Paleoflora of the Messel Formation explained

This is an overview of the of the Eocene Messel Formation as explored by the Messel Pit excavations in Germany. A former quarry and UNESCO World Heritage Site, the Messel Formation preserves what once were a series of lakes surrounded by sub-tropical rainforest during the Middle Eocene, approximately 47 Ma.^[1]

Several major monographs on the paleoflora have been published since the early 1900s, with producing the first leaf monograph in 1922, followed by Heidemarie Thiele-Pfeiffer in 1988 who provided the first in-depth palynological revision and expansion. Most recently 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.

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.^[2]

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.

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.^[3] 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.^[2]

Family	Genus	Species	Authority	Notes	Image
Botryococcaceae	Botryococcus	Botryococcus braunii^[4]	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.

Conifers

Family	Genus	Species	Authority	Notes	Image
Doliostrobaceae	†Doliostrobus	†Doliostrobus taxiformis^[6]	(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^[7]	Wilde	A plum yew species

Basal angiosperms

Nymphaeales

Family	Genus	Species	Authority	Notes	Image
Nymphaeaceae	†Nuphaea	†Nuphaea engelhardtii^[8]	Gee & Taylor, 2018	A transitional nymphaealean water lily genus
Incertae sedis	Incertae sedis	Incertae sedis		Seeds of possible nymphaealean affinity Not identified to species

Magnoliids

Laurales

Family	Genus	Species	Authority	Notes	Image
Lauraceae	†Laurocarpum	†Laurocarpum sp.		Seeds of lauraceous affinity 3 unnamed morphospecies
	†Laurophyllum	†Laurophyllum lanigeroides^[9]	(Engelhardt) Wilde	A lauraceous leaf morphotaxon host to two forms of asphondyliine tribe midge galls
	incertae sedis	incertae sedis		Seeds of lauraceous affinity not identified to genus/species

Magnoliales

Family	Genus	Species	Authority	Notes	Image
Magnoliaceae	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

Family	Genus	Species	Authority	Notes	Image
Araceae	†Araciphyllites	†Araciphyllites tertiarius^[10]	(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

Family	Genus	Species	Authority	Notes	Image
Arecaceae	Arecocaryon	Arecocaryon messelensis^[11]	(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
	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
Smilacaceae	Smilax	†Smilax grandifolia	Unger	A greenbrier
		†Smilax lingulata	Heer	A greenbrier
		†Smilax ovata	Wess.	A greenbrier
		†Smilax reticulata	Heer	A greenbrier

Poales

Family	Genus	Species	Authority	Notes	Image
Cyperaceae	†Volkeria	†Volkeria messelensis^[13]	Smith et al., 2009	A mapanioid sedge
Poaceae	Phragmites	†Phragmites oeningensis	A.Braun ex Heer	A grass foliage

Basal eudicots

Ranunculales

Family	Genus	Species	Authority	Notes	Image
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
	†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	incertae sedis		menispermaceous endocarp of Tinosporeae subfamily affinity 3 morphotypes not identified to genus/species

Superasterids

Cornales

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^[14]	Heer	A dogwood species
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
	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

Family	Genus	Species	Authority	Notes	Image
Pentaphylacaceae	Cleyera	incertae sedis		A Cleyera species. Not identified to species
Symplocaceae	Symplocos	†Symplocos minutula^[15]	(Sternberg) Kirchheimer	A Symplocos species fruit.
Theaceae	†Camelliacarpoidea	†Camelliacarpoidea messelensis	Collinson, Manchester, & Wilde	A theaceous species. A dry fruit with persistent calyx

Lamiids

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
Icacinaceae	†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
	†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.
	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

Family	Genus	Species	Authority	Notes	Image
Cannabaceae	Aphananthe	Cf. †Aphananthe tenuicostata		A hemp relative fruit. Similar to Aphananthe tenuicostata
Celastraceae	Celastrus	†Celastrus elaenus^[16]	Unger	A Celastrus species.
		†Celastrus friedrichi	Wonnacott	A Celastrus species.
		†Celastrus illicinoides	Engelhardt	A Celastrus species.
		†Celastrus oxyphyllus	Unger	A Celastrus species.
	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.
Fabaceae	†Mimosites	†Mimosites spiegeli	Engelhardt	A caesalpinioid legume species pod.
	†Leguminocarpon	†Leguminocarpon herendeenii	Collinson, Manchester, & Wilde	A legume species pod.
		†Leguminocarpon sp.		Legume species pods. 2 mophotypes noted not identified to species
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^[17]	Unger	A cherry relative
Ulmaceae	†Cedrelospermum	†Cedrelospermum leptospermum	(Ettingshausen) Manchester emend. Wilde & Manchester	An ulmaceous elm relative fruit.

