Lourinhã Formation
At its upper limit, the formation includes some Early Cretaceous (Berriasian) strata, but it mostly consists of Late Jurassic (Kimmeridgian/Tithonian) rocks. The stratigraphy of the formation is generally complex and controversial, with the classification and grouping of its lithostratigraphic units varying between different stratigraphers.
The fossil biota present there is very similar to that of the Morrison Formation in the United States and, to a lesser extent, the Tendaguru Formation in Tanzania. There are also similarities to the Villar del Arzobispo Formation in Spain, and the neardy Alcobaça Formation.
The formation is named after the municipality of Lourinhã, the geographic area it mostly occupies.
Lithology and depositional history
Depositional history
The Lourinhã Formation is located within the Lusitanian Basin, a mostly onshore North South orientated rift basin within western Portugal, formed during the Opening of the North Atlantic Ocean, with sediment deposition beginning during the Late Triassic-Early Jurassic. It primarily consists of syn-rift near-coastal continental siliciclastic sediments, with several marine intercalations. The primary flow direction was North to South, originating from Galicia and flowing between the Iberian landmass to the east and the now largely submerged Berlengas horst, a north–south oriented ridge, to the west.
Stratigraphy
The unit was first formally proposed by Hill in 1988.
The stratigraphy of the Lourinhã Formation is complex and varies between sub-basins with several competing stratigraphic proposals and there is currently no consensus on the matter, one of the most recent stratigraphies, divides the formation into three members which are from oldest to youngest the Praia da Amoreira-Porto Novo Member, Praia Azul Member, and the Assenta Member.
Praia da Amoreira-Porto Novo Member
The Praia da Amoreira-Porto Novo Member is composed of the Priaia de Amoreira Member, which consists of massive mudrock-sand with metre thick sandstone lenses, with massive mudrock with calcrete. The overlying Poto Novo Mb. consists of massive bodies of sandstone, often cross bedded. The environment of deposition is interpreted as a meandering fluvial system, while the Porto Novo Mb is interpreted as a deltaic deposit. It is interpreted to be latest Kimmeridgian in age, and overlies the Consolacao Unit at the top of the Aulacostephanus eudoxus ammonite zone.
Praia Azul Member
The Praia Azul Member, formerly known as the Sobral unit/member is 80 to 130 metres thick and consists of tabular marls and mudstones, with rare sandstones bodies. There are three distinct laterally extensive (>20 km) thin shelly carbonate horizons within this member, indicating brief marine transgressions. South of Santa Cruz primarily consists of sandstone with rare conglomerate. The age is considered to be latest Kimmeridgian to earliest Tithonian, correlated to the ammonite zones of Hybonoticeras beckeri and Hybonoticeras hybonotum.
Santa Rita Member + Assenta Member
The Santa Rita Member in the Consolação sub-basin and its lateral equivalent in the Turcifal Basin the Assenta Member is around 300 metres thick and predominantly consists of mudstones with frequent layers of caliche. Near the top of the member several layers of tens of metres thick nodular and marly bioclastic limestones are present, containing marine benthic forams, the nodularity is derived from intense Thalassinoides burrowing. It is predominately late Tithonian in age, with the last few metres probably being earliest Berriasian, with the top of the formation roughly correlative with the base of the magnetochron M18n. (~144.7 Ma) The environment of deposition is interpreted as being an upper fluvial-dominated delta to meandering fluvial systems flowing on a paralic plain.
Fauna
Dinosaurs
In a 2003 study, an analysis of all Portuguese dinosaurs was published. The study created a cladogram showing the possible relations of all Portuguese dinosaurs, including those at the time known from the Lourinhã Formation.
