Paleoflora of the Messel Formation
This is an overview of the
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
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Lenz et al., 2007 |
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.[7]
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Undescribed[7] |
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.[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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Asplenium sp.[2] |
Fern pinnuels attributed to Asplenium |
Conifers
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
Nymphaeales
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
Laurales
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
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
Alismatales
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
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Smilacaceae | Smilax |
Unger |
A greenbrier |
||
Heer |
A greenbrier |
||||
Wess. |
A greenbrier |
||||
Heer |
A greenbrier |
Pandanales
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
S. Y. Smith, Collinson & Rudall, 2008 |
A Cyclanthus species |
Poales
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Smith et al., 2009 |
|||||
A.Braun ex Heer |
A grass foliage |
Basal eudicots
Proteales
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Meliosma sp.[5] |
A Meliosma species seed |
Ranunculales
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
Campanulids
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Manchester) Manchester |
A Toricellia species. |
Cornales
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
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
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
Fabids
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 |
||||
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
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Decodon
|
Incertae sedis[5] |
Seeds of similar morphology to Decodon[5] Not identified to species |
Saxifragales
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
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 |
coprolites
|
Incertae sedis
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
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
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 S. L. affinity
|
||
(Potonié & Venitz) Thomson & Pflug |
A palynospecies of possible S. L. affinity
|
||||
†Leiotriletes |
Kedves |
Fern spores of S. L. affinity
|
|||
Krutzsch |
Fern spores of S. L. affinity
|
||||
Krutzsch |
Fern spores of S. L. affinity
|
||||
Murriger & Pflug |
Fern spores of S. L. affinity
|
||||
Fern spores of S. L. affinityA palynomorph component in zone 1 of the Messel Formation. |
Lycophyte spores
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Selaginellaceae
|
Kedves |
A spikemoss palynospecies
|
Conifer pollens
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer |
A waterlily family palynospecies |
Chloranthalean pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer |
A chloranthaceous palynomorph |
Magnoliid pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Krutzsch |
A magnolia family palynospecies |
Monocot pollens
Alismatalean pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Thiele-Pfeiffer |
A frogbit palynomorph |
Arecalean pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Thomson & Pflug |
A palm family palynospecies |
Commelinid pollens
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
Campanulid pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Aquifoliaceae
|
†Ilexpollenites |
(Potonié) Thiergart |
A Ilex
|
||
(Potonié) Thiergart |
A holly family pollen
Possibly affiliated Ilex mucronata |
||||
(Potonié) Potonié |
A holly family pollen
Possibly affiliated Ilex mucronata |
||||
Araliaceae | †Araliaceoipollenites |
(Potonié) Potonié |
An araliaceous pollen |
||
Thiele-Pfeiffer |
An araliaceous pollen |
Cornalean pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Potonié |
A nyssaceous palynospecies. |
Ericalean pollens
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Potonié |
Santalalean pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Olacaceae |
(Potonié) Erdtman |
An anacoloseous palynospecies |
|||
An olacaceous palynomorph |
Superrosid pollens
Fabid pollens
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
(Potonié) Thomson & Pflug |
A Alnus
|
||||
Cannabaceae | †Celtipollenites |
(Krutzsch & Vanhoorne) Thiele-Pfeiffer |
A hemp family palynospecies |
||
Thiele-Pfeiffer |
A hemp family palynospecies |
||||
Juglandaceae | †Caryapollenites |
(Pflug) Krutzsch |
A Carya
|
||
A Walnut family palynospecies |
|||||
Kedves |
A Walnut family palynospecies |
||||
†Momipites |
(Potonié) Nagy |
A Engelhardtia, Oreomunnea, or Aifaroa
|
|||
(Potonié) Nichols |
A Engelhardtia
|
||||
(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 Ulmus or Zelkova
|
||||
(Wolf) Thomson & Pflug |
An elm family palynospecies |
Malvid pollens
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
Family | Genus | Species | Authority | Notes | Image |
---|---|---|---|---|---|
Lythraceae | †Lythraceaepollenites |
Thiele-Pfeiffer |
A myrtle family palynospecies |
||
†Lythraceaepollenites sp.[3] |
A Woodfordia Described but not named |
Incertae sedis pollens
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
- ^ 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.
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