Malvids

Family	Genus	Species	Authority	Notes	Image
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^[18]	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
	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
Burseraceae	†Bursericarpum	†Bursericarpum species		A Bursericarpum species endocarp. Not identified to species
	Canarium	Canarium sp.		A Canarium species endocarp. Not identified to species
Malvaceae	†Byttneriopsis	†Byttneriopsis daphnogenes^[19]	(Ettingshausen) Kvaček & Wilde	A malvaceous leaf morphotaxon
		†Byttneriopsis spiegelii	(Ettingshausen) Kvaček & Wilde	A malvaceous leaf morphotaxon
		†Byttneriopsis steuerii	Kvaček & Wilde	A malvaceous leaf morphotaxon
Rutaceae	†Rutaspermum	†Rutaspermum chandleri	Collinson & Gregor	A citrus family seed.
	Toddalia	†Toddalia ovata^[20]	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
Sapindaceae	Dodonaea	†Dodonaea salicites^[21]	Ettingshausen	A Dodonaea species
	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.

Saxifragales

Family	Genus	Species	Authority	Notes	Image
Altingiaceae	†Steinhauera	†Steinhauera subglobosa	Presl emend Mai, 1968	A sweet gum species. Raceme with at least 7 infructecences
Hamamelidaceae	Mytilaria	†Mytilaria boglei	Collinson, Manchester, & Wilde	A Mytilaria species infructecence.
	Corylopsis	†Corylopsis maii	Collinson, Manchester, & Wilde, 2012	A winter hazel species infructecence.
		†Corylopsis waltheri	Collinson, Manchester, & Wilde, 2012	Isolated winter hazel species seeds.

Vitaceae

Family	Genus	Species	Authority	Notes	Image
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
†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^[22]		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)^[23] with follow-up papers in 1957^[24] and coauthored with P. Thomas in 1953.^[25] The pollen and spore record was revised and expanded by Heidemarie Thiele-Pfeiffer (1988)^[26] who provided the largest in depth palynological work focusing exclusively on the Messel Formation.

Pteridophyte spores

Family	Genus	Species	Authority	Notes	Image
Osmundaceae	†Baculatisporites	†Baculatisporites primarius^[27]	(Wolff) Thomson & Pflug	A royal fern palynospecies
Polypodiaceae	†Polypodiaceoisporites	†Polypodiaceoisporites gracillimus	Nagy	Fern spores polypodiaceous affinity
		†Polypodiaceoisporites lusaticus	Krutzsch	Fern spores polypodiaceous affinity
Schizaeaceae	†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
incertae sedis	†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
	†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.

Conifer pollens

Family	Genus	Species	Authority	Notes	Image
Cupressaceae	†Cupressacites	†Cupressacites insulipapillatus	(Trevisan) Krutzsch	A taxodioid cypress family palynospecies
	†Inaperturopollenites	†Inaperturopollenites concedipites	(Wodehouse) Krutzsch	A taxodioid cypress family palynospecies
		†Inaperturopollenites verrupapillatus	Trevisan	A taxodioid cypress family palynospecies
Pinaceae	†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.

Monocot pollens

Commelinid pollens

Family	Genus	Species	Authority	Notes	Image
Cyperaceae	†Cyperaceaepollis	†Cyperaceaepollis germanicus	Krutzsch	A sedge family palynospecies Affiliated with Carex or possibly Lepironia
		Unnamed	Krutzsch	A sedge family palynomorph of uncertain generic affinity
Restionaceae	†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
Aquifoliaceae	†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
Araliaceae	†Araliaceoipollenites	†Araliaceoipollenites euphori	(Potonié) Potonié	An araliaceous pollen
		†Araliaceoipollenites reticuloides	Thiele-Pfeiffer	An araliaceous pollen

Ericalean pollens

Family	Genus	Species	Authority	Notes	Image
Ericaceae	†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

Santalalean pollens

Family	Genus	Species	Authority	Notes	Image
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
Betulaceae	†Polyvestibulopollenites	†Polyvestibulopollenites verus	(Potonié) Thomson & Pflug	A betulaceous palynospecies Affiliated with Alnus
Cannabaceae	†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
Juglandaceae	†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
	†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
	†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.
	†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
Ulmaceae	†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
Malvaceae	†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
	†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

Myrtalean pollens

Family	Genus	Species	Authority	Notes	Image
Lythraceae	†Lythraceaepollenites	†Lythraceaepollenites minimus	Thiele-Pfeiffer	A myrtle family palynospecies Possibly affiliated with Rotala or Ammannia
		†Lythraceaepollenites sp.		A myrtle family palynospecies Possibly affiliated with Rotala, Ammannia, or Woodfordia Described but not named

