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

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

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 present in the oldest Messel formation stata Considered almost identical to the modern Ctenodaphnia magna.
Daphniidae	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
? Palaemonidae	†Bechleja	†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

Araneae

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

Family	Genus	Species	Authority	Notes	Image
Undescribed	Undescribed	Undescribed^[6]^[7]		Six harvestmen awaiting description.

Insects

Coleoptera

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	†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

Family	Genus	Species	Authority	Notes	Image
Blaberidae	Morphna	†Morphna cenozoica^[15]	Šmídová, Vidlička & Wedmann, 2021	A Blaberid giant cockroach.

Dipterans

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
Aradidae	Aneurus?	Aneurus? incertus^[20]	Wappler, Heiss & Wedmann, 2015	A species of flat bug.
Aradidae	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

Family	Genus	Species	Authority	Notes	Image
Apidae	†Pygomelissa	†Pygomelissa lutetia^[25]	Wappler & Engel, 2003	A bee of uncertain tribal placement.
Apidae	†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	†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	†Pseudectatomma striatula^[27]	Dlussky & Wedmann, 2012	An ectatommine ant
	†Titanomyrma	†Titanomyrma gigantea^[30]	(Lutz, 1986)	A formiciine subfamily ant	Titanomyrma simillima holotype
	†Titanomyrma	†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	†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

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

Family	Genus	Species	Authority	Notes	Image
Mantispidae	†Symphrasites	†Symphrasites eocenicus^[35]	Wedman & Makarkin, 2007	A symphrasine mantidfly Described from a isolated forewing

Odonata

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

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"

Amiiformes

Family	Genus	Species	Authority	Notes	Image
Amiidae	†Cyclurus	†Cyclurus kehreri^[38]	(Andreae, 1893)	An amiine bowfin relative	Cyclurus kehreri

Anguilliformes

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

Family	Genus	Species	Authority	Notes
Lepisosteidae	Atractosteus	Atractosteus messelensis^[39]	Grande, 2010	One of three Messel gar species.
	Atractosteus	†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

Family	Genus	Species	Authority	Notes
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	A 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^[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

Caudata

Family	Genus	Species	Authority	Notes	Image
Salamandridae	Chelotriton	Chelotriton robustus^[47]	Westphal, 1980	A newt, possibly related to ribbed newts and crocodile newts.
Salamandridae	Chelotriton	Chelotriton sp.^[48]	Undescribed	A newt, possibly related to ribbed newts and crocodile newts.

Anura

Family	Genus	Species	Authority	Notes
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

Clade	Genus	Species	Authority	Notes
Anguidae	Ophisauriscus	Ophisauriscus quadrupes^[52]	Kuhn, 1940	A serpentine reptile with vestigial limbs. It shows evidence of different osteoderm morphology after tail regeneration.
Anguidae	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).
Eolacertidae	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.
Messelopythonidae	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.
Palaeovaranidae	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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

Palaeognathae

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

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

Galliformes

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

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

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

Strisores

Family	Genus	Species	Authority	Notes
	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

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

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

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

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

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

Family	Genus	Species	Authority	Notes
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 (?)

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
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

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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

Apatotheria

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

Family	Genus	Species	Authority	Notes
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

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

Family	Genus	Species	Authority	Notes
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

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

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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

Family	Genus	Species	Authority	Notes
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

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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[Mazancourt2022-4] Mazancourt, V.; Wappler, T.; Wedmann, S. (2022-10-27). "Exceptional preservation of internal organs in a new fossil species of freshwater shrimp (Caridea: Palaemonoidea) from the Eocene of Messel (Germany)". Scientific Reports. 12 (1): 18114. Bibcode:2022NatSR..1218114D. doi:10.1038/s41598-022-23125-9. ISSN 2045-2322. PMC 9613706. PMID 36302944.

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[PAABlaberidae-15] Šmídová, L.; Vidlička, L.; Wedmann, S. (2021). "Appearance of the family Blaberidae (Insecta: Blattaria) during the Cretaceous and a review of fossils of this family". Palaeontographica Abteilung A. 321 (1–6): 71–79. doi:10.1127/pala/2021/0109. S2CID 243467634.

