume=7 \|issue=4 \|pages=20–128 \|url=https://www.biodiversitylibrary.org/bibliography/7648}} \| \| Fern pinnuels attributed to Asplenium Not identified to species \|

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Family

! Genus

! Species

! Authority

! Notes

! Image

Aspleniaceae

Asplenium

Asplenium sp.{{cite journal |last1=Engelhardt |first1=H. |year=1922 |title=Die alttertiäre Flora von Messel bei Darmstadt |journal=Abhandlungen der Hessischen Geologischen Landesanstalt zu Darmstadt |volume=7 |issue=4 |pages=20–128 |url=https://www.biodiversitylibrary.org/bibliography/7648}}

Fern pinnuels attributed to Asplenium
Not identified to species

Conifers

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Family ! Genus ! Species ! Authority ! Notes ! Image
Doliostrobaceae \| †Doliostrobus \| †Doliostrobus taxiformis{{cite journal \|last1=Collinson \|first1=M. \|last2=Manchester \|first2=S. \|last3=Wilde \|first3=V. \|year=2012 \|title=Fossil fruits and seeds of the Middle Eocene Messel biota, Germany. \|journal=Abhandlungen der Senckenberg Gesellschaft für Naturforschung \|volume=570 \|pages=1–249 \|isbn=978-3-510-61400-4 \|url=}} \| (Sternberg) Z. Kvaček emend. Kunzmann \| A conifer of possible cupressaceous affinities \|
Podocarpaceae \| †Podozamites \| †Podozamites eocaenica \| Engelhardt \| Possible podocarpaceous foliage First suggested as a cyacad species \|
Taxaceae \| Cephalotaxus \| †Cephalotaxus messelensis{{cite book \|title=Fossilium Catalogus. II. Plantae. \|chapter=Pars 105. Gymnospermae (6) Ginkgophyta et Coniferae.\|last1=Dijkstra \|first1=S. \|last2=van Amerom \|first2=H. \|last3=Schultze-Motel \|first3=J. \|editor-last1=van Amerom \|editor-first1=H. \|publisher=Backhuys Publishers \|location=Leiden, Netherlands \|date=2002}} \| Wilde \| A plum yew species \|

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Family

! Genus

! Species

! Authority

! Notes

! Image

Doliostrobaceae

†Doliostrobus

†Doliostrobus taxiformis{{cite journal |last1=Collinson |first1=M. |last2=Manchester |first2=S. |last3=Wilde |first3=V. |year=2012 |title=Fossil fruits and seeds of the Middle Eocene Messel biota, Germany. |journal=Abhandlungen der Senckenberg Gesellschaft für Naturforschung |volume=570 |pages=1–249 |isbn=978-3-510-61400-4 |url=}}

(Sternberg) Z. Kvaček emend. Kunzmann

A conifer of possible cupressaceous affinities

Podocarpaceae

†Podozamites

†Podozamites eocaenica

Engelhardt

Possible podocarpaceous foliage
First suggested as a cyacad species

Taxaceae

Cephalotaxus

†Cephalotaxus messelensis{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 105. Gymnospermae (6) Ginkgophyta et Coniferae.|last1=Dijkstra |first1=S. |last2=van Amerom |first2=H. |last3=Schultze-Motel |first3=J. |editor-last1=van Amerom |editor-first1=H. |publisher=Backhuys Publishers |location=Leiden, Netherlands |date=2002}}

Wilde

A plum yew species

Basal angiosperms

=Nymphaeales=

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Family ! Genus ! Species ! Authority ! Notes ! Image
Nymphaeaceae \| †Nuphaea \| †Nuphaea engelhardtii{{cite journal \|last1=Gee \|first1=C. \|last2=Taylor \|first2=D. \|year=2019 \|title=An Extinct Transitional Leaf Genus of Nymphaeaceae from the Eocene Lake at Messel, Germany: Nuphaea engelhardtii Gee et David W. Taylor gen. et sp. nov. \|journal=International Journal of Plant Sciences \|volume=180 \|issue=7 \|pages=724–736 \|doi=10.1086/704376\|s2cid=201211546 }} \| Gee & Taylor, 2018 \| A transitional nymphaealean water lily genus \|
Incertae sedis \| Incertae sedis \| Incertae sedis \| \| Seeds of possible nymphaealean affinity Not identified to species

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Family

! Genus

! Species

! Authority

! Notes

! Image

Nymphaeaceae

†Nuphaea

†Nuphaea engelhardtii{{cite journal |last1=Gee |first1=C. |last2=Taylor |first2=D. |year=2019 |title=An Extinct Transitional Leaf Genus of Nymphaeaceae from the Eocene Lake at Messel, Germany: Nuphaea engelhardtii Gee et David W. Taylor gen. et sp. nov. |journal=International Journal of Plant Sciences |volume=180 |issue=7 |pages=724–736 |doi=10.1086/704376|s2cid=201211546 }}

Gee & Taylor, 2018

A transitional nymphaealean water lily genus

Incertae sedis

Seeds of possible nymphaealean affinity
Not identified to species

Magnoliids

=Laurales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=3 | Lauraceae

†Laurocarpum

†Laurocarpum sp.

