Land vertebrate faunachrons (LVFs) are biochronological units used to correlate and date terrestrial sediments and fossils based on their tetrapod faunas. [1] First formulated on a global scale by Spencer G. Lucas in 1998, LVFs are primarily used within the Triassic Period (252 - 201 Ma), [1] though Lucas later designated LVFs for other periods as well. [2] Eight worldwide LVFs are defined for the Triassic. The first two earliest Triassic LVFs, the Lootsbergian and Nonesian, are based on South African synapsids and faunal assemblage zones estimated to correspond to the Early Triassic. These are followed by the Perovkan and Berdyankian, based on temnospondyl amphibians and Russian assemblages estimated to be from the Middle Triassic. The youngest four Triassic LVFs, the Otischalkian, Adamanian, Revueltian, and Apachean, are based on aetosaur and phytosaur reptiles common in the Late Triassic of the southwestern United States. [1] [3] [4]
The LVF system, though widely used, is also a controversial application of biostratigraphy, as many Triassic tetrapods are rife with complications which endanger their utility as index fossils. Limited occurrences, inaccurate age estimates, overlapping LVF faunas, or taxonomic disagreement may jeopardize global correlations between Triassic tetrapods. This could render some LVFs as misleading assessments of Triassic faunal change through time. [5] [6] [7] [8] Regardless, Late Triassic phytosaurs are considered to have strong biostratigraphic utility even among detractors of Lucas's system. [9]
Tetrapod biostratigraphy has been used for the Triassic of South Africa since 1906 and Argentina since 1966, [6] but without much connection to global faunas. [1] Starting in 1993, New Mexico Museum of Natural History and Science paleontologist Spencer G. Lucas and his colleagues began to define tetrapod biostratigraphy intervals in the Triassic of China [10] and eastern [11] and western [12] North America. These named biostratigraphic intervals were inspired by the Land Mammal Age (LMA) system already in use for Cenozoic faunal assemblages.
Triassic tetrapod biozones, under the term "land vertebrate faunachrons" (LVFs) were formalized on a global level by Lucas in 1998. They were diagnosed by a primary index fossil (a particular genus of widespread time-constrained tetrapod) and characterized by a faunal type assemblage (distinguishing collection of taxa) from a fossiliferous geological formation. Together, the defining index fossil and assemblage could be used to correlate fossil assemblages worldwide. [1] Updates to this system have been published continuously for Triassic LVFs, which remain a heavily-discussed topic in the study of Triassic chronology. [13] [3] [14] [4] Lucas has also defined LVFs for the Permian, [2] [15] [16] Jurassic, [17] and Carboniferous, [18] though these are not as widely used as his Triassic LVFs.
Later authors characterized Lucas's LVFs as "interval eubiochrons". This means that they correspond to a segment of time (and strata) between two paleobiological events: the first appearance datum (FAD) of one index taxon and the FAD of another. [9] A first appearance datum is a point in the geological record with the earliest known fossil of a given animal, which can estimate when that animal speciates or evolves into existence. As an example, the Lootsbergian LVF is defined as the period of time between the FAD (estimated speciation) of Lystrosaurus and the FAD (estimated speciation) of Cynognathus . Some taxa which are index fossils for one stage may persist into a later stage. [1]
LVFs of the Triassic Period from youngest to oldest:
Several paleontologists have independently questioned the validity of Lucas’s system, criticizing its inconsistent and often contradictory approach to taxonomy and faunal correlations. [5] [6] [7] [8]
Many index taxa are very rare or endemic to a single continent, and have no relevance to a global biostratigraphy system. These include Doswellia , Longosuchus , Typothorax , “ Pseudopalatus ” ( Machaeroprosopus ), Redondasaurus , and Redondasuchus , among others. [5] [6] [7] [21] For the Berdyankian LVF, very few species are shared between the index assemblage (the Bukobay Formation of Russia) and other correlated assemblages. Direct relationships between Russian, German, and South American dicynodonts are conjectural and based on undiagnostic European fragments. [5] [7]
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Index taxa for a given LVF often range into the succeeding LVF, blurring the distinction between the two time periods. Angistorhinus , Hyperodapedon , Paleorhinus / Parasuchus (all Otischalkian index taxa) range into the Adamanian, fossils referred to Rutiodon (an Adamanian index taxon) range into the Revueltian, and Metoposaurus can be found throughout the Otischalkian, Apachian, and Revueltian LVFs. [5] [6] [7] [21]