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[WY08-17] Wedmann, S.; Yeates, D.K. (2008). "Eocene records of bee flies (Insecta, Diptera, Bombyliidae, Comptosia): their palaeobiogeographic implications and remarks on the evolutionary history of bombyliids". Palaeontology. 51 (1): 231–240. Bibcode:2008Palgy..51..231W. doi:10.1111/j.1475-4983.2007.00745.x. S2CID 128576916.

[WG07-18] Wedmann, S.; Richter, G. (2007). "The ecological role of immature phantom midges (Diptera: Chaoboridae) in the Eocene Lake Messel, Germany". African Invertebrates. 48 (1): 59–70.

[WHEZG21-19] Wedmann, S.; Hörnschemeyer, T.; Engel, M. S.; Zetter, R.; Grímsson, F. (2021). "The last meal of an Eocene pollen-feeding fly". Current Biology. 31 (9): 2020–2026.e4. Bibcode:2021CBio...31E2020W. doi:10.1016/j.cub.2021.02.025. PMID 33705719.

[PZMesselflatbugs-20] Torsten Wappler; Ernst Heiss; Sonja Wedmann (2015). "New flatbug (Hemiptera: Heteroptera: Aradidae) records from the Middle Eocene Messel Maar, Germany". Paläontologische Zeitschrift. 89 (3): 653–660. Bibcode:2015PalZ...89..653W. doi:10.1007/s12542-014-0237-1. S2CID 83668045.

[planthoppers2006-21] Szwedo, J.; Wappler, T. (2006). "New planthoppers (Insecta: Hemiptera: Fulgoromorpha) from the Middle Eocene Messel Maar". Annales Zoologici. 56 (3): 555–566. doi:10.3161/000345406778811907 (inactive 1 November 2024).{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)

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[Lutetiacader2006-24] Wappler, T. (2006). "Lutetiacader, a puzzling new genus of cantacaderid lace bugs (Heteroptera: Tngidae) from the Middle Eocene Messel Maar, Germany". Palaeontology. 49 (2): 435–444. Bibcode:2006Palgy..49..435W. doi:10.1111/j.1475-4983.2006.00547.x. S2CID 86692157.

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[Dlussky2012-26] Dlussky, G.M. (2012). "New Fossil Ants of the Subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany". Paleontological Journal. 46 (3): 288–292. Bibcode:2012PalJ...46..288D. doi:10.1134/s0031030111050054. S2CID 83891156.

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[Dlussky2009-28] Dlussky, GM; Wappler, T; Wedmann, S (2009). "Fossil ants of the genus Gesomyrmex Mayr (Hymenoptera, Formicidae) from the Eocene of Europe and remarks on the evolution of arboreal ant communities". Zootaxa. 2031: 1–20. doi:10.11646/zootaxa.2031.1.1. S2CID 56253714.

[Dlussky-etal-2015-29] Dlussky, G.M.; Rasnitsyn, A.P.; Perfilieva, K.S. (2015). "The Ants (Hymenoptera: Formicidae) of Bol'shaya Svetlovodnaya (Late Eocene of Sikhote-Alin, Russian Far East)". Caucasian Entomological Bulletin. 11 (1): 131–152. doi:10.23885/1814-3326-2015-11-1-131-152.

[Archibald_et_al_(2011)-30] Archibald, S. Bruce; Johnson, Kirk R.; Mathewes, Rolf W.; Greenwood, David R. (2011). "Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals". Proceedings of the Royal Society B. 278 (1725): 3679–3686. doi:10.1098/rspb.2011.0729. PMC 3203508. PMID 21543354.

[Mesornatus-31] ^ ^a ^b ^c ^d ^e ^f ^g Tamara Spasojevic; Sonja Wedmann; Seraina Klopfstein (2018). "Seven remarkable new fossil species of parasitoid wasps (Hymenoptera, Ichneumonidae) from the Eocene Messel Pit". PLOS ONE. 13 (6): e0197477. Bibcode:2018PLoSO..1397477S. doi:10.1371/journal.pone.0197477. PMC 5991363. PMID 29874268.

[32] Wedmann, S.; Pouillon, J.; Nel, A. (2014). "New Palaeogene horntail wasps (Hymenoptera, Siricidae) and a discussion of their fossil record". Zootaxa. 3869 (1): 33–43. doi:10.11646/zootaxa.3869.1.3. PMID 25283896.

