Grit, not grass hypothesis
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 is the primary driver of hypsodont tooth development, not the silica-rich composition of grass, as was previously thought.[1]
Traditional Coevolution 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. During the Cretaceous Period (145-66 million years ago), the Great Plains were covered by a shallow inland sea called the Western Interior Seaway which began to recede during the Late Cretaceous to the Paleocene (65-55 million years ago), leaving behind thick marine deposits and a relatively flat terrain. During the Miocene and Pliocene epochs (25 million years), the continental climate became favorable to the evolution of grasslands. Existing forest biomes declined and grasslands became much more widespread. The grasslands provided a new niche for mammals, including many ungulates 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.[citation needed]
Modern Evolutionary Hypothesis
Early Evidence
In 2006 Strömberg examined the independent acquisition of high-crowned cheek teeth (hypsodonty) in several ungulate lineages (e.g., camelids, equids, 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 (≥21.9 Million years ago), at least 4 million years prior to the emergence of hypsodonty in Equidae.[2] 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 artiodactyls and perissodactyls. The results suggested that high-crowned teeth represent are adapted for a particular feeding environment, not diet preference. [3]
Coevolution
Even more recent research has led to a reexamination of the traditional coevolution hypothesis for several reasons. Broadly, the existence of coevolution has become contentious,[4] and more specifically studies on morphology, biogeography, and analogous taxa and have shown a mismatch between the timing in changes in teeth morphology and grassland proliferation. Though coevolution is known to operate on small scales, there is little evidence it to support coevolutionary shifts on large scales.[4]
Morphology
Further, examination of mammalian teeth suggests that it is the open, gritty habitat and not the grass itself which is linked to diet changes.[3][5] 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. That the relationship between high-crowned mammals and the source of tooth wear in the fossil record may not be straightforward, hence the spread of grasslands in South America, traditionally linked with the development of notoungulate hypsodonty, is in question.[5]
Observations of hypsodonty development in Cenozoic mammals is also supported by the evolution of hypsodonty in other groups. For example, hadrosaurs, a group of herbivorous dinosaurs, 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.[6]
Temporal Discontinuity
Most importantly, evidence has shown, that the development of hypsodonty is out of sync with the flourishing of grasslands both in North America and South America, where grasslands spread 10 millions years earlier.[1][5]
Hypsodonty
Hypsodonty is observed both in the fossil record and the modern world. It is a characteristic of large clades (equids) as well as subspecies level specialization. For example, the Sumatran rhinoceros and the Javan rhinoceros both have brachydont, lophodont cheek teeth whereas the Indian rhinoceros, has hypsodont dentition. A mammal may have exclusively hypsodont molars or have a mix of dentitions. Hypsodont dentition is characterized by:[7][8]
- High-crowned teeth
- A rough, flattish occlusal surface adapted for crushing and grinding
- Cementum both above and below the gingival 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
- ^ a b Jardine, Phillip E.; Janis, Christine M.; Sahney, Sarda; Benton, Michael J. (2012). "Grit not grass: Concordant patterns of early origin of hypsodonty in Great Plains ungulates and Glires". Palaeogeography, Palaeoclimatology, Palaeoecology. 365–366: 1–10. doi:10.1016/j.palaeo.2012.09.001.
- ^ Caroline A. E. Strömberg (2006). "Evolution of hypsodonty in equids: testing a hypothesis of adaptation" (PDF). Paleobiology. 32: 236–258. doi:10.1666/0094-8373(2006)32[236:eohiet]2.0.co;2.
- ^ a b Mendoza, M.; Palmqvist, P. (February 2008). "Hypsodonty in ungulates: an adaptation for grass consumption or for foraging in open habitat?" (PDF). Journal of Zoology. 274 (2): 134–142. doi:10.1111/j.1469-7998.2007.00365.x.
- ^ a b Sahney, S., Benton, M.J. and Ferry, P.A. (2010). "Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land" (PDF). Biology Letters. 6 (4): 544–547. doi:10.1098/rsbl.2009.1024. PMC 2936204. PMID 20106856.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c Billet, Blondel, and Muizon (2009), "Dental microwear analysis of notoungulates (Mammalia) from Salla (Late Oligocene, Bolivia) and discussion on their precocious hypsodonty", Palaeogeography, Palaeoclimatology, Palaeoecology, doi:10.1016/j.palaeo.2009.01.004
{{citation}}: CS1 maint: multiple names: authors list (link) - ^ "Hadrosaur chowdown — grind, grind, grind", Associated Press, 2009
- ^ Flynn, John J.; Wyss, André R.; Charrier, Reynaldo (May 2007). "South America's Missing Mammals". Scientific American. 296 (5): 68–75. doi:10.1038/scientificamerican0507-68.
- ^ Kwan, Paul W.L. (2007). "Digestive system I" (PDF). Tufts University. Retrieved May 2013.
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