Paleofauna of the Messel Formation

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

Sponges edit

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.

 
Ephydatia gutenbergiana gemmule

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.

 
Lutetiospongilla heili Holotype
gemmules and large radial gemmuloscleres

"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 edit

Family Genus Species Authority Notes Image

Viviparidae

Viviparus

Undescribed[2]

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

Crustaceans edit

Family Genus Species Authority Notes Image
Daphniidae

Ctenodaphnia

Undescribed[3]

A water flea
present in the oldest Messel formation stata
Considered almost identical to the modern Ctenodaphnia magna.

Daphnia

Unidentified[3]

A Daphnia subgenus Daphnia water flea
First thought to have had single egged ephippia, but this was later proven to be false.
Notably similar 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.

? Palaemonidae Bechleja

Bechleja brevirostris[4]

de Mazancourt, Wappler & Wedmann, 2022

A palaemonid? freshwater shrimp
known from exceptionally preserved remains.

 
Bechleja brevirostri

Bechleja sp.[4]

A palaemonid? freshwater shrimp
A single fossil with a longer upcurved rostrum than B. brevirostris
not named pending more fossils found.[4]

incertae sedis

incertae sedis

incertae sedis[2]

An undescribed freshwater shrimp discovered in 2005.

Arachnids edit

Araneae edit

Family Genus Species Authority Notes Image

Araneidae

Undescribed

Undescribed[5]

An orb-weaver spider
first mentioned by Wunderlich (1986).

?Cybaeidae

Lutetiana

Lutetiana neli[6]

Selden & Wappler, 2019

A possible cybaeid ground spider
A Marronoidea based on leg length and claws
The first described Messel arachnid.

 
Lutetiana neli holotype

Hersiliidae

Undescribed

Undescribed[7]

A well preserved tree trunk spider.

Opiliones edit

Family Genus Species Authority Notes Image

Undescribed

Undescribed

Undescribed[6][7]

Six harvestmen awaiting description.

Insects edit

Coleoptera edit

Family Genus Species Authority Notes Image

Attelabidae

Palaeoalatorostrum

Palaeoalatorostrum schaali[8]

Rheinheimer, 2007

An Attelabine leaf-rolling weevil

Cupedidae Cupes

Cupes messelensis[9]

(Tröster, 1993)

A reticulated beetle species

Cupes nabozhenkoi[9]

Kirejtshuk, 2020

A reticulated beetle species

 
Cupes nabozhenkoi holotype

Cupes wedmannae[9]

Kirejtshuk, 2020

A reticulated beetle species

 
Cupes wedmannae holotype
Curculionidae Palaeocrassirhinus

Palaeocrassirhinus messelensis[8]

Rheinheimer, 2007

A Brachyderine weevil

Palaeocrassirhinus rugosithorax[8]

Rheinheimer, 2007

A Brachyderine weevil

Palaeocneorhinus

Palaeocneorhinus messelensis[8]

Rheinheimer, 2007

A Brachyderine weevil

Elateridae Macropunctum

Macropunctum angulosum[10]

Tröster, 1999

A click beetle species.

Macropunctum angustiscutellurn[11]

Tröster, 1994

A click beetle species.

Macropuncturn latiscutellurn[11]

Tröster, 1994

A click beetle species.

Macropunctum messelense[11]

Tröster, 1994

A click beetle species.

Macropunctum minuturn[11]

(Meunier, 1921)

A click beetle species.

Macropunctum rebugense[11]

Tröster, 1994

A click beetle species.

Macropunctum senckenbergi[11]

Tröster, 1994

A click beetle species.

Lucanidae

Protognathinus

Protognathinus spielbergi[12]

Chalumeau et al. 2001

A 55 mm (2.2 in) long stag beetle with preserved coloration.

Psephenidae

Unnamed

Unnamed[13]

An eubrianacine water-penny beetle
Known from larval remains.

Tenebrionidae

Ceropria?

Ceropria? messelense[14]

Hornschemeyer, 1994

A tenebrionine darkling beetle

Dictyopterans edit

Family Genus Species Authority Notes Image

Blaberidae

Morphna

Morphna cenozoica[15]

Šmídová, Vidlička & Wedmann, 2021

A Blaberid giant cockroach.