[33] Abels, J.; Wedmann, S. (2021). "A fossil wasp (Hymenoptera: Vespidae: Vespula? hassiaca) from the Eocene of Messel". Palaeontographica Abteilung A. 323 (1–3): 105–117. doi:10.1127/pala/2021/0122. S2CID 245248645.

[Leps2011-34] McNamara, M.; Briggs, D.; Orr, P.; Wedmann, S.; Noh, H.; Cao, H. (2011). "Fossilized biophotonic nanostructures reveal the original colors of 47-million-year-old moths". PLOS Biology. 9 (11): 1–8. doi:10.1371/journal.pbio.1001200. PMC 3217029. PMID 22110404.

[Mantispidae2007-35] Wedmann, S.; Makarkin, V. (2007). "A new genus of Mantispidae (Insecta: Neuroptera) from the Eocene of Germany, with a review of the fossil record and palaeobiogeography of the family". Zoological Journal of the Linnean Society. 149 (4): 701–716. doi:10.1111/j.1096-3642.2007.00273.x. S2CID 13701556.

[Pseudostenolestes-36] Romain Garrouste; André Nel (2015). "New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata)". Zootaxa. 4028 (3): 354–366. doi:10.11646/zootaxa.4028.3.2. PMID 26624314.

[WBR07-37] Wedmann, S.; Bradler, S; Rust, J. (9 January 2007). "The first fossil leaf insect: 47 million years of specialized cryptic morphology and behavior". Proceedings of the National Academy of Sciences. 104 (2): 565–9. Bibcode:2007PNAS..104..565W. doi:10.1073/pnas.0606937104. PMC 1766425. PMID 17197423.

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[SS20-54] Scanferla, A.; Smith, K.T. (2020). "Exquisitely Preserved Fossil Snakes of Messel: Insight into the Evolution, Biogeography, Habitat Preferences and Sensory Ecology of Early Boas". Diversity. 12 (3): 100. doi:10.3390/d12030100. hdl:11336/145425.

[SSS16-55] Agustín Scanferla; Krister T. Smith; Stephan F. K. Schaal (2016). "Revision of the cranial anatomy and phylogenetic relationships of the Eocene minute boas Messelophis variatus and Messelophis ermannorum (Serpentes, Booidea)". Zoological Journal of the Linnean Society. 176 (1): 182–206. doi:10.1111/zoj.12300. hdl:11336/77086.

[S09-56] Smith, K.T. (2009). "Eocene Lizards of the Clade Geiseltaliellus from Messel and Geiseltal, Germany, and the Early Radiation of Iguanidae (Reptilia: Squamata)". Bulletin of the Peabody Museum of Natural History. 50 (2): 219–306. doi:10.3374/014.050.0201. S2CID 84085505.

[M01-57] Müller, J. (2001). "Osteology and relationships of Eolacerta robusta, a lizard from the Middle Eocene of Germany (Reptilia, Squamata)". Journal of Vertebrate Paleontology. 21 (2): 261–278. doi:10.1671/0272-4634(2001)021[0261:oaroer]2.0.co;2. S2CID 86300545.

[C17-58] Čerňanský, A.; Smith, K. T. (2017). "Eolacertidae: a new extinct clade of lizards from the Palaeogene; with comments on the origin of the dominant European reptile group – Lacertidae". Historical Biology. 30 (7): 994–1014. doi:10.1080/08912963.2017.1327530. S2CID 49546941.

[SS21-59] Smith, K. T.; Scanferla, A. (2021). "A nearly complete skeleton of the oldest definitive erycine boid (Messel, Germany)". Geodiversitas. 43 (1): 1–24. doi:10.5252/geodiversitas2021v43a1. S2CID 231666355.

[Messelbook-60] Smith, K.T.; Čerňanský, A.; Scanferla, A. (2018). "10.1 Lizards and snakes-Warmth-loving sunbathers". Messel-An Ancient Greenhouse Ecosystem. Senckenberg. pp. 123–147.

[C16b-61] Cadena, E. (2016). "Microscopical and elemental FESEM and Phenom ProX-SEM-EDS analysis of osteocyte- and blood vessel-like microstructures obtained from fossil vertebrates of the Eocene Messel Pit, Germany". PeerJ. 4: e1618. doi:10.7717/peerj.1618. PMC 4727973. PMID 26819855.

