Hornerstown Formation

Hornerstown Formation
Hornerstown Formation
Stratigraphic range: latest Maastrichtian-Danian ; 66.5–62 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Type	Geological formation
Unit of	Rancocas Group
Underlies	Vincentown Formation
Overlies	New Egypt Formation, Navesink Formation and Tinton Formation
Lithology
Primary	Marl, greensand
Location
Region	New Jersey
Country	United States
Type section
Named for	Hornerstown, New Jersey

Last updated January 31, 2025

The Hornerstown Formation is a latest Cretaceous to early Paleocene-aged geologic formation in New Jersey. It preserves a variety of fossil remains, including those of dinosaurs, and contains direct evidence of the mass mortality that occurred at the Cretaceous-Paleogene boundary.^[2]^[3]^[4]

Age & significance

The age of the Hornerstown deposits have been controversial. While most fossils are of animal taxa known from the earliest Cenozoic era, several fossils of otherwise exclusively Cretaceous age have been found. These include remains of the shark Squalicorax , several types of non-avian dinosaurs, the teleost fish Enchodus , several species of ammonite, and marine lizards referred to the genus Mosasaurus . Some of these remains show signs of severe abrasion and erosion, however, implying that they may be re-worked from older deposits. Most of these fossils are restricted to the lowest point in the formation, one rich in fossils and known as the Main Fossiliferous Layer, or MFL. Other explanations for the out-of-place fossils in the MFL is that they represent a time-averaged assemblage that built up and remained unburied during a time of low sediment deposition, or that they were stirred up from deeper in the sediment and deposited together during a tsunami.^[6] Biochemical analyses done on mosasaur bones from the Hornerstown Formation and the underlying, purely Cretaceous New Egypt Formation have found differing chemical signatures in the content of rare earth elements depending on whether the bones derive from the New Egypt or the Hornerstown Formation. This provides evidence against the idea that the presence of these remains in the Hornerstown is just the result of reworking, and supports the Hornerstown Formation including Cretaceous strata.^[7]

To account for these Cretaceous fossils, the Hornerstown Formation is generally treated as including the last portion of the Maastrichtian shortly before the Cretaceous-Paleogene boundary, and is thus divided into three sections: the section below the MFL (entirely Maastrichtian), the MFL itself (at the K-Pg boundary), and the section above the MFL (entirely Danian). However, other studies continue to treat the Hornerstown Formation as a Paleocene formation that saw significant reworking of Cretaceous fossils into itself.^[8] In addition, recent studies have found evidence of high iridium concentration and shocked quartz within the MFL, suggesting that the MFL represents a thanatocoenosis formed from an ecosystem collapse during the Cretaceous-Paleogene extinction event. This makes the Hornerstown Formation one of the few geological formations to contain direct evidence of the immediate impact of the extinction event. The Hornerstown Formation is important paleontologically, as it shows the impact of the K-Pg extinction on the coastal waters of eastern Appalachia, contains many of the last known records of taxa that went extinct at the K-Pg boundary, while also providing evidence of survivorship for the taxa that managed to survive the extinction event.^[3]

Vertebrate paleofauna

Cartilaginous fishes

Based on Case (1996) and Boles et al (2024):^[3]^[9]

Chimaeras

Chimaeras of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Edaphodon	E. agassizi		Danian		Tooth plates	A callorhinchid chimaera.
	E. mantelli		Danian
	E. mirificus		Maastrichtian
	E. stenobyrus		Maastrichtian
Ischyodus	I. bifurcatus		Maastrichtian, potentially Danian		Tooth plates, jaw elements	A callorhinchid chimaera.
	I. thurmanni		Maastrichtian
	I. williamsae		Danian