Dipterans edit

Family Genus Species Authority Notes Image

Bibionidae

Plecia

Plecia acourti[16]

Plecia hoffeinsorum[16]

Unnamed species [16]

Cockerell, 1921

Skartveit, 2009

Several species of march fly.

Bombyliidae

Comptosia

Comptosia pria[17]

Wedmann & Yeates, 2008

A species of bee fly.

Chaoboridae

Chaoborus

Unnamed[18]

Unnamed midges known from aquatic larval form.

Nemestrinidae

Hirmoneura

Hirmoneura messelense[19]

Wedmann et al., 2021

A pollen feeding species of tangle-veined fly.

Hemiptera edit

Family Genus Species Authority Notes Image
Aradidae

Aneurus?

Aneurus? incertus[20]

Wappler, Heiss & Wedmann, 2015

A species of flat bug.

Mezira

Mezira parapetrificata[20]

Mezira petrificata[20]

Wappler, Heiss & Wedmann, 2015

Two species of flat bug.

Dictyopharidae

Wedelphus

Wedelphus dichopteroides[21]

Szwedo & Wappler, 2006

A dictyopharid planthopper

Eurybrachidae

Amalaberga

Amalaberga ostrogothiorum[21]

Szwedo & Wappler, 2006

An eurybrachid planthopper

Lophopidae

Baninus

Baninus thuringiorum[21]

Szwedo & Wappler, 2006

A lophopid planthopper

Pentatomidae

Eospinosus

Eospinosus peterkulkai[22]

Wedmann et al., 2021

A species of shield bug exhibiting prominent spiky protrustions possibly used in defense or camouflage.

Tingidae

Chorotingiotes

Chorotingiotes prisca[23]

Wappler, 2003

A ypsotingine tingine lacebug

Exmesselensis

Exmesselensis disspinosus[23]

Wappler, 2003

A phatnomine canatacaderine lacebug

Lutetiacader

Lutetiacadeir petrefactus[24]

Wappler, 2006

A cantacaderine canatacaderine lacebug

Oblongomorpha

Oblongomorpha lutetia[23]

Wappler, 2003

A litadeine tingine lacebug

Hymenoptera edit

Family Genus Species Authority Notes Image
Apidae

Pygomelissa

Pygomelissa lutetia[25]

Wappler & Engel, 2003

A bee of uncertain tribal placement.

Protobombus

Protobombus messelensis[25]

Wappler & Engel, 2003

An electrapinae bee.

Formicidae

Archimyrmex

Archimyrmex wedmannae[26]

Dlussky, 2012

A myrmeciine ant with long mandibles.

 
Archimyrmex wedmannae
holotype

Casaleia

Casaleia eocenica[27]

Dlussky & Wedmann, 2012

An amblyoponine ant.

 
Casaleia eocenica
holotype
Cephalopone

Cephalopone grandis[27]

Dlussky & Wedmann, 2012

A ponerine Poneromorph ant

 
Cephalopone grandis
holotype

Cephalopone potens[27]

Dlussky & Wedmann, 2012

A ponerine Poneromorph ant

 
Cephalopone potens
holotype
Cyrtopone

Cyrtopone curiosa[27]

Dlussky & Wedmann, 2012

A ponerine ant

Cyrtopone elongata[27]

Dlussky & Wedmann, 2012

A ponerine ant

Cyrtopone microcephala[27]

Dlussky & Wedmann, 2012

A ponerine ant

Cyrtopone striata[27]

Dlussky & Wedmann, 2012

A ponerine ant

 
Cyrtopone striata
holotype

Gesomyrmex

Gesomyrmex pulcher[28]

Dlussky, Wappler, & Wedmann, 2009

A formicine subfamily ant

 
Gesomyrmex pulcher
holotype

Messelepone

Messelepone leptogenoides[27]

Dlussky & Wedmann, 2012

A poneromorph ant.

