Paleoflora of the Messel Formation
This is an overview of the paleoflora 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 Hermann Engelhardt producing the first leaf monograph in 1922,[2] followed by Heidemarie Thiele-Pfeiffer in 1988[3][4] 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.[5]
Dinoflagellates
[edit]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.[6]
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Lenz et al., 2007 |
Charophyta
[edit]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.[7]
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Undescribed[7] | A minor Charophyte green algae component of the Messel Formation. |
Chlorophyta
[edit]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.[7] 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.[8][6]
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Kützing | |||||
(A. Braun) Hansgirg | A chlorophyte green algae |
Pteridophytes
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Asplenium sp.[2] | Fern pinnuels attributed to Asplenium |
Conifers
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
†Doliostrobus taxiformis[5] | (Sternberg) Z. Kvaček emend. Kunzmann | A conifer of possible cupressaceous affinities | |||
Engelhardt | Possible podocarpaceous foliage | ||||
Wilde | A plum yew species |
Basal angiosperms
[edit]Nymphaeales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
†Nuphaea engelhardtii[10] | Gee & Taylor, 2018 | A transitional nymphaealean water lily genus | |||
Incertae sedis | Incertae sedis | Incertae sedis[5] | Seeds of possible nymphaealean affinity |
Magnoliids
[edit]Laurales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Lauraceae | †Laurocarpum sp.[5] | Seeds of lauraceous affinity | |||
(Engelhardt) Wilde | A lauraceous leaf morphotaxon | ||||
incertae sedis | incertae sedis[5] | Seeds of lauraceous affinity |
Magnoliales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Magnoliaceae | Magnolia | Magnolia sp.[5] | A Magnolia species fruit | ||
Magnolia sp.[5] | Magnolia species seeds | ||||
Myristicacarpum sp.[5] | A Myristicacarpum species seed |
Monocots
[edit]Alismatales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Araceae | †Araciphyllites | †Araciphyllites tertiarius[12] | (Engelhardt) Wilde, Z. Kvaček & Bogner, 2005 | An araceous leaf | |
†Araciphyllites schaarschmidtii[12] | Wilde, Z. Kvaček & Bogner, 2005 | An araceous leaf | |||
†Caladiosoma messelense[12] | Wilde, Z. Kvaček & Bogner, 2005 | An araceous leaf |
Arecales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Arecaceae | (Collinson, Manchester, & Wilde) Doweld | Palm species fruits | |||
A Palm leaf | |||||
Incertae sedis | incertae sedis[5] | Pinnate palm leaves of possible lepidocaryoid tribal affinity. | |||
incertae sedis[5] | Palmate palm leaves of uncertain tribal affinity. |
Liliales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Smilacaceae | Smilax | Unger | A greenbrier | ||
Heer | A greenbrier | ||||
Wess. | A greenbrier | ||||
Heer | A greenbrier |
Pandanales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
S. Y. Smith, Collinson & Rudall, 2008 | A Cyclanthus species |
Poales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Smith et al., 2009 | |||||
A.Braun ex Heer | A grass foliage |
Basal eudicots
[edit]Proteales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Meliosma sp.[5] | A Meliosma species seed |
Ranunculales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Menispermaceae | Collinson, Manchester, & Wilde, 2012 | A Cocculus species endocarp | |||
Collinson, Manchester, & Wilde, 2012 | A Diploclisia species seed | ||||
