Biology:Paleofauna of the Messel Pit

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This is an overview of the fossil fauna of the Eocene Messel Pit in Germany . A former quarry and UNESCO world heritage site, the Messel Pit preserves what once were a series of lakes surrounded by sub-tropical rainforst during the Eocene, approximately 47 Ma.

Sponges

Family Genus Species Authority Notes Image

Spongillidae

Ephydatia

Ephydatia gutenbergiana[1]

Müller et al. 1982

A possibly rheophile species sponge, it grew dominant under favorable conditions and gradually replaced the native population of Lutetiospongilla until a change to the environment. Decrease in freshwater supply likely diminished the sponges population.

Palaeospongillidae

Lutetiospongilla

Lutetiospongilla heili[1]

Richter & Wuttke 1999

A permanent resident of the Messel lake, L. heili was already present when Ephydatia arrived in the lake and differed from the other species by possibly being rheophobe. Following the later species decline, Lutetiospongilla became the second "mass species" of the lake.

"Type 3"[1]

An unnamed sponge, informally referred to as "Type 3" by Heil (1964). It may have colonized Messel lake after Lutetiospongilla had replaced Ephydatia as the dominant sponge species.[1]

Molluscs

Family Genus Species Authority Notes Image

Viviparidae

Viviparus

Undescribed[2]

A freshwater snail found in certain layers of the Messel formation.

Crustaceans

Family Genus Species Authority Notes Image

Daphniidae

Ctenodaphnia

Undescribed[3]

A water flea found in the oldest layers of the Messel formation.
It is almost identical to the living Ctenodaphnia magna.

Daphniidae

Daphnia

Unidentified[3]

A water flea originally thought to have had ephippia containing only one egg, this was later proven to be false.
It is almost identical to the living Daphnia pulex.

Moinidae

Moina

Undescribed[3]

A water flea reported by Lutz from ephippia containing two eggs.
They resemble the living Moina macrocopa.

incertae sedis

incertae sedis

incertae sedis[2]

An undescribed freshwater shrimp discovered in 2005.

Arachnids

Araneae

Family Genus Species Authority Notes Image
Araneidae[4] Undescribed Undescribed An orb-weaver spider first mentioned by Wunderlich in 1986.
Hersiliidae[5] Undescribed Undescribed A well preserved tree trunk spider.
Cybaeidae Lutetiana Lutetiana neli[6] Selden & Wappler, 2019 A possible cybaeide spider and the first arachnid named from Messel. It was a ground spider that shares some traits with the Lycosidae, however assignment to this family is unlikely as wolf spiders are hypothesized to have only become groundwellers following the spread of grassland during the Miocene.

Opiliones

Family Genus Species Authority Notes Image
Undescribed Undescribed[6][5] Six harvestmen awaiting description.

Insects

Coleoptera

Family Genus Species Authority Notes Image
Cupedidae Cupes Cupes nabozhenkoi[7]

Cupes wedmannae[7]

Kirejtshuk, 2020

Kirejtshuk, 2020

Elateridae Macropunctum Macropunctum angulosum[8]

Macropunctum angustiscutellurn[9]

Macropuncturn latiscutellurn[9]

Macropunctum messelense[9]

Macropunctum minuturn[9]

Macropunctum rebugense[9]

Macropunctum senckenbergi[9]

Tröster, 1999

Tröster, 1994

Tröster, 1994

Tröster, 1991

(Meunier, 1921)

Tröster, 1994

Tröster, 1994

A genus of click beetle.
Lucanidae Protognathinus Protognathinus spielbergi[10] Chalumeau et al. 2001 A 55 mm (2.2 in) long stag beetle that still preserves the colors of its exoskeleton.
Psephenidae Unnamed Unnamed[11] An eubrianacine water-penny beetle known from larval remains.

Dictyopterans

Family Genus Species Authority Notes Image
Blaberidae Morphna Morphna cenozoica[12] Šmídová, Vidlička & Wedmann, 2021 An extinct relative of modern giant cockroaches of the family Blaberidae.

Dipterans

Family Genus Species Authority Notes Image
Chaoboridae Chaoborus Chaoborus sp.[13] A genus of midget known from its aquatic larval form.
Bombyliidae Comptosia Comptosia pria[14] Wedmann & Yeates, 2008 A genus of bee fly.
Nemestrinidae Hirmoneura Hirmoneura messelense[15] Wedmann et al., 2021 A genus of tangle-veined fly found to have fed on pollen.
Bibionidae Plecia Plecia acourti[16]

Plecia hoffeinsorum[16]

Unnamed species [16]

Cockerell, 1921

Skartveit, 2009

A genus of march fly.