Ornithischia
Color key
|
Notes Uncertain or tentative taxa are in small text; |
Ornithopods
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
| Ankylopollexia indet. | Indeterminate |
|
Appendicular, axial, and cranial elements | ||
|
Intermediate |
Limb material |
Now referred to its own genus, Draconyx, along with some other material. |
|||
|
D. loureiroi |
Praia Azul Member |
One partial skeleton |
 | ||
| Dryosauridae indet. | Indeterminate | Praia Azul Member | Appendicular, axial, and cranial elements | ||
|
D. sp. |
Praia Azul Member |
 | |||
| Eousdryosaurus | E. nanohallucis | Praia da Amoreira-Porto Novo Formation | A partial postcranial skeleton. | ||
| Hesperonyx | H. martinhotomasorum | Porto Novo Member | Bones from the fore- and hindlimbs | A dryomorphan iguanodontian with uncertain affinities |  |
|
Ornithopoda indet. |
Intermediate |
A single track |
Gigantic track indicating an ornithopod with a hip height of 2.5 meters. No known Jurassic Ornithopod reaches this size; only known evidence for such sizes in this group at the time. Found alongside Deltapodus print. |
||
|
P. henkelli. |
Two teeth |
Color key
|
Notes Uncertain or tentative taxa are in small text; |
Thyreophorans
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
|
D. armatus |
A stegosaurid |  | |||
|
D. brodricki |
Eleven tracks; Nine pes and two manus prints. |
The tracks can be separated into three different morphologies, though all fall within range of the Ornithopodsociation of the pes and manus tracks to the same taxon cannot be directly supported. Preserve various well preserved skin impressions. Largest prints are larger than those from the type horizon. The tracks are individually represented and do not form any sort of trackway, thought one print is associated with giant ornithopod track, potentially representing that the creatures were traveling together or were otherwise going to a similar location. Another is similarly associated with theropod and sauropod prints. |
|||
| Dracopelta | D. zbyszewskii | Assenta Member | An ankylosaur. | ||
| Miragaia | M. longicollum |
|
Holotype, neck, partial skull, forelimbs, ribs. Tentative juvenile specimen assigned to this taxon. | Stegosaur with unusually long neck of 17 cervicals, with more neck vertebrae than most sauropods. Probably a junior synonym of Dacentrurus. |  |
Color key
|
Notes Uncertain or tentative taxa are in small text; |
Other Ornithiscians
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
|
T. cuneatus |
Amoreira-Porto Novo Member |
3 isolated teeth. | |||
|
A. kuehnei |
Saurischia
Color key
|
Notes Uncertain or tentative taxa are in small text; |
Sauropods
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
| D. lourinhanensis | Praia da Amoreira-Porta Novo Member | One specimen. Vertebrae; potentially other parts of the body. | A diplodocid. Tschopp et al. (2015) sunk the genus into Supersaurus. |    | |
|
Diplodocidae indet. |
Intermediate |
One Dorsal Vertebra |
Regarded by Mannion et al. (2012) as being unique from Dinheirosaurus and possibly indicating another diplodocid in the formation, but being non-diagnostic it doesn't warrant description. | ||
|
L. alenquerensis |
Praia Azul Member | A partial postcranial skeleton. | Possibly a Camarasaurid Macronarian. | ||
|
L. atalaiensis |
Praia Azul Member |
Fragmentary material. |
A large brachiosaur, a close relative of Brachiosaurus proper. | ||
| Oceanotitan | O. dantasi | Praia da Amoreira-Porta Novo Member | scapula, almost all of the pelvis, a complete leg sans the toes, and nine caudals. | A titanosauriform | |
| Zby | Z. atlanticus | Amoreira-Porto Novo Member | Holotype: Tooth, cervical neutral arch, forelimb, various other fragments. | A turiasaur. |
Theropods
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
| Abelisauridae indet. | Intermediate | Praia da Amoreira-Porto Novo Member | Teeth; ML 966, Ml 327. | Potentially diagnostic abelisaur teeth. | |
|
A. europaeus |
Praia Azul Member |
Two specimens, covering much of the body. |
