Paleobiota of the Solnhofen Formation

Solnhofen Limestone
Solnhofen Limestone
Stratigraphic range: Kimmeridgian-Tithonian[1][2]
	A brittle star fossil from the Solnhofen Formation.
Type	Geological formation
Underlies	Usseltal Formation
Overlies	Torleite Formation
Lithology
Primary	Lithographic limestone[1]
Location
Region	Bavaria
Country	Germany
Type section
Named for	Solnhofen

The Solnhofen Plattenkalk, a collective term of multiple Late Jurassic lithographic limestones in southeastern Germany, is famous for its well preserved fossil flora and fauna dating to the late Jurassic.

Ardeosaurus brevipes

Platychelys oberndorferi

A. leptospondylus tail

Alligatorellus beaumonti

Scaphognathus crassirostris

Rhamphorhynchus muensteri

Cycnorhamphus suevicus

Ctenochasma elegans

Reptiles
Group	Genus	Species
Lizards	Ardeosaurus	A. brevipes
	Ardeosaurus	A. digitalellus
	Bavarisaurus	B. macrodactylus
	Eichstaettisaurus	E. schroederi
	Schoenesmahl	S. dyspepsia[3]
Turtles	Eurysternum	E. wagleri [4]
	Cyrtura	C. temnospondyla
	Idiochelys	I. fitzingeri
	Palaeomedusa	P. testa
	Platychelys	P. oberndorferi
	Solnhofia	S. parsonsi
Rhynchocephalia	Homoeosaurus	H. maximillioni
	Sapheosaurus	S. laticeps
	Kallimodon	K. pulchellus
	Oenosaurus	O. muehlheimensis
	Vadasaurus	V. herzogi
	Pleurosaurus	P. goldfussi[5]
Ichthyosauria	Aegirosaurus	A. leptospondylus[6]
Ichthyosauria	"Ichthyosaurus"	"I." posthumus
Crocodylomorpha	Alligatorellus	A. beaumonti[7]
	Cricosaurus	C. elegans [8]
	Dakosaurus	D. maximus[8][9]
	Geosaurus	G. giganteus[10]
	Rhacheosaurus	R. gracilis[10][8]
Dinosauria	Archaeopteryx	A. siemensii [11]
	Archaeopteryx	A. lithographica[12]
	Compsognathus	C. longipes[12]
Pterosauria	Aerodactylus	A. scolopaciceps[13]
	Anurognathus	A. ammoni
	Ardeadactylus	A. longicollum[14]
	Aurorazhdarcho	A. micronyx
	Ctenochasma	C. elegans
	Cycnorhamphus	C. suevicus
	Germanodactylus	G. cristatus
	Germanodactylus	G. rhamphastinus
	Gnathosaurus	G. subulatus
	Pterodactylus	P. antiquus[15]
	Rhamphorhynchus	R. muensteri[16]
	Scaphognathus	S. crassirostris[17]

Lepidotes maximus

Lepidotes notopterus

Propterus

Gyrodus circularis

Gyrodus hexagonus

Mesturus verrucosus

Hypsocormus

Ophiopsis

Aspidorhynchus

Pachythrissops

Thrissops

Living Elops individual.