Hemiptera

Family Genus Species Authority Notes Image
Eurybrachidae Amalaberga Amalaberga ostrogothiorum[17] Szwedo & Wappler, 2006 An eurybrachid planthopper
Lophopidae Baninus Baninus thuringiorum[17] Szwedo & Wappler, 2006 A lophopid planthopper
Aradidae Aneurus? Aneurus? incertus[18] Wappler, Heiss & Wedmann, 2015 A species of flat bug.
Tingidae Chorotingiotes Chorotingiotes prisca[19] Wappler, 2003 A ypsotingine tingine lacebug
Pentatomidae Eospinosus Eospinosus peterkulkai[20] Wedmann et al., 2021 A species of shield bug exhibiting prominent spiky protrustions possibly used in defense or camouflage.
Tingidae Exmesselensis Exmesselensis disspinosus[19] Wappler, 2003 A phatnomine canatacaderine lacebug
Tingidae Lutetiacader Lutetiacadeir petrefactus[21] Wappler, 2006 A cantacaderine canatacaderine lacebug
Aradidae Mezira Mezira parapetrificata[18]

Mezira petrificata[18]

Wappler, Heiss & Wedmann, 2015 A species of flat bug.
Tingidae Oblongomorpha Oblongomorpha lutetia[19] Wappler, 2003 A litadeine tingine lacebug
Dictyopharidae Wedelphus Wedelphus dichopteroides[17] Szwedo & Wappler, 2006 A dictyopharid planthopper

Hymenoptera

Family Genus Species Authority Notes Image
Formicidae Archimyrmex Archimyrmex wedmannae[22] Dlussky, 2012 A myrmeciine ant with long mandibles.
Archimyrmex wedmannae Smfmei2016 01.jpg
Formicidae Casaleia Casaleia eocenica[23] An amblyoponine ant.
Casaleia eocenica SMFMEI5565.jpg
Formicidae Cephalopone Cephalopone grandis[23]

Cephalopone potens[23]

Poneromorph ants
Cephalopone grandis holotype SMFMEI5361.jpg
Formicidae Cyrtopone Cyrtopone curiosa[23]

Cyrtopone elongata[23]

Cyrtopone microcephala[23]

Cyrtopone striata[23]

Ponerinae subfamily ants
Cyrtopone striata holotype SMFMEI1392.jpg
Formicidae Gesomyrmex Gesomyrmex pulcher[24] A formicine subfamily ant
Gesomyrmex pulcher SMFMEI10999.jpg
Mesornatus Mesornatus markovici[25]
Formicidae Messelepone Messelepone leptogenoides[23] A poneromorph ant.
Messelepone leptogenoides holotype SMFMEI4808.jpg
Formicidae Pachycondyla Pachycondyla eocenica[23]

Pachycondyla lutzi[23]

Pachycondyla? messeliana[23]

Pachycondyla parvula[23][26]

Pachycondyla petiolosa[23]

Pachycondyla petrosa[23]

Pachycondyla lutzi SMFMEI11958.jpg
Polyhelictes Polyhelictes bipolarus[25]
Formicidae Protopone Protopone? dubia[23]

Protopone germanica[23]

Protopone magna[23]

Protopone oculata[23]

Protopone sepulta[23]

Protopone vetula[23]

Ponerine subfamily ants
Protopone magna holotype SMFMEI8790.jpg
Formicidae Pseudectatomma Pseudectatomma eocenica[23]

Pseudectatomma striatula[23]

Ectatommine subfamily ants
Pseudectatomma eocenica SMFMEI400.jpg
Rhyssella Rhyssella vera[25]
Scambus Scambus fossilobus[25]
Formicidae Titanomyrma Titanomyrma gigantea[27]

Titanomyrma simillima[27]

Formiciine subfamily ants
Titanomyrma simillima SMFMEI01006 01.jpg
Trigonator Trigonator macrocheirus[25]
Vespula? Vespula? hassiaca[28]
Xanthopimpla Xanthopimpla messelensis[25]

Xanthopimpla praeclara[25]

File:Xanthopimpla messelensis sp. nov., holotype SF MeI 16988.tiff
Xoanon? Xoanon? eocenicus[29]

Neuroptera

Family Genus Species Authority Notes Image

Mantispidae

Symphrasites

Symphrasites eocenicus[30]

Wedman & Makarkin, 2007

A symphrasine mantidfly
Described from a isolated forewing

Symphrasites eocenicus holotype fig2A.png

Odonata

Family Genus Species Authority Notes Image

†Dysagrionidae

Petrolestes

Petrolestes messelensis[31]

Garrouste & Nel, 2015

A damselfly
Related to the North American P. hendersoni.

center|frameless

†Pseudostenolestidae

Pseudostenolestes

Pseudostenolestes bechlyi[31]

Garrouste & Nel, 2015

A damselfly species.

Phasmatodea

Family Genus Species Authority Notes Image

Phasmatodea

Eophyllium

Eophyllium messelensis[32]

Wedmann, Bradler & Rust, 2007

A Phylliidae leaf insect,
The camouflage adaptions of modern Phylliids were already present in E. messelensis.

"Fish"

Amiiformes

Family Genus Species Authority Notes Image
Amiidae Cyclurus Cyclurus kehreri[33] (Andreae, 1893) An amiine bowfin relative known to have performed cannibalism. Cyclurus kehreri

Anguilliformes

Family Genus Species Authority Notes Image
Anguillidae Anguilla Anguilla ignota[2] Micklich, 1985 The presence of Anguilla ignota indicates that the lake was likely connected to a larger drainage system connecting to the ocean.