Incertae sedis pollens

Genus	Species	Authority	Notes	Image
†Brosipollis	†Brosipollis striatobrosus	(Krutzsch) Krutzsch	Possible affinity is to Gomphandra (Icacinaceae) or Burseraceae
†Compositoipollenites	†Compositoipollenites minimus	(Krutzsch & Vanhoorne)	A palynomorph species of uncertain affiliation.
†Labrapollis	†Labrapollis labraferus	(Potonié) Krutzsch	A palynomorph species of uncertain affiliation.
†Multiporopollenites	†Multiporopollenites microreticulatus	Krutzsch	A palynomorph species of uncertain affiliation.
	†Multiporopollenites Sp.1		A palynomorph species of uncertain affiliation.
	†Multiporopollenites Sp.2		A palynomorph species of uncertain affiliation.
†Pentapollenites	†Pentapollenites pentangulus	(Pflug) Krutzsch	A palynomorph species of uncertain affiliation. Two known subspecies P. p. subsp foveostriatus & P. p. subsp pentangulus
	†Pentapollenites punctoides	Krutzsch	A palynomorph species of uncertain affiliation.
	†Pentapollenites Sp.1		A palynomorph species of uncertain affiliation.
	†Pentapollenites Sp.2		A palynomorph species of uncertain affiliation.
†Pollenites	†Pollenites setarius	Potonié	A palynomorph species of uncertain affiliation.
†Pompeckjoidaepollenites	†Pompeckjoidaepollenites subhercynicus	(Krutzsch) Krutzsch	A palynomorph species of uncertain affiliation.
†Tricolporopollenites	†Tricolporopollenites messelensis	Thiele-Pfeiffer	A palynomorph with suggested affinity to Oleaceae or Rutaceae A component in zone 1 of the Messel Formation.
	†Tricolporopollenites solé de portai	Thiele-Pfeiffer	A palynomorph with suggested affinity to Rosaceae A component in zone 1 of the Messel Formation.