 
Messelepone leptogenoides
holotype
Pachycondyla

Pachycondyla eocenica[27]

Dlussky & Wedmann, 2012

A ponerine ant

Pachycondyla lutzi[27]

Dlussky & Wedmann, 2012

A ponerine ant

 
Pachycondyla lutzi
holotype

Pachycondyla? messeliana[27]

Dlussky & Wedmann, 2012

A ponerine ant

Pachycondyla parvula[27][29]

Dlussky et al., 2015

A ponerine ant

Pachycondyla petiolosa[27]

Dlussky & Wedmann, 2012

A ponerine ant

Pachycondyla petrosa[27]

Dlussky & Wedmann, 2012

A ponerine ant

Protopone

Protopone? dubia[27]

Dlussky & Wedmann, 2012

A ponerine ant

Protopone germanica[27]

Dlussky & Wedmann, 2012

A ponerine ant

Protopone magna[27]

Dlussky & Wedmann, 2012

A ponerine ant

 
Protopone magna
holotype

Protopone oculata[27]

Dlussky & Wedmann, 2012

A ponerine ant

Protopone sepulta[27]

Dlussky & Wedmann, 2012

A ponerine ant

Protopone vetula[27]

Dlussky & Wedmann, 2012

A ponerine ant

Pseudectatomma

Pseudectatomma eocenica[27]

Dlussky & Wedmann, 2012

An ectatommine ant

 
Pseudectatomma eocenica
holotype

Pseudectatomma striatula[27]

Dlussky & Wedmann, 2012

An ectatommine ant

Titanomyrma

Titanomyrma gigantea[30]

(Lutz, 1986)

A formiciine subfamily ant

 
Titanomyrma simillima
holotype

Titanomyrma simillima[30]

(Lutz, 1986)

A formiciine subfamily ant

 
Titanomyrma simillima
holotype
Ichneumonidae

Mesornatus

Mesornatus markovici[31]

Spasojevic, Wedmann & Klopfstein, 2018

An ichneumon wasp of uncertain subfamilial affinity

 
Mesornatus markovici
holotype

Polyhelictes

Polyhelictes bipolarus[31]

Spasojevic, Wedmann & Klopfstein, 2018

An ichneumon wasp of uncertain subfamilial affinity

 
Polyhelictes bipolarus
holotype

Rhyssella

Rhyssella vera[31]

Spasojevic, Wedmann & Klopfstein, 2018

A rhyssine ichneumon wasp

 
Rhyssella vera
holotype

Scambus

Scambus fossilobus[31]

Spasojevic, Wedmann & Klopfstein, 2018

An ephialtine pimplinae ichneumon wasp

 
Scambus fossilobus
holotype

Trigonator

Trigonator macrocheirus[31]

Spasojevic, Wedmann & Klopfstein, 2018

A labenine ichneumon wasp

 
Trigonator macrocheirus
holotype
Xanthopimpla

Xanthopimpla messelensis[31]

Spasojevic, Wedmann & Klopfstein, 2018

A pimpline pimplinae ichneumon wasp

 
Xanthopimpla messelensis
holotype

Xanthopimpla praeclara[31]

Spasojevic, Wedmann & Klopfstein, 2018

A pimpline pimplinae ichneumon wasp

 
Xanthopimpla praeclara
holotype

Siricidae

Xoanon?

Xoanon? eocenicus[32]

Wedmann, Pouillon, & Nel, 2014

A siricid horntail wasp
Uncertainly placed into Xoanon

Vespidae

Vespula?

Vespula? hassiaca[33]

Abels & Wedmann, 2021

A vespid wasp
Tentatively placed with the yellowjackets

Lepidoptera edit

Family Genus Species Authority Notes Image

?Zygaenidae

undescribed

undescribed[34]

Two undescribed moth taxa
Possibly belonging to Zygaenidae
wings scales with preserved structural coloration

 
Undescribed moth

Neuroptera edit

Family Genus Species Authority Notes Image

Mantispidae

Symphrasites

Symphrasites eocenicus[35]

Wedman & Makarkin, 2007

A symphrasine mantidfly
Described from a isolated forewing

 

Odonata edit

Family Genus Species Authority Notes Image

Dysagrionidae

Petrolestes

Petrolestes messelensis[36]

Garrouste & Nel, 2015

A damselfly
Related to the North American P. hendersoni.

 

Pseudostenolestidae

Pseudostenolestes

Pseudostenolestes bechlyi[36]

Garrouste & Nel, 2015

A damselfly species.