Seeds of lauraceous affinity
3 unnamed morphospecies

†Laurophyllum

†Laurophyllum lanigeroides{{cite journal |last1=Wappler |first1=T. |last2=Tokuda |first2=M. |last3=Yukawa |first3=J. |last4=Wilde |first4=V. |year=2010 |title=Insect herbivores on Laurophyllum lanigeroides (Engelhardt 1922) Wilde: the role of a distinct plant-insect associational suite in host taxonomic assignment |journal=Palaeontographica, Palaeontographica, Abt.B: Palaeobotany–Palaeophytology |volume=283 |issue=4–6 |pages=137–155}}

(Engelhardt) Wilde

A lauraceous leaf morphotaxon
host to two forms of asphondyliine tribe midge galls

incertae sedis

Seeds of lauraceous affinity
not identified to genus/species

=Magnoliales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=2| Magnoliaceae

| rowspan=2| Magnolia

Magnolia sp.

A Magnolia species fruit
Not identified to species

Magnolia sp.

Magnolia species seeds
not identified to species

Myristicaceae

†Myristicacarpum

Myristicacarpum sp.

A Myristicacarpum species seed
Not identified to species

Monocots

=Alismatales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=3 | Araceae

| rowspan=2 | †Araciphyllites

†Araciphyllites tertiarius{{cite journal |last1=Wilde |first1=V. |last2=Kvaček |first2=Z. |last3=Bogner |first3=J. |title=Fossil leaves of the Araceae from the European Eocene and notes on other aroid fossils |journal=International Journal of Plant Sciences |date=2005 |volume=166 |issue=1 |pages=157–183 |doi=10.1086/425673|s2cid=85215526 }}

(Engelhardt) Wilde, Z. Kvaček & Bogner, 2005

An araceous leaf

†Araciphyllites schaarschmidtii

Wilde, Z. Kvaček & Bogner, 2005

An araceous leaf

†Caladiosoma

†Caladiosoma messelense

Wilde, Z. Kvaček & Bogner, 2005

An araceous leaf

=Arecales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=4 | Arecaceae

Arecocaryon

Arecocaryon messelensis{{Cite journal|last=Doweld |first=A. B. |year=2021 |title=Fossil Alloceltidoxylon, Allonymphaea, Arecocaryon, Paralnoxylon and Paranyssa and extant Komaroviopsis, Marcanodendron, and Papyrocactus (Magnoliophyta), new replacement generic names |journal=Phytotaxa |volume=524 |issue=2 |pages=92–98 |doi=10.11646/phytotaxa.524.2.3 |s2cid=243482734 }}

(Collinson, Manchester, & Wilde) Doweld

Palm species fruits
First described as Friedemannia messelensis, a genus jr homonym
Moved to Arecocaryon messelensis in 2021
The most abundant fruit at Messel
known from a full flower to mature fruit ontogenetic series

†Calamopsis

Cf. †Calamopsis bredana

A Palm leaf
Possibly affiliated with C. bredana

rowspan=2| Incertae sedis

incertae sedis

Pinnate palm leaves of possible lepidocaryoid tribal affinity.
Not identified to species

incertae sedis

Palmate palm leaves of uncertain tribal affinity.
Not identified to species

=Liliales=

Family ! Genus ! Species ! Authority ! Notes ! Image
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rowspan=4 \| Smilacaceae \| rowspan=4 \| Smilax \| †Smilax grandifolia \| Unger \| A greenbrier \|
†Smilax lingulata \| Heer \| A greenbrier \|
†Smilax ovata \| Wess. \| A greenbrier \|
†Smilax reticulata \| Heer \| A greenbrier \|

=Pandanales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Cyclanthaceae

Cyclanthus

Cyclanthus messelensis{{cite journal |last1=Smith |first1=S. |last2=Collinson |first2=M. |last3=Rudall |first3=P. |title=Fossil Cyclanthus (Cyclanthaceae, Pandanales) from the Eocene of Germany and England |journal=American Journal of Botany |date=2008 |volume=95 |issue=6 |pages=688–699 |doi=10.3732/ajb.2007390 |pmid=21632394}}

S. Y. Smith, Collinson & Rudall, 2008

A Cyclanthus species
Described from fruits and seeds

=Poales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Cyperaceae

†Volkeria

†Volkeria messelensis{{cite journal |last1=Smith |first1=S. |last2=Collinson |first2=M. |last3=Simpson |first3=D. |last4=Rudall |first4=P. |last5=Marone |first5=F. |last6=Stampanoni |first6=M. |year=2009 |title=Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe |journal=American Journal of Botany |volume=96 |issue=8 |pages=1506–1518 |doi=10.3732/ajb.0800427|pmid=21628296 |url=https://www.zora.uzh.ch/id/eprint/27030/11/Collinson.pdf }}

Smith et al., 2009

A mapanioid sedge

Poaceae

Phragmites

†Phragmites oeningensis

A.Braun ex Heer

A grass foliage

Basal eudicots

=Proteales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Sabiaceae

Meliosma

Meliosma sp.