Collinson, Manchester, & Wilde, 2012 | A menispermaceous endocarp | ||||
Collinson, Manchester, & Wilde, 2012 | A menispermaceous genus seed | ||||
†Palaeosinomenium | Collinson, Manchester, & Wilde, 2012 | A menispermaceous endocarp | |||
Chandler, 1961 | A menispermaceous endocarp | ||||
†Palaeosinomenium sp.[5] | A menispermaceous endocarp | ||||
A menispermaceous endocarp | |||||
?Pericampylus sp.[5] | A tentative Pericampylus species endocarp. | ||||
Collinson, Manchester, & Wilde, 2012 | A Stephania species seed | ||||
Collinson, Manchester, & Wilde, 2012 | A menispermaceous endocarp | ||||
†Wardensheppeya sp.[5] | A menispermaceous endocarp | ||||
incertae sedis | incertae sedis[5] | menispermaceous endocarp of Tinosporeae subfamily affinity |
Superasterids
[edit]Campanulids
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Manchester) Manchester | A Toricellia species. |
Cornales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Alangium sp.[5] | An Alangium species likely belonging to Alangium sect. Marlea. | ||||
Heer | A dogwood species | ||||
Nyssaceae | incertae sedis[5] | A germination valve similar to Beckettia | |||
An endocarp similar to Eomastixia rugosa | |||||
incertae sedis[5] | A possible Mastixia endocarp and mesocarp | ||||
Kirchheimer | A nyssaceous endocarp | ||||
Nyssa | (R. Ludw.) Kirchheimer | A nyssaceous endocarp with germination valve | |||
Unger | A nyssaceous species | ||||
Unger | A nyssaceous species |
Ericales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
incertae sedis[5] | A Cleyera species. | ||||
(Sternberg) Kirchheimer | A Symplocos species fruit. | ||||
†Camelliacarpoidea messelensis[5] | Collinson, Manchester, & Wilde | A theaceous species. |
Lamiids
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
incertae sedis[5] | A Cypselites species | ||||
A catalpa species. | |||||
Icacinaceae | †Palaeohosiea | (Ettingshausen) Kvaček & Bůžek | A phytocrenaceous species. | ||
incertae sedis[5] | A Palaeohosiea endocarp. | ||||
incertae sedis[5] | Collinson, Manchester, & Wilde | A Cleyera species. | |||
†Icacinicarya | Collinson, Manchester, & Wilde | An icacinaceous species. | |||
Collinson, Manchester, & Wilde | An icacinaceous species. | ||||
incertae sedis[5] | An Icacinicarya endocarp. | ||||
incertae sedis[5] | An endocarp similar to Natsiatum. | ||||
incertae sedis[5] | A Palaeophytocrene endocarp. | ||||
Collinson, Manchester, & Wilde | A Phytocrene species. | ||||
Pyrenacantha | "Species 1"[5] | Collinson, Manchester, & Wilde | An unnamed Pyrenacantha species. | ||
"Species 2"[5] | Collinson, Manchester, & Wilde | An unnamed Pyrenacantha species. | |||
incertae sedis[5] | Collinson, Manchester, & Wilde | An endocarp similar to Pyrenacantha. |
Superrosids
[edit]Fabids
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
A hemp relative fruit. | |||||
Celastraceae | Celastrus | Unger | A Celastrus species. | ||
Wonnacott | A Celastrus species. | ||||
Engelhardt | A Celastrus species. | ||||
Unger | A Celastrus species. | ||||
Elaeodendron | (Unger) Ettingshausen | An Elaeodendron species. | |||
Ettingshausen | An Elaeodendron species. | ||||
Engelhardt | An Elaeodendron species. | ||||
Ettingshausen | A Euonymus species. | ||||
Collinson, Manchester, & Wilde | A Sloanea species fruit. | ||||
Collinson, Manchester, & Wilde | Trilocular euphorb fruits. | ||||
Fabaceae | Engelhardt | A caesalpinioid legume species pod. | |||
†Leguminocarpon | Collinson, Manchester, & Wilde | A legume species pod. | |||
†Leguminocarpon sp.[5] | Legume species pods. | ||||
Juglandaceae | †Cruciptera schaarschmidtii[5] | Manchester, Collinson, & Goth | A juglandaceous winged fruit. | ||
