Argentodites Coloniensis
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''Argentodites'' is a possible
multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
mammal from the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
of Argentina. The single species, ''Argentodites coloniensis'', is known from a single blade-like fourth lower
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
(p4) from the
La Colonia Formation The La Colonia Formation is a geological Formation (geology), formation in Argentina whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel et al., 2004, pp.600- ...
, which is mostly or entirely
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
(latest Cretaceous) in age. The p4 is 4.15 mm long and bears eight cusps on its upper margin and long associated ridges on both sides. The enamel consists of prisms that are completely or partly surrounded by a sheath and that are on average 6.57 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
apart.
Zofia Kielan-Jaworowska Zofia Emilia Kielan-Jaworowska (25 April 1925 – 13 March 2015) was a Polish paleobiologist. In the mid-1960s, she led a series of Polish-Mongolian paleontological expeditions to the Gobi Desert. She was the first woman to serve on the executi ...
, who described and named the fossil in 2007, regarded it as a multituberculate, perhaps a
cimolodonta Cimolodonta is a clade of Multituberculata, multituberculate mammals that lived from the Cretaceous to the Eocene. They probably lived something of a rodent-like existence until their ecological niche was assumed by true rodents. The more basal ...
n—and thus, a member of a mostly
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
n (northern) group and an immigrant to Argentina from North America—on the basis of the shape of the tooth and features of its enamel. In 2009, however, two teams argued that ''Argentodites'' may in fact be close to or identical with ''
Ferugliotherium ''Ferugliotherium'' is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period (Late Cretaceous; around 70 million years ago) of Argentina. It contains a single species, ''Ferugliotherium windhaus ...
'', a member of the small
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n (southern) group
Gondwanatheria Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa, and Antarctica during the Upper Cretaceous through the Miocene (and possibly much earlier, if '' Allostaffia' ...
; although their relationships are disputed, gondwanatheres may themselves be multituberculates.


Discovery and context

''Argentodites'' is known from a single
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
tooth, MPEF 604, in the collections of the Museo Paleontológico "Egidio Feruglio" in
Trelew Trelew (, from "town" and the name of the founder, Lewis Jones) is a city in the eastern part of the Chubut Province of Argentina, 21km away from the coast. Located in Patagonia, the city is the largest and most populous in the low valley of the ...
, Argentina. It is from the middle part of the
La Colonia Formation The La Colonia Formation is a geological Formation (geology), formation in Argentina whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel et al., 2004, pp.600- ...
of
Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...
, Argentina, which is
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
(
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
and perhaps partly
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. Campa ...
) in age.Kielan-Jaworowska et al., 2007, p. 258. The premolar was described in 2007 by
Zofia Kielan-Jaworowska Zofia Emilia Kielan-Jaworowska (25 April 1925 – 13 March 2015) was a Polish paleobiologist. In the mid-1960s, she led a series of Polish-Mongolian paleontological expeditions to the Gobi Desert. She was the first woman to serve on the executi ...
and colleagues as a new
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, ''Argentodites coloniensis''. The generic name, ''Argentodites'', combines "Argentina" with the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''hodites'' "traveler", in reference to the animal's presumed migration from North America to Argentina, and the
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''coloniensis'', refers to the La Colonia Formation.Kielan-Jaworowska et al., 2007, p. 257.


Description

The single known example of ''Argentodites'' is a blade-like fourth lower premolar (p4). It has a length of 4.15 mm, height of 2.10 mm, and width of 1.35 mm. The crown is nearly complete, but the roots are largely missing. Kielan-Jaworowska considered two possible orientations of the tooth—one with the back margin nearly vertically, and the other with the margin inclined backward—but preferred the former, which made for more natural placement of the roots. Although the left and right sides of the tooth are almost identical, they believed the tooth is most likely a left p4, as this would make the lingual (inner) side the more convex one, as is usual in the p4 of
cimolodonta Cimolodonta is a clade of Multituberculata, multituberculate mammals that lived from the Cretaceous to the Eocene. They probably lived something of a rodent-like existence until their ecological niche was assumed by true rodents. The more basal ...
n multituberculates with a large p4. The front root is larger than the back one. In side view, the upper and back margins are straight, but the front margin is convex. There are eight cusps arranged in a row on the upper margin, the first of which is located about one-third of the tooth's length from the front margin. Only the last is slightly worn, indicating that the tooth is from a young animal.Kielan-Jaworowska et al., 2007, p. 260. Long ridges extend from the cusps diagonally toward the front on both the lingual and labial (outer) sides of the tooth. The first seven ridges on both sides are connected to the respective cusps, but the eighth ridges do not quite reach their cusp. On both sides of the tooth, there is also a small ridge behind the eighth ridge that extends to the back margin; these ridges are called the posterolabial and posterolingual ridge. An even smaller ridge is located below the ridge on the lingual side. The
tooth enamel Tooth enamel is one of the four major Tissue (biology), tissues that make up the tooth in humans and many animals, including some species of fish. It makes up the normally visible part of the tooth, covering the Crown (tooth), crown. The other ...
is well preserved over most of the tooth. It consists of prisms—bundles of
hydroxyapatite Hydroxyapatite (International Mineralogical Association, IMA name: hydroxylapatite) (Hap, HAp, or HA) is a naturally occurring mineral form of calcium apatite with the Chemical formula, formula , often written to denote that the Crystal struc ...
crystals—with an average diameter of 3.8 μm. Most are entirely surrounded by a sheath, but in some the sheath is open. The prisms are slightly curved toward the outer surface of the tooth. Between the prisms, and oriented at an angle of about 45° to them, are crystals of interprismatic material. On average, the prisms are 6.57 μm apart, so that there are about 27,247 per mm2.