Pholidophorus

Fish
Group	Genus	Species
Chondrichthyes	Asteracanthus	A. ornatissimus [18]
Chondrichthyes	Paracestracion	P. falcifer
Chondrostei	Coccolepis	C. bucklandi[19]
Halecostomi	Lepidotes	L. decoratus[20]
		L. intermedius[20]
		L. maximus[20]
		L. notopterus[20]
		L. oblongus[20]
		L. pustulosus[20]
		L. subovatus[20]
		L. unguiculatus[20]
	Histionotus	H. oberndorferi[21]
	Macrosemius	M. rostratus[21]
	Notagogus	N. decoratus[21]
	Notagogus	N. denticulatus
	Propterus	P. elongatus[21]
	Propterus	P. microstomus[21]
	Eusemius	E. beatae[21]
	Eomesodon	E. gibbosus[22]
	Gyrodus	G. circularis[22]
	Gyrodus	G. hexagonus[22]
	Mesturus	M. verrucosus[22]
	Macromesodon	M. heckeli[22]
	Macromesodon	M. macropterus[22]
	Piranhamesodon	P. pinnatomus[23]
	Proscinetes	P. elegans[22]
	Proscinetes	P. formosus[22]
	Asthenocormus	A. titanius[24]
	Hypsocormus	H. insignis[24]
	Hypsocormus	H. macrodon[24]
	Orthocormus	O. cornutus[24]
	Pseudoasthenocormus	P. retrodorsalis[25]
	Sauropsis	S. depressus[24]
	Sauropsis	S. longimanus[24]
	Heterostrophus	H. latus[26]
Halecomorphi	Amblysemius	A. bellicianus[27]
	Amblysemius	A. pachyurus[27]
	Caturus	C. furcatus[27]
	Caturus	C. giganteus[27]
	Liodesmus	L. gracilis[27]
	Liodesmus	L. sprattiformis[27]
	Ionoscopus	I. cyprinoides[28]
		I. esocinus[28]
		I. münsteri[28]
		I. striatissimus[28]
	Furo	F. angustus[29]
		F. latimanus[29]
		F. longiserratus[29]
		F. microlepidotes[29]
	Amiopsis	A. lepidota[30]
	Ophiopsis	O. muensteri
	Ophiopsiella	O. attenuta[30]
	Ophiopsiella	O. procera[30]
	Solnhofenamia	S. elongata[30]
	Brachyichthys	B. radiato-punctatus[31]
	Brachyichthys	B typicus[31]
	Callopterus	C. agassizzi[31]
	Lophiurus	L. minutus[31]
Teleostei	Pleuropholis[32]	P. laevissima[32]
		P. pompecki[32]
		P. wagneri[32]
	Aspidorhynchus[33]	A. acutirostris[33]
	Belonostomus[33]	B. münsteri[33]
	Belonostomus[33]	B. tenuirostris[33]
	Ankylophorus[34]	Indeterminate.[34]
	Eurycormus[34]	E. speciosus[34]
	Pholidophoristion[34]	P. micronyx[34]
	Pholidophoristion[34]	P. ovatus[34]
	Pholidophorus[34]	P. armatus[34]
		P. diagonalis[34]
		P. falcifer[34]
		P. macrocephalus[34]
		P. microps[34]
		P. sculptus[34]
	Allothrissops[35]	A. mesogaster[35]
	Allothrissops[35]	A. salmoneus[35]
	Pachythrissops[35]	P. propterus[35]
	Thrissops[35]	T. formosus[35]
	Thrissops[35]	T. subovatus[35]
	Leptolepides[36]	L. haerteisi[36]
	Leptolepides[36]	L. sprattiformis[36]
	Orthogonikleithrus[36]	O. hoelli[36]
	Orthogonikleithrus[36]	O. leichi[36]
	Anaethalion[37]	A. angustissimus[37]
		A. angustus[37]
		A. knorri[37]
		A. subovatus[37]
	Ascalabos[38]	A. voithii[38]
	Daitingichthys[38]	D. tischlingeri[38]
	Eichstättia[38]	E. mayri[38]
	Elops[38]	cf. E. sp. 1[38]
		cf. E. sp. 2[38]
		cf. E. sp. 3[38]
	Tharsis[38]	T. dubius[38]
	Tischlingerichtys[38]	T. viohli[38]

Saccocoma tenella

Belemnotheutis

Mesolimulus

Mesurupetala - Houston Museum of Natural Science

Invertebrates
Group	Genus	Species
Crinoids	Saccocoma[39]
Cephalopods	Belemnotheutis mayri[40]
	Plesioteuthis
	Physodoceras[41]
	Fontannesiella[41]
Horseshoe Crabs	Mesolimulus walchi
Thylacocephalans	Clausocaris	C. lithographica
	Dollocaris	D. michelorum
	Mayrocaris	M. bucculata
	Falcatacaris	F. bastelbergeri
Insects	Mesurupetala