[MHH11-62] Müller, Johannes; Christy A. Hipsley; Jason J. Head; Nikolay Kardjilov; André Hilger; Michael Wuttke; Robert R. Reisz (2011). "Eocene lizard from Germany reveals amphisbaenian origins". Nature. 473 (7347): 364–367. Bibcode:2011Natur.473..364M. doi:10.1038/nature09919. PMID 21593869. S2CID 205224382.

[ZS20-63] Zaher, H.; Smith, K. T. (2020). "Pythons in the Eocene of Europe reveal a much older divergence of the group in sympatry with boas". Biology Letters. 16 (12). doi:10.1098/rsbl.2020.0735. PMC 7775975. PMID 33321065.

[SS21b-64] Smith, K. T.; Scanferla, A. (2022). "More than one large constrictor snake lurked around paleolake Messel". Palaeontographica Abteilung A. 323 (1–3): 75–103. Bibcode:2022PalAA.323...75S. doi:10.1127/pala/2021/0119. S2CID 245089117.

[SH21-65] Smith, K. T.; Habersetzer, J. (2021). "The anatomy, phylogenetic relationships, and autecology of the carnivorous lizard "Saniwa" feisti Stritzke, 1983 from the Eocene of Messel, Germany". Comptes Rendus Palevol. 20 (23): 441–506. doi:10.5852/cr-palevol2021v20a23.

[Sm17-66] Smith, K. T. (2017). "First crocodile-tailed lizard (Squamata: Pan-Shinisaurus) from the Paleogene of Europe". Journal of Vertebrate Paleontology. 37 (3): e1313743. Bibcode:2017JVPal..37E3743S. doi:10.1080/02724634.2017.1313743. S2CID 89730027.

[W04-67] Weber, S. (2004). "Ornatocephalus metzleri gen. et spec. nov. (Lacertilia, Scincoida) - Taxonomy and Paleobiology of a basal Scincoid Lizard from the Messel Formation (Middle Eocene: basal Lutetian, Geiseltalium), Germany". Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft. 561. ISBN 978-3-510-61373-1.

[JMSS12-68] Joyce, W.G.; Micklich, N.; Schaal, S.F.K.; Scheyer, T.M. (2012). "Caught in the act: the first record of copulating fossil vertebrates". Biology Letters. 8 (5): 846–848. doi:10.1098/rsbl.2012.0361. PMC 3440985. PMID 22718955. S2CID 478288.

[H04-69] Hervet, S. (2004). "Systématique du groupe "Palaeochelys sensu lato – Mauremys" (Chelonii, Testudinoidea) du Tertiaire d'Europe occidentale: principaux résultats". Annales de Paléontologie. 90 (1): 13–78. doi:10.1016/j.annpal.2003.12.002. ISSN 0753-3969.

[ACJ21-70] Ascarrunz, E.; Claude, J.; Joyce, WG. (2021). "The phylogenetic relationships of geoemydid turtles from the Eocene Messel Pit Quarry: a first assessment using methods for continuous and discrete characters". PeerJ. 9: e11805. doi:10.7717/peerj.11805. PMC 8349520. PMID 34430073.

[C15-71] Cadena, E. (2015). "A global phylogeny of Pelomedusoides turtles with new material of Neochelys franzeni Schleich, 1993 (Testudines, Podocnemididae) from the middle Eocene, Messel Pit, of Germany". PeerJ. 3: e1221. doi:10.7717/peerj.1221. PMC 4556147. PMID 26336649.

[S93-72] Schleich, H. H. (1993). "New reptile material from the German Tertiary. 11. Neochelys franzeni n. sp., the first pleurodiran turtle from Messel". Kaupia. 3: 15–21.

[C16-73] Cadena, E. (2016). "Palaeoamyda messeliana nov. comb. (Testudines, Pan-Trionychidae) from the Eocene Messel Pit and Geiseltal localities, Germany, taxonomic and phylogenetic insights". PeerJ. 4: e2647. doi:10.7717/peerj.2647. PMC 5088588. PMID 27812431. S2CID 37745074.

[DS12-74] Delfino, M.; Smith, T. (2012). "Reappraisal of the morphology and phylogenetic relationships of the middle Eocene alligatoroid Diplocynodon deponiae (Frey, Laemmert, and Riess, 1987) based on a three-dimensional specimen". Journal of Vertebrate Paleontology. 32 (6): 1358–1369. doi:10.1080/02724634.2012.699484. S2CID 84977303.