Notes and References

1. Mertz . D. . Renne . P. . 2005 . A numerical age for the Messel fossil deposit (UNESCO World Heritage Site) derived from ⁴⁰Ar/³⁹Ar dating on a basaltic rock fragment . Courier Forschungsinstitut Senckenberg . 255 . 7–75.
2. Lenz . . O. . Wilde . V. . Riegel . W. . Heinrichs . T. . Distribution and paleoecologic significance of the freshwater dinoflagellate cyst Messelodinium thielepfeifferae gen. et sp. nov from the Middle Eocene of Lake Messel, Germany . Palynology . 2007 . 31 . 1 . 119–134 . 10.2113/gspalynol.31.1.119.
3. Lenz . O. . Wilde . V. . Riegel . W. . 2011 . Short-term fluctuations in vegetation and phytoplankton during the Middle Eocene greenhouse climate: a 640-kyr record from the Messel oil shale (Germany) . International Journal of Earth Sciences . 100 . 8 . 1851–1874. 10.1007/s00531-010-0609-z . 2011IJEaS.100.1851L . 52968593 . free .
4. Adam . P. . Schaeffer . P. . Albrecht . P. . 2006 . 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) . Organic Geochemistry . 37 . 5 . 584–596. 10.1016/j.orggeochem.2006.01.001 .
5. Engelhardt . H. . 1922 . Die alttertiäre Flora von Messel bei Darmstadt . Abhandlungen der Hessischen Geologischen Landesanstalt zu Darmstadt . 7 . 4 . 20–128 .
6. Collinson . M. . Manchester . S. . Wilde . V. . 2012 . Fossil fruits and seeds of the Middle Eocene Messel biota, Germany. . Abhandlungen der Senckenberg Gesellschaft für Naturforschung . 570 . 1–249 . 978-3-510-61400-4 .
7. Book: Fossilium Catalogus. II. Plantae. . Pars 105. Gymnospermae (6) Ginkgophyta et Coniferae.. Dijkstra . S. . van Amerom . H. . Schultze-Motel . J. . van Amerom . H. . Backhuys Publishers . Leiden, Netherlands . 2002.
8. Gee . C. . Taylor . D. . 2019 . 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. . International Journal of Plant Sciences . 180 . 7 . 724–736 . 10.1086/704376. 201211546 .
9. Wappler . T. . Tokuda . M. . Yukawa . J. . Wilde . V. . 2010 . Insect herbivores on Laurophyllum lanigeroides (Engelhardt 1922) Wilde: the role of a distinct plant-insect associational suite in host taxonomic assignment . Palaeontographica, Palaeontographica, Abt.B: Palaeobotany–Palaeophytology . 283 . 4–6 . 137–155.
10. Wilde . V. . Kvaček . Z. . Bogner . J. . Fossil leaves of the Araceae from the European Eocene and notes on other aroid fossils . International Journal of Plant Sciences . 2005 . 166 . 1 . 157–183 . 10.1086/425673. 85215526 .
11. Doweld . A. B. . 2021 . Fossil Alloceltidoxylon, Allonymphaea, Arecocaryon, Paralnoxylon and Paranyssa and extant Komaroviopsis, Marcanodendron, and Papyrocactus (Magnoliophyta), new replacement generic names . Phytotaxa . 524 . 2 . 92–98 . 10.11646/phytotaxa.524.2.3 . 243482734 .
12. Smith . S. . Collinson . M. . Rudall . P. . Fossil Cyclanthus (Cyclanthaceae, Pandanales) from the Eocene of Germany and England . American Journal of Botany . 2008 . 95 . 6 . 688–699 . 10.3732/ajb.2007390 . 21632394.
13. Smith . S. . Collinson . M. . Simpson . D. . Rudall . P. . Marone . F. . Stampanoni . M. . 2009 . Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe . American Journal of Botany . 96 . 8 . 1506–1518 . 10.3732/ajb.0800427. 21628296 .
14. Book: Fossilium Catalogus. II. Plantae. . Pars 23. Umbellifloraea: Cornaceae. Kirchheimer F. . Jongmans, W. . Verlag Gustav Feller . Neubrandenburg . 2 July 1938.
15. Book: Fossilium Catalogus. II. Plantae. . Pars 26. Symplocaceae.. Kirchheimer F.. Jongmans, W. . Verlag Gustav Feller . 14 Jul 1950.
16. Book: Fossilium Catalogus. II. Plantae. . Pars 29. Celastraceae.. Wonnacott . F. . Jongmans . W. . Uitgeverij Dr. W. Junk . 's-Gravenhague . 28 September 1955.
17. Book: Fossilium Catalogus. II. Plantae. . Pars 25. Rosaceae.. Kirchheimer . F. . Jongmans . W. . Verlag Gustav Feller . 14 Jul 1942.
18. Book: Fossilium Catalogus. II. Plantae. . Pars 20. Anacardiaceae.. Edwards . W. . Wonnacott . F. . Jongmans . W. . W. Junk . 's-Gravenhague . 28 August 1935.
19. Kvacek . Z. . Wilde . V. . 2010 . Foliage and seeds of malvalean plants from the Eocene of Europe . Bulletin of Geosciences . 85 . 1 . 163–182 . 10.3140/bull.geosci.1170. free .
20. Winkler . I. . Labandeira . C. . Wappler . T. . Wilf . P. . 2010 . Distinguishing Agromyzidae (Diptera) leaf mines in the fossil record: new taxa from the Paleogene of North America and Germany and their evolutionary implications . Journal of Paleontology . 84 . 5 . 935–954 . 10.1666/09-163.1. 56378236 .
21. Book: Fossilium Catalogus. II. Plantae. . Pars 14. Sapindaceae.. Edwards . W. . Wonnacott . F. . Jongmans . W. . Verlag Gustav Feller . 14 Jul 1950.
22. Tang . H. . Liu . J. . Wu . F.-X. . Spicer . T. . Spicer . R. . Deng . W.-Y.-D. . Xu . C.-L. . Zhao . F. . Huang . J. . Li . S.-F. . Su . T. . Zhou . Z.-K. . 2019 . Extinct genus Lagokarpos reveals a biogeographic connection between Tibet and other regions in the Northern Hemisphere during the Paleogene . Journal of Systematics and Evolution . 57 . 6 . 670–677 . 10.1111/jse.12505 . free .
23. Pflug . H.D. . 1952 . Palynologie und Stratigraphie der eozänen Braunkohlen von Helmstedt (Palynology and stratigraphy of the Eocene Brown-Coal of Helmstedt) . Paläontologische Zeitschrift . 26 . (1/2) . 112–137 . 10.1007/BF03041729 . 140715968 . German.
24. Pflug . H.D. . 1957 . Zur Altersfolge und Faziesgliederung mitteleuropäischer (insbesondere hessischer) Braunkohlen (On the age succession and facies correlation of Middle European, (in particular Hessen) Brown Coal.) . Notizbl Hess Landesamt Bodenforsch Wiesbaden . 85 . 152–178 . German.
25. Thomson . P. . Pflug . H. . 1953 . Pollen und Sporen des mitteleuropäischen Tertiärs (Pollen and spores of the Middle European Tertiary) . Palaeontographica Abteilung B . 94 . 1–138 . German.
26. Web site: White . J M . Thiele-Pfeiffer, H . Palynodata . Palynodata Inc. . 2 July 2022.
27. Thiele-Pfeiffer . H. . 1988 . The microflora from the Middle Eocene oil shale of Messel near Darmstadt West Germany . Palaeontographica Abteilung B . 211 . 1–86.