References

Rauhut, O. W., Heyng, A. M., López-Arbarello, A., & Hecker, A. (2012). A new rhynchocephalian from the Late Jurassic of Germany with a dentition that is unique amongst tetrapods. PLoS ONE, 7(10): e46839.
Schweigert, G. 2007. Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions." Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 245(1): 117-125. doi:10.1127/0077-7749/2007/0245-0117
Conrad, Jack L. (2018-06-28). "A new lizard (Squamata) was the last meal of Compsognathus (Theropoda: Dinosauria) and is a holotype in a holotype". Zoological Journal of the Linnean Society. 183 (3): 584–634. doi:10.1093/zoolinnean/zlx055. ISSN 0024-4082.
JOYCE, W. G. (2003). A new Late Jurassic turtle specimen and the taxonomy of Palaeomedusa testa and edusa testa Eurysternum wagleri.
Dupret, V. (2004). The pleurosaurs: anatomy and phylogeny. Revue de Paléobiologie, Vol spéc, 9, 61-80.
Bardet, N., & Fernández, M. (2000). A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria. Journal Information, 74(3).
Schwarz‐Wings, D., Klein, N., Neumann, C., & Resch, U. (2011). A new partial skeleton of Alligatorellus (Crocodyliformes) associated with echinoids from the Late Jurassic (Tithonian) lithographic limestone of Kelheim, S‐Germany. Fossil Record, 14(2), 195-205.
De Andrade, M. B., Young, M. T., Desojo, J. B., & Brusatte, S. L. (2010). The evolution of extreme hypercarnivory in Metriorhynchidae (Mesoeucrocodylia: Thalattosuchia) based on evidence from microscopic denticle morphology. Journal of Vertebrate Paleontology, 30(5), 1451-1465.
Young MT, Brusatte SL, de Andrade MB, Desojo JB, Beatty BL, et al. (2012) The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe. PLoS ONE 7(9): e44985. doi:10.1371/journal.pone.0044985
Young, Mark T., and Marco Brandalise de Andrade, 2009. "What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern, Germany." Zoological Journal of the Linnean Society, 157: 551-585.
Mayr, G.; Phol, B.; Hartman, S.; Peters, D. S. (2007). "The tenth skeletal specimen of Archaeopteryx". Zoological Journal of the Linnean Society. 149: 97–116. doi:10.1111/j.1096-3642.2006.00245.x.
Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Jurassic, Europe)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 545–549. ISBN 0-520-24209-2.
Vidovic SU, Martill DM (2014) Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny. PLoS ONE 9(10): e110646. doi:10.1371/journal.pone.0110646
Bennett, S.C. (2002). "Soft tissue preservation of the cranial crest of the pterosaur Germanodactylus from Solnhofen." Journal of Vertebrate Paleontology, 22(1): 43-48.
Jouve, S. (2004). "Description of the skull of a Ctenochasma (Pterosauria) from the latest Jurassic of eastern France, with a taxonomic revision of European Tithonian Pterodactyloidea." Journal of Vertebrate Paleontology, 24(3): 542-554.
Bennett, S. C. (1995). "A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: Year-classes of a single large species." Journal of Paleontology, 69: 569-580.
"Scaphognathus." In: Cranfield, Ingrid (ed.). The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 308-309.
Stumpf, Sebastian; López‐Romero, Faviel A.; Kindlimann, René; Lacombat, Frederic; Pohl, Burkhard; Kriwet, Jürgen (2021). "A unique hybodontiform skeleton provides novel insights into Mesozoic chondrichthyan life". Papers in Palaeontology. 7 (3): 1479–1505. doi:10.1002/spp2.1350. S2CID 234204226.
"Table 1; Chondrostei; Coccolepidae" in Lambers (1999) pg. 216.
"Table 1; Halecostomi; Semionotidae" in Lambers (1999) pg. 216.
"Table 1; Halecostomi; Macrosemiidae" in Lambers (1999) pg. 216.
"Table 1; Halecostomi; Pycnodontiformes" in Lambers (1999) pg. 216.
Kölbl-Ebert; Ebert; Bellwood; Schulbert, Martina; Martin; David R.; Christian (October 18, 2018). "A Piranha-like Pycnodontiform Fish from the Late Jurassic". Current Biology. 28 (21): 3516–3521.e2. doi:10.1016/j.cub.2018.09.013. PMID 30344113. S2CID 53045425.{{cite journal}}: CS1 maint: multiple names: authors list (link)
"Table 1; Halecostomi; Pachycormidae" in Lambers (1999) pg. 216.
Note: Lambers abbreviated Pseudoasthenocormus as "Pseudoasth.," "Table 1; Halecostomi; Pachycormidae" in Lambers (1999) pg. 216.
"Table 1; Halecostomi; 'Halecostomi inc. sed.'" in Lambers (1999) pg. 216.
"Table 1; Halecomorphi; Caturidae" in Lambers (1999) pg. 216.
"Table 1; Halecomorphi; Ionoscopidae" in Lambers (1999) pg. 216.
"Table 1; Halecomorphi; Furidae" in Lambers (1999) pg. 216.
"Table 1; Halecomorphi; 'Amiidae' and 'Ophiopsidae'" in Lambers (1999) pg. 216.
"Table 1; Halecomorphi; Halecomorphi incertae cedis" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Pleuropholidae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Aspidorhynchidae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Pholidophoridae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Allothrissopidae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Orthogonikleithridae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; Anaethalionidae" in Lambers (1999) pg. 216.
"Table 1; Teleostei; 'Elopidae,' 'Ostariophysi,' 'Clupeocephala inc. sed.,' 'Elopocephala inc. sed.,' and 'Teleostei inc. sed'" in Lambers (1999) pg. 216.
Clare V. Milsom. (1994) Saccacoma, a benthic crinoid from the Jurassic Solnhofen Limestone, Germany Archived 2011-07-16 at the Wayback Machine. Palaeontology 37 (1) 121-129.
Lomax, Dean R. (2010). "Am Ichthyosaurus (Reptilia, Ichthyosauria) with gastric contents from Charmouth, England: First report of the genus from the Pliensbachian" (PDF). Paludicola. Rochester Institute of Vertebrate Paleontology. 8 (1): 22–36. Archived from the original (PDF) on July 26, 2012. Retrieved February 8, 2011.
Keupp, Helmut (July 2007). "Complete ammonoid jaw apparatuses from the Solnhofen plattenkalks: Implications for aptychi function and microphagous feeding of ammonoids". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 245: 93–101. doi:10.1127/0077-7749/2007/0245-0093.