[W35-75] Weitzel, K. (1935). "Hassiacosuchus haupti n. g n. s. ein durophages Krokodil aus dem Mitteleozän von Messel". Notizblatt des Vereins für Erdkunde und der hessischen geologischen Landesanstalt zu Darmstadt. 16: 40–49.

[CAB04-76] Brochu, Christopher A. (2004). "Alligatorine phylogeny and the status of Allognathosuchus Mook, 1921". Journal of Vertebrate Paleontology. 24 (4): 857–873. doi:10.1671/0272-4634(2004)024[0857:APATSO]2.0.CO;2. ISSN 0272-4634. S2CID 85050852.

[RRO00-77] Rossmann, T.; Rauhe, M.; Ortega, F. (2000). "Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions". Paläontologische Zeitschrift. 74 (3): 379–392. doi:10.1007/BF02988108. S2CID 129507618.

[DS09-78] Delfino, M.; Smith, T. (2009). "A reassessment of the morphology and taxonomic status of 'Crocodylus'depressifronsBlainville, 1855 (Crocodylia, Crocodyloidea) based on the Early Eocene remains from Belgium". Zoological Journal of the Linnean Society. 156: 140–167. doi:10.1111/j.1096-3642.2008.00478.x.

[B13-79] Brochu, C. A. (2013). "Phylogenetic relationships of Palaeogene ziphodont eusuchians and the status of Pristichampsus Gervais, 1853". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 521–550. doi:10.1017/S1755691013000200. S2CID 128920027.

[R-80] Rossmann, T. (2002). "Studies on Cenozoic Crocodiles: 10. First Evidence of a Tomistomid Crocodilian (Eusuchia: Tomistomidae) from the Middle Eocene (Geiseltalien, MP11) of Grube Messel, Germany". Neues Jahrb. Geol. Paläontol. Monatsh. 2002 (3): 129–146. doi:10.1127/njgpm/2002/2002/129.

[list-81] Mayr, G. (2017). "The early Eocene birds of the Messel fossilsite: a 48 million-year-old bird community adds a temporal perspective to the evolution of tropical avifaunas". Biol. Rev. 92 (2): 1174–1188. doi:10.1111/brv.12274. PMID 27062331. S2CID 3464510.

[Li08-82] Mayr, G. (2008). "First substantial Middle Eocene record of the Lithornithidae (Aves): A postcranial skeleton from Messel (Germany)". Annales de Paléontologie. 94 (1): 29–37. doi:10.1016/j.annpal.2007.12.004.

[M14-83] Mayr, G. (2014). "The middle Eocene European "ratite" Palaeotis (Aves, Palaeognathae) restudied once more". Paläontologische Zeitschrift. 89 (3): 503–514. doi:10.1007/s12542-014-0248-y. S2CID 83787063.

[Hellmund2013-84] Hellmund M (2013). "Reappraisal of the bone inventory of Gastornis geiselensis (Fischer, 1978) from the Eocene Geiseltal Fossillagerstatte (Saxony-Anhalt, Germany)". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 269 (2): 203–220. doi:10.1127/0077-7749/2013/0345.

[85] Angst D.; Lécuyer C.; Amiot R.; Buffetaut E.; Fourel F.; Martineau F.; Legendre S.; Abourachid A.; Herrel A. (2014). "Isotopic and anatomical evidence of an herbivorous diet in the Early Tertiary giant bird Gastornis. Implications for the structure of Paleocene terrestrial ecosystems". Naturwissenschaften. 101 (4): 313–322. Bibcode:2014NW....101..313A. doi:10.1007/s00114-014-1158-2. PMID 24563098. S2CID 18518649.

[MG06-86] Mayr, Gerald (2006). "New specimens of the early Eocene stem group galliform Paraortygoides (Gallinuloididae), with comments on the evolution of a crop in the stem lineage of Galliformes". Journal of Ornithology. 147 (1): 31–37. Bibcode:2006JOrni.147...31M. doi:10.1007/s10336-005-0006-8. S2CID 30402641.

[Mayr14-87] Mayr, G. (2014). "The Eocene Juncitarsus – its phylogenetic position and significance for the evolution and higher-level affinities of flamingos and grebes". Comptes Rendus Palevol. 13 (1): 9–18. Bibcode:2014CRPal..13....9M. doi:10.1016/j.crpv.2013.07.005. ISSN 1631-0683.