Lepisosteiformes

Family Genus Species Authority Notes Image
Lepisosteidae Atractosteus Atractosteus messelensis[34]

Atractosteus strausi[35]

Grande, 2010

(Kinkelin, 1884)

Two species of gars in the extant genus Actractosteus. Atractosteus strausi - Huston Museum Nat Sci - DSC01960.jpg
Lepisosteidae Masillosteus Masillosteus kelleri[36] Micklich & Klappert, 2001 A rarer, blunt snouted species of gar that likely fed on small invertebrates. It may not have been a permanent resident of the Messel lake.[2]

Perciformes

Family Genus Species Authority Notes Image
Percichthydae[37] Amphiperca Amphiperca multiformis[38] Weitzel, 1933 Amphiperca likely was a thrust or ambush hunter catching prey from near-shore hiding spots. Amphiperca multiformis 01.jpg
Moronidae Palaeoperca Palaeoperca proxima[39] Micklich, 1978 Palaeoperca may have been open water fish. Palaeoperca proxima.jpg
incertae sedis Rhenanoperca Rhenanoperca minuta[40] Gaudant & Micklich, 1990 An small perciform with teeth suited for a durophagous diet. Fossils show they at least occasionally fed on smaller members of their species.[2]

Thaumaturidae

Family Genus Species Authority Notes Image
Thaumaturidae Thaumaturus Thaumaturus intermedius[41] Weitzel, 1933[38] Thaumaturus is only known from juvenile specimens, ranging in length from 3–6 cm (1.2–2.4 in).[2] It's relationship with other groups of fish remains unknown. Thaumaturus intermedius.jpg

Amphibians

Caudata

Family Genus Species Authority Notes Image
Salamandridae Chelotriton Chelotriton robustus[42]

Chelotriton sp.[43]

Westphal, 1980

Undescribed

A salamander, possibly related to modern ribbed newts and crocodile newts.

Anura

Family Genus Species Authority Notes Image
Pelobatidae Eopelobates Eopelobates wagneri[44] Weitzel, 1938 A relative to the modern spadefoot toad, it was likely a primarily terrestrial animal that entered the lake to spawn, as shown by preserved eggs. Eopelobates wagneri 2.jpg
Incertae sedis Lutetiobatrachus Lutetiobatrachus gracilis[45] Wuttke, 1998 Eventhough the name was coined in 1988, it was long deemed a nomen nudum until its redescription in 2012.
Palaeobatrachidae "Messelobatrachus" "Messelobatrachus tobieni"[46] Nomen nudum Although referred to as Messelobatrachus in literature, the name is a nomen nudum. One of the fossils preserves what is thought to be the tadpole of "Messelobatrachus". Messelobatrachus tobieni 2.jpg

Reptiles

Crocodyliformes

Family Genus Species Authority Notes Image
Tomistominae Tomistominae indet. Tomistominae indet.[47]
Crocodyloidea "Asiatosuchus" "Asiatosuchus" germanicus[48] Berg, 1966 A large and well known species of the paraphyletic genus Asiatosuchus. Asiatosuchus germanicus - skull.jpg
Bergisuchidae Bergisuchus Bergisuchus dietrichbergi[49] Kuhn, 1968 Member of the terrestrial Sebecosuchians with ziphodont teeth. Bergisuchus dietrichbergi - mandibula fragment.jpg
Planocraniidae Boverisuchus Boverisuchus magnifrons[50] Kuhn, 1938 A terrestrial Crocodilian previously assigned to the now dubious Pristichampsus Boverisuchus magnifrons.jpg
Alligatoroidea Diplocynodon Diplocynodon darwini

Diplocynodon deponaie[51]

(Ludvig, 1877)

(Frey, Laemmert & Riess, 1987)

The two species can be differentiated by the presence of well-developed osteoderms on the tail of Diplocynodon deponaie. 2011-07-09 gasometer 37.JPG
Alligatoridae Hassiacosuchus Hassiacosuchus haupti [52] Weitzel, 1935 A small species of alligatorid, some consider it a species of Allognathosuchus. Christopher Brochu argues for the continued use of Hassiacosuchus however.[53] Hassiacosuchus haupti.jpg