General

Lambers, P. H. (1999). The actinopterygian fish fauna of the Late Kimmeridgian and Early Tithonian 'Plattenkalke' near Solnhofen (Bavaria, Germany): state of the art. Geologie en Mijnbouw 78:215-229.

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[rhyncho2012-1] Rauhut, O. W., Heyng, A. M., López-Arbarello, A., & Hecker, A. (2012). A new rhynchocephalian from the Late Jurassic of Germany with a dentition that is unique amongst tetrapods. PLoS ONE, 7(10): e46839.

[ammonitedate-2] Schweigert, G. 2007. Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions." Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 245(1): 117-125. doi:10.1127/0077-7749/2007/0245-0117

[:0-3] Conrad, Jack L. (2018-06-28). "A new lizard (Squamata) was the last meal of Compsognathus (Theropoda: Dinosauria) and is a holotype in a holotype". Zoological Journal of the Linnean Society. 183 (3): 584–634. doi:10.1093/zoolinnean/zlx055. ISSN 0024-4082.

[turtles-4] JOYCE, W. G. (2003). A new Late Jurassic turtle specimen and the taxonomy of Palaeomedusa testa and edusa testa Eurysternum wagleri.

[pleurosaurs-5] Dupret, V. (2004). The pleurosaurs: anatomy and phylogeny. Revue de Paléobiologie, Vol spéc, 9, 61-80.

[aegirosaurus-6] Bardet, N., & Fernández, M. (2000). A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria. Journal Information, 74(3).

[7] Schwarz‐Wings, D., Klein, N., Neumann, C., & Resch, U. (2011). A new partial skeleton of Alligatorellus (Crocodyliformes) associated with echinoids from the Late Jurassic (Tithonian) lithographic limestone of Kelheim, S‐Germany. Fossil Record, 14(2), 195-205.

[hypercarnivory-8] De Andrade, M. B., Young, M. T., Desojo, J. B., & Brusatte, S. L. (2010). The evolution of extreme hypercarnivory in Metriorhynchidae (Mesoeucrocodylia: Thalattosuchia) based on evidence from microscopic denticle morphology. Journal of Vertebrate Paleontology, 30(5), 1451-1465.

[9] Young MT, Brusatte SL, de Andrade MB, Desojo JB, Beatty BL, et al. (2012) The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe. PLoS ONE 7(9): e44985. doi:10.1371/journal.pone.0044985

[Young&Andrade2009-10] Young, Mark T., and Marco Brandalise de Andrade, 2009. "What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern, Germany." Zoological Journal of the Linnean Society, 157: 551-585.

[10th_find-11] Mayr, G.; Phol, B.; Hartman, S.; Peters, D. S. (2007). "The tenth skeletal specimen of Archaeopteryx". Zoological Journal of the Linnean Society. 149: 97–116. doi:10.1111/j.1096-3642.2006.00245.x.

[latejurassicdistribution-12] Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Jurassic, Europe)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 545–549. ISBN 0-520-24209-2.

[13] Vidovic SU, Martill DM (2014) Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny. PLoS ONE 9(10): e110646. doi:10.1371/journal.pone.0110646

[bennett2002-14] Bennett, S.C. (2002). "Soft tissue preservation of the cranial crest of the pterosaur Germanodactylus from Solnhofen." Journal of Vertebrate Paleontology, 22(1): 43-48.

[jouve2004-15] Jouve, S. (2004). "Description of the skull of a Ctenochasma (Pterosauria) from the latest Jurassic of eastern France, with a taxonomic revision of European Tithonian Pterodactyloidea." Journal of Vertebrate Paleontology, 24(3): 542-554.

[bennett1995-16] Bennett, S. C. (1995). "A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: Year-classes of a single large species." Journal of Paleontology, 69: 569-580.

