The grit, not grass hypothesis is an evolutionary hypothesis that explains the evolution of high-crowned teeth, particularly in New World mammals. The hypothesis is that the ingestion of gritty

soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...

is the primary driver of

hypsodont Hypsodont is a pattern of dentition characterized by with high crowns, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritty, fibrous material. The oppos ...

tooth development, not the

silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant f ...

-rich composition of

grass Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family (biology), family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and spe ...

, as was previously thought.

Traditional co-evolution hypothesis

Since the morphology of the hypsodont tooth is suited to a more abrasive diet, hypsodonty was thought to have evolved concurrently with the spread of

grasslands A grassland is an area where the vegetation is dominated by grasses (Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes such as clover, and other herbs. Grasslands occur ...

. During 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 ...

Period (145-66 million years ago), the Great Plains were covered by a shallow inland sea called the

Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...

which began to recede during the

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 ...

to the

Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

(65-55 million years ago), leaving behind thick marine deposits and relatively flat terrain. During the

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

and

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58niche for mammals, including many

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 that switched from browsing diets to grazing diets. Grass contains silica-rich phytoliths (abrasive granules), which wear away dental tissue more quickly. So the spread of grasslands was linked to the development of high-crowned (hypsodont) teeth in grazers.

Modern evolutionary hypothesis

Early evidence

In 2006 Strömberg examined the independent acquisition of high-crowned cheek teeth (hypsodonty) in several ungulate lineages (e.g.,

camelid Camelids are members of the biological family (biology), family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant taxon, extant members of this group are: dromedary, dromedary camels, Bactrian camels, wild Bac ...

equid Equidae (commonly known as the horse family) is the Taxonomy (biology), taxonomic Family (biology), family of Wild horse, horses and related animals, including Asinus, asses, zebra, zebras, and many extinct species known only from fossils. The fa ...

s, rhinoceroses) from the early to middle Miocene of North America, which had been classically linked to the spread of grasslands. She showed habitats dominated by C3 grasses (cool-season grasses) were established in the Central Great Plains by early late

Arikareean The Arikareean North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 30,600,000 to 20,800,000 years BP, a period of . It is usuall ...

(≥21.9 Million years ago), at least 4 million years prior to the emergence of hypsodonty in

Equidae Equidae (commonly known as the horse family) is the Taxonomy (biology), taxonomic Family (biology), family of Wild horse, horses and related animals, including Asinus, asses, zebra, zebras, and many extinct species known only from fossils. The fa ...

. In 2008 Mendoza and Palmqvist determined the relative importance of grass consumption and open habitat foraging in the development of hypsodont teeth using a dataset of 134 species of

artiodactyl Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other t ...

s and

perissodactyl Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...

s. The results suggested that high-crowned teeth represent are adapted for a particular feeding environment, not diet preference.

Morphology

More recent examination of mammalian teeth suggests that it is the open, gritty habitat and not the grass itself which is linked to diet changes. Analysis of dental microwear patterns of hypsodont notoungulates from the Late Oligocene Salla Beds of Bolivia showed shearing movements are associated with a diet rich in tough plants, not necessarily grasses. Hence the relationship between high-crowned mammals and the source of tooth wear in the fossil record may not be straightforward and the spread of grasslands in South America, traditionally linked with the development of notoungulate hypsodonty, was called into question.

Temporal discontinuity

Most importantly, evidence has shown, that the development of hypsodonty in Cenozoic mammals is out of sync with the flourishing of grasslands both in North America and South America, where grasslands spread 10 million years earlier. Observations of this temporal discontinuity between the spread grasslands and the development of hypsodonty in mammals is also supported by earlier evidence of hypsodonty in dinosaurs. For example, hadrosaurs, a group of herbivorous

dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

, likely grazed on low-lying vegetation and microwear patterns show that their diet contained an abrasive material, such as grit or silica. Grasses had evolved by the Late Cretaceous, but were not particularly common, so this study concluded that grass probably did not play a major component in the hadrosaur's diet.

Modern examples of hypsodonty

Hypsodonty is observed both in the fossil record and the modern world. It is a characteristic of large

clades In biology, a clade (), also known as a monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach to taxonomy ...

(equids) as well as subspecies level specialization. For example, the

Sumatran rhinoceros The Sumatran rhinoceros (''Dicerorhinus sumatrensis''), also known as the Sumatran rhino, hairy rhinoceros or Asian two-horned rhinoceros, is a rare member of the family Rhinocerotidae and one of five extant species of rhinoceros; it is the o ...

and the Javan rhinoceros both have brachydont, lophodont cheek teeth whereas the

Indian rhinoceros The Indian rhinoceros (''Rhinoceros unicornis''), also known as the greater one-horned rhinoceros, great Indian rhinoceros or Indian rhino, is a species of rhinoceros found in the Indian subcontinent. It is the second largest living rhinocer ...

, has hypsodont dentition. A mammal may have exclusively hypsodont molars or have a mix of dentitions. Hypsodont dentition is characterized by: * High-crowned

teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...

* A rough, flattish occlusal surface adapted for crushing and grinding *

Cementum Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament. Structure The cells of cementum are ...

both above and below the

gingiva The gums or gingiva (: gingivae) consist of the mucosal tissue that lies over the mandible and maxilla inside the mouth. Gum health and disease can have an effect on general health. Structure The gums are part of the soft tissue lining of the ...

l line * Enamel which covers the entire length of the body and likewise extends past the gum line * The cementum and the enamel invaginate into the thick layer of dentine

References

{{reflist, 2 Evolutionary biology Evolution of mammals Biological hypotheses