A Meliosma species seed
Not identified to species

=Ranunculales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=13 | Menispermaceae

Cocculus

†Cocculus lottii

Collinson, Manchester, & Wilde, 2012

A Cocculus species endocarp

Diploclisia

†Diploclisia rugulosa

Collinson, Manchester, & Wilde, 2012

A Diploclisia species seed

†Karinschmidtia

†Karinschmidtia rotulae

Collinson, Manchester, & Wilde, 2012

A menispermaceous endocarp

†Martinmuellera

†Martinmuellera tuberculata

Collinson, Manchester, & Wilde, 2012

A menispermaceous genus seed

rowspan=3 | †Palaeosinomenium

†Palaeosinomenium ornamentum

Collinson, Manchester, & Wilde, 2012

A menispermaceous endocarp

†Palaeosinomenium venablesii

Chandler, 1961

A menispermaceous endocarp
First described from the English Eocene (1961)

†Palaeosinomenium sp.

A menispermaceous endocarp
Not identified to species

Parabaena

†Cf. Parabaena europaea

A menispermaceous endocarp
Fossil similar to P. europaea
Not identified to species

?Pericampylus

?Pericampylus sp.

A tentative Pericampylus species endocarp.
not described to species

Stephania

†Stephania hootae

Collinson, Manchester, & Wilde, 2012

A Stephania species seed

†Tinomiscoidea

†Tinomiscoidea jacquesii

Collinson, Manchester, & Wilde, 2012

A menispermaceous endocarp

†Wardensheppeya

†Wardensheppeya sp.

A menispermaceous endocarp
Not identified to species

incertae sedis

menispermaceous endocarp of Tinosporeae subfamily affinity
3 morphotypes not identified to genus/species

Superasterids

=Campanulids=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Torricelliaceae

Toricellia

†Toricellia bonesii

(Manchester) Manchester

A Toricellia species.
Endocarp with germination valve

=Cornales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Alangiaceae

Alangium

Alangium sp.

An Alangium species likely belonging to Alangium sect. Marlea.
Not identified to species
Identified from single endocarp, others likely in Messel collections

Cornaceae

Cornus

†Cornus orbifera{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 23. Umbellifloraea: Cornaceae|author=Kirchheimer F. |editor=Jongmans, W. |publisher=Verlag Gustav Feller |location=Neubrandenburg |date=2 July 1938}}

Heer

A dogwood species

rowspan=7 | Nyssaceae

Cf. †Beckettia

incertae sedis

A germination valve similar to Beckettia
Not identified to species

†Eomastixia

Cf. Eomastixia rugosa

An endocarp similar to Eomastixia rugosa
Not identified to species

?Mastixia

incertae sedis

A possible Mastixia endocarp and mesocarp
Possibly included in a coprolite
Not identified to species

†Mastixiopsis

†Mastixiopsis nyssoides

Kirchheimer

A nyssaceous endocarp

rowspan=3 | Nyssa

†Nyssa disseminata

(R. Ludw.) Kirchheimer

A nyssaceous endocarp with germination valve

†Nyssa europaea

Unger

A nyssaceous species

†Nyssa ornithobroma

Unger

A nyssaceous species

=Ericales=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Pentaphylacaceae

Cleyera

incertae sedis

A Cleyera species.
Not identified to species

Symplocaceae

Symplocos

†Symplocos minutula{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 26. Symplocaceae.|author=Kirchheimer F.|editor=Jongmans, W. |publisher=Verlag Gustav Feller |date=14 Jul 1950}}

(Sternberg) Kirchheimer

A Symplocos species fruit.

Theaceae

†Camelliacarpoidea

†Camelliacarpoidea messelensis

Collinson, Manchester, & Wilde

A theaceous species.
A dry fruit with persistent calyx

=Lamiids=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Apocynaceae

†Cypselites

incertae sedis

A Cypselites species
Elongated seed with wind dispersal hairs.
Not identified to species, 11 specimens known

Bignoniaceae

†Darmstadtia

Darmstadtia biseriata

A catalpa species.
Pods with dry seeds lacking wind dispersal wings
Described from2 specimens

rowspan=12| Icacinaceae

| rowspan=2| †Palaeohosiea

†Palaeohosiea bilinica

(Ettingshausen) Kvaček & Bůžek

A phytocrenaceous species.
known from endocarps

incertae sedis

A Palaeohosiea endocarp.
Not identified to species

Cleyera

incertae sedis

Collinson, Manchester, & Wilde

A Cleyera species.
Not identified to species

rowspan=3| †Icacinicarya

Icacinicarya densipunctata

Collinson, Manchester, & Wilde

An icacinaceous species.