Squamata

Clade Genus Species Authority Notes Image
Lacertidae relative Undescribed Undescribed [54] An undescribed arboreal lizard relative.
Gekkota Undescribed Undescribed [54] A partly disarticulated skeleton of a gekko
Pan-Shinisaurus Pan Shinisaurus indet. Pan Shinisaurus indet.[55] The shed tail of a crocodile lizard, showing adaptation for swimming.
Polychrotidae Undescribed Undescribed [54] A possible relative of the modern bush anole.
Scincidae Scincidae indet. Scincidae indet.[2]
Lacertibaenia Cryptolacerta Crytolacerta hassiaca[56] Müller et al., 2011 Phylogenetic analysis indicates that it may be a close relative of the limbless Amphisbaenia. Only two specimens are known, one of which was found in the stomach of the varanoid Paranecrosaurus
Boidae Eoconstrictor Eoconstrictor fischeri[57] (Schaal, 2004) An early species of boa previously known as Palaeopython fischeri.
Palaeopython fischeri - Naturmuseum Senckenberg - DSC02249.JPG
Eolacertidae Eolacerta Eolacerta robusta[58] Nöth, 1940 A large species of lizard, reaching a length of 30 cm (12 in) and a weight of 1 kg (2.2 lb).
Helodermatidae Eurheloderma Eurheloderma sp.[59] A relative of the extant gila monster, it was likely already venomous.
Corytophanidae Geiseltaliellus Geiseltaliellus maarius[60] Smith, 2009 An extinct, possibly arboreal, relative of the modern basilisk lizard. Geiseltaliellus-Geiseltal.jpg
Boidae Messelophis Messelophis variatus[61] Baszio, 2004 A minute boa
Messelopythonidae Messelopython Messelopython freyi[62] Zaher & Smith, 2020 A medium sized pythonoid.
Anguidae Ophisauriscus Ophisauriscus quadrupes[63] Kuhn, 1940 A serpentine reptile with vestigial limbs. It shows evidence of different osteoderm morphology after tail regeneration.
Scincoidea Ornatocephalus Ornatocephalus metzleri[64] Weber, 2004 An arboreal lizard with long claws and a prehensile tail, it was found with plants remains in its guts.
Messelopythonidae Palaeopython Palaeopython schaali[65] Smith & Scanferla, 2022 A pythonoid snake of similar size to Eoconstrictor. Several traits of the skull suggest it may have been arboreal.
Palaeovaranidae Palaeovaranus Palaeovaranus sp.[54] Identified as Necrosaurus by Smith, Čerňanský and Scanferla, it is smaller than Paranecrosaurus.
Palaeovaranidae Paranecrosaurus Paranecrosaurus feisti[66] (Stritzke, 1983) First described as "Saniwa" feisti, it was found with the skull of Cryptolacerta in its guts. This indicates that this varanoid was at least partly carnivorous. Paranecrosaurus feisti - Milan img1.jpg
Glyptosaurinae Placosauriops "Placosauriops abderhaldeni"[67] Kuhn, 1940 A type of melanosaurinin Glyptosaurine of dubious assignment, as the type material has not been adequately diagnosed.
Erycinae Rageryx Rageryx schmidi[68] Smith & Scanferla, 2021 A non-burrowing sandboa that reached a length of 52 cm (20 in).
Boidae Rieppelophis Rieppelophis ermannorum[61] (Schaal & Baszio, 2004) A minute boa previously regarded as a species of Messelophis. center
Eolacertidae Stefanikia Stefanikia siderea[69] Čerňanský & Smith, 2017 A close relative of Eolacerta, the two genera were recovered as a sister-group to modern lizards.

Testudinata

Family Genus Species Authority Notes Image
Carettochelyidae Allaeochelys Allaeochelys crassesculptata[70] (Harrassowitz, 1922) A relative of the modern pig-nosed turtle, one fossil preserves two specimens while mating. Allaeochelys crassesculptata - Naturmuseum Freiburg - DSC06737.jpg
Geoemydidae Euroemys Euroemys kehreri[71] Staesche, 1928 A species of pond turtles of uncertain relationship, it was previously thought to be a species of Ocadia and Palaeoemys.[72] Euroemys kehreri 01.jpg
Geoemydidae Francellia Francellia messeliana[71] Staesche, 1928 A species of pond turtles of uncertain relationship, it was previously thought to be a species of Ocadia and Palaeoemys.[72]
Geoemydidae Juvemys Juvemys sp.[2] Hervet, 2003 A species of pond turtles.
Podocnemididae Neochelys Neochelys franzeni[73] Schleich, 1993 [74] A podocnemidid side-necked turtle Neochelys franzeni 2.jpg
Trionychidae Palaeoamyda Palaeoamyda messeliana [75] (Reinach, 1900) A species of softshell turtle Trionyx sp 354.jpg

Birds

Anseriformes

Family Genus Species Authority Notes Image
Gastornithidae Gastornis Gastornis geiselensis[76] Fischer, 1978 A large, herbivorous anseriform.[77]
Gastornis geiselensis.jpg

Cariamiformes (?)

Several groups of Messel birds share characteristics with the modern seriemas, which has lead to them being placed within the clade Cariamae in the past. However, this placement typically occured under the assumption that they are a group within gruiformes, which has been disputed by more recent analysis. Instead more recent publications consider Cariamae (or Cariamiformes) as basal members of Australaves.[78]

Family Genus Species Authority Notes Image
Idiornithidae Dynamopterus Dynamopterus cf. itardiensis[79]

Dynamopterus tuberculata[80]

(Mourer-Chauviré, 1983)

(Peters, 1995)

Previously known as Idiornis, it was later found to be synonymous with Dynamopterus. Idiornis tuberculata
Salmilidae Salmila Salmila robusta[81] Mayr, 2000 Salmila was originally described as a part of Cariamae within Gruiformes, but later analysis suggested that it was a sister taxon to the group instead. It shares characteristics with both seriemas and trumpeter birds. Salmila robusta
Ameghinornithidae Strigogyps Strigogyps sapea[82] (Peters, 1987) Previously known as Aenigmavis, stomach contents indicate it was a herbivore. The phylogenetic position of Strigogyps is uncertain. Strigogyps sapea

Charadriiformes

Family Genus Species Authority Notes Image
Jacanidae? Vanolimicola Vanolimicola longihallucis[83] Mayr, 2017 A small wading bird with elongated hallux. It is thought to either be a jacanid or a rail-relative of the family Songziidae, which would make it a Gruiform.