†Hooleya sp.[5] | A juglandaceous fruiting raceme. | ||||
†Palaeocarya sp.[5] | A juglandaceous fruit. | ||||
Collinson, Manchester, & Wilde | |||||
Unger | A cherry relative | ||||
(Ettingshausen) Manchester emend. Wilde & Manchester | An ulmaceous elm relative fruit. |
Malvids
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Anacardiaceae | Manchester, Wilde & Collinson | A cashew species. | |||
Collinson, Manchester, & Wilde | A Lannea species. | ||||
Ettingshausen | A mango relative. | ||||
(Reid & Chandler) Manchester | A spondiadeous cashew relative. | ||||
(Reid & Chandler) Manchester | A Pleiogynium species. | ||||
Rhus | Heer | A sumac relative. | |||
Pilar | A sumac relative. | ||||
Engelhardt | A sumac relative. | ||||
(Reid & Chandler) Mai | A Tapiscia species fruit. | ||||
Burseraceae | †Bursericarpum species[5] | A Bursericarpum species endocarp. | |||
Canarium sp.[5] | A Canarium species endocarp. | ||||
Malvaceae | †Byttneriopsis | (Ettingshausen) Kvaček & Wilde | A malvaceous leaf morphotaxon | ||
(Ettingshausen) Kvaček & Wilde | A malvaceous leaf morphotaxon | ||||
Kvaček & Wilde | A malvaceous leaf morphotaxon | ||||
Rutaceae | Collinson & Gregor | A citrus family seed. | |||
Wilde | A Toddalia leaf morphospecies | ||||
incertae sedis[5] | A Seed similar to Toddalia. | ||||
Sapindaceae | Ettingshausen | A Dodonaea species | |||
Sapindus | Ettingshausen | A Sapindus species | |||
Unger | A Sapindus species | ||||
Engelhardt | A Sapindus species | ||||
Unger | A Sapindus species | ||||
Engelhardt | A Thouinia species | ||||
Unger | An Ailanthus seed. |
Myrtales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Incertae sedis[5] | Seeds of similar morphology to Decodon[5] Not identified to species |
Saxifragales
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Presl emend Mai, 1968 | A sweet gum species. | ||||
Hamamelidaceae | Collinson, Manchester, & Wilde | A Mytilaria species infructecence. | |||
Corylopsis | Collinson, Manchester, & Wilde, 2012 | A winter hazel species infructecence. | |||
Collinson, Manchester, & Wilde, 2012 | Isolated winter hazel species seeds. |
Vitaceae
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Vitaceae | Ampelopsis sp.[5] | A peppervine species seed. | |||
(Gregor) Chen & Manchester | A Cayratia species seed. | ||||
(Chen & Manchester) Collinson, Manchester, & Wilde | Vitaceous species fruits and seeds. | ||||
†Palaeovitis sp.[5] | A vitaceous species seed. | ||||
(Heer) Chandler, 1957 | Virginia creeper species fruits and seeds. | ||||
Collinson, Manchester, & Wilde, 2012 | Grape species fruits and seeds. |
Incertae sedis
[edit]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 | Engelhardt | Doubtfully referred to Anacardiaceae per Edwards & Wonnacott (1935) | ||
(Engelhardt) | A Carpolithus species with thick woody tissues | |||
Engelhardt | A fruit of possible rosaceous affinity | |||
Engelhardt | A Carpolithus species fruit | |||
Engelhardt | A Carpolithus species fruit | |||
Engelhardt | A Carpolithus species fruit | |||
Species 1–62[5] | various distinct morphotypes | |||
incertae sedis[24] | A Lagokarpos species winged fruit | |||
A Saportaspermum species winged seed | ||||
†incertae sedis[5] | A Spirellea species seed |
Pollen and spores
[edit]The first foray into Messel palynology was produced by H. D. Pflug (1952)[25] with follow-up papers in 1957[26] and coauthored with P. Thomas in 1953.[27] The pollen and spore record was revised and expanded by Heidemarie Thiele-Pfeiffer (1988)[4] who provided the largest in depth palynological work focusing exclusively on the Messel Formation.