Icacinicarya tiffneyi

Collinson, Manchester, & Wilde

An icacinaceous species.
endocarp with a preserved layer of soft tissue

incertae sedis

An Icacinicarya endocarp.
Not identified to species

Cf. Natsiatum

incertae sedis

An endocarp similar to Natsiatum.
Not identified to species

†Palaeophytocrene

incertae sedis

A Palaeophytocrene endocarp.
Not identified to species

Phytocrene

†Phytocrene punctilinearis

Collinson, Manchester, & Wilde

A Phytocrene species.

rowspan=2| Pyrenacantha

"Species 1"

Collinson, Manchester, & Wilde

An unnamed Pyrenacantha species.
endocarp with a preserved layer of soft tissue

"Species 2"

Collinson, Manchester, & Wilde

An unnamed Pyrenacantha species.
endocarp without a preserved layer of soft tissue

Cf. Pyrenacantha

incertae sedis

Collinson, Manchester, & Wilde

An endocarp similar to Pyrenacantha.
not identified to species

Superrosids

=Fabids=

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Family

! Genus

! Species

! Authority

! Notes

! Image

Cannabaceae

Aphananthe

Cf. †Aphananthe tenuicostata

A hemp relative fruit.
Similar to Aphananthe tenuicostata

rowspan=8| Celastraceae

| rowspan=4| Celastrus

†Celastrus elaenus{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 29. Celastraceae.|last1=Wonnacott |first1=F. |editor-last1=Jongmans |editor-first1=W. |publisher=Uitgeverij Dr. W. Junk |location='s-Gravenhague |date=28 September 1955}}

Unger

A Celastrus species.

†Celastrus friedrichi

Wonnacott

A Celastrus species.

†Celastrus illicinoides

Engelhardt

A Celastrus species.

†Celastrus oxyphyllus

Unger

A Celastrus species.

rowspan=3| Elaeodendron

†Elaeodendron degener

(Unger) Ettingshausen

An Elaeodendron species.

†Elaeodendron dryadum

Ettingshausen

An Elaeodendron species.
Similar to the living E. orientale

†Elaeodendron obovatifolium

Engelhardt

An Elaeodendron species.

Euonymus

†Euonymus heerii

Ettingshausen

A Euonymus species.

Elaeocarpaceae

Sloanea

†Sloanea messelensis

Collinson, Manchester, & Wilde

A Sloanea species fruit.

Euphorbiaceae

†Euphorbiotheca

†Euphorbiotheca gothii

Collinson, Manchester, & Wilde

Trilocular euphorb fruits.

rowspan=3| Fabaceae

†Mimosites

†Mimosites spiegeli

Engelhardt

A caesalpinioid legume species pod.

rowspan=2| †Leguminocarpon

†Leguminocarpon herendeenii

Collinson, Manchester, & Wilde

A legume species pod.

†Leguminocarpon sp.

Legume species pods.
2 mophotypes noted
not identified to species

rowspan=3| Juglandaceae

†Cruciptera

†Cruciptera schaarschmidtii

Manchester, Collinson, & Goth

A juglandaceous winged fruit.

†Hooleya

†Hooleya sp.

A juglandaceous fruiting raceme.
Not identified to species.

†Palaeocarya

†Palaeocarya sp.

A juglandaceous fruit.
Not identified to species.

Rhamnaceae

Berchemia

†Berchemia mellerae

Collinson, Manchester, & Wilde

A Berchemia buckthorn species fruit.

Rosaceae

Prunus

†Prunus pereger{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 25. Rosaceae.|last1=Kirchheimer |first1=F. |editor-last1=Jongmans |editor-first1=W. |publisher=Verlag Gustav Feller |date=14 Jul 1942}}

Unger

A cherry relative

Ulmaceae

†Cedrelospermum

†Cedrelospermum leptospermum

(Ettingshausen) Manchester emend. Wilde & Manchester

An ulmaceous elm relative fruit.

=Malvids=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=9| Anacardiaceae

Anacardium

†Anacardium germanicum

Manchester, Wilde & Collinson

A cashew species.
The first European record of the genus
Known from at least 16 fruits and hypocarps

Lannea

†Lannea hessenensis

Collinson, Manchester, & Wilde

A Lannea species.
Fruits are unilocular, separating them from Lannea europaea
23 specimens in the type series

Mangifera

†Mangifera tertiaria{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 20. Anacardiaceae.|last1=Edwards |first1= W. |last2=Wonnacott |first2=F. |editor-last1=Jongmans |editor-first1=W. |publisher=W. Junk |location='s-Gravenhague |date=28 August 1935}}

Ettingshausen

A mango relative.

†Pentoperculum

†Pentoperculum minimus

(Reid & Chandler) Manchester

A spondiadeous cashew relative.
Rounded-pentangular fruits in transverse cross-section
Identified from 7 fossils

Pleiogynium

†Pleiogynium mitchellii

(Reid & Chandler) Manchester

A Pleiogynium species.
A larger spondiadeous fruit with 10 germination valves
Described from the holotype only

rowspan=3| Rhus

†Rhus meriani

Heer

A sumac relative.

†Rhus saportana

Pilar

A sumac relative.

†Rhus ternata

Engelhardt

A sumac relative.

Tapiscia

†Tapiscia pusilla

(Reid & Chandler) Mai

A Tapiscia species fruit.
Some fruits possibly preserved in coprolites

rowspan=2| Burseraceae

†Bursericarpum

†Bursericarpum species

A Bursericarpum species endocarp.
Not identified to species

Canarium

Canarium sp.