[MG16-88] Mayr, G. (2016). "Fragmentary but distinctive: three new avian species from the early Eocene of Messel, with the earliest record of medullary bone in a Cenozoic bird". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 279 (3): 273–286. doi:10.1127/njgpa/2016/0555.

[Quasi-89] Gerald, M. (2004). "New specimens of Hassiavis laticauda (Aves: Cypselomorphae) and Quasisyndactylus longibrachis (Aves: Alcediniformes) from the Middle Eocene of Messel, Germany" (PDF). Cour. Forsch.-Inst. Senckenberg. 252: 23–28.

[PKM01-90] Mayr, G. (2001). "Comments on the osteology of Masillapodargus longipes Mayr 1999 and Paraprefica major Mayr 1999, caprimulgiform birds from the Middle Eocene of Messel (Hessen, Germany)". Neues Jahrbuch für Geologie und Paläontologie - Monatshefte. 2001 (2): 65–76. doi:10.1127/njgpm/2001/2001/65.

[GM03-91] Mayr, G. (2003). "A new Eocene swift-like bird with a peculiar feathering". Ibis. 145 (3): 382–391. doi:10.1046/j.1474-919x.2003.00168.x.

[FHotOW-92] Mayr, G. (2005). "Fossil hummingbirds in the Old World" (PDF). Biologist. 52 (1): 12–16.

[PKM05-93] Mayr, G. (2005). "The Palaeogene Old World potooParapreficaMayr, 1999 (Aves, Nyctibiidae): Its osteology and affinities to the New World Preficinae Olson, 1987". Journal of Systematic Palaeontology. 3 (4): 359–370. Bibcode:2005JSPal...3..359M. doi:10.1017/s1477201905001653. S2CID 85093005.

[PCMK-94] Gerald, M. (2005). "A New Cypselomorph Bird from the Middle Eocene of Germany and the Early Diversification of Avian Aerial Insectivores". The Condor. 107 (2): 342–352. doi:10.1093/condor/107.2.342.

[SSZ15-95] Mayr, G. (2015). "Skeletal morphology of the middle Eocene swift Scaniacypselus and the evolutionary history of true swifts (Apodidae)". J Ornithol. 156 (2): 441–450. Bibcode:2015JOrni.156..441M. doi:10.1007/s10336-014-1142-9. S2CID 14282557.

[MG17-96] Mayr, G. (2017). "A small, "wader-like" bird from the Early Eocene of Messel (Germany)". Annales de Paléontologie. 103 (2): 141–147. Bibcode:2017AnPal.103..141M. doi:10.1016/j.annpal.2017.01.001.

[M04-97] Morlo, M. (2004). "Diet of Messelornis (Aves: Gruiformes), an Eocene bird from Germany" (PDF). Courier Forschungsinstitut Senckenberg. 252: 29–33.

[MGM2002-98] Mayr, G. (2002). "A skull of a new pelecaniform bird from the Middle Eocene of Messel, Germany". Acta Palaeontologica Polonica. 47 (3).

[M02-99] Mayr, G. (2002). "A contribution to the osteology of the Middle Eocene ibis Rhynchaeites messelensis (Aves: Threskiornithidae: Rhynchaeitinae nov. subfam.)". Neues Jahrbuch für Geologie und Paläontologie - Monatshefte. 2002 (8): 501–512. doi:10.1127/njgpm/2002/2002/501.

[P92-100] Peters, D.S. (1992). "A new species of owl (Aves:Strigiformes) from the Middle Eocene Messel oil shale" (PDF). Papers in Avian Paleontology Honoring Pierce Brodkorp. 36: 161–169.

[MG05M-101] Mayr, G. (2005). "A new eocene Chascacocolius-like mousebird (Aves: Coliiformes) with a remarkable gaping adaptation". Organisms Diversity & Evolution. 5 (3): 167–171. Bibcode:2005ODivE...5..167M. doi:10.1016/j.ode.2004.10.013.

[MG00-102] Mayr, G.; Mourer-Chauviré, C. (2000). "Rollers (Aves: Coraciiformes s.s.) from the Middle Eocene of Messel (Germany) and the Upper Eocene of the Quercy (France)". Journal of Vertebrate Paleontology. 20 (3): 533–546. doi:10.1671/0272-4634(2000)020[0533:racssf]2.0.co;2. S2CID 56065258.

