Musturzabalsuchus
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''Musturzabalsuchus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of allodaposuchid
eusuchia Eusuchia is a clade of crocodylomorphs that first appeared in the Early Cretaceous with '' Hylaeochampsa''. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-T extinction. Since the other two clades ...
n
crocodyliform Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
. The type and only
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
is ''Musturzabalsuchus buffetauti''.


Etymology

The generic name means "broadened rostrum crocodile", with "Musturzabal" meaning "broadened rostrum" in
Basque Basque may refer to: * Basques, an ethnic group of Spain and France * Basque language, their language Places * Basque Country (greater region), the homeland of the Basque people with parts in both Spain and France * Basque Country (autonomous co ...
and "suchus" meaning "crocodile" in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
. The type and only
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
is ''M. buffetauti'', named after the French paleoherpetologist
Eric Buffetaut The given name Eric, Erich, Erikk, Erik, Erick, or Eirik is derived from the Old Norse name ''Eiríkr'' (or ''Eríkr'' in Old East Norse due to monophthongization). The first element, ''ei-'' may be derived from the older Proto-Norse ''* ain ...
.


Discovery

The material first assigned to ''Musturzabalsuchus'' in 1997 has been found from the locality of Laño in
Condado de Treviño Condado de Treviño ("County of Treviño") is a municipality in the province of Burgos, autonomous community of Castile and León, Spain. This municipality and the geographically smaller La Puebla de Arganzón make up the enclave of Treviño. ...
, northern
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. Although dating back to the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
, the exact age of the strata in which material of ''Musturzabalsuchus'' occurs in the locality is not known: it is either Late
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...
or very Early
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
. Despite the unusually high quantity of remains belonging to the genus, the only skeletal elements known from ''Musturzabalsuchus'' are the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
and
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
. Some fragments of these bones have been found from the locality of Armuña in the province of
Segovia Segovia ( , , ) is a city in the autonomous community of Castile and León, Spain. It is the capital and most populated municipality of the Province of Segovia. Segovia is in the Inner Plateau (''Meseta central''), near the northern slopes of th ...
that were previously referred to an unnamed trematochampsid. Like the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
and
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
material found from Laño, these
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s, known collectively as UPUAM-502, are Campano-Maastrichtian in age. Another specimen (MHNM 10834.0) from the Fuvelian Lignites of
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
has been referred to ''Musturzabalsuchus'' in 1999. However, the characteristics used to assign the material to better-known specimens of ''Musturzabalsuchus'' from Spain were questioned in a later 2008 study. Material from ''Musturzabalsuchus'' has been found more recently from
Valencia Valencia ( va, València) is the capital of the Autonomous communities of Spain, autonomous community of Valencian Community, Valencia and the Municipalities of Spain, third-most populated municipality in Spain, with 791,413 inhabitants. It is ...
, Spain, being slightly older in age than specimens from other localities, dating back to the Early or Middle Campanian.


Classification

''Musturzabalsuchus'' was initially assigned to
Alligatoroidea Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more c ...
on the basis of several characters, including a lateral displacement of the foramen aereum of the
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
. Additionally, the enlarged fourth mandibular tooth occludes into a pit in the rostrum. Only the latter feature is present in the more derived alligatorids, and thus it is excluded from the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
. The pit that the fourth mandibular occludes into is placed posterior to the last
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
ry tooth, similar to what is seen in ''"
Diplocynodon ''Diplocynodon'' is an extinct genus of alligatoroid that lived during the Paleocene to Middle Miocene in Europe. It looked very similar to the modern caiman in that it was small and had bony armour scutes covering its neck, back, belly, and tail ...
" hantoniensis'', another early alligatoroid. A figure used in an 1869 study by P. Matheron of crocodylian remains from the Fuveau Lignites (''illustration'') pictured a mandible similar in profile to that of ''Musturzabalsuchus'', but was labeled as belonging to the crocodylian ''Crocodilus affuvelensis''. In 1997 it was suggested that due to this apparent similarity, along with the vagueness of descriptions in the 1869 paper and the loss of the
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
, that ''C. affuvelensis'' could be reassigned to ''Musturzabalsuchus'', although it was also acknowledged that the lack of posterior cranial material in the genus made detailed comparisons difficult. A recent 2008 study on Matheron's specimens concluded that they belonged to a new genus distinct from ''
Crocodylus ''Crocodylus'' is a genus of true crocodiles in the family Crocodylidae. Taxonomy The generic name, ''Crocodylus'', was proposed by Josephus Nicolaus Laurenti in 1768. ''Crocodylus'' contains 13–14 extant (living) species and 5 extinct species ...
'' and different from ''Musturzabalsuchus'', named ''
Massaliasuchus ''Massaliasuchus'' is an extinct monospecific genus of allodaposuchid eusuchian crocodyliform that is known from fossils found in Santonian–Campanian-age Upper Cretaceous rocks of southeastern France. Systematics ''Massaliasuchus'' was first ...
''. Recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies now recover ''Musturzabalsuchus'' as a member of
Allodaposuchidae Allodaposuchidae is an extinct clade of eusuchians that lived in Europe during the Late Cretaceous (Santonian-Maastrichtian). Systematics The type genus, ''Allodaposuchus'', was originally described in 1928 by Nopcsa from the Maastrichtian-age ...
, outside
Alligatoroidea Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more c ...
. However, ''Musturzabalsuchus'' is usually omitted from
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses due to the lack of sufficient anatomical information.


Paleobiogeography

''Musturzabalsuchus'' and ''
Acynodon ''Acynodon'' is an extinct genus of eusuchian crocodylomorph from the Late Cretaceous, with fossils found throughout Southern Europe. Classification The genus ''Acynodon'' contains three species: ''A. iberoccitanus'', ''A. adriaticus'', and ''A. ...
'', a contemporary crocodilian also common from Laño, are thought to have been closely related to Paleolaurasian alligatoroids. The latter genus is the only taxon not known from
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
to be related to the more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
Late Cretaceous tribodont alligatorids. It is clear that ''Musturzabalsuchus'' was
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, and was most likely restricted to Ibero-Armorica Island,Ibero-Armorica consisted of what is now the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
and southern France
as the genus is absent from Northern and Eastern European localities from which other alligatoroid fossils have been found.


References

{{Taxonbar, from=Q3755474 Neosuchians Late Cretaceous reptiles of Europe Late Cretaceous crocodylomorphs of Europe Prehistoric pseudosuchian genera