A Canarium species endocarp.
Not identified to species

rowspan=3| Malvaceae

| rowspan=3| †Byttneriopsis

†Byttneriopsis daphnogenes{{cite journal |last1=Kvacek |first1=Z. |last2=Wilde |first2=V. |year=2010 |title=Foliage and seeds of malvalean plants from the Eocene of Europe |journal=Bulletin of Geosciences |volume=85 |issue=1 |pages=163–182 |doi=10.3140/bull.geosci.1170|doi-access=free }}

(Ettingshausen) Kvaček & Wilde

A malvaceous leaf morphotaxon

†Byttneriopsis spiegelii

(Ettingshausen) Kvaček & Wilde

A malvaceous leaf morphotaxon

†Byttneriopsis steuerii

Kvaček & Wilde

A malvaceous leaf morphotaxon

rowspan=3| Rutaceae

†Rutaspermum

†Rutaspermum chandleri

Collinson & Gregor

A citrus family seed.

Toddalia

†Toddalia ovata{{cite journal |last1=Winkler |first1=I. |last2=Labandeira |first2=C. |last3=Wappler |first3=T. |last4=Wilf |first4=P. |year=2010 |title=Distinguishing Agromyzidae (Diptera) leaf mines in the fossil record: new taxa from the Paleogene of North America and Germany and their evolutionary implications |journal=Journal of Paleontology |volume=84 |issue=5 |pages=935–954 |doi=10.1666/09-163.1|s2cid=56378236 }}

Wilde

A Toddalia leaf morphospecies
Host to Phytomyzites schaarschmidti trace fossils -
leaf mines attributed to agromyzid flies

Cf. Toddalia

incertae sedis

A Seed similar to Toddalia.
Not identified to genus/species

rowspan=6| Sapindaceae

Dodonaea

†Dodonaea salicites{{cite book |title=Fossilium Catalogus. II. Plantae. |chapter=Pars 14. Sapindaceae.|last1=Edwards |first1=W. |last2=Wonnacott |first2=F. |editor-last1=Jongmans |editor-first1=W. |publisher=Verlag Gustav Feller |date=14 Jul 1950}}

Ettingshausen

A Dodonaea species

rowspan=4| Sapindus

†Sapindus cupanioides

Ettingshausen

A Sapindus species

†Sapindus dubius

Unger

A Sapindus species

†Sapindus firmifolius

Engelhardt

A Sapindus species

†Sapindus heliconius

Unger

A Sapindus species

Thouinia

†Thouinia occidentalis

Engelhardt

A Thouinia species

Simaroubaceae

Ailanthus

†Ailanthus confucii

Unger

An Ailanthus seed.

=Myrtales=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Lythraceae

Cf. Decodon

Incertae sedis

Seeds of similar morphology to Decodon Not identified to species

=Saxifragales=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Altingiaceae

†Steinhauera

†Steinhauera subglobosa

Presl emend Mai, 1968

A sweet gum species.
Raceme with at least 7 infructecences

rowspan=3| Hamamelidaceae

Mytilaria

†Mytilaria boglei

Collinson, Manchester, & Wilde

A Mytilaria species infructecence.

rowspan=2| Corylopsis

†Corylopsis maii

Collinson, Manchester, & Wilde, 2012

A winter hazel species infructecence.

†Corylopsis waltheri

Collinson, Manchester, & Wilde, 2012

Isolated winter hazel species seeds.

=Vitaceae=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=6| Vitaceae

Ampelopsis

Ampelopsis sp.

A peppervine species seed.
not identified to species

Cayratia

†Cayratia jungii

(Gregor) Chen & Manchester

A Cayratia species seed.

†Crassivitisemen

†Crassivitisemen wildei

(Chen & Manchester) Collinson, Manchester, & Wilde

Vitaceous species fruits and seeds.
Fruits with soft tissue preservation

†Palaeovitis

†Palaeovitis sp.

A vitaceous species seed.
not identified to species

Parthenocissus

†Parthenocissus britannica

(Heer) Chandler, 1957

Virginia creeper species fruits and seeds.

Vitis

†Vitis messelensis

Collinson, Manchester, & Wilde, 2012

Grape species fruits and seeds.
Some specimens preserved in coprolites

''Incertae sedis''

Carpolitus is a plant morphogenus circumscribed for fossil fruits and seeds that are distinct but not identifiable to defined taxa. Collinson, Manchester, & Wilde (2012) described but did not name 63 distinct Carpolithus morphotypes from the Messel Formation.