[BPZL21-103] Babarović, F.; Puttick, M. N.; Zaher, M.; Learmonth, E.; Gallimore, E.-J.; Smithwick, F. M.; Mayr, G.; Vinther, J. (2019). "Characterization of melanosomes involved in the production of non-iridescent structural feather colours and their detection in the fossil record". Journal of the Royal Society Interface. 16 (155). doi:10.1098/rsif.2018.0921. PMC 6597762. PMID 31238836. S2CID 195352503.

[MB98-104] Mayr, G.; Peters, D. S. (1998). "The mousebirds (Aves: Coliiformes) from the Middle Eocene of Grube Messel (Hessen, Germany)". Senckenbergiana Lethaea. 78 (1–2): 179–197. doi:10.1007/bf03042768. S2CID 82480792.

[GMGM01-105] Mayr, G. (2001). "A New Specimen of the Tiny Middle Eocene Bird Gracilitarsus Mirabilis (New Family: Gracilitarsidae)". The Condor. 103 (1): 78–84. doi:10.1093/condor/103.1.78.

[MT09-106] Mayr, G. (2009). "A well-preserved second trogon skeleton (Aves, Trogonidae) from the middle Eocene of Messel, Germany". Palaeobiodiversity and Palaeoenvironments. 89 (1–2): 1–6. Bibcode:2009PdPe...89....1M. doi:10.1007/s12549-009-0001-9. S2CID 73722121.

[Mayr2000-107] Mayr, G. (2000). "Tiny Hoopoe-Like Birds From the Middle Eocene of Messel (Germany)". The Auk. 117 (4): 964–970. doi:10.1093/auk/117.4.964.

[Mayr2006-108] Mayr, G. (2006). "New specimens of the Eocene Messelirrisoridae (Aves: Bucerotes), with comments on the preservation of uropygial gland waxes in fossil birds from Messel and the phylogenetic affinities of Bucerotes". Paläontol. Z. 80 (4): 390–405. doi:10.1007/BF02990211. S2CID 51778381.

[W06-109] Weidig, I. (2006). "The first New World occurrence of the Eocene birdPlesiocathartes (Aves: ?Leptosomidae)". Paläontologische Zeitschrift. 80 (3): 230–237. doi:10.1007/bf02988439. S2CID 140684204.

[MMCW-110] Mayr, G.; Mourer-Chauviré, C.; Weidig, I. (2004). "Osteology and systematic position of the eocene primobucconidae (aves, coraciiformes sensu stricto), with first records from Europe". Journal of Systematic Palaeontology. 2 (1): 1–12. Bibcode:2004JSPal...2....1M. doi:10.1017/S1477201903001093. S2CID 34962540.

[Selmes-111] Mayr, G. (2001). "New specimens of the Middle Eocene fossil mousebird Selmes absurdipes Peters 1999". Ibis. 143 (4): 427–434. doi:10.1111/j.1474-919x.2001.tb04943.x.

[Jarvis2014-112] Jarvis, E. D.; Mirarab, S.; Aberer, A. J.; Li, B.; Houde, P.; Li, C.; Ho, S. Y. W.; Faircloth, B. C.; Nabholz, B.; Howard, J. T.; Suh, A.; Weber, C. C.; Da Fonseca, R. R.; Li, J.; Zhang, F.; Li, H.; Zhou, L.; Narula, N.; Liu, L.; Ganapathy, G.; Boussau, B.; Bayzid, M. S.; Zavidovych, V.; Subramanian, S.; Gabaldon, T.; Capella-Gutierrez, S.; Huerta-Cepas, J.; Rekepalli, B.; Munch, K.; et al. (2014). "Whole-genome analyses resolve early branches in the tree of life of modern birds" (PDF). Science. 346 (6215): 1320–1331. Bibcode:2014Sci...346.1320J. doi:10.1126/science.1253451. hdl:10072/67425. PMC 4405904. PMID 25504713. Archived from the original (PDF) on 2015-02-24. Retrieved 2015-08-29.

[M2000-113] Mayr, G. (2000). "New or previously unrecorded avian taxa from the Middle Eocene of Messel (Hessen, Germany)". Foss. Rec. 3: 207–219. doi:10.1002/mmng.20000030110.

