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Grit, not grass hypothesis

From Wikipedia, the free encyclopedia

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]

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Transcription

You remember back in the days right after the Permian-Triassic extinction event, when that giant flaming asteroid and those methane explosions killed almost all of the organisms on the planet? No, of course you don't because that happened 252 million years ago, and mammals weren't a thing yet. But that's kind of the point of this episode. That asteroid was a...a disturbance to the ecology of the planet. The flora, fauna and soils were largely wiped out, leaving a blank canvas for the organisms that survived, and there really weren't all that many of them, to fill in as they could. What happened after the Permian-Triassic "disturbance" is a dramatic example of ecological succession, how the makeup of a community changes over time, starting from, like, the day after a disturbance. Just, usually, the disturbance is a little less disturbing. The study of how ecological communities change doesn't just look at huge-long periods of time, or the effects of some apocalypse. Succession can easily happen over a season in a park, or in just a few days in a patch of land as small as your garden. And, this might come as a surprise, but disturbances that shake up the status quo within a community actually serve to make that community better in the long run. Because much like life, and the entire universe, succession is all about change. And change is how a universe full of nothing but hydrogen came to include a planet full of life. Disturbances happen in ecosystems all the time, every day: a wildfire, a flood, a windstorm. After these unpredictable events, ecologists kept seeing predictable, even orderly changes in the ecosystem. How life deals with these disturbances is an important key to understanding ecosystems. First, let's note that a tree falling in the forest and a comet falling in the forest, while both disturbances, are different levels of disturbance. Likewise there are a couple of different types of succession. The first type, the one that happens after the asteroid hits or the glacier plows over the landscape or the forest fire-slash-volcano burns the verdant ecosystem into pure desolation, that's called a primary succession: when organisms populate an area for the first time. The jumping off point for primary succession is your basic, lifeless, post- apocalyptic wasteland. You're probably thinking, that place sounds terrible! Who would ever want to live there? Well, actually, there is one tremendous advantage of to desolate wastelands...no competition. A lot of organisms don't mind settling down in the more inhospitable nooks and crannies of the planet. These pioneer species are often prokaryotes or protists, followed by nonvascular plants, then maybe some extra super hardy vascular plants. There are tons of organisms that make their living colonizing dead places. It's their thing. Like before the Permian-Triassic extinction, there were these dense forests of gymnosperms, probably full of species a lot like the conifers, gingkos and cycads we still have today. But after the asteroid hit, the big forests died and were replaced by lycophytes, simpler vascular plants like the now-extinct scale trees and today's club mosses. While they might have had a hard time competing with the more complicated plants during the good times. The rest of the Paleozoic flora barely survived extinction, of all the dozens of species of gingko that were around back then, only one still exists, completely genetically isolated, a living fossil. It's important to remember that when we talk about primary ecological succession, we're talking about plants, pretty much exclusively. Because plants rule the world, remember? Without plants, the animals in a community don't stand a chance, and primary successional species are often plants that have windborne seeds, like lycophytes, or mosses and lichens that have spores that blow in and colonize the area. And the outcome of a primary successional landscape is to build, or rebuild, soils, which develop over time as the mosses, grasses and tiny little plants grow, die and decompose. Once the soils are ready, slightly bigger plants can move in, at which point, we move onto secondary succession. And then it's game on: a whole redwood forest could develop out of that. But primary succession takes a long, long time: like hundreds, maybe thousands of years in some places. In fact, the recovery of these big gymnosperm forests after the Permian-Triassic extinction event took about 4 or 5 million years. Dirt may seem unglamorous to you, but it is alive and beautiful and complicated, and making good soil takes time! Now, secondary succession isn't just the next act after primary succession has made a place livable after some disaster. It's usually the first response after a smaller disturbance like a flood or a little fire has knocked back the plants that have been ruling the roost for a while. Even a disturbance as small as a tree crashing down in the woods can make a tiny patch of forest more like it was 50 years ago, before that one tree got so huge and shady: In that tiny area, there will suddenly be a different microclimate than in the rest of the forest, which might have more sunlight, slightly higher temperatures, less protection from weather, etc. And just like every other ecosystem on earth, this tiny patch of forest will be affected by temperature and precipitation the most, which will be different in different parts of the forest. So, as a result of the fallen tree, the soils will become different, the mix of plants will become different, and different animals will want to do business there because that little niche suits their needs better than other little niches. So the question becomes, when does succession stop and things get back to normal? Never. Because change doesn't end. Change is the only constant people...you know who said that? Heraclitus...in 500 BC. So it's been true since at least then. Consider it a lesson in life. And as ideas in ecology go, it's actually a pretty new way of looking at things. See, back in the early 20th century, ecologists noticed the tendency of communities to morph over time. But they also saw succession in terms of a community changing until it ultimately ended in what they called a climax community, which would have a predictable assemblage of species that would remain stable until the next big disturbance. Well, maybe that's what seemed to be happening, but ecological succession is actually a lot more complicated than that. For starters, there's a little thing called stochasticity or randomness which prevents us from ever knowing exactly what a community is going to look like 100 years after a disturbance. Stochasticity is basically your element of unpredictable variability in anything. So, you can predict with some accuracy what plants are going to take over a glacial moraine after the ice has receded, because the seeds of some colonizer species typically make it there first. But unpredictable things like weather conditions during the early stages of succession can end up favoring another species. The point is, scientists' attempts to predict what a community ends up looking like in 100 years should always be thought of as probabilities, not certainties. Another difficulty with the whole model of a climax community has to do with the idea of an ecosystem eventually stabilizing. That word, "stable"? Whenever it's used in a sentence that also includes the word "ecology", you can pretty much be sure it's being used wrong. Because stability never happens. There are always disturbances happening all the time, in every ecosystem. A small portion of the forest might burn, a windstorm might take out a bunch of trees, some yeehaw might rent himself a backhoe one weekend and clear himself a little patch of heaven on the mountain beside his house because he's got nothing better to do. Who knows! Stuff happens. So instead of ending in some fixed, stable climax community, we now know that an ecosystem is in later successional stages if it has high biodiversity. Lots and lots of biodiversity. The only way biodiversity could be high is if there are tons of little niches for all those species to fit into. And the only way there could be that many niches is if, instead of a single community, an ecosystem was actually made up of thousands of tiny communities, a mosaic of habitats where specific communities of different organisms lived. Such mosaics of niches are created by disturbances over time, with everything always changing here and there. But it's important that these disturbances be of the right kind, and the right scale. Because it turns out that the kind of disturbances that have the greatest effect on biodiversity are the most moderate disturbances. When ecologists figured this out, they decided to call it the Intermediate Disturbance Hypothesis. Because, it hypothesizes that intermediate disturbances, not too big and not too little, are ideal. See, just a little disturbance, like a falling tree or something, isn't enough to really change the game. On the other hand, a really severe disturbance, like getting covered with lava, would take the community all the way back to asteroid wipe-out- level primary succession. But every nice mid-level disturbance creates its own habitat at its own stage of succession with its own unique niches. More niches means more biodiversity, and more biodiversity means more stability and healthier ecosystems. Even if two disturbances happen in two different areas with roughly the same climate at the same time, the stochastic nature of ecosystems mean that the two areas might recover in completely different ways, leading to even more niches and more biodiversity! Now, this does not mean that you should go rent a backhoe tomorrow and cut a swath into the wilderness. It just suggesting that medium-level of disturbance is natural and normal and good for an ecosystem. Keeps everybody on their toes. And, like I said, disturbance happens. And by and large we should let it happen. This, too, is a relatively new idea in ecology. In fact, for most of the history of public land management in the U.S., great swaths of forests were not allowed to burn. People considered the "purpose" of forests to be wood production. And you don't want to burn down some trees that are gonna make you a bunch of money. But because of the lack of intermediate disturbances over a long period of time, we ended up with catastrophic fires like the one that torched Yellowstone National Park back in 1988. A single lightning strike totally annihilated almost 800,000 acres of public forest because the ecosystem hadn't been allowed to indulge in a nice leisurely burn every now and then. But now those forests have undergone more than 20 years of succession, and some parts have even re-burned at a more intermediate level, creating a nice, high-biodiversity mosaic of habitats. And it's gorgeous, you should come visit it sometime. And that is ecological succession for you... how destruction and disturbance lead to beauty and diversity. Just remember what my main man Heraclitus said and you'll be good: the only constant is change. Thank you for watching this episode of Crash Course Ecology. And thank you to everyone who helped us put this episode together. If you want to review any of the concepts we studied today, there's a table of contents over there. And if you have any questions, ideas or comments, we're on Facebook, Twitter and of course, down in the comments below. We'll see you next time.