Lucas's approach to correlating LVFs with global marine stages has been met with criticism. The Triassic timescale is under constant revision from a series of age dating methods, including magnetostratigraphy, cyclostratigraphy, radiometric dating, and biozones of marine invertebrates such as conodonts and ammonoids. [8] [21] However, there are only a few areas where fossils of Triassic land tetrapods and marine organisms overlap, mostly restricted to coastal sediments in central Europe. Palynomorph and conchostracan biozones can help correlate terrestrial strata to an extent. One complication is that Lucas's view of the Late Triassic time scale contradicts the consensus established by other biostratigraphers. Most paleontologists estimate that the three stages of the Late Triassic (Carnian, Norian, and Revueltian) are strongly unequal in size, with the Norian far longer than the Carnian. Under this consensus "long-Norian" hypothesis, the Carnian-Norian boundary is close to 228 Ma. Lucas, on the other hand, prefers a "short-Norian" perspective, with a lengthier Carnian stage and a Carnian-Norian boundary at around 220 Ma. [8]
For example, Lucas has maintained that the lower part of the Chinle Formation (the Blue Mesa Member and equivalent units) is Carnian (>220 Ma) in age. This was justified by the assumption that fossils of Stagonolepis , a European aetosaur, can be found in North and South America, allowing correlation between these regions. However, this proposed widespread occurrence of Stagonolepis is a debatable, as many species assigned to the genus may not be closely related (see below). [1] [3] [6] [21]
According to the "short-Norian" interpretation, these lower Chinle Formation, and other strata of the Adamanian LVF, would be firmly pre-Norian in age, suggesting that any taxonomic change between the Adamanian and Revueltian represents a Carnian-Norian extinction event. However, the consensus "long-Norian" interpretation firmly places Adamanian strata of North America into the Norian stage (<228 Ma). The Norian age of the lower Chinle Formation has been independently confirmed by U-Pb dating and magnetostratigraphic correlations to global time scales. [8] [27] [28] Conversely, other "Adamanian" strata, such as fossiliferous layers in the lower Ischigualasto Formation of Argentina, can be assigned to the late Carnian (~231 Ma). [6] [29] This supports the conclusion that LVFs such as the Adamanian are fraught with uncertain time estimates brought on by weak correlations on a global scale. [6] [8]
Some authors have elected to ignore LVFs in favor of older and more localized biostratigraphic units. Named tetrapod assemblages zones (AZs) were well-established for the Triassic of Gondwana prior to the LVF, and recent updates have helped to constrain these units with greater clarity and agreement than global correlations. In Argentina, Bonaparte (1966) established the Chanarian (named after the Chañares Formation) and the Ischigualastian (named after the Ischigualasto Formation). Equivalents faunas are easily traced across Brazil, Africa, and India. These two biostratigraphic zones correlate with Lucas's Berdyankian, Otischalkian, and Apachean LVFs, but do not precisely overlap in time with those LVFs. Moreover, aetosaurs and phytosaurs, which are common in the Northern Hemisphere, are rarer and more scattered in the Southern Hemisphere. As a result, Gondwanan assemblage zones are defined by more common Southern taxa. For example, the Ischigualastian zone is defined by the rhynchosaur Hyperodapedon and the cynodont Exaeretodon, as well as the aetosaur Aetosauroides and herrerasaurid dinosaurs. [6]
Some correlations are based on connections between fragmentary or poorly-constrained taxa rather than direct correlations between type assemblages or LVF-defining index taxa. For example, the Ermaying Formation of China is correlated with the Moenkopi Formation of the United States via a tenuous (and likely unjustifiable) comparison between proposed erythrosuchid fossils. The primary index fossil of the Perovkan LVF, Eocyclotosaurus , is absent from China. [5] [7]
One particularly contradictory index fossil is Mastodonsaurus , the defining index fossil of the Berdyankian LVF. Fossils referable to this genus can be found across Ladinian-age Europe, but the proposed Russian species (M. torvus) may be unrelated to the endemic German type species (M. giganteus). Moreover, if one approaches Mastodonsaurus from a broader taxonomic perspective (as expected if M. torvus is included), they must also incorporate Anisian and Carnian material referred to the genus, including the small species “Heptasaurus” cappelensis . This precludes any reason to use Mastodonsaurus as a time-constrained index taxon. [5] [7]