Genus ! Species ! Authority ! Notes ! Image
class="wikitable" align="center" width="100%"
rowspan=7\| †Carpolithus \| †Carpolithus anacardiaceus \| Engelhardt \| Doubtfully referred to Anacardiaceae per Edwards & Wonnacott (1935) \|
†Carpolithus callosaeoides \| (Engelhardt) \| A Carpolithus species with thick woody tissues \|
†Carpolithus drupaceus \| Engelhardt \| A fruit of possible rosaceous affinity \|
†Carpolithus euphoriaeoides \| Engelhardt \| A Carpolithus species fruit Suggested to be similar to Euphoria \|
†Carpolithus sapindiformis \| Engelhardt \| A Carpolithus species fruit Suggested to possibly be affiliated to Sapindus leaves \|
†Carpolithus sapindoides \| Engelhardt \| A Carpolithus species fruit Suggested to possibly be comparable to Sapindus fruits \|
Species 1–62 \| \| various distinct morphotypes \|
†Lagokarpos \| incertae sedis{{Cite journal\|last1=Tang \|first1=H. \|last2=Liu \|first2=J. \|last3=Wu \|first3=F.-X. \|last4=Spicer \|first4=T. \|last5=Spicer \|first5=R. \|last6=Deng \|first6=W.-Y.-D. \|last7=Xu \|first7=C.-L. \|last8=Zhao \|first8=F. \|last9=Huang \|first9=J. \|last10=Li \|first10=S.-F. \|last11=Su \|first11=T. \|last12=Zhou \|first12=Z.-K. \|year=2019 \|title=Extinct genus Lagokarpos reveals a biogeographic connection between Tibet and other regions in the Northern Hemisphere during the Paleogene \|journal=Journal of Systematics and Evolution \|volume=57 \|issue=6 \|pages=670–677 \|doi=10.1111/jse.12505 \|doi-access=free }} \| \| A Lagokarpos species winged fruit Not identified to species \|
†Saportaspermum \| †Saportaspermum kovacsiae \| \| A Saportaspermum species winged seed Suggested to have malvaceous affinity \|
†Spirellea \| †incertae sedis \| \| A Spirellea species seed Not identified to species \|

Pollen and spores

The first foray into Messel palynology was produced by H. D. Pflug (1952){{cite journal |last1=Pflug |first1=H.D. |year=1952 |title=Palynologie und Stratigraphie der eozänen Braunkohlen von Helmstedt (Palynology and stratigraphy of the Eocene Brown-Coal of Helmstedt) |journal=Paläontologische Zeitschrift |volume=26 |issue=(1/2) |pages=112–137 |doi=10.1007/BF03041729 |s2cid=140715968 |language=German}} with follow-up papers in 1957{{cite journal |last1=Pflug |first1=H.D. |year=1957 |title=Zur Altersfolge und Faziesgliederung mitteleuropäischer (insbesondere hessischer) Braunkohlen (On the age succession and facies correlation of Middle European, (in particular Hessen) Brown Coal.) |journal=Notizbl Hess Landesamt Bodenforsch Wiesbaden |volume=85 |pages=152–178 |language=German}} and coauthored with P. Thomas in 1953.{{cite journal |last1=Thomson |first1=P. |last2=Pflug |first2=H. |year=1953 |title=Pollen und Sporen des mitteleuropäischen Tertiärs (Pollen and spores of the Middle European Tertiary) |journal=Palaeontographica Abteilung B |volume=94 |pages=1–138 |language=German}} The pollen and spore record was revised and expanded by Heidemarie Thiele-Pfeiffer (1988){{cite web |last1=White |first1=J M |title=Thiele-Pfeiffer, H |url=https://paleobotany.ru/palynodata/publication/15550 |website=Palynodata |publisher=Palynodata Inc. |access-date=2 July 2022}} who provided the largest in depth palynological work focusing exclusively on the Messel Formation.

=Pteridophyte spores=

Family ! Genus ! Species ! Authority ! Notes ! Image
class="wikitable" align="center" width="100%"
Osmundaceae \| †Baculatisporites \| †Baculatisporites primarius{{cite journal \|last1=Thiele-Pfeiffer \|first1=H. \|year=1988 \|title=The microflora from the Middle Eocene oil shale of Messel near Darmstadt West Germany \|journal=Palaeontographica Abteilung B \|volume=211 \|pages=1–86}} \| (Wolff) Thomson & Pflug \| A royal fern palynospecies \|
rowspan=2\| Polypodiaceae \| rowspan=2\| †Polypodiaceoisporites \| †Polypodiaceoisporites gracillimus \| Nagy \| Fern spores polypodiaceous affinity \|
†Polypodiaceoisporites lusaticus \| Krutzsch \| Fern spores polypodiaceous affinity \|
rowspan=3\| Schizaeaceae \| rowspan=2\| †Cicatricosisporites \| †Cicatricosisporites dorogensis \| Potonié & Gelletich \| A schizaeaceous fern palynospecies \|
†Cicatricosisporites paradorogensis \| Krutzsch \| A schizaeaceous fern palynospecies \|
†Microfoveolatosporis \| †Microfoveolatosporis granuloides \| (Krutzsch) Krutzsch \| A schizaeaceous fern palynospecies \|
?Schizaeaceae \| †Ischyosporites \| †Ischyosporites tertiarius \| Krutzsch \| A palynospecies of possible schizaeaceous fern affinity \|
rowspan=7\| incertae sedis \| rowspan=2\| †Laevigatosporites \| †Laevigatosporites discordatus \| Pflug \| A palynospecies of possible Schizaeaceae or Polypodiaceae S. L. affinity \|
†Laevigatosporites haardtii \| (Potonié & Venitz) Thomson & Pflug \| A palynospecies of possible Schizaeaceae or Polypodiaceae S. L. affinity \|
rowspan=5\| †Leiotriletes \| †Leiotriletes kopeckii \| Kedves \| Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity \|
†Leiotriletes maxoides \| Krutzsch \| Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity \|
†Leiotriletes microadriennis \| Krutzsch \| Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity \|
†Leiotriletes triangulus \| Murriger & Pflug \| Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity \|
†Leiotriletes sp. \| \| Fern spores of Schizaeaceae and Polypodiaceae S. L. affinity A palynomorph component in zone 1 of the Messel Formation. \|