Identity

Kielan-Jaworowska and colleagues identified ''Argentodites'' as a
multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
, a diverse fossil group from the northern continents (
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
) that is also known from a few questionable or fragmentary records from the southern continents (
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
). They tentatively allocated it to the multituberculate subgroup Cimolodonta on the basis of its enamel microstructure, which particularly recalls
Ptilodontoidea Ptilodontoidea is a group of extinct mammals from the Northern Hemisphere. They were generally small, somewhat rodent-like creatures of the extinct order Multituberculata. Some of these genera have a great many species, though remains are gene ...
(one of the subgroups of Cimolodonta), and the convex front margin of the tooth. On the other hand, the straight back margin resembles the condition in the other major subgroup of multituberculates, the "
plagiaulacida Plagiaulacida is a group of extinct Multituberculata, multituberculate mammals. Multituberculates were among the most common mammals of the Mesozoic, "the age of the dinosaurs". Plagiaulacids are a paraphyletic grouping, containing all multituber ...
ns", and it does not have the highly vaulted upper margin of p4 that is characteristic of Ptilodontoidea. They regarded ''Argentodites'' as distinct from MACN-RN 975, a fragmentary fossil
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
(lower jaw) with p4 from the Late Cretaceous
Los Alamitos Formation The Los Alamitos Formation is a geological Formation (geology), formation of the North Patagonian Massif in Rio Negro Province, northwestern Patagonia, Argentina, whose strata date back to the Late Cretaceous (Campanian, Late Campanian to Maastri ...
of Argentina, which they identified as a "plagiaulacidan" multituberculate,Kielan-Jaworowska et al., 2007, p. 262. because the p4 of MACN-RN 975 is rectangular in shape and has fewer cusps. They believed ''Argentodites'' to represent an immigrant from North America, but could not determine precisely in which part of the Cretaceous it reached South America. In a 2009 paper on the affinities of Gondwanatheria, Yamila Gurovich and Robin Beck argue that the difference in shape between MACN-RN 975 and ''Argentodites'' is due to extensive wear on the former specimen; they write that the parts of the p4 that are not worn are virtually identical to the equivalents parts of the ''Argentodites'' p4. In addition, MACN-RN 975 is said to have as many ridges as ''Argentodites'' and to be of approximately similar size—about 15% larger. They allocate MACN-RN 975 to the gondwanathere ''
Ferugliotherium ''Ferugliotherium'' is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period (Late Cretaceous; around 70 million years ago) of Argentina. It contains a single species, ''Ferugliotherium windhaus ...
'' and consequently, they argue that ''Argentodites'' most likely represents either ''Ferugliotherium'' or some related species.Gurovich and Beck, 2009, p. 32. In the same year, Guillermo Rougier and colleagues also suggested ferugliotherian affinities of ''Argentodites'' in a paper on the mammals of the
Allen Formation The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian.Salgado et al., 2007 Dinosaur remains are among the fossils that have been recovered from the forma ...
, another Cretaceous rock unit of Argentina. ''Ferugliotherium'' is a
gondwanathere Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa, and Antarctica during the Upper Cretaceous through the Miocene (and possibly much earlier, if '' Allostaffia' ...
from Late Cretaceous Argentinean deposits. Gondwanatheres are a small and enigmatic group from the late Cretaceous and
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
of South America, Antarctica, Madagascar, India, and perhaps Tanzania. Although the evolutionary affinities of gondwanatheres are controversial, both teams that identified ''Argentodites'' as gondwanathere believe gondwanatheres are likely themselves multituberculates or closely related to them.Gurovich and Beck, 2009, p. 25; Rougier et al., 2009, p. 233.


References


Bibliography

* * * * * {{Good article Gondwanatheria Maastrichtian life Late Cretaceous mammals of South America Cretaceous Argentina Fossils of Argentina Cañadón Asfalto Basin Fossil taxa described in 2007 Taxa named by Zofia Kielan-Jaworowska Taxa named by Edgardo Ortiz−Jaureguizar Taxa named by Carolina Vieytes Taxa named by Rosendo Pascual Taxa named by Francisco J. Goin