Pteridophyte spores
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Wolff) Thomson & Pflug | A royal fern palynospecies | ||||
Polypodiaceae | †Polypodiaceoisporites | Nagy | Fern spores polypodiaceous affinity | ||
Krutzsch | Fern spores polypodiaceous affinity | ||||
Schizaeaceae | †Cicatricosisporites | Potonié & Gelletich | A schizaeaceous fern palynospecies | ||
Krutzsch | A schizaeaceous fern palynospecies | ||||
(Krutzsch) Krutzsch | A schizaeaceous fern palynospecies | ||||
Krutzsch | A palynospecies of possible schizaeaceous fern affinity | ||||
incertae sedis | †Laevigatosporites | Pflug | A palynospecies of possible Schizaeaceae or Polypodiaceae S. L. affinity | ||
(Potonié & Venitz) Thomson & Pflug | A palynospecies of possible Schizaeaceae or Polypodiaceae S. L. affinity | ||||
†Leiotriletes | Kedves | Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity | |||
Krutzsch | Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity | ||||
Krutzsch | Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity | ||||
Murriger & Pflug | Fern spores of Schizaeaceae or Polypodiaceae S. L. affinity | ||||
Fern spores of Schizaeaceae and Polypodiaceae S. L. affinity |
Lycophyte spores
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Kedves | A spikemoss palynospecies |
Conifer pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Cupressaceae | (Trevisan) Krutzsch | A taxodioid cypress family palynospecies | |||
†Inaperturopollenites | (Wodehouse) Krutzsch | A taxodioid cypress family palynospecies | |||
Trevisan | A taxodioid cypress family palynospecies | ||||
Pinaceae | †Pityosporites | (Potonié) Thomson & Pflug | A pine family palynomorph. | ||
(Potonié) Thomson Pflug | A pine family palynomorph. |
Basal angiosperm pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer | A waterlily family palynospecies |
Chloranthalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer | A chloranthaceous palynomorph |
Magnoliid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Krutzsch | A magnolia family palynospecies |
Monocot pollens
[edit]Alismatalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer | A frogbit palynomorph |
Arecalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Thomson & Pflug | A palm family palynospecies |
Commelinid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Cyperaceae | †Cyperaceaepollis | Krutzsch | A sedge family palynospecies | ||
Unnamed[3] | Krutzsch | A sedge family palynomorph | |||
Restionaceae | †Milfordia | (Thomson & Pflug) Krutzsch | A restiad family palynospecies | ||
Krutzsch | A restiad family palynospecies |
Superasterid pollens
[edit]Campanulid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Aquifoliaceae | †Ilexpollenites | (Potonié) Thiergart | A holly family pollen | ||
(Potonié) Thiergart | A holly family pollen | ||||
(Potonié) Potonié | A holly family pollen | ||||
Araliaceae | †Araliaceoipollenites | (Potonié) Potonié | An araliaceous pollen | ||
Thiele-Pfeiffer | An araliaceous pollen |
Cornalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Potonié | A nyssaceous palynospecies. |
Ericalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Ericaceae | †Ericipites | (Potonié) Krutzsch | A heather/heath family palynomorph | ||
(Potonié) Potonié | A heather/heath family palynomorph | ||||
Thiele-Pfeiffer | A sweetleaf family palynospecies. |
Lamiid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Potonié |
Santalalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Olacaceae | (Potonié) Erdtman | An anacoloseous palynospecies | |||
An olacaceous palynomorph |
Superrosid pollens
[edit]Fabid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Thomson & Pflug | A betulaceous palynospecies | ||||
Cannabaceae | †Celtipollenites | (Krutzsch & Vanhoorne) Thiele-Pfeiffer | A hemp family palynospecies | ||
Thiele-Pfeiffer | A hemp family palynospecies | ||||
Juglandaceae | †Caryapollenites | (Pflug) Krutzsch | A Walnut family palynospecies | ||
A Walnut family palynospecies | |||||
Kedves | A Walnut family palynospecies | ||||
†Momipites | (Potonié) Nagy | A Walnut family palynospecies | |||