=Lycophyte spores=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Selaginellaceae

†Echinatisporis

†Echinatisporis hungaricus

Kedves

A spikemoss palynospecies

=Conifer pollens=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=3| Cupressaceae

†Cupressacites

†Cupressacites insulipapillatus

(Trevisan) Krutzsch

A taxodioid cypress family palynospecies

rowspan=2| †Inaperturopollenites

†Inaperturopollenites concedipites

(Wodehouse) Krutzsch

A taxodioid cypress family palynospecies

†Inaperturopollenites verrupapillatus

Trevisan

A taxodioid cypress family palynospecies

rowspan=2| Pinaceae

| rowspan=2| †Pityosporites

†Pityosporites labdacus

(Potonié) Thomson & Pflug

A pine family palynomorph.
A component in zone 1–3 of the Messel Formation.

†Pityosporites microalatus

(Potonié) Thomson Pflug

A pine family palynomorph.

=Basal angiosperm pollens=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Nymphaeaceae

†Monocolpopollenites

†Monocolpopollenites crassiexinus

Thiele-Pfeiffer

A waterlily family palynospecies
Possibly affiliated with Nymphaea

=Chloranthalean pollens=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Chloranthaceae

†Emmapollis

†Emmapollis pseudoemmaensis

Thiele-Pfeiffer

A chloranthaceous palynomorph
Possibly affiliated with Ascarina or Ascarinopsis

=Magnoliid pollens=

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Magnoliaceae

†Magnolipollis

†Magnolipollis magnolioides

Krutzsch

A magnolia family palynospecies
Possible affiliated with Magnolia or Michelia

=Monocot pollens=

==Alismatalean pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Hydrocharitaceae

†Punctilongisulcites

†Punctilongisulcites microechinatus

Thiele-Pfeiffer

A frogbit palynomorph
A component in zone 1 of the Messel Formation.
Possibly affiliated with Blyxa, Ottelia, or Stratiotes

==Arecalean pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Arecaceae

†Monocolpopollenites

†Monocolpopollenites tranquillus

(Potonié) Thomson & Pflug

A palm family palynospecies
Possibly affiliated with Phoenix

==Commelinid pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=2| Cyperaceae

| rowspan=2| †Cyperaceaepollis

†Cyperaceaepollis germanicus

Krutzsch

A sedge family palynospecies
Affiliated with Carex or possibly Lepironia

Unnamed

Krutzsch

A sedge family palynomorph
of uncertain generic affinity

rowspan=2| Restionaceae

| rowspan=2| †Milfordia

†Milfordia incerta

(Thomson & Pflug) Krutzsch

A restiad family palynospecies
Possibly affiliated with Hypolaena, Leptocarpus, Lepyrodia, or Restio

†Milfordia minima

Krutzsch

A restiad family palynospecies
Possibly affiliated with Lyginia

=Superasterid pollens=

==Campanulid pollens==

Family ! Genus ! Species ! Authority ! Notes ! Image
class="wikitable" align="center" width="100%"
rowspan=3\| Aquifoliaceae \| rowspan=3\| †Ilexpollenites \| †Ilexpollenites iliacus \| (Potonié) Thiergart \| A holly family pollen Affiliated with Ilex \|
†Ilexpollenites margaritatus \| (Potonié) Thiergart \| A holly family pollen Possibly affiliated Ilex mucronata \|
†Ilexpollenites propinquus \| (Potonié) Potonié \| A holly family pollen Possibly affiliated Ilex mucronata \|
rowspan=2\| Araliaceae \| rowspan=2\| †Araliaceoipollenites \| †Araliaceoipollenites euphori \| (Potonié) Potonié \| An araliaceous pollen \|
†Araliaceoipollenites reticuloides \| Thiele-Pfeiffer \| An araliaceous pollen \|

==Cornalean pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Nyssaceae

†Nyssapollenites

†Nyssapollenites kruschii

Potonié

A nyssaceous palynospecies.
Two described subspecies N. k. subsp. accessorius & N. k. subsp. analepticus

==Ericalean pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=2| Ericaceae

| rowspan=2| †Ericipites

†Ericipites callidus

(Potonié) Krutzsch

A heather/heath family palynomorph
Of uncertain generic affiliation

†Ericipites ericius

(Potonié) Potonié

A heather/heath family palynomorph
Of uncertain generic affiliation

Symplocaceae

†Porocolpopollenites

†Porocolpopollenites rarobaculatus

Thiele-Pfeiffer

A sweetleaf family palynospecies.
Affiliated with Symplocos

==Lamiid pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

Icacinaceae

†Compositoipollenites

†Compositoipollenites rhizophorus

(Potonié) Potonié

A moonseed palynomorph affiliated with Iodes.