Coraciimorphae

Family Genus Species Authority Notes Image
Sandcoleidae Chascacocolius Chascacocolius cacicirostris[84] Mayr, 2005 A coliiform bird with a conical beak.
Eocoraciidae Eocoracias Eocoracias brachyptera[85] Mayr, 2000 An early species of roller, analysis of the plumage gave indication as to what its colours would have been in life.[86]
Eocoracias brachyptera.jpg
Sandcoleidae Eoglaucidium Eoglaucidium pallas [87]

Eoglaucidium sp. indet.[88]

Mayr & Peters, 1998 Originally described as an owl from the Geiseltal, it was later found to be related to mousebirds. It's tail is notably shorter than that of its extant relatives. It is possible that the indetermined species instead belongs to Anneavis.
Eoglaucidium sp.jpg
Gracilitarsidae Gracilitarsus Gracilitarsus mirabilis[89] Mayr, 1998 The shape of the beak indicates that Gracilitarsus was either an insectivore or fed on nectar.
Coliidae Masillacolius Masillacolius brevidactylus[87] Mayr & Peters, 1998 An extinct species of mousebird.
Trogonidae Masillatrogon Masillatrogon pumilio[90] Mayr, 2009 An extinct species of trogon that supports the hypothesis that the family originated in the Old World.
Dans l'ombre des dinosaures - Masillatrogon - 018.jpg
Messelirrisoridae Messelirrisor Messelirrisor grandis[91]

Messelirrisor halycrostris[92]

Messelirrisor parvus[92]

Mayr, 2000

Mayr, 1998

Mayr, 1998

Relatives of the modern hoopoe and wood hoopoe, they were birds with long, pointed beaks and diminutive body size. The tail feathers of one specimen show they were barred, preserving a pattern of dark and light stripes.
Messelirrisor halcyrostris.jpg
Leptosomidae? Plesiocathartes Plesiocathartes kelleri[93] Mayr, 2002 A possible relative of the modern cuckoo-roller.
Primobucconidae Primobucco Primobucco frugilegus[94]

Primobucco perneri

Mayr, Mourer-Chaviré & Weidig, 2004 The first recorded species of Primobucco in Europe.
Primobucco.jpg
Incertae sedis Quasisyndactylus Quasisyndactylus longibrachis[95] Mayr, 1998 Quasisyndactylus is thought to be a relative of modern kingfishers, todies and motmots.
Quasisyndactylus longibrachis 1.jpg
Coliidae [88] Selmes Selmes absurdipes [96] Peters, 1999 A genus of mousebird.

Cuculiformes

Family Genus Species Authority Notes Image
Foratidae? Unnamed Unnamed Known from an isolated skull similar to the larger Foro panarium.

Falconiformes

Family Genus Species Authority Notes Image
Masillaraptoridae Masillaraptor Masillaraptor parvunguis[97][98] Mayr, 2006 A long-legged, possibly more terrestrial bird. It shares several traits with modern falcons, but may be a more basal member of Falconiformes.

Galliformes

Clade Genus Species Authority Notes Image
Gallinuloididae Paraortygoides Paraortygoides messelensis[99] Mayr, 2000 One of the earliest known galliforms, the lack of a crop indicates they fed on only soft plant material.

Gruimorphae

Family Genus Species Authority Notes Image
Messelornithidae Messelornis Messelornis cristata[100] Hesse, 1988 The most common fossil bird found at Messel, it was likely a terrestrial animal capable of only short bursts of flight due to its short wings. It may have had a comb or crest atop its head.
Messelornis sp 563.JPG

Incertae sedis

Family Genus Species Authority Notes Image
Eopachypterygidae Eopachypteryx Eopachypteryx praeterita

Eopachypteryx ? sp.[101]

Mayr, 2015
Incertae sedis Lapillavis Lapillavis incubarens[102] Mayr, 2016
Incertae sedis Perplexicervix Perplexicervix microcephalon[103] Mayr, 2010 The genus is named for the bony tubercles present on the cervical vertebrae. Although its relationship is not fully understood, it was found to not be a member of Idiornithidae and shared traits with Anhimidae and Cathartidae.