Phasmatodea edit

Family Genus Species Authority Notes Image

Phasmatodea

Eophyllium

Eophyllium messelensis[37]

Wedmann, Bradler & Rust, 2007

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

"Fish" edit

Amiiformes edit

Family Genus Species Authority Notes Image

Amiidae

Cyclurus

Cyclurus kehreri[38]

(Andreae, 1893)

An amiine bowfin relative

 
Cyclurus kehreri

Anguilliformes edit

Family Genus Species Authority Notes Image

Anguillidae

Anguilla

Anguilla ignota[2]

Micklich, 1985

Anguilla ignota fossils suggest a likely aquatic link to an ocean.

 

Lepisosteiformes edit

Family Genus Species Authority Notes Image
Lepisosteidae Atractosteus

Atractosteus messelensis[39]

Grande, 2010

One of three Messel gar species.

Atractosteus strausi[40]

(Kinkelin, 1884)

One of three Messel gar species.

 

Masillosteus

Masillosteus kelleri[41]

Micklich & Klappert, 2001

A rare, blunt snouted gar that likely fed on small invertebrates.
It may not have been a permanent resident of the Messel lake.[2]

Perciformes edit

Family Genus Species Authority Notes Image

Moronidae

Palaeoperca

Palaeoperca proxima[42]

Micklich, 1978

Palaeoperca may have been open water fish.

 

Percichthydae[43]

Amphiperca

Amphiperca multiformis[44]

Weitzel, 1933

Amphiperca likely was a thrust or ambush hunter catching prey from near-shore hiding spots.

 

incertae sedis

Rhenanoperca

Rhenanoperca minuta[45]

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 edit

Family Genus Species Authority Notes Image
Thaumaturidae Thaumaturus Thaumaturus intermedius[46] Weitzel, 1933[44] Thaumaturus is only known from juvenile specimens, ranging in length from 3–6 cm (1.2–2.4 in).[2] Its relationship with other groups of fish remains unknown.  

Amphibians edit

Caudata edit

Family Genus Species Authority Notes Image
Salamandridae Chelotriton

Chelotriton robustus[47]

Westphal, 1980

A newt,
possibly related to ribbed newts and crocodile newts.

Chelotriton sp.[48]

Undescribed

A newt,
possibly related to ribbed newts and crocodile newts.

Anura edit

Family Genus Species Authority Notes Image

Palaeobatrachidae

"Messelobatrachus"

"Messelobatrachus tobieni"[49]

Nomen nudum

Messelobatrachus and M. tobieni were first coined in a 1988 PhD. dissertation, but never formally published.[43]
One fossil preserves a possible tadpole of "Messelobatrachus".

 

Pelobatidae

Eopelobates

Eopelobates wagneri[50]

Weitzel, 1938

A spadefoot toad relative
It was likely primarily terrestrial and entered the lake to spawn, as shown by preserved eggs.

 

Pipidae

Undescribed

Undescribed[43]

Two undescribed specimens of Xenopinae clawed frogs.

Incertae sedis

Lutetiobatrachus

Lutetiobatrachus gracilis[51]

Wuttke, 1998

First coined in 1988, but nomen nudum until redescription in 2012.

Squamata edit

Clade Genus Species Authority Notes Image
Anguidae Ophisauriscus Ophisauriscus quadrupes[52] Kuhn, 1940 A serpentine reptile with vestigial limbs. It shows evidence of different osteoderm morphology after tail regeneration.
Placosauriops "Placosauriops abderhaldeni"[53] Kuhn, 1940 A type of melanosaurinin Glyptosaurine of dubious assignment, as the type material has not been adequately diagnosed.
Boidae Eoconstrictor Eoconstrictor fischeri[54] (Schaal, 2004) An early species of boa previously known as Palaeopython fischeri.
 
Messelophis Messelophis variatus[55] Baszio, 2004 A minute boa
Rieppelophis Rieppelophis ermannorum[55] (Schaal & Baszio, 2004) A minute boa previously regarded as a species of Messelophis.
 