Sharks

Sharks of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Araloselachus	A. cuspidatus		Maastrichtian		Teeth	A sand shark.^[10]
Carcharias	C. teretidens		Danian			A relative of the sand tiger shark.
Cretalamna	C. appendiculata		Maastrichtian & Danian			A megatooth shark.
Heptranchias	H. howellii		Maastrichtian			A relative of the sharpnose sevengill shark.
Hexanchus	H. microdon		Danian			A sixgill shark.
Hexanchus	H. sp.		Maastrichtian & Danian			A sixgill shark.
Otodus	O. obliquus		Danian			A megatooth shark.
Odontaspis	O. sp.		Danian			A sand shark.
Palaeocarcharodon	P. sp.		Danian			A megatooth shark.
Palaeogaleus	P. vincenti		Maastrichtian & Danian			A houndshark.
Palaeohypotodus	P. rutoti		Danian			A sand shark.
Pseudocorax	P. affinis		Maastrichtian			A pseudocoracid shark.
Scapanorhynchus	S. texanus		Maastrichtian			A goblin shark.^[11]
Scyliorhinus	S. gilberti		Danian			A catshark.
Sphenodus	S. lundgreni		Maastrichtian			A orthacodontid shark.
Squalicorax	S. pristodontus		Maastrichtian			A crow shark.
Squalus	S. minor		Maastrichtian & Danian			A spurdog.
Squalus	S. sp		Maastrichtian & Danian			A spurdog.
Squatina	S. sp.		Maastrichtian			An angelshark.
Weltonia	W. ancistrodon		Danian			A cow shark.
Xampylodon	X. brotzeni		Maastrichtian			A cow shark.

Rays

Rays of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Aetomylaeus	A. striatus		Maastrichtian		Teeth	An eagle ray.^[10]
Dasyatis	D. crosswickense		Danian			A whiptail stingray. Type locality for species.
Hypolophites	H. hutchinsi		Danian			A whiptail stingray. Type locality for species.
Hypolophodon	H. sylvestris		Danian			A whiptail stingray.
Ischyrhiza	I. mira		Maastrichtian			A sclerorhynchid sawskate.^[10]
Rhinoptera	R. sp.		Maastrichtian			A cownose ray.
Rhombodus	R. laevis		Maastrichtian			A rhombodontid ray.
Viperecucullus	V. kuehnei		Danian			A whiptail stingray, type locality for the species and genus.

Ray-finned fishes

Actinopterygii of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Acipenser	A. cf. albertensis		Maastrichtian			A sturgeon.^[3]
Anomoeodus	A. phaseolus		Maastrichtian, potentially Danian		8 teeth	A pycnodont. One tooth is known from the Danian section of the formation, which would be the latest record of this genus and prove its occurrence in the Cenozoic; however, it may have been reworked from lower layers.^[3]
Atractosteus	A. sp.		Maastrichtian & Danian		11 teeth	A gar. First record of gars from eastern North America during the Paleocene.^[3]
cf. Bananogmius	cf. B. sp.		Maastrichtian			A plethodid tselfatiiform.^[3]^[10]
Dercetidae indet.			Maastrichtian & Danian		27 flank scales	A dercetid aulopiform. The first evidence in eastern North America of the Dercetidae surviving the K-Pg extinction event.^[3]
Enchodus	E. ferox		Maastrichtian			An enchodontid aulopiform. The E. gladiolus remains are the first record of the species in eastern North America.^[3]
Enchodus	E. gladiolus		Maastrichtian		8 teeth
Iridopristis	I. parrisi	Sewell, New Jersey	Danian		3 partial articulated specimens, including a near-complete skull.^[8]	A stem-lineage member of Holocentridae. The earliest known definitive holocentrid. Type locality for genus and species.^[8]
Paralbula	P. marylandica		Maastrichtian & Danian		23 teeth	A phyllodontid elopomorph. First evidence of P. marylandica existing during the Cretaceous, indicating that it survived the extinction event.^[3]
Phyllodus	P. paulkatoi		Maastrichtian		1 tooth plate	A phyllodontid elopomorph. First known occurrence in eastern North America.^[3]
Saurocephalus	S. lanciformis		Danian		1 tooth	A saurodontid ichthyodectiform. First known occurrence in eastern North America and first known occurrence of ichthyodectiforms as a whole in the Paleocene and Cenozoic.^[3]

Reptiles

Birds

A number of fossil birds are known from the greensands of the formation. The Hornerstown serves as the type locality for all these genera and species:

Birds of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Notes
Anatalavis	A. rex		?earliest Paleocene^[12]^[13]		A waterfowl potentially related to the magpie-goose.^[3]^[14]
Graculavus	G. velox				A potential wader.^[3]^[14]
Laornis	L. edwardsianus				A laornithid wading bird.^[3]^[14]
Novacaesareala	N. hungerfordi				A potential tropicbird.^[3]^[14]
Palaeotringa	P. littoralis				A potential wader.^[3]^[14]
Palaeotringa	P. vagans				A potential wader.^[3]^[14]
Telmatornis	T. priscus				A potential wader.^[3]^[14]
Tytthostonyx	T. glauconiticus				A potential seabird.^[3]^[14]