Some LVFs are based on evolutionary grades as index taxa. This ignores the potential for high diversity and long temporal ranges within a given grade, and may lead to arbitrary and subjective inclusion or exclusion of descendant taxa. “ Stagonolepis ”, in its broadest form, is a wastebasket taxon of basal aetosaurs ranging through the Otischalkian and Apachean. Lucas’s usage of Stagonolepis lumps in many genera separated by other authors, such as Aetosauroides and Calyptosuchus . [6] [7] A similar situation occurs in Paleorhinus/Parasuchus, which has historically been used as a persistent grade of early phytosaurs. On the other hand, the characteristic phytosaur (Redondasaurus) and aetosaur (Redondasaurus) genera of the Apachean LVF are very similar to, and perhaps synonymous with, index taxa of the underlying Revueltian LVF: “Pseudopalatus” (Machaeroprosopus) and Typothorax, respectively. [5] [7]
Although the utility of a global LVF system is questionable, LVF-derived biostratigraphy may be useful in limited circumstances. Phytosaurs in particular have played a large role in the tetrapod biostratigraphy of the Chinle and Dockum Group of the southwest United States. A revision of the LVF system in this narrow context was undertaken by Jeff Martz and Bill Parker (2017), retaining several names and concepts previously used by Lucas and colleagues. [9]
Martz and Parker argued that the term "faunachron" was misleading and redundant, as each "faunachron" is bound by a single taxon rather than an assemblage (fauna) of multiple taxa. They preferred using a specific type of interval biozone known as a teilzone, referring to a local interval of strata equivalent to an interval of time. [9] [30] The base of each teilzone was marked by the Lowest known Occurrence (LOk) of a particular category of phytosaur, i.e. the oldest layer where fossils of that category are found in the study area. LOks are local points in time and stratigraphy, disregarding occurrences in other regions or the estimated time of speciation. For the Otischalkian, Adamanian, and Revueltian, the top of each teilzone is marked by the LOk of a more exclusive subgroup of phytosaurs. The top of the Apachean is marked by the LOk of Protosuchus , an Early Jurassic crocodylomorph, as with Lucas's system. [9]
"Faunachrons" could also be defined beyond the constraints of teilzones; other biozonation categories include holochronozones (a stratigraphic interval, involving multiple study areas) and holochrons (an estimated time interval, involving the time of speciation or immigration into the region). Each phytosaur-based "faunachron" could be considered a teilzone (in local biostratigraphy), an estimated holochronozone (in regional chronostratigraphy), or an estimated holochron (in regional biochronology). [9]
One complication in defining biozones based on phytosaurs is instability in phytosaur systematics. Many proposed phytosaur taxa are dubious, paraphyletic (such as Leptosuchus and Machaeroprosopus) or have unclear relationships to each other. Nevertheless, a series of nested clades is apparent in most recent overviews. Rather than relying on a single index taxon per biozone, Martz and Parker allowed multiple representatives per a given stage of phytosaur evolution. These representatives were chosen based on their occurrence in the southwest United States, commonness, and relatively stable phylogenetic position despite paraphyly in some circumstances. [9]
Although most LVFs or equivalent concepts are not marked by major biotic changes, one exception is apparent in the southwest United States. The boundary between the Adamanian and Revueltian zones is marked by a faunal turnover, an event where several tetrapod species quickly disappear from the fossil record as others appear for the first time. At Petrified Forest National Park, the event occurs in the Jim Camp Wash beds. This sediment layer is positioned in the middle of the Chinle Formation's Sonsela Member, and would have been deposited around 215 million years ago. Trilophosaurus , Poposaurus , Desmatosuchus , dicynodonts, and non-mystriosuchin phytosaurs are extirpated from the area around this time, while metoposaurs and allokotosaurs [31] as a whole decline in abundance. New species of aetosaurs and phytosaurs replaced losses across the purported boundary event. Palynomorph assemblages overturn to more dry adapted species, and a higher concentration of pedogenic carbonate nodules may also support increasing aridity. [21] [31]