Mirandornithes

Family Genus Species Authority Notes Image
Juncitarsus Juncitarsus merkeli[104] Olson & Feduccia, 1980 A wading bird thought to be basal to both flamingos and grebes. Both the beak shape and the presence of gastroliths show that it was not yet adapted to filderfeed and instead fed on hard items.
Juncitarsus merkeli 1.jpg

Palaeognathae

Family Genus Species Authority Notes Image
Lithornithidae Lithornis Lithornis sp.[88][105] Its size was intermediate between Lithornis plebius and Lithornis hookeri. Additionally, it represents the last known lithornithid in the fossil record.
Incertae sedis Palaeotis Palaeotis weigelti[106] Lambrecht, 1928 A flightless palaeognath, previous analysis considered it to be a relative to ostriches or rheas. More recent research however doubts this. Palaeotis weigelti 2.jpg

Pelecaniformes

Family Genus Species Authority Notes Image
Threskiornithidae Rhynchaeites Rhynchaeites messelensis[107] Wittich, 1898 Also known as the "snipe-rail" for its similarities to rails and painted snipes, it was actually a type of ibis.
Rhynchaeites messelensis 28.jpg

Psittacopasserae

Family Genus Species Authority Notes Image
Messelasturidae Messelastur Messelastur gratulator Peters, 1994 Messelastur is a bird with a poorly understood ecology. Although shown to be closely related to Tynsyka, its relationship to other groups is not fully understood. Older publications suggest it may have been related to parrots or various birds of prey (including owls, hawks and falcons). One recent publication positions them at the base of Psittacopasserae alongside the Pseudasturidae (=Halcyornithidae).[108]
Messelastur gratulator (cropped).jpg
Quercypsittidae? Palaeopsittacus Palaeopsittacus cf. georgei[109] Harrison, 1982 Although initially assigned to Psittaciformes, later research instead suggested the idea that it was a type of frogmouth instead.
Zygodactylidae Primozygodactylus Primozygodactylus ballmanni[110]

Primozygodactylus danielsi[110]

Primozygodactylus eunjooae[111]

Primozygodactylus longibrachium[110]

Primozygodactylus mayor[110]

Primozygodactylus quintus[110]

Mayr, 1998

Mayr, 1998

Mayr and Zelenkov, 2009

Mayr, 2016

Mayr, 1998

Zygodactyl birds originally thought to be relatives of woodpeckers, later research placed them as a sister group to the Passeriformes. Diet varries between species, with the type species likely being an insectivore while the larger Primozygodactylus mayor was found with grapes in its stomach. The wings were short and the legs long, indicating a maneuverable animal foraging in the undergrowth. At least in some species the central two tail feathers were greatly elongated and possibly used in display.
Primozygodactylus eunjooae.jpg
Pseudasturidae Pseudorasturidae indet. Pseudorasturidae indet.[79] A poorly preserved pseudasturid of smaller size than Serudaptus. It may be a different species of the same genus, a juvenile, a member of the opposite sex or a unique genus.
Psittacopedidae Psittacopes Psittacopes lepidus[112] Mayr & Daniels, 1998 A bird originally classified as a parrot, later research indicated that it may instead be closer to passeriforms.
Psittacopedidae[88] Pumiliornis Pumiliornis tessellatus[113] Mayr, 1999 A bird the size of a small wren.
Pseudasturidae Serudaptus Serudaptus pohli[79] Mayr, 2000 A bird with zygodactyl feet and long claws suited for climbing. It's exact phylogenetic position is poorly understood, with some research indicating it may be a basal member of the clade containing Passeriformes and Psittaciformes.
Serudaptus NHMW.jpg

Strigiformes

Family Genus Species Authority Notes Image
Palaeoglaucidae Palaeoglaux Palaeoglaux artophoron[114] Peters, 1992 A small owl preserving ribbon-like feathers arranged in a dense layer on its back.
Palaeoglaux artophoron.JPG

Strisores

Family Genus Species Authority Notes Image
Cypseloramphus Cypseloramphus dimidius[102] Mayr, 2016 A tentative apodiform.
Archaeotrogonidae Hassiavis Hassiavis laticauda[95] Mayr, 1998 The skull of Hassiavis closely resembles that of modern owlet-nightjars, being broad and short.
Hassiavis laticauda 1.jpg
Masillapodargus Masillapodargus longipes[115] Mayr, 2001 A relative of the modern frogmouths.
Jungornithidae? Parargornis Parargornis messelensis[116] Mayr, 2003 A swift-like bird with short wings and long tail feathers, phylogenetic analysis indicate that it was a basal type of hummingbird.[117]
Parargornis messelensis.JPG
Nyctibiidae Paraprefica Paraprefica kelleri[118]

Paraprefica major[115]

Mayr, 1999 An old-world genus of potoo. The two species can clearly be differentiated by size.
Paraprefica kelleri.jpg
Protocypselomorphus Protocypselomorphus manfredkelleri[119] Mayr, 2005 An aerial insectivore with reduced feet, it was recovered as a sister-taxon to all other Strisores.
Pan-Apodidae Scaniacypselus Scaniacypselus szarskii[120] (Peters, 1985) An early genus of swift, its reduced legs and less developed wings indicate that it wasn't as aerial as its modern relatives and likely nested in trees.
Dans l'ombre des dinosaures - Scaniacypselus - 02.jpg

Suliformes

Family Genus Species Authority Notes Image
Sulidae? Masillastega Masillastega rectirostris[121] Mayr, 2002 A bird tentatively referred to Sulidae, it was approximately the size of a modern European shag. It was likely piscivorous.
Masillastega rectirostris holotype sp140b.jpg