==Santalalean pollens==

class="wikitable" align="center" width="100%"

Family

! Genus

! Species

! Authority

! Notes

! Image

rowspan=2| Olacaceae

†Anacolosidites

†Anacolosidites efflatus

(Potonié) Erdtman

An anacoloseous palynospecies

†Olaxipollis

Cf. †Olaxipollis matthesi

An olacaceous palynomorph
Possibly affiliated with O. matthesi

=Superrosid pollens=

==Fabid pollens==

Family ! Genus ! Species ! Authority ! Notes ! Image
class="wikitable" align="center" width="100%"
Betulaceae \| †Polyvestibulopollenites \| †Polyvestibulopollenites verus \| (Potonié) Thomson & Pflug \| A betulaceous palynospecies Affiliated with Alnus \|
rowspan=2\| Cannabaceae \| rowspan=2\| †Celtipollenites \| †Celtipollenites intrastructurus \| (Krutzsch & Vanhoorne) Thiele-Pfeiffer \| A hemp family palynospecies Affiliated with Celtis \|
†Celtipollenites laevigatus \| Thiele-Pfeiffer \| A hemp family palynospecies Affiliated with Celtis \|
rowspan=15\| Juglandaceae \| rowspan=2\| †Caryapollenites \| †Caryapollenites circulus \| (Pflug) Krutzsch \| A Walnut family palynospecies Affiliated with Carya \|
Cf. †Caryapollenites triangulus \| \| A Walnut family palynospecies Possibly affiliated with Caryapollenites triangulus \|
†Labraferoidaepollenites \| †Labraferoidaepollenites menatensis \| Kedves \| A Walnut family palynospecies Possibly affiliated with Engelhardia \|
rowspan=2\| †Momipites \| †Momipites punctatus \| (Potonié) Nagy \| A Walnut family palynospecies Possibly affiliated with Engelhardtia, Oreomunnea, or Aifaroa \|
†Momipites quietus \| (Potonié) Nichols \| A Walnut family palynospecies Possibly affiliated with Engelhardtia \|
†Nudopollis \| †Nudopollis terminalis \| (Thomson & Pflug) Pflug \| A Walnut family palynospecies One named subspecies N. terminalis subsp hastaformis \|
rowspan=4\| †Platycaryapollenites \| †Platycaryapollenites miocaenicus \| Nagy \| A Walnut family palynospecies Affiliated with Platycarya \|
†Platycaryapollenites platycaryoides \| (Roche) Kedves \| A Walnut family palynospecies Affiliated with Platycarya \|
†Platycaryapollenites semicyclus \| (Krutzsch & Vanhoorne) Thiele-Pfeiffer \| A Walnut family palynospecies Possibly affiliated with Platycarya \|
†Platycaryapollenites sp. \| \| A Walnut family palynospecies Affiliated with Platycarya A component in zone 1–3 of the Messel Formation. \|
rowspan=4\| †Plicatopollis \| †Plicatopollis hungaricus \| Kedves \| A Walnut family palynospecies \|
†Plicatopollis plicatus \| (Potonié) Krutzsch \| A Walnut family palynospecies \|
†Plicatopollis pseudoexcelsus \| Krutzsch \| A Walnut family palynospecies One named subspecies P. p. subsp. microturgidus'' Two unnamed subspecies \|
†Plicatopollis Sp. \| \| A Walnut family palynospecies Not identified to species \|
†Pterocaryapollenites \| †Pterocaryapollenites stellatus \| (Potonié) Thiergart \| A Walnut family palynospecies Affiliated with Pterocarya \|
?Juglandaceae \| †Plicatopollis \| †Plicatopollis lunatus \| Kedves \| A palynospecies of possible juglandaceous affinity \|
rowspan=3\| Ulmaceae \| rowspan=3\| †Polyporopollenites \| †Polyporopollenites eoulmoides \| Krutzsch & Vanhoorne \| An elm family palynospecies \|
†Polyporopollenites undulosus \| (Wolf) Thomson & Pflug \| An elm family palynospecies Possibly affiliated with Ulmus or Zelkova \|
†Polyporopollenites verrucatus \| (Wolf) Thomson & Pflug \| An elm family palynospecies Possibly affiliated with Hemiptelea \|

==Malvid pollens==

Family ! Genus ! Species ! Authority ! Notes ! Image
class="wikitable" align="center" width="100%"
rowspan=5\| Malvaceae \| rowspan=3\| †Bombacacidites \| †Bombacacidites tilioides \| Krutzsch \| A bombacoid palynospecies Possibly affiliated with Bombax \|
†Bombacacidites Sp.1 \| \| A bombacoid palynomorph Described but not formally named Affiliated with Bombax \|
†Bombacacidites Sp.2 \| \| A bombacoid palynomorph Described but not formally named Affiliated with Bombax \|
rowspan=2\| †Intratriporopollenites \| †Intratriporopollenites minimus \| Mai \| A tilioid palynospecies Of uncertain generic affinity \|
Cf. †Intratriporopollenites maxoides \| \| A tilioid palynomorph Possibly affiliated with I. maxoides \|
Thymelaeaceae \| †Pseudospinaepollis \| †Pseudospinaepollis pseudospinosus \| Krutzsch \| A thymelaeaceous palynospecies Possibly affiliated with Stellera or Wikstroemia \|