Non-avian dinosaurs

Dinosaurs of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Notes
Hadrosauridae indet.			Maastrichtian		A hadrosaurid ornithischian.^[3]

The tyrannosauroid Dryptosaurus is sometimes referred to this formation, as its remains were found in the New Egypt Formation, which is sometimes considered a part of the Hornerstown.^[15]

Crocodylomorphs

Crocodylomorphs of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Borealosuchus	B. threeensis		Maastrichtian		Lower jaw, postcranial remains	A eusuchian. Type locality of species.^[3]^[16]
Bottosaurus	B. harlani		Maastrichtian & Danian		Remains including lower jaw of a juvenile individual^[17]	An early caiman.^[3]^[17]
Bottosaurus	B. tuberculatus^[10]		Maastrichtian		Remains including lower jaw of a juvenile individual^[17]	An early caiman.^[3]^[17]
Hyposaurus	H. rogersii		Maastrichtian & Danian			A dyrosaurid.^[3]^[18]
cf. Procaimanoidea	P. sp.		Maastrichtian			A caiman.^[3]
Thoracosaurus	T. neocesariensis		Maastrichtian & Danian			A gavialoid.^[3]

Plesiosaurs

Plesiosaurs of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
" Plesiosaurus "	"P." brevifemur		Maastrichtian			An indeterminate plesiosaur known from a well-documented specimen, now lost.^[19]^[20]

Turtles

Testudines of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Notes	Images
Adocus	A. agilis^[19]		Maastrichtian		An adocid.^[3]
	A. beatus
	A. syntheticus^[19]
Agomphus	A. pectoralis		Maastrichtian		A kinosternoid related to Dermatemys .^[3]^[21]
Bothremys	B. sp.		Maastrichtian		A bothremydid side-necked turtle.^[3]
Euclastes	E. wielandi		Maastrichtian & Danian		A sea turtle.^[3]
Lytoloma	L. jeanesii		Maastrichtian		A sea turtle.^[19]
Osteopygis	O. emarginatus		Maastrichtian		A macrobaenid.^[19]
Peritresius	P. ornatus		Maastrichtian		A sea turtle.
Taphrosphys	T. sulcatus		Maastrichtian		A bothremydid side-necked turtle.^[3]
Taphrosphys	T. strenuus		Maastrichtian		A bothremydid side-necked turtle.^[3]

Mosasaurs

Mosasaurs of the Hornerstown Formation
Genus	Species	Location	Stratigraphic position	Abundance	Material	Notes	Images
Halisaurus	H. platyspondylus		Maastrichtian			A halisaurine. Type locality of the genus and species.
Mosasaurus	M. hoffmanni (=Nectoportheus validus)^[19]		Maastrichtian		Pterygoid, teeth, vertebrae^[3]^[7]	A mosasaurine. This formation contains some of the last known remains, comprising individuals killed during the mass mortality event that formed the MFL.^[3] A very large specimen is known.^[7]
Plioplatecarpus	P. sp.		Maastrichtian			A plioplatecarpine.