Only European species of Allosaurus. |
 | |
|
A. fragilis |
Praia Azul Member |
Two specimens, covering much of the body. |
Now thought to represent a specimen of A. europaeus. |
||
|
C. sp. |
Praia da Amoreira-Porto Novo Member |
Four specimens; teeth, a femur. |
Potentially a synonym of the type species, C. nasicornis. Sometimes referred to as C. sp., giving indication of possible distinctiveness or of being intermediate. |
 | |
| Dendroolithidae | Indeterminant | Fragments of multiple eggs in a clutch, with associated embryonic remains. | Probably eggs of Torvosaurus. | ||
| Lusovenator | L. santosi |
|
Two partial postcranial skeletons. | Earliest known Carcharodontosaurian from Laurasia. | |
|
L. antunesi |
Praia Azul Member |
Three individuals, one largely complete; over 100 eggs with significant amount of skeletal material. |
Has come out in various places in the tree, erroneously said to be a megalosaur, mostly accepted to be a carnosaur, probably allosauroid, or basal coelurosaur. Currently unstable on the tree. |
  | |
|
M. insignis |
Teeth. |
Invalid. Teeth belong to various other theropod taxa. |
|||
|
M. pombali |
Teeth. |
Invalid. Teeth belong to various other theropod taxa |
|||
|
M. sp. |
Tooth fragment. |
Invalid; Dubious. |
|||
|
R. aff. gilmorei |
Tooth; ML 939 |
Only definite record of this taxon is from the Late Cretaceous of North America, despite erroneous and referrals from other sites in Portugal. Probably a close relative of Richardoestesia and not an actual representation of the taxon. |
|||
|
T. gurneyi |
|
Maxilla, Teeth, Femur; Egg clutch and embryos. |
Largest known European theropod. Previous known as Portugal populations of the type species, or as T. sp., before description in early 2014. |
 | |
|
T. tanneri |
Praia da Amoireira-Porto Novo |
Maxilla, Teeth, Femur; Probable embryo. |
Now described as a distinct species of Torvosaurus, T. gurneyi. Sometimes referred to as T. sp. in the past. |
Pterosaurs
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
| Dsungaripteroidea indet. | Indeterminate | Proximal right femur | |||
| Rhamphorhynchidae | Indeterminate | Tooth | |||
| Lusognathus | L. almadrava | Skull and cervical vertebrae | Ctenochasmatid |  | |
| Pteraichnus | Partial tracks |
Mammaliaformes
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
|
Haldanodon expectatus |
Partial skeleton and isolated bones |
Semi-aquatic forager. |
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
|
Nanolestes drescherae |
Right lower molar. |
Amphitheriidae; small omnivore or insectivore. | ||
|
Guimarotodus inflatus |
Right mandible. |
Dryolestidae; insectivore or omnivore. | ||
|
Krebsotherium lusitanicum |
Left mandible. |
Dryolestidae; insectivore or omnivore. | ||
|
Drescheratherium acutum |
Upper jaw. |
Paurodontidae; herbivore. |
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
| Kuehneodon | K. hahni | A member of the family Paulchoffatiidae |
Amphibans
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
| Celtedens | Indeterminate | Porto Novo/Praia da Amoreira, Praia Azul | Frontal bones, along with other parts of the skull and limbs | An albanerpetontid. |
| Urodela | Indeterminate | Atlas vertebra | A salamander, suggested to belong to Scapherpetontidae. | |
| "Discoglossidae" | Indeterminate | Partial left humerus | A primitive frog |
Squamates
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
| Paramacellodidae | Indeterminate | Frontal and dentary bones | Scincomorph lizard |
Crocodyliformes
| Genus | Species | Member | Material | Notes | |
|---|---|---|---|---|---|
| Ophiussasuchus | O. paimogonectes | Praia Azul Member | Partial skull | Goniopholidid | |
| Goniopholididae | Indeterminate | Teeth and partial skeleton. | An aquatic neosuchian | ||
| Bernissartiidae | Tooth | ||||
| Lusitanisuchus | Teeth, and partial skull and jaw fragments | A mesoeucrocodylian of uncertain placement | |||
| Atoposauridae | Teeth |  | |||
| Mesoeucrocodylia | Teeth | Distinct from Lusitanisuchus |