Corytophanidae Geiseltaliellus Geiseltaliellus maarius[56] Smith, 2009 An extinct, possibly arboreal, relative of the modern basilisk lizard.  
Eolacertidae Eolacerta Eolacerta robusta[57] Nöth, 1940 A large species of lizard, reaching a length of 30 cm (12 in) and a weight of 1 kg (2.2 lb).
Stefanikia Stefanikia siderea[58] Čerňanský & Smith, 2017 A close relative of Eolacerta, the two genera were recovered as a sister-group to modern lizards.
Erycinae Rageryx Rageryx schmidi[59] Smith & Scanferla, 2021 A non-burrowing sandboa that reached a length of 52 cm (20 in).
Cf. Lacertidae Undescribed Undescribed [60] An undescribed arboreal lizard relative.
Gekkota Undescribed Undescribed [60] A partly disarticulated skeleton of a gekko
Helodermatidae Eurheloderma Eurheloderma sp.[61] A relative of the extant gila monster, it was likely already venomous.
Lacertibaenia Cryptolacerta Crytolacerta hassiaca[62] 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
Messelopythonidae Messelopython Messelopython freyi[63] Zaher & Smith, 2020 A medium sized pythonoid.
Palaeopython Palaeopython schaali[64] 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.[60] Identified as Necrosaurus by Smith, Čerňanský and Scanferla, it is smaller than Paranecrosaurus.
Paranecrosaurus Paranecrosaurus feisti[65] (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.  
Pan-Shinisaurus Pan Shinisaurus indet. Pan Shinisaurus indet.[66] The shed tail of a crocodile lizard, showing adaptation for swimming.
Polychrotidae Undescribed Undescribed [60] A possible relative of the modern bush anole.
Scincidae Scincidae indet. Scincidae indet.[2]
Scincoidea Ornatocephalus Ornatocephalus metzleri[67] Weber, 2004 An arboreal lizard with long claws and a prehensile tail, it was found with plants remains in its guts.

Testudinata edit

Family Genus Species Authority Notes Image
Carettochelyidae Allaeochelys Allaeochelys crassesculptata[68] (Harrassowitz, 1922) A relative of the modern pig-nosed turtle, one fossil preserves two specimens while mating.  
Geoemydidae Euroemys Euroemys kehreri[69] Staesche, 1928 A species of pond turtles of uncertain relationship, it was previously thought to be a species of Ocadia and Palaeoemys.[70]  
Francellia Francellia messeliana[69] Staesche, 1928 A species of pond turtles of uncertain relationship, it was previously thought to be a species of Ocadia and Palaeoemys.[70]
Juvemys Juvemys sp.[2] Hervet, 2003 A species of pond turtles.
Podocnemididae Neochelys Neochelys franzeni[71] Schleich, 1993 [72] A podocnemidid side-necked turtle  
Trionychidae Palaeoamyda Palaeoamyda messeliana [73] (Reinach, 1900) A species of softshell turtle  

Crocodyliformes edit

Family Genus Species Authority Notes Image
Alligatoroidea Diplocynodon Diplocynodon darwini

Diplocynodon deponaie[74]

(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.  
Alligatoridae Hassiacosuchus Hassiacosuchus haupti [75] Weitzel, 1935 A small species of alligatorid, some consider it a species of Allognathosuchus. Christopher Brochu argues for the continued use of Hassiacosuchus however.[76]  
Bergisuchidae Bergisuchus Bergisuchus dietrichbergi[77] Kuhn, 1968 Member of the terrestrial Sebecosuchians with ziphodont teeth.  
Crocodyloidea "Asiatosuchus" "Asiatosuchus" germanicus[78] Berg, 1966 A large and well known species of the paraphyletic genus Asiatosuchus.  
Planocraniidae Boverisuchus Boverisuchus magnifrons[79] Kuhn, 1938 A terrestrial Crocodilian previously assigned to the now dubious Pristichampsus  
Tomistominae Tomistominae indet. Tomistominae indet.[80]

Birds edit

Palaeognathae edit

Family Genus Species Authority Notes Image
Lithornithidae Lithornis Lithornis sp.[81][82] 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[83] Lambrecht, 1928 A flightless palaeognath, previous analysis considered it to be a relative to ostriches or rheas. More recent research however doubts this.  

Anseriformes edit

Family Genus Species Authority Notes Image
Gastornithidae Gastornis Gastornis geiselensis[84] Fischer, 1978 A large, herbivorous anseriform.[85]
 

Galliformes edit

Clade Genus Species Authority Notes Image
Gallinuloididae Paraortygoides Paraortygoides messelensis[86] Mayr, 2000 One of the earliest known galliforms, the lack of a crop indicates they fed on only soft plant material.