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References

↑ Gentry, A.D.; Kiernan, C.R.; Parham, J.F. (2022). "A large non-marine turtle from the Upper Cretaceous of Alabama and a review of North American "Macrobaenids"". The Anatomical Record. 306 (6): 1411–1430. doi:10.1002/ar.25054. PMID 37158131.
↑ Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 Boles, Zachary; Ullmann, Paul; Putnam, Ian; Ford, Mariele; Deckhut, Joseph (2024-04-12). "New vertebrate microfossils expand the diversity of the chondrichthyan and actinopterygian fauna of the Maastrichtian–Danian Hornerstown Formation in New Jersey". Acta Palaeontologica Polonica. doi: 10.4202/app.01117.2023 .
↑ Staron, Richard M.; Grandstaff, Barbara S.; Gallagher, William B.; Grandstaff, David E. (2001). "REE Signatures in Vertebrate Fossils from Sewell, NJ: Implications for Location of the K-T Boundary". PALAIOS. 16 (3): 255–265. doi:10.2307/3515603. ISSN 0883-1351.
↑ Voegele, Kristyn K.; Ullmann, Paul V.; Lonsdorf, Tara; Christman, Zachary; Heierbacher, Michael; Kibelstis, Brian J.; Putnam, Ian; Boles, Zachary M.; Walsh, Shane; Lacovara, Kenneth J. (2021-11-01). "Microstratigraphic Analysis of Fossil Distribution in the Lower Hornerstown and Upper Navesink Formations at the Edelman Fossil Park, NJ". Frontiers in Earth Science. 9: 1033. Bibcode:2021FrEaS...9.1033V. doi: 10.3389/feart.2021.756655 . ISSN 2296-6463.
↑ Gallagher, W.B. (2005). "Recent mosasaur discoveries from New Jersey and Delaware, USA: stratigraphy, taphonomy and implications for mosasaur extinction." Netherlands Journal of Geosciences — Geologie en Mijnbouw, 84(3): 241-245. Archived 2012-09-04 at the Wayback Machine
1 2 3 Gallagher, W. B. (2015). "Greensand mosasaurs of New Jersey and the Cretaceous–Paleogene transition of marine vertebrates". Netherlands Journal of Geosciences. 94 (1): 87–91. Bibcode:2015NJGeo..94...87G. doi:10.1017/njg.2014.30. ISSN 0016-7746.
1 2 3 Andrews, J. V.; Schein, J. P.; Friedman, M. (2023). "An earliest Paleocene squirrelfish (Teleostei: Beryciformes: Holocentroidea) and its bearing on the timescale of holocentroid evolution". Journal of Systematic Palaeontology. 21 (1). Bibcode:2023JSPal..2168571A. doi: 10.1080/14772019.2023.2168571 .
↑ Case, Gerard R. (1996-12-19). "A new Selachian Fauna from the Lower Hornerstown Formation (Early Paleocene/Montian) of Monmouth County, New Jersey". Palaeontographica Abteilung A. 242 (1–3): 1–14. Bibcode:1996PalAA.242....1C. doi:10.1127/pala/242/1996/1.
1 2 3 4 5 "PBDB Collection". paleobiodb.org. Retrieved 2024-11-12.
↑ "PBDB Collection". paleobiodb.org. Retrieved 2024-11-12.
↑ Olson, S. (1994). "A giant Presbyornis (Aves: Anseriformes) and other birds from the Paleocene Aquia Formation of Maryland and Virginia". Proceedings of the Biological Society of Washington. 107: 429–435. hdl:10088/6493.
↑ Brocklehurst, Neil; Upchurch, Paul; Mannion, Philip D.; O'Connor, Jingmai (2012-06-25). "The Completeness of the Fossil Record of Mesozoic Birds: Implications for Early Avian Evolution". PLOS ONE. 7 (6): e39056. Bibcode:2012PLoSO...739056B. doi: 10.1371/journal.pone.0039056 . ISSN 1932-6203. PMC 3382576 . PMID 22761723.
1 2 3 4 5 6 7 "3.22 New Jersey, United States; 7. Hornerstown Formation," in Weishampel, et al. (2004). Page 586.
↑ Olsson, R.K. (1960). "Foraminifera of latest Cretaceous and earliest Tertiary age in the New Jersey coastal plain". Journal of Paleontology. 34: 1–58.
↑ Brochu, Christopher A.; Parris, David C.; Grandstaff, Barbara Smith; Denton, Robert K.; Gallagher, William B. (2012). "A new species of Borealosuchus (Crocodyliformes, Eusuchia) from the Late Cretaceous–early Paleogene of New Jersey". Journal of Vertebrate Paleontology. 32 (1): 105–116. Bibcode:2012JVPal..32..105B. doi:10.1080/02724634.2012.633585. ISSN 0272-4634.
1 2 Cossette, A. P., and C. A. Brochu. 2018. A new specimen of the alligatoroid Bottosaurus harlani and the early history of character evolution in alligatorids. Journal of Vertebrate Paleontology. DOI:10.1080/02724634.2018.1486321.
↑ Morgan, Donald J (December 2018). "PRESENCE OF A DYROSAURID NEOSUCHIAN IN THE SEVERN/BRIGHTSEAT FORMATION OF MARYLAND". The Journal of the Delaware Valley Paleontological Society. X: 91 to 104.
1 2 3 4 5 6 "PBDB Collection". paleobiodb.org. Retrieved 2024-11-12.
↑ Parris, David C. (1974). "Additional Records of Plesiosaurs from the Cretaceous of New Jersey". Journal of Paleontology. 48 (1): 32–35. ISSN 0022-3360. JSTOR 1303102.
↑ "PBDB Taxon". paleobiodb.org. Retrieved 2024-11-11.