Fish
| Genus | Species | Member | Material | Notes |
|---|---|---|---|---|
| Hybodus | H. cf. reticulatus | Teeth | A hybodontid shark | |
| Pycnodontiformes | Indeterminate | |||
| Lepidotes sensu lato | A ginglymodian | |||
| Caturus | An amiiform |
Flora
| Genus | Species | Member | Material | Notes | Images |
|---|---|---|---|---|---|
| Pterophyllum | P. mondeguensis | Bennettitales leaf | |||
| Otozamites | Bennettitales leaf | ||||
| Cupressinocladus | Conifer leaves | ||||
| Protocupressinoxylon | Conifer wood | ||||
| Prototaxoxylon | |||||
| Classopollis | Pollen of Cheirolepidiaceae conifers |
Correlation
See also
- Camadas de Alcobaça
- Camadas de Guimarota
- List of fossil sites
- Museu da Lourinhã
- List of dinosaur bearing rock formations
References
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- ^ Araújo, R., Castanhinha R., Martins R. M. S., Mateus O., Hendrickx C., Beckmann F., Schell N., & Alves L. C. (2013). "Filling the gaps of dinosaur eggshell phylogeny: Late Jurassic Theropod clutch with embryos from Portugal" (PDF). Scientific Reports. 3: 1924. doi:10.1038/srep01924. PMC 3667465. PMID 23722524.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Milàn, J; Christiansen, P; Mateus, O. "A three-dimensionally preserved sauropod manus impression from the Upper Jurassic of Portugal: implications for sauropod manus shape and locomotor mechanics". Kaupia. 14: 47–52.
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- ^ Ogg, J. G. (2012-01-01), Gradstein, Felix M.; Ogg, James G.; Schmitz, Mark D.; Ogg, Gabi M. (eds.), "Chapter 5 - Geomagnetic Polarity Time Scale", The Geologic Time Scale, Boston: Elsevier, pp. 85–113, ISBN 978-0-444-59425-9, retrieved 2021-12-17
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{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Rotatori, Filippo Maria; Moreno-Azanza, Miguel; Mateus, Octávio (2022-05-07). "Reappraisal and new material of the holotype of Draconyx loureiroi (Ornithischia: Iguanodontia) provide insights on the tempo and modo of evolution of thumb-spiked dinosaurs". Zoological Journal of the Linnean Society. 195 (1): 125–156. doi:10.1093/zoolinnean/zlab113. ISSN 0024-4082.
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{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Escaso, F.; Ortega, F.; Dantas, P.; Malafaia, E.; Silva, B.; Sanz, J.L. (2007). "Elementos postcraneales de Dacentrurus (Dinosauria: Stegosauria) del Jurásico Superior de Moçafaneira (Torres Vedras, Portugal)" (PDF). Libro de resúmenes V Encuentro de Jóvenes Investigadores en Paleontología: 157–172.
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{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Mateus, O., Maidment S., & Christiansen N. (2009). "A new long-necked 'sauropod-mimic' stegosaur and the evolution of the plated dinosaurs". Proceedings of the Royal Society B: Biological Sciences. 276 (1663): 1815–21. doi:10.1098/rspb.2008.1909. PMC 2674496. PMID 19324778.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Costa, Francisco; Mateus, Octávio (2019-11-13). "Dacentrurine stegosaurs (Dinosauria): A new specimen of Miragaia longicollum from the Late Jurassic of Portugal resolves taxonomical validity and shows the occurrence of the clade in North America". PLOS ONE. 14 (11): e0224263. Bibcode:2019PLoSO..1424263C. doi:10.1371/journal.pone.0224263. ISSN 1932-6203. PMC 6853308. PMID 31721771.
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- ^ Mateus, O., Mannion P. D., & Upchurch P. (2014). "Zby atlanticus, a new turiasaurian sauropod (Dinosauria, Eusauropoda) from the Late Jurassic of Portugal". Journal of Vertebrate Paleontology. 34 (3): 618–634. Bibcode:2014JVPal..34..618M. doi:10.1080/02724634.2013.822875. S2CID 59387149.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Tschopp, E.; Mateus, O. V.; Benson, R. B. J. (2015). "A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda)". PeerJ. 3: e857. doi:10.7717/peerj.857. PMC 4393826. PMID 25870766.