Mirandornithes edit

Family Genus Species Authority Notes Image
Juncitarsus Juncitarsus merkeli[87] 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.
 

Cuculiformes edit

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

Strisores edit

Family Genus Species Authority Notes Image
Cypseloramphus Cypseloramphus dimidius[88] Mayr, 2016 A tentative apodiform.
Archaeotrogonidae Hassiavis Hassiavis laticauda[89] Mayr, 1998 The skull of Hassiavis closely resembles that of modern owlet-nightjars, being broad and short.
 
Masillapodargus Masillapodargus longipes[90] Mayr, 2001 A relative of the modern frogmouths.
Jungornithidae? Parargornis Parargornis messelensis[91] Mayr, 2003 A swift-like bird with short wings and long tail feathers, phylogenetic analysis indicate that it was a basal type of hummingbird.[92]
 
Nyctibiidae Paraprefica Paraprefica kelleri[93]

Paraprefica major[90]

Mayr, 1999 An old-world genus of potoo. The two species can clearly be differentiated by size.
 
Protocypselomorphus Protocypselomorphus manfredkelleri[94] Mayr, 2005 An aerial insectivore with reduced feet, it was recovered as a sister-taxon to all other Strisores.
Pan-Apodidae Scaniacypselus Scaniacypselus szarskii[95] (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.
 

Charadriiformes edit

Family Genus Species Authority Notes Image

Jacanidae?

Vanolimicola

Vanolimicola longihallucis[96]

Mayr, 2017

A small wading bird with elongated hallux.
It is possibly a jacanid or a Songziidae rail-relative Gruiform.

Gruimorphae edit

Family Genus Species Authority Notes Image
Messelornithidae Messelornis Messelornis cristata[97] 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.
 

Suliformes edit

Family Genus Species Authority Notes Image
Sulidae? Masillastega Masillastega rectirostris[98] Mayr, 2002 A bird tentatively referred to Sulidae, it was approximately the size of a modern European shag. It was likely piscivorous.
 

Pelecaniformes edit

Family Genus Species Authority Notes Image
Threskiornithidae Rhynchaeites Rhynchaeites messelensis[99] Wittich, 1898 Also known as the "snipe-rail" for its similarities to rails and painted snipes, it was actually a type of ibis.
 

Strigiformes edit

Family Genus Species Authority Notes Image
Palaeoglaucidae Palaeoglaux Palaeoglaux artophoron[100] Peters, 1992 A small owl preserving ribbon-like feathers arranged in a dense layer on its back.
 

Coraciimorphae edit

Family Genus Species Authority Notes Image
Sandcoleidae Chascacocolius Chascacocolius cacicirostris[101] Mayr, 2005 A coliiform bird with a conical beak.
Eocoraciidae Eocoracias Eocoracias brachyptera[102] Mayr, 2000 An early species of roller, analysis of the plumage gave indication as to what its colours would have been in life.[103]
 
Sandcoleidae Eoglaucidium Eoglaucidium pallas [104]

Eoglaucidium sp. indet.[81]

Mayr & Peters, 1998 Originally described as an owl from the Geiseltal, it was later found to be related to mousebirds. Its tail is notably shorter than that of its extant relatives. It is possible that the indetermined species instead belongs to Anneavis.
 
Gracilitarsidae Gracilitarsus Gracilitarsus mirabilis[105] Mayr, 1998 The shape of the beak indicates that Gracilitarsus was either an insectivore or fed on nectar.
Coliidae Masillacolius Masillacolius brevidactylus[104] Mayr & Peters, 1998 An extinct species of mousebird.
Trogonidae Masillatrogon Masillatrogon pumilio[106] Mayr, 2009 An extinct species of trogon that supports the hypothesis that the family originated in the Old World.
 
Messelirrisoridae Messelirrisor Messelirrisor grandis[107]

Messelirrisor halycrostris[108]

Messelirrisor parvus[108]

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.
 
Leptosomidae? Plesiocathartes Plesiocathartes kelleri[109] Mayr, 2002 A possible relative of the modern cuckoo-roller.
Primobucconidae Primobucco Primobucco frugilegus[110]

Primobucco perneri

Mayr, Mourer-Chaviré & Weidig, 2004 The first recorded species of Primobucco in Europe.
 
