LACM 149371
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LACM 149371 (
Natural History Museum of Los Angeles County Nature is an inherent character or constitution, particularly of the ecosphere or the universe as a whole. In this general sense nature refers to the laws, elements and phenomena of the physical world, including life. Although humans are par ...
specimen 149371)Goin et al., 2004, p. 145 is an enigmatic fossil
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ian tooth from the
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 ...
(66 to 23 million years ago, mya) of Peru. It is from the
Santa Rosa Santa Rosa is the Italian, Portuguese, and Spanish name for Saint Rose. Santa Rosa may also refer to: Places Argentina *Santa Rosa, Mendoza, a city * Santa Rosa, Tinogasta, Catamarca * Santa Rosa, Valle Viejo, Catamarca *Santa Rosa, La Pampa * S ...
fossil site, which is of uncertain age but possibly late
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(55 to 34 mya) or
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
(34 to 23 mya). The tooth is poorly preserved and may have been degraded by acidic water or because it passed through a predator's digestive tract. Its largest dimension is 2.65 mm. It is triangular in shape and bears six cusps that surround the middle of the tooth, where there are three basins (fossae). Crests connects the cusps and separate the fossae. The microscopic structure of the enamel is poorly preserved. LACM 149371 was described in 2004 by Francisco Goin and colleagues, who tentatively interpreted the tooth as a left last upper molar. Although they saw similarities with South American
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s, some early
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s, and
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 ...
s, they believed the tooth was most likely of 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' ...
. Among gondwanatheres—a small and poorly known group otherwise known 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 ...
through
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
of some of 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 thought the Cretaceous Argentinian ''
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 ...
'' to be the most similar.


Discovery and context

LACM 149371 was discovered in 1998 at the
Santa Rosa Santa Rosa is the Italian, Portuguese, and Spanish name for Saint Rose. Santa Rosa may also refer to: Places Argentina *Santa Rosa, Mendoza, a city * Santa Rosa, Tinogasta, Catamarca * Santa Rosa, Valle Viejo, Catamarca *Santa Rosa, La Pampa * S ...
fossil site in the
Ucayali Region Ucayali () is an inland department and region of Peru. Located in the Amazon rainforest, its name is derived from the Ucayali River. Its capital is the city of Pucallpa. It is the second largest department in Peru, after Loreto, and it is s ...
of Peru. The Santa Rosa fauna also contains fossils of various unique species of
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s and
hystricognath The Hystricognathi are an infraorder of rodents, distinguished from other rodents by the bone structure of their skulls. The masseter medialis (a jaw muscle) passes partially through a hole below each eye socket (called the infraorbital foram ...
rodents, a possible
bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
, and some
notoungulate Notoungulata is an extinct order of ungulates that inhabited South America from the early Paleocene to the end of the Pleistocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms re ...
s. The fauna was published in a volume of the ''Science Series'' of the
Natural History Museum of Los Angeles County Nature is an inherent character or constitution, particularly of the ecosphere or the universe as a whole. In this general sense nature refers to the laws, elements and phenomena of the physical world, including life. Although humans are par ...
in 2004, which included a paper by Francisco Goin and colleagues that described and discussed LACM 149371. The age of the Santa Rosa fauna remains highly uncertain, as the
outcrop An outcrop or rocky outcrop is a visible exposure of bedrock or ancient superficial deposits on the surface of the Earth and other terrestrial planets. Features Outcrops do not cover the majority of the Earth's land surface because in most p ...
where the fossils were found cannot easily be placed in a known
stratigraphical Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostrati ...
unit, and the fossils are so distinct from other known fossil faunas that
biostratigraphy Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. "Biostratigraphy." ''Oxford Reference: Dictionary of Biology ...
cannot provide a precise estimate. In a summary of the 2004 volume, Kenneth Campbell tentatively referred Santa Rosa to the
Mustersan The Mustersan age is a period of geologic time (48.0–42.0 Mya (unit), Ma) within the Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Casamayoran and precedes th ...
South American Land Mammal Age The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as age ...
(SALMA), which he placed near the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
boundary, around 35 million years ago. However, Mario Vucetich and colleagues suggested in 2010 that the Santa Rosa fauna may be substantially later—perhaps as young as the
Deseadan The Deseadan () age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It follows the Tingu ...
SALMA (late Oligocene, around 25 million years ago). According to Campbell, the Santa Rosa mammals likely lived in a
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
habitat that contained rivers.