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{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Mateus, O., Walen A. and Antunes M. T. (2006). "The large theropod fauna of the Lourinhã Formation (Portugal) and its similarity to the Morrison Formation, with a description of a new species of Allosaurus". New Mexico Museum of Natural History and Science Bulletin. 36: 123–129.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Malafaia, E.; Ortega, F.; Escaso, F.; Dantas, P.; Pimentel, N.; Gasulla, J. M.; Ribeiro, B.; Barriga, F.; Sanz, J. L. (2010-12-10). "Vertebrate fauna at the Allosaurus fossil-site of Andrés (Upper Jurassic), Pombal, Portugal". Journal of Iberian Geology. 36 (2): 193–204. doi:10.5209/rev_jige.2010.v36.n2.7.
- ^ Allosauruseuropaeus. archosaur.us
- ^ Mateus, O. and Antunes M. T. (2000). Ceratosaurus sp. (Dinosauria: Theropoda) in the Late Jurassic of Portugal. Abstract volume of the 31st International Geological Congress. Rio de Janeiro, Brazil
- ^ Malafaia, Elisabete; Ortega, Francisco; Escaso, Fernando; Silva, Bruno (2015-10-03). "New evidence of Ceratosaurus (Dinosauria: Theropoda) from the Late Jurassic of the Lusitanian Basin, Portugal". Historical Biology. 27 (7): 938–946. doi:10.1080/08912963.2014.915820. S2CID 129349509.
- ^ Elisabete Malafaia; Pedro Mocho; Fernando Escaso; Francisco Ortega (2020). "A new carcharodontosaurian theropod from the Lusitanian Basin: evidence of allosauroid sympatry in the European Late Jurassic". Journal of Vertebrate Paleontology. 40: e1768106. doi:10.1080/02724634.2020.1768106.
- ^ Lourinhanosaurusantunesi. archosaur.us
- ^ Hendrickx, Christophe; Mateus, Octávio (2014). "Torvosaurus gurneyi n. sp., the Largest Terrestrial Predator from Europe, and a Proposed Terminology of the Maxilla Anatomy in Nonavian Theropods". PLOS ONE. 9 (3): e88905. Bibcode:2014PLoSO...988905H. doi:10.1371/journal.pone.0088905. PMC 3943790. PMID 24598585.
- ^ Malafaia, E.; Mocho, P.; Escaso, F.; Ortega, F. (2017-03-01). "New data on the anatomy of Torvosaurus and other remains of megalosauroid (Dinosauria, Theropoda) from the Upper Jurassic of Portugal". Journal of Iberian Geology. 43 (1): 33–59. doi:10.1007/s41513-017-0003-9. ISSN 1886-7995. S2CID 132198140.
- ^ Torvosaurusgurneyi. archosaur.us
- ^ Bertozzo, Filippo; Camilo da Silva, Bruno; Martill, David; Vorderwuelbecke, Elsa; Aureliano, Tito; Schouten, Remmert; Aquino, Pedro (2021). "A large pterosaur femur from the Upper Jurassic (Lusitanian Basin) of Portugal". Acta Palaeontologica Polonica. 66. doi:10.4202/app.00858.2020. ISSN 0567-7920.
- ^ Guillaume, Alexandre Renaud Daniel Microvertebrates of the Lourinhã Formation (Late Jurassic, Portugal) (2018) PhD thesis https://run.unl.pt/handle/10362/58236
- ^ Mateus (2010). "First records of crocodyle and pterosaur tracks in the Upper Jurassic of Portugal". New Mexico Museum of Natural History and Science Bulletin. 51: 83–87.
- ^ Guillaume, Alexandre R. D.; Natário, Carlos; Mateus, Octávio; Moreno-Azanza, Miguel (2023-04-03). "Plasticity in the morphology of the fused frontals of Albanerpetontidae (Lissamphibia; Allocaudata)". Historical Biology. 35 (4): 537–554. doi:10.1080/08912963.2022.2054712. ISSN 0891-2963.
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