Incertae sedis Quasisyndactylus Quasisyndactylus longibrachis[89] Mayr, 1998 Quasisyndactylus is thought to be a relative of modern kingfishers, todies and motmots.
 
Coliidae [81] Selmes Selmes absurdipes [111] Peters, 1999 A genus of mousebird.

Cariamiformes (?) edit

Several groups of Messel birds share characteristics with the modern seriemas, which has led to them being placed within the clade Cariamae in the past. However, this placement typically occurred 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.[112]

Family Genus Species Authority Notes Image
Idiornithidae Dynamopterus Dynamopterus cf. itardiensis[113]

Dynamopterus tuberculata[114]

(Mourer-Chauviré, 1983)

(Peters, 1995)

Previously known as Idiornis, it was later found to be synonymous with Dynamopterus.  
Salmilidae Salmila Salmila robusta[115] 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.  
Ameghinornithidae Strigogyps Strigogyps sapea[116] (Peters, 1987) Previously known as Aenigmavis, stomach contents indicate it was a herbivore. The phylogenetic position of Strigogyps is uncertain.  

Falconiformes edit

Family Genus Species Authority Notes Image
Masillaraptoridae Masillaraptor Masillaraptor parvunguis[117][118] 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.

Psittacopasserae edit

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).[119]
 
Quercypsittidae? Palaeopsittacus Palaeopsittacus cf. georgei[120] 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[121]

Primozygodactylus danielsi[121]

Primozygodactylus eunjooae[122]

Primozygodactylus longibrachium[121]

Primozygodactylus mayor[121]

Primozygodactylus quintus[121]

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.
 
Pseudasturidae Pseudorasturidae indet. Pseudorasturidae indet.[113] 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[123] Mayr & Daniels, 1998 A bird originally classified as a parrot, later research indicated that it may instead be closer to passeriforms.
Psittacopedidae[81] Pumiliornis Pumiliornis tessellatus[124] Mayr, 1999 A bird the size of a small wren.
Pseudasturidae Serudaptus Serudaptus pohli[113] Mayr, 2000 A bird with zygodactyl feet and long claws suited for climbing. Its exact phylogenetic position is poorly understood, with some research indicating it may be a basal member of the clade containing Passeriformes and Psittaciformes.
 

Incertae sedis edit

Family Genus Species Authority Notes Image
Eopachypterygidae Eopachypteryx Eopachypteryx praeterita

Eopachypteryx ? sp.[125]

Mayr, 2015
Incertae sedis Lapillavis Lapillavis incubarens[88] Mayr, 2016
Incertae sedis Perplexicervix Perplexicervix microcephalon[126] 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.

Mammals edit

Apatotheria edit

Family Genus Species Authority Notes Image
Apatemyidae Heterohyus Heterohyus nanus[127] 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]
 

Artiodactyla edit

Family Genus Species Authority Notes Image
Dichobunidae Aumelasia Aumelasia cf. gabineaudi[128] Sudré, 1980 A lightly built early artiodactyl with a noticeably long tail. Aumelasia is referred to the Diacodexeidae by some researchers.[43]
 
Dichobunidae Messelobunodon Messelobunodon schaeferi[129] Franzen, 1980 An early species of even-toed ungulate.
 
Choeropotamidae Masillabune Masillabune martini[130] Tobien, 1980 An early species of even-toed ungulate.
 

Chiroptera edit

Family Genus Species Authority Notes Image
Archaeonycteridae Archaeonycteris Archaeonycteris pollex[131]

Archaeonycteris trigonodon[132]

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[133]

Hassianycteris messelensis[133]

Hassianycteris? revilliodi[43]

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[43]

Palaeochiropteryx tupaiodon[43]

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]
 
Emballonuridae Tachypteron Tachypteron franzeni[134] Storch, Sigé & Habersetzer, 2002 A fast flying species that inhabited open airspaces, Tachypteron resembled the modern black-bearded tomb bat in proportions.[2]

Cimolesta edit

Family Genus Species Authority Notes Image
Pantolestidae Buxolestes Buxolestes piscator

Buxolestes minor[135]

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.
 