The cause and relevance of this turnover is debatable, as it may indicate only a small localized extinction. The Manicouagan Impact, the second-largest bolide impact of the Mesozoic Era (besides the Chicxulub Impact which caused the K-Pg Mass Extinction at 66 Ma), is dated to around 215.4 Ma. [32] [33] While certainly large enough to momentarily devastate areas near the impact point in Quebec, broader environmental effects of the Manicouagan impact are mostly conjectural. [8] Besides the Adamanian-Revueltian turnover, the impact has also been linked to a minor marine extinction in eastern Panthalassa. [34]
Alternatively, the Adamanian-Revueltian turnover may be a consequence of the gradual aridification of western Pangea as it drifted north into arid latitudes. [31] Comparative estimates of extinction rates and occurrences find little support for a synchronized Adamanian-Revueltian turnover, and instead support a model where extinctions are stretched out over several million years. For most species, extinction probabilities are "decoupled" in time from other species, as well as geological or climatological drivers. The only plausible correlation is between the Manicouagan Impact and palynomorph turnover, and even then the probability of synchronicity is only about 34%. [35]
Phytosaurs are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. and are sometimes referred to as parasuchians. Phytosauria, Parasuchia, Parasuchidae, and Phytosauridae have often been considered equivalent groupings containing the same species. Some recent studies have offered a more nuanced approach, defining Parasuchidae and Phytosauridae as nested clades within Phytosauria as a whole. Phytosaurs were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodilians in size, appearance, and lifestyle, as an example of convergence or parallel evolution. The name "phytosaur" means "plant reptile", as the first fossils of phytosaurs were mistakenly thought to belong to plant eaters.
The Carnian is the lowermost stage of the Upper Triassic Series. It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by the Norian. Its boundaries are not characterized by major extinctions or biotic turnovers, but a climatic event occurred during the Carnian and seems to be associated with important extinctions or biotic radiations. Another extinction occurred at the Carnian-Norian boundary, ending the Carnian age.
Aetosaurs are heavily armored reptiles belonging to the extinct order Aetosauria. They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs more closely related to crocodilians than to birds and other dinosaurs. All known aetosaurs are restricted to the Late Triassic, and in some strata from this time they are among the most abundant fossil vertebrates. They have small heads, upturned snouts, erect limbs, and a body ornamented with four rows of plate-like osteoderms. Aetosaur fossil remains are known from Europe, North and South America, parts of Africa, and India. Since their armoured plates are often preserved and are abundant in certain localities, aetosaurs serve as important Late Triassic tetrapod index fossils. Many aetosaurs had wide geographic ranges, but their stratigraphic ranges were relatively short. Therefore, the presence of particular aetosaurs can accurately date a site in which they are found.
The Chinle Formation is an Upper Triassic continental geological formation of fluvial, lacustrine, and palustrine to eolian deposits spread across the U.S. states of Nevada, Utah, northern Arizona, western New Mexico, and western Colorado. In New Mexico, it is often raised to the status of a geological group, the Chinle Group. Some authors have controversially considered the Chinle to be synonymous to the Dockum Group of eastern Colorado and New Mexico, western Texas, the Oklahoma panhandle, and southwestern Kansas. The Chinle Formation is part of the Colorado Plateau, Basin and Range, and the southern section of the Interior Plains. A probable separate depositional basin within the Chinle is found in northwestern Colorado and northeastern Utah. The southern portion of the Chinle reaches a maximum thickness of a little over 520 meters (1,710 ft). Typically, the Chinle rests unconformably on the Moenkopi Formation.
Acaenasuchus is an extinct genus of pseudosuchian, endemic to what would be presently be known as Arizona during the Late Triassic, specifically during the Carnian and Norian stages of the Triassic. Acaenasuchus had a stratigraphic range of approximately 11.5 million years. Acaenasuchus is further categorized as one of the type fauna that belong to the Adamanian LVF, based on the fauna of the Blue Mesa Member of the Chinle Petrified Forest Formation of Arizona, where Acaenasuchus was initially discovered.