Mammals

Apatotheria

Family Genus Species Authority Notes Image
Apatemyidae Heterohyus Heterohyus nanus[122] Teilhard de Chardin, 1921 A mammal with elongated fingers and strong teeth that resembles the modern aye-aye and likely filled a similar niche.[2]
Heterohyus 4939.jpg

Artiodactyla

Family Genus Species Authority Notes Image
Dichobunidae Aumelasia Aumelasia cf. gabineaudi[123] Sudré, 1980 A lightly built early artiodactyl with a noticably long tail. Aumelasia is referred to the Diacodexeidae by some researchers.[37]
Aumelasia sp.jpg
Dichobunidae Messelobunodon Messelobunodon schaeferi[124] Franzen, 1980 An early species of even-toed ungulate.
Messelobunodon schaeferi franzen.jpg
Choeropotamidae Masillabune Masillabune martini[125] Tobien, 1980 An early species of even-toed ungulate.
Masillabune Geiseltal.jpg

Chiroptera

Family Genus Species Authority Notes Image
Archaeonycteridae Archaeonycteris Archaeonycteris pollex[126]

Archaeonycteris trigonodon[127]

Storch & Habersetzer, 1988

Revilliod, 1917

A basal genus among the Messel bats, Archaeonycteris resembles modern vesper bats in wing morphology and likely preferred more open spaces in the upper trunk areas to hunt. Stomach contents indicate a preference for beetles.[2]
Archaeonycteris trigonodon
Hassianycterididae Hassianycteris Hassianycteris magna[128]

Hassianycteris messelensis[128]

Hassianycteris? revilliodi[37]

Smith & Storch, 1981

Smith & Storch, 1981

(Russell & Sigé, 1970)

Hassianycteris was a somewhat heavy bat with narrow wings. Based on its anatomy, which resembles modern free-tailed bats, it was a fast flying animal hunting above the treetops.[2]
Archaeonycteris trigonodon
Palaeochiropterygidae Palaeochiropteryx Palaeochiropteryx spiegeli[37]

Palaeochiropteryx tupaiodon[37]

Revilliod, 1917 The wing morphology of Palaeochiropteryx indicates that it flew slowly but agile relatively close to the ground to catch insects such as moths and butterflies.[2]
Palaeochiropteryx Muséum Grenoble 03082017.jpg
Emballonuridae Tachypteron Tachypteron franzeni[129] Storch, Sigé & Habersetzer, 2002 A fast flying species that inhabited open airspaces, Tachypteron resembled the modern black-bearded tomb bat in proportions.[2]

Chimolesta

Family Genus Species Authority Notes Image
Pantolestidae Buxolestes Buxolestes piscator

Buxolestes minor[130]

Koenigswald 1980

Pfretzschner 1999

Buxolestes is a member of a basal group of mammals. The proportions of Buxolestes piscator are similar to modern otters and its stomach contents indicate a diet of fish and small vertebrates. B. minor shows adaptations towards digging and was found with plant material in its stomach, but likely also fed on various animals.
Buxolestes.jpg
Paroxyclaenidae Kopidodon Kopidodon macrognathus[2] (Wittich, 1902) Kopidodon was the first mammal discovered in Messel and grew to a length of 115 cm (45 in). It shows some adaptations towards a climbing lifestyle.
Kopidodon macrognathus - National Museum of Nature and Science, Tokyo - DSC07781.JPG
Paroxyclaenidae "Paroxyclaenus" "Paroxyclaenus" sp.[37] An undescribed relative of Kopidodon differing through the lack of a pronounced sagittal crest.

Eulipotyphla

Family Genus Species Authority Notes Image
Amphilemuridae Macrocranion Macrocranion tenerum[131]

Macrocranion tupaiodon[132]

Tobien, 1962

Weitzel, 1949

Macrocranion were small, likely nocturnal, animals living on the forest floor. M. tupaiodon reached a size of up to 30 cm (12 in) and was possibly omnivorous with woolly fur.[2] The smaller M. tenerum meanwhile was inferred to be insectivorous and coverd in bristles. Some researchers suggest it may have moved like modern springhares.[2][133]
Macrocranion tupaiodon 2.jpg
Amphilemuridae Pholidocercus Pholidocercus hassiacus[134] von Koenigswald & Storch, 1983 A larger relative of Macrocranion, Pholidocercus was an animal 40 cm (16 in). It's proportions indicate that it was a quadrupedal animal that lived on the forest floor. The tail was covered in overlapping scales and the head was similarily covered in a hardened structure while the torso was covered in thick bristles.[2]
Pholidocercus hassiacus, hedgehog, Late Early Eocene, Darmstadt, Germany - Houston Museum of Natural Science - DSC01950.JPG

Leptictida

Family Genus Species Authority Notes Image
Pseudorhyncocyonidae Leptictidium Leptictidium auderiense[37]

Leptictidium nasutum[135]

Leptictidium tobieni[136]

Tobien, 1962

Storch & Lister, 1985

Von Koenigswald & Storch, 1987

Known from Messel in the form of three species and multiple specimens, Leptictidium was a eutherian mammal of uncertain relationships. It's hindlimbs are noticably larger than its forelimbs, but its means of locomotion are not agreed upon.
Leptictidium auderiense.JPG