Description

LACM 149371 is a poorly preserved molar-like tooth that largely lacks a recognizable enamel surface and shows many small grooves and holes on the
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, parti ...
surface.Goin et al., 2004, p. 146 This suggests the tooth may have been chemically degraded, perhaps by acidic water or because it passed through the digestive tract of a predator.Goin et al., 2004, p. 149 The roots are broken off, but remaining
pulp Pulp may refer to: * Pulp (fruit), the inner flesh of fruit * Pulp (band), an English rock band Engineering * Pulp (paper), the fibrous material used to make paper * Dissolving pulp, highly purified cellulose used in fibre and film manufacture ...
cavities suggest the presence of four main roots, which are partially joined into two pairs. A smaller pulp cavity between those roots suggests the likely presence of a fifth root and a slight depression in the tooth may represent another root. The crown of the tooth is triangular and contains six cusps, connected by low crests, that surround two prominent, low-lying fossae (basins) and a third, smaller fossa. Because of the complexity of the crown, Goin and colleagues interpreted it as a molar; because of the number of roots, the arrangement of the cusps, and the shape of the tooth, as an upper molar; and because it tapers towards the end, as a last molar. One side, the longest, is flat and low compared to the others, suggesting it is the labial (outer) face. This would imply that the tooth is from the left jaw. Under this interpretation, the length of the tooth is 2.65 mm, width is 2.20 mm, height at the labial side is 1.05 mm, and height at the lingual side is 1.30 mm. For convenience, Goin and colleagues designated the six cusps as A through F: A on the front labial corner of the tooth; B on the labial face; C on the back corner; D on the lingual (inner) face; E on the front lingual corner; and F on the front face. The large front fossa is located between cusps A, B, D, E, and F; the smaller intermediate fossa is between cusps B and D; and the much smaller back fossa is just in front of cusp C. All three are nearly round. Cusp A, the largest cusp, is triangular in shape and is separated from the smaller, rounded B by a deep valley; a low crest connects the two cusps further lingually, separating the valley from the front fossa.Goin et al., 2004, p. 147 At its back, B connects to a long crest that reaches the back fossa and behind it the small cusp C, which has a groove on its labial side. A valley separates it from cusp D. D itself is crest-shaped and forms the lingual wall of the intermediate fossa; it is described as "very odd", and may in fact consist of two fused, triangular cusps.Goin et al., 2004, p. 148 A crest issuing from D separates the back from the intermediate fossa, and another, larger crest separates the front from the back fossa and nearly reaches cusp B. Cusp E is triangular and separated from cusps F and D by valleys, which are bordered internally by crests connecting the cusps. F is rounded. The microstructure of 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 not clearly recognizable, evidently because the tooth is degraded, though structures resembling
enamel prism An enamel prism, or enamel rod, is the basic unit of tooth enamel. Measuring 3-6 μm in diameter in primates, enamel prism are tightly packed hydroxyapatite crystals structures. The hydroxyapatite crystals are hexagonal in shape, providing rigidit ...
s (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) and Hunter-Schreger bands are recognizable.


Identity

Because of the complexity of the crown, Goin and colleagues identified the tooth as a mammal; although some non-mammalian groups, like
crocodylia Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
ns, may have complex teeth, none approach the level of complexity seen in LACM 149371. They could find no resemblance to
australosphenida The Australosphenida are a clade of mammals, containing mammals with tribosphenic molars, known from the Jurassic to Mid-Cretaceous of Gondwana. Although they have often been suggested to have acquired tribosphenic molars independently from those ...
ns including
monotreme Monotremes () are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...
s,
metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as wel ...
ns including
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s,
xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a superorder and major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and ...
ns, and some related groups.Goin et al., 2004, p. 151 They did see some general resemblances to the upper
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 ...
s of the early South American
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s, but the cusp arrangement is different from that of any ungulate.Goin et al., 2004, pp. 151–152 There are also some resemblances to the early
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s '' Ivanantonia'' from Asia and '' Nonomys'' from North America, but ''Ivanantonia'' has a central groove and lacks fossae, and ''Nonomys'' has a prominent cingulum (shelf) at the edges of the tooth and also lacks the fossae of LACM 149371.Goin et al., 2004, p. 152 The tooth resembles
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 ...
s—a large group of extinct mammals with many-cusped teeth—in the shapes of the valleys and crests, but multituberculates lack fossae and usually have quadrangular teeth with two longitudinal rows of cusps separated by a central valley. In the same features, LACM 149371 resembles
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' ...
s, a small and enigmatic group of mammals 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 ...
through
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
of the southern (
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) continents that may be related to multituberculates. In particular, ''
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 ...
'' from the late
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 has similarly formed cusps and also has crests that connect the cusps to the center of the tooth. However, the upper molars are unknown, and the low-crowned teeth of ''Ferugliotherium'' lack deep fossae. Members of the higher-crowned gondwanathere family
Sudamericidae Sudamericidae is a family of gondwanathere mammals that lived during the late Cretaceous to Miocene. Its members include '' Lavanify'' and '' Vintana'' from the Cretaceous of Madagascar, '' Bharattherium'' (=''Dakshina'') from the Cretaceous of I ...
do have fossae. Goin and colleagues conclude that LACM 149371 most likely represents a member of the gondwanathere family
Ferugliotheriidae Ferugliotheriidae is one of three known families in the order Gondwanatheria, an enigmatic group of extinct mammals. Gondwanatheres have been classified as a group of uncertain affinities or as members of Multituberculata, a major extinct mammali ...
; if so, it would be among the youngest known gondwanatheres.


References


Bibliography

* Campbell, K.E., Jr. 2004. The Santa Rosa local fauna: A summary. Science Series, Natural History Museum of Los Angeles County 40:155–163. * Goin, F.J., Vieytes, E.C., Vucetich, M.G., Carlini, A.A. and Bond, M. 2004. Enigmatic mammal from the Paleogene of Perú. Science Series, Natural History Museum of Los Angeles County 40:145–153. * Vucetich, M.G., Vieytes, E.C., Pérez, M.E. and Carloni, A.A. 2010
The rodents from La Cantera and the early evolution of caviomorphs in South America
pp. 193–205 in Madden, R.H., Carlini, A.A., Vucetich, M.G. and Kay, R.F. (eds.). The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge University Press, 458 pp. Paleogene mammals of South America Mammal fossils Paleogene fossil record Paleogene Peru Fossils of Peru Taxa named by Francisco J. Goin {{Good article Undescribed vertebrate species