The Dockum is a Late Triassic geologic group found primarily on the Llano Estacado of western Texas and eastern New Mexico with minor exposures in southwestern Kansas, eastern Colorado, and Oklahoma panhandle. The Dockum reaches a maximum thickness of slightly over 650 m but is usually much thinner. The Dockum rests on an unconformity over the Anisian aged Anton Chico Formation.
In paleontology, biochronology is the correlation in time of biological events using fossils. In its strict sense, it refers to the use of assemblages of fossils that are not tied to stratigraphic sections. Collections of land mammal ages have been defined for every continent except Antarctica, and most are correlated with each other indirectly through known evolutionary lineages. A combination of argon–argon dating and magnetic stratigraphy allows a direct temporal comparison of terrestrial events with climate variations and mass extinctions.
Redondasaurus is an extinct genus or subgenus of phytosaur from the Late Triassic of the southwestern United States. It was named by Hunt & Lucas in 1993, and contains two species, R. gregorii and R. bermani. It is the youngest and most evolutionarily-advanced of the phytosaurs.
The Cooper Canyon Formation is a geological formation of Norian age in Texas and New Mexico. It is one of several formations encompassed by the Dockum Group.
Ebrachosaurus is an extinct genus of aetosaur. It was named after the town of Ebrach, Germany, near an outcrop of the Blasensandstein Formation where the original fossils have been found. Other Blasensandstein fauna include the temnospondyl Metoposaurus and the phytosaur Francosuchus. The genus has often been considered synonymous with the closely related Stagonolepis. The holotype specimen was lost during World War II, so its relationships within Stagonolepididae remain indeterminant.
Adamanasuchus is an extinct genus of aetosaur. Fossils have been found from several localities from the Chinle Group in Arizona and date back to the late Carnian stage of the Late Triassic. The locality from which it was named after also lends its name to the Adamanian LVF.
Leptosuchus is an extinct genus of leptosuchomorph phytosaur with a complex taxonomical history. Fossils have been found from the Dockum Group and lower Chinle Formation outcropping in Texas, New Mexico, and Arizona, USA, and date back to the Carnian stage of the Late Triassic.
Lucasuchus is an extinct genus of aetosaur. Fossils have been found from the Bull Canyon Formation of the Dockum Group outcropping in the Revuelto Creek locality in Quay County, New Mexico. All specimens date back to the Norian stage of the Late Triassic. The genus was named in 1995 after the American paleontologist Spencer G. Lucas.
Tecovasuchus is an extinct genus of aetosaur. It is known primarily from osteoderms found from the Tecovas Formation in Texas, which is Late Triassic in age, dating back to the lower Norian. Material is also known from several other localities of the Chinle Group in New Mexico and Arizona, such as older Carnian outcrops and younger Rhaetian outcrops. Specimens of Tecovasuchus have been collected from the Tecovas Formation, the Bluewater Creek Formation, and the Los Esteros Member of the Santa Rosa Formation.
The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to 208.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.
The Manda Formation is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassic, including some of the earliest dinosauromorph archosaurs. The formation is often considered to be Anisian in age according to general tetrapod biochronology hypotheses and correlations to the Cynognathus Assemblage Zone of South Africa. However, some recent studies cast doubt to this age, suggesting that parts deposits may actually be younger (Carnian) in age.
The Colorado City Formation is a Late Triassic geologic formation in the Dockum Group of Texas, United States. It has previously been known as the Iatan Member, Colorado City Member or 'Pre-Tecovas Horizon'.
The Bluewater Creek Formation is a geologic formation in west-central New Mexico. It preserves fossils dating back to the late Triassic period.
Vivaron is a genus of rauisuchid known from the Late Triassic Chinle Formation in New Mexico. It is the second rauisuchid known from the southwestern United States, and it highlights the wide biogeographic range similar rauisuchid taxa occupied during the Late Triassic across Pangaea, despite the varied faunal assemblages at different latitudes.