(Potonié) Nichols | A Walnut family palynospecies | ||||
(Thomson & Pflug) Pflug | A Walnut family palynospecies | ||||
†Platycaryapollenites | Nagy | A Walnut family palynospecies | |||
(Roche) Kedves | A Walnut family palynospecies | ||||
(Krutzsch & Vanhoorne) Thiele-Pfeiffer | A Walnut family palynospecies | ||||
†Platycaryapollenites sp.[7] | A Walnut family palynospecies | ||||
†Plicatopollis | Kedves | A Walnut family palynospecies | |||
(Potonié) Krutzsch | A Walnut family palynospecies | ||||
Krutzsch | A Walnut family palynospecies | ||||
†Plicatopollis Sp.[3] | A Walnut family palynospecies | ||||
(Potonié) Thiergart | A Walnut family palynospecies | ||||
Kedves | A palynospecies of possible juglandaceous affinity | ||||
Ulmaceae | †Polyporopollenites | Krutzsch & Vanhoorne | An elm family palynospecies | ||
(Wolf) Thomson & Pflug | An elm family palynospecies | ||||
(Wolf) Thomson & Pflug | An elm family palynospecies |
Malvid pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Malvaceae | †Bombacacidites | Krutzsch | |||
†Bombacacidites Sp.1[3] | A bombacoid palynomorph | ||||
†Bombacacidites Sp.2[3] | A bombacoid palynomorph | ||||
†Intratriporopollenites | Mai | A tilioid palynospecies | |||
A tilioid palynomorph | |||||
Krutzsch | A thymelaeaceous palynospecies |
Myrtalean pollens
[edit]Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Lythraceae | †Lythraceaepollenites | Thiele-Pfeiffer | A myrtle family palynospecies | ||
†Lythraceaepollenites sp.[3] | A myrtle family palynospecies |
Incertae sedis pollens
[edit]Genus | Species | Authority | Notes | Image |
---|---|---|---|---|
(Krutzsch) Krutzsch | Possible affinity is to Gomphandra (Icacinaceae) or Burseraceae | |||
(Krutzsch & Vanhoorne) | A palynomorph species of uncertain affiliation. | |||
(Potonié) Krutzsch | A palynomorph species of uncertain affiliation. | |||
†Multiporopollenites | Krutzsch | A palynomorph species of uncertain affiliation. | ||
†Multiporopollenites Sp.1[3] | A palynomorph species of uncertain affiliation. | |||
†Multiporopollenites Sp.2[3] | A palynomorph species of uncertain affiliation. | |||
†Pentapollenites | (Pflug) Krutzsch | A palynomorph species of uncertain affiliation. | ||
Krutzsch | A palynomorph species of uncertain affiliation. | |||
†Pentapollenites Sp.1[3] | A palynomorph species of uncertain affiliation. | |||
†Pentapollenites Sp.2[3] | A palynomorph species of uncertain affiliation. | |||
Potonié | A palynomorph species of uncertain affiliation. | |||
(Krutzsch) Krutzsch | A palynomorph species of uncertain affiliation. | |||
†Tricolporopollenites | Thiele-Pfeiffer | A palynomorph with suggested affinity to Oleaceae or Rutaceae | ||
Thiele-Pfeiffer | A palynomorph with suggested affinity to Rosaceae |
References
[edit]- ^ 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.
- ^ a b c d e f g h i Engelhardt, H. (1922). "Die alttertiäre Flora von Messel bei Darmstadt". Abhandlungen der Hessischen Geologischen Landesanstalt zu Darmstadt. 7 (4): 20–128.
- ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk bl bm bn bo bp bq br bs bt bu bv bw bx by bz ca Thiele-Pfeiffer, H. (1988). "The microflora from the Middle Eocene oil shale of Messel near Darmstadt West Germany". Palaeontographica Abteilung B. 211: 1–86.
- ^ a b White, J M. "Thiele-Pfeiffer, H". Palynodata. Palynodata Inc. Retrieved 2 July 2022.
- ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk bl bm bn bo bp bq br bs bt bu bv bw bx by bz ca cb cc cd ce cf cg ch ci 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. ISBN 978-3-510-61400-4.
- ^ a b c Lenz ., O.; Wilde, V.; Riegel, W.; Heinrichs, T. (2007). "Distribution and paleoecologic significance of the freshwater dinoflagellate cyst Messelodinium thielepfeifferae gen. et sp. nov from the Middle Eocene of Lake Messel, Germany". Palynology. 31 (1): 119–134. doi:10.2113/gspalynol.31.1.119.
- ^ a b c d e f g h i j k l m n o p q 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)" (PDF). International Journal of Earth Sciences. 100 (8): 1851–1874. Bibcode:2011IJEaS.100.1851L. doi:10.1007/s00531-010-0609-z. S2CID 52968593.
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