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.
 
Paroxyclaenidae "Paroxyclaenus" "Paroxyclaenus" sp.[43] An undescribed relative of Kopidodon differing through the lack of a pronounced sagittal crest.

Eulipotyphla edit

Family Genus Species Authority Notes Image
Amphilemuridae Macrocranion Macrocranion tenerum[136]

Macrocranion tupaiodon[137]

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 covered in bristles. Some researchers suggest it may have moved like modern springhares.[2][138]
 
Amphilemuridae Pholidocercus Pholidocercus hassiacus[139] von Koenigswald & Storch, 1983 A larger relative of Macrocranion, Pholidocercus was an animal 40 cm (16 in). Its 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 similarly covered in a hardened structure while the torso was covered in thick bristles.[2]
 

Leptictida edit

Family Genus Species Authority Notes Image
Pseudorhyncocyonidae Leptictidium Leptictidium auderiense[43]

Leptictidium nasutum[140]

Leptictidium tobieni[141]

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. Its hindlimbs are noticeably larger than its forelimbs, but its means of locomotion are not agreed upon.
 

Metatheria edit

Family Genus Species Authority Notes Image
Herpetotheriidae Amphiperatherium Amphiperatherium goethei[142]

Amphiperatherium cf. maximum[143]

Crochet, 1979 A metatherian mammal, it was larger than Peradectes but with a proportionally shorter tail.
 
Herpetotheriidae Herpetotheriidae indet. Either Amphiperatherium or Peratherium.[43]
Peradectidae "Peradectes" "Peradectes sp."[143] A small arboreal mammal similar to Peradectes with a prehensile tail.
 

Pan-Carnivora edit

Family Genus Species Authority Notes Image
Proviverridae Lesmesodon Lesmesodon behnkeae[144]

Lesmesodon edingeri[145]

(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.
 
Miacidae Messelogale Messelogale kessleri[146] (Springhorn, 1982)[145] 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[147] Springhorn, 1980 An arboreal miacid.
 

Pholidota edit

Family Genus Species Authority Notes Image
Eomanidae[148] 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.[149]
 
Euromanis Euromanis krebsi[150] (Storch & Martin, 1994) A type of scaleless pangolin originally described as a species of Eomanis.
 
Eurotamanduidae Eurotamandua Eurotamandua joresi[151] 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.
 

Perissodactyla edit

Family Genus Species Authority Notes Image
Palaeotheriidae Eurohippus Eurohippus parvulus[152]

Eurohippus messelensis[153]

(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.
 
Hallensia Hallensia matthesi[154] Franzen & Haubold, 1986 A small species of primitive horse.
 
Hyrachyidae Hyrachyus Hyrachyus minimus [155] Fischer, 1829 An early rhinoceratoid or tapir, its discovery played a rolle in making Messel a protected site.
 
Palaeotheriidae Propalaeotherium Propalaeotherium hassiacum[156] Haupt, 1925 An early species of horse reaching a shoulder height of 55–60 cm (22–24 in), making it larger than Eurohippus.
 

Primates edit

Family Genus Species Authority Notes Image
Adapidae Darwinius Darwinius masillae[157] 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.
 
Adapidae Europolemur Europolemur kelleri[158]

Europolemur koenigswaldi[158]

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 embedded into it.[2]
 
Adapidae Godinotia Godinotia neglecta[159] (Thalmann et al., 1989) Preserved stomach contents of Godinotia suggest that it primarily fed on leaves and fruit.[160]
 

Rodentia edit

Family Genus Species Authority Notes Image

Alagomyidae[43]

Ailuravus

Ailuravus macrurus[161]

Weitzel, 1949

An 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]

 

Gliridae

Eogliravus

Eogliravus wildi[162]

Hartenberger, 1971

The oldest known dormouse
E. wildi had a long, bushy tail
It was likely herbivorous, feeding on seeds and plant buds.[citation needed]

Ischyromyidae Masillamys

Masillamys beegeri[163]

Masillamys krugi[163]

Masillamys parvus[163]

Tobien, 1954

M. parvus was sometimes placed in Hartenbergeromys,[2]
However a 2019 study found the asserted differences to the type species insufficient to warrant this split.
Its tail was only sparsely covered in fur.[163]

 
Masillamys

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