Bibliography

Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. 861 pp. ISBN 0-520-24209-2.

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Text is available under the CC BY-SA 4.0 license; additional terms may apply.
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[GKP22-1] Gentry, A.D.; Kiernan, C.R.; Parham, J.F. (2022). "A large non-marine turtle from the Upper Cretaceous of Alabama and a review of North American "Macrobaenids"". The Anatomical Record. 306 (6): 1411–1430. doi:10.1002/ar.25054. PMID 37158131.

[dinosaurdistribution-2] Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607.

[:0-3] 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 Boles, Zachary; Ullmann, Paul; Putnam, Ian; Ford, Mariele; Deckhut, Joseph (2024-04-12). "New vertebrate microfossils expand the diversity of the chondrichthyan and actinopterygian fauna of the Maastrichtian–Danian Hornerstown Formation in New Jersey". Acta Palaeontologica Polonica. doi: 10.4202/app.01117.2023 .

[4] Staron, Richard M.; Grandstaff, Barbara S.; Gallagher, William B.; Grandstaff, David E. (2001). "REE Signatures in Vertebrate Fossils from Sewell, NJ: Implications for Location of the K-T Boundary". PALAIOS. 16 (3): 255–265. doi:10.2307/3515603. ISSN 0883-1351.

[5] Voegele, Kristyn K.; Ullmann, Paul V.; Lonsdorf, Tara; Christman, Zachary; Heierbacher, Michael; Kibelstis, Brian J.; Putnam, Ian; Boles, Zachary M.; Walsh, Shane; Lacovara, Kenneth J. (2021-11-01). "Microstratigraphic Analysis of Fossil Distribution in the Lower Hornerstown and Upper Navesink Formations at the Edelman Fossil Park, NJ". Frontiers in Earth Science. 9: 1033. Bibcode:2021FrEaS...9.1033V. doi: 10.3389/feart.2021.756655 . ISSN 2296-6463.

[gallagher2005-6] Gallagher, W.B. (2005). "Recent mosasaur discoveries from New Jersey and Delaware, USA: stratigraphy, taphonomy and implications for mosasaur extinction." Netherlands Journal of Geosciences — Geologie en Mijnbouw, 84(3): 241-245. Archived 2012-09-04 at the Wayback Machine

[:1-7] 1 2 3 Gallagher, W. B. (2015). "Greensand mosasaurs of New Jersey and the Cretaceous–Paleogene transition of marine vertebrates". Netherlands Journal of Geosciences. 94 (1): 87–91. Bibcode:2015NJGeo..94...87G. doi:10.1017/njg.2014.30. ISSN 0016-7746.

[EarliestPaleoceneSquirrelfish-8] 1 2 3 Andrews, J. V.; Schein, J. P.; Friedman, M. (2023). "An earliest Paleocene squirrelfish (Teleostei: Beryciformes: Holocentroidea) and its bearing on the timescale of holocentroid evolution". Journal of Systematic Palaeontology. 21 (1). Bibcode:2023JSPal..2168571A. doi: 10.1080/14772019.2023.2168571 .

[:3-9] Case, Gerard R. (1996-12-19). "A new Selachian Fauna from the Lower Hornerstown Formation (Early Paleocene/Montian) of Monmouth County, New Jersey". Palaeontographica Abteilung A. 242 (1–3): 1–14. Bibcode:1996PalAA.242....1C. doi:10.1127/pala/242/1996/1.

[:4-10] 1 2 3 4 5 "PBDB Collection". paleobiodb.org. Retrieved 2024-11-12.

[11] "PBDB Collection". paleobiodb.org. Retrieved 2024-11-12.

[12] Olson, S. (1994). "A giant Presbyornis (Aves: Anseriformes) and other birds from the Paleocene Aquia Formation of Maryland and Virginia". Proceedings of the Biological Society of Washington. 107: 429–435. hdl:10088/6493.

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