[CMM13-114] Mourer-Chauviré, Cécile (2013). "Idiornis Oberholser, 1899 (Aves, Gruiformes, Cariamae, Idiornithidae): a junior synonym of Dynamopterus Milne-Edwards, 1892 (Paleogene, Phosphorites du Quercy, France)". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 270 (1): 13–22. doi:10.1127/0077-7749/2013/0355.

[MG02-115] Mayr, G. (2002). "A new specimen ofSalmila robusta (Aves: Gruiformes: Salmilidae n. fam.) from the Middle Eocene of Messel". Paläontologische Zeitschrift. 76 (2): 305–316. Bibcode:2002PalZ...76..305M. doi:10.1007/BF02989866. S2CID 84345501.

[MR11-116] Mayr, G.; Richter, G. (2011). "Exceptionally preserved plant parenchyma in the digestive tract indicates a herbivorous diet in the Middle Eocene bird Strigogyps sapea (Ameghinornithidae)". Paläontologische Zeitschrift. 85 (3): 303–307. doi:10.1007/s12542-010-0094-5. S2CID 84479974.

[MGM06-117] Mayr, G. (2006). "A new raptorial bird from the Middle Eocene of Messel, Germany". Historical Biology. 18 (2): 99–106. doi:10.1080/08912960600640762. S2CID 34895565.

[MGM09-118] Mayr, G. (2009). "A well-preserved skull of the "falconiform" bird Masillaraptor from the middle Eocene of Messel (Germany)" (PDF). Palaeodiversity. 2: 315–320.

[M21-119] Mayr, Gerald (2021-06-01). "A partial skeleton of a new species of Tynskya Mayr, 2000 (Aves, Messelasturidae) from the London Clay highlights the osteological distinctness of a poorly known early Eocene "owl/parrot mosaic"". PalZ. 95 (2): 337–357. doi:10.1007/s12542-020-00541-8. ISSN 1867-6812.

[WDM06-120] Waterhouse, D. M. (2006). "Parrots in a nutshell: The fossil record of Psittaciformes (Aves)". Historical Biology. 18 (2): 227–238. doi:10.1080/08912960600641224. S2CID 83664072.

[PLQ-121] Mayr, G. (2016). "New species of Primozygodactylus from Messel and the ecomorphology and evolutionary significance of early Eocene zygodactylid birds (Aves, Zygodactylidae)". Historical Biology. 29 (7): 875–884. doi:10.1080/08912963.2016.1261135. S2CID 89599089.

[MZ09-122] Mayr, G.; Zelenkov, N. (2009). "New Specimens of Zygodactylid Birds from the Middle Eocene of Messel, with Description of a New Species of Primozygodactylus". Acta Palaeontologica Polonica. 54 (1): 15–20. doi:10.4202/app.2009.B103. S2CID 55370152.

[GM2015-123] Gerald Mayr (2015). "A reassessment of Eocene parrotlike fossils indicates a previously undetected radiation of zygodactyl stem group representatives of passerines (Passeriformes)". Zoologica Scripta. 44 (6): 587–602. doi:10.1111/zsc.12128. S2CID 85599482.

[ZS2015-124] Mayr, G. (2008). "Pumiliornis tessellatus Mayr, 1999 revisited — new data on the osteology and possible phylogenetic affinities of an enigmatic Middle Eocene bird". Paläontologische Zeitschrift. 82 (3): 247–253. doi:10.1007/bf02988891. S2CID 55757331.

[eopa16-125] Mayr, G. (2016). "Towards completion of the early Eocene aviary: A new bird group from the Messel oil shale (Aves, Eopachypterygidae, fam. nov.)". Zootaxa. 4013 (2): 252–264. doi:10.11646/zootaxa.4013.2.6. PMID 26623896.

[MG10-126] Mayr, G. (2010). "A New Avian Species with Tubercle-bearing Cervical Vertebrae from the Middle Eocene of Messel (Germany)". Records of the Australian Museum. 62: 21–28. doi:10.3853/j.0067-1975.62.2010.1537.

[VK90-127] Von Koenigswald, W. (1990). "Die Paläobiologie der Apatemyiden (Insectivora sl) und die Ausdeutung der Skelettfunde von Heterohyus nanus aus dem Mitteleozän von Messel bei Darmstadt". Palaeontographica. Abteilung A, Paläozoologie, Stratigraphie. 210 (1–3): 41–77.

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