Metatheria

Family Genus Species Authority Notes Image
Herpetotheriidae Amphiperatherium Amphiperatherium goethei[137]

Amphiperatherium cf. maximum[138]

Crochet, 1979 A metatherian mammal, it was larger than Peradectes but with a proportionally shorter tail.
Amphiperatherium.JPG
Herpetotheriidae Herpetotheriidae indet. Either Amphiperatherium or Peratherium.[37]
Peradectidae "Peradectes" "Peradectes sp."[138] A small arboreal mammal similar to Peradectes with a prehensile tail.
Peradectes sp 1.jpg

Pan-Carnivora

Family Genus Species Authority Notes Image
Proviverridae Lesmesodon Lesmesodon behnkeae[139]

Lesmesodon edingeri[140]

(Morlo & Habersetzer, 1999)

(Springhorn, 1982)

Agile predators of the forest floor, at least one species (L. behnkeae) is known to have had a bushy tail. The exact position hyaenodontids like Lesmesodon have within mammals is debated.
Lesmesodon edingeri.jpg
Miacidae Messelogale Messelogale kessleri[141] (Springhorn, 1982)[140] Originally thought to be a species of Miacis, it was elevated to its own genus in 2000. Unlike Lesmesodon it was an arboreal hunter.
Miacidae Paroodectes Paroodectes feisti[142] Springhorn, 1980 An arboreal miacid.
Paroodectes feisti.JPG

Pholidota

Family Genus Species Authority Notes Image
Eomanidae[143] Eomanis Eomanis waldi Storch, 1978 Eomanis is a more derived member of Pholidota than the contemporary genera found in Messel and already shows the classic scales this group is known for.[144]
Eomanis waldi.jpg
Euromanis Euromanis krebsi[145] (Storch & Martin, 1994) A type of scaleless pangolin originally described as a species of Eomanis.
Euromanis krebsi.jpg
Eurotamanduidae Eurotamandua Eurotamandua joresi[146] Storch, 1981 Described as an anteater based on the enlarged claws and elongated skull, later research concluded that it was more likely to be a type of pangolin.
Eurotamandua joresi 1.jpg

Perissodactyla

Family Genus Species Authority Notes Image
Palaeotheriidae Eurohippus Eurohippus parvulus[147]

Eurohippus messelensis[148]

(Laurillard, 1849)

(Haupt, 1925)

Originally species of Propalaeotherium and Lophiotherium, they were eleveted to a distinct genus in 2006. Several specimens were preserved with their fetus, showing that they only carried a single foal.
Eurohippus with fetus.PNG
Hallensia Hallensia matthesi[149] Franzen & Haubold, 1986 A small species of primitive horse.
Hallensia matthesi.jpg
Hyrachyidae Hyrachyus Hyrachyus minimus [150] Fischer, 1829 An early rhinoceratoid or tapir, its discovery played a rolle in making Messel a protected site.
Hyrachyus minimus.jpg
Palaeotheriidae Propalaeotherium Propalaeotherium hassiacum[151] Haupt, 1925 An early species of horse reaching a shoulder height of 55–60 cm (22–24 in), making it larger than Eurohippus.
Propalaeotherium hassiacum.jpg

Primates

Family Genus Species Authority Notes Image
Adapidae Darwinius Darwinius masillae[152] Franzen et al., 2009 An early primate known from a single specimen preserved in two slabs. The counterslab includes several elements fabricated to make it appear more complete. The classification of Darwinius has been subject to continued debate.
Darwinius masillae PMO 214.214.jpg
Adapidae Europolemur Europolemur kelleri[153]

Europolemur koenigswaldi[153]

Franzen, 2000

Franzen, 1987

An early primate primarily known from partial remains. One lower jaw was found in the coprolite of Buxolestes while another fossil was found with the tooth of a crocodile embeded into it.[2]
Europolemur koenigswaldi 843.jpg
Adapidae Godinotia Godinotia neglecta[154] (Thalmann et al., 1989) Preserved stomach contents of Godinotia suggest that it primarily fed on leaves and fruit.[155]
Godinotia neglecta GMH L-2.jpg

Rodentia

Family Genus Species Authority Notes Image
Alagomyidae[37] Ailuravus Ailuravus macrurus[156] Weitzel, 1949 A large arboreal rodent that resembles modern squirrels in proportion. It reached a total length of 1 m (3 ft 3 in), most of which consisting of the bushy tail, and primarily fed on leaves.[2]
Ailuravus.jpg
Gliridae Eogliravus Eogliravus wildi[157] Hartenberger, 1971 The oldest known dormouse, it had a long and bushy tail and fed on seeds and plant buds.
Masillamys Masillamys beegeri[158]

Masillamys krugi[158]

Masillamys parvus[158]

Tobien, 1954 Masillamys parvus is at times referred to the genus Hartenbergeromys,[2] but a 2019 study finds the differences to the type species insufficient to warrant this referal. It's tail was only sparsely covered in fur.[158]
Masillamys Senckenberg 2007-01.JPG

References

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