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. 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 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.[2]

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.[3] 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. [4]

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. [1][4][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. 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.[5]

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.[1][5] 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, 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]

Modern Examples of 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

  1. ^ a b c 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. Bibcode:2012PPP...365....1J. doi:10.1016/j.palaeo.2012.09.001.
  2. ^ Stirton, R. A. (1947). "Observations on Evolutionary Rates in Hypsodonty". Evolution. 1 (1–2): 32–41. doi:10.1111/j.1558-5646.1947.tb02711.x. S2CID 87532435.
  3. ^ Caroline A. E. Strömberg (2006). "Evolution of hypsodonty in equids: testing a hypothesis of adaptation" (PDF). Paleobiology. 32 (2): 236–258. doi:10.1666/0094-8373(2006)32[236:eohiet]2.0.co;2. S2CID 12338144.
  4. ^ 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.
  5. ^ 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, 274 (1–2): 114–124, Bibcode:2009PPP...274..114B, doi:10.1016/j.palaeo.2009.01.004{{citation}}: CS1 maint: multiple names: authors list (link)
  6. ^ "Hadrosaur chowdown — grind, grind, grind", Associated Press, 2009
  7. ^ Flynn, John J.; Wyss, André R.; Charrier, Reynaldo (May 2007). "South America's Missing Mammals". Scientific American. 296 (5): 68–75. Bibcode:2007SciAm.296e..68F. doi:10.1038/scientificamerican0507-68. PMID 17500416.
  8. ^ Kwan, Paul W.L. (2007). "Digestive system I" (PDF). Tufts University.
This page was last edited on 12 September 2023, at 21:46
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