[cranfield-scaphognathus-17] "Scaphognathus." In: Cranfield, Ingrid (ed.). The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 308-309.

[18] Stumpf, Sebastian; López‐Romero, Faviel A.; Kindlimann, René; Lacombat, Frederic; Pohl, Burkhard; Kriwet, Jürgen (2021). "A unique hybodontiform skeleton provides novel insights into Mesozoic chondrichthyan life". Papers in Palaeontology. 7 (3): 1479–1505. doi:10.1002/spp2.1350. S2CID 234204226.

[lamberstable-chondrostei-19] "Table 1; Chondrostei; Coccolepidae" in Lambers (1999) pg. 216.

[lamberstable-semionotidae-20] "Table 1; Halecostomi; Semionotidae" in Lambers (1999) pg. 216.

[lamberstable-macrosemiidae-21] "Table 1; Halecostomi; Macrosemiidae" in Lambers (1999) pg. 216.

[lamberstable-pycnodontiformes-22] "Table 1; Halecostomi; Pycnodontiformes" in Lambers (1999) pg. 216.

[:1-23] Kölbl-Ebert; Ebert; Bellwood; Schulbert, Martina; Martin; David R.; Christian (October 18, 2018). "A Piranha-like Pycnodontiform Fish from the Late Jurassic". Current Biology. 28 (21): 3516–3521.e2. doi:10.1016/j.cub.2018.09.013. PMID 30344113. S2CID 53045425.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[lamberstable-pachycormidae-24] "Table 1; Halecostomi; Pachycormidae" in Lambers (1999) pg. 216.

[lamberstable-pseudoasth-25] Note: Lambers abbreviated Pseudoasthenocormus as "Pseudoasth.," "Table 1; Halecostomi; Pachycormidae" in Lambers (1999) pg. 216.

[lamberstable-halecostomi-is-26] "Table 1; Halecostomi; 'Halecostomi inc. sed.'" in Lambers (1999) pg. 216.

[lamberstable-caturidae-27] "Table 1; Halecomorphi; Caturidae" in Lambers (1999) pg. 216.

[lamberstable-ionoscopidae-28] "Table 1; Halecomorphi; Ionoscopidae" in Lambers (1999) pg. 216.

[lamberstable-furidae-29] "Table 1; Halecomorphi; Furidae" in Lambers (1999) pg. 216.

[lamberstable-others-30] "Table 1; Halecomorphi; 'Amiidae' and 'Ophiopsidae'" in Lambers (1999) pg. 216.

[lamberstable-halecomorphi-ic-31] "Table 1; Halecomorphi; Halecomorphi incertae cedis" in Lambers (1999) pg. 216.

[lamberstable-pleuropholidae-32] "Table 1; Teleostei; Pleuropholidae" in Lambers (1999) pg. 216.

[lamberstable-aspidorhynchidae-33] "Table 1; Teleostei; Aspidorhynchidae" in Lambers (1999) pg. 216.

[lamberstable-pholidophoridae-34] "Table 1; Teleostei; Pholidophoridae" in Lambers (1999) pg. 216.

[lamberstable-allothrissopidae-35] "Table 1; Teleostei; Allothrissopidae" in Lambers (1999) pg. 216.

[lamberstable-orthogonikleithridae-36] "Table 1; Teleostei; Orthogonikleithridae" in Lambers (1999) pg. 216.

[lamberstable-anaethalionidae-37] "Table 1; Teleostei; Anaethalionidae" in Lambers (1999) pg. 216.

[lamberstable-teleostei-others-38] "Table 1; Teleostei; 'Elopidae,' 'Ostariophysi,' 'Clupeocephala inc. sed.,' 'Elopocephala inc. sed.,' and 'Teleostei inc. sed'" in Lambers (1999) pg. 216.

[saccocoma-milsom-39] Clare V. Milsom. (1994) Saccacoma, a benthic crinoid from the Jurassic Solnhofen Limestone, Germany Archived 2011-07-16 at the Wayback Machine. Palaeontology 37 (1) 121-129.

[lomax-40] Lomax, Dean R. (2010). "Am Ichthyosaurus (Reptilia, Ichthyosauria) with gastric contents from Charmouth, England: First report of the genus from the Pliensbachian" (PDF). Paludicola. Rochester Institute of Vertebrate Paleontology. 8 (1): 22–36. Archived from the original (PDF) on July 26, 2012. Retrieved February 8, 2011.

[:2-41] Keupp, Helmut (July 2007). "Complete ammonoid jaw apparatuses from the Solnhofen plattenkalks: Implications for aptychi function and microphagous feeding of ammonoids". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 245: 93–101. doi:10.1127/